EvolutionPDF generated using the open source mwlib toolkit. See http://code.pediapress.com/ for more information. PDF generated at: Wed, 27 Oct 2010 05:33:11 UTC Contents Articles Overview Introduction to evolution Evolution Evolution as theory and fact Evolutionary history of life Timeline of evolution 1 1 16 47 53 83 92 92 114 121 122 126 154 158 160 162 172 172 176 179 183 212 212 223 233 236 246 260 268 268 History History of evolutionary thought Lamarckism Saltationism Orthogenesis On the Origin of Species Darwinism The Genetical Theory of Natural Selection Neo-Darwinism Modern evolutionary synthesis Base concepts Heredity Fitness Common descent Evidence of common descent Mechanisms Adaptation Genetic drift Gene flow Mutation Natural selection Speciation Phylogenetics Phylogenetics Cladistics Cladogram Molecular phylogenetics 274 284 287 290 290 296 299 316 332 332 359 384 406 436 Fields Evolutionary developmental biology Molecular evolution Human evolution Evolutionary psychology Controversy and social impacts Creation–evolution controversy Objections to evolution Creationism Intelligent design Social effect of evolutionary theory References Article Sources and Contributors Image Sources, Licenses and Contributors 444 454 Article Licenses License 458 1 Overview Introduction to evolution Artist's depiction of a T. rex. Natural selection does not lead to perfection; dramatic changes in the environment often lead to mass extinctions, as in the case of the dinosaurs nearly 65 million years ago. Overview Life forms reproduce to make offspring. The offspring differs from the parent in minor random ways. If the differences are helpful, the offspring is more likely to survive and reproduce. This means that more offspring in the next generation will have the helpful difference. These differences accumulate resulting in changes within the population. Over time, populations branch off to become new species as they become geographically separated and genetically isolated. This process is responsible for the many diverse life forms in the world today. Haeckel's Paleontological Tree of Vertebrates (c. 1879). The evolutionary history of species has been described as a "tree", with many branches arising from a single trunk. While Haeckel's tree is somewhat outdated, it illustrates clearly the principles that more complex modern reconstructions can obscure. Evolution is the process of change in all forms of life over generations, and evolutionary biology is the study of how evolution occurs. The biodiversity of life evolves by means of natural selection, mutations and genetic drift. The process of natural selection is based on three conditions. First, every individual is supplied with hereditary material in the form of genes that are received from their parents; then, passed on to their offspring. Second, organisms tend to produce more offspring than the environment can support. Third, there are variations among offspring as a consequence of either the random introduction of new genes via mutations or reshuffling of existing genes during sexual reproduction.[1] [2] [3] Natural selection will occur when these facts of nature (heredity, overproduction of offspring, and variation) hold true. Natural selection means individuals do not have equal chances of reproductive success. As a consequence, some individuals produce more offspring and thus have a higher degree of fitness. Traits that ensure organisms are Introduction to evolution better adapted to their living conditions become more common in descendant populations.[2] [3] For this reason, populations will never remain exactly the same over successive generations. The forces of evolution are most evident when populations become isolated, either through geographic distance or by mechanisms that prevent genetic exchange. Over time, isolated populations can branch off into new species.[4] [5] Random genetic drift describes another natural process that regulates the evolution of minor mutations in the genes, leading to changes in allele frequencies over time. These smaller mutations occur with regular frequency in and among populations. The vast majority of genetic mutations neither assist, change the appearance of, nor bring harm to individuals. These mutated genes are neutrally sorted among populations and survive across generations by chance alone. When species migrate they carry different genetic varieties to different places. When organisms mate they exchange genetic material and new individuals are born. The outward expression of each unique genetic mixture in different environments, the phenotype, comprises a diversity of traits that can be measured and observed as individuals grow and develop. In contrast to genetic drift, natural selection is not a random process because it acts on traits that become adaptive for their functional utilities that are necessary for survival.[6] Natural selection and random genetic drift are forever constant and dynamic parts of life. More than 99.9% of all species have become extinct since life began over 3.5 billion years ago. Evolution is more death than survival and over time this has shaped the branching structure in the tree of life.[7] The modern understanding of evolution began with the 1859 publication of Charles Darwin's On the Origin of Species. In addition, Gregor Mendel's work with plants helped to explain the hereditary patterns of genetics.[8] Fossil discoveries in paleontology, advances in population genetics and a global network of scientific research have provided further details into the mechanisms of evolution. Scientists now have a good understanding of the origin of new species (speciation) and have observed the speciation process in the laboratory and in the wild. Evolution is the principal theory that biologists use to understand life and is used in many disciplines, including medicine, psychology, conservation biology, anthropology, forensics, agriculture and other social-cultural applications. 2 Darwin's idea: evolution by natural selection In the 19th century, natural history collections and museums were a popular pastime. The European expansion and naval expeditions employed naturalists and curators of grand museums showcasing preserved and live specimens of the varieties of life. Charles Darwin was an English graduate who was educated and trained in the disciplines of natural history science. Such natural historians would collect, catalogue, describe and study the vast collections of specimens stored and managed by curators at these museums. Charles Darwin served as a ship's naturalist on board the HMS Beagle, assigned to a five-year research expedition around the world. During his voyage, Darwin observed and collected an abundance of organisms, being very interested in the diverse forms of life along the coasts of South America and the neighboring Galapagos Islands.[9] [10] Charles Darwin gained extensive experience as he collected and studied the natural history of life forms from distant places. Through his studies, Darwin formulated the idea that each species had developed from ancestors with similar features. In 1838, he described how a process he called natural selection would make this happen.[11] Darwin's idea of how evolution works relied on the following observations:[12] • If all the individuals of a species reproduced successfully, the population of that species would increase uncontrollably. • Populations tend to remain about the same size from year to year. • Environmental resources are limited. • No two individuals in a given species are exactly alike. • Much of this variation in a population can be passed on to offspring. Introduction to evolution 3 Charles Darwin proposed the theory of evolution by natural selection. Darwin noted that orchids exhibited a variety of complex adaptations to ensure pollination; all derived from basic floral parts. Darwin deduced that since organisms produce more offspring than their environment could possibly support, there must be a competitive struggle for survival—only a few individuals can survive out of each generation. Darwin realized that it was not chance alone that determined survival. Instead, survival depends on the traits of each individual and if these traits aid or hinder survival and reproduction. Well-adapted, or "fit", individuals are likely to leave more offspring than their less well-adapted competitors. Darwin realized that the unequal ability of individuals to survive and reproduce could cause gradual changes in the population. Traits that help an organism survive and reproduce would accumulate over generations. On the other hand, traits that hinder survival and reproduction would disappear. Darwin used the term natural selection to describe this process.[13] Natural selection is commonly equated with survival of the fittest, but this expression originated in Herbert Spencer's Principles of Biology in 1864, after Charles Darwin published his original works. Survival of the fittest describes the process of natural selection incorrectly, because natural selection is not only about survival and it is not always the fittest that survives.[14] Observations of variations in animals and plants formed the basis of the theory of natural selection. For example, Darwin observed that orchids and insects have a close relationship that allows the pollination of the plants. He noted that orchids have a variety of structures that attract insects, so that pollen from the flowers gets stuck to the insects’ bodies. In this way, insects transport the pollen from a male to a female orchid. In spite of the elaborate appearance of orchids, these specialized parts are made from the same basic structures that make up other flowers. In Fertilisation of Orchids Darwin proposed that the orchid flowers did not represent the work of an ideal engineer, but were adapted from pre-existing parts, through natural selection.[15] Darwin was still researching and experimenting with his ideas on natural selection when he received a letter from Alfred Wallace describing a theory very similar to his own. This led to an immediate joint publication of both theories. Both Wallace and Darwin saw the history of life like a family tree, with each fork in the tree’s limbs being a common ancestor. The tips of the limbs represented modern species and the branches represented the common ancestors that are shared amongst many different species. To explain these relationships, Darwin said that all living things were related, and this meant that all life must be descended from a few forms, or even from a single common Introduction to evolution ancestor. He called this process descent with modification.[12] Darwin published his theory of evolution by natural selection in On the Origin of Species in 1859. His theory means that all life, including humanity, is a product of continuing natural processes. The implication that all life on Earth has a common ancestor has met with objections from some religious groups who believe even today that the different types of life are due to special creation.[16] Their objections are in contrast to the level of support for the theory by more than 99 percent of those within the scientific community today.[17] 4 Source of variation Darwin’s theory of natural selection laid the groundwork for modern evolutionary theory, and his experiments and observations showed that the organisms in populations varied from each other, that some of these variations were inherited, and that these differences could be acted on by natural selection. However, he could not explain the source of these variations. Like many of his predecessors, Darwin mistakenly thought that heritable traits were a product of use and disuse, and that features acquired during an organism's lifetime could be passed on to its offspring. He looked for examples, such as large ground feeding birds getting stronger legs through exercise, and weaker wings from not flying until, like the ostrich, they could not fly at all.[18] This misunderstanding was called the inheritance of acquired characters and was part of the theory of transmutation of species put forward in 1809 by Jean-Baptiste Lamarck. In the late 19th century this theory became known as Lamarckism. Darwin produced an unsuccessful theory he called pangenesis to try to explain how acquired characteristics could be inherited. In the 1880s August Weismann's experiments indicated that changes from use and disuse could not be inherited, and Lamarckism gradually fell from favor.[19] The missing information needed to help explain how new features could pass from a parent to its offspring was provided by the pioneering genetics work of Gregor Mendel. Mendel’s experiments with several generations of pea plants demonstrated that inheritance works by separating and reshuffling hereditary information during the formation of sex cells and recombining that information during fertilization. This is like mixing different hands of cards, with an organism getting a random mix of half of the cards from one parent, and half of the cards from the other. Mendel called the information factors; however, they later became known as genes. Genes are the basic units of heredity in living organisms. They contain the information that directs the physical development and behavior of organisms. Genes are made of DNA, a long molecule that carries information. This information is encoded in the sequence of nucleotides in the DNA, just as the sequence of the letters in words carries information on a page. The genes are like short instructions built up of the "letters" of the DNA alphabet. Put together, the entire set of these genes gives enough information to serve as an "instruction manual" of how to build and run an organism. The instructions spelled out by this DNA alphabet can be changed, however, by mutations, and this may alter the instructions carried within the genes. Within the cell, the genes are carried in chromosomes, which are packages for carrying the DNA, with the genes arranged along them like beads on a string. It is the reshuffling of the chromosomes that results in unique combinations of genes in offspring. Although such mutations in DNA are random, natural selection is not a process of chance: the environment determines the probability of reproductive success. The end products of natural selection are organisms that are adapted to their present environments. Natural selection does not involve progress towards an ultimate goal. Evolution does not necessarily strive for more advanced, more intelligent, or more sophisticated life forms.[20] For example, fleas (wingless parasites) are descended from a winged, ancestral scorpionfly, and snakes are lizards that no longer require limbs - although pythons still grow tiny structures that are the remains of their ancestor's hind legs.[21] [22] Organisms are merely the outcome of variations that succeed or fail, dependent upon the environmental conditions at the time. Rapid environmental changes typically cause extinctions.[23] Of all species that have existed on Earth, 99.9 percent are now extinct.[24] Since life began on Earth, five major mass extinctions have led to large and sudden drops in the variety of species. The most recent, the Cretaceous–Tertiary extinction event, occurred 65 million years ago, and has Introduction to evolution attracted more attention than all others because it killed the dinosaurs.[25] 5 Modern synthesis The modern evolutionary synthesis was the outcome of a merger of several different scientific fields into a cohesive understanding of evolutionary theory. In the 1930s and 1940s, efforts were made to merge Darwin's theory of natural selection, research in heredity, and understandings of the fossil records into a unified explanatory model.[26] The application of the principles of genetics to naturally occurring populations, by scientists such as Theodosius Dobzhansky and Ernst Mayr, advanced understanding of the processes of evolution. Dobzhansky's 1937 work Genetics and the Origin of Species was an important step in bridging the gap between genetics and field biology. Mayr, on the basis of an understanding of genes and direct observations of evolutionary processes from field research, introduced the biological species concept, which defined a species as a group of interbreeding or potentially interbreeding populations that are reproductively isolated from all other populations. The paleontologist George Gaylord Simpson helped to incorporate fossil research, which showed a pattern consistent with the branching and non-directional pathway of evolution of organisms predicted by the modern synthesis. The modern synthesis emphasizes the importance of populations as the unit of evolution, the central role of natural selection as the most important mechanism of evolution, and the idea of gradualism to explain how large changes evolve as an accumulation of small changes over long periods of time. Evidence for evolution Scientific evidence for evolution comes from many aspects of biology, and includes fossils, homologous structures, and molecular similarities between species' DNA. Fossil record Research in the field of paleontology, the study of fossils, supports the idea that all living organisms are related. Fossils provide evidence that accumulated changes in organisms over long periods of time have led to the diverse forms of life we see today. A fossil itself reveals the organism's structure and the relationships between present and extinct species, allowing paleontologists to construct a family tree for all of the life forms on Earth.[27] Modern paleontology began with the work of Georges Cuvier (1769–1832). Cuvier noted that, in sedimentary rock, each layer contained a specific group of fossils. The deeper layers, which he proposed to be older, contained simpler life forms. He noted that many forms of life from the past are no longer present today. One of Cuvier’s successful contributions to the understanding of the fossil record was establishing extinction as a fact. In an attempt to explain extinction, Cuvier proposed the idea of “revolutions” or catastrophism in which he speculated that geological catastrophes had occurred throughout the Earth’s history, wiping out large numbers of species.[28] Cuvier's theory of revolutions was later replaced by uniformitarian theories, notably those of James Hutton During the voyage of the Beagle, naturalist Charles Darwin collected fossils in South America, and found fragments of armor which he thought were like giant versions of the scales on the modern armadillos living nearby. On his return, the anatomist Richard Owen showed him that the fragments were from gigantic extinct glyptodons, related to the armadillos. This was one of the patterns of distribution that helped Darwin to [11] develop his theory. Introduction to evolution and Charles Lyell who proposed that the Earth’s geological changes were gradual and consistent.[29] However, current evidence in the fossil record supports the concept of mass extinctions. As a result, the general idea of catastrophism has re-emerged as a valid hypothesis for at least some of the rapid changes in life forms that appear in the fossil records. A very large number of fossils have now been discovered and identified. These fossils serve as a chronological record of evolution. The fossil record provides examples of transitional species that demonstrate ancestral links between past and present life forms.[30] One such transitional fossil is Archaeopteryx, an ancient organism that had the distinct characteristics of a reptile (such as a long, bony tail and conical teeth) yet also had characteristics of birds (such as feathers and a wishbone). The implication from such a find is that modern reptiles and birds arose from a common ancestor.[31] 6 Comparative anatomy The comparison of similarities between organisms of their form or appearance of parts, called their morphology, has long been a way to classify life into closely related groups. This can be done by comparing the structure of adult organisms in different species or by comparing the patterns of how cells grow, divide and even migrate during an organism's development. Taxonomy Taxonomy is the branch of biology that names and classifies all living things. Scientists use morphological and genetic similarities to assist them in categorizing life forms based on ancestral relationships. For example, orangutans, gorillas, chimpanzees, and humans all belong to the same taxonomic grouping referred to as a family – in this case the family called Hominidae. These animals are grouped together because of similarities in morphology that come from common ancestry (called homology).[32] Strong evidence for evolution comes from the analysis of homologous structures: structures in different species that no longer perform the same task but which share a similar structure.[33] Such is the case of the forelimbs of mammals. The forelimbs of a human, cat, whale, and bat all have strikingly similar bone structures. However, each of these four species' forelimbs performs a different task. The same bones that construct a bat's wings, which are used for flight, also construct a whale's flippers, which are used for swimming. Such a "design" makes little sense if they are unrelated and uniquely constructed for their particular tasks. The theory of evolution explains these homologous structures: all four animals shared a common ancestor, and each has undergone change over many generations. These changes in structure have produced forelimbs adapted for different tasks.[34] Embryology In some cases, anatomical comparison of structures in the embryos of two or more species provides evidence for a shared ancestor that may not be obvious in the adult forms. As the embryo develops, these homologies can be lost to view, and the structures can take on different functions. Part of the basis of classifying the vertebrate group (which includes humans), is the presence of a tail (extending beyond the anus) and pharyngeal slits. Both structures appear during some stage of embryonic development but are not always obvious in the adult form.[35] Because of the morphological similarities present in embryos of different species during development, it was once assumed that organisms re-enact their evolutionary history as an embryo. It was thought that human embryos passed through an amphibian then a reptilian stage before completing their development as mammals. Such a re-enactment, (often called Recapitulation theory), is not supported by scientific evidence. What does occur, however, is that the first stages of development are similar in broad groups of organisms.[36] At very early stages, for instance, all vertebrates appear extremely similar, but do not exactly resemble any ancestral species. As development continues, specific features emerge from this basic pattern. . Vestigial refers to anatomical parts that are of minimal. yet are only distantly related – sharks are fish and dolphins are mammals. if any. Thus. Organisms that share similar environments will often develop similar physical features. Such similarities are a result of both populations being exposed to the same selective pressures. as not all anatomical similarities indicate a close relationship. value to the organism that possesses them. and the semilunar fold in the corner of the eye. a process known as convergent evolution.[39] 7 The bird and the bat wing are examples of convergent evolution. A bat is a mammal and its forearm bones have been adapted for flight. they developed similar appearances (morphology). Such is the case in whales. the wisdom teeth. changes that aid swimming have been favored.Introduction to evolution Vestigial structures Homology includes a unique group of shared structures referred to as vestigial structures. These apparently illogical structures are remnants of organs that played an important role in ancestral forms. over time. body hair (including goose bumps). which have small vestigial bones that appear to be remnants of the leg bones of their ancestors which walked on land. Both sharks and dolphins have similar body forms.[37] Humans also have vestigial structures. even though they are not closely related. Within both groups. the tail bone.[38] Convergent evolution Anatomical comparisons can be misleading. including the ear muscles. the appendix. the more distantly related two organisms are. If two organisms are closely related. the plant has evolved swollen thorns that the ants use as shelter and special flower parts that the ants eat. a plant that the ant uses for food and shelter. and they influence the properties of an organism. brothers are closely related and have very similar DNA. A section of DNA Co-evolution Co-evolution is a process in which two or more species influence the evolution of each other.[40] On the other hand. the more differences they will have. Genes determine an individual's general appearance and to some extent their behavior. The ant defends the acacia against herbivores and clears the forest floor of the seeds from competing plants. while cousins share a more distant relationship and have far more differences in their DNA. and the acacia. The relationship between the two is so intimate that it has led to the evolution of special structures and behaviors in both organisms. . These comparisons have allowed biologists to build a relationship tree of the evolution of life on Earth. across a population small genetic changes in both ant and tree benefited each. a type of ant.Introduction to evolution 8 Molecular biology Every living organism (with the possible exception of RNA viruses) contains molecules of DNA. their DNA will be very similar. In response. which carries genetic information. Over time. Rather. in co-evolution there is evidence that genetically determined traits in each species directly resulted from the interaction between the two organisms. For example. Genes are the pieces of DNA that carry this information. Similarities in DNA are used to determine the relationships between species in much the same manner as they are used to show relationships between individuals. All organisms are influenced by life around them. comparing chimpanzees with gorillas and humans shows that there is as much as a 96 percent similarity between the DNA of humans and chimps. For example. The benefit gave a slightly higher chance of the characteristic being passed on to the next generation.[44] Such co-evolution does not imply that the ants and the tree choose to behave in an altruistic manner.[41] [42] The field of molecular systematics focuses on measuring the similarities in these molecules and using this information to work out how different types of organisms are related through evolution.[43] They have even allowed scientists to unravel the relationships between organisms whose common ancestors lived such a long time ago that no real similarities remain in the appearance of the organisms. Comparisons of DNA indicate that humans and chimpanzees are more closely related to each other than either species is to gorillas. however.[40] An extensively documented case of co-evolution is the relationship between Pseudomyrmex. successive mutations created the relationship we observe today. 000 years ago. Humans determine which animal or plant will reproduce and which of the offspring will survive. For speciation to occur. which reproduce asexually. (The members of a species cannot produce viable. and enough time. The process of artificial selection has had a significant impact on the evolution of domestic animals. Allopatric speciation begins when a population becomes geographically separated.[46] Prior to domestication.[33] Geological processes. the formation of canyons. thus.Introduction to evolution 9 Artificial selection Artificial selection is the controlled breeding of domestic plants and animals. There are numerous species of cichlids that demonstrate dramatic variations in morphology. The differences in size between the Chihuahua and the Great Dane are the result of artificial selection.[48] Darwin carefully observed the outcomes of artificial selection in animals and plants to form many of his arguments in support of natural selection. Today The Maize Genetics Cooperation • Stock Center maintains a collection of more than 10. could produce the differences seen in living things today. but does not apply to organisms such as bacteria.[13] A widely accepted method of speciation is called allopatric speciation. they and all other dogs evolved from a few wolves domesticated by humans in what is now China less than 15. then natural selection. while in natural selection the surviving species is the one with random mutations useful to it in its non-human environment.000 years ago in central Mexico. Scientists have struggled to find a precise and all-inclusive definition of species. separation must be substantial. For example.[49] Much of his book On the Origin of Species was based on these observations of the many varieties of domestic pigeons arising from artificial selection. the edible portion of the wild form was small and difficult to collect. such as the emergence of mountain ranges. recent genetic evidence suggests that domestication occurred 10. Artificial selection has produced a wide variety of plants. fertile offspring. or the flooding of land bridges by changes in sea level may result in separate populations. given millions of years. Darwin proposed that if humans could achieve dramatic changes in domestic animals in short periods.[45] The results of artificial selection: a Chihuahua mix and a Great Dane. Ernst Mayr (1904–2005) defined a species as a population or group of populations whose members have the potential to interbreed naturally with one another to produce viable. so that genetic exchange between the two . evolution leads to the emergence of new species. Despite their dramatically different physical appearance.000 genetic variations of maize that have arisen by random mutations and chromosomal variations from the original wild type. fertile offspring with members of other species). and the underlying genetic processes are essentially the same. people have produced different types of dogs by controlled breeding. they determine which genes will be passed on to future generations. Speciation is the lineage-splitting event that results in two separate species forming from a single common ancestral population. In the case of maize (corn). In both natural and artificial selection the variations are a result of random mutations.[50] Mayr's definition has gained wide acceptance among biologists. Species Given the right circumstances.[47] In artificial selection the new breed or variety that emerges is the one with random mutations attractive to humans. there are conflicting opinions as to which is the primary unit of evolutionary change – the . the slight modifications introduced in the new species have changed the mate selection process and the five forms that arose could not be convinced to interbreed. The accumulated genetic changes may result in separated populations that can no longer interbreed if they are reunited. serving to link the diverse specialty areas of biology. If interbreeding is no longer possible. The significance of evolutionary theory is best described by the title of a paper by Theodosius Dobzhansky (1900–1975). the rate at which evolution occurs is still under discussion. In their separate environments. These fish have complex mating rituals and a variety of colorations.Introduction to evolution populations is completely disrupted. However speciation has been observed in present day organisms. In addition. thus direct observations within human life-spans are rare. occurring over very long time spans. the genetically isolated groups follow their own unique evolutionary pathways.[55] The evidence for speciation in this case was morphology (physical appearance) and lack of natural interbreeding.[51] Usually the process of speciation is slow. then they will be considered different species. and past speciation events are recorded in fossils. For example. There is much discussion within the scientific community concerning the mechanisms behind the evolutionary process. published in American Biology Teacher. Each group will accumulate different mutations as well as be subjected to different selective pressures.[57] Nevertheless. "Nothing in Biology Makes Sense Except in the Light of Evolution".[13] Barriers that prevent interbreeding are either prezygotic (prevent mating or fertilization) or postzygotic (barriers that occur after fertilization).[17] Evolution provides the field of biology with a solid scientific base.[56] 10 Different views on the mechanism of evolution James Hutton Stephen Jay Gould Richard Dawkins The theory of evolution is widely accepted among the scientific community. the theory of evolution is not static.[52] [53] [54] Scientists have documented the formation of five new species of cichlid fishes from a single common ancestor that was isolated fewer than 5000 years ago from the parent stock in Lake Nagubago. like hands of cards soon after they are dealt. When we have [60] served our purpose we are cast aside. both living and long dead. Evolution can be directly observed in artificial selection. by chance. for example. There are genetic variations within a population of individuals. While mutations are random. then persisting in that form for long periods. Evolutionary trees are based on the idea that profound differences in species are the result of many small changes that accumulate over long periods. The offspring might inherit the useful feature. The paleontologist Stephen Jay Gould (1941–2002) developed a model that suggests that evolution. Chromosomes too are shuffled to oblivion.[58] Unit of change A common unit of selection in evolution is the organism. a model called "punctuated equilibrium" which explains the fossil record without contradicting Darwin's ideas. alternating with long periods of relative stability. Such a view can seem to contradict the fossil record. The view that evolution is gradual had its basis in the works of the geologist James Hutton (1726–1797) and his theory called "gradualism".[61] Summary Several basic observations establish the theory of evolution. that natural selection operates as an evolutionary mechanism on genes as well as organisms. Hutton's theory suggests that profound geological change was the cumulative product of a relatively slow continuing operation of processes which can still be seen in operation today.Introduction to evolution organism or the gene. The organism view has been challenged by a variety of biologists as well as philosophers. .[59] In his 1976 book The Selfish Gene. Darwin and his contemporaries viewed evolution as a slow and gradual process. They are the replicators and we are their survival machines. Some individuals. Of course they march on. Evolution is an inevitable result of imperfectly copying. although a slow process in human terms. they merely change partners and march on. Fossils. Stephen Jay Gould called for a hierarchical perspective on selection. The outcome of evolution is not a perfectly designed organism. But the cards themselves survive the shuffling. dogs. not just at a single level of organism or gene. have features that allow them to survive and thrive better than their kind. natural selection is not. the genetic code. he explains: “ Individuals are not stable things. Richard Dawkins (born 1941) proposes that much insight can be gained if we look at evolution from the gene's point of view. 11 Rate of change Two views exist concerning the rate of evolutionary change. and agricultural plants serve as examples of evolution. ” Others view selection working on many levels. and the peculiar distribution of life on Earth provide a record of evolution and demonstrate the common ancestry of all organisms. self-replicating organisms reproducing over billions of years under the selective pressure of the environment. A uniformitarian perspective was adopted for biological changes. which shows evidence of new species appearing suddenly. The diverse breeds of cats. which explains the variety and relationship of all living things. Evolution is not a random process. they are fleeting. The outcome is simply an individual that can survive better and reproduce more successfully than its neighbors in a particular environment. Natural selection occurs when the reproductive success of an individual is improved or reduced by an inherited characteristic. The individuals that survive will be more likely to have offspring of their own. as opposed to catastrophism which promoted the idea that sudden changes had causes which can no longer be seen at work. But genes are denizens of geological time: genes are forever. that is. The cards are the genes. and reproductive success is measured by the number of an individual's surviving offspring. That is their business. undergoes periods of relatively rapid change over only a few thousand or million years. The genes are not destroyed by crossing-over. the selective breeding for certain traits of domestic animals and plants. horses. (September/October 1994). John van (2002). PMC 2424310. ncbi. . [12] Wyhe. ISBN 978-0393309270. PMID 18519614. J. ISBN 0-8018-6389-9. uk/ darwin. nationalgeographic. html). org/ RC/ AB/ BC/ Gregor_Mendel. Extinction: bad genes or bad luck. 1.75 million different species in existence today. Charles (1859). The Textbook League. 136.) Evolutionary Science and Society: Educating a New Generation. . Retrieved 2008-01-24. Retrieved 2007-12-23. CBE Life Sci Educ. . National Geographic. Access Excellence. org/ ). "Nonsense in schoolbooks: 'The Imaginary Lamarck'" (http:/ / www. "Confessions of a Darwinist" (http:/ / www. Baltimore and London: The Johns Hopkins University Press. org. ncbi. Berkeley. and Bybee R. . W. PMID 6085987. [3] Gregory. (http:/ / books.) Effects of the increased Use and Disuse of Parts. doi:10. American Institute of Biological Sciences.1187/cbe. "Fertilisation of Orchids" (http:/ / darwin-online. org. Retrieved 2008-01-07. . W. "Was Darwin Wrong?" (http:/ / ngm.). nih.). 1433. (2005). nlm.[5] [62] 12 See also • • • • • Creation-evolution controversy Evidence of common descent Evolution as theory and fact Level of support for evolution Misconceptions about evolution Notes [1] Darwin. with an estimated 1. html#top). od. . [2] Gould. google. M. London: John Murray. The Indelible Stamp: The Evolution of an Idea (http:/ / books. Klymkowsky. The Complete Works of Charles Darwin. uk/ content/ frameset?itemID=F373& viewtype=text& pageseq=16) (1st ed. "Gregor Mendel" (http:/ / www. xiii. 9780674006133. .2006. ISBN 13-978-0-7624-2136-7. University of Cambridge. com/ ?id=8klou91MwJoC& printsec=frontcover& dq=Extinction:+ bad+ genes+ or+ bad+ luck& cd=1#v=onepage& q=). . David (2004). pp. L. . 210. uk/ content/ frameset?itemID=F381& viewtype=text& pageseq=20) (3rd ed. J. "Cell evolution and Earth history: stasis and revolution" (http:/ / www. org. gov/ pubmed/ 16754610). D. Wilson (http:/ / books. the evidence of observation and experiments over a hundred years by thousands of scientists supports evolution. Michael T. Philadelphia and London: Running Press. The Nature of Natural Selection. NIH Record (National Institutes of Health). Cynthia (2006). doi:10. Retrieved 2007-12-21. Clio Med 19 (2): 136–55. "Understanding Randomness and its Impact on Student Learning: Lessons Learned from Building the Biology Concept Inventory (BCI)" (http:/ / www. O. R.. Philos Trans R Soc Lond B Biol Sci 361 (1470): 969–1006. 108 (http:/ / darwin-online. Stephen J. T. On the Origin of Species (http:/ / darwin-online. W. org. gov/ newsletters/ 2006/ 07_28_2006/ story03.07-08-0063. On the Origin of Species (http:/ / darwin-online. htm). textbookleague. org. com/ content/ 2331741806807x22/ ). Finding Order in Nature: The Natualist Tradition from Linnaeus to E. com/ ngm/ 0411/ feature1/ fulltext. .: Norton and Co. "Understanding Natural Selection: Essential Concepts and Common Misconceptions" (http:/ / www. Darwin. ISBN 0674006135. Ch. pp. pages 93-98 in Cracraft. W. (Eds. p. berkeley. [13] Quammen. [8] Rhee. 24) . John van (2002). . The University of California Museum of Paleontology. Related earlier ideas were acknowledged in Darwin. "The enigma of Darwin". google. com/ ?id=tyG4pfKJ8WEC& printsec=frontcover& dq=Finding+ Order+ in+ Nature:+ The+ Natualist+ Tradition+ from+ Linnaeus+ to+ E.. accessexcellence. K. uk/ content/ frameset?itemID=F391& viewtype=text& pageseq=136). . National Geographic Magazine. as controlled by Natural Selection [19] Ghiselin. P. org/ 54marck. Retrieved 2008-01-23 [20] (Gould (a) 1981. [9] Farber. (2000). edu/ evolibrary/ article/ 0_0_0/ evo_01). p. . p. html). 7 (2): 227–233. (1992). 8. com/ ?id=LEWfWf0mUJIC& printsec=frontcover& dq=Darwin:+ The+ Indelible+ Stamp:+ The+ Evolution+ of+ an+ Idea& cd=1#v=onepage& q=). University of Cambridge. Charles (1861). Harvard University Press. (2009). gov/ pmc/ articles/ PMC2424310/ ). pp. The Textbook Letter (http:/ / www. J. The Virginia Quarterly Review: 32–53. [4] "An introduction to evolution" (http:/ / evolution. (2008). M. 2008. The Complete Work of Charles Darwin Online. National Health Museum. springerlink. [7] Raup. nlm. google. textbookleague.[16] The result of four billion years of evolution is the diversity of life around us. [11] Eldredge. D. 1257. . berkeley. Sue Yon (1999). [18] (Darwin 1872. nih. + Wilson& cd=1#v=onepage& q=). PMID 16754610. Understanding Evolution: your one-stop source for information on evolution (http:/ / evolution. pp. Retrieved 2008-01-23.Introduction to evolution Although some groups raise objections to the theory of evolution. [16] DeVries A (2004).. . Evolution: Education and Outreach 2 (2): 156–175.1842.1007/s12052-009-0128-1. [10] Watson. html). (2005). . "Finding the Evolution in Medicine" (http:/ / nihrecord. "Charles Darwin: gentleman naturalist" (http:/ / darwin-online. Retrieved 2008-01-23 [5] Cavalier-Smith T (2006). org/ articles/ 2006/ spring/ eldredge-confessions-darwinist/ ). [15] Wyhe. Retrieved 2008-01-16. [6] Garvin-Doxas.1098/rstb. Retrieved 2008-01-05. Niles (Spring 2006). New York. + O. uk/ EditorialIntroductions/ Freeman_FertilisationofOrchids. [17] Delgado. The Structure of Evolutionary Theory. PMC 1578732. D. [14] Futuyma. (2002). nih. edu/ evolibrary/ article/ 0_0_0/ evo_02) (web resource). vqronline. doi:10. London: John Murray. htm). PMID 12492145. htm). 2006-06-21. php?record_id=12026& page=10). [40] Kennedy. html). Diewert V.Life's Epic" (http:/ / www. [37] Theobald. com/ km/ evol/ embryos/ Haeckel. Retrieved 2008-01-06. National Academy of Sciences. Retrieved 2008-04-04. 'On Science' column in St. html). [44] Janzen. . Daniel (1974). talkorigins. pbs. html) (web resource). "The tangled web of biological science" (http:/ / books. [41] Lovgren. "Origin of dogs traced" (http:/ / news. Evolution Resources.. edu/ readingroom/ books/ evolution98/ evol3. Evolution: a jouney into where we're from and where we're going (http:/ / www. ISSN 1367-4803. org/ wgbh/ evolution/ library/ glossary/ index. pbs. nap. pdf) (pdf). p. p. org/ wgbh/ evolution/ extinction/ massext/ index.zool.1123061. TalkOrigins Archive. Doerks T. Smithsonian Contributions to Biology. org/ pdf/ comdesc.I: Staging criteria and macroscopic skeletal morphogenesis of the head and limbs". [39] Johnson. p.x. pp. [45] McGourty. nap. WGBH Educational Foundation.005. Evolution and the nature of science. "29+ Evidences for Macroevolution Part 2: Past History" (http:/ / www. A. doi:10. . [23] Drummond. Snel B. "Embryonic development of Python sebae . "Chimps. html). Bork P (2006). Wang. George (2002). "Better Books by Trial and Error. Department of Agriculture. Richman JM (2007). Grenier & Weatherbee 2000) [43] Ciccarelli FD. uk/ 2/ hi/ science/ nature/ 2498669. . Zoology (Jena) 110 (3): 212–30. stm). pp. php?record_id=11876& page=22). nationalgeographic. 8) [36] Miller. National Academy of Sciences and Institute of Medicine of the National Academies (2008). html). com/ news/ 2005/ 08/ 0831_050831_chimp_genes. "Origination. . ISSN 1367-4803. html) (web resource). nap. Evol. George (2002). [48] Silverman. .K. 16) [33] Drummond. bbc. "Evolution Library" (http:/ / www. "Teaching about evolution and the nature of science" (http:/ / www. org/ wgbh/ evolution/ index. Creevey CJ. [46] Hall. Strimmer.C. K (Jul 2001). Retrieved 2008-01-23.2. . "Swollen-Thorn Acacias of Central America" (http:/ / www.CO. cropsci. Retrieved 2007-12-19. [28] (Tattersall 1995. Hardy. Bioinformatics (Oxford. Retrieved 2008-01-23. A. and mass depletions of marine diversity" (http:/ / findarticles. Dev. [31] (Gould (b) 1995. txtwriter. net/ ~lifebook/ evolve. pbs. sil. .01. txtwriter. Kenneth (1997). and Creationism" (http:/ / books. [25] Bambach. txtwriter. html). 5–6) [29] (Lyell 1830.02033. The Evidence for Evolution (http:/ / www. "Science. ca/ transgene-escape-are-traditional-corn-varieties-in-mexico-threatened-by-transgenic-corn-crops/ ?).H. 76) [30] Committee on Revising Science and Creationism: A View from the National Academy of Sciences. Retrieved 2008-01-06. . . The role of theory in advancing 21st Century Biology:Catalyzing Transformation Research. edu/ smithsoniancontributions/ Botany/ pdf_hi/ sctb-0013. [27] "The Fossil Record . [47] "The Maize Genetics Cooperation • Stock Center" (http:/ / maizecoop. extinction. com/ backgrounders/ Evolution/ EVpage14. comcast. "Limbs in whales and limblessness in other vertebrates: mechanisms of evolutionary and developmental transformation and loss". php). Retrieved 2007-12-14.1126/science.Introduction to evolution [21] Bejder L.S. Frequently Asked Questions About Evolution (http:/ / www. Humans 96 Percent the Same. National Research Council. html) (web resource). com/ p/ articles/ mi_qa4067/ is_200410/ ai_n9458414/ pg_1). "Haeckel and his Embryos" (http:/ / www. Strimmer. Retrieved 2007-12-23." (http:/ / home.. 4 (6): 445–58. . 'On Science' column in St. co. The TalkOrigins Foundation. Retrieved 2007-12-14. doi:10. R. U. . ubc. The National Academy of Science. PMID 11448888.2007. . E. [42] (Carroll. England) (WGBH Educational Foundation) 17 (7): 662–3. Gene Study Finds" (http:/ / news. com/ backgrounders/ index. (December 2004). PMID 11448888. edu/ openbook. Evolution. Science 311 (5765): 1283–87. National Geographic. html). The Virtual Fossil Museum. Stefan (2005-08-31). scq. [22] Boughner JC.1046/j. . txtwriter. Fu K. "Transgene Escape: Are Traditioanl Corn Varieties In Mexico Threatened by Transgenic Corn Crops" (http:/ / www. Hallgrímsson B. Donald. fossilmuseum.1666/0094-8373(2004)030<0522:OEAMDO>2. "Convergent and Divergent Evolution" (http:/ / www. Scientific Creative Quarterly. si. David (2002). edu/ openbook. Hall BK (2002). net/ fossilrecord. National Geographic News. Knoll. millerandlevine. Louis Post Dispatch. (Working group on teaching evolution) (1998). . pbs. . 13 . PMID 16513982. . A. [38] Johnson. [34] (Mayr 2001. org/ wgbh/ evolution/ library/ faq/ cat03. 360) [32] (Diamond 1992. Retrieved 2008-01-23. html). "Vestigial Structures" (http:/ / www. com/ backgrounders/ index. PMID 17499493. Christine (2002-11-22). von Mering C.2002. The Evidence for Evolution (http:/ / www. England) (WGBH Educational Foundation) 17 (7): 662–3.1525-142X. Smithsonian Institution.0. html). org/ wgbh/ evolution/ library/ faq/ cat03. Retrieved 2007-09-02. edu/ mgc-info. doi:10. Retrieved 2008-01-27. Retrieved 2008-01-23. p. Glossary (http:/ / www. Retrieved 2007-12-30. com/ backgrounders/ Evolution/ EVpage12. Retrieved 2007-08-31. Buchtová M. Douglas (2004). . S. Bioinformatics (Oxford. BBC News. html). Louis Post Dispatch. National Plant Germplasm. 25–27) [35] (Weichert & Presch 1975. "Toward automatic reconstruction of a highly resolved tree of life". Paleobiology 30 (4): 522–42. doi:10. . pbs. K (Jul 2001). Retrieved 2008-01-23 [24] "Roundtable: Mass Extinction" (http:/ / www. html). Retrieved 2008-01-24 [26] Committee on Defining and Advancing the Conceptual Basis of Biological Sciences (1989). pdf) (pdf). 2001. Retrieved 2007-08-31.1016/j. uiuc. . ISBN 0674690109 • Mayr. Jared (1992). Species. Oxford: Blackwell Publishing. [52] Jiggins CD. Bridle JR (2004). com/ issues/ 2005/ december/ darwin. "Evidence for Rapid Speciation Following a Founder Event in the Laboratory" (http:/ / jstor.org. org/ faqs/ faq-speciation. Norton & Company. org/ evolution/ education/ cans-cants-teaching-evolution). Dinosaur in a Haystack. [55] (Mayr 1970. PMC 18385. Ernst (2001).1073/pnas. Charles (1830). html). Evolution (Evolution. Starczak VR. Frank J (December 2005).2307/2407990. Stud Hist Philos Biol Biomed Sci. .com/doc/104123/ Richard-Dawkins-The-Selfish-Gene-Original-Ed) (1st ed.2-Z& size=LARGE& origin=JSTOR-enlargePage). J. 37 (1): 26–40. The Panda's Thumb: More Reflections in Natural History. Oxford University Press. Charles. TalkOrigins Archive. org/ article. ISBN 0192860925 • Diamond. doi:10.008.002. ISBN 0195061012 • Weichert. U. 33. ISBN 0393308197 • Gould (b).tree. enviroliteracy. Sewall (September 1980). The Selfish Gene (http://www.org/books/lyell/principles/facsimile/). Charles (1872). . . "Darwin's artificial selection as an experiment". 348) [56] (Mayr 1970) [57] "NCSE Resource" (http:/ / ncseweb. org/ stable/ 2409766). PMID 16473266. Ian (1995). doi:10.2003.Introduction to evolution [49] Wilner A. Evol.). John (1995). Stephen Jay Gould Archive. Presch. 5) 34 (5): 825. ISBN 0060183071 • Gould (a). 4) 46 (4): 1214–20. jstor. Stephen Jay (1995). Natl.uk/content/ frameset?itemID=F391&viewtype=text&pageseq=1)]] (6th ed. Smithsonian Institution. CO. William (1975).scribd. ISBN 0070690081 . Weatherbee. "Genic and Organismic Selection" (http:/ / links. Roger (2007). pp.). Trends Ecol. [53] Boxhorn. The Fossil Trail: How We Know What We Think We Know About Human Evolution. Lloyd EA (1999). Richard (1976). Natural History. org/ sici?sici=0014-3820(198009)34:5<825:GAOS>2. PMID 16701238. Grenier. Smithsonian Magazine. php/ 58. 14 References • Carroll. [58] Gould. talkorigins. Principles of geology (http://www.96. . Retrieved 2007-08-31. The Third Chimpanzee: the evolution and future of the human animal. ISBN 0-465-04425-5 • Tattersall.A. html). No. pp. 34. 0. No. 2001-02-13. 165–69) [51] Sulloway.12. From DNA to Diversity: Molecular Genetics and the Evolution of Animal Design (2nd ed. Stephen Jay (1991). . Retrieved 2007-05-10. Retrieved 2007-08-31. Jorg.esp. . New York: W. Retrieved 2007-12-23. and Evolution.2307/2409766. MA: Belknap Press of Harvard University Press. [54] Weinberg JR. [60] (Dawkins 1976. pnas. (Amst. "Opus 200" (http:/ / www. Proc. org/ cgi/ pmidlookup?view=long& pmid=10518549). 46. 96 (21): 11904–9. ISBN 014043528X • Mayr. "Individuality and adaptation across levels of selection: how shall we name and generalize the unit of Darwinism?" (http:/ / www. New York: Oxford University Press. doi:10. (2006). p. New York: Basic Books. php?page=2). .S. ISBN 0517703939 • Lyell. "How many species are there?" (http:/ / www. Retrieved 2008-01-05. [[On the Origin of Species|The Origin of Species (http://darwin-online. Evolution (Evolution. What evolution is.1016/j.11904.21. New York: HarperCollins. "The Evolution of Charles Darwin" (http:/ / www. doi:10. SB.shpsc. [50] (Mayr 2001. ISBN 1-4051-1950-0 • Darwin. "Speciation in the apple maggot fly: a blend of vintages?".1016/j. "Observed Instances of Speciation" (http:/ / www. Vol. Elements of Chordate Anatomy. [62] Sedjo. Retrieved 2008-01-01. Retrieved 2008-01-18.12. 35) [61] Gould SJ. D (1992). [59] Wright.) 19 (3): 111–14. Acad. Populations. Retrieved 2008-01-24. smithsonianmagazine. PMID 10518549. New York: Penguin Books. Sci. p. stephenjaygould. SD (2000). London: John Murray • Dawkins.W. Cans and Can`ts of Teaching Evolution. Cambridge. Environmental Literacy Council.). html). Stephen Jay (1981). New York: Harmony Books. Vol. doi:10. Ernst (1970). org/ library/ gould_opus200. New York: McGraw-Hill. . National Center for Science Education.2005. Carl Sagan on evolution (http://video. wellcome.cotch. • Greg Krukonis (2008). Quammen. • Carl Sagan.youtube. (2001-11-20) (web resource). Mark (2009). Marshall. New York: Farrar Straus Giroux. Evolution: a journey into where we're from and where we're going (http://www.pbs. How Science Works: Evolution. ed.html).wellcome. Retrieved 2008-01-24. Evolution For Dummies (For Dummies (Math & Science)). Brian (2003). Deborah. R.Introduction to evolution 15 Further reading • Liam Neeson (narrator). (2006-07-06) (Google video).edu/units/basics/tour) (web resource). ISBN 9048131820 External links • Brain. ISBN 0-470-11773-7. Retrieved 2008-01-24 • (web resource) Understanding Evolution: your one-stop source for information on evolution (http://evolution. Retrieved 2008-01-24. Wellcome Trust. ASIN: B00005RG6J. Springer. On the Origin of Species: The Illustrated Edition.pdf). The Rough Guide to Evolution. January 2007. Retrieved 2008-01-24. Retrieved 2008-01-24 • Carl Sagan. The tree of life: a book depicting the life of Charles Darwin.edu/evolibrary/article/0_0_0/evo_01). Carl (2009). Charlesworth. "How Stuff Works: Evolution Library" (http://science. • Charlesworth. Leslie Alan (2002). Berkeley. ISBN 1402756399 • Pallan. Retrieved 2008-01-23 • "The Talk Origins Archive: Exploring the Creation/Evolution Controversy" (http://www.com/ watch?v=E1Y5zMo74cY). Oxford: Oxford University Press. htm) (web resource).com/evolution-channel.net. Retrieved 2010-06-01 . VT: WGBH Boston / PBS television series Nova.com. ISBN 0192805045. ISBN 0-374-45628-3.howstuffworks.google.net/taevo. John (2010). [DVD]. utah. Fossils: a very short introduction. ISBN 0192802518. • Thomson. Charles (2008). Indianapolis: Alpha Books. Keith Stewart (2005). South Burlington.vectorsite.ac. Howstuffworks. Oxford: Oxford University Press.org/) (web resource). Retrieved 2008-01-24 • "Introduction To Evolution" (http://www. Evolution: a very short introduction.com.net/) (web resource).uk/Professional-resources/Education-resources/Big-Picture/Evolution/). ISBN 1858289467 • Zimmer. The Big Picture Series (http://www.ac. ISBN 0981519474 • Ellis. [streaming video]. Theory of Evolution Explained (http://www. Peter (2003). (2006-10-21) (Youtube video). David. .org/wgbh/evolution/index.talkorigins. berkeley. The University of California Museum of Paleontology.com/ videoplay?docid=-522726029201501667&q=carl+sagan). Retrieved 2008-01-24 • "University of Utah Genetics Learning Center animated tour of the basics of genetics" (http://learn. • Darwin.uk/stellent/groups/corporatesite/ @msh_publishing_group/documents/web_document/wtd026042. geologist & thinker.html) (web resource). For Dummies.genetics. Rough Guides.Age level: Grade 7+ • Horvitz. The complete idiot's guide to evolution. ISBN 0028642260. • Sis. Retrieved 2008-01-24 • "The Big Picture on Evolution (PDF)" (http://www. Sterling. Roberts and Company Publishers. Google. • "Evolution Education Wiki: EvoWiki" (http://wiki. naturalist. vectorsite. Youtube. The Tangled Bank: An Introduction to Evolution. Howstuffworks. [streaming video]. Another cause of evolution is genetic drift. This powerful explanatory and predictive theory has become the central organizing principle of modern biology. variation can also be increased by the transfer of genes between species.[14] leading to the overwhelming acceptance of evolution among scientists. variants with particular traits become more. This iterative process adjusts traits so they become better suited to an organism's environment: these adjustments are called adaptations. while traits that hinder survival and reproduction become rarer. but significant. all living (and extinct) species are descended from a common ancestor via a long series of speciation events. forming the modern evolutionary synthesis. directing research and providing a unifying explanation for the history and diversity of life on Earth. Evolutionary biologists document the fact that evolution occurs. Genetic drift is most important when traits do not strongly influence survival—particularly so in small populations. Darwin's seminal work On the Origin of Species brought the new theory of evolution by natural selection to a wide audience.[16] [17] [20] Evolution is applied and studied in fields as diverse as agriculture. Ultimately.Evolution 16 Evolution Evolution (also known as biological. The study of evolutionary biology began in the mid-nineteenth century. The main source of variation is mutation. Under certain circumstances. conservation biology.[6] However. ecology. in which a single ancestral species splits and diversifies into multiple new species. These changes are heritable (can be passed on through reproduction). genetic and other similarities between groups of organisms. and also develop and test theories which explain its causes.[2] [3] and by the extremely rare. Natural selection occurs because only a small proportion of individuals in each generation will survive and reproduce. A trait is a particular characteristic—anatomical. wholesale incorporation of genomes through endosymbiosis. heritable variation in traits is filtered by natural selection and the beneficial changes are successively retained through differential survival and reproduction. Darwinian natural selection became understood in combination with Mendelian inheritance.[15] [16] [17] [18] In the 1930s. There are several modes through which this occurs. As a result.[9] [10] [11] [12] This is visible in anatomical. anthropology. Another source of variation is genetic recombination. In 1859. which leads to random changes in how common traits are in a population.[19] which connected the substrate of evolution (inherited genetics) and the mechanism of evolution (natural selection). One is natural selection. biochemical or behavioural—that is the result of gene–environment interaction. since resources are limited and organisms produce many more offspring than their environment can support. geographical distribution of related species. medicine.5 billion years during which life has existed on Earth. which introduces genetic changes. and other processes that remove it. through which traits that aid survival and reproduction become more common. which shuffles the genes into new combinations which can result in organisms exhibiting different traits.[1] This change results from interactions between processes that introduce variation into a population. in which chance plays a disproportionate role in the frequency of traits passed on to the next generation. philosophy. and psychology along with other specific topics in the previous listed fields. common. A key process in evolution is speciation. and plays a role in the molecular clocks that are used in phylogenetic studies. when research into the fossil record and the diversity of living organisms convinced most scientists that species changed over time.[13] The mechanism driving these changes remained unclear until the theory of natural selection was independently proposed by Charles Darwin and Alfred Wallace.[4] [5] Two main processes cause variants to become more common or rarer in a population. . the fossil record and the recorded genetic changes in living organisms over many generations. These events stretch back in a diverse "tree of life" which has grown over the 3. not all change is adaptive. genetic or organic evolution) is the change in the inherited traits of a population of organisms through successive generations. paleontology. and may give rise to alternative traits in organisms. Over many generations.[7] [8] Genetic drift is important in the neutral theory of molecular evolution. or less. whose specialist training in particular organisms addressed general questions in evolution. it began to draw more widely from the biological sciences. Charles Darwin formulated his idea of natural selection in 1838 and was still developing his theory in 1858 when Alfred Russel Wallace sent him a similar theory. the identification of DNA as the genetic material by Oswald Avery and colleagues and the subsequent publication of the structure of DNA by James Watson and Francis Crick in 1953. Sewall Wright.S. their rediscovery of and subsequent work on genetics eventually provided a solid basis on which the theory of evolution stood even more convincingly than when it was originally proposed.[26] In the 1880s. with the Greek philosopher Anaximander. but were rejected by mainstream scientists. August Weismann's experiments indicated that changes from use and disuse were not heritable. ecology. Around 1854 Charles Darwin began writing out what became On the Origin of Species. and Darwin could not explain the source of the heritable variations which would be acted on by natural selection. and particularly Ronald Fisher. philosophy and .[19] In its early history. and evolutionary concepts are used in even more distant disciplines such as psychology.B.[19] Currently the study of evolutionary biology involves scientists from fields as diverse as biochemistry. That variation is the main fuel used by natural selection to shape the wide variety of adaptive traits observed in organic life. the growth of modern biology out of natural history is fairly recent. Haldane. and both were presented to the Linnean Society of London in separate papers. particularly after the development of the modern evolutionary synthesis. demonstrated the physical basis for inheritance.[31] In the 1940s.[23] At the end of 1859.[27] [28] More significantly.[25] a theory which was subsequently dubbed Lamarckism. and Lamarckism gradually fell from favour. genetics and molecular biology have become core parts of evolutionary biology and have revolutionised the field of phylogenetics. Darwin could not account for how traits were passed down from generation to generation. The end result was a combination of evolution by natural selection and Mendelian inheritance. evolutionary biology primarily drew in scientists from traditional taxonomically oriented disciplines. he still thought that parents passed on adaptations acquired during their lifetimes. As biological knowledge grew in the 18th century. genetics and physiology. meaning "to unroll like a scroll") appeared in English in the 17th century.[30] The apparent contradiction between Darwin's theory of evolution by natural selection and Mendel's work was reconciled in the 1920s and 1930s by evolutionary biologists such as J.[22] The ideas of the biologist Jean-Baptiste Lamarck about transmutation of species influenced radicals. disagreements over the rate of evolution predicted by early geneticists and biometricians led to a rift between the Mendelian and Darwinian models of evolution. Since then. In 1865 Gregor Mendel found that traits were inherited in a predictable manner. proto-evolutionary ideas were set out by a few natural philosophers such as Pierre Maupertuis in 1745 and Erasmus Darwin in 1796.[29] When Mendel's work was rediscovered in 1900s. Darwin's publication of On the Origin of Species explained natural selection in detail and presented evidence leading to increasingly wide acceptance of the occurrence of evolution. Debate about the mechanisms of evolution continued.[21] However.[24] Like Lamarck.Evolution 17 History of evolutionary thought The roots of naturalistic thinking on biology can be dated to at least the 6th century BCE. As evolutionary biology expanded as an academic discipline. medicine. Even though Hugo de Vries and other early geneticists rejected gradual natural selection. The word evolution (from the Latin evolutio. Yet it was the rediscovery of Gregor Mendel's pioneering work on the fundamentals of genetics (of which Darwin and Wallace were unaware) by Hugo de Vries and others in the early 1900s that provided the impetus for a better understanding of how variation occurs in plant and animal traits. the modern evolutionary synthesis. who set the foundations for the establishment of the field of population genetics. most traits are more complex and are controlled by multiple interacting genes. suntanned skin comes from the interaction between a person's genotype and sunlight. the new allele may affect the trait that the gene controls.[35] DNA is a long polymer composed of four types of bases.[36] As a result. current research in evolutionary biology deals with several areas where the modern evolutionary synthesis may need modification or extension. the different forms of this sequence are called alleles. Epigenetics is when a trait is inherited without there being any change in gene sequences. For example. Bases are in the center.[35] The complete set of observable traits that make up the structure and behavior of an organism is called its phenotype. producing new alleles. many aspects of an organism's phenotype are not inherited. DNA sequences can change through mutations. These traits come from the interaction of its genotype with the environment.Evolution computer science. such as assessing the relative importance of various ideas on the unit of selection and evolvability and how to fully incorporate the findings of evolutionary developmental biology. different genes have different sequences of bases. the long strands of DNA form condensed structures called chromosomes.[32] [33] 18 Heredity Evolution in organisms occurs through changes in heritable traits – particular characteristics of an organism. Before a cell divides. due to differences in their genotype. In the 21st century. However.[38] [39] The study of such complex traits is a major area of current genetic research. a molecule that encodes genetic information. Within cells. the DNA is copied. In humans. altering the phenotype of the organism.[37] Heritable traits are passed from one generation to the next via DNA. eye colour is an inherited characteristic and an individual might inherit the "brown-eye trait" from one of their parents. If the DNA sequence at a locus varies between individuals. Portions of a DNA molecule that specify a single functional unit are called genes. surrounded by phosphate–sugar chains in a double helix. so that each of the resulting two cells will inherit the DNA sequence. The specific location of a DNA sequence within a chromosome is known as a locus. If a mutation occurs within a gene.[40] . Another unsolved question in genetics is whether or not epigenetics is important in evolution. a striking example are people with the inherited trait of albinism. DNA structure. some people tan more easily than others. The sequence of bases along a particular DNA molecule specify the genetic information. while this simple correspondence between an allele and a trait works in some cases. suntans are not passed on to people's children. However. thus. for example. in a manner similar to a sequence of letters spelling out a sentence. who do not tan at all and are very sensitive to sunburn.[34] Inherited traits are controlled by genes and the complete set of genes within an organism's genome is called its genotype. with these fragments then recombining to form new combinations with new functions.[44] Mutation Random mutations constantly occur in the genomes of organisms. which can introduce extra copies of a gene into a genome. domains act as modules Duplication of part of a chromosome .[58] [59] The creation of new genes can also involve small parts of several genes being duplicated. these mutations create genetic variation. most of the genome of a species is identical in all individuals of that species.[42] Despite the constant introduction of variation through these processes. becoming more or less prevalent relative to other forms of that gene. The frequency of one particular allele will fluctuate.[52] Extra copies of genes are a major source of the raw material needed for new genes to [53] evolve. This process is easier once a gene has been duplicated because it increases the redundancy of the system. and thus evade the defensive responses of e. and hybridization in plants.[45] [46] [47] These mutations involve several different types of change in DNA sequences.Evolution 19 Variation An individual organism's phenotype results from both its genotype and the influence from the environment it has lived in. Variation also comes from exchanges of genes between different species.g.[39] The modern evolutionary synthesis defines evolution as the change over time in this genetic variation. such as DNA repair enzymes. Evolutionary forces act by driving these changes in allele frequency in one direction or another. organisms have evolved mechanisms such as DNA repair to remove mutations. these can either have no effect. A substantial part of the variation in phenotypes in a population is caused by the differences between their genotypes. through horizontal gene transfer in bacteria. the optimal mutation rate for a species is a trade-off between costs of a high mutation rate. and the reshuffling of genes through sexual reproduction. migration between populations (gene flow).[49] Viruses that use RNA as their genetic material have rapid mutation rates. with about 70 percent of these mutations having damaging effects. Mutations are changes in the DNA sequence of a cell's genome and are caused by radiation. the human eye uses four genes to make structures that sense light: three for colour vision and one for night vision.[54] For example. viruses.[48] Due to the damaging effects that mutations can have on cells. and the metabolic costs of maintaining systems to reduce the mutation rate. this will probably be harmful.[60] [61] When new genes are assembled from shuffling pre-existing parts. This is important because most new genes evolve within gene families from pre-existing genes that share common ancestors.[51] Mutations can involve large sections of a chromosome becoming duplicated (usually by genetic recombination).[50] which can be an advantage since these viruses will evolve constantly and rapidly. the human immune system.[43] However.[45] Therefore. chimpanzees and humans differ in only about 5% of their genomes.[55] New genes can be created from an ancestral gene when a duplicate copy mutates and acquires a new function. alter the product of a gene. all four are descended from a single ancestral gene. as well as errors that occur during meiosis or DNA replication. such as deleterious mutations. one gene in the pair can acquire a new function while the other copy continues to perform its original function. Variation disappears when a new allele reaches the point of fixation — when it either disappears from the population or replaces the ancestral allele entirely. for example. even relatively small changes in genotype can lead to dramatic changes in phenotype: for example. or prevent the gene from functioning. Studies in the fly Drosophila melanogaster suggest that if a mutation changes a protein produced by a gene. and the remainder being either neutral or weakly beneficial. transposons and mutagenic chemicals.[56] [57] Other types of mutation can even create entirely new genes from previously noncoding DNA.[41] Variation comes from mutations in genetic material. such as transposons. A set of alleles that is usually inherited in a group is called a haplotype.[79] [80] that has prompted ideas such as the Red Queen hypothesis. and they retain these separate chromosomes.[62] For example. a phenomenon known as linkage. producing offspring with new combinations of alleles. as they cannot mix with genes of other organisms during reproduction. the offspring of sexual organisms contain random mixtures of their parents' chromosomes that are produced through independent assortment. In contrast. can cause mutations and may separate beneficial combinations of genes. which pass intact from fathers to sons – harmful mutations accumulate.[65] Sequences of DNA that can move about the genome. sexual organisms exchange DNA between two matching chromosomes.[66] For example. asexuality might allow the two sets of alleles in their genome to diverge and gain different functions. more than a million copies of the Alu sequence are present in the human genome.[46] 20 Sex and recombination In asexual organisms.[63] Changes in chromosome number may involve even larger mutations. recombination and reassortment can produce individuals with new and advantageous gene combinations. and may have been important in the evolution of genomes. they can mutate or delete existing genes and thereby produce genetic diversity. the rate of recombination is low (approximately two events per chromosome per generation).[74] This tendency is measured by finding how often two alleles occur together on a single chromosome.[70] [71] However. which can be mixed together creating new combinations with new and complex functions.[68] Recombination and reassortment do not alter allele frequencies.[69] Sex usually increases genetic variation and may increase the rate of evolution.[78] In addition. two chromosomes in the Homo genus fused to produce human chromosome 2. they contain up to one hundred independent domains that each catalyze one step in the overall process. This can be important when one allele in a particular haplotype is strongly beneficial: natural selection can drive a selective sweep that will also cause the other alleles in the haplotype to become more common in the population. this fusion did not occur in the lineage of the other apes. and genes that are close together tend to be inherited together.[67] Another effect of these mobile DNA sequences is that when they move within a genome.[78] The reasons for the evolution of sexual reproduction are therefore unclear and this question is still an active area of research in evolutionary biology. make up a major fraction of the genetic material of plants and animals. As a result. like a step in an assembly line.[73] Recombination allows even alleles that are close together in a strand of DNA to be inherited independently. genes close together on a chromosome may not always be shuffled away from each other.[75] When alleles cannot be separated by recombination – such as in mammalian Y chromosomes. which is called their linkage disequilibrium.[72] Here. the most important role of such chromosomal rearrangements may be to accelerate the divergence of a population into new species by making populations less likely to interbreed. However. For example. but instead change which alleles are associated with each other. genes are inherited together. and thereby preserving genetic differences between these populations.[76] [77] By breaking up allele combinations. sexual reproduction allows the removal of harmful mutations and the retention of beneficial mutations.Evolution with simple independent functions. this effect is called genetic hitchhiking. In a related process called homologous recombination. asexuality is advantageous in some environments as it can evolve in previously sexual animals. where segments of the DNA within chromosomes break and then rearrange. and these sequences have now been recruited to perform functions such as regulating gene expression.[64] In evolution. polyketide synthases are large enzymes that make antibiotics. or linked.[81] . These positive effects are balanced by the fact that sex reduces an organism's reproductive rate. the allele frequency measures the fraction of the gene pool composed of a single allele (for example. A single gene in this population may have several alternate forms.[84] .Evolution 21 Population genetics White peppered moth Black morph in peppered moth evolution From a genetic viewpoint. Evolution occurs when there are changes in the frequencies of alleles within a population of interbreeding organisms.[82] A population is a localised group of individuals belonging to the same species. An example might be a gene for colouration in moths that has two alleles: black and white. The Hardy-Weinberg principle states that the frequencies of alleles (variations in a gene) in a sufficiently large population will remain constant if the only forces acting on that population are the random reshuffling of alleles during the formation of the sperm or egg. To understand the mechanisms that cause a population to evolve. what fraction of moth colouration genes are the black allele). evolution is a generation-to-generation change in the frequencies of alleles within a population that shares a common gene pool. all of the moths of the same species living in an isolated forest represent a population. which account for variations between the phenotypes of the organisms. it is not evolving. the allele for black colour in a population of moths becoming more common. for example. A gene pool is the complete set of alleles for a gene in a single population. and the random combination of the alleles in these sex cells during fertilization. For example.[83] Such a population is said to be in Hardy-Weinberg equilibrium. it is useful to consider what conditions are required for a population not to evolve. [100] Viruses can also carry DNA between organisms. with the two different sets of chromosomes each being able to pair with an identical partner during meiosis. Conversely. although cases have been seen in many types of animals. this contributes to the spread of antibiotic resistance. but hybrids will be selected against and the species will remain distinct. Migration into or out of a population can change allele frequencies. gene flow may slow this process by spreading genetic differences between the populations. due to the two different sets of chromosomes being unable to pair up during meiosis.[95] Horizontal gene transfer is the transfer of genetic material from one organism to another organism that is not its offspring. Immigration may add new genetic material to the established gene pool of a population. closely related species may regularly interbreed. as when one bacteria acquires resistance genes it can rapidly transfer them to other species.[90] with the gray tree frog being a particularly well-studied example. oceans and deserts or even man-made structures such as the Great Wall of China. as with horses and donkeys mating to produce mules. which allows them to avoid inbreeding depression in small populations. however. In this case. which are usually of the same species. male lions leave the pride where they were born As barriers to reproduction between two diverging and take over a new pride to mate. it may still be possible for them to produce offspring. this is most common among bacteria.[94] Polyploids also have more genetic diversity. and plants. Gene flow is hindered by mountain ranges.[86] Examples of gene flow within a species include the migration and then breeding of organisms. or the exchange of pollen. allowing transfer of genes even across biological domains.Evolution 22 Gene flow Gene flow is the exchange of genes between populations.[96] In medicine. fungi. populations are required for the populations to become new species. When they mature. an important means of speciation in plants.[97] Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces cerevisiae and the adzuki bean beetle Callosobruchus chinensis may also have occurred. which has hindered the flow of plant genes.[87] Depending on how far two species have diverged since their most recent common ancestor.[88] Such hybrids are generally infertile.[102] .[91] Hybridization is. viable hybrids are occasionally formed and these new species can either have properties intermediate between their parent species. since polyploidy (having more than two copies of each chromosome) is tolerated in plants more readily than in animals. during the acquisition of chloroplasts and mitochondria. or possess a totally new phenotype.[101] Large-scale gene transfer has also occurred between the ancestors of eukaryotic cells and prokaryotes. Gene transfer between species includes the formation of hybrid organisms and horizontal gene transfer. causing gene flow between [85] prides. as well as introducing genetic variation into a population.[98] [99] An example of larger-scale transfers are the eukaryotic bdelloid rotifers. However.[92] [93] Polyploidy is important in hybrids as it allows reproduction. emigration may remove genetic material.[89] The importance of hybridization in creating new species of animals is unclear. which appear to have received a range of genes from bacteria. [106] If an allele increases fitness more than the other alleles of that gene.[1] However. result in a more uniform population. fitness is not the same as the total number of offspring: instead fitness is indicated by the proportion of subsequent generations that carry an organism's genes. organisms with traits that give them an advantage over their Natural selection of a population for dark colouration. then with each generation this allele will become more common within the population. It has often been called a "self-evident" mechanism because it necessarily follows from three simple facts: • Heritable variation exists within populations of organisms.[41] Natural selection Natural selection is the process by which genetic mutations that enhance reproduction become. Population bottlenecks.[103] Natural selection usually predominates in large populations. Consequently.[106] Fitness is measured by an organism's ability to survive and reproduce. while traits that do not confer an advantage are not passed on to the next generation. Genetic drift is the random change in the frequency of alleles. These conditions produce competition between organisms for survival and reproduction.[104] As a result. if an organism could survive well and reproduce rapidly. caused by the random sampling of a generation's genes during reproduction.[106] However. traits that were lost in the past may not re-evolve in an identical form (see Dollo's law). Conversely. more common in successive generations of a population. which is the number of individuals capable of breeding. where the population shrinks temporarily and therefore loses genetic variation. even if the direction of selection does reverse in this way. previously neutral or harmful traits may become beneficial and previously beneficial traits become harmful. the fitness of an allele is not a fixed characteristic.[109] [110] . The relative importance of natural selection and genetic drift in a population varies depending on the strength of the selection and the effective population size. Natural selection is the process which favors genes that aid survival and reproduction. These traits are said to be "selected for".[108] Importantly. The dominance of genetic drift in small populations can even lead to the fixation of slightly deleterious mutations. whereas genetic drift dominates in small populations.Evolution 23 Mechanisms The two main mechanisms that produce evolution are natural selection and genetic drift. • Organisms produce more offspring than can survive. • These offspring vary in their ability to survive and reproduce. and remain. this organism would make little genetic contribution to future generations and would thus have low fitness.[107] For example. and increased fecundity. but its offspring were all too small and weak to survive.[105] The central concept of natural selection is the evolutionary fitness of an organism. the lower fitness caused by having a less beneficial or deleterious allele results in this allele becoming rarer — they are "selected against". competitors pass these advantageous traits on. Examples of traits that can increase fitness are enhanced survival. if the environment changes. which determines the size of its genetic contribution to the next generation. changing population size can dramatically influence the course of evolution. in stabilizing selection there is selection against extreme trait values on both ends. as discussed below. which causes a decrease in variance around the average value and less diversity. that is. sexually selected traits. biotic diversity. which can replicate and spread throughout a genome. which is selection for any trait that increases mating success by increasing the attractiveness of an organism to potential mates. organisms slowly getting taller. for example. This broad understanding of nature enables scientists to delineate specific forces which. with distinct relationships to other parts of the system. and its geographic range. are more or less likely to survive.Evolution 24 Natural selection within a population for a trait that can vary across a range of values. which is a shift in the average value of a trait over time — for example. groups of organisms and species.Directional selection antlers. such as height.[121] . a founder of ecology.[117] [118] None of these are mutually exclusive and selection can act on multiple levels simultaneously. The first is directional selection. or position.in a given area interacting with the physical environment so that a flow of energy leads to clearly defined trophic structure. An active area of research is the unit of selection.Stabilizing selection 3. Finally.Disruptive selection some animal species. may allow the evolution of co-operation.. cells. A special case of natural selection is sexual selection. together.[119] An example of selection occurring below the level of the individual organism are genes called transposons. disruptive selection is selection for extreme trait values and often results in two different values becoming most common.[113] Traits that evolved through sexual selection are particularly prominent in males of A chart showing three types of selection. This would be when either short or tall organisms had an advantage. but not those of medium height..[115] Natural selection most generally makes nature the measure against which individuals.[111] Secondly. can be categorised into three different types. decreasing the survival of individual males. a system in which organisms interact with every other element.[120] Selection at a level above the individual. physical as well as biological. mating calls or bright colours that attract predators."[116] Each population within an ecosystem occupies a distinct niche. and individual traits. 1. with natural selection being proposed to work at the level of genes. Eugene Odum. its position in the food chain. "Nature" in this sense refers to an ecosystem. comprise natural selection.[105] [112] This would. such as group selection. individual organisms. in their local environment. defined an ecosystem as: "Any unit that includes all of the organisms. despite traits such as cumbersome 2. These relationships involve the life history of the organism. with selection against the average value.[114] This survival disadvantage is balanced by higher reproductive success in males that show these hard to fake. cause organisms to slowly become all the same height. and material cycles (ie: exchange of materials between living and nonliving parts) within the system. In mathematical terms. avoiding predators or attracting mates.[124] An example when genetic drift is probably of central importance in determining a trait is the loss of pigments from animals that live in caves. such as extinction and speciation.[123] The precise measure of population that is important is called the effective population size.[130] [131] However. Most prominent are the specific behavioral and physical adaptations that are the outcome of natural selection. which is evolution that occurs at or above the level of species.[128] Hence.[127] These investigations were prompted by the neutral theory of molecular evolution.[105] Simulation of genetic drift of 20 unlinked alleles in populations of 10 (top) and 100 (bottom). and the lower probability of animals that live far apart managing to mate with each other. These outcomes of evolution are sometimes divided into macroevolution. which is smaller evolutionary changes.[125] However. Genetic drift may therefore eliminate some alleles from a population due to chance alone.[122] The time for an allele to become fixed by genetic drift depends on population size. the number of animals that are too old or young to breed. As a result. This drift halts when an allele eventually becomes fixed. evolution produces new species through splitting ancestral populations of organisms into new groups that cannot or will not interbreed. Even in the absence of selective forces. a change that produces no obvious advantage or disadvantage in complete darkness. and microevolution. These adaptations increase fitness by aiding activities such as finding food. . when selective forces are absent or relatively weak. allele frequencies tend to "drift" upward or downward randomly (in a random walk).Evolution 25 Genetic drift Genetic drift is the change in allele frequency from one generation to the next that occurs because alleles in offspring are a random sample of those in the parents. most genetic changes in a population are the result of constant mutation pressure and genetic drift. it is usually difficult to measure the relative importance of selection and drift. or replacing the other alleles entirely. with fixation occurring more rapidly in smaller populations. in this model. In the longer term. Outcomes Evolution influences every aspect of the form and behavior of organisms. where most mutations only have small effects on fitness.[129] This form of the neutral theory is now largely abandoned. since it does not seem to fit the genetic variation seen in nature. usually by aiding their relatives or engaging in mutually beneficial symbiosis. either by disappearing from the population. Drift to fixation is more rapid in the smaller population. as well as from the role that chance plays in determining whether a given individual will survive and reproduce. Organisms can also respond to selection by co-operating with each other.[126] so the comparative importance of these two forces in driving evolutionary change is an area of current research. a more recent and better-supported version of this model is the nearly neutral theory. alleles are subject to sampling error. genetic drift can cause two separate populations that began with the same genetic structure to drift apart into two divergent populations with different sets of alleles. which proposed that most evolutionary changes are the result of the fixation of neutral mutations that do not have any immediate effects on the fitness of an organism. The effective population is always smaller than the total population since it takes into account factors such as the level of inbreeding. while a wide geographic range increases the chance of speciation.[141] and constitute the vast majority of Earth's biodiversity.[135] [136] [137] A common misconception is that evolution has goals or long-term plans. In this sense. evolution has no long-term goal and does not necessarily produce greater complexity. or the ability of horses to run fast and escape predators. in macroevolution. For example. realistically however. 1. with complex life only appearing more diverse because it is more noticeable. the traits of the entire species may be important.[147] [148] Also. the overwhelming majority of species are microscopic prokaryotes. the adaptation of horses' teeth to the grinding of grass.[138] [139] Although complex species have evolved. Adaptations are produced by natural selection.[150] 2. By using the term adaptation for the evolutionary process.[143] Indeed. versus macroevolutionary processes such as species selection acting on entire species and affecting their rates of speciation and extinction. with genetic changes causing antibiotic resistance by both modifying the target of the drug.[155] [156] and the soil bacterium Sphingobium evolving an entirely new metabolic pathway that degrades the synthetic pesticide pentachlorophenol. For instance. which form about half the world's biomass despite their small size.[157] [158] An interesting but still controversial idea is that some adaptations might increase the ability of organisms to generate genetic diversity and adapt by natural selection (increasing organisms' evolvability).and macroevolution is not a fundamental one – the difference is simply the time involved.[133] Thus. macroevolution is regarded as the outcome of long periods of microevolution.[152] Adaptation may cause either the gain of a new feature. the evolution of microorganisms is particularly important to modern evolutionary research. the distinction between micro.[154] Flavobacterium evolving a novel enzyme that allows these bacteria to grow on the by-products of nylon manufacturing. lessening the chance of it going extinct.[142] Simple organisms have therefore been the dominant form of life on Earth throughout its history and continue to be the main form of life up to the present day. the term adaptation may refer to a trait that is important for an organism's survival. within a species or population.[149] The following definitions are due to Theodosius Dobzhansky. Adaptation is the evolutionary process whereby an organism becomes better able to live in its habitat or habitats. Adaptedness is the state of being adapted: the degree to which an organism is able to live and reproduce in a given set of habitats.[132] In general.[151] 3.[159] [160] . the two senses of the word may be distinguished. by making it more likely that part of the population will become isolated. An adaptive trait is an aspect of the developmental pattern of the organism which enables or enhances the probability of that organism surviving and reproducing. An example that shows both types of change is bacterial adaptation to antibiotic selection.[134] However.[140] For example. and adaptive trait for the product (the bodily part or function).[153] Other striking examples are the bacteria Escherichia coli evolving the ability to use citric acid as a nutrient in a long-term laboratory experiment.[144] [145] 26 Adaptation Adaptation is one of the basic phenomena of biology. microevolution and macroevolution might involve selection at different levels – with microevolution acting on genes and organisms. since their rapid reproduction allows the study of experimental evolution and the observation of evolution and adaptation in real time. or the loss of an ancestral feature.Evolution such as adaptations. a large amount of variation among individuals allows a species to rapidly adapt to new habitats. or increasing the activity of transporters that pump the drug out of the cell.[146] and is the process whereby an organism becomes better suited to its habitat. they occur as a side effect of the overall number of organisms increasing. and simple forms of life still remain more common in the biosphere. the head has become so flattened that it assists in gliding from tree to tree—an exaptation.[172] [173] [174] [175] However. which developed an extremely flat head for hiding in crevices. or the limbs and wings of arthropods and vertebrates. within bat wings. many traits that appear to be simple adaptations are in fact exaptations: structures originally adapted for one function. can depend on a common set of homologous genes that control their assembly and function.[176] Within cells. squid and vertebrate eyes.[167] the non-functional remains of eyes in blind cave-dwelling fish. which were adapted from front leg bones: while c indicates vestigial leg bones. a and b label flipper bones.[161] Examples of vestigial structures in humans include wisdom teeth. natural selection will tend to force species to adapt to different ecological niches. due to the descent of all these structures from a common mammalian ancestor.[164] [165] During adaptation. but which coincidentally became somewhat useful for some other function in the process. Consequently.[132] Another example is the recruitment of enzymes from glycolysis and xenobiotic metabolism to serve as structural proteins called crystallins within the lenses of organisms' eyes. this is called deep homology. However. This may mean that. since all living organisms are related to some extent.[182] An area of current investigation in evolutionary developmental biology is the developmental basis of adaptations and exaptations. or other closely related species. such as embryonic bone structures that develop into the jaw in other animals instead forming part of the middle ear in mammals. such as a mutation in chickens causing embryos to grow teeth similar to those of crocodiles. The bones [161] sea.[171] and primitive reflexes. This is the result of a single ancestral structure being adapted to A baleen whale skeleton.[184] These studies have shown that evolution can alter development to create new structures.[185] It is also possible for structures that have been lost in evolution to reappear due to changes in developmental genes.[179] [180] A critical principle of ecology is that of competitive exclusion: no two species can occupy the same niche in the same environment for a long time. Examples include pseudogenes. structures with similar internal organization may have different functions in related organisms. for example. yet have a clear function in ancestral species. in this species.[169] and the presence of hip bones in whales and snakes. are very similar to those in mice feet and primate hands. two species of cichlid fish adapt to live in different habitats.[181] Consequently. suggesting an adaptation from land to function in different ways.Evolution 27 Adaptation occurs through the gradual modification of existing structures. some structures may lose their original function and become vestigial structures.[166] Such structures may have little or no function in a current species.[176] One example is the African lizard Holaspis guentheri.[162] However.[166] the vermiform appendix. molecular machines such as the bacterial flagella[177] and protein sorting machinery[178] evolved by the recruitment of several pre-existing proteins that previously had different functions.[168] wings in flightless birds.[170] the coccyx.[166] and other behavioral vestiges such as goose bumps.[183] This research addresses the origin and evolution of embryonic development and how modifications of development and developmental processes produce novel features. such as arthropod. which will minimise the competition between them for food. for example.[187] . as can be seen by looking at its near relatives.[163] even organs that appear to have little or no structural similarity.[186] It is now becoming clear that most alterations in the form of organisms are due to changes in a small set of conserved genes. [194] Other processes that may promote cooperation include group selection. allowing them to exchange nutrients with their hosts. which is where one organism acts to help raise a relative's offspring. these species can develop matched sets of adaptations.[190] Many cases of mutually beneficial interactions have evolved. the fungi actually grow inside plant cells. their uncontrolled growth causes cancer.[191] This is a reciprocal relationship as the plants provide the fungi with sugars from photosynthesis.[195] . When the interaction is between pairs of species. it is likely that its kin will also contain these alleles and thus those alleles will be passed on. This cycle of selection and response is called co-evolution. For instance. the evolution of one species causes adaptations in a second species. Here.[193] This activity is selected for because if the helping individual contains alleles which promote the helping activity.Evolution 28 Co-evolution Interactions between organisms can produce both conflict and co-operation. On an even smaller scale. the somatic cells that make up the body of an animal limit their reproduction so they can maintain a stable organism. in turn. termites and ants. cause new adaptations in the first species.[188] An example is the production of tetrodotoxin in the rough-skinned newt and the evolution of tetrodotoxin resistance in its predator. not all interactions between species involve conflict. which then supports a small number of the animal's germ cells to produce offspring. while sending signals that suppress the plant immune system. where cooperation provides benefits to a group of organisms. an extreme cooperation exists between plants and the mycorrhizal fungi that grow on their roots and aid the plant in absorbing nutrients from the soil. Here.[189] Common garter snake (Thamnophis sirtalis sirtalis) which has evolved resistance to tetrodotoxin in its amphibian prey. somatic cells respond to specific signals that instruct them whether to grow. such as bees. where sterile insects feed and guard the small number of organisms in a colony that are able to reproduce. the common garter snake. If cells ignore these signals and multiply inappropriately. or die.[45] Such cooperation within species may have evolved through the process of kin selection. an evolutionary arms race has produced high levels of toxin in the newt and correspondingly high levels of toxin resistance in the snake. such as a pathogen and a host. In this predator-prey pair. remain as they are. These changes in the second species then. or a predator and its prey. Here. Co-operation However.[192] Coalitions between organisms of the same species have also evolved. An extreme case is the eusociality found in social insects. Although properties such as genetics and morphology are used to help separate closely related lineages. which is the evolution of traits that promote mating within a species. This view is counterintuitive since the classical idea of species is still widely held. which can undergo parapatric speciation in response to localised metal pollution from mines. which is when two species become more distinct in appearance. which eventually produced complete reproductive isolation. which occurs when small populations of organisms become isolated in a new environment.[205] Here. speciation results from reproductive isolation followed by genealogical divergence. and this is called the species problem. Instead. which occurs in populations initially isolated geographically. the exact definition of the term "species" is still The four mechanisms of speciation. an observable similar group of individuals.[199] Typically the actual focus on biological study is the population. Here. but the definition used is a pragmatic choice that depends on the particularities of the species concerned. Selection under these conditions can produce very rapid changes in the appearance and behaviour of organisms. plants evolve that have resistance to high levels of metals in the soil.[204] The third mechanism of speciation is parapatric speciation. the founder effect causes rapid speciation through both rapid genetic drift and selection on a small gene pool. One example is the grass Anthoxanthum odoratum. with a species seen as a class of organisms exemplified by a "type specimen" that bears all the traits common to this species. There are four mechanisms for speciation.[206] . rather than a species.[196] Generally this occurs when there has been a drastic change in the environment within the parental species' habitat.[203] The second mechanism of speciation is peripatric speciation. Speciation has been observed multiple times under both controlled laboratory conditions and in nature.[201] [202] As selection and drift act independently on populations isolated from the rest of their species.Evolution 29 Speciation Speciation is the process where a species diverges into two or more descendant species. Biologists have proposed a range of more precise definitions. speciation results from the evolution of mechanisms that reduce gene flow between the two populations. controversial. particularly in [198] [199] prokaryotes.[200] In sexually reproducing organisms. Instead. This differs from allopatric speciation in that the isolated populations are numerically much smaller than the parental population. an observable interacting group of organisms. a species is now defined as a separately evolving lineage that forms a single gene pool.[196] Evolutionary biologists view species as statistical phenomena and not categories or types. such as by habitat fragmentation or migration. separation may eventually produce organisms that cannot interbreed. Indeed. this definition has fuzzy [197] boundaries. Selection against interbreeding with the metal-sensitive parental population produced a gradual change in the flowering time of the metal-resistant plants. as well as character displacement. This is similar to peripatric speciation in that a small population enters a new habitat. but differs in that there is no physical separation between these two populations. Selection against hybrids between the two populations may cause reinforcement. The most common in animals is allopatric speciation. to form polyploids. with natural selection and genetic drift acting most strongly on organisms undergoing speciation in novel habitats or small populations. since each parent's chromosomes are represented by a pair already.[218] These extinctions have happened continuously throughout the history of life. and .[208] One type of sympatric speciation involves cross-breeding of two related species to produce a new hybrid species.[211] This happened about 20.[217] and extinction appears to be the ultimate fate of all species.[207] Generally. which accounts for the pattern in the fossil record of short "bursts" of evolution interspersed with relatively long periods of stasis. Extinction is not an unusual event.000 years ago. as half the doubled chromosomes will be unmatched when breeding with undoubled organisms. This form is rare since even a small amount of gene flow may remove genetic differences between parts of a population.[216] Nearly all animal and plant species that have lived on Earth are now extinct. to allow reproductive isolation to evolve. during which the non-avian dinosaurs went extinct. and therefore rarely being preserved as fossils. is Tyrannosaurus rex.[93] Speciation events are important in the theory of punctuated equilibrium. with approximately Cretaceous period. but the earlier Permian–Triassic Cretaceous–Tertiary extinction event at the end of the extinction event was even more severe. the periods of stasis in the fossil record correspond to the parental population. it is more Geographical isolation of finches on the common in plants because plants often double their number of Galápagos Islands produced over a dozen new species. However.[219] The Holocene extinction event is an ongoing mass extinction associated with humanity's expansion across the globe over the past few thousand years.[214] In this theory. 96 percent of species driven to extinction.[209] This allows the chromosomes from each parental species to form matching pairs during meiosis. As a result.[219] The Cretaceous–Tertiary extinction event. Non-avian dinosaurs died out in the the most well-known.[210] An example of such a speciation event is when the plant species Arabidopsis thaliana and Arabidopsis arenosa cross-bred to give the new species Arabidopsis suecica. chromosome doubling within a species may be a common cause of reproductive isolation. as species regularly appear through speciation. where species remain relatively unchanged. although the rate of extinction spikes in occasional mass extinction events. which allows the study of the genetic mechanisms involved in this process. This is because during meiosis the homologous chromosomes from each parent are from different species and cannot successfully pair. This is not common in animals as animal hybrids are usually sterile. speciation and rapid evolution are linked.[213] Indeed. in sympatric speciation species diverge without geographic isolation or changes in habitat. Present-day extinction rates are 100–1000 times greater than the background rate. sympatric speciation in animals requires the evolution of both genetic differences and non-random mating. chromosomes. and disappear through extinction.[215] Extinction Extinction is the disappearance of an entire species.Evolution 30 Finally. and the organisms undergoing speciation and rapid evolution are found in small populations or geographically restricted habitats.[212] and the speciation process has been repeated in the laboratory. with their diversity the product of a long series of speciation and extinction events. this could produce species selection.[222] The role of extinction in evolution is not very well understood and may depend on which type of extinction is considered. this approach is most successful for organisms that had hard body parts. may be the result of competition between species for limited resources (competitive exclusion).[223] 31 Evolutionary history of life Origin of life The origin of life is a necessary precursor for biological evolution. modern research has suggested that. but proposals include self-replicating molecules such as RNA. bones or teeth.[219] The causes of the continuous "low-level" extinction events. due to horizontal gene transfer. but instead of acting as a selective force. their fossils do not provide information on their ancestry. However. or the identity and nature of any last universal common ancestor or ancestral gene pool. they have geographic distributions that cannot be explained by local adaptation. constitute the morphological. the structure of the first living things. Fossils. vestigial traits with no clear purpose resemble functional ancestral traits. such as shells. Second.[232] [233] Past species have also left records of their evolutionary history.[224] The current scientific consensus is that the complex biochemistry that makes up life came from simpler chemical reactions. they drastically reduce diversity in a nonspecific manner and promote bursts of rapid evolution and speciation in survivors.[163] [230] Current species are a stage in the process of evolution. that organisms can be classified using these similarities into a hierarchy of nested groups – similar to a family tree.[117] The intermittent mass extinctions are also important.[228] and the assembly of simple cells.[231] The common descent of organisms was first deduced from four simple facts about organisms: First.[234] By comparing the anatomies of both modern and extinct species. which form the majority of extinctions. this "tree of life" may be more complicated than a simple branching tree since some genes have spread independently between distantly related species.[225] Not much is certain about the earliest developments in life. along with the comparative anatomy of present-day organisms. with the fitter species surviving and the other species being driven to extinction. but it is unclear how this occurred. the diversity of life is not a set of completely unique organisms. but understanding that evolution occurred once organisms appeared and investigating how this happens does not depend on understanding exactly how life began. there is no scientific consensus on how life began. record. Further. . or anatomical. paleontologists can infer the lineages of those species. as prokaryotes such as bacteria and archaea share a limited set of common morphologies. and finally.[19] If one species can out-compete another.[226] [227] Consequently. The hominoids are descendants of a common ancestor.[14] However. Third.Evolution up to 30 percent of species may be extinct by the mid 21st century. but organisms that share morphological similarities.[221] global warming may further accelerate it in the future.[220] Human activities are now the primary cause of the ongoing extinction event.[229] Common descent All organisms on Earth are descended from a common ancestor or ancestral gene pool. These came from ancient bacteria being engulfed by the ancestors of eukaryotic cells. Snel B. all living cells use the same basic set of nucleotides and amino acids.[240] The eukaryotes were the next major change in cell structure. a remarkable amount of biological diversity appeared over approximately 10 million years. and eukaryotes red.[247] About 500 million years ago.Ciccarelli FD. plants and fungi colonised the land.[142] . The three domains are bacteria evolving into either coloured.[242] An independent second engulfment of cyanobacterial-like organisms led to the formation of chloroplasts in algae and plants.[235] The development of molecular genetics has revealed the record of evolution left in organisms' genomes: dating when species diverged through the molecular clock produced by mutations.[249] Amphibians first appeared around 300 million years ago. and archaea until about 610 million years ago when multicellular organisms began to appear in the oceans in the Ediacaran period. with the majority of both biomass and species being prokaryotes. the majority of types of modern animals appeared in the fossil record.Evolution More recently. evidence for common descent has come from the study of biochemical similarities between organisms. slime moulds and myxobacteria. with the 1283–87. in a cooperative association called Evolutionary tree showing the divergence of modern species from their common ancestor endosymbiosis. von Mering C.1126/science. then mammals around 200 million years ago and birds around 100 million years ago (both from "reptile"-like lineages). brown algae.[238] [239] No obvious changes in morphology or cellular organization occurred in these organisms over the next few billion years. Science 311 (5765): underwent co-evolution.[245] Soon after the emergence of these first multicellular organisms. as well as unique lineages that subsequently became extinct. smaller organisms similar to the types that evolved early in this process continue to be highly successful and dominate the Earth. bacteria and the host cell then "Toward automatic reconstruction of a highly resolved tree of life". PMID 16513982. mitochondria or hydrogenosomes. followed by early amniotes. For example.[238] [244] The evolution of multicellularity occurred in multiple independent events.7 billion years ago.[243] It is unknown when the first eukaryotic cells appeared though they first emerged between 1. archaea green. it is generally accepted that prokaryotes inhabited the Earth from approximately 3–4 billion years ago. and were soon followed by arthropods and other animals.[237] 32 Evolution of life Despite the uncertainty on how life began.[236] For example. Here. in organisms as diverse as sponges. with bacteria blue. Bork P (2006).[248] Insects were particularly successful and even today make up the majority of animal species. doi:10.[102] [241] The engulfed in the center. including the accumulation of oxygen in the atmosphere from photosynthesis.6 – 2. prokaryotes. cyanobacteria. Creevey CJ. The history of life was that of the unicellular eukaryotes.[246] Various triggers for the Cambrian explosion have been proposed. these DNA sequence comparisons have revealed that humans and chimpanzees share 96% of their genomes and analyzing the few areas where they differ helps shed light on when the common ancestor of these species existed. in an event called the Cambrian explosion.1123061. despite the evolution of these large animals. Doerks T. However. Nowadays. dramatic increases in the power of computers allowed practical applications. since different mutations had occurred in the isolated populations that had evolved in different caves. . social and religious implications of evolution. such selection has become a vital part of genetic engineering. the fact that organisms evolve is uncontested in the scientific literature and the modern evolutionary synthesis is widely accepted by scientists. Breeding together different populations of this blind fish produced some offspring with functional eyes.[257] Artificial evolution became a widely recognised optimization method as a result of the work of Ingo Rechenberg in the 1960s and early 1970s.[256] As evolution can produce highly optimised processes and networks. simulations of evolution using evolutionary algorithms and artificial life started with the work of Nils Aall Barricelli in the 1960s.[259] As academic interest grew.[263] While various religions and denominations have reconciled their beliefs with evolution through concepts such as theistic evolution.[19] However. which lacks red blood cells.[253] For example.[254] This helped identify genes required for vision and pigmentation. it has many applications in computer science.[251] More recently. and in particular the understanding of how organisms evolve through natural selection.[260] Evolutionary algorithms are now used to solve multi-dimensional problems more efficiently than software produced by human designers. and also to optimise the design of systems. comparing the genome of the Antarctic icefish. which is the intentional selection of certain traits in a population of organisms. It is also possible to use repeated rounds of mutation and selection to evolve proteins with particular properties. the Mexican tetra is an albino cavefish that lost its eyesight during evolution. are not distinctly As evolution became widely accepted in the 1870s.[252] Understanding the changes that have occurred during organism's evolution can reveal the genes needed to construct parts of the body.Evolution 33 Applications Evolutionary biology. Here. particularly after the publication of On the Origin of Species in 1859. the most controversial aspect of evolutionary biology is the implication of human evolution that human mental and moral faculties. who published a series of papers on simulation of artificial selection. which had been thought purely spiritual. there are creationists who believe that evolution is contradicted by the creation myths found in their respective religions and who raise various objections to evolution. and was extended by Alex Fraser. is an area of science with many practical applications.[132] [264] [265] As had been demonstrated by responses to the publication of Vestiges of the Natural History of Creation in 1844. evolution remains a contentious concept for some theists.[255] Similarly. such as crystallins and the melanocortin 1 receptor. caricatures of Charles Darwin with an ape or monkey body symbolised [262] evolution.[258] Genetic algorithms in particular became popular through the writing of John Holland. Humans have used artificial selection for thousands of years in the domestication of plants and animals. including the automatic evolution of computer programs. in a process called directed evolution. genes which may be involved in human genetic disorders.[250] A major technological application of evolution is artificial selection. who used evolution strategies to solve complex engineering problems. with selectable markers such as antibiotic resistance genes being used to manipulate DNA in molecular biology. such as modified enzymes or new antibodies. the idea that life had evolved was an active source of academic debate centered on the philosophical.[261] Social and cultural responses In the 19th century. to close relatives such as the Antarctic rockcod revealed genes needed to make these blood cells. Acad.W. sdsc. but it was gradually re-introduced about a generation later and legally protected with the 1968 Epperson v. Retrieved 2007-03-18. Palmer (January 2007). doi:10. Evolution. but it returned in the form of intelligent design. Rivera. Cradle of life: the discovery of Earth's earliest fossils. On the Origin of Species (http:/ / darwin-online.1073/pnas. UK. J. Lynn (1998). American Association for the Advancement of Science. (2005). Nature 465 (7295): 219–222.L.F. Princeton. The Scopes Trial decision of 1925 caused the subject to become very rare in American secondary biology textbooks for a generation. gov/ articlerender.[271] [272] [273] 34 See also • Current research in evolutionary biology References [1] Futuyma. Charles (1859). Oxford University Press.[269] Another example somewhat associated with evolutionary theory that is now widely regarded as unwarranted is "Social Darwinism".95.S. PMID 17494753. a derogatory term associated with the 19th century Malthusian theory developed by Whig philosopher Herbert Spencer. PMID 9618502. "Darwin's greatest discovery: design without designer" (http:/ / www. (February. and contemporary scientists and philosophers consider these ideas to be neither mandated by evolutionary theory nor supported by data. the competing religious belief of creationism was legally disallowed in secondary school curricula in various decisions in the 1970s and 1980s. edu/ ~shindyal/ ejc121304.96. M.3801. pubmedcentral.[266] While other scientific fields such as cosmology[267] and Earth science[268] also conflict with literal interpretations of many religious texts.0701072104. . PMID 10097118. Massachusetts: Sinauer Associates. doi:10. Aaron O. doi:10. London: John Murray.1093/jxb/erl148. edu/ courses/ mb437_537_2004_fall/ docs/ uprooting. Sci. fcgi?tool=pmcentrez& artid=22660). PMID 17030541.1038/nature09014. a religious conflict focusing on politics and public education." (http:/ / www. shtml) from the Understanding Evolution University of California at Berkeley [8] Evolution 101: Peripatric Speciation (http:/ / evolution. Journal of Experimental Botany 58 (1): 1–9. It was later expanded by others into ideas about "survival of the fittest" in commerce and human societies as a whole. PMID 20463738. [11] Theobald. . London: John Murray. Evolution:The History of an Idea. .A. C. uk/ content/ frameset?itemID=F381& viewtype=text& pageseq=20) (3rd ed. The symbiotic planet: a new look at evolution. J. org/ content/ 104/ suppl. Proc Natl Acad Sci U S A 96 (7): 3801–6. edu/ evosite/ evo101/ IIID2Genesdrift. [10] Woese. nih.[270] However.1073/pnas. "Horizontal gene transfer among genomes: the complexity hypothesis. uk/ content/ frameset?itemID=F373& viewtype=text& pageseq=16) (1st ed. xiii.C. "AAAS Resolution: Present Scientific Status of the Theory of Evolution" (http:/ / archives. (2003). [13] Bowler. 1. pdf). R. ISBN 0-87893-187-2. .7. Dover Area School District case. Natl. to be excluded once again in the 2005 Kitzmiller v. U. D. On the Origin of Species (http:/ / darwin-online. Evolution by association: a history of symbiosis. ISBN 0801413192. PMC 1876431. evolutionary biology experiences significantly more opposition from religious literalists. edu/ evosite/ evo101/ VC1cPeripatric. doi:10.). Scientific American 282 (2): 90–95. fcgi?tool=pmcentrez& artid=22375).1073/pnas. [15] AAAS Council (December 26.).6854. [6] Ayala FJ (2007). 2000).. edu/ evosite/ evo101/ IIID1Samplingerror. org. London. 1/ 8567. ISBN 0-52023693-9. (1994). Since then. [4] Margulis. Charles (1861). (1998). pnas. p. "The Universal Ancestor" (http:/ / www. and led to claims that social inequality. shtml) and Effects of Genetic Drift (http:/ / evolution. Douglas J. org/ docs/ resolutions. and Jeffrey D. pubmedcentral. php?doc_id=450). Proc.. University of California Press.. racism. pdf). PMC 22660. PMC 22375. (1999). (1999).12. shtml) from the Understanding Evolution webpages made by the University of California at Berkeley [9] Schopf. msu. gov/ articlerender. . these ideas contradict Darwin's own views. PMID 10710791. "Horizontal gene transfer in plants" (http:/ / www. Arkansas decision. Inc. . Related earlier ideas were acknowledged in Darwin. The teaching of evolution in American secondary school biology classes was uncommon in most of the first half of the 20th century. W. montana. [2] Jain. [14] Darwin. Peter J.[13] In some countries—notably the United States—these tensions between science and religion have fueled the current creation-evolution controversy. [3] Richardson. org. and imperialism were justified. sexism. berkeley. . berkeley. PNAS 95 (12): 6854–6859. full). Sunderland. berkeley.A. Weidenfeld & Nicolson. [5] Sapp. Lake. 1922). [12] Doolittle. 104 (Suppl 1): 8567–73. nih. [7] Evolution 101:Sampling Error and Evolution (http:/ / evolution. doi:10. J. ISBN 0801413192. "A formal test of the theory of universal common ancestry". aaas.Evolution separated from those of other animals. "Uprooting the tree of life" (http:/ / shiva. (2010). Turner P (2006).W. . Joint statement issued by the national science academies of 67 countries. doi:10. PMID 18319743. Genet. "Do we need an extended evolutionary synthesis?".. The illustrated origin of species. Rev. PMC 1692205. "Genetics: what is a gene?". [20] "Special report on evolution" (http:/ / www. fcgi?tool=pmcentrez& artid=2689140). PMID 15241603. Lin M (2006). File/ Master/ 6/ 150/ Evolution statement. . as controlled by Natural Selection" (http:/ / darwin-online. pubmedcentral. 108. The Mendelian Revolution: The Emergence of Hereditarian Concepts in Modern Science and Society. net/ Object. J.2004. uk/ content/ frameset?itemID=F350& viewtype=text& pageseq=1). including the United Kingdom's Royal Society [17] Board of Directors. Lond. A History of the Life Sciences (http:/ / books. [37] Oetting WS.. [36] Visscher PM. Nature 446 (7133): 279–83. Frudakis TN (2004). Acad. Norton & Company. nih. org. National Center for Science Education. Genet.1998.1099/mic. Baltimore: Johns Hopkins University Press. Evolution 61 (12): 2743–9. "Historical study: Johann Gregor Mendel 1822–1884". American Association for the Advancement of Science.00246.. (2006). PMID 16485021. "The clinical spectrum of albinism in humans". com/ ?id=YKJ6gVYbrGwC& printsec=frontcover#v=onepage& q) (Third ed. Philos.1038/nrg1804. PMID 17924956. "On the Tendency of Species to form Varieties. "Hybrid speciation". [40] Richards EJ (May 2006). pubmedcentral.1016/j. . google. [26] Leakey. Genet. King RA (1996). 7 (3): 229–37. [34] Sturm RA. Niklas K (2004). .1558-5646. textbookleague.29013-0. [30] Quammen.1007/s00114-004-0515-y.1073/pnas. 20 (8): 327–32.x. Retrieved 2007-04-25. Rev. (http:/ / www. PMC 2575274.. Charles (1979). "Epistasis—the essential role of gene interactions in the structure and evolution of genetic systems" (http:/ / www. "Effects of the increased Use and Disuse of Parts. [22] Terrall. "Factors affecting levels of genetic diversity in natural populations" (http:/ / www. [23] Wallace. and on the Perpetuation of Varieties and Species by Natural Means of Selection" (http:/ / darwin-online.1038/nrg1834. 2006. (1989).it is the steady accumulation of beneficial differences which has given rise to. "DNA secretion and gene-level selection in bacteria". "Eye colour: portals into pigmentation genes and ancestry". Am. .S.0171. ISBN 0-571-14586-8. nytimes.1111/j.0200. nih. S (1984). . Nat. pubmedcentral. "Heritability in the genomics era—concepts and misconceptions". org. Darwin. Microbiology (Reading. A (1858).1038/nrg2452. Evolution and the Genetics of Populations.1016/1357-4310(96)81798-9. org/ ). doi:10.0711445105. ISBN 0-226-91038-5. Charles (1872).010. doi:10. *Mallet J (2007). doi:10. 105 (33): 11833–8.1002/ajmg. "Functional mapping – how to map and study the genetic architecture of dynamic complex traits". doi:10. Trans. doi:10. p. 9 (4): 255–66. [19] Kutschera U. [32] Pigliucci M (December 2007). textbookleague. PMID 9533122. The University of Chicago Press. htm). Retrieved 2008-01-23. Med. The University of Chicago Press.1098/rstb. . PMID 16946263. Wray NR (April 2008).. fcgi?tool=pmcentrez& artid=2575274). [35] Pearson H (2006). Nat. doi:10. Sci. com/ 2006/ 08/ 27/ books/ review/ Desmond.tig.1038/nrg2322.0. [42] Draghi J. The Man who flattened the Earth: Maupertuis and the sciences in the Enlightenment. [41] Harwood AJ. p. gov/ articlerender. Nat.Evolution [16] "IAP Statement on the Teaching of Evolution" (http:/ / www. (September/October 1994). nih. Darwin: An intimate portrait of Charles Darwin and the making of his theory of evolution. ISBN 9780203911006. doi:10.1038/nature05706. Peter J. [28] Magner. [21] Wright. Retrieved 2007-12-28. U. 9 (11): 855–67. doi:10. "Systemic darwinism" (http:/ / www. B. PMID 18852697. t. doi:10. Trends Genet. "Statement on the Teaching of Evolution" (http:/ / www. Michael T. The Textbook League. aaas. The reluctant Mr. NY: W. CRC Press. "The modern theory of biological evolution: an expanded synthesis". ISBN 978-0226793610. Richard E. ISBN 978-0801838880.1038/441398a. "Nonsense in schoolbooks: 'The Imaginary Lamarck'" (http:/ / www. PMID 16724031. New Scientist.2006. Harwood. Retrieved 2007-05-13.A. Soc. D. Proc. Volume 1: Genetic and Biometric Foundations.1098/rsnr. html?n=Top/ Reference/ Times Topics/ People/ D/ Darwin. Rev.) 152 (Pt 9): 2683–8. Journal of the Proceedings of the Linnean Society of London. pdf) (PDF). American Association for the Advancement of Science (2006-02-16). 35 . newscientist. Nature 441 (7092): 398–401. gov/ articlerender. PMC 2689140. Naturwissenschaften 91 (6): 255–76. doi:10.). doi:10. gov/ articlerender. PMID 16534512. doi:10. J (1998). org/ news/ releases/ 2006/ pdf/ 0219boardstatement.06. PMID 18697926. (2002). The Textbook Letter (http:/ / www. Genet. uk/ content/ frameset?viewtype=text& itemID=F391& pageseq=136). 353 (1366): 177–86. Molecular medicine today 2 (8): 330–5. [29] Weiling F (1991). 17-18 [27] Ghiselin. The Interacademy Panel on International Issues. org/ 54marck. Marcel Dekker. Lois N. from the world's largest general scientific society [18] "Statements from Scientific and Scholarly Organizations" (http:/ / ncse. M (2002). Rev. pdf) (PDF). 6th edition. . doi:10. 7 (5): 395–401. "Inherited epigenetic variation—revisiting soft inheritance". Zoology 3: 53–62. interacademies. 2008-01-19. Sci. doi:10. Hill WG. com/ media/ voices/ science). [31] Bowler. PMID 1887835. Nat. discussion 26. com/ topic/ evolution). PMID 17361174. [38] Phillips PC (November 2008). [39] Wu R. [24] The chapter on variation in 'Origin of Species' ends "Whatever the cause may be of each slight difference between the offspring and their parents.1320400103. John Murray.. fcgi?tool=pmcentrez& artid=1692205). PMID 8796918. The Origin of Species. Engl. Charles Robert) New York. PMID 15262401. Genet. Natl. London: Faber. 40 (1): 1–25. [33] Winther RG (August 2008).2007. Biol.the modifications of structure" Origin ch 5 [25] Darwin. Brilliant MH. R. PMC 2765273.2006. "Homologous pairing and strand exchange in genetic recombination". [59] Siepel A (October 2009).200700751.1016/S0098-2997(00)00007-8. "Prevalence of positive selection among nearly neutral amino acid replacements in Drosophila" (http:/ / www. long). gov/ articlerender. [61] Long M.1007/s00239-006-0045-7. PMID 16950096. full). doi:10.1002/1521-1878(200012)22:12<1057::AID-BIES3>3. 1/ 6535. doi:10. org/ content/ 19/ 10/ 1693.1016/j. R. Lond. Strub K (2006). Genome Res. Rev. cshlp.1002/cbic.CO.07.1126/science. PMID 17075697. Wang X.1016/S0022-2836(02)00109-2. doi:10. Genet. Aspects Med. Drosophila. doi:10. [49] Sniegowski P. [62] Wang M. 74: 867–900.1146/annurev. 2 (7): E206.008. B.1093/nar/gkl706. [63] Weissman KJ. Genet. Waldor M (2004). 353 (1366): 187–98.0501847102.0020206. html). "Testing the chromosomal speciation hypothesis for humans and chimpanzees" (http:/ / www. PMID 11173079.2008. and mosquitoes" (http:/ / www.1073/pnas. Natl. Tyler DM (2008). J. Thornton K. Biol. "Levels of genetic polymorphism: marker loci versus quantitative traits" (http:/ / www. [70] Peters AD.1101/gr. "Rapid evolution of RNA genomes". doi:10.13910.pbio.002201. [54] Harrison P.1101/gr. gov/ articlerender. England) 12 (Pt 3b): 541–7. [53] Carroll SB. [52] Hastings. Science 309 (5735): 764–7. full). Sci. Macpherson JM. Parsch J. JR. [47] Burrus V. PMC 449868.1073/pnas. Annu. [55] Bowmaker JK (1998). Tregenza T (1998). Gerrish P. com/ cr/ journal/ v18/ n10/ full/ cr2008278a. Genome Res. "Comparative genomic analysis of human and chimpanzee indicates a key role for indels in primate evolution". SM. Holland JJ (1999). VandePol S (1982). Microbiol. Philos. Petrov DA (2005). [64] Zhang J. "Alu elements as regulators of gene expression" (http:/ / www.0701572104. Sci.1126/science. pubmedcentral. PMID 19797681. [57] Hurles M (July 2004). 63 (5): 682–90. pubmedcentral. doi:10. doi:10. Sci.1371/journal. P J.1038/cr. doi:10. Coluzzi M (2005). Gerstein M (2002). "The role of selfish genetic elements in eukaryotic evolution". Spindler K. U. PLoS Biol.2008. Genome Res. Res. Trans. PMC 1871816. doi:10. . 18 (10): 985–96. Shaver A (2000). nih. PMID 17020921. Genetics 10 (8): 551–564. Second Edition.1038/nrg1204. "Gene duplication: the genomic trade in spare parts" (http:/ / www.1038/35084545. "The evolution of mutation rates: separating causes from consequences". "The molecular biology of cancer". .11. pnas.A. doi:10. [50] Drake JW. [65] Ayala FJ. Acad.96.2004. [51] Holland J.cub. Genet.4. Cell Res. ISBN 1-4051-1950-0.1002/bies. Zhang Z. doi:10. nih. Hartl DL (2007). doi:10. Cavelier L. nature. PMID 10207154. fcgi?tool=pmcentrez& artid=1692210). Acad. "Studying genomes through the aeons: protein families. org/ content/ 102/ suppl. PMID 16051794. [46] Aminetzach YT. Caetano-Anollés G (2009). gov/ articlerender. G (2009). "Protein families and their evolution-a structural perspective". PMID 19597530. PMID 15123584.120182.A. pubmedcentral.317 (inactive 2009-11-14). . 36 . doi:10.7041255. PMC 479111. "Mechanisms of change in gene copy number" (http:/ / www.A. PMID 9533123.1112699. pubmedcentral. Otto SP (2003).1016/j. "Pesticide resistance via transposition-mediated adaptive gene truncation in Drosophila". PMID 12766942. Biochem. PMID 19141283. 104 (16): 6504–10. PMC 1131864. PMID 10570172.0. nih. [67] Häsler J. Rev. doi:10.74. nih. Eye (London. "Protein-protein interactions in multienzyme megasynthetases". Ira. "Chromosome speciation: humans. org/ content/ 9/ 4/ 317. Proc. Rev. [45] Bertram J (2000). Grenier J. Acad. Nature Reviews. pseudogenes and proteome evolution". Okamura K. pubmedcentral.1101/gr. PMC 1636486.133029. Lupski. doi:10. 96 (24): 13910–3. Natl.ge.1891104. doi:10. Rev. Mol. Nucleic Acids Res. fcgi?tool=pmcentrez& artid=449868). Betrán E. Bioessays 22 (12): 1057–66.098376. doi:10. PMID 18711447. [58] Liu N. [56] Gregory TR. Nat. "Mutation rates among RNA viruses" (http:/ / www.Evolution [43] Butlin RK. [66] Hurst GD. J Mol Biol 318 (5): 1155–74. Annu. PMID 11483984. Mol. "Evolution of colour vision in vertebrates". PMC 2712117. [48] Sawyer SA. PMID 15252449.082803. Proc.1146/annurev. nih. Oxford: Blackwell Publishing. Nichol S. fcgi?tool=pmcentrez& artid=479111).012. gov/ articlerender. doi:10. doi:10. Horodyski F. Sevov M.biochem. "Shaping bacterial genomes with integrative and conjugative elements". Evol. Soc. Bergström TF (2006). From DNA to Diversity: Molecular Genetics and the Evolution of Animal Design.10291.9. Johnson T. . PMID 15207870.1016/j. [68] Radding C (1982).str. pnas. Science 215 (4540): 1577–85.. 102 (Suppl 1): 6535–42. 2 (8): 597–606. doi:10. Chembiochem 9 (6): 826–48.2-W. full).01. pubmedcentral. Nat. PMC 2864001. "Evolution of sex: why do organisms shuffle their genotypes?". cshlp. PMID 15954844.0201.S. [60] Orengo CA. org/ content/ 96/ 24/ 13910. Bioessays 25 (6): 533–7. U. fcgi?tool=pmcentrez& artid=2864001). "The evolution and functional diversification of animal microRNA genes" (http:/ / www.1098/rstb. PMID 18357594. PMC 1692210. PMID 7041255. 155 (5): 376–86. "Liberating genetic variance through sex". .resmic.16.1998.109. 19 (10): 1693–5. 21 (6): 167–223. Werren JH (2001). Proc.S. 16: 405–37. 9 (4): 317–24. 14 (5): 845–51. doi:10. [69] Agrawal AF (2006). Podlaha O (2004). 34 (19): 5491–7. 4 (11): 865–75. Thornton JM (2005). Wang W (November 2003). "Darwinian alchemy: Human genes from noncoding DNA" (http:/ / genome. PMID 14634634. "The origin of new genes: glimpses from the young and old". Müller R (April 2008). Grabau E. PMID 15851677. fcgi?tool=pmcentrez& artid=1871816). doi:10. Biol. PMID 9775215. 16 (17): R696. Sci.278. Weatherbee SD (2005). Curr. doi:10. PMID 6297377. Hebert PD (1999). U. doi:10. "The evolutionary mechanics of domain organization in proteomes and the rise of modularity in the protein world". gov/ articlerender.1038/nrg2593. doi:10.1073/pnas.24. gov/ articlerender. doi:10. PMID 17409186. [44] Wetterbom A.S. nih.063. PMID 12083509. fcgi?tool=pmcentrez& artid=1636486). PMID 11084621. Rosenberg. Natl. Structure 17 (1): 66–78. PMC 24164. "The modulation of DNA content: proximate causes and ultimate consequences" (http:/ / genome. "Evidence for heterogeneity in recombination in the human pseudoautosomal region: high resolution analysis by sperm typing and radiation-hybrid mapping" (http:/ / www. 97 (13): 7051–7. doi:10. PMID 14616063. Plant Mol. "The state of affairs in the kingdom of the Red Queen". html).. Wang J. nih. Biological Sciences 355 (1403): 1553–1562. gov/ articlerender. uk/ 2/ hi/ science/ nature/ 7039478. pubmedcentral. gov/ articlerender.1144363. fcgi?tool=pmcentrez& artid=2517139). "Genetics and the fitness of hybrids". pubmedcentral. umt. [84] Bright. O'Brieni SJ. doi:10. Kerry (2006). Chen Z. Retrieved 2008-01-06. dbs. PMC 34383. "The role of genetic and genomic attributes in the success of polyploids" (http:/ / www. PMID 17932297. edu/ synthetic/ synth_2.0050087.6800237. nih. Series B. fcgi?tool=pmcentrez& artid=1288109).1038/351562a0.hdy. Hum. Charlesworth D (November 2000). Philos. Curr.2000. "The contribution of the mule to scientific thought".pbio. doi:10. "The advantages and disadvantages of being polyploid". PMID 15800622. gov/ articlerender.084247. Genetics 164 (3): 1099–118. doi:10.1038/sj. fcgi?tool=pmcentrez& artid=1692900). [81] Salathé M. [86] Morjan C. PLoS Biol. pubmedcentral. fcgi?tool=pmcentrez& artid=2600545). Mutat. doi:10. Boschetti C (April 2007). Genet. PMID 12871918. Lond.1371/journal. Pound GE.1098/rstb. Sci.1098/rstb.007. Res. "Lateral gene transfer and the origins of prokaryotic groups". Ecol. Trends Ecol.010. doi:10. Retrieved 2007-12-30. 5 (4): e87.cub. doi:10.09. "Hardy-Weinberg Equilibrium Model" (http:/ / anthro. 37 .1038/nrg749. Science 318 (5848): 268–71. palomar. Rieseberg L (2004).1073/pnas. doi:10.gde. [83] O'Neil. Biol. doi:10.1016/j. [88] Short RV (1975). nih.1525-142X. . Curr Opin Genet Dev 17 (6): 505–12.102401. (June 1991). PMID 10860970. Genet. nih. Bonhoeffer S (August 2008).2007. PMC 2517139. B. [80] Butlin R (April 2002). PMID 14195748. 13 (6): 1341–56. "Independently evolving species in asexual bdelloid rotifers" (http:/ / www. doi:10.7051. Sci. [79] Doncaster CP. (Amst. [77] Charlesworth B.tree. [96] Boucher Y. Lay summary (http:/ / www. "Polyploid hybrids: multiple origins of a treefrog species". PMID 10749210. Nature 434 (7033): 636–40. [75] Barton. Biol.A. "Evolution of sex: The costs and benefits of sex: new insights from old asexual lineages". Lay summary (http:/ / news. PMID 18597889. "The Great Wall of China: a physical barrier to gene flow?".02164. PMID 11967555. Rev. physorg. Suppl.050503.97. Philosophical Transactions of the Royal Society of London. The synthetic theory of evolution: An introduction to modern evolutionary concepts and theories. PMID 11127900. Mol.37. bbc. doi:10. PMID 17373857. [92] Wendel J (2000). gov/ picrender. Biol. [85] Packer C. 96 (3): 241–52. nih.1038/35005078. stm). Douady CJ.1146/annurev. 3 (4): 311–7.1111/j. . "The degeneration of Y chromosomes" (http:/ / www. 42 (1): 225–49. doi:10. Godfray HC. PMC 2600545. htm#hardy).1016/j. "The ecological cost of sex". Reprod. PMID 12634804. PMID 11700276.03. [76] Muller H (1964). J. Cox SJ (March 2000). Pusey AE. Papke RT. Boudreau ME. [78] Otto S (1 July 2003). PMC 1692896. Proc. Dev. Rev. PMC 1288109.1126/science. 8 (3): 304–17. Rev. fcgi?tool=pmcentrez& artid=34383). PMID 10688139. gov/ articlerender. nih. Rappold G. Gilbert DA. PMC 1462613. nih.04. pubmedcentral. Nature 351: 562–65. [93] Sémon M. Wolfe KH (2007). Teach Evolution and Make It Relevant. N H (2000). Annu. Heredity 90 (3): 212–9.2000. Walsh DA. [74] Lien S.x. "The relation of recombination to mutational advance". 6 (11): 836–46. Szyda J. gov/ articlerender. PMID 15140081. Doolittle WF (2003).35. 355 (1403): 1563–72. "How species evolve collectively: implications of gene flow and selection for the spread of advantageous alleles" (http:/ / www. PMID 18006297. 35: 31–52. Qu L. Hered. "Causes of evolution" (http:/ / evoled.2008. Nesbo CL. Schechinger B.x. [73] Pouchkina-Stantcheva NN. gov/ articlerender.085719. PMID 10677316. Evol. nih. edu/ lessons/ causes. Soc. htm). doi:10. Herniou EA.1365-294X. PMC 1828144. Genet. J.005. PMID 16581499. Burt A (2005). Case RJ. pubmedcentral. doi:10. Palomar College. Behavioral Sciences Department.) 23 (8): 439–45. Natl. Dennis (2008).genet.0717. pubmedcentral. Kouyos RD. "Sex increases the efficacy of natural selection in experimental yeast populations". fcgi?tool=pmcentrez& artid=1462613). "Functional divergence of former alleles in an ancient asexual invertebrate". R. [89] Gross B.2004.1023/A:1006392424384. doi:10. Rieseberg L (2005).genet. Am. Evol. doi:10. "Genome evolution in polyploids". doi:10. "Consequences of genome duplication". [91] Vrijenhoek RC (2006). Genet. Zhang Z. Arnold ML (2001). "A molecular genetic analysis of kinship and cooperation in African lions". co. Nat. Nature 404 (6775): 281–5. doi:10. PMC 1692900.1016/j. [87] Su H. Trans. Fertil. [94] Comai L (2005). com/ news93597385. PMID 16304599. (23): 359–64.1093/jhered/esi026. He K. 106: 2–9. J. gov/ articlerender. [72] Fontaneto D. [95] Soltis P.00101.2006. fcgi?tool=pmcentrez& artid=1828144). National Science Foundation.1038/nature03405. fcgi?artid=1692896& blobtype=pdf). Gu H (2003). PMID 1107543.2006.1038/nrg1711.S. Arnheim N (February 2000). "Mutationism and the dual causation of evolutionary change". 16 (7): R245. Acad. 66 (2): 557–66. pubmedcentral. [90] Burke JM. [82] Stoltzfus A (2006). doi:10. . Boschetti C (October 2007).1146/annurev.Evolution [71] Goddard MR. PMID 16686641.1086/302754. "Genetic hitchhiking" (http:/ / www. U. PMID 15618301. pubmedcentral. McGee BM.0716. doi:10.1111/j. "The advantages of segregation and the evolution of sex" (http:/ / www. Nat. Soltis D (June 2000). doi:10. Annu Rev Genet 37: 283–328. PMID 11127901. Retrieved 2009-09-23.13. "The ecological genetics of homoploid hybrid speciation" (http:/ / www. "Genome fragment of Wolbachia endosymbiont transferred to X chromosome of host insect" (http:/ / www.00505.2020. fcgi?tool=pmcentrez& artid=104298). Symp.1073/pnas. [124] Charlesworth B (March 2009). org/ cgi/ content/ abstract/ 252626899v1). PMID 19204717. Bussière L (2004). Trends in Ecology & Evolution 23 (11): 602–609.1098/rspb. Biol. nih. gov/ articlerender. doi:10.2008.1038/nrg2506.2006. PMC 138577. Bentsen C. "Combinatorial genetic evolution of multiresistance". [104] Ohta T (2002). Philos.tree. pubmedcentral. PMC 1692213. PMID 1334911. Bioessays 29 (1): 74–84. [110] Collin. fcgi?tool=pmcentrez& artid=55389). "The objects of selection" (http:/ / www. gov/ articlerender. Berrigan D.A.x. U. "The units of selection". PMC 137875. Genet. Lloyd EA (1999). nih. Emma E. [100] Gladyshev EA. PMID 12386340. U. fcgi?tool=pmcentrez& artid=1208011). [122] Lande R (1989). fcgi?tool=pmcentrez& artid=1691039). nih. nih. doi:10. Sci. doi:10. 10 (8): 531–9. Whitlock M (1 June 1997). Nat. doi:10. Brooks R. [115] Hunt J. Virol. doi:10. Genetics 146 (2): 723–33. [123] Otto S. "Fundamental concepts in genetics: Effective population size and patterns of molecular evolution and variation". PMC 1208011. Natl. doi:10. U. [113] Andersson M. pnas. PMID 2687093. "Individuality and adaptation across levels of selection: how shall we name and generalise the unit of Darwinism?" (http:/ / www. PMID 18814933.6. Genet. [98] Kondo N. Shimada M. Arnold SJ (1983).) 21 (6): 296–302. [109] Goldberg. [103] Whitlock M (1 June 2003). [99] Sprague G (1991). nih. nih. PMID 10518549.A. Evolution 62 (11): 2727–2741.. doi:10. gov/ articlerender.1156407. PMC 2753274. Sci. Acad. "High-quality male field crickets invest heavily in sexual display but die young". [120] Hickey DA (1992). doi:10. PMID 17187354. 10 (2): 83–93. org/ stable/ 2408842). doi:10. [118] Mayr E (1997).A.1998.1002/bies. gov/ articlerender. "Fitness and its role in evolutionary genetics" (http:/ / www. 98 (16): 9157–60. [116] Odum. Hill C. EP (1971) Fundamentals of ecology. pubmedcentral.1038/nature03084. PMID 9653725. "The theory of natural selection today". "The sexual selection continuum" (http:/ / www. "Evolution and horizontal transfer of dUTPase-encoding genes in viruses and their hosts" (http:/ / www.1111/j. gov/ articlerender. Sci.2006. gov/ articlerender. gov/ articlerender. doi:10. Sci. pubmedcentral.1007/BF00133725.1558-5646. Sci. 213: 203–11. Nikoh N.1016/j. Natl. Sci. Genet. nih. Genet. Acad. gov/ articlerender. "Evaluating hypotheses for the origin of eukaryotes". PMC 1691039. PMC 104298. Biol.1016/j. Rachel. Proc. Saunders New York [117] Gould SJ (February 1998). gov/ articlerender. Genetics 164 (2): 767–79. fcgi?tool=pmcentrez& artid=33654).Evolution [97] Walsh T (2006). Hoekstra J. "The probability of fixation in populations of changing size" (http:/ / www. Genetics 93 (3): 773–95. gov/ articlerender.2008. nih. "Fisherian and Wrightian theories of speciation". doi:10. doi:10. doi:10.1038/nrg2526. PMID 13644170. PMC 18385. Proc. 38 . PMID 16942901. pubmedcentral.08. PMC 1462574.96. "Near-neutrality in evolution of genes and gene regulation" (http:/ / www. 1 (4): 530–3.11904. doi:10. third edition. nih.2091. Beerli P. fcgi?tool=pmcentrez& artid=1692213). Science 320 (5880): 1210–3. J. [119] Maynard Smith J (1998). pubmedcentral. PMID 19119264.1016/j. Igić. Genetica 86 (1–3): 269–74. "Strength and tempo of directional selection in the wild" (http:/ / www. 73 (9): 7710–21. Curr.222228199. PMID 11470913. Nat. fcgi?tool=pmcentrez& artid=137875). Simmons L (2006).252626899. [111] Hoekstra H. gov/ articlerender. 99 (22): 14280–5. Proc. PMID 18511688. nih. Rev.06. nih. (Amst. "Excursions along the Interface between Disruptive and Stabilizing Selection" (http:/ / www. "Evolutionary dynamics of transposable elements in prokaryotes and eukaryotes". Vignieri S. Ijichi N. doi:10. PMID 12461171. doi:10. Nature 183 (4663): 710–3. [112] Felsenstein (1 November 1979). . Kingsolver J (July 2001). Penny D (2007). Genome 31 (1): 221–7. Sci. PMID 16769428. PMID 15616562. fcgi?tool=pmcentrez& artid=2753274). [114] Kokko H. Hoang A. Dev. [121] Gould SJ. Acad. Trends Ecol. PMID 10438861. "Fixation probability and time in subdivided populations" (http:/ / www. PMID 9533127. Microbiol. PMID 17248980. "Massive horizontal gene transfer in bdelloid rotifers". pubmedcentral.009. Novartis Found. PMID 12079655. Miglietta. Fukatsu T (2002).015. Opin. Natl. Jennions M.013.tree. "The measurement of selection on correlated characters" (http:/ / jstor. pubmedcentral. 94 (6): 2091–4.A.S. R. B (2008). [107] Haldane J (1959). pubmedcentral. doi:10.2002. 10 (3): 195–205. fcgi?tool=pmcentrez& artid=1214112). Smith M. 99 (25): 16134–7. Meselson M. Opin.1073/pnas. "Genetic exchange between kingdoms". PMID 1822285. pubmedcentral. pubmedcentral. PMID 9178020. McClure M (1 September 1999). PMID 12807795. U. Trans.S.20516. Nat. 96 (21): 11904–9. . Rev. PMC 33654.161281098.03.mib. U. "Gulliver's further travels: the necessity and difficulty of a hierarchical theory of selection" (http:/ / www.A.1038/183710a0. "Reversing opinions on Dollo's Law". PMID 18764918. Acad. fcgi?tool=pmcentrez& artid=18385). doi:10. PMC 55389. Acad. pubmedcentral. PMC 1214112. doi:10.1038/nrg2603. doi:10.2307/2408842. Arkhipova IR (May 2008).0211. [102] Poole A. Natl. Curr. Natl. [105] Hurst LD (February 2009). Proc. [108] Lande R. "On phylogenetic tests of irreversible evolution".S. Evolution 37 (6): 1210–26.S. 353 (1366): 307–14. Proc. Soc.94. Lond.S. "Fundamental concepts in genetics: genetics and the understanding of selection". "Sexual selection and mate choice". fcgi?tool=pmcentrez& artid=1462574). discussion 211–7. 269 (1498): 1331–40. Evol.1073/pnas.1016/S0959-437X(05)80203-5.1073/pnas. [106] Orr HA (August 2009).21. MP (2008).1126/science.1073/pnas. B. PMID 19546856. Proc. 9 (5): 476–82.1098/rstb. Rev. doi:10. Nature 432 (7020): 1024–7. McNamara J. Houston A (2002). Brooks R. [101] Baldo A. PMID 9122151. U. edu/ ~ambuj/ Courses/ bioinformatics/ neutral-theory).051. Nature 457 (7231): 824–9. Handelsman J (December 2004). discussion 683. [142] Schloss P. doi:10. Microbiol Mol Biol Rev 68 (4): 686–91. Biol. Matzke NJ (May 2007).1073/pnas. PMID 17306543. Proc Natl Acad Sci U S a 95 (12): 6578–83. doi:10. [130] Kreitman M (August 1996). [145] Elena SF. PMID 15716908. "Selectionism and neutralism in molecular evolution" (http:/ / www.CO. "High-level fluoroquinolone resistance in Pseudomonas aeruginosa due to interplay of the MexAB-OprM efflux pump and the DNA gyrase mutation" (http:/ / www. 4 (6): 457–69. Wiebe W (1998). PMC 539005. . 1998.1093/molbev/msi242.1038/35059227.4. Mol. and became instead a continuing dynamic process.A. Thomas. Sugimoto Y. 82–92. pubmedcentral.367. [150] Dobzhansky T.2007. 66 (4): 367–86. Meredith. Jpn. "Neutral theory: a historical perspective.15. Natl. "Evolutionary adaptation is a phenomenon of pervasive importance in biology. DB (2007). nih. "The Beagle in a bottle". Genet. [149] Orr H (2005). doi:10. "The scale independence of evolution". com/ article. Nat. PMC 1513187.1525-142x. Genetics of the evolutionary process.68. "Biological design in science classrooms" (http:/ / www. doi:10. doi:10. Astyanax mexicanus" (http:/ / www. PMID 9618454. "Regressive evolution in the Mexican cave tetra.1038/nrg1523. 22 (12): 2318–42. "An introduction to microevolution: rate. Brockhurst MA. Proc. . nih.." [148] The Oxford Dictionary of Science defines adaptation as "Any change in the structure or functioning of an organism that makes it better suited to its environment". could no longer be considered a static condition.x. bioone. "Evolution experiments with microorganisms: the dynamics and genetic bases of adaptation". Acad. fcgi?tool=pmcentrez& artid=1876445).Y. Conrad. fcgi?tool=pmcentrez& artid=1513187). PMID 15590780. Willis. Evolution 10. Evolutionary biology 2.cub. [144] Buckling A. JB. [143] Nealson K (1999). gov/ articlerender. Harvard. Evol. doi:10. doi:10. doi:10. gov/ articlerender. 1970. Immunol. nih. Gross.S. "Tempo and mode in the macroevolutionary reconstruction of Darwinism" (http:/ / www. pattern. [131] Leigh E. J. doi:10. [129] Kimura M (1989). 1956. org/ perlserv/ ?request=get-abstract). gov/ articlerender. pubmedcentral. html) [140] Carroll SB (2001). doi:10. Colegrave N (February 2009). p4–6.1038/nrg1088. [135] Gould 2002. "The genetic theory of adaptation: a brief history". Nakae T (2002). "Post-Viking microbiology: new approaches. go. (Jr) (2007). PMC 1876445. 1–34.1023/A:1013368628607. Sci. PMID 12776215. new insights". Dougherty. Evol. Rev. Columbia. ucsb. PMID 11234024. 46 (6): 391–5. doi:10. doi:10. Natl.1111/j. D. doi:10.66. Kinnison MT (2001). nih. Borowsky. doi:10. org/ indexcc/ CB/ CB932. Long live the neutral theory" (http:/ / www.1038/nrg2207. jst. doi:10. jst. Microbiol.0701505104.1038/nature07892. Is the human race evolving or devolving? (http:/ / www.1666/0094-8373(2000)26[15:MAMSAH]2. [127] Nei M (2005). Yoneyama H.1073/pnas.G. . "The neutral theory of molecular evolution and the world view of the neutralists".1128/MMBR.1016/j. Orig Life Evol Biosph 29 (1): 73–93.1420-9101. pubmedcentral. sciam.12. pp. Genetic changes in populations of Drosophila pseudoobscura and Drosphila persimilis in some locations in California. go.A. 104 Suppl 1: 8669–76. pubmedcentral. talkorigins. gov/ articlerender. [139] TalkOrigins Archive response to Creationist claims – Claim CB932: Evolution of degenerate forms (http:/ / www. cfm?id=is-the-human-race-evolvin) Scientific American July 20. Sci.00044.2004. 2 (2): 67–77. gov/ articlerender. PMC 33863.686-691. fcgi?tool=pubmed& pubmedid=17306543#R2). .1046/j.1266/jjg. 39 . jp/ article/ mandi/ 46/ 6/ 46_391/ _article/ -char/ en). jp/ article/ jjg/ 66/ 4/ 66_367/ _article). PMID 17494747. "Status of the microbial census" (http:/ / www. PMID 11838760.S. [128] Kimura M (1991). [137] Jablonski. (2000). PMID 11258392.01410. pnas. Genet. PMC 44281. a product of a creative past. pubmedcentral. doi:10. M. Coleman D. PMC 2570642. [134] Leroi AM (2000). U. [126] Mitchell-Olds. PMID 2687096. Nat.and macroevolution: scale and hierarchy in evolutionary biology and paleobiology" (http:/ / www. 91 (15): 6764–71. [138] Michael J. process". . Lenski RE (June 2003). Proc. jstage. Genome 31 (1): 24–31.1073/pnas. . Rev. 1966. Current Biology 17 (5): 452–454. PMID 17943192. cs.95. 1968. On some fundamental concepts of evolutionary biology.1002/bies. Paleobiology 26 (sp4): 15–52. Nature 409 (6823): 1102–9.".2007.01. "The neutral theory of molecular evolution: a review of recent evidence" (http:/ / www. Bioessays 18 (8): 678–83. 6 (2): 119–27. doi:10. [146] Williams. [132] Scott EC. PMID 11536899. N. R (2007). Adaptation and natural selection: a critique of some current evolutionary thought. "Prokaryotes: the unseen majority" (http:/ / www. org/ cgi/ pmidlookup?view=long& pmid=8041695). jstage.6578. PMID 1954033. Tabin. Craig Maclean R. 84–87 [152] Dobzhansky T. C. PMID 12153116. fcgi?tool=pmcentrez& artid=33863).6764. PMID 17956380..2000. nih.91." p5 [147] Mayr.Evolution [125] Protas. [133] Hendry AP. George C.2. Princeton. The growth of biological thought. fcgi?tool=pmcentrez& artid=539005).0. "Chance and necessity: the evolution of morphological complexity and diversity". Dev. Acad. 79–82. [141] Whitman W. PMID 19212400. Nature Reviews Genetics 8 (11): 845–856. Genetica 112–113: 1–8. doi:10. "The neutral theory is dead. Genetics of natural populations XXV. PMID 8760341.1023/A:1006515817767.950180812. [151] Dobzhansky T. [153] Nakajima A. p483: "Adaptation. Genet. "Which evolutionary processes influence natural genetic variation for phenotypic traits?". new data. "Micro. JH. PMID 8041695. Journal of Evolutionary Biology 20 (6): 2075–91. PMID 16120807. 657–8 [136] Gould SJ (July 1994). Ernst 1982. Goldstein.x. PMID 16526515. (1872) The Expression of the Emotions in Man and Animals John Murray. pdf). New York. Gerhart J (July 1998). [169] Maxwell EE. Penguin Group.1046/j. doi:10.1016/S0959-437X(99)00020-9. doi:10. Dev.2421. Charles.1073/pnas.A. doi:10. Trends Biochem. PMID 12492145. "Osteology and myology of the wing of the Emu (Dromaius novaehollandiae). Nat Rev Micro 4 (10): 784–790. 105 (23): 7899–906. doi:10. PMID 10607605.15. 26: 249–68. Science 131: 1292–7. pp. J.S.1017/S1464793102006097. 25 (6): 261–5. Sci. PMID 16953248. Ann. 1-56. "Vestigialization and Loss of Nonfunctional Characters". VA. .1292.95. Evol. "Large-scale analysis of pseudogenes in the human genome". com/ hdy/ journal/ v100/ n5/ full/ 6801095a. Worth Publishers. Acad. fcgi?tool=pmcentrez& artid=33871). Sci. "Adaptive evolution of eye degeneration in the Mexican blind cavefish".1038/nature07891. PMID 17123025. S. PMID 19212399. (2009). Brian K (2003). [170] Silvestri AR. Nature 457 (7231): 818–823. PMID 8236445. [173] Behavior Development in Infants (http:/ / books. Hall BK (2002). 40 . [167] Zhang Z. .06. Why Evolution is True.1038/nrmicro1493. Negoro S.es. 95 (15): 8420–7. "The recent evolution of pentachlorophenol (PCP)-4-monooxygenase (PcpB) and associated pathways for bacterial degradation of PCP".1093/jhered/esi028. pp. pubmedcentral. "The reducible complexity of a mitochondrial molecular machine" (http:/ / www.1525-142X. Sci. Journal of the American Dental Association (1939) 134 (4): 450–5. Natl. [181] Hardin G (April 1960). Genet. doi:10. doi:10. Hallgrímsson. ada. Vol. "Lens crystallins: gene recruitment and evolutionary dynamism". [176] Gould 2002.1554/05-233.1146/annurev. Proceedings of the National Academy of Sciences 106 (37): 15791–15795. J. doi:10. [156] Ohno S (April 1984). "Inaugural Article: Historical contingency and the evolution of a key innovation in an experimental population of Escherichia coli" (http:/ / www.S. nih.gde. . Abigail. Carroll.. internally repetitious coding sequence" (http:/ / www.110195. 18 (8): 301–6. PMC 2747197.26. "Limbs in whales and limblessness in other vertebrates: mechanisms of evolutionary and developmental transformation and loss". 9 (6): 672–77. org/ doi/ abs/ 10. [166] Fong D. PMID 15261647. org/ cgi/ pmidlookup?view=long& pmid=18524956).1126/science. "The competitive exclusion principle" (http:/ / www. 85–86. . [157] Copley SD (June 2000). Proc. PMID 18212804.6801095. Nakamura S (1983). Bursac. citing a study "Reflexes and other motor activities in newborn infants: a report of 125 cases as a preliminary study of infant behavior" published in the Bull. PMID 14558591. [161] Bejder L.1016/j. Neurol.hdy.A. gov/ articlerender. 268 (5): 423–41. 1932. "Experimental evolution: experimental evolution and evolvability" (http:/ / www.1073/pnas.8. Genet. U. ISBN 978-0-670-02053. 1554/ 05-233. [178] Clements.001341. pp.1038/sj. [175] Anthony Stevens (1982).2004. Perry. PMC 345072. "Deep homology and the origins of evolutionary novelty". 14 (4): 328–35. Mark J. 2. bioone. Larsson HC (2007). "The nature of the last universal common ancestor". [155] Okada H. PMID 15653557.001341. Natl. Trends Biochem. long). AJ. [168] Jeffery WR (2005). Routledge & Kegan Paul. Kimura H. "Birth of a unique enzyme from an alternative reading frame of the preexisted. doi:10.003. PMID 19717453. Sci. doi:10. google. nature. Heredity 100 (5): 464–70. Sci. Culver D (1995). nl/ ~gkorthof/ pdf/ Pallen_Matzke. org/ cgi/ content/ full/ 134/ 4/ 450). "Evolutionary adaptation of plasmid-encoded enzymes for degrading nylon oligomers". Rev. [162] Young.2002. Archetype: A Natural History of the Self. "Evolvability" (http:/ / www. Acad.0803151105.02033. PMID 8032155. Biodegradation 18 (5): 525–39. B (2005).131. S (2009).A. "Recruitment of enzymes and stress proteins as lens crystallins". PMID 12733778. Singh I (2003). Hered. [172] Peter Gray (2007). Celik. doi:10. PMID 10838562. doi:10. 66. ISBN 0-7167-0617-2. 87.1146/annurev. Psychology (fifth ed. .1073/pnas. . Tomarev SI (1994). S et al. Nature 306 (5939): 203–6. [158] Crawford RL.1073/pnas. Gatsos. Civciristov.1016/S0968-0004(00)01562-0. . Natl. C. PMC 33871. Kane T. [164] Hall. EXS 71: 241–50.. doi:10.110195. Evolution 59 (12): 2691. Neil. Inst. pnas. org/ cgi/ pmidlookup?view=long& pmid=14399717). and its bearing on the evolution of vestigial structures". pnas. org/ cgi/ pmidlookup?view=long& pmid=6585807). ISBN 0-7100-0980-1. Gopal-Srivastava R.81. Tabin. PMID 17390336. "Descent with modification: the unity underlying homology and homoplasy as seen through an analysis of development and evolution". Matzke. Ecol. Jung CM. PMID 9671692. Likic. doi:10. Acad. Lenski RE (June 2008). doi:10. N. pp. London. PMID 6585807. doi:10. Kantorow M. org/ content/ 106/ 37/ 15791. Proc. 1235–6 [177] Pallen. Curr. Syst. Proc. [174] Jerry Coyne (2009). [165] Shubin. sciencemag. Curr. PMID 18524956. Morphol.1038/306203a0. Gerstein M (August 2004). pnas. NJ (2006-10). doi:10.). X.1016/0968-0004(93)90041-K.1. U. html). doi:10. U. "Serial homology and the evolution of mammalian limb covariation structure" (http:/ / www.0908264106.es. Poole A (1999). [179] Piatigorsky J.26. [159] Colegrave N. Biological Reviews of the Cambridge Philosophical Society 78 (3): 409–433. [171] Darwin. Strap JL (October 2007). PMID 6646204. 1). D.8420. 81 (8): 2421–5. . Collins S (May 2008). PMC 2430337. doi:10. Retrieved 2009-09-18.10527. Retrieved 2009-09-24. 96 (3): 185–96. [180] Wistow G (August 1993). pp.S. doi:10. planet.x. [163] Penny D. "The unresolved problem of the third molar: would people be better off without it?" (http:/ / jada. PMID 14399717.Evolution [154] Blount ZD. Retrieved 2009-09-18.1007/s10532-006-9090-6. Borland CZ. doi:10. Poggio. "From The Origin of Species to the origin of bacterial flagella" (http:/ / home. doi:10. Opin. Opin.3409.1002/jmor. [160] Kirschner M. Dev. 4 (6): 445–58. "Evolution of a metabolic pathway for degradation of a toxic xenobiotic: the patchwork approach". com/ books?id=OzL2Bgvf9Q0C& pg=PA122& dq="chancy+ and+ mcgraw+ (31)+ investigated+ the+ reflex+ in+ the+ 125+ infants"") (via Google Books) by Evelyn Dewey. Nathan M. Dev. Evolution 56 (10): 2067–82. doi:10.1007/s00425-004-1436-x. [191] Paszkowski U (2006).1038/nrg2031. Huyghe. 147 (4): 403–37.1159388. Genet. gov/ articlerender. Cell 134 (1): 25–36. es/ web/ paper. doi:10. "Speciation in the apple maggot fly: a blend of vintages?". J. [199] de Queiroz K (May 2005). "Five rules for the evolution of cooperation". Garcia-Fernàndez J (2003). Planta 221 (2): 184–96.0502030102. PMID 17517608. 104 (23): 9736–40.J. doc). Hasso SM. pnas. PMC 2290806.tree..05. Ridenhour BJ. Porter AH (2001).R. Hanage WP (February 2009). PMID 16701238. talkorigins.cell.. .1126/science. PMC 1224642.2006. "Remarkable Creatures – Clues to Toxins in Deadly Delicacies of the Animal Kingdom" (http:/ / www. doi:10. Biol.. doi:10. I. *Weinberg JR. J.. PMID 16910971. ehu. Plant Biol. D (1992). discussion 1426–36. "Mutualism and parasitism: the yin and yang of plant symbioses".2307/2410209. nih.S. "Evidence for Rapid Speciation Following a Founder Event in the Laboratory" (http:/ / jstor. and the Synthesis of Evolutionary and Developmental Biology" (http:/ / philsci-archive. PMC 1131873.1023/A:1023940220348. [184] Baguñà J. PMID 11838782. 102 (Suppl 1): 6600–7. Science 314 (5805): 1560–3. "Adaptive evolution and explosive speciation: the cichlid fish model" (http:/ / hcgs. doi:10. "Cooperation within and among species".2003. Hamilton W (2005). Brodie ED (2002). unh. "Perspective: models of speciation: what have we learned in 40 years?". 8 (3): 185–95. PMID 17279094. doi:10. Hölldobler B (2007). [197] De Queiroz K (December 2007). Proc Natl Acad Sci U S A. Alm EJ. [189] Geffeney S. doi:10. [200] Rice. "The development of archosaurian first-generation teeth in a chicken mutant". New York Times. PMID 12449493. Fester T (2005). "The evolutionary response of predators to dangerous prey: hotspots and coldspots in the geographic mosaic of coevolution between garter snakes and newts".06. [194] Axelrod R. "The bacterial species challenge: making sense of genetic and ecological diversity". 19 (5): 1415–8. doi:10. "Evolution of the mammalian middle ear".1002/jmor.01152. K.1073/pnas. "Evo-Devo: the long and winding road" (http:/ / www. 16 (4): 371–7. 5 (4): 288–98. TalkOrigins Archive. Proceedings of the National Academy of Sciences 105 (12): 4792–5. 102 (38): 13367–71.1126/science. org/ cgi/ pmidlookup?view=long& pmid=15851674). pitt. Rev. Rev. Carroll (December 21. U.1133755. Curr. (Amst. doi:10. *Jiggins CD. fcgi?tool=pmcentrez& artid=1887545). PMID 14628909. [190] Sachs J (2006). Opin. [195] Wilson EO. doi:10. . Science 323 (5915): 741–6. Jorg.S. PMID 15851674. "Toward a new synthesis: population genetics and evolutionary developmental biology". PMID 14756346. PMID 18614008. doi:10. Evol. Int. doi:10. Fallon JF (2006).12. nytimes. Science 297 (5585): 1336–9. 2009).cub. PMID 15131652.pbi. *Love AC. org/ stable/ 2409766). Sci. org/ faqs/ faq-speciation.1073/pnas.) 19 (3): 111–4. "Laboratory experiments on speciation: what have we learned in 40 years".7466396.Evolution [182] Kocher TD (April 2004). "The co-evolutionary genetics of ecological communities". [187] Carroll SB (July 2008). "Ernst Mayr and the modern concept of species" (http:/ / www. D.1554/02-727. 47 (7–8): 705–13. B. Polz MF. "Eusociality: origin and consequences" (http:/ / www. *Sean B.x. Brodie ED. PMC 1887545. "The emergence of a superorganism through intergroup competition" (http:/ / www. "Rapid large-scale evolutionary divergence in morphology and performance associated with exploitation of a different dietary resource" (http:/ / www. Acad. Syst. Nat. Proc. *Boxhorn. nih. U.0703466104. "Observed Instances of Speciation" (http:/ / www. Dev. pubmedcentral.2008. [198] Fraser C. J (1995). "Evo-devo and an expanding evolutionary synthesis: a genetic theory of morphological evolution". Nat. doi:10. *Nowak M (2006). Ferguson MW. Biol. pdf). "Mechanisms of adaptation in a predator-prey arms race: TTX-resistant sodium channels". gov/ articlerender. Spratt BG. Trends Ecol. doi:10. PMID 7466396. PMID 16157878. PMID 18344323. Science 211 (4489): 1390–6. . fcgi?tool=pmcentrez& artid=1224642). [193] Reeve HK. PMID 15871030.1038/nrg1316. Vanhooydonck. 41 .1016/j. Bridle JR (2004).1111/j. pubmedcentral.1126/science.12. PMID 18027281. Breugelmans.A.1073/pnas.008. Genet. com/ 2009/ 12/ 22/ science/ 22creature.1074310. Curr.1073/pnas. Natl. PMID 19197054. [188] Wade MJ (2007). PMID 16713732.1051470404. Sci. . Evol. PMID 17158317. J. Biol. [192] Hause B.1420-9101. R. Biology and Philosophy 18 (2): 309–345. doi:10.030. edu/ staff/ kocher/ pdfs/ Kocher2004. html?hpw). edu/ archive/ 00000375/ 00/ LondonPaper. 9 (4): 364–70. gov/ articlerender.1016/j. [201] Herrel. [186] Harris MP.. Retrieved 2008-12-26. Ruben PC.047. Hölldobler B (2005). doi:10.1126/science. pubmedcentral. [183] Johnson NA.. (2003). *Brodie ED.2006. Evolution 47 (6): 1637–1653. fcgi?tool=pmcentrez& artid=2290806). "Species concepts and species delimitation". Evolution 46 (4): 1214–20. Hostert (1993). Genetica 112–113: 45–58. ijdb. doi:10. php?doi=14756346). doi:10. Irschick.1016/j.A.1023/A:1013371201773. (2008).1016/j. doi:10.. Morphol. W. . doi:10. doi:10.0505858102. Proc. Acad. A.2307/2409766. PMID 1202224. "Evolutionary Morphology. Brodie ED (2002). Grbac.. PMID 12193784. Evolution 57 (10): 2197–215. . doi:10. Backeljau. Van Damme. [185] Allin EF (1975).2005. . doi:10. nih. [196] Gavrilets S (2003). "Molecular and cell biology of arbuscular mycorrhizal symbiosis". T. 56 (6): 879–86. K. Natl. "The evolution of cooperation".008.0711998105.1080/10635150701701083. Biol. PMID 16488870. Starczak VR. html). Innovation. Genetics 94 (4): 1011–38.S. Schopf. "Punctuated equilibria: an alternative to phyletic gradualism" (http:/ / www. 91 (15): 6764–71. doi:10.1101476. (1997). PMID 16829570. 106 (33): 13875–9. Nature 437 (7063): 1353–356.1038/nature04004.7701342. "Genomic clues to the evolutionary success of polyploid plants". T. nih.A.1016/j. doi:10.I. doi:10. present. "Climate change.0604181103. doi:10. Lind-Halldén C. Ayala FJ.0802504105. "Evolution in closely adjacent plant populations X: long-term persistence of prereproductive isolation at a mine boundary" (http:/ / www.1016/j. "Diversification and extinction in the history of life".1093/molbev/msk006. doi:10. "Sympatric speciation in Nicaraguan crater lake cichlid fish". Higgle M.1007/s10709-006-0013-6. nih.1073/pnas. [221] Pimm S. genetics.S. pnas.15. "Adaptive differentiation following experimental island colonization in Anolis lizards". Biol. doi:10. fcgi?tool=pmcentrez& artid=33235). nih. 105 (Suppl 1): 11453–7. 82–115. .2003.A. PMC 1831839. "Tempo and mode in the macroevolutionary reconstruction of Darwinism" (http:/ / www. Hubbell SP.1016/j. PMID 18492478. Mol. 16 (5): 1019–29. Natl. species-area curves and the extinction crisis" (http:/ / www.S.6758. 1/ 11453. PMC 1214177. Mark (2005).15. Genetica 129 (3): 309–27. [213] Bomblies K. nih. Biol. Sci.1126/science. Shabel AB (2004). . PMID 15459379. ed. "Arabidopsis-a model genus for speciation".. Warhelt. PMID 16549398. org/ cgi/ reprint/ 94/ 4/ 1011). html).1073/pnas. full). Sci. html). doi:10. "Sympatric speciation in palms on an oceanic island". Takebayashi N. Sci.S. San Francisco: Freeman Cooper. Models in Paleobiology. nih. fcgi?tool=pmcentrez& artid=1544153). Natl. [218] Avise JC. Salzburger W. Acad.S.J. Sci. Hiscock SJ (2008).1038/nature04325. doi:10.101092598. PMID 16467788.1073/pnas. Acad. Schoener. U. Hutton I. Powell MP. [220] Novacek MJ. [224] Isaak. Sci. "Biological extinction in Earth history". PMID 16639420. Clarkson JJ. gov/ articlerender. "The current biodiversity extinction event: scenarios for mitigation and recovery" (http:/ / www.03. Raven P. Tavaré S (2006).gde. . pubmedcentral. PMID 16467837. Peterson A. Evol. fcgi?tool=pmcentrez& artid=44280). fcgi?tool=pmcentrez& artid=44281). Halldén C (2003). Gries R.043. doi:10. Anstett M-C. PMID 8041695. Acad. [219] Raup DM (1994). Salamin N. "The role of extinction in evolution" (http:/ / www. Sci. Natl. PMC 44280. "Assessing the causes of late Pleistocene extinctions on the continents". PMID 16900317. Philos.S. [207] Savolainen V. asp) In T. 1972. Greenspoon PB. [217] Raup DM (1986).1073/pnas.hdy. PMID 18006296. PMID 7701342. Stölting KN.09. Reprinted in N. Natl. U. "The Maynard Smith model of sympatric speciation". Nature 441 (7090): 210–3. [214] Niles Eldredge and Stephen Jay Gould. Koch PL. Barker MS.1712. doi:10. Rieseberg LH (August 2009). [223] Jablonski D (May 2001). McDonald KR. Gries G (2007). PMID 18695213. U. gov/ articlerender. TalkOrigins Archive.006. Feranec RS. B.041. doi:10. Lond. *Barluenga M. U. PMID 8041694. gov/ articlerender. Proc. pubmedcentral. PMID 11344295.cub. PMID 6777243. gov/ articlerender. "The frequency of polyploid speciation in vascular plants" (http:/ / www. Sci.A. 23 (6): 1217–31. Theor. Proc. 239 (2): 172–82. Proc. PMC 2728988. 98 (10): 5393–8. "Human impacts on the rates of recent. 91 (15): 6758–63. doi:10. Science 306 (5693): 70–5.6764.A. Evol. Natl. Proc. "Chloroplast DNA indicates a single origin of the allotetraploid Arabidopsis suecica".91.1046/j. Meyer A (2006). Press. Natl. [208] Gavrilets S (2006). Baker WJr (2006). doi:10. PMID 16251964. Acad.0811575106. blackwellpublishing. 1985 [215] Gould SJ (1994). doi:10. org/ indexcc/ CB/ CB090. gov/ articlerender. pubmedcentral. 98 (10): 5466–70. Proc. fcgi?tool=pmcentrez& artid=1831839).S. "A unique recent origin of the allotetraploid species Arabidopsis suecica: Evidence from nuclear DNA markers". gov/ articlerender. Current Biology 18 (10): 435–44. doi:10.1073/pnas. Natl. Weigel D (2007). Princeton: Princeton Univ.1038/nature04566. Sekercioglu CH.2005. doi:10. Mayrose I. Science 268 (5207): 52–8. PMID 11344284. doi:10. Trans.. Proc. doi:10. pubmedcentral. fcgi?tool=pmcentrez& artid=2728988).B.11542058. [209] Wood TE. PMC 2556414.M. Eldredge Time frames.A. talkorigins.091093698.1038/sj.2008. com/ hdy/ journal/ v97/ n1/ full/ 6800835a. fcgi?tool=pmcentrez& artid=33224). [206] Nosil P. PMID 16553315. Biol. 42 . Acad. nih. Lexer C. J. Jakobsson M. doi:10. Cleland EE (2001). "Reinforcement drives rapid allopatric speciation". [211] Jakobsson M. "Claim CB090: Evolution without abiogenesis" (http:/ / www. Ehrlich PR (2006). PMC 44281. K. Moritz C (2005). [216] Benton MJ (1995). "In the light of evolution II: Biodiversity and extinction" (http:/ / www. Springate D. Wing SL. Nature 439 (7077): 719–23. [212] Säll T.91. "Natural selection and divergence in mate preference during speciation". pp. U. 361 (1465): 163–71. Retrieved 2008-12-26. doi:10. PMC 1544153.jtbi.6800835. U. PMID 11542058. [204] Templeton AR (1 April 1980).2007. doi:10. Acad.1126/science. gov/ articlerender.Evolution [202] Losos.08.x. doi:10.1073/pnas. Curr Opin Genet Dev 17 (6): 500–4. [205] Antonovics J (2006).1038/387070a0. PMC 33224. Muschick M. .2005. J. Biol. Nature 387 (6628): 70–3.1126/science. Crespi B. 103 (29): 10941–6. Hagenblad J. "Lessons from the past: evolutionary impacts of mass extinctions" (http:/ / www. pubmedcentral. pubmedcentral.W. and future bird extinctions" (http:/ / www. Soc. *Barnosky AD. R.1098/rstb.1420-9101. Sci. [210] Hegarty Mf.A. nih. Science 231: 1528–33. PMC 33235. "The theory of speciation via the founder principle" (http:/ / www. nature. com/ ridley/ classictexts/ eldredge. pubmedcentral. [222] Lewis OT (2006). org/ content/ 105/ suppl. PMID 19667210. PMID 16242727.00554. Proc. Heredity 97 (1): 33–7. Norup MV. J.A. [203] Hoskin CJ. U. Acad. PMID 14635917. (August 2008).1073/pnas. Evol. Pace N (2001). doi:10. Biol. Acad. Burger G (1999). Evol. Proc Natl Acad Sci U S a 91 (15): 6735–42.1080/106351501750435040. Jablonski D (2003). Ouzounis CA (2005). "The antiquity of RNA-based evolution". 15 (12): 1746–58.1574-6976.2114.1038/311019a0.1842.08. im. [230] Theobald. (2010). .resmic. Int.090145. doi:10. doi:10. J. Sci. Genome Res.1016/j. Nature 418 (6894): 214–21.1834. .15. Grishin NV. .1016/j. Cell Biol. [238] Cavalier-Smith T (2006). Kazmierczak J (2003). "Ancient invasions: from endosymbionts to organelles". doi:10.1016/j. [244] DeLong E.x. [239] Schopf J (2006).2006. PMID 11700279. PMID 9071008.1007/PL00000055. doi:10. [235] Mason SF (1984).tim. PMID 12175808. "Environmental diversity of bacteria and archaea".cellbi. gov/ articlerender. "The missing link between hydrogenosomes and mitochondria".35. [233] Kunin V. Stano P (2006). 43 . doi:10. Psenner R (2001).07. 13 (6): 256–61. Syst Biol 50 (4): 470–8.2004. PMID 10381868. "Origins of biomolecular handedness". [247] Ohno S (1997). Altheide TK (2005). "Molecular adaptation and the origin of land plants".1098/rstb. [227] Trevors JT. 18 (9): 472–79.genet. "Controversies on the origin of life" (http:/ / www. Science 304 (5668): 253–57. 13 (10): 457–59. J. doi:10. Johnson P (2004).2001. Benachenhou-Lahfa N. "Genome trees and the tree of life". [229] Trevors JT. es/ web/ paper. "Disparate rates. doi:10. PMID 10607659. Mol. doi:10.1038/nature09014. ehu. nih. [237] Varki A.1469-185X.Evolution [225] Peretó J (2005).08.1073/pnas. "Does the 'Ring of Life' ring true?". BioSystems 28 (1–3): 15–32. Philos Trans R Soc Lond B Biol Sci 361 (1470): 869–85. Development 126 (5): 851–9. Mol.ympev. PMID 17261804. [243] Lang B. PMID 15936656.2006.00018. Ferri F. "A formal test of the theory of universal common ancestry".1111/j. "Fossil evidence of Archaean life" (http:/ / www. php?doi=14756327). . Rogozin IB. PMID 8041691. Trends Microbiol. Brown M.1016/S0168-9525(02)02744-0. doi:10. Rev. PMID 14756327. PMID 15965028.2003. [240] Schopf J (1994). 47 (7–8): 517–22. "From self-assembly of life to present-day bacteria: a possible role for nanocells". Trends Genet. [246] Valentine JW. fcgi?tool=pmcentrez& artid=44277).1073/pnas. 8 (1): 23–31. microbios. Curr Opin Plant Biol 2 (6): 513–19. "Why are there so many insect species? Perspectives from fossils and phylogenies". Darzentas N.03. biologists. "The future of the fossil record". *Valentine J.Forterre P.1098/rstb. [249] Mayhew PJ (August 2007). [245] Kaiser D (2001). "Fossils. org/ cgi/ reprint/ 126/ 5/ 851). PMID 16292523.2005. doi:10. [226] Luisi PL.284. 28 (11): 729–39. PMID 15906258. PMID 14596897. doi:10. Labedan B (1992). Int.1016/j. .1111/j. "Cell evolution and Earth history: stasis and revolution" (http:/ / www. PMID 11742692. doi:10. PMC 1172039.1101/gr. FEMS Microbiol. fcgi?tool=pmcentrez& artid=1578735).x.006.1.1038/418214a. PMID 12116647. nih. Abel DL (2004). 29 (3): 456–63. *Altermann W. Proc.1094884. Genome Res. Annu Rev Genet 33: 351–97. PMID 20463738.1126/science.018. Gray M. Int.tb00592. Confalonieri F. pdf) (PDF). Philos Trans R Soc Lond B Biol Sci 361 (1470): 969–1006. PMID 9927587.2005. "Building a multicellular organism".genet. doi:10. U. pubmedcentral. [248] Waters ER (2003). [241] Dyall S. "Chance and necessity do not explain the origin of life".A. 35: 103–23. Nature 465 (7295): 219–22. [242] Martin W (2005). genome. PMID 16754610. Rev.1146/annurev. PMID 12110897. DL. fcgi?tool=pmcentrez& artid=1578732). PMC 44277.1007/s00114-005-0056-z. Elie C. doi:10. still open questions". [231] Bapteste E. Microbiol. doi:10.2003. *McFadden G (1999). PMID 17624962. "Approaches to semi-synthetic minimal cells: a review".S. org/ cgi/ pmidlookup?view=long& pmid=17261804). [236] Wolf YI. org/ 0801/ 0801023.1016/S1369-5266(99)00025-4.005. doi:10. "Morphological and developmental macroevolution: a paleontological perspective" (http:/ / www. ijdb. "Pattern pluralism and the Tree of Life hypothesis" (http:/ / www.6735. Trends Microbiol. PMID 16754604. org/ cgi/ pmidlookup?view=long& pmid=15965028). Jablonski D.012. PMID 10690412. 104 (7): 2043–9. Nature 311 (5981): 19–23. doi:10.33. 15 (7): 954–9. doi:10. PMID 15073369.1146/annurev. Goldovsky L.351. doi:10. doi:10.1101/gr. PMID 6472461.3666505. pubmedcentral.1126/science. Annu. Bapteste E (February 2007). PMID 15563395. "The nature of the last universal ancestor and the root of the tree of life. Duguet M.102401.06.0610699104. PMC 1578735. "The net of life: reconstructing the microbial phylogenetic network" (http:/ / www. 44 Suppl 1: S23–7. PMID 1337989. "Comparing the human and chimpanzee genomes: searching for needles in a haystack". gov/ articlerender. doi:10. PMID 16339373. Walsh DA (2005). differing fates: tempo and mode of evolution changed from the Precambrian to the Phanerozoic" (http:/ / www. PMC 1578732. Dev.2007. Science 284 (5423): 2114–16. Erwin DH (1 March 1999). nih.3737405. Natl. gov/ articlerender.tim. Biol Rev Camb Philos Soc 82 (3): 425–54. molecules and embryos: new perspectives on the Cambrian explosion" (http:/ / dev. Naturwissenschaften 93 (1): 1–13. "The reason for as well as the consequence of the Cambrian explosion in animal evolution".5423. doi:10. PMC 1892968. PMID 14615186.91. Koonin EV (2002). "Mitochondrial genome evolution and the origin of eukaryotes". "Endosymbiosis and evolution of the plant cell". Res Microbiol 154 (9): 611–17. [228] Joyce GF (2002). Genet. doi:10. doi:10. Phylogenet. [232] Doolittle WF.006. doi:10.1016/j. 25 (5): 573–82. pnas.1016/0303-2647(92)90004-I. "Archean microfossils: a reappraisal of early life on Earth". pubmedcentral. [234] Jablonski D (1999). PMID 16109488. pgen. [260] Koza. Borowsky R. doi:10. Fromman-Holzboog. see Midgley. PMID 12952685. "bloodthirsty. Parker SK. "History of Science: Origins of Evolutionary Theory" (http:/ / records. Nature 181 (4603): 208–9.37. London: Pimlico. Nature 409 (6817): 175–78. M. Biol.025.04. p. (2003). "Intelligent design and biological complexity". M. doi:10.1016/j.32. 5 (1): e1000326. The Myths we Live By. Liberal Studies Department. Nolta. Mathematical. PMID 19360058. Annu Rev Ecol Syst 32: 183–217. viu. [273] Gowaty. Retrieved 2007-05-24.1016/j. "Protein design by directed evolution". Journal of Geoscience Education 53 (3): 319. V.1038/458695a.114020. 44 . [267] Spergel.ecolsys. [253] Maher B. doi:10. *Bowler. religious. Komatsu.006. Tabin CJ (January 2009).ydbio.. [265] Hameed. "Tools for intelligent control: fuzzy controllers. PMC 2603666.4 Gyr ago". Malaspina University College. R. "Who Believes What? Clearing up Confusion over Intelligent Design and Young-Earth Creationism" (http:/ / www. Harvard University Press. edu/ ~sahCS/ Hameed-Science-Creationism. Smith BD (2006). Feminism and evolutionary biology: boundaries. Routledge.1371/journal. Adaptation in Natural and Artificial Systems. H. 62. G. Wichman HA (2001). Janet (2003). doi:10. PMID 17190597. PJ (2003).2007.1000326. Patricia Adair (1997). Retrieved 2008-04-28. Philosophical transactions. doi:10. [261] Jamshidi M (2003). doi:10. Zhou Y. 18 (1): R23–4. Public acceptance of evolution". Bennett...1126/science.11. "Applied evolution". Hinshaw.cell..biophys. ISBN 978-0674445574. PMID 15293515. Okamoto. Retrieved 2009. E. MIT Press. doi:10. "A novel role for Mc1r in the parallel evolution of depigmentation in independent populations of the cavefish Astyanax mexicanus" (http:/ / www. "Evidence from detrital zircons for the existence of continental crust and oceans on the Earth 4. PMID 18177707. (1992). physical. PMID 19074331. et al. [263] For an overview of the philosophical. Cornell CN. M (2004). doi:10.023. Charles Darwin: The Power of Place.006. Dev. Ingo (1973) (in German). "Restoring sight in blind cavefish".1016/j. pubmedcentral. Completely Revised and Expanded.gene.1126746. [269] Understanding Creationism after Kitzmiller (http:/ / www. Detrich HW (July 2005). History and philosophy of the life sciences 25 (1): 51–79. doi:10. neural networks and genetic algorithms". nagt. org/ files/ nagt/ jge/ abstracts/ Ross_v53n3p319. "Evolutionary ethics from Darwin to Moore". D (1998). . doi:10. Nature 458 (7239): 695–8. Simon & Schuster. [254] Borowsky R (January 2008). ca/ ~johnstoi/ darwin/ sect3. PMID 16902112.. ISBN 978-0415340779. Salman (2008-12-12). PMID 19119422. Gaut BS. see: Johnston. ISBN 978-0684824710. The Astrophysical Journal Supplement Series 148: 175–94. S (2006). "First-Year Wilkinson Microwave Anisotropy Probe (WMAP) Observations: Determination of Cosmological Parameters". Evolution: The History of an Idea. Gene 385: 2–18. Verde. pdf) (PDF).1163672. see: Dennett. Evolutionsstrategie – Optimierung technischer Systeme nach Prinzipien der biologischen Evolution (PhD thesis). [264] Ross. "Science communication. 1641/ B570313) 2007 [270] On the history of eugenics and evolution. bioone.Evolution [250] Bull JJ. N.1038/35051550.1146/annurev. Scott. D.1098/rsta. (2005). And Still We Evolve. Third Edition. D (1995). [259] Holland. and cosmological controversies. [268] Wilde SA. "Science and Religion: Bracing for Islamic Creationism" (http:/ / helios. Halpern. In the Name of Eugenics: Genetics and the Uses of Human Heredity.1126/science.cub.2005. University of California Press. see Kevles. L. ISBN 978-0520236936. PMID 13504138. .2006. "Monte Carlo analyses of genetic models". Kast P. PMID 11196637. [258] Rechenberg. Ian C.1225. nih. University of Michigan Press.1146/annurev. Curr.R. PMID 15890331. Genetic Programming. htm). 283 (1): 97–112. "The molecular genetics of crop domestication".. Annu Rev Biophys 37: 153–73. doi:10. M. [252] Jäckel C. pdf).1038/181208a0. (1975). PMID 17011142. doi:10. 376–379.1080/03919710312331272945. "Evolution: Biology's next top model?".2006.1016/j. ISBN 0-7126-6837-3. *Zuckerkandl E (2006). PMID 18573077. [256] Yergeau DA.. [251] Doebley JF. (April 2009). Hilvert D (2008). C. Peiris. [255] Gross JB. PLoS Genet. hampshire.081501.1086/377226.03. Science 313 (5788): 765–66. Valley JW. L. [272] Allhoff F (2003).2003.125832. . and frontiers. pp. [266] Spergel D. and engineering sciences 361 (1809): 1781–808. Series A..032807. intersections. an RBCC/TRIM gene required for erythropoiesis in zebrafish". Biol. Peck WH. [262] Browne. [257] Fraser AS (1958). *For the scientific and social reception of evolution in the 19th and early 20th centuries. ISBN 0262111705. N. [271] Darwin strongly disagreed with attempts by Herbert Spencer and others to extrapolate evolutionary ideas to all possible subjects. gov/ articlerender. London: Chapman & Hall.12. Cell 127 (7): 1309–21. Graham CM (2001). EC. doi:10.. Darwin's Dangerous Idea: Evolution and the Meanings of Life. doi:10. John H. ISBN 0-412-07361-7. doi:10. John R. ISBN 0262581116. Science 322 (5908): 1637–1638. org/ doi/ full/ 10. fcgi?tool=pmcentrez& artid=2603666). ISBN 0-465-04426-3. Cambridge: Harvard University Press. Advanced reading • Barton. ISBN 978-1-59102-479-8. The Top 10 Myths about Evolution. Brian Keith (2003). (2009). Speciation.B. Sylvia S.). J.A. Evolution: The Triumph of an Idea. The Theory of Evolution: Canto Edition. • Gould. The Rough Guide to Evolution. C. Wendy. C.W. Almost Like a Whale: The Origin of Species Updated. The Selfish Gene: 30th Anniversary Edition. • Larson. • Pallen. ISBN 0-879-69684-2. Oxfordshire: Oxford University Press. J. ISBN 0-521-45128-0. ISBN 0-198-50294-X. M. N.A. (2001). ISBN 0-345-42277-5. • Olson. (2003). D. (2001). On the Origin of Species (http://darwin-online. Evolution. (2005).B. The Structure of Evolutionary Theory.uk/content/ frameset?itemID=F373&viewtype=text&pageseq=1) (1st ed. (1993). • Coyne.E. and Charlesworth. C. (2004). . ISBN 0-393-30700-X. McGraw Hill. Evolution: The Remarkable History of a Scientific Theory. ISBN 0-878-93187-2. (2001). D. Pendarvis (9th ed. ISBN 9780073258393. (2007). Norton. ISBN 0-679-64288-9. ISBN 0-878-93089-2. (2006). C.J.W. Goldstein. E. ISBN 0-674-00613-5. • Dawkins. ISBN 0-393-06016-0. • Charlesworth. ISBN 0-674-02240-8. New York: Basic Books. S.H. and Sullivan. ISBN 0801413192. E. London: John Murray. Endless Forms Most Beautiful. • Jones. R. Oxford University Press. J. and Szathmáry. S. Eisen.Evolution 45 Further reading Introductory reading • Carroll. H. • Zimmer. Cambridge University Press. What Evolution Is.. and Orr. Oxfordshire: Oxford University Press. Cold Spring Harbor Laboratory Press. New York: Ballantine Books. Wonderful Life: The Burgess Shale and the Nature of History. The Major Transitions in Evolution. S. (2007).G. Charles (1859).H. • Futuyma. Prometheus Books. D. Sunderland: Sinauer Associates. Evolution. Norton. New York: W. (2005). J. • Maynard Smith. (1997)..org. • Maynard Smith. (American title: Darwin's Ghost). Hall. N. Rough Guides. E. (2002). ISBN 0199291152.B.J. Sunderland: Sinauer Associates. • Mader. New York: Modern Library.J.).J. • Smith.J. ISBN 978-1-85828-946-5. Evolution. S. History of evolutionary thought • Darwin.. (1989). Keywords and concepts in evolutionary developmental biology. (2007). and Patel.A. D. • Gould. Biology. (2004). ISBN 0-060-19906-7. Cambridge: Belknap Press (Harvard University Press). Briggs. Murray P. New York: W. London: HarperCollins. • Mayr. ISBN 0-192-80251-8. bbc.net/ articles/understanding_evolution. Carroll .htm) On-line lectures • What Genomes Can Tell Us About the Past (http://ascb. and Implications (http://www. Berkeley (http://evolution.co.bbc. Theory.uk/programmes/p00545gl) on In Our Time at the BBC.org/evolution/) • Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and Theories (http://anthro.cfm) – lecture by Sydney Brenner • The Origin of Vertebrates (http://ascb.uk/iplayer/console/p00545gl/In_Our_Time_Evolution)) • Everything you wanted to know about evolution by New Scientist (http://www.howstuffworks.cfm) – lecture by Marc Kirschner • The Making of the Fittest (http://www.edu/) • Evolution of Evolution .org/ibioseminars/kirschner/kirschner1.rationalrevolution.com/topic/ evolution) • Howstuffworks.org.co. palomar.html) – lecture by Sean B.htm) • National Academies Evolution Resources (http://nationalacademies.molbio.wisc.com/evolution/evolution.jsp) History of evolutionary thought • The Complete Work of Charles Darwin Online (http://darwin-online.150 Years of Darwin's "On the Origin of Species" (http://www.uk/) • Understanding Evolution: History.berkeley.Evolution 46 External links General information • Evolution (http://www. ( listen now (http:// www.edu/carroll/Fittest.edu/synthetic/) • Understanding Evolution from University of California.newscientist.nsf.org/ibioseminars/brenner/brenner1.gov/news/ special_reports/darwin/textonly/index. Evidence.com — How Evolution Works (http://science. Now in this use of the term fact. Evolution Evolution is usually defined simply as changes in trait or gene frequency in a population of organisms from one generation to the next. hypothesis. principle.[12] [13] [14] [15] "Fact" is also used in a wider sense to mean any theory for which there is overwhelming evidence. "fact". is also used more broadly to incorporate processes such as natural selection and genetic drift. This illustrates a confusion in terminology that hampers discussion. and dandelions. bees. and act as if it were true. the only proper one. There is no sharp line between speculation. "evolution" is often used to include the following additional claims: 1. One meaning is empirical: evolution can be observed through changes in allele frequencies or traits of a population over successive generations. evolution is said to be a fact in the same way as the Earth revolving around the Sun is a fact. but only a difference along a sliding scale. they mean it is a fact that all living organisms have descended from a common ancestor (or ancestral gene pool) [8] even though this cannot be directly observed. giraffes. When we say a thing is a fact. fact and theory Evolution has been described as "fact and theory". it embraces everything from slight changes in the proportion of different alleles within a population (such as those determining blood types) to the successive alterations that led from the earliest proto-organism to snails. When scientists say "evolution is a fact". in the degree of probability of the idea.[9] [10] The meanings of the terms "evolution". This implies more tangibly that it is a fact that humans share a common ancestor with other primates.[16] [17] The following quotation from H. Evolution. 2. they are using one of two meanings of the word "fact". According to Douglas Futuyma: Biological evolution may be slight or substantial. J. we only mean that its probability is an extremely high one: so high that we are not bothered by doubt about it and are ready to act accordingly. —Douglas Futyuma[16] Evolution is a fact in the sense that it is overwhelmingly validated by the evidence. and fact. All living organisms alive today have descended from a common ancestor (or ancestral gene pool). When scientists say evolution is a fact in this sense. A fact is a hypothesis that is so firmly supported by evidence that we assume it is true. "One Hundred Years Without Darwin Are Enough" explains the point. Differences in trait composition between isolated populations over many generations may result in the origin of new species.Evolution as theory and fact 47 Evolution as theory and fact The statement "evolution is both a theory and a fact" is often seen in biological literature. Frequently.[3] . However. especially when referred to as a "theory".[11] The term "evolution". Another way "fact" is used is to refer to a certain kind of theory. theory. then. Muller. "fact not theory". and "theory" are described below. not a fact". evolution is a fact. Fact Fact is often used by scientists to refer to experimental or empirical data or objective verifiable observations. and "only a theory.[1] [2] [3] [4] [5] [6] [7] Evolution is a "theory" in the scientific sense of the term "theory". it is an established scientific model that explains observations and makes predictions through mechanisms such as natural selection. one that has been so powerful and productive for such a long time that it is universally accepted by scientists. just as gravity is both a fact and a theory. that masses attract one another) and the theory used to explain the facts (the reason why masses attract one another). People throughout history have wondered what causes this effect. for example fruit flies. The scientific definition of the word "theory" is different from the colloquial sense of the word. Colloquially. The "theory of evolution" is the framework that best explains observed changes of species over time and best predicts the new observations that continue to be made in evolutionary biology and related sciences.. based on a model that explains all the available data and observations (and provides testable predictions).) The word "gravity". for example. a current theory is a theory that has no equally acceptable or more acceptable alternative theory. However. the meaning of theory is more rigorous: a theory must be based on observed facts and make testable predictions. evolutionary change — has been observed in the laboratory. an opinion. transmutationism and orthogenesis were all non-Darwinian theories that attempted to explain the observations of species and fossils. In this way. and so forth.[21] The observation of fruit fly populations changing over time is also an example of a fact. In science. In this sense all facts are provisional. realized that the fact of gravity can be extended to the tendency of any two masses to attract one another.[1] [19] 48 Theory A scientific theory is a well-supported body of interconnected statements that explains observations and can be used to make testable predictions. . But scientists can also use fact to mean something that has been tested or observed so many times that there is no longer a compelling reason to keep testing or looking for examples. the process of change from generation to generation — that is. can be used to refer to the observed facts (i. Thus.S. and Einstein have all developed useful models of gravity. Many explanations have been proposed over the centuries. Scientists no longer question whether descent with modification occurred because the evidence is so strong. Evolution compared with gravity The application of the terms "fact" and "theory" to evolution is similar to their use in describing gravity. Galileo. The occurrence of evolution in this sense is fact. In science. therefore. gravity is both a theory and a fact.[20] The most obvious fact of gravity is that objects in our everyday experience tend to fall downwards when not otherwise prevented from doing so. Lamarckism.[18] Philosophers of science argue that we do not know anything with absolute certainty: even direct observations may be "theory laden" and depend on assumptions about our senses and the measuring instruments used. the modern theory of evolution is the explanation for all relevant observations regarding the development of life.e. Newton. Scientific theories describe the coherent framework into which observable data fit.Evolution as theory and fact The National Academy of Science (U. "theory" can mean a conjecture. (Newton. Aristotle. In the study of biological species. or a speculation that does not have to be based on facts or make testable predictions. So evolution is a fact just as the observations of gravity are a fact. as well as other evidence. the remains of extinct species in the fossil record. In species that rapidly reproduce. some very similar to each other and some very dissimilar. the facts include the existence of many different species in existence today. evolution is not only a fact but also a theory.) makes a similar point: Scientists most often use the word "fact" to describe an observation. There have been many attempts to explain these biological observations over the years. each of which constitutes a theory of gravity. I will grant you that in an absolute sense evolution is not a fact. J. Theories are structures of ideas that explain and interpret facts. and for pity's sake."[24] • Douglas Futuyma writes in his Evolutionary Biology book. kangaroos. biologist Richard Lenski says. in this context. "Scientific understanding requires both facts and theories that can explain those facts in a coherent manner.. among biologists there is a consensus that evolution is a fact: • American zoologist and paleontologist George Simpson stated that "Darwin. "biologists rarely make reference to 'the theory of evolution. Ryan Gregory says. let's stop confusing the philosophically naive by calling it so.. then. starfish. It is also a fact."[3] • Kenneth R. Evolution. but apples did not suspend themselves in mid-air. that it is no more a fact than that you are hearing or reading these words. And humans evolved from ape-like ancestors whether they did so by Darwin's proposed mechanism or by some other yet to be discovered. "evolution is as much a fact as anything we know in science.. the changes that occur within species over generations. "The statement that organisms have descended with modifications from common ancestors—the historical reality of evolution—is not a theory. not theory. It is an incontrovertible fact that organisms have changed. Einstein's theory of gravitation replaced Newton's. "It is time for students of the evolutionary process. is both a fact and a theory. as fully as the fact of the earth's revolution about the sun. the increasingly sophisticated body of explanations for the fact of evolution). "Evolution is a theory. the fact of descent with modification) or 'evolutionary theory' (i. It is a fact that we are cousins of gorillas. That evolution is a theory in the proper scientific sense means that there is both a fact of evolution to be explained and a well-supported mechanistic framework to account for it. especially those who have been misquoted and used by the creationists."[7] Evolution as fact and theory Commonly "fact" is used to refer to the observable changes in organisms' traits over generations while the word "theory" is reserved for the mechanisms that cause these changes: • Paleontologist Stephen Jay Gould writes."[6] • Biologist T. Miller writes. or evolved. "If you like."[23] • Ernst Mayr observed. Evolution is a ."[2] • Similarly.' referring instead simply to 'evolution' (i.Evolution as theory and fact 49 Evolution as theory and fact in the literature The confusion over the word evolution and the distinction between "fact" and "theory" is largely due to authors using evolution to refer to three related yet distinct ideas: first. C.e. second. finally and definitely established evolution as a fact. However. "The basic theory of evolution has been confirmed so completely that most modern biologists consider evolution simply a fact. focusing on the changes in species over generations and in some cases common ancestry have stressed that evolution is a fact to emphasize the weight of supporting evidence while denying it is helpful to use the term "theory": • R. And facts and theories are different things. to state clearly that evolution is a fact. not rungs in a hierarchy of increasing certainty. and bacteria. or rather. Evolution is as much a fact as the heat of the sun."[11] • Richard Dawkins says. Muller wrote. and third. during the history of life on Earth. the concept of common descent. Lewontin wrote. the mechanism thought to drive change. Facts do not go away when scientists debate rival theories to explain them."[20] Evolution as fact not theory Other commentators. How else except by the word evolution can we designate the sequence of faunas and floras in precisely dated geological strata? And evolutionary change is also simply a fact owing to the changes in the content of gene pools from generation to generation. "One thing all real scientists agree upon is the fact of evolution itself. It is not a theory. Facts are the world's data. And biologists have identified and investigated mechanisms that can explain the major patterns of change."[22] • H. It is a fact. pending the outcome.e.. It has been predicted by the theory of evolution that the differences in such DNA sequences between two organisms should roughly resemble both the biological difference between them according to their anatomy and the time that had passed since these two organisms have separated in the course of evolution. In rocks more than 385 million years old they could only find fish."[4] 50 Predictive power A central tenet in science is that a scientific theory is supposed to have predictive power.[26] • Some DNA sequences are shared by very different organisms. Evolutionary theory predicted that since amphibians evolved from fish. Well-tested explanations are called theories. The rate of accumulating such changes should be low for some sequences. These results have been experimentally confirmed. Since this prediction was made. and verification of predictions are seen as an important and necessary support for the theory. and DNA sequences coding for fibrinopeptides (amino acid chains that are discarded during the formation of fibrin). ears. coli. Two examples are DNA sequences coding for rRNA. Very well-established "theories" that rely on a simple principle are often called scientific "laws". this provides just such a mechanism. Such an intermediate form should have many fish-like characteristics. Richard Lenski observed that E. it is important to understand that the current questions about how life evolves in no way implies any disagreement over the fact of evolution. coli evolved the ability to metabolize citrate. Gould: In science. and an increase in the information of the DNA of the E. an intermediate form should be found in rock dated between 365 and 385 million years ago. "the law of natural selection". The term theory is no longer appropriate except when referring to the various models that attempt to explain how life evolves. In the words of Stephen J. and four legs. Proof only exists in formal sciences. such as mathematics. In 2004. which has a mutation rate of roughly 10−9 per nucleotide per cell division. which are highly non-conserved.[28] In an ongoing experiment.[27] • Evolutionary theory predicts that novel inventions can arise. paleontologists had found fossils of amphibians with necks.Evolution as theory and fact fact. or the "laws of evolution. conserved from 385 million years ago or more.. namely those that code for critical RNA or proteins. but for every specific sequence.[29] Related concepts and terminology • Speculative or conjectural explanations are called hypotheses. Experimental observation of the predictions made by a hypothesis or theory is called validation. The theory of evolution has provided such predictions. Four examples are: • Genetic information must be transmitted in a molecular way that will be almost exact but permit slight changes. which constitutes a novel invention. the rate of change should be roughly constant over time. as seen in fossil evidence.. while creationists predict that new "information" cannot arise. and high for others that code for less critical RNA or proteins. in rock no older than 365 million years old.[26] • Prior to 2004. which is highly conserved. For example. • A scientific law is a concept related to a scientific theory. an expedition to islands in the Canadian arctic searching specifically for this fossil form in rocks that were 375 million years old discovered fossils of Tiktaalik."[25] • Neil Campbell wrote in his 1990 biology textbook. nearly all biologists acknowledge that evolution is a fact. it is common to encounter reference to "the law of gravity"." . without these amphibian characteristics. but also have many amphibian characteristics as well. biologists have discovered the existence of DNA. and that the Second Law of Thermodynamics only allows for "information" to be lost."[2] • "Proof" of a theory does not exist in natural sciences. "fact" can only mean "confirmed to such a degree that it would be perverse to withhold provisional assent. • "Fact" does not mean "absolute certainty". "Today. [8] Cavalier-Smith T (2006). Second Edition (1999). • . nih. Ernst (1988). php?record_id=6024& page=28) [19] Wilkins. Phoenix AZ: ORYX Press.Evolution as theory and fact 51 See also • • • • • Epistemology Evidence of common descent Evolution is just a theory. H. Reprinted in: • Zetterberg. doi:10. ISBN 0897740610. (2000). [23] "Miller. Stephen Jay (1981-05-01). delphiforums. talkorigins. and What Does 'Science' Mean?" (http:/ / www. Cambridge: Harvard University Press... Retrieved 2007-10-18. org). .1959. nps. J. 1175.1038/230289a0. Pavlovsky O (1971). Evolution Versus Creationism: The Public Education Controversy.) (1983-05-01).) (1983-05-01). html). htm).) (1982). Richard E. Finding Darwin's God: A Scientist's Search for Common Ground Between God and Evolution (P. Facts. htm#F) [14] Webster's New Millennium Dictionary of English. actionbioscience. religioustolerance. [6] Lenski. "Experimentally created incipient species of Drosophila". Peter (ed. . reference. ISBN 0-674-89666-1. Jane B. htm). [7] Mayr. [4] Campbell. . Ryan (2007). html). stephenjaygould.A. doi:10. Benjamin Cummings. (ed. [5] Dobzhansky. something known to be true: 'Scientists gather facts about plant growth. Retrieved 2009-08-17. Herbert F. discovery. Kenneth S. org/ library/ futuyma_theory. doi:10. • Gould. "Evolution as Fact and Theory" (http:/ / www. co. csicop. Reece. edu/ openbook.Y: Harper Perennial. org/ library/ gould_fact-and-theory. (http:/ / books. gov/ views/ System/ Glossary. html). Speak Out Against The New Right. p. B. com/ search?q=scientific+ fact& r=66) [15] Webster's Encyclopedic Unabridged Dictionary of the English Language (1996) gives a third meaning of the word "fact" as (3) A truth known by actual experience or observation. (2007). "Is the theory of evolution merely a "theory"?" (http:/ / www. Evolution Versus Creationism: The Public Education Controversy. The TalkOrigins Archive (http:/ / www. "Evolution and Philosophy:Is Evolution Science. [3] Muller. ISBN 0807004863. LLC (http:/ / dictionary. 2006. skepticfiles. org/ faqs/ evolphil/ falsify. Philos Trans R Soc Lond B Biol Sci 361 (1470): 969–1006. N. Jerry Coyne (http:/ / www. New York. "One hundred years without Darwin are enough" (http:/ / www. p. (2002-02-05). [20] Gregory. "Nothing in biology makes sense except in the light of evolution" (http:/ / people. stephenjaygould. National Academy Press. "Evolution as Fact. Discover 2 (5): 34–37. Fact (in Creation-evolution controversy) Notes [1] Moran. [9] Is "Evolution" a "Theory" or "Fact" or Is This Just a Trivial Game of Semantics? (http:/ / www. JS (1997). and Path". [2] Gould. "Evolution: Fact and Theory" (http:/ / www. . edu/ perl/ webwn?s=scientific fact) [13] United States National Park Service Glossary (http:/ / www2. Phoenix AZ: ORYX Press. PMID 16754610. [12] Wordnet entry for phrase "scientific fact" (http:/ / wordnet. pubmedcentral. Evolution: Education and Outreach 1 (1): 46–52. (2005-08-30). Reprinted in: Zetterberg.6). nature.1842.9. (1997). . org/ evolut/ 100pcnts. com/ lordorman/ light. nap. Peter (ed.1949-8594. Washington DC. org/ evolution/ lenski.1098/rstb. ISBN 0393017168. "Cell evolution and Earth history: stasis and revolution" (http:/ / www. org/ a/ 6401) by Casey Luskin [10] Committee for Skeptical Inquiry — Evolution & Creationism: Terminology in Conflict (http:/ / www. [21] Dobzhansky T. ISBN 0897740610. research) [18] Science and Creationism: A View from the National Academy of Sciences. . html).).2006. html) [17] Guardian article by Richard Dawkins. ISBN 0878931899. Reprinted in: • Vetter.1007/s12052-007-0001-z. American Institute of Biological Sciences. Hen's Teeth and Horse's Toes. htm). princeton. org/ ev_stat. Biology 6th ed. —Douglas Futyuma (http:/ / www. . 751. PMID 5549403. New York: Norton. Laurence (1993-01-22). gov/ articlerender. ISBN 0805366245. org/ faqs/ evolution-fact.tb08235. Theory. doi:10. Retrieved 2007-10-18.S. T. Beacon Press. uk/ science/ 2005/ sep/ 01/ schools. Toward a New Philosophy of Biology: Observations of an Evolutionist. National Academy of Sciences (NAS). not a fact (in objections to evolution) Misconceptions about evolution (in List of misconceptions) Theory vs. Theodosius (1973-03-01). guardian. talkorigins. Nature 230 (5292): 289–92. org/ specialarticles/ show/ evolution_amp_creationism_terminology_in_conflict/ ) by Richard Joltes [11] Futuyma.' [16] Hypotheses. Evolutionary Biology. School Science and Mathematics 59: 304–305. Preview Edition (v 0. Douglas J.1111/j. and the Nature of Science.x. Neil A. Retrieved 2007-10-18. [22] Robinson. "Evolution is a Fact and a Theory" (http:/ / www. (1959). . fcgi?tool=pmcentrez& artid=1578732). PMC 1578732. Sinauer Associates. talkorigins.. American Biology Teacher 35. ISBN 0061233501. Stephen Jay (1994-04-01).origins. 3rd ed. Copyright © 2003–2006 Lexico Publishing Group. Talk. org/indexcc/CB/CB910. Martin Raff. Springer Science + Business Media.P.com/ content/fr258627q2x3t378/fulltext. Phoenix AZ: Oryx Press. (1981). ISBN 9780375424472. Dept. talkorigins.com) Discredits the assertion that evolution is "just a theory". Grahamstown.). Your Inner Fish. "The biology of four haplochromine species of Lake Kivu (Zaire) with evolutionary implications. (1983-05-01). C. ed. with an explanation of the meaning of the word 'theory' in a scientific context. Halifax. Evo Edu Outreach (2008) 1:287–289. Neil.Evolution as theory and fact [24] Lewontin. Alexander Johnson. (2008). Louise S. Losseau-Hoebeke. Peter.. Peter Walter (March." Annual Meeting Canadian Federation of Biological Societies. org/ faqs/ thermo/ probability. Bioscience 31: 559. (1990).talkorigins. [27] "Shubin. • Talk Origins (http://www. Franck. Mead (2008-06-06). Rhodes University. html) [29] NS:bacteria make major evolutionary shift in the lab (http:/ / www. Reprinted in: Zetterberg.notjustatheory. "Evolution of a satellite DNA family in tilapia. . External links • Not Just a Theory (http://www. 2002). Ichthyology. naturalhistorymag. Julian Lewis. • M. "“Theory” in Theory and Practice" (http://www. Evolution Versus Creationism: the public education controversy.pdf) (pdf). "Evolution/Creation Debate: a time for truth". • Glenn Branch. R. htm) • 52 References • J. Routledge. et al. com/ channel/ life/ dn14094-bacteria-make-major-evolutionary-shift-in-the-lab. Molecular Biology of the Cell (4th ed. [28] TalkOrigins. (1992)." Thesis. newscientist. [25] Natural History article : The Illusion of Design (http:/ / www.org (http:/ / www. Pantheon. ISBN 0897740610.html) Response to the claim that no examples of speciation have been observed.springerlink. Keith Roberts. ISBN 0-8153-3218-1. Retrieved 2008-07-21. com/ features/ 101500/ the-illusion-of-design) by Richard Dawkins [26] Bruce Alberts. [14] Invertebrate animals appear during the Vendian period. archosaurs became the most abundant land vertebrates.0 [11] million years ago.[21] One archosaur group. multicellular organisms began to appear.[5] The earliest evidence of eukaryotes (complex cells with organelles). Social insects appeared around the same time as flowering plants. including the ancestors of mammals. with differentiated cells performing specialised functions. around 1700 [9] million years ago. there are signs of a steady increase in brain size after about 3 [31] million years ago. The similarities between all present day organisms indicate the presence of a common ancestor from which all known species have diverged through the process of evolution.Evolutionary history of life 53 Evolutionary history of life The evolutionary history of life on Earth traces the processes by which living and fossil organisms evolved.[17] During the Permian period.[23] After the Cretaceous–Tertiary extinction event 65 [24] million years ago killed off the non-avian dinosaurs[25] mammals increased rapidly in size and diversity.[22] with the ancestors of mammals surviving only as small insectivores. Although they occupy only small parts of the insect "family tree". around 3500 [1] million years ago. thought to be over 3500 [1] million years ago. Flowering plants and marine phytoplankton are still the dominant producers of organic matter.[2] Microbial mats of coexisting bacteria and archaea were the dominant form of life in the early Archean and many of the major steps in early evolution are thought to have taken place within them. dominated the Jurassic and Cretaceous periods.[12] though evidence suggests that algal scum formed on the land as early as 1200 [13] million years ago. synapsids. beginning around 2400 [4] million years ago. It stretches from the origin of life on Earth.[20] During the recovery from this catastrophe. probably helped by coevolution with pollinating insects.[3] The evolution of oxygenic photosynthesis. between 130 [28] million years ago and 90 [29] million years ago. Later. the dinosaurs. dominated the land. dates from 1850.0 [19] million years ago came close to wiping out all complex life.[26] Such mass extinctions may have accelerated evolution by providing opportunities for new groups of organisms to diversify.0 [6] million years ago. their diversification accelerated when they started using oxygen in their metabolism.[18] but the Permian–Triassic extinction event 251.[27] Fossil evidence indicates that flowering plants appeared and rapidly diversified in the Early Cretaceous.[10] The earliest land plants date back to around 450. they now form over half the total mass of insects.[7] [8] and while they may have been present earlier. eventually led to the oxygenation of the atmosphere. Although early members of this lineage had chimp-sized brains. Earliest history of Earth History of Earth and its life . Land plants were so successful that they are thought to have contributed to the late Devonian extinction event. to the present day.[15] while vertebrates originated about 525 [16] million years ago during the Cambrian explosion. displacing therapsids in the mid-Triassic. Humans evolved from a lineage of upright-walking apes whose earliest fossils date from over 6 [30] million years ago. Evolutionary history of life 54 Hadean Archean Protero -zoic Phanero -zoic Eo Paleo Meso Neo Paleo Meso Neo Paleo Meso Ceno Scale: Millions of years The oldest meteorite fragments found on Earth are about 4540 [32] million years old.[33] About 40 million years later a planetoid struck the Earth. coupled primarily with the dating of ancient lead deposits. has put the estimated age of Earth at around that time. throwing into orbit the material that formed the Moon.[34] . this. interpreted as fossilized bacteria.Evolutionary history of life Until recently the oldest rocks found on Earth were about 3800 [35] million years old.. The three domains are colored.. the oldest unchallenged evidence for life is geochemical signatures from rocks deposited 3400 [49] million years ago. whose name means "hellish". they named this part of Earth's history the Hadean eon.[45] with geochemical evidence also seeming to show the presence of life 3800 [35] million years ago.[42] 55 Earliest evidence for life on Earth The earliest identified organisms were minute and relatively featureless. was "seeded" from elsewhere in the universe.[36] However analysis of zircons formed 4400 to 4000 [37] million years ago indicates that Earth's crust solidified about 100 million years after the planet's formation and that the planet quickly acquired oceans and an atmosphere.. C. and the Earth should have experienced an even heavier bombardment due to its stronger gravity. which are very difficult to tell apart from structures that arise through abiotic physical processes.[38] Evidence from the Moon indicates that from 4000 to 3800 [39] million years ago it suffered a Late Heavy Bombardment by debris that was left over from the formation of the Solar system. doi:10. von Mering. in this case gases and water from comet impacts may have contributed to their replacement. which fall into two main groups: 1) that life arose spontaneously on Earth or 2) that it Evolutionary tree showing the divergence of modern species from their common ancestor in the center.. Origins of life on Earth Biologists reason that all living organisms on Earth must share a single last universal ancestor.[51] [52] As previously mentioned the earliest organisms for which fossil evidence is available are bacteria. C. although volcanic outgassing on Earth would have contributed at least half.[33] leading scientists to believe for decades that Earth's surface had been molten until then. because it would be virtually impossible that two or more separate lineages could have independently developed the many complex biochemical mechanisms common to all living organisms. and non-biological processes were found which could produce all of the "signatures of life" that had been reported.[47] [48] While this does not prove that the structures found had a non-biological origin. Accordingly.1126/science. PMID 16513982. Science 311 (5765): 1283–7. Doerks. with bacteria blue.[44] Other finds in rocks dated to about 3500 [1] million years ago have been interpreted as bacteria. Currently. which may have been capable of supporting life. and eukaryotes red. Creevey. T.J. dates to 3000 [43] million years ago.[46] However these analyses were closely scrutinized.[53] The lack of fossil or geochemical evidence for earlier organisms has left plenty of scope for hypotheses.D. archaea green. "Toward automatic reconstruction of a highly resolved tree of life". et al (2006). F.1123061. there is no reason to think that the Earth was not also affected by this late heavy bombardment.Ciccarelli. The oldest undisputed evidence of life on Earth. cells far too complex to have arisen directly from non-living materials.[44] [50] although these statements have not been thoroughly examined by critics.[36] [40] While there is no direct evidence of conditions on Earth 4000 to 3800 [39] million years ago.[41] This event may well have stripped away any previous atmosphere and oceans. . they cannot be taken as clear evidence for the presence of life. their fossils look like small rods. its ability to feed and repair itself. and lack water's other advantages.Evolutionary history of life 56 Life "seeded" from elsewhere The idea that life on Earth was "seeded" from elsewhere in the universe dates back at least to the fifth century BCE.[58] Scientists are divided over the likelihood of life arising independently on Mars.[54] In the twentieth century it was proposed by the physical chemist Svante Arrhenius. would be more difficult for organisms to extract. possibly as a result of accidental contamination by micro-organisms that they brought with them.[61] Research on how life might have emerged unaided from non-living chemicals focuses on three possible starting points: self-replication.[56] and by molecular biologist Francis Crick and chemist Leslie Orgel.[58] Independent emergence on Earth Life on earth is based on carbon and water. which would have been replaced later by RNA. in which case the only credible source is Mars. since theoretical and empirical approaches are only beginning to make contact with each other. silicon. such as ammonia. DNA's structure and replication systems are far more complex than those of the original replicator. because most of its compounds are solids.[60] Organisms based on alternative biochemistry may however be possible on other planets. especially from carbon dioxide. an organism's ability to produce offspring that are very similar to itself.[55] [58] Experiments suggest that some micro-organisms can survive the shock of being catapulted into space and some can survive exposure to radiation for several days. Water is an excellent solvent and has two other useful properties: the fact that ice floats enables aquatic organisms to survive beneath it in winter.[72] [73] In 2003 it was proposed that porous metal sulfide precipitates would assist RNA synthesis at about 100 °C (212 °F) and ocean-bottom pressures near hydrothermal vents. which enables it to form a wider range of compounds than other solvents can.[66] RNA would later have been replaced by DNA. The only other element with similar chemical properties. TNA or GNA.[55] by the astronomers Fred Hoyle and Chandra Wickramasinghe. Carbon provides stable frameworks for complex chemicals and can be easily extracted from the environment.[59] or on other planets in our galaxy. and its molecules have electrically negative and positive ends. Other good solvents. expanding the range of capabilities a single organism can have. This system is far too complex to have emerged directly from non-living materials.[57] and from outside the Solar system but by natural means. forms much less stable structures and.[58] by alien visitors.[53] The discovery that some RNA molecules can catalyze both their own replication and the construction of proteins led to the hypothesis of earlier life-forms based entirely on RNA. [53] Even the simplest members of the three modern domains of life use DNA to record their "recipes" and a complex array of RNA and protein molecules to "read" these instructions and use them for growth. are liquid only at such low temperatures that chemical reactions may be too slow to sustain life. which allow food to enter and waste products to leave.[69] Although short self-replicating RNA molecules have been artificially produced in laboratories.[70] doubts have been raised about where natural non-biological synthesis of RNA is possible. and external cell membranes. In this hypothesis lipid membranes would be the last major cell . maintenance and self-replication. metabolism. modern cells' "protein factories". but exclude unwanted substances. but there is no proof that they can survive in space for much longer periods.[63] [64] Replication first: RNA world The replicator in virtually all known life is deoxyribonucleic acid.[66] [67] [68] Ribozymes remain as the main components of ribosomes. which is more stable and therefore can build longer genomes.[62] Research on abiogenesis still has a long way to go.[65] These ribozymes could have formed an RNA world in which there were individuals but no species.[57] There are three main versions of the "seeded from elsewhere" hypothesis: from elsewhere in our Solar system via fragments knocked into space by a large meteor impact.[71] The earliest "ribozymes" may have been formed of simpler nucleic acids such as PNA. as mutations and horizontal gene transfers would have meant that the offspring in each generation were quite likely to have different genomes from those that their parents started with. The formation of the earliest cells may have been aided by similar processes.[81] . have properties that make them plausible accelerators for the emergence of an RNA world: they grow by self-replication of their crystalline pattern. they would be subject to natural selection for longevity and reproduction. and then reproduce themselves. notably montmorillonite. although one stage required 250 °C (482 °F) and a pressure equivalent to that found under 7 kilometres (4.[77] The clay theory RNA is complex and there are doubts about whether it can be produced non-biologically in the wild.[76] Experiments that simulated the conditions of the early Earth have reported the formation of lipids.[71] Some clays. Most of the steps required temperatures of about 100 °C (212 °F) and moderate pressures.Evolutionary history of life components to appear and until then the proto-cells would be confined to the pores.[80] A similar hypothesis presents self-replicating iron-rich clays as the progenitors of nucleotides. lipids and amino acids. Hence it was suggested that self-sustaining synthesis of proteins could have occurred near hydrothermal vents. and these can spontaneously form liposomes.[75] Membranes first: Lipid world 57 = water-attracting heads of lipid molecules = water-repellent tails Cross-section through a liposome. double-walled "bubbles". they are subject to an analog of natural selection.[74] Metabolism first: Iron-sulfur world A series of experiments starting in 1997 showed that early stages in the formation of proteins from inorganic materials including carbon monoxide and hydrogen sulfide could be achieved by using iron sulfide and nickel sulfide as catalysts. and they can catalyze the formation of RNA molecules. These "bubbles" can then grow by absorbing additional lipids and then divide.3 mi) of rock.[78] Although this idea has not become the scientific consensus. Although they are not intrinsically information-carriers as nucleic acids are.[79] Research in 2003 reported that montmorillonite could also accelerate the conversion of fatty acids into "bubbles". and that the "bubbles" could encapsulate RNA attached to the clay. Nucleic acids such as RNA might then have formed more easily within the liposomes than they would have outside. it still has active supporters. as the clay "species" that grows fastest in a particular environment rapidly becomes dominant. It has been suggested that double-walled "bubbles" of lipids like those that form the external membranes of cells may have been an essential first step. and then downwards as it resumed on the next day.[47] In 2006 another find of stromatolites was reported from the same part of Australia as previous ones.[83] It is estimated that the appearance of oxygenic photosynthesis by bacteria in mats increased biological productivity by a factor of between 100 and 1. from which all multicellular organisms are built.[3] .[91] [92] the oxygenation of the atmosphere was a prerequisite for the evolution of the most complex eukaryotic cells.[83] Stromatolites are stubby pillars built as microbes in mats slowly migrate upwards to avoid being smothered by sediment deposited on them by water. Western Australia. multi-species colonies of bacteria and other organisms that are generally only a few millimeters thick.000. possibly via endosymbiosis.[90] Although eukaryotes may have been present much earlier.[85] In modern underwater mats the top layer often consists of photosynthesizing cyanobacteria which create an oxygen-rich environment.[3] Cyanobacteria have the most complete biochemical "toolkits" of all the mat-forming organisms. Hence they are the most self-sufficient of the mat organisms and were well-adapted to strike out on their own both as floating mats and as the first of the phytoplankton. providing the basis of most marine food chains.[82] There has been vigorous debate about the Modern stromatolites in Shark Bay. which is much more plentiful than the geologically-produced reducing agents required by the earlier non-oxygenic photosynthesis.[93] The boundary between oxygen-rich and oxygen-free layers in microbial mats would have moved upwards when photosynthesis shut down overnight. in rocks dated to 3500 [1] million years ago. but still contain a wide range of chemical environments.Evolutionary history of life 58 Environmental and evolutionary impact of microbial mats Microbial mats are multi-layered. each of which favors a different set of micro-organisms. as the by-products of each group of micro-organisms generally serve as "food" for adjacent groups. while the bottom layer is oxygen-free and often dominated by hydrogen sulfide emitted by the organisms living there.[82] To some extent each mat forms its own food chain. The reducing agent used by oxygenic photosynthesis is water. This would have created selection pressure for organisms in this intermediate zone to acquire the ability to tolerate and then to use oxygen. where one organism lives inside another and both of them benefit from their association.[88] [89] Oxygen became a significant component of Earth's atmosphere about 2400 [4] million years ago.[86] From this point onwards life itself produced significantly more of the resources it needed than did geochemical processes. validity of alleged fossils from before 3000 [43] million years ago.[87] Oxygen is toxic to organisms that are not adapted to it. but greatly increases the metabolic efficiency of oxygen-adapted organisms.[84] with critics arguing that so-called stromatolites could have been formed by non-biological processes. probably an archaean.[100] There is a debate about when eukaryotes first appeared: the presence of steranes in Australian shales may indicate that eukaryotes were present 2700 [101] million years ago.[91] but most modern eukaryotes require oxygen.[110] .[108] The earliest known fossils of fungi date from 1430 [109] million years ago. a process which sometimes involved transfer of genes between them.Evolutionary history of life 59 Diversification of eukaryotes Apusozoa Archaeplastida (Land plants. which their mitochondria use to fuel the production of ATP. After each endosymbiosis began.[93] In the 1970s it was proposed and. but the victim survived inside the attacker and the new combination became the ancestor of plants. and glaucophytes) Bikonta Chromalveolata Rhizaria Excavata Eukaryotes Amoebozoa Unikonta Opisthokonta Metazoa (Animals) Choanozoa Eumycota (Fungi) One possible family tree of eukaryotes[94] [95] Eukaryotes may have been present long before the oxygenation of the atmosphere. widely accepted that eukaryotes emerged as a result of a sequence of endosymbioses between "procaryotes".or hydrogen-metabolising endosymbionts.[96] [97] [98] Another hypothesis proposes that mitochondria were originally sulfur. after much debate. and the attacker took up residence and evolved into the first of the mitochondria. but the attack was neutralized. green algae. For example: a predatory micro-organism invaded a large procaryote. and indicates that eukaryotes with organelles had already evolved. red algae.[92] however an analysis in 2008 concluded that these chemicals infiltrated the rocks less than 2200 [102] million years ago and prove nothing about the origins of eukaryotes. the partners would have eliminated unproductive duplication of genetic functions by re-arranging their genomes.[103] Fossils of the alga Grypania have been reported in 1850.[105] A diverse collection of fossil algae were found in rocks dated between 1500 [106] million years ago and 1400 [107] million years ago. one of these chimeras later tried to swallow a photosynthesizing cyanobacterium. the internal energy supply of all known cells. and became oxygen-consumers later.0 [6] million-year-old rocks (originally dated to 2100 [104] million years ago but later revised[8] ). and so on.[99] On the other hand mitochondria might have been part of eukaryotes' original equipment. most of which lose the opportunity to reproduce themselves.[119] The disadvantages of sexual reproduction are well-known: the genetic reshuffle of recombination may break up favorable combinations of genes. The mold (yellow) explored and filled the maze (left).Evolutionary history of life 60 Multicellular organisms and sexual reproduction Multicellularity The simplest definitions of "multicellular". There is strong evidence that sexual reproduction arose early in the history of eukaryotes and that the genes controlling it have changed very little since then.[115] The available evidence indicates that eukaryotes evolved much earlier but remained inconspicuous until a rapid diversification around 1000 [116] million years ago. rogue cells which retain the ability to reproduce may take over and reduce the organism to a mass of undifferentiated cells. brown algae. Sexual reproduction eliminates such rogue cells from the next generation and therefore appears to be a prerequisite for complex multicellularity.[115] the ability to create an internal environment that gives protection against the external one.[114] A slime mold solves a maze.[111] These features would also have provided opportunities for other organisms to diversify. many of which attacked by engulfing.[121] . the benefits of which include resistance to antibiotics and other toxins.[115] Evolution of sexual reproduction The defining characteristic of sexual reproduction is recombination. and sexual reproduction enabled eukaryotes to exploit that advantage by producing organisms with multiple cells that differed in form and function. The only respect in which eukaryotes clearly surpass bacteria and archaea is their capacity for variety of forms. could include colonial cyanobacteria like Nostoc.[117] Nevertheless the great majority of animals.[118] However conjugation is not a means of reproduction. in which each of the offspring receives 50% of its genetic inheritance from each of the parents.[120] How sexual reproduction evolved and survived is an unsolved puzzle. and is not limited to members of the same species – there are cases where bacteria transfer DNA to plants and animals. Even a professional biologist's definition such as "having the same genome but different types of cell" would still include some genera of the green alga Volvox. an asexual population can out-breed and displace in as little as 50 generations a sexual population that is equal in every other respect. In an asexual multicellular organism.[114] and even the opportunity for a group of cells to behave "intelligently" by sharing information. fungi and protists reproduce sexually. and the ability to utilize new metabolites. and since males do not directly increase the number of offspring in the next generation. plants. the ability to reach upwards to filter-feed or to obtain sunlight for photosynthesis. cyanobacteria. for example "having multiple cells". The initial advantages of multicellularity may have included: increased resistance to predators. plants. although this approach to the evolution of complexity could be regarded as "rather anthropocentric". by creating more varied environments than flat microbial mats could. which have cells that specialize in reproduction.[115] Multicellularity with differentiated cells is beneficial to the organism as a whole but disadvantageous from the point of view of individual cells. fungi. the mold concentrated most of its mass there and left only the most efficient connection between the two [111] points (right).[117] Bacteria also exchange DNA by bacterial conjugation.[112] Multicellularity evolved independently in organisms as diverse as sponges and other animals.[8] [113] For the sake of brevity this article focuses on the organisms that show the greatest specialization of cells and variety of cell types. slime moulds and myxobacteria. the ability to resist currents by attaching to a firm surface. When the researchers placed sugar (red) at two separate points. as it transfers DNA by means of such a "selfish gene". If so. sexual recombination of genes will reduce the harm that bad mutations do to offspring and at the same time eliminate some bad mutations from the gene pool by isolating them in individuals that perish quickly because they have an above-average number of bad mutations.[124] • It may have evolved from cannibalism. However the evidence suggests that the MDH's assumptions are shaky.[121] Other combinations of hypotheses that are inadequate on their own are also being examined. as one of the clones may survive the attacks of parasites for long enough to out-breed the sexual species.[117] The random nature of recombination causes the relative abundance of alternative traits to vary from one generation to another.[123] 61 Horodyskia apparently re-arranged itself into fewer but larger main masses as the sediment [8] grew deeper round its base. and swapping plasmids. and there is some experimental evidence for this. The meiosis stage of sexual reproduction may then have evolved as a way of removing the symbiotes.[118] However conjugation is not a means of reproduction and is not limited to members of the same species. This genetic drift is insufficient on its own to make sexual reproduction advantageous. and there are cases where bacteria transfer DNA to plants and animals.[123] • Sexual reproduction may have evolved from ancient haloarchaea through a combination of jumping genes. . and the ability to utilize new metabolites. but a combination of genetic drift and natural selection may be sufficient. because many species have on average less than one harmful mutation per individual and no species that has been investigated shows evidence of synergy between harmful mutations.[122] The most primitive form of sex may have been one organism repairing damaged DNA by replicating an undamaged strand from a similar organism. On the other hand the benefits of good traits are neutralized if they appear along with bad traits. Sexual recombination gives good traits the opportunities to become linked with other good traits. When chance produces combinations of good traits. the benefits of which include resistance to antibiotics and other toxins.Evolutionary history of life The Red Queen Hypothesis suggests that sexual reproduction provides protection against parasites.[117] The Mutation Deterministic Hypothesis assumes that each organism has more than one harmful mutation and the combined effects of these mutations are more harmful than the sum of the harm done by each individual mutation.[119] Nevertheless it may be an example of the "selfish genetic element" hypothesis.[117] The following hypotheses attempt to explain how and why sex evolved: • It may have enabled organisms to repair genetic damage. natural selection gives a large advantage to lineages in which these traits become genetically linked. where some of the victim's DNA was incorporated into the cannibal organism. because it is easier for parasites to evolve means of overcoming the defenses of genetically identical clones than those of sexual species that present moving targets. and mathematical models suggest this may be more than enough to offset the disadvantages of sexual reproduction.[125] • Or it may have evolved as a form of vaccination in which infected hosts exchanged weakened symbiotic copies of parasitic DNA as protection against more virulent versions.[126] Bacteria also exchange DNA by bacterial conjugation. However there is still doubt about whether it would explain the survival of sexual species if multiple similar clone species were present. the F-plasmid.[123] • Sexual reproduction may have originated from selfish parasitic genetic elements propagating themselves by transfer to new hosts. 0 [130] million years ago.[8] however it has also been interpreted as a colonial foraminiferan. annelids.[131] Animals are multicellular eukaryotes. sea anemones and hydras). tardigrades. hemichordates.[] They may have had differentiated cells. brachiopods. The red alga called Bangiomorpha. which transmits and processes signals. is the earliest known organism which certainly has differentiated. specialized cells. nematodes.[128] Another early multicellular fossil.[115] The 1430 [109] million-year-old fossils interpreted as fungi appear to have been multicellular with differentiated cells. hydras) Ctenophora (comb jellies) Placozoa Porifera (sponges): Calcarea Porifera: Hexactinellida & Demospongiae Choanoflagellata Mesomycetozoea A family tree of the animals. echinoderms) Bilaterians Protostomes Ecdysozoa (arthropods. including muscles. may have been an early metazoan. sea anemones.[135] The earliest widely-accepted animal fossils are rather modern-looking cnidarians (the group that includes jellyfish.) Acoelomorpha Cnidaria (jellyfish. dated at 1200 [13] million years ago. although fossils from the Doushantuo Formation can only be dated approximately. which move parts of the animal by contracting. [8] or a colonial foraminiferan [127] The Francevillian Group Fossil.[127] Emergence of animals Deuterostomes (chordates. dated to 2100 [104] million years ago. Qingshania. possibly from around 580 [136] million years ago. algae. etc. and is also the oldest known sexually-reproducing organism. found in rocks dated from 1500 [106] million years ago to 900. is the earliest known fossil organism that is clearly multicellular.[110] The "string of beads" organism Horodyskia.[133] All animals are motile.[129] dated to 1700 [9] million years ago. and nerve tissue. etc.) Lophotrochozoa (molluscs.[132] and are distinguished from plants.Evolutionary history of life 62 Fossil evidence for multicellularity and sexual reproduction Horodyskia may have been an early metazoan.[134] if only at certain life stages.[137] . Their presence implies that the cnidarian and bilaterian lineages had already diverged. appears to consist of virtually identical cells. All animals except sponges have bodies differentiated into separate tissues. and fungi by lacking cell walls. about 385 to 359 [160] million years ago.[154] and Haikouichthys had distinct vertebrae. how and why it happened and why it appears unique in the history of animals. Some tiny Early Cambrian shells almost certainly belonged to molluscs. such as Opabinia. and that it may have been closely related to the modern tardigrades. One deuterostome group. first appeared in the Late Ordovician. if so.[146] A re-analysis of fossils from the Burgess Shale lagerstätte increased interest in the issue when it revealed animals. many of which have hard calcite "shells".[138] were the first animals more than a very few centimeters long. are generally absent in the Ediacarans. however. which flourished for the last 40 million years before the start of the Cambrian. However there seems little doubt that Kimberella was at least a triploblastic bilaterian animal.[139] Others. such as the Acanthodians. such as similarities to living organisms. mainly because the diagnostic features which allow taxonomists to classify more recent organisms. Halkieria and Microdictyon. a lagerstätte in China.[146] Other deuterostome groups are soft-bodied. respiration and gas exchange systems have to change. reproductive systems cannot depend on water to carry eggs and sperm towards each other.Evolutionary history of life The Ediacara biota. in other words significantly more complex than cnidarians. were eventually identified when more complete specimens were found in Cambrian lagerstätten that preserved soft-bodied animals. Cloudina. the echinoderms.[157] [158] Although the earliest good evidence of land plants and animals dates back to the Ordovician period (488 to 444 [159] million years ago). Many were flat and had a "quilted" appearance. At the time these were interpreted as evidence that the modern phyla had evolved very rapidly in the "Cambrian explosion" and that the Burgess Shale's "weird wonders" showed that the Early Cambrian was a uniquely experimental period of animal Opabinia made the largest single evolution. shows signs of successful defense against predation and may indicate the start of an evolutionary arms race. modern land ecosystems only appeared in the late Devonian. while the owners of some "armor plates". been interpreted as early molluscs (Kimberella[140] [141] ).[155] Vertebrates with jaws. are fairly common from the Early Cambrian small shelly fauna onwards.[143] Parvancorina[144] ). one of the two main groups of complex animals. and arthropods (Spriggina.[148] Later discoveries of similar animals and the development of contribution to modern interest in the [147] Cambrian explosion.[156] 63 Colonization of land Adaptation to life on land is a major challenge: all land organisms need to avoid drying-out and all those above microscopic size have to resist gravity. echinoderms (Arkarua[142] ). The earliest. new theoretical approaches led to the conclusion that many of the "weird wonders" were evolutionary "aunts" or "cousins" of modern groups[149] – for example that Opabinia was a member of the lobopods. There is still debate about the classification of these specimens.[150] Nevertheless there is still much debate about whether the Cambrian explosion was really explosive and. Vendozoa. which may have been slightly mineralized. and most of the significant Cambrian deuterostome Acanthodians were among the earliest vertebrates with [152] jaws fossils come from the Chengjiang fauna.[151] Most of the animals at the heart of the Cambrian explosion debate are protostomes.[145] The small shelly fauna are a very mixed collection of fossils found between the Late Ediacaran and Mid Cambrian periods.[161] .[146] In the 1970s there was already a debate about whether the emergence of the modern phyla was "explosive" or gradual but hidden by the shortage of Pre-Cambrian animal fossils. a group which includes the ancestors of the arthropods. which did not fit into any known phylum.[153] The Chengjiang fossils Haikouichthys and Myllokunmingia appear to be true vertebrates. and seemed so strange that there was a proposal to classify them as a separate kingdom. a vascular plant from the Silurian. did not exist.[161] Lichens. which are not plants but use the same photosynthesis mechanisms. trees such as Archaeopteris were so abundant that they changed river systems from mostly braided to mostly meandering.[165] By the Late Devonian 370 [168] million years ago. Water and any nutrients in it would have drained away very quickly. which are symbiotic combinations of a fungus (almost always an ascomycete) and one or more photosynthesizers Lichens growing on concrete (green algae or cyanobacteria).[169] In fact they caused a "Late Devonian wood crisis". and are attributed to annelids ("worms") or arthropods. the group that includes trees. Mat-forming cyanobacteria could have gradually evolved resistance to desiccation as they spread from the seas to tidal zones and then to land. This suggests that microbial mats may have been the first organisms to colonize dry land.[161] Soil formation would have been very slow until the appearance of burrowing animals.[171] In later ecosystems the carbon Reconstruction of Cooksonia.[161] Burrows have been found in Ordovician sediments. roots were required in order to extract water from the ground. because their roots bound the soil firmly. In Mid Silurian rocks 430 [167] million years ago there are fossils of actual plants including clubmosses such as Baragwanathia. have been found in Mid Ordovician rocks dated to about 476 [166] million years ago.[162] The earliest known ascomycete fossils date from 423 to 419 [163] million years ago in the Silurian.[161] and their ability to break down rocks contributes to soil formation in situations where plants cannot survive. possibly in the Precambrian.9 in) high.[157] [165] Spores of land plants. almost all cells are capable of photosynthesies and are nearly independent. and some appear closely related to vascular plants. a combination of mineral particles and decomposed organic matter. the parts in between became supports and transport systems for water and nutrients. possibly rather like liverworts.[161] Films of cyanobacteria. Life on land required plants to become internally more complex and specialized: photosynthesis was most efficient at the top. soil. . reducing the greenhouse effect and thus causing an ice age in the Carboniferous period. most were under 10 centimetres (3. have been found in modern deserts.Evolutionary history of life 64 Evolution of soil Before the colonization of land.[162] are also important colonizers of lifeless environments.[161] [164] Plants and the Late Devonian wood crisis In aquatic algae. which mix the mineral and organic components of soil and whose feces are a major source of the organic components. and only in areas that are unsuitable for vascular plants. Land surfaces would have been either bare rock or unstable sand produced by weathering.[170] because: • They removed more carbon dioxide from the atmosphere. [179] The fossil record of other major invertebrate groups on land is poor: none at all for non-parasitic flatworms. because their existing jointed exoskeletons provided protection against desiccation.[174] which are thought to be evolutionary "aunts" of myriapods. and most land animals developed internal fertilization of their eggs. about 415 [177] million years ago. On the other hand in some ways movement and breathing became easier.[172] • The increasing depth of plants' roots led to more washing of nutrients into rivers and seas by rain. The difference in refractive index between water and air required changes in their eyes.[158] Some trace fossils from the Cambrian-Ordovician boundary about 490.Evolutionary history of life 65 dioxide "locked up" in wood is returned to the atmosphere by decomposition of dead wood. the earliest fossils of gastropods on land date from the Late Carboniferous.[171] Fossilized trees from the Mid-Devonian Gilboa fossil forest. This caused algal blooms whose high consumption of oxygen caused anoxic events in deeper waters. but it is thought that insects developed the ability to fly in the Early Carboniferous or even Late Devonian. but they may still have been aquatic animals. and there is clear evidence of numerous arthropods on coasts and alluvial plains shortly before the Silurian-Devonian boundary. Land invertebrates Animals had to change their feeding and excretory systems.[180] About 99% of modern insect species fly or are descendants of flying species. and this group may have had to wait until leaf litter became abundant enough to provide the moist conditions they need.[181] Land vertebrates Acanthostega changed views about the early evolution [182] of tetrapods .[175] Other trace fossils from the Late Ordovician a little over 445 [176] million years ago probably represent land invertebrates.[178] Arthropods were well pre-adapted to colonise land.0 [173] million years ago are interpreted as the tracks of large amphibious arthropods on coastal sand dunes. including signs that some arthropods ate plants. support against gravity and a means of locomotion that was not dependent on water. and a means of escape from predators and from unfavorable changes in the environment. nematodes or nemerteans. and the better transmission of high-frequency sounds in air encouraged the development of hearing. and may have been made by euthycarcinoids. However the earliest fossil evidence of fungi that can decompose wood also comes from the Late Devonian. some parasitic nematodes have been fossilized in amber. This gave them a wider range of ecological niches for feeding and breeding. annelid worm fossils are known from the Carboniferous.[158] The earliest confirmed fossils of flying insects date from the Late Carboniferous. increasing the extinction rate among deep-water animals. in ways that enabled it to raise its head to breathe air while its body remained submerged. Some of these look like early relatives of modern amphibians. the head is not joined to the shoulder girdle and it has a distinct neck. this would have attracted grazing invertebrates and small fish that preyed on them. possibly in order to go from a pool that was drying out to one that was deeper. However in 1987 nearly-complete fossils of Acanthostega from about 363 [186] million years ago showed that this Late Devonian transitional animal had legs and both lungs and gills. vertebrates with four limbs. the bones at the back of its skull are locked together.3 ft) long.[182] Later discoveries revealed earlier transitional forms between Acanthostega and completely fish-like animals.[182] The Devonian proliferation of land plants may help to explain why air-breathing would have been an advantage: leaves falling into streams and rivers would have encouraged the growth of aquatic vegetation. providing strong attachment points for muscles that raised its head.Evolutionary history of life 66 Osteolepiformes ("fish") Panderichthyidae Obruchevichthidae Acanthostega Ichthyostega "Fish" Tulerpeton Early amphibians Anthracosauria Amniotes Family tree of tetrapods[183] Tetrapods. while others are . air-breathing would have been necessary because these waters would have been short of oxygen. its ribs were too short to prevent its lungs from being squeezed flat by its weight. The current hypothesis is that Acanthostega. evolved from other rhipidistians over a relatively short timespan during the Late Devonian. Its skeleton differed from that of most fish. which was about 1 metre (3. most of which need to keep their skins moist and to lay their eggs in water. but could never have survived on land: its limbs and its wrist and ankle joints were too weak to bear its weight. its fish-like tail fin would have been damaged by dragging on the ground. including: its jaws show modifications that would have enabled it to gulp air. since warm water holds less dissolved oxygen than cooler marine water and since the decomposition of vegetation would have used some of the oxygen. between 370 [168] million years ago and 360 [184] million years ago.[187] Unfortunately there is then a gap of about 30 million years between the fossils of ancestral tetrapods and Mid Carboniferous fossils of vertebrates that look well-adapted for life on land.[185] From the 1950s to the early 1980s it was thought that tetrapods evolved from fish that had already acquired the ability to crawl on land. was a wholly aquatic predator that hunted in shallow water. they would have been attractive prey but the environment was unsuitable for the big marine predatory fish. whose water-proof skins and eggs enable them to live and breed far from water. birds and mammals Early synapsids (extinct) Extinct pelycosaurs Synapsids Pelycosaurs Therapsids Mammaliformes Extinct therapsids Extinct mammaliformes Mammals Anapsids.Evolutionary history of life accepted as early relatives of the amniotes.[183] 67 Dinosaurs. whether turtles belong here is debated [188] Captorhinidae and Protorothyrididae Araeoscelidia (extinct) Amniotes Squamata (lizards and snakes) Extinct archosaurs Crocodilians Pterosaurs (extinct) Sauropsids Diapsids Archosaurs Dinosaurs Theropods Extinct theropods Birds Sauropods (extinct) Ornithischians (extinct) . [189] [190] The synapsid pelycosaurs and their descendants the therapsids are the most common land vertebrates in the best-known Permian fossil beds.[208] However fossil teeth discovered in Madagascar indicate that true mammals existed at least 167 [209] million years ago. This apparent set-back may actually have promoted the evolution of mammals. probably evolved in the Late Carboniferous period. The earliest fossils of the two surviving amniote groups.[211] Mammals throughout the time of the dinosaurs had been restricted to a narrow range of taxa. However at the time these were all in temperate zones at middle latitudes.[199] but by the Mid Triassic archosaurs were the dominant land vertebrates.[197] During the slow recovery from this catastrophe. mainly nocturnal insectivores. between 330 [191] million years ago and 314 [192] million years ago. synapsids and sauropsids.[195] The Permian-Triassic extinction wiped out almost all land vertebrates.[201] During the Late Jurassic. drier environments nearer the Equator were dominated by sauropsids and amphibians.0 [19] million years ago.[205] By 195 [206] million years ago in the Early Jurassic there were animals that were very nearly mammals. predatory theropod dinosaurs. and became the dominant land vertebrates of the Jurassic and Cretaceous periods. birds evolved from small.[196] as well as the great majority of other life.[212] [213] with bats taking to the air within 13 million years.[215] 68 .[207] Unfortunately there is a gap in the fossil record throughout the Mid Jurassic.Evolutionary history of life Possible family tree of dinosaurs. between 229. between 199 [200] million years ago and 65 [24] million years ago. birds and mammals[189] [190] Amniotes. whose eggs can survive in dry environments.[214] and cetaceans to the sea within 15 million years. but increased rapidly in size and diversity after the extinction. bony tails from their dinosaur ancestors.[202] The first birds inherited teeth and long. Dinosaurs distinguished themselves from other archosaurs in the Late Triassic.[204] While the archosaurs and dinosaurs were becoming more dominant in the Triassic.[202] but some developed horny. the dinosaurs perished 65 [24] million years ago in the Cretaceous–Tertiary extinction along with many other groups of organisms.[198] a previously obscure sauropsid group became the most abundant and diverse terrestrial vertebrates: a few fossils of archosauriformes ("shaped like archosaurs") have been found in Late Permian rocks. toothless beaks by the very Late Jurassic[203] and short pygostyle tails by the Early Cretaceous. and there is evidence that hotter. the mammaliform successors of the therapsids could only survive as small. date from around 313 [193] million years ago.0 [194] million years ago and 251. estimated to be 30M years.[210] After dominating land vertebrate niches for about 150 million years. sizes and shapes. for example nocturnal life may have accelerated the development of endothermy ("warm-bloodedness") and hair or fur. [216] [217] and that their rise was associated with that of pollinating insects. Another possible family tree.[216] [217] Social insects The social insects are remarkable because the great majority of individuals in each colony are sterile. [217] The 250.[218] . although some eusocial lineages have diversified into several families.000 to 400. There is fossil evidence that flowering plants diversified rapidly in the Early Cretaceous. and it seems that eusociality has evolved independently only 12 times among arthropods. In fact there are very few eusocial insect species: only 15 out of approximately 2.600 living families of insects contain eusocial species. they form over 50% of the total mass of insects.Evolutionary history of life 69 Flowering plants Angiosperms (flowering plants) Gnetales (gymnosperm) Welwitschia (gymnosperm) Gymnosperms Ephedra (gymnosperm) Gymnosperms Gingko Cycads (gymnosperm) Bennettitales Bennettitales Angiosperms (flowering plants) Gnetales (gymnosperm) Conifers (gymnosperm) One possible family tree of flowering [216] plants.000 species of flowering plants outnumber all other ground plants combined. and are the dominant vegetation in most terrestrial ecosystems.[217] Among modern flowering plants Magnolias are thought to be close to the common ancestor of the group.[216] However paleontologists have not succeeded in identifying the earliest stages in the evolution of flowering plants. for example although ants and termites account for only about 2% of known insect species. This appears contrary to basic concepts of evolution such as natural selection and the selfish gene. Nevertheless social insects have been spectacularly successful. between 130 [28] million years ago and 90 [29] million years ago. Their ability to control a territory appears to be the foundation of their success. while the species to which they are assigned has only slight influence. apparently by Australopithecus garhi.000 years ago and. The Mazon Creek lagerstätten These termite mounds have survived a bush fire. from the Late Carboniferous. after colonies have established this security. which then migrated all over the world less than 200. eusociality evolves only in species that are under strong pressure from predators and competitors. and advanced social bees have been found in Late Cretaceous rocks but did not become abundant until the Mid Cenozoic. and indicate that insects had occupied their main modern ecological niches as herbivores. some gigantic by modern standards. Social termites and ants first appear in the Early Cretaceous. but there has been a fourfold increase in the last 3 million years. it is based on kin selection. In support of this explanation they cite the appearance of eusociality in bathyergid mole rats. if so. and were found near animal bones that bear scratches made by these tools. include about 200 species.[228] There is also debate about whether anatomically-modern humans had an intellectual.[227] There is a long-running debate about whether modern humans evolved all over the world simultaneously from existing advanced hominines or are descendants of a single small population in Africa.000 years ago and replaced previous hominine species. but there is no evidence of nepotism in colonies that have multiple queens. which has the paradoxical consequence that two sterile worker daughters of the same queen share more genes with each other than they would with their offspring if they could breed. they write. Instead. a statistical analysis suggests that hominine brain sizes depend almost completely on the date of the fossils.[223] Humans Modern humans evolved from a lineage of upright-walking apes that has been traced back over 6 [30] million years ago to Sahelanthropus. about 300 [222] million years ago.[219] However Wilson and Hölldobler argue that this explanation is faulty: for example.Evolutionary history of life 70 The sacrifice of breeding opportunities by most individuals has long been explained as a consequence of these species' unusual haplodiploid method of sex determination.[220] The earliest fossils of insects have been found in Early Devonian rocks from about 400 [221] million years ago.[218] which are not haplodiploid.5 [225] million years ago.[229] . cultural and technological "Great Leap Forward" under 100. detritivores and insectivores.[224] The first known stone tools were made about 2.[226] The earliest hominines had chimp-sized brains. which preserve only a few varieties of flightless insect. whether this was due to neurological changes that are not visible in fossils. but in environments where it is possible to build "fortresses". they gain other advantages though co-operative foraging. as reconstructed from the fossil record. Both of these phenomena could be explained in one or more ways:[233] • The oceans may have become more hospitable to life over the last 500 million years and less vulnerable to mass extinctions: dissolved oxygen became more widespread and penetrated to greater depths. So paleontologists have mistakenly assigned parts of the same organism to different genera which were often defined solely to accommodate these finds – the story of Anomalocaris is an example of this. and marine ecosystems became more diversified so that food chains were less likely to be disrupted.Evolutionary history of life 71 Mass extinctions K–T Tr–J P–Tr Late D O–S Millions of years ago Apparent extinction intensity. Many of the "superfluous" genera are represented by fragments which are not found again and the "superfluous" genera appear to become extinct very quickly. it is rarely because the new dominant group is "superior" to the old and usually because an extinction event eliminates the old dominant group and makes way for the new one.[233] . and complete fossils are rarest in the oldest rocks. the fraction of genera going extinct at any given time. mass extinctions have sometimes accelerated the evolution of life on earth.[234] [235] • Reasonably complete fossils are very rare. (Graph not meant to include recent epoch of Holocene extinction event) Life on earth has suffered occasional mass extinctions at least since 542 [230] million years ago. The risk of this mistake is higher for older fossils because these are often unlike parts of any living organism. the development of life on land reduced the run-off of nutrients and hence the risk of eutrophication and anoxic events. most extinct organisms are represented only by partial fossils.[231] [232] The fossil record appears to show that the gaps between mass extinctions are becoming longer and the average and background rates of extinction are decreasing. When dominance of particular ecological niches passes from one group of organisms to another. Although they are disasters at the time.e. i. and possibly at an accelerating rate over the last 40.Evolutionary history of life 72 All genera "Well-defined" genera Trend line "Big Five" mass extinctions Other mass extinctions Million years ago Thousands of genera Phanerozoic biodiversity as shown by the fossil record Biodiversity in the fossil record.[243] . in which the devastating Permian–Triassic extinction event is an important factor.000 years. a slight decline from 400 to 200 [238] million years ago. Well-known examples include the changes in coloration of the peppered moth over the last 200 years and the more recent appearance of pathogens that are resistant to antibiotics. those whose first occurrence predates and whose last occurrence postdates that time"[236] shows a different trend: a fairly swift rise from 542 to 400 [237] million years ago. and a swift rise from 200 [239] million years ago to the present. and phytoplankton are the dominant marine producers. which is "the number of distinct genera alive at any given time.[241] [242] There is even evidence that humans are still evolving. that is.[240] The processes that drive evolution are still operating.[236] The present Oxygenic photosynthesis accounts for virtually all of the production of organic matter from non-organic ingredients. Production is split about evenly between land and marine plants. Retrieved 2007-10-22.0195. org/ pdf_files/ TJB. ISBN 0-632-05614-2. nationalgeographic. com/ news/ 2007/ 06/ 070620-mammals-dinos.Y.1038/46965.1998. doi:10. php?Ma=1850 [7] Knoll. Hu. [24] http:/ / toolserver. Han. edu/ phyla/ metazoafr. doi:10. Gordon. PMC 1578724. Massachusetts: Sinuer Associates. ... and Cohen. T.. php?Ma=450 [12] "The oldest fossils reveal evolution of non-vascular plants by the middle to late Ordovician Period (~450-440 m. [23] "Amniota . php?Ma=65 [25] Fastovsky DE. S-X. Inc. . doi:10. doi:10.2517/prpsj. J. R. Retrieved 2009-03-08. Zhu. ISBN 0-87893-187-2. . Andrew H. 2007. org/ ~verisimilus/ Timeline/ Timeline. vend. gov/ articlerender.7. Retrieved 2008-09-02. 1130/ 1052-5173(2005)015<4:TEOTDI>2. Marshall Space Flight Center. fcgi?tool=pmcentrez& artid=1578724).CO. N.. Conway Morris. D. org/ web/ 20071025225841/ http:/ / nmnaturalhistory. [15] "Metazoa: Fossil Record" (http:/ / www. php?Ma=1200 [14] Algeo. berkeley. [26] "Dinosaur Extinction Spurred Rise of Modern Mammals" (http:/ / news.Evolutionary history of life 73 See also • • • • Evolution Evolutionary history of plants Timeline of evolution History of evolutionary thought Footnotes [1] http:/ / toolserver. edu/ users/ pciesiel/ gly3150/ plant. [3] Nisbet.1130/1052-5173(2005)015<4:TEOTDI>2. html). [27] Van Valkenburgh. Retrieved 2007-05-18.1098/rstb. pdf) on October 25. 2009. M. Donald F. H-L. org/ doi/ abs/ 10. [28] http:/ / toolserver. palaeos. Hewitt. Philosophical Transactions of the Royal Society of London. 1146/ annurev.0934. php?Ma=1700 [10] Bonner. NASA. (1998) The origins of multicellularity. html) [13] http:/ / toolserver. Douglas J. 1999). Sunderland.. S. gov/ headlines/ y2002/ 28jan_extinction.1140325. paleo. [6] http:/ / toolserver. Nature 402: 42–46. org/ ~verisimilus/ Timeline/ Timeline. and Fowler. [21] Tanner LH. Science and Technology Directorate. csupomona.) 317 (5846): 1903–1906. PMC 1690475.. . "Archaean metabolic evolution of microbial mats" (http:/ / www. (1997). Zhang. Sheehan PM (2005). "The origin of the Metazoa in the light of the Proterozoic fossil record" (http:/ / www.1126/science...a. php?Ma=3 . et al. fcgi?tool=pmcentrez& artid=1690475). . . B. . pdf) (PDF). org/ pdf_files/ TJB. E. B.T. PMID 17901330. et al. Patrick L. L.earth. Integr. Arnold. "Major patterns in the history of carnivorous mammals" (http:/ / arjournals. [8] Fedonkin. pdf) (PDF).J. org/ ~verisimilus/ Timeline/ Timeline. nih.1999. gov/ articlerender. Retrieved March 26.1. 1. "The extinction of the dinosaurs in North America" (http:/ / www. Earth-Science Reviews 65 (1-2): 103–139. org/ ~verisimilus/ Timeline/ Timeline. . Vertebrate Paleontology. org/ ~verisimilus/ Timeline/ Timeline. [18] Hoyt. Kendall. (November 4. J. 1. pubmedcentral. Kaufman.. clas. Blackwell Publishers.2. E. earth.) on the basis of fossil spores" Transition of plants to land (http:/ / www. . annualreviews.9. "Assessing the record and causes of Late Triassic extinctions" (http:/ / web.. M.G. Philosophical Transactions of the Royal Society B: Biological Sciences 353 (1365): 113–130. ucmp. Proceedings of the Royal Society: Biology 266 (1436): 2375. gsajournals. (2005). (March 2003). php?Ma=251 [20] Barry.0. archive.. A. News. xii-452.2). doi:10. "Eukaryotic organisms in Proterozoic oceans" (http:/ / www. php?Ma=525 [17] Shu et al. org/ ~verisimilus/ Timeline/ Timeline. CO.1098/rstb.. PMID 16754612. and marine anoxic events". P. . (December 7. GSA Today 15 (3): 4–10.1843. Lucas SG & Chapman MG (2004).Palaeos" (http:/ / www. [9] http:/ / toolserver. (1999). doi:10.com. 27. weathering processes. "Lower Cambrian vertebrates from south China". [22] Benton.463. doi:10.M. org/ perlserv/ ?request=get-document& doi=10. A. "Synapsid Reptiles" (http:/ / www. (2006). Annual Review of Earth and Planetary Sciences 26: 463–493.nationalgeographic.1098/rspb. org/ ~verisimilus/ Timeline/ Timeline.
[email protected]. htm). Javaux. A. nasa. E.. Luo. 27–36 [11] http:/ / toolserver. [16] http:/ / toolserver. Chen.y. org/ ~verisimilus/ Timeline/ Timeline. Paleontological Research 7 (1): 9–41. "The Great Dying" (http:/ / science. (1998). Science (New York. (2007). ru/ pub/ Fedonkin_2003. Archived from the original (http:/ / nmnaturalhistory.R. Scheckler.. nih. Creaser.abstract with link to free full content (PDF) [4] http:/ / toolserver. "A whiff of oxygen before the great oxidation event?". org/ ~verisimilus/ Timeline/ Timeline. html). Part B 361 (1470): 1023–38. php?Ma=3500 [2] Futuyma. 2002). 1999). doi:10. (January 28. pubmedcentral. 463). org/ ~verisimilus/ Timeline/ Timeline. (2004). Duan. php?Ma=2400 [5] Anbar.1016/S0012-8252(03)00082-5. edu/ ~dfhoyt/ classes/ zoo138/ SYNAPSID.J. org/ Amniota). [19] http:/ / toolserver.27. HTML). php?Ma=6 [31] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. doi:10. 0. G. php?Ma=90 [30] http:/ / toolserver. php?Ma=130 [29] http:/ / toolserver. G. org/ ~verisimilus/ Timeline/ Timeline.1146/annurev. T. "Terrestrial-marine teleconnections in the Devonian: links between the evolution of land plants. org/ ~verisimilus/ Timeline/ Timeline.. M. Lyons. Biol. C.. S. ufl. Y. X-L. Evolution. org/ cgi/ content/ abstract/ 190/ 1/ 205). html). King. org/ ~verisimilus/ Timeline/ Timeline. Akilia. uci. (January 2005). [49] http:/ / toolserver. K.H.1144/GSL. A. • Dalrymple. A. and Rothman. doi:10.04. Space.1126/science. Icarus 148 (2): 508–512. "Archean microfossils: a reappraisal of early life on Earth". and Fedo. "Metasomatic Origin of Quartz-Pyroxene Rock. (December 2000). pdf) (PDF). Special Publications 190: 205–221.2. Geological Society. Retrieved 2008-08-30.2003. S. mcmaster. "The origin of life on the earth" (http:/ / courses. edu/ news/ releases/ 2005/ 438. .R. S. [46] Mojzsis. S. M. [52] Orgel. Earth and Planetary Science Letters 235 (3-4): 663–681. • Lepland. PMID 11099411. (2006). doi:10.com. html). (1991). "A fresh look at the fossil evidence for early Archaean cellular life" (http:/ / physwww. geology. edu/ zircon/ Valley2002Cool_Early_Earth. G. "Origin of the Earth-Moon System". and the Edinburgh Ion Microprobe Facility (July 15. pubmedcentral.M. php?Ma=3800 [36] Cohen. org/ ~verisimilus/ Timeline/ Timeline.. (December 2000).L. PMC 1578727.1098/rstb. W.1006/icar.M.1038/scientificamerican1094-76.J.5108. doi:10. Retrieved 2008-08-30. doi:10. php?Ma=3000 [44] Brasier. html).. C. org/ cgi/ content/ abstract/ 290/ 5497/ 1754). (2003).resmic.0. and Krivtsov. • Altermann.A.. and Wacey.. J. and Wilde. Retrieved 2008-09-13.E. colorado.W. (May 2002). php?Ma=4540 Dalrymple. [47] Grotzinger..+1038/383423a0. (2000). "A cool early Earth" (http:/ / www. [51] Mason.1835. washington. doi:10. 2005). PMID 12029129. Schopf.J. (July 2007). T.006.R. com/ nature/ journal/ v384/ n6604/ abs/ 384055a0. N.. Retrieved 2008-08-30. A. php?Ma=4000–3800 [40] Britt. USGS. com/ science?_ob=ArticleURL& _udi=B6V61-4GDKB05-3& _coverDate=07/ 15/ 2005& _alid=382434001& _rdoc=1& _fmt=& _orig=search& _qd=1& _cdi=5801& _sort=d& view=c& _acct=C000050221& _version=1& _urlVersion=0& _userid=10& md5=be47e49c535d059be188b66c6e596dd5). E.M. Harrison. Green. B.1126/science.J.. Nature 384 (6604): 55–59. "Origins of biomolecular handedness".290. 2005-11-17. 6. Earth Syst. . and Marty. London. pdf) (PDF).).2006. .M.6489. PMID 11539831. lyellcollection.1130/0091-7613(2002)030<0351:ACEE>2.1754. doi:10. G. nih. and Whitehouse. D. PMID 6472461. Valley.. PMID 7524147. • • 74 [39] http:/ / toolserver. and Kazmierczak. (April 1993).CO. (June 2006). Newman. wisc.. p. doi:10. ca/ ~higgsp/ 3D03/ BrasierArchaeanFossils. Science 290 (5497): 1754–1756. [43] http:/ / toolserver. PMID 14596897.5497. F. "The age of the Earth in the twentieth century: a problem (mostly) solved" (http:/ / sp. Geology 30 (4): 351–354. "Questioning the evidence for Earth's earliest life — Akilia revisited" (http:/ / geology. edu/ biol354/ The Origin of Life on Earth.1016/j.M. sciencemag.B.1007/BF02715942. J.2001. M.epsl. Scientific American 271 (4): 76–83. PMID 16754604. and Friend.. org/ cgi/ content/ abstract/ 260/ 5108/ 640).1130/G20890. • • . G.A. "Fossil evidence of Archaean life" (http:/ / www. O. A. Blackwell Science. doi:10. Nutman. University of Colorado.028. . Fedo. M. W. Also available as a web page (http:/ / proxy. pdf) (PDF). php?Ma=3400 [50] Schopf. (December 2005). Peck. space. org/ cgi/ content/ abstract/ 296/ 5572/ 1448). History of Life (3rd ed.14.. doi:10. Retrieved 2008-08-31. Greenland. arts. PMC 1578735. fcgi?tool=pmcentrez& artid=1578735).J. Science 260 (5108): 640–646. C.. nature. A.Evolutionary history of life [32] http:/ / toolserver. com/ scienceastronomy/ planetearth/ earth_bombarded_020724. .1016/j. (1984). C. J. org/ ~verisimilus/ Timeline/ Timeline. S.1070336. R. T. . D. Res Microbiol 154 (9): 611–7. Publications Services. and Kring. . J.A. The Age of the Earth. Retrieved 2007-09-20. "Support for the Lunar Cataclysm Hypothesis from Lunar Meteorite Impact Melt Ages" (http:/ / www. ias. Retrieved 2008-08-29. usgs. . PMID 8900275. L. "An abiotic model for stomatolite morphogenesis".B. J. . . "Evidence for life on Earth before 3.D. N. gov/ articlerender.M.800 million years ago" (http:/ / www. org/ cgi/ content/ abstract/ 33/ 1/ 77). html) [53] Cowen. Swindle.260.2005. Philosophical Transactions of the Royal Society: Biology 361 (1470): 887–902. (October 1994). (November 1996).W. California: Stanford University Press. ISBN 0632044446. doi:10. org/ ~verisimilus/ Timeline/ Timeline. Whitehouse..1038/384055a0. Science 296 (5572): 1448–1452. [37] http:/ / toolserver. "The Late Asteroidal and Cometary Bombardment of Earth as Recorded in Water Deuterium to Protium Ratio". edu/ ~nideffer/ Hawking/ early_proto/ orgel. McKeegan. [41] Valley. org/ ~verisimilus/ Timeline/ Timeline. doi:10.1038/311019a0. doi:10.1834. geoscienceworld. and Implications for Earth's Earliest Life" (http:/ / www.P. ac. ISBN 0-8047-1569-6. Nature 383: 423–425. [34] Galimov.D. Retrieved 2009-01-11. McLoughlin.640. (2001). sciencemag. (http:/ / www. html). sciencedirect. J. Arrhenius. Wilde. .1.1098/rstb.F.A.190. "Age of the Earth" (http:/ / pubs. D. doi:10. doi:10. . E. org/ ~verisimilus/ Timeline/ Timeline.. Robert. Arrhenius. Retrieved 2008-08-30. (April 2002). Cavosie. G. Retrieved 2008-08-30.01. "Microfossils of the Early Archean Apex Chert: New Evidence of the Antiquity of Life" (http:/ / www. PMID 16754605. gov/ gip/ geotime/ age. (1996). Sci.2006. sciencemag.2000. W. Retrieved 2008-08-30. in/ jessci/ dec2005/ ilc-3.08. doi:10. (2002-07-24).SP.P. "Magmatic δ18O in 4400-3900 Ma detrital zircons: A record of the alteration and recycling of crust in the Early Archean" (http:/ / www. 114 (6): 593–600. R.. Philosophical Transactions of the Royal Society of London: B Biological Sciences 361 (1470): 869–85. Geology 33 (1): 77–79. Retrieved 2006-04-15. B.L. M. doi:10. [42] Dauphas. J. van Zuilen. . php?Ma=4400–4000 • • "Early Earth Likely Had Continents And Was Habitable" (http:/ / www.H. W. pdf) [35] http:/ / toolserver. Nature 311 (5981): 19–23.1126/science. "Evidence for Ancient Bombardment of Earth" (http:/ / www. ISBN 0198557043. A simpler nucleic acid". cfm?articleID=00073A97-5745-1359-94FF83414B7F0000& pageNumber=1& catID=2). .5481. doi:10. Deamer.. pnas. Scientific American: 64–71. (1992).1183. N. The Quest for Extraterrestrial Life. (2004). M. F. com. Retrieved 2008-09-01.tb00592. doi:10. Natl. PMID 11296516. and Lancet. doi:10. 8 (1): 23–31. and Miller.7933. [69] Cech. "The ribosome is a ribozyme" (http:/ / www. Acad. PMC 18108. T. and Wickramasinghe. W. doi:10. PMID 11358999.1038/418214a. org/ cgi/ pmidlookup?view=long& pmid=7667262). . [75] Wächtershäuser. "From self-assembly of life to present-day bacteria: a possible role for nanocells". [78] Cairns-Smith. htm). • Larralde. Nature 418 (6894): 214–21. Elie. still open questions".. S. "Evolution without speciation but with selection: LUCA. pp. U. pdf) (PDF). E. com/ nature/ journal/ v433/ n7025/ full/ 433469a. G. D. Astrophysics and Space Science 66: 77–90.1073/pnas. 25 (5): 573–82. D. Proc. iUniverse. sciam. 92 (18): 8158–60.2004. R. B. Die Umschau volume=7. and Psenner. "On the Nature of Interstellar Grains". PMC 41115. and Labedan.1111/j.J. D. Robertson. [56] Hoyle. Retrieved 2008-08-30. PMID 15011140. "Controversies on the origin of life" (http:/ / www. (July 1998).1016/0019-1035(73)90110-3. pdf)) [67] Trevors. Irwin. PMID 15690023. J. ed. Philosophical Transactions of the Royal Society: Biological 358 (1429): 59–85. D. Towards a Theoretical Biology. fcgi?tool=pmcentrez& artid=1693102). (2006). 57–66 • 75 . Int. (November 2005). H.1038/362709a0. FEMS Microbiol. Sci. funpecrp.cellbi... PMID 15563395. "Life: In search of the simplest cell" (http:/ / www. PMID 10979855. Levy. M. org/ cgi/ content/ short/ 289/ 5481/ 878). "Origin of life.A. PMID 10760258. [76] Trevors. Proc. and Weiss. In Waddington. (2008). F (1973). Benachenhou-Lahfa. the Last Universal Common Ancestor in Gilbert’s RNA world" (http:/ / www. (1968). J. and Abel.1007/s00114-005-0078-6. F. . M.L. L.1098/rstb. PMID 7667262. doi:10. Retrieved 2008-09-01. Duguet. com/ article. (December 2003)). and Russell. .006..14. U.1007/s00114-005-0056-z. [62] Peretó. L.1126/science. "Rates of decomposition of ribose and other sugars: implications for chemical evolution" (http:/ / www. 1. space. P.92. "Claim of Martian Life Called 'Bogus'" (http:/ / www. ISBN 0691135495. ac. J. (August 2000). doi:10. . doi:10. L. Ben-Eli.R.S. PMID 16292523. Origins of Life and Evolution of Biospheres 2001 31 (1-2): 119–45. S. Science (journal) 290 (5495): 1306–7.1016/0303-2647(92)90004-I. "The antiquity of RNA-based evolution". pp. . Sci. Rev. U.x. and Miller. "The Propagation of Life in Space". K.1060786. doi:10.A. (November 2000). (April 2000). [59] Ker. Than (August 2007). br/ gmr/ year2003/ vol4-2/ gmr0070_full_text. com/ ?id=lEQKnip7St4C& pg=PA84& dq=life+ earth+ carbon+ water). . S.A. doi:10. html).1126/science. PMID 12594918. (August 1995). doi:10. J. [55] Arrhenius.. [60] Bennett. PMID 10960319. Science 289 (5481): 878–9. nature. and from prokaryotes to nucleated cells" (http:/ / www. pnas.2002.95. pdf) (PDF). C. Levy. (February 2005). Natl.97. Life as we don't know it". Reprinted in Goldsmith. "Peptide nucleic acids rather than RNA may have been the first genetic molecule" (http:/ / www. Sci.1007/BF00648361. (April 2006). G. PMID 16525788.F. Retrieved 2009-01-11. PMID 15906258. Retrieved 2008-09-02. T. Edinburgh University Press. and Stano. br/ gmr/ year2003/ vol4-2/ / pdf/ gmr0070.. Nature 362 (6422): 709–15. D.8158. N.H. Retrieved 2008-09-01.. org/ cgi/ pmidlookup?view=long& pmid=9653118). Proc. (August 2000). D. 28 (11): 729–39.8. doi:10.. weizmann. T. K. (2003). S. M. [68] Forterre.S. • Lindahl. google.1023/A:1006746807104. ISBN 0595495966.289.Evolutionary history of life [54] O'Leary.E.. doi:10. (1903). B. pnas. (February–April 2001). [65] Joyce. Nature 433 (7025): 469–470. "Directed Panspermia". Microbiol. "Instability and decay of the primary structure of DNA". L. (April 1993). D. gov/ articlerender. 97 (8): 3868–71.com. Genetic and Molecular Research 2 (4): 366–375. space. Beyond UFOs: The Search for Extraterrestrial Life and Its Astonishing Implications for Our Future.L. PMID 12110897. [70] Johnston.. [77] Segré. [72] Orgel.06. Retrieved 2007-10-07. "The stability of the RNA bases: implications for the origin of life" (http:/ / www. D. [61] Schulze-Makuch. (2002). University Science Books. PMID 1337989. .1574-6976. Confalonieri. (2008). doi:10. org/ cgi/ pmidlookup?view=long& pmid=10760258). R. [74] Martin.G. C. nih.2001. doi:10. Science 292 (5520): 1319–1325.. Anaxagoras and the Origin of Panspermia Theory. Acad. Naturwissenschaften 93 (4): 155–72. (2005). [64] Luisi.1016/j. O.878. (2001). M. Inc. pubmedcentral. [57] Crick.(also available as PDF (http:/ / www. . "RNA-Catalyzed RNA Polymerization: Accurate and General RNA-Templated Primer Extension". P. Icarus 19: 341–348. doi:10.3868. "Approaches to semi-synthetic minimal cells: a review". F. BioSystems 28 (1-3): 15–32.18. sciencemag. microbios. "Did Life Come From Another World?" (http:/ / www. Natl. [73] Nelson. im. "The prospect of alien life in exotic forms on other worlds". html). A. il/ Segre_Lipid_World. "An approach to a blueprint for a primitive organism". PMC 1693102. M. Int. W. com/ news/ 070823_mars_life.T. [63] Szathmáry. Science (journal) 289 (5483): 1307–8.1073/pnas. and Miller. Ferri. PMID 8469282. funpecrp. [58] Warmflash. P. "On the origins of cells: a hypothesis for the evolutionary transitions from abiotic geochemistry to chemoautotrophic prokaryotes. PMID 11185405. doi:10.S. L. et al (2001). Naturwissenschaften 93 (1): 1–13. Cell Biol. Princeton University Press.R. org/ 0801/ 0801023. "What is life?" (http:/ / books. . P. doi:10. "The Lipid World" (http:/ / ool. [66] Hoenigsberg.1073/pnas.1038/433469a. Retrieved 2008-09-02.. (1979). com.. 82–85. Acad. PMID 11742692.. "Origin of life. PMC 20907. . 95 (14): 7933–8. PMID 9653118. "Chance and necessity do not explain the origin of life". "The nature of the last universal ancestor and the root of the tree of life. C. .1038/35085554. . R. Logan. Retrieved 2008-07-14. pnas.. "A molecular timescale of eukaryote evolution and the rise of complex multicellular life" (http:/ / www. [93] Hedges. (June 1999). sciencedirect. (December 2006). the difference between eukaryotic and prokaryotic cells" (http:/ / www.. pdf) (PDF). L (January 2004). com/ science?_ob=ArticleURL& _udi=B6VJ1-429XTFM-H& _user=10& _rdoc=1& _fmt=& _orig=search& _sort=d& view=c& _version=1& _urlVersion=0& _userid=10& md5=8370ca16bcde45bfa1c050068a2d6e19). 1–28. "Molecular evidence for the evolution of photosynthesis" (http:/ / www. Paterson.. "Toxic oxygen: The radical life-giver" (http:/ / www.. Archived from the original (http:/ / trc. (2003). ISBN 0716712563. A. Vol. (1999). and Summons. [80] Hanczyc. . . Retrieved 2008-07-14. Kamber. L..10173. N. fcgi?tool=pmcentrez& artid=1713255). E. A. PLoS ONE 2 (8): e790.1038/35085554.. org/ cgi/ content/ abstract/ 302/ 5645/ 618). (2001).0000790.1371/journal. [86] Blankenship. net/ abstracts/ EGU06/ 00770/ EGU06-J-00770. pp. R. Kluwer Academic. San Francisco: W. G.E. Retrieved 2008-07-14. (7 November 2002). PMID 10446042. J. [97] Vellai. gov/ articlerender. and Zavarzin. Global Mechanism. doi:10. PMID 10467746. Retrieved 2008-07-14.M. D. Geophysical Research Abstracts 8 (00770). (PDF). and Godelle. Gerdes. doi:10. 106. Fujikawa. G. Å.pgen. doi:10. gov/ articlerender. . R. and Palinska. Bebout.J.M. doi:10. 1994).1126/science. A. Levit. com/ content/ t1n325268n01217k/ ). 3) 196 (3): 311–314.H. 33/ krumbein/ htdocs/ Archive/ 397/ Krumbein_397. J. Retrieved 2008-07-14. sciencemag. Des Marais. K. BMC Evolutionary Biology 4 (2): 2. 33/ krumbein/ htdocs/ Archive/ 397/ Krumbein_397. L. PMID 7937858.J. W. "Stromatolite reef from the Early Archaean era of Australia" (http:/ / www.1038/420027a. [92] Brocks. (1981). "The Last Universal Common Ancestor: emergence. University of California. Evolution: A Molecular Point of View (Biological Bulletin.91. html) on October 29.J. M-A. J. . doi:10. nih. Trends in Plant Science 6 (1): 4–6. Venturi. PMID 12422197. and Labedan. Retrieved 2008-09-01. Growth. Xu. html). M. nih. edu/ biosci10v/ bis10v/ week3/ 06aerobicrespirintro. "Biofilm. 76 .. Biomat Microbialites.. pdf). D. . Brehm. Fossil and Recent Biofilms: A Natural History of Life on Earth. "Archaean molecular fossils and the rise of eukaryotes" (http:/ / www. PLoS Genetics 2 (12): e220. A.1016/S0169-5347(00)02084-X. I. org/ web/ 20070106201614/ http:/ / 134.. PMID 10390828. Proceedings of the National Academy of Sciences 91 (21): 10173–10177. [94] Burki. C.1371/journal. (October 11. I. G. S. fcgi?tool=pmcentrez& artid=1949142). S. PMC 44980. W. Jack W.1186/1745-6150-3-29. B. "Reducing the genome size of organelles favours gene transfer to the nucleus" (http:/ / www. Retrieved 2008-09-02. Origins of Life and Evolution of Biospheres 28 (4–6): 512–521. "Evaluating Support for the Current Classification of Eukaryotic Diversity" (http:/ / www. gov/ articlerender.A. "Photosynthesis and the Origin of Life" (http:/ / www. [89] "Introduction to Aerobic Respiration" (http:/ / web.. . M.2307/1542957. org/ web/ 20071029120120/ http:/ / trc. springerlink. Retrieved 2009-01-10. M. archive. Abert. jstor. B.. Biology Direct 3 (29): 29. nih.. 242. (1999). and Division" (http:/ / www. PMC 1713255. Biodictyon. Lenton. Retrieved 2008-09-01. pdf) on January 6. G.. PMID 11460161. [84] (the editor) (June 2006)). 2007. Parahistology" (http:/ / web. (2006). com/ nature/ journal/ v420/ n6911/ full/ 420027a. .1126/science. cosis. Skjæveland. . com/ 1471-2148/ 4/ 2/ abstract/ ). [87] Hoehler. pubmedcentral. nature. B. Patterson. D. Trends in ecology & evolution 16 (3): 135–141. Retrieved 2008-07-09 [83] Risatti. PMC 2478661. Lasser. and Stahl. T. S.M. T. doi:10. et al.1089904. D. PMID 16760969. PMID 15005799. Shalchian-Tabrizi.4 Ga as a bistability in atmospheric oxygen due to UV shielding by ozone" (http:/ / www. [88] Abele.1073/pnas.A. pubmedcentral. nature. BM. D. R. Archived from the original (http:/ / 134. Buick. doi:10. PMID 11460161. and Katz.21. F. and Szostak. "The Great Oxidation at ~2. Proceedings of the Royal Society: Biology 266 (1428): 1571–1577.. 242..1033. Gorbushina. (October 1998). Retrieved 2008-08-31. nature.M. sciencemag. org/ pss/ 1542957). Biological Bulletin. pdf) (PDF). . Davis.A.1038/nature04764. fcgi?tool=pmcentrez& artid=2478661). doi:10. Bebout. Nikolaev. (June 2006)). "Phylogenomics Reshuffles the Eukaryotic Supergroups" (http:/ / www. html). Science 302 (5645): 618–622. Capman. In Krumbein. U. fcgi?tool=pmcentrez& artid=1690172). "Experimental Models of Primitive Cellular Compartments: Encapsulation. E. [95] Parfrey. and Shoe. .E. com/ nature/ journal/ v412/ n6844/ full/ 412324a0. doi:10. ucdavis. 106. org/ cgi/ content/ abstract/ 285/ 5430/ 1033). "The origin of eukaryotes. html). E. Nature 420 (27): 27. [82] Krumbein.0020220. J. No. Retrieved 2008-09-02. doi:10. 196. Marshall. Bhattacharya. Blair. gov/ articlerender.pone. Barbero. D. P.. html). [91] Glansdorff. "Editor's Summary: Biodiversity rocks" (http:/ / www. Dunthorn. E. doi:10. Symbiosis in cell evolution.M. Geophysiology.. B. DJ (1 January 2001).. Retrieved 2008-09-01. K. A. "Prebiotic Synthesis on Minerals: Bridging the Prebiotic and RNA Worlds" (http:/ / www.1186/1471-2148-4-2. pubmedcentral. (2008). PMC 341452. T.. .P.1999. Oolites. B.M. . [81] Hartman.285. edu/ biosci10v/ bis10v/ week3/ 06aerobicrespirintro.. (October 2003). and Vida. L. doi:10..A. W. H. org/ content/ 91/ 21/ 10173. com/ nature/ journal/ v441/ n7094/ abs/ nature04764. Retrieved 2008-07-09. [90] Goldblatt. PMID 18613974.. L. Walter. PMC 1949142. C. nature. biomedcentral. C. W. PMID 17726520. constitution and genetic legacy of an elusive forerunner" (http:/ / www. Stromatolites. B.1098/rspb. B. W. Freeman. J. sciencedirect. and Burch. S. "Community structure of a microbial mat: the phylogenetic dimension" (http:/ / www. doi:10.5430. M. (19 July 2001). Nature 441. [85] Allwood..Evolutionary history of life [79] Ferris. ucdavis.0817.. Retrieved 2008-09-01.. Nature 441 (7094): 714–718. Minge. .. com/ nature/ journal/ v441/ n7094/ edsumm/ e060608-01. [98] Selosse. ISBN 1402015976. PMC 1690172. J. archive. pubmedcentral. html). full. and Des Marais.. M. [96] Margulis. doi:10. (2007). PMID 14576428.. Y. 2007. Nature 412 (6844): 324–327. com/ science?_ob=ArticleURL& _udi=B6TD1-424KK4J-3& _user=10& _rdoc=1& _fmt=& _orig=search& _sort=d& view=c& _acct=C000050221& _version=1& _urlVersion=0& _userid=10& md5=6f38f9f1d29b24fc90d0145837338b9e). "The role of microbial mats in the production of reduced gases on the early Earth" (http:/ / www... C. nih. and Watson.E. Science 285 (5430): 1033–1036. G. PMID 17194223. M.. gov/ books/ bv. PMID 1631544.R.1016/S0022-5193(84)80178-2. "Supertrees disentangle the chimerical origin of eukaryotic genomes". geoscienceworld.F. php?Ma=2100 [105] Han. T.G.090145. org/ cgi/ content/ abstract/ 26/ 3/ 386).5407. org/ cgi/ content/ abstract/ 257/ 5067/ 232). [116] http:/ / toolserver.. (April 2001). PMID 12151256. [104] http:/ / toolserver. (October 2008).K. ISBN 0-8050-6331-5. "Why have sex? The population genetics of sex and recombination" (http:/ / www. (September 2004). Molecular Microbiology 40 (22): 294–305. 1002/ (SICI)1520-6602(1998)1:1<27::AID-INBI4>3.tb01500. [124] Hickey. Nature 407 (6803): 470. Encyclopedia of Life Sciences. bioinfo. S-B.M. B. John Wiley & Sons. pp. J. evidence for sex in Giardia and an early eukaryotic origin of meiosis" (http:/ / euplotes. "Sex: Advantage".. I.. org/ ~verisimilus/ Timeline/ Timeline. [120] Ramesh. [100] Gray. com/ nature/ journal/ v407/ n6803/ abs/ 407470a0. doi:10. doi:10. org/ cgi/ content/ abstract/ 31/ 1/ 165).1038/npg. "A phylogenomic inventory of meiotic genes. G. Paleobiology 31 (1): 165–182. nih. W. fcgi?highlight=conjugation& rid=mmed. Malik. [106] http:/ / toolserver. D.02302.. J. doi:10. biochemsoctrans. "Megascopic eukaryotic algae from the 2.2005. sp. "Genetics: Exchange of Genetic Information" (http:/ / www. J.O. G. Burger. doi:10. PMID 11700279. doi:10.0001716 [118] Holmes. R. PMID 15668177. doi:10. (2006). Dr.. E. doi:10. org/ ~verisimilus/ Timeline/ Timeline. [101] http:/ / toolserver. (2001). Retrieved 2008-12-22. M.2-6). and Kilburn. "Selfish DNA: a sexually-transmitted nuclear parasite" (http:/ / www. sciencedirect. P. S.00027. doi:10. .1038/nature07381. H. nlm. php?Ma=1000 [117] Jokela. and Runnegar. and Michod. com/ journal/ 118814335/ abstract).003. and Tóth. Genetics 101 (3–4): 519–31. geoscienceworld. and Logsdon.0. M. Baron's Medical Microbiology (4th ed.: implications for the evolution of sex. com/ journal/ 119168103/ abstract).1146/annurev.1095-8312. [123] Judson. and Jobling. P. php?Ma=1500 [107] http:/ / toolserver. Michigan" (http:/ / www.102401. . Fletcher. In Baron. Retrieved 2008-09-02. . uiowa. Retrieved 2008-09-02. American Scientist 95: 224–231. D. doi:10.1111/j.0. . J. (1996). Tatiana's sex advice to all creation. "Reassessing the first appearance of eukaryotes and cyanobacteria". doi:10.1042/BST0340519. PMID 16856849.2004.O. "Extreme Microbes". Mol Biol Evol. CO.1997. J. [125] DasSarma. gen. Retrieved 2008-09-03. (1968). org/ cgi/ content/ abstract/ 283/ 5407/ 1476). Science 257 (5067): 232–235. .. gov/ articlerender.1365-2958.els. Retrieved 2008-09-02. H. htm). Retrieved 2008-08-02.1111/j. Jr. 468). interscience.. A. pubmedcentral. PMID 10066161. "Size and complexity among multicellular organisms" (http:/ / www3. Retrieved 2008-09-02. Retrieved 2008-09-03. section. A. fcgi?tool=pmcentrez& artid=1201875). . Biochemical Society Transactions 34 (Pt 4): 519–522.1-billion-year-old negaunee iron-formation.x. Science 283 (5407): 1476–1481. and Mooers. Annual Review of Genetics 35: 103–123. doi:10.. (1982). 77 . J. Retrieved 2008-12-22. 233–4. doi:10. M. H. Integrative Biology 1 (1): 27–36. D. biology. (2002). php?Ma=1430 [110] Butterfield. (2005)..R. (2007). B.01.x. Retrieved 2008-09-02. (July 1992). com/ 10.1472-4677.1126/science. Byerly.R. nature..A. (January 1999). B.1476. T. [109] http:/ / toolserver.genet. and the Mesoproterozoic/Neoproterozoic radiation of eukaryotes" (http:/ / paleobiol.2002.x. org/ bst/ 034/ 0519/ bst0340519. (2007). nih. 0. PMID 6293914. .2. org/ ~verisimilus/ Timeline/ Timeline. Hopf. O. wiley. J. php?Ma=1400 [108] Javaux. doi:10.1046/j.. A. "Origin of sex". [114] Bonner. R. N. (January 2005). Paleobiology 26 (3): 386–404. "Building a multicellular organism".Evolutionary history of life [99] Pisani. PMID 18948954. pl/ meid:10183). "Mitochondrial evolution" (http:/ / www. . M. Retrieved 2008-09-02 [119] Christie. (September 2000).CO. ncbi. Ltd. doi:10.1093/molbev/msm095. PMID 17504772. [112] Bell. [122] Bernstein.35. com/ science?_ob=ArticleURL& _udi=B6WMD-46DM0JD-1& _user=10& _rdoc=1& _fmt=& _orig=search& _sort=d& view=c& _version=1& _urlVersion=0& _userid=10& md5=3f5b76ba0b7fff999ee670479f06e5af).. M. Journal of Theoretical Biology 110 (3): 323–351. Biological Journal of the Linnean Society 60 (3): 345–363. J. pdf) (PDF).2-6.1016/j. Retrieved 2008-09-02. (1999). org/ ~verisimilus/ Timeline/ Timeline.1631544. F. and Walter. . org/ ~verisimilus/ Timeline/ Timeline. . php?Ma=2200 [103] Rasmussen.3008. org/ ~verisimilus/ Timeline/ Timeline. J. C.1002/(SICI)1520-6602(1998)1:1<27::AID-INBI4>3. sciencemag. wiley. New York: Metropolitan Books. (2001). doi:10. . wiley. (1984).2001. Nature 455 (7216): 1101–1104. R. php?Ma=2700 [102] http:/ / toolserver. interscience.CO.1126/science.CO.0. J. "Bangiomorpha pubescens n. Knoll. "Intelligence: Maze-solving by an amoeboid organism" (http:/ / www. Journal of Theoretical Biology 216 (4): 387–96. (September 2000). Current Biology 15 (2): 185–91. org/ ~verisimilus/ Timeline/ Timeline. H. Cotton. . and McInerney. html).1006/jtbi. [115] Butterfield. PMID 11028990. Retrieved 2008-09-03. "The Origins of Multicellularity" (http:/ / doi. "Type IV secretion: intercellular transfer of macromolecules by systems ancestrally related to conjugation machines" (http:/ / lib. doi:10. doi:10. Á. ISBN 0-9631172-1-1. . n. multicellularity. and Lang. and Gerstein. Extreme Microbes. (2002). A. Brocks.). S. "Probable Proterozoic fungi" (http:/ / paleobiol. Yamada.283. PMC 1201875.1666/0094-8373(2000)026<0386:BPNGNS>2. "On the origin of sex as vaccination" (http:/ / www. [126] Sterrer. Geobiology 2 (3): 121–132.W.1038/35035159. "TEM evidence for eukaryotic diversity in mid-Proterozoic oceans" (http:/ / www3. [111] Nakagaki. PMID 6209512. edu/ web/ jmlpubls/ rml05. T. sciencemag.1666/0094-8373(2005)031<0165:PPF>2. N.2. Galveston: University of Texas Medical Branch. [113] Kaiser. 24 (8): 1752–60.cub. [121] Otto.. S. J. doi:10. doi:10. 34: 355–384. [136] http:/ / toolserver. doi:10. doi:10. [139] Seilacher. Gwyneth. . .1144/gsjgs. [135] Hinde.earth.. .0714. D.. E.G. A. doi:10. Paleobiology 30: 203.Cambrian Biological Revolutions. [150] Budd. htm). 103001?journalCode=earth). Retrieved 2007-11-24. [146] Bengtson. E. . doi:10.157. Annu. Gehling. pp. html). D. (2006). . Zhao.1472-4669. org/ cgi/ content/ abstract/ 43/ 1/ 157) (Free full text). (2000-05-05). (2004).031504.. Retrieved 2008-09-03. S (2003).Evolutionary history of life [127] Dong. pdf). Peng. org/ ~verisimilus/ Timeline/ Timeline.. (PDF). html).00.1006/dbio. pdf) (PDF). Science 288 (5467): 841. Fedonkin. earth. J. and Zhou. doi:10.. J. confex.1601858.E.4.33.841. Abstracts with Programs (Geological Society of America) 35 (6): 105. htm) (abstract).1142024.. "Early skeletal fossils" (http:/ / www. Studer. (2001). "Silicified Horodyskia and Palaeopascichnus from upper Ediacaran cherts in South China: tentative phylogenetic interpretation and implications for evolutionary stasis" (http:/ / findarticles. edu/ cells/ animalcell. E. . J. Philippe. June 30th. (1989). Retrieved 2008-07-15.J. Padilla-gamiño. [151] Marshall. (2004). (1992). doi:10. J. Y. doi:10.els. B. Weersing. Norton & Company. Oliveri. and Waggoner. Encyclopedia of Life Sciences. Li. edu/ GG/ FACULTY/ GAIDOS/ geobiology2007. . Evans.L. magnet. and Kirschvink.1038/npg. Washington. [140] Martin.x. Dunford.. [148] Gould. (2006). [141] Fedonkin. White Sea. John's University. S. [152] Janvier. Retrieved 2008-09-03. [149] Budd. P.288. edu/ SSAUPE/ biol116/ Zoology/ digestion. A fossil fish. Zhang. 1146/ annurev... S.. A. but are now thought to be heavily modified members of the Cnidaria: Jímenez-Guri. Bioessays 28 (12): 1161–6. (1998).1093/icb/43. S. Shen.1126/science. ISBN 0195513681. Retrieved 2008-07-18 [147] Gould. P.. V. "The Cambrian Fossil Record and the Origin of the Phyla" (http:/ / intl-icb. B. M. Hu. org/ ~verisimilus/ Timeline/ Timeline. Retrieved 2008-09-03. "Explaining the Cambrian “Explosion” of Animals" (http:/ / arjournals. cosmonova. Science 317 (116): 116–118.Y. Integrative and Comparative Biology 43 (1): 157–165. org/ cgi/ content/ abstract/ 288/ 5467/ 841) (abstract). .. Dubuc. Journal of the Geological Society.1111/j.. (2007). College of St. "A Parvancorina-like arthropod from the Cambrian of South China". American Zoologist 38: 965–974. [142] Mooi. F. PMID 12142030. edu/ tietjen/ Evolution/ once_we_were_worms.tb01831. . Benedict / St. uwm. Florida State University. PMID 10797002.2002. H. ISBN 0091742714. "Hooking some stem-group "worms": fossil lophotrochozoans in the Burgess Shale".1. "Buddenbrockia is a cnidarian worm" (http:/ / www.L. org/ download/ 18. Bottjer. and Stanley. M.2007. Science News..-Y. (2003). "The Cnidaria and Ctenophora".1146/annurev. 28–57. "Vertebrata (Vertebrates)".W..G. J. S. Retrieved 2007-11-06. (July 2007). hawaii. Retrieved 2008-09-03.. M. . "The morphology of Opabinia regalis and the reconstruction of the arthropod stem-group".CO. [143] McMenamin.. uk/ site/ story/ 0. M. P. and Waggoner. Rev. . Wiley InterScience. London 149 (4): 607–613. "Once we were worms" (http:/ / cas. B.. Retrieved 2008-09-03. oxfordjournals. Yuan. com/ nature/ journal/ v388/ n6645/ abs/ 388868a0. L. (August 2002). A. org/ cgi/ reprint/ 38/ 6/ 965. sciencemag. 67–78. Oxford University Press. "Vendobionta and Psammocorallia: lost constructions of Precambrian evolution" (http:/ / jgs. html) (abstract). R. D. E. "Precambrian Animal Life: Probable Developmental and Adult Cnidarian Forms from Southwest China" (http:/ / www. "African fossils suggest complex life arose early" (http:/ / media. New Scientist 179 (2406): 34.. lyellcollection. (2006). (January 2008). co. T. "Concepts of Biology" (http:/ / employees.1038/42242. edu/ ~sdornbos/ PDF's/ Chen et al... W. Retrieved 2008-09-05. doi:10. M. 2010. Earth Planet. D. soest. (August 2. org/ cgi/ content/ abstract/ 317/ 5834/ 116).1080/08912960500508689. org/ cgi/ content/ abstract/ 149/ 4/ 607) (abstract). S. D. Dornbos.. guardian. 124–136 and many others. [144] Lin. has also been named Qingshania. Babcock. Geobiology 5: 351. In Anderson. doi:10. A. P.1126/science. A. Retrieved 2008-09-02.. Grazhdankin.. K. [138] Grazhdankin. S. "Animal Cell Structure" (http:/ / micro.1502-3931. C-W. PMID 17120226. (1999). Nature 388: 868–871. php?Ma=900 [131] Gaidos.149. Bowring.. Wonderful Life: The Burgess Shale and the Nature of History. Russia: Implications for Metazoan Evolution" (http:/ / www.14173. pdf) (PDF). 78 .103001. ISSN 0094–8373.. 031504. . 2003). Retrieved 2008-07-03.col1). Hutchinson Radius. scienceonline. and Davidson.. J. also from China. Retrieved 2007-11-24. J." (2000). htm). W. B. [130] http:/ / toolserver.. Developmental Biology 248 (1): 182–196. P. 33. McAndrew. J.X. Okamura. Neoproterozoic . [128] Dickey.2. com/ gsa/ 2003AM/ finalprogram/ abstract_62056. J. S.W.1111/j. nature. Retrieved 2007-06-21. Lethaia 29 (1): 1–14. fsu. Gon. csbsju. "Spriggina is a trilobitoid ecdysozoan" (http:/ / gsa.. B. Sci.1144/0016-76492007-074. [145] Butterfield. 10. (1997)... D. L. M..M. G. "The Precambrian emergence of animal life: a geobiological perspective" (http:/ / www.W.0607. S. Wednesday. doi:10. Historical Biology 18 (1): 33–45. . G. X. S. Retrieved on 2010-07-02. R. [132] Myxozoa were thought to be an exception. [133] Davidson. S. ISBN 039330700X. D..T. A. (1996). 4e32c81078a8d9249800021554/ Bengtson2004ESF. doi:10.H. doi:10. M.0. Gao. 2002. In Lipps. php?Ma=580 [137] Chen. Wonderful Life. . Invertebrate Zoology. and Holland.x. PMID 17615357.1996. "Patterns of distribution in the Ediacaran biotas: facies versus biogeography and evolution".1666/0094-8373(2004)030<0203:PODITE>2. bellarmine. L. "Age of Neoproterozoic Bilaterian Body and Trace Fossils. com/ p/ articles/ mi_qa3721/ is_200801/ ai_n24394476/ pg_1?tag=artBody.L. The name of one of these will have to change.00125. Journal of the Geological Society 165: 367–378.H. ISSN 0016–7649. .Q.5467. Xiao. annualreviews. H. C.M. oxfordjournals.1002/bies. S.20507. pdf) (PDF). D. [134] Saupe. N. Yu. Retrieved 2008-07-03. pp.C. and Bruno.. org/ doi/ abs/ 10. R. "The late Precambrian fossil Kimberella is a mollusc-like bilaterian organism" (http:/ / www.0001531 [153] Conway Morris. (1989).. T. E. [129] Name given as in Butterfield's paper "Bangiomorpha pubescens . J. "Evolution within a bizarre phylum: Homologies of the first echinoderms" (http:/ / icb. C. Braddy. D. org/ ~verisimilus/ Timeline/ Timeline..1130/0091-7613(2002)030<0391:FSOLAT>2. J. [158] Selden. T. . . Han. and Escudero. 79 . PMC 1692181.. 71–74. php?Ma=370 [169] Scheckler. (2001). Retrieved 2008-09-05. Cambridge University Press. nature. 155–160. Feakes. Evolution of the Insects. Blackwell. . PMID 17769314. (1987). p. and Engel.0. niu. com/ ?id=AHsrhGOTRM4C& pg=PA69& lpg=PA69& dq=devonian+ meandering+ plants+ trees).. D. D..CO. and Smith. org/ ~verisimilus/ Timeline/ Timeline. and Scheckler. Li. 156–171.1038/nature01264. (2005).. doi:10. com/ Vertebrates/ Units/ 150Tetrapoda/ 150. M.1016/0034-6667(95)00086-0. P. "The Early Development of Terrestrial Ecosystems" (http:/ / books. Yasui.235. K. "Terrestrial-marine teleconnections in the Devonian: links between the evolution of land plants. botany. nih. html).. Retrieved 2008-09-05. E. (1998). Review of Paleobotany and Palynology 90: 249–262. Mangano. P. org/ ~verisimilus/ Timeline/ Timeline.. E... ISBN 0521821495. Encyclopedia of Life Sciences. Taylor and Francis. ISBN 0521821495. ""Terrestrialization of Animals"" (http:/ / books. In Gee. Retrieved 2008-09-05 [170] The phrase "Late Devonian wood crisis" is used at "Palaeos – Tetrapoda: Acanthostega" (http:/ / www. T. 67–70. D. D.. Conway Morris.-L.A. PMID 15282604. G. pubmedcentral. gov/ articlerender. (June 1998).. org/ cgi/ content/ abstract/ 30/ 5/ 391). Retrieved 2009-01-11. 169–184. (2004).R. S. Retrieved 2008-09-05. sciencedirect. doi:10. A. Blackwell. Major Events in Early Vertebrate Evolution.. pp. M. . Zhu. palaeos. . Cole. J. L. (2001). J. Zhang. [171] Algeo. History of Life (3rd ed. doi:10. pdf) (PDF). X-L. John Wiley & Sons. Y. Retrieved 2009-01-11. Retrieved 2008-09-09 [162] Hawksworth.R.. G. H-L. (2000). com/ ?id=Ql6Jl6wKb88C& dq="Evolution+ of+ the+ Insects"+ grimaldi& printsec=frontcover). M. Edgecombe. E. P. . weathering processes. and Chen. PALAEOS: The Trace of Life on Earth. Retrieved 2008-09-05..Evolutionary history of life [154] Shu. pp. com/ ?id=Ql6Jl6wKb88C& dq="Evolution+ of+ the+ Insects"+ grimaldi& printsec=frontcover). Retrieved 2008-09-05. doi:10. . edu/ davis/ bios458/ Shu1. Li. L. Liu. P. D-G. Nature 402: 42–46. [175] Vaccari. Janvier. R. . N. and marine anoxic events" (http:/ / www. N. X. geoscienceworld. org/ ~verisimilus/ Timeline/ Timeline. com/ article. html). evolutionary trends in environmental expansion. F. D.G. PALAIOS (PALAIOS. S. php?Ma=476 [167] http:/ / toolserver. S. P.-N. Retrieved 2008-09-05 [159] http:/ / toolserver.-Q. com/ science?_ob=ArticleURL& _udi=B6V6W-454YDFK-7& _user=10& _rdoc=1& _fmt=& _orig=search& _sort=d& view=c& _acct=C000050221& _version=1& _urlVersion=0& _userid=10& md5=9d5d008d99d044684e947ad74b05514d). H. Conway Morris. C. edu/ courses/ botany_940/ 06EvidEvol/ papers/ KendrickCrane1997. doi:10.. J. P. doi:10. and Lukie. and Crowther. ISBN 0632044446..E. Canada" (http:/ / geology. [182] Clack. P. A.0000368 [163] http:/ / toolserver. History of Life (3rd ed. [156] Sansom I. University of Chicago Press. [155] Shu. 150.E. [172] Taylor T. org/ cgi/ content/ abstract/ 13/ 3/ 217). google. "The Ordovician radiation of vertebrates". Scientific American. Nature 430 (6999): 554–557. [180] Grimaldi.4784. No. In Ahlberg.. doi:10.. T. sepmonline. R. D.. "Insects Take to the Skies" (http:/ / books. J. pdf) (PDF). Ltd. J. Evolution of the Insects. .-F. [179] Cowen. (November 1999). "Lichens". D.. [181] Grimaldi.G. Zhang. Retrieved 2008-09-05. bios. and Crane. D. p. A. S-X. Cambridge University Press. org/ ~verisimilus/ Timeline/ Timeline. In Briggs.. "Head and backbone of the Early Cambrian vertebrate Haikouichthys" (http:/ / www. Han. google. Vol.-G. J. M. php?Ma=445 [177] http:/ / toolserver.1038/37918. M.J. google. (January 2003). and behavioral complexity" (http:/ / palaios. . Briggs.. Retrieved 2008-09-05. "First steps on land: Arthropod trackways in Cambrian-Ordovician eolian sandstone. [173] http:/ / toolserver. and Taylor. and Liu. M.). com/ ?id=AHsrhGOTRM4C& pg=PA71& lpg=PA71& dq="Terrestrialization+ of+ Animals"+ selden). ISBN 0632051493.R.1038/46965. "Diversity of evolution" (http:/ / books. [176] http:/ / toolserver.. Shaking the Tree: Readings from Nature in the History of Life.els. doi:10. org/ ~verisimilus/ Timeline/ Timeline. G. and Crowther. 126. doi:10. L. H. and Engel.. P. (2000). E. M. M. 120–122. In Briggs. J. (November. Chen. doi:10. Retrieved 2008-09-06. ISBN 0632051493. php?Ma=490 [174] MacNaughton. Y. N. Luo. R.L. (2000). W. php?Ma=385–359 [161] Shear.2. 12. J... . and Osborn J..61. php?Ma=430 [168] http:/ / toolserver. fcgi?tool=pmcentrez& artid=1692181). cfm?id=getting-a-leg-up-on-land). S.. Palaeobiology II: A Synthesis. Z. W. Nature 421 (6922): 526–529. pp. com/ nature/ journal/ v421/ n6922/ abs/ nature01264. G.. "Trace Fossil Evidence for Late Ordovician Animals on Land". B. M. ISBN 0226284964. ISBN 0632044446. "The origin and early evolution of plants on land" (http:/ / botit. R. . php?Ma=423–419 [164] Retallack. pp. Smith.1126/science. Dalrymple. google. (2005). Science 235 (4784): 61–63. (1996). [165] Kenrick.E.. C. "The ichnologic record of the continental invertebrate invasion. (May 2002). (2001).1038/nature02705.2307/3515447. "Cambrian origins and affinities of an enigmatic fossil group of arthropods".1998. php?Ma=415 [178] Buatois. ISBN 0-415-23370-4 [157] Cowen. Philosophical Transactions of the Royal Society: Biology 353: 113–130.. "Getting a Leg Up on Land" (http:/ / www. com/ ?id=ZJe_Dmdbm-QC& pg=PA233& dq=evolution+ flowering+ plant+ angiosperm). Nature 389: 33. Blackwell Science. Palaeobiology II: A Synthesis. Hu. 2005). org/ ~verisimilus/ Timeline/ Timeline.. 13. php?Ma=488–444 [160] http:/ / toolserver. ""Afforestation – the First Forests"" (http:/ / books. [166] http:/ / toolserver. pp. ecospace utilization.). (September 1997).1098/rstb. org/ ~verisimilus/ Timeline/ Timeline.0195. southeastern Ontario. 3) 13 (3): 217–240. pp. org/ ~verisimilus/ Timeline/ Timeline. . R. Zhang. S. sciam. Geology 30 (5): 391–394. "The importance of fungi in shaping the paleoecosystem" (http:/ / www. "Lower Cambrian vertebrates from south China" (http:/ / www. google. PMID 12556891.. L-Z. J. wisc. Genise. Chen. (2001). Blackwell Science.1038/npg. doi:10. Bremer K. T.. E. R. Berkeley: University of California Press. J. Zhou. D. A-L. doi:10.tb01280. R.. org/ ~verisimilus/ Timeline/ Timeline. fr/ ~boudouresque/ Publications_DOM_2006_2007/ Daeschler_et_al_2006_Nature. pdf). F. (May 1990). J. and Jenkins. L. (2007). doi:10. W. Nature 434 (7035): 886–889. pdf) (PDF). uk/ Benton/ reprints/ 1983success. J. (2005). Physiological and Biochemical Zoology 77 (5): 697–699. R. E. Retrieved 2008-09-07. "Tetrapod Phylogeny" (http:/ / si-pddr. pdf) (PDF). and Feduccia. Parrish.. PMID 16598249. com/ nature/ journal/ v434/ n7035/ full/ nature03393. com/ nature/ journal/ v368/ n6471/ abs/ 368507a0. sciencemag.. M. P. Retrieved 2008-09-08. oxfordjournals. pp. doi:10. interscience. . php?Ma=199 [201] Benton. (September–October 2005). edu/ dspace/ bitstream/ 10088/ 4689/ 1/ VZ_1989GauthieretalHierLife. Damiani. Larsson.CO. American Zoologist 40 (4): 585–596. and Zhang. Martin. J. Retrieved 2008-09-07.1086/425182. O. H. . (March 1983). S. Retrieved 2008-09-06. Smith. (April 1994). 80 . C.x.2007. Rakotosamimanana. . T. "Recovery from the most profound mass extinction of all time" (http:/ / journals.1038/43420.. (2006). univ-mrs. M. PMID 17047029. R. W. C.. ac. J. . Thames & Hudson. C. A.. "A New Mammaliaform from the Early Jurassic and Evolution of Mammalian Characteristics" (http:/ / www. de Queiroz. M. H. interscience. O'Keefe.. L. ISBN 0-520-24209-2. [209] http:/ / toolserver. Zoological Journal of the Linnean Society 120 (3): 281–354. Retrieved 2008-09-08. Z-X. N.1038/377616a0.. doi:10. "The Origin and Early Diversification of Tetrapods" (http:/ / www. org/ ~verisimilus/ Timeline/ Timeline. .1111/j. .4. PMID 18198148. html). . php?Ma=363 [187] Daeschler.1016/j.2005. . com/ science?_ob=ArticleURL& _udi=B6X1G-4GYH7VN-1& _user=10& _rdoc=1& _fmt=& _orig=search& _sort=d& view=c& _version=1& _urlVersion=0& _userid=10& md5=add24b0622f2aff0b41e7c42a3160fa7). D. J. [188] Debraga. [189] Benton M. bris.1111/j. H. Kluge. Simpson. (2000). When Life Nearly Died: The Greatest Mass Extinction of All Time. doi:10. doi:10. php?Ma=195 [207] Luo. Journal of Molecular Evolution 30 (5): 409–424. si. (April 2005). PMID 2111854. Retrieved 2008-09-08 [200] http:/ / toolserver. B.Evolutionary history of life [183] Ahlberg. M.. and Sun. com/ nature/ journal/ v401/ n6748/ abs/ 401057a0. (May 2001). F. 210–231. . Comptes Rendus Palevol 4 (6-7): 623–636. Journal of Anatomy 208 (3): 287–308. org/ ~verisimilus/ Timeline/ Timeline. and Maga. [191] http:/ / toolserver. Shubin.1058476. David B.1126/science. and Wyss.005. The Dinosauria (Second ed. org/ content/ qq5un1810k7605h5/ fulltext. J. Nature 368: 507–514. Z. org/ cgi/ content/ full/ 40/ 4/ 585).J. doi:10. A. Quarterly Review of Biology 58 (1). .. M. php?Ma=313 [194] http:/ / toolserver. The Hierarchy of Life. G. "Reptile phylogeny and the interrelationships of turtles" (http:/ / www3. org/ ~verisimilus/ Timeline/ Timeline.00534. P. php?Ma=314 [193] http:/ / toolserver. In Weishampel. S. Retrieved 2008-09-06. S. "A Devonian tetrapod-like fish and the evolution of the tetrapod body plan" (http:/ / www. "Dinosaur Success in the Triassic: a Noncompetitive Ecological Model" (http:/ / palaeo. (April 2006). A.. org/ cgi/ content/ full/ 24/ 1/ 26). html). gly. [184] http:/ / toolserver. nature. p. & Osmólska. html). Halszka (eds. . [198] Sahney.). Journal of Paleontology 75: 1214. php?Ma=229 [195] Sidor.1093/icb/40. K.L. M. "Early mammalian radiations" (http:/ / findarticles. (October 2002). nature.1997. "Revisiting the Vertebrate Invasion of the Land". (2008). Elsevier Science. A. Steyer. . . J. Dodson. (July 1997). B. "A beaked bird from the Jurassic of China" (http:/ / www. Z. doi:10. K. Peter. PMID 16533313. Proceedings of the Royal Society: Biological 275 (1636): 759. "The recovery of terrestrial vertebrate diversity in the South African Karoo Basin after the end-Permian extinction" (http:/ / www. P. nature. (1989).2. royalsociety.1098/rspb. R. Crompton. php?Ma=167 [210] Flynn. org/ ~verisimilus/ Timeline/ Timeline. J.. F. . A.1007/BF02101113. . M. A.. (September 1999). Nature 401: 57–60.0. Retrieved 2008-09-08.. html). "Paleontological Evidence to Date the Tree of Life" (http:/ / mbe. J. and Rowe. B. 345. Retrieved 2008-09-08. wiley. (September/October 2004). O. (November 2001). B.585. A. J. org/ ~verisimilus/ Timeline/ Timeline. Ide. Graham.. Retrieved 2008-09-08. doi:10. (2004).07. E. Retrieved 2008-09-08 [202] Padian. D. org/ ~verisimilus/ Timeline/ Timeline. org/ cgi/ content/ full/ 292/ 5521/ 1535). and Milner.1370. doi:10. com/ journal/ 119830935/ abstract). and Botha.1038/nature03393. sciencedirect. Retrieved 2008-09-07. php?Ma=360 [185] Gordon. doi:10. "Permian tetrapods from the Sahara show climate-controlled endemism in Pangaea" (http:/ / www. T. and Donoghue.1096-3642. . In B. nature.. [196] Smith. [197] Benton. PMC 2596898.1666/0022-3360(2001)075<1214:EMR>2. org/ ~verisimilus/ Timeline/ Timeline. "Selective Factors Associated with the Origin of Fur and Feathers" (http:/ / icb... "A Middle Jurassic mammal from Madagascar" (http:/ / www. "Phylogeny of the Major Tetrapod Groups: Morphological Data and Divergence Dates" (http:/ / www.). [203] Hou. Fernholm. C. Molecular Biology and Evolution 24 (1): 26–53. [208] Cifelli. Nature 440 (7085): 757–763. [190] Benton. [186] http:/ / toolserver. (PDF). [199] Gauthier. F. J. com/ journal/ 118559634/ abstract?CRETRY=1& SRETRY=0). wiley. A. PMC 2100246. J.1038/nature04639. pdf) (PDF). "Basal Avialae". and Benton. [206] http:/ / toolserver. doi:10. [204] Clarke. A. com/ nature/ journal/ v377/ n6550/ abs/ 377616a0.1469-7580. J. doi:10.. C. Nature 377: 616–618. PMID 11375489.crpv. Sereno.R. com. Science 292 (5521): 1535–1540.2006. php?Ma=330 [192] http:/ / toolserver. A. PMID 15829962. Zhou. "Insight into the evolution of avian flight from a new clade of Early Cretaceous ornithurines from China and the morphology of Yixianornis grabaui" (http:/ / www3. ISBN 978-0500285732. [205] Ruben. and Wang. org/ ~verisimilus/ Timeline/ Timeline. H.1093/molbev/msl150.x. and Jörnvall. R. com/ content/ k152294003652458/ )... oxfordjournals. doi:10. Cannatella. springerlink. and Jones. and Rieppel.1038/368507a0. com/ p/ articles/ mi_qa3790/ is_200111/ ai_n8958762/ pg_6). M. . Gondwana Alive: Biodiversity and the Evolving Biosphere. and Henneberg.1146/annurev.1073/pnas. [234] Martin. Habersetzer. . sciencemag. [230] http:/ / toolserver.. Retrieved 2008-09-09. A. doi:10.earth. F. (1995).. E. sciencemag..). doi:10. N... an Eocene cetacean (Mammalia) from Pakistan". (2000). Behavioral Ecology and Sociobiology 28 (1): 37–45. Witwatersrand University Press. M. M. [211] MacLeod. org/ cgi/ content/ abstract/ 320/ 5880/ 1213).Smith.. HOMO .. ISBN 0226284964. pdf+ html). PMC 34068. 463). and Pedersen. I. com/ science?_ob=ArticleURL& _udi=B7GW4-4DPCHXC-2& _user=10& _rdoc=1& _fmt=& _orig=search& _sort=d& view=c& _version=1& _urlVersion=0& _userid=10& md5=aef79dbca1f189c885cfe9f36636b131). com/ p/ articles/ mi_qa3721/ is_199703/ ai_n8738406/ print). santafe. and Ratnieks. "Environment and Behavior of 2. PMID 12078635. php?Ma=2. E. B.0510792103. PMID 18270539.. Retrieved 2008-09-24. D. White. Global and Planetary Change 11 (1): 1.1016/0921-8181(94)00011-2. J. A. (January 1991). doi:10. Retrieved 2008-09-09. .5-Million-Year-Old Bouri Hominids" (http:/ / www. Milner.000 years ago?" (http:/ / www. Morris. Evans. D. C. (2000).. Culver. full.1038/nature00879. M. [219] Hughes. Proceedings of the National Academy of Sciences 97 (24): 12939–12941. Madar. G. [229] Mellars. ISBN 0465053130.1080/106351599260472.. S. asp?ref=0632056371). PMC 1480416. P. . F. C. 81 . org/ content/ 97/ 24/ 12939. Courier Forschungsinstitut Senckenberg 191: 1–86. L.1126/science. "The Cretaceous–Tertiary biotic transition" (http:/ / findarticles. com/ content/ k4n52v522l816125/ ). nature.0505858102. and de Wit. ISBN 978-0-632-05637-8. N. (1996). doi:10. J.284. E. S. Richard (1994). [214] Simmons. T. html). doi:10..1156108. S.463. B. Rosen.. "Quantitative Analysis of the Timing of the Origin and Diversification of Extant Placental Orders" (http:/ / www. Retrieved 2008-09-09. P. success. org/ cgi/ content/ full/ sci. New York. G. Journal of the Geological Society 154 (2): 265–292. C. .. PMC 1224642. (PDF). (1999). Reptiles Of The Triassic" (http:/ / www. pnas. [227] De Miguel.2. T. . (2001). (2004). NY: Basic Books. R. "A new hominid from the Upper Miocene of Chad. M. [213] Archibald. K. F. M. L. doi:10. R. Chambers.1126/science. Banner. D. Nature 451 (7180): 818–821.1023/A:1011317930838. M. J..97. J.1038/nature06549. [225] http:/ / toolserver.Evolutionary history of life Retrieved 2008-09-08. pdf). doi:10. J.1. [232] Van Valkenburgh.. D. . [233] MacLeod. van Wyk. H. annualreviews. Science (American Association for the Advancement of Science) 320 (5880): 1213–1216.. Science 284 (5414): 625–629. P. Blackwell.. Retrieved 2008-09-11. Science Masters Series. R. pnas.284/ 5414/ 625). "Why did modern human populations disperse from Africa ca. (June 2001). 1146/ annurev. and Eble. K. Lord. pp. . and relationships: Darwin’s abominably "perplexing phenomenon"" (http:/ / www. Retrieved 2008-09-09. [216] Crane. 5 [226] de Heinzelin. "Ambulocetus natans.5414. L. doi:10. html). A. et al (April 1999). "The evolution of eusociality in molerats (Bathyergidae): a question of risks. University of Chicago Press. doi:10. H.. K. . Milner. H. "The fossil record of North American mammals: evidence for a Paleocene evolutionary radiation". P. 87–89. A. D. P..... F. W. org/ ~verisimilus/ Timeline/ Timeline. . Retrieved 2008-09-09 [217] Crepet. Beekman. Rawson. J.625. com/ nature/ journal/ v418/ n6894/ abs/ nature00879. [212] Alroy. Guy. 233–250. W. google. sciencedirect. "Primitive Early Eocene bat from Wyoming and the evolution of flight and echolocation". Proceedings of the National Academy of Sciences 102 (38): 13367–13371. "Major patterns in the history of xarnivorous mammals" (http:/ / arjournals.1144/gsjgs. PMID 12110880.. org/ ~verisimilus/ Timeline/ Timeline. B. T. et al (July 2002). "Cyclic and secular variation in microfossil biomineralization: clues to the biogeochemical evolution of Phanerozoic oceans". C. doi:10. The Origin of Humankind. [215] Thewissen... Pilbeam. 60. .1078/0018-442X-00019. In Anderson. . ingentaconnect. PMID 18511689.1073/pnas. [218] Wilson. Nature 418 (6894): 145–151. M. Systematic biology 48 (1): 107–18. B. "Eusociality: Origin and consequences" (http:/ / www. B. T. and Young. Central Africa" (http:/ / www. pdf+ html). J. "The Origin and Early Diversification of Angiosperms" (http:/ / books. PMID 11087846. E. Urquhart. and Deutschman. (March 1999). R.. (November 2000). ISBN 978-3-510-61084-6. Jeffery. J. pnas. G. Annual Review of Earth and Planetary Sciences 26: 463–493. org/ content/ 102/ 38/ 13367. Paul (2006). Seymour. "Ancestral Monogamy Shows Kin Selection Is Key to the Evolution of Eusociality" (http:/ / www. PMID 16157878. com/ home/ articles/ earth/ extinction-page-2-1_1258. Burnett. L.. "Variation in hominid brain size: How much is due to method?" (http:/ / www.1007/BF00172137. doi:10. O.. [220] Lovegrove. php?Ma=300 [223] Labandeira.154. . and Hussain. M. Taylor... and Gunnell. J. Forey. Thackeray. "Extinction!" (http:/ / www. springerlink. org/ doi/ abs/ 10. org/ ~verisimilus/ Timeline/ Timeline.27. Boudagher-Fadel. In Gee. B. B. [221] http:/ / toolserver. Oldroyd. Vertebrate Palaeontology (3rd ed. E. 27. Clark. E. (2001-01-06). org/ cgi/ reprint/ 0510792103v1). Kaminski. and costs" (http:/ / www. Bown. R. "Progress in understanding angiosperm history.. A. W.Journal of Comparative Human Biology 52 (1): 3–58. Mackaye.. doi:10. full. org/ ~verisimilus/ Timeline/ Timeline. firstscience. (2008-05-30). com/ content/ klu/ jomm/ 2001/ 00000008/ 00000002/ 00342277). Retrieved 2008-08-04. M. (February 2008). (September 2005). "6. . 1.12939. PMID 10213682. P. Proceedings of the National Academy of Sciences 103 (25): 9381. N. numbers. . pp.24. G. H.. Journal of Mammalian Evolution 8 (2): 107–124. php?Ma=542 [231] Benton. Shaking the Tree: Readings from Nature in the History of Life. Friis. "The Fossil Record of Insect Diversity and Disparity" (http:/ / www.1073/pnas. P. R. J. com/ ?id=ZJe_Dmdbm-QC& pg=PA233& dq=evolution+ flowering+ plant+ angiosperm). and Hölldobler. blackwellpublishing. [228] Leakey. php?Ma=400 [222] http:/ / toolserver. Retrieved 2008-09-07 [224] Brunet. F. Retrieved 2008-09-07. J. PMID 16772383. doi:10.0265. Retrieved 2008-09-07. edu/ research/ publications/ workingpapers/ 00-08-044. F. earth. doi:10. (1997).. com/ book. R.. B. R. . O. N. A. Owen. S. htm) . R. (1996)... E. Genetics 154 (3): 985–997. A. genetics.berkeley. PMC 1460977. (March 2005). Retrieved 2008-09-11.. and Falkowski. Oxford University Press. and Muller.fossilmuseum. php?Ma=542–400 [238] http:/ / toolserver.htm) Everything you wanted to know about evolution by New Scientist (http://www. V. "Primary Production of the Biosphere: Integrating Terrestrial and Oceanic Components" (http:/ / www. and Implications (http://www.uk) • Understanding Evolution: History. John Maynard.howstuffworks. External links General information • • • • • General information on evolution. B. (2004). doi:10. gov/ papers/ Rohde-Muller-Nature. PMID 15758998. [243] Hawks. No. (2002). "Recent acceleration of human adaptive evolution" (http:/ / www. org/ ~verisimilus/ Timeline/ Timeline. [236] Rohde. Science 281 (5374): 237–240. pdf) (PDF). oxfordjournals. Boston: Houghton Mifflin Company.1073/pnas.htm) Understanding Evolution from University of California. . pnas. Vol. Wang. doi:10. Eörs Szathmáry (1997). org/ cgi/ content/ abstract/ 93/ 2/ 86). Cochran. J. The Selfish Gene. .com/evolution. and Moyzis. Nature 434 (7030): 208–210. doi:10. G. lbl. Retrieved 2008-09-11. "Compensatory Mutations. C. (http://www. . Theory. palomar. Journal of Heredity 93 (2): 86–90. • Richard Dawkins. L. S. PMID 9657713. C.rationalrevolution. R. A.edu/) National Academies Evolution Resources (http://nationalacademies.com/channel/life/ evolution) • Howstuffworks. N. full). Retrieved 2008-09-11. H. The Major Transitions in Evolution.). PMID 12140267. org/ cgi/ content/ full/ sci.237. 11. and diversity of the marine biosphere" (http:/ / jstor. 82 References Further reading • Cowen. R. org/ ~verisimilus/ Timeline/ Timeline. Retrieved 2008-09-13.2. ISBN 978-1405117562. and Wiseman. doi:10. .1093/jhered/93. ISBN 0-618-00583-8. php?Ma=400–200 [239] http:/ / toolserver. Berkeley (http://evolution. Blackwell Publishing Limited. (December 2007). Perrot.net/ articles/understanding_evolution. 2000). R.E. org/ content/ 104/ 52/ 20753.281/ 5374/ 237). org/ stable/ 3515230). J. M. B. Randerson. R. php?Ma=200 [240] Field.com/tree_of_life.86. and Walker. (March 1. L. Oxfordshire: Oxford University Press. History of Life (4th ed. [241] Grant. • Smith. sciencemag. Retrieved 2008-09-22. M. (July 1998). T. [237] http:/ / toolserver. Harpending. ISBN 0-198-50294-X..Fossil Museum nav. (1990).htm/printable) • Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and Theories (http://anthro.newscientist.281.. PMC 2410101. ISSN 1465-7333. . org/ ~verisimilus/ Timeline/ Timeline. R. ISBN 0192860925. doi:10. "Secular increase in nutrient levels through the Phanerozoic: Implications for productivity. J.Evolutionary history of life [235] Martin. 2004.2307/3515230. "Cycles in fossil diversity" (http:/ / muller. 3) 11 (3): 209–219. Antibiotic Resistance and the Population Genetics of Adaptive Evolution in Bacteria" (http:/ / www. org/ cgi/ content/ abstract/ 154/ 3/ 985).0707650104. PMID 18087044..com — How Evolution Works (http://science. Behrenfeld. Proceedings of the National Academy of Sciences 104 (52): 20753–20758. . Evidence. T.5374.1038/nature03339. P. A Pilgrimage to the Dawn of Life. biomass.org.edu/synthetic/) History of evolutionary thought • The Complete Work of Charles Darwin Online (http://darwin-online. B.1126/science. "Recent History of Melanism in American Peppered Moths" (http:/ / jhered. [242] Levin..net/Evolution.org/evolution/) Evolution poster. PALAIOS (PALAIOS.PDF format "tree of life" (http://tellapallet. • The Ancestor's Tale. K. PMID 10757748. 200. 3 billion years of photosynthesis. 65 million years since the non-avian dinosaurs died out.000 years since humans started looking like they do today. .5 billion year old Earth. Its occurrence over large stretches of time explains the origin of new species and ultimately the vast diversity of the biological world. 475 million years of land plants. 25. evolution is the process by which populations of organisms acquire and pass on novel traits from generation to generation. 400 million years of insects and seeds. For a thorough explanatory context. Detailed timeline Ma.5 million years since the appearance of the genus Homo.8 billion years of simple cells (prokaryotes).Timeline of evolution 83 Timeline of evolution This timeline of the evolution of life outlines the major events in the development of life on the planet Earth (See Organism). 2. 600 million years of simple animals. 570 million years of arthropods (ancestors of insects. with (very approximate) dates: • • • • • • • • • • • • • • • • • • • 3. 200 million years of mammals. products of evolution and speciation over billions of years. and geologic time scale.000 years since Neanderthals died out. 1 billion years of multicellular life. Contemporary species are related to each other through common descent. arachnids and crustaceans). ka means "thousand years ago" and ya means "years ago" Hadean Eon 3800 Ma and earlier. 300 million years of reptiles. ("megaannum") means "million years ago". 550 million years of complex animals. 150 million years of birds. 2 billion years of complex cells (eukaryotes). 500 million years of fish and proto-amphibians. 360 million years of amphibians. see the history of Earth. Basic timeline The basic timeline is a 4. 130 million years of flowers. In biology. The dates given in this article are estimates based on scientific evidence. (See Oxygen catastrophe) 2700 Timeframe of cyanobacteria evolution suggested by more recent research. a set of chemical reactions that free the 2500 Ma energy of organic molecules such as glucose and store it in the chemical bonds of ATP. or possibly not. The atmosphere and the oceans form.[19] More recent Ma research. suggests a later time of 2700 Ma. Ma Proterozoic Eon 2500 Ma – 542 Ma .Timeline of evolution 84 Date Event 4600 Ma The planet Earth forms from the accretion disc revolving around the young Sun. unchanged. Somewhere Cells resembling prokaryotes appear. creating iron ore. sending a very large number of moonlets into [1] orbit around the young Earth. Later. 3900 Ma Late Heavy Bombardment: peak rate of impact events upon the inner planets by meteors. thereby producing oxygen as waste product. Between 4500 The earliest life appears. These organisms generate ATP by exploiting a proton gradient. or life might have been transported to Earth by a meteor. they use water as a reducing agent. which soon become limited. DNA molecules then take over as the main replicators and these archaic genomes soon develop inside enclosing membranes which provide a stable physical and chemical [7] [8] [9] environment conducive to their replication: proto-cells. resulting in competition. possibly derived from self-reproducing RNA molecules. The oxygen concentration in the atmosphere subsequently rises. 3000 Photosynthesizing cyanobacteria evolve.[12] These first organisms are chemoautotrophs: they use carbon dioxide as a carbon source between 3900 . a mechanism still used in virtually all organisms. These moonlets eventually coalesce to form the Moon.[17] Ma [18] Bacteria develop primitive forms of photosynthesis which at first do not produce oxygen.and oxidize inorganic materials to extract energy. 4100 Ma The surface of the Earth cools enough for the crust to solidify. may have led to the RNA world of competing organic compounds. The oxygen initially oxidizes dissolved iron in the oceans.[4] and iron sulfide synthesis along deep ocean platelet boundaries. Glycolysis (and ATP) continue to be [13] [14] used in almost all organisms. to this day. 4500 Ma According to one plausible theory. These extreme mixing influences are thought to stimulate evolutionary processes. space.[3] PAH infall. prokaryotes evolve glycolysis. the planet Earth and the planet Theia collide. and smaller building blocks. as some early microbes could have survived in hydrothermal [10] [11] vents below the Earth's surface.[5] [6] The replication of these organisms and 3500 Ma requires resources like energy. with natural selection favouring those molecules which are more efficient at replication. The earth is continually wracked by hurricane force winds. acting as a poison for many bacteria.[15] [16] the split between bacteria and archaea occurs. The gravitational pull of the new Moon [2] stabilises the Earth's fluctuating axis of rotation and sets up the conditions in which life formed. however. The moon is still very close to the earth and causes tides 1000 feet (305 m) high. Archean Eon 3800 Ma – 2500 Ma Date Event 3500 Lifetime of the last universal ancestor. This constant disturbance may have obliterated any life that had evolved to that point. [32] 525 Earliest graptolites.[34] They are accompanied Ma by fungi. literally the "period of well-displayed life". Ma 440 First agnathan fishes: Heterostraci. . Paleozoic Era 542 Ma – 251.g. the Paleozoic.Timeline of evolution 85 Date Event By 1850 Eukaryotic cells appear. arthropods (e.[31] This blocks ultraviolet radiation. etc. Ma brachiopods.although their affinities remain a subject of debate. mollusks. shell-forming and/or trace-making organisms. which are divided by major mass extinctions.[25] [26] [27] 580–542 The Ediacaran biota represent the first large.0 Ma Date Event 535 Major diversification of living things in the oceans: chordates. (See Endosymbiosis) By 1200 Sexual reproduction first appears.[23] [24] Opinion is divided on whether it increased or decreased biodiversity or the rate of Ma evolution. echinoderms. Ma 450 Land arthropod burrows (millipedes) appear. which may have aided the colonisation of land through symbiosis. 850–630 A global glaciation may have occurred. Mesozoic and Cenozoic.[22] Ma 1200 Ma Simple multicellular organisms evolve. 530 The first known footprints on land date to 530 Ma.[33] having evolved from green algae living along the edges of lakes. complex multicellular organisms . increasing the rate of evolution. Galeaspida. Ma 485 First vertebrates with true bones (jawless fishes). permitting the Ma colonisation of the land.[28] Ma 580–500 Most modern phyla of animals begin to appear in the fossil record during the Cambrian explosion. and Pituriaspida. mostly consisting of cell colonies of limited complexity. Ma 510 First cephalopods (Nautiloids) and chitons. Ma 434 The first primitive plants move onto land. It is subdivided into three eras. foraminifers and radiolarians.[31] 560 Ma Earliest fungi Phanerozoic Eon 542 Ma – present The Phanerozoic Eon. marks the appearance in the fossil record of abundant. along with the first complete conodonts and echinoids. crustaceans).[29] [30] Ma 580–540 The accumulation of atmospheric oxygen allows the formation of an ozone layer. indicating that early animal explorations may have predated the development of Ma terrestrial plants. Ma 505 Fossilization of the Burgess Shale.[20] [21] Eukaryotes contain membrane-bound organelles with diverse functions. trilobites. probably derived from Ma prokaryotes engulfing each other via phagocytosis. Insects roamed the land and would soon take to the skies.[35] and vegetation covered the land. Ma 245 Ma Earliest ichthyosaurs. diversity in cycads. with seed-bearing plants and forests soon to flourish. Earliest harvestman. Mesozoic Era Date Event From The Mesozoic Marine Revolution begins: increasingly well-adapted and diverse predators pressurise sessile marine groups. Four-limbed tetrapods gradually gain adaptations which will help them occupy a terrestrial life-habit. Terrestrial organisms were not as seriously affected as Ma the marine biota. but life on land took 30M years to completely [36] recover. seed plants and conifers diversify while lepidodendrids and sphenopsids decrease. and land scorpions. trigonotarbid arachnids. the Earth begins to be recognisable. Terrestrial temnospondyl amphibians Ma and pelycosaurs (e. 240 Ma Increase in diversity of gomphodont cynodonts and rhynchosaurs.g. Ma 395 First lichens. and ammonoids. hexapods (springtails). lycophytes. 360 First crabs and ferns.g. and trimerophytes. First teleost fishes. 215 Ma First mammals (e.4 The Permian-Triassic extinction event eliminates over 90-95% of marine species. Petrolacosaurus). stoneworts. and conifers. first cardiid bivalves. Eozostrodon). Ma 280 Earliest beetles.g. Dimetrodon) diversify in species. Land flora dominated by seed ferns. Ma 340 Diversification of amphibians. Ma 305 Earliest diapsid reptiles (e. 251. 225 Ma Earliest dinosaurs (prosauropods).Timeline of evolution 86 420 Earliest ray-finned fishes. minor vertebrate extinctions occur .4 of power" in the oceans shifts dramatically as some groups of prey adapt more rapidly and effectively than others. Ma 330 First amniote vertebrates (Paleothyris). Ma 410 First signs of teeth in fish. bennettitaleans. Ma sharks swam the oceans as top predators. Earliest nautiid nautiloids. Ma 363 By the start of the Carboniferous Period. the "balance 251. Ma 350 First large sharks. and hagfish. This "clearing of the slate" may have led to an ensuing diversification. ratfishes. mites. verticordiid. 68 Ma Tyrannosaurus. 165 Ma First rays and glycymeridid bivalves. or may be a more recent phenomenon. along with triconodontid and symmetrodont mammals. irregular echinoids. Extensive development of sponge reefs. and cladotherian mammals. Diversity in stegosaurian and theropod dinosaurs. and thyasirid bivalves. appears in the fossil record. 106 Ma Spinosaurus. Earliest examples of ticks. Cynodonts become extinct while sauropod dinosaurs diversify. First Coelophysoid dinosaurs 200 Ma The first accepted evidence for viruses (at least. Earliest limopsid. monocots. First freshwater pelomedusid turtles. ornithischian dinosaurs: heterodontosaurids. 176 Ma First members of the Stegosauria group of dinosaurs 170 Ma Earliest salamanders. toothed diving birds. the largest theropod dinosaur. Earliest snakes and nuculanid bivalves. appears in the fossil record. 100 Ma Earliest bees. 161 Ma Ceratopsian dinosaurs appear in the fossil record (Yinlong) 155 Ma First blood-sucking insects (ceratopogonids). 80 Ma First ants and termites. 130 Ma The rise of the Angiosperms: These flowering plants boast structures that attract insects and other animals to spread pollen. and cheilosome bryozoans. the group Geminiviridae) exists. 70 Ma Multituberculate mammals increase in diversity. among the earliest dinosaurs.. and tubulipore bryozoans. and scelidosaurids. herbivores grow to huge sizes in order to accommodate the large guts necessary to digest the nutrient-poor plants. the largest terrestrial predator of North America appears in the fossil record. Archaeopteryx. corbulid bivalves. First yoldiid bivalves. rudist bivalves. First species of Triceratops. 190 Ma Pliosaurs appear in the fossil record. Earliest examples of Ankylosaurian dinosaurs 195 Ma First pterosaurs with specialized feeding (Dorygnathus). Large diversification in angiosperms: magnoliids. and ginger. cryptoclidid & elasmosaurid plesiosaurs. Diversification in small. . First sauropod dinosaurs. fabrosaurids. hermit crabs. 115 Ma First monotreme mammals. This innovation causes a major burst of animal evolution through co-evolution. a possible ancestor to the birds. 110 Ma First hesperornithes. modern starfish. Gymnosperm forests dominate the land. First lepidopteran insects (Archaeolepis).[37] Viruses are still poorly understood and may have arisen before "life" itself. hamamelidids. Major extinctions in terrestrial vertebrates and large amphibians. appeared in the fossil record 230 million years ago. 90 Ma Extinction of ichthyosaurs. newts.Timeline of evolution 87 220 Ma Eoraptor. rosids. first flies and turtles (Odontochelys). brontotheres. and the first eagles and hawks. appeared in the fossil record. The first short-faced bears (Arctodus) appear. Stupendemys appears in the fossil record as the largest freshwater turtle. Ardipithecus) 5 Ma First tree sloths and hippopotami. Extinction of Gastornis. the largest terrestrial mammal that ever lived. 40 Ma Modern type butterflies and moths appear. swifts. is alive. along with mammals becoming the dominant species. opossums. and amphibians. 6. carnivorous and lipotyphlan mammals. burrowing rodents.these species are unrelated to modern-type felines 35 Ma Grasses evolve from among the angiosperms.Timeline of evolution Cenozoic Era 65.5 Ma First hominin (Sahelanthropus). The ancestors of the carnivorous mammals (miacids) were alive. loons.5 The Cretaceous–Tertiary extinction event eradicates about half of all animal species. most planktic foraminifers. edentates. armadillos. and small carnivores. Slight increase in diversity of cold-tolerant ostracods and foraminifers. first bovids and kangaroos. 56 Ma Gastornis. Many modern mammal groups begin to appear: first glyptodonts. diversity in Australian megafauna. The ancestor (according to theory) of the species in Carcharodon. birds. appearance of sirenians. tapirs. parrots. diversification of grazing herbivores.7 Ma Evolution of Paranthropus 2. dogs. peccaries. becoming an apex predator at the time. pterosaurs. 4 Ma Evolution of Australopithecus. first whale (Himalayacetus). flightless bird appears in the fossil record. Rapid diversification in ants. 25 Ma First deer. large carnivorous mammals. saber-toothed cats.5 Ma The earliest species of Smilodon evolve . tapirs. flightless birds. earliest pigs and cats. extinction of embrithopod and brontothere mammals. increase in diversity of anomalodesmatan and heteroconch bivalves. perissodactyl and artiodactyl mammals in the fossil record. kangaroos. and vampire bats traveled to North America while horses. woodpeckers). earliest rodents. 52 Ma First bats appear (Onychonycteris). rudist and inoceramid bivalves. and owls. an important group of carnivorous mammals. 63 Ma Evolution of the creodonts. decrease in the number of perissodactyl mammals. Ma ammonites. 55 Ma Modern bird groups diversify (first song birds. and camels appear in the fossil record. diversification of primates. 28 Ma Paraceratherium appears in the fossil record. and deer entered South America. 50 Ma Peak diversity of dinoflagellates and nanofossils. proboscideans. increase in bird diversity. 33 Ma Evolution of the thylacinid marsupials (Badjcinus). vultures increase in size. plesiosaurs. diversity in insects. 10 Ma Grasslands and savannas are established.5 Ma – present Date Event 88 65. First psammobiid 65 Ma bivalves. 37 Ma First Nimravid carnivores ("False Saber-toothed Cats") . Armadillos. Earliest true primates. the early mako shark Isurus hastalis.8 Ma Mammoths appear in the fossil record. grasslands begin to expand. horses increase in body size and develop high-crowned teeth. ground sloths. major diversification in grassland mammals and snakes. Basilosaurus. reptiles. 20 Ma First giraffes and giant anteaters. a large. along with the first semelid bivalves. where various land and freshwater faunas migrated between North and South America. Extinction of Nimravid carnivores 4. 6 Ma Australopithecines diversify (Orrorin. 30 Ma First balanids and eucalypts. 2. belemnites. one of the first of the giant whales. especially ants and termites. and all of the dinosaurs excluding their descendants [38] the birds From Rapid dominance of conifers and ginkgos in high latitudes. Diversity in toothed and baleen whales. along with major extinctions of gastropods. hummingbirds. 60 Ma Diversification of large. rhinoceroses. lagomorphs. including mosasaurs. 15 Ma Mammut appears in the fossil record. 3 Ma The Great American Interchange. Angiosperms diversify. All Equidae become extinct in North America 10 ka The Holocene Epoch starts 10. for a list of ancestors common to humans and other living species References [1] Planetary Science Institute page (http:/ / www. psi." Making the Moon (http:/ / saturn. The eventual ancestor of cattle. 40 ka The last of the giant monitor lizards (Megalania) die out 30 ka Extinction of Neanderthals 15 ka The last Woolly rhinoceros (Coelodonta) are believed to have gone extinct 11 ka The giant short-faced bears (Arctodus) vanish from North America.000 years before present they start colonising the other continents. replacing the Neanderthals in Europe and other hominins in Asia. Hartmann and Davis belonged to the PSI. the last member of the family Thylacinidae See also • • • • • • • • • Evolutionary history of life Evolutionary history of plants Extinction events Geologic time scale History of Earth Natural history Sociocultural evolution Timeline of human evolution Timeline of plant evolution Further reading • The Ancestor's Tale by Richard Dawkins. sometime in the first 700 million years of its existence. (URL accessed on August 7.Timeline of evolution 89 2 Ma First members of the genus Homo appear in the fossil record. html) on the Giant Impact Hypothesis.7 Ma Extinction of australopithecines. [2] "Because the Moon helps stabilize the tilt of the Earth's rotation. htm) (URL ." How the Oceans Formed (http:/ / www.[39] [40] [41] Around 50.2 Ma Evolution of Homo antecessor. and the Earth entered a new phase of development. The last mainland species of Woolly mammoth (Mammuthus primigenius) die out. Without the Moon. This page also contains several paintings of the impact by Hartmann himself.000[42] years ago after the Late Glacial Maximum. edu/ projects/ moon/ moon. oceansonline. a giant relative of the orangutan dies out from Asia 200 ka Anatomically modern humans appear in Africa. seasonal shifts would likely outpace even the most adaptable forms of life. once the Earth cooled sufficiently. astrobio. The last members of Paranthropus die out. 600 ka Evolution of Homo heidelbergensis 350 ka Evolution of Neanderthals 300 ka Gigantopithecus. com/ ocean_form. clouds began to form in the atmosphere. 2010) [3] "However. as does the last Smilodon species 6 ka Small populations of American Mastodon die off in places like Utah and Michigan 4500 The last members of a dwarf race of Woolly Mammoths vanish from Wrangel Island near Alaska ya 383 ya The last wild Aurochs die out 37 ya The Thylacine goes extinct in a Tasmanian zoo. Diversification of conifers in high latitudes. it prevents the Earth from wobbling between climatic extremes. net/ pressrelease/ 50/ making-the-moon) Astrobiology Magazine. 1. with the last Giant Ground Sloths dying out. Bos primigenius evolves in India 1. [11] " Between about 3. Platts PAH_World. Genetic and Molecular Research 2 (4): 366–375.A. doi:10. and Abel. Ford (February. Philosophical Transactions of the Royal Society of London. "Late Proterozoic low-latitude global glaciation: The snowball Earth" (http:/ / www. org/ pdf/ Hoffman_Science1998. [13] Romano AH. queensu. funpecrp. PMC 1578724. A. "Enzyme-catalyzed phosphoryl transfer reactions".004305. [22] Nicholas J. geoscienceworld. vend.cellbi. pdf). PMID 16754612. doi:10.2004. [18] Olson JM (May 2006). Reuters. P. C. paleo.. Queen's University. " When did eukaryotic cells (cells with nuclei and other internal organelles) first evolve? What do we know about how they evolved from earlier life-forms? (http:/ / www. Res. Hewitt.palaeo. Retrieved 2007-03-10. J Peter Gogarten.J. reuters.2008. net/ media/ resources/ 2007/ 06/ 21/ richtimhywelfinal. sciencemag. Cell Biol.. 2005) [12] Carl Woese. PMC 153566. [21] Fedonkin. PMID 1337989. and Cohen.S. Reuters.(also available as PDF (http:/ / www. Retrieved 2007-06-28. still open questions". D. Hoenigsberg. (1996) Evolution of carbohydrate metabolic pathways. no place in the solar system was safe from the huge arsenal of asteroids and comets left over from the formation of the planets. 363 (1504): 2731–43. "Bangiomorpha pubescens n.1016/0303-2647(92)90004-I. pdf) (PDF). F..0041. Retrieved 2008-09-02. P. Full online article (pdf 260 Kb) (http:/ / www. N. "Eukaryotic organisms in Proterozoic oceans" (http:/ / www.. D. PMID 9721097. D.. gov/ nai2005/ abstracts/ 616 . L. pubmedcentral. [20] Knoll. . D. (2003-04-15). Retrieved 2008-08-30.1016/j.070180. (1998-08-28). caltech. Kaufman. msu.. Proceedings of the National Academy of Sciences 100 (8): 4399–4404. Philos. 2009. Confalonieri. A. pubmedcentral.9.F.06.M. Ying Xu & Bernard Labedan: The Last Universal Common Ancestor : emergence. doi:10. PMID 6250450.1098/rstb. "The antiquity of RNA-based evolution".P. pp. org/ web/ 20060907081933/ http:/ / shiva. doc.com. pdf)) [8] Trevors. Photosyn. .. Palaeoecology 232 (232): 114–130. Rev.N. html) 90 . "Traces of Archaebacteria in ancient sediments". "The nature of the last universal ancestor and the root of the tree of life. doi:10.bi. "Study turns back clock on origins of life on Earth" (http:/ / www. Cambridge University Press. constitution and genetic legacy of an elusive forerunner. . 49: 877–919. R. [28] Narbonne.7. B. 88 (2): 109–17. edu/ ~jkirschvink/ pdfs/ firstsnowball. M. [9] Forterre. (2002).1098/rstb. gov/ articlerender. doi:10. [17] Hahn. archive. Department of Geological Sciences and Geological Engineering. [27] Corsetti. Res Microbiol. Trans. Walter (February 1986). Annu. (2004).Timeline of evolution accessed on January 9. sciam. [19] Buick R (August 2008). fcgi?tool=pmcentrez& artid=2606769).030. htm). [15] Doolittle. Biol. M. doi:10. ucmp. snowballearth.1007/s11120-006-9040-5. multicellularity. sp. Retrieved 2007-05-04.P. Joyce. 1999. Pat Haug (1986).10. . (December 2003). "The biotic response to Neoproterozoic Snowball Earth".5 billion years ago. and Labedan. J. 28 (11): 729–39. Pierce. pdf) [24] Kirschvink. T. Biology Direct 2008. "The RNA World".. doi:10.F. ru/ pub/ Fedonkin_2003.2005. B.. [29] The Cambrian Period (http:/ / www.2006. "Photosynthesis in the Archean era". . Uprooting the tree of life (http:/ / web. Olcott. doi:10. montana. Elie. pdf [26] Corsetti. Benachenhou-Lahfa. org/ cgi/ content/ abstract/ 100/ 8/ 4399). Palaeogeography. N. Paleontological Research 7 (1): 9–41. nasa. F. PMID 18468984. Duguet. S. edu/ courses/ mb437_537_2004_fall/ docs/ uprooting. pnas.. the Last Universal Common Ancestor in Gilbert’s RNA world" (http:/ / www. (2006). W. PMID 12110897. USA" (http:/ / www. Science 281 (5381): 1342. Guy (June 2006). Soc. 51–52. JW. PMID 15011140. and Klein.281.49. 2009). edu/ news/ 1998/ december2/ marsunder122. 2005) * The 'PAH World' (http:/ / nai.1146/annurev. Part B 361 (1470): 1023–38. nih. G. nih. Conway T. Palaeoclimatology. br/ gmr/ year2003/ vol4-2/ gmr0070_full_text. gen. Lond. n. ca/ people/ narbonne/ recent_pubs1. html) (URL accessed on January 9. com. A. pdf) Gilbert.1126/science. Javaux. BioSystems 28 (1-3): 15–32. "Evolution without speciation but with selection: LUCA. Awramik.006.1038/319618a0. C. com/ askexpert_question. Julie (May 21. . com. (PDF). gov/ articlerender.1038/418214a. Sci. [16] Nicolas Glansdorff. C. G.J. H. Cambridge.L. J. Bakermans. and the Mesoproterozoic/Neoproterozoic radiation of eukaryotes" (http:/ / paleobiol.. Death Valley. (2006). E. doi:10. Sleep and Zahnle calculate that Earth was probably hit repeatedly by objects up to 500 kilometers across" Geophysicist Sleep: Martian underground may have harbored early life (http:/ / news-service. 3:29. stanford. berkeley. "The Origin and Early Evolution of Animals" (http:/ / geol. P. 147(6-7):448-55 PMID 9084754 [4] [5] [6] [7] [14] Knowles JR (1980). (1992). html). The Proterozoic Biosphere: A Multidisciplinary Study. System Applied Microbiology 7 (Archaebacteria '85 Proceedings): 178–83.. Schrag. Retrieved May 21. .. PMC 2606769.. funpecrp. Nature 418 (6894): 214–21. com/ article/ scienceNews/ idUSTRE54J5PX20090520?pageNumber=1& virtualBrandChannel=0).1843.5381. "The origin of the Metazoa in the light of the Proterozoic fossil record" (http:/ / www. edu/ cambrian/ camb. "When did oxygenic photosynthesis evolve?" (http:/ / www. fcgi?tool=pmcentrez& artid=1578724). In Schopf. doi:10. "Chance and necessity do not explain the origin of life".1073/pnas. Int. br/ gmr/ year2003/ vol4-2/ / pdf/ gmr0070. Scientific American 282 (6): 90–95. org/ cgi/ content/ abstract/ 26/ 3/ 386) [23] Hoffman. PMID 16453059.2517/prpsj. 2000). cfm?articleID=000C32DD-60E1-1C72-9EB7809EC588F2D7& catID=3& topicID=3)" Scientific American.8 billion and 4. [10] Steenhuysen. Nature 319: 618. [25] http:/ / researchpages. pdf). October 21.0730560100. "A Neoproterozoic Snowball Earth" (http:/ / www.1342.: implications for the evolution of sex. Butterfield. gps. F. PMID 15563395. doi:10. doi:10. (March 2003).A. (1992). Jürgen. Biochem. Andrew H. "A complex microbiota from snowball Earth times: Microfossils from the Neoproterozoic Kingston Peak Formation. Halverson. org/ cgi/ content/ full/ 281/ 5381/ 1342?ijkey=48d78da67bab492803c333f50c0dd84fbbef109c).1016/j. PMID 12682298. Previous fossils indicated that animals didn't take this step until 40 million years later. 2009 [40] Alemseged. html) [33] "The oldest fossils reveal evolution of non-vascular plants by the middle to late Ordovician Period (~450–440 Ma) on the basis of fossil spores" Transition of plants to land (http:/ / www.150517. royalsociety. [42] "International Stratigraphic Chart" (http:/ / www. (2002).Sequence of Plant Evolution (http://sci. Lobster-sized.ac.10032. . S.Timeline of evolution [30] The Cambrian Explosion – Timing (http:/ / palaeo.ensemble. edu/ users/ pciesiel/ gly3150/ plant.nz/evolution/plantEvolution. chem. uct. gly. htm) . and Benton. during the successive waves of colonisation of the terrestrial environment. "The Mesozoic Radiation of Birds". bb/ bcs/ bl14apl/ conq.palaeos.1098/rspb. Proceedings of the Royal Society: Biological 275 (1636): 759." Origins of Viruses (http:/ / www. [41] Stoneking. ac. brown.ucmp.from the University of Cambridge Ensemble Project . com/ html/ reports/ earth_sciences/ report-9641. mcb.explore complete phylogenetic tree interactively • A more compact timeline (http://www.talkorigins.ac. doi:10. This means that viruses also probably emerged from the waters with their different hosts. org/ cheu.ac.. Soodyall. & Dyke. doi:10.waikato. doi:10. edu. org/ content/ qq5un1810k7605h5/ fulltext. Mark.iinet. "Hominid cranium from Homo: Description and taxonomy of Homo-323-1976-896". shtml) • Graphical Timeline of evolution (http://www.org/Life_on_Earth/1) . doi:10. Y.1016/S0959-437X(96)80028-1. Coppens.com. . PMC 2596898.1002/ajpa. [39] The Oldest Homo Sapiens: (http:/ / www. PMID 11815945.1146/annurev. elasmo-research..uk/projects/plantsci/timeline) . Retrieved 2009-02-03. htm) [35] "The ancestry of sharks dates back more than 200 million years before the earliest known dinosaur.org/origins/geo_timeline.. shtml) • University of Waikato . ac.URL retrieved May 15. fungi and bacteria/archaea—probably evolved with their hosts in the seas. clas. more specifically green algae. za/ tutorial/ virorig. org/ education/ evolution/ evol_s_predator.org/?page=segments) from Planck Time to the lifespan of the universe • Interactive Plant Evolution Timeline (http://www.The Trace of Life on Earth (http://www.J. edu/ myl/ ct7/ ozone/ ozone_formation. as suggested by certain common biochemical traits" The first land plants (http:/ / scitec.au/~drage/) • Exploring Time (http://exploringtime. html) (URL accessed on January 9. Current Opinion in Genetics & Development 6 (6): 731–6. Geraads." Oldest fossil footprints on land (http:/ / www. [37] "Viruses of nearly all the major classes of organisms—animals. 91 External links • Berkeley Evolution (http://www.org/en/wiki/Evolution_on_Earth) • History of Life on Earth (http://members. bris. innovations-report. Am J Phys Anthropol 117 (2): 103–12.berkeley. D. com/ releases/ 2005/ 02/ 050223122209. ufl.edu) • Tree of Life Web Project (http://tolweb.33. sciencedaily. geologic time and age determination (http:/ / www. International Commission on Stratigraphy. Luis M. htm) [36] Sahney.wikitimescale. PMID 18198148.com) • University of Waikato . uk/ Palaeofiles/ Cambrian/ timing/ timing. plants. html) [34] "The land plants evolved from the algae. html) [32] "The oldest fossils of footprints ever found on land hint that animals may have beaten plants out of the primordial seas.1370. M. "Recovery from the most profound mass extinction of all time" (http:/ / journals. 2005) [38] Chiappe.2007. pdf) (PDF). (2008). Z. "Human evolution and the mitochondrial genome".html) at the TalkOrigins Archive • Palaeos .Sequence of Animal Evolution (http://sci. Introduction to shark evolution. centipede-like or slug like animals such as Protichnites and Climactichnites made the prints wading out of the ocean and scuttling over sand dunes about 530 million years ago. stratigraphy. (2002).010802. pdf). html) [31] Formation of the Ozone Layer (http:/ / jcbmac. Annual Review of Ecology & Systematics 33: 91–124.ecolsys. given that most of the evolution of life on this planet has occurred there. Gareth J.nz/evolution/AnimalEvolution. Himla (1996). uwichill.waikato. the conception that species change over time. and it synthesized a broad range of evidence from animal husbandry. The debate over Darwin's work led to the rapid acceptance of the general concept of evolution. population genetics became integrated with other biological fields. and sudden large mutations (saltationism). Jean-Baptiste Lamarck proposed his theory of the transmutation of species. geology. combined with biogeography and systematics.92 History History of evolutionary thought Evolutionary thought. Alternatives to natural selection suggested during the eclipse of Darwinism included inheritance of acquired characteristics (neo-Lamarckism). an innate drive for change (orthogenesis). Throughout the 1930s and 1940s. biogeography. natural selection. Naturalists began to focus on the variability of species. morphology. the emergence of paleontology with the concept of extinction further undermined the static view of nature. has roots in antiquity. Charles Darwin and Alfred Russel Wallace published a new evolutionary theory that was explained in detail in Darwin's On the Origin of Species (1859). This began to change during the Enlightenment when evolutionary cosmology and the mechanical philosophy spread from the physical sciences to natural history. In 1858. until the 18th century. based on protein sequences and immunological tests. Darwin proposed common descent and a branching tree of life. led to sophisticated mathematical and causal models of evolution. and later incorporating RNA and DNA studies. resulting in a widely applicable theory of evolution that encompassed much of biology—the modern evolutionary synthesis. and embryology. in the ideas of the ancient Greeks. However. Paleontology and comparative anatomy allowed more detailed reconstructions of the history of life. After the rise of molecular genetics in the 1950s. The synthesis of natural selection with Mendelian genetics during the 1920s and 1930s founded the new discipline of population genetics. the first fully formed scientific theory of evolution. Western biological thinking was dominated by essentialism. and Chinese as well as in medieval Islamic science. The theory was based on the idea of natural selection. Following the establishment of evolutionary biology. but the specific mechanism he proposed. the field of molecular evolution developed. . Before that time most biologists argued that other factors were responsible for evolution. studies of mutation and variation in natural populations. In the early 19th century. The gene-centered view of evolution rose to The Tree of Life as depicted by Ernst Haeckel in The Evolution of Man (1879) illustrates the 19th-century view that evolution was a progressive process leading towards man. was not widely accepted until it was revived by developments in biology that occurred during 1920s through the 1940s. Romans. Unlike Lamarck. the belief that every species has essential characteristics that are unalterable. History of evolutionary thought prominence in the 1960s, followed by the neutral theory of molecular evolution, sparking debates over adaptationism, the units of selection, and the relative importance of genetic drift versus natural selection. In the late 20th century, DNA sequencing led to molecular phylogenetics and the reorganization of the tree of life into the three-domain system. In addition, the newly recognized factors of symbiogenesis and horizontal gene transfer introduced yet more complexity into evolutionary history. 93 Antiquity Greeks Several ancient Greek philosophers discussed ideas that involved change in living organisms over time. Anaximander (c.610–546 BC) proposed that the first animals lived in water and animals that live on land were generated from them.[1] Empedocles (c. 490–430 BC) wrote of a non-supernatural origin for living things,[2] suggesting that adaptation did not require an organizer or final cause. Aristotle summarized his idea: "Wherever then all the parts came about just what they would have been if they had come to be for an end, such things survived, being organized spontaneously in a fitting way; whereas those which grew otherwise perished and continue to perish ..." although Aristotle himself rejected this view.[3] Plato (c. 428–348 BC) was, in the words of biologist and historian Ernst Mayr, "the great antihero of evolutionism",[4] as he established the philosophy of essentialism, which he called the Theory of Forms. This theory holds that objects observed in the real world are only reflections of a limited number of essences (eide). Variation merely results from an imperfect reflection of these constant essences. In his Timaeus, Plato set forth the idea that the Demiurge had created the cosmos and everything in it because, being good, and hence, "... free from jealousy, He desired that all things should be as like Himself as they could be". The creator created all conceivable forms of life, since "... without them the universe will be incomplete, for it will not contain every kind of animal which it ought to contain, if it is to be perfect". This "plenitude principle"—the idea that all potential forms of life are essential to a perfect creation—greatly influenced Christian thought.[5] Plato (left) and Aristotle (right), a detail of The School of Athens Aristotle (384–322 BC), one of the most influential of the Greek philosophers, is the earliest natural historian whose work has been preserved in any real detail. His writings on biology resulted from his research into natural history on and around the isle of Lesbos, and have survived in the form of four books, usually known by their Latin names, De anima (on the essence of life), Historia animalium (inquiries about animals), De generatione animalium (reproduction), and De partibus animalium (anatomy). Aristotle's works contain some remarkably astute observations and interpretations—along with sundry myths and mistakes—reflecting the uneven state of knowledge during his time.[6] However, for Charles Singer, "Nothing is more remarkable than [Aristotle's] efforts to [exhibit] the relationships of living things as a scala naturæ."[6] This scala naturæ, described in Historia animalium, classified organisms in relation to a hierarchical "Ladder of Life" or "Chain of Being", placing them according to their complexity of structure and function, with organisms that showed greater vitality and ability to move described as "higher organisms".[5] History of evolutionary thought 94 Chinese Ideas on changing biologic species were expressed by ancient Chinese thinkers such as Zhuangzi (Chuang Tzu), a Taoist philosopher who lived around the 4th century BC. According to Joseph Needham, Taoism explicitly denies the fixity of biological species and Taoist philosophers speculated that species had developed differing attributes in response to differing environments.[7] Humans, nature and the heavens are seen as existing in a state of "constant transformation" known as the Tao, in contrast with the more static view of nature typical of Western thought.[8] Romans Titus Lucretius Carus (d. 50 BC), the Roman philosopher and atomist, wrote the poem On the Nature of Things (De rerum natura), which provides the best surviving explanation of the ideas of the Greek Epicurean philosophers. It describes the development of the cosmos, the Earth, living things, and human society through purely naturalistic mechanisms, without any reference to supernatural involvement. On the Nature of things would influence the cosmological and evolutionary speculations of philosophers and scientists during and after the Renaissance.[9] [10] Augustine of Hippo In line with earlier Greek thought, the 4th century bishop and theologian, St. Augustine of Hippo, wrote that the creation story in Genesis should not be read too literally. In his book De Genesi ad litteram ("On The Literal Interpretation of Genesis"), he stated that in some cases new creatures may have come about through the "decomposition" of earlier forms of life.[11] For Augustine, "plant, fowl and animal life are not perfect ... but created in a state of potentiality", unlike what he considered the theologically perfect forms of angels, the firmament and the human soul.[12] Augustine's idea 'that forms of life had been transformed "slowly over time"' prompted Father Giuseppe Tanzella-Nitti, Professor of Theology at the Pontifical Santa Croce University in Rome, to claim that Augustine had suggested a form of evolution.[13] [14] Middle Ages Islamic philosophy and the struggle for existence Although Greek and Roman evolutionary ideas died out in Europe after the fall of the Roman Empire, they were not lost to Islamic philosophers and scientists. In the Islamic Golden Age of the 8th to the 13th centuries, philosophers explored ideas about natural history. These ideas included transmutation from non-living to living: "from mineral to plant, from plant to animal, and from animal to man".[15] The first Muslim biologist and philosopher to publish detailed speculations about natural history, the Afro-Arab writer al-Jahiz, wrote in the 9th century. In the Book of Animals, he considered the effects of the environment on an animal's chances for survival, and described the struggle for existence.[16] Al-Jahiz also wrote descriptions of food chains.[17] Al-Jahiz speculated on the influence of the environment on animals and considered the effects of the environment on the likelihood of an animal to survive. For example, Al-Jahiz's wrote in his Book of Animals: "All animals, in short, can not exist without food, neither can the hunting animal escape being hunted in his turn. Every weak animal devours those weaker than itself. Strong animals cannot escape being devoured by other animals stronger than they."[16] History of evolutionary thought 95 Christian philosophy and the great chain of being During the Early Middle Ages, Greek classical learning was all but lost to the West. However, contact with the Islamic world, where Greek manuscripts were preserved and expanded, soon led to a massive spate of Latin translations in the 12th century. Europeans were re-introduced to the works of Plato and Aristotle, as well as to Islamic thought. Christian thinkers of the scholastic school, in particular Abelard and Thomas Aquinas, combined Aristotelian classification with Plato's ideas of the goodness of God, and of all potential life forms being present in a perfect creation, to organize all inanimate, animate, and spiritual beings into a huge interconnected system: the scala naturæ, or great chain of being.[5] [18] Within this system, everything that existed could be placed in order, from "lowest" to "highest", with Hell at the bottom and God at the top—below God, an angelic hierarchy marked by the orbits of the planets, mankind in an intermediate position, and worms the lowest of the animals. As the universe was ultimately perfect, the great chain was Drawing of the great chain of being from also perfect. There were no empty links in the chain, and no link was Rhetorica Christiana (1579) by Diego Valades represented by more than one species. Therefore no species could ever move from one position to another. Thus, in this Christianized version of Plato's perfect universe, species could never change, but remained forever fixed, in accordance with the text of Genesis. For humans to forget their position was seen as sinful, whether they behaved like lower animals or aspired to a higher station than was given them by their Creator.[5] Creatures on adjacent steps were expected to closely resemble each other, an idea expressed in the saying: natura non facit saltum ("nature does not make leaps").[5] This basic concept of the great chain of being greatly influenced the thinking of Western civilization for centuries (and still has an influence today). It formed a part of the argument from design presented by natural theology. As a classification system, it became the major organizing principle and foundation of the emerging science of biology in the 17th and 18th centuries.[5] Thomas Aquinas on creation and natural processes While the development of the great chain of being and the argument from design by Christian theologians contributed to the view that the natural world fit into an unchanging designed hierarchy, some theologians were more open to the possibility that the world might have developed through natural processes. Thomas Aquinas went even farther than Augustine of Hippo in arguing that scriptural texts like Genesis should not be interpreted in a literal way that conflicted with or constrained what natural philosophers learned about the workings of the natural world. He felt that the autonomy of nature was a sign of God's goodness and that there was no conflict between the concept of a divinely created universe, and the idea that the universe may have evolved over time through natural mechanisms.[19] However, Aquinas disputed the views of those like the ancient Greek philosopher Empodocles who held that such natural processes showed that the universe could have developed without an underlying purpose. Rather holding that: "Hence, it is clear that nature is nothing but a certain kind of art, i.e., the divine art, impressed upon things, by which these things are moved to a determinate end. It is as if the shipbuilder were able to give to timbers that by which they would move themselves to take the form of a ship."[20] History of evolutionary thought 96 Renaissance and Enlightenment In the first half of the 17th century, René Descartes's mechanical philosophy encouraged the use of the metaphor of the universe as a machine, a concept that would come to characterise the scientific revolution.[21] Between 1650 and 1800, some evolutionist theories supported the view that the universe, including life on Earth, had developed mechanically, entirely without divine guidance. In contrast, most contemporary theories of evolution, such of those of Gottfried Leibniz and J. G. Herder, held that evolution was a fundamentally spiritual process.[22] In 1751, Pierre Louis Maupertuis veered toward more materialist ground. He wrote of natural modifications occurring during reproduction and accumulating over the course of many generations, producing races and even new species, a description that anticipated in general terms the concept of natural selection.[23] Pierre Belon compared the skeletons of birds and humans in his Book of Birds (1555). Later in the 18th century, the French philosopher G. L. L. Buffon, one of the leading 18th century naturalists, suggested that what most people referred to as species were really just well-marked varieties, modified from an original form by environmental factors. For example, he believed that lions, tigers, leopards and house cats might all have a common ancestor. He further speculated that the 200 or so species of mammals then known might have descended from as few as 38 original animal forms. Buffon's evolutionary ideas were limited; he believed each of the original forms had arisen through spontaneous generation and that each was shaped by "internal moulds" that limited the amount of change. Buffon's works, Natural History and The Epochs of Nature, containing well developed theories about a completely materialistic origin for the Earth and his ideas questioning the fixity of species, were extremely influential.[24] [25] Between 1767 and 1792, James Burnett, Lord Monboddo included in his writings not only the concept that man had descended from primates, but also that, in response to the environment, creatures had found methods of transforming their characteristics over long time intervals.[26] Charles Darwin's grandfather, Erasmus Darwin, published Zoönomia in 1796, which suggested that "all warm-blooded animals have arisen from one living filament".[27] In his 1802 poem Temple of Nature, he described the rise of life from minute organisms living in mud to all of its modern diversity.[28] History of evolutionary thought 97 Early 19th century Paleontology and geology In 1796, Georges Cuvier published his findings on the differences between living elephants and those found in the fossil record. His analysis demonstrated that mammoths and mastodons were distinct species, different from any living animal, effectively ending a long-running debate over whether the extinction of a species was possible.[29] In 1788, James Hutton described gradual geological processes operating continuously over deep time.[30] In the 1790s William Smith began the process of ordering rock strata by examining fossils in the layers while he worked on his geologic map of England. Independently, in 1811, Georges Cuvier and Alexandre Brongniart published an influential study of the geologic history of the region around Paris, based on the stratigraphic succession of rock layers. These works helped establish the antiquity of the Earth.[31] Cuvier advocated catastrophism to explain the patterns of extinction and faunal succession revealed by the fossil record. Knowledge of the fossil record continued to advance rapidly during the first few decades of the 19th century. By the 1840s, the outlines of the Diagram of the geologic timescale from an 1861 book by Richard Owen showing the appearance geologic timescale were becoming clear, and in 1841 John Phillips of major animal types named three major eras, based on the predominant fauna of each: the Paleozoic, dominated by marine invertebrates and fish, the Mesozoic, the age of reptiles, and the current Cenozoic age of mammals. This progressive picture of the history of life was accepted even by conservative English geologists like Adam Sedgwick and William Buckland; however, like Cuvier, they attributed the progression to repeated catastrophic episodes of extinction followed by new episodes of creation.[32] Unlike Cuvier, Buckland and some other advocates of natural theology among British geologists made efforts to explicitly link the last catastrophic episode proposed by Cuvier to the biblical flood.[33] [34] From 1830 to 1833, Charles Lyell published his multi-volume work Principles of Geology, which, building on Hutton's ideas, advocated a uniformitarian alternative to the catastrophic theory of geology. Lyell claimed that, rather than being the products of cataclysmic (and possibly supernatural) events, the geologic features of the Earth are better explained as the result of the same gradual geologic forces observable in the present day—but acting over immensely long periods of time. Although Lyell opposed evolutionary ideas (even questioning the consensus that the fossil record demonstrates a true progression), his concept that the Earth was shaped by forces working gradually over an extended period, and the immense age of the Earth assumed by his theories, would strongly influence future evolutionary thinkers such as Charles Darwin.[35] History of evolutionary thought 98 Transmutation of species Jean-Baptiste Lamarck proposed, in his Philosophie Zoologique of 1809, a theory of the transmutation of species. Lamarck did not believe that all living things shared a common ancestor but rather that simple forms of life were created continuously by spontaneous generation. He also believed that an innate life force drove species to become more complex over time, advancing up a linear ladder of complexity that was related to the great chain of being. Lamarck recognized that species were adapted to their environment. He explained this by saying that the same innate force driving increasing complexity caused the organs of an animal (or a plant) to change based on the use or disuse of those organs, just as muscles are affected by exercise. He argued that these changes would be inherited by the next generation and produce slow adaptation to the environment. It was this secondary mechanism of adaptation through the inheritance of acquired characteristics that would become known as Lamarckism and would influence discussions of evolution into the 20th century.[36] [37] A radical British school of comparative anatomy that included the anatomist Robert Grant was closely in touch with Lamarck's French school of Transformationism. One of the French scientists who Diagram from Vestiges of the Natural History of influenced Grant was the anatomist Étienne Geoffroy Saint-Hilaire, Creation (1844) by Robert Chambers shows a whose ideas on the unity of various animal body plans and the model of development where fish (F), reptiles (R), and birds (B) represent branches from a path homology of certain anatomical structures would be widely influential leading to mammals (M). and lead to intense debate with his colleague Georges Cuvier. Grant became an authority on the anatomy and reproduction of marine invertebrates. He developed Lamarck's and Erasmus Darwin's ideas of transmutation and evolutionism, and investigated homology, even proposing that plants and animals had a common evolutionary starting point. As a young student Charles Darwin joined Grant in investigations of the life cycle of marine animals. In 1826 an anonymous paper, probably written by Robert Jameson, praised Lamarck for explaining how higher animals had "evolved" from the simplest worms; this was the first use of the word "evolved" in a modern sense.[38] [39] In 1844, the Scottish publisher Robert Chambers anonymously published an extremely controversial but widely read book entitled Vestiges of the Natural History of Creation. This book proposed an evolutionary scenario for the origins of the Solar System and life on Earth. It claimed that the fossil record showed a progressive ascent of animals with current animals being branches off a main line that leads progressively to humanity. It implied that the transmutations lead to the unfolding of a preordained plan that had been woven into the laws that governed the universe. In this sense it was less completely materialistic than the ideas of radicals like Robert Grant, but its implication that humans were only the last step in the ascent of animal life incensed many conservative thinkers. The high profile of the public debate over Vestiges, with its depiction of evolution as a progressive process, would greatly influence the perception of Darwin's theory a decade later.[40] [41] Ideas about the transmutation of species were associated with the radical materialism of the Enlightenment and were attacked by more conservative thinkers. Georges Cuvier attacked the ideas of Lamarck and Geoffroy Saint-Hilaire, agreeing with Aristotle that species were immutable. Cuvier believed that the individual parts of an animal were too closely correlated with one another to allow for one part of the anatomy to change in isolation from the others, and argued that the fossil record showed patterns of catastrophic extinctions followed by re-population, rather than gradual change over time. He also noted that drawings of animals and animal mummies from Egypt, which were History of evolutionary thought thousands of years old, showed no signs of change when compared with modern animals. The strength of Cuvier's arguments and his scientific reputation helped keep transmutational ideas out of the mainstream for decades.[42] In Britain the philosophy of natural theology remained influential. William Paley's 1802 book Natural Theology with its famous watchmaker analogy had been written at least in part as a response to the transmutational ideas of Erasmus Darwin.[43] Geologists influenced by natural theology, such as Buckland and Sedgwick, made a regular practice of attacking the evolutionary ideas of Lamarck, Grant, and The This 1847 diagram by Richard Owen shows his Vestiges of the Natural History of Creation.[44] [45] Although the conceptual archetype for all vertebrates. geologist Charles Lyell opposed scriptural geology, he also believed in the immutability of species, and in his Principles of Geology (1830–1833), he criticized Lamarck's theories of development.[35] Idealists such as Louis Agassiz and Richard Owen believed that each species was fixed and unchangeable because it represented an idea in the mind of the creator. They believed that relationships between species could be discerned from developmental patterns in embryology, as well as in the fossil record, but that these relationships represented an underlying pattern of divine thought, with progressive creation leading to increasing complexity and culminating in humanity. Owen developed the idea of "archetypes" in the Divine mind that would produce a sequence of species related by anatomical homologies, such as vertebrate limbs. Owen led a public campaign that successfully marginalized Robert Grant in the scientific community. Darwin would make good use of the homologies analyzed by Owen in his own theory, but the harsh treatment of Grant, and the controversy surrounding Vestiges, showed him the need to ensure that his own ideas were scientifically sound.[39] [46] [47] 99 Anticipations of natural selection Several writers anticipated aspects of Darwin's theory, and in the third edition of On the Origin of Species published in 1861 Darwin named those he knew about in an introductory appendix, An Historical Sketch of the Recent Progress of Opinion on the Origin of Species, which he expanded in later editions.[48] In 1813, William Charles Wells read before the Royal Society essays assuming that there had been evolution of humans, and recognising the principle of natural selection. Charles Darwin and Alfred Russel Wallace were unaware of this work when they jointly published the theory in 1858, but Darwin later acknowledged that Wells had recognised the principle before them, writing that the paper "An Account of a White Female, part of whose Skin resembles that of a Negro" was published in 1818, and "he distinctly recognises the principle of natural selection, and this is the first recognition which has been indicated; but he applies it only to the races of man, and to certain characters alone."[49] When Darwin was developing his theory, he was influenced by Augustin de Candolle's natural system of classification, which laid emphasis on the war between competing species.[50] [51] Patrick Matthew wrote in the obscure book Naval Timber & Arboriculture (1831) of "continual balancing of life to circumstance. ... [The] progeny of the same parents, under great differences of circumstance, might, in several generations, even become distinct species, incapable of co-reproduction."[52] Charles Darwin discovered this work after the initial publication of the Origin. In the brief historical sketch that Darwin included in the 3rd edition he says "Unfortunately the view was given by Mr. Matthew very briefly in an Appendix to a work on a different subject ... He clearly saw, however, the full force of the principle of natural selection."[53] It is possible to look through the history of biology from the ancient Greeks onwards and discover anticipations of almost all of Darwin's key ideas. However, as historian of science Peter J. Bowler says, "Through a combination of bold theorizing and comprehensive evaluation, Darwin came up with a concept of evolution that was unique for the time." Bowler goes on to say that simple priority alone is not enough to secure a place in the history of science; someone has to develop an idea and convince others of its importance to have a real impact.[54] T. H. Huxley said in his essay on the reception of the Origin of Species: History of evolutionary thought The suggestion that new species may result from the selective action of external conditions upon the variations from their specific type which individuals present and which we call spontaneous because we are ignorant of their causation is as wholly unknown to the historian of scientific ideas as it was to biological specialists before 1858. But that suggestion is the central idea of the Origin of Species, and contains the quintessence of Darwinism.[55] 100 Natural selection The biogeographical patterns Charles Darwin observed in places such as the Galapagos islands during the voyage of the Beagle caused him to doubt the fixity of species, and in 1837 Darwin started the first of a series of secret notebooks on transmutation. Darwin's observations led him to view transmutation as a process of divergence and branching, rather than the ladder-like progression envisioned by Lamarck and others. In 1838 he read the new 6th edition of An Essay on the Principle of Population, written in the late 18th century by Thomas Malthus. Malthus' idea of population growth leading to a struggle for survival combined with Darwin's knowledge on how breeders selected traits, led to the inception of Darwin's theory of natural Darwin's first sketch of an evolutionary tree from his First Notebook selection. Darwin did not publish his ideas on evolution on Transmutation of Species (1837) for 20 years. However he did share them with certain other naturalists and friends, starting with Joseph Hooker, with whom he discussed his unpublished 1844 essay on natural selection. During this period he used the time he could spare from his other scientific work to slowly refine his ideas and, aware of the intense controversy around transmutation, amass evidence to support them. In September 1854 he began full time work on writing his book on natural selection.[47] [56] [57] Unlike Darwin, Alfred Russel Wallace, influenced by the book Vestiges of the Natural History of Creation, already suspected that transmutation of species occurred when he began his career as a naturalist. By 1855 his biogeographical observations during his field work in South America and the Malay Archipelago made him confident enough in a branching pattern of evolution to publish a paper stating that every species originated in close proximity to an already existing closely allied species. Like Darwin, it was Wallace's consideration of how the ideas of Malthus might apply to animal populations that led him to conclusions very similar to those reached by Darwin about the role of natural selection. In February 1858 Wallace, unaware of Darwin's unpublished ideas, composed his thoughts into an essay and mailed them to Darwin, asking for his opinion. The result was the joint publication in July of an extract from Darwin's 1844 essay along with Wallace's letter. Darwin also began work on a short abstract summarising his theory, which he would publish in 1859 as On the Origin of Species.[58] History of evolutionary thought 101 1859–1930s: Darwin and his legacy By the 1850s whether or not species evolved was a subject of intense debate, with prominent scientists arguing both sides of the issue.[59] However, it was the publication of Charles Darwin's On the Origin of Species (1859) that fundamentally transformed the discussion over biological origins.[60] Darwin argued that his branching version of evolution explained a wealth of facts in biogeography, anatomy, embryology, and other fields of biology. He also provided the first cogent mechanism by which evolutionary change could persist: his theory of natural selection.[61] One of the first and most important naturalists to be convinced by Origin of the reality of evolution was the British anatomist Thomas Henry Huxley. Huxley recognized that unlike the earlier transmutational ideas of Lamarck and Vestiges, Darwin's theory provided a mechanism for evolution without supernatural involvement, even if Huxley himself was not completely convinced that natural selection was the key evolutionary mechanism. Huxley would make advocacy of evolution a Diagram by O.C. Marsh of the evolution of horse feet cornerstone of the program of the X Club to reform and and teeth over time as reproduced in T.H Huxley's 1876 book Professor Huxley in America professionalise science by displacing natural theology with naturalism and to end the domination of British natural science by the clergy. By the early 1870s in English-speaking countries, thanks partly to these efforts, evolution had become the mainstream scientific explanation for the origin of species.[61] In his campaign for public and scientific acceptance of Darwin's theory, Huxley made extensive use of new evidence for evolution from paleontology. This included evidence that birds had evolved from reptiles, including the discovery of Archaeopteryx in Europe, and a number of fossils of primitive birds with teeth found in North America. Another important line of evidence was the finding of fossils that helped trace the evolution of the horse from its small five-toed ancestors.[62] However, acceptance of evolution among scientists in non-English speaking nations such as France, and the countries of southern Europe and Latin America was slower. An exception to this was Germany, where both August Weismann and Ernst Haeckel championed this idea: Haeckel used evolution to challenge the established tradition of metaphysical idealism in German biology, much as Huxley used it to challenge natural theology in Britain.[63] Haeckel and other German scientists would take the lead in launching an ambitious programme to reconstruct the evolutionary history of life based on morphology and embryology.[64] Darwin's theory succeeded in profoundly altering scientific opinion regarding the development of life and in producing a small philosophical revolution.[65] However, this theory could not explain several critical components of the evolutionary process. Specifically, Darwin was unable to explain the source of variation in traits within a species, and could not identify a mechanism that could pass traits faithfully from one generation to the next. Darwin's hypothesis of pangenesis, while relying in part on the inheritance of acquired characteristics, proved to be useful for statistical models of evolution that were developed by his cousin Francis Galton and the "biometric" school of evolutionary thought. However, this idea proved to be of little use to other biologists.[66] They agreed that humans shared a common ancestor with apes. to modern humans for them to be convincing intermediates between humans and other primates. the scientific community believed that. a series of archaeological discoveries in the 1840s and 1850s showed stone tools associated with the remains of extinct animals. On the other hand. especially in the critical characteristic of cranial capacity. He argued that all the differences between humans and apes were explained by a combination of the selective pressures that came from our ancestors moving from the trees to the plains. For most of the first half of the 19th century.[67] Therefore the debate that immediately followed the publication of The Origin of Species centered on the similarities and differences between humans and modern apes. This illustration was the frontispiece of Thomas Henry Huxley's although geology had shown that the Earth and life book Evidence as to Man's Place in Nature (1863). Thomas Henry Huxley sought to demonstrate a close anatomical relationship between humans and apes. The only human fossils found before the discovery of Java man in the 1890s were either of anatomically modern humans or of Neanderthals that were too close.[67] In 1871. but questioned whether any purely materialistic mechanism could account for all the differences between humans and apes. Another viewpoint was advocated by Charles Lyell and Alfred Russel Wallace. especially some aspects of the human mind. For example. which contained his views on human evolution. at that time there was no fossil evidence to demonstrate human evolution. he viewed morality as a natural outgrowth of instincts that were beneficial to animals living in social groups. and Selection in Relation to Sex. Despite this precaution. were very old. The debate over human origins. and over the degree of human uniqueness continued well into the 20th century. This view of human history was more compatible with an evolutionary origin for humanity than was the older view. However. Darwin argued that the differences between the human mind and the minds of the higher animals were a matter of degree rather than of kind. Huxley summarized his argument in his highly influential 1863 book Evidence as to Man's Place in Nature. Therefore he almost completely ignored the topic of human evolution in The Origin of Species. and sexual selection. In one famous incident. On the other hand. it had become widely accepted that humans had existed during a prehistoric period – which stretched many thousands of years before the start of written history.[68] Richard Owen vigorously defended the classification suggested by Cuvier and Johann Friedrich Blumenbach that placed humans in a separate order from any of the other mammals. the issue featured prominently in the debate that followed the book's publication. which by the early 19th century had become the orthodox view. human beings had appeared suddenly just a few thousand years before the present. Huxley showed that Owen was mistaken in claiming that the brains of gorillas lacked a structure present in human brains.[67] . Darwin published The Descent of Man. as summarized in Charles Lyell's 1863 book Geological Evidences of the Antiquity of Man. Carolus Linnaeus had been criticised in the 18th century for grouping humans and apes together as primates in his ground breaking classification system.History of evolutionary thought 102 Application to humans Charles Darwin was aware of the severe reaction in some parts of the scientific community against the suggestion made in Vestiges of the Natural History of Creation that humans had arisen from animals by a process of transmutation. By the early 1860s. William Bateson. and would remain the position of some naturalists well into the 20th century. this idea gradually fell out of favor among scientists.[69] [70] In the late 19th century. They considered Lamarckism to be philosophically superior to Darwin's idea of selection acting on random variation. the term neo-Lamarckism came to be associated with the position of naturalists who viewed the inheritance of acquired characteristics as the most important evolutionary mechanism. but the acceptance of natural selection as its driving mechanism was much less widespread. orthogenesis. . which Osborn claimed was an example of an orthogenic trend in evolution. Morgan. Saltationism was the idea that new species arise as a result of large mutations. Inheritance of acquired characteristics was part of Haeckel's recapitulation theory of evolution. in a unilinear fashion. and was popular with early geneticists such as Hugo de Vries. although it retained a strong popular following. Despite these criticisms. neo-Lamarckism remained the most popular alternative to natural selection at the end of the 19th century. and saltationism. and thought he found. However. such as the German biologist August Weismann and Alfred Russel Wallace. Orthogenesis was popular among some paleontologists.[69] [70] This photo from Henry Fairfield Osborn's 1918 book Origin and Evolution of Life shows models depicting the evolution of Titanothere horns over time.[69] [70] Orthogenesis was the hypothesis that life has an innate tendency to change. and the American paleontologist Edward Drinker Cope. theistic evolution had largely disappeared from professional scientific discussions. who believed that the fossil record showed a gradual and constant unidirectional change. neo-Lamarckism. It was seen as a much faster alternative to the Darwinian concept of a gradual process of small random variations being acted on by natural selection. the German biologist Ernst Haeckel. Cope looked for. H. pointed out that no one had ever produced solid evidence for the inheritance of acquired characteristics. as they became more and more committed to the idea of methodological naturalism and came to believe that direct appeals to supernatural involvement were scientifically unproductive. towards ever-greater perfection. and its proponents included the Russian biologist Leo Berg and the American paleontologist Henry Fairfield Osborn. By 1900. and early in his career. which held that the embryological development of an organism repeats its evolutionary history. T.History of evolutionary thought 103 Alternatives to natural selection The concept of evolution was widely accepted in scientific circles within a few years of the publication of Origin. patterns of linear progression in the fossil record. It became the basis of the mutation theory of evolution.[69] [70] Critics of neo-Lamarckism. Advocates of this position included the British writer and Darwin critic Samuel Butler. The four major alternatives to natural selection in the late 19th century were theistic evolution. It had a significant following in the 19th century. Theistic evolution (a term promoted by Darwin's greatest American advocate Asa Gray) was the idea that God intervened in the process of evolution to guide it in such a way that the living world could still be considered to be designed. History of evolutionary thought 104 Mendelian genetics. who had focused on measurement and statistical analysis Diagram from T. Their opponents were the biometricians. but the Mendelians maintained that the variations measured by biometricians were too insignificant to account for the evolution of new species. such as a change in eye color.[71] [72] 1920s–1940s Biston betularia f. Instead. could not explain the continuous range of variation melanogaster seen in real populations. Morgan began experimenting with breeding the fruit fly Drosophila melanogaster. They were led by William Bateson (who coined the word genetics) and Hugo de Vries (who coined the word mutation). carbonaria is the black-bodied form of the peppered moth. the work at his lab between 1910 and 1915 reconfirmed Mendelian genetics and provided solid experimental evidence linking it to chromosomal inheritance. Their leaders. Weldon's work with crabs and snails provided evidence that selection pressure from the environment could shift the range of variation in wild populations. Karl Pearson and Walter Frank Raphael Weldon. In one camp were the Mendelians. of the white-eyed mutation in Drosophila such as genes.[71] [72] When T. typica is the white-bodied form of the peppered moth. who were interested in the continuous variation of characteristics within populations. and mutation The so-called rediscovery of Gregor Mendel's laws of inheritance in 1900 ignited a fierce debate between two camps of biologists. he was a saltationist who hoped to demonstrate that a new species could be created in the lab by mutation alone. mutations served to increase variation within the existing population. Biston betularia f. H. His work also demonstrated that most mutations had relatively small effects. who were focused on discrete variations and the laws of inheritance. showing the sex-linked inheritance Mendelian genetics on the basis that discrete units of heredity. The biometricians rejected Basis of Heredity. followed in the tradition of Francis Galton. Morgan's 1919 book The Physical of variation within a population. . and that rather than creating a new species in a single step.H. biometrics. H. which emphasized the importance of allopatric speciation in the formation of new species.[73] [74] [76] [77] Evolutionary biologist Ernst Mayr was influenced by the work of the German biologist Bernhard Rensch showing the influence of local environmental factors on the geographic distribution of sub-species and closely related species. Mayr also formulated the biological species concept that defined a species as a group of interbreeding or potentially interbreeding populations that were reproductively isolated from all other populations. This integrated natural selection with Mendelian genetics. continued throughout the 1930s and 1940s to demonstrate the power of selection due to ecological factors including the ability to maintain genetic diversity through genetic polymorphisms such as human blood types. these populations had large amounts of genetic diversity.S. But as the field of genetics continued to develop. He helped to bridge the divide between the foundations of microevolution developed by the population geneticists and the patterns of macroevolution observed by field biologists. with marked differences between sub-populations. another British geneticist. This form of speciation occurs when the geographical isolation of a sub-population is followed by the development of mechanisms for reproductive isolation. relatively isolated populations that exhibited genetic drift. In 1932. isolated sub-population away from an adaptive peak. and the effects of inbreeding on small.History of evolutionary thought 105 Population genetics The Mendelian and biometrician models were eventually reconciled with the development of population genetics. resulting in evolution.A. A key step was the work of the British biologist and statistician R. which helped to integrate botany into the synthesis. Mayr followed up on Dobzhansky's work with the 1942 book Systematics and the Origin of Species. George Gaylord Simpson showed that the fossil record was consistent with the irregular non-directional pattern predicted by the developing evolutionary synthesis. and that the linear trends that earlier paleontologists had claimed supported orthogenesis and neo-Lamarckism did not hold up to closer examination.[73] [74] The American biologist Sewall Wright. The work of Fisher. which was the critical first step in developing a unified theory of how evolution worked. Haldane and Wright founded the discipline of population genetics. applied statistical analysis to real-world examples of natural selection.B. Ledyard Stebbins published Variation and Evolution in Plants. had been influenced by the work on genetic diversity by Russian geneticists such as Sergei Chetverikov. with his 1937 book Genetics and the Origin of Species. The emerging cross-disciplinary consensus on the workings of evolution would be known as the modern evolutionary synthesis. contrary to the assumptions of the population geneticists. Wright introduced the concept of an adaptive landscape and argued that genetic drift and inbreeding could drive a small. such as the evolution of industrial melanism in peppered moths. G.B. Morgan's lab. the pioneer of ecological genetics.[73] [74] Modern evolutionary synthesis In the first few decades of the 20th century. and showed that natural selection worked at an even faster rate than Fisher assumed.[75] Theodosius Dobzhansky. Ford's work would contribute to a shift in emphasis during the course of the modern synthesis towards natural selection over genetic drift.[73] [74] . allowing natural selection to drive it towards different adaptive peaks. Haldane. Dobzhansky examined the genetic diversity of wild populations and showed that. In a series of papers beginning in 1924. a postdoctoral worker in T. and that natural selection could change gene frequencies in a population.[73] [74] [78] In the 1944 book Tempo and Mode in Evolution. The book also took the highly mathematical work of the population geneticists and put it into a more accessible form. In Great Britain E. most field naturalists continued to believe that Lamarckian and orthogenic mechanisms of evolution provided the best explanation for the complexity they observed in the living world. In a series of papers starting in 1918 and culminating in his 1930 book The Genetical Theory of Natural Selection. In 1950. Ford. J. focused on combinations of interacting genes. Fisher. who had a background in animal breeding experiments. It received its name from the book Evolution: The Modern Synthesis by Julian Huxley. those views became less tenable. Fisher showed that the continuous variation measured by the biometricians could be produced by the combined action of many discrete genes. thus making sexual reproduction common.[84] [85] Late 20th century Gene-centered view In the mid-1960s. but not all. Motoo Kimura and others provided a theoretical basis for the molecular clock. Leigh Van Valen proposed the term "Red Queen". and macroevolution was simply considered the result of extensive microevolution.[82] Studies of protein differences within species also brought molecular data to bear on population genetics by providing estimates of the level of heterozygosity in natural populations.[87] Models of the period showed that group selection was severely limited in its strength.[89] [90] The gene-centric view has also led to an increased interest in Darwin's old idea of sexual selection. At the same time. Such explanations were largely replaced by a gene-centered view of evolution. natural selection acting on genetic variation was virtually the only acceptable mechanism of evolutionary change (panselectionism). Hamilton. such as protein electrophoresis and sequencing. to protein sequences. Price and John Maynard Smith. D. is responsible for a large portion of genetic change: the neutral theory of molecular evolution. which he took from Through the Looking-Glass by Lewis Carroll. By 1969. biological disciplines. biochemists Linus Pauling and Emile Zuckerkandl proposed the molecular clock hypothesis: that sequence differences between homologous proteins could be used to calculate the time since two species diverged. Hamilton.[88] In 1973. increasingly powerful techniques for analyzing proteins.[86] This viewpoint would be summarized and popularized in the influential 1976 book The Selfish Gene by Richard Dawkins. rather than natural selection. through the genetic code. despite the tremendous cost from the gene-centric point of view of a system where only half of an organism's genome is passed on during reproduction. Simpson. Theodosius Dobzhansky and G. and with it an understanding of the chemical nature of genes as sequences of DNA and their relationship. Established evolutionary biologists—particularly Ernst Mayr. spawning the neutralist-selectionist debate over the relative importance of drift and selection. In the early 1960s. which continued into the 1980s without a clear resolution. natural selection and Mendelian population genetics—that tied together many. to describe a scenario where a species involved in one or more evolutionary arms races would have to constantly change just to keep pace with the species with which it was co-evolving. G. George R. Williams and others suggested that this idea might explain the evolution of sexual reproduction: the increased genetic diversity caused by sexual reproduction would help maintain resistance against rapidly evolving parasites.[91] and more recently in topics such as sexual conflict and intragenomic conflict.[83] From the early 1960s. brought biochemical phenomena into realm of the synthetic theory of evolution. three of the architects of the modern synthesis—were extremely skeptical of molecular approaches. . Williams strongly critiqued explanations of adaptations worded in terms of "survival of the species" (group selection arguments). George C. especially when it came to the connection (or lack thereof) to natural selection. It helped establish the legitimacy of evolutionary biology. a primarily historical science. though newer models do admit the possibility of significant multi-level selection. arguing that—at the molecular level at least—most genetic mutations are neither harmful nor helpful and that genetic drift.[80] [81] 106 1940s–1960s: Molecular biology and evolution The middle decades of the 20th century saw the rise of molecular biology. in a scientific climate that favored experimental methods over historical ones. molecular biology was increasingly seen as a threat to the traditional core of evolutionary biology.History of evolutionary thought The evolutionary synthesis provided a conceptual core—in particular. epitomized by the kin selection arguments of W.[79] The synthesis also resulted in a considerable narrowing of the range of mainstream evolutionary thought (what Stephen Jay Gould called the "hardening of the synthesis"): by the 1950s. The molecular clock hypothesis and the neutral theory were particularly controversial. This was due to the paucity of morphological traits and the lack of a species concept in microbiology. and endosymbiosis Microbiology was largely ignored by early evolutionary theory. Archaea green. particularly amongst prokaryotes.[103] Microbiology. some derived from game theory.[109] More recently. to explore the taxonomy and evolution of these organisms. [98] genome data. Hamilton's work on kin selection contributed to the emergence of the discipline of sociobiology. horizontal gene transfer. Other theories followed. such as reciprocal altruism. The existence of altruistic behaviors has been a difficult problem for evolutionary theorists from the beginning. produced by the comparative ease of microbial genomics. behavior. including work on other aspects of the altruism problem. work has continued in sociobiology and the related discipline of evolutionary psychology. including Stephen Jay Gould and Richard Lewontin. E. Niles Eldredge and Stephen Jay Gould proposed that there was a pattern of fossil species that remained largely unchanged for long periods (what they termed stasis). Eukaryotes of evolutionary theories using this huge amount of are colored red.O. Discoveries in biotechnology now allow the modification of entire genomes.[96] [97] Improvements in sequencing methods resulted in a large increase of sequenced genomes.[99] [100] These genomic analyses have produced fundamental changes in the understanding of the evolutionary history of life.[108] This transfer of genetic material between different species of bacteria was first recognized because it played a major role in the spread of antibiotic resistance.[102] One of the results has been an exchange of ideas between theories of biological evolution and the field of computer science known as evolutionary computation. which showed how eusociality in insects (the existence of sterile worker classes) and many other examples of altruistic behavior could have evolved through kin selection. allowing the testing and refining A phylogenetic tree showing the three-domain system.History of evolutionary thought 107 Sociobiology W.[93] In 1975. Wilson published the influential and highly controversial book Sociobiology: The New Synthesis which claimed evolutionary theory could help explain many aspects of animal. as knowledge of genomes has . and Bacteria blue. They also claimed that the theories of sociobiologists often reflected their own ideological biases. claimed that sociobiology greatly overstated the degree to which complex human behaviors could be determined by genetic factors.[94] [95] Evolutionary paths and processes One of the most prominent debates arising during the 1970s was over the theory of punctuated equilibrium. interspersed with relatively brief periods of rapid change during speciation.[101] Advances in computational hardware and software allow the testing and extrapolation of increasingly advanced evolutionary models and the development of the field of systems biology.[92] Significant progress was made in 1964 when Hamilton formulated the inequality in kin selection known as Hamilton's rule. such as the proposal of the three-domain system by Carl Woese. which attempts to mimic biological evolution for the purpose of developing new computer algorithms. advancing evolutionary studies to the level where future experiments may involve the creation of entirely synthetic organisms. Comparisons between these genomes provide insights into the molecular mechanisms of speciation and adaptation.[106] [107] One particularly important outcome from studies on microbial evolution was the discovery in Japan of horizontal gene transfer in 1959. Despite these criticisms.[105] These studies are revealing unanticipated levels of diversity amongst microbes. evolutionary researchers are taking advantage of their improved understanding of microbial physiology and ecology. Critics of sociobiology. including human. D.[104] Now. and emphasized the self-organizing processes of development as factors directing the course of evolution.[119] Such perspectives influenced the disciplines of phylogenetics.[110] These high levels of horizontal gene transfer have led to suggestions that the family tree of today's organisms. It had been suggested in the late 19th century when similarities between mitochondria and bacteria were noted. calling them "exaptations". and animals. had descended from independent bacteria that came to live symbiotically within other cells. as part of the endosymbiotic theory for the origin of organelles. for example.[123] [124] Some have suggested that in some cases a form of . plants. and spawned the new discipline of evolutionary developmental biology.[120] More recent work in this field by Mary Jane West-Eberhard has emphasized phenotypic and developmental plasticity. to account for the generation of novel forms.[122] Experimental and theoretical research on these and related ideas have been presented in the multi-authored volume Origination of Organismal Form. but from changes in the deployment of a small set of proteins that were common to all animals. paleontology and comparative developmental biology.[117] Molecular data regarding the mechanisms underlying development accumulated rapidly during the 1980s and '90s. There was a renewal of structuralist themes in evolutionary biology in the work of biologists such as Brian Goodwin and Stuart Kauffman. could arise incidentally as an accidental result of selection on another structure.[118] These proteins became known as the "developmental toolkit". They called such incidental structural changes "spandrels" after an architectural feature. rather than through direct selection for that particular adaptation. it has been suggested that lateral transfer of genetic material has played an important role in the evolution of all organisms. with the evolutionary biologist Richard Lewontin. The resulting forms were later stabilized by natural selection. but largely dismissed until it was revived and championed by Lynn Margulis in the 1960s and 70s. It became clear that the diversity of animal morphology was not the result of different sets of proteins regulating the development of different animals. the so-called "tree of life". a form of horizontal gene transfer has been a critical step in the evolution of eukaryotes such as fungi. By the first decade of the 21st century it had become accepted that in some cases such environmental factors could affect the expression of genes in subsequent generations even though the offspring were not exposed to the same environmental factors.[111] [112] Indeed. The evolutionary biologist Stephen Jay Gould revived earlier ideas of heterochrony.[116] Later. Gould and Vrba discussed the acquisition of new functions by novel structures arising in this fashion. such as differential cell adhesion and biochemical oscillation.[121] It has been suggested. alterations in the relative rates of developmental processes over the course of evolution. or even a structural novelty. the study of how environmental factors affect the way genes express themselves during development. wrote an influential paper in 1979 suggesting that a change in one biological structure. that the rapid emergence of basic animal body plans in the Cambrian explosion was due in part to changes in the environment acting on inherent material properties of cell aggregates. and. This shows that in some cases non genetic changes to an organism can be inherited and it has been suggested that such inheritance can help with adaptation to local conditions and affect evolution. and there had been no genetic changes. is more similar to an interconnected web or net.[115] 108 Evolutionary developmental biology In the 1980s and 1990s the tenets of the modern evolutionary synthesis came under increasing scrutiny. Margulis was able to make use of new evidence that such organelles had their own DNA that was inherited independently from that in the cell's nucleus.History of evolutionary thought continued to expand. which incorporated ideas from cybernetics and systems theory.[113] [114] The endosymbiotic theory holds that organelles within the cells of eukorytes such as mitochondria and chloroplasts. 21st century Epigenetic inheritance Yet another area where developmental biology has led to the questioning of some tenets of the modern evolutionary synthesis is in the field of epigenetics. "The Vatican claims Darwin's theory of evolution is compatible with Christianity" (http:/ / www. edu/ entries/ lucretius/ ). [9] Sedley. ca/ RAS lecture on daoism and nature. 2004). [10] Simpson. ii. htm#H4).[125] 109 Unconventional evolutionary theory Omega Point Pierre Teilhard de Chardin's metaphysical Omega Point Theory describes the gradual development of the universe from subatomic particles to human society. utm. . 89–90 [12] Gill 2005. . possibly in the form of a technological singularity. uk/ news/ newstopics/ religion/ 4588289/ The-Vatican-claims-Darwins-theory-of-evolution-is-compatible-with-Christianity. stanford. [6] Singer 1931 [7] Needham & Ronan 1995. R. . uk/ tol/ comment/ faith/ article5705331. p. David (August 4. "Lucretius" (http:/ / www. pdf) (PDF).uk. "Lucretius" (http:/ / plato. Retrieved 2008-07-15. "Daoism and Nature" (http:/ / www. Malaspina University College. Royal Asiatic Society. iep. Retrieved 2009-02-12. ca/ ~johnstoi/ darwin/ sect3. who proposed that the living and nonliving parts of Earth can be viewed as a complex interacting system with similarities to a single organism. viu. .[128] This modified hypothesis postulates that all living things have a regulatory effect on the Earth's environment that promotes life overall. utm. Retrieved 2008-07-24. jamesmiller.History of evolutionary thought Lamarckian evolution may occur. "Section Three: The Origins of Evolutionary Theory" (http:/ / records. mit. [11] Augustine 1982. London: Telegraph. . . edu/ anaximan/ #H8). transhumanists often view such technological evolution itself as a goal in their philosophy. Retrieved 2010-02-27. 101 [8] Miller. . p. Retrieved 2008-07-24. [4] Mayr 1982. p. . utm. Gordon. 2. [2] Campbell. 2008). Retrieved 2008-07-15. timesonline. "Anaximander" (http:/ / www. James (January 8. Internet Encyclopedia of Philosophy. htm). Retrieved 2007-08-11. "Empedocles" (http:/ / www. Gaia hypothesis Teilhard de Chardin's ideas have been seen as being connected to the more specific Gaia theory by James Lovelock. See also • Faith and rationality • Galápagos Islands • The Voyage of the Beagle Notes [1] Couprie. And Still We Evolve: A Handbook on the History of Modern Science. edu/ Aristotle/ physics. Chris (2009-02-12). London: Times Online. iep. Ian (1999). . The Internet Encyclopedia of Philosophy. Stanford Encyclopedia of Philosophy. Transhumanism Futurists have often viewed scientific and technological progress as a continuation of biological evolution.[126] The Gaia hypothesis has also been viewed by Lynn Margulis[127] and others as an extension of endosymbiosis and exosymbiosis. edu/ l/ lucretiu. co. R. K. co. ece). "Vatican buries the hatchet with Charles Darwin" (http:/ / www. pp.. which he viewed as its final stage and goal. html). [3] Hardie. Dirk L. htm). Gaye.. 304 [5] Johnston.P. Internet Encyclopedia of Philosophy... telegraph. David (2006). [14] Irvine. edu/ e/ empedocl.co. Retrieved 2008-07-15. Liberal Studies Department. "Physics by Aristotle" (http:/ / classics. Retrieved 2009-02-12. iep. 251 [13] Owen. . . Richard (2009-02-11). html). Among these. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=77) [52] Matthew. 142–143 [42] Larson 2004. 5–24 [43] Bowler 2003. 138 [46] Larson 2004. William E. "obliterates decades of labor by teachers. However. pp. org/ articles/ science/ sc0035.0171. pp. in his study of the impact of Vestiges of the Natural History of Creation. . htm).1098/rsnr. 158 [55] Huxley. 86–94 [37] Larson 2004. pp. Retrieved 2010-03-20. "Mind the gap: Did Darwin avoid publishing his theory for many years?" (http:/ / darwin-online. editors.. The Complete Works of Charles Darwin Online. Fathers of the English Dominican Province. pp. [16] Conway Zirkle (1941). uk/ content/ view/ 110/ 104/ ). [21] Bowler 2003. pp. he argues. Retrieved 2010-03-23. p. Retrieved 2009-11-17. amnh. 129–134 [36] Bowler 2003. 120–129 [40] Bowler 2003. . pp. 110 . xiv (http:/ / darwin-online. catholiceducation. p. 33–38 [22] Schelling. Thomas Henry (1895). 71–123. p. pp. . 29–38 [33] Bowler 2003. pp. xiv (http:/ / darwin-online. pp. uk/ content/ frameset?itemID=A143& viewtype=text& pageseq=1). argues that in some ways Vestiges had as much or more impact than Origin. Retrieved 2007-11-02. 42–46 [47] van Wyhe. org. org. htm#14). 50 [60] The centrality of Origin of Species in the rise of widespread evolutionary thinking has been has long been accepted by historians of science. org. 37–38 [45] Bowler 2003. Retrieved 11-3-2010. uk/ content/ frameset?itemID=F381& viewtype=text& pageseq=20) [49] Darwin 1866. 40 [39] Bowler 2003. p. Bulletin of the Ecological Society of America. pp. html). Focusing so much on Darwin and Origin. . pp. and Thomas Aquinas" (http:/ / www. 151 [51] Darwin 1859. xiii (http:/ / darwin-online. org/ education/ resources/ rfl/ web/ essaybooks/ earth/ p_hutton." [31] Bowler 2003. pp. James A. technicians. Natural Selection before the "Origin of Species". Project Gutenberg. pp. Catholic Education Resource Center. darwinproject. edu/ history/ medieval. 73–75 [24] Bowler 2003. 134–138 [41] Bowler & Morus 2005. [56] Bowler & Morus 2005. 103–104 [44] Larson 2004. Secord. org. [35] Bowler 2003. 129–149 [57] Larson 2004. 67–80 [19] Carroll. "Physica Book 2. html). Patrick (1860). 62 (http:/ / darwin-online. "A History of the Ecological Sciences. Lecture 14" (http:/ / dhspriory. 38–41 [38] Desmon & Moore 1993. doi:10. 55–71 [58] Bowler 2003. p. org. uk/ content/ frameset?viewtype=text& itemID=F385& pageseq=21) [50] Bowler 2003. ac. 7 [30] American Museum of Natural History (2000). "The Reception of the Origin of Species" (http:/ / infomotions. pp. Evolution. [20] Thomas Aquinas. Retrieved 2008-01-17. 177–205. Gardeners' Chronicle and Agricultural Gazette" (http:/ / darwin-online. p. pp. [48] Darwin 1861. 15 [29] Larson 2004. p. pp. theologians.History of evolutionary thought [15] "Medieval and Renaissance Concepts of Evolution and Paleontology" (http:/ / www. org/ thomas/ Physics2. org. John (27 March 2007). uk/ content/ frameset?itemID=F381& viewtype=text& pageseq=21) [54] Bowler 2003. April 2002: 142–146 [143] [18] Lovejoy 1936. pp. . 75–80 [25] Larson 2004. ucmp. University of California Museum of Paleontology.2006. at least into the 1880s. com/ etexts/ gutenberg/ dirs/ etext00/ oroos10. some scholars have recently begun to challenge this idea. p. and other researchers. 1800 [23] Bowler 2003. pp. pp. pp. 14–15 [26] Henderson 2000 [27] Darwin 1818. p. berkeley. pp. printers. 173–176 [59] Larson 2004. Darwin Correspondence Project. pp. [53] Darwin 1861. Egerton. Vol I section XXXIX [28] Darwin 1825. "we find no vestige of a beginning. Earth: Inside and Out. 113 [32] Larson 2004. p. . Proceedings of the American Philosophical Society 84 (1). . [17] Frank N. Retrieved 2007-11-01. uk/ content/ frameset?viewtype=text& itemID=A544& pageseq=1). p. "James Hutton: The Founder of Modern Geology" (http:/ / www. "Nature's law of selection. "Creation. pp. System of Transcendental Idealism. no prospect of an end. . html). Part 6: Arabic Language Science – Origins and Zoological". 115–116 [34] "Darwin and design: historical essay" (http:/ / www. PMC 33654. 109–110 [64] Bowler 2003. 338–341 [78] Mayr & Provine 1998. Princeton: Princeton Univ. paralogs. pp. Cambridge University Press. "Five rules for the evolution of cooperation". pp. Acad. [83] Powell. Reprinted in N. . 97–188 [80] Sapp 2003. pp. PMC 1692213. ed. (1998-03-01). [99] Koonin EV (2005). 49–51 [69] Larson 2004. [87] Bowler 2003. 207–216 [68] Bowler 2003. U. 196–253 [71] Bowler 2003. JB (1999). "Naturalists. pp. "Cooperation within and among species". 402.1023/A:1004257523100. "Gulliver's further travels: the necessity and difficulty of a hierarchical theory of selection" (http:/ / www. [100] Hegarty MJ. PMID 17158317. pp. Sci. p. [93] Nowak M (2006). [86] Mayr E (1997). B. asp)" In T. discussion 1426–36. Models in Paleobiology. Schierwater. org/ library/ gould_synthesis. doi:10.1126/science. 1972. Jeffrey R (1994) [1994]. pnas. pp. 279 [90] Bowler 2003. ISBN 3-7643-2942-4. doi:10." Secord 2000. 82–115. PMID 16910971. PMID 11110893. Genet. Natl. pp. E§year=1988. org/ cgi/ content/ full/ 17/ 12/ 1776). pnas. Biol. [85] Hagen. "The objects of selection" (http:/ / www. 190–191 [65] Bowler 2003. "A case for evolutionary genomics and the comprehensive examination of sequence biodiversity" (http:/ / mbe. [77] Mayr & Provine 1998.6. Streit.1133755. 139–40 [63] Larson 2004.1073/pnas. com/ ridley/ classictexts/ eldredge. 39: 309–38. and R. Acad. 1985 [97] Gould SJ (1994). Journal of the History of Biology 27 (1): 21–59. PMID 11619919. 515–518 [61] Larson 2004. 353 (1366): 307–14. In B. 358–359 [92] Sachs J (2006).S. Doggett NA. 295–298.1998. doi:10. pp. Hiscock SJ (2005).1420-9101.94. "Hybrid speciation in plants: new insights from molecular studies". Journal of the History of Biology 32 (2): 321–341.x.1146/annurev. p.2006. 358 [91] Bowler 2003. Molecular Ecology and Evolution: Approaches and Applications. pp.J. p.01253. 416 [76] Mayr. 152–156 [81] Gould. pp. P. html). 270–278 [95] Bowler 2003. Press. Wagner..1098/rstb. Trans. PMID 8041695. 79–111 [62] Larson 2004. and evolutionary genomics".1073/pnas. pp. Evol.2004. Harvard University Press. pp.A. pubmedcentral. PMC 44281. U. doi:10. Science 314 (5805): 1560–63. Biol. PMID 15720652.2091. pp.1023/A:1004660202226. J. pp. Journal of the History of Biology 31 (1): 85–111. doi:10. pp. New Phytol.114725.History of evolutionary thought whose work has made evolutionary debates so significant during the past two centuries. B.x. pp. "Tempo and mode in the macroevolutionary reconstruction of Darwinism" (http:/ / www. 121–123.A. 19 (5): 1415–8. fcgi?tool=pubmed& pubmedid=9533127). Michael R. Stephen Jay (1983).M. 361 [88] Gould SJ (1998). Eldredge Time frames. Lond. Cummings MP (1 December 2000). blackwellpublishing. Annu. Natl. "Molecular techniques in population genetics: A brief history".. " Punctuated equilibria: an alternative to phyletic gradualism (http:/ / www.1469-8137. Proc. (1994-03-01). 152–157 [67] Bowler 2003. Mol. . stephenjaygould. 177–223 [66] Larson 2004. Soc. Philos. [82] Dietrich. 111 . and the Challenges of Molecular Evolution". PMID 16285863. Schopf.0211. 94 (6): 2091–94. San Francisco: Freeman Cooper. [98] Pollock DD. nih.39. pp.genet. Rev. Molecular Biologists. oxfordjournals. p. "The hardening of the modern synthesis" (http:/ / www. pp. Biol. 359–361 [96] Niles Eldredge and Stephen Jay Gould. "The origins of the neutral theory of molecular evolution". 17 (12): 1776–88. . pp. 153–174 [73] Bowler 2003. gov/ articlerender. doi:10. 91 (15): 6764–71. pp. pp. "Paradox and Persuasion: Negotiating the Place of Molecular Evolution within Evolutionary Biology". Sci. org/ cgi/ reprint/ 91/ 15/ 6764).). doi:10. 33–34 [79] Smocovitis 1996. doi:10. doi:10. Evol. 256–273 [72] Larson 2004. "Orthologs. 165 (2): 411–23. pp. . doi:10. G. 131–156. In Marjorie Grene. Birkhäuser Verlag. org/ cgi/ content/ full/ 94/ 6/ 2091). 221–243 [75] Mayr & Provine 1998. PMID 11624208. PMID 9122151. . 105–129 [70] Bowler 2003.6764.01152. 325–339 [74] Larson 2004. Michael R. Proc. PMID 11639258.15. Towards a new philosophy of biology: observations of an evolutionist. [89] Larson 2004. [94] Larson 2004. De Salle (eds. Sci.91. pp.1111/j. pp.1111/j. R. Eisen JA. pp.S. [84] Dietrich. Dimensions of Darwinism.1007/BF01058626. PMID 9533127.073003. [122] Newman SA. fcgi?tool=pubmed& pubmedid=15590780#r6). Yokoi H.cellbio. Rev. nih.History of evolutionary thought [101] Woese C. Evol. Proc. [126] Lovelock J (2003). doi:10. Lawrence JG.A. doi:10. Gevers D. "Status of the microbial census" (http:/ / www. Brown M. [110] de la Cruz F. "Horizontal gene transfer and the origin of species: lessons from bacteria".1038/nrg1637.4576.559. [116] Gould SJ (1997). [117] Gould SJ and Vrba ES (1982). PMID 15073369. . Nature 426 (6968): 769–70. Developmental Plasticity and Evolution. 102 Suppl 1: 6630–5. Oxford University Press. Acad.1073/pnas. Proc Natl Acad Sci USA 87 (12): 4576–79.coli strains". "The net of life: reconstructing the microbial phylogenetic network" (http:/ / www. me. com/ biomedicine/ 22061/ ). [103] Benner SA.0610699104. Swings J (2005). .1073/pnas. M-J (2003). Acad. (in Japanese) [109] "Lateral gene transfer and the nature of bacterial innovation" (http:/ / www. pnas. doi:10. Retrieved 2010-02-21. "Evo-Devo: the long and winding road" (http:/ / www.95." (http:/ / jrsm.CO. interscience. 8 (3): 128–33. Dev. 6 (7): 533–43. "Epigenetics and Plant Evolution" (http:/ / www3.0501984102. Trends Microbiol. Nat. . B Mol. "Evolutionary developmental biology and genomics". Wheelis M (1990). org/ cgi/ reprint/ 87/ 12/ 4576).1073/pnas. gov/ articlerender. Annu. U. "Gaia: the living Earth". Sci. New Phytologist. Müller GB (2000). [107] Schloss P.140619. Retrieved 2010-02-21. com/ christinalrichards/ Portfolio/ Epigenetics. [119] Cañestro C. Science 304 (5668): 253–7.1094884. PMID 15995697. PMC 1079665. PMID 12142278. 112 . Jonathan. [115] "Endosymbiosis: Lynn Margulis" (http:/ / evolution.12. Microbiol Mol Biol Rev 68 (4): 686–91.1038/nrg2226. Cohan FM. . Nat. . J. 47 (7–8): 467–77. Ouzounis CA (2005). doi:10.1146/annurev. PMID 14756346.1126/science.2-G.1002/bies. [106] Whitman W. "Symbiogenesis. 47 (7–8): 705–13. "Gene co-option in physiological and morphological evolution". "The exaptive excellence of spandrels as a term and prototype" (http:/ / www.1258/jrsm. Int.686-691. Davies J (2000). Sismour AM (2005). 2000.004. gov/ articlerender. Genome Res.1016/S0966-842X(00)01703-0.1073/pnas. [105] Coenye T. Rev.S. org/ cgi/ pmidlookup?view=long& pmid=17261804). "Epigenetic mechanisms of character origination". stat. [127] Margulis. Wiebe W (1998). Kandler O. J. edge. Bioessays 29 (1): 74–84. [125] Singer. "Towards a prokaryotic genomic taxonomy". org/ documents/ ThirdCulture/ n-Ch. PMC 1892968. Natl. Cell Dev. 29 (2): 147–67. PMID 15590780. University of California Berkely. Bacteria. "Towards a natural system of organisms: proposal for the domains Archaea.94. Johnson P (2004). pnas. Int. "Opinion: Re-evaluating prokaryotic species". doi:10. doi:10. Wendell. . doi:10. nih. PMID 15808739. [128] Fox. Proc. Robin (2004).18. Retrieved 2007-09-01.4. The Third Culture. PMID 15574850. (2005). PMID 16138101.2004. "Evaluating hypotheses for the origin of eukaryotes". . [120] Baguñà J. fcgi?tool=pubmed& pubmedid=11038582). "A Comeback for Lamarckian Evolution?" (http:/ / www. 94 (20): 10750–5. Retrieved 2007-09-30. wiley. edu/ evolibrary/ article/ _0_0/ history_24). Biol. Retrieved 2010-02-21. pubmedcentral. pubmedcentral. doi:10. doi:10. Nat Rev Genet 8 (12): 932–942. U.10750. PMC 1172039. genome.020402. technologyreview. Journal of the royal society of medicine 97 (12): 559. Postlethwait JH (2007). [111] Kunin V. PMID 14685210. Ryan.20516. "Ancient invasions: from endosymbionts to organelles". FEMS Microbiol. [124] Rapp. Proc Natl Acad Sci USA 95 (12): 6578–83. Christina. org/ cgi/ content/ full/ 95/ 12/ 6578). rice. doi:10. "Prokaryotes: the unseen majority" (http:/ / www. .11. "Exaptation—a missing term in the science of form".1016/j. . Garcia-Fernàndez J (2003). com/ journal/ 118647390/ abstract). gov/ articlerender.87. "The morphogenesis of evolutionary developmental biology". org/ cgi/ pmidlookup?view=long& pmid=15965028). . Handelsman J (2004). et al. Dev. Nature Vol 405. [121] West-Eberhard. Yamanaka T. PMID 15965028. Rev. [108] Ochiai K. PMC 23474. PMID 17261804. . [118] True JR. pubmedcentral.20. Goldovsky L. fcgi?tool=pubmed& pubmedid=15851668).2004. 3 (9): 733–9. "Inheritance of drug resistance (and its transfer) between Shigella strains and Between Shigella and E.97. Biol. Coleman D. doi:10. 288 (4): 304–17. ehu. [102] Medina M (2005). *Gilbert SF (2003). PMC 54159. Zool. . Microbiol. [114] Dyall S. Exp. pnas. . Biol. html). es/ web/ paper. Vandamme P.A. PMID 11038582. PMID 14756322. PMID 15851668. Carroll SB (2002). Kimura K Sawada O (1959). Van de Peer Y. Genet. PMID 11144279. Proc. phylogeny. Rev. Lynn (1995). html). Sci. Sci.S.1128/MMBR.1038/426769a. "Gaia Is a Tough Bitch" (http:/ / www. and Eucarya" (http:/ / www. ijdb. . "Epigenetics and Evolution" (http:/ / web.1002/1097-010X(20001215)288:4<304::AID-JEZ3>3. Emily (2009). doi:10. Retrieved 2010-02-20. doi:10.1038/nrmicro1236. pdf) (PDF). doi:10. PMID 2112744. Natl. "Synthetic biology". U. [113] Poole A. edu/ ~mathbio/ Ochman2000. nih.1073/pnas. rsmjournals. com/ cgi/ content/ full/ 97/ 12/ 559).3666505. and evolutionary systems biology" (http:/ / www. PMC 1131869. . Natl.femsre. doi:10. php?doi=14756346). 7. South Florida University.68. PMID 9618454. [112] Doolittle WF. doi:10. J. Acad. doi:10.0. PMID 18007650. 104 (7): 2043–9. 18: 53–80.1101/gr. Penny D (2007). Bapteste E (February 2007). Develop. MIT Technology Review. 15 (7): 954–9. PMID 10707066. [123] Roberts. May 18.12. "Pattern pluralism and the Tree of Life hypothesis" (http:/ / www. [104] Gevers D. Hihon Iji Shimpor 1861: 34. Paleobiology 8 (1): 4–15. Darzentas N.A. PMC 33863. "Genomes.12. doi:10.6578. PMC 539005.S. PMID 17187354. berkeley. • Darwin. Princeton University Press. • Darwin. Ernst (1982). ISBN 0-679-64288-9. 9780809103263. The University of Chicago Press. (2000). • Desmond. John Murray. ISBN 0-691-03343-9. (2004). Clarendon Press. On the Origin of Species by Means of Natural Selection. ISBN 0-19-515618-8. Unifying Biology: The Evolutionary Synthesis and Evolutionary Biology. John Hammond Taylor. Vassiliki Betty (1996). Adrian. or the Preservation of Favoured Races in the Struggle for Life (1st ed. Reception. Cambridge University Press. (2003).). and Secret Authorship of Vestiges of the Natural History of Creation. The Growth of Biological Thought: Diversity. The Structure of Evolutionary Theory. The Emperor's Kilt: The Two Secret Histories of Scotland. On the Origin of Species by Means of Natural Selection. • Needham. or the Preservation of Favoured Races in the Struggle for Life (6th ed. James (1994). Evolution: The Remarkable History of a Scientific Theory. Cyril (1959). On the Origin of Species by Means of Natural Selection.. W. or the Preservation of Favoured Races in the Struggle for Life (4th ed. Jan-Andrew (2000). ISBN 0521292867. trans. University of Chicago Press. • Secord. Oxford University Press. University of California Press. .google.com/?id=AksE4jKI8yQC). Harvard University Press. • Singer. ISBN 0521832144. Stephen Jay (2002). B. ISBN 0393311503. • Gould. W. • Gill. Darwin's place in history. Peter J. Edward J. Making Modern Science. ISBN 0-674-36153-9. or The Origin of Society (http://books. Blackwell. Ernst. Evolution: The History of an Idea (3rd ed. Norton & Company. The Great Chain of Being: A Study of the History of an Idea. ISBN 0-226-06861-7. • Henderson. John Murray.com/ ?id=oAl9y-0FSJQC&dq=Erasmus+Darwin+Temple). Iwan Rhys (2005). p85. Victorian Sensation: The Extraordinary Publication. Charles (1866). and Inheritance. Ronan. Augustine in the Italian Renaissance: Art and Philosophy from Petrarch to Michelangelo (http://books. ISBN 0-674-27225-0.. Harvard University Press. The Temple of Nature.htm).gutenberg. • Darlington. London. London. Jan (2003). • Lovejoy. The Literal Meaning of Genesis (http://books. Oxford. John Murray. Modern Library. John Murray. The Evolutionary Synthesis: Perspectives on the Unification of Biology. ISBN 0-674-00613-5.). or the Preservation of Favoured Races in the Struggle for Life (3rd ed. Charles (1872). Cambridge University Press. Peter J. Arthur (1936). The Newman Press. Meredith Jane (2005).History of evolutionary thought 113 References • Augustine (1982).com/?id=_s0kIgD0nCcC). Morus.google. Evolution.). • Bowler.org/files/15707/15707-h/15707-h. • Darwin. ISBN 9780521832144. Provine.google. Mainstream Publishing. Belknap Press of Harvard University Press. Erasmus (1818). Jones & Company. • Smocovitis. W. Darwin: The Life of a Tormented Evolutionist. Colin Alistair (1995). The Shorter Science and Civilisation in China: An Abridgement of Joseph Needham's Original Text. Charles (1861). London. Vol. • Mayr. • Sapp. Moore. On the Origin of Species by Means of Natural Selection. • Larson.). eds (1998). ISBN 0-52023693-9. Joseph. 1. London. ISBN 0-226-74410-8. • Darwin. Genesis: The Evolution of Biology. A Short History of Biology. Charles (1859). ISBN 0809103265. • Mayr. ISBN 0-674-36445-7. • Bowler. James A.). Zoonomia (http://www. Erasmus (1825). Charles (1931). The Belknap Press of Harvard University Press. • Darwin. • Darwin. Mendelian genetics supplanted the notion of inheritance of acquired traits. eventually leading to the development of the modern evolutionary synthesis. He is often incorrectly cited as the founder of soft inheritance. authored by Robert Chambers in St Andrews and published anonymously in England in 1844. . He incorporated this mechanism into his thoughts on evolution. and the general abandonment of the Lamarckian theory of evolution in biology. and worked with him on marine creatures. who incorporated the action of soft inheritance into his evolutionary theories.. Lamarck founded a school of French Transformationism which Jean-Baptiste Lamarck included Étienne Geoffroy Saint-Hilaire. History Between 1794 and 1796 Erasmus Darwin wrote Zoönomia suggesting "that all warm-blooded animals have arisen from one living filament. which proposes that individual efforts during the lifetime of the organisms were the main mechanism driving species to adaptation. In addition. with each round of "improvements" being inherited by successive generations. It is named after the French biologist Jean-Baptiste Lamarck (1744–1829). seeing it as resulting in the adaptation of life to local environments. • The Alfred Russel Wallace Page (http://www.Origins Archive. After publication of Charles Darwin's theory of natural selection. proposed a theory which combined radical phrenology with Lamarckism. As a young student.org/faqs/darwin-precursors. soft inheritance is of use when examining the evolution of cultures and ideas. Robert Jameson. as they supposedly would acquire adaptive changes and pass them on to offspring. with the power of acquiring new parts" in response to stimuli.edu/~smithch/index1. In a wider context. Lamarck" for explaining how the higher animals had "evolved" from the "simplest worms" – this was the first use of the word "evolved" in a modern sense. causing political controversy for its radicalism and unorthodoxy. and is related to the theory of Memetics. html).berkeley.wku. Charles Darwin was tutored by Grant. but exciting popular interest and preparing a huge and prosperous audience for Darwin. Part of the Talk. The Vestiges of the Natural History of Creation. and which corresponded with a radical British school of anatomy based in the extramural anatomy schools in Edinburgh which included the surgeon Robert Knox and the comparative anatomist Robert Edmund Grant. Later.htm) Lamarckism Lamarckism (or Lamarckian inheritance) is the idea that an organism can pass on characteristics that it acquired during its lifetime to its offspring (also known as heritability of acquired characteristics or soft inheritance).talkorigins.edu/evolibrary/article/history_01) • Darwin's precursors and influences by John Wilkins (http://www.History of evolutionary thought 114 External links • UC Berkeley's History of Evolutionary Thought (http://evolution. Subsequently Jean-Baptiste Lamarck repeated in his Philosophie Zoologique of 1809 the folk wisdom that characteristics which were "needed" were acquired (or diminished) during the lifetime of an organism then passed on to the offspring.. the Regius Professor of Natural History. the importance of individual efforts in the generation of adaptation was considerably diminished. was the probable author of an anonymous paper in 1826 praising "Mr. in line with Lamarck's overall evolutionary theory. throw off 'gemmules' or 'pangenes' which travelled around the body (though not necessarily in the bloodstream). and Darwin believed that they eventually accumulated in the germ cells where they could pass on to the next generation the newly acquired characteristics of the parents. essentially maintaining the principle that genetic changes can be influenced and directed by environmental factors. Francis Galton carried out experiments on rabbits. particularly the midwife toad. was based on the idea that somatic cells would. experiments by Paul Kammerer on amphibians. with Darwin's cooperation. Lamarckism largely fell from favor. and explained it in the final chapter of his book Variation in Plants and Animals under Domestication. Darwin's half-cousin.inheritance of acquired characters .[1] Darwin called his Lamarckian hypothesis Pangenesis. Gould advocated defining "Lamarckism" more broadly. 115 Lamarck's theory The identification of Lamarckism with the inheritance of acquired characteristics is regarded by some as an artifact of the subsequent history of evolutionary thought. and truly a discredit to the memory of a man and his much more comprehensive system"[4] . but Darwin objected. The evolution of necks is often used as the example in explanations of Lamarckism. that he had done nothing of the sort. in response to environmental stimulation (use and disuse). cast aside the guts of it . repeated in textbooks without analysis. which have no blood. Pangenesis. in a letter to Nature. after describing numerous examples to demonstrate what he considered to be the inheritance of acquired characteristics. Since 1988 certain scientists have produced work proposing that Lamarckism could apply to single celled organisms. in which he transfused the blood of one variety of rabbit into another variety in the expectation that its offspring would show some characteristics of the first. and Galton declared that he had disproved Darwin's hypothesis of Pangenesis. as. They did not."[3] He argued that "the restriction of "Lamarckism" to this relatively small and non-distinctive corner of Lamarck's thought must be labelled as more than a misnomer. These pangenes were microscopic particles that supposedly contained information about the characteristics of their parent cell. since he had never mentioned blood in his writings. appeared to find evidence supporting Lamarckism. In The Case of the Midwife Toad Arthur Koestler surmised that the specimens had been faked by a Nazi sympathiser to discredit Kammerer for his political views. in his day considered to be generally true: . A version of Lamarckian acquisition in higher order animals is still posited in certain branches of psychology. This ideologically driven research influenced Soviet agricultural policy which in turn was later blamed for crop failures. which he emphasised was a hypothesis. but did not rule out a variant of Lamarckism as a supplementary mechanism. Lamarck incorporated two ideas into his theory of evolution. in the Jungian racial memory. A form of Lamarckism was revived in the Soviet Union of the 1930s when Trofim Lysenko promoted Lysenkoism which suited the ideological opposition of Joseph Stalin to Genetics. Stephen Jay Gould wrote that late 19th century evolutionists "re-read Lamarck.Lamarckism Darwin's Origin of Species proposed natural selection as the main mechanism for development of species.to a central focus it never had for Lamarck himself.. for example. Neo-Lamarckism is a theory of inheritance based on a modification and extension of Lamarckism.[2] With the development of the modern synthesis of the theory of evolution and a lack of evidence for either a mechanism or even the heritability of acquired characteristics. In the 1920s. but his specimens with supposedly-acquired black foot-pads were found to have been tampered with. and elevated one aspect of the mechanics .. He pointed out that he regarded Pangenesis as occurring in Protozoa and plants. [5] In essence. which may have been transmitted to the offspring by means of epigenetic mechanisms. provided that the acquired modifications are common to both sexes. In every animal which has not passed the limit of its development. 116 Current views on "Lamarckism" Interest in Lamarckism has recently increased. All the acquisitions or losses wrought by nature on individuals. resulting in change in behavior. none of these views were original to Lamarck. leading to ever more complex construction regardless of the organ's use or disuse. Use and disuse – Individuals lose characteristics they do not require (or use) and develop characteristics that are useful. 2. Sweden. This link has been shown before in studies of human populations . through his work. This would drive organisms from simple to complex forms. through the influence of the environment in which their race has long been placed. Le pouvoir de la vie (a complexifying force) . while the permanent disuse of any organ imperceptibly weakens and deteriorates it. • A blacksmith. Lamarck's contribution was a systematic theoretical framework for understanding evolution. develops and enlarges that organ. specialising them for their environment.. a more frequent and continuous use of any organ gradually strengthens. and gives it a power proportional to the length of time it has been so used. which have looked at foraging behavior in chickens as well as stress factors[8] . genetic exchange without reproduction between single-celled organisms has been well-researched and is termed horizontal gene transfer. This may have prepared the offspring for coping with an unpredictable environment. and hence through the influence of the predominant use or permanent disuse of any organ. and progressively diminishes its functional capacity. 1. bringing change in form over time — and thus the gradual transmutation of the species. all these are preserved by reproduction to the new individuals which arise. This would take organisms sideways off the path from simple to complex. a change in the environment brings about change in "needs" (besoins). He saw evolution as comprising two processes. Some recent notable studies include those made by the University of Linköping. strengthens the muscles in his arms. L'influence des circonstances (an adaptive force) .in which the use and disuse of characters led organisms to become more adapted to their environment. as historians of science such as Michael Ghiselin and Stephen Jay Gould have pointed out. Examples of what is traditionally called "Lamarckism" would include: • Giraffes stretching their necks to reach leaves high in trees (especially Acacias). 2. strengthen and gradually lengthen their necks. The group of researchers at The University of Linköping again highlighted the apparent link between food intake and cross-generational inheritance of acquired traits. The conclusion of the referenced study is as follows: Our findings suggest that unpredictable food access caused seemingly adaptive responses in feeding behavior. With this in mind.in which the natural. until it finally disappears. alchemical movements of fluids would etch out organs from tissues. Inheritance of acquired traits – Individuals inherit the traits of their ancestors.[8] On a much smaller scale. His sons will have similar muscular development when they mature. or at least to the individuals which produce the young.. bringing change in organ usage and development. including regulation of immune genes.[6] [7] On the contrary. These giraffes have offspring with slightly longer necks (also known as "soft inheritance").. Lamarck has been credited in some textbooks and popular culture with developing two laws: 1. 2. Transmissions of information across generations which does not involve traditional inheritance of DNA-sequence alleles is often referred to as soft inheritance [9] or 'Lamarckian inheritance'. as several studies in the field of epigenetics have highlighted the possible inheritance of behavioral traits acquired by the previous generation. However.Lamarckism 1. He claimed that with each generation. The findings provide support for a 200-year-old theory of evolution that has been largely dismissed: Lamarckian evolution. where epigenetic factors have altered the functioning of genes[10] . like Lamarck. "The effects of an animal's environment during adolescence can be passed down to future offspring . little more than indirect evidence was ever acquired to support it. than to any coherent body of theoretical work. claimed to have observed a similar phenomenon in animals being subject to conditioned reflex experiments. which states that acquired characteristics can be passed on to offspring. one conducted by researchers at MIT and another by researchers at the Tufts University School of Medicine. Several recent studies. in which he simply followed the accepted wisdom of his time. lacked a plausible alternative mechanism of inheritance . proposed a neo-Lamarckian mechanism to try to explain why homologous DNA sequences from the VDJ gene regions of parent mice were found in their germ cells and seemed to persist in the offspring for a few generations. Soma to germ-line feedback In the 1970s the immunologist Ted Steele. These changed epigenetic factors appear to show traits in the next generation such as an increased occurrence of diabetes. which in turn led to obesity for the daughter. and not their genetics (or specific genes). have rekindled the debate once again. The mRNA products of these somatically novel genes were captured by retroviruses endogenous to the B-cells and were then transported through the blood stream where they could breach the soma-germ barrier and retrofect (reverse transcribe) the newly acquired genes into the cells of the germ line.. if Lamarck had been aware of Darwin's proposed mechanism of natural selection. Although Steele was advocating this theory for the better part of two decades. far more than for the mechanism of evolution..Lamarckism who have experienced starvation. Its full significance was not appreciated until the Modern evolutionary synthesis in the early 1920s. The process of methylation is thought to be behind such changes. caused by poor experimental controls." 117 Neo-Lamarckism Unlike neo-Darwinism. formerly of the University of Wollongong. According to Stephen Jay Gould. However. Harvard University researcher William McDougall studied the abilities of rats to correctly solve mazes. the term neo-Lamarckism refers more to a loose grouping of largely heterodox theories and mechanisms that emerged after Lamarck's time. who was also a Lamarckist.[14] [15] [16] [17] [18] At around the same time. He found that children of rats that had learned the maze were able to run it faster.the particulate nature of inheritance was only observed by Gregor Mendel somewhat later. which Darwin’s own theory lacked. and published in 1866. Several historians have argued that Lamarck's name is linked somewhat unfairly to the theory that has come to bear his name. Ivan Pavlov. and that Lamarck deserves credit for being an influential early proponent of the concept of biological evolution. McDougall attributed this to some sort of Lamarckian evolutionary process. Note also that Darwin. and colleagues. An interesting attribute of this idea is that it strongly resembles Darwin's . Lamarck died 30 years before the first publication of Charles Darwin's Origin of Species. prior to the conception of the daughter. An important point in its favour at the time was that Lamarck's theory contained a mechanism describing how variation is maintained. A "paternal high-fat diet" was shown to cause cell dysfunction in the daughter. the animals became easier to condition. The mechanism involved the somatic selection and clonal amplification of newly acquired antibody gene sequences that were generated via somatic hyper-mutation in B-cells. Pavlov never suggested a mechanism to explain these observations. but after a few generations it was down to 20. In October 2010. there is no reason to assume he would not have accepted it as a more likely alternative to his own mechanism. The study strongly suggested that fathers can transfer a propensity for obesity to their daughters as a result of the fathers' food intake. further evidence linking food intake to traits inherited by the offspring were shown in a study of rats conducted by several Australian universities[11] . The first rats would get it wrong 165 times before being able to run it perfectly each time. In the 1920s.[13] Oscar Werner Tiegs and Wilfred Eade Agar later showed McDougall's results to be incorrect. As reported [12] in MIT's Technology Review in February 2009. though Cairns later chose to distance himself from such a position. Although the reality of epigenetic inheritance is not doubted (as countless experiments have validated it). These are considered "Lamarckian" in the sense that they are responsive to environmental stimuli and can differentially affect gene expression adaptively. Cairns. the trait reappeared in some of the resulting offspring. If bacteria that had overcome their own inability to consume lactose passed on this "learned" trait to future generations.[22] The group took a mutated strain of E. When the researchers mated the yeast cells with cells not containing the prion. and selection remains Darwinian. N. England. among others. its significance to the evolutionary process is uncertain. (see Horizontal Gene Transfer) In 1988. involving the duplication of genetic material and its segregation during meiosis. dubbed the process adaptive mutation. where the effects of cultural niche construction are transmissible from one generation to the next. as in addition to being subject to . discovered that in yeast cells containing a specific prion protein Sup35. They observed over time that mutations occurred within the colony at a rate that suggested the bacteria were overcoming their handicap by altering their own genes. coli that was unable to consume the sugar lactose and placed it in an environment where lactose was the only food source. There has been some research into Lamarckism and prions. with no potential to introduce evolutionary novelty into a species lineage. it could be argued as a form of Lamarckism. both of which regulate the activity of genes.[23] More typically. Lamarckism and societal change Jean Molino (2000) has proposed that Lamarckian evolution may be accurately applied to cultural evolution. indicating that some information indeed was passed down. some scientists controversially argue that it can be observed among microorganisms. except in the soma to germ line feedback theory. and a group of other scientists renewed the Lamarckian controversy (which at that point had been a dead debate for many years). Most neo-Darwinians consider epigenetic inheritance mechanisms to be little more than a specialized form of phenotypic plasticity. some of which gave them new abilities such as resistance to a particular herbicide.[21] Whether such mutations are directed or not also remains in contention. Laland and colleagues have recently suggested that human culture can be looked upon as an ecological niche like phenomena. This was also previously suggested by Peter Medawar (1959) and Conrad Waddington (1961). pangenes are replaced with realistic retroviruses. While this does not directly confer advantage to the organism on the organismal level.[24] Finally. for example.[19] Epigenetic inheritance Forms of 'soft' or epigenetic inheritance within organisms have been suggested as neo-Lamarckian in nature by such scientists as Eva Jablonka and Marion J. Lamb. though whether or not the information is genetic is debatable: trace prion amounts in the cells may be passed to their offspring. A group of researchers. with phenotypic results that can persist for many generations in certain organisms. the yeast were able to gain new genetic material. John Cairns at the Radcliffe Infirmary in Oxford.Lamarckism own theory of pangenesis.[20] 118 Lamarckism and single-celled organisms While Lamarckism has been discredited as an evolutionary influence for larger lifeforms. there are other hereditary elements that pass into the germ cells also. While the acquisition of new genetic traits is random. These include things like methylation patterns in DNA and chromatin marks. it makes sense at the gene-evolution level. giving the appearance of a new genetic trait where there is none. K. the active process of identifying the necessity to mutate is considered to be Lamarckian. there is growing evidence that cells can activate low-fidelity DNA polymerases in times of stress to induce mutations. it might be viewed as a form of ontogenic evolution. In addition to 'hard' or genetic inheritance. One interpretation of the Meme theory is that memes are both Darwinian and Lamarckian in nature. Harvard: Belknap Harvard. Marion J. A Weekly Illustrated Journal of Science. 148. U. "Oscar Werner Tiegs". Vol. Nat Rev Genet 7(5): 395–401. "First report on a test of McDougall's Lamarckian experiment on the training of rats". [17] W E Agar. A. [15] W E Agar. Steele. Edward J. The Structure of Evolutionary Theory. Quote from page 66. 1984.Lamarckism selection pressures based on their ability to differentially influence Human minds. 1371/ journal. & Moore. F H Drummond (1942). inist. "Pangenesis" (http:/ / darwin-online. html [12] http:/ / www. F H Drummond (1935). com/ nature/ journal/ v467/ n7318/ full/ nature09491. Journal of Experimental Biology 12: 191. 1998 [20] Epigenetic Inheritance and Evolution: The Lamarckian Dimension. "Third report on a test of McDougall's Lamarckian experiment on the training of rats". M M Gunson (1954). Robert V. F H Drummond (1948). Journal of Experimental Biology 25: 103. Perseus Books. Lamb. [10] http:/ / cat. org. ISBN 0-674-00613-5. Richard Dawkins notes (in Blackmore 2000: The Meme machine. Chicago Press. fr/ ?aModele=afficheN& cpsidt=3596539 [11] http:/ / www. [3] Gould. org/ article/ info:doi/ 10. (2002). S. April 1998 . Stephen J. Biographical Memiors of Fellows of the Royal Society (The Royal Society) 3: 247. pp. org/ 54marck. "Shades of Lamarck". page 13).617 "Darwin was loathe to let go of the notion that a well-used and strengthened organ could be inherited" [2] Charles Darwin (27 April 1871). Eva Jablonka.113 [6] The Imaginary Lamarck: a look at bogus "history" in schoolbooks (http:/ / www. technologyreview. pone. Lindley. 1938. nature. pp. Robyn A. 0006405 [9] Richards EJ (2006) Inherited epigenetic variation–revisiting soft inheritance. . [18] W E Agar.J. plosone. 119 See also • • • • • • • • • • • • • • • • Jean-Baptiste Lamarck Baldwinian evolution Darwinism Epigenetic inheritance Epigenetics Evolution Inheritance of acquired characters Lysenkoism Memetics Obsolete scientific theories Orthogenesis Marcus Pembrey Racial memory Ted Steele History of evolutionary thought Eclipse of Darwinism Notes [1] Desmond. com/ biomedicine/ 22061 [13] McDougall. memes can be modified and the effects of that modification passed on. 1995 [21] Adaptive mutation Genetics. that Memes can be copied in a Lamarckian way (copying of the product) or in a Weismann-type evolutionary way (copying of the instruction) which is much more resistant against changes. (1991) Darwin Penguin Books p. Hugh Elliot. "Second report on a test of McDougall's Lamarckian experiment on the training of rats". J. Oxford University Press. 502–503. [4] Gould.65-71. [19] Lamarck's Signature: how retrogenes are changing Darwin's natural selection paradigm. [16] W E Agar. "Fourth (final) report on a test of McDougall's Lamarckian experiment on the training of rats". p. (2002) The Structure of Evolutionary Theory [8] http:/ / www. Journal of Experimental Biology 31: 308. Blanden. uk/ content/ frameset?viewtype=text& itemID=F1751& pageseq=1). F H Drummond. Journal of Experimental Biology 19: 158. Retrieved 2007-11-08. 177–178. Nature. C F A (November 1957). htm) by Michael Ghiselin [7] Gould. Stephen J. [5] Jean-Baptiste Lamarck Zoological Philosophy trans. British Journal of Psychology 28:321-345 [14] Pantin. textbookleague. reprinted in The Panda's Thumb (1980) pp. A History of Evolutionary Thought. • Hall Barry G.) and realised by CNRS (France .uk/ science/2010/mar/19/evolution-darwin-natural-selection-genes-wrong). • Guralnick. 2009.pdf) External links • Nonsense in Schoolbooks . Conrad (1961). "The human evolutionary system". Take 2 (http://www.edu/history/ lamarck. • Vetsigian K. Issue #2461 120 Further references • Burkeman. et.guardian. ca/ biochem/ courses/ 4103/ topics/ adaptive_mutation. html Adaptive mutation in bacteria [23] Adaptive mutation in E. This web site contents all books.). 186. • Cairns. 15 [24] Lamarckism and prions. Woese C. ISBN 0-262-23206-5. University of California Museum of Paleontology. Merker & Wallin (Eds.ucmp. 6.fr/?lang=en): an English/French web site edited by Pietro Corsi (Oxford Univ.). manuscripts and the lamarck's herbarium. Peter (1959).newsweek. • Medawar.co. al (2006). and S. Chicago: University of Chicago Press. The Guardian. In Brown. texts. • Jean-Baptiste Lamarck : works and heritage (http://www. "The threat and the glory". Adrian (1989).genetics.berkeley. J. I. Overbaugh. "Collective Evolution and the Genetic Code.IT team of CRHST). . March 19. • Gould. • Molino. BBC Reith Lectures No. • The Sins of the Fathers. 2006.org/54marck. Harvard: Belknap Harvard. 1988. Science.The Imaginary Lamarck (http://www. Darwinism and the Study of Society. New Scientist.lamarck.org/cgi/reprint/120/4/887.Lamarckism [22] http:/ / www. Elizabeth. and Reform in Radical London. Ghiselin recounts Lamarck's times and writings. No. Oliver. From the magazine issue dated January 26. The Politics of Evolution: Morphology.html). Medicine. August 2004. Newsweek. 265:318." PNAS 103: 10696-10701. Journal of Bacteriology. Mutations Involving an Insertion Sequence (http://www.textbookleague. Rob. • Waddington. pp. • Desmond. Why everything you've been told about evolution is wrong (http://www. 170–197 on Lamarck. Vol.com/id/180103/output/print): "At tributes to Darwin.. mun. ISBN 0-226-14374-0. Miller. "Toward an Evolutionary Theory of Music and Language". Nature 335: 142-145 • Culotta. 2010. Stephen J. The Structure of Evolutionary Theory. (2002). J." By Sharon Begley. 21 August 2004. In: Michael Banton (Ed. Lamarckism — inheritance of acquired traits — will be the skunk at the party. The Origins of Music.. London: Tavistock. 1994. coli. "A Boost for 'Adaptive' Mutation". Adaptive Evolution That Requires Multiple Spontaneous Mutations.htm):Michael T. "Jean-Baptiste Lamarck (1744-1829)" (http://www. Retrieved 3 July 2010.cnrs. Jean (2000). ISBN 0-674-00613-5. Goldenfeld N. but in complex organisms gave way to the generally accepted Darwinian mechanisms. but they are not part of the germ line. but there are some prominent proponents. suggested that the absence of RNA signature continuum between domains of bacteria. Pop culture In popular culture. See also • • • • • • Catastrophism Phyletic gradualism Rapid modes of evolution The Blind Watchmaker History of evolutionary thought Eclipse of Darwinism . a form of saltation appears to have emerged from misconceptions over currently accepted theories of evolution (the X-men and its various spin-offs being the most egregious examples). a case when a child belongs to a different species than its parents . punctuated equilibrium refers instead to a pattern of evolution where most speciation occurs relatively rapidly from a geological perspective (tens of thousands of years instead of millions of years). Punctuated Equilibrium It is a popular misconception that punctuated equilibrium is a saltationist theory. including Carl Woese. in comparison with the usual variation of an organism. or violate. that is large. not during a generation. It was also fictionalized in Greg Bear's novel. Woese.Saltationism 121 Saltationism In biology."[3] However. The Teenage Mutant Ninja Turtles ( and many other. Darwin's Radio. not by saltations. nongradual changes (especially single-step speciation) that are atypical of. often mistaken for Richard Goldschmidt's hypothesis of "Hopeful Monsters.gradualism . saltus. Saltation does not fit into contemporary evolutionary theory. archaea. Mammalian liver cells are typically polyploidal. The term is used for occasionally hypothesized. and colleagues.involved in neo-Darwinian evolution. and eukarya constitutes a primary indication that the three primary organismal lineages materialized via one or more major evolutionary saltations from some universal ancestral state involving dramatic change in cellular organization that was significant early in the evolution of life. saltation (from Latin. even though most polyploid individuals are sterile. "leap") is a sudden change from one generation to the next.between generations.[1] Polyploidy (most common in plants but not unknown in animals) is considered a type of saltation [2] . similar science-fiction pieces ) are not examples of saltationism. but through neo-Darwinian evolution. however. A saltation would be a substantial change that takes place during reproduction. standard concepts . or very large. Polyploidy meets the basic criteria of saltation in that a significant change (in gene numbers) results in speciation in just one generation. N. full?sid=f3651397-00e9-4a57-802b-f41c6ef6cf5a). Both scientists proposed mechanisms whereby evolution proceeded in unilinear fashion. for a time. stephenjaygould. Retrieved 2008-05-05. This is important because similar flaws occur recurrently at the fringes of science. The distinction can be seen when we recognize that orthogenesis is inherent in the theories of German biologist Ernst Haeckel and American paleontologist Richard Swann Lull." Orthogenesis Orthogenesis. org/ pss/ 49988). Proceedings of the Royal Society. . progressive evolution or autogenesis. the paleontologist Henry Fairfield Osborn. however. punctuated equilibrium became a saltational theory wedded to Goldschmidt's hopeful monsters as a mechanism. 1006–1021. philosopher Henri Bergson and. The term orthogenesis was popularised by Theodor Eimer. Those who accepted orthogenesis in this way. Jean-Baptiste Lamarck himself accepted the idea. and theories of speciation for a rectilinear model of guided evolution acting on discrete species with "essences". Many sources mix this heterodox view of evolution with another—. Paul D. "Molecular signatures of ribosomal evolution" (http:/ / www. did not . the hypothesised mechanism of which resembled the "mysterious inner force" of orthogenesis. though many of the ideas are much older (Bateson 1909).g. Harvard University Press. The hypothesis is based on essentialism and cosmic teleology and proposes an intrinsic drive which slowly transforms species. and Zaida Luthey-Schulten (May 19. html). it does not have a goal. orthogenetic evolution. typically taking the form of mysterious molecular drives that supposedly are pushing phenotypic evolution in certain directions or forcing the formation of new species. Herbert (1994). org/ content/ 105/ 37/ 13953. [3] Gould. it is important to recognize that these are in fact two separate ideas that are rejected by mainstream science. Anurag Sethi†.that evolution is proceeding to some long term or ultimate goal. While it is true that early and famous examples of orthogenesis often conflated these two ideas (e. Stephen Jay. is the hypothesis that life has an innate tendency to move in an unilinear fashion due to some internal or external "driving force". Woese. and it had a central role in his theory of inheritance of acquired characteristics. "[T]he urban legend rests on the false belief that . org/ library/ gould_structure. the result are definitions that state "orthogenesis proposes that evolution moves in a unilinear fashion towards a perfect goal". "Hybridization and origins of polyploidy" (http:/ / www. jstor. Origins The orthogenesis hypothesis had a significant following in the 19th century when a number of evolutionary mechanisms. Carl R. Though it proceeds in a linear fashion driven by some internal mechanism. . Proceedings of the National Academy of Sciences.[2] No Goal Orthogenesis does not postulate a "goal" for evolution.. "Punctuated Equilibrium's Threefold History" (http:/ / www. pp.Saltationism 122 Notes and references [1] Elijah Roberts. and in invertebrate paleontology thought there was a gradual and constant directional change. such as Lamarckism. Orthogenesis was particularly accepted by paleontologists who saw in their fossils a directional change. Jonathan Montoya. George Gaylord Simpson (1953) in an attack on orthogenesis called this mechanism "the mysterious inner force". I have labored to refute this nonsensical charge from the day I first heard it. . The Structure of Evolutionary Theory.. Retrieved 2010-05-06.[1] Classic proponents of orthogenesis have rejected the theory of natural selection as the organising mechanism in evolution. 2008). [2] France Dufresne. Other proponents of orthogenesis included Leo Berg. pnas. but neither saw goals (instead they made pseudo-scientific appeals to unknown genetic driving processes). Jean-Baptiste Lamarck's theory of evolution). were being proposed. the latter idea of goal-oriented evolution is better understood as a form of teleology. Depends upon source quoted. i. The modern evolutionary synthesis. Status Prevailing in modified form as modern evolutionary synthesis. No. died hard. The orthogenetic hypothesis. Common descent Yes. that fortuitous variations can furnish the material for the precise and balanced adaptations that all nature reveals? Selection cannot create the materials on which it is supposed to operate. As more was understood about these mechanisms it became obvious that there was no possible naturalistic way in which the newly discovered mechanism of heredity could be far-sighted or have a memory of past trends. the course of evolution. Moreover. From the Butlerian side. in every direction. a step or . of the minute dovetailing of structure. no other guidance or aim. both in the production of variations and their selection. natural selection adopted by some in latter years. refuted the hypothesis for good. in which the genetic mechanisms of evolution were discovered. Lamarckism Intrinsic drive towards perfection and inheritance of acquired characteristics (both are Lamarckian principles). pointed to the existence of orthogenetic series. i. seemed to him to imply the existence of an originative. speciation rejected or considered unimportant in long term trends. for which he invented the term bathmism (Gr. because in Darwin's time. and this had been common usage since at least 1647. and the theory of evolution by natural selection became the prevailing theory of evolution. How is it possible. but apparently a steady and progressive march in one direction. Collapse of the hypothesis The orthogenesis hypothesis began to collapse when it became clear that it could not explain the patterns found by paleontologists in the fossil record.Orthogenesis necessarily accept that the mechanism that drove orthogenesis was teleological.[4] Edward Drinker Cope put such a line of argument in the most cogent fashion.e. but in absence of a mechanism some still rejected the idea. Characters produced may be totally non-adaptive. however. many naturalists. the initial stages of new functions cannot be supposed to have been useful. natural selection unimportant.e. too. of long lines of ancestry. function and environment. Refuted by Charles Darwin's Origin of Species and the modern evolutionary synthesis. In fact. Orthogenesis Intrinsic drive towards perfection. Darwin certainly was impressed with the view that natural selection and variation together formed a mechanism. the central product of which was adaptation. have no survival value. came the most urgent opposition to Darwinism. Selected traits are adaptive. it was said. especially those concerned with palaeontology. conscious and directive force. Even Darwin was at first not opposed to orthogenic thinking. spontaneous generation of new species resulting in parallel evolution. and regarded the organic world as almost a miracle of adaptation. have some survival value. βαθμ. though the mechanism was not refuted until the modern evolutionary synthesis in which it was established that the mechanism does not exist.[3] 123 Comparison of Theories Comparison of different theories of evolution Darwinism Mechanism Short-sighted Natural Selection sorting random genetic variation. which displayed not a sporadic differentiation. which was non-linear with many complications. Darwin himself rarely used the term "evolution" now so commonly used to describe his theory. Declined after the Origin. Signs that species shared a common ancestor were detected before Darwin. The hypothesis was generally abandoned when no mechanism could be found that would account for the process. new species coming into existence by speciation events. evolution usually was associated with some sort of progressive process like orthogenesis. the beginnings of new organs. as this quote from the 1911 Encyclopedia Britannica demonstrates: Darwin and his generation were deeply imbued with the Butlerian tradition. Obviously variations depend on the constitution of the varying organism. as it refuted the idea of teleology as first postulated by Aristotle and accepted by Immanuel Kant. . must inevitably display orthogenesis to some extent. by Jacobs et al. in one sense it is a mere coincidence if a particular variation turn out to be useful. This form of orthogenesis has now also been abandoned as more about evolutionary processes has been discovered (Wilkins 1997). were distinct from the processes of microevolution (genetic variation and natural selection) which were by then well understood and it was known they could not behave in an orthogenetic manner. it is not true that evolution never proceeds in a linear way. 1995 & Woodley 2006) but are not strictly orthogenetic.. and that science was the study of God's creation. reinforcing characteristics. The apparent opposition between the conflicting schools is more acute than the facts justify. but such examples are entirely consistent with the modern neo-Darwinian theory of evolution. in The Phenomenon of Man (a book influential among non-scientists that was published four years after his death in 1959) argued for evolution aiming for the "Omega Point".. von Kitzing 1998). and simply appear as linear and constant changes because of environmental and molecular constraints on the direction of change. The past history of the organic world displays many successful series and these. a Jesuit paleontologist. sustained environmental change. during a period of slow. who had greatly influenced many scientists. On the other hand. The refutation of orthogenesis had some ramifications in the field of philosophy. is a modification of an already definite and limited structure.. These examples have sometimes been referred to as orthogenetic (e. 124 Modern co-opted usage Though linear teleological evolution has been refuted.g. while putting man at the center of the universe and accounting for original sin (Dennett 1995.. the long term trends in evolution.. but it also displays many failures which indeed may be regarded as showing that the limitation of variation has been such that the organisms have lost the possibility of successful response to a new environment.. The term Chardin used for this was "directed additivity". whether it be large or small. A continuous environment both from the point of view of production of variation and selection of variation would appear necessarily to result in a series with the appearance of orthogenesis.. for example. a modification.Orthogenesis beginning).[4] A few hung on to the orthogenesis hypothesis as late as the 1950s by claiming that the processes of macroevolution. Teilhard de Chardin. dislike of mystical interpretations of natural facts has driven many capable naturalists to another extreme and has led them to insist on the all-powerfulness of natural selection and on the complete indefiniteness of variation.. Before the scientific and philosophical revolution that began with Charles Darwin's ideas. The refutation of these concepts have led to a shift in what science and scientists are perceived to be. the prevailing philosophy was that the world was teleological and purposeful. as they have survived. there is no connection between the appearance of the variation and the use to which it may be put. in certain lineages at times. But there are several directions in which the field of variation appears to be not only limited but defined in a certain direction. Syst.. Phylogenetic Relationships and Orthogenetic Evolution of Coat Color Among Tamarins (Genus Saguinus). Oct 13 2004) 8. p. 44(4):515—532. 2. Simon & Schuster. Allan Larson & James M. Copyright in the United States of America by the Encyclopaedia Britannica Company. 4.C. Cambridge University Press. [2] The evolutionary future of man: A biological view of progress (http:/ / www. 5. p. ac. 1911. ox.. Life of the Past: An Introduction to Paleontology. Yale University Press. an essay in Ever Since Darwin: Reflections in Natural History.edu/systbiol/issues/44_4/44_4abstracts. 1942. May 31. 6. John. E-book (http://www.html#Jacobs). George G. Sciences. The Limits of Ecology: New Perspectives from a Theoretical Borderland. Ernst. Bateson. 1995. Woodley.. TalkOrigins Archive (http://www. Abstract (http:// hydrodictyon. London: Weidenfeld and Nicolson. shtml) [3] Darwin's Dilemma: The Odyssey of Evolution (http:/ / web. 3. . Daniel. The Modern Evolutionary Synthesis. Dennett. simonyi. What Evolution Is.Orthogenesis 125 See also • • • • • Facilitated variation Evolution of complexity Eclipse of Darwinism History of evolutionary thought Law of Complexity/Consciousness References [1] George Gaylord Simpson. htm#evolution).gutenberg. Seward ed. Jacobs. Simpson. 1977. Susan C. 125. Huxley. Sources 1. Masters and Scholars of the University of Cambridge. Norton.eeb. Julian. W. Chapter V. 1909. ISBN 0-393-06425-5 [4] The Encyclopaedia Britannica: A Dictionary of Arts. archive. 1953. 1957. Stephen Jay Gould. Heredity and variation in modern lights. Michael A. New Haven. Life Of The Past: Introduction to Paleontology. Copyright in all countries subscribing the Berne Convention by the Chancellor. org/ web/ 20051216025526/ http:/ / wordorigins.talkorigins. 1995. Darwin's Dangerous Idea.119. 2002. in Darwin and Modern Science (A. Yale University Press. London.org/faqs/ macroevolution. What is macroevolution?. Wilkins. org/ wordore. William. London: George Allen and Unwin.uconn. uk/ dawkins/ WorldOfDawkins-archive/ Dawkins/ Work/ Articles/ 1993-09-11future.net/etext/1909). Abramis Academic Press.). Eleventh Edition. Mayr. 2006.html) (14:08 UTC. 7. 1997. W. Literature and General Information. Cheverud. Biol. Various evolutionary ideas had already been proposed to explain new findings in biology. but transmutation was not accepted by the scientific mainstream. Its full title was On the Origin of Species by Means of Natural Selection. correspondence. . and experimentation. Ideas about the transmutation of species were controversial as they conflicted with the beliefs that species were unchanging parts of a designed hierarchy and that humans were unique. and on the Perpetuation of Varieties and Species by Natural Means of Selection Fertilisation of Orchids [2] Charles Darwin's On the Origin of Species. Darwin's book introduced the scientific theory that populations evolve over the course of generations through a process of natural selection. There was growing support for such ideas among dissident anatomists and the general public. while science was part of natural theology. is a work of scientific literature which is considered to be the foundation of evolutionary biology.''On the Origin of Species'' 126 On the Origin of Species On the Origin of Species The title page of the 1859 edition [1] of On the Origin of Species Author Country Language Subject(s) Genre(s) Publisher Publication date Media type Pages ISBN Preceded by Followed by Charles Darwin United Kingdom English Natural selection Evolutionary biology science. The political and theological implications were intensely debated. It presented a body of evidence that the diversity of life arose by common descent through a branching pattern of evolution. unrelated to animals. or the Preservation of Favoured Races in the Struggle for Life. biology John Murray 24 November 1859 Print (Hardback) 502 N/A On the Tendency of Species to form Varieties. Darwin included evidence that he had gathered on the Beagle expedition in the 1830s and his subsequent findings from research. published on 24 November 1859. the short title was changed to The Origin of Species. but during the first half of the 19th century the English scientific establishment was closely tied to the Church of England. For the sixth edition of 1872. • Resources such as food are limited and are relatively stable over time (fact).''On the Origin of Species'' The book was written for non-specialist readers and attracted widespread interest upon its publication. now the unifying concept of the life sciences. 127 Summary of Darwin's theory Darwin's theory of evolution is based on key facts and the inferences drawn from them. populations remain roughly the same size (fact). • A struggle for survival ensues (inference). which biologist Ernst Mayr summarised as follows:[3] • Every species is fertile enough that if all offspring survived to reproduce the population would grow (fact). philosophical. Nature was widely believed to be unstable and capricious. Background Developments before Darwin's theory In later editions of the book. Darwin's concept of evolutionary adaptation through natural selection became central to modern evolutionary theory. these variations accumulate over time to form new species (inference). and religious discussion. • Individuals in a population vary significantly from one another (fact). and spontaneous generation of life. but scientists were slow to give natural selection the significance that Darwin thought appropriate. with monstrous births from union between species. • Individuals less suited to the environment are less likely to survive and less likely to reproduce. individuals more suited to the environment are more likely to survive and more likely to reproduce and leave their inheritable traits to future generations. Darwin pictured shortly before publication • This slowly effected process results in populations changing to adapt to their environments. his findings were taken seriously and the evidence he presented generated scientific. various other mechanisms of evolution were given more credit. • Much of this variation is inheritable (fact).[6] organisms were described by their mythological and heraldic significance as well as by their physical form.[5] Early Christian Church Fathers and Medieval European scholars interpreted the Genesis creation myth allegorically rather than as a literal historical account.[4] the text he cites is a summary by Aristotle of the ideas of the earlier Greek philosopher Empedocles.H. With the development of the modern evolutionary synthesis in the 1930s and 1940s.[7] . Darwin traced evolutionary ideas as far back as Aristotle. and ultimately. The debate over the book contributed to the campaign by T. had occurred. • Despite periodic fluctuations. As Darwin was an eminent scientist. which produces the process of natural selection (inference). with a branching pattern of common descent. Huxley and his fellow members of the X Club to secularise science by promoting scientific naturalism. Within two decades there was widespread scientific agreement that evolution. During the "eclipse of Darwinism" from the 1880s to the 1930s. ''On the Origin of Species'' 128 The Protestant Reformation inspired a literal interpretation of the Bible. Wernerians thought strata were deposits from shrinking seas. and read Paley. in his taxonomy. This process was later called Lamarckism. but by the 1780s geologists assumed a much older world. Georges Cuvier strongly disputed such ideas. their adaptation and complexity designed by God. which he explained by local catastrophes. initially identifying the most recent catastrophe as the biblical flood.[13] Some anatomists such as Robert Grant were influenced by Lamarck and Geoffroy. All naturalists in English universities were Church of England clergymen. the Royal Society wanted to show that science did not threaten religious and political stability. followed by repopulation of the affected areas by other species. Georges Buffon suggested that some similar species. but James Hutton proposed a self-maintaining infinite cycle. Charles Darwin's grandfather Erasmus Darwin outlined a hypothesis of transmutation of species in the 1790s. he studied catastrophist geology with Adam Sedgwick. adapting to the environment by inheriting changes in adults caused by use or disuse.[9] Geoffroy contended that embryonic development recapitulated transformations of organisms in past eras when the environment acted on embryos.[11] In Britain. The biological classification introduced by Carolus Linnaeus in 1735 also viewed species as fixed according to the divine plan. carnivores caused mercifully swift death. anticipating [8] uniformitarianism. Both envisaged that spontaneous generation produced simple forms of life that progressively developed greater complexity. might be varieties descended from a common ancestor. species were static and fixed. In 1766. and that animal structures were determined by a constant plan as demonstrated by homologies. The Ussher chronology of the 1650s had calculated creation at 4004 BC. Lamarck thought there was an inherent progressive tendency driving organisms continuously towards greater complexity. William Paley's Natural Theology saw adaptation as evidence of beneficial "design" by the Creator acting through natural laws.[15] At Cambridge University starting in 1827. In his second year he neglected his medical studies for natural history and spent four months assisting Robert Grant's research into marine invertebrates. and science became a search for these laws. fixed species showed similarities that reflected a design for functional needs. in parallel but separate lineages with no extinction. holding that unrelated. Cuvier's 1799 paper on living and fossil elephants helped establish the reality of extinction.[14] Inception of Darwin's theory Darwin went to Edinburgh University in 1825 to study medicine.[16] [17] . John Ray developed an influential natural theology of rational order. such as horses and asses. After the turmoil of the English Civil War. In God's benevolent design. and leopards. and Jean-Baptiste Lamarck published a more developed theory in 1809. or lions. but the suffering caused by parasitism was a puzzling problem. but most naturalists regarded their ideas of transmutation as a threat to divinely appointed social order. Filled with zeal for science. with concepts of creation that conflicted with the findings of an emerging science seeking explanations congruent with the mechanical philosophy of René Descartes and the empiricism of the Baconian method.[12] Geologists adapted catastrophism to show repeated worldwide annihilation and creation of new fixed species adapted to a changed environment. Darwin learnt science as natural theology from botanist John Stevens Henslow. but Darwin rejected it. John Herschel and Alexander von Humboldt. tigers. and varieties showed minor differences caused by local conditions.[10] His paleontological work in the 1790s had established the reality of extinction. Grant revealed his enthusiasm for the transmutation of species. Darwin discovered fossils resembling huge armadillos. Jago. and around July sketched a evolutionary tree. genealogical branching of a single evolutionary tree.[24] In late September 1838. he had noted a similarity between the act of breeders selecting traits and a Malthusian Nature selecting among variants thrown up by "chance" so that "every part of newly acquired structure is fully practical and perfected". at St. Darwin asked questions of fancy pigeon and animal breeders as well as established scientists.[33] . he joined the Beagle expedition as a geologist and naturalist. At the zoo he had his first sight of an ape. and on page 36 wrote "I think" above his first another" to explain these findings.[29] [30] Further development Darwin continued to research and extensively revise his theory while focusing on his main work of publishing the scientific results of the Beagle voyage. so that the survivors would pass on their form and abilities.[25] [26] By December 1838.[20] As the Beagle neared England in 1836.[29] He tentatively wrote of his ideas to Lyell in January 1842.[19] and he no longer saw an unbridgeable gap between humans and animals. he started reading Thomas Malthus's An Essay on the Principle of Population with its statistical proof that human populations breed beyond their means and struggle to survive. on the possibility that "one species does change into notebook on Transmutation of Species. and noted the geographical distribution of modern species in hope of finding their "centre of creation". yet to Darwin their relatives on the island seemed "miserable. and was profoundly impressed by how human the orangutan seemed. he immediately envisioned "a force like a hundred thousand wedges" pushing well-adapted variations into "gaps in the economy of nature".[21] Richard Owen showed that fossils of extinct species Darwin found in South America were allied to living species on the same continent. and that several distinct birds from those islands were all classified as finches. to be expanded with his research results and published if he died prematurely.[27] Darwin now had the framework of his theory of natural selection "by which to work". Darwin related this to the struggle for existence among wildlife and botanist de Candolle's "warring of the species" in plants. and by July had rounded out his "sketch" into a 230-page "Essay". that mockingbirds collected on the Galápagos Islands represented three separate species each unique to a particular island. found Lyell's uniformitarianism a key to the geological history of landscapes.''On the Origin of Species'' 129 In December 1831. In March 1837. and unfavourable variations would be destroyed.[22] Darwin began speculating. in a series of In mid-July 1837 Darwin started his "B" notebooks.[23] Unconventionally.[28] but he was fully occupied with his career as a geologist and held off writing a sketch of his theory until his book on The Structure and Distribution of Coral Reefs was completed in May 1842. ornithologist John Gould announced that Darwin's Rhea was a separate species from the previously described rhea (though their territories overlapped). degraded savages". discarding Lamarck's independent lineages progressing to higher forms. he noted that species might not be fixed.[31] then in June he roughed out a 35-page "Pencil Sketch" of his theory.[32] Darwin began correspondence about his theorising with the botanist Joseph Dalton Hooker in January 1844.[18] The three Fuegian missionaries the expedition returned to Tierra del Fuego were friendly and civilised. He read Charles Lyell's Principles of Geology and from the first stop ashore. Reminded of his lack of expertise in taxonomy. including Adam Sedgwick. with a mass of supporting evidence. While few naturalists were willing to consider transmutation. and scorned its amateurish geology and zoology.''On the Origin of Species'' 130 In November 1844. he completed the last part of his Beagle-related writing and began working full-time on evolution. He conducted empirical research focusing on difficulties with his theory. he saw increasing specialisation within large stable populations as continuously exploiting new ecological niches. In 1854. widened public interest in the concept of transmutation of species. became actively involved in fancy pigeon breeding.[35] Vestiges had significant influence on public of Plants and Animals Under Domestication of opinion. His thinking changed from the view that species formed in isolated populations only.[36] Hooker was persuaded to take away a copy of the "Essay" in January 1847. Vestiges used evidence from the fossil record and embryology to support the claim that living things had progressed from the simple to the more complex over time. By 1856. acceptance of the more scientifically sophisticated Origin by moving evolutionary speculation into the mainstream. as on islands. and he found an intermediate stage in the evolution of distinct sexes. Herbert Spencer became an active proponent of Lamarckism and progressive development in the 1850s. the anonymously published popular science book Vestiges of the Natural History of Creation. to an emphasis on speciation without isolation. he discovered homologies showing that slightly changed body parts served different functions to meet new conditions. attacked its morality and Darwin researched how the skulls of different pigeon breeds varied. Darwin began an eight year study of barnacles. Using his theory. He studied the developmental and anatomical differences between different breeds of many domestic animals.[37] [38] . written by Scottish journalist Robert Chambers. becoming the leading expert on their classification.[37] Darwin's barnacle studies convinced him that variation arose constantly and not just in response to changed circumstances. that is. and eventually sent a page of notes giving Darwin much needed feedback. his theory was much more sophisticated.[34] but he carefully reviewed his own arguments after leading scientists. and experimented (with the help of his son Francis) on ways that plant seeds and animals might disperse across oceans to colonise distant islands. Darwin read it soon after publication. and the intense debate helped to pave the way for the 1868. But it proposed a linear progression rather than the branching common descent theory behind Darwin's work in progress. and it ignored adaptation. as shown in his Variation scientific errors. with the title page adding by Means of Natural Selection. On 14 May 1856. and focused on ecological pressures that kept different varieties adapted to local conditions. will be smashed".[39] Charles Lyell recognised the implications of Wallace's paper and its possible connection to Darwin's work. of the cabbage" and proceeds to a discussion of "the hereditary varieties or races of our domestic animals and plants". a response to Darwin's recent encouragement. although Darwin did not. and by July had decided to produce a full technical treatise on species. While Darwin considered Wallace's idea to be identical to his concept of natural selection. Wallace drew no comparison to selective breeding. the papers entitled On the Tendency of Species to form Varieties. and emphasised competition between individuals.[40] Wallace (1823–1913) taken in Darwin wrote to Lyell that "your words have come true with a vengeance. while on holiday at Sandown.[51] ) Darwin had initially decided to call it An abstract of an Essay on the Origin of Species and Varieties Through natural selection. by Wallace and Darwin respectively. were read out but drew little reaction.[50] (eventually Murray paid £180 to Darwin for the 1st edition and by Darwin's death in 1882 the book was in its 6th edition. and an offer to Darwin of 2 ⁄3 of the profits.[46] and wrote parts of it from memory. It enclosed twenty pages describing an evolutionary mechanism. closely related species.. whatever it may amount to. at once write and offer to send [it] to any journal" that Wallace chose. and in the spring of 1856 Lyell urged Darwin to publish his theory to establish priority.[40] Darwin was hard at work on his "big book" on Natural Selection. . Darwin described natural selection as being analogous to the artificial selection practised by animal breeders.[47] Lyell discussed arrangements with publisher John Murray III. he began a "sketch" account. Darwin was torn between the desire to set out a full and convincing account and the pressure to quickly produce a short paper. and on 1 July 1858. Darwin decided to write "an abstract of my whole work". but with Murray's persuasion it was eventually changed to the snappier title: On the Origin of Species.[42] [43] [44] After the meeting. written by Alfred Russel Wallace. historians have pointed out differences. earning Darwin nearly £3000. He decided he did not want to expose his ideas to review by an editor as would have been required to publish in an academic journal. or the Preservation of Favoured Races in the Struggle for Life.''On the Origin of Species'' 131 Publication Events leading to publication An 1855 paper on the "introduction" of species.[41] Lyell and Hooker agreed that a joint paper should be presented at the Linnean Society. for instance. Singapore in 1862 forestalled" and he would "of course. The mechanism was similar to Darwin's own theory..[45] He started work on 20 July 1858. claimed that patterns in the geographical distribution of species and fossils could be explained if every new species always came into existence near an already existing. and on the Perpetuation of Varieties and Species by Natural Means of Selection.[48] who responded immediately to Darwin's letter of 31 March 1859[49] with an agreement to publish the book without even seeing the manuscript. when on 18 June 1858 he received a parcel from Wallace. adding that "all my originality.[2] Here the term "races" is used as an alternative for "varieties" and does not carry the modern connotation of human races—the first use in the book refers to "the several races. with a request to send it on to Lyell if Darwin A photograph of Alfred Russel thought it worthwhile. of the publishing house John Murray.[52] . who was working in Borneo. and notes Darwin's large output of books and busy family life during that time. The book had been offered to booksellers at Murray's autumn sale on Tuesday 22 November. he told many people about his interest in transmutation without causing outrage. and concern about upsetting his clergymen naturalist friends or his pious wife Emma. around 1. He firmly intended to publish.[56] During Darwin's lifetime the book went through six editions.[54] Publication and subsequent editions On the Origin of Species was first published on Thursday 24 November 1859. The third edition came out in 1861. While he was researching. with a number of sentences rewritten or added and an introductory appendix. when Wallace's letter arrived on 18 June 1858. and he may have wanted to be sure he was correct.[57] while the fourth in 1866 had further revisions. Book sales increased from 60 to 250 per month.[58] In January 1871. Reasons suggested have included fear of religious persecution or social disgrace if his views were revealed. It was long thought that Darwin avoided or delayed making his ideas public for personal reasons. which had been coined by the philosopher Herbert Spencer in his Principles of Biology (1864). and Darwin's contemporaries thought the time he took was reasonable. Miscellaneous objections.250 copies were printed but after deducting presentation and review copies. 1. An Historical Sketch of the Recent Progress of Opinion on the Origin of Species.[2] The second edition of 3. His estimate that writing his "big book" would take five years was optimistic.170 copies were available for sale. and all available copies had been taken up immediately. Darwin always finished one book before starting another. Dallas. Darwin was still not ready to publish his theory. It includes a glossary compiled by W.S. published on 10 February 1869.[55] and incorporated numerous corrections as well as a response to religious objections by the addition of a new epigraph on page ii. using the word "evolution" for the first time. The fifth edition. In total. His paper on Glen Roy had proved embarrassingly wrong.[53] A more recent study by science historian John van Wyhe has determined that the idea that Darwin delayed publication only dates back to the 1940s. the price was halved to 7s 6d by printing in a smaller font. David Quammen has suggested all these factors may have contributed.''On the Origin of Species'' 132 Time taken to publish Darwin had his basic theory of natural selection "by which to work" by December 1838. incorporated more changes and for the first time included the phrase "survival of the fittest". priced at fifteen shillings. George Jackson Mivart's On the Genesis of Species listed detailed arguments against natural selection. to address Mivart's arguments. but it was not until September 1854 that he could work on it full time. and the phrase "by the Creator" amended to the closing sentence.[61] . yet almost twenty years later. and claimed it included false metaphysics. with cumulative changes and revisions to deal with counter-arguments raised. Charles Darwin's illness caused repeated delays. Darwin had told Murray of working men in Lancashire clubbing together to buy the 5th edition at fifteen shillings and wanted it made more widely available. a quotation from Charles Kingsley.[59] Darwin made revisions to the sixth edition of the Origin.[61] The sixth edition was published by Murray on 19 February 1872 with "On" dropped from the title.[60] and added a new chapter VII.000 copies was quickly brought out on 7 January 1860. and five for Stationers' Hall copyright. Origin was published in Swedish in 1869. adding controversial themes that Darwin had deliberately omitted. Bronn produced a second edition based on the third English edition and Darwin's suggested additions. Bronn translated "favoured races" as "perfected races". but he had difficulty getting her to remove her notes and was troubled by these editions. Spanish in 1877 and Serbian in 1878. as well as numerous explanatory notes giving her own answers to doubts that Darwin expressed.[64] who got their edition out in mid January 1860. but it is not clear if this referred to the first printing only as there were four that year. Hungarian in 1873–1874.[72] .[68] Darwin corresponded closely with Julius Victor Carus. additional translations had appeared in Italian and Russian. but learnt that two New York publishing firms were already planning to exploit the absence of international copyright to print Origin. and added essays on issues including the origin of life.[69] Darwin's attempts to find a translator in France fell through.[2] [65] The book was widely translated in Darwin's life time.[71] By 1864. In a May letter.[66] Darwin distributed presentation copies in France and Germany. but the German translation published in 1860 imposed Bronn's own ideas. it had appeared in an additional 18 languages. and asked Gray to keep any profits. Asa Gray negotiated with a Boston publisher for publication of an authorised American version. By 1977.[62] Darwin was delighted by the popularity of the book. Polish in 1873. Danish in 1872.[66] In Darwin's lifetime.[68] [70] He remained unsatisfied until a translation by Edmond Barbier was published in 1876. but then died of a heart attack. but problems arose with translating concepts and metaphors.[63] Gray managed to negotiate a 5% royalty with Appleton's of New York. as well as a final chapter on religious implications partly inspired by Bronn's adherence to Naturphilosophie. and some translations American botanist Asa Gray (1810–1888) were biased by the translator's own agenda.[2] A Dutch translation was published in 1860. Darwin mentioned a print run of 2. who published an improved translation in 1867.''On the Origin of Species'' 133 Publication outside Great Britain In the United States. He welcomed the distinguished elderly naturalist and geologist Heinrich Georg Bronn.500 copies. Darwin corresponded with Royer about a second edition published in 1866 and a third in 1870. hoping that suitable applicants would come forward. as translators were expected to make their own arrangements with a local publisher. and the translation by Clémence Royer published in 1862 added an introduction praising Darwin's ideas as an alternative to religious revelation and promoting ideas anticipating social Darwinism and eugenics. and the other two withdrew.[67] In 1862. and sets out the essence of his theory: As many more individuals of each species are born than can possibly survive. and (2) ubiquitous small differences (example: slightly shorter or longer bill of pigeons). any selected variety will tend to propagate its new and modified form. with experts disagreeing and changing their decisions when new forms were found.[78] Starting with the third edition. unknown to Wallace or himself.M. and the arrival of Wallace at the same conclusion. He concludes that "a well-marked variety .[80] in the fourth edition he mentioned that William Charles Wells had done so as early as 1813. anticipated the concept of natural selection in an appendix to a book published in 1831.[81] Variation under domestication and under nature Chapter I covers animal husbandry and plant breeding.' as naturalist. it follows that any being. Darwin discusses contemporary opinions on the origins of different breeds under cultivation to argue that many have been produced from common ancestors by selective breeding. and as.[83] noting that "[t]he diversity of the breeds is something astonishing". and thus be naturally selected. there is a frequently recurring struggle for existence. 'Beagle. in a nod to the religious concerns of his oldest friends. going back to ancient Egypt.[73] harmonising science and religion in accordance with Isaac Newton's belief in a rational God who established a law-abiding cosmos. and to that of the Galápagos tortoises and mockingbirds. under the complex and sometimes varying conditions of life. which led him to "publish this Abstract" of his incomplete work. However.[56] The Introduction establishes Darwin's credentials as a naturalist and author.[74] In the second edition.[77] John Gould's illustration of Darwin's Rhea was published in 1841. consequently. for Darwin the small changes were most important in evolution.S.[79] In that sketch he acknowledged that Patrick Matthew had. yet all were descended from one species of rock pigeon. He outlines his ideas. Darwin refers specifically to the distribution of the species rheas.[75] then refers to John Herschel's letter suggesting that the origin of species "would be found to be a natural in contradistinction to a miraculous process":[76] WHEN on board H. Darwin specifies that the distinction between species and varieties is arbitrary. as it has been called by one of our greatest philosophers. From the strong principle of inheritance.[84] Darwin saw two distinct kinds of variation: (1) rare abrupt changes he called "sports" or "monstrosities" (example: ancon sheep with short legs). He mentions his years of work on his theory. The existence of two rhea species with overlapping ranges influenced Darwin.[85] Both types of hereditary changes can be used by breeders. Darwin added an epigraph from Joseph Butler affirming that God could work through scientific laws as much as through miracles.[82] As an illustration of artificial selection.''On the Origin of Species'' 134 Content Title pages and introduction Page ii contains quotations by William Whewell and Francis Bacon on the theology of natural laws. Darwin prefaced the introduction with a historical sketch that traced the development of evolutionary ideas. he describes fancy pigeon breeding. These facts seemed to me to throw some light on the origin of species—that mystery of mysteries. In Chapter II. if it vary however slightly in any manner profitable to itself. and in the geological relations of the present to the past inhabitants of that continent. I was much struck with certain facts in the distribution of the inhabitants of South America. will have a better chance of surviving. saying: But how..''On the Origin of Species'' may be justly called an incipient species" and that "species are only strongly marked and permanent varieties". and so be enabled to increase in numbers. by the term of Natural Selection. I have called this principle. any variation. can any analogous principle apply in nature? I believe it can and does apply most efficiently. however slight and from whatever cause proceeding. it may be asked." He discusses checks to such increase including complex ecological interdependencies. and suggests that natural selection might do the same. Darwin asks how varieties "which I have called incipient species" become distinct species.[88] in the fifth edition he adds. constitution. if useful. is preserved. nor that the favoured animals were better or higher. P. Darwin emphasises that he used the phrase "struggle for existence" in "a large and metaphorical sense. immigration of others and.[92] Historians have remarked that here Darwin anticipated the modern concept of an ecological niche. if it be in any degree profitable to an individual of any species.[86] He argues for the ubiquity of variation in nature. in its infinitely complex relations to other organic beings and to external nature. by so much will they be better enabled to seize on many and widely diversified places in the polity of nature. descendants of some species became adapted to new conditions. that archetype being a fixed ideal in the mind of God. He remarks that the artificial selection practised by animal breeders frequently produced sharp divergence in character between breeds. he gives examples ranging from plants struggling against drought to plants competing for birds to eat their fruit and disseminate their seeds.[90] Chapter IV details natural selection under the "infinitely complex and close-fitting . natural selection. and will generally be inherited by its offspring .. and is sometimes equally convenient.[83] 135 Struggle for existence. including dependence of one being on another". will tend to the preservation of that individual.[87] Historians have noted that naturalists had long been aware that the individuals of a species differed from one another. is more accurate. by which each slight variation. Darwin and Wallace made variation among individuals of the same species central to understanding the natural world. from the simple circumstance that the more diversified the descendants from any one species become in structure. but merely more adapted to their surroundings. of the Survival of the Fittest."[89] Owing to this struggle for life. "But the expression often used by Mr. and in answer introduces the key concept he calls "natural selection". where suitable variations occurred.[91] Darwin takes as an example a country where a change in conditions led to extinction of some species.[88] He notes that both A. and divergence In Chapter III. and notes that competition is most severe between closely related forms "which fill nearly the same place in the economy of nature"... mutual relations of all organic beings to each other and to their physical conditions of life".[93] He did not suggest that every favourable variation must be selected. He describes the struggle resulting from population growth: "It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms. but had generally considered such variations to be limited and unimportant deviations from the archetype of each species. . Herbert Spencer. and habits. de Candolle and Charles Lyell had stated that all organisms are exposed to severe competition. in order to mark its relation to man's power of selection. bird songs. "The laws governing inheritance are quite unknown. He describes branches falling off as extinction occurred.. he struggled with the theoretical difficulty that novel individual variations would tend to blend into a populationa. and disuse diminishes them. but given the effectiveness of artificial selection. one thing was clear: whatever the exact nature and causes of new variations. and that this also applied in nature. inherited variation could be seen. and accepted this process as increasing the variation available for natural selection. such as the loss of functional wings in some island dwelling insects. Darwin also admitted ignorance of the source of inheritable variations. However.[95] Variation and heredity In Darwin's time there was no agreed-upon model of heredity. which may be effected in the long course of time by nature's power of selection". and Selection in Relation to Sex (1871)."[97] He accepted a version of the inheritance of acquired characteristics (which after Darwin's death came to be called Lamarckism). Using a tree diagram and calculations.[96] in Chapter I Darwin admitted.[99] [100] However. Although Darwin had privately questioned blending inheritance. and similar patterns of variation in distinct species were explained by Darwin as demonstrating common descent.[85] The observation that selection works in domestic animals is not destroyed by lack of understanding of the underlying hereditary mechanism. is the only illustration in the coadaptations between all organic Origin of Species. which tried to explain heredity through his hypothesis of pangenesis. he wrote that he thought "there can be little doubt that use in our domestic animals strengthens and enlarges certain parts. deer antlers. and Chapter V discusses what he called the effects of use and disuse.[101] More detail was given in Darwin's 1868 book on The Variation of Animals and Plants under Domestication. In later additions of Origin. He recounted how Lord Morton's mare apparently demonstrated telegony.''On the Origin of Species'' 136 Darwin proposes sexual selection. and the bright plumage of some male birds.[94] He analysed sexual selection more fully in The Descent of Man. might be produced by natural selection.. driven by competition between males for mates. while new branches formed in "the great Tree of life .[102] and Darwin's concept of selection working on a population with a range of small variations was workable. Darwin knew from observation and experiment that breeders were able to select such variations and produce huge differences in many generations of selection. peacock tails. and that such modifications are inherited". to explain sexually dimorphic features such as lion manes. or tending to revert to an ancestral form. Natural selection was expected to work very slowly in forming new species. but speculated they might be produced by environmental factors. he indicates the "divergence of character" from original species into new species and genera. with its ever branching and beautiful ramifications". Darwin expanded the role attributed to the inheritance of acquired characteristics. he could "see no limit to the amount of change. offspring inheriting characteristics of a previous mate of the female parent. used to show the divergence of species.[98] Darwin stated that some changes that were commonly attributed to use and disuse.[103] It was not until the modern evolutionary synthesis in . beings. Breeding of animals and plants showed related varieties varying in similar ways. to the beauty and infinite complexity of the This tree diagram. one with another and with their physical conditions of life. which could not possibly have been formed by numerous. and that often features seemed non-adaptive because their function was unknown. under pressure from natural selection to economise wax. and the viability and fertility of the hybrids. Darwin said that. slight modifications. —the larvæ of ichneumonidæ feeding within the live bodies of caterpillars. He said some such features could have been by-products of adaptive changes to other features. and. He suggested that bees that make hexagonal cells evolved in steps from bees that made round cells. organs of trifling importance. His examples included two he had investigated experimentally: slave-making ants and the construction of hexagonal cells by honey bees. it may not be a logical deduction. but even rather to support the view. But I can find out no such case. such as the tail of a giraffe. namely. the first being that often no intermediate forms between closely related species are found. but as small consequences of one general law. and in other cases what were considered to be mere varieties of the same species could only be crossed with difficulty. though the theory implies such forms must have existed.[107] He discussed various simple eyes found in invertebrates. flatfish with both eyes on one side and the camouflage of stick insects could not have . leading to the advancement of all organic beings. could have been formed by the modification of some animal with wholly different habits? Can we believe that natural selection could produce. for instance. as examples of how the vertebrate eye could have evolved. organs of such wonderful structure. starting with nothing more than an optic nerve coated with pigment.''On the Origin of Species'' the 1930s and 1940s that a model of heredity became completely integrated with a model of variation. successive. Sometimes what were widely considered to be separate species produced fertile hybrid offspring freely. and he observed that many ant species will collect and store the pupae of other species as food. Darwin concluded: Finally. is it possible that an animal having. —ants making slaves. far from being constant."[110] In the sixth edition Darwin inserted a new chapter VII (renumbering the subsequent chapters) to respond to criticisms of earlier editions. the structure and habits of a bat. vary. varied greatly. of which we hardly as yet fully understand the inimitable perfection?[106] His answer was that in many cases animals exist with intermediate structures that are functional.[104] 137 Difficulties for the theory Chapter VI begins by saying the next three chapters will address possible objections to the theory. often leading to extinction of such forms. He thought it reasonable that species with an extreme dependency on slave workers had evolved in incremental steps. the facts briefly given in this chapter do not seem to me opposed to.[109] Chapter VIII addresses the idea that species had special characteristics that prevented hybrids from being fertile in order to preserve separately created species. Darwin noted that some species of slave-making ants were more dependent on slaves than others. —not as specially endowed or created instincts. Darwin said: Secondly. combined with the small number of individuals of intermediate forms. on the one hand. including his claim that features such as baleen filters in whales. that there is no fundamental distinction between species and varieties.[105] The rest of the chapter deals with whether natural selection could produce complex specialised structures. on the other hand. especially among plants. He presented flying squirrels. when it would be difficult to imagine how intermediate forms could be functional. the difficulty in producing hybrids of related species. as the eye. but to my imagination it is far more satisfactory to look at such instincts as the young cuckoo ejecting its foster-brothers. Much of the chapter responds to George Jackson Mivart's criticisms. and flying lemurs as examples of how bats might have evolved from non-flying ancestors."[108] Chapter VII (of the first edition) addresses the evolution of instincts. then. including the objection that many features of organisms were not adaptive and could not have been produced by natural selection. which serves as a fly-flapper. Darwin concluded: "Finally. as shown by his book on Fertilisation of Orchids that explained how their elaborate structures facilitated pollination by insects. and the behaviours to use them. Darwin concludes: "If it could be demonstrated that any complex organ existed. Darwin attributed this to the competition between different forms. let the strongest live and the weakest die. my theory would absolutely break down. multiply. with related groups of species having a continued existence until extinction. but not at the same rate – some organisms such as Lingula were unchanged since the earliest fossils. Darwin proposed scenarios for the incremental evolution of each feature.[117] Geographic distribution Chapter XI deals with evidence from biogeography. The pace of natural selection would depend on variability and change in the environment.[121] .[117] This distanced his theory from Lamarckian laws of inevitable progress. Recently extinct species were more similar to living species than those from earlier eras. Darwin noted that barriers to migration played an important role in the differences between the species of different regions. but those regions have very different plants and animals. but would still be related to species found on the continent. and William Clift had shown in Australia. many of which he had investigated experimentally. he again cited Principles of Geology and other observations based on sedimentation and erosion. Darwin had no doubt that earlier seas had swarmed with living creatures. Africa.[112] [113] To show that there had been enough time for natural selection to work slowly. but continuing discoveries gave him well founded hope that new finds would occasionally reveal transitional forms.[113] [118] but other scholars have preferred to emphasise Darwin's commitment to gradualism. fossils from recent geological periods resembled species still living in the same area. as is so commonly and notoriously the case. there could only be fragmentary knowledge of geological formations. starting with the observation that differences in flora and fauna from separate regions cannot be explained by environmental differences alone. The coastal sea life of the Atlantic and Pacific sides of Central America had almost no species in common even though the Isthmus of Panama was only a few miles wide. spread over vast periods of time. and fossil collections were very poor. and even families are confined to the same areas. He went on to say: "On this principle of inheritance with modification. and were followed by increasingly diverse and specialised forms. without the innumerable transitional fossils expected from gradual changes. These species would become modified over time. including an estimate that erosion of The Weald had taken 300 million years. extending the history of life back for billions of years. and Australia all have regions with similar climates at similar latitudes. Evolved local varieties which migrated into a wider area would seem to be the sudden appearance of a new species.[114] The initial appearance of entire groups of well developed organisms in the oldest fossil-bearing layers. His explanation was a combination of migration and descent with modification. and Darwin observed that this was a common pattern. Darwin did not expect to be able to reconstruct evolutionary history."[120] Darwin explained how a volcanic island formed a few hundred miles from a continent might be colonised by a few species from that continent. now known as the Cambrian explosion. since few areas had been geologically explored. South America. and as he had seen in South America. Darwin discussed ways that species could be dispersed across oceans to colonise islands. Darwin expected species to change slowly. Darwin borrowed Charles Lyell's argument in Principles of Geology that the record is extremely imperfect as fossilisation is a very rare occurrence.''On the Origin of Species'' evolved through natural selection because intermediate stages would not have been adaptive.[112] It has been argued that this anticipated the punctuated equilibrium hypothesis. posed a problem. matching the branching of common descent from an ancestor.[116] Chapter X examines whether patterns in the fossil record are better explained by common descent and branching evolution through natural selection than by the individual creation of fixed species.[119] He cited Richard Owen's findings that the earliest members of a class were a few simple and generalised species with characteristics intermediate between modern forms. then not reappearing.[115] Fossil evidence of pre-Cambrian life has since been found. but stated that he had no satisfactory explanation for the lack of fossils.[111] 138 Geologic record Chapter IX deals with the fact that the geologic record appears to show forms of life suddenly arising. The species found in one area of a continent are more closely allied with species found in other regions of that same continent than to species found on other continents.[112] Patterns of extinction matched his theory. we can understand how it is that sections of genera. whole genera. . "What can be more curious than that the hand of a man. from the war of nature.. formed for grasping. having been originally breathed into a few forms or into one. on the view that the natural system is founded on descent with modification. and that. namely.. here he ventures a cautious hint that psychology would be put on a new foundation and that "Light will be thrown on the origin of man". Thus. After a brief discussion of freshwater species... whether of land or water. and. for example on some islands roles played by mammals on continents were played by other animals such as flightless birds or reptiles.. We can thus understand the localisation of sub-genera. genera. in the same relative positions?"[124] He notes that animals of the same class often have extremely similar embryos. as I have endeavoured to show. and families. He says. the paddle of the porpoise. in so far. and how it is that under different latitudes. that community of descent is the hidden bond which naturalists have been unconsciously seeking.[123] Darwin discusses morphology. are found only in embryonic stages. and are being. and should include the same bones. it returns to oceanic islands and their peculiarities. embryology. all true classification is genealogical. so different from each other. have all been produced by laws acting around us .. but of these a great number should be endemic or peculiar.. We can thus understand the high importance of barriers.[122] 139 Classification. and deserts. from famine and death. which separate our several zoological and botanical provinces.''On the Origin of Species'' Chapter XII continues the discussion of biogeography. such as the wings of flightless birds and the rudiments of pelvis and leg bones found in some snakes.. and with worms crawling through the damp earth. why oceanic islands should have few inhabitants.. the production of the higher animals. evolved. . of the forests. with birds singing on the bushes. are in so mysterious a manner linked together by affinity. morphology. including the importance of homologous structures. the most exalted object which we are capable of conceiving. Darwin concludes: All the foregoing rules and aids and difficulties in classification are explained. He remarks that some rudimentary organs. with various insects flitting about. and to reflect that these elaborately constructed forms. the leg of the horse. whilst this planet has gone cycling on according to the fixed law of gravity. that of a mole for digging. clothed with many plants of many kinds. together with subsequent modification and the multiplication of new forms. from so simple a beginning endless forms most beautiful and most wonderful have been. should all be constructed on the same pattern. and are likewise linked to the extinct beings which formerly inhabited the same continent . and Darwin concludes by hoping that his theory might produce revolutionary changes in many fields of natural history. marshes.. Darwin discusses rudimentary organs. The summary of both chapters says: . rudimentary organs Chapter XIII starts by observing that classification depends on species being grouped together in a multilevel system of groups and sub groups based on varying degrees of resemblance. are those which have been inherited from a common parent. the inhabitants of the plains and mountains. On these same principles. such as teeth in baleen whales. Although he avoids the controversial topic of human origins in the rest of the book so as not to prejudice readers against his theory.[126] . After discussing classification issues. if I do not greatly deceive myself. I think all the grand leading facts of geographical distribution are explicable on the theory of migration (generally of the more dominant forms of life). we can understand. with its several powers. for instance in South America. There is grandeur in this view of life.[125] Darwin ends with a passage that became well known and much quoted: It is interesting to contemplate an entangled bank. directly follows. and dependent on each other in so complex a manner. Concluding remarks The final chapter reviews points from earlier chapters. that the characters which naturalists consider as showing true affinity between any two or more species. and the wing of the bat. "[128] [134] While the book was readable enough to sell. and many parts of the book abound in information. "Much of Mr.[129] [130] Later chapters provide evidence that evolution has occurred. All. its ability to produce the effects of interest. is by no means of this description. Unlike the still-popular Vestiges.[128] Therefore the first four chapters lay out his case that selection in nature. Quammen advised that later editions were weakened by Darwin making concessions and adding details to address his critics. is analogous to the selection of variations under domestication. and to show that natural selection had been the chief agent of change.''On the Origin of Species'' 140 Structure and style Nature and structure of Darwin's argument Darwin's aims were twofold: to show that species had not been separately created. its dryness ensured that it was seen as aimed at specialist scientists and could not be dismissed as mere journalism or imaginative fiction. it was more approachable than the big book on natural selection Darwin had been working on.[127] He knew that his readers were already familiar with the concept of transmutation of species from Vestiges. and recommended the first edition. in other places his writing was beautiful. but noted that Darwin's literary style was uneven: in some places he used convoluted sentences that are difficult to read. and the case studies and observations are presented in a narrative style unusual in serious scientific books. writing which to comprehend requires concentrated attention and some preparation for the task.[130] [131] [132] The structure of Darwin's argument showed the influence of John Herschel. which broadened its audience. however. Darwin presents supporting facts drawn from many disciplines. showing that his theory could explain a myriad of observations from many fields of natural history that were inexplicable under the alternate concept that species had been individually created. Darwin had long been immersed in the literary forms and practices of specialist science. and its ability to explain a wide range of observations.[128] David Quammen has described the book as written in everyday language for a wide audience. though the closing sentence clearly hinted at cosmic progression. whose philosophy of science maintained that a mechanism could be called a vera causa (true cause) if three things could be demonstrated: its existence in nature.[133] Literary style The Examiner review of 3 December 1859 commented. and that the accumulation of adaptive variations provides a scientifically testable mechanism for evolutionary speciation. easy to comprehend and both instructive and entertaining.' that is.[136] . and made effective use of his skills in structuring arguments. Darwin's volume is what ordinary readers would call 'tough reading. and his introduction ridicules that work as failing to provide a viable mechanism. caused by the struggle for existence. adaptive evolution without directly proving that selection is the mechanism.[135] James T. it avoided the narrative style of the historical novel and cosmological speculation. but parts are almost lyrical. supporting the idea of branching. which would have been encumbered by scholarly footnotes and much more technical detail. Costa said that because the book was an abstract produced in haste in response to Wallace's essay. He added that parts of Origin are dense. other mechanisms implying more progress or purpose were more acceptable. but the transmutational ideas of Lamarck and the vague "law of development" of Vestiges had not found scientific favour. He popularised the terms evolution and survival of the fittest.[138] but had influenced a wide public readership into believing that nature and human society were governed by natural laws. Its proponents made full use of a surge in the publication of review journals.[141] Human evolution became central to the debate and was strongly argued by Huxley who featured it in his popular "working-men's lectures". though it failed to match the continuing sales of Vestiges. Herbert Spencer had already incorporated Lamarckism into his popular philosophy of progressive free market human society.[143] British cartoonists presented Darwin's theory in an unthreatening way. evolutionism was triumphant. while opponents held to the idealist school of William Whewell's Philosophy of the Inductive Sciences. and by 1875. and many thought Spencer was central to evolutionary thinking. Darwin had avoided the subject of human evolution. Despite this. Alfred Russel Wallace discussed evidence from his Malay archipelago research. In the 1870s iconic caricatures of Darwin with an ape or monkey body emphasised his significance in transforming ideas. not just his own ideas. including an 1864 paper with an evolutionary explanation for the Wallace line. By the mid 1870s.[25] The Origin of Species as a book of wide general interest became associated with ideas of social reform. with no sharp line between scientific issues and ideological. which had been rejected by scientists. Henry Walter Bates presented research in 1861 that explained insect mimicry using natural selection. and it was given more popular attention than almost any other scientific work.[140] Darwin's book legitimised scientific discussion of evolutionary mechanisms. but Darwin had to be taken seriously as a prominent and respected name in science. Darwin presented natural selection as a scientifically testable mechanism while accepting that other mechanisms such as inheritance of acquired characters were possible. Darwin's scientific method was also disputed.[142] The naturalism of natural selection conflicted with presumptions of purpose in nature and while this could be reconciled by theistic evolution. His strategy established that evolution through natural laws was worthy of scientific study. Darwin did not publish his own views on this until 1871. with his proponents favouring the empiricism of John Stuart Mill's A System of Logic.[139] Much of the initial reaction was hostile.[145] .[139] With the exception of a brief hint in the final chapter.''On the Origin of Species'' 141 Reception The book aroused international interest[138] and a widespread debate. Impact on the scientific community Scientific readers were already aware of arguments that species changed through processes that were subject to laws of nature. and the newly coined term Darwinism was used to cover the whole range of evolutionism. social and religious implications. in which investigation could begin with the intuitive truth that species were fixed objects created by design.[144] Early support for Darwin's ideas came from the findings of field naturalists studying biogeography and ecology. and contributed to widespread identification of evolutionism with [137] Darwinism. There was much less controversy than had greeted the 1844 publication Vestiges of Creation. the first review claimed it made a creed of the "men from monkeys" idea from Vestiges. including Joseph Dalton Hooker in 1860. and Asa Gray in 1862. most scientists accepted that evolution occurred but few thought natural selection was significant. but higher mental faculties could not have evolved through a purely material process. with new species appearing by natural birth. as yet. Huxley had emphasised anatomical similarities between apes and humans.[155] Others. the German morphologist Ernst Haeckel would convince Huxley that comparative anatomy and palaeontology could be used to reconstruct evolutionary genealogies. Huxley strongly supported Darwin on evolution.[156] . though he called for experiments to show whether natural selection could form new species. Their disagreement over human origins came to the fore at the British Association for the Advancement of Science meeting featuring the legendary 1860 Oxford evolution debate.[154] In two years of acrimonious public dispute that Charles Kingsley satirised as the "Great Hippocampus Question" and parodied in The Water-Babies as the "great hippopotamus test". thought that humans shared a common ancestor with apes.[147] Despite this. and initiated the X Club. Huxley showed that Owen was incorrect in asserting that ape brains lacked a structure present in human brains.[150] Huxley gained influence.[153] Since 1858. an idealist who had shifted to the view in the 1850s that the history of life was the gradual unfolding of a divine plan. hardly penetrated" and coining the term "Darwinism" as part of his efforts to secularise and professionalise science. Hooker and Darwin. and his article in the Huxley used illustrations to show that humans and apes had the same basic skeletal [146] April 1860 Westminster Review promoted structure. included the Duke of Argyll who explained beauty in plumage by design. which used the journal Nature to promote evolution and naturalism.[148] [149] praising Darwin for "extending the domination of Science over regions of thought into which she has. and questioned if Darwin's gradualism was sufficient without sudden leaps to cause speciation. but also signalled acceptance of a kind of evolution as a teleological plan in a continuous "ordained becoming". shaping much of late Victorian science. contesting Owen's view that humans were a separate sub-class.''On the Origin of Species'' 142 Evolution had less obvious applications to anatomy and morphology. Huxley wanted science to be secular. Darwin published his own explanation in the Descent of Man (1871). scientific naturalism over natural theology. without religious interference. and at first had little impact on the research of the anatomist Thomas Henry Huxley. Others that rejected natural selection.[152] Owen's review of the Origin in the April 1860 Edinburgh Review bitterly attacked Huxley. but supported "creation by birth". including Charles Lyell and Alfred Russel Wallace. Later.[147] [151] The leading naturalist in Britain was the anatomist Richard Owen. including Karl von Nägeli's insistence that a trivial characteristic with no adaptive advantage could not be developed by selection. Mivart supported directed evolution. were accepted by German biologists accustomed to ideas of homology in morphology from Goethe's Metamorphosis of Plants and from their long tradition of comparative anatomy. Darwin tried to meet these objections in the 5th edition. aiming to synthesise Darwin's ideas with those of Lamarck and Goethe while still reflecting the spirit of Naturphilosophie. Edward Drinker Cope and Alpheus Hyatt reconciled this view with evolutionism in a form of neo-Lamarckism involving recapitulation theory. French speaking naturalists in several countries showed appreciation of the much modified French translation by Clémence Royer.''On the Origin of Species'' Impact outside Great Britain Evolutionary ideas. Darwin made considerable changes to the sixth edition.[157] Asa Gray promoted and defended Origin against those American naturalists with an idealist approach. Darwin was cautious about such histories. unlike [157] Darwin's branching evolutionary tree. In response. where any scientists supporting evolutionary ideas opted for a form of Lamarckism. Ernst Haeckel was particularly ardent. but relegated natural selection to a minor role as they believed evolution was purposeful and progressive. but Darwin's ideas had little impact in France. although the Malthusian aspects were felt to be relatively unimportant. most scientists accepted evolution.[158] 143 Haeckel showed a main trunk leading to mankind with minor branches to various animals. Darwin accepted blending inheritance. but Fleeming Jenkin calculated that as it mixed traits. classifying as species what others considered merely varieties. and scientists were quick to take it into account.[66] Challenges to natural selection There were serious scientific objections to the process of natural selection as the key mechanism of evolution.[67] [158] Their ambitious programme to reconstruct the evolutionary history of life was joined by Huxley and supported by discoveries in palaeontology. but enthused political radicals. and compiled scientific and religious objections to natural selection. The political economy of struggle was criticised as a British stereotype by Karl Marx and by Leo Tolstoy.[59] [159] By the mid 1870s. although not natural selection. Darwin conceded that these could be linked to adaptive characteristics. which embodied a progressive. who had the character Levin in his novel Anna Karenina voice sharp criticism of the morality of Darwin's views. forms of orthogenesis claiming that species had an inherent tendency to change in a particular direction. natural selection could not accumulate useful traits. was called neo-Darwinism. that natural selection was the only mechanism. The range of evolutionary theories during "the eclipse of Darwinism" included forms of "saltationism" in which new species were thought to arise through "jumps" rather than gradual adaptation. The problems of the age of the Earth and heredity were only resolved in the 20th century. Haeckel used embryology extensively in his recapitulation theory. Bronn's alterations in his German translation added to the misgivings of conservatives. and forms of neo-Lamarckism in which inheritance of acquired characteristics led to progress. almost linear model of evolution. notably Louis Agassiz who viewed every species as a distinct fixed unit in the mind of the Creator. The minority view of August Weismann.[70] The intelligentsia in Russia had accepted the general phenomenon of evolution for several years before Darwin had published his theory. and had already noted that von Baer's laws of embryology supported his idea of complex branching. His estimate that the age of the Earth allowed gradual evolution was disputed by William Thomson (later awarded the title Lord Kelvin). It was thought that the . who calculated that it had cooled in less than 100 million years. some liberal clergymen interpreted natural selection as an instrument of God's design. Eventually it was realised that supernatural intervention could not be a scientific explanation. having been originally breathed by the Creator into a few forms or into one".[56] Darwin's view at the time was of God creating life through the laws of nature.[181] [182] The liberal theologian Baden Powell defended evolutionary ideas by arguing that the introduction of new species should be considered [165] a natural rather than a miraculous process.[160] [161] 144 Religious attitudes The book produced a wide range of religious responses at a time of changing ideas and increasing secularisation. defined narrowly as including rejection of design.[162] but defence of the argument from design and natural theology was central to debates over the book in the English speaking world.[176] [177] Asa Gray responded that this charge misrepresented Darwin's text. which from then on read "life. pointed out that early Church Fathers had not interpreted Genesis literally in this area. four noted authors of The Fundamentals were explicitly open to the possibility that God created through evolution.[166] [167] In the second edition of January 1860.''On the Origin of Species'' rediscovery of Mendelian inheritance invalidated Darwin's views.[162] In the Church of England. Developments in geology meant that there was little opposition based on a literal reading of Genesis. Charles Hodge argued in his 1874 critique "What is Darwinism?" that "Darwinism". it quickly became central to the debate as mental and moral qualities were seen as spiritual aspects of the immaterial soul. or modus operandi.[166] [174] [175] Theistic evolution became a popular compromise. and St. starting with Mivart. Natural Selection is not inconsistent with Natural Theology. Darwin quoted Kingsley as "a celebrated cleric". but other Catholic writers. The issues raised were complex and there was a large middle ground. was atheism though he accepted that Asa Gray did not reject design. George Jackson Mivart was among those accepting evolution but attacking Darwin's naturalistic mechanism. or viewing evolution as a purposeful and progressive ascent to mankind's position at the head of nature.[171] Baden Powell praised "Mr Darwin's masterly volume [supporting] the grand principle of the self-evolving powers of nature". Some conservative Roman Catholic writers and influential Jesuits opposed evolution in the late 19th and early 20th century. This conflict could be reconciled by supposing there was some supernatural intervention on the path leading to humans.[162] Even though the book had barely hinted at human evolution.[162] While many conservative theologians accepted evolution. and while some such as Louis Agassiz were strongly opposed to the ideas in the book. with the cleric Charles Kingsley seeing it as "just as noble a conception of Deity". design.[179] but fundamentalism inspired the American creation–evolution controversy that began in the 1920s. . of the first cause.[178] By the early 20th century. and it was believed that animals did not have spiritual qualities. which stated that evolution was not inconsistent with Catholic teaching.[163] [164] Natural theology was not a unified doctrine. with its several powers.[169] [170] and even in the first edition there are several references to "creation". and added the phrase "by the Creator" to the closing sentence.[168] While some commentators have taken this as a concession to religion that Darwin later regretted.[180] The Vatican stated its official position in a 1950 papal encyclical.[173] and published a pamphlet defending the book in terms of theistic evolution. Asa Gray argued that evolution is the secondary effect.[172] In America. and naturalistic mechanisms such as neo-Lamarckism were favoured over natural selection as being more compatible with purpose. others sought a reconciliation in which evolution was seen as purposeful. • Transmutation of species • Modern evolutionary synthesis • History of evolutionary thought • History of biology . about his life and work. published in 1871 and his second great book on evolutionary theory. biogeography. an exhaustive external index published in 1981. cladistics. The text of Origin itself has been subject to much analysis including a variorum. phylogenetics and other fields.''On the Origin of Species'' 145 Modern influence Various alternative evolutionary mechanisms favoured during "the eclipse of Darwinism" became untenable as more was learned about inheritance and mutation.[189] See also • Charles Darwin bibliography • The Complete Works of Charles Darwin Online • The Descent of Man. Darwin's theory of evolution by natural selection. The full significance of natural selection was at last accepted in the 1930s and 1940s as part of the modern evolutionary synthesis. The theory explains the diversity of living organisms and their adaptation to the environment. Fisher. and scholars have generated an extensive literature. a religion-based political controversy has developed over how evolution is taught in schools.[187] Worldwide commemorations of the 150th anniversary of the publication of On the Origin of Species and the bicentenary of Darwin's birth were scheduled for 2009[188] . vestigiality. parallels in embryonic development. first published in 1959.B.[185] Interest in Darwin's writings continues.S. During that synthesis biologists and statisticians.[161] A modern phylogenetic tree based on genome analysis shows the three-domain Modern evolutionary theory continues to system. biological homologies. detailing the changes made in every edition. with unrivalled explanatory power. They celebrate the ideas which "over the last 150 years have revolutionised our understanding of nature and our place within it". It makes sense of the geologic record. Haldane. including R. the Darwin Industry. A. with its tree like model of branching common descent. has become the unifying theory of the life sciences. and Selection in Relation to Sex. it has also become essential to applied sciences such as medicine and agriculture. develop.[186] and a concordance. merged Darwinian selection with a statistical understanding of Mendelian genetics.[183] [184] Despite the scientific consensus. Sewall Wright and J. especially in the United States. uk/ content/ frameset?viewtype=side& itemID=F373& pageseq=8) Freeman 1977 Mayr 1982. J. 47–54 [11] Bowler 2003. org. pp. uk/ content/ frameset?viewtype=side& itemID=CUL-DAR121. D. 67. [46] Darwin 2006. and Gayle. C. pp. . 27–36. R. pp. 77–80 [9] Bowler 2003. ii. 186–187 (http:/ / darwin-online. 39–42. 148–149 [17] Browne 1995. pp. 134–135 (http:/ / darwin-online. p. 74 (http:/ / darwin-online. 1–13. and Geology 16: 184–196. p. van Wyhe 2008. translated by Hardie. 436–437 [33] van Wyhe 2007. uk/ entry-2303). . 7–10 (http:/ / darwin-online. 461–465 [36] Bowler 2003. 133–140 [18] Larson 2004. org. pp. 240–244 [25] van Wyhe 2009 [26] Larson 2004. Eldredge 2006 [22] Quammen 2006. pp. org. C. retrieved 2008-11-24 [35] Browne 1995. html#back-mark-814. 59–62 [19] Darwin 1845. pp. 117–121 (http:/ / darwin-online. 44. pp. 188 [34] Darwin Correspondence Project – Letter 814—Darwin. 129 [13] Bowler 2003. . f5). org. org. Darwin Correspondence Project. 26. p. pp.. -& pageseq=112). including Zoology. Darwin's Notebook D: Transmutation of species. 24–25 [23] Herbert 1980. pp. pp. uk/ content/ frameset?viewtype=text& itemID=CUL-DAR123. 436 [31] Desmond & Moore 1991. 2. 34–35 [15] Browne 1995. darwinproject. 5 July (1858)" (http:/ / www. retrieved 2009-04-08 [27] Darwin's Notebook E: Transmutation of species. pp. to Hooker. 175–176 [45] "Letter 2303 — Darwin. mit. 57–62. D.''On the Origin of Species'' 146 Notes [1] [2] [3] [4] [5] Darwin 1859. pp. pp. pp. . pp. 45 [8] Bowler 2003. R. . org. pp.. Alfred R. pp. 1-76& pageseq=70) . uk/ content/ frameset?viewtype=text& itemID=F1497& pageseq=121) [38] Quammen 2006. org. 205–208 (http:/ / darwin-online. pp. xix–xx (http:/ / darwin-online. org. html). Retrieved 2010-09-07. pp. 70. pp. 43. p. 27. 84–90 [10] Desmond 1989. uk/ content/ frameset?viewtype=text& itemID=F1583e& pageseq=9). . (7 Jan 1845) (http:/ / www. ac. 135–158 [41] Darwin Correspondence Project – Letter 2285—Darwin to Lyell (June 1858) (http:/ / www. org. uk/ darwinletters/ calendar/ entry-814. 115–117 [14] Desmond & Moore 1991. pp. iii (http:/ / darwin-online. darwinproject. Botany. 120 (http:/ / darwin-online. xiii (http:/ / darwin-online. Annals and Magazine of Natural History. -& pageseq=63). K. J. pp. org. P. R. retrieved 2008-03-15 [42] Larson 2004. pp. uk/ content/ frameset?viewtype=text& itemID=A544& pageseq=10) [30] Browne 1995. Darwin's Notebook B: Transmutation of species. Physics (http:/ / classics. p. 479–480 Darwin 1872. retrieved 2009-03-14 [28] Darwin 1958. 91. 244–250 [21] Keynes 2000. org. pp. to Hooker. retrieved 2009-03-16 [24] Desmond & Moore 1991. -& pageseq=1). 138–142 [39] Wallace. retrieved 2009-04-23 [6] Forster & Marston 1999. edu/ Aristotle/ physics. 80–88 [16] Bowler 2003. pp. 135–140 [37] Bowler 2003. 26–27 [7] Bowler 2003. . "On the law which has regulated the introduction of new species" (http:/ / darwin-online. pp. 36 verso (http:/ / darwin-online. 292 [32] Browne 1995. p. and hosted by MIT's Internet Classics Archive. org. pp. ac. pp. p. (1855). pp. 169–173 Darwin 1958. html). uk/ content/ frameset?viewtype=text& itemID=F391& pageseq=18) Aristotle. 111–114 [12] Browne 1995. p. uk/ content/ frameset?viewtype=text& itemID=CUL-DAR124. uk/ content/ frameset?viewtype=text& itemID=F1840& pageseq=22). pp. uk/ darwinletters/ calendar/ entry-2285. uk/ content/ frameset?viewtype=text& itemID=F1497& pageseq=124) [29] van Wyhe 2007. uk/ content/ frameset?itemID=F14& viewtype=text& pageseq=218) [20] Browne 1995. 75 (http:/ / darwin-online. uk/ content/ frameset?viewtype=text& itemID=A11& pageseq=1). [40] Quammen 2006. darwinproject. 162–163 [44] Bowler 2003. R. 36. org. pp. pp. p. 66–70. ac. uk/ content/ frameset?viewtype=side& itemID=CUL-DAR158. pp. 74–75 [43] Quammen 2006. to Gray. 256–259 [67] Browne 2002. 20–28 (http:/ / darwin-online. p. R. p.1345. 95–96 [57] Darwin 1861. C. uk/ darwinletters/ calendar/ entry-2665. p. org. com/ files/ aT0Fpi7QEAJXZty6RRr0*ayjT5aN-CLIR1HAnom7DRBt*Z64Ko2B*I88LPx*h2otwLvz5Vp*IKVqHQunTD*sItKN3P7MUMve/ TeylerWinklerDarwin. Principles of Geology vol 2. pp. uk/ charles-darwin-and-john-murray). [61] Desmond & Moore 1991. darwinproject. darwinproject. Darwin. pp. . ning." Spencer 1864. pp. Retrieved 2010-09-07. darwinproject. uk/ content/ frameset?viewtype=text& itemID=A1& pageseq=102) [73] Darwin 1859. or the preservation of favoured races in the struggle for life. Freeman 1977 [62] Darwin Correspondence Project – Letter 2592—Darwin. pp. uk/ books?id=SRkRAAAAYAAJ& pg=PA444& ci=84. [79] Darwin & Costa 2009. 492 [66] Browne 2002. 7 [83] Quammen 2006. ac. R. by T. 28 Jan (1860) (http:/ / www. [86] Darwin 1859. uk/ content/ view/ 28/ 38/ ). p. 15 (http:/ / darwin-online. C. Winkler.. Darwin Correspondence Project. uk/ content/ frameset?viewtype=text& itemID=F385& pageseq=21). p. pp. to Gray. p. Retrieved 2010-09-07. II (http:/ / darwin-online. Charles. ac. google. ac. org.C. doc) Lecture by Marijn van Hoorn (http:/ / teylersmuseum. 23 April 2009 [72] Freeman 1977. 510 [76] van Wyhe 2007. 592–593. 1 (http:/ / darwin-online. p. ning. darwinproject. Princeton University Press. 84–92 [54] van Wyhe 2007 [55] Darwin 1958. org. uk/ content/ frameset?itemID=F373& viewtype=text& pageseq=20). xiii (http:/ / darwin-online. . org. p. [81] Darwin 1866. darwinproject. R. uk/ darwinletters/ calendar/ entry-2592. to Murray. pp.57& source=bookclip) [59] Mivart 1871 [60] Used in the transformist sense by Charles Lyell in 1832. uk/ content/ frameset?itemID=F373& viewtype=text& pageseq=75) 147 . volume 14: 1866 (http:/ / www. retrieved 2008-12-06 [63] Darwin Correspondence Project – Letter 2665—Darwin. 1 [80] Darwin 1861. org. xiv-xv (http:/ / darwin-online. uk/ content/ frameset?itemID=F373& viewtype=text& pageseq=16). p. Darwin (http:/ / api. 21 Dec (1859) (http:/ / www. C. retrieved 2009-03-06 [69] Darwin Correspondence Project – The correspondence of Charles Darwin. org. 31 Mar (1859)" (http:/ / www. html#back-mark-2665. retrieved 2008-12-06 [64] Darwin Correspondence Project – Letter 2706—Gray. . 2010. D. pp. darwinproject. darwinproject. 142–144 [71] Ch. ac. darwinproject. uk/ content/ frameset?itemID=F373& viewtype=side& pageseq=7). ii (http:/ / darwin-online. 582. R. p. [52] Darwin 1859. uk/ entry-2437). transl. uk/ content/ view/ 32/ 38/ ). xiii (http:/ / darwin-online. 44–59 Chap. 20 Feb 1860 (http:/ / www. C. John (b). uk/ content/ frameset?itemID=F1497& viewtype=text& pageseq=126) [56] Browne 2002. Het ontstaan der soorten van dieren en planten door middel van de natuurkeus of het bewaard blijven van bevoorregte rassen in de strijd des levens. darwinproject. [78] Darwin 1859. pp. org. f3). R. uk/ content/ frameset?itemID=F381& viewtype=text& pageseq=20) [58] "This survival of the fittest. volume 10: 1862 (http:/ / www. [50] "Letter 2443 — Murray. Asa to Darwin. is that which Mr. . Prague. 577. Retrieved 2010-09-07. Retrieved 2010-09-07. org. 197 [77] Darwin 1859. 44 [88] Darwin 1859. org. uk/ entry-2443). uk/ darwinletters/ calendar/ entry-2706. [74] Phipps 1983 [75] Secord 2000. 15 Mar (1859)" (http:/ / www. p. C.. and previously by Darwin in The Descent of Man in 1871. J. darwinproject. Retrieved 2010-09-07. org. Darwin Correspondence Project.817. Darwin Correspondence Project. Winkler (Haarlem 1860) Source: Teyler. John (b) to Darwin.. p. com/ profiles/ blogs/ 142-teyler-winkler-darwin) MA at the Congress of the European Botanical and Horticultural Libraries Group. 101–111 (http:/ / darwin-online. html#mark-2592. ac. pp. ac. 11. [87] Darwin & Costa 2009. III (http:/ / darwin-online. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=59). [48] "Letter 2437 — Darwin. . pp. Darwin Correspondence Project. retrieved 2009-03-06 [70] Browne 2002. Asa. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=30) [53] Quammen 2006.49. . C. pp. 590. [82] Darwin & Costa 2009. 122 (http:/ / darwin-online. uk/ content/ frameset?viewtype=text& itemID=F381& pageseq=20). . 5 (http:/ / darwin-online. pp. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=35) [85] David Reznick (2009) The Origin Then and Now. 140–142 [68] Darwin Correspondence Project – The correspondence of Charles Darwin. html). co. . ac. [51] "Charles Darwin and his publisher" (http:/ / www. . R. 1 Apr 1859" (http:/ / www. p. which I have here sought to express in mechanical terms. Darwin has called 'natural selection'. Darwin Correspondence Project. to Hooker. p. 444–445 (http:/ / books. f9). org. [49] "Letter 2441 — Darwin. uk/ entry-2432). p. uk/ entry-2441). . 184–186 [84] Darwin 1859. to Lyell. p. 28 Mar (1859)" (http:/ / www. R. ac. Asa. C. 60–61 Chap.''On the Origin of Species'' [47] "Letter 2432 — Darwin. org. ac. ac. retrieved 2008-12-06 [65] Desmond & Moore 1991. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=330) [118] Rhodes 1987 [119] Darwin & Costa 2009. pp. 312–345 (http:/ / darwin-online. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=324) [116] Schopf 2000 [117] Darwin 1859. pp. . uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=205) [109] Darwin 1859. 171–178 (http:/ / darwin-online. org. 196 Darwin 1859. co. org. uk/ content/ frameset?itemID=F373& viewtype=text& pageseq=133). p. pp. 88–89 [133] Darwin & Costa 2009. [99] Larson 2004. org. 87–101 (http:/ / darwin-online. "An early flowering of genetics | Books" (http:/ / www. 200–201 [104] Bowler 1989 [105] Darwin 1859. 92 Darwin & Costa 2009. org. ix [137] Browne 2002. pp. 346–382 (http:/ / darwin-online. 420 (http:/ / darwin-online. p. Punctuated Equilibria (http:/ / www. 159–167 Darwin 1859. 488 (http:/ / darwin-online. 182 [113] Wesley R. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=300) [115] Darwin 1859. pp. pp. org. 187–190 (http:/ / darwin-online. pp. p. 62–76 (http:/ / darwin-online. pp. org. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=426) [123] Darwin 1859. 200–201 [132] Larson 2004. pp. 177–180 [140] Browne 2002. 80 (http:/ / darwin-online. pp. pp. p. org. evolution). pp. 171–172 (http:/ / darwin-online. [103] Bowler 2003. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=189) [107] Darwin 1859. p. p. 190. [96] Larson 2004. 282–287 (http:/ / darwin-online. 306–308 (http:/ / darwin-online. org. org. 134 (http:/ / darwin-online. p. 183–188 [130] Bowler 2003. org. 1& pageseq=165) [128] Secord 2000. org. pp. org. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=77) [91] Darwin 1859. p. org. p. org. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=152). pp. 434 (http:/ / darwin-online. pp. org. pp. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=149) [101] Quammen 2006. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=189) [106] Darwin 1859. 189 [94] Darwin 1859. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=452) [125] Browne 2002. 245–278 (http:/ / darwin-online. 176–181 [136] Darwin & Costa 2009. 488 Quammen 2006. 108 [120] Darwin 1859. pp. p. pp. org. pp. 180–181 (http:/ / darwin-online. p. pp. Retrieved 2010-10-24. pp. pp. pp. pp. talkorigins. 13 (http:/ / darwin-online. pp. org. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=95) [92] Darwin 1859. 376–379 [138] van Wyhe 2008. uk/ content/ frameset?viewtype=text& itemID=F391& pageseq=196) [112] Bowler 2003. 102–103 148 . org. pp. org. pp. uk/ content/ frameset?viewtype=text& itemID=F937. xvii [134] Crawford 1859 [135] Quammen 2006. 117–130 (http:/ / darwin-online. p. pp. pp. p. p. org. . uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=368) [121] Darwin 1859. 408–409 (http:/ / darwin-online. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=506) [126] Darwin 1859. org. pp. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=364) [122] Darwin 1859. 86–87 [100] Darwin 1859. 152 (http:/ / darwin-online. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=261) [110] Darwin 1859. uk/ content/ frameset?itemID=F373& viewtype=text& pageseq=102) [95] Darwin 1859. 350–351 (http:/ / darwin-online. org. p. p. The Guardian. p. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=177) [102] Richard Dawkins (8 February 2003). 508–511 [129] Quammen 2006. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=438) [124] Darwin 1859. org. 48–49 [139] Bowler 2003. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=507) [127] Darwin 1871. html#pe-vs-pg). 112 (http:/ / darwin-online. org. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=198) [108] Darwin 1859. guardian. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=263) [111] Darwin 1872. uk/ content/ frameset?itemID=F373& viewtype=text& pageseq=28) [98] Darwin 1859. 168–205 (http:/ / darwin-online. org. 85 [97] Darwin 1859. pp. org. org. 72 (http:/ / darwin-online. 131–150 (http:/ / darwin-online. org/ faqs/ punc-eq. pp. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=127) [93] Quammen 2006.''On the Origin of Species'' [89] Darwin 1869. uk/ books/ 2003/ feb/ 08/ peopleinscience. 159–167 (http:/ / darwin-online. pp. uk/ content/ frameset?viewtype=text& itemID=F387& pageseq=101) [90] Darwin 1859. pp. 60–61 Larson 2004. Elsberry (1996). 243–244 (http:/ / darwin-online. retrieved 2009-04-30 [114] Darwin 1859. 489–490 (http:/ / darwin-online. 180–181 [131] Quammen 2006. org. R. [187] Quammen 2006. p. pp. 183–184. p. 271–283 [188] The ISTC of On the Origin of Species is A02-2009-00000001-4. 184–185 [148] Browne 2002. 197–198 [145] Bowler 2003. 464–465. Darwin Correspondence Project. 295 [185] Darwin & Costa 2009. vatican. 512 [154] Lucas 1979 Desmond & Moore 1991. p. C. 89–92 [165] Bowler 2003. 545–547 Secord 2000. p. pp. p. Vatican. retrieved 2009-04-11 [176] Forster & Marston 1999. this work has been the first one to be registered by The International ISTC Agency. . html). Asa (1860). 493–499 [155] Browne 2002. ac. 179. p. 26 [164] Larson 2004. 208–211. . p. p. "Natural Selection is not inconsistent with Natural Theology" (http:/ / www. . 149 . pp. 186–187. p. pp. pp. . p. va/ holy_father/ pius_xii/ encyclicals/ documents/ hf_p-xii_enc_12081950_humani-generis_en. pp. pp. 139 [166] Darwin and design: historical essay (http:/ / www. As a tribute to its influence. 27 [174] Miles 2001 [175] Gray. ac. Atlantic Monthly (Darwin Correspondence Project – Essay: Natural selection & natural theology). p. 177 [178] Gray. pp. 179–180. The Complete Works of Charles Darwin Online. 205–234 [162] Bowler 2003. p. [172] Desmond & Moore 1991. 237. . 189 [146] Bowler 2003. 41–43 [180] Bolwer 2003. 119 [170] Moore 2006 [171] Barlow 1963. 323–324 [181] Pius XII (1950). ac. retrieved 2009-04-23 [184] Larson 2004. pp. 18 Nov 1859 (http:/ / www. darwinproject. retrieved 2009-06-08 [182] Kreeft 2001. pp. 234–236 [160] Bowler 2003. pp. . uk/ content/ frameset?viewtype=text& itemID=F376& pageseq=508). pp.''On the Origin of Species'' [141] Browne 2002. p. pp. 490 (http:/ / darwin-online. x [186] Peckham 1959. darwinproject. 21 January 2009. pp. darwinproject. 108 [152] Bowler 2003. html). recently reprinted. Humani Generis (http:/ / www. pp. Asa (28 May 1874). retrieved 2009-03-03 [179] Forster & Marston 1999. pp. retrieved 2009-04-11 [168] Darwin 1860. uk/ content/ frameset?viewtype=text& itemID=F1577& pageseq=9). 198–200. 169–170. 220–222 [144] Bowler 2003. 2007. 207 (http:/ / darwin-online. 160–161 [156] Bowler 2003. 490–491. pp. pp. 105–106 [149] Huxley 1860 [150] Bowler 2003. Charles to Darwin. "What is Darwinism?" (http:/ / www. pp. uk/ content/ view/ 110/ 104/ ). org. html). uk/ content/ view/ 84/ 69/ ). 203–207. 124–126 [153] Desmond & Moore 1991. pp. 241 [159] Bowler 2003. ac. 190–192 [158] Bowler 2003. org. uk/ biography. p. retrieved 2008-09-17 [167] Darwin Correspondence Project – Letter 2534—Kingsley. [169] Quammen 2006. darwinproject. pp. uk/ content/ view/ 86/ 71/ ). 214–216 [157] Bowler 2003. 225 [161] Quammen 2006. org. 500 [173] Dewey 1994. pp. The Nation (Darwin Correspondence Project).. p. 208 [147] Bowler 2003. 49 [183] Biography (http:/ / darwin-online. 184 [151] Larson 2004. 202–208 [163] Dewey 1994. 87 Leifchild 1859 [142] Bowler 2003. 207–209 Huxley 1863 [143] Bowler 2003. 37–40 [177] Hodge 1874. pp. 287–288. uk/ darwinletters/ calendar/ entry-2534. 487–488. Nora. ISBN 0-520-23693-9 • Browne. London: John Murray. retrieved 2009-04-22 • Darwin. retrieved 2009-01-09 . retrieved 2009-01-09 • Darwin. Journal of Researches Into the Natural History and Geology of the Countries Visited During the Voyage of H. retrieved 2009-02-22 • Darwin. Charles (1859).. On the Origin of Species by Means of Natural Selection.). Beagle Round the World.nytimes. darwin200. Baltimore: Johns Hopkins University Press. The Natural History Museum.org. Charles Darwin: Vol. The Origin of Species by Means of Natural Selection. London: Collins. On the Origin of Species by Means of Natural Selection.).uk/content/frameset?itemID=F376&viewtype=text& pageseq=1) (2nd ed. Nora.S. retrieved 2009-02-22 • Darwin. (1859). London: Jonathan Cape. Charles (1866).org. retrieved 2009-06-10 • Bowler. 1 Voyaging. retrieved 2009-01-09 • Darwin. London: Jonathan Cape. Janet (2002). 2 The Power of Place.). London: John Murray. London: John Murray.50&viewtype=text&pageseq=1). E. Charles (1861).html)]] (1st ed. London: John Murray. uk/EditorialIntroductions/Freeman_LifeandLettersandAutobiography. . (http:// darwin-online.com/packages/images/nytint/docs/ charles-darwin-on-the-origin-of-species/original.). Peter J.). Published anonymously. org/ what-is. 7): 201–278. or the Preservation of Favoured Races in the Struggle for Life (http://darwin-online.uk/content/ frameset?itemID=CUL-DAR226. ISBN 0-485-11375-9 • Bowler. Janet (1995). ISBN 0-7126-6837-3 • Crawford. London: John Murray.org. ed.uk/ EditorialIntroductions/Freeman_TheDescentofMan.org. or the Preservation of Favoured Races in the Struggle for Life (http://darwin-online. With the Original Omissions Restored.N. Charles Darwin: Vol. E.). On the Origin of Species by Means of Natural Selection. London: John Murray. or the Preservation of Favoured Races in the Struggle for Life (http://darwin-online. retrieved 2009-01-09 • Darwin. or the Preservation of Favoured Races in the Struggle for Life (http://darwin-online. London: John Murray. retrieved 2009-04-23 150 References • Barlow. ed.M. and Selection in Relation to Sex (http://darwin-online. Bulletin of the British Museum (Natural History) Historical Series 2 (No.org. (1989).org. Charles (1871). Charles (1872).org.). Barlow.org. (1963). On the Origin of Species by Means of Natural Selection. J. London: John Murray.1. Charles (1869).uk/content/frameset?itemID=F391&viewtype=text& pageseq=1) (6th ed. html).org. retrieved 2009-04-29 • Darwin. Peter J. • Darwin. or the Preservation of Favoured Races in the Struggle for Life (http://darwin-online. Charles (1845).).uk/content/frameset?itemID=F385&viewtype=text& pageseq=1) (4th ed.''On the Origin of Species'' [189] Darwin 200: Celebrating Charles Darwin's bicentenary – What is Darwin200? (http:/ / www. [[The Descent of Man. "(Review of) On the Origin of Species" (http://darwin-online. Examiner: 722–723.pdf) 1st ed. University of California Press.org. ISBN 1-84413-314-1 • Browne.uk/content/frameset?viewtype=text&itemID=F14&pageseq=1) (2nd ed. Evolution: The History of an Idea (3rd ed. Edited and with Appendix and Notes by his Granddaughter Nora Barlow]. or the Preservation of Favoured Races in the Struggle for Life (http://darwin-online. On the Origin of Species by Means of Natural Selection.uk/content/frameset?itemID=F373&viewtype=text& pageseq=1) ( Full image view (http://graphics8. The Mendelian Revolution: The Emergence of Hereditarian Concepts in Modern Science and Society. Charles (1860).uk/content/ frameset?viewtype=text&itemID=F1577&pageseq=1).). retrieved 2009-01-09 • Darwin. Under the Command of Captain Fitz Roy. [[The Autobiography of Charles Darwin (http://darwin-online. "Darwin's Ornithological Notes" (http://darwin-online.uk/content/frameset?itemID=F381&viewtype=text& pageseq=1) (3rd ed. R. Charles (1958).uk/content/frameset?itemID=F387&viewtype=text& pageseq=1) (5th ed.html)] 1809–1882.org. (2003). ISBN 0-87975-853-8 • Eldredge.uk/EditorialIntroductions/vanWyhe_JournalDAR158. England: Belknap Press of Harvard University Press.txt). Medicine.ac. ed.1. retrieved 2006-12-15 • Mayr. Richard B. "Confessions of a Darwinist" (http://www.M.ox. • Huxley. (2004). Costa.S.html). retrieved 2007-01-14 • Huxley. retrieved 2008-11-22 • Lucas. Charles. ISBN 1-85424-441-8. retrieved 2008-11-22 • Malthus. San Francisco: Ignatius Press. Edward J. retrieved 2006-12-15 • Keynes.org. Niles (2006).1-76&pageseq=1). The Politics of Evolution: Morphology. "Journal" (http://darwin-online.html) (6th ed. Ernst (1982). "On the Origin of Species" (http://darwin-online.uk/content/ frameset?itemID=A32&viewtype=text&pageseq=1). CUL-DAR158.asa3.org.org/details/bulletinofbritis07brit) • Hodge. ISBN 978-0-674-03281-1 • Desmond. Penguin Group.org. Darwin. England: Dawson. Costa.org/group/group.uk/~jrlucas/ legend. Great Essays in Science.uk/content/ frameset?viewtype=side&itemID=F1583e&pageseq=1). London: John Murray.uk/content/frameset?viewtype=side& itemID=CUL-DAR158. Adrian. The Historical Journal 22 (2): 313–330.org/ ASA/PSCF/2001/PSCF9-01Miles.org/articles/2006/spring/ eldredge-confessions-darwinist/). "Wilberforce and Huxley: A Legendary Encounter" (http://users. Darwin Online.1017/S0018246X00016848. (1979). retrieved 2009-03-24 • Freeman.. Bulletin of the British Museum (Natural History) Historical Series 7: 1–164 Also available here (http://www. Chicago: University of Chicago Press. The Virginia Quarterly Review (Spring 2006): 32–53. retrieved 2009-02-22 • Herbert. Beagle. (2000). Adrian (1989). Evolution: The Remarkable History of a Scientific Theory. Thomas Henry (1860). "The Influence of Darwinism on Philosophy". Published anonymously. Sandra.html). John (1994). "Review of 'Origin'" (http://darwin-online. Catholic Christianity. retrieved 2008-11-04 • Forster. ed. ISBN 0-89870-798-6 • Larson. "Darwin on the Origin of Species" (http://darwin-online. "Genesis Through History" (http://www.org.uk/EditorialIntroductions/ Freeman_OntheOriginofSpecies.vqronline. ISBN 0-8129-6849-2 • Leifchild (1859).).1-76. Perspectives on Science and Christian Faith 53: 196–201. New York: Modern Library. Folkestone. Roger.org. in van Wyhe. (1977). England: Monarch Books.html)].html).ivycottage. ISBN 0-674-36446-5 • Miles. Moore. in Martin Gardner. Richard. The Works of Charles Darwin: An Annotated Bibliographical Handlist (2nd ed. aspx?id=6826). Sara Joan (2001). ISBN 0-7181-3430-3 • Dewey. Dr Paul (1999). and Reform in Radical London. Thomas Robert (1826). What is Darwinism? (http://www. with an Inquiry into Our Prospects Respecting the Future Removal or Mitigation of the Evils which It Occasions (http://www. Darwiniana) (http://aleph0. retrieved 151 . John.clarku. Reason Science and Faith (Ivy Cottage: E-Books ed. "Charles Darwin and Asa Gray Discuss Teleology and Design" (http://www. Cambridge. Cambridge University Press. Charles (2006). doi:10. The Growth of Biological Thought. [Darwin's personal 'Journal' (1809-1881) (http:/ /darwin-online. ISBN 052167350X • Kreeft. Massachusetts.). 19 November 1859). 1673.econlib. Peter (2001). Charles (1874). Athenaeum (No. James T. Westminster Review 17 (April 1860): 541–570. Prometheus Books. ISBN 0712907408.org/files/19192/19192-8. Chester. Charles Darwin's Zoology Notes & Specimen Lists from H. London: Michael Joseph.html).8&pageseq=1). (1980). ISBN 0-226-14374-0 • Desmond. retrieved 2010-09-07 • Darwin. James (1991).org/library/Malthus/malPlong. and London.). Marston.gutenberg. "The Red Notebook of Charles Darwin" (http://darwin-online.uk/content/frameset?viewtype=image& itemID=CUL-DAR226.archive. Harvard University Press.org.edu/ huxley/CE2/Phen. An Essay on the Principle of Population: A View of its Past and Present Effects on Human Happiness. John R.''On the Origin of Species'' • Darwin. Six Lectures to Working Men "On Our Knowledge of the Causes of the Phenomena of Organic Nature" (Republished in Volume II of his Collected Essays. Scribner Armstrong. The Annotated Origin: A Facsimile of the First Edition of On the Origin of Species Annotated by James T. Thomas (1863). (2009). 2006.shtml). retrieved 2009-01-05 • van Wyhe. George Jackson (1871). retrieved 2009-06-06 Further reading • Janet Browne (2007). Marijn (2009). 1987). (2000). Herbert (1864). org. publicradio.). 1983): 809–811. The Complete Works of Charles Darwin Online. London: Andre Deutsch.1098/rsnr. St.0171.ning.org. James (2006). retrieved 2009-04-21 • Secord. ISBN 0-233-00251-0 • van Wyhe. 1 (http://books. Published anonymously. Reznick (2009) The Origin Then and Now: An Interpretive Guide to the Origin of Species ISBN 978-0-691-12978-5. New York: Appleton Moore. and Secret Authorship of Vestiges of the Natural History of Creation. Arnold B. the Scientific Creationist" (http://www. (1983). (June. "Darwinian Gradualism and Its Limits: The development of Darwin's Views on the Rate and Pattern of Evolutionary Change" (http://www. William. asp?title=1681).pubmedcentral. Victorian Sensation: The Extraordinary Publication. PMID 10860955. On the Genesis of Species. Evolution and Wonder – Understanding Charles Darwin (http://speakingoffaith. William (2000). American Public Media. Charles Darwin: Gentleman Naturalist: A Biographical Sketch (http://darwin-online.doc).6947. ISBN 0-226-74411-6 • Spencer. The Reluctant Mr.com/?id=SRkRAAAAYAAJ).org/showarticle. ISBN 978-0-8122-1954-8 Quammen. New York: Atlas Books. J. 23 April 2009) (http://api. 6 November 2004. Morse (ed. Proceedings of the National Academy of Sciences of the USA 97 (13): 6947–6953. The Origin and Evolution of Intelligence.uk/content/ frameset?itemID=A17&viewtype=text&pageseq=1). J. Chicago: University of Chicago Press. Speaking of Faith (Radio Program). . Contemporary reviews • Carpenter. Winkler. William Benjamin (1859). Scheibel.13. Teyler Net (Weblog of the Teylers Museum.html). Vol.google. "Solution to Darwin's dilemma: Discovery of the missing Precambrian record of life" (http://www. Haarlem). Saturday. doi:10.com/files/ aT0Fpi7QEAJXZty6RRr0*ayjT5aN-CLIR1HAnom7DRBt*Z64Ko2B*I88LPx*h2otwLvz5Vp*IKVqHQunTD*sItKN3P7MUMv TeylerWinklerDarwin. John (2009).''On the Origin of Species'' 2008-11-22 Mivart. Social Sciences and Law (Springer Netherlands) 20 (2): 139–157.religion-online. The Principles of Biology.gov/articlerender.com/content/p7u22v726028j1t7/). Boston: Jones and Bartlett. ISBN 0-7637-0365-6 152 • • • • • • • • Schopf. Reception. Christian Century (September 14–21.springerlink. doi:10. "Mind the gap: Did Darwin Avoid Publishing his Theory for Many Years?" (http:// darwin-online. Darwin's Origin of Species: A Biography. retrieved 2008-11-22 Phipps.org. London: Williams and Norgate • van Hoorn. Darwin: The Story of the Man and His Theories of Evolution. National Review 10 (December 1859): 188–214. "Darwin. PMC 34368. Journal of the History of Biology. Humanities. ISBN 0-393-05981-2 Rhodes.fcgi?artid=34368).nih. "Darwin on the Origin of Species" (http://darwin-online. James A. David (2006). The Origin of Species: a variorum text (2006 reprint ed. William E.1073/pnas. John (2007). Philadelphia: University of Pennsylvania Press. doi:10. Darwin (Lecture given at the Congress of the European Botanical and Horticultural Libraries Group.97. retrieved 2010-04-27 • van Wyhe.1007/BF00138435 Schopf.uk/darwin..) (1959).T. Teyler. Frank H. (1997). Darwin.uk/content/frameset?viewtype=text&itemID=A544&pageseq=1). John (2008). retrieved 2007-01-11 Peckham.org/programs/darwin/transcript. ISBN 978-0871139535 • David N. Notes and Records of the Royal Society 61: 177–205. Prague. uk/content/ frameset?itemID=A24&viewtype=text&pageseq=1).org. North British Review 46 (June 1867): 277–318.org.uk/Variorum/index. "(Review of) The Origin of Species" (http://darwin-online.org.''On the Origin of Species'' • Gray. Richard (1860).uk/content/ frameset?itemID=A213&viewtype=text&pageseq=1). Macmillan's Magazine 1: 142–148. Published anonymously. html).php?query=(origin species) AND creator:(darwin)) • • • • • .org/science/texts. 10 March 2009. Athenaeum (1710: 4 August 1860): 161. German.org.html). • Online Variorum (http://darwin-online.uk/content/ frameset?itemID=A30&viewtype=text&pageseq=1).html).org.org (http://www.uk/reviews. by means of Natural Selection. • Owen.org. • For further reviews. full text with embedded audio. Proceedings of the Royal Society of Edinburgh 4: 274–291. "Review of Darwin's Origin of Species" (http://darwin-online.org. the first American edition. or the Preservation of Favoured Races in the Struggle for Life" (http://darwin-online. bibliography of On the Origin of Species (http://darwin-online.uk/EditorialIntroductions/Freeman_OntheOriginofSpecies. Asa (1860). French.org. see Darwin Online: Reviews & Responses to Darwin (http://darwin-online.org/search. Samuel (1860). Russian and Spanish. Victorian Science Texts (http://www.uk/ content/frameset?itemID=A14&viewtype=text&pageseq=1).uk/content/ frameset?itemID=A19&viewtype=text&pageseq=1). Origin of Species (http://publicliterature.net/taorgin. Extract from Proceedings of the American Academy of Arts and Sciences 4 (1860): 411–415. Cambridge. "On Mr Darwin's Theory of the Origin of Species" (http://darwin-online. A short. PDF scans at Archive.uk/ content/frameset?itemID=A43&viewtype=image&pageseq=1). "Time and Life: Mr Darwin's "Origin of Species"" (http://darwin-online. The Origin of Species: An Outline (http://www. • Jenkin.uk/contents.php). showing every change between the six British editions.vectorsite. Darwin Online.html). the 6th edition with additions and corrections (final text). Polish. • Huxley.uk).victorianweb. retrieved 2009-06-18 153 External links • The Complete Works of Charles Darwin Online: • Table of contents (http://darwin-online.uk/content/ frameset?itemID=A166&viewtype=text&pageseq=1).org/books/origin_of_species/xaa. "Darwin on the Origin of Species" (http://darwin-online. The Times (26 December 1859): 8–9.archive. "(Review of) On the Origin of Species. Dutch.org. Thomas Henry (1859). University Library.darwinproject.html) – links to text and images of all six British editions of The Origin of Species.org. accessible outline of the book. and translations into Danish. Fleeming (1867). "(Review of) The Origin of Species" (http://darwin-online. Quarterly Review 108 (215: July 1860): 225–264. • Murray. Thomas Henry (1859). • Huxley. Published anonymously.org. Published anonymously. • Wilberforce. Published anonymously. Andrew (1860).ac.html) Darwin Correspondence Project Home Page (http://www. Edinburgh Review 3 (April 1860): 487–532. it is increasingly regarded as an inappropriate description of modern evolutionary theory.[4] Charles Darwin in 1868 While the term has remained in use amongst scientific authors. then around 1900 it was eclipsed by Mendelism until the modern evolutionary synthesis unified Darwin's and Gregor Mendel's ideas.[6] and was used to describe evolutionary concepts.[13] Having hailed the book as. In the late 19th century it came to mean the concept that natural selection was the sole mechanism of evolution. not explicable by natural selection? Twenty years hence naturalists may be in a position to say whether this is.Darwinism 154 Darwinism Darwinism is a set of movements and concepts related to ideas of transmutation of species or evolution. in contrast to Lamarckism. being freely used as a short hand for evolutionary theory. and praising the usefulness of Darwin's ideas while expressing professional reservations about Darwin's gradualism and doubting if it could be proved that natural selection could form new species. the case. or is not.[5] The term was coined by Thomas Henry Huxley in April 1860. . and knew nothing of genetic drift. caricatures of Charles Darwin with an ape [12] or monkey body symbolised evolution. including ideas with no connection to the work of Charles Darwin.[10] having as a result only a vague and inaccurate understanding of heredity. Darwinism is often used by creationists as a pejorative term but in the United Kingdom the term has no negative connotations. "a veritable Whitworth gun in the armoury of liberalism" promoting scientific naturalism over theology.[11] Conceptions of Darwinism While the term Darwinism had been used previously to refer to the work of Erasmus Darwin in the late 18th century. including earlier concepts such as Malthusianism and Spencerism.[7] [8] [9] For example. and varies depending on who is using the term.[1] [2] [3] The meaning of Darwinism has changed over time. here and there. but in either event they will owe the author of "The As "Darwinism" became widely accepted in the 1870s.[4] In the United States.[14] Huxley compared Darwin's achievement to that of Copernicus in explaining planetary motion: What if the orbit of Darwinism should be a little too circular? What if species should offer residual phenomena. the term has been associated at times with specific ideas. the term as understood today was introduced when Charles Darwin's 1859 book On the Origin of Species was reviewed by Thomas Henry Huxley in the April 1860 issue of the Westminster Review. As modern evolutionary theory has developed. Darwin was unfamiliar with the work of Gregor Mendel. in its day. the higher intellectual development. such as those by Huxley. in his book Mutual Aid: A Factor of Evolution.[6] Another important evolutionariy theorist of the same period was Peter Kropotkin who. and was not centred on natural selection at all. any work has appeared calculated to exert so large an influence. and the prevalent interpretations of Darwinism. and it was used by opponents and proponents of Darwin's biological theory alike to mean whatever they wanted it to in a larger context.Darwinism Origin of Species" an immense debt of gratitude. and voluntary corrective measures would be desirable—the foundation of eugenics. as yet.. its extension. on the contrary. one notably favoured by Darwin's half-cousin Francis Galton. economic and social theories of the time. secure the maintenance of the species. and the most open to further progress. the most prosperous. Another interpretation. What is now called "Social Darwinism" was. synonymous with "Darwinism" — the application of Darwinian principles of "struggle" to society. like Spencer Haeckel's "Darwinism" had only a rough resemblance to the theory of Charles Darwin. and the further growth of sociable habits.. – Peter Kropotkin. Mutual Aid: A Factor of Evolution (1902).. His conception was centred around what he saw as the widespread use of cooperation as a survival mechanism in human societies and animals. hardly penetrated. are invariably the most numerous. of course. and Ernst Haeckel developed what was known as Darwinismus in Germany. and its further progressive evolution. And viewed as a whole. He used biological and sociological arguments in an attempt to show that the main factor in facilitating evolution is cooperation between individuals in free-associated societies and groups.. which was later taken to be emblematic of Darwinism even though Spencer's own understanding of evolution was more similar to that of Jean-Baptiste Lamarck than to that of Darwin. advocated a conception of Darwinism counter to that of Huxley. and predated the publication of Darwin's theory. was that Darwinism implied that because natural selection was apparently no longer working on "civilised" people it was possible for "inferior" strains of people (who would normally be filtered out of the gene pool) to overwhelm the "superior" strains. which was where the [15] maker was. Kropotkin's conception of Darwinism could be summed up by the following quote: In the animal world we have seen that the vast majority of species live in societies. The mutual protection which is obtained in this case. and that they find in association the best arms for the struggle for life: understood. not only on the future of Biology. although. in its wide Darwinian sense – not as a struggle for the sheer means of existence. 155 19th Century usage "Darwinism" soon came to stand for an entire range of evolutionary (and often revolutionary) philosophies about both biology and society.. . One of the more prominent approaches was that summed in the phrase "survival of the fittest" by the philosopher Herbert Spencer. which provided a rationalisation for the dominant political. Conclusion. but as a struggle against all natural conditions unfavourable to the species. in which individual struggle has been reduced to its narrowest limits. This was in order to counteract the conception of fierce competition as the core of evolution. who is targeted as an opponent by Kropotkin. the possibility of attaining old age and of accumulating experience. The ideas had international influence. The animal species. “ [Both] a Darwinian 'left' and a Darwinian 'right' were in place before most people had grasped the Darwinian middle. we do not believe that. since the publication of Von Baer's "Researches on Development. ” In Darwin's day there was no rigid definition of the term "Darwinism". usually in support of anti-philanthropic political agendas. are doomed to decay." thirty years ago. but in extending the domination of Science over regions of thought into which she has. The unsociable species. and the practice of mutual aid has attained the greatest development. [22] Casting evolution as a doctrine or belief. in comparison to more recent mechanisms such as genetic drift and gene flow. starting in New York and going via Boston.[18] However. before answering the question posed in the book's title by concluding: "It is Atheism."[19] [20] [21] Creationists use the term Darwinism. as well as a pseudo-religious ideology like Marxism. in 1886 Alfred Russel Wallace went on a lecture tour across the United States.[24] See also • • • • • • • • Darwinism (book) Modern evolutionary synthesis Neo-Darwinism Neural Darwinism Social Darwinism Darwin Awards Pangenesis . Darwinism is also used neutrally within the scientific community to distinguish modern evolutionary theories from those first proposed by Darwin. In the United Kingdom the term retains its positive sense as a reference to natural selection. as an epithet to attack evolution as though it were an ideology (an "ism") of philosophical naturalism. Washington. However. unlike Johnson. Phillip E. that as a scientist he is a Darwinist. Iowa and Nebraska to California. whom they cast as dogmatic and inflexible in their belief. lecturing on what he called Darwinism without any problems. and for example Richard Dawkins wrote in his collection of essays A Devil's Chaplain. often pejoratively. Kansas. notably by leading members of the intelligent design movement.Charles Darwin's hypothetical mechanism for heredity Universal Darwinism .[23] bolsters religiously motivated political arguments to mandate equal time for the teaching of creationism in public schools. published in 2003. Darwinism may be used to refer to Darwin's proposed mechanism of natural selection. Hodge confined the term to exclude those like Asa Gray who combined Christian faith with support for Darwin's natural selection theory. or atheism.[16] 156 Other uses The term Darwinism is often used in the United States by promoters of creationism.[17] For example. It may also refer specifically to the role of Charles Darwin as opposed to others in the history of evolutionary thought — particularly contrasting Darwin's results with those of earlier theories such as Lamarckism or later ones such as the modern synthesis.Darwinism While the reaction against Darwin's ideas is nowadays often thought to have been widespread immediately. as well as by historians to differentiate it from other evolutionary theories from around the same period. Johnson makes this accusation of atheism with reference to Charles Hodge's book What Is Darwinism?. For example. to imply that the theory has been held as true only by Darwin and a core group of his followers. 152. docstoc. nytimes. com/ ?id=SHWaeRiRD-cC& printsec=frontcover& dq="michael+ ruse"+ darwinism). shtml). . impactpress. interview about 2003 book. Retrieved 18 July 2008. (2008). . [17] Scott.Darwinism 157 Notes [1] John Wilkins (1998). . uk/ content/ frameset?viewtype=side& itemID=A32& pageseq=29). com/ 2008/ 07/ 15/ lets-get-rid-of-darwinism/ ). Retrieved 19 June 2008. [20] Hodge. pp. publicradio. Journal of Biosciences (Bangalore. [16] "Evolution and Wonder . [18] Johnson. html). . . . html). doi:10.H. "ART. "What is Darwinism?" (http:/ / www. ISBN 0-393-33073-7. Retrieved 29 June 2008. com/ content/ n47h34357743w4p0/ ?p=e3b030036a4d442a8ce393291fe0688f& pi=9). talkorigins. 376–379 [13] "The Huxley File § 4 Darwin's Bulldog" (http:/ / aleph0. [12] Browne 2002. com/ docs/ 20835072/ Darwinism-should-be-allowed-to-collapse-and-end-up-on-the-ash-heap-of-history). . Branch. T. htm). clarku. org/ faqs/ anti-darwin. beliefnet. org/ faqs/ darwinism. Retrieved 4 January 2007. Laura. com/ papers/ Winter1997/ CharlesHodge. Charles. TalkOrigins Archive. . ISSN 1936-6434. "What is Darwinism?" (http:/ / www. Retrieved 27 July 2007. What is Darwinism?. Retrieved 27 June 2008. html). net/ news/ index. p.. . TalkOrigins Archive. VIII. "How to be Anti-Darwinian" (http:/ / www. In Godfrey. MA: Harvard University Press. talkorigins. "What if the orbit of Darwinism should be a little too circular?" [7] John Wilkins (1998). 541–70. Retrieved 19 June 2008. Retrieved 3 January 2009. [22] Sullivan. pp. . Impact Press. "Let’s Get Rid of Darwinism" (http:/ / judson. Retrieved 4 January 2007. blogs. astrobio. arn. [23] "Darwinism should be allowed to collapse and end up on the ash heap of history" (http:/ / www. com/ story/ 136/ story_13688_1. . Ropp. pp. Glenn (16 January 2009). "Charles Hodge and His Objection to Darwinism" (http:/ / www. [3] based on an European Southern Observatory release (9 December 2006). 293. ucf. Armstrong. Retrieved 22 December 2008.com. Andrew (June 2006). Norton. [6] Huxley. . [19] Matthew. and Company. [14] Browne 2002. . org. New York: W.earth science evolution distribution Origin of life universe . html).Darwin on the origin of Species" (http:/ / darwin-online. com/ index. Michael (2003). org/ docs/ johnson/ wid. springerlink. [2] "Expelled Exposed: Why Expelled Flunks » …on what evolution explains" (http:/ / www. org/ programs/ darwin/ transcript. Phillip E. Retrieved 4 January 2007. "How to be Anti-Darwinian" (http:/ / www. php/ contest/ on-what-evolution-explains). . [10] Sclater. (April 1860). google. doi:10. . [8] Ruse. php?name=News& file=article& sid=2169& theme=Printer). edu/ huxley/ guide4. htm)... com/ articles/ spring05/ sullivanspring05. . Darwin and Design: Does Evolution Have a Purpose? (http:/ / books. Andrew J. Eugenie C. edu/ ~clp/ Courses/ seminar/ papers/ 07-Scott-scientists_confront-cs_lite. 105–106 [15] Gopnik 2009. Westminster Review. "Don’t Call it “Darwinism”" (http:/ / www. Scribner. Retrieved 17 November 2009. Laurie R.1007/BF02703910..Understanding Charles Darwin . html). "Random Genetic Drift" (http:/ / www. TalkOrigins Archive. India: Springer India / Indian Academy of Sciences) 31 (2): 191–193. expelledexposed. OCLC 11489956. . New York Times. Petto. org/ faqs/ anti-darwin. springerlink. pp. Cambridge. [21] Hodge. "The extent of Charles Darwin’s knowledge of Mendel" (http:/ / www. [4] Joel Hanes. "What is Darwinism?" (http:/ / www. [5] Scott. org/ files/ 19192/ 19192-8. talkorigins. pdf).1007/s12052-008-0111-2. National Center for Science Education. "Creation Science Lite: "Intelligent Design" as the New Anti-Evolutionism" (http:/ / biology. Evolution: Education and Outreach (New York: Springer) 2 (1): 90. Eugenie C. talkorigins. Charles (1874). 72. Scientists Confront Creationism: Intelligent Design and Beyond. W. Retrieved 27 June 2008. . "Galactic Darwinism :: Astrobiology Magazine . M (2005). . BeliefNet. PMID 16809850. html). "From the Beagle to the School Board: God Goes Back to School" (http:/ / www. txt). [11] Laurence Moran (1993).. Retrieved 18 September 2008. Retrieved 22 December 2008. gutenberg. pp. Religion: For Dummies (http:/ / www. org/ faqs/ genetic-drift. [9] Olivia Judson (15 July 2008).life beyond :: Astrobiology is study of earth science evolution distribution Origin of life in universe terrestrial" (http:/ / www. html).Speaking of Faith from American Public Media" (http:/ / speakingoffaith. theropps. Retrieved 19 June 2008. TalkOrigins Archive. [24] Sheahen. ISBN 0674016319. com/ content/ w112307246x77t37/ ). ISBN 9781847249296. Conditions of Permanent Civilization . and Modern Life. Man and Society 9.html) • The Darwinian Revolution (http://www.ru/lib/book3542/) (Дарвинизм. Lincoln. A Critical Study (http://new. Angels and Ages: A Short Book About Darwin. Reproduction in Relation to Social Class 11.runivers. slightly revised edition was republished in 1958. edited by Henry Bennett. Charles Darwin: Vol. notes and alterations accidentally omitted from the second edition was published. Chapters It contains the following chapters: 1.htm) The Genetical Theory of Natural Selection The Genetical Theory of Natural Selection is a book by R. Janet (2002). The Nature of Inheritance [2] 2. Editions A second. It is one of the most important books of the modern evolutionary synthesis[1] and is commonly cited in biology books. Критическое исследование) at Runivers.talkorigins. Mimicry 8. Adam (2009).edu/entries/darwinism/) • What is Darwinism (http://www.org/faqs/darwinism. a third variorum edition (ISBN 0-19-850440-3). London: Jonathan Cape. The Inheritance of Human Fertility 10.info/history.com) • (Russian) Nikolai Danilevsky. 2 The Power of Place.Darwinism 158 References • Browne. In 1999. The Fundamental Theorem of Natural Selection 3.A. Fisher first published in 1930 by Clarendon.ru in DjVu format • Stanford Encyclopedia of Philosophy entry (http://plato.universaldarwinism. The Evolution of Dominance 4. External links • Universal Darwinism (http://www. ISBN 0712668373.evolution-of-man. with the original 1930 text. • Gopnik. Variation etc 6. Social Selection of Fertility 12. Variation as determined by Mutation and Selection 5. annotated with the 1958 alterations. Sexual Reproduction and Sexual Selection [3] 7. London: Quercus. E. 1885-1889 Darwinism.stanford. "In gratitude for the encouragement. I weighed as of equal importance to the entire rest of my undergraduate Cambridge BA course and. Unlike in 1958. The publication of the variorum edition in 1999 led to renewed interest in the work and reviews by Laurence Cook ("This is perhaps the most important book on evolutionary genetics ever written").F. Fisher considers the nature of inheritance..W. The second chapter introduces Fisher's fundamental theorem of natural selection. by discussing many of the problems dealt with in this book". during the last fifteen years. The work had a great effect on W. however. Most chapters took me weeks. For a book that I rate only second in importance in evolution theory to Darwin's Origin (this as joined with its supplement Of Man).[4] Sewall Wright.[9] Brian Charlesworth. . Fisher's friend.. will I have understood all that is true in this book and will I get a First? I doubt it. I think it notched down my degree. By the time of my ultimate graduation. this brilliant. natural selection has become part of the syllabus of our intellectual life and the topic is certainly included in every decent course in biology.''The Genetical Theory of Natural Selection'' 159 Contents In the preface. who discovered it as an undergraduate at Oxford[8] and noted on the rear cover of the 1999 variorum edition: This is a book which. The third considers the evolution of dominance. In some ways some of us have overtaken Fisher. The last five chapters (8-12) include Fisher's more idiosyncratic views on eugenics. however. Haldane described it as "brilliant". rejecting blending inheritance in favour of particulate inheritance. and that the then-recent advances in the field of genetics (see history of genetics) now allowed this.[6] Reginald Punnett was negative.And little modified even by molecular genetics.B. which Fisher believed was strongly influenced by modifiers. Dedication The book is dedicated to Major Leonard Darwin.. who had many disagreements with Fisher.[10] Jim Crow[11] and A.D. through the time I spent on it. and also rate as undoubtedly one of the greatest books of the twentieth century the appearance of a variorum edition is a major event. and in particular the fundamental theorem was misunderstood. daring man is still far in front. some months. reviewed the book and wrote that it was "certain to take rank as one of the major contributions to the theory of evolution". Reviews Henry Bennett gave an account of the writing and reception of Fisher's Genetical Theory.[5] J. Fisher's logic and ideas still underpin most of the ever broadening paths by which Darwinism continues its invasion of human thought. correspondent and son of Charles Darwin. In the first chapter.. Hamilton.[7] ' The Genetical Theory was largely overlooked for 40 years. given to the author. including that there must be an understanding of natural selection distinct from that of evolution.S. in many. Edwards[12] . Fisher considers some general points. as a student. ''The Genetical Theory of Natural Selection'' 160 References [1] Grafen, Alan; Ridley, Mark (2006). Richard Dawkins: How A Scientist Changed the Way We Think. New York, New York: Oxford University Press. p. 69. ISBN 0199291160. [2] http:/ / www. blackwellpublishing. com/ ridley/ classictexts/ fisher1. pdf [3] http:/ / www. blackwellpublishing. com/ ridley/ classictexts/ fisher2. pdf [4] http:/ / digital. library. adelaide. edu. au/ coll/ special/ fisher/ natsel/ tp_intro. pdf [5] Wright, S., 1930 The Genetical Theory of Natural Selection: a review (http:/ / jhered. oxfordjournals. org/ cgi/ reprint/ 21/ 8/ 349). J. Hered. 21:340-356. [6] Haldane, J.B.S., 1932 The Causes of Evolution. Longman Green, London. [7] Punnett, R.C. 1930, A review of The Genetical Theory of Natural Selection, Nature 126: 595-7 [8] Grafen, A. 2004. 'William Donald Hamilton’ (http:/ / users. ox. ac. uk/ ~grafen/ cv/ WDH_memoir. pdf). Biographical Memoirs of Fellows of the Royal Society, 50, 109-132 [9] Cook, L. 2000 Book reviews. The Genetical Theory of Natural Selection — A Complete Variorum Edition. R. A. Fisher (edited by Henry Bennett) (http:/ / www. nature. com/ hdy/ journal/ v84/ n3/ full/ 6887132a. html). Heredity 84 (3) , 390–39 [10] Charlesworth, B. 2000 The Genetical Theory of Natural Selection. A Complete Variorum Edition. By R. A. Fisher (edited with foreword and notes by J. H. Bennett). Oxford University Press. 1999. ISBN 0-19-850440-3. xxi+318 pages. (http:/ / journals. cambridge. org/ action/ displayAbstract?fromPage=online& aid=52675) Genetics Research 75: 369-373 [11] Crow, J.F. 2000 Second only to Darwin - The Genetical Theory of Natural Selection. A Complete Variorum Edition by R.A. Fisher (http:/ / www. sciencedirect. com/ science?_ob=ArticleURL& _udi=B6VJ1-405BR68-M& _user=10& _rdoc=1& _fmt=& _orig=search& _sort=d& view=c& _acct=C000050221& _version=1& _urlVersion=0& _userid=10& md5=3a8c3a64ce75187ab109f4b0b9d18ba9) Trends in Ecology and Evolution, Volume 15, Number 5, 1 May 2000 , pp. 213-214(2) [12] Edwards, A.W.F. 2000 The Genetical Theory of Natural Selection (http:/ / www. genetics. org/ cgi/ content/ full/ 154/ 4/ 1419) Genetics, Vol. 154, 1419-1426, April 2000 External links • Full text of 1930 edition (http://www.openlibrary.org/details/geneticaltheoryo031631mbp), Open Library Neo-Darwinism Neo-Darwinism is a term used to describe the 'modern synthesis' of Darwinian evolution through natural selection with Mendelian genetics, the latter being a set of primary tenets specifying that evolution involves the transmission of characteristics from parent to child through the mechanism of genetic transfer, rather than the 'blending process' of pre-Mendelian evolutionary science. Neo-Darwinism also separates Darwin's ideas of natural selection from his hypothesis of Pangenesis as a Lamarckian source of variation involving blending inheritance.[1] As part of the disagreement about whether natural selection alone was sufficient to explain speciation, George Romanes coined the term neo-Darwinism to refer to the version of evolution advocated by Alfred Russel Wallace and August Weismann with its heavy dependence on natural selection.[2] Weismann and Wallace rejected the Lamarckian idea of inheritance of acquired characteristics, something that Darwin had not ruled out.[3] The term was first used in 1895 to explain that evolution occurs solely through natural selection, in other words, without any mechanism involving the inheritance of acquired characteristics resulting from use or disuse.[4] These two scientists' complete rejection of Lamarckism came from Weismann's germ plasm theory. Weismann realised that the cells that produce the germ plasm, or gametes (such as sperm and egg in animals), separate from the somatic cells that go on to make other body tissues at an early stage in development. Since he could see no obvious means of communication between the two he asserted that the inheritance of acquired characteristics was therefore impossible; a conclusion now known as Weismann's barrier.[5] From the 1880s to the 1930s the term continued to be applied to the panselectionist school of thought, which argued that natural selection was the main and perhaps sole cause of all evolution.[6] From then until around 1947 the term was used for the panselectionist followers of R. A. Fisher. Neo-Darwinism 161 Modern evolutionary synthesis Following the development, from about 1937 to 1950, of the modern evolutionary synthesis, now generally referred to as the synthetic view of evolution or the modern synthesis, the term neo-Darwinian is often used to refer to contemporary evolutionary theory.[7] However, such usage has been described by some as incorrect;[8] [1] [4] with Ernst Mayr writing in 1984: "...the term neo-Darwinism for the synthetic theory is wrong, because the term neo-Darwinism was coined by Romanes in 1895 as a designation of Weismann's theory."'[9] Despite this, publications such as the Encyclopaedia Britannica,[10] [11] use this term to refer to current evolutionary theory. This term is also used in the scientific literature, with the academic publishers Blackwell Publishing referring to "neo-Darwinism as practised today",[12] and some figures in the study of evolution like Richard Dawkins[13] and Stephen Jay Gould,[14] using the term in their writings and lectures. See also • Modern evolutionary synthesis • The Origin of Species • Developmental systems theory • Evolutionary developmental biology • Neo-Lamarckism References [1] Kutschera U, Niklas KJ (2004). "The modern theory of biological evolution: an expanded synthesis". Naturwissenschaften 91 (6): 255–76. doi:10.1007/s00114-004-0515-y. PMID 15241603. [2] Gould The Structure of Evolutionary Theory p. 216 [3] Kutschera U. 2003. A comparative analysis of the Darwin-Wallace papers and the development of the concept of natural selection. Theory in Biosciences 122, 343-359 (http:/ / www. springerlink. com/ content/ f6131358k265g3u4/ ) [4] Reif W-E. Junker T. Hoßfeld U. (2000). "The synthetic theory of evolution: general problems and the German contribution to the synthesis". Theory in Biosciences 119 (1): 41–91(51). doi:10.1078/1431-7613-00004. [5] Barbieri FD (1989). "The origin of Metazoa and Weismann's germ line theory". Riv. Biol. 82 (1): 61–74. PMID 2665023. [6] "How to be Anti-Darwinian" (http:/ / www. talkorigins. org/ faqs/ anti-darwin. html). . Retrieved 2007-09-19. [7] "The Modern Synthesis of Genetics and Evolution" (http:/ / www. talkorigins. org/ faqs/ modern-synthesis. html). . Retrieved 2007-09-19. [8] Pigliucci, M. (2007). "Do We Need An Extended Evolutionary Synthesis?" (http:/ / www. bioone. org/ perlserv/ ?request=get-abstract). Evolution 61 (12): 2743–2749. doi:10.1111/j.1558-5646.2007.00246.x. PMID 17924956. . [9] Mayr E. (1984). "What is Darwinism Today?" (http:/ / links. jstor. org/ sici?sici=0270-8647(1984)1984<145:WIDT>2. 0. CO;2-5). Proceedings of the Biennial Meeting of the Philosophy of Science Association 2: 145–156. . [10] "neo-Darwinism" (http:/ / www. britannica. com/ eb/ article-9373225/ neo-Darwinism). Britannica.com. . Retrieved 2007-09-19. [11] "neo-Darwinism" (http:/ / encyclopedia. farlex. com/ neo-Darwinism). Hutchinson Encyclopedia. . Retrieved 2007-09-19. [12] "A-Z Browser of Evolution" (http:/ / www. blackwellpublishing. com/ ridley/ a-z/ Neo-Darwinism. asp). Blackwell Publishing. . Retrieved 2007-09-19. [13] "Lecture on Neo-Darwinism" (http:/ / richarddawkins. net/ article,1345,Lecture-on-Neo-Darwinism,Richard-Dawkins). RichardDawkins.net : The Official Richard Dawkins Website. . Retrieved 2007-09-19. [14] "Challenges to Neo-Darwinism and Their Meaning for a Revised View of Human Consciousness" (http:/ / www. tannerlectures. utah. edu/ lectures/ documents/ gould85. pdf). Cambridge University: The Tanner Lectures on Human Values. . Retrieved 2009-03-05. Neo-Darwinism 162 External links • The Variation of Animals and Plants Under Domestication (http://www.esp.org/books/darwin/variation/ facsimile/title3.html) • Challenges to neo-Darwinism - Stephen Jay Gould (http://web.archive.org/web/20060924065722/http:// www.tannerlectures.utah.edu/lectures/gould85.pdf) at the Wayback Machine (archived September 24, 2006). • Neodarwinism - Richard Dawkins (http://video.google.com/videoplay?docid=4633079169415752395) Modern evolutionary synthesis The modern evolutionary synthesis is also referred to as the new synthesis, the modern synthesis, the evolutionary synthesis and the neo-darwinian synthesis. It is a union of ideas from several biological specialties which provides a widely accepted account of evolution. The synthesis reflects the current overwhelming consensus.[1] The synthesis was produced over a decade (1936–1947). The previous development of population genetics (1918–1932) was a stimulus, as it showed that Mendelian genetics was consistent with natural selection and gradual evolution. The synthesis is still, to a large extent, the current paradigm in evolutionary biology.[2] Julian Huxley invented the term, when he produced his book, Evolution: The Modern Synthesis (1942). Other major figures in the modern synthesis include R. A. Fisher, Theodosius Dobzhansky, J.B.S. Haldane, Sewall Wright, E.B. Ford, Ernst Mayr, Bernhard Rensch, Sergei Chetverikov, George Gaylord Simpson, and G. Ledyard Stebbins. The modern synthesis solved difficulties and confusions caused by the specialisation and poor communication between biologists in the early years of the 20th century. Discoveries of early geneticists were difficult to reconcile with gradual evolution and the mechanism of natural selection. The synthesis reconciled the two schools of thought, while providing evidence that studies of populations in the field were crucial to evolutionary theory. It drew together ideas from several branches of biology that had become separated, particularly genetics, cytology, systematics, botany, morphology, ecology and paleontology. Summary of the modern synthesis The modern synthesis bridged the gap between experimental geneticists and naturalists, and between both and palaeontologists. It states that:[3] [4] [5] 1. All evolutionary phenomena can be explained in a way consistent with known genetic mechanisms and the observational evidence of naturalists. 2. Evolution is gradual: small genetic changes, recombination ordered by natural selection. Discontinuities amongst species (or other taxa) are explained as originating gradually through geographical separation and extinction (not saltation). 3. Natural selection is by far the main mechanism of change; even slight advantages are important when continued. The object of selection is the phenotype in its surrounding environment. 4. The role of genetic drift is equivocal. Though strongly supported initially by Dobzhansky, it was downgraded later as results from ecological genetics were obtained. 5. Thinking in terms of populations, rather than individuals, is primary: the genetic diversity existing in natural populations is a key factor in evolution. The strength of natural selection in the wild is greater than previously expected; the effect of ecological factors such as niche occupation and the significance of barriers to gene flow are all important. 6. In palaeontology, the ability to explain historical observations by extrapolation from microevolution to macroevolution is proposed. Historical contingency means explanations at different levels may exist. Gradualism does not mean constant rate of change. Modern evolutionary synthesis The idea that speciation occurs after populations are reproductively isolated has been much debated. In plants, polyploidy must be included in any view of speciation. Formulations such as 'evolution consists primarily of changes in the frequencies of alleles between one generation and another' were proposed rather later. The traditional view is that developmental biology ('evo-devo') played little part in the synthesis,[6] but an account of Gavin de Beer's work by Stephen J. Gould suggests he may be an exception.[7] 163 Developments leading up to the synthesis 1859–1899 Charles Darwin's The Origin of Species was successful in convincing most biologists that evolution had occurred, but was less successful in convincing them that natural selection was its primary mechanism. In the 19th and early 20th centuries, variations of Lamarckism, orthogenesis ('progressive' evolution), and saltationism (evolution by jumps) were discussed as alternatives.[8] Also, Darwin did not offer a precise explanation of how new species arise. As part of the disagreement about whether natural selection alone was sufficient to explain speciation, George Romanes coined the term neo-Darwinism to refer to the version of evolution advocated by Alfred Russel Wallace and August Weismann with its heavy dependence on natural selection.[9] Weismann and Wallace rejected the Lamarckian idea of inheritance of acquired characteristics, something that Darwin had not ruled out.[10] Weismann's idea was that the relationship between the hereditary material, which he called the germ plasm (de: Keimplasma), and the rest of the body (the soma) was a one-way relationship: the germ-plasm formed the body, but the body did not influence the germ-plasm, except indirectly in its participation in a population subject to natural selection. Weismann was translated into English, and though he was influential, it took many years for the full significance of his work to be appreciated.[11] Later, after the completion of the modern synthesis, the term neo-Darwinism came to be associated with its core concept: evolution, driven by natural selection acting on variation produced by genetic mutation, and genetic recombination (chromosomal crossovers).[9] 1900–1915 Gregor Mendel's work was re-discovered by Hugo de Vries and Carl Correns in 1900. News of this reached William Bateson in England, who reported on the paper during a presentation to the Royal Horticultural Society in May 1900.[12] It showed that the contributions of each parent retained their integrity rather than blending with the contribution of the other parent. This reinforced a division of thought, which was already present in the 1890s.[13] The two schools were: • Saltationism (large mutations or jumps), favored by early Mendelians who viewed hard inheritance as incompatible with natural selection[14] • Biometric school: led by Karl Pearson and Walter Weldon, argued vigorously against it, saying that empirical evidence indicated that variation was continuous in most organisms, not discrete as Mendelism predicted. The relevance of Mendelism to evolution was unclear and hotly debated, especially by Bateson, who opposed the biometric ideas of his former teacher Weldon. Many scientists believed the two theories substantially contradicted each other.[15] This debate between the biometricians and the Mendelians continued for some 20 years and was only solved by the development of population genetics. T. H. Morgan began his career in genetics as a saltationist, and started out trying to demonstrate that mutations could produce new species in fruit flies. However, the experimental work at his lab with Drosophila melanogaster, which helped establish the link between Mendelian genetics and the chromosomal theory of inheritance, demonstrated that rather than creating new species in a single step, mutations increased the genetic variation in the population.[16] Modern evolutionary synthesis 164 The foundation of population genetics The first step towards the synthesis was the development of population genetics. R.A. Fisher, J.B.S. Haldane, and Sewall Wright provided critical contributions. In 1918, Fisher produced the paper "The Correlation Between Relatives on the Supposition of Mendelian Inheritance",[17] which showed how the continuous variation measured by the biometricians could be the result of the action of many discrete genetic loci. In this and subsequent papers culminating in his 1930 book The Genetical Theory of Natural Selection, Fisher was able to show how Mendelian genetics was, contrary to the thinking of many early geneticists, completely consistent with the idea of evolution driven by natural selection.[18] During the 1920s, a series of papers by J.B.S. Haldane applied mathematical analysis to real world examples of natural selection such as the evolution of industrial melanism in peppered moths.[] Haldane established that natural selection could work in the real world at a faster rate than even Fisher had assumed.[19] Sewall Wright focused on combinations of genes that interacted as complexes, and the effects of inbreeding on small relatively isolated populations, which could exhibit genetic drift. In a 1932 paper he introduced the concept of an adaptive landscape in which phenomena such as cross breeding and genetic drift in small populations could push them away from adaptive peaks, which would in turn allow natural selection to push them towards new adaptive peaks.[] Wright's model would appeal to field naturalists such as Theodosius Dobzhansky and Ernst Mayr who were becoming aware of the importance of geographical isolation in real world populations.[19] The work of Fisher, Haldane and Wright founded the discipline of population genetics. This is the precursor of the modern synthesis, which is an even broader coalition of ideas.[] [19] [20] The modern synthesis Theodosius Dobzhansky, a Ukrainian emigrant, who had been a postdoctoral worker in Morgan's fruit fly lab, was one of the first to apply genetics to natural populations. He worked mostly with Drosophila pseudoobscura. He says pointedly: "Russia has a variety of climates from the Arctic to sub-tropical... Exclusively laboratory workers who neither possess nor wish to have any knowledge of living beings in nature were and are in a minority".[21] Not surprisingly, there were other Russian geneticists with similar ideas, though for some time their work was known to only a few in the West. His 1937 work Genetics and the Origin of Species was a key step in bridging the gap between population geneticists and field naturalists. It presented the conclusions reached by Fisher, Haldane, and especially Wright in their highly mathematical papers in a form that was easily accessible to others. It also emphasized that real world populations had far more genetic variability than the early population geneticists had assumed in their models, and that genetically distinct sub-populations were important. Dobzhansky argued that natural selection worked to maintain genetic diversity as well as driving change. Dobzhansky had been influenced by his exposure in the 1920s to the work of a Russian geneticist named Sergei Chetverikov who had looked at the role of recessive genes in maintaining a reservoir of genetic variability in a population before his work was shut down by the rise of Lysenkoism in the Soviet Union.[] [19] Edmund Brisco Ford's work complemented that of Dobzhansky. It was as a result of Ford's work, as well as his own, that Dobzhansky changed the emphasis in the third edition of his famous text from drift to selection.[22] Ford was an experimental naturalist who wanted to test natural selection in nature. He virtually invented the field of research known as ecological genetics. His work on natural selection in wild populations of butterflies and moths was the first to show that predictions made by R.A. Fisher were correct. He was the first to describe and define genetic polymorphism, and to predict that human blood group polymorphisms might be maintained in the population by providing some protection against disease.[23] Ernst Mayr's key contribution to the synthesis was Systematics and the Origin of Species, published in 1942. Mayr emphasized the importance of allopatric speciation, where geographically isolated sub-populations diverge so far that reproductive isolation occurs. He was sceptical of the reality of sympatric speciation believing that geographical isolation was a prerequisite for building up intrinsic (reproductive) isolating mechanisms. Mayr also introduced the biological species concept that defined a species as a group of interbreeding or potentially interbreeding populations Modern evolutionary synthesis that were reproductively isolated from all other populations.[] [19] [24] Before he left Germany for the United States in 1930, Mayr had been influenced by the work of German biologist Bernhard Rensch. In the 1920s Rensch, who like Mayr did field work in Indonesia, analyzed the geographic distribution of polytypic species and complexes of closely related species paying particular attention to how variations between different populations correlated with local environmental factors such as differences in climate. In 1947, Rensch published Neuere Probleme der Abstammungslehre: die Transspezifische Evolution (English translation 1959: Evolution above the Species level). This looked at how the same evolutionary mechanisms involved in speciation might be extended to explain the origins of the differences between the higher level taxa. His writings contributed to the rapid acceptance of the synthesis in Germany.[25] [26] George Gaylord Simpson was responsible for showing that the modern synthesis was compatible with paleontology in his book Tempo and Mode in Evolution published in 1944. Simpson's work was crucial because so many paleontologists had disagreed, in some cases vigorously, with the idea that natural selection was the main mechanism of evolution. It showed that the trends of linear progression (in for example the evolution of the horse) that earlier paleontologists had used as support for neo-Lamarckism and orthogenesis did not hold up under careful examination. Instead the fossil record was consistent with the irregular, branching, and non-directional pattern predicted by the modern synthesis.[] [19] The botanist G. Ledyard Stebbins was another major contributor to the synthesis. His major work, Variation and Evolution in Plants, was published in 1950. It extended the synthesis to encompass botany including the important effects of hybridization and polyploidy in some kinds of plants.[] 165 Further advances The modern evolutionary synthesis continued to be developed and refined after the initial establishment in the 1930s and 1940s. The work of W. D. Hamilton, George C. Williams, John Maynard Smith and others led to the development of a gene-centric view of evolution in the 1960s. The synthesis as it exists now has extended the scope of the Darwinian idea of natural selection to include subsequent scientific discoveries and concepts unknown to Darwin, such as DNA and genetics, which allow rigorous, in many cases mathematical, analyses of phenomena such as kin selection, altruism, and speciation. A particular interpretation most commonly associated with Richard Dawkins, author of The Selfish Gene, asserts that the gene is the only true unit of selection.[27] Dawkins further extended the Darwinian idea to include non-biological systems exhibiting the same type of selective behavior of the 'fittest' such as memes in culture. The synthesis continues to undergo regular review.[28] (See also Current research in evolutionary biology). Modern evolutionary synthesis 166 After the synthesis There are a number of discoveries in earth sciences and biology which have arisen since the synthesis. Listed here are some of those topics which are relevant to the evolutionary synthesis, and which seem soundly based. Understanding of Earth history The Earth is the stage on which the evolutionary play is performed. Darwin studied evolution in the context of Charles Lyell's geology, but our present understanding of Earth history includes some critical advances made during the last half-century. • The age of the Earth has been revised upwards. It is now estimated at 4.56 billion years, about one-third of the age of the universe. It is worth noting that the Phanerozoic only occupies the last 1/9th of this period of time.[29] • The triumph of Alfred Wegener's idea of continental drift came around 1960. The key principle of plate tectonics is that the lithosphere exists as separate and distinct tectonic plates, which ride on the fluid-like (visco-elastic solid) asthenosphere. This discovery provides a unifying theory for geology, linking phenomena such as volcanos, earthquakes, orogeny, and providing data for many paleogeographical questions.[30] One major question is still unclear: when did plate tectonics begin?[31] • Our understanding of the evolution of the Earth's atmosphere has progressed. The substitution of oxygen for carbon dioxide in the atmosphere, which occurred in the Proterozoic, caused probably by cyanobacteria in the form of stromatolites, caused changes leading to the evolution of aerobic organisms.[32] [33] • The identification of the first generally accepted fossils of microbial life was made by geologists. These rocks have been dated as about 3.465 billion years ago.[34] Walcott was the first geologist to identify pre-Cambrian fossil bacteria from microscopic examination of thin rock slices. He also thought stromatolites were organic in origin. His ideas were not accepted at the time, but may now be appreciated as great discoveries.[35] • Information about paleoclimates is increasingly available, and being used in paleontology. One example: the discovery of massive ice ages in the Proterozoic, following the great reduction of CO2 in the atmosphere. These ice ages were immensely long, and led to a crash in microflora.[36] See also Cryogenian period and Snowball Earth. • Catastrophism and mass extinctions. A partial reintegration of catastrophism has occurred,[37] and the importance of mass extinctions in large-scale evolution is now apparent. Extinction events disturb relationships between many forms of life and may remove dominant forms and release a flow of adaptive radiation amongst groups that remain. Causes include meteorite strikes (K–T junction; Upper Devonian); flood basalt provinces (Deccan traps at K/T junction; Siberian traps at P–T junction); and other less dramatic processes.[38] [39] Conclusion: Our present knowledge of earth history strongly suggests that large-scale geophysical events influenced macroevolution and megaevolution. These terms refer to evolution above the species level, including such events as mass extinctions, adaptive radiation, and the major transitions in evolution.[40] [41] The structure of evolutionary biology. The history and causes of evolution (center) are subject to various subdisciplines of evolutionary biology. The areas of segments give an impression of the contributions of subdisciplines to the literature of evolutionary biology. Modern evolutionary synthesis Symbiotic origin of eukaryotic cell structures Once symbiosis was discovered in lichen and in plant roots (rhizobia in root nodules) in the 19th century, the idea arose that the process might have occurred more widely, and might be important in evolution. Anton de Bary invented the concept of symbiosis;[42] several Russian biologists promoted the idea;[43] Edwin Wilson mentioned it in his epic text The Cell;[44] a book was published in the U.S.A. in 1927 with the title Symbionticism and the origin of species;[45] and there was a brief mention by Julian Huxley in 1930;[46] all in vain because sufficient evidence was lacking. Symbiosis as a major evolutionary force was not discussed at all in the evolutionary synthesis.[47] The role of symbiosis in cell evolution was revived partly by Joshua Lederberg,[48] and finally brought to light by Lynn Margulis in a series of papers and books.[49] [50] It turns out that some cell organelles are of microbial origin: mitochondria and chloroplasts definitely, cilia, flagella and centrioles possibly, and perhaps the nuclear membrane and much of the chromosome structure as well. What is now clear is that the evolution of eukaryote cells is either caused by, or at least profoundly influenced by, symbiosis with bacterial and archaean cells in the Proterozoic. The origin of the eukaryote cell by symbiosis in several stages was not part of the evolutionary synthesis. It is, at least on first sight, an example of megaevolution by big jumps. However, what symbiosis provided was a copious supply of heritable variation from microorganisms, which was fine-tuned over a long period to produce the cell structure we see today. This part of the process is consistent with evolution by natural selection.[51] Trees of life The ability to analyse sequence in macromolecules (protein, DNA, RNA) provides evidence of descent, and permits us to work out genealogical trees covering the whole of life, since now there are data on every major group of living organisms. This project, begun in a tentative way in the 1960s, has become a search for the universal tree or the universal ancestor, a phrase of Carl Woese.[52] [53] The tree that results has some unusual features, especially in its roots. There are two kingdoms of prokaryotes: bacteria and archaea, both of which contributed genetic material to the eukaryotes, mainly by means of symbiosis. Also, since bacteria can pass genetic material to other bacteria, their relationships look more like a web than a tree. Once eukaryotes were established, their sexual reproduction produced the traditional branching tree-like pattern, the only diagram Darwin put in the Origin. The last universal ancestor (LUA) would be a prokaryotic cell before the split between the bacteria and archaea. LUA is defined as most recent organism from which all organisms now living on Earth descend (some 3.5 to 3.8 billion years ago, in the Archean era).[54] This technique may be used to clarify relationships within any group of related organisms. It is now a standard procedure, and examples are published regularly. April 2009 sees the publication of a tree covering all the animal phyla, derived from sequences from 150 genes in 77 taxa.[55] Early attempts to identify relationships between major groups were made in the 19th century by Ernst Haeckel, and by comparative anatomists such as Thomas Henry Huxley and E. Ray Lankester. Enthusiasm waned: it was often difficult to find evidence to adjudicate between different opinions. Perhaps for that reason, the evolutionary synthesis paid surprisingly little attention to this activity. It is certainly a lively field of research today. Evo-devo What once was called embryology played a modest role in the evolutionary synthesis,[56] mostly about evolution by changes in developmental timing (allometry and heterochrony).[57] Man himself was, according to Bolk, a typical case of evolution by retention of juvenile characteristics (neoteny). He listed many characters where "Man, in his bodily development, is a primate foetus that has become sexually mature".[58] Unfortunately, his interpretation of these ideas was non-Darwinian, but his list of characters is both interesting and convincing.[59] Modern interest in Evo-devo springs from clear proof that development is closely controlled by special genetic systems, and the hope that comparison of these systems will tell us much about the evolutionary history of different groups.[60] [61] In a series of experiments with the fruit-fly Drosophila, Edward B. Lewis was able to identify a 167 Modern evolutionary synthesis complex of genes whose proteins bind to the cis-regulatory regions of target genes. The latter then activate or repress systems of cellular processes that accomplish the final development of the organism.[62] [63] Furthermore, the sequence of these control genes show co-linearity: the order of the loci in the chromosome parallels the order in which the loci are expressed along the anterior-posterior axis of the body. Not only that, but this cluster of master control genes programs the development of all higher organisms.[64] [65] Each of the genes contains a homeobox, a remarkably conserved DNA sequence. This suggests the complex itself arose by gene duplication.[66] [67] [68] In his Nobel lecture, Lewis said "Ultimately, comparisons of the [control complexes] throughout the animal kingdom should provide a picture of how the organisms, as well as the [control genes] have evolved". The term deep homology was coined to describe the common origin of genetic regulatory apparatus used to build morphologically and phylogenetically disparate animal features.[69] It applies when a complex genetic regulatory system is inherited from a common ancestor, as it is in the evolution of vertebrate and invertebrate eyes. The phenomenon is implicated in many cases of parallel evolution.[70] A great deal of evolution may take place by changes in the control of development. This may be relevant to punctuated equilibrium theory, for in development a few changes to the control system could make a significant difference to the adult organism. An example is the giant panda, whose place in the Carnivora was long uncertain.[71] Apparently, the giant panda's evolution required the change of only a few genetic messages (5 or 6 perhaps), yet the phenotypic and lifestyle change from a standard bear is considerable.[72] [73] The transition could therefore be effected relatively swiftly. Fossil discoveries In the past thirty or so years there have been excavations in parts of the world which had scarcely been investigated before. Also, there is fresh appreciation of fossils discovered in the 19th century, but then denied or deprecated: the classic example is the Ediacaran biota from the immediate pre-Cambrian, after the Cryogenian period. These soft-bodied fossils are the first record of multicellular life. The interpretation of this fauna is still in flux. Many outstanding discoveries have been made, and some of these have implications for evolutionary theory. The discovery of feathered dinosaurs and early birds from the Lower Cretaceous of Liaoning, N.E. China have convinced most students that birds did evolve from coelurosaurian theropod dinosaurs. Less well known, but perhaps of equal evolutionary significance, are the studies on early insect flight, on stem tetrapods from the Upper Devonian,[74] [75] and the early stages of whale evolution.[76] A shaft of light has been thrown on the evolution of flatfish (pleuronectiformes), such as plaice, sole, turbot and halibut, by recent work. Flatfish are interesting because they are one of the few vertebrate groups with external asymmetry. Their young are perfectly symmetrical, but the head is remodelled during a metamorphosis, which entails the migration of one eye to the other side, close to the other eye. Some species have both eyes on the left (turbot), some on the right (halibut, sole); all living and fossil flatfish to date show an 'eyed' side and a 'blind' side.[77] The lack of an intermediate condition in living and fossil flatfish species had led to debate about the origin of such a striking adaptation. The case was considered by Lamark,[78] who thought flatfish precursors would have lived in shallow water for a long period, and by Darwin, who predicted a gradual migration of the eye, mirroring the metamorphosis of the living forms. Darwin's long-time critic St. George Mivart thought that the intermediate stages could have no selective value,[79] and in the 6th edition of the Origin, Darwin made a concession to the possibility of acquired traits.[80] Many years later the geneticist Richard Goldschmidt put the case forward as an example of evolution by saltation, bypassing intermediate forms.[81] [82] A recent examination of two fossil species from the Eocene has provided the first clear picture of flatfish evolution. The discovery of stem flatfish with incomplete orbital migration refutes Goldschmidt's ideas, and demonstrates that "the assembly of the flatfish bodyplan occurred in a gradual, stepwise fashion".[83] There are no grounds for thinking that incomplete orbital migration was maladaptive, because stem forms with this condition ranged over two geological stages, and are found in localities which also yield flatfish with the full cranial asymmetry. The evolution 168 503–518 [21] Mayr & Provine 1998 p. 2006. Massimo 2007. Harvard. springerlink. Ontogeny and phylogeny. 1942. 271–272 [17] Transactions of the Royal Society of Edinburgh. Chapman and Hall. 298–299. [26] Mayr and Provine 1998 pp. p. Cambridge. treated with special regard to discontinuity in the origin of species. Columbia University Press N. Evolution: the modern synthesis. Ecological genetics. 325–339 [20] Gould The Structure of Evolutionary Theory pp. blackwell-synergy. What evolution is. 1951.Modern evolutionary synthesis of flatfish falls squarely within the evolutionary synthesis. The age of the Earth in the twentieth century: a problem (mostly) solved. John Innes Centre. 33–34 [25] Smith. DC: The National Academy of Sciences. UK. 1111/ j. wku. ISBN 0199291160. Richard Dawkins: How A Scientist Changed the Way We Think. p270 [3] Huxley J. Betty. 231 [22] Dobzhansky T. 2nd ed 1963. Western Kentucky University. x?cookieSet=1& journalCode=evo) Evolution 61 12. Norwich. 2743–2749. 2003. Mark (2006). [23] Ford E. "Mendel's Peas" (http:/ / www. A comparative analysis of the Darwin-Wallace papers and the development of the concept of natural selection. Harvard 1977. [4] Mayr & Provine 1998 [5] Mayr E. Geological Society of London 190. p221-2 [8] Bowler P. htm). The division of thought was between gradualists of the Darwinian school. [16] Bowler pp. com/ content/ f6131358k265g3u4/ ) [11] Bowler pp. Special Publications. 343-359 (http:/ / www. 253–256 [12] Mike Ambrose. London. Genetic Resources Unit. Genetics and the Origin of Species. 1999. London. 3rd ed. V. 2nd ed. 2003. 205–221. ac. Alan. htm). and saltationists such as Bateson. Cambs. 2009. Washington. 221–243 [19] Bowler Evolution: The history of an Idea pp. The growth of biological thought: diversity. edu/ openbook. [14] Larson pp. The start of the world as we know it. . . Ridley. uk/ germplas/ pisum/ zgs4f. Charles H. pp236–256 [9] Gould The Structure of Evolutionary Theory p. . 1964. [13] Bateson. Brent 2001. 2007. Unifying Biology: the evolutionary synthesis and evolutionary biology. p. jic. Retrieved 2007-09-22. New York: Oxford University Press. 1558-5646. p192 [7] Gould S.).B. Bernhard (Carl Emmanuel) (Germany 1900–1990)" (http:/ / www.S. 157–166 [15] Grafen. New views on an old planet: a history of global change. php?record_id=6024& page=27#p200064869970027001) (php). evolution & inheritance. "Appendix: Frequently Asked Questions" (http:/ / www. Mutations (as 'sports') and polymorphisms were well known long before the Mendelian recovery. nap. .361 [28] Pigliucci. 4th edn 1975. Ernst 2002. William 1894. G. 1996. 00246. [2] Mayr. Retrieved 2007-09-22. [31] Witz A. Do we need an extended evolutionary synthesis? (http:/ / www. [29] Dalrymple. edu/ openbook. 69.Y. 3rd ed 1974. p128. 52:399–433 [18] Larson Evolution: The Remarkable History of a Scientific Theory pp. Princeton University Press. Science and Creationism: A View from the National Academy of Sciences (http:/ / books. London. Allen & Unwin. "Rensch. com/ doi/ abs/ 10. 28. 416 [27] Bowler p. p567 et seq.. Evolution: the history of an idea. Retrieved September 24. Theory in Biosciences 122. Materials for the study of variation. 216 [10] Kutschera U. edu/ ~smithch/ chronob/ RENS1900. Tjeerd 1994. [6] Smocovitis. nap. 1982. New York.[77] 169 See also • • • • • • • • Developmental systems theory Evolution Gene-centered view of evolution History of evolutionary thought Particulate inheritance theory Polymorphism (biology) Population genetics The Origin of Species Footnotes [1] "The scientific consensus around evolution is overwhelming". [24] Mayr and Provine 1998 pp. Weidenfeld & Nicolson. php?record_id=6024& page=28) (Second ed. Nature 442.J. [30] Van Andel. ISBN ISBN-0-309-06406-6.J. 2nd ed 1940 as Embryos and ancestors. London. 1944. (ed) 2008. 1930. Symbiosis as a source of evolutionary innovation. The origin of the eukaryote cell is one of the seven major transitions. N. [65] Lewis E. according to these authors. N. [55] Dunn.Y. p505. The bithorax complex. Murray. The Universal Ancestor. [77] Janvier. 4115. Uprooting the tree of life. 170 . N. The cell in development and heredity . Oxford. Here we use the terms as part of the evolutionary synthesis: they do not imply any change in mechanism. From Embryology to Evo-Devo: a history of developmental evolution. Paris. Clusters of master control genes regulate the development of higher organisms. Fossils. Evolution above the species level.Y. 1927. 1976. H. 2124–2128.C. 1992. [57] de Beer. 3rd ed 1958. [36] Knoll A.J. Macroevolution: pattern & process. The major transitions in evolution. San Francisco. 403–430. Freeman. Columbia. 2009. N. Oxygen: the molecule that made the world. 1996. [54] Doolittle. Berlin. 1984. [69] Shubin N. Ford 1999. Fieldiana Memoires (Zoology) 3. 1992. same title. 1987. Macmillan. 1947. [38] Hallam A. Ontogeny and phylogeny. Kent State. PNAS USA 81. Master control systems in development and evolution: the homeobox story. The Proterozoic biosphere: a multi-disciplinary study. Harvard.uk/jac/ |title=Jenny Clack homepage}} [76] Both whale evolution and early insect flight are discussed in Raff R. Charles Doolittle Walcott: paleontologist. Symbiosis in cell evolution: microbial communities in the Archaean and Proterozoic eras. Der Problem der Menschwerdung. 2002. ultrabithorax and fushi tarazu loci of Drosophila. genes and evolution: the developmental-genetic basis of evolutionary changes. 285–319. and Klein (eds) 1992. 1926. Macmillan. p186–188. 1979. Structural relationships among genes that control developmental sequence homology between the antennipedia. London. The science of life. MIT. 2007. 1995. Bloomington. Charles 1872.Y. Norton. [66] McGinnis W. Blackwell. [50] Margulis L. [44] Wilson E. January 1994. Tempo and mode in evolution. The major transitions in evolution. Baltimore. The shape of life. [70] Shubin N. Phylogenetic classification and the Universal Tree. p357 [60] Raff R. [45] Walin I. [48] Lederberg J. Cradle of life: the discovery of Earth's earliest fossils. Jenny A.B. [49] Margulis L and Fester R (eds) 1991. Gaining Ground: the origin and evolution of tetrapods. and Holland H. Oxford. Yale. The giant panda: a morphological study of evolutionary mechanisms. Oxford.G. 1993. [72] Davies D. Strassburg. Indiana. Physiological Reviews 32. Gavin 1930. and Wignall P. Tabin C and Carroll S. p818–823. Nature 388. Cell genetics and hereditary symbiosis.D. London vol 2. Philip 2008. [46] Wells H. et al 2009. Medical Assoc. et al 1984. World Scientific. Columbia. 267. Endless forms most beautiful: the new science of Evo-Devo and the making of the animal kingdom. 1925. 1997. and Szathmáry E. [37] Huggett. A conserved DNA sequence in homeotic genes of the Drosophila antennipedia and bithorax complexes. new ed. The making of a fly. and Wells G. Concepts of symbiogenesis: a historical and critical study of the research of Russian scientists. Broad phylogenetic sampling improves resolution of the animal tree of life. Cambridge University Press. [43] Khakhina. Nature 245. Verso. 1952. [39] Elewa A. 2005.M. [59] short-list of 25 characters reprinted in Gould. [51] Maynard Smith J. Nature 457. W. 1871. N. Huxley J. This section (The ABC of genetics) was written by Huxley.P. They were also used by some non-Darwinian evolutionists such as Yuri Filipchenko and Richard Goldschmidt. Nick 2002. [73] Stanley Steven M. 1524–1531. Macmillan. Casey W. [80] Darwin.theclacks. The panda is a bear. [68] Gehring W. Nobel Prize lecture. 1999. Rev. Freeman. Stephen Jay 1977.B. Effects of past climates upon life. and Szathmáry E. [58] Bolk. J. 1995. Ohio. and Kaufman C. genes and the evolution of animal limbs. Nature 454. [33] Lane. Die Erscheinung der Symbiose.D. W. Am.B. 1997. The whole of Chapter 7 in this edition is taken up with answering critics of natural selection.A. 1997. [41] Maynard Smith J.E..B. Physiology or Medicine. L. Springer. [63] Lawrence P. Chicago. [64] Duncan I. Liya Nikolaevna 1992. The bithorax complex: the first fifty years. Jena. 169 [78] Lamark J. 1997.H.Y. Mass extinctions and their aftermath. In National Research Council. p157 [74] Clack. 1964. Ford 2000.A. [53] Doolittle. ISBN 0-253-34054-3 [75] ||cite web |url=[http://www.T. Nobel lectures. 6th ed. and Weiner A. Nature 452. [47] Sapp. 1809. [56] Laubichler M. and Rensch B. and Maienschein J. 1998. 1–339. Princeton. [40] The terms (or their equivalents) were used as part of the synthesis by Simpson G.B. Repr. Ellis L. 1999. Deep homology and the origins of evolutionary novelty. Carl 1998. Oxford. Williams & Wilkins. 1997. [79] Mivart St G. 1983. [35] Yochelson. Symbionticism and the origin of species.A. Embryology and evolution.P. The origin of species. 639–648. Fischer. Ann. The genesis of species.Modern evolutionary synthesis [32] Schopf J. N. 6854–6859. Philosophie zoologique. National Academy. [42] de Bary. These discussions provide a welcome synthesis of evo-devo and paleontology. Singapore. Oxford. Embryos.Y. 428–433. [62] Lewis E. [52] Woese. MIT.G. (ed) 1997. Oxygen and Proterozoic evolution: an update. [67] Scott M.org. Scientific American 282 (6): 90–95. 1879. Nature 308. Squint of the fossil flatfish. [34] Schopf J. Genetics 21. Washington D. Sean B. Richard J. 218–220.W.W. Science 284. Mass extinctions. Oxford. [71] Sarich V.Y. [61] Carroll. in Ringertz N. PNAS 95. Tabin C and Carroll S. 745–749. Evolution by association: a history of symbiosis. Catastrophism. 1986. W. Evolutionary Biology. 1978 ISBN 0-691-08200-6 • Bowler. Longman. R. Nature 454. 1942 ISBN 0-02-846800-7 • Larson. T. [82] Goldschmidt R. The Evolutionary Synthesis: Perspectives on the Unification of Biology. (2003). ISBN 0-691-02442-1 • Huxley. Systematics and the Origin of Species. 2:30. 1998 ISBN 0-674-27225-0 • Simpson. ISBN 0-52023693-9. . The Genetical Theory of Natural Selection. S. The new biology: beyond the Modern Synthesis (http://www. J. ISBN 0-674-00613-5. Lynn and Dorion Sagan. S. Peter J. Biology Direct 2007. Stephen Jay (2002). Thomas Hunt Morgan: The Man and His Science. Allen and Unwin. Some aspects of evolution. V. Harvard University Press reprint ISBN 0-674-86250-3 • Mayr. Perseus Books Group. Evolution: The Modern Synthesis. The New Systematics. Princeton University Press reprint. D. Princeton University Press.pdf). Betty. Green and Co. A review of biology in light of recent innovations since the initiation of modern synthesis. S. and W. Clarendon Press. 1937 ISBN 0-231-05475-0 • Fisher. The evolutionary origin of flatfish asymmetry. Provine. Yale. p. 1944 ISBN 0-231-05847-0 • Smocovitis.com/ content/pdf/1745-6150-2-30. The Blind Watchmaker. 1942. 209–212. 12 0-87-893189-9 • Gould. 1940. 1940 ISBN 0-403-01786-6 • Huxley. Modern Library. 171 References • Allen. Oakley TH. • Margulis. 1932. E. J. B. Evolution:The Remarkable History of a Scientific Theory. 2002 ISBN 0-465-04391-7 • Mayr. Tempo and Mode in Evolution. Harvard University Press. Genetics and the Origin of Species. Science 78. ISBN 0-679-64288-9. Garland. "Acquiring Genomes: A Theory of the Origins of Species". J. The Structure of Evolutionary Theory. Columbia University Press. • Dawkins. E. [83] Friedman. Sinauer Associates. The material basis of evolution. External links • Rose MR. 1996 ISBN 0-691-27226-9 • Wright. Columbia University Press. Belknap Press of Harvard University Press. University of California Press. 1931. Norton and Company.. eds. 1930 ISBN 0-19-850440-3 • Futuyma. Oxford University Press. B. Columbia University Press. S.J.Modern evolutionary synthesis [81] Goldschmidt R. Genetics 16: 97–159. Edward J. Matt 2008. ed. (2004). G. 539–547.. A. Reissue Edition 1996 ISBN 0-393-31570-3 • Dobzhansky. The Causes of Evolution. • Haldane.W. Unifying Biology: The Evolutionary Synthesis and Evolutionary Biology. Princeton University Press. Richard. Evolution:The History of an Idea. "Evolution in Mendelian populations".biology-direct. G. " "bloodline. Abu al-Qasim al-Zahrawi (known as Albucasis in the West). and first described the struggle for existence. In this work. An opposing school of thought. in his Al-Tasrif. Aeschylus. and Aristotle thought that male and female semen mixed at conception. These pangenes subsequently moved through their blood to the genitals and then to the children.[1] [2] His ideas on the struggle for existence in the Book of Animals have been summarized as follows: "Animals engage in a struggle for existence. Environmental factors influence organisms to develop new characteristics to ensure survival. History The ancients had a variety of ideas about heredity: Theophrastus proposed that male flowers caused female flowers to ripen. Animals that survive to breed can pass on their successful characteristics to offspring. The inheritance of acquired traits also formed a part of early Lamarckian ideas on evolution. proposed the male as the parent." Various hereditary mechanisms were envisaged without being properly tested or quantified. the Arab physician. and prenatal influences of the womb. Ovists thought women carried eggs containing boy and girl children. which includes the field of epigenetics. The study of heredity in biology is called genetics. a hereditary genetic disorder." "full-blooded. experimentally tested and disproved pangenesis during the 1870s. and that sperm merely stimulated the growth of the egg. the Afro-Arab writer Al-Jahiz considered the effects of the environment on the likelihood of an animal to survive. with the female as a "nurse for the young life sown within her. the ovists. The terms "blood relative. The concept originated with the ancient Greeks. Nevertheless. Hippocrates speculated that "seeds" were produced by various body parts and transmitted to offspring at the time of conception. Francis Galton. These scientists formed a school of thought known as the "spermists. Through heredity. Pangenesis was an idea that males and females formed "pangenes" in every organ.172 Base concepts Heredity Heredity is the passing of traits to offspring (from its parent or ancestors). to avoid being eaten and to breed.[4] During the 18th century. in 458 BC." They contended the only contributions of the female to the next generation were the womb in which the homunculus grew." and "royal blood" are relics of pangenesis. for resources."[3] In 1000 AD. believed that the future human was in the egg. he wrote of an Andalusian family whose males died of bleeding after minor injuries. Charles Darwin's cousin. This is the process by which an offspring cell or organism acquires or becomes predisposed to the characteristics of its parent cell or organism. variations exhibited by individuals can accumulate and cause a species to evolve. and that the gender of the offspring was determined well before conception. thus transforming into new species. In the 9th century AD. wrote the first clear description of haemophilia. Some scientists speculated they saw a "little man" (homunculus) inside each sperm. These included blending inheritance and the inheritance of acquired traits. people were able to develop domestic breeds of animals as well as crops through artificial selection. and influenced biology until as recently as a century ago. . Dutch microscopist Antonie van Leeuwenhoek (1632–1723) discovered "animalcules" in the sperm of humans and other animals. Correlation genotype–phenotype • Dominant • Intermediate (also called "codominant") • Recessive These three categories are part of every exact description of a mode of inheritance in the above order. Additionally.. cryptorchism) Inheritance through the maternal line (in case of mitochondrial DNA loci) Inheritance through the paternal line (in case of Y-chromosomal loci) 6. The effects of this allele are only seen when it is present in both chromosomes. Since the allele for green pods is dominant. Sex-linked interactions • • • • Sex-linked inheritance (gonosomal loci) Sex-limited phenotype expression (e. Locus–locus interactions • Epistasis with other loci (e. For example. pea plants with the pair of alleles GG (homozygote) or Gg (heterozygote) will have green pods.. G. Number of involved loci • Monogenetic (also called "simple") – one locus • Oligogenetic – few loci • Polygenetic – many loci 2. The allele for yellow pods is recessive.g. more specifications may be added as follows: 4. g. overdominance) • Gene coupling with other loci (also see crossing over) • Homozygotous lethal factors • Semi-lethal factors .Heredity 173 Types of heredity Dominant and recessive An allele is said to be dominant if it is always expressed in the appearance of an organism (phenotype).g. Coincidental and environmental interactions • Penetrance • Complete • Incomplete (percentual number) • Expressivity • Invariable • Variable • Heritability (in polygenetic and sometimes also in oligogenetic modes of inheritance) • Maternal or paternal imprinting phenomena (also see epigenetics) 5. gg (homozygote). Involved chromosomes • Autosomal – loci are not situated on a sex chromosome • Gonosomal – loci are situated on a sex chromosome • X-chromosomal – loci are situated on the X chromosome (the more common case) • Y-chromosomal – loci are situated on the Y chromosome • Mitochondrial – loci are situated on the mitochondrial DNA 3. is dominant to that for yellow pods. The description of a mode of biological inheritance consists of three main categories: 1. in peas the allele for green pods. The primacy of population thinking: the genetic diversity carried in natural populations is a key factor in evolution. recombination ordered by natural selection. Gregor Mendel: father of modern genetics The idea of particulate inheritance of genes can be attributed to the Moravian[5] monk Gregor Mendel who published his work on pea plants in 1865. Historical contingency means explanations at different levels may exist. . etc. The strength of natural selection in the wild was greater than expected. Darwin believed in a mix of blending inheritance and the inheritance of acquired traits (pangenesis). even slight advantages are important when continued. Galton rejected the aspects of Darwin's pangenesis model which relied on acquired traits. 4. and then heavily modified by. the effect of ecological factors such as niche occupation and the significance of barriers to gene flow are all important. his work was not widely known and was rediscovered in 1901. 3. However. The inheritance of acquired traits was shown to have little basis in the 1880s when August Weismann cut the tails off many generations of mice and found that their offspring continued to develop tails. the ability to explain historical observations by extrapolation from micro to macro-evolution is proposed. one of its major problems was the lack of an underlying mechanism for heredity. work by Fisher and others resulted in a combination of Mendelian and biometric schools into the modern evolutionary synthesis. who laid the framework for the biometric school of heredity. All evolutionary phenomena can be explained in a way consistent with known genetic mechanisms and the observational evidence of naturalists. The role of genetic drift is equivocal.Heredity Determination and description of a mode of inheritance is primarily achieved through statistical analysis of pedigree data. and between both and palaeontologists. Blending inheritance would lead to uniformity across populations in only a few generations and thus would remove variation from a population on which natural selection could act. though strongly supported initially by Dobzhansky. The object of selection is the phenotype in its surrounding environment. 174 Charles Darwin: theory of evolution When Charles Darwin proposed his theory of evolution in 1859. In case the involved loci are known.A. 5. methods of molecular genetics can also be employed. such as those seen by Mendel in his pea plants—and the idea of additive effect of (quantitative) genes was not realised until R. his cousin Francis Galton.) rather than suggesting mechanisms. It was initially assumed the Mendelian inheritance only accounted for large (qualitative) differences. Darwin's primary approach to heredity was to outline how it appeared to work (noticing that traits could be inherited which were not expressed explicitly in the parent at the time of reproduction. it was downgraded later as results from ecological genetics were obtained. Gradualism does not mean constant rate of change. that certain traits could be sex-linked." Modern development of genetics and heredity In the 1930s. stating that:[6] [7] 1. Darwin's initial model of heredity was adopted by. "The Correlation Between Relatives on the Supposition of Mendelian Inheritance. Selection is overwhelmingly the main mechanism of change. In palaeontology. Evolution is gradual: small genetic changes. Discontinuities amongst species (or other taxa) are explained as originating gradually through geographical separation and extinction (not saltation). 2. Fisher's (1918) paper. This led to Darwin adopting some Lamarckian ideas in later editions of On the Origin of Species and his later biological works. The modern synthesis bridged the gap between experimental geneticists and naturalists. Trofim Lysenko however caused a backlash of what is now called Lysenkoism in the Soviet Union when he emphasised Lamarckian ideas on the inheritance of acquired traits. p221-2 External links • Stanford Encyclopedia of Philosophy entry on Heredity and Heritability (http://plato. net. Formulations such as 'evolution consists primarily of changes in the frequencies of alleles between one generation and another' were proposed rather later. The growth of biological thought: diversity. Robin Marantz (2000). Encyclopedia of Alternative Medicine [5] Henig. co. There is no doubt. [4] Patricia Skinner (2001). Intelligent Design (http:/ / www. php) [3] Gary Dargan. The traditional view is that developmental biology ('evo-devo') played little part in the synthesis. Proceedings of the American Philosophical Society 84 (1).J. uk/ knowledge/ al-jahiz. This movement affected agricultural research and led to food shortages in the 1960s and seriously affected the USSR. Harvard. Natural Selection before the "Origin of Species". In plants. It cleared up many confusions. written by an obscure Moravian monk named Gregor Mendel" [6] Mayr & Provine 1998 [7] Mayr E. ABC. 71-123. 1982. but an account of Gavin de Beer's work by Stephen Jay Gould suggests he may be an exception. and was directly responsible for stimulating a great deal of research in the post-World War II era. Encounter. The Monk in the Garden : The Lost and Found Genius of Gregor Mendel. The Islamic Quarterly. Cambs. p567 et seq. "The article.edu/entries/ heredity/) . however. 1983). the Father of Genetics. with varying degrees of success. p. polyploidy must be included in any view of speciation.Heredity The idea that speciation occurs after populations are reproductively isolated has been much debated. Ontogeny and phylogeny. that the synthesis was a great landmark in evolutionary biology. Harvard 1977. Unani-tibbi (http:/ / findarticles. 175 See also • Genetics • Hard inheritance • Heritability Notes and references [1] Conway Zirkle (1941). abc. ISBN 0-395-97765-7.stanford. htm). Houghton Mifflin. evolution & inheritance. (http:/ / www. salaam. [8] Gould S. London. au/ rn/ encounter/ stories/ 2006/ 1656361. "Al-Jahiz And the Rise of Biological Evolutionism". com/ p/ articles/ mi_g2603/ is_0007/ ai_2603000716).[8] Almost all aspects of the synthesis have been challenged at times. [2] Mehmet Bayrakdar (Third Quarter. including 0. a ratio smaller than 1. the alleles with higher fitness become more common. Relative fitness can therefore take any nonnegative value. Absolute fitness Absolute fitness ( ) of a genotype is defined as the ratio between the number of individuals with that genotype after selection to those before selection. it is equal to the average contribution to the gene pool of the next generation that is made by an average individual of the specified genotype or phenotype.e. If differences between alleles at a given gene affect fitness. but depend on the environment in which the individuals live. As phenotype is affected by both genes and environment. However. It is calculated for a single generation and may be calculated from absolute numbers or from frequencies.Fitness 176 Fitness Fitness (often denoted in population genetics models) is a central idea in evolutionary theory. which is weighted by genotype frequencies. By taking the logarithm of fitness each term may be added rather than multiplied. Because fitness is a coefficient. Measures of fitness There are two commonly used measures of fitness. one genotype is normalized at and the fitnesses of other genotypes are measured with respect to that genotype. the fitnesses of different individuals with the same genotype are not necessarily equal. absolute fitness and relative fitness. When the fitness is larger than 1. and both of them are equivalent when they are divided by the mean fitness. . the genotype increases in frequency. Absolute fitness for a genotype can also be calculated as the product of the proportion survival times the average fecundity. absolute fitness is more difficult. i. since the fitness of the genotype is an averaged quantity. biologists may work with "log fitness" (particularly so before the advent of computers). in preference to individuals with a different allele. This process is called natural selection. then the frequencies of the alleles will change over generations.0. It is often difficult to determine how many individuals of a genotype there were immediately after reproduction.0 indicates a decrease in frequency. Relative fitness Relative fitness is quantified as the average number of surviving progeny of a particular genotype compared with average number of surviving progeny of competing genotypes after a single generation. it will reflect the reproductive outcomes of all individuals with that genotype. An individual's fitness is manifested through its phenotype. Inclusive fitness differs from individual fitness by including the ability of an allele in one individual to promote the survival and/or reproduction of other individuals that share that allele. and a variable may be multiplied by it several times. It can be defined either with respect to a genotype or to a phenotype. While researchers can usually measure relative fitness. In either case. One mechanism of inclusive fitness is kin selection. The two concepts are related. Fitness 177 Maynard-Smith's Definition As another example we may mention the definition of fitness given by Maynard Smith in the following way: "Fitness is a property.keeping the mean fitness constant . In the case N is a Gaussian it is fairly easily proved that the average information (information entropy. this would not prove the new genotype to have low fitness. that the individual will be included among the group selected as parents of the next generation. and perhaps more accurately. Thus the phrase ’expected number of offspring’ means the average number.having an array x of phenotypes . the mean fitness may be determined as a mean over the set of individuals in a large population. disorder. History The British sociologist Herbert Spencer coined the phrase "survival of the fittest" (though originally. and m is its centre of gravity. If the first human infant with a gene for levitation were struck by lightning in its pram.D. not of an individual. The British biologist J. The next further advance was the introduction of the concept of inclusive fitness by the British biologist W. diversity) of a large population may be maximized by Gaussian adaptation . Peaks correspond to local fitness maxima. Haldane was the first to quantify fitness. s(x). This can result in suboptimal local maxima becoming stable. and each of the other dimensions represents allele identity for a different gene." [1] Hartl's Definition Yet another possible measure has been formulated: "The fitness of the individual . Hamilton in 1964 in his paper on The Evolution of Social Behavior. first conceptualized by Sewall Wright. This measure is a suitable basis of a model of an evolution selecting individuals.B. but only that the particular child was unlucky. the central limit theorem. It may in principle take even the stroke of the lightning into consideration. "survival of the best fitted") in his 1851 work Social Statics and later used it to characterise what Charles Darwin had called natural selection." Then.S. because natural selection cannot return to the less-fit "valleys" of the landscape on the way to reach higher peaks. This is in contrast to Fisher's fundamental theorem of natural selection. is a way of visualising fitness in terms of a high-dimensional surface. in which height indicates fitness. it is often said that natural selection always progresses uphill but can only do so locally.in accordance with recapitulation. the Hardy-Weinberg law and the second law of thermodynamics. not the number produced by some one individual. [2] where N is the probability distribution function of phenotypes in the population. in terms of the modern evolutionary synthesis of Darwinism and Mendelian genetics starting with his 1924 paper A Mathematical Theory of Natural and Artificial Selection. . but of a class of individuals – for example homozygous for allele A at a particular locus. Fitness landscape A fitness landscape.is the probability. Thinking about Evolution: Historical. D. and Political Perspectives. PMID 19546856. Singh.fcgi?tool=pmcentrez&artid=2753274).309-321. The Two Faces of Fitness. Krimbas.blackwellpublishing. PMC 2753274. 10 (8): 531–9.wisc. and J.edu/entries/fitness/) . External links • Evolution A-Z: Fitness (http://www. E. D.Fitness 178 Genetic load Genetic load measures the average fitness of a population of individuals. pp. Nat.com/ridley/a-z/Fitness. See also • • • • • Gene-centered view of evolution Inclusive fitness Natural selection Reproductive success Selection coefficient Notes [1] Maynard-Smith.edu/sober/tff.).pdf) • Orr HA (August 2009).gov/ articlerender. Philosophical. In R. C. J. relative to a hypothetical population in which the most fit genotype has become fixed. Full text (http://philosophy. doi:10. (2001). (1981) A Primer of Population Genetics ISBN 0878932712 Further reading • Sober.nih.stanford.asp) • Stanford Encyclopedia of Philosophy entry (http://plato. Genet. Beatty (Eds. Paul. (1989) Evolutionary Genetics ISBN 0198542151 [2] Hartl. Rev. Cambridge University Press. L.pubmedcentral. "Fitness and its role in evolutionary genetics" (http://www.1038/nrg2603. sensations. it is not to be wondered at that this fitness is present in all the species that are currently in existence? Chance. Richard Dawkins coined the word concestor. and associations.. there was neither fitness nor order: all of these latter have perished. This new word is very gradually entering scientific parlance. Immanuel Kant wrote in Kritik der Urtheilskraft (Critique of Judgement) that the analogy of animal forms implies a common original type.[6] History In the 1740s. one would say. since the earth began to exist. in the fortuitous combinations of the productions of nature. the most recent common ancestor of all currently living organisms.[9] In 1790. Charles Darwin's grandfather.[2] Charles Darwin proposed the theory of universal common descent through an evolutionary process in On the Origin of Species.[10] In 1795.9 billion years ago. by a formal test"[1] for the theory that all living organisms on Earth are descended from a common ancestor. Maupertuis noted: Could one not say that. others lacking reproductive organs could not perpetuate themselves . and thus possessing the faculty of continuing to improve by its own inherent activity. directed by irritations. "There is strong quantitative support. asked: [W]ould it be too bold to imagine. Erasmus Darwin. produced an innumerable multitude of individuals. having been originally breathed into a few forms or into one.[3] The last universal ancestor (LUA) (or last universal common ancestor.[1] is believed to have appeared about 3. with its several powers. LUCA). in another infinitely greater number. world without end?[11] In 1859.[7] [8] In Essai de Cosmologie. that all warm-blooded animals have arisen from one living filament. having been originally breathed into a few forms or into one". with its several powers. Charles Darwin's The Origin of Species was published. with the power of acquiring new parts attended with new propensities. a small number found themselves constructed in such a manner that the parts of the animal were able to satisfy its needs. Pierre-Louis Moreau de Maupertuis made the first known suggestion in a series of essays that all organisms may have had a common ancestor. volitions.[4] [5] In The Ancestor's Tale.. into which life was first breathed. The species we see today are but the smallest part of what blind destiny has produced . "Therefore I should infer from analogy that probably all the organic beings which have ever lived on this earth have descended from some one primordial form. would it be too bold to imagine. which the great First Cause endued with animality.Common descent 179 Common descent In evolutionary biology. The views about common descent expressed therein were that it was possible that there was only one progenitor for all life forms. Animals lacking a mouth could not live. saying.. and thus a common parent."[3] (p 490) . "There is a grandeur in this view of life. that in the great length of time. perhaps millions of ages before the commencement of the history of mankind. and of delivering down those improvements by generation to its posterity. and that they had diverged through random variation and natural selection. as there must be some characterized by a certain relation of fitness which are able to subsist. that is. a group of organisms have common descent if they have a common ancestor." [3] (p 484) Darwin's famous closing sentence describes the "grandeur in this view of life. as a substitute for common ancestor or most recent common ancestor.. [12] A statistical comparison of various alternative hypotheses has shown that universal common ancestry is significantly more probable than models involving multiple origins. The universality of this code is generally regarded by biologists as definitive evidence in favor of the theory of universal common descent. transcribed into RNA. rather than universal common descent and therefore resulted in convergent evolution. . any irrelevant similarity between the sequences is evidence for common descent.[1] for the unity of life. and the fact that all amino acids found in proteins are left-handed. In 2010 an analysis of available genetic data. such as appearance. It is possible that these similarities resulted because of the laws of physics and chemistry.there is now strong quantitative support. Such evidence has come from two areas: amino acid sequences and DNA sequences. . etc. these trees have been built using morphological methods.. with ATP. Analysis of the small differences in the genetic code has also provided support for universal common descent. Furthermore. A phylogenetic tree based on rRNA genes. if two species use the same codon at the same place to specify an amino acid that can be represented by more than one codon. NADH and others as energy sources. These similarities include the energy carrier adenosine triphosphate (ATP).. In certain cases. "genealogic trees" of species) mapping out the proposed divisions and common ancestors of all living species. through the effect of protein. etc. gave "firm quantitative support for the unity of life. mapping them to phylogenetic trees. and therefore they provide compelling support for the theory of universal common descent.[13] Selectively neutral similarities Similarities which have no adaptive relevance cannot be explained by convergent evolution. Phylogenetic trees Another important piece of evidence is that it is possible to construct detailed phylogenetic trees (that is. from bacteria to humans. there are several codons (DNA triplets) that code for the same amino acid. the genetic code (the "translation table" according to which DNA information is translated into proteins) is nearly identical for all known lifeforms.Common descent 180 Evidence of universal common descent Common biochemistry and genetic code All known forms of life are based on the same fundamental biochemical organisation: genetic information encoded in DNA. that is evidence for a recent common ancestor. Thus.[2] Traditionally. then translated into proteins by (highly similar) ribosomes.and RNA-enzymes. by a formal test. Other similarities The universality of many aspects of cellular life is often pointed to as supportive evidence to the more compelling evidence listed above. Proteins with the same three-dimensional structure need not have identical amino acid sequences. embryology. Wild cabbage Early farmers cultivated many popular vegetables from the Brassica oleracea (wild cabbage) by artificially selecting for certain attributes. kale.[14] 181 Illustrations of common descent Artificial selection Artificial selection demonstrates the diversity that can exist among organisms that share a relatively recent common ancestor. even though that most genetic variation has no influence over external morphology. having descended from wolves. Great Dane. which did not undergo artificial selection. In artificial selection. cauliflower. Basset Hound. Cabbage was created by selecting for short petioles. Wild wolves. are relatively uniform in comparison. broccoli. Humans selectively bred them to enhance specific characteristics. That phylogenetic trees based on different types of information agree with each other is strong evidence of a real underlying common descent. and Poodle. such as color and length or body size. The Chihuahua mix and Great Dane illustrate the range of sizes among dog breeds.[15] Brussels sprouts were created by artificially selecting for large bud size. Dog breeding The diversity of domesticated dogs is an example of the power of artificial selection. Artificial selection was successful long before science discovered the genetic basis. Broccoli was bred by selecting for large flower stalks. Kale was bred by selecting for large leaves. These characteristics become increasingly well-developed in successive generations. based on similarities and differences between genetic and protein sequences. one species is bred selectively at each generation. Wild Cabbage plant . it has been possible to construct these trees using molecular data. All these methods produce essentially similar results. Common vegetables such as cabbage. Pug. All breeds share common ancestry. allowing only those organisms that exhibit desired characteristics to reproduce. kohlrabi and Brussels sprouts are all descendants of the wild cabbage plant.Common descent Recently. This created a range of breeds that include the Chihuahua. Darwin observed 13 species of finches that are closely related and differ most markedly in the shape of their beaks. [14] Theobald. their slender beaks allowed the birds to be better equipped for pulling out the insects from their tiny hiding places. p. ISBN 9780872200258. suggesting that beak shapes evolved by natural selection. com/ ?id=XRUaAAAAMAAJ& printsec=frontcover& q=).1038/nature09014. PMID 20463738. 1981. 1970. ISBN 9780416723700..1038/35047500. London.. com/ books?id=OOgzAAAAIAAJ& pg=PA62& dq=Maupertuis+ "for+ the+ first+ time"& cd=1#v=onepage& q=Maupertuis "for the first time"& f=false). Leon Harris. [6] Singham. Darwin's finches During Charles Darwin's studies on the Galápagos Islands. genesis and revelations. "The Origin of Species by Means of Natural Selection. Nature 465 (7295): 219–222. 304: "Despite all the variety among these forms. "A formal test of the theory of universal common ancestry. The making of modern Europe. et al. 1985. com/ articles/ LookingForDeep. pg. msu. com/ books?id=y5wQD-ioaUUC& pg=PA304& dq=Critique+ of+ Judgment+ "original+ mother"& cd=1#v=onepage& q=& f=false). ISBN 9780873954877. C. edu/ courses/ mb437_537_2004_fall/ docs/ uprooting. produced by a common original mother. Slender beaks were found on the finches which found insects to be the best source of food on the island they inhabited. CUP Archive. Pluhar. (13 May 2010). 2009. they seem to have been produced according to a common archetype. p. 2006). Biology Direct 2008. ISBN 9780521075244. Zoonomia. (1859) p. John Murrary. "Generation" (http:/ / books. Ying Xu & Bernard Labedan: The Last Universal Common Ancestor : emergence. Taylor & Francis. Douglas L. with readings from Empedocles to Wilson (http:/ / books. Retrieved November 19. Retrieved November 20. 2009. [12] Knight. 29+ Evidences for Macroevolution. Robin. [7] J. W. D.8]. google. TalkOrigins Foundation. or the Laws of Organic Life. Evolution. org/ faqs/ comdesc/ section1.L.Common descent 182 Natural selection Natural selection is the evolutionary process by which heritable traits that increase an individual's fitness become more common. html). Machines Like Us. 62-63. 142 [9] C. google. "Prediction 1. google. "Looking for Deep Ancestors" (http:/ / machineslikeus. [8] Geoffrey Russell Richards Treasure. The finch is also found on the main land and it is thought that they migrated to the islands and began adapting to their environment through natural selection. google. The new Cambridge modern history: The rise of Great Britain and Russia. PMID 11253070. References [1] Theobald. 490 [4] Doolittle.. Critique of Judgment (http:/ / books. or the Preservation of Favoured Races in the Struggle For Life". google. . 2000).". com/ books?id=Fkqg3TIkBJwC& pg=PA107& dq=Essai+ de+ Cosmologie+ "blind+ destiny"& cd=1#v=onepage& q=Essai de Cosmologie "blind destiny"& f=false). . Scientific American 282 (6): 90–95. Uprooting the tree of life (http:/ / shiva. and heritable traits that decrease an individual's fitness become less common. 1648-1780 (http:/ / books. [5] Nicolas Glansdorff. & Penny. Bromley. D. pgs. Large beaks were found on the Darwin's finches islands where the primary source of food for the finches is nuts and therefore the large beaks allowed the birds to be better equipped for opening the nuts and staying well nourished. Douglas (2004). Nature 465. Retrieved November 20. 219-222. constitution and genetic legacy of an elusive forerunner. 1 (4th American ed. 3:29. pdf). Philadelphia: Edward Earle. 397 [§ 39. "Rewiring the Keyboard: Evolvability of the Genetic Code". [13] Theobald." [11] Darwin. .4. doi:10. com/ books?id=C94OAAAAQAAJ& pg=PA142& dq="blind+ destiny+ has+ produced"& cd=6#v=onepage& q="blind destiny has produced"& f=false). and this analogy among them reinforces our suspicion that they are actually akin.168 (2010) [3] Darwin. 1688-1715/25 (http:/ / books. 2009. . SUNY Press. S. doi:10. Erasmus (1818) [1795]. 107 [10] Immanuel Kant and Werner S. Nature Reviews Genetics 2 (1): 49–58. Nature 465. Ford (February.). html#independent_convergence). M. (January 2001). Mano (October 13. (2010) [2] Steel. 1987. The beak of each species is suited to the food available in its particular environment. Hackett Publishing.3: Consilience of Independent Phylogenies" (http:/ / www. pg. talkorigins. montana. (2005). Further evidence comes from the field of biogeography because evolution with common descent provides the best and most thorough explanation for a variety of facts concerning the geographical distribution of plants and animals across the world. Universal biochemical organization and molecular variance patterns in all organisms also show a direct correlation with common descent. Further evidence for common descent comes from genetic detritus such as pseudogenes. ISBN 0716762846.html) Evidence of common descent A large body of evidence of common descent of living things has been discovered by scientists working in a variety of fields over many years. OCLC 183148564.). As fossilization is an uncommon occurrence. bones and teeth is especially scarce. This evidence has demonstrated and verified the occurrence of evolution and provided a wealth of information on the natural processes by which the variety and diversity of life on Earth developed. ed. the fossil record only provides sparse and intermittent information about the evolution of life. Combined with the theory of plate tectonics common descent provides a way to combine facts about the current distribution of species with evidence from the fossil record to provide a logically consistent explanation of how the distribution of living organisms has changed over time. The comparative study of the anatomy of groups of animals reveals structural features that are fundamentally similar or homologous. clearly demonstrating phylogenetic and ancestral relationships with other organism. but are no longer active and appear to be undergoing a steady process of degeneration. This is especially obvious in the field of island biogeography. Vestigial structures and comparisons in embryonic development are largely a contributing factor in anatomical resemblance in concordance with common descent. testing hypotheses. most especially when compared with fossils of ancient extinct organisms. Comparison of the genetic sequence of organisms has revealed that organisms that are phylogenetically close have a higher degree of sequence similarity than organisms that are phylogenetically distant. Evolutionary biologists document the fact of common descent: making testable predictions. like the spread of pesticide resistant forms of plants and insects provides evidence that evolution due to natural selection is an ongoing process in the natural world. Furthermore. Multiple forms of such have been described and documented as examples for individual modes of speciation. Freeman. so it is possible to determine when certain metabolic processes appeared by comparing the traits of the descendants of a common ancestor. From the TalkOrigins Archive • The Tree of Life Web Project (http://tolweb.org/tree/phylogeny. Biology of Plants (7th rev. The development and spread of antibiotic resistant bacteria. a member of the mustard family. Speciation has been observed directly and indirectly in the lab and in nature. Alongside this. New York: W. Evidence of organisms prior to the development of hard body parts such as shells. research into the evolution of the basic cellular processes is done largely by comparison of existing organisms’ physiology and biochemistry. Many lineages diverged at different stages of development. usually requiring hard body parts and death near a site where sediments are being deposited. and developing theories that illustrate and describe its causes.H." 183 External links • 29+ Evidences for Macroevolution: the Scientific Case for Common Descent (http://www.Common descent [15] Raven. evidence of .org/faqs/ comdesc/).. Since metabolic processes do not leave fossils. "[These vegetables were] all produced from a single species of plant (Brassica oleraca). but exists in the form of ancient microfossils. are observed instances of the separation of populations of species into sets of new species (speciation).talkorigins. et al. Peter H. Fossils are important for estimating when various lineages developed in geologic time. This evidence supports the modern evolutionary synthesis. the current scientific theory that explains how and why life changes over time. as well as impressions of various soft-bodied organisms. regions of DNA that are orthologous to a gene in a related organism. including repeats. Notably they will have inherited mutations unique to that ancestor. or RNA for viruses). the evolutionary history of extinct organisms). Such reconstructions. if there is one best way to produce a hoof. However. the product of shared genetic elements. Perhaps most tellingly. and mutations in protein-coding sequences that do not result in changes in amino-acid sequence. all hoofed creatures will share a genetic basis even if they are not related). The simplest and most powerful evidence is provided by phylogenetic reconstruction. polymers of amino acids) by highly conserved ribosomes. many new to science. For example. transcribed into RNA. More closely-related species will have a greater fraction of identical sequence and will have shared substitutions when compared to more distantly-related species. This evidence does not support the rival hypothesis that genetic similarity of two species is the product of common functional or structural requirements. This article explains the different types of evidence for evolution with common descent along with many specialized examples of each. While a minority of these elements might later be found to harbor function. in fact. Charles Darwin collected numerous specimens. 184 Evidence from comparative physiology and biochemistry Genetics Although it has only recently become available. Comparative sequence analysis examines the relationship between the DNA sequences of different species. The most detailed reconstructions have been performed on the basis of the mitochondrial genomes shared by all eukaryotic organisms. in aggregate they demonstrate that identity must be the product of common descent rather than common function. Universal biochemical organisation and molecular variance patterns All known extant organisms are based on the same fundamental biochemical organisation: genetic information encoded as nucleic acid (DNA. transposons. While on board HMS Beagle. These reconstructed phylogenies recapitulate the relationships established through morphological and biochemical studies. A deeper understanding of developmental biology shows that common morphology is. If the hypothesis of common descent is true. the Genetic Code (the "translation table" between DNA and amino acids) is the same for almost every organism.Evidence of common descent common descent extends from direct laboratory and experimentation with the artificial selection of organisms—historically and currently—and other controlled experiments involving many of the topics in the article. meaning that a piece of DNA in a bacterium codes for the same amino acid as in a human cell. one of the strongest evidences for common descent comes from the study of gene sequences. and not common descent (for example. phylogenetic relationships also extend to a wide variety of nonfunctional sequence elements. ATP is used as energy currency by all extant life. which are short and easy to sequence.[1] producing several lines of evidence that confirm Darwin's original hypothesis of common descent. especially when done using slowly-evolving protein sequences. the broadest reconstructions have been performed either using the sequences of a few very ancient proteins or by using ribosomal RNA sequence. are often quite robust and can be used to reconstruct a great deal of the evolutionary history of modern organisms (and even in some instances such as the recovered gene sequences of mammoths. pseudogenes. rex. then species that share a common ancestor will have inherited that ancestor's DNA sequence. then translated into proteins (that is. which supported his later theory of evolution by natural selection. although camera-like eyes are believed to have evolved independently on many . Neanderthals or T. they share a common set of light-sensing proteins (opsins). Other mechanisms There is also a large body of molecular evidence for a number of different mechanisms for large evolutionary changes. 1. Vital proteins. was used to find the broad phylogenetic relationships between all extant life. This is an example of a vestige since reproducing these genes uses energy and is a waste. Other overarching similarities between all lineages of extant organisms. resulted in the three-domain system. suggesting a common point of origin for all sighted creatures. this theory offers a mechanism for a sudden evolutionary leap by incorporating the genetic material and biochemical composition of a separate species. RNA. whose structure is controlled by the homeobox (Hox) family of genes. Further evidence for reconstructing ancestral lineages comes from junk DNA such as pseudogenes. the simplest hypothesis is that the choice was made randomly by early organisms and passed on to all extant life through common descent. and RNA polymerase. and 6. RNA. or dead DNA is the extra DNA that is in our genome but does not get transcribed into RNA to produce protein. horizontal gene transfer. as Junk DNA. Rather than evolving eukaryotic organelles slowly. and amino acids is conserved across all known life. such as DNA. and the lipid bilayer. arguing for two major splits in the early evolution of life. "dead" genes which steadily accumulate mutations. when it disappears. yet 99% of our genome is these (1% working DNA). the process of transferring genetic material to another cell that is not an organism's offspring. which are organelles of eukaryotic cells. it does not affect fitness. The analysis.[4] [5] The sequence of the 16S ribosomal RNA gene. among them: genome and gene duplication. Proteins The proteomic evidence also supports the universal ancestry of life. DNA sequencing Comparison of the DNA sequences allows organisms to be grouped by sequence similarity. such as the ribosome. and the resulting phylogenetic trees are typically congruent with traditional taxonomy. and recombination. largely affecting the regulation and protein-protein interaction of the core. originally done by Carl Woese.[2] Another noteworthy example is the familiar vertebrate body plan. a vital gene encoding a part of the ribosome. give support to the theory of common descent. As there is no functional advantage to right. For example.[3] Genetic sequence evidence thus allows inference and quantification of genetic relatedness between humans and other apes. Evidence supporting this mechanism has recently been found in the protist Hatena: as a predator it engulfs a green algae cell.or left-handed molecular chirality.g. The chirality of DNA. Higher organisms have evolved additional protein subunits. Sequence comparison is considered a measure robust enough to be used to correct erroneous assumptions in the phylogenetic tree in instances where other evidence is scarce. neutral human DNA sequences are approximately 1. and are often used to strengthen or correct taxonomic classifications.Evidence of common descent separate occasions. The first split led to modern Bacteria and the subsequent split led to modern Archaea and Eukaryotes. amino acids.[7] The way a pseudogene is produced is a gene that is no longer used will accumulate mutations that make it non-functional but since it is not used. allowing for species to acquire beneficial genes from each other. the chimpanzee. as the incorporation of an ancient prokaryotic cell into ancient eukaryotic cell. DNA polymerase. serving similar functions. 185 .6% from gorillas.[6] Pseudogenes Pseudogenes also known. chloroplasts).6% from baboons. The core part of the protein is conserved across all lineages of life. The Endosymbiotic theory explains the origin of mitochondria and plastids (e. are found in everything from the most primitive bacteria to the most complex mammals. capable of reassorting large numbers of different alleles and of establishing reproductive isolation.2% divergent (based on substitutions) from those of their nearest genetic relative. which facilitates rapid evolution by providing substantial quantities of genetic material under weak or no selective constraints. this leads to the conclusion that the two species diverged N years ago. the common telomeres. The closest human relative. Normally a chromosome has just one centromere. nourishing Hatena.Evidence of common descent which subsequently behaves as an endosymbiont. which in turn loses its feeding apparatus and behaves as an autotroph. the number of amino acids making up the cytochrome c protein in monkeys differ by one from that of humans. According to researcher J. The same is true of the more distant gorilla and orangutan. has near-identical DNA sequences to human chromosome 2. all living organisms have it. All Hominidae with the exception of humans have 24 pairs of chromosomes.1 and c29B is the relic of an ancient telomere-telomere fusion and marks the point at which two ancestral ape chromosomes fused to give rise to human chromosome 2. These are normally found only at the ends of a chromosome.[8] [9] Since metabolic processes do not leave fossils. IJdo. The variance of cytochrome c of different organisms is measured in the number of differing amino acids. and a vestigial centromere chimpanzee. If each differing amino acid is assumed to be the result of one base pair substitution. Humans have only 23 pairs. but they are found in two separate chromosomes. it can be calculated how long ago the two species diverged by multiplying the number of base pair substitutions by the estimated time it takes for a substituted base pair of the cytochrome c gene to be successfully passed on. because it performs very basic life functions) protein Cytochrome c in living cells. but in chromosome 2 there are additional telomere sequences in the middle. W.e. The primary structure of cytochrome c consists of a chain of about 100 amino acids.[15] Chromosome 2 thus presents very strong evidence in favour of the common descent of humans and other apes. Many higher order organisms possess a chain of 104 amino acids. research into the evolution of the basic cellular processes is done largely by comparison of existing organisms.[12] [13] • The presence of a vestigial centromere. 186 Specific examples Chromosome 2 in humans Clear evidence for the evolution of Homo sapiens from a common ancestor with chimpanzees is the number of chromosomes in human as compared to all other members of Hominidae. "We conclude that the locus cloned in cosmids c8. each differing amino acid being a result of a base pair substitution. and it is theoretically possible to determine when certain metabolic processes appeared by comparing the traits of the descendants of a common ancestor or by detecting their physical manifestations. Human chromosome 2 is widely accepted to be a result of an end-to-end fusion of two ancestral chromosomes. Many lineages diverged when new metabolic processes appeared. For example.[16] . a mutation. the appearance of oxygen in the earth's atmosphere is linked to the evolution of photosynthesis.[10] [11] The evidence for this includes: • The correspondence of chromosome 2 to two ape Fusion of ancestral chromosomes left distinctive remnants of chromosomes. but in chromosome 2 there are remnants of a second centromere. if the average time it takes for a base pair of the cytochrome c gene to mutate is N years.[14] • The presence of vestigial telomeres."[15] Cytochrome c A classic example of biochemical evidence for evolution is the variance of the ubiquitous (i. As an example. Both chicken and turkeys have identical sequence homology (amino acid for amino acid). Direct examination of the genetic structure of modern populations via DNA sequencing has recently allowed verification of many of these predictions.[20] [21] Darwin described how common descent could provide a logical basis for classification:[22] “ All the foregoing rules and aids and difficulties in classification are explained. Both humans and chimpanzees share the identical molecule. as do pigs. B. while rhesus monkeys share all but one of the amino acids:[17] the 66th amino acid is isoleucine in the former and threonine in the latter. For example. Most existing species can be organized rather easily in a nested hierarchical classification. style and ovary. several genera can be grouped together into one family. which states that modern humans developed in Africa and a small sub-population migrated out (undergoing a population bottleneck).[19] 187 Evidence from comparative anatomy Comparative study of the anatomy of groups of animals or plants reveals that certain structural features are basically similar. Specifically.[16] What makes these homologous similarities particularly suggestive of common ancestry in the case of cytochrome C. that the characters which naturalists consider as showing true affinity between any two or more species. S.. but he showed that his theory of evolution with its branching pattern of common descent could explain them. Based on shared derived characters. page 577 . Both of these predictions are borne out by actual data from a number of studies. and. The different existing configurations of amino acids do not significantly affect the functionality of the protein. yet the size. number of parts and specific structure are different for each individual species. in so far. but the result of random mutations that aren't subject to selection. petals. Ronald Fisher and J. .. the Out of Africa theory of human origins. ” —Charles Darwin. etc. are those which have been inherited from a common parent. cows and sheep. pioneered by the likes of Sewall Wright. several families can be grouped together into an order. implies that modern populations should show the signatures of this migration pattern. that community of descent is the hidden bond which naturalists have been unconsciously seeking. post-bottleneck populations (Europeans and Asians) should show lower overall genetic diversity and a more uniform distribution of allele frequencies compared to the African population. on the view that the natural system is founded on descent with modification. colour. On the Origin of Species. if I do not greatly deceive myself. in addition to the fact that the phylogenies derived from them match other phylogenies very well. all true classification is genealogical. For example. closely related organisms can be placed in one group (such as a genus).[18] Out of Africa hypothesis of human evolution Mathematical models of evolution. allow predictions about the genetic structure of evolving populations. This is evident from the Linnaean classification scheme. which indicates that the base pair substitutions are not part of a directed design.Evidence of common descent The cytochrome c molecule has been extensively studied for the glimpse it gives into evolutionary biology. is the high degree of functional redundancy of the cytochrome C molecule.[20] The existence of these nested hierarchies was recognized by many biologists before Darwin. Nested hierarchies and classification Taxonomy is based on the fact that all organisms are related to each other in nested hierarchies based on shared characteristics. the basic structure of all flowers consists of sepals. stigma. Haldane and extended via diffusion theory by Motoo Kimura. those that are smaller and simpler in structure than corresponding parts in the ancestral species. are called vestigial organs. and thus likely to be derived from a common ancestor. haltere (hind wings) of flies and mosquitos. When a group of organisms share a homologous structure which is specialized to perform a variety of functions in order to adapt different environmental conditions and modes of life are called adaptive radiation. They are usually degenerated or underdeveloped. • In cases where the similar structures serve different functions in adults. A large variety of organism’s genomes contain a small fraction of genes that control the organisms development. cactus) and parasitic plants (e. The degree of resemblance between two organisms should indicate how closely related they are in evolution: • Groups with little in common are assumed to have diverged from a common ancestor much earlier in geological history than groups which have a lot in common. • In deciding how closely related two animals are. terrestrial frogs and salamanders passing through the larval stage within the egg—with features of typically aquatic larvae—but hatch ready for life on land. even though they may serve different functions in the adult. “ The most reasonable conclusion to draw is that these creatures descended from creatures in which these parts were functional.[24] the appearance of transitions from fish to amphibians to reptiles and then to mammals in all mammal embryos.g. which in turn indicates that most (or indeed all) creatures descended from common ancenstors. dodder). . and the leaves of some xerophytes (e. Vestigial structures A strong and direct evidence for common descent comes from vestigial structures. Such structures are described as homologous and suggest a common origin. The Challenge of Creation. page 262 Evolutionary developmental biology and embryonic development Evolutionary developmental biology is the biological field that compares the developmental process of different organisms to determine ancestral relationships between species. a comparative anatomist looks for structures that are fundamentally similar. it may be necessary to trace their origin and embryonic development. Those organs are thought to be functional in the ancestral species but are now either nonfunctional or repurposed. Remains of ancestral traits often appear and disappear in different stages of the embryological development process. development and degeneration of a yolk sac. however.Evidence of common descent 188 Homologous structures and divergent (adaptive) evolution If widely separated groups of organisms are originated from a common ancestry. wings of flightless birds such as ostriches.[25] and the appearance of gill-like structures (pharyngeal arch) in vertebrate embryo development including. they are expected to have certain basic features in common. The existence of vestigial organs can be explained in terms of changes in the environment or modes of life of the species.[23] Rudimentary body parts. For example the halteres in dipterists help balance the insect while in flight and the wings of ostriches are used in mating rituals. The gradual spreading of organisms with adaptive radiation is known as divergent evolution. Note that in fish the arches become gills while in humans for example. become the pharynx. Examples are the pelvic girdles of whales. Examples such as hair growth and loss (lanugo) during human development. It must be noted. Hox genes are an example of these types of nearly universal genes in organisms pointing to an origin of common ancestry.g. that vestigial structures may have had their original function replaced with another. ” —Natan Slifkin. Embryological evidence stems from the development of organisms at the embryological level with the comparison of different organisms embryos similarity. A similar developmental origin suggests they are the same structure. [31] reemergence of sexual reproduction in Hieracium pilosella and Crotoniidae.Evidence of common descent 189 Atavisms An atavism is an evolutionary throwback. .[26] extra nipples.[30] chicken's teeth. such as traits reappearing which had disappeared generations ago.[26] Atavisms occur because genes for previously existing phenotypical features are often preserved in DNA.[28] and large canine teeth. The third metacarpal is shaded throughout. the shoulder is crossed-hatched. even though the genes are not expressed in some or most of the organisms possessing them. All conform to the basic pentadactyl pattern but are modified for different usages.[32] and humans with tails.[28] Specific examples Figure 5a: The principle of homology illustrated by the adaptive radiation of the forelimb of mammals.[26] [28] [29] the extra toes of ungulates that do not even reach the ground.[27] Some examples of this are hind-legged snakes or whales. labrum. from amphibians to mammals). 5a). 190 Figure 5c: Adaptation of insect mouthparts: a. indicating that they originated from a common ancestor. antennae. For example: • In the monkey. and the second and fifth digits are reduced. the forelimbs are much elongated to form a grasping hand for climbing and swinging among trees. Throughout the tetrapods.e. mandibles. • In the bat. followed by five series of metacarpals (palm bones) and phalanges (digits). • In the whale. It can even be traced back to the fins of certain fossil fishes from which the first amphibians are thought to have evolved such as tiktaalik. They have become superficially different and unrelated structures to serve different functions in adaptation to different environments and modes of life. This phenomenon is clearly shown in the forelimbs of mammals. md.Evidence of common descent Figure 5b: Illustration of the Eoraptor lunensis pelvis of the saurischian order and the Lesothosaurus diagnosticus pelvis of the ornithischian order in the Dinosauria superorder. • The anteater uses its enlarged third digit for tearing down ant hills and termite nests. a series of carpals (wrist bones). The remaining two digits are longer and stouter than the rest and bear a hoof for supporting the body. spade-like forelimbs for burrowing. . while the hook-like first digit remains free for hanging from trees. • In the horse. the forelimbs have turned into wings for flying by great elongation of four digits. labrium. maxillae. • The mole has a pair of short. lr. these fundamental structures have been modified. The limb has a single proximal bone (humerus). the first digit is lost. the fundamental structures of pentadactyl limbs are the same. The cladogram is shown to illustrate the distance of divergence between the two species. the forelimbs are adapted for support and running by great elongation of the third digit bearing a hoof. The parts of the pelvis show modification over time. Pentadactyl limb The pattern of limb bones called pentadactyl limb is an example of homologous structures (Fig. the forelimbs become flippers for steering and maintaining equilibrium during swimming. mx. lb. c. compound eye. two distal bones (radius and ulna). But in the course of evolution. • In the pig. It is found in all classes of tetrapods (i. Strong mandibles and maxillae for manipulating food. a hypopharynx (floor of mouth).g. which is a cranial nerve. (C) Sucking: e. the earliest dinosaurs split into two distinct orders—the saurischia and ornithischia. Labrum reduced.g. mandibles lost.Evidence of common descent Pelvic structure of dinosaurs Similar to the pentadactyl limb in mammals. it is clear that early saurischians resembled early ornithischians. 191 The path of the recurrent laryngeal nerve in giraffes. despite its optimal route being a distance of just several inches. mandibles form piercing stylets. Parallel developments are seen in some arachnids: The anterior pair of legs may be modified as analogues of antennae. with the duplicates modified in different directions. others are reduced and lost. and a labium. These structures are enlarged and modified. a pair of maxillae. honey bee. They are classified as one or the other in accordance with what the fossils show. passes through the neck down to heart. rounds the dorsal aorta and returns up to the larynx. grasshopper. but for giraffes it becomes quite suboptimal. labrum grooved to hold other parts. butterfly. The modifications enable the insects to exploit a variety of food materials (Fig. and passed near the gill arch. (D) Piercing and sucking. modern birds show a similarity to ancient saurischian pelvic structures therefore indicating the evolution of birds from dinosaurs. Since then gills have evolved and the gill arch has become a dorsal aorta. In mammals its path is extraordinary: as a part of the vagus nerve. Recurrent laryngeal nerve in giraffes The recurrent laryngeal nerve is a fourth branch of the vagus nerve. The laryngeal nerve is compensated for by subsequent tinkering from natural selection. e. Each order of dinosaur has slightly differing pelvis bones providing evidence of common descent. which walk on six legs. which had no neck. This path looks not too optimal even for human. including a labrum (upper lip). particularly in whip scorpions. female mosquito. The pattern of the pelvis in all species of dinosaurs is an example of homologous structures. In Figure 5b. 5c): (A) Primitive state — biting and chewing: e.. (B) Ticking and biting: e. These developments provide support for the theory that complex modifications often arise by duplication of components. a pair of mandibles. Other arthropod appendages Insect mouthparts and antennae are considered homologues of insect legs. the recurrent laryngeal nerve may be up to 4m long (13 ft). Due to the lengths of their necks. Additionally. Insect mouthparts The basic structures are the same. Labrum and maxillae form tube. Labium long to lap up nectar. maxillae long forming sucking tube. mandibles chew pollen and mould wax. Therefore the nerve's route was quite reasonable: it was a relatively short nerve that innervated one gill slit. it comes from the brain.[33] [34] . again through the neck. Its route is explained by the evolution of mammals from fishes.g.g. Evidence of common descent Route of the vas deferens Similar to the laryngeal nerve in giraffes, the vas deferens is part of the male anatomy of many vertebrates; it transports sperm from the epididymis in anticipation of ejaculation. In humans, the vas deferens routes up from the testicle, looping over the ureter, and back down to the urethra and penis. It has been suggested that this is due to the descent of the testicles during the course of human evolution—likely associated with temperature. As the testicles descended, the vas deferens lengthened to accommodate the accidental “hook” over the ureter.[34] [35] 192 Route of the vas deferens from the testis to the penis Evidence from paleontology When organisms die, they often decompose rapidly or are consumed by scavengers, leaving no permanent evidences of their existence. However, occasionally, some organisms are preserved. The remains or traces of organisms from a past geologic age embedded in rocks by natural processes are called fossils. They are extremely important for understanding the evolutionary history of life on Earth, as they provide direct evidence of evolution and detailed information on the ancestry of organisms. Paleontology is the study of past life based on fossil records and their relations to different geologic time periods. For fossilization to take place, the traces and remains of organisms must be quickly buried so that weathering and decomposition do not occur. Skeletal structures or other hard parts of the organisms are the most commonly occurring form of fossilized remains (Paul, 1998), (Behrensmeyer, 1980) and (Martin, 1999). There are also some trace "fossils" showing moulds, cast or imprints of some previous organisms. An insect trapped in amber. As an animal dies, the organic materials gradually decay, such that the bones become porous. If the animal is subsequently buried in mud, mineral salts will infiltrate into the bones and gradually fill up the pores. The bones will harden into stones and be preserved as fossils. This process is known as petrification. If dead animals are covered by wind-blown sand, and if the sand is subsequently turned into mud by heavy rain or floods, the same process of mineral infiltration may occur. Apart from petrification, the dead bodies of organisms may be well preserved in ice, in hardened resin of coniferous trees (amber), in tar, or in anaerobic, acidic peat. Fossilization can sometimes be a trace, an impression of a form. Examples include leaves and footprints, the Evidence of common descent fossils of which are made in layers that then harden. 193 Fossil record It is possible to find out how a particular group of organisms evolved by arranging its fossil records in a chronological sequence. Such a sequence can be determined because fossils are mainly found in sedimentary rock. Sedimentary rock is formed by layers of silt or mud on top of each other; thus, the resulting rock contains a series of horizontal layers, or strata. Each layer contains fossils which are typical for a specific time period during which they were made. The lowest strata contain the oldest rock and the earliest fossils, while the highest strata contain the youngest rock and more recent fossils. A succession of animals and plants can also be seen from fossil records. By studying the number and complexity of different fossils at different stratigraphic levels, it has been shown that older fossil-bearing rocks contain fewer types of fossilized organisms, and they all have a simpler structure, whereas younger rocks contain a greater variety of fossils, often with increasingly complex structures. In the past, geologists could only roughly estimate the ages of various strata and the fossils found. They did so, for instance, by estimating the time for the formation of sedimentary rock layer by layer. Today, by measuring the proportions of radioactive and stable elements in a given rock, the ages of fossils can be more precisely dated by scientists. This technique is known as radiometric dating. Throughout the fossil record, many species that appear at an early stratigraphic level disappear at a later level. This is interpreted in evolutionary terms as indicating the times at which species originated and became extinct. Geographical regions and climatic conditions have varied throughout the Earth's history. Since organisms are adapted to particular environments, the constantly changing conditions favoured species which adapted to new environments through the mechanism of natural selection. Extent of the fossil record Fossil trilobite. Trilobites were hard-shelled arthropods, related to living horseshoe crabs and spiders, that first appeared in significant numbers around 540 mya, dying out 250 mya. Charles Darwin collected fossils in South America, and found fragments of armor which he thought were like giant versions of the scales on the modern armadillos living nearby. The anatomist Richard Owen showed him that the fragments were from gigantic extinct glyptodons, related to the armadillos. This was one of the patterns of distribution that helped Darwin to develop his theory.[36] Evidence of common descent 194 Despite the relative rarity of suitable conditions for fossilization, approximately 250,000 fossil species are known.[37] The number of individual fossils this represents varies greatly from species to species, but many millions of fossils have been recovered: for instance, more than three million fossils from the last Ice Age have been recovered from the La Brea Tar Pits in Los Angeles.[38] Many more fossils are still in the ground, in various geological Cynognathus, a Eucynodont, one of a grouping of Therapsids ("mammal-like reptiles") that is ancestral to formations known to contain a high fossil density, allowing all modern mammals. estimates of the total fossil content of the formation to be made. An example of this occurs in South Africa's Beaufort Formation (part of the Karoo Supergroup, which covers most of South Africa), which is rich in vertebrate fossils, including therapsids (reptile/mammal transitional forms).[39] It has been estimated that this formation contains 800 billion vertebrate fossils.[40] Limitations The fossil record is an important source for scientists when tracing the evolutionary history of organisms. However, because of limitations inherent in the record, there are not fine scales of intermediate forms between related groups of species. This lack of continuous fossils in the record is a major limitation in tracing the descent of biological groups. Furthermore, there are also much larger gaps between major evolutionary lineages. When transitional fossils are found that show intermediate forms in what had previously been a gap in knowledge, they are often popularly referred to as "missing links". There is a gap of about 100 million years between the beginning of the Cambrian period and the end of the Ordovician period. The early Cambrian period was the period from which numerous fossils of sponges, cnidarians (e.g., jellyfish), echinoderms (e.g., eocrinoids), molluscs (e.g., snails) and arthropods (e.g., trilobites) are found. The first animal that possessed the typical features of vertebrates, the Arandaspis, was dated to have existed in the later Ordovician period. Thus few, if any, fossils of an intermediate type between invertebrates and vertebrates have been found, although likely candidates include the Burgess Shale animal, Pikaia gracilens, and its Maotianshan shales relatives, Myllokunmingia, Yunnanozoon, Haikouella lanceolata, and Haikouichthys. Some of the reasons for the incompleteness of fossil records are: • In general, the probability that an organism becomes fossilized is very low; • Some species or groups are less likely to become fossils because they are soft-bodied; • Some species or groups are less likely to become fossils because they live (and die) in conditions that are not favourable for fossilization; • Many fossils have been destroyed through erosion and tectonic movements; • Most fossils are fragmentary; • Some evolutionary change occurs in populations at the limits of a species' ecological range, and as these populations are likely to be small, the probability of fossilization is lower (see punctuated equilibrium); • Similarly, when environmental conditions change, the population of a species is likely to be greatly reduced, such that any evolutionary change induced by these new conditions is less likely to be fossilized; • Most fossils convey information about external form, but little about how the organism functioned; • Using present-day biodiversity as a guide, this suggests that the fossils unearthed represent only a small fraction of the large number of species of organisms that lived in the past. Evidence of common descent 195 Specific examples Evolution of the horse Due to an almost-complete fossil record found in North American sedimentary deposits from the early Eocene to the present, the horse provides one of the best examples of evolutionary history (phylogeny). This evolutionary sequence starts with a small animal called Hyracotherium (commonly referred to as Eohippus) which lived in North America about 54 million years ago, then spread across to Europe and Asia. Fossil remains of Hyracotherium show it to have differed from the modern horse in three important respects: it was a small animal (the size of a fox), lightly built and adapted for running; the limbs were short and slender, and the feet elongated so that the digits were almost vertical, with four digits in the forelimbs and three digits in the hindlimbs; and the incisors were small, the molars having low crowns with rounded cusps covered in enamel. The probable course of development of successive rock strata. The foot diagrams are all front views of the left forefoot. The horses from Hyracotherium to Equus (the third metacarpal is shaded throughout. The teeth are shown in longitudinal section. modern horse) involved at least 12 genera and several hundred species. The major trends seen in the development of the horse to changing environmental conditions may be summarized as follows: • Increase in size (from 0.4 m to 1.5 m — from 15in to 60in); • • • • Lengthening of limbs and feet; Reduction of lateral digits; Increase in length and thickness of the third digit; Increase in width of incisors; Evolution of the horse showing reconstruction of the fossil species obtained from • Replacement of premolars by molars; and • Increases in tooth length, crown height of molars. Fossilized plants found in different strata show that the marshy, wooded country in which Hyracotherium lived became gradually drier. Survival now depended on the head being in an elevated position for gaining a good view of the surrounding countryside, and on a high turn of speed for escape from predators, hence the increase in size and the replacement of the splayed-out foot by the hoofed foot. The drier, harder ground would make the original splayed-out foot unnecessary for support. The changes in the teeth can be explained by assuming that the diet changed from soft vegetation to grass. A dominant genus from each geological period has been selected to show the slow alteration of the horse lineage from its ancestral to its modern form. Evidence of common descent 196 Evidence from geographical distribution Data about the presence or absence of species on various continents and islands (biogeography) can provide evidence of common descent and shed light on patterns of speciation. Continental distribution All organisms are adapted to their environment to a greater or lesser extent. If the abiotic and biotic factors within a habitat are capable of supporting a particular species in one geographic area, then one might assume that the same species would be found in a similar habitat in a similar geographic area, e.g. in Africa and South America. This is not the case. Plant and animal species are discontinuously distributed throughout the world: • Africa has Old World monkeys, apes, elephants, leopards, giraffes, and hornbills. • South America has New World monkeys, cougars, jaguars, sloths, llamas, and toucans. • Deserts in North and South America have native cacti, but deserts in Africa, Asia, and Australia have succulent native euphorbs that resemble cacti but are very different, even though in some cases cacti have done very well (for example in Australian deserts) when introduced by humans.[41] Even greater differences can be found if Australia is taken into consideration, though it occupies the same latitude as much of South America and Africa. Marsupials like kangaroos, bandicoots, and quolls make up about half of Australia's indigenous mammal species.[42] By contrast, marsupials are today totally absent from Africa and form a smaller portion of the mammalian fauna of South America, where opossums, shrew opossums, and the monito del monte occur. The echidnas and platypus, the only living representatives of primitive egg-laying mammals (monotremes), can be found only in Australia and are totally absent in the rest of the world. On the other hand, Australia is missing many groups of placental mammals that are common on other continents (carnivorans, artiodactyls, shrews, squirrels, lagomorphs), although it does have indigenous bats and murine rodents; many other placentals, such as rabbits and foxes, have been introduced there by humans. Other animal distribution examples include bears, located on all continents excluding Africa and Australia, and the polar bear only located nearest to the North Pole. Penguins are located only around the South Pole despite similar weather conditions at the North Pole. Families of sirenians are distributed exclusively around the earth’s waters, where manatees are located in western Africa waters, northern South American waters, and West Indian waters only while the related family, the Dugongs, are located only in Oceanic waters north of Australia, and the coasts surrounding the Indian Ocean. The same kinds of fossils are found from areas known to be adjacent to one another in the past but which, through the process of continental drift, are now in widely divergent geographic locations. For example, fossils of the same types of ancient amphibians, arthropods and ferns are found in South America, Africa, India, Australia and Antarctica, which can be dated to the Paleozoic Era, at which time these regions were united as a single landmass called Gondwana.[43] Sometimes the descendants of these organisms can be identified and show unmistakable similarity to each other, even though they now inhabit very different regions and climates. Evidence of common descent 197 Island biogeography Types of species found on islands Evidence from island biogeography has played an important and historic role in the development of evolutionary biology. For purposes of biogeography, islands are divided into two classes. Continental islands are islands like Great Britain, and Japan that have at one time or another been part of a continent. Oceanic islands, like the Hawaiian islands, the Galapagos islands and St. Helena, on the other hand are islands that have formed in the ocean and never been part of any continent. Oceanic islands have distributions of native plants and animals that are unbalanced in ways that make them distinct from the biotas found on continents or continental islands. Oceanic islands do not have native Four of the 13 finch species found on the Galápagos terrestrial mammals (they do sometimes have bats and seals), Archipelago, are thought to have evolved by an amphibians, or fresh water fish. In some cases they have terrestrial adaptive radiation that diversified their beak shapes to adapt them to different food sources. reptiles (such as the iguanas and giant tortoises of the Galapagos islands) but often (for example Hawaii) they do not. This despite the fact that when species such as rats, goats, pigs, cats, mice, and cane toads, are introduced to such islands by humans they often thrive. Starting with Charles Darwin, many scientists have conducted experiments and made observations that have shown that the types of animals and plants found, and not found, on such islands are consistent with the theory that these islands were colonized accidentally by plants and animals that were able to reach them. Such accidental colonization could occur by air, such as plant seeds carried by migratory birds, or bats and insects being blown out over the sea by the wind, or by floating from a continent or other island by sea, as for example by some kinds of plant seeds like coconuts that can survive immersion in salt water, and reptiles that can survive for extended periods on rafts of vegetation carried to sea by storms.[] Endemism Many of the species found on remote islands are endemic to a particular island or group of islands, meaning they are found nowhere else on earth. Examples of species endemic to islands include many flightless birds of New Zealand, lemurs of Madagascar, the Komodo dragon of Komodo ,[44] the Dragon’s blood tree of Socotra ,[45] Tuatara of New Zealand,[46] [47] and others. However many such endemic species are related to species found on other nearby islands or continents; the relationship of the animals found on the Galapagos Islands to those found in South America is a well-known example.[] All of these facts, the types of plants and animals found on oceanic islands, the large number of endemic species found on oceanic islands, and the relationship of such species to those living on the nearest continents, are most easily explained if the islands were colonized by species from nearby continents that evolved into the endemic species now found there.[48] Other types of endemism do not have to include, in the strict sense, islands. Islands can mean isolated lakes or remote and isolated areas. Examples of these would include the highlands of Ethiopia, Lake Baikal, Fynbos of South Africa, forests of New Caledonia, and others. Examples of endemic organisms living in isolated areas include the Kagu of New Caledonia,[49] cloud rats of the Luzon tropical pine forests of the Philippines,[50] [51] the boojum tree (Fouquieria columnaris) of the Baja California peninsula,[52] the Baikal Seal[53] and the omul of Lake Baikal. Evidence of common descent Adaptive radiations Oceanic islands are frequently inhabited by clusters of closely related species that fill a variety of ecological niches, often niches that are filled by very different species on continents. Such clusters, like the Finches of the Galapagos, Hawaiian honeycreepers, members of the sunflower family on the Juan Fernandez Archipelago and wood weevils on St. Helena are called adaptive radiations because they are best explained by a single species colonizing an island (or group of islands) and then diversifying to fill available ecological niches. Such radiations can be spectacular; 800 species of the fruit fly family Drosophila, nearly half the world's total, are endemic to the Hawaiian islands. Another illustrative example from Hawaii is the Silversword alliance, which is a group of thirty species found only on those islands. Members range from the Silverswords that flower spectacularly on high volcanic slopes to trees, shrubs, vines and mats that occur at various elevations from mountain top to sea level, and in Hawaiian habitats that vary from deserts to rainforests. Their closest relatives outside Hawaii, based on molecular studies, are tarweeds found on the west coast of North America. Interestingly, these tarweeds have sticky seeds that facilitate distribution by migrant birds. Continental islands have less distinct biota, but those that have been long separated from any continent also have endemic species and adaptive radiations, such as the 75 lemur species of Madagascar, and the eleven extinct moa species of New Zealand.[] [54] 198 Ring Species In biology, a ring species is a connected series of neighboring populations that can interbreed with relatively closely related populations, but for which there exist at least two "end" populations in the series that are too distantly related to interbreed. Often such non-breeding-though-genetically-connected populations co-exist in the same region thus creating a "ring". Ring species provide important evidence of evolution in that they illustrate what happens over time as populations genetically diverge, and are special because they represent in living populations what normally happens over time between long deceased ancestor populations and living populations. If any of the populations intermediate between the two ends of the ring were gone they would not be a continuous line of reproduction and each side would be a different species.[55] [56] Specific examples Figure 6a: A dymaxion map of the world showing the distribution of present species of camelid. The solid black lines indicate migration routes and the blue represents current camel locations. Evidence of common descent 199 Figure 6b: Current distribution of Glossopteris placed on a Permian map showing the connection of the continents. (1, South America; 2, Africa; 3, Madagascar; 4, India; 5, Antarctica; and 6, Australia) Figure 6c: Present day distribution of marsupials. (Distribution shown in blue. Introduced areas shown in green.) Migration, isolation, and distribution of the Camel The history of the camel provides an example of how fossil evidence can be used to reconstruct migration and subsequent evolution. The fossil record indicates that the evolution of camelids started in North America (see figure 6a), from which 6 million years ago they migrated across the Bering Strait into Asia and then to Africa, and 3.5 million years ago through the Isthmus of Panama into South America. Once isolated, they evolved along their own lines, giving rise to the Bactrian camel and Dromedary in Asia and Africa and the llama and its relatives in South America. Camelids then went extinct in North America at the end of the last ice age.[57] Distribution of Glossopteris The combination of continental drift and evolution can sometimes be used to make predictions about what will be found in the fossil record. Glossopteris is an extinct species of seed fern plants from the Permian. Glossopteris appears in the fossil record around the beginning of the Permian on the ancient continent of Gondwana. Continental drift explains the current biogeography of the tree. Present day Glossopteris fossils are found in Permian strata in southeast South America, southeast Africa, all of Madagascar, northern India, all of Australia, all of New Zealand, and scattered on the southern and northern edges of Antarctica. During the Permian, these continents were connected as Gondwana (see figure 6b) in agreement with magnetic striping, other fossil distributions, and glacial scratches pointing away from the temperate climate of the South Pole during the Permian.[48] Evidence of common descent Distribution of marsupials The history of marsupials also provides an example of how the theories of evolution and continental drift can be combined to make predictions about what will be found in the fossil record. The earliest marsupial fossils are about 80 million years old and found in North America; by 40 million years ago fossils show that they could be found throughout South America, but there is no evidence of them in Australia, where they now predominate, until about 30 million years ago. The theory of evolution predicts that the Australian marsupials must be descended from the older ones found in the Americas. The theory of continental drift says that between 30 and 40 million years ago South America and Australia were still part of the Southern hemisphere super continent of Gondwana and that they were connected by land that is now part of Antarctica. Therefore combining the two theories scientists predicted that marsupials migrated from what is now South America across what is now Antarctica to what is now Australia between 40 and 30 million years ago. This hypothesis led paleontologists to Antarctica to look for marsupial fossils of the appropriate age. After years of searching they found, starting in 1982, fossils on Seymour Island off the coast of the Antarctic Peninsula of more than a dozen marsupial species that lived 35-40 million years ago.[41] 200 Evidence from observed natural selection Examples for the evidence for evolution often stems from direct observation of natural selection in the field and the laboratory. Scientists have observed and documented a multitude of events where natural selection is in action. The most well known examples are antibiotic resistance in the medical field along with better-known laboratory experiments documenting evolution's occurrence. Natural selection is tantamount to common descent in the fact that long-term occurrence and selection pressures can lead to the diversity of life on earth as found today. All adaptations—documented and undocumented changes concerned—are caused by natural selection and genetic drift. The examples below are only a small fraction of the actual experiments and observations. Specific examples of natural selection in the lab and in the field Antibiotic and pesticide resistance The development and spread of antibiotic resistant bacteria, like the spread of pesticide resistant forms of plants and insects is evidence for evolution of species, and of change within species. Thus the appearance of vancomycin resistant Staphylococcus aureus, and the danger it poses to hospital patients is a direct result of evolution through natural selection. The rise of Shigella strains resistant to the synthetic antibiotic class of sulfonamides also demonstrates the generation of new information as an evolutionary process.[58] Similarly, the appearance of DDT resistance in various forms of Anopheles mosquitoes, and the appearance of myxomatosis resistance in breeding rabbit populations in Australia, are all evidence of the existence of evolution in situations of evolutionary selection pressure in species in which generations occur rapidly. E. coli long-term evolution experiment Experimental evolution uses controlled experiments to test hypotheses and theories of evolution. In one early example, William Dallinger set up an experiment shortly before 1880, subjecting microbes to heat with the aim of forcing adaptive changes. His experiment ran for around seven years, and his published results were acclaimed, but he did not resume the experiment after the apparatus failed.[59] The E. coli long-term evolution experiment begun in 1988 under the leadership of Richard Lenski is still in progress, and has shown adaptations including the evolution of a strain of E. coli that was able to grow on citric acid in the growth media. with recent findings demonstrating the population which is at risk of the severe debilitating disease kuru has significant over-representation of an immune variant of the prion protein gene G127V versus non-immune alleles. The color of the moth has gone from light to dark to light again over the course of a few hundred years due to the appearance and later disappearance of pollution from the Industrial Revolution in England.[63] However. This is seen as a good example of evolution through mutation and natural selection that has been observed as it occurs. seeds that float far will often land on infertile concrete. the weed Crepis sancta. Wangiella dermatitidis. in non-dairy consuming societies. and a variety of insects . lactase production usually drops about 90% during the first four years of life. baboons in South Africa.[69] Research at the Albert Einstein College of Medicine showed that three melanin-containing fungi. increased in biomass and accumulated acetate faster in an environment in which the radiation level was 500 times higher than in the normal environment. although the exact drop over time varies widely. Radiotrophic fungus Recently evolved Radiotrophic fungi are fungi which appear to use the pigment melanin to convert gamma radiation into chemical energy for growth[69] [70] and were first discovered in 2007 as black molds growing inside and around the Chernobyl Nuclear Power Plant. The normal mammalian condition is for the young of a species to experience reduced lactase production at the end of the weaning period (a species-specific length of time). whereas the fluffy seeds float further away on the wind. certain human populations have a mutation on chromosome 2 which eliminates the shutdown in lactase production. heavy and fluffy.[65] [66] [67] [68] 201 Peppered moth One classic example of adaptation in response to selection pressure is the case of the peppered moth. has two types of seed. This appears to be an evolutionarily recent adaptation to dairy consumption.[71] [72] Other examples of urban wildlife are rock pigeons and species of crows adapting to city environments around the world. These types of environments can exert selection pressures on organism. found in France. African penguins in Simons Town. In urban environments. There is scientific consensus that the capacity to synthesize nylonase most probably developed as a single-step mutation that survived because it improved the fitness of the bacteria possessing the mutation. and has occurred independently in both northern Europe and east Africa in populations with a historically pastoral lifestyle. Cladosporium sphaerospermum.[60] [61] Other reported evolutionary trends in other populations include a lengthening of the reproductive period. reduction in cholesterol levels. and Cryptococcus neoformans. Scientists postulate one of the reasons for the rapid selection of this genetic variant is the lethality of the disease in non-immune persons. blood glucose and blood pressure.[62] Lactose intolerance in humans Lactose intolerance is the inability to metabolize lactose. often leading to new adaptations. making it possible for members of these populations to continue consumption of fresh milk and other dairy products throughout their lives without difficulty. Within about 5-12 generations. For example. In humans. the weed has been found to evolve to produce significantly more heavy seeds than its rural relatives do.[64] Nylon-eating bacteria Nylon-eating bacteria are a strain of Flavobacterium that is capable of digesting certain byproducts of nylon 6 manufacture.Evidence of common descent Humans Natural selection is being observed in contemporary human populations. Urban wildlife Urban wildlife is wildlife that is able to live or thrive in urban environments. because of a lack of the required enzyme lactase in the digestive system. The heavy ones land nearby to the parent plant. The smaller blackcap population only recently rerouted to Spain. Sylvia atricapillab. reproductive isolation was produced. After 25 generations of this mating test. Scientists have observed numerous examples of speciation in the laboratory and in nature. The examples shown below provide strong evidence for common descent and are only a small fraction of the instances observed. Stuart Bearhop from the University of Exeter reported that birds wintering in England tend to mate only among themselves. The flies had three choices to take in finding food.Evidence of common descent living in human habitations. (Bearhop et al. etc). Pupae from the two strains were then placed together in the maze and allowed to mate at the food site. lives in Germany and flies southwest to Spain while a smaller group flies northwest to Great Britain during the winter. They collected a population of Drosophila melanogaster from Davis. attracted to light and ethanol. California and placed the pupae into a habitat maze. sympatric. The differences found have arisen in about 30 generations. Specific examples Blackcap The bird species. allopatric. evolution has produced far more species than an observer would consider necessary. both explained in an earlier section. Newborn flies had to investigate the maze to find food.[73] Great examples of observed speciation come from the observations of island biogeography and the process of adaptive radiation. This entailed that none of their gametes would pass on to the next generation. The other strain emerged late and immediately flew downward.[75] [76] [77] . A selective penalty was imposed on the female flies that switched habitats. One of the strains emerged early. Gregor Rolshausen from the University of Freiburg found that the genetic separation of the two populations is already in progress. Rice and George W. 2005) It is still inference to say that the populations will become two different species. immediately flying upward in the dark attracted to the acetaldehyde.g. They then cultured two strains that chose opposite habitats.000 described species of beetles. A common fruit fly (Drosophila melanogaster). however. They then were collected. This eventually divided the flies into 42 spatio-temporal habitats. there are well over 350. commonly referred to as Blackcaps. polyploidization. and the scent of acetaldehyde and the scent of ethanol (chemotaxis) were the three options. Speciation can occur from a variety of different causes and are classified in various forms (e. up and down (geotaxis). Light and dark (phototaxis). With DNA sequencing. and not usually with those wintering in the Mediterranean.[74] Drosophila melanogaster William R. 202 Evidence from observed speciation Speciation is the evolutionary process by which new biological species arise. For example. Salt found experimental evidence of sympatric speciation in the common fruit fly. but experts deduce that it is expected due to the continued genetic and geographic separation. They repeated the experiment again without creating the penalty against habitat switching and the result was the same. it showed reproductive isolation between the two strains. the individuals can be assigned to a correct group with an 85% accuracy. placed them in large plastic bottles. were introduced. however.e. and bites rats. Japan. This mosquito.[79] London Underground mosquito The London Underground mosquito is a species of mosquito in the genus Culex found in the London Underground. Michael Tobler from the Texas A&M University has studied the fish for years and found that two distinct populations of mollies—the dark interior fish and the bright surface water fish—are becoming more genetically divergent. consistent with genetic drift during a founder event. A distinct population emerged in North America in the 19th century some time after apples. rather than the population at each station being related to the nearest above-ground population (i. the cave-adapted fish endured the most damage. the situation was the opposite. Culex pipiens molestus. Tobler plans on further 203 . These widely separated populations are distinguished by very minor genetic differences. and bites only birds. and that there is little evidence of interbreeding (researchers have documented a 4-6% hybridization rate) suggests that this is occurring.[82] are extremely difficult to mate. has been found in underground systems around the world. although first discovered in the London Underground system.[83] More specifically. The current hawthorn feeding population does not normally feed on apples.Evidence of common descent Hawthorn fly One example of evolution at work is the case of the hawthorn fly. with four out of every five stab-wounds from the water bugs sharp mouthparts. After a day. a non-native species. breeds all-year round.[78] Different populations of hawthorn fly feed on different fruits. in contrast to the above ground species Culex pipiens that is cold tolerant. but not in the other ones. (Tobler et al. When the two varieties were cross-bred the eggs were infertile suggesting reproductive isolation. and humans. in the light. The species have very different behaviours. which appears to be undergoing sympatric speciation. is cold intolerant. that hawthorn flies mature later in the season and take longer to mature than apple flies. such as the fact that six out of thirteen allozyme loci are different. The mollies’ senses can detect a predator’s threat in their own habitats. Byrne and Nichols' working hypothesis was that adaptation to the underground environment had occurred locally in London once only.[84] a single fixed microsatellite difference in populations spanning Europe. also known as the apple maggot fly. the middle East and Atlantic islands. Australia. the pipiens form). it was found that. this mosquito.[81] The evidence for this mosquito being a different species from Culex pipiens comes from research by Kate Byne and Richard Nichols. In the dark. which suggest that the molestus form developed recently: a single mtDNA difference shared among the underground populations of ten Russian cities. Rhagoletis pomonella. mice.[81] Madeira House Mouse The Madeira House mouse is a new species of mouse descended from the house mouse (Mus Musculus) that went through a recent speciation event.[85] Mollies The Shortfin Molly—Poecilia Mexicana—is a small fish that lives in the Sulfur Caves of Mexico. It is thought to have evolved from the overground species Culex pipiens recently. and put them back in the cave. Some evidence. it was found that the mollies are hunted by a large water bug (Belostoma spp). hibernates in the winter.[80] although more recent evidence suggests that it is a southern mosquito variety related to Culex pipiens that has adapted to the warm underground spaces of northern cities. It is suggested that it may have adapted to human-made underground systems since the last century from local above-ground Culex pipiens.[80] and with different allele frequency. 2009) The populations have no obvious barrier separating the two. Tobler collected the bug and both types of mollies. Moving from one habitat to the other significantly increases the risk of dying. This apple-feeding population normally feeds only on apples and not on the historically preferred fruit of hawthorns.[80] [82] The fundamental results still stands: the genetic data indicate that the molestus form in the London Underground mosquito appeared to have a common ancestry. blend in—the change in fur color—and to stay warm in the arctic environment. feet covered with heavy matting to protect the bottoms from intense cold and provide traction. Arabidopsis thaliana. mouse-ear cress or Arabidopsis). and the ability to fast for up to nine months while recycling their urea. 2007) To test this. Bomblies discovered that only two genes were required to cause the autoimmune response. When the genes are passed down. but they evolved separately to react defectively when combined. smaller ears to reduce the loss of heat. eyelids that act like sunglasses. reducing their chances of successful mating and turning distinct strains into separate species. the hybrids developed normally. a large stomach capacity to enable opportunistic feeding.[86] Thale cress Kirsten Bomblies et al. (Bomlies et al. the 20 plants also shared a comparable collection of genetic activity in a group of 1. Bomblies crossed 280 genetically different strains of Arabidopsis in 861 distinct ways and found that 2 per cent of the resulting hybrids were necrotic. In conjunction with this. these incompatibilities could create divisions between different plant strains.Evidence of common descent experiments. Along with allocating the same indicators. but it is believed that it is a good example of the rise of a new species. (Bomlies et al. Bomblies looked at one hybrid in detail and found that one of the two genes belonged to the NB-LRR class. In the parents.080 genes. In almost all of the cases. Additionally. which though clearly related to the brown bear (Ursus arctos) by virtue of the fact that though separate species they can still interbreed and produce fertile offspring[88] it has also obviously acquired significant physiological differences from the brown bear. 2007) 204 Arabidopsis thaliana (colloquially known as thale cress. finding species that are able reproduce successfully or create hybrids between two different species infers that their relationship is close. a common group of disease resistance genes involved in recognizing new infections.[89] [90] . The examples provided are only a small fraction of the observed instances of speciation through hybridization. It is worth noting that plants are often subject to the creation of a new species though hybridization. the elongation of the neck makes it easier to keep their heads above water while swimming and the oversized webbed feet that act as paddles when swimming. to accommodate their all-meat diet. These differences allow the polar bear to comfortably survive in conditions that the brown could not including the ability to swim sixty miles or more at a time in freezing waters. from the Max Planck Institute for Developmental Biology discovered that two genes passed down by each parent of the thale cress plant. Over successive generations. hybridization has been found to be a precursor to the creation of new species by being a source of new genes for a species. Polar bear A specific example of large-scale evolution is the polar bear (Ursus maritimus). When Bomblies removed the problematic gene. it ignites a reaction in the hybrid plant that turns its own immune system against it.[87] Interspecies fertility or hybridization Understood from laboratory studies and observed instances of speciation in nature. the genes were not detrimental. The polar bear has also evolved small papillae and vacuole-like suction cups on the soles to make them less likely to slip on the ice. Two other fertile forms of Raphanobrassica are known. Raparadish. such as S. S. Like S. x baxteri. Tragopogon porrifolius 205 Welsh groundsel is an allopolyploid. pratensis). Plants of this parentage are now known as radicole. Other fertile hybrids are also known. squalidus was introduced from Sicily in the early 18th century. Tragopogon mirus. indicating that strong breed barriers exist between this new hybrid and its parents. eboracensis is self-compatible. is a tetraploid hybrid of T. The Raphanobrassica is a fascinating plant. and the oyster plant (T. etc). mirus population. it shows little or no natural crossing with its parent species. vulgaris is native to Britain. while S. Welsh groundsel Purple Salsify. and the allohexaploid S. dubius and T. This has led some botanists to propose that the accidental hybridization of a flower by pollen of another species in nature could be a mechanism of speciation common in higher plants. S. These species. is also an allopolyploid. hibernicus. are now common weeds in urban wastelands. Tragopogon miscellus. the meadow salsify (T. It is thought to have resulted from a backcrossing of the F1 hybrid of its parents to S. vulgaris. the western salsify (T. "Raphanofortii" is the allopolyploid hybrid between Brassica tournefortii and Raphanus caudatus. The T.Evidence of common descent Raphanobrassica Raphanobrassica includes all intergeneric hybrids between the genera Raphanus (radish) and Brassica (cabbages. vulgaris var. cambrensis. One new species. which according to molecular evidence probably originated independently at least three times in . × baxteri plant led to the formation of a new fertile species. mirus population grows mainly by reproduction of its own members. Other hybrids descended from the same two parents are known. dubius and T. pratensis. porrifolius. now common in Britain. where the three already known species overlapped. S. a plant which contains sets of chromosomes originating from two different species. Its ancestor was Senecio × baxteri. and is therefore reproductively isolated. The other species. an accidental doubling of the number of chromosomes in an S. but its ancestors were T. however. In the early 20th century. but additional episodes of hybridization continue to add to the T. dubius). These new species are usually referred to as "the Ownbey hybrids" after the botanist who first described them. eboracensis has speciated from those two species within the last 300 years. In the 1950s. humans introduced three species of goatsbeard into North America. vulgaris. therefore. including S. because—in spite of its hybrid nature—it is not sterile. botanists found two new species in the regions of Idaho and Washington.[91] [92] The Raphanobrassica is an allopolyploid cross between the radish (Raphanus sativus) and cabbage (Brassica oleracea). Some are infertile. Salsify Salsifies are one example where hybrid speciation has been observed. an allopolyploid hybrid between Raphanus sativus and Brassica rapa is grown as a fodder crop. porrifolius). an infertile hybrid which can arise spontaneously when the closely-related groundsel (Senecio vulgaris) and Oxford ragwort (Senecio squalidus) grow alongside each other. Sometime in the early 20th century.[93] [94] York groundsel The York groundsel (Senecio eboracensis) is a hybrid species of the self-incompatible Senecio squalidus (also known as Oxford ragwort) and the self-compatible Senecio vulgaris (also known as Common groundsel). [98] For example. David J. Christoph Adami et al. flower breeding. genomic complexity is forced to increase.[95] 206 Evidence from artificial selection Artificial selection demonstrates the diversity that can exist among organisms that share a relatively recent common ancestor. but the electronic computer is now powerful enough to simulate evolution"[97] assisting bioinformatics in its attempt to solve biological problems.[99] . The evolution of specific cellular mechanisms like spliceosomes that can turn the cell's genome into a vast workshop of billions of interchangeable parts that can create tools that create tools that create tools that create us can be studied for the first time in an exact way. are explained by selection for evolvability. complexity needs to be both rigorously defined and measurable." within a fixed environment. For example. These characteristics become increasingly well-developed in successive generations. Deem make this point in Evolvability is a selectable trait: Not only has life evolved. We show that. eboracensis as separate from other known hybrids.[96] and others. Many observations within evolutionary biology. genetically modified food. "It has taken more than five decades. This selection is not forbidden by causality and is strongest on the largest-scale moves within the mutational hierarchy. heretofore considered evolutionary happenstance or accidents. That is. but life has evolved to evolve. Sensibly. correlations within protein structure have evolved. allowing only those organisms that exhibit desired characteristics to reproduce. Morphological and genetic evidence support the status of S. A recent information-theoretic (but intuitively evident) definition identifies genomic complexity with the amount of information a sequence stores about its environment. A similar driving force for biased codon usage as a result of productively high mutation rates is observed in the hemagglutinin protein of influenza A. Examples of artificial selection would be dog breeding. Artificial selection was successful long before science discovered the genetic basis. rapid or extreme environmental change leads to selection for greater evolvability. The rates at which the various events within the hierarchy of evolutionary moves occur are not random or arbitrary but are selected by Darwinian evolution. and mechanisms to manipulate these correlations have evolved in tandem. We investigate the evolution of genomic complexity in populations of digital organisms and monitor in detail the evolutionary transitions that increase complexity. Earl and Michael W. In artificial selection. Evidence from computation and mathematical iteration The Chihuahua mix and Great Dane illustrate the range of sizes among dog breeds. the vertebrate immune system shows that the variable environment of antigens has provided selective pressure for the use of adaptable codons and low-fidelity polymerases during somatic hypermutation. For example. one species is bred selectively at each generation. providing evidence for the hidden causes of known evolutionary events. cultivation of foods such as wild cabbage.Evidence of common descent different locations. allowing a mathematical understanding of the nature of the processes behind evolution. because natural selection forces genomes to behave as a natural "Maxwell Demon. Computer science allows the iteration of self changing complex systems to be studied. make this point in Evolution of biological complexity: To make a case for or against a trend in the evolution of complexity in biological evolution. J Hered. [5] The New York Times report Still Evolving. "Pseudogene evolution and natural selection for a compact genome" (http:/ / jhered. Warthmann. and selection of the fittest of individual molecules (proteins. pdf [3] Two sources: Chen FC.1115661 (HTML abstract) Supporting Online Material [104] • Bomblies. Jason. PMC 1235277. DNA. PMID 11170892.1086/318206.0050236 • Davis. recombination. html) is literally a picture of a link in humans that links two separate chromosomes in the nonhuman apes creating a single chromosome in humans. Am J Hum Genet. Gabriel J. ch/ research/ noll/ publ/ Dev_Cell_2003_5_773_785. [7] http:/ / www. (2004).1064503.pbio. this too is a trace from the past corresponding to some living beings which when alive were the physical embodiment of this link. Smithsonian Books (in association with the Natural History Museum of London). [4] The picture labeled "Human Chromosome 2 and its analogs in the apes" in the article Comparison of the Human and Great Ape Chromosomes as Evidence for Common Ancestry (http:/ / www. nov. also called directed evolution or in vitro molecular evolution involves the iterated cycle of mutation..011.C."[100] Evolutionary molecular engineering. Brudno M.Evidence of common descent "Computer simulations of the evolution of linear sequences have demonstrated the importance of recombination of blocks of sequence rather than point mutagenesis alone. Sidow A (May 2003). Fossil Plants. imls. Also.2006. PMC 430923. . fcgi?tool=pmcentrez& artid=1235277). Paul and Kenrick. . Repeated cycles of point mutagenesis.3. Inouye I (October 2006). while the term originally referred to fossil evidence. based on A Map of Recent Positive Selection in the Human Genome (http:/ / biology.[100] [101] [102] [103] 207 See also • Nothing in Biology Makes Sense Except in the Light of Evolution Sources • Bearhop. PLoS Biol 5: e236 doi:10.1126/science. and selection should allow in vitro molecular evolution of complex sequences. D.. NY. doi:10. Paul. doi:10. 2007. doi:10. Li WH (February 2001). Green ED.1098/rsbl. 68 (2): 444–56. PMID 16891155.1371/journal. Votier. Science 310 (5746): 287. Lanz. "Quantitative estimates of sequence divergence for comparative analyses of mammalian genomes" (http:/ / www. a katablepharid undergoing probable plastid acquisition". et sp. Waldron.221. Berthold. and the ape-human connection is of particular interest. 0040072).protis. Washington. Does a predatory insect contribute to the divergence between cave. Micheal (2009). "Hatena arenicola gen. Bowen. Epple.. 13 (5): 813–20. Genome Res. Natural evolution can be relived showing us possible paths from catalytic cycles based on proteins to based on RNA to based on DNA. Hartl DL (2000).0272 References [1] Mount DM. . [6] Petrov DA.1101/gr.1016/j. doi:10. Wolfgang. Robert W. genomesize. PMID 16224014.). Newton. Stuart. Protist 157 (4): 401–19. and RNA). 1371/ journal. nytimes. "Genomic divergences between humans and other hominoids and the effective population size of the common ancestor of humans and chimpanzees" (http:/ / www. Batzoglou S. ISBN 0-87969-608-7. Human Genes Tell New Story (http:/ / www.. doi:10. Bioinformatics: Sequence and Genome Analysis (2nd ed. genome. doi:10. "A secondary symbiosis in progress". Cold Spring Harbor Laboratory Press: Cold Spring Harbor. Susan. pubmedcentral. pbio. plosjournals. Inouye I (2005). 2004. Keith (2005): Assortative mating as a mechanism for rapid evolution of a migratory divide. states the International HapMap Project is "providing the strongest evidence yet that humans are still evolving" and details some of that evidence. net/ ~rwms/ hum_ape_chrom. gate. Autoimmune response as a mechanism for a Dobhansky-Muller-Type incompatibility syndrome in plants. It is considered a missing link.05.2009. Science 310(5747): 502-504. Cooper GM.. 91 (3): 221–7. [2] http:/ / www. nih. com/ [8] Okamoto N. Lempe. PMID 12727901.1093/jhered/91. uzh. gov/ articlerender. [9] Okamoto N. ISBN 1-58834-156-9 • Tobler. Peter & Farnsworth. html). org/ perlserv/ ?request=get-document& doi=10. Fiedler. Stephen C.1116125.and surface-adapted fish populations? Biology Letters doi:10. PMID 10833048. org/ cgi/ pmidlookup?view=long& pmid=10833048).1126/science. Dangl & Weigel. Furness. com/ 2006/ 03/ 07/ science/ 07evolve. oxfordjournals. org/ cgi/ content/ full/ 13/ 5/ 813). multiplication with recombination. such as proteins. Richard (2009). 208 . pp.1007/BF00217134. . edu/ ~pgore/ geology/ historical_lab/ preservationlab. [22] Charles Darwin (1859). Why Evolution is True. Retrieved 2009-03-08 [32] Domes. "Mutant Chicken Grows Alligatorlike Teeth" (http:/ / www. A. "29+ Evidences for Macroevolution: Part 2" (http:/ / www. Rough Guides. [16] Amino acid sequences in cytochrome c proteins from different species (http:/ / www. Retrieved 2006-11-10. Reeders. "Evidence for an ancestral alphoid domain on the long arm of human chromosome 2". Incorporated. pp. com/ content/ mr73x80w2x7744jp/ fulltext. vqronline. gpc. [27] http:/ / animals. htm). Roy A. pp.. John P. (1992). [43] http:/ / biology. org/ content/ 104/ 17/ 7139). ca/ I10-10-snake. Patrick T. Science and Earth History. [48] Coyne. ISBN 978-0-670-02053-9. Blackwell Publishing. [11] Evidence of Common Ancestry: Human Chromosome 2 (http:/ / www. pdf). (July 2004). org/ faqs/ comdesc/ section4.. Viking. terranature. 91–99. Baldini. ISBN 0-19-550870-X. Mark. Berry. htm [44] Trooper Walsh. [45] Burdick.. socotra. htm). Retrieved 2009-04-08 [33] Mark Ridley (2004). [18] 29+ Evidences for Macroevolution. springerlink..: Smithsonian Books. Schadewald. Caiulo. fcgi?tool=pmcentrez& artid=52649). PMID 1587535. ISBN 0-87893-189-9. pp. 79. Jerry A. t. Human Genetics 89 (2): 247–9. "Reevolution of sexuality breaks Dollo's law" (http:/ / www.C. O. Niles Eldredge. M (1992).). html [30] Tyson. PNAS 104 (17): 7139–7144. "Origin of human chromosome 2: an ancestral telomere-telomere fusion" (http:/ / www. page 348. talkorigins. A. ISBN 1405103450. . (2009). pdf [25] Douglas J. ISBN 1-4051-5089-0. ALAN BURDICK. org/ education/ guide/ index. (2009). [42] Menkhorst.Evidence of common descent [10] Human Chromosome 2 is a fusion of two ancestral chromosomes (http:/ / www. us/ whales/ atavisms. cfm?id=mutant-chicken-grows-alli). Maraun. Clifford R. com/ ?id=b-HGB9PqXCUC& lpg=RA1-PA281& pg=RA1-PA282#v=onepage& q=) (3rd ed. clc. gate. Katja. [19] Pallen.. [14] Avarello et al. . edu/ ~ensiweb/ lessons/ molb.. pp. Bantam Press. nz/ templates/ page. Reid. Mark (2009). html#nested_hierarchy [21] Coyne. . doi:10. 122. PMC 1855408. com/ Paleozoic/ Permian/ Karroo. [17] Lurquin PF.D. Zoo Torah. Department of Conservation. htm) [40] Six Flood Arguments Creationists Can't Answer (http:/ / www. Knight.20. 420. The Challenge of Creation. Ward. Proceedings of the National Academy of Sciences 88 (20): 9051–5. net/ ~rwms/ hum_ape_chrom. Oxford Uniersity Press.88. PMID 17438282. pubmedcentral. 200–206. PMC 52649. Peter. Natural selection: domains. . [13] Human and Ape Chromosomes (http:/ / www. pp. org/ faqs/ comdesc/ section2.7063861. by Pamela J. "The origin of man: a chromosomal pictorial legacy". [47] "Facts about tuatara" (http:/ / www. Oxford Press. html#atavisms). jpg [29] http:/ / www. Yemen" (http:/ / travel. Alan (2007-03-25).1073/pnas. [46] "Tuatara" (http:/ / www. Genes. Norton. O (1982). ISBN 0-19-506932-3.. Colahan. uc. . A. RA (1991). Ph. DC. 258–9. p. Arthur. htm [28] http:/ / universe-review.). palaeos.. Jerry A. doi:10. talkorigins. and Human Evolution: A Synthesis (http:/ / books. Frank (2001). 1997. Scheu. org/ tuatara. D. 364–365. com/ 2007/ 03/ 25/ travel/ tmagazine/ 03well. Graham. adapted from Strahler. Culture. W. nih. org/ articles/ 2006/ spring/ eldredge-confessions-darwinist/ ). Futuyma (1998). pnas. ST.9051. ISBN 1-58834-073-2.. lhup. gov/ articlerender. accessed 18 May 2006. Washington. doc. Zuffardi. accessed 8 September 2007. p. Scientific American. html). sciam. PMID 1924367. Conservation: Native Species. 8–11. [24] http:/ / www. Douglas Theobald. Jerry A.Fossil Preservation (http:/ / facstaff. html) [39] (http:/ / www. 14. Threatened Species Unit. Science 215 (4539): 1525–1530. Colomba De LA Panouse (2002). Georgia Perimeter College [38] (http:/ / www. 2004. James Jerome. Why Evolution is True. G. ISBN 978-0-670-02053-9.. html). com/ article. indiana. Stone L (2006). (2009). Retrieved 2010-07-08.1073/pnas. Stefan (2007-04-24). Retrieved 2007-02-10. [15] IJdo et al. Gore. google. and challenges. evolutionpages. Wells. pp. Protein functional redundancy (http:/ / www. [20] http:/ / www. Murphy.1126/science. com/ ?id=zdeWdF_NQhEC& pg=PA79& lpg=PA79& dq=chimpanzee+ rhesus+ cytochrome+ c). edwardtbabinski. com/ animal-facts/ atavism. aspx?id=33163). Fraccaro. On the Origin of Species. com/ chromosome_2. html#protein_redundancy). [23] Natan Slifkin (2006). ISBN 978-1-85828-946-5. Claudio Ciofi. John Murray. [26] TalkOrigins Archive. [35] Williams. "The Wonder Land of Socotra. Evolution (http:/ / books. . A Field Guide to the Mammals of Australia. David (2006-02-22). ws. Rough Guide to Evolution. by Robert J. edu/ ~dsimanek/ 6flood. edu/ courses/ bio303/ contdrift. google. New Zealand Ecology: Living Fossils. ISBN 978-0-670-02053-9. The Greatest Show on Earth: The Evidence for Evolution. levels. htm) by Alec MacAndrew. Pedicini. 1982 [41] Coyne. ISBN 978-1-4165-9478-9. [37] Laboratory 11 . Sinauer Associates Inc. tarpits. govt. nytimes. Blackwell Publishing. Viking. 99–110. org/ faqs/ comdesc/ section1. howstuffworks. pp. com/ watch?v=x-WAHpC0Ah0) (video) 2007 [12] Yunis and Prakash. TerraNature Trust. ISBN 1-933143-15-0. Government of New Zealand. Why Evolution is True.0700034104.C. PMID 7063861. Retrieved 2006-11-08. php). talkorigins. Evolutionary Biology (3rd ed. Veterinary Radiology & Ultrasound 45 (4): 315–317 [31] Biello. doi:10. . Komodo Dragons: Biology and Conservation (Zoo and Aquarium Biology and Conservation Series). Viking. pp. Retrieved 2010-06-22. Prakash. "Skeletal Atavism in a Miniature Horse". . [36] "Confessions of a Darwinist" (http:/ / www. doi:10. [34] Dawkins. 282. youtube. 23. archive. 45.: Early Advocate of Theistic Evolution and Foe of Spontaneous Generation" (http:/ / www. [71] Cheptou. abstract). Salt (1990).1073/pnas. uk/ doc_WTX038968. "Drug-resistance and its transferability of Shigella strains isolated in 1986 in Japan". aspx) Wayback: Soy Nutrition [64] Coles Harriet (2007-01-20). html).Evidence of common descent [49] "New Caledonia's most wanted" (http:/ / www. msn. PMID 17520016. com/ releases/ 2009/ 11/ 091120091959. [59] J.. James (27 July 2001). Ker Than. 107–117. "The Evolution of Reproductive Isolation as a Correlated Character Under Sympatric Conditions: Experimental Evidence". . "Evolution in Health and Medicine Sackler Colloquium: Natural selection in a contemporary human population" (http:/ / www. Wellcome Trust. Horns. . T.. H. "Ionizing radiation changes the electronic properties of melanin and enhances the growth of melanized fungi" (http:/ / www. Schweitzer AD. 2009)). S. talkorigins. (2009). PMID 19858476. asp [56] http:/ / evolution. 0000457). PMC 2868295. 5. H. Creation Evolution Journal (National Center for Science Education (NCSE)) 5 (2): 1–3. htm). ucsb.3. Retrieved 2010-05-23. [63] Soy and Lactose Intolerance (http:/ / web. [61] Mead. asp). html). Vol. pp. et al. com/ stories/ 2001/ 07/ 27/ 106.1056/NEJMoa0809716. Niedbalski J. William R. . pubmedcentral. org/ giant-bushy-tailed-cloud-rat/ crateromys-schadenbergi/ info. html). . Retrieved 2009-11-22. Poulter." (http:/ / www. Stearns. "The lactase gene in Africa: Do you take milk?" (http:/ / genome.. html). doi:10. pone. com/ ridley/ a-z/ Ring_species. Genetics 163 (3): 939–53. com/ notrocketscience/ 2009/ 12/ british_birdfeeders_split_blackcaps_into_two_genetically_dis.0906199106. tusks. P.. Proceedings of the National Academy of Sciences 105 (10): 3796–9. p.. S. org/ articles/ 20070526/ fob5. Evolution. Science Daily (online) (Science News). McHugh in Resh. Huang X. Retrieved 2010-07-08. Society for the Study of Evolution 44. M. P. sciencedaily. The Human Genome.. J. A. PMC 1866175. htm) [67] Why scientists dismiss 'intelligent design' (http:/ / www. doi:10. S. Whitfield.. php). 2009). Ewbank. and flippers: the evolution of hoofed mammals. George W. Casadevall A. 171.. Dr. V. Retrieved 2010-07-08. Proceedings of the National Academy of Sciences 107: 1787. R. edu/ evolibrary/ article/ 0_0_0/ devitt_02 [57] Prothero. Robert M. PMID 12663534.5 Sympatric Speciation in Drosophila melanogaster" (http:/ / www. [60] Medical Research Council (UK) ((November 21. Shah. Dark Power: Pigment seems to put radiation to good use (http:/ / www. MSNBC.1371/journal. Schoch. lifesci.. Sept. [72] "Evolution in the urban jungle" (http:/ / theoystersgarter. org/ news/ features/ 2006/ 05/ new_caledonia. . The New England journal of medicine 361 (21): 2056–2065. Moadel T. Al-Dujaily. PMC 2268839. org/ ASA/ PSCF/ 2000/ PSCF6-00Haas. [50] "Giant bushy-tailed cloud rat (Crateromys schadenbergi)" (http:/ / www. JHU press. org/ web/ 20071215230655/ http:/ / www. Only a Theory: Evolution and the Battle for America's Soul (2008) pp. wellcome. (June 2000). Aisen P. Rough Guides. D. O. . 209 . D. doi:10. Cantarel. com/ 2008/ 03/ 12/ evolution-in-the-urban-jungle/ ).. Retrieved 2010-06-15. . com/ cej/ 5/ 2/ new-proteins-without-gods-help). . S. Jr. H. .S.0708446105. ac. Kenneth R. "Evidence for inversion polymorphism related to sympatric host race formation in the apple maggot fly.. [77] "Observed Instances of Speciation.S. J. asa3. [62] Byars. [75] William R.. Romero-Severson J (1 March 2003). 87. (2002). genetics. (2008). com/ SoyHealth/ SoyLactoseIntolerance. Retrieved 2007-09-27. Rice. html). plosone. "New Proteins Without God's Help" (http:/ / ncse. "British birdfeeders split blackcaps into two genetically distinct groups : Not Exactly Rocket Science" (http:/ / scienceblogs. gov/ articlerender. . msnbc. Guide to Philippine Flora and Fauna. . G. Rhagoletis pomonella" (http:/ / www. nmsr. [76] "he Evolution of Reproductive Isolation as a Correlated Character Under Sympatric Conditions: Experimental Evidence" (http:/ / www. [65] Thwaites WM (Summer 1985). William H. D. Liebherr and Joseph V.0000457. Cardé (Editors) 2003. Roethele JB. pnas. 80-82 [69] Science News. "Lake Baikal’s Vanishing Nerpa Seal" (http:/ / www. Campbell. (2007). "The Rev. Rough Guide to Evolution. [74] Ed Yong (December 3. [51] Rabor. org/ article/ fetchArticle. org/ nylon.. arkive. Hummerich. 1371/ journal. Week of May 26. Rouifed. C. PMC 1462491. Mark (2009). ISBN 0801871352. "Rapid evolution of seed dispersal in an urban environment in the weed Crepis sancta. Perspectives on Science and Christian Faith 52. Nosanchuk JD. Humphrey. birdlife. PMID 18316722. W. Carrue. Natural Resources Management Centre. soynutrition. Govindaraju. 325 by Davide Castelvecchi [70] Dadachova E. No. pdf). Joseph Boxhorn. nih. 2007. Encyclopedia of Insects. Retrieved 2008-07-18. [52] Robert R. Hashimoto H. doi:10. The Moscow Times. & R. (2009).". [54] Pallen. Retrieved 2010-05-23. The Boojum and its Home [53] Schofield. Rice. . Kansenshogaku Zasshi 63 (1): 15–26. Haas. edu/ eemb/ faculty/ rice/ publications/ pdf/ 25. blackwellpublishing. pp. berkeley. Filchak K. "A Novel Protective Prion Protein Variant that Colocalizes with Kuru Exposure. Academic Press. (1986). Uphill. org/ faqs/ faq-speciation. PMID 2501419. PMID 19923577. org/ content/ 105/ 10/ 3796. ISBN 978-1-85828-946-5. Bryan RA.pone. html). [55] http:/ / www. . 21 . [73] James K. ScienceBlogs. com/ id/ 9452500/ page/ 2/ ). . [58] Tanaka T. [66] Evolution and Information: The Nylon Bug (http:/ / www. Retrieved 2010-07-08. Ministry of Natural Resources and University of the Philippines. "Brain Disease 'Resistance Gene' Evolves in Papua New Guinea Community. Salt. pp. org/ cgi/ pmidlookup?view=long& pmid=12663534). Retrieved 2010-05-21. 2005 [68] Miller.. themoscowtimes. sciencenews.1073/pnas.. (1989). George W. Could Offer Insights Into CJD" (http:/ / www.. T. [78] Feder JL. Dallinger F. Donald R. action?articleURI=info:doi/ 10.R. fcgi?tool=pmcentrez& artid=2868295). PLoS ONE 2 (5): e457. . [87] "New plant species arise from conflicts between immune system genes" (http:/ / scienceblogs. pnas. Collier TC (April 2000). "Evolvability is a selectable trait" (http:/ / www. Bush GL (1969). doi:10. com/ Biology/ Zoology/ Adaptive-Traits-of-the-Polar-Bear-Ursus-Maritimus. "An electrophoretic analysis of Rhagoletis (Diptera: Tephritidae) phylogeny" (http:/ / jstor. et al.47. pnas.21. Lowe.10747. . Keyghobadi N. html [98] Adami C. 1998-08-26. html).. doi:10. [81] Fonseca DM. doi:10. . [95] Missouri Botanical Garden. springerlink.1073/pnas. org/ cgi/ content/ full/ 101/ 32/ 11531). Roethele JB. pp. kaist.ento. org/ pss/ 2446125). [96] Raven. Berlocher SH. . PMID 15001783. Bull. .94. Nomenclatural and Specimen Data Base. Crossing between Brassico-raphanus and B. html) [103] In Vitro Molecular Evolution (http:/ / www. ac. 91 (22): 10747–51. org/ bear-facts/ polar-bear-evolution/ ) [90] Ron Rayborne Accepts Hovind's Challenge (http:/ / www. ISBN 0716762846. org/ cgi-bin/ search_pick?name=Senecio+ vulgaris). . "Culex pipiens in London Underground tunnels: differentiation between surface and subterranean populations". Nichols RA (January 1999). physiological and molecular characteristics of underground populations of the urban mosquito Culex pipiens Linnaeus f. php). [92] Terasawa. org/ files/ pdfs/ wr380. Peter H. . molestus Forskål (Diptera: Culicidae) from several areas of Russia European Mosquito Bulletin. 101 (32): 11531–6. PMC 23485. org/ gecco-2005/ free-tutorials. Ed Yong.6884120. Journal of the European Mosquito Control Association ISSN1460-6127 [85] http:/ / www. . com/ p/ articles/ mi_m1134/ is_1_110/ ai_70770157). G.1126/science. Oecologia 76 (1): 138–47. doi:10. PMID 11729091. scienceray. doi:10. Berlocher SH (October 1997). polarbearsinternational. com/ notrocketscience/ 2009/ 05/ giant_insect_splits_cavefish_into_distinct_populations. Biology of Plants (7th rev.D. [100] Stemmer WP (October 1994). Science 303 (5663): 1535–8. org/ cgi/ pmidlookup?view=long& pmid=7938023). Polyploid hybrids of Raphanus sativus X Brassica oleracea L.. ch/ gentech/ 1998/ Jul-Sep/ msg00188. . doi:10.145312. Bot. "Sympatric speciation in phytophagous insects: moving beyond controversy?". pdf). American Journal of Botany 83 (10): 1365–1372. Annu Rev Entomol.1038/sj. OCLC 183148564. PMID 15289608. trnmag. 97 (9): 4463–8. [84] Vinogradova EB and Shaikevich EV Morphometric. Freeman.H. Systematic Zoology 31 (2): 136–55. doi:10. Proc Natl Acad Sci USA.310/ 5747/ 502/ DC1 210 . Richard J. "Selective maintenance of allozyme differences among sympatric host races of the apple maggot fly" (http:/ / www. PMID 10200079. Malcolm CA. "Evolution of biological complexity" (http:/ / www. com/ 250K/ ron. Coyne (2009).1073/pnas. ed..1007/BF00379612 (inactive 2010-01-08). isgec. "Insect From the Underground — London.22. Retrieved 2010-05-22. Missouri State Library.2307/2413033. org/ cgi/ content/ full/ sci. Proc Natl Acad Sci USA. "Origins of the New Allopolyploid Species ‘’Senecio camrensis (asteracea)’’ and its Relationship to the Canary Islands Endemic ‘’Senecio tenerifae’’" (http:/ / www. gene. (2005).1094247. Evolution 23 (2): 237–51. sciencemag. Deem MW (August 2004). [88] Adaptive Traits of the Polar Bear (Ursus Maritimus) (http:/ / www. Wlazlo B. mosquitocatalog. 22(2007). PMID 11038585. org/ cgi/ content/ full/ 94/ 21/ 11417). Prokopy RJ. Proc Natl Acad Sci USA. pnas. Heredity 82 (1): 7–15. org/ stable/ 2413033). doi:10. [94] Jerry A. ""Accelerated Evolution" Converts RNA Enzyme to DNA Enzyme In Vitro" (http:/ / www. Ofria C. 207777) [89] Polar Bear Evolution (http:/ / www. (March 2004). [82] Alan Burdick (2001). 17-24. et al. [83] Byrne K.91. 94 (21): 11417–21. htm) [91] Karpechenko.2307/2406788. PMC 45099. PMC 18257. Appl. kr/ ~jsrhee/ research03. "Emerging vectors in the Culex pipiens complex" (http:/ / www.).9. .Evidence of common descent [79] Berlocher SH. "Behavioral evidence for host races in Rhagoletis pomonella flies" (http:/ / www. . TSRI – News & Views 6 (11). "TROPICOS Web display Senecio vulgaris L. doi:10. 187 – 189. [80] "London underground source of new insect forms" (http:/ / www.hdy. html#ivme) [104] http:/ / www. Japanese Journal of Genetics. jstor. 8(4): 229–230 (1933). 47: 773–815. scripps. . chinensis and Raphanus sativus.1073/pnas. The Times. Ed Yong. Bush GL (1982). PMC 511006. New York: W. "Sympatric host race formation and speciation in frugivorous flies of the genus Rhagoletis (Diptera: Tephritidae)" (http:/ / jstor.2307/2446125.1146/annurev. Diehl SR. doi:10. [102] (http:/ / bio. Cooley SS (June 1988). "DNA shuffling by random fragmentation and reassembly: in vitro recombination for molecular evolution" (http:/ / www. kent-hovind." (http:/ / mobot. Retrieved 2008-02-01. PMID 10781045. England Underground home to different species of mosquitos" (http:/ / findarticles. php). [93] Andrew J. Natural History. ISBN 978-0-670-02053-9.091201. Retrieved 2010-05-22. pnas. [99] Earl DJ.1073/pnas. Y. 17:305-408 (1927). doi:10. html).97. Feder JL (2002). Feder JL. mobot. com/ Stories/ 2003/ 052103/ Simulated_evolution_gets_complex_052103. org/ stable/ 2406788). org/ cgi/ content/ full/ 97/ 9/ 4463). . Why Evolution is True. com/ nature/ journal/ v403/ n6766/ full/ 403158a0. [101] Sauter E (March 27. com/ notrocketscience/ 2009/ 08/ new_plant_species_arise_from_conflicts_between_immune_system. . edu/ newsandviews/ e_20060327/ evo. nature. com/ content/ p1716r36n2164855/ ?p=d8018d5a59294c2984f253b7152445b7& pi=20). Proc Natl Acad Sci USA. . 2006). Penguin Group.11417. Abbott (1996). PMID 7938023.4463.0404656101. [97] http:/ / www. html [86] "Giant insect splits cavefish into distinct populations" (http:/ / scienceblogs. • Sober.com/nature/newspdf/evolutiongems.php) • Evolution by Natural Selection (http://www. Endler.edu/books/ 0309063647/html/) • Evolution (http://www. Cambridge.chainsofreason.com — How Evolution Works (http://science. YK (2004). Murray. London: J. What evolution is. ISBN 0-691-08387-8. Elliott (2008).berkeley. Coyne. ISBN 0-517-12320-7 Dawkins. (dated 1998. Taphonomy: a process approach. London: John Murray. html) • Understanding Evolution: Your one-stop source for information on evolution (http://evolution. New York: Oxford University Press. Richard (2009).org/faqs/comdesc/) • TalkOrigins Archive . ISBN 0-521-33115-3. Oxford.howstuffworks. Fossils in the making: vertebrate taphonomy and paleoecology. D. Manhattan Press. ISBN 0-7195-7552-4. Chicago: University of Chicago Press. The Greatest Show on Earth: The Evidence for Evolution. New York: Basic Books. 1859.shtml) • National Academy Press: Teaching About Evolution and the Nature of Science (http://www. Evidence and Evolution: The logic behind the science. The adequacy of the fossil record.org/categories/index/ genome/evolution. Cambridge University Press. Natural selection in the wild. 502 pages. 1998. Donovan SK (1998). Reprinted: Gramercy (May 22. • • • • • • Ho.29+ Evidences for Macroevolution: The Scientific Case for Common Descent (http:// www. Darwin. Advanced-level Biology for Hong Kong. External links • National Academies Evolution Resources (http://nationalacademies.nature.html) — Provided by PBS.pdf) . The Empire of chance: how probability changed science and everyday life. UK: Cambridge University Press. Futuyma. ISBN 962-990-635-X • Martin RE (1999). Cambridge.htm/printable) • 15 Evolutionary Gems (http://www.Evidence of common descent 211 Further reading • • • • Biological science. Clegg CJ (1998).pbs. (1986). Jerry A. ISBN 978-0-19-923084-6. ISBN 0-465-04426-3. Behrensmeyer AK (1980). Evolutionary Biology.edu/ evosite/evo101/index. On the Origin of Species by means of Natural Selection or the Preservation of Favoured Races in the Struggle for Life. Gerd (1989). New Jersey: Princton University Press.nap. New York: John Wiley. UK: Cambridge University Press.com/evolution. ISBN 0-226-04169-7. ISBN 978-0-521-87188-4. Albemarle Street. ISBN 978-1-4165-9478-9.Transitional Vertebrate Fossils FAQ (http://www.org/wgbh/evolution/index.talkorigins. Hill A.org/faqs/faq-transitional. 1995). • Paul CRC. Genetics & evolution. • Evolution News from Genome News Network (GNN) (http://www. Charles November 24. (2009). Why Evolution is True. John A.genomenewsnetwork. Sinauer Associates. ISBN 0-521-59833-8. Ernst (2001).org/wiki/Chain_3) — An introduction to the logic of the theory of evolution by natural selection • Howstuffworks. published 1997) ISBN 0-87893-189-9 Gigerenzer. ISBN 0-471-96988-5. 3rd ed. Bantam Press. • Mayr.J. 2002.talkorigins.org/evolution/) • TalkOrigins Archive . Massachusetts. Sunderland. temperature regulation. vocalizations. by natural selection. By using the term adaptation for the evolutionary process. Adaptation. is the requirement that each genetic and phenotypic change during evolution should be relatively small. . it is not clear what "relatively small" should mean. Behavioural adaptations are composed of inherited behaviour chains and/or the ability to learn: behaviours may be inherited in detail (instincts). or their ability to run fast and escape predators. From this we see that adaptation is not just a matter of visible traits: in such parasites critical adaptations take place in the life-cycle. secreting slime.[11] [12] Adaptation is not always a simple matter. making venom. The other is speciation (species-splitting or cladogenesis). where the bodily structure is greatly simplified. This places constraints on the evolution of development. The main constraint. the adaptation of horses' teeth to the grinding of grass. because developmental systems are so complex and interlinked. the two senses of the word may be distinguished.[13] The origin of the symbiosis of multiple micro-organisms to form a eukaryota is a more exotic example. adaptation is often used for the product: those features of a species which result from the process.[5] For example. An organism must be viable at all stages of its development and at all stages of its evolution.[9] [10] A favourite example used today to study the interplay of adaptation and speciation is the evolution of cichlid fish in African lakes. Julian Huxley[6] Adaptation is. Adaptation is one of the two main processes that explain the diverse species we see in biology. body covering. and also the internal organization).[3] and is one of the basic phenomena of biology. first of all. as a practical term. Such adaptations are produced in a variable population by the better suited forms reproducing more successfully.[1] [2] This process takes place over many generations. but also more general functions such as growth and development. a process. then.[8] However. General principles The significance of an adaptation can only be understood in relation to the total biology of the species.212 Mechanisms Adaptation Adaptation is the evolutionary process whereby a population becomes better suited to its habitat. ionic balance and other aspects of homeostasis. behaviour and structure of organisms. rather than a physical part of a body. Physiological adaptations permit the organism to perform special functions (for instance. such as the different species of Darwin's finches. where the ideal phenotype evolves for a given external environment. armament. which is often quite complex. However. where the question of reproductive isolation is much more complex.[7] The distinction may be seen in an internal parasite (such as a fluke). but nevertheless the organism is highly adapted to its unusual environment. affects all aspects of the life of an organism. though there are always some features whose function is in doubt. behavioral or physiological.[15] These adaptive traits may be structural. Many aspects of an animal or plant can be correctly called adaptations. that is. Examples: searching for food. phototropism). over which there has been much debate. caused by geographical isolation or some other mechanism. mating. Structural adaptations are physical features of an organism (shape.[4] The term adaptation may also refer to a feature which is especially important for an organism's survival. for example polyploidy in plants is a reasonably common large genetic change.[14] All adaptations help organisms survive in their ecological niches. and adaptive trait for the bodily part or function (the product). or a tendency for learning may be inherited (see neuropsychology). This can sometimes be tested through a reciprocal transplant experiment. however. Aristotle.[25] The Bridgewater Treatises are a product of natural theology. did believe in final causes.Adaptation 213 Definitions The following definitions are mainly due to Theodosius Dobzhansky. the one does not imply the other: a phenotype with high adaptedness may not have high fitness. whereas fitness predicts a trait's future. Adaptedness is the state of being adapted: the degree to which an organism is able to live and reproduce in a given set of habitats. though some of the authors managed to present their work in a fairly neutral manner. insofar as they are heritable.[17] 3.[23] William Paley believed that organisms were perfectly adapted to the lives they lead. 2. The average contribution to the next generation by a phenotype or a class of phenotypes. Absolute fitness.[26] . The extent to which a phenotype fits its local ecological niche. adaptation was interpreted as the work of a deity. Adaptation is the evolutionary process whereby an organism becomes better able to live in its habitat or habitats.[16] 2. an argument that shadowed Leibniz. Adaptedness. and Hume also argued against design. relative to the contributions of other phenotypes in the population. Dobzhansky mentioned the example of the Californian redwood. A few of the most significant ideas:[22] • Empedocles did not believe that adaptation required a final cause (~ purpose). Differences in fitness between genotypes predict the rate of evolution by natural selection. This is also known as Darwinian fitness. who had argued that God had brought about the best of all possible worlds. it is clear that there is a relationship between adaptedness and fitness (a key population genetics concept). Brief history Adaptation as a fact of life has been accepted by all the great thinkers who have tackled the world of living organisms.[16] Elliott Sober commented that adaptation was a retrospective concept since it implied something about the history of a trait.[18] Adaptedness and fitness From the above definitions.[20] 1. but "came about naturally. Natural selection changes the relative frequencies of alternative phenotypes. which is highly adapted. The series was lampooned by Robert Knox. as the Bilgewater Treatises. 1. The absolute contribution to the next generation by a phenotype or a class of phenotypes. who held quasi-evolutionary views. Darwin broke with the tradition by emphasising the flaws and limitations which occurred in the animal and plant worlds. It is their explanations of how adaptation arises that separates these thinkers. Also known as the Malthusian parameter when applied to the population as a whole. even as evidence for the existence of God. • In natural theology. Relative fitness.[19] Although the two are connected. since such things survived". selective coefficient. and other terms. but a relic species in danger of extinction.[21] 3. Voltaire's Dr Pangloss[24] is a parody of this optimistic idea. An adaptive trait is an aspect of the developmental pattern of the organism which enables or enhances the probability of that organism surviving and reproducing. Lawrence). Niles Eldredge[31] Changes in habitat Before Darwin. The relative numbers of species in a given habitat are always changing. this is the typical response of flying insects or oceanic organisms. When the habitat changes.[33] Genetic change Genetic change is what occurs in a population when natural selection acts on the genetic variability of the population. accepted adaptation. Also.[27] He proposed a tendency for organisms to become more complex. Also. habitats are subject to changes in their biota: for example. invasions of species from other areas. moving up a ladder of progress. and especially by Weismann. It is now clear that habitats and biota do frequently change. the population adapts genetically to its circumstances.[36] [37] . This was known even before Mendel by medical men interested in human races (Wells. though much depends on the speed and degree of the change. the process affects every species in a particular ecosystem. but was a much stronger force than had previously been thought. usually expressed as use and disuse. for putting forward a mechanism whose significance had only been glimpsed previously. it may happen that the environment changes little.Adaptation 214 • Lamarck. and then the species becomes less and less well adapted. but especially when the population cannot or does not move to another. less hostile area. plus "the influence of circumstances". which goes on all the time to some extent. By this means. adaptation is a genetic tracking process. This illustrates the real merit of Darwin and Wallace.[34] Genetic changes may result in visible structures. Many other students of natural history. and some also accepted evolution. it follows that the process of adaptation is never finally complete. It was not appreciated that as the climate changed. In fact. to a greater or lesser extent. it may happen that changes in the environment occur relatively rapidly. and the species comes to fit its surroundings better and better. and as the habitat changed. Habitat tracking When a habitat changes. and secondary figures such as Lamarck Bates. all three things may occur in sequence.[35] Over time. the most common thing to happen is that the resident population moves to another locale which suits it.[32] This common response is called habitat tracking. fail because they cannot be reconciled with heredity. On the other hand. only genetic change brings about adaptation. genetic change or extinction. experimental field studies and breeding experiments by such as Ford and Dobzhansky produced evidence that natural selection was not only the 'engine' behind adaptation. who have wide (though not unlimited) opportunity for movement. so did the habitat. such as Buffon. His is a proto-evolutionary theory of the inheritance of acquired traits. Of these three effects.[28] [29] [30] Types of adaptation Adaptation is the heart and soul of evolution. It is one explanation put forward for the periods of apparent stasis in the fossil record (the punctuated equilibrium thesis). adaptation was seen as a fixed relationship between an organism and its habitat. so did the biota. without voicing their opinions as to the mechanism. or may adjust physiological activity in a way that suits the changed habitat. His evolutionary ideas. whose main purpose is to explain adaptations by natural means. Change is the rule. Seen like this. and those of Geoffroy. Therefore. three main things may happen to a resident population: habitat tracking. A century later. There are many examples of this.[38] . and may continue on a trajectory for millions of years. wing structures. and mouthpart morphologies of fossilized beetles and flies suggest that they acted as early pollinators. but with the life of other species. new or 'improved' adaptations which occur in one species are often followed by the appearance and spread of corresponding features in the other species. 215 Intimate relationships: co-adaptations In co-evolution. competing species had to constantly adapt to maintain their relative standing. These relationships are intrinsically dynamic. as has the relationship between flowering plants and insects (pollination). The evolution of nectaries in late Cretaceous flowers signals the beginning of the mutualism between hymenopterans and angiosperms. where the existence of one species is tightly bound up with the life of another species.Adaptation Van Valen thought that even in a stable environment. This became known as the Red Queen hypothesis. as can be seen by the colour of the pollen in their baskets: • • • • • • • • Co-extinction Infection-resistance Mimicry Mutualism Parasite-host Pollination syndrome Predator-prey Symbiosis The gut contents. not just with the physical environment. The association between beetles and angiosperms during the early Cretaceous period led to parallel radiations of angiosperms and insects into the late Cretaceous. the idea emphasises that the life and death of living things is intimately connected. Pollinator constancy: these honeybees selectively visit flowers from only one species. Such mimicry does not need to be perfect to improve the survival of the palatable species. . is a story of waste. Such adaptations may have significant consequences. A common example seen in temperate gardens is the hover-fly.[41] All aspects of this situation can be.[40] Bates. Compromise and conflict between adaptations It is a profound truth that Nature does not know best. mammals' hair helps temperature.. Adaptations serving different functions may be mutually destructive. many of which – though bearing no sting – mimic the warning colouration of hymenoptera (wasps and bees). Here the risk to life is counterbalanced by the necessity for reproduction.[45] Such adaptations may make the clade (monophyletic group) more viable in a wide range of habitats. compromise and blunder. Selection pressures pull in different directions. would be temperature regulation. or improvements in brain function.[49] Another example: camouflage to avoid detection is destroyed when vivid colors are displayed at mating time. but they can't scratch their backs.[43] • More on mimicry: Warning Colour and Mimicry [44] Lecture outline from University College London 216 A and B show real wasps. Peter Medawar[46] All adaptations have a downside: horse legs are great for running on grass. it is impossible to improve simultaneously all aspects of the phenotype to the same degree.[39] This is the mimicry by a palatable species of an unpalatable or noxious species. the subject of research. not perfection. the topic connects strongly to speciation. and to score victories in the annual rut. The acquisition of such major adaptations has often served as the spark for adaptive radiation. but offers a niche for ectoparasites. Their size during the last glacial period presumably depended on the relative gain and loss of reproductive capacity in the population of elks during that time.. Ernst Mayr. and have been. (often supposed to be far too large. makeshift. a view which is now standard amongst biologists. An example in plants would be the development of the reproductive system in flowering plants.Adaptation Mimicry Henry Walter Bates' work on Amazonian butterflies led him to develop the first scientific account of mimicry. the only flying penguins do is under water. and huge success for long periods of time for a whole group of animals or plants. But they are costly in terms of resource. in vertebrates. or an effective immune system. that genetical evolution.[42] Field and experimental work on these ideas continues to this day. genetics and development. Wallace and Müller believed that Batesian and Müllerian mimicry provided evidence for the action of natural selection. and the adaptation that results is some kind of compromise. Examples. the rest are mimics: three hoverflies and one beetle.[47] Since the phenotype as a whole is the target of selection.[48] Consider the antlers of the Irish elk. Obviously antlers serve positively for defence against predators. The basic machinery: internal adaptations There are some important adaptations to do with the overall coordination of the systems in the body. Compromise and make-shift occur widely. especially the kind of mimicry which bears his name: Batesian mimicry. in deer antler size has an allometric relationship to body size). also. and is hugely conspicuous.[57] White Leghorn fowl are markedly more resistant to vitamin B deficiency than other breeds. by the selective advantage of heterozygotes. with a big natural range in India. Much of the problem comes from our upright bipedal stance.[59] In diploid eukaryotes. For example. this is a consequence of the system of sexual reproduction. the blue peafowl Pavo cristatus is a pretty successful species. That is the result of the birth compromise. (which cannot be larger than about 400ccs. An Indian Peacock's train in full display The conflict between the size of the human foetal brain at birth."[50] The kind of sexual selection represented by the peacock is called 'mate choice'. the polyploid rice-grass Spartina townsendii is better adapted than either of its parent species to their own habitat of saline marsh and mud-flats. .[60] [61] We owe this esoteric knowledge to the comparative anatomists. Co-option of existing traits: exaptation The classic example is the ear ossicles of mammals. Neanderthals had a similar problem. its growth costs food resources. The first experimental evidence of the pre-adaptive nature of genetic variants in micro-organisms was provided by Salvador Luria and Max Delbrück who developed fluctuation analysis. a century ago. speech) just have to wait while the brain grows and matures. but on a restricted diet this preadaptation could be decisive.Adaptation 217 The peacock's ornamental train (grown anew in time for each mating season) is a famous adaptation. and further back still were part of the gill arches of early fish. a method to show the random fluctuation of pre-existing genetic changes that conferred resistance to phage in the bacterium Escherichia coli. Micro-organisms.[58] On a plentiful diet there is no difference.[51] The recognition of sexual selection was for a long time in abeyance. without which our pelvis could be shaped more suitably for birth. with their huge populations. with an implication that the process selects the more fit over the less fit. where mutant alleles get partially shielded. Julian Huxley[56] Pre-adaptations This occurs when a species or population has characteristics which (by chance) are suited for conditions which have not yet arisen. in exclusive relation to reproduction. else it will not get through the mother's pelvis) and the size needed for an adult brain (about 1400ccs). for example. and so has survival value. so the overall outcome of their mating system is quite viable. also carry a great deal of genetic variability. means the brain of a newborn child is quite immature.[52] In practice. were at the cutting edge of evolutionary studies. Pre-adaptation may occur because a natural population carries a huge quantity of genetic variability. It must reduce his maneuverability and flight.[63] The origin of wings from feathers that were originally used for temperature regulation is a more recent discovery (see feathered dinosaurs). but has been rehabilitated.[53] [54] [55] Shifts in function Adaptation and function are two aspects of one problem. which are surprisingly common in evolutionary history. The most vital things in human life (locomotion.[62] The word exaptation was coined to cover these shifts in function. Darwin's explanation of its advantage was in terms of sexual selection: "it depends on the advantage which certain individuals have over other individuals of the same sex and species. who. which we know from palaeontological and embrological studies originated in the upper and lower jaws and the hyoid of their Synapsid ancestors. but a change in habitats caused what used to be an adaptation to become unnecessary or even a hindrance (maladaptations). it is a gradual process caused by natural selection which changes the genetic make-up of a population so the collective performs better in its niche. In order to evolve to another. there is also co-extinction. Because genes have pleiotropic effects. With the loss of function goes the loss of positive selection. for changes during life which improve the performance of individuals. a trait may have been adaptive at some point in an organism's evolutionary history. it is certainly an insidious one". then obviously. on the other hand. Co-extinction can also occur when a flowering plant loses its pollinator. an advantage may accrue from their elimination once they are not functional.) exceeds the birth rate for a long enough period for the species to disappear. While co-extinction may not be the most important cause of species extinctions. . the extinction of parasitic insects following the loss of their hosts. learning is the term used for improvement in behavioral performance during life. Species extinction occurs when the death rate over the entire species (population. Adaptation. and are either not functional or reduced in functionality. the loss of structure in endoparasites.[66] Co-extinction Just as we have co-adaptation.. Examples: wisdom teeth in humans. Acclimatization is a term used for automatic physiological adjustments during life.. at least in that locale. and the subsequent accumulation of deleterious mutations. or through the disruption of a food chain.[64] Fitness landscapes Sewall Wright proposed that populations occupy adaptive peaks on a fitness landscape. acclimatization. occurs over many generations. Extinction If a population cannot move or change sufficiently to preserve its long-term viability. Such adaptations are termed vestigial. spandrels). It was an observation of Van Valen that groups of species tend to have a characteristic and fairly regular rate of extinction. Co-extinction refers to the loss of a species due to the extinction of another.Adaptation 218 Related issues Non-adaptive traits Some traits do not appear to be adaptive. for example. higher peak. As a result of changes in lifestyle the organs became redundant. not all traits may be functional (i.[68] Flexibility. Vestigial organs Many organisms have vestigial organs. a population would first have to pass through a valley of maladaptive intermediate stages.. In biology these terms are preferred.. Alternatively. the loss of pigment and functional eyes in cave fauna.[67] "Species co-extinction is a manifestation of the interconnectedness of organisms in complex ecosystems . it will become extinct. not adaptation. learning Flexibility deals with the relative capacity of an organism to maintain themselves in different habitats: their degree of specialization. they appear to have a neutral or even deleterious effect on fitness in the current environment. which are the remnants of fully functional structures in their ancestors. The species may or may not survive in other locales. These adjustments are not inherited genetically by the next generation. Since any structure represents some kind of cost to the general economy of the body.e. gene pool . that is.[65] A given population might be "trapped" on a peak that is not optimally adapted. Gradually the whole population becomes adapted to the new conditions. obviously. A heart pumps blood: that is its function. Now. that does not involve genetic transmission except to the extent that the capacities are inherited. but not what is learnt. The heart has a history (which may be well or poorly understood). and varies to some extent between individuals. rats. It would be quite wrong to use the word adaptation about a trait which arose as a by-product.[71] Once again. which is just an ancillary side-effect. on the other hand. and is possible for quite a few mammals and birds: of course. all living things can adjust to circumstances. and that history is about how natural selection formed and maintained the heart as a pump. A biologist would be conscious that sometime in the remote past feathers on a small dinosaur . although that is their normal function. The tendency to behave in a specialized or exploratory manner is inherited – it is an adaptation. The rapid colour changes in some flatfish. Similarly. chameleons are examples. A highly specialized animal or plant lives only in a well-defined habitat. respiration and physical exertion become a problem. A generalist. Over a longer period of time. or to its move to a different habitat.[74] [75] It is widely regarded as unprofessional for a biologist to say something like "A wing is for flying". but not the acclimatization itself.Adaptation Flexibility Populations differ in their phenotypic plasticity. The ability to acclimatize is an adaptation. It also says something about its history: how it has come about. and the capacities to be flexible are inherited. the capacity to learn is an inherited adaptation. an adaptation must have a functional history: therefore we expect it must have undergone selection caused by relative survival in its habitat. They will contribute more heavily to later generations. It also emits sound. This we know takes place. even when the new arrivals have had time to make physiological adjustments. but not the details of language. eats a specific type of food. there are huge differences between species. eats a range of food. Fecundity goes down. Acclimatization If humans move to a higher altitude. Every aspect of an organism that has a function has a history.[72] Some kinds of acclimatization happen so rapidly that they are better called reflexes.[69] [70] To a greater or lesser extent. 219 Function and teleonomy Adaptation raises some issues concerning how biologists use key terms such as function. That is not its function. Examples are humans. and can survive in many different conditions. crabs and many carnivores. some people will reproduce better at these high altitudes than others. but after spending time in high altitude conditions they acclimatize to the pressure by increasing production of red blood corpuscles. and cannot survive if its needs are not met. Function To say something has a function is to say something about what it does for the organism. the capacity for human speech is inherited. and pandas which require bamboo. Rather different is developmental flexibility: "An animal or plant is developmentally flexible if when it is raised or transferred to new conditions it develops so that it is better fitted to survive in the new circumstances". cephalopods. The degree of flexibility is inherited. which is the ability of an organism with a given genotype to change its phenotype in response to changes in its habitat. but deaths from some tropical diseases also goes down. because the performance of long-term communities at higher altitude is significantly better than the performance of new arrivals.[73] Learning Social learning is supreme for humans. extreme examples are koalas which depend on eucalyptus. Many herbivores are like this. 1980. Evolution the modern synthesis. the biologist would rather say that the wings on a bird or an insect usually had the function of aiding flight. they flaunt it. purpose and foresight. p449 [7] Mayr. The following is a definition for its use in biology: Teleonomy: The hypothesis that adaptations arise without the existence of a prior purpose. penguins. p1 [4] Williams. [10] Mayr. Natural History Museum. Oxford. California. It is contrasted with Aristotle's teleology. could no longer be considered a static condition." [8] Price P. So. London.W. Stansfield W. 7th ed. ostriches). Ernst 1982.g. Williams and Jacques Monod picked up the term and used it in evolutionary biology. and became instead a continuing dynamic process. 220 Teleonomy Teleonomy is a term invented to describe the study of goal-directed functions which are not guided by the conscious forethought of man or any supernatural entity.J. "Teleology is like a mistress to a biologist: he cannot live without her but he’s unwilling to be seen with her in public". p483: "Adaptation. George C. Ernest Nagel analysed the concept of goal-directedness in biology. a product of a creative past. adaptation hoards hindsight rather than foresight. [2] Bowler P. Princeton. but by the action of natural selection on genetic variability. Today the mistress has become a lawfully wedded wife. p10 [3] Patterson C. evolution and inheritance. Allen & Unwin. The only concession which they make to its disreputable past is to rename it ‘teleonomy’. 2006. Harvard.D.[77] it grew out of cybernetics and self-organising systems. Harvard. [9] Mayr E. Ernst 1982. p562–566 . 1999. Princeton. and that later many wings were not used for flying (e. Julian 1942. The growth of biological thought.C. Evolution: the history of an idea.[76] The term may have been suggested by Colin Pittendrigh in 1958. which has connotations of intention.Adaptation had the function of retaining heat. London. Evolution. The growth of biological thought: diversity. The evolutionary biology of parasites. and Mulligan P.[83] See also • • • • • • • • • • Adaptive radiation Co-adaptation Co-evolution Ecological trap Evolvability Exaptation Intragenomic conflict Maladaptation Mimicry Polymorphism (biology) References [1] The Oxford Dictionary of Science defines adaptation as "Any change in the structure or functioning of an organism that makes it better suited to its environment". [6] Huxley. Biologists no longer feel obligated to apologize for their use of teleological language. 1966.[82] and David Hull commented on the use of teleology and teleonomy by biologists: Haldane can be found remarking. Ernst Mayr. 2003. George C. A dictionary of genetics.[78] [79] [80] [81] Philosophers of science have also commented on the term. That would carry the connotation of being an adaptation with a history of evolution by natural selection. "Evolutionary adaptation is a phenomenon of pervasive importance in biology.. Animal species and evolution. Evolution is teleonomic. Harvard.. Adaptation and natural selection: a critique of some current evolutionary thought. 1963." p5 [5] Both uses of the term 'adaptation' are recognized by King R. World University Library. Symbiosis as a source of evolutionary innovation: speciation and morphogenesis MIT.antler size and skull size in 'Irish Elk'. evolution and inheritance.Y. [46] Medawar. Elliott 1993. The genetic theory of adaptation: a brief history. Am J. [25] Sober. Charles. Variation and evolution in plants. com/ content/ pdf/ 1471-2148-5-17. 1. 1974. G. p33 [32] Eldredge. p33–51: 'Fitness and adaptation'. Murray. University of Chicago Press. [48] Mayr. Methuen. Gould. ac. & Griffiths P. Ledyard. Natural selection in the wild. of course.1186/1471-2148-5-17. 2005. explosive speciation.C. .E. warning signals and mimicry. Harvard. p217 ISBN O-226-77304-3 [37] Freeman S.R. [44] http:/ / www. William 1986. The evolution of modern human childbirth.I. Columbia. Peter H. Ledyard. Smith. [20] Sober. Chapter 2 [26] Darwin. Oxford. N. Chapman and Hall. Evolution: the history of an idea. [30] Orr H. doi:10. 31. London. Pearson Education. Megaloceros giganteus. Genetics. Philosophy of biology. Princeton. N. Human Evolution (Florence) 9: 331-342. [31] Eldredge. Jr. 1968. Harvard. org/ doi/ abs/ 10. [43] Mallet. PMID 15723698. PMC 554777. 163). [22] references and details in their articles [23] Desmond. Genetics of natural populations XXV. Mimicry in plants and animals. 1986. Genetical theory of natural selection. html [45] Stebbins. John A. 2007. London. 2005. The niche is the central concept in evolutionary ecology. [40] Wickler W. p136. 82–92. 4th ed. 1930. 119–127. for example. Oxford. 887-8. James 2001. The future of Man. Nature Rev.Y. key-innovations and phylogeography of the haplochromine cichlid fishes" (http:/ / www. p33–51: 'Fitness and adaptation'. ecolsys. Wiley N. Leicestershire Museums. London. [39] Carpenter GDH and Ford EB 1933. Sherratt TN and Speed MP 2004. Ledyard. 2.A.1146/annurev. On some fundamental concepts of evolutionary biology. "Out of Tanganyika: Genesis. Of glaciers and beetles. Ernst 1982.T. pdf) (PDF). Stephen J. Genetics 6. Reinventing Darwin: the great evolutionary debate. see especially part II The niche: an abstractly inhabited hypervolume. Niles 1986. Adrian 1989. 1974. Cambridge. Reinventing Darwin: the great evolutionary debate. especially "Whatever the true nature of Lamark's theory. [51] The case was treated by Fisher R. Jr. Lynn (ed) 1991. Flowering plants: evolution above the species level. Charles 1871. Evolution 10. Henry Walter Bates FRS 1825-1892: explorer. Ernst 1982. J Evolutionary Biology 14. Methuen. Contains an extensive analysis of the evolution of adaptations in the radiation of Angiosperms. Irv. [19] Endler. p86–95. 1999. Columbia. [27] see. The growth of biological thought: diversity. p364 ISBN 0-13-227584-8 [38] Stebbins. 1970. p119–127. "African Cichlid Fishes: Model Systems for Evolutionary Biology" (http:/ / arjournals. p481 (and sequence) tells how Darwin's ideas on adaptation developed as he came to appreciate it as "a continuing dynamic process" (bottom p483). Genetic changes in populations of Drosophila pseudoobscura and Drosphila persimilis in some locations in California. The politics of evolution. Annual Review of Ecology and Systematics 31: 163. Science 196 1161–1166. 1986. Meyer A. John A. 6th edition. 84–87 [18] Dobzhansky T. Nancy & Jordan. The ecological theatre and the evolutionary play. not possible to test selective pressures on extinct populations in any direct way. ou l'optimisme. [13] Stebbins. [47] Jacob. 1872.ecolsys.Adaptation [11] Salzburger W. p589 [49] It is. [12] Kornfield. [52] Cronin.1. Oxford. biomedcentral. Nature Rev. 1968.. [14] Margulis. London. 1–34. (2005). [54] Friedlander. [29] Ford E. Niles 1995. The speciation revolution.. Mimicry. [36] Sterelny K. [24] In Candide. chapters 8 and 9. Chicago. Evelyn 1965. p4–6. doi:10. Evolution 28(2): 191-220. Harvard. [35] Mayr. p397: Rudimentary. p134–139. 6. Genetics of the evolutionary process. ISBN 0-262-13269-9 [15] Hutchinson G. The growth of biological thought: diversity. the discussion in Bowler. atrophied and aborted organs. it was his mechanism of adaptation that caught the attention of later naturalists".163. 2005. Niles 1995. [41] Moon H. 1146/ annurev. Mack T. uk/ ~ucbhdjm/ courses/ b242/ Mimic/ Mimic. p31/32. p26–78 [16] Dobzhansky T. Natural selection in the wild. [17] Dobzhansky T. Avoiding attack: the evolutionary ecology of crypsis. The origin of species. BMC Evolutionary Biology 5 (1): 17. Evolutionary biology 2. Leicester. Sewall Wright and evolutionary biology. The genetic theory of adaptation: a brief history. Elliott 1984. 2003.31.B. [21] following discussion in Endler. p64 [34] Orr H. & Herron J. Obstetric implications of Neanderthal robusticity and bone density. Flowering plants: evolution above the species level. Jr. 3rd ed. The Descent of Man and selection in relation to sex. [53] Rosenberg K. 1995. Harvard.Y. [42] Ruxton GD. footnote 18. Physical Anthropology 35. Princeton. Wiley. G. Francois 1977. [33] Eldredge. David K. 221 . Verheyen E. doi:10. .. G. Evolutionary analysis. Harvard. University of Chicago Press. annualreviews. (1974): Origin and function of 'bizarre' structures . London. p89–124. The ant and the peacock: altruism and sexual selection from Darwin to the present day. 1976. California. 79–82. Peter (November 2000). Ecological genetics. Sex and death: an introduction to philosophy of biology. The nature of selection: a philosophical enquiry. evolution and inheritance.P. Peter 1960. Evolution and tinkering. 1950. M. (p90) [28] Provine. Polyploidy. ucl. scientist and darwinian. Helen 1991. 1975. 1956.2307/2407322 (First page text) [50] Darwin. Yale. Time frames: the rethinking of Darwinian evolution and the theory of punctuated equilibria. Provine WB and Wallace B. George C. Qvarnström A & Irwin DE 2003. Chapter 4 Homology and the evidence for evolution. Lian Pih. a critique of some current evolutionary thought. 1966. red clover (Trifolium pratense. [72] Moore Lorna G. 1958. p33–50.S. The theory of evolution. and Regensteiner Judith G. 1632-1634. Springer-Verlag. Chapter 10. Adaptation and natural selection. Knopf.B. Sci. pages 587-614 in The evolutionary biology of hearing edited by Douglas B. p417 [57] Huskins C. 1973. Jacques 1971. p8–10 [76] "The hypothesis that adaptations arise without the existence of a prior purpose. [65] Wright. [83] Hull D. Journal of Philosophy 74: 261–301.5-billion-year history of the human body Pantheon Books 2008. 1939.H. Columbia University Press N. Alec. Biol. Geoffrey 2007. p32. [70] Price T. 1966. E. [80] Williams. Breed differences in resistance to a deficiency in vitamin B1 in the fowl. Sewall 1932. Chapter 28. Princeton. But one might question the word chance. Lerner (ed) Cause and effect. Moore JA. Classification.Adaptation [55] Miller.A. 531. Fay. Chapter 3 "The multiple meanings of teleological". Richard R. Section IV (Mammals). Evolution of the auditory system in Synapsida ("Mammal-like reptiles" and primitive mammals) as seen in the fossil record. Roe and George Gaylord Simpson (eds) Behavior and evolution. since natural selection.W. 1. Cambridge. 4–15. Cambridge. Science. In Proceedings of the Sixth International Congress on Genetics. J Exp Biol. [63] Gould. humble-bees (bumblebees). Webster. Dobzhansky's genetics of natural populations. inbreeding. 307–315. Phenotypic plasticity. 1993. and Simpson J. Free Press. Genetica 12. L. Charles Darwin's notebooks (1836–1844). George C. Fløistad (ed) Philosophy of Science Nijhoff. [59] [Dobzhansky T. ISBN 978-0-375-42447-2. [78] Mayr. London. 6th edition. eds Lewontin RC. Cambridge. Evolution the modern synthesis. and selection in evolution. In A. p285–304. [81] Monod. [77] Pittendrigh C. Teleology revisited: goal-directed processes in biology. The origin of Spartina townsendii. 2004. (eds) The evolution of mind: fundamental questions and controversies. New York. Adaptation to high altitude. Vrba 1982. 222 . 3rd ed. [61] Neil Shubin 2008. Chapter 9. Exaptation – a missing term in the science of form. Princeton. evolution and the nature of biology.Y. 1993. ISBN 0-387-97588-8. p85–86 [75] Williams. 270 p1433–1440. Hopson 1992. Agric. In Gangestad S. sexual selection and the evolution of colour patterns." Oxford Dictionary of Zoology. [69] Price TD. Allin and James A.F and Hutt F. Ann. The role of phenotypic plasticity in driving genetic evolution. [79] Mayr. Paleobiology 8. p57. Rev. 2006. ISBN 0-394-46615-2 [82] Nagel. is not a random process (it may be a stochastic or probabilistic process. In D. [66] Van Valen L. however). Washington 58. Chance and necessity: an essay on the natural philosophy of modern biology. Adaptation and natural selection: a critique of some current evolutionary thought. Cambridge. [60] Egdar F. mice and cats.L. crossbreeding. 1981. p33. Brain evolution. [67] Darwin in the Origin of Species tells the story of "a web of complex relations" involving heartsease (Viola tricolor). New York. 1992. [73] Maynard Smith uses the term physiologically versatile for such animals. [68] Koh. Res. Julian 1942. "Ears" [62] Panchen. 10 September 2004. Evolutionary Theory 1. 1–30. 5690. [56] Huxley. p355–366. 1983. [74] Sober. Allen & Unwin. Oxford. [58] Lamoreux W. 1931. 209 p2368–2376 [71] Maynard Smith J. Guildford. Adaptation. (ed) 1987. Stephen Jay and Elizabeth S. A new evolutionary law. Ernst 1988. Philosophy of biology. 3rd ed. Philosophy and biology. The theory of evolution. by its operation in particular habitats. natural selection and behavior. Maynard Smith J. Your Inner Fish: a journey into the 3. Proc. Elliott 1993. and Arthur N. Yale. 305. Anthropology 12. J. [64] Charles Darwin was the first to put forward such ideas: Barrett P. Toward a new philosophy of biology. but by chance may change the fitness of an organism. Ernst 1965. In G. 1977. Popper. Cause and effect in biology. The roles of mutation. Origin.D.] 1981. and may be beneficial. fixation can occur in just a few generations. In small populations. Repeat this process a number of times. .[2] the changes due to genetic drift are not driven by environmental or adaptive pressures. there will have been a random shift in the allele frequencies. which leads to changes in allele frequencies over time.) Repeat this process until there are 20 new marbles in the second jar. The effect of genetic drift is larger in small populations. The numbers of red and blue marbles picked each generation will fluctuate: sometimes more red. sometimes more blue. consisting of 20 marbles of various colors. and both colors correspond to two different alleles of one gene in the population. It may cause gene variants to disappear completely. In this example.[1] Genetic drift is an important evolutionary process. A population's allele frequency is the fraction of the copies of one gene that share a particular form. In contrast to natural selection. and this remained the dominant view for several decades. while the remaining blue allele has become fixed: all future generations will be entirely blue.[3] Analogy with marbles in a jar The process of genetic drift can be illustrated using 20 marbles in a jar to represent 20 organisms in a population. if no red marbles are selected the jar representing the new generation will contain only blue offspring.Genetic drift 223 Genetic drift Genetic drift or allelic drift is the change in the frequency of a gene variant (allele) in a population due to random sampling. Unless the second jar contains exactly 10 red and 10 blue marbles. The second jar will then contain a second generation of "offspring". In 1968 Motoo Kimura rekindled the debate with his neutral theory of molecular evolution which claims that most of the changes in the genetic material (although not necessarily changes in phenotypes) are caused by genetic drift. neutral. randomly select a marble from the original jar and deposit a new marble with the same color as its "parent" into a new jar. The alleles in the offspring are a sample of those in the parents. Vigorous debates wage among scientists over the relative importance of genetic drift compared with natural selection. (The selected marble remains in the original jar. This fluctuation is genetic drift – a change in the population's allele frequency resulting from a random variation in the distribution of alleles from one generation to the next. meaning they have no offspring. In each new generation the organisms reproduce at random.[4] Half of them are red and half blue. which makes gene variants more common or less common depending on their reproductive success. and smaller in large populations. or detrimental to reproductive success. and thereby reduce genetic variation. Ronald Fisher held the view that genetic drift plays at the most a minor role in evolution. If this happens. randomly reproducing each generation of marbles to form the next. the red allele has been lost permanently in the population. To represent this reproduction. and chance has a role in determining whether a given individual survives and reproduces. It is even possible that in any one generation no marbles of a particular color will be chosen. (B-B-B-A). The drop of solution then shrinks until it has only enough food to sustain four bacteria. (B-B-B-B). the law of large numbers predicts little change taking place over time when the population is large. (A-A-A-B). Among the four who survive. (A-B-B-B). The alleles are labeled A and B. there are sixteen possible combinations for the A and B alleles: (A-A-A-A). which in this example is the total number of surviving bacteria). Genetic drift is therefore considered to be a consequential mechanism of evolutionary change primarily within small. (B-A-A-A). (B-A-B-A). we get the following table. Thus both A and B have allele frequency 1/2. however. the effects of sampling error can alter the allele frequencies significantly. (A-A-B-B). This being the case. Probability and allele frequency In probability theory. (A-B-A-A). The probabilities are calculated with the slightly faulty premise that the peak population size was infinite. A and B are neutral alleles. (B-A-B-B).Genetic drift 224 In this simulation. (A-B-A-B). A B Combinations Probability 4 0 1 3 1 4 2 2 6 1 3 4 0 4 1 1/16 4/16 6/16 4/16 1/16 The probability of any one possible combination is where 1/2 (the probability of the A or B allele for each surviving bacterium) is multiplied four times (the total sample size. (B-A-A-B).[5] This effect can be illustrated using a simplified example. (B-B-A-A). Half of the bacteria have allele A and the other half have allele B. The bacteria are genetically identical except for a single gene for which there are two different alleles. When the reproductive population is small. meaning they do not affect the bacteria's ability to survive and reproduce. all bacteria in this colony are equally likely to survive and reproduce. . All the others die without reproducing. isolated populations. (A-B-B-A). there is fixation in the blue "allele" within five generations. (A-A-B-A). Consider a very large colony of bacteria which are isolated within a drop of solution. If each of the combinations with the same number of A and B respectively are counted. (B-B-A-B). where N is the effective population size. Each copy of the gene found in the new generation is drawn independently at random from all copies of the gene in the old generation. the total number of possible combinations to have an equal (conserved) number of A and B alleles is six.Genetic drift As seen in the table. computer simulations are usually easier to perform using the Wright-Fisher model. There is a higher probability that this surviving population will undergo drift (10/16) than the probability it will remain the same (6/16). The probability for any one of the possible combinations can be calculated with the formula 225 where N is the number of bacteria and k is the number of A (or B) alleles in the combination. The original colony began with an equal distribution of A and B alleles but chances are that the remaining population of four members has an unequal distribution. it takes N timesteps to get through one generation. a phenomenon known as population bottleneck. In diploid populations consisting of N individuals there are 2N copies of each gene. On the other hand. This expression can also be formulated using the binomial coefficient. which means that mathematical solutions are easier for the Moran model than for the Wright-Fisher model. and its probability is 6/16. The formula to calculate the probability of obtaining k copies of an allele that had frequency p in the last generation is then[7] where the symbol "!" signifies the factorial function. Each bacterium has a single allele. and the probability of unequal number of A and B alleles is 10/16. go down by one. We can call the frequency of one allele p and the frequency of the other q. annual plants have exactly one generation per year. Using the formula to calculate the probability that between them the surviving four bacteria have two A alleles and two B alleles. The Wright-Fisher model assumes that generations do not overlap. The total number of possible combinations can be represented as binomial coefficients and they can be derived from Pascal's triangle.[6] Genetic drift occurs when a population's allele frequencies change due to random events. Wright-Fisher model The example given uses bacteria to illustrate the effect of sampling and probability in genetic drift. In practice. or can stay the same. . This means that the transition matrix is tridiagonal. the Moran model and Wright-Fisher model give qualitatively similar results. the number of copies of a given allele can go up by one. The function '()' signifies the binomial coefficient and can be expressed as "N choose k". In the Wright-Fisher model. In the Moran model. because fewer time steps need to be calculated. one individual is chosen to reproduce and one individual is chosen to die. it takes just one. A or B. In this example the population contracted to just four random survivors. At each time step. but genetic drift runs twice as fast in the Moran model. For example. So in each timestep. An individual can have two copies of the same allele or two different alleles. The total number of possible alternative combinations is ten. Moran model The Moran model assumes overlapping generations. Thus even while genetic drift is a random. the direction of the random change in allele frequency each generation is strongly autocorrelated instead of independent over time. there is no residual influence on this probability from the frequency distribution of alleles in the grandparent. and because random declines or increases in allele frequency will influence the expected allele distributions for the next following generation. directionless process.[8] One important alternative source of stochasticity. When an allele reaches a frequency of 1 (100%) it is said to be "fixed" in the population and when an allele reaches a frequency of 0 (0%) it is lost.[12] . This means that the mathematical probabilities associated with the distribution of alleles in any generation are only derived from the distribution of alleles in the generation immediately prior. Genetic draft has very different mathematical properties. or selection. Because random sampling can remove but not replace an allele.Genetic drift 226 Random effects other than sampling error Random changes in allele frequencies can also be caused by other effects than sampling error. population—only that of the parent population.[11] Populations do not gain new alleles from the random sampling of alleles passed to the next generation. or any earlier.[10] However. perhaps more important than genetic drift. it acts to eliminate genetic variation over time. and the allele frequency cannot change unless a new allele is introduced in the population via mutation or gene flow. randomly mating populations the allele frequencies will tend to remain constant from one generation to the next unless the equilibrium is disturbed by migration. but the sampling can cause an existing allele to disappear.[9] Genetic draft describes the way the fate of a new mutation depends on the luck of which other alleles happen to be nearby at linked loci. is genetic draft. genetic mutation. Thus when random fluctuations result in a change of the allele frequency from the parent generation to the offspring generation. genetic drift comes to a halt. Once an allele becomes fixed. that deviation establishes new expected values for the allele distributions in the next generation to follow. Drift and fixation The Hardy–Weinberg principle states that within sufficiently large. unlike genetic drift. for example random changes in selection pressure. The predicted distribution of alleles among the offspring is a memory-less probability as described in the Markov property. genetic drift drives a population towards genetic uniformity over time. for fixation to occur can be estimated with probability. In estimating the drift of a neutral mutation in a large population. according to the Wright-Fisher model. is Ten simulations of random genetic drift of a single given allele with an initial frequency distribution 0. and p is the initial frequency for the given allele. and . Ne is the effective population size. and is proportional to the population size. where T is the number of generations. which is measured by the number of generations.[16] The expected time for the neutral allele to be lost through genetic drift can be calculated as[17] These calculations are also used to understand the impact genetic drift will have on a new allele introduced through genetic mutation. repeated in three reproductively synchronous populations of different sizes. The effective population size (Ne) takes into account factors such as the level of inbreeding. alleles drift to loss or fixation (frequency of 0.[14] Normally the effective population size. the number of organisms that are too old or young to breed. is used to determine these probabilities.5 measured over the course of 50 generations. which is smaller than the total population. the formulas can be simplified to[18] for average number of generations needed for fixation of a neutral mutation. at any given point in time the probability that an allele will eventually become fixed in the population is simply its frequency in the population at that given point in time. The result is the number of generations expected to pass before fixation occurs for a given allele in a population with given size (Ne) and allele frequency (p).0) significantly faster in smaller populations.[13] For example.[15] One forward-looking formula used for approximating the time needed for a neutral allele to become fixed through genetic drift. and the lower probability that two organisms that live far apart will reproduce.0 or 1. if the frequency p for allele A is 75% and the frequency q for allele B is 25%. such that fixation is predicted to occur much more rapidly in smaller populations. In general. (which can be assumed to begin as a single occurrence of the allele and thus its initial frequency will be negligible).Genetic drift 227 Time to fixation or loss Assuming genetic drift is the only evolutionary force acting upon an allele. The time needed. then given unlimited time the probability A will ultimately become fixed in the population is 75% and the probability that B becomes fixed is 25%. drift acts upon the genotypic frequencies within a population without regard their relationship to the phenotype. As a consequence. drift will predominate. also varies.[23] In a large population. selection lowers the frequencies for alleles that cause unfavorable traits. and thus affects genes differentially. and ignores those which are neutral. While natural selection is directioned. Selection indirectly rewards the alleles that develop adaptively advantageous phenotypes. For example. That is. genetic drift has no direction and is guided only by the mathematics of chance. natural selection acts directly on the phenotype and indirectly on its underlying genotype. . Drift is blind with respect to any advantage or disadvantage the allele may bring. Even weak selection forces acting upon an allele will push its frequency upwards or downwards (depending on whether the allele's influence is beneficial or harmful). When populations are very small.[21] Most mutations have a clear negative selective effect and cause the gametes that they occur in to disappear after a few generations. new advantageous mutations are almost as vulnerable to loss through genetic drift as are neutral mutations.[20] As a result. genetic drift and natural selection do not act in isolation. With a higher recombination rate. This means that mutation and selection in combination. Changes to the genotype caused by genetic drift may or may not result in changes to the phenotype. times the total population (f0 is between zero and one). some of the occurrencies will be removed because they are linked to such negative mutations. the degree to which alleles are affected by drift or selection varies according to population size. and may preserve unfavorable alleles and eliminate favorable ones (this means purifying selection has a stronger effect in species with a larger effective population[22] ). Alternatively. the effective population size is reduced by the factor f0. in cases where the allele frequency is very small. they are located in chromosomes that are removed because of selection against a mutation in another part of the same chromosome.[19] In cases where Ne and N are assumed to be equal. yet the fluctuations in their allele frequencies are all driven in a quantitatively similar manner. whereas drift is regarded as a sampling procedure across successive generations without regard to fitness pressures imposed by the environment. is said to be a factor f0. while disadvantageous mutations are usually eliminated quickly in large populations.Genetic drift 228 for the average number of generations needed for the loss of a neutral mutation. Selection responds specifically to the adaptive advantage or disadvantage presented by a phenotypic trait. and thereby genetic drift. even over many generations. both forces are always at play. the equilibrium frequency of non-deleterious alleles. This is because natural selection emblematizes the ecological interaction of a population. The size of the remaining population. effective population size. with an increase in reproductive success for the phenotype comes an increase in allele frequency. In drift each allele in a population is randomly and independently affected. genetic drift operates randomly while natural selection functions non-randomly.[21] In natural populations. However. as the small changes in frequency they would produce are overshadowed by drift. drift can also overpower selection—even in large populations. However. It is not until the allele frequency for the advantageous mutation reaches a certain threshold that genetic drift will have little effect. Because strength of genetic linkage varies along the chromosome. guiding evolution by impelling heritable adaptations to the environment. the probability of sampling error is small and little change to the allele frequencies is expected. It is possible to calculate how many percent of each generation will be removed by such mutations. the ratio of time-to-fixation over time-to-loss is[16] Genetic drift versus natural selection Although both processes drive evolution. Weak selective effects may not be seen at all. When a neutral mutation spreads by drift in a population. causes the drift to have more effect. By the same token. The statistical effect of sampling error during the reproduction of alleles is much greater in small populations than in large ones. The declines in population resulted from hunting and habitat destruction.[27] 229 Population bottleneck A population bottleneck is when a population contracts to a significantly smaller size over a short period of time due to some random environmental event. Currently the greater prairie chicken is experiencing low reproductive success. And its impact can be sustained. DNA analysis comparing birds from the mid century to birds in the 1990s documents a steep decline in the genetic variation in just in the latter few decades. but the random consequence has been a loss of most of the species' genetic diversity.[30] [31] There have been many known cases of population bottleneck in the recent past. a radical reduction in population size increases the likelihood of further allele fluctuation from drift in generations to come. and are not improved by any particular inherent genetic advantage. A population's genetic variation can be greatly reduced by a bottleneck. North American prairies were habitat for millions of greater prairie chickens. and even beneficial adaptations may be permanently eliminated.[33] . the odds for survival of any member of the population are purely random. Their resulting decline in genetic variation can be deduced by comparing it to that of the southern elephant seal which were not so aggressively hunted.[26] and negative correlation between gene density and diversity at noncoding sites.[24] [25] This effect is visible in molecular data as a correlation between local recombination rate and genetic diversity. Even when the allele frequency of the original population is carried forward in the surviving population.[32] Over-hunting also caused a severe population bottleneck in the northern elephant seal in the 19th century.[29] The loss of variation leaves the surviving population vulnerable to any new selection pressures such as disease. even when the bottleneck is caused by a Changes in a population's allele frequency following a population bottleneck: the rapid one-time event such as a natural and radical decline in population size has reduced the population's genetic variation. completely independent of selection. because adapting in response to environmental changes requires sufficient genetic variation in the population for natural selection to take place. The bottleneck can result in radical changes in allele frequencies. their numbers plummeted from about 100 million birds in 1900 to about 50 birds in the 1990s. catastrophe. climate change or shift in the available food source.[28] In a true population bottleneck.Genetic drift linkage decreases and with it this local effect on drift. Prior to the arrival of Europeans. In Illinois alone. [37] However. or natural selection.Genetic drift 230 Founder effect The founder effect is a special case of genetic drift. genetic drift) a significant driving force in the evolution of new species. When a newly formed colony is small. occurring when a small group in a population splinters off from the original population and forms a new one. In the figure this results in fixation of the red allele. newly isolated populations with his shifting balance theory of speciation. gene flow and mutation will all contribute to this divergence. His first use of the term "drift" was in 1929. though he was never entirely comfortable to see his name given to it.[39] though at the time he was using it in the sense of a directed process of change. He preferred "drifting at random". The random sample of alleles in the just formed new colony is expected to grossly misrepresent the original population in at least some respects. scientists were just beginning to blend the new science of population genetics with Charles Darwin's theory of natural selection. Sewall Wright was the first to attach this significance to random drift and small. the two separated populations may become distinct. increases. It came to be known as the "Sewall-Wright effect". or genetic distance. both genetically and phenetically. the founder effect occurs. make-up far into the future. Later that year he used it to refer to a purely random process. Wright focused on the effects of inbreeding on small relatively isolated populations.[34] It is even possible that the number of alleles for some genes in the original population is larger than the number of gene copies in the founders. Working within this new framework.[38] History of the concept The concept for genetic drift was first introduced by one of the founders in the field of population genetics. the small population experiences intensive drift. For a period after the foundation. As a result of many generations of inbreeding.[21] [35] The difference in gene frequencies between the original population and colony may also trigger the two groups to diverge significantly over the course of many generations. A well documented example is found in the Amish migration to Pennsylvania in 1744. This potential for relatively rapid changes in the colony's gene frequency led most scientists to consider the founder effect (and by extension. although not only genetic drift but also natural selection. Members of the colony and their descendants tend to be religious isolates and remain relatively insular.[36] Following after Wright. As the difference. or change due to the effects of sampling error. Ernst Mayr created many persuasive models to show that the decline in genetic variation and small population size following the founder effect were critically important for new species to develop. making complete representation impossible. there is much less support for this view today since the hypothesis has been tested repeatedly through experimental research and the results have been equivocal at best. its founders can strongly affect the population's genetic When very few members of a population migrate to form a separate new population. and "drift" came to be adopted as a technical term in the stochastic sense exclusively. Sewall Wright. Two members of the new colony shared the recessive allele for Ellis–van Creveld syndrome.[40] In the early days of the modern evolutionary synthesis. Ellis-van Creveld syndrome is now much more prevalent among the Amish than in the general population. He introduced the concept of an adaptive landscape in which phenomena such as cross breeding and genetic drift in small populations could push . Theoretical Aspects of Population Genetics. berkeley. 680. L. . 112.[41] Wright thought smaller populations were more suited for natural selection because "inbreeding was sufficiently intense to create new interaction systems through random drift but not intense enough to cause random nonadaptive fixation of genes. ISBN 0-87893-189-9. Retrieved 2009-11-17. Sinauer Associates. 4th edition."[40] Wright's views on the role of genetic drift in the evolutionary scheme were controversial almost from the very beginning.1038/nrg2526. Dan Graur (1991).J. steady. p. University of California Berkeley.[42] population geneticist Motoo Kimura rekindled the debate with his neutral theory of molecular evolution. [18] Wen-Hsiung Li. actionbioscience.[3] 231 See also • Allopatric speciation • Antigenic drift • Gene pool • Small population size Notes [1] [2] [3] [4] Futuyma. ActionBioscience. Hill MM. doi:10. ISBN 978-0-87893-308-2. gov/ articlerender. Leland (2004). Genetics of Populations. 29. Princeton University Press. Philip W. Daniel (2007). Douglas (1998). Natl. Avers. ISBN 0763747726. org/ content/ 104/ 13/ 5698. pp. [12] Li. [11] Hartl. 737. pp. Fisher conceded genetic drift played some role in evolution. Sidow A (March 2007). "Fundamental concepts in genetics: Effective population size and patterns of molecular evolution and variation". Process and Pattern in Evolution.Genetic drift them away from adaptive peaks. . [6] Walker J. Dan Graur (1991). Sinauer Associates. Proc. [7] Hartl. Genetics 146 (2): 723–33. [14] Otto S..1073/pnas. John H. 413. pnas. [13] Futuyma. 364. PMID 9178020. [15] Charlesworth B (March 2009). Princeton. Nat. Wen-Hsiung.[2] In 1968. George Gaylord (1967). PMC 1838466. Charlotte (1989). Alberto (1996). Whitlock M (1 June 1997). shtml).A. Dan Graur (1991). Andrew Clark (2007). Genet. 320. Ohta. p.S. Sinauer Associates. 104 (13): 5698–703. ISBN 978-0-87893-452-9. pubmedcentral. p. Douglas (1998). The history and geography of human genes. Paolo. ISBN 0-87893-452-9. Menozzi. which would in turn allow natural selection to push them towards new adaptive peaks. Sinauer Associates. pp. PMID 11794777. fourmilab. 102. Genetics: From Genes to Genomes. [22] Small KS. [19] Kimura. U. [21] Cavalli-Sforza. html). . To Fisher. ISBN 0-07-246248-5. which claims that most of the genetic changes are caused by genetic drift acting on neutral mutations.). [5] Zimmer. 300. see answer to question Is natural selection the only mechanism of evolution?)" (http:/ / www. ISBN 0691080984. pp. Retrieved 2009-11-01. [20] "Natural Selection: How Evolution Works (An interview with Douglas Futuyma. One of the most vociferous and influential critics was colleague Ronald Fisher. 102. viewing the process of evolution as a long. But the debates have continued between the "gradualists" and those who lean more toward the Wright model of evolution where selection and drift together play an important role. Rev. PMID 19204717. Retrieved 2009-11-24. p. Sinauer Associates. p. Fundamentals of Molecular Evolution. "The probability of fixation in populations of changing size" (http:/ / www. [8] Li. The RetroPsychoKinesis Project.: Princeton University Press. [23] Simpson. The Meaning of Evolution (Second ed. ISBN 0-06-095850-2. ISBN 0-691-02905-9. Carl (2002). ISBN 978-0-87893-308-2. . Futuyma.0700890104. p. [10] Hartwell el al. p. Evolutionary Biology. Fisher has been accused of misunderstanding Wright's views because in his criticisms Fisher seemed to argue Wright had rejected selection almost entirely. Jones and Bartlett Publishers. p. html). [17] Daniel Hartl. fcgi?tool=pubmed& pubmedid=9178020). PMID 17372217. long). Fourmilab. New York. p. ISBN 0-87893-452-9. Sinauer Associates. Yale University Press. adaptive progression was the only way to explain the ever increasing complexity from simpler forms. . McGraw Hill. Fundamentals of Molecular Evolution. [16] Hedrick. 33. Tomoko (2001). Oxford University Press. "Evolution 101:Sampling Error and Evolution" (http:/ / evolution. Daniel. 28. edu/ evosite/ evo101/ IIID1Samplingerror. org/ evolution/ futuyma. Wen-Hsiung. Principles of Population Genetics. ISBN 0-87893-189-9. N. Principles of Population Genetics. (2001). . Sinauer Associates. "Extreme genomic variation in a natural population" (http:/ / www. 2nd edition. Brudno M. ISBN 0-87893-189-9. Glossary. "Introduction to Probability and Statistics" (http:/ / www. Sinauer Associates. [9] Gillespie. Andrew Clark (2007). Evolution : The Triumph of an Idea. PMC 1208011.org. L. but an insignificant one. p. 10 (3): 195–205. Douglas (1998). Piazza. NY: Perennial. Evolution 55 (11): 2161–2169. 232. Evolutionary Biology. "Is the population size of a species relevant to its evolution?". Principles of Population Genetics. Sci. doi:10. ISBN 978-0-87893-308-2. Motoo. Evolutionary Biology. nih. ch/ rpkp/ experiments/ statistics. Acad. (2004). Sinauer Associates. Fundamentals of Molecular Evolution. com/ research/ population-bottleneck-gen-03/ ) [29] Futuyma. doi:10. Evolutionary Biology. Daniel J. p.. Craig H. PMC 1205596. Campbell (1996). Walter M. 3 (7): e196. (2000).). pp. Retrieved 2009-04-07. 470. Life: The Science of Biology. "The evolution of dominance". Jason. Springer.. (2004). Dictionary of Concepts in Physical Anthropology. Public Broadcast System. David E. Steward H. Gordon H. ISBN 0-313-24756-0.1038/217624a0. Biology.065. Systematics and the Origin of Species: on Ernst Mayr's 100th anniversary (Illustrated ed. Wolf.1. PMID 8375663. Francisco José Ayala (2005). [31] Cornuet JM.1371/journal. Edward J. . esajournals. . edu/ evosite/ evo101/ IIID3Bottlenecks. National Academies Press. PMC 1135296. 1251. H. Curr. "Climate change and the molecular ecology of arctic marine mammals" (http:/ / www. 1890/ 06-0795. p. [33] "Evolution 101: Bottlenecks and Founder Effects" (http:/ / evolution. 46. Brodie. 367. pubmedcentral. Biol. [25] Charlesworth B. PMID 5637732. .talkorigins. Douglas (1998). Modern Library. PMC 1207747. 52–57". Berkeley. Non-neutral evolution: theories and molecular data. ISBN 978-0-412-05391-7. ISBN 0-19-512806-0.. [26] Presgraves DC (September 2005).org/content/free/figures/ch15. Morgan MT. Diversity and Ecology. [39] Wright S (1929). PMID 15907155. org/ cgi/ reprint/ 134/ 4/ 1289). Epistasis and the evolutionary process. Luikart G (1996). 423. [35] "Genetic Drift and the Founder Effect" (http:/ / www. Hu TT. Ecological Applications 18 (2 Suppl): S56–S76.cub. [42] Kimura M (1968). doi:10. Berlocher. fcgi?tool=pmcentrez& artid=1135296). 1). "Recombination enhances protein adaptation in Drosophila melanogaster".evolution-textbook. ISBN 9780309095365. p. (2006). [34] Neill. fcgi?tool=pmcentrez& artid=1207747). 330. html). 303–304. Purves. Westport. Ernst. "The pattern of polymorphism in Arabidopsis thaliana" (http:/ / www. San Francisco: W.1016/j. The American Naturalist 63 (689): 556–61. Ishino Y. nih.. doi:10. pp. . "The effect of deleterious mutations on neutral molecular variation" (http:/ / www. ISBN 978-0679642886.2005. "Description and power analysis of two tests for detecting recent population bottlenecks from allele frequency data" (http:/ / www. Genetics 144 (4): 2001–14. David M. berkeley. The Benjamin/Cummings Publishing Company.1890/06-0795. pbs. gov/ articlerender. genetics. [40] Stevenson. pubmedcentral. PLoS Biol.1086/280290. (1998). [27] Nordborg M. "Chs. Evolution: The Remarkable History of a Scientific Theory. Heller. shtml). p.html) • Genetic drift illustrations in Barton et al. 21–33.0030196. Michael S. [41] Larson. Sinauer Associates. (July 2005). Edmund D. Freeman..07.Genetic drift [24] Golding B (1994). 232 External links • The TalkOrigins Archive (http://www. [28] Population Bottleneck | Macmillan Genetics (http:/ / www. Nature 217 (5129): 624–26. Conn: Greenwood Press. Charlesworth D (August 1993). bookrags. Jody Hey. html) .. [38] Howard. ISBN 0-8053-1940-9. PMID 16169487. 15 (18): 1651–6. [37] Mayr. Sadava. Fourth edition. [32] Hillis. Fitch. II: Evolution. doi:10. PMID 8978083. pp. org/ wgbh/ evolution/ library/ 06/ 3/ l_063_03.pbio. [30] O'Corry-Crowe G (2008). Endless Forms (Illustrated ed. ISBN 0-87893-189-9. 1.). Orians. [36] Wade. PMID 18494363. ISBN 9780195109016. Evolution. (http://www. Genetics 134 (4): 1289–303. org/ doi/ full/ 10. Retrieved 2009-04-07. gov/ articlerender. et al. William K. ISBN 9780716798569. nih. University of California. "Evolutionary rate at the molecular level". doi:10. United States: Oxford University Press.org/faqs/genetic-drift. Oxford [Oxfordshire]: Oxford University Press. Joan C. (1991). For example. they can be artificial. Maintained gene flow between two populations can also lead to a combination of the two gene pools. It was also found that this gene flow between European and West African groups is much greater in the Northern U. then the alleles in the pollen have effectively been able to move from the population on one side of the highway to the other. By measuring the frequencies of the West African and European groups. For example. pollen is likely to be transported from one side to the other and vice versa. because there is little to no gene flow to provide recombination of the gene pools. man-made barriers. It is for this reason that gene flow strongly acts against speciation. Barriers to gene flow need not always be physical. One of the most significant factors is mobility. repairing the developing differences in genetic variation that would have led to full speciation and creation of daughter species. in the United States. oceans. In some cases. Immigration may also result in the addition of new genetic variants to the established gene pool of a particular species or population. Barriers to gene flow Physical barriers to gene flow are usually. and thus. because malaria is almost non-existent. such as the Great Wall of China. In West Africa. yet show very limited gene flow due to limited hybridization or hybridization yielding unfit hybrids. but not always. by recombining the gene pools of the groups. Samples of the same species which grow on either side have been shown to have developed genetic differences. Migration into or out of a population may be responsible for a marked change in allele frequencies (the proportion of members carrying a particular variant of a gene). gene flow (also known as gene migration) is the transfer of alleles of genes from one population to another. Animals tend to be more mobile than plants. and this allele is thus present in nearly all of the West African population. the Duffy antigen provides some resistance to the disease. where malaria is prevalent. There are a number of factors that affect the rate of gene flow between different populations. If this pollen is able to fertilize the plant where it ends up and produce viable offspring. gene flow was observed between a white European population and a black West African population. if a species of grass grows on both sides of a highway. natural.S. . They may include impassable mountain ranges. In contrast. Species can live in the same environment. Europeans have either the allele Fya or Fyb. scientists found that the allele frequencies became mixed in each population because of movement of individuals.Gene flow 233 Gene flow In population genetics. reducing the genetic variation between the two groups. than in the South. Gene flow in humans Gene flow has been observed in humans. as greater mobility of an individual tends to give it greater migratory potential. or vast deserts. although pollen and seeds may be carried great distances by animals or wind. which were recently brought together. which has hindered the gene flow of native plant populations[1] . introgression and genetic swamping. hybridization with or without introgression may threaten a rare species' existence nonetheless. naturally-evolved.g. evolutionarily constructive processes." [4] Biologist Gogarten suggests "the original metaphor of a tree no longer fits the data from recent genome research".[9] [10] . Models of gene flow Models of gene flow can be derived from population genetics. These phenomena can be especially detrimental for rare species coming into contact with more abundant ones. Gene transfer. wild species can be threatened with extinction[7] through the process of genetic pollution. it is difficult to trace organismal phylogeny in the presence of HGT [horizontal gene transfer]. their genetic modification from reaching other conventionally hybridized or wild native plant and animal populations by using gene flow mitigation usually through unintentional cross pollination and crossbreeding." [6] Genetic pollution Purebred. and can exchange plasmids across species boundaries [3]. [5] "Using single genes as phylogenetic markers. Each contemporary molecule has its own history and traces back to an individual molecule cenancestor. Combining the simple coalescence model of cladogenesis with rare HGT [horizontal gene transfer] events suggest there was no single last common ancestor that contained all of the genes ancestral to those shared among the three domains of life. Reasons to limit gene flow may include biosafety or agricultural co-existence. . antigenic shift. defined as the movement of genetic material across species boundaries. which includes horizontal gene transfer. it becomes necessary to prevent "genetic pollution" i. "Sequence comparisons suggest recent horizontal transfer of many genes among diverse species including across the boundaries of phylogenetic "domains". Some degree of gene flow may be due to normal. either through hybridization or gene transfer from bacteria or virus to new hosts. However. and all constellations of genes and genotypes cannot be preserved. Nonnative species can bring about a form of extinction of native plants and animals by hybridization and introgression either through purposeful introduction by humans or through habitat modification. region-specific. That being said. potentially causing uncontrolled hybridization. Thus determining the phylogenetic history of a species can not be done conclusively by determining evolutionary trees for single genes. Sewall Wright's neighborhood model. The extent of this facet of gene flow is not always apparent from morphological (outward appearance) observations alone. and reassortment is sometimes an important source of genetic variation. These processes can lead to homogenization or replacement of local genotypes as a result of either a numerical and/or fitness advantage of introduced plant or animal[8] . exchange genes with living bacteria. Bacteria can incorporate genes from other dead bacteria. Biologists [should] instead use the metaphor of a mosaic to describe the different histories combined in individual genomes and use the metaphor of an intertwined net to visualize the rich exchange and cooperative effects of horizontal gene transfer. Interbreeding between the species can cause a 'swamping' of the rarer species' gene pool.e. Viruses can transfer genes between species [2]. Gene flow mitigation When cultivating genetically modified (GM) plants or livestock. creating hybrids that drive the originally purebred native stock to complete extinction. these molecular ancestors were likely to be present in different organisms at different times. e.Gene flow 234 Gene flow between species Gene flow can occur between species. Wright's island model and the stepping stone model. in which GM and non-GM cropping systems work side by side. bringing previously isolated species into contact. May 8. and Daniel Simberloff. esalenctr. USDA Forest Service. Erickson. by H. CO. fcgi?artid=33232) [8] Glossary: definitions from the following publication: Aubry. pdf [4] http:/ / opbs. any modification located in plastids cannot be transmitted by the pollen. 2001. Among these programmes are Transcontainer. html [5] http:/ / www. 1146/ annurev. In transplastomic plants. E. 7. edu/ ~melcher/ MG/ MGW3/ MG334. 1. Department of Wildlife Ecology. org/ lj/ community/ ref_courses/ 3484/ enmicro. and use on national forests and grasslands in the Pacific Northwest. pdf [7] Hybridization and Introgression. A. 27. okstate. nih. preventing the formation of pollen. it remains unclear how reliable cleistogamy is for gene flow mitigation: a Co-Extra research project on rapeseed investigating the matter has published preliminary results which cast doubt on the attainment of a high degree of reliability. 104/ search?q=cache:tpICVNWaTbgJ:non. which focuses on the biosafety of genetically modified plants. doi: 10.1146/annurev. edu/ gogarten/ articles/ TIG2004_cladogenesis_paper.1. which are cellular compartments outside the nucleus. Maine 04469. Mooney and E. 563 SW Jefferson Ave. consequently. Therefore.dyndns. An example for plastids are chloroplasts. nature. there are three approaches to gene flow mitigation: keeping the genetic modification out of the pollen.. Department of Biological Science. Florida 32306. This trait is called cleistogamy and occurs naturally in some plants. 102. Cleland" Proc Natl Acad Sci U S A. the modified DNA is not situated in the cell's nucleus but is present in plastids. nativeseednetwork. SIGMEA. Annual Review of Ecology and Systematics. However. The National Academy of Sciences (http:/ / www. Institute for Applied Ecology. 44 pages.98(10). edu/ ~bah/ BIO471/ Reader/ Pennisi_2003. Shoal and V. org/ display/ confpage. Proc Natl Acad Sci U S A. Male sterile plants are unable to produce functioning flowers and therefore cannot release viable pollen. Native Seed Network (NSN). 27.org [13] References [1] http:/ / www. 0.091093398. org/ sici?sici=0066-4162(1996)27<83:EBHAI>2. from "The evolutionary impact of invasive species. USA. Corvallis. Extinctions. Florida State University. html [2] http:/ / 66. uconn. • The first approach requires transplastomic plants. Cytoplasmic male sterile plants are known to produce higher yields. jstor. ecolsys. USA (http:/ / www.2-A#abstract) . Rhymer . USA. 2005. University of Maine.. the pollen does not contain plastids and. development. pubmedcentral. C. by Judith M. 83). Grass cultivars: their origins. OR 97333. Pages 83-109 (doi: 10. 98(10): 5446–5451. In some plants. • The second approach relies on male sterile plants. nau. researchers are trying to introduce this trait to genetically modified crops. com/ hdy/ journal/ v90/ n3/ full/ 6800237a. Orono. in which photosynthesis occurs. fiction. plus appendices. Vol. org/ article_view?id=13) [9] EXTINCTION BY HYBRIDIZATION AND INTROGRESSION.27. Cleistogamous plants produce flowers which either open only partly or not at all. and Co-Extra. annualreviews. November 1996.Gene flow Scientists in several large research programmes are investigating methods of limiting gene flow in plants. which studies the co-existence of GM and non-GM product chains. Generally.ecolsys. and keeping the pollen inside the flower. which investigates methods for biocontainment. cfm?confid=10& pageid=105& pgtype=1 [6] http:/ / web. v.83) (http:/ / arjournals. 235 See also • Biological dispersal • Genetic erosion • Genetic admixture External links • Co-Extra research on gene flow mitigation [11] • Transcontainer research on biocontainment [12] • SIGMEA research on the biosafety of GMOs: http://sigmea. Tallahassee. gov/ articlerender. R.1073/pnas. org/ doi/ abs/ 10. 2001 May 8. (http:/ / links. pdf+ sex+ evolution+ %22Horizontal+ gene+ transfer%22+ -human+ Conjugation+ RNA+ DNA& hl=en [3] http:/ / www2. • The third approach works by preventing the flowers from opening. gov.[9] Most genes belong to larger families of genes of shared ancestry.[6] which can be an advantage since these viruses will evolve constantly and rapidly. by Brad M. ISBN 0 642 58336 6. each with a particular and independent function. eu/ research_themes/ topics188. these can either have no effect. that can be mixed together to produce genes encoding new proteins with novel properties. Barbour. rirdc. usually through genetic recombination. Potts.[14] Another advantage of duplicating a gene (or even an entire genome) is that this increases redundancy. Mutations are caused by radiation. organisms have evolved mechanisms such as DNA repair to remove mutations.[19] In evolution. by cellular processes such as hypermutation. and thereby preserving genetic differences between these populations. Robert C.[17] [18] Changes in chromosome number may involve even larger mutations. RIRDC Project No CPF . or by recombining parts of different genes to form new combinations with new functions. alter the product of a gene. org 236 • Su.[20] .3A. pdf) [11] http:/ / www. ISSN 1440-6845. such as deleterious mutations. the human eye uses four genes to make structures that sense light: three for color vision and one for night vision.[8] These duplications are a major source of raw material for evolving new genes. Hingston. (2003) "The Great Wall of China: a physical barrier to gene flow?. html [12] http:/ / www. Australian Government. H et al. and they retain these separate chromosomes.[13] For example. with tens to hundreds of genes duplicated in animal genomes every million years.[1] Therefore. the optimal mutation rate for a species is a trade-off between costs of a high mutation rate. with about 70 percent of these mutations having damaging effects. RIRDC Publication No 01/114. and the remainder being either neutral or weakly beneficial. viruses.[10] Novel genes are produced by several methods. or prevent the gene from functioning properly or completely. as well as errors that occur during meiosis or DNA replication. coextra. Mutation can result in several different types of change in DNA sequences. transcontainer. transposons and mutagenic chemicals. this allows one gene in the pair to acquire a new function while the other copy performs the original function. the human immune system. For example.[11] [12] Here. and the metabolic costs of maintaining systems to reduce the mutation rate. September 2001. Volume 9 Pages 212-219 Mutation Mutations are changes in a genomic sequence: the DNA sequence of a cell's genome or the DNA or RNA sequence of a virus. two chromosomes in the Homo genus fused to produce human chromosome 2.[7] Description Mutations can involve large sections of DNA becoming duplicated.[4] Due to the damaging effects that mutations can have on cells. Rural Industrial Research and Development Corporation (http:/ / www. where segments of the DNA within chromosomes break and then rearrange. Andrew B. org/ UK [13] http:/ / sigmea. Joint Venture Agroforestry Program. all four arose from a single ancestral gene. domains act as modules.g. and thus evade the defensive responses of e. this fusion did not occur in the lineage of the other apes. commonly through the duplication and mutation of an ancestral gene. dyndns." Heredity.[5] Viruses that use RNA as their genetic material have rapid mutation rates. such as DNA repair enzymes. the most important role of such chromosomal rearrangements may be to accelerate the divergence of a population into new species by making populations less likely to interbreed.Gene flow [10] Genetic Pollution from Farm Forestry using eucalypt species and hybrids.[1] [2] [3] They can also be induced by the organism itself. Studies in the fly Drosophila melanogaster suggest that if a mutation changes a protein produced by a gene. this will probably be harmful. au/ reports/ AFT/ 01-114.[15] [16] Other types of mutation occasionally create new genes from previously noncoding DNA. A report for the RIRDC/L&WA/FWPRDC. DNA repair mechanisms are able to mend most changes before they become permanent mutations. This is a somatic mutation that may also be passed on in the germ line. For example. If this color change is advantageous. mutations can be subdivided into germ line mutations. The majority of these mutations will have no effect. making it harder (or easier) for predators to see. The source of the mutation is unrelated to the consequence. For example. and over time the number of butterflies with this mutation may form a larger percentage of the population. in less fit offspring.[22] Another effect of these mobile DNA sequences is that when they move within a genome. while other "more favorable" mutations may accumulate and result in adaptive evolutionary changes. the chance of this butterfly surviving and producing its own offspring are a little better. These can accumulate over time due to genetic drift. although the consequences are related to which cells were mutated. more than a million copies of the Alu sequence are present in the human genome. It is believed that the overwhelming majority of mutations have no significant effect on an organism's fitness. Neutral mutations are defined as mutations whose effects do not influence the fitness of an individual.[2] In multicellular organisms with dedicated reproductive cells. and somatic mutations (also called acquired mutations)[23] . but one might change the color of one of the butterfly's offspring. they can mutate or delete existing genes and thereby produce genetic diversity. A mutation has caused this garden moss rose to produce flowers of different colors. that tend to die out. . a butterfly may produce offspring with new mutations. and many organisms have mechanisms for eliminating otherwise permanently mutated somatic cells. Also. Mutation is generally accepted by biologists as the mechanism by which natural selection acts. make up a major fraction of the genetic material of plants and animals. and may have been important in the evolution of genomes.[21] For example. such as transposons. The abundance of some genetic changes within the gene pool can be reduced by natural selection. and these sequences have now been recruited to perform functions such as regulating gene expression. which can be passed on to descendants through their reproductive cells. generating advantageous new traits that survive and multiply in offspring as well as disadvantageous traits. 237 Nonlethal mutations accumulate within the gene pool and increase the amount of genetic variation[24] . If the organism can reproduce asexually through mechanisms such as cuttings or budding the distinction can become blurred.Mutation Sequences of DNA that can move about the genome. plants can sometimes transmit somatic mutations to their descendants asexually or sexually where flower buds develop in somatically mutated parts of plants. A new mutation that was not inherited from either parent is called a de novo mutation. which involve cells outside the dedicated reproductive group and which are not usually transmitted to descendants. BrdU) • Alkylating agents (e. 5-methylcytosine. and 5MeC (5-methylcytosine) → T. Each of these classes of chemical mutagens has certain effects that then lead to transitions. Examples include C → U and A → HX (hypoxanthine). UV radiation. N-ethyl-N-nitrosourea) These agents can mutate both replicating and non-replicating DNA. Mutation rates also vary across species. a base analog can only mutate the DNA when the analog is incorporated in replicating the DNA. A hotspot can be at an unusual base. • Slipped strand mispairing . • Depurination – Loss of a purine base (A or G) to form an apurinic site (AP site). • Agents that form DNA adducts (e. Spontaneous mutations on the molecular level can be caused by: • Tautomerism – A base is changed by the repositioning of a hydrogen atom. where mutations occur up to 100 times more frequently than the normal mutation rate. ethidium bromide) • DNA crosslinkers • Oxidative damage • Nitrous acid converts amine groups on A and C to diazo groups. ochratoxin A metabolites)[26] • DNA intercalating agents (e. the major mutagen in [25] tobacco smoke. which is less likely to be detected as a mutation because thymine is a normal DNA base. • Deamination – Hydrolysis changes a normal base to an atypical base containing a keto group in place of the original amine group. This can lead to insertions or deletions. • Ionizing radiation • Viral infections[28] DNA has so-called hotspots. UV light can induce adjacent pyrimidine bases in a DNA strand to become covalently joined as a pyrimidine dimer.g. which can be corrected by DNA repair mechanisms. followed by renaturation in a different spot ("slipping"). can also cause oxidative damage to DNA[27] .g. e.. and DNA • Ultraviolet radiation (nonionizing radiation). altering the hydrogen bonding pattern of that base resulting in incorrect base pairing during replication.Mutation 238 Causes Two classes of mutations are spontaneous mutations (molecular decay) and induced mutations caused by mutagens.g. or deletions. particularly longer-wave UVA. Two nucleotide bases in DNA – cytosine and thymine – are most vulnerable to radiation that can change their properties.g. In contrast.Denaturation of the new strand from the template during replication. altering their hydrogen bonding patterns which leads to incorrect base pairing during replication. transversions. because they allow organisms to evolve and therefore adapt more quickly to their . Induced mutations on the molecular level can be caused by: • Chemicals • Hydroxylamine NH2OH • Base analogs (e. • Radiation A covalent adduct between benzo[a]pyrene. Evolutionary biologists have theorized that higher mutation rates are beneficial in some situations.g. exchange a single nucleotide for another[30] . 239 Classification of mutation types By effect on structure The sequence of a gene can be altered in a number of ways. For example. repeated exposure of bacteria to antibiotics. and selection of resistant mutants.Mutation environments. An example of a transversion is adenine (A) being converted into a cytosine (C). base mis-pairing. can result in the selection of bacteria that have a much higher mutation rate than the original population (mutator strains). or mutagenic base analogs such as 5-bromo-2-deoxyuridine (BrdU). Most common is the transition that exchanges a purine for a purine (A ↔ G) or a pyrimidine for a pyrimidine. including: • Point mutations. A transition can be caused by nitrous acid. Point mutations that occur within the protein coding region of a gene may be classified into three kinds. . which exchanges a purine for a pyrimidine or a pyrimidine for a purine (C/T ↔ A/G). in a standard table of the genetic code of [29] amino acids. such as those affecting a small gene in one or a few nucleotides. Less common is a transversion. Gene mutations have varying effects on health depending on where they occur and whether they alter the function of essential proteins. in which the nucleotide is changed back to its original state (true reversion) or by second-site reversion (a complementary mutation elsewhere that results in regained gene functionality). These changes are classified as transitions or transversions[31] . (C ↔ T). Mutations in the structure of genes can be classified as: • Small-scale mutations. A point mutation can be reversed by another point mutation. Selection of disease-causing mutations. often caused by chemicals or malfunction of DNA replication. depending upon what the erroneous codon codes for: Illustrations of five types of chromosomal mutations. increasing the dosage of the genes located within them. the defective glycoprotein product of the fibrillin gene (FBN1) antagonizes the product of the normal allele. including: • Amplifications (or gene duplications) leading to multiple copies of all chromosomal regions. • A back mutation or reversion is a point mutation that restores the original sequence and hence the original phenotype. When the allele has a complete loss of function (null allele) it is often called an amorphic mutation. For example. a type of brain tumor. • Interstitial deletions: an intra-chromosomal deletion that removes a segment of DNA from a single chromosome. • Large-scale mutations in chromosomal structure. potentially bringing together separate genes to form functionally distinct fusion genes (e. Marfan syndrome is an example of a dominant negative mutation occurring in an autosomal dominant disease. In humans. They are usually caused by transposable elements. These include: • Chromosomal translocations: interchange of genetic parts from nonhomologous chromosomes. These mutations usually have dominant phenotypes. AT repeats). leading to loss of the genes within those regions. were found to have a chromosomal deletion removing sequences between the "fused in glioblastoma" (fig) gene and the receptor tyrosine kinase "ros". • Mutations whose effect is to juxtapose previously separate pieces of DNA. These mutations usually result in an altered molecular function (often inactive) and are characterised by a dominant or semi-dominant phenotype. • Nonsense mutations: which code for a stop and can truncate the protein. in an organism that previously had two different alleles. cells isolated from a human astrocytoma. either by a deletion or recombination event. transposable elements able to revert a very short deletion (say 1–2 bases) in any location are either highly unlikely to exist or do not exist at all. • Deletions remove one or more nucleotides from the DNA. • Chromosomal inversions: reversing the orientation of a chromosomal segment. both of which can significantly alter the gene product.g. or when the reduced dosage of a normal gene product is not enough for a normal phenotype (this is called haploinsufficiency). Like insertions. Note that a deletion is not the exact opposite of an insertion: the former is quite random while the latter consists of a specific sequence inserting at locations that are not entirely random or even quite narrowly defined. or cause a shift in the reading frame (frameshift). or errors during replication of repeating elements (e. thereby apposing previously distant genes. bcr-abl).g. Insertions in the coding region of a gene may alter splicing of the mRNA (splice site mutation). Exceptions are when the organism is haploid.[32] . 240 By effect on function • Loss-of-function mutations are the result of gene product having less or no function. • Deletions of large chromosomal regions. producing a fusion protein (FIG-ROS). • Missense mutations: which code for a different amino acid. • Insertions add one or more extra nucleotides into the DNA. • Lethal mutations are mutations that lead to the death of the organisms which carry the mutations. • Gain-of-function mutations change the gene product such that it gains a new and abnormal function. Insertions can be reverted by excision of the transposable element. Phenotypes associated with such mutations are most often recessive. The abnormal FIG-ROS fusion protein has constitutively active kinase activity that causes oncogenic transformation (a transformation from normal cells to cancer cells). They are generally irreversible: though exactly the same sequence might theoretically be restored by an insertion. • Loss of heterozygosity: loss of one allele. • Dominant negative mutations (also called antimorphic mutations) have an altered gene product that acts antagonistically to the wild-type allele.Mutation • Silent mutations: which code for the same amino acid. Often called a neomorphic mutation. these mutations can alter the reading frame of the gene. In this condition. Due to the triplet nature of gene expression by codons. • A homozygous mutation is an identical mutation of both the paternal and maternal alleles. or the grouping of the codons. sickle-cell disease. In reality. genetic drift is the basis for most variation at the molecular level. a change from AAA to AGA will encode lysine. but chemically similar. and thus increases the fitness of the organism. In the neutral theory of molecular evolution. resulting in a completely different translation from the original. p. 39). • Missense mutations or nonsynonymous mutations are types of point mutations where a single nucleotide is changed to cause substitution of a different amino acid. viewing the fitness effects of mutations in these discrete categories is an oversimplification. the more altered the protein produced is. the current distribution is still uncertain. However. • non inheritable somatic (e. This in turn can render the resulting protein nonfunctional. For example. or a nonsense codon in the transcribed mRNA.[34] • A wildtype or homozygous non-mutated organism is one in which neither allele is mutated. and possibly a truncated. • A harmful mutation is a mutation that decreases the fitness of the organism. and SOD1 mediated ALS (Boillée 2006. • A deleterious mutation has a negative effect on the phenotype.[33] By inheritance • inheritable generic in pro-generic tissue or cells on path to be changed to gametes.. Attempts have been made to infer the distribution of fitness effects using mutagenesis experiments or theoretical models applied to molecular sequence data. • A heterozygous mutation is a mutation of only one allele. The similarity between the two is enough that little or no change is often rendered in the protein. In theoretical population genetics. Such mutations are responsible for diseases such as Epidermolysis bullosa. The earlier in the sequence the deletion or insertion occurs. • A nonsense mutation is a point mutation in a sequence of DNA that results in a premature stop codon. although most nearly neutral mutations are slightly deleterious. • An advantageous mutation has a positive effect on the phenotype. • A nearly neutral mutation is a mutation that may be slightly deleterious or advantageous. or which promotes traits that are desirable. and thus decreases the fitness of the organism. it is more usual to speak of such mutations as deleterious or advantageous. • A neutral mutation is a mutation that occurs in an amino acid codon which results in the use of a different. (Just not a mutation) By impact on protein sequence • A frameshift mutation is a mutation caused by insertion or deletion of a number of nucleotides that is not evenly divisible by three from a DNA sequence. amino acid. carcinogenic mutation) • non inheritable post mortem aDNA mutation in decaying remains. By pattern of inheritance The human genome contains two copies of each gene – a paternal and a maternal allele.Mutation 241 By effect on fitness In applied genetics it is usual to speak of mutations as either harmful or beneficial.g. • A beneficial mutation is a mutation that increases fitness of the organism. and often nonfunctional protein product. forming the basis for the molecular clock. the insertion or deletion can disrupt the reading frame. a chemically similar molecule to the . • Compound heterozygous mutations or a genetic compound comprises two different mutations in the paternal and maternal alleles. Such mutations occur at a steady rate. • A neutral mutation has no harmful or beneficial effect on the organism. and some aspects of the distribution likely vary between species. To function correctly. Note that for mutations in RNA. the adenine at the 76th position was replaced by a thymine. For example. and thus cause cancer[37] . ΔF508) – The Greek letter Δ (delta) indicates a deletion. or they may occur within a codon in a manner that does not alter the final amino acid sequence. each cell depends on thousands of proteins to function in the right places at the right times. Harmful mutations Changes in DNA caused by mutation can cause errors in protein sequence.g. • Amino acid deletion (e. thereby leading to a changed protein.100g>c if the mutation occurred in RNA. A condition caused by mutations in one or more genes is called a genetic disorder. Nomenclature Nomenclature of mutations specify the type of mutation and base or amino acid changes.100G>C if the mutation occurred in mitochondrial DNA.[35] However. • Nucleotide substitution (e. D111E) – The first letter is the one letter code of the wild type amino acid. a mutation may occur in a somatic cell of an organism. the first letter represents the wild type nucleotide. this will probably be harmful.g. In the given example. a silent mutation in the exon/intron border may lead to alternative splicing by changing the splice site (see Splice site mutation). and the remainder being either neutral or weakly beneficial. The letter refers to the amino acid present in the wild type and the number is the position from the N terminus of the amino acid were it to be present as in the wild type. a medical condition can result. They may occur in a region that does not code for a protein. occurring only within exons.The number is the position of the nucleotide from the 5' end. and the second letter represents the nucleotide which replaced the wild type. For example. 76A>T) . • Silent mutations are mutations that do not result in a change to the amino acid sequence of a protein. however. synonymous mutations are a subcategory of the former. When a mutation alters a protein that plays a critical role in the body. Nonsense mutations are represented with an X for the second amino acid (e. mitochondrial DNA. it can give rise to offspring that carries the mutation in all of its cells. the nucleotide code is written in lower case.g. studies in yeast have shown that only 7% of mutations that are not in genes are harmful. D111X). if the 100th base of a nucleotide sequence mutated from G to C. Studies of the fly Drosophila melanogaster suggest that if a mutation does change a protein. The name silent could be a misnomer.g. The phrase silent mutation is often used interchangeably with the phrase synonymous mutation. This is the case in hereditary diseases. with about 70 percent of these mutations having damaging effects. Some mutations alter a gene's DNA base sequence but do not change the function of the protein made by the gene.Mutation intended arginine.[36] If a mutation is present in a germ cell. and the second letter is the one letter code of the amino acid present in the mutation. but might have no deleterious consequences at a lower temperature (permissive condition). On the other hand. For example. a simple convention is used. • Amino acid substitution (e. Such mutations will be present in all descendants of this cell within the same organism. or r. • If it becomes necessary to differentiate between mutations in genomic DNA. m. creating partially or completely non-functional proteins. and RNA. the number is the position of the amino acid from the N terminus. .100G>C if the mutation occurred in genomic DNA. 242 Special classes • Conditional mutation is a mutation that has wild-type (or less severe) phenotype under certain "permissive" environmental conditions and a mutant phenotype under certain "restrictive" conditions. then it would be written as g. a temperature-sensitive mutation can cause cell death at high temperature (restrictive condition). and certain mutations can cause the cell to become malignant. One possible explanation of the etiology of the relatively high frequency of CCR5-Δ32 in the European population is that it conferred resistance to the bubonic plague in mid-14th century Europe. One-third of all indigenous inhabitants of Sub-Saharan Africa carry the gene[41] . thus its frequency in the population increased. 243 Beneficial mutations Although most mutations that change protein sequences are neutral or harmful. Because DNA can be damaged or mutated in many ways. is Sickle cell disease which is a blood disorder in which the body produces an abnormal type of the oxygen-carrying substance hemoglobin in the red blood cells. where the bubonic plague never reached. there is a survival value in carrying only a single sickle-cell gene (sickle cell trait).[40] Another example. gene mutations that could cause a genetic disorder are repaired by the DNA repair system of the cell. the process of DNA repair is an important way in which the body protects itself from disease. In this case.Mutation Often.[39] This theory could explain why this mutation is not found in southern Africa.[42] Those with only one of the two alleles of the sickle-cell disease are more resistant to malaria. Each cell has a number of pathways through which enzymes recognize and repair mistakes in DNA. the mutation may enable the mutant organism to withstand particular environmental stresses better than wild-type organisms. or reproduce more quickly. For example. Prion mutation Prions are proteins and do not contain genetic material. People with this mutation were more likely to survive infection. some mutations have a positive effect on an organism. However.[38] The CCR5 mutation is more common in those of European descent. a specific 32 base pair deletion in human CCR5 (CCR5-Δ32) confers HIV resistance to homozygotes and delays AIDS onset in heterozygotes. since the infestation of the malaria plasmodium is halted by the sickling of the cells which it infests.[43] See also • • • • • • • • • • • • • • Aneuploidy Antioxidant Budgerigar colour genetics Homeobox Macromutation Muller's morphs Mutant Mutagenesis Polyploidy Robertsonian translocation Signature tagged mutagenesis Site-directed mutagenesis TILLING (molecular biology) Trinucleotide repeat expansion . A newer theory suggests that the selective pressure on the CCR5 Delta 32 mutation was caused by smallpox instead of the bubonic plague. In these cases a mutation will tend to become more common in a population through natural selection. prion replication has been shown to be subject to mutation and natural selection just like other forms of replication. because in areas where malaria is common. "The evolutionary mechanics of domain organization in proteomes and the rise of modularity in the protein world". [17] Liu N. "UVA radiation is highly mutagenic in cells that are unable to repair 7. [10] Harrison P.24. PMC 1131864.008.11.A..4.13910. nih. Hartl DL (2007). doi:10. doi:10. Rev.1073/pnas. Acad. PMID 15851677. Grabau E. doi:10. Elie C.S. [3] Burrus V. Horodyski F. pubmedcentral. PMC 2864001. pubmedcentral. "Prevalence of positive selection among nearly neutral amino acid replacements in Drosophila" (http:/ / www. Podlaha O (2004). 102 (38): 13538–43. 2 (7): E206. "Evolution of colour vision in vertebrates". Nat. PMID 19597530. "The distribution of fitness effects of new mutations". Werren JH (2001). [2] Aminetzach YT. fcgi?tool=pmcentrez& artid=1636486). "Ochratoxin A: An overview on toxicity and carcinogenicity in animals and humans". "The role of selfish genetic elements in eukaryotic evolution". [21] Hurst GD. 6 (8): 2977–83. Royal A (1 August 1986). PMID 15123584.1038/35084545. Mol.1038/nrg2146. "Mutation rates among RNA viruses" (http:/ / www.0701572104. . full).1101/gr. Genetics 10 (8): 551–564. Natl.1073/pnas.0504497102. Bioessays 22 (12): 1057–66. gov/ articlerender. [8] Hastings. PMID 14634634.1891104.1038/nrg1204. nih.74.1016/j. "Alu elements as regulators of gene expression" (http:/ / www. Eye (London. Cell. nature. . org/ content/ 19/ 10/ 1693. PMC 24164. ISSN 1471-0056. doi:10. 34 (19): 5491–7. doi:10. gov/ articlerender. org/ pdb/ cgi/ explore. 155 (5): 376–86. Res.0501847102.1101/gr. [18] Siepel A (October 2009). Cell Res.1016/j. From DNA to Diversity: Molecular Genetics and the Evolution of Animal Design.1038/cr.317 (inactive 2009-11-14). Retrieved 2010-06-06. P J. U.1002/mnfr. Langelier Y. Hebert PD (1999). org/ cgi/ pmidlookup?view=long& pmid=16157879).str. Tyler DM (2008). [16] Hurles M (July 2004). "The modulation of DNA content: proximate causes and ultimate consequences" (http:/ / genome. html). pnas. nih. PMID 11173079. [28] Pilon L. Acad.082803. doi:10. Strub K (2006). Thornton JM (2005). Sci. Genet. Betrán E. "Protein families and their evolution-a structural perspective". A. doi:10. [23] "Genome Dictionary" (http:/ / www. [6] Drake JW. Shaver A (2000). Biochem. "Testing the chromosomal speciation hypothesis for humans and chimpanzees" (http:/ / www. "Shaping bacterial genomes with integrative and conjugative elements". Boiteux S.S. doi:10. [12] Long M.A. Acad. "Gene duplication: the genomic trade in spare parts" (http:/ / www. Microbiol. "The origin of new genes: glimpses from the young and old". Macpherson JM. P. Natl. "Chromosome speciation: humans. Nucleic Acids Res. Wang W (November 2003). [14] Bowmaker JK (1998). PLoS Biol. Manderville RA (January 2007). gov/ articlerender. Genome Res. full). PMID 10570172. Aspects Med. doi:10. Keightley.1016/S0022-2836(02)00109-2.pbio. PMID 15252449. doi:10.278. pubmedcentral. org/ content/ 96/ 24/ 13910.1038/nrg2593. 18 (10): 985–96. doi:10. Slezak G.1016/S0098-2997(00)00007-8. PMID 15207870. 4 (11): 865–75. Gerrish P. doi:10. cshlp. Sci. ISBN 1-4051-1950-0. PMID 19797681. [19] Zhang J. 19 (10): 1693–5. long).Mutation 244 References [1] Bertram J (2000). doi:10. .1073/pnas.1126/science.1126/science. nih. PMID 17409186. 74: 867–900.0020206.2008. [11] Orengo CA. Parsch J.2008. . PMID 10207154. doi:10. doi:10. Genetics 8 (8): 610–618. [9] Carroll SB.098376. Genet.012. 1/ 6535. "Studying genomes through the aeons: protein families. PMID 11084621. "Rapid evolution of RNA genomes".A.8-dihydro-8-oxoguanine in Saccharomyces cerevisiae" (http:/ / www. (Aug 2007). Weatherbee SD (2005). doi:10. gov/ articlerender. org/ content/ 9/ 4/ 317. "The evolution and functional diversification of animal microRNA genes" (http:/ / www.200600137. G (2009). PMID 16051794. de Rycke Y.01. PMC 1871816. Grenier J. doi:10. asm.7041255. doi:10. PMID 17637733. Johnson T.1112699. [22] Häsler J. org/ cgi/ pmidlookup?view=long& pmid=3023954). Proc. doi:10. [27] Kozmin S. Natl. Nature Reviews. Wang X. doi:10. England) 12 (Pt 3b): 541–7. [24] Eyre-Walker.1073/pnas. fcgi?tool=pmcentrez& artid=1871816). org/ content/ 102/ suppl. [7] Holland J. VandePol S (1982). PMID 17195275. "The molecular biology of cancer". Sci..S. PMID 11483984.1146/annurev. Okamura K.133029. "The evolution of mutation rates: separating causes from consequences". Nat. [4] Sawyer SA. fcgi?tool=pmcentrez& artid=2864001). [15] Gregory TR. Waldor M (2004). Spindler K. cgi?pdbId=1JDG) [26] Pfohl-Leszkowicz A. PMC 2712117. Thornton K. Sage E (September 2005). PMID 19141283. JR.1093/nar/gkl706. Lupski. fcgi?tool=pmcentrez& artid=449868). pnas. .S. Nature reviews. 9 (4): 317–24.2-W. 2 (8): 597–606. Reynaud-Angelin A. [5] Sniegowski P. PMID 9775215.9. PMC 1636486.2004. Sci. theodora. Mol Nutr Food Res 51 (1): 61–99. PMC 479111. Oxford: Blackwell Publishing. Drosophila. Proc. gov/ articlerender. Science 215 (4540): 1577–85. Gerstein M (2002). Structure 17 (1): 66–78. Petrov DA (2005).1101/gr. Biol. Rev. .96. Acad. doi:10. Proc. pubmedcentral. PMC 1224634. [20] Ayala FJ.A. . SM.1002/1521-1878(200012)22:12<1057::AID-BIES3>3. Zhang Z. doi:10. PMID 7041255. PMC 449868. PMID 18711447. . J Mol Biol 318 (5): 1155–74. 21 (6): 167–223.1371/journal. full).CO.biochem. and mosquitoes" (http:/ / www. PMID 16157879. PMC 2765273. pseudogenes and proteome evolution". pnas. Coluzzi M (2005). 102 (Suppl 1): 6535–42.resmic. Nichol S. fcgi?tool=pmcentrez& artid=479111). nih. rcsb. pubmedcentral. "Darwinian alchemy: Human genes from noncoding DNA" (http:/ / genome. PMID 12083509. 96 (24): 13910–3. Science 309 (5735): 764–7. Natl. 14 (5): 845–51. PMID 15954844. U. "Mechanisms of change in gene copy number" (http:/ / www. [25] Created from PDB 1JDG (http:/ / www. U. Genome Res.109. Genome Res. Second Edition.0. U. com/ cr/ journal/ v18/ n10/ full/ cr2008278a. com/ genetics/ #somaticcellgeneticmutation). doi:10. Ira. Rosenberg. 104 (16): 6504–10. Caetano-Anollés G (2009). Mol. Holland JJ (1999). "Pesticide resistance via transposition-mediated adaptive gene truncation in Drosophila". "Herpes simplex virus type 2 mutagenesis: characterization of mutants induced at the hprt locus of nonpermissive XC cells" (http:/ / mcb. Annu. [13] Wang M. cshlp. Proc. Rev. PMID 17020921. org/ content/ 98/ 18/ 10214. ac. U. de/ link/ service/ journals/ 00439/ bibs/ 1108002/ 11080167. bbc.S. pdf). [31] Freese. Hartl DL (2007). . Ernst (April 1959). Acad. Swain D.net/~rwms/EvoMutations. doi:10.2435085100.wikibooks. PMID 11281456.1371/journal. gov/ articlerender. 45 (4): 622–33. PMC 1871816.A. PLoS Genet. PMID 8505985. . full). doi:10. doi:10.co. "Ubiquitous somatic mutations in simple repeated sequences reveal a new mechanism for colonic carcinogenesis". Malkhosyan S.4. [39] "PBS:Secrets of the Dead. Slatkin M (2003).com (http:/ / www.1007/s004390000447.md (http:/ / sicklecell. 4 (8): e1000183. asp?articlekey=33675) [35] Sawyer SA. [34] Medterms. pbs. 1: 87–105. html). J.1016/S0022-2836(59)80038-3. gov/ articlerender.0701572104.181325198.1038/nrg2146. "The Specific Mutagenic Effect of Base Analogues on Phage T4". "Evaluating plague and smallpox as historical selective pressures for the CCR5-Δ32 HIV-resistance allele" (http:/ / www. pnas. doi:10. PMC 222607. "A catalog of neutral and deleterious polymorphism in yeast" (http:/ / www. Keightley PD (August 2007). htm). Sci. medterms. [29] References for the image are found in Wikimedia Commons page at: Commons:File:Notable mutations. "The Difference between Spontaneous and Base-Analogue Induced Mutations of Phage T4" (http:/ / www. nih. bbc. Nature 8 (8): 610–8. Nature 363 (6429): 558–61. [32] Ellis NA.1073/pnas.1038/363558a0.S. Perucho M (1993). PMC 2515631. [36] Doniger SW. PMC 56941. lifesci.stanford. (2001). nih. [40] Galvani A. fcgi?tool=pmcentrez& artid=299980). PMID 17409186.in/variationcentral) • The mutations chapter of the WikiBooks General Biology textbook (http://en. Natl.pgen.html) from the Huntington's Disease Outreach Project for Education at Stanford • Central Locus Specific Variation Database at the Institute of Genomics and Integrative Biology (http://miracle. doi:10. nih.45. PMID 18769710. doi:10.1073/pnas. Zhang Z. igib. pubmedcentral. [38] Sullivan. . "The distribution of fitness effects of new mutations" (http:/ / www. .Mutation PMID 3023954. org/ wnet/ secrets/ previous_seasons/ case_plague/ clues. com/ script/ main/ art. PNAS 95 (18): 10214–10219. "The coreceptor mutation CCR5Δ32 influences the dynamics of HIV epidemics and is selected for by HIV" (http:/ / www. doi:10. Parsch J. fcgi?tool=pmcentrez& artid=1871816). Hum Genet 108 (2): 167–73.ram)]] . German J (2001).html) • Correcting mutation by gene therapy (http://www. springer. PMID 17637733.org/wiki/ General_Biology/Genetics/Mutation#Mutation) • Examples of Beneficial Mutations (http://www.genetherapynet.A. Biol. PMID 16590424. sussex. PMID 14645720. doi:10. "Prevalence of positive selection among nearly neutral amino acid replacements in Drosophila" (http:/ / www. Ciocci S. Shibata D. (August 2008). uk/ 1/ hi/ health/ 8435320. md/ faq. Sci. [33] Eyre-Walker A. gov/ articlerender. Ernst (1959). nih.md FAQ (http:/ / sicklecell.1073/pnas.edu/causes/mutation/q0. fcgi?tool=pmcentrez& artid=222607). Kim HS. doi:10. Amy D. et al.uk/radio4/history/inourtime/rams/inourtime_20071206. pubmedcentral. md) [42] Sicklecell.res. gov/ articlerender. PMC 367868. asp#q1): "Why is Sickle Cell Anaemia only found in Black people? [43] 'Lifeless' prion proteins are 'capable of evolution' (http:/ / news.1000183.svg#References. Natl.GENETIC MUTATION . [37] Ionov Y. . co. Case File: Mystery of the Black Death" (http:/ / www. 104 (16): 6504–10. fcgi?tool=pmcentrez& artid=2515631). [41] Sicklecell. pubmedcentral. Acad. "Back mutation can produce phenotype reversion in Bloom syndrome somatic cells" (http:/ / link.streaming audio . [30] Freese. uk/ home/ Adam_Eyre-Walker/ Website/ Publications_files/ EWNRG07. Mol.gate. stm) 245 External links • "All About Mutations" (http://hopes.com) • BBC Radio 4 In Our Time . Peinado MA. Proc. pubmedcentral.622. Proc. Proc Natl Acad Sci USA 100 (25): 15276–9. PMC 299980.1073/pnas. et al. U. PMID 11517319.with [[Steve Jones (biologist)|Steve Jones (http://www. then there will be a slightly higher proportion of fast rabbits or efficient algae in the next Darwin's illustrations of beak variation in the generation. in natural selection the environment acts as a sieve through which only certain variations can pass. but some differences may improve the chances of survival of a particular individual. or the observable characteristics of an organism. The union of traditional Darwinian evolution with subsequent discoveries in classical and molecular genetics is termed the modern evolutionary synthesis. If the traits that give these individuals a reproductive advantage are also heritable. In this way the natural environment of an that beak shapes evolved by natural selection.[1] in which natural selection was described as analogous to artificial selection. at the time of Darwin's writing. Factors which affect reproductive success are also important. organism "selects" for traits that confer a reproductive advantage. Natural selection is one of the cornerstones of modern biology. suggesting exponential growth. due to species is suited to its preferred food. This effect was first described and named by Charles Darwin. A rabbit that runs faster than others may be more likely to escape from predators. this process can result in adaptations that specialize populations for particular ecological niches and may eventually result in the emergence of new species. Many of these differences do not affect survival (such as differences in eye color in humans). but during the industrial revolution many of the trees on which the moths rested became blackened by soot. Natural selection remains the primary explanation for adaptive evolution. This gave dark-colored moths a better chance of surviving to produce dark-colored offspring. The beak of each heritable advantage will become dominant in the population. passed from parent to child. natural selection is an important process (though not the only process) by which evolution takes place within a population of organisms. As opposed to artificial selection. This is known as differential reproduction. nothing was known of modern genetics. the peppered moth exists in both light and dark colors in the United Kingdom. Natural selection acts on the phenotype. and algae that are more efficient at extracting energy from sunlight will grow faster. The term was introduced by Darwin in his influential 1859 book On the Origin of Species. In other words.Natural selection 246 Natural selection Natural selection is the process by which traits become more or less common in a population due to consistent effects upon the survival or reproduction of their bearers. For example. but the genetic (heritable) basis of any phenotype which gives a reproductive advantage will become more common in a population (see allele frequency). Individuals that have better odds for survival also have better odds for reproduction. . causing gradual changes or evolution of life. Even if the finches of the Galápagos Islands. in which humans favor specific traits. that is. over many generations any in the shape of their beaks. The concept of natural selection was originally developed in the absence of a valid theory of heredity. an issue which Charles Darwin developed in his ideas on sexual selection. The natural genetic variation within a population of organisms may cause some individuals to survive and reproduce more successfully than others in their current environment. and in just a few generations the majority of the moths were dark. General principles Natural variation occurs among the individuals of any population of organisms. It is a key mechanism of evolution. a process by which animals and plants with traits considered desirable by human breeders are systematically favored for reproduction. Over time. giving the dark-colored moths an advantage in hiding from predators. which hold 13 closely related species that differ most markedly reproductive advantage is very slight. examples include many human genetic disorders like cystic fibrosis. natural selection is "blind" in the sense that changes in phenotype (physical and behavioral characteristics) can give a reproductive advantage regardless of whether or not the trait is heritable (non heritable traits can be the result of environmental factors or the life experience of the organism). the precise meaning of the term is much more subtle.Natural selection The concept of natural selection predates the understanding of genetics. scientists define "natural selection" specifically as "those mechanisms that contribute to the selection of individuals that reproduce". Modern evolutionary theory defines fitness not by how long an organism lives. because these are the traits that directly participate in evolution. This is the link between natural selection and genetics. Although a complete theory of evolution also requires an account of how genetic variation arises in the first place (such as by mutation and sexual reproduction) and includes other evolutionary mechanisms (such as gene flow). In modern times. natural selection is still understood as a fundamental mechanism for evolution. However. 247 Nomenclature and usage The term natural selection has slightly different definitions in different contexts. as with natural selection above. . which is the study of heredity. This is sometimes referred to as "phenotypic natural selection". and Richard Dawkins manages in his later books to avoid it entirely. it has high fitness value in populations which have high malaria infection rates. The fitness of a particular genotype corresponds to the average effect on all individuals with that genotype. Broadly. it is also possible that a favorable mutation arises in an individual that does not survive to adulthood for unrelated reasons. Selection for a trait may also result in the selection of other correlated traits that do not themselves directly influence reproductive advantage. to discussing the various senses in which the term is used). when this distinction is important. its genes will become more common in the adult population of the next generation. it is understood that selection acts on an organism's phenotype. Conditions like sickle-cell anemia may have low fitness in the general human population. The Extended Phenotype. The phenotype is the result of the genotype and the environment in which the organism lives (see Genotype-phenotype distinction).[2] [3] It is sometimes helpful to explicitly distinguish between selection's mechanisms and its effects. If an organism lives half as long as others of its species. individuals which are more "fit" have better potential for survival. without regard to whether the basis of the selection is heritable. Though natural selection acts on individuals. as described in the modern evolutionary synthesis. Following Darwin's primary usage[1] the term is often used to refer to both the evolutionary consequence of blind selection and to its mechanisms. but it is the organism's genetic make-up or genotype that is inherited. but by how successful it is at reproducing. as in the well-known phrase "survival of the fittest". whereas those that reduce success are selected against. It is most often defined to operate on heritable traits. This may occur as a result of pleiotropy or gene linkage. However. Since fitness is an averaged quantity. (He devotes a chapter of his book. but because the sickle-cell trait confers immunity from malaria. the effects of chance mean that fitness can only really be defined "on average" for the individuals within a population.[5] Fitness The concept of fitness is central to natural selection.[4] Traits that cause greater reproductive success of an organism are said to be selected for. Fitness also depends crucially upon the environment. or observable characteristics. Very low-fitness genotypes cause their bearers to have few or no offspring on average. but has twice as many offspring surviving to adulthood. as in cases of competition among individuals of the same sex in a population. intrasexual selection is often associated with sexual dimorphism. and hence the outcome of natural selection. aggression between members of the same sex is sometimes associated with very distinctive features. The unit of selection can be the individual or it can be another level within the hierarchy of biological organisation. including differences in body size between males and females of a species. in a process called intragenomic conflict.g. When individuals can reproduce more than once. The viability of produced gametes can differ. Finally. a longer survival in the reproductive phase increases the number of offspring. and kin groups. called survival selection. intrasexual selection involves male–male competition and intersexual selection involves female choice of suitable males. The life cycle of a sexually reproducing organism. while "sexual selection" refers specifically to competition for mates. Sexual selection It is useful to distinguish between "ecological selection" and "sexual selection". An individual organism must survive until adulthood before it can reproduce. natural selection often results in the maintenance of the status quo by eliminating less fit variants. The fecundity of both females and males (for example. More generally. non-kin group. though likely examples have also been found in amphibian and bird species. and selection of those that reach this stage is called viability selection. some species exhibit sex-role reversed behavior in which it is males that are most selective in mate choice. and infanticide). and selective pressure can be produced by any aspect of the environment. the extravagant plumage of some male birds. giant sperm in certain species of Drosophila)[7] can be limited via "fecundity selection".Natural selection 248 Types of selection Natural selection can act on any phenotypic trait. kin selection. Selection at a different level such as the gene can result in an increase in fitness for that gene. competition. e.[10] . such as the antlers of stags. including sexual selection and competition with members of the same species.[9] Some features that are confined to one sex only of a particular species can be explained by selection exercised by the other sex in the choice of a mate. or intersexual. while at the same time reducing the fitness of the individuals carrying that gene. There is still debate about whether natural selection acts at the level of groups or species to produce adaptations that benefit a larger. Natural selection occurs at every life stage of an individual. However. such as genes. cells. this does not imply that natural selection is always directional and results in adaptive evolution.[8] Sexual selection can be intrasexual. while intragenomic conflicts such as meiotic drive between the haploid gametes can result in gametic or "genic selection". as in cases where one sex controls reproductive access by choosing among a population of available mates. Overall. due to the generally greater investment of resources for a female than a male in a single offspring. for example. the union of some combinations of eggs and sperm might be more compatible than others. this is termed compatibility selection. However. the combined effect of all selection pressures at various levels determines the overall fitness of an individual. the best-known examples of this pattern occur in some fishes of the family Syngnathidae. Most commonly. and success in this competition determines who will parent the next generation. which are used in combat with other stags. Similarly. Ecological selection covers any mechanism of selection as a result of the environment (including relatives. Various components of natural [6] selection are indicated for each life stage. In many species. adults must compete with each other for mates via sexual selection. most changes in the genetic material were considered neutral or close to neutral because they occurred in noncoding DNA or resulted in a synonymous substitution. However. shape. these individuals are more likely to survive when next confronted with that antibiotic. considerable variation in their genetic material. At the same time. and repeated exposure to the antibiotic. populations of bacteria are large enough that a few individuals will have beneficial mutations. A similar situation occurs with pesticide resistance in plants and insects. creating a new population of resistant bacteria.[14] [15] Although both mutation . size and internal arrangement of the chromosomes).[12] This is an example of what is known as an evolutionary arms race. Any of these changes might have an effect that is highly advantageous or highly disadvantageous. Due to the elimination of the maladapted individuals in the past generation. but some may have mutations that make them slightly less susceptible. stronger antibiotics. these individuals will survive the treatment. most bacteria die quickly. and advantageous mutations are even rarer. If a new mutation reduces their susceptibility to an antibiotic. among their vast numbers of individual members. Evolution by means of natural selection A prerequisite for natural selection to result in adaptive evolution. The widespread use and misuse of antibiotics has resulted in increased microbial resistance to antibiotics in clinical use. to the point that the methicillin-resistant Staphylococcus aureus (MRSA) has been described as a "superbug" because of the threat it poses to health and its relative invulnerability to existing drugs. Given enough time. is the presence of heritable genetic variation that results in fitness differences. recombinations and alterations in the karyotype (the number. This selective elimination of maladapted individuals from a population is natural selection. When exposed to antibiotics. but large effects are very rare. it is not necessarily optimally adapted any more to the old antibiotic free environment. contributing new genetic variation to the existing genetic variation. This new changed population of antibiotic-resistant bacteria is optimally adapted to the context it evolved in. If the exposure to antibiotics is short. whose offspring then inherit the resistance. a well-documented example involves the spread of a gene in the butterfly Hypolimnas bolina suppressing male-killing activity by Wolbachia bacteria parasites on the island of Samoa. while both perform substandard in the other environment. Genetic variation is the result of mutations. where the spread of the gene is known to have occurred over a period of just five years [13] Resistance to antibiotics is increased though the survival of individuals which are immune to the effects of the antibiotic. Since the discovery of penicillin in 1928 by Alexander Fleming. At the same time. this population contains more bacteria that have some resistance against the antibiotic. a population of antibiotic-resistant bacteria will emerge. novel traits and speciation. while medical researchers continue to develop new antibiotics that can kill them. antibiotics have been used to fight bacterial diseases. Spontaneous mutations are very rare. primarily as the result of mutations. producing the next generation. in which bacteria continue to develop strains that are less susceptible to antibiotics. new mutations occur. These surviving bacteria will then reproduce again. new strains of MRSA have recently emerged that are resistant even to these drugs. recent research suggests that many mutations in non-coding DNA do have slight deleterious effects. Natural populations of bacteria contain. Arms races are not necessarily induced by man. In the past.[11] Response strategies typically include the use of different. The end result of natural selection is two populations that are both optimally adapted to their specific environment. however.Natural selection 249 Examples of natural selection A well-known example of natural selection in action is the development of antibiotic resistance in microorganisms. However. but not all. until they dominate the population. 2. mutations in regulatory genes result in non-viable zygotes. Speciation Speciation requires selective mating. In every generation. a change in the physical environment (geographic isolation by an extrinsic barrier) would follow number 1. 250 Some mutations occur in so-called regulatory genes. or 3. either because of differences in selection pressures on the different subgroups. be present in only one or other of two subgroups when they first become separated. one species of deer shifts location and therefore changes its "rut") for number 3. estimates from data in humans have found that a majority of mutations are slightly deleterious. Changes in these can have large effects on the phenotype of the individual because they regulate the function of many other genes. This peacock is an albino. Geographic isolation. which result in a reduced gene flow. natural selection will favor these phenotypes and the novel trait will spread in the population. Examples of nonlethal regulatory mutations occur in HOX genes in humans. enhancing these traits over successive generations. alleles which on average result in greater fitness become more abundant in the next generation. traits that have high fitness in one environmental context may be much less fit if environmental conditions change. As a result. each new generation will be enriched by the increasing abundance of alleles that contribute to those traits that were favored by selection. by chance. Most. is believed to retain function in photoperiod perception. If the selection forces remain the same for many generations. or because different mutations arise spontaneously in the different populations. also called evolutionary baggage. A lesser-known mechanism of speciation occurs via hybridization.Natural selection rates and average fitness effects of mutations are dependent on the organism.[16] By the definition of fitness. possibly resulting in a vestigial manifestation of the trait. A famous example of a vestigial structure. Over time. In many circumstances. which can result in a cervical rib[17] or polydactyly. Established traits are not immutable. producing a new spectrum of phenotypes. selection against albinos in nature is intense because they are easily spotted by predators or are unsuccessful in competition for mates. while individuals with lesser fitness are more likely to die early or fail to reproduce.e. which suggests that evolutionary change and particularly speciation typically happens quickly after interrupting long periods of stasis. new mutations and re-combinations arise spontaneously. while alleles which generally reduce fitness become rarer. For example. or because of founder effects – some potentially beneficial alleles may. individuals with greater fitness are more likely to contribute offspring to the next generation. well-documented in plants and occasionally observed in species-rich groups of animals such as cichlid fishes.[20] Such mechanisms of rapid speciation can reflect a mechanism of evolutionary change known as punctuated equilibrium. In the absence of natural selection to preserve such a trait. Selective mating can be the result of 1. or may be co-opted for other advantageous traits in a phenomenon known as preadaptation. while alleles with a lesser fitness disappear. these subgroups might diverge radically to become different species. Therefore. Temporal isolation. it will become more variable and deteriorate over time. an increase in the number of fingers or toes. X-ray of the left hand of a ten year old boy with polydactyly.[18] When such mutations result in a higher fitness.. a change in camouflage for number 2 or a shift in mating times (i.[19] The exuberant tail of the peacock is thought to be the result of sexual selection by females. Behavioral isolation. the eye of the blind mole rat. beneficial alleles become more and more abundant. . the apparently vestigial structure may retain a limited functionality. [27] Similar ideas were later expressed by Nasir al-Din Tusi[28] and Ibn Khaldun. As long as there is some variation between them. then fixation of these genes in the respective subgroups will lead to two reproductively isolated populations. and that only those creatures survive that manage to provide for themselves and reproduce successfully.[35] .Natural selection Genetic changes within groups result in increasing incompatibility between the genomes of the two subgroups. who argued that environmental factors influence animals to develop new characteristics to ensure survival. these will be two different species. While these forerunners had an influence on Darwinism.[32] This theory has come to be known as Lamarckism and was an influence on the anti-genetic ideas of the Stalinist Soviet biologist Trofim Lysenko. apparently randomly. Charles Darwin set out his theory of evolution by natural selection as an explanation for adaptation and speciation. virtually imperceptible changes in successive generations could produce consequences on the scale of differences between species. there will be an inevitable selection of individuals with the most advantageous variations. is preserved".[22] while related ideas were later refined by Aristotle.[24] [25] [26] Abu Rayhan Biruni described the idea of artificial selection and argued that nature works in much the same way. thus reducing gene flow between the groups. Until the early 19th century. and populations that evolve to be sufficiently different eventually become different species. 251 Historical development Pre-Darwinian theories Several ancient philosophers expressed the idea that nature produces a huge variety of creatures.[23] The struggle for existence was later described by Al-Jahiz. eventually causing transmutation of species. Early 19th century evolutionists such as Jean Baptiste Lamarck suggested the inheritance of acquired characteristics as a mechanism for evolutionary change. then differential reproductive success will lead to a progressive evolution of particular populations of a species.[34] The concept was simple but powerful: individuals best adapted to their environments are more likely to survive and reproduce. Lucretius. weak forces could act continuously over long periods of time to produce radical changes in the Earth's landscape. they later had little influence on the trajectory of evolutionary thought after Charles Darwin. but a negative effect when they occur together. well-known examples include Empedocles[21] and his intellectual successor. Gene flow will effectively cease when the distinctive mutations characterizing each subgroup become fixed. including Charles Darwin's grandfather Erasmus Darwin. The success of this theory raised awareness of the vast scale of geological time and made plausible the idea that tiny. The modern theory of natural selection derives from the work of Charles Darwin in the nineteenth century. However. According to the biological species concept. If the variations are inherited. As few as two mutations can result in speciation: if each mutation has a neutral or positive effect on fitness when they occur separately. He defined natural selection as the "principle by which each slight variation [of a trait]. if useful. the prevailing view in Western societies was that differences between individuals of a species were uninteresting departures from their Platonic idealism (or typus) of created kinds.[33] Darwin's theory In 1859. adaptive traits acquired by an organism during its lifetime could be inherited by that organism's progeny.[29] [30] Such classical arguments were reintroduced in the 18th century by Pierre Louis Maupertuis[31] and others. the theory of uniformitarianism in geology promoted the idea that simple. noted that population (if unchecked) increases exponentially whereas the food supply grows only arithmetically.[44] The fifth edition of On the Origin of Species published in 1869 included Spencer's phrase as an alternative to natural selection. This principle of preservation. owing to the high geometrical powers of increase of each species. I have called. Darwin regretted the use of the term "Natural Selection". with credit given: "But the expression often used by Mr. the Reverend Thomas Malthus. and by the work of a political economist. But if variations useful to any organic being do occur. causing an infinite diversity in structure. but had neither developed them nor presented them in notable scientific publications. Herbert Spencer introduced the term survival of the fittest. Lyell and Joseph Dalton Hooker decided (without Wallace's knowledge) to present his essay together with unpublished writings which Darwin had sent to fellow naturalists. Natural Selection. which became a popular summary of the theory. who in An Essay on the Principle of Population.[41] For Darwin and his contemporaries. and from the strong principle of inheritance they will tend to produce offspring similarly characterised. and On the Tendency of Species to form Varieties.[36] When Darwin read Malthus in 1838 he was already primed by his work as a naturalist to appreciate the "struggle for existence" in nature and it struck him that as population outgrew resources. for the sake of brevity. At the time. and is sometimes equally convenient. and Darwin believed that selection was likely only part of the story: "I am convinced that [it] has been the main. "favourable variations would tend to be preserved. thus inevitable limitations of resources would have demographic implications. in the same way as so many variations have occurred useful to man. to be advantageous to them. some thinkers enthusiastically embraced natural selection. natural selection remained controversial as a mechanism. which he called "artificial selection". but not exclusive means of modification. constitution. and on the Perpetuation of Varieties and Species by Natural Means of Selection was read to the Linnean Society announcing co-discovery of the principle in July 1858. a severe struggle for life. season."[45] Although the phrase is still often used by 252 . other mechanisms of evolution such as evolution by genetic drift were not yet explicitly formulated. preferring the term "Natural Preservation". Darwin was meticulous about gathering and refining evidence as his "prime hobby" before making his idea public. and unfavourable ones to be destroyed. assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life. I think it would be a most extraordinary fact if no variation ever had occurred useful to each being's own welfare. leading to a "struggle for existence".[42] a response that has been characterized as the single most significant impediment to the idea's acceptance. considering the infinite complexity of the relations of all organic beings to each other and to their conditions of existence.[39] Darwin thought of natural selection by analogy to how farmers select crops or livestock for breeding. after reading Darwin.Natural selection Darwin's ideas were inspired by the observations that he had made on the Beagle voyage. natural selection was essentially synonymous with evolution by natural selection. and this certainly cannot be disputed. In the 3rd edition of 1861 Darwin acknowledged that others — notably William Charles Wells in 1813. Once he had his theory "by which to work". or year. The result of this would be the formation of new species. if there be. After the publication of On the Origin of Species. then. at some age.[43] However."[37] Here is Darwin's own summary of the idea."[40] In a letter to Charles Lyell in September 1860.[38] Darwin published a detailed account of his evidence and conclusions in On the Origin of Species in 1859. Herbert Spencer. However. of the Survival of the Fittest. He was in the process of writing his "big book" to present his researches when the naturalist Alfred Russel Wallace independently conceived of the principle and described it in an essay he sent to Darwin to forward to Charles Lyell. organic beings vary at all in the several parts of their organisation. and Patrick Matthew in 1831 — had proposed similar ideas. educated people generally accepted that evolution had occurred in some form. partly because it was perceived to be too weak to explain the range of observed characteristics of living organisms. and I think this cannot be disputed. and habits. is more accurate. in his early manuscripts he referred to a 'Nature' which would do the selection. which can be found in the fourth chapter of the Origin: If during the long course of ages and under varying conditions of life. and partly because even supporters of evolution balked at its "unguided" and non-progressive nature. the struggle for existence.[53] Interpretation of natural selection as necessarily 'progressive'. Although the Austrian monk Gregor Mendel. which the economists celebrate as the highest historical achievement. he used the theory as a justification for policies of the Nazi state. Social and psychological theory The social implications of the theory of evolution by natural selection also became the source of continuing controversy. so different from each other. as well as broader sociopolitical positions now described as Social Darwinism. selection for toughness. where it remains today. in writings that he subsequently disowned. This claim inspired some of Darwin's most ardent supporters—and provoked the most profound opposition.[51] lay in its power to "dethrone some of the deepest and most traditional comforts of Western thought". by creating genetic diversity. Friedrich Engels. Wilhelm Roux. In recent years. supplied the raw material for natural selection: see Genetics and the Origin of Species). stochasticity at the molecular level generates diversity in cell types whereas cell interactions impose a characteristic order on the developing embryo. leading to increasing 'advances' in intelligence and civilisation.[48] Theodosius Dobzhansky (who established the idea that mutation. This synthesis cemented natural selection as the foundation of evolutionary theory. is the normal state of the animal kingdom". the father of modern genetics. was a contemporary of Darwin's. was used as a justification for colonialism and policies of eugenics. occurring at all levels.S. modern biologists avoid it because it is tautological if "fittest" is read to mean "functionally superior" and is applied to individuals rather than considered as an averaged quantity over populations. and . had a profound influence on 19th century thought. a German political philosopher and co-originator of the ideology of communism.[47] Sewall Wright (who elucidated the nature of selection and adaptation). heroism.B.[46] 253 Modern evolutionary synthesis Natural selection relies crucially on the idea of heredity. his work would lie in obscurity until the early 20th century. The radicalism of natural selection. along with those of Adam Smith and Karl Marx.. Ernst Mayr (who recognised the key importance of reproductive isolation for speciation: see Systematics and the Origin of Species)[50] and many others formed the modern evolutionary synthesis. Haldane (who introduced the concept of the "cost" of natural selection). it challenged long-standing beliefs in such concepts as a special and exalted place for humans in the natural world and a benevolent creator whose intentions were reflected in nature's order and design. in Germany in 1940. but it was developed long before the basic concepts of genetics.Natural selection non-biologists. He wrote ". Perhaps the most radical claim of the theory of evolution through natural selection is that "elaborately constructed forms.[2] J. Impact of the idea Darwin's ideas. Only after the integration of Darwin's theory of evolution with a complex statistical appreciation of Gregor Mendel's 're-discovered' laws of inheritance did natural selection become generally accepted by scientists. The work of Ronald Fisher (who developed the required mathematical language and The Genetical Theory of Natural Selection). a founder of modern embryology.[49] William Hamilton (who conceived of kin selection). In particular. wrote in 1872 that "Darwin did not know what a bitter satire he wrote on mankind when he showed that free competition.. according to Stephen Jay Gould. a modern version of this theory has been proposed by Jean-Jacques Kupiec. and dependent on each other in so complex a manner" evolved from the simplest forms of life by a few simple principles. Cell and molecular biology In the 19th century. However. from molecules to organs. Konrad Lorenz won the Nobel Prize in Physiology or Medicine in 1973 for his analysis of animal behavior in terms of the role of natural selection (particularly group selection). wrote a book entitled « Der Kampf der Teile im Organismus » (The struggle of parts in the organism) in which he suggested that the development of an organism results from a Darwinian competition between the parts of the embryo. According to this cellular Darwinism [52]. must be accomplished by some human institution. A typical example is that certain combinations of genes for eye color in humans which. first described in this form by Richard Dawkins[57] and subsequently expanded upon by philosophers such as Daniel Dennett as explanations for complex cultural activities. Alfred Lotka proposed that natural selection might be understood as a physical principle which could be described in terms of the use of energy by a system. and this state is stable over time. when all the organisms in a population share the same allele for a particular trait. a class of heuristic optimization algorithms known as genetic algorithms.[55] More recently.) . Phenotype is determined by an organism's genetic make-up (genotype) and the environment in which the organism lives. for instance. in default of selective factors. or physical characteristics.[61] For example..[62] identify optimal solutions by simulated reproduction and mutation of a population of solutions defined by an initial probability distribution. which is the basis of natural selection.[63] Such algorithms are particularly useful when applied to problems whose solution landscape is very rough or has many local minima. such as "soft" artificial life. Goldberg.[58] Extensions of the theory of natural selection to such a wide range of cultural phenomena have been distinctly controversial and are not widely accepted. the allele is said to be fixed in that population. (On the other hand.[60] a concept that was later developed by Howard Odum as the maximum power principle whereby evolutionary systems with selective advantage maximise the rate of useful energy transformation. pioneered by John Holland in the 1970s and expanded upon by David E. have been hypothesized to have similar origins as adaptations to the early environment in which modern humans evolved. the concept of memes – "units of cultural transmission". is the hypothesis that the human brain is adapted to acquire the grammatical rules of natural language. Natural selection acts on an organism's phenotype. natural selection acts on specific traits of an individual. each of these versions is known as an allele.[59] 254 Information and systems theory In 1922. When different organisms in a population possess different versions of a gene for a certain trait. or culture's equivalents of genes undergoing selection and recombination – has arisen. from specific cultural norms such as incest avoidance to broader patterns such as gender roles.[56] Other aspects of human behavior and social structures."[54] Others have developed ideas that human societies and culture evolve by mechanisms that are analogous to those that apply to evolution of species. work among anthropologists and psychologists has led to the development of sociobiology and later evolutionary psychology.Natural selection social utility. notably advanced in the early work of Noam Chomsky and later by Steven Pinker. give rise to the phenotype of blue eyes. if mankind. It is this genetic variation that underlies phenotypic traits.. Often. The principles of natural selection have inspired a variety of computational techniques. that simulate selective processes and can be highly efficient in 'adapting' entities to an environment defined by a specified fitness function. The racial idea as the basis of our state has already accomplished much in this respect. Genetic basis of natural selection The idea of natural selection predates the understanding of genetics. Such concepts are sometimes relevant in the study of applied thermodynamics. The most prominent such example. Genotype and phenotype See also: Genotype-phenotype distinction. is not to be ruined by domestication-induced degeneracy. and the terms phenotype and genotype are used narrowly to indicate these specific traits. including human consciousness. a field that attempts to explain features of human psychology in terms of adaptation to the ancestral environment. We now have a much better idea of the biology underlying heritability. By analogy to the action of natural selection on genes. Purifying selection results in functional genetic features.[68] [69] Selection and genetic variation A portion of all genetic variation is functionally neutral in that it produces no phenotypic effect or significant difference in fitness. which favors genotypes that depart from the average in either direction (that is.[64] 255 Directionality of selection When some component of a trait is heritable. balancing selection can occur through frequency-dependent selection. This can occur in diploid species (that is. selection will alter the frequencies of the different alleles. these genes can produce a continuum of possible phenotypic values. those that have two pairs of chromosomes) when heterozygote individuals. This process can continue until the allele is eliminated from the population. and can result in a bimodal distribution of trait values. This is called heterozygote advantage or overdominance.Natural selection Some traits are governed by only a single gene. Finally. where the fitness of one particular phenotype depends on the distribution of other phenotypes in the population. the genetic variation at these sites will be eliminated due to genetic drift. but most traits are influenced by the interactions of many genes.[65] However.[65] Directional selection occurs when a certain allele has a greater fitness than others. the opposite of overdominance). after a period with no new mutation. have a higher fitness than homozygote individuals that have two of the same alleles. At the same time. being conserved over time due to selective pressure against deleterious variants. particularly in the study of kin selection and the evolution of reciprocal altruism. who have different alleles on each chromosome at a single genetic locus. Mutation selection balance Natural selection results in the reduction of genetic variation through the elimination of maladapted individuals and consequently of the mutations that caused the maladaptation. A variation in one of the many genes that contributes to a trait may have only a small effect on the phenotype. It is directional selection that is illustrated in the antibiotic resistance example above. on the basis of its effect on allele frequencies. selection cannot directly affect the frequency of such variation. together. which lowers the frequency of alleles that have a deleterious effect on the phenotype – that is. such as protein-coding genes or regulatory sequences. Finally. a number of forms of balancing selection exist. which do not result in fixation. Consequently. resulting in a mutation-selection balance. which is a function of how unfavorable the mutation proves to be. but maintain an allele at intermediate frequencies in a population. The exact outcome of the two processes depends both on the rate at which new mutations occur and on the strength of the natural selection. or variants of the gene that produces the variants of the trait. produce organisms of lower fitness. of which the best-known example is the malarial resistance observed in heterozygous humans who carry only one copy of the gene for sickle cell anemia. Far more common is stabilizing selection (which is commonly confused with purifying selection[66] [67] ). changes in the mutation rate or the selection pressure will result in a different mutation-selection balance. new mutations occur. When genetic variation does not result in differences in fitness. resulting in an increase of its frequency. Maintenance of allelic variation can also occur through disruptive or diversifying selection. . The principles of game theory have been applied to understand the fitness distributions in these situations. the genetic variation at those sites will be higher than at sites where variation does influence fitness. As a result. This process can continue until the allele is fixed and the entire population shares the fitter phenotype. the hypothesis that this variation accounts for a large fraction of observed genetic diversity is known as the neutral theory of molecular evolution and was originated by Motoo Kimura. Selection can be divided into three classes. 1993) The Nature of Selection: Evolutionary Theory in Philosophical Focus University of Chicago Press ISBN 0-226-76748-5 . Proceedings of the 9th International Congress of Genetics. Sunderland. Massachusetts. the presence of a block of strong linkage disequilibrium might indicate that there has been a 'recent' selective sweep near the center of the block. and none of the haplotypes will dominate the population. This is called genetic hitchhiking. uk/ ): On the origin of species by means of natural selection. Selective sweeps occur when an allele becomes more common in a population as a result of positive selection. through this mechanism. However.. Background selection is the opposite of a selective sweep. Natural Selection in the Wild. Therefore. London. selection can have a strong influence on patterns of variation in the genome. uk/ content/ frameset?itemID=F373& viewtype=side& pageseq=2). During the formation of gametes. the chance that such a reshuffle occurs between two alleles depends on the distance between those alleles. Adaptation and Natural Selection. Princeton. Evolution 37:1210-26 Futuyma DJ (2005) Evolution. However. [4] Works employing or describing this usage: Lande R & Arnold SJ (1983) The measurement of selection on correlated characters. it does not produce clear blocks of linkage disequilibrium. producing a region in the genome of low overall variability. modern reprint Charles Darwin. ISBN 0-691-00057-3. New Jersey: Princeton University Press.B. 1953. 1: 480-487 [5] Sober E (1984. the closer the alleles are to each other. Consequently. On The Origin of Species.Natural selection Genetic linkage Genetic linkage occurs when the loci of two alleles are linked. and this can be used to identify sites recently under selection. A strong selective sweep results in a region of the genome where the positively selected haplotype (the allele and its neighbours) are essentially the only ones that exist in the population. whether they are neutral or even slightly deleterious. Oxford [3] Works employing or describing this usage: Endler JA (1986). although with low recombination it can still lead to slightly negative linkage disequilibrium overall. Whether a selective sweep has occurred or not can be investigated by measuring linkage disequilibrium. the less likely it is that such a reshuffle will occur. The measurement of natural selection. org. [2] Fisher RA (1930) The Genetical Theory of Natural Selection Clarendon Press. or the preservation of favoured races in the struggle for life (http:/ / darwin-online. Sinauer Associates. Inc.[70] 256 See also • • • • • • Artificial selection Co-evolution Evolvability Gene-centered view of evolution Negative selection Unit of selection References [1] Darwin C (1859) On the Origin of Species by Means of Natural Selection. recombination of the genetic material results in reshuffling of the alleles. genetic recombination results in a reshuffling of the different alleles within a haplotype. Signet Classics. ISBN 0-87893-187-2 Haldane. J. Normally. this automatically results in selection of the other allele as well.S. when selection targets one allele. Oxford University Press. or whether a given haplotype is overrepresented in the population. As the prevalence of one allele increases. Published online at The complete work of Charles Darwin online (http:/ / darwin-online. selection for a specific allele will also result in selection of neighbouring alleles. Williams GC (1966). Because background selection is a result of deleterious new mutations. If a specific site experiences strong and persistent purifying selection. or in close proximity to each other on the chromosome. ISBN 0-451-52906-5. org. during a selective sweep. linked alleles can also become more common. Julian Huxley (2003). or the Preservation of Favoured Races in the Struggle for Life John Murray. linked variation will tend to be weeded out along with it. which can occur randomly in any haplotype. org. Nora Barlow (1958). PMID 11918795 [21] Empedocles. Book II [22] Lucretius. Proc Natl Acad Sci USA 94:13695-700 [19] Sanyal S. In: Evolutionary ecology (ed. Shorrocks B) pp65–79. M (1995). Roderick. De rerum natura (http:/ / classics. edu/ texts/ presoc/ emp. "Al-Jahiz And the Rise of Biological Evolutionism". J Exp Zool 285:19-26 [18] Zakany J. Physics (http:/ / classics. html). com/ news/ MRSASuperbug). Histoire de l'academie des sciences et belle lettres de Berlin 1746: 267–294. D. Sex-role reversal in vertebrates: behavioural and endocrinological accounts. com/ aiweb/ categories/ magazine/ 92_folder/ 92_articles/ 92_tusi. muslimphilosophy. p. PMID 16445718. Azerbaijan International 9 (2). inboxrobot. Jansen HG. 2006. Alfred Russel (1870) Contributions to the Theory of Natural Selection New York: Macmillan & Co. (1959). The eye of the blind mole rat. (http:/ / www.1111/j. Chapter 6. xiii (http:/ / darwin-online. ed. Blackwell Scientific. Nevo E. PMID 15884733 [11] "MRSA Superbug News" (http:/ / www. htm) [31] Maupertuis. edu/ Aristotle/ physics. org. Schmidt S & Sunyaev S (2005) Small fitness effect of mutations in highly conserved non-coding regions. org. "The autobiography of Charles Darwin 1809-1882" (http:/ / darwin-online. PMID 11074317 [10] Barlow GW. blackwell-synergy. (2005). 1111/ j. mit. uk/ content/ frameset?itemID=F373& viewtype=text& pageseq=20) [36] T. com/ doi/ abs/ 10.1126/science. Human Molecular Genetics 14:2221-9 [15] Bejerano G. Kent WJ. rsu. Pitnick S (1996) Investment in testes and the cost of making long sperm in Drosophila. Springer. pp. doi:10. PMID 16547091 [17] Galis F (1999) Why do almost all mammals have seven cervical vertebrae? developmental constraints. ISBN 1402088655 [27] Jan Z. p. Retrieved 2006-05-06. Rudiment with hidden function? Invest Ophthalmol Vis Sci. "Extraordinary flux in sex ratio". Phelps T. de Monet PA (1809) Philosophie Zoologique [33] Joravsky D. Muqaddimah.1469-0691. George K. faculty. [38] Wallace. New Jersey: Princeton University Press. edu/ cgi/ t/ text/ text-idx?c=moa& idno=AJP5195. com/ ik/ Muqaddimah/ Chapter6/ Ch_6_05. umich. 0001. A 13th-Century Darwin? Tusi's Views on Evolution (http:/ / azer. html). Agutter & Denys N. Book V [23] Aristotle. Genetics 173(2):891-900. de Jong WW. hanover. Oxford by adding survival selection in the reproductive phase [7] Pitnick S & Markow TA (1994) Large-male advantage associated with the costs of sperm production in Drosophila hydei. mit. Spalax ehrenbergi. Emily A.Natural selection [6] Modified from Christiansen FB (1984) The definition and measurement of fitness. Retrieved 2008-11-03. London. v. Pierre Louis (1748). "An Essay on the Principle of Population" (http:/ / www. [29] Franz Rosenthal and Ibn Khaldun. Hurst (2007). Sexual Selection. Muqaddimah. 1990 31(7):1398-404. p. How Do We Decide that a Species is Sex-Role Reversed? The Quarterly Review of Biology 80(1):28–35. (1990). 61 (http:/ / darwin-online. Baric S.1143369. Science 317 (5835): 214. Pinxten R. On Nature (http:/ / history. Makunin I. Rogers State University. Warot X & Duboule D (1997) Regulation of number and size of digits by posterior Hox genes: a dose-dependent mechanism with potential evolutionary implications. doi:10. org. Wilczynski (December 1959). PMID 17626876. [12] Schito GC (2006). p. htm). Hox genes. [9] Eens M.2006. Journal of the History of Ideas 20(1):85-104. ii. Fullard. edu/ Carus/ nature_things. (http:/ / www. Part 5 (http:/ / www. com/ ik/ Muqaddimah/ Chapter6/ Ch_6_29. hti. [32] Chevalier de Lamarck J-B. html). htm) [30] Franz Rosenthal and Ibn Khaldun. Speciation via introgressive hybridization in East African cichlids? Mol Ecol 11(3): 619–625. Proc Natl Acad Sci USA 91:9277-81. Woolfit M. 2. Princeton. Book II. 5 (http:/ / darwin-online. uk/ content/ frameset?itemID=F373& viewtype=side& pageseq=76) [35] Darwin 1859. Part 29 (http:/ / www. 01343. "On the Presumed Darwinism of Alberuni Eight Hundred Years before Darwin". Mattick JS & Haussler D (2004) Ultraconserved elements in the human genome. . James H. Wheatley (2008). de Grip WJ. [37] Charles Darwin. [25] Mehmet Bayrakdar (Third Quarter. edu/ ~felwell/ Theorists/ Malthus/ Essay. . 1469-0691. and cancer. p. Clin Microbiol Infect 12 Suppl 1: 3–8. Science 304:1321-5 [16] Eyre-Walker A. Pheasant M. uk/ content/ frameset?itemID=F381& viewtype=text& pageseq=20) 257 . Proceedings of the American Philosophical Society 84 (1). htm#112). muslimphilosophy. x) [13] Sylvain Charlat. Am Nat 148:57-80 [8] Andersson. [26] Paul S. uk/ content/ frameset?viewtype=text& itemID=F1497& pageseq=124). ISBN 0-691-00057-3. 5. PMID 2142147 [20] Salzburger W. 120. Neil Davies.01343. 1983). [14] Kryukov GV. Hornett. Stephen S. Natural Selection before the "Origin of Species". Isis 50 (4): 459–466. (2000). a species with giant sperm. Nina Wedell & Gregory D. [34] Darwin 1859. Chapter 6. Chapters 4 and 8 [24] Conway Zirkle (1941). 001& view=toc) [39] Darwin 1861. (2002). (2006). Robert Malthus (1798).1086/348801 [28] Farid Alakbarov (Summer 2001). doi:10. . 71-123. London: Collins.x. Sturmbauer C. Retrieved 2008-11-03. "The importance of the development of antibiotic resistance in Staphylococcus aureus". FromentalRamain C. The distribution of fitness effects of new deleterious amino acid mutations in humans. Thinking about Life: The History and Philosophy of Biology and Other Sciences. Soviet Marxism and Biology before Lysenko. The Islamic Quarterly. "Derivation of the laws of motion and equilibrium from a metaphysical principle (Original French text)". 43. Behav Processes 51(1-3):135-147. Stucke. (1958). Illinois. ISBN 0-87893-702-1 See esp. (2000). see Rose H. ISBN 978-0-521-73071. PMID 16957730. USA. Darwin Correspondence Project. New York: Oxford University Press ISBN 0-19-507951-5 [62] Goldberg DE. blackwellpublishing. [47] Haldane JBS (1932) The Causes of Evolution. UK ISBN 0-582-24302-5 [65] Rice SH. J Genet 55:511-24( (http:/ / www. org/ archive/ marx/ works/ 1883/ don/ index. darwinproject. org/ cgi/ content/ full/ 162/ 9/ 1760) [55] e. Darwin has called ‘natural selection’. com/ ridley/ classictexts/ wright. ISBN 0-316-18066-1 [59] For example. htm) [54] Quoted in translation in Eisenberg L (2005) Which image for Lorenz? Am J Psychiatry 162:1760 (http:/ / ajp. 28 September 1860" (http:/ / www. Retrieved 2010-01-12. R. wrote “This survival of the fittest.g. Consciousness Explained. Maurice E. USA. The Phylogenetic Handbook. ISBN 0-226-45808-3 [44] "Letter 5145 — Darwin. org. p. ISBN 0-19-286092-5 [58] Dennett DC. Massachusetts. 2006) [52] http:/ / www. nature. HarperCollins: New York. [1976] (1989). ac. fcgi?artid=1085053& blobtype=pdf) [PDF] Proc Natl Acad Sci USA 8:151–4 [61] Kauffman SA (1993) The Origin of order. (1996). The Language Instinct: How the Mind Creates Language. Harlow. com/ cgi/ viewcontent. uk/ darwinletters/ calendar/ entry-2931. ISBN 0-609-60513-5 [60] Lotka AJ (1922a) Contribution to the energetics of evolution (http:/ / www.. html). Optimization and Machine Learning. (2004). ch. ISBN 0-674-86250-3 [51] The New York Review of Books: Darwinian Fundamentalism (http:/ / www. The Propensity Interpretation of Fitness. and the Nature of Society. nybooks. 5 July (1866)" (http:/ / www. (2nd ed. p. psychiatryonline. Originally in Philosophy of Science (1979) 46: 263-286. . 258 . Darwin's Century: Evolution and the Men Who Discovered It. MIT Press: Cambridge. bepress. The Selfish Gene. C. org. A. [48] Wright S (1932) The roles of mutation. (1971). Rose SPR.”" [45] Darwin 1872. Essex. The genetical evolution of social behaviour I and II. Philippe. (1989). darwinproject. pubmedcentral. NY. MIT Press. ISBN 0-226-90134-3 [56] Pinker S. Melanie. uk/ content/ frameset?itemID=F391& viewtype=text& pageseq=70). Little. The Structure of Scientific Revolution 3rd ed. 3rd edn. com/ ridley/ classictexts/ haldane2. The evolution of reciprocal altruism. to Wallace. ISBN 0-262-69162-0. vol. "Better Competition Advocacy" (http:/ / works. University of Chicago Press. Alas. Addison-Wesley: Boston. New York. ISBN 0-06-097651-9 [57] Dawkins R. asp) Proc 6th Int Cong Genet 1:356–66 [49] Dobzhansky Th (1937) Genetics and the Origin of Species Columbia University Press. NY. USA [63] Mitchell. MA. An Introduction to Genetic Algorithms. ed. uk/ content/ frameset?itemID=F373& viewtype=side& pageseq=21) [41] "Darwin Correspondence Online Database: Darwin. . [70] Keightley PD. [1979] (1994). nih. "Interference among deleterious mutations favours sex and recombination in finite populations". is that which Mr. com/ Cellular_Darwinism_stochastic_gene_expression_in_cell_differentiation_and_embryo_development. fcgi?artid=1085052& blobtype=pdf) [PDF] Proc Natl Acad Sci USA 8:147–51 Lotka AJ (1922b) Natural selection as a physical principle (http:/ / www. gov/ picrender. PMID 5875341 PMID 5875340 [69] Trivers RL.1038/nature05049. Marco Salemi. Self-organization and selection in evolution. scitopics. 1941. 5 and 6 for a quantitative treatment. [67] http:/ / www. which I have here sought to express in mechanical terms. f3). Wilson. ac. pp3-23. New York. 1964 (http:/ / www. Moscow: Progress. USA. [46] Mills SK. com/ scitable/ topicpage/ Negative-Selection-1136 [68] Hamilton WD.. nih. com/ articles/ 1151) (accessed May 6. Jencks C.192. University of Chicago Press: Chicago. p. [42] Eisley L. NY. [43] Kuhn TS. html [53] Engels F (1873-86) Dialectics of Nature 3d ed. Anne-Mieke Vandamme (2009). Haldane JBS (1957) The cost of natural selection. [64] Falconer DS & Mackay TFC (1996) Introduction to Quantitative Genetics Addison Wesley Longman. R. Doubleday & Co: New York. Charles. and Otto SP (2006). 1951) [50] Mayr E (1942) Systematics and the Origin of Species Columbia University Press. Elliott Sober. Cambridge. Retrieved 2007-08-29. (1991). Evolutionary Theory: Mathematical and Conceptual Foundations. "Herbert Spencer in his Principles of Biology of 1864. gov/ picrender. Poor Darwin: Arguments Against Evolutionary Psychology. Religion. Harmony Books. USA. [1962] (1996). blackwellpublishing. [1994] (1995). Journal of Theoretical Biology 7: 1-16 and 17-52. and Co: New York. p. cgi?article=1000& context=maurice_stucke). Q Rev Biol 46: 35-57. USA. marxists. 444. Genetic Algorithms in Search.. uk/ entry-5145#mark-5145. republished in Conceptual Issues in Evolutionary Biology 2nd ed. 49 (http:/ / darwin-online. doi:10. Cambridge University Press. [66] Lemey. C. Retrieved 2006-05-10. crossbreeding and selection in evolution (http:/ / www. DS (2002) Darwin's Cathedral: Evolution. MA. Oxford University Press: New York. Nature 443 (7107): 89–92.Natural selection [40] Darwin 1859. . USA. 1. USA. (1964). inbreeding. pdf). Beatty JH. Sinauer Associates: Sunderland. to Lyell. R. Brown. Massachusetts. 6 (http:/ / darwin-online. pubmedcentral. or the preservation of favoured races in the struggle for life. • Kohm M (2004) A Reason for Everything: Natural Selection and the English Imagination. George C. University of Wisconsin • Natural Selection (http://evolution. • Dennett.wcer. Ballantine Books ISBN 0-345-42277-5. Doubleday. Simon & Schuster ISBN 0-684-82471-X. • Williams. • For general audiences • Dawkins. • Williams George C. 71-123. Richard (1996) Climbing Mount Improbable.springerlink. ISBN 0-521-45128-0. Karl (1978) Natural selection and the emergence of mind. Ryan Gregory: Understanding Natural Selection: Essential Concepts and Common Misconceptions (http:// www. Natural Selection before the "Origin of Species". London: Faber and Faber. Dialectica 32:339-55.Natural selection 259 Further reading • For technical audiences • Gould.com/content/2331741806807x22/fulltext. ISBN 0-670-85018-7. Oxford University Press.Natural Selection • Natural Selection (http://www. The Structure of Evolutionary Theory. John (1993).com/research/karl-popper-on-the-scientific-status-of-darwins-theory-of-evolution/) • Sober.php?topic_id=53) from University of Berkeley education website • T.org/authors/darwin-charles/ the-origin-of-species/chapter-04. Steve (2001) Darwin's Ghost: The Origin of Species Updated. see (http://human-nature. (1992) Natural Selection: Domains. • Jones.Modeling for Understanding in Science Education.html) – Chapter 4. ISBN 0-674-00613-5. p. Proceedings of the American Philosophical Society 84 (1). • Maynard Smith. Vintage Books. (1966) Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. The Theory of Evolution: Canto Edition. Norton.com/nibbs/05/wyhe.edu/ncisla/muse/naturalselection/index. Jonathan (1994) The Beak of the Finch: A Story of Evolution in Our Time. Stephen Jay (1997) Ever Since Darwin: Reflections in Natural History. Richard (1978) Adaptation. • Historical • Zirkle C (1941).html) Evolution: Education and Outreach .wisc.html).html) van Wyhe J (2005) Human Nature Review 5:1-4 External links • On the Origin of Species by Charles Darwin (http://www. • Lewontin. • Popper. Oxford University Press. Cambridge University Press. ISBN 0-393-06425-5. • Gould. Elliott (1984) The Nature of Selection: Evolutionary Theory in Philosophical Focus. Harvard University Press. Stephen Jay (2002). ISBN 1-86230-025-9. ISBN 0-571-22392-3. Levels and Challenges.literature. Also published in Britain under the title Almost like a whale: the origin of species updated. ISBN 0-679-73337-X. See (http:// mertsahinoglu. For review. Scientific American 239:212-30 • Weiner. Penguin Books.edu/evolibrary/search/topicbrowse2. Daniel (1995) Darwin's Dangerous Idea: Evolution and the Meanings of Life.berkeley. University of Chicago Press. peripatric. changes in the number or size of their bony plates. and sympatric.[4] One example of natural speciation is the diversity of the three-spined stickleback.[3] Natural speciation All forms of natural speciation have taken place over the course of evolution. Speciation may also be induced artificially. and color differences. the sticklebacks show structural differences that are greater than those seen between different genera of fish including variations in fins. parapatric. a view known as punctuated equilibrium. however it still remains a subject of debate as to the relative importance of each mechanism in driving biodiversity.[1] [2] Whether genetic drift is a minor or major contributor to speciation is the subject of much ongoing discussion. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis. a marine fish that. some palaeontologists such as Niles Eldredge and Stephen Jay Gould have argued that species usually remain unchanged over long stretches of time. The biologist Orator F. variable jaw structure. Over an estimated 10. Observed examples of each kind of speciation are provided throughout. While some evolutionary biologists claim that speciation events have remained relatively constant over time. through animal husbandry or laboratory experiments. has undergone speciation into new freshwater colonies in isolated lakes and streams. and that speciation occurs only over relatively brief intervals.[5] There is debate as to the rate at which speciation events occur over geologic time. based on the extent to which speciating populations are geographically isolated from one another: allopatric.000 generations.' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages.Speciation 260 Speciation Speciation is the evolutionary process by which new biological species arise. after the last ice age. There are four geographic modes of speciation in nature. The three-spined stickleback (Gasterosteus aculeatus) . a subform of allopatric speciation. Evidence for its speciation include genetic divergence. a population splits into two geographically isolated populations (for example. they have evolved such that they are reproductively isolated and are no longer capable of exchanging genes. has been observed in many circumstances. (b) they independently undergo genetic drift. but it was only nine months later that he reflected that such facts could show that species were changeable. the tendency of small. and difficulty in mating.Speciation 261 Allopatric During allopatric (from the ancient Greek allos. The Galápagos islands are particularly famous for their influence on Charles Darwin. Comparison of allopatric. by habitat fragmentation due to geographical change such as mountain building or social change such as emigration). Observed instances • Mayr bird fauna • The Australian bird Petroica multicolor • Reproductive isolation occurs in populations of Drosophila subject to population bottlenecking The London Underground mosquito is a variant of the mosquito Culex pipiens that entered in the London Underground in the nineteenth century. since small populations often undergo bottlenecks. It is related to the concept of a founder effect. and noticed that Mockingbirds differed from one island to another. new species are formed in isolated. parapatric and sympatric speciation. not just varieties. Genetic drift is often proposed to play a significant role in peripatric speciation. "other" + Greek patrā. more recent research has shown the birds now known as Darwin's finches to be a classic case of adaptive evolutionary radiation. During his five weeks there he heard that Galápagos tortoises could be identified by island.[6] Peripatric In peripatric speciation. peripatric. (c) different mutations arise in the two populations. smaller peripheral populations that are prevented from exchanging genes with the main population. and famously that other differing Galápagos birds were all species of finches. Observed instances Island genetics. behavioral differences. "fatherland") speciation. isolated genetic pools to produce unusual traits. The isolated populations then undergo genotypic and/or phenotypic divergence as: (a) they become subjected to dissimilar selective pressures. for example on Komodo. When he returned to England. his speculation on evolution deepened after experts informed him that these were separate species. When the populations come back into contact. including insular dwarfism and the radical changes among certain famous island chains. Though the finches were less important for Darwin.[7] . • The Greenish Warbler (Phylloscopus trochiloides). A niche must be available in order for a new species to be successful. connected habitat acting as a source of natural selection rather than the effects of isolation of habitats produced in peripatric and allopatric speciation. that have been studied by Dolph Schluter (who received his Ph. freshwater fishes. Ecologists refer to parapatric and peripatric speciation in terms of ecological niches. a Canadian benthic prefers a Japanese benthic over its close limnetic cousin from . there has a been a dearth of hard evidence that supports this form of speciation. The most widely accepted example of sympatric speciation is that of the cichlids of Lake Nabugabo in East Africa. which form a ring round the Central Valley in California. species diverge while inhabiting the same place.D. the existence of sympatric speciation as a mechanism of speciation is still hotly contested. there is only partial separation of the zones of two diverging populations afforded by geography. aquarium tests showed: • the benthic species from one lake will spawn with the benthic species from the other lakes and • likewise the limnetic species from the different lakes will spawn with each other. In sympatric speciation. around the Himalayas. for his work on Darwin's finches with Peter J. Schluter and colleagues found: • Two different species of three-spined sticklebacks in each of five different lakes • a large benthic species with a large mouth that feeds on large prey in the littoral zone • a smaller limnetic species — with a smaller mouth — that feeds on the small plankton in open water • DNA analysis indicates that each lake was colonized independently. Often-cited examples of sympatric speciation are found in insects that become dependent on different host plants in the same area.Speciation 262 Parapatric In parapatric speciation. Parapatric speciation is modelled on continuous variation within a 'single'. Grant) and his current colleagues in British Columbia. People have argued that the evidences of sympatric speciation are in fact examples of micro-allopatric. • The Ensatina salamanders. But there has been at least one recent study that suggests that sympatric speciation has occurred in Tennessee cave salamanders. or heteropatric speciation. neither mates with the other. • the grass Anthoxanthum has been known to undergo parapatric speciation in such cases as mine contamination of an area. Until recently. were once thought to provide an intriguing example best explained by sympatric speciation. • However. which is thought to be due to sexual selection.[8] [9] However. presumably by a marine ancestor. after the last ice age • DNA analysis also shows that the two species in each lake are more closely related to each other than they are to any of the species in the other lakes • The two species in each lake are reproductively isolated. individuals of each species may come in contact or cross habitats from time to time. Observed instances • Ring species • The Larus gulls form a ring species around the North Pole. Sympatric Sympatric speciation refers to the formation of two or more descendant species from a single ancestral species all occupying the same geographic location. but reduced fitness of the heterozygote leads to selection for behaviours or mechanisms that prevent their inter-breeding.[10] The three-spined sticklebacks. • These benthic and limnetic species even display their mating preferences when presented with sticklebacks from Japanese lakes. with a general feeling that interbreeding would soon eliminate any genetic differences that might appear. that is. 263 Hawthorn fly One example of evolution at work is the case of the hawthorn fly. The three-spine stickleback provides an example of how molecular biogeographic studies that rely solely on mtDNA can be misleading. Rhagoletis pomonella. that hawthorn flies mature later in the season and take longer to mature than apple flies. reproduction is often by parthenogenesis since polyploid animals are often sterile. also known as the apple maggot fly. and that consideration of the genealogical history of alleles from multiple unlinked markers (i. nuclear genes) is necessary to infer speciation histories. Sympatric speciation driven by ecological factors may also account for the extraordinary diversity of crustaceans living in the depths of Siberia's Lake Baikal.[11] using genetic markers from the nuclear genome. and that there is little evidence of interbreeding (researchers have documented a 4-6% hybridization rate) suggests that sympatric speciation is occurring. [needs further editing—not true in plants]. Some evidence. a non-native species. Not all polyploids are reproductively isolated from their parental plants.[13] Different populations of hawthorn fly feed on different fruits. Polyploidy is observed in many species of both plants and animals. for example. tetraploid zygote. which self-fertilize to produce a undergone an event of polyploidization in their evolutionary history. • The fact that this pattern of speciation occurred the same way on three separate occasions suggests strongly that ecological factors in a sympatric population can cause speciation. However.e. so an increase in chromosome number may not result in the complete cessation of gene flow between the incipient polyploids and their parental diploids (see also hybrid speciation). Speciation via polyploidization Polyploidy is a mechanism often attributed to causing some speciation events in sympatry. A more recent study. were introduced. the DNA evidence cited above is from mitochondrial DNA (mtDNA). The three-spine stickleback is now usually considered an example of "double invasion" (a form of allopatric speciation) in which repeated invasions of marine forms have subsequently differentiated into benthic and limnetic forms. but most often result in prenatal death. Rare instances of polyploid mammals are known. and results in rapid speciation since offspring of. • Their conclusion: in each lake. A distinct population emerged in North America in the 19th century some time after apples. This apple-feeding population normally feeds only on apples and not on the historically preferred fruit of hawthorns. producing diploid proposed that many of the existing plant and most animal species have gametes. tetraploid x Speciation via polyploidy: A diploid cell diploid matings result in triploid sterile progeny. The current hawthorn feeding population does not normally feed on apples. which appears to be undergoing sympatric speciation. such as the fact that six out of thirteen allozyme loci are different. what began as a single population faced such competition for limited resources that: • disruptive selection — competition favoring fishes at either extreme of body size and mouth size over those nearer the mean — coupled with: • assortative mating — each size preferred mates like it — favored a divergence into two subpopulations exploiting different food in different parts of the lake. The emergence of the . shows that limnetic forms in different lakes are more closely related to each other (and to marine lineages) than to benthic forms in the same lake.[12] It has been undergoes failed meiosis.Speciation its own lake. However. which can often move easily between closely related species ("introgression") when they hybridize. W. the two groups and their offspring were isolated reproductively because of their strong habitat preferences: they mated only within the areas they preferred.Speciation new hawthorn fly is an example of evolution in progress. Each generation was placed into the maze. and no longer produce viable offspring with Ovis orientalis.. then further mating between the populations will produce hybrids. which are usually created by diverged species coming into secondary contact. can be considered the same species as several varieties of wild ox. After thirty-five generations. starch and maltose.[20] Diane Dodd was also able to show how reproductive isolation can develop from mating preferences in Drosophila pseudoobscura fruit flies after only eight generations using different food types. on the other hand. Without reinforcement. and so did not mate with flies that preferred the other areas.[21] . and the groups of flies that came out of two of the eight exits were set apart to breed with each other in their respective groups. one species from which they are descended. Salt bred fruit flies.[15] It may occur after two populations of the same species are separated and then come back into contact. For example. etc. but the initial dates and methods of the initiation of such species are not clear. This is sometimes called the Wallace effect after the evolutionary biologist Alfred Russel Wallace who suggested in the late 19th century that it might be an important factor in speciation. using a maze with three different choices of habitat such as light/dark and wet/dry.g.[17] Domestic cattle.) The reasoning behind this is that if the parents of the hybrid offspring each have naturally selected traits for their own certain environments. called their "hybrid zone. If their reproductive isolation is incomplete. then the populations will merge back into the same species within the area they are in contact. Reinforcement is required for both parapatric and sympatric speciation. the geographic area of contact between different forms of the same species. Drosophila melanogaster. If their reproductive isolation was complete. 264 Reinforcement (Wallace effect) Reinforcement is the process by which natural selection increases reproductive isolation. Reinforcement may be induced in artificial selection experiments as described below. Hybrid zones are regions where diverged populations meet and interbreed. then there will be no further reproductive isolation and speciation has essentially occurred (e." will not develop into a boundary between the different species. the hybrid offspring will bear traits from both. then they will have already developed into two separate incompatible species. therefore would not fit either ecological niche as well as either parent. as in horses and donkeys.[16] If the hybrid offspring are more fit than their ancestors.. The low fitness of the hybrids would cause selection to favor assortative mating. Hybrid offspring are very common in these regions. Without reinforcement the two species would have uncontrollable inbreeding. yak.[19] The history of such attempts is described in Rice and Hostert (1993). or fertile but less fit than their ancestors.[18] The best-documented creations of new species in the laboratory were performed in the late 1980s. William Rice and G. as they readily produce fertile offspring with them. Artificial speciation New species have been created by domesticated animal husbandry.[14] Speciation via hybrid formation See Hybrid speciation section under the Genetics heading below. gaur. which would control hybridization. domestic sheep were created by hybridisation. If the hybrids are infertile. which may or may not be fertile. Hybridization without change in chromosome number is called homoploid hybrid speciation. However. his idea was sound. it may lead to a separate species. In 2008 a speciation gene causing reproductive isolation was reported. especially in plant species. Gene transposition as a cause Theodosius Dobzhansky. reproductive isolation between hybrids and their parents is particularly difficult to achieve and thus hybrid speciation is considered an extremely rare event. Those mobile sections can cause sterility in inter-species hybrids. who studied fruit flies in the early days of genetic research in 1930s.[26] This validates the reproductive isolation mechanism.Speciation 265 Dodd's experiment has been easy for many others to replicate. but scientists long debated whether it actually happened in nature.[23] It causes hybrid sterility between related subspecies. Eventually a competing theory involving the gradual accumulation of mutations was shown to occur in nature so often that geneticists largely dismissed the moving gene hypothesis. a key component of speciation.[27] . is a more common phenomenon. which can act as a speciation pressure. 2006 research shows that jumping of a gene from one chromosome to another can contribute to the birth of new species. This phenotype can also be fitter than the parental lineage and as such natural selection may then favor these individuals. In theory.[25] However. including with other kinds of fruit flies and foods.[22] Genetics Few speciation genes have been found. speculated that parts of chromosomes that switch from one location to another might cause a species to split into two different species. They usually involve the reinforcement process of late stages of speciation. He mapped out how it might be possible for sections of chromosomes to relocate themselves in a genome. Polyploid speciation. Hybrid speciation Hybridization between two different species sometimes leads to a distinct phenotype. The Mariana Mallard is known to have arisen from hybrid speciation. which involves changes in chromosome number. Eventually. if reproductive isolation is achieved. It is considered very rare but has been shown in Heliconius butterflies [24] and sunflowers. [28] See also • Extinction • Species problem • Heteropatry • Chronospecies • Koinophilia References [1] [2] [3] [4] Cook. BENJAMIN M. BRIAN T. PMC 1690834. J. 2008. McPHAIL (2000). Bush GL (1982). mechanisms. org/ cgi/ pmidlookup?view=long& pmid=12663534). Forbes. nih.1098/rspb. Sympatric speciation in phytophagous insects: moving beyond controversy? Annual Review of Entomology 47:773-815 [10] MATTHEW L. "Historical contingency and determinism interact to prime speciation in sticklebacks" (http:/ / www. O.2000.1294.Speciation 266 Interspersed repeats Interspersed repetitive DNA sequences function as isolating mechanisms. The importance is that this allows the splitting of a gene pool without requiring physical isolation of the organisms harboring those gene sequences. Sulloway (1982). doi:10. available online (http:/ / darwin-online. com/ hdy/ journal/ v82/ n1/ full/ 6884120a. A. L. PMC 1462491. nature. F. J. gov/ articlerender. x) [11] E. p. W. doi:10. D.2307/2413033.. jstor. . and Bush and the complexities of sympatric speciation in Rhagoletis.. and rates of polyploid formation in flowering plants. 1111/ j. 1906. uk/ content/ frameset?viewtype=text& itemID=A86& pageseq=2) [7] Katharine Byrne and Richard A Nichols (1999) "Culex pipiens in London Underground tunnels: differentiation between surface and subterranean populations" (http:/ / www. "Evidence for inversion polymorphism related to sympatric host race formation in the apple maggot fly. Pathways. Factors of species-formation. F. pubmedcentral. 2005.M. J.2-1& origin=ISI) available online [12] Ramsey.D. Cook. and M. Roethele JB.B. Dambroski. "Recent divergence with gene flow in Tennessee cave salamanders (Plethodontidae: Gyrinophilus) inferred from gene genealogies".1 million years ago. Xie. (January 2009) "From Atoms to Traits. O. Annual Review of Ecology and Systematics 29:467-501 [13] Feder JL. Bulletin of the British Museum (Natural History) Zoology Series 43 (2): 49–58. org/ faqs/ faq-speciation. Molecular Ecology 17 (9): 2258–2275. Filchak K. Retrieved 8 June 2009. both within gene families and also allelic forms of a gene. Analysis of genetic drift and recombination using a Markov model suggests humans and chimpanzees speciated apart 4. 1908. B. Proceedings of the National Academy of Sciences. Romero-Severson J (1 March 2003). Human speciation Humans have genetic similarities with chimpanzees and gorillas. Baker (2005). Filchak. USA 1902:6573-6580 [9] Berlocher. PMID 12663534.2005. Dobzhansky. These repeats protect newly evolving gene sequences from being overwritten by gene conversion. "Adaptive speciation: The role of natural selection in mechanisms of geographic and non-geographic speciation". Aluja. J. American Naturalist 42:727-731. Science 23:506-507. (http:/ / www. 0. "The Beagle collections of Darwin's finches (Geospizinae)". 03750. [5] Kingsley. html) by Joseph Boxhorn. X. talkorigins. NIEMILLER. Proceedings of the Royal Society of London Series B 267 (1460): 2375–2384. This barrier allows nascent novel genes to evolve without being overwritten by the progenitors of these genes. 1365-294X. suggesting common ancestors. com/ ridley/ tutorials/ Speciation15. 1998. K. Studies in History and Philosophy of Biological and Biomedical Sciences 36 (2): 303–326. Leung. blackwellpublishing. Feder. and J. PMID 19260194. Genetics 163 (3): 939–53. 57 [6] Frank J. TAYLOR. org/ sici?sici=0962-8452(200012)267:1460<2375:HCAEDI>2. M. available online (http:/ / www. 2002. .shpsc. due to the creation of non-homologies between otherwise homologous DNA sequences. com/ doi/ abs/ 10. [14] Berlocher SH. E. html) [8] Feder. Evolution without isolation. doi:10. S. and D. asp) . Systematic Zoology 31 (2): 136–55. fcgi?tool=pmcentrez& artid=1690834). CO. Schemske.1016/j.. [15] Ridley. genetics. Rull. Mayr. "An electrophoretic analysis of Rhagoletis (Diptera: Tephritidae) phylogeny" (http:/ / jstor. H.M. Observed Instances of Speciation (http:/ / www. This uncoupling allows the evolution of new genes. S. Velez. org/ stable/ 2413033).005." Scientific American.03. FITZPATRICK. PMID 11133026. H. L. org. MILLER (2008). J. Niedbalski J. blackwell-synergy. (2003) "Speciation — What is the role of reinforcement in speciation?" adapted from Evolution 3rd edition (Boston: Blackwell Science) tutorial online (http:/ / www. Rhagoletis pomonella" (http:/ / www. The non-homologies create barriers to gene conversion. [25] University of Rochester Press Releases (http:/ / www.net/weblog/topics/phylogeny/speciation. Allen Orr (September 2006).R. C. com/ article. doi:10. J. Linares. org/ stable/ 2410209).Speciation [16] Ollerton. Animal Species and Evolution. sciam. • White. New York: Columbia University Press.D. 1086/ 338370) [23] http:/ / www. edu/ doi/ abs/ 10. Chimpanzee.info/speciation. [22] Kirkpatrick. and H. Schierup MH (2007) "Genomic Relationships and Speciation Times of Human. J. & Orr. J." (http:/ / genetics. The American Naturalist 131: 911–917. (1981). ISBN 0-231-05113-1.edu/evolibrary/article/0_0_0/evo_45) from Understanding Evolution (http://evolution. D. Hostert (1993). 1371/ journal. php?id=2603) [26] Masly. plosjournals. P. Mohamed A. (1978).royalsociety.B. Oxford: Elsevier. Harvard University Press. Sunderlands. and • Evidence for Speciation (http://evolution.php) by the University of California Museum of Paleontology • Speciation (http://johnhawks.1038/nature04738. (Baltimore: Johns Hopkins University Press) [19] Rice. • Dedicated issue of Philosophical Transactions B on Speciation in microorganisms is freely available. R. PMID 16778888.org/origins/faqs-qa. 2006) "Wandering Fly Gene Supports New Model of Speciation" (http:/ / www.net/weblog/topics/phylogeny/) • Speciation in the context of evolution (http://evolution-of-man. Noor. Retrieved 2007-03-18." Evolution 43:1308–1311. Allopatry. John P. Salcedo. (September 8.0030007) 267 Further reading • Coyne. et al. pgen.edu/evolibrary/article/0_0_0/evo_40). . doi:10. ISBN 978-0-444-52033-3. M. August 2005. H. (http:// publishing. "Laboratory experiments on speciation: What have we learned in forty years?" (http:/ / jstor.. (1989) "Reproductive isolation as a consequence of adaptive divergence in Drosophila pseudoobscura.berkeley.2307/2410209. 0030007) PLoS Genet 3(2): e7 (doi:10.. northampton.1086/284831. [27] Minkel. A. J. [21] Dodd. org/ perlserv/ ?request=get-document& doi=10. Evolution 47 (6): 1637–1653.R. rochester. E. Retrieved 2007-05-22. "Flowering time and the Wallace Effect" (http:/ / oldweb. org/ cgi/ content/ short/ 323/ 5912/ 376 [24] Mavarez. [20] W. journals. . ISBN 0-674-03750-2 • Marko. Freeman and Company. ed.berkeley. edu/ news/ show. Christensen OF. Salazar. and G.htm) .berkeley. J.W. D. F. M.talkorigins. Speciation. C. Mailund T. (1963).E. E. and V. cfm?chanID=sa003& articleID=000A84DB-CA3B-1501-8A3B83414B7F0000) Science News [28] Hobolth A.. genetics. W. Ravigné (2002) "Speciation by Natural and Sexual Selection: Models and Experiments" The American Naturalist 159:S22–S35 DOI (http:/ / www. . PMID 16960009.html) from the Talk. A.org/faqs/faq-speciation.pgen. H. B.talkorigins.A. sciencemag. uk/ aps/ env/ lbrg/ journals/ papers/ OllertonHeredityCommentary2005. M. and evolution (http://johnhawks.1126/science. (2006). Heredity. (2002) "Molecular analysis of wild and domestic sheep questions current nomenclature and provides evidence for domestication from two different subspecies" Proceedings of the Royal Society B: Biological Sciences 269:893-904 [18] Nowak. doi:10. Nature 441 (7095): 868.. Salt (1988). (2008). pdf) (PDF).1371/journal. Jones. org/ cgi/ content/ short/ 313/ 5792/ 1448).. "Speciation by hybridization in Heliconius butterflies". sciencemag.1128721. and Gorilla Inferred from a Coalescent Hidden Markov Model. San Francisco.paleoanthropology. (2004). "Speciation via disruptive selection on habitat preference: experimental evidence". (1999) Walker's Mammals of the World 6th ed.M.Origins Frequently Asked Questions (http://www. John Locke. • Grant. [17] Hiendleder S. ISBN 0-87893-089-2.). Modes of Speciation. Plant Speciation (2nd Edit. Bermingham. "Gene Transposition as a Cause of Hybrid Sterility in Drosophila" (http:/ / www. • Mayr. California: W. Inc. Jiggins. Corbin D. V. ac. Science 313 (5792): 1448–1450.html) from John Hawks' Anthropology Weblog . ..html) • Speciation (http://evolution. uchicago.R. C. doi:10. Massachusetts: Sinauer Associates. ISBN 0-716-70284-3.edu/evolibrary/home. Rice and E.org/microgenesis) External links • Observed Instances of Speciation (http://www. and MCMC-based Bayesian inference. hybridize together. The most commonly-used methods to infer phylogenies include parsimony. Phenetics. has been richly informed by phylogenetics but remains methodologically and logically distinct. phylogenetics is the study of evolutionary relatedness among various groups of organisms (for example.[2] In biological systematics as a whole. which is discovered through molecular sequencing data and morphological data matrices.[3] A phylogenetic tree represents a hypothesis of the order in which evolutionary events are assumed to have occurred." and genetikos (γενετικός). identification.268 Phylogenetics Phylogenetics In biology. which represent groups of lineage-connected individuals. race.[1] The fields overlap however in the science of phylogenetic systematics – often called "cladism" or "cladistics" – where only phylogenetic trees are used to delimit taxa. . meaning "tribe. maximum likelihood. wherein the characters are aligned nucleotide or amino acid sequences. Taxonomy. All methods depend upon an implicit or explicit mathematical model describing the evolution of characters observed in the species included. phylogenetic analyses have become essential in researching the evolutionary tree of life. the classification. and are usually used for molecular phylogeny. or terminate by extinction. Construction of a phylogenetic tree Evolution is regarded as a branching process. This may be visualized in a phylogenetic tree. meaning "relative to birth" from genesis (γένεσις. species or populations). The problem posed by phylogenetics is that genetic data are only available for living taxa. popular in the mid-20th century but now largely obsolete. The term phylogenetics is of Greek origin from the terms phyle/phylon (φυλή/φῦλον). uses distance matrix-based methods to construct trees based on overall similarity. "birth"). Cladistics is the current method of choice to infer phylogenetic trees. which is often assumed to approximate phylogenetic relationships. and the fossil records (osteometric data) contains less data and more-ambiguous morphological characters. and naming of organisms. whereby populations are altered over time and may speciate into separate branches. paraphyletic. Phylogenetic groups. RedToL aims at reconstructing the Red Algal Tree of Life. "Modern reptile" (cyan in the diagram) is a grouping that contains a common ancestor. but does not contain all descendants of that ancestor (birds are excluded). or taxa. however. A grouping such as warm-blooded animals would include only mammals and birds (red/orange in the diagram) and is called polyphyletic because the members of this grouping do not include the most recent common ancestor. can be monophyletic. all birds and reptiles are believed to have descended from a single common ancestor. are used to construct phylogenetic trees. For instance. . often anatomical characteristics.[4] The overall goal of National Science Foundation's Assembling the Tree of Life activity (AToL) is to resolve evolutionary relationships for large groups of organisms throughout the history of life. biologists tend to be restricted with analysing present-day organisms to identify their evolutionary relationships. or polyphyletic. with the research often involving large teams working across institutions and disciplines. Therefore. fossil records are often too poor to be of good help. Fossils can aid with the reconstruction of phylogenies. This is an example of a paraphyletic group. which includes protein and DNA sequences. algorithm development and dissemination in computational phylogenetics and phyloinformatics.Phylogenetics 269 Grouping of organisms There are some terms that describe the nature of a grouping in such trees. Due to the fact that evolution takes place over long periods of time that cannot be observed directly. Today. analysis. biologists must reconstruct phylogenies by inferring the evolutionary relationships among present-day organisms. Investigators are typically supported for projects in data acquisition. For example. Molecular phylogenetics The evolutionary connections between organisms are represented graphically through phylogenetic trees. molecular data. so this taxonomic grouping (yellow in the diagram below) is called monophyletic. Phylogenetic relationships in the past were reconstructed by looking at phenotypes. i. organisms can inherit genes in two ways: vertical gene transfer and horizontal gene transfer. Carl Woese came up with the three-domain theory of life (eubacteria. Williamson suggested that larvae and embryos represented adults in other taxa that have been transferred by hybridization (the larval transfer theory). This may be partly due to the breaking up of long branches. Vertical gene transfer is the passage of genes from parent to offspring. Therefore. it has been proposed that it is more important to increase the number of taxa in the matrix than to increase the number of characters. a common phenomenon in prokaryotes. archaea and eukaryotes) based on his discovery that the genes encoding ribosomal RNA are ancient and distributed over all lineages of life with little or no horizontal gene transfer. the development of an organism exactly mirrors the evolutionary development of the species. it has become feasible to gather large amounts of data (DNA or amino acid sequences) to infer phylogenetic hypotheses. Most modern biologists recognize numerous connections between ontogeny and phylogeny. or view them as supporting evidence for that theory. For example. Haeckel's early version of this hypothesis [that the embryo mirrors adult evolutionary ancestors] has since been rejected. rRNAs are commonly recommended as molecular clocks for reconstructing phylogenies. because the more taxa the more robust is the resulting phylogenetic tree [9] . and inconsistencies in phylogeny have been reported among specific groups of organisms depending on the genes used to construct evolutionary trees. He was accused by five professors of falsifying his images of embryos (See Ernst Haeckel). Donald I.[8] Gene transfer Genealogical tree suggested by Haeckel (1866) In general. This theory was often expressed as "ontogeny recapitulates phylogeny".e. explain them using evolutionary theory.Phylogenetics 270 Ernst Haeckel's recapitulation theory During the late 19th century. or biogenetic law. and there is a significant body of evidence against the larval transfer theory. This has been particularly useful for the phylogeny of microorganisms. Horizontal gene transfer has complicated the determination of phylogenies of organisms. Ernst Haeckel's recapitulation theory.[5] [6] However. Williamson's views do not represent mainstream thought in molecular biology[7] . Taxon sampling and phylogenetic signal Owing to the development of advanced sequencing techniques in molecular biology. was widely accepted. However. and the hypothesis amended as the embryo's development mirroring embryos of its evolutionary ancestors.000 nucleotides. in many animals). It has been argued that this is an important reason to incorporate data . to which the species concept does not apply and which are too morphologically simple to be classified based on phenotypic traits. it is not rare to find studies with character matrices based on whole mitochondrial genomes (~16. and horizontal gene transfer or lateral gene transfer occurs when genes jump between unrelated organisms. for instance. Even so. 271 Importance of missing data In general. such characters should be given less weight in the reconstruction of a tree.[14] Unfortunately the only objective way to determine convergence is by the construction of a tree – a somewhat circular method. the more accurate and reliable the resulting tree will be. Of course. weighting homoplasious characters does indeed lead to better-supported trees. however. as they can provide information in sparse areas of trees. Another important factor that affects the accuracy of tree reconstruction is whether the data analyzed actually contain a useful phylogenetic signal.[14] Further refinement can be brought by weighting changes in one direction higher than changes in another. there is no way to measure whether a particular phylogenetic hypothesis is accurate or not.[15] See also • • • • • • • • • Bauplan Bioinformatics Biomathematics Cladistics Coalescent theory Computational phylogenetics EDGE of Existence Programme Important publications in phylogenetics Language family • Maximum parsimony • Molecular phylogeny • PhyloCode . although its impact is greatest when most of the missing data is in a small number of taxa. it is sometimes difficult to incorporate fossil data into phylogenies.[11] Ultimately. rather than DNA data. the inclusion of fossils is invaluable. Missing data is no less detrimental than simply having less data. The fewer characters that have missing data. thus. although because wings are often lost secondarily. the presence of thoracic wings almost guarantees placement among the pterygote insects. phylogenetic data that include fossil taxa are generally based on morphology. the better. fossil taxa contribute as much to tree resolution as modern taxa. the more data that is available when constructing a tree. concentrating the missing data across a small number of character states produces a more robust tree. However. logically. unless the "true" relationships among the taxa being examined are already known. The best result an empirical systematist can hope to attain is a tree with branches well-supported by the available evidence.Phylogenetics from fossils into phylogenies where possible.[12] Role of fossils Because many morphological characters involve embryological or soft-tissue characters that cannot be fossilized. despite these limitations. a term that is used generally to denote whether related organisms tend to resemble each other with respect to their genetic material or phenotypic traits. Using simulations. Derrick Zwickl and David Hillis[10] found that increasing taxon sampling in phylogenetic inference has a positive effect on the accuracy of phylogenetic analyses. their absence does not exclude a taxon from the group. breaking up long branches and constraining intermediate character states. and the interpretation of fossils is more ambiguous than living taxa.[13] Homoplasy weighting Certain characters are more likely to be evolved convergently than others. M. doi:10. pdf) [12] Prevosti. pp. Systematics Assoc. [13] Cobbett. "Missing data and the design of phylogenetic analyses"..2006.2008. (2008). Edwards. 9 November 2009 [8] Michael W.x. doi:10. doi:10. S.1111/j. ISBN 978-0716-77928-5. UC Berkeley. Cavalli-Sforza LL Phylogenetics is that branch of life science. Springer. . Proceedings of the National Academy of the Sciences. .1096-0031. Wilkinson. (1967). M. "xviii". No.1080/10635150290102339. F. html). edu/ glossary/ glossary_1.x. "Fossils impact as hard as living taxa in parsimony analyses of morphology". org/ stable/ 2406616).. Wills. Grosberg. org/ content/ early/ 2009/ 10/ 22/ 0910229106). doi:10.001. Hillis DM (2002). [15] Goloboff. Genetics: A conceptual Approach (3rd ed. "Caterpillars did not evolve from onychophorans by hybridogenesis" (http:/ / www.00289. doi:10. [4] Pierce.04. Freeman. ARS Technica. W.0910229106) [9] Wiens J (2006). and Richard K. A. "Examining science on the fringes: vital. Vrian (1998).1096-0031. Reconstruction of evolutionary trees.x. ISBN 978-1402-01514-4. A. "Testing for phylogenetic signal in comparative data: behavioral traits are more labile". [5] Williamson DI (2003-12-31). A.00236. The Origins of Larvae (2nd ed.1111/j. P. doi:10.which deals with the study of evolutionary relation among various groups of organisms. (1997).1073/pnas. Arias. Zoological Journal of the Linnean Society 148: 585–602.. Carpenter. (2007-12-17).). M.Phylogenetics • • • • • • • • • • • Joe Felsenstein Systematics Phylogenetic tree Phylogenetic network Phylogenetic nomenclature Phylogenetics software Phylogenetic tree viewers Phylogeography Phylodynamics Phylogenetic comparative methods Microbial phylogenetics 272 References [1] Edwards AWF. 6: Phenetic and Phylogenetic Classification. Benjamin A. J. J. PMID 15922672. F.1096-3642. Systematic Biology 51 (4): 588–598. berkeley. Ives AR (2003).2009. 261.00209. (2009). Systematic biology 56 (5): 753–766. Retrieved 2008-03-22. PMID 17886145. ars).2005. W.1111/j. "Phylogenetic Analysis: Models and Estimation Procedures" (http:/ / jstor.through molecular sequencing data. Evolution 21 (3): 550–570. �A. (2007). edu/ people/ faculty/ Garland/ BlomEA03.. "Increased taxon sampling greatly reduces phylogenetic error". A. pp.tb00317. Chemisquy.1997. pnas. [2] Speer. [6] Williamson DI (2006). A. [10] Zwickl DJ. [7] John Timmer. M. Garland T Jr. "The impact of missing data on real morphological phylogenies: influence of the number and distribution of missing entries". PMID 12778543. Hart.2307/2406616.1016/j. [11] Blomberg SP. Publ.1096-0031. PDF (http:/ / www. Journal of Biomedical Informatics 39 (1): 34–42. 67–76. Esquivel. ed. [3] Cavalli-Sforza. but generally wrong" (http:/ / arstechnica.x. (1964). Cladistics 13: 225. "Weighting against homoplasy improves phylogenetic analysis of morphological data sets". "UCMP Glossary: Phylogenetics" (http:/ / www. com/ science/ news/ 2009/ 11/ examining-science-on-the-fringes-vital-but-generally-wrong. doi:10.. H. "Hybridization in the evolution of animal form and life-cycle". biology. L. Evolution 57 (4): 717–745. Cladistics 26: 326.jbi. L. "Self-Weighted Optimization: Tree Searches and Character State Reconstructions under Implied Transformation Costs". [14] Goloboff. Cladistics 24: 758. ucr. P. D.1111/j.).1080/10635150701627296. . M. 30 October 2009 (doi: 10. R.. PMID 12228001.. J. doi:10. ucmp. geneious.cfm) – lecture on phylogenetics by Sydney Brenner • Mikko's Phylogeny Archive (http://www. and A.org in Spanish (http://www.html) • What Genomes Can Tell Us About the Past (http://ascb. program for computing phylogenetic trees and unrooted phylogenetic networks • Dendroscope (http://www.gov/About/primer/phylo. T. Brower.helsinki.trex.Phylogenetics 273 Further reading • Schuh.Dendroscope.nih.org/tree/learn/concepts/whatisphylogeny.nlm. R. 2009.org) Filogenetica.org).berkeley.org/) UCMP Exhibit Halls: Phylogeny Wing (http://www.pdf) .embl.uqam. Z.cladistics. program for drawing phylogenetic trees and rooted phylogenetic networks • Phylogenetic inferring on the T-REX server (http://www. V.) ISBN 978-0-8014-4799-0 External links • • • • • • • • • The Tree of Life (http://tolweb.html) • Geneious Pro (http://www.ru.edu/~moret/poincare.SplitsTree. Biological Systematics: principles and applications (2nd edn.ncbi.ucmp.cs.filogenetica.org).html) Willi Hennig Society (http://www.org/ibioseminars/brenner/brenner1.org/mesquite/mesquite.unm.html) Interactive Tree of Life (http://itol. Phylogenetic Patterns (http://www.ca) • Mesquite (http://mesquiteproject.edu/exhibit/phylogeny.cmbi.edu/phylocode/) ExploreTree (http://exploretree.org) PhyloPat.fi/~mhaaramo/) • Phylogenetic Reconstruction from Gene-Order Data (http://www.com) all-in-one phylogenetics software • NCBI – Systematics and Molecular Phylogenetics (http://www.de) PhyloCode (http://www.nl/phylopat) SplitsTree (http://www.ohiou. [4] Cladists use cladograms. crocodiles. to represent the monophyletic relationships of species.[2] In the terms of biological systematics. The technique and sometimes the name have been successfully applied in other disciplines: for example. However in the 1990s it rapidly became the dominant method of classification in evolutionary biology. to determine the relationships between the surviving manuscripts of the Canterbury Tales. cladistic classifications (usually trees called cladograms) are intended to reflect the relative recency of common ancestry or the sharing of homologous features. who referred to it as "phylogenetic systematics" (also the name of his 1966 book).[5] Hennig referred to his own approach as phylogenetic systematics. cladistic by Cain and Harrison in 1960. Cladistics can be distinguished from other taxonomic systems. dinosaurs. a clade is a single "branch" on the "tree of life". the use of the terms "cladistics" and "clade" was popularized by other researchers. termed sister-group relationships. a monophyletic group. From the time of his original formulation until the end of the 1980s cladistics remained a minority approach to classification. This is interpreted as representing phylogeny. birds. History of cladistics The term clade was introduced in 1958 by Julian Huxley. and all descendants (living or extinct) of their most recent common ancestor form a clade. For example. by its focus on shared derived characters (synapomorphies). Computers made it possible to process large quantities of data about organisms and their characteristics. or evolutionary relationships. genetic sequencing data and computational phylogenetics are now very commonly used in the generation of cladograms. Although traditionally such cladograms were generated largely on the basis of morphological characters. such as phenetics. which consist of 1) all the descendants of an ancestral organism and 2) the ancestor itself.[3] Cladistics originated in the work of the German entomologist Willi Hennig. Cladistics is also distinguished by its emphasis on parsimony and hypothesis testing (particularly falsificationism). rather than subjective decisions that some other taxonomic systems rely upon.Cladistics 274 Cladistics Cladistics[1] is a method of classifying species of organisms into groups called clades. diagrams which show ancestral relations between species. families and other higher level classification.[6] . Previous systems usually employed overall morphological similarity to group species into genera. At about the same time the development of effective polymerase chain reaction techniques made it possible to apply cladistic methods of analysis to biochemical and molecular genetic features of organisms as well as to anatomical ones. and cladist (for an adherent of Hennig's school) by Mayr in 1965. or morphological evidence might suggest that fish and amphibians share a common ancestor that was not an ancestor of the other tetrapods. an apomorphy of one clade is the plesiomorphy of another contained within it. thus. The new information would cause us to define a fish-and-amphibian clade. the blue group (reptiles) is paraphyletic. In a hypothetical example. These terms are equivalent to but more precise than the homology. further genetic The yellow group (sauropsids) is monophyletic. it is hypothesized that the first vertebrate species is the common ancestor of all vertebrate species. This can be seen as a predecessor[7] to some methods of today's cladistics (namely distance matrix methods such as neighbor-joining). including fishes (Pisces). the presence of a backbone (shared by all vertebrates) can be hypothesized to have existed in the common vertebrate ancestor. An important caution is that any cladogram is a provisional hypothesis. Terminology The following terms are used to identify shared or distinct characters among groups:[8] [9] [10] • Plesiomorphy ("close form") or ancestral state. altering the cladogram. For example. mammals and birds.Cladistics 275 Cladistics as a successor to phenetics For some decades in the mid to late twentieth century. reptiles. and the red group (warm-blooded animals) is polyphyletic. having four limbs is an apomorphy for vertebrates but a plesiomorphy for tetrapods. in the tree shown. The tetrapod clade is thus nested within the vertebrate clade. They are thus relative terms. In the diagram. For example. The first tetrapod is the ancestor of all tetrapods. a commonly used methodology was phenetics ("numerical taxonomy"). including amphibians. The tetrapod ancestor was a descendant of the original vertebrate ancestor. all vertebrates are monophyletic. also symplesiomorphy ("shared plesiomorphy"). Clades A clade is a group of taxa consisting of an ancestor taxon and all of its descendant taxa. only similarities. is a characteristic that is present at the base of the tree. The application of the above terms to a group depends on one's perspective in the tree. For example. Three main types of groups can be identified by plotting their relationships in cladograms:[8] [9] [10] • Monophyletic groups are groups containing only taxa descended from a given ancestor taxon and all those descendants. • Apomorphy ("separate form") or derived state is a characteristic believed to have evolved within the tree. allowing one to express the hierarchical relationships among different homologies. since all the taxa are descended from a single . all tetrapods have four limbs. but made no attempt to resolve phylogeny. but is not an ancestor of any fish living today. In the diagram provided. • Synapomorphy ("shared apomorphy") is an apomorphy which is shared between taxa. is incorrectly applied. Paraphyletic groups are typically diagnosed on the basis of shared plesiomorphy and the exclusion of groups that are diagnosable by a synapomorphy. but the Mammalian ancestor came from the Amniotes. Polyphyletic groups are recognized by the homoplasy relation (that is. For example. The bird clade is nested within the reptilian clade. Monophyly is diagnosed by the synapomorphy relation. with the proviso that cladistic taxa always denote a clade. birds are warm-blooded. the vertebrate clade is basal to the tetrapod and fish clades. a grouping of birds and mammals based on their warm-bloodedness is not monophyletic. see wastebasket taxon). . with such a deficit being taken as an indication of primitiveness. excluding birds from the sample cladogram would create a paraphyletic group. For instance. • Paraphyletic groups are groups excluding one or more descendant taxa of the common ancestor. The plesiomorphy is the amniotic fluid and the synapomorphy is warm-bloodedness. The three ways to define a clade synapomorphically) with that trait in A. (This type of definition was originally called "stem-based". which because of that exclusion is a paraphyletic group. and all descendants of that ancestor. and all descendants of that ancestor. Crown groups are a type of node-based clade. the Aves ancestor came from the Archosaurs. but this was changed to avoid confusion with the term "stem group". In this case. 276 Three definitions of clade There are three major ways to define a clade for use in a cladistic taxonomy. • Branch-based: the first ancestor of A which is not also an ancestor of C. The warm-bloodedness is not a plesiomorphy but is a homoplasy. • Apomorphy-based: the first ancestor of A to possess derived trait M homologously (that is. and thus contain some but not all of its descendants. reptiles.Cladistics ancestor (the common vertebrate species) and there are no others. For instance. or convergence. • Polyphyletic groups are groups containing taxa from two or more different monophyletic groups. Clades relate to each other in these ways: • A clade is basal to another clade if it contains that other clade as a subset within it. and all descendants of that ancestor.) Total groups are a type of branch-based clade.[11] • A clade located within a clade is said to be nested within that clade. the latest common ancestors are not the same and therefore the group is polyphyletic. Although there is an ultimate common ancestor among the Tetrapods. The use of "basal" to mean a clade that is less species-rich than a sister clade. a group is polyphyletic because it is diagnosed by a character that actually forms a homoplasy. In the diagram.[12] • Node-based: the last common ancestor of A and B. excluding them from being reptiles. as birds are also descended from amniotes. The process of identifying and naming groups based on apomorphies is the method that most resembles classical systematics. Many cladograms are not of this type.g. or other data characterizing those species.[21] and it is appropriate to include extinct species in the group of organisms being analyzed. morphological data in the absence of fossils or other dating information) • When the characteristic data are DNA/RNA sequences. 1 cm = 1 million years). If a cladogram represents N species. The starting point of cladistic analysis is a group of species and molecular. if there are 32 species of deer.[19] with time running forward from the base of the tree to the leaves of the tree. all species lie at the leaves. because the various branches of the cladogram will not be uniformly deep.135 2.[16] Each subtree.*(2N-3) This superexponential growth of the number of possible cladograms explains why manual creation of cladograms becomes very difficult when the number of species is large. molecular sequencing data and computational phylogenetics are now very commonly used in the generation of cladograms. especially . This formula gives a lower limit. Such cladograms are called scaled cladograms. the time axis of the cladogram could be assigned a time scale (e. with a common ancestor and no additional descendents. A cladogram tree has an implicit time axis.459.395 135. and the forks of the tree could be graphically located along the time axis. Cladograms that are based on DNA/RNA generally do not include extinct species because DNA/RNA samples from extinct species are rare. although this is really just the lower limit. Cladograms are subject to revision as additional data become available. could be about 23 levels deep. the number of distinct cladograms that can be drawn (ignoring which cladogram best matches the species characteristics) is:[17] Number of species 2 3 4 5 6 7 8 9 10 N Number of cladograms 1 3 15 105 945 10. diagrams which show ancestral relations between taxa.[13] or sometimes a dendrogram (Greek for "tree drawing"). expressed as millions of years ago) of all the evolutionary forks were known.027. A cladogram representing the complete tree of life. Thus. The two taxa on either side of a split. Conversely. For a given set of species.[14] The cladogram graphically represents a hypothetical evolutionary process. those dates could be captured in the cladogram.g. The terms "evolutionary tree". Although traditionally such cladograms were generated largely on the basis of morphological characters.425 1*3*5*7*. morphological. for a variety of reasons: • They are built from species characteristics that cannot be readily dated (e. Cladograms based on morphology. are called "sister taxa" or "sister groups".025 34. with the actual depth generally a larger value. and sometimes "phylogenetic tree" are often used synonymously with cladogram. but converting those ages into actual years requires a significant approximation of the rate of change[20] • Even when the dating information is available. The end result is a tree-like relationship diagram called a cladogram. is called a "clade". it is feasible to use sequence differences to establish the relative ages of the forks. In cladograms. positioning the cladogram's forks along the time axis in proportion to their dates may cause the cladogram to become difficult to understand or hard to fit within a human-readable format Cladistics makes no distinction between extinct and extant species.[18] For example. Some cladograms include 3-way or 4-way forks when there are insufficient data to resolve the forking to a higher level of detail (see under phylogenetic tree).. the number of levels (the "depth") in the cladogram is on the order of log2(N). a cladogram representing deer could be around 5 levels deep (because 25 = 32). the depth may be shallower if forks larger than 2-way forks are permitted. with about 10 million species. Many cladists require that all forks in a cladogram be 2-way forks.. whether it contains only two or a hundred thousand items.[15] but others treat phylogenetic tree as a broader term that includes trees generated with a nonevolutionary emphasis. to represent the evolutionary tree of life. If the approximate date (for example.Cladistics 277 Cladograms Cladists use cladograms. " Willi Hennig's pioneering work provoked a spirited debate[24] about the relative merits of phylogenetic nomenclature versus Linnaean or evolutionary taxonomy.and lower-level taxa (i.[25] however Hennig did not advocate abandoning the Linnaean nomenclatural system. . because the evolutionary tree is so deep and so complex that it is inadvisable to set a fixed number of levels. also may subjectively consider similarity and has a fixed hierarchy of taxonomic ranks. which has continued down to the present. and its taxa are not required to correspond to clades.e. In contrast. and family. Phylogenetic nomenclature does not feature those terms. neo-Darwinism (summed up in the "evolutionary" systematics of Mayr and Simpson) not only lacked a definable investigatory method. The concept of phylogeny as mere genealogy should be terminologically distinguished as cladistics. such as kingdom. biological species and natural selection.Cladistics morphological characteristics that are preserved in fossils. Linnaean taxonomists have generally attempted to make at least family. class. The term phylogeny should be retained for the broad concept of phylogeny. phylum. Some of the debates in which the cladists were engaged had been running since the 19th century. while since the advent of evolutionary theory following phylogeny. automatically generated tree of life based on [22] completely sequenced genomes "It would seem to me to be quite evident that the two concepts of phylogeny (and their role in the construction of classifications) are sufficiently different to require terminological distinction. Linnaean taxonomy allows both monophyletic and paraphyletic groups as taxa. 278 Cladistics in taxonomy Phylogenetic nomenclature contrasted with traditional taxonomy Most taxonomists have used the traditional approaches of Linnaean taxonomy and later Evolutionary taxonomy to organize life forms. Nelson. but came to depend. Since the early 20th century. To lump the two concepts together terminologically could not help but produce harmful equivocation. are more likely to include extinct species. promoted by Darwin and adopted by most students of phylogeny in the ensuing 90 years.[26] as can be seen from the Foreword to Hennig (1979) by Rosen. those regulated by the codes of nomenclature) monophyletic. and Patterson:[27] "Encumbered with vague and slippery ideas about adaptation. Evolutionary taxonomy insists that groups reflect phylogenies. fitness. Linnaean taxonomy." Phylogenetic nomenclature strictly and exclusively follows phylogeny and has arbitrarily deep trees with binary branching: each taxon corresponds to a clade. Ernst Mayr in 1985 drew a distinction between the terms cladistics and phylogeny:[23] A highly resolved. on consensus or authority. order. These approaches use several fixed levels of a hierarchy. both for evolutionary interpretation and classification. but they were renewed fervor. as a viewpoint zooms into the tree of life. This definition gives us the Crown group tetrapods (or Crown-Tetrapoda). because they detract from cladistics' emphasis on clades (monophyletic groups).[29] Every single speciation event. that is. such as snakes). and it dissolves into many smaller lineages. In this definition. branching events are relatively regular. Look closely at the 'lineage' of a phylogeny . ecology." The overall shape of a dichotomous (bifurcating) tree is recursive. that both paraphyletic groups and clades are valuable notions with separate purposes. The tree of life is a quasi-self-similar fractal. and so on. Mammalia. which hypothesizes long periods of stability followed by punctuations of rapid speciation.[31] At this level Eldredge's and Gould's punctuated equilibrium applies. the early vertebrates that would someday evolve into the family Hominidae cannot be placed in any other monophyletic family. A few primitive four legged ancestors (the Ichthyostegalia) fall outside Crown-Tetrapoda. Reptilia. Linnaean taxonomy contains the taxon Tetrapoda.Cladistics 279 Paraphyletic groups discouraged Many cladists discourage the use of paraphyletic groups in classification of organisms. the deep reconstruction is not as regular as the shallow reconstruction. it is often possible to estimate the times between them. PhyloCode approach to naming species A formal code of phylogenetic nomenclature. proponents of the use of paraphyletic groups argue that any dividing line in a cladogram creates both a monophyletic section above and a paraphyletic section below. complete cladogram of the tree of life. based on the fossil record. the complexity and depth of the tree can be very large. the same type of tree appears no matter what the scale. and by deep the more ancient branches further back. Complexity of the Tree of Life The cladistic tree of life is a fractal:[28] "The tree of life is inherently fractal-like in its complexity. In the deep part of the tree. In the shallow part of the tree. or life history – in short.[33] An alternative is to define tetrapoda as all animals more closely related to mammals than to lungfish (our nearest living non-tetrapod relatives).[32] is currently under development.. When extinct species are considered (both known and unknown). In several instances (see for example Hesperornithes) it has been employed to clarify uncertainties in Linnaean systematics so that in combination they yield a taxonomy that unambiguously places problematic groups in the evolutionary tree in a way that is consistent with current knowledge. whose living members can be classified phylogenically as "the clade defined by the common ancestor of amphibians and mammals". It is intended for use by both those who would like to abandon Linnaean taxonomy and those who would like to use taxa and clades side by side. Aves. In contrast. "homology assessments" are "difficult" and the times vary widely. They also argue that paraphyletic taxa provide information about significant changes in organisms' morphology. Phylogenetic nomenclature also contains the taxon Tetrapoda (see the diagram under Clades above). Example For example. a series of events called fractal evolution.[30] By shallow Mishler means the most recent branching toward and at the tips. and divides into the classes Amphibia. which are harder to reconstruct and are missing unknown extinct lines. including all the species that are now extinct. represents an additional fork on the hypothetical. together with a host of fossil animals usually classed as . defined morphologically as vertebrates with four limbs (as well as animals with four-limbed ancestors.. They also contend that paraphyletic taxa are necessary for classifying earlier sections of the tree – for instance. the ichthyostegalians are included. or more precisely the clade defined by the common ancestor of a specific amphibian and mammal (or bird or snake).. ... down to a very fine scale. the PhyloCode. Moreover the tree continues to recreate itself by bifurcation. which is often given the rank of superclass. regardless of evolutionary mechanism . and Kingdoms Summary of criticisms of phylogenetic nomenclature Critics of phylogenetic nomenclature include Ashlock. Amniota is divided into the two clades Sauropsida (which contains birds and all living amniotes other than mammals. the latter of which the amiotes is a sub-clade.[38] Williams. parvorder.Cladistics crossopterygian fish. Orders. Non of the phylogenetic taxa as described above have a rank. Aves and Mammalia are subclades.g. contained in the subclade Amniota. infraorder.[39] Some of their criticisms include: Phylogenetic Nomenclature Limited to entities related by evolution or ancestry Linnaean Taxonomy Supports groupings without reference to evolution or ancestry Includes a process for giving unique names to species Taxa definitions based on tangible characteristics Permits clearly defined groups such as reptiles Straightforward process to determine if a given species is in a taxon or not Does not include a process for naming species Clade definitions emphasize ancestry at the expense of descriptive characteristics Ignores sensible. while Reptilia and Amphibia are paraphyletic taxa. A third option is to define Tetrapoda according to their apomorphy (their unique trait. not clades. All the subclades are contained within one another. This wider definition is termed Pan-Tetrapoda. including all living traditional reptiles) and Theropsida (mammals and the extinct mammal-like reptiles). Amphibia can be split into the Batrachomorpha (fossil amphibians more closely related to modern amphibians) and Reptiliomorpha. clearly defined paraphyletic groups such as reptiles Difficult to determine if a given species is in a clade or not (e. magnorder) to accommodate new discoveries.[35] Ichthyostegalians and other Stem-tetrapods represent sister groups from splits predating the Batrachomorpha/Reptilopmorpha split. not applicable to organisms that evolved via complex gene sharing or lateral transfer Applicable to all organisms.[37] Mayr.[34] Instead of classifying non-mammalian.e. then the only way to determine if species Y is in the clade is to perform a complex evolutionary analysis) Limited to organisms that evolved by inherited traits. Biased towards trees about 4 to 12 levels deep. The clades are not divided into several non-overlapping taxa (as in traditional taxonomy). non-avian amniotes as reptiles. suborder. having legs rather than fins). and neither do its subtaxa. if clade X is defined as "most recent common ancestor of A and B along with its descendants".[33] 280 Summary of advantages of phylogenetic nomenclature Proponents of phylogenetic nomenclature enumerate key distinctions between phylogenetic nomenclature and Linnaean taxonomy as follows:[36] Phylogenetic Nomenclature Handles arbitrarily deep trees. With regards to the traditional classes. i. a definition that yield the same group as the Linnaean taxon. Acceptable to name and use paraphyletic groups Discourages naming or use of groups that are not monophyletic Primary goal is to reflect actual process of evolution Assumes that the shape of the tree will change frequently with new discoveries Primary goal is to group species based on morphological similarities New discoveries often require renaming or releveling of Classes. Linnaean Taxonomy Often must invent new level names (such as superorder. a process repeated throughout. rather the clade is split into two clades at the first branching. Similarly. . Oxford. F. 164. P. 78. 221 [8] Patterson 1982. "Homology in classical and molecular biology". .[41] Compares cultures or artifacts using groups of cultural traits or artifact features. (2005).Cladistics 281 Application to other disciplines The comparisons used to acquire data on which cladograms can be based are not limited to the field of biology. doi:10. "Toward a phylogenetic system of biological nomenclature". . doi:10.1023/A:1006037525704. "Natural History Collections: Cladistics" (http:/ / www. p. C. Retrieved 4 July 2009. [17] Lowe. [10] de Pinna.G. "branch") [2] Glossary entry "clade" (http:/ / evolution. com/ ?id=LalmQ4346O0C& dq=Nielsen.G (1991). [7] Mayr 1982. p. pp. 3–14. In Koenemann. php?page=236. J. University of California Museum of Paleontology. pp.tb00045. 45. "Concepts and tests of homology in the cladistic paradigm". Science. canterburytalesproject. Retrieved 2008-10-15. 13. doi:10. (February 1998). + 2001+ "Animal+ evolution"+ Chelicerata). pp.[45] Compares animal species using behavioral traits presumed hereditary. Analysis. ISBN 1405100338. P. J. [5] Dupuis 1984 [6] Baron. [11] Krell.2004. ISBN 0387946233. ac. CRC Press. Andrew (2004). 555. Blackwell Publishing.1991. ISBN 1578083516.1111/j.00262.[43] [44] Compares manuscripts of the same work (original lost) using groups of distinctive copying errors.. [18] Aldous. 203–4. and to which a set of common characteristics may or may not apply. ..x.. edu/ evolibrary/ glossary/ glossary_popup. M. Gauthier. nhc. "Evolution and systematics of the Chelicerata".1096-0031. Jenner. Høeg. New York: Norton. hypothesized to have a common ancestor. Assembling the Tree of Life. berkeley. England: Oxford University Press.x. "Probability Distributions on Cladograms".0307-6970. uk/ index. Plant Systematics: An Integrated Approach. • Textual criticism or Stemmatics.-T.A. Recent attempts to use cladistic methods outside of biology address the reconstruction of lineages in: • Anthropology and archeology. 357. Experimental and Applied Acarology 22 (2): 63–79. "Which side of the tree is more basal?". [13] Cracraft.1016/0169-5347(94)90231-3. David (1996). Molecular Biology and Evolution 5 (6): 603–625. Cranston. • Ethology. pp.+ C. Steven Jay (1983). 273.[40] Any group of individuals or classes. 444). Systematic Entomology 29 (3): 279–281. Gurcharan (2004). "Gould. R. Cladograms can be used to depict the hypothetical descent relationships within groups of items in many different academic realms. [3] Phillips. ISBN 0849334985. Hen's Teeth and Horse's Toes. Ecological Genetics: Design. eds (2004). [4] "Canterbury Tales Project" (http:/ / www. Cladistics 7: 367–394. [15] Singh.1111/j. [16] Gould. php?word=clade) Understanding Evolution. 2010.T. google. Random Discrete Structures. pp. Retrieved 4 July 2009. (2004). Crustacea and Arthropod Relationships. Michael J. Donaghue. (1994). and Application.. S. Joel. The only requirement is that the items have characteristics that can be identified and measured. ed. klados. ISBN 0393017168. Colin (1988). doi:10.[42] Compares languages using groups of linguistic features. Scharm and the paleontologocal perspective in evolutionary biology" (http:/ / books. PMID 3065587. University of Edinburgh. can be compared pairwise. Springer. K. [14] Weygoldt... Trends in Research in Ecology and Evolution 9 (1): 27–31. • Linguistics. 21–74 [9] Patterson. p. Ben (2000). [12] de Queiroz. org). See also • • • • • • • • Bauplan Bioinformatics Biomathematics Coalescent theory Dendrogram Evolution of Mollusca Last common ancestor • • • • • • • Important publications in phylogenetics • Language family Maximum parsimony Molecular phylogeny PhyloCode Phylogenetics Phylogenetic network Phylogenetics software • • • • • • • Phylogenomics Phylogeography Phylogenetic comparative methods Scientific Classification Stratocladistics Systematics Tree structure Three-taxon analysis Computational phylogenetics • References [1] (Ancient Greek: κλάδος. doi:10. [39] Williams. p.2307/2412223. html). (1974). "Willi Hennig's impact on taxonomic thought". P... Scott (1998). ..org/stable/2412223). Kaufman. Quentin (2000). 83. "Cladistic analysis of an Old Norse manuscript tradition" (http:/ / rjohara. Peter D.2307/2413350. .05. Robert J. Science as a Process. Ruth. org/ stable/ 2413350). Evolutionary Analysis. Blackwell Science Ltd 3rd ed. uk/ benton/ vertclass.. Oxford: Oxford University Press. 1. M. WJ (2002). Biological Reviews 75 (4): 633–648. p. (1996). [42] Oppenheimer. Annual Review of Ecology and Systematics 15: 1–24.1093/bioinformatics/btl529.5537.. doi:10. [35] Benton.. 290–300. • Ashlock. [40] Mace. Collard. ix [28] Mishler 2005. 62: "In deep reconstruction problems. Claude (1984). ISBN 0521340861. (2001).1126/science.W. Species Concepts and Phylogenetic Theory: A Debate.). [30] Mishler 2005. [33] Clarck. crown clades. (2000).2307/2412559. doi:10.1469-185X. Harry J. Ashlock 1972. p. Vertebrate Paleontology. p.. Bork. "On Theory in Comparative Psychology". Journal of Zoological Systematics and Evolutionary Research 40: 169–194. Available free online at http://gallica.org/stable/ 2412559). "Darwin and the Definition of Phylogeny" (http:/ / jstor.org/stable/2412435). p. (1992). ISBN 0226360512. doi:10. This is the paper credited by Hennig (1979) for the first use of the term 'clade'. p. Peter D.1146/annurev. Science 293 (5537): 1979–1980. "An evolutionary systematist’s view of classification" (http://jstor. eds (2005). E (1985).. ISBN 0-19-860426-2. [38] Mayr 1974. The Variety of Life: A survey and a Celebration of all the Creatures that Have Ever Lived. N. "Evolutionary Biology: Preparing the Ground for a Modern 'Tree of Life'". World Scientific. Piscataway: Transaction Publishers. E. Carl. In Sternberg. Bart D.2307/2412435. Lucien (1940). according to Benton 2004 [36] Hennig 1975. Michael J. Colin (2002). • Ashlock. org/ Ichthyostega). ISSN 0066-4162. Systematic Zoology 34 (1): 97–98. Cambridge University Press.00211. (1979). [23] Mayr.. O’Hara. The text of the New Testament: its tranmission.x. James C. [37] Ashlock 1971. Robert Lynn (1997). ISBN 0231101430. 57 [29] Gordon.. P. (2000).293. p. R.Cladistics [19] Freeman. 83.2002. Prentice Hall.. ISBN 9810222688. doi:10. (1990).1439-0469. (1997): Ichthyostega (http:/ / tolweb. Mark et al. relative lengths of . p. Bock. Cambridge University Press. David (1988).1111/j. (1972). Shennan. Comptes Rendus de l'Académie des Sciences de Paris 210: 23–27. • Cuénot. "Monophyly again" (http://jstor. [21] Scott-Ram. Peter D. Systematic Zoology 20 (1): 63–69. Inc. [24] Wheeler. nodes. doi:10. Taxonomy and Evolution. I. "Classifications and other ordering systems". Clare. Robert J. M.1046/j. Systematic Zoology 21 (4): 430–438. 380. Bioinformatics 23 (1): 127–8. 232–276. [25] Benton. "Confusion in cladism". ISBN 0123852501. Columbia University Press.2000. 664. doi:10. 80.tb00055. 254. doi:10.A. net/ cv/ 1996-rhc). [43] Ehrman. Mahwah. 340–56. p. the branch events at issue happened a relatively long time ago and the . Stephen (2006).. ISBN 0135680239. J. 207 ff. [26] Hull. "Monophyly and associated terms" (http://jstor.000501. "The uses of cladistics". "Stems. [22] Letunic. [44] Robinson. gly. p. . ISSN 0066-4162. 2004 – see also taxonomic hierarchy of the vertebrates (http:/ / palaeo. bris.es.A.. [20] Carroll.fr (No direct URL). (2005). pp. (2003).. Mayr 1978. Synthese 01: 135–132. University of Chicago Press.. O'Brien. NJ: Lawrence Erlbaum Associates. The Origins of the British. 282 Bibliography • Ashlock. p. • Ashlock. The evolution of intelligence. Peter D. Cavendish Publishing. J. Research in Humanities Computing 4: 115–137. London: Robinson. PMID 11557858. [27] Hennig 1979.." [32] Pennisi. 2nd Ed. Peter M. branches are often quite different. E. Annual Review of Ecology and Systematics 5: 81–99. (1971). and rank-free lists: is Linnaeus dead?".x. Clare J. p. Mapping Our Ancestors: Phylogenetic Approaches in Anthropology and Prehistory. ISBN 052147809X. corruption and restoration (4th ed. Richard (1999). [41] Lipo. Mayr.1979. doi:10. ISBN 0786718900.bnf. ac. • Dupuis. New York: Oxford University Press. "Interactive Tree Of Life (iTOL): an online tool for phylogenetic tree display and annotation" (Pubmed). (2007). eds (2006). Ashlock 1979. 62 [31] Mishler 2005. pp. [45] Jerison. Patterns and Processes of Vertebrate Evolution. Transformed Cladistics. Systematic Zoology 28 (4): 441–450. PMID 17050570.110174. PMID 11117201. The Evolution of Cultural Diversity: A Phylogenetic Approach. The Hierarchical Genome and Differentiation Waves. "Remarques sur un essai d'arbre généalogique du règne animal". Ashlock 1974. Tree of Life website [34] Tudge. Stephen. IL: Univ. Parsimony. Problems in Phylogenetic Reconstruction. A. ISBN 0-252-06814-9. • Schuh. of Illinois Press (reprinted 1979 and 1999).org/faqs/comdesc/phylo.berkeley. Joe.. "Cladistic analysis or cladistic classification?". The Compleat Cladist: A Primer of Phylogenetic Procedures. ISBN 0-674-36446-5. Urbana. Friday. 19. MA: Harvard University Press. D. Ernst (1978). Willi (1979). Translated from manuscript and so never published in German." This is the paper to which Hennig 1975 is a response. Brent D. The TalkOrigins Archive.edu/clad/clad4. "Morphological characters and homology". • Felsenstein. "Report on the Textual Criticism Challenge 1991" (http://rjohara.. Retrieved 21 January 2010. doi:10. ISBN 0-12-391250-4. Includes a reprint of Mayr's 1974 anti-cladistics paper at pp. Peter.2307/2412765. "Origin and history of some terms in systematic and evolutionary biology" (http://jstor. • Mayr. Phylogenetic systematics (3rd edition of 1966 book). Seattle: University of Washington. Cambridge.Cladistics • Hennig. Retrieved 21 January 2010. Ernst (1982).palomar. MA: Harvard University Press. • Theobald. Guralnick. • Robinson. 433–476. html). • Hull. Dwight Davis and Rainer Zangerl).2307/2412818. Systematic Zoology 28 (4): 416–440. Brower. "Classification of Living Things" (http://anthro. • Mishler. Allen G. University of California Museum of Paleontology. "Chapter 1: Introduction. (1992).amnh. Colin (1982). "Phylogeny Programs" (http://evolution. Kenneth A.org/ stable/2412765).org/stable/2412558).talkorigins.2307/2412558. • Mayr. V. Biological Systematics: principles and applications (2nd edn. "'Cladistic analysis or cladistic classification': a reply to Ernst Mayr" (http://jstor. Oxford University Press. Z. • Hennig.washington. In Albert. Systematic Zoology 24 (2): 244–256. . (1979).O. Funk.ucmp. Victor. London: Academic Press.R.edu/animal/default. "The logic of the data matrix in phylogenetic analysis". Retrieved 21 January 2010. Willi (1966).. Phylogeny. Robert J.. Dave (1994-2005).html). ISBN 0893380350.org/ stable/2412818). D. • O'Neil. ISBN 0-252-06814-9. and Genomics. Retrieved 21 January 2010. 2009. ISBN 978-0-8014-4799-0 283 External links • Collins. Willi (1975).net. O'Hara. Cambridge. Urbana: University of Illinois Press.pdf).gs.. Smith. "Phylogenetics Primer" (http://www. In Joysey. Dennis (1998-2008). Evolution and the diversity of life (Selected essays). Douglas (1999-2004). rjohara. The growth of biological thought: diversity. The paper to which he was responding is reprinted in Mayr 1976. ISBN 0-674-27105-X. Systematic Zoology 27 (1): 83–88. Systematics Association Special Volume 21. (1991). • Mayr. • Patterson. Zeitschrift fűr Zoologische Systematik und Evolutionforschung 12: 94–128.A. evolution and inheritance.html).org/learn/pd/fish_2/pdf/compleat_cladist. (2005). Ernst (1974).net/ darwin/files/bmcr). htm). ISBN 0199297304. American Museum of Natural History. E. pp. doi:10. San Marcos CA: Palomar College. David L. Terms and Concepts" (http://www. "Journey into Phylogenetic Systematics" (http:// www. Rob. D.) Cornell University Press. Brooks. Phylogenetic systematics (tr. doi:10. "The limits of cladism" (http://jstor. "Cladistic analysis or cladistic classification. E. • Wiley. Reissued 1997 in paperback. • Mayr. 57–70. Randall T. and Andrew V. The University of Kansas Museum of Natural History Special Publication No. Siegel-Causey.edu/phylip/software. • Hennig. Retrieved 21 January 2010. Ernst (1976). As DNA sequencing has become cheaper and easier. the ancestor at the bottom . which incorporate explicit models of sequence evolution. maximum likelihood and Bayesian inference. or other genetic information). warm blooded. Consider possible cladograms 3. are non-Hennigian ways to evaluate sequence data. etc. skeletal anatomy.Cladogram 284 Cladogram A cladogram is a diagram used in cladistics which shows ancestral relations between organisms. if analyzing 20 species of birds. to represent the evolutionary tree of life. molecular systematics has become a more and more popular way to reconstruct phylogenies. They are also generally assumed to have a low incidence of homoplasies A vertical cladogram. which are thought to be less prone to the problem of reversion that plagues sequence data. with the ancestor at the top Two vertical cladograms. all cladistic analysis used morphological data. DNA and RNA sequencing data and computational phylogenetics are now very commonly used in the generation of cladograms. Gather and organize data 2. Select best cladogram Step 1 A cladistic analysis begins with the following data: • a list of species to be organized • a list of characteristics to be compared • for each species.[2] Using a parsimony criterion is only one of several methods to infer a phylogeny from molecular data. Another powerful method of reconstructing phylogenies is the use of genomic retrotransposon markers.) or molecular (DNA.[1] Prior to the advent of DNA sequencing. the value of each of the listed characteristics or character states For example. the data might be: • the list of 20 species • characteristics such as genome sequence. biochemical processes. its particular genome sequence. unicellular. and feather coloration • for each of the 20 species. and feather coloration Molecular versus morphological data The characteristics used to create a cladogram can be roughly categorized as either morphological (synapsid skull. Although traditionally such cladograms were generated largely on the basis of morphological characters. A horizontal cladogram. with the ancestor (not named) to the left Generating a cladogram A greatly simplified procedure for generating a cladogram is:[1] 1. skeletal anatomy. RNA. biochemical processes. notochord. and insects serve the same function. The choice of an outgroup is a crucial step in cladistic analysis because different outgroups can produce trees with profoundly different topologies. Though the wings of birds.[3] Typically. Plesiomorphies and synapomorphies The researcher must decide which character states were present before the last common ancestor of the species group (plesiomorphies) and which were present in the last common ancestor (synapomorphies) and does so by comparison to one or more outgroups. possibly resulting in a false evolutionary scenario. but character reversions that are unrecognizable as such are more common in the latter (see long branch attraction). a homoplasy would be introduced into the dataset. "presence of wings". molecular. If a bird. "presence of wings". Note that only synapomorphies are of use in characterizing clades. morphological. Morphological homoplasies can usually be recognized as such if character states are defined with enough attention to detail. bat. and this would confound the analysis. as otherwise the characters might not apply at all to a large number of taxa. Ideally. this seems at least sometimes not to be the case. Use of homoplasies when building a cladogram is sometimes unavoidable but is to be avoided when possible. When analyzing "supertrees" (datasets incorporating as many taxa of a suspected clade as possible). no method of evaluation is likely to produce a phylogeny representing the evolutionary reality.Cladogram because it was once thought that their integration into the genome was entirely random. as can be seen by their anatomy. and a winged insect were scored for the character. and possibly other phylogenies should be combined into an analysis of total evidence: All have different intrinsic sources of error. however. character convergence (homoplasy) is much more common in morphological data than in molecular sequence data. 285 Apomorphy in cladistics . bats. For example. Cautious choice and definition of characters thus is another important element in cladistic analyses. Avoid homoplasies A homoplasy is a character that is shared by multiple species due to some cause other than common ancestry. homoplasies occur due to convergent evolution. to continue with the "wings" example. each evolved independently. it may become unavoidable to introduce character definitions that are imprecise. A well known example of homoplasy due to convergent evolution would be the character. as most of these don't have wings at all. Homoplasies can often be avoided outright in morphological datasets by defining characters more precisely and increasing their number. the presence of wings would hardly be a useful character if attempting a phylogeny of all Metazoa. With a faulty outgroup or character set. [6] Most algorithms use a metric to measure how consistent a candidate cladogram is with the data. Evaluate the candidate cladograms by comparing them to the characteristic data 2. In general. Step 3 Simple cladistics There are several algorithms available to identify the "best" cladogram. Repeat these steps until the cladograms stop getting better Computer programs that generate cladograms use algorithms that are very computationally intensive. A typical cladistic program begins by using heuristic techniques to identify a small number of candidate cladograms.Cladogram 286 Step 2 When there are just a few species being organized. but most cases require a computer program. Many cladistic programs then continue the search with the following repetitive steps: 1. Because the total number of possible cladograms grows exponentially with the number of species. other algorithms are useful only when the characteristic data are morphological. the user should input the data in various orders and compare the results. and Bayesian inference. In these situations. parsimony. There are scores of computer programs available to support cladistics.[5] because the cladogram problem is NP-hard. neighbor-joining. cladogram generation algorithms must be implemented as computer programs. Algorithms for cladograms include least squares. Inputting the data in various orders can cause the same algorithm to produce different "best" cladograms. it is impractical for a computer program to evaluate every individual cladogram. Biologists sometimes use the term parsimony for a specific kind of cladogram generation algorithm and sometimes as an umbrella term for all cladogram algorithms. Use heuristics to create several new candidate cladograms unrelated to the prior candidates 5. Create additional candidates by creating several variants of each of the best candidates from the prior step 4. although some algorithms can be performed manually when the data sets are trivial (for example. Most cladogram algorithms use the mathematical techniques of optimization and minimization.[7] Algorithms that perform optimization tasks (such as building cladograms) can be sensitive to the order in which the input data (the list of species and their characteristics) is presented. . Other algorithms can be used when the characteristic data includes both molecular and morphological data. it is possible to do this step manually. maximum likelihood.[4] See phylogenetic tree for more information about tree-generating computer programs. Some algorithms are useful only when the characteristic data are molecular (DNA. just a few species and a couple of characteristics). Identify the best candidates that are most consistent with the characteristic data 3. Using different algorithms on a single data set can sometimes yield different "best" cladograms. RNA). because each algorithm may have a unique definition of what is "best". 1038/361603a0. pars. Stewart. West-Eberhard. Oxford University Press. Nature 361 (6413): 603–607. David (1996). Dessauer (herpetology). DNA-DNA hybridization was the dominant technique.[8] To help solve this problem. followed by Allan C. for example. Sibley (birds). Developmental Plasticity and Evolution.Cladogram Because of the astronomical number of possible cladograms. html). Birkhauser. Linus Pauling and Walter M. is the use of the structure of molecules to gain information on an organism's evolutionary relationships. Techniques in Molecular Systematics and Evolution. Conserved sequences. many cladogram algorithms use a simulated annealing approach to increase the likelihood that the selected cladogram is the optimal one. Press. In the period of 1974–1986. (1999). The result of a molecular phylogenetic analysis is expressed in a phylogenetic tree. ISBN 0878932828. ISBN 376436257X. they provided tantalizing hints that long-held notions of the classifications of birds. Caro-Beth (1993). and Morris Goodman (primates). PMID 8437621. 61–128. Peter (1993). p. and John C. Herbert C. genetics. pp. "The Powers and Pitfalls of Parsimony". as well as the emergence of a new branch of criminal forensics focused on evidence known as genetic fingerprinting. Hillis. Oxford Univ. Oxford Univ. 66. algorithms cannot guarantee that the solution is the overall best solution. washington.x. Another application of the techniques that make this possible can be seen in the very limited field of human genetics. Mary Jane (2003). Emanuel Margoliash.1111/j. History of molecular phylogenetics The theoretical frameworks for molecular systematics were laid in the 1960s in the works of Emile Zuckerkandl. Molecular Systematics. Ian (1998).1096-0031. and proteins. The most common approach is the comparison of homologous sequences for genes using sequence alignment techniques to identify similarity. Another application of molecular phylogeny is in DNA barcoding.[9] 287 References [1] [2] [3] [4] [5] [6] [7] [8] [9] DeSalle. such as the ever more popular use of genetic testing to determine a child's paternity. Avise (who studied various groups). where the species of an individual organism is identified using small sections of mitochondrial DNA. Closely related organisms generally have a high degree of agreement in the molecular structure of these substances. Rob (2002). are expected to accumulate mutations over time. Although the results were not quantitative and did not initially improve on morphological classification. A nonoptimal cladogram will be selected if the program settles on a local minimum rather than the desired global minimum. Molecular phylogeny uses such data to build a "relationship tree" that shows the probable evolution of various organisms. Cladistics: The Theory and Practice of Parsimony Analysis. Robert K. has it been possible to isolate and identify these molecular structures. ISBN 0195122356. Cladistics 15: 407–414. doi:10. such as mitochondrial DNA. needed substantial revision. 353–376. Foley. "The Parsimony Ratchet: a new method for rapid parsimony analysis". Work with protein electrophoresis began around 1956. CRC Press. Trevor (2006). .tb00277. Press. RNA.[2] Techniques and applications Every living organism contains DNA. Sinaur.[1] Applications of molecular systematics were pioneered by Charles G. Wilson. Nixon K. Molecular phylogenetics Molecular phylogenetics. Not until recent decades. Hodkinson. Kitching. and assuming a constant rate of mutation provide a molecular clock for dating divergence. Fitch. . however. also known as molecular systematics. while the molecules of organisms distantly related usually show a pattern of dissimilarity. Cladistics: A Practical Course in Systematics. p. C. edu/ phylip/ software. ISBN 0849395798. ISBN 0198577664. Reconstructing the Tree of Life: Taxonomy and Systematics of Species Rich Taxa. ISBN 0198501382.1999. "List of Cladistics Software Programs" (http:/ / evolution. Selander. doi:10. haplotypes from a smaller number of individuals from a definitely different taxon are determined: these are referred to as an out group. The base sequences for the haplotypes are then compared. since there are four base types. the difference between two haplotypes is assessed by counting the number of locations where they have different bases: this is referred to as the number of substitutions (other kinds of differences between haplotypes can also occur. rather than on particular sections of DNA. and this has been done for only a few species. this assumption could be wrong if either some genotypic modification acted to prevent interbreeding between two groups of organisms. Usually the difference between organisms is re-expressed as a percentage divergence. the resulting triangular matrix of differences is submitted to some form of statistical cluster analysis. Any group of haplotypes that are all more similar to one another than any of them is to any other haplotype may be said to constitute a clade. taxa are also determined. In the simplest case. for organisms within a particular species or in a group of related species. with 1000 base pairs. However. Molecular systematics often uses the molecular clock assumption that quantitative similarity of genotype is a sufficient measure of the recency of genetic divergence. The advantage claimed for using hybridisation rather than gene sequencing was that it was based on the entire genotype. These have been largely replaced in recent times by DNA sequencing which produces the exact sequences of nucleotides or bases in either DNA or RNA segments extracted using different techniques. In a molecular systematic analysis. Limitations of molecular systematics Molecular systematics is an essentially cladistic approach: it assumes that classification must correspond to phylogenetic descent. Particularly in relation to speciation. and that all valid taxa must be monophyletic. Statistical techniques such as bootstrapping and jackknifing help in providing reliability estimates for the positions of haplotypes within the evolutionary trees. and the resulting dendrogram is examined in order to see whether the samples cluster in the way that would be expected from current ideas about the taxonomy of the group. it has been found empirically that only a minority of sites show any variation at all and most of the variations that are found are correlated. but supposedly different. carbohydrates and other molecules which were separated and characterized using techniques such as chromatography. An older and superseded approach was to determine the divergences between the genotypes of individuals by DNA-DNA hybridisation. In principle. or genetic modification proceeded at different rates in different subgroups of the organisms. At present it is still a long and expensive process to sequence the entire DNA of an organism (its genome). or not. ideally a substantial sample of individuals of the target species or other taxon are used however many current studies are based on single individuals. These are generally considered superior for evolutionary studies since the actions of evolution are ultimately reflected in the genetic sequences. Haplotypes of individuals of closely related. the bases found in a given position may vary between organisms. the haplotypes are determined for a defined area of genetic material. However it is quite feasible to determine the sequence of a defined area of a particular chromosome. Modern sequence comparison techniques overcome this objection by the use of multiple sequences.Molecular phylogenetics 288 Theoretical background Early attempts at molecular systematics were also termed as chemotaxonomy and made use of proteins. Typical molecular systematic analyses require the sequencing of around 1000 base pairs. we could have 41000 distinct haplotypes. so that the number of distinct haplotypes that are found is relatively small. by dividing the number of substitutions by the number of base pairs analysed: the hope is that this measure will be independent of the location and length of the section of DNA that is sequenced. The particular sequence found in a given organism is referred to as its haplotype. enzymes. Finally. Once the divergences between all pairs of samples have been determined. for example the insertion of a section of nucleic acid in one haplotype that is not present in another). . At any location within such a sequence. J. Dordrecht. (1992) Molecular systematics of plants. Blackwell Science. • Soltis. ISBN 0-87893-282-8. Soltis.Molecular phylogenetics In animals. objectivity. ISBN 0-86542-889-1.html) . and Doyle. R. vol. 2nd ed.S.J. P.. edu/ sora/ Auk/ v116n03/ index.J.gov/About/primer/phylo. ISBN 0-87893-177-5. D. no. 1998..S. and Doyle. & Moritz. D. Phil. E. & Biomed. J. and the construction of molecular phylogenies. it is often convenient to use mitochondrial DNA for molecular systematic analysis. ISBN-0-41211-131-4. because in mammals mitochondria are inherited only from the mother.nlm. 289 Further reading • Felsenstein.ncbi. See also • • • • molecular evolution computational phylogenetics PhyloCode Microbial phylogenetics References [1] Edna Suárez-Díaz & Victor H. Chapman & Hall. Oxford. & Holmes. New York. Sibley: A commentary on 30 years of collaboration. (1998) Molecular Systematics of Plants II: DNA Sequencing. London. • Hillis. 3 (July 1999). However.. php). Anaya-Muñoz (2008) History. Jon E.nih. C. M. P. Kluwer Academic Publishers Boston. D. Molecular evolution: a phylogenetic approach. Sinauer Associates Incorporated. 2004. Inferring phylogenies. 39:451–468 [2] Ahlquist.. The Auk. Sinauer Associates Incorporated.E.Systematics and Molecular Phylogenetics (http://www. 1999: Charles G.E. External links • NCBI . 1996. Molecular systematics. • Page. J. • Soltis. Soltis. Biol. A PDF or DjVu version of this article can be downloaded from the issue's table of contents page (http:/ / elibrary. M.. D. ISBN-0-41202-231-1. unm. C. 116. because inheritance in the paternal line might not be detected. this is not fully satisfactory. Sci. Hist. Stud. Evo-devo seeks the genetic and evolutionary basis for the division of the embryo into distinct modules. Often these parts are repeated. it is well understood how genetic mutation occurs. Currently. how ecology impacts in development and evolutionary change. and to discover how developmental processes evolved. and vertebrates. however. and for the partly independent development of such modules.290 Fields Evolutionary developmental biology Evolutionary developmental biology (evolution of development or informally. the origins of evolutionary developmental biology come from both an improvement in molecular biology techniques as applied to development. General hypotheses remain hard to test because organisms differ so much in shape and form. It addresses the origin and evolution of embryonic development. . Others.[1] the role of developmental plasticity in evolution. how modifications of development and developmental processes lead to the production of novel features. such as fingers. but can be distinguished from earlier approaches to evolutionary theory by its focus on a few crucial ideas. Evolutionary developmental biology studies how the dynamics of development determine the phenotypic variation arising from genetic variation and how that affects phenotypic evolution (specially its direction). Evolutionary developmental biology is not yet a unified discipline. One of these is modularity: as has been long recognized. and the developmental basis of homoplasy and homology. At the same time evolutionary developmental biology also studies how development itself evolves. It was not until the 1980s and 1990s. biologists incorporated Gregor Mendel's principles of genetics to explain both. and variation. when more comparative molecular sequence data between different kinds of organisms was amassed and detailed.[2] Although interest in the relationship between ontogeny and phylogeny extends back to the nineteenth century. heredity. Thus. such as the evolution of feathers. and the full appreciation of the limitations of classic neo-Darwinism as applied to phenotypic evolution. evo-devo) is a field of biology that compares the developmental processes of different organisms to determine the ancestral relationship between them. drawing on findings of discordances between genotype and phenotype and epigenetic mechanisms of development. developmental mechanisms are not understood sufficiently to explain which kinds of phenotypic variation can arise in each generation from variation at the genetic level.[4] [5] Initially the major interest has been in the evidence of homology in the cellular and molecular mechanisms that regulate body plan and organ development. insects. plants and animal bodies are modular: they are organized into developmentally and anatomically distinct parts. ribs.[6] Basic principles Charles Darwin's theory of evolution is based on three principles: natural selection. At the time that Darwin wrote. however. resulting in the modern synthesis. the principles underlying heredity and variation were poorly understood. However.[3] Nevertheless. evo-devo demonstrates that evolution alters developmental processes (genes and gene networks) to create new and novel structures from the old gene networks (such as bone structures of the jaw deviating to the ossicles of the middle ear) or will conserve (molecular economy) a similar program in a host of organisms such as eye development genes in molluscs. In the 1940s. are mounting an explicit challenge to neo-Darwinism. it now appears that just as evolution tends to create new genes from parts of old genes (molecular economy). However more modern approaches include developmental changes associated with speciation. that an understanding of the molecular basis of the developmental mechanisms has arisen. and body segments. Some evo-devo researchers see themselves as extending and enhancing the modern synthesis by incorporating into it findings of molecular genetics and developmental biology. the contemporary field of evo-devo has gained impetus from the discovery of genes regulating embryonic development in model organisms. Jean-Pierre Changeux and François Jacob discovered within the bacterium Escherichia coli a gene that functioned only when "switched on" by an environmental stimulus.[9] In the 1850s Owen began to support an evolutionary view that the history of life was the gradual unfolding of a teleological divine plan. scientists discovered specific genes in animals. organismal form can be influenced by mutations in promoter regions of genes. and the way in which von Baer's branching pattern matched his own idea of descent with modification[12] : . some of which act as structural components of cells and others as enzymes that regulate various biochemical pathways within an organism. just regulating them differently. but was opposed by Karl Ernst von Baer's embryology of divergence in which embryonic parallels only applied to early stages where the embryo took a general form. evolution can proceed by a "phenotype-first" route. not only protein-specifying sequences. the genes of the "developmental-genetic toolkit". also called the biogenetic law or embryological parallelism. see below).[11] In On the Origin of Species (1859). that act as switches for other genes. Darwin recognised the importance of embryonic development in the understanding of evolution. The modification of existing. The anatomist Richard Owen used this to support his idealist concept of species as showing the unrolling of a divine plan from an archetype. this finding suggested that the crucial distinction between different species (even different orders or phyla) may be due less to differences in their content of gene products than to differences in spatial and temporal expression of conserved genes. notably by Robert Edmond Grant. including a subgroup of the genes which contain the homeobox DNA motif. Jacques Monod. Similarly. The case for this was argued for by Mary Jane West-Eberhard in her 2003 book Developmental plasticity and evolution.[7] Later. rather than initiating.. however. the basis of the recognition that organismal phenotypes are not uniquely determined by their genotypes. Genes specify proteins. and could be induced by other gene products. biochemical pathways (and. and dependent on external or environmental inputs. called Hox genes. the evolution of new species of organisms) depended on specific genetic mutations.Evolutionary developmental biology Another central idea is that some gene products function as switches whereas others act as diffusible signals. and became a prominent part of his version of Lamarckism leading to disagreements with Georges Cuvier. and in the 1830s attacked the transmutation of species proposed by Lamarck. It was widely supported in the Edinburgh and London schools of higher anatomy around 1830. Charles Darwin proposed evolution through natural selection.[8] 291 History An early version of recapitulation theory. was put forward by Étienne Serres in 1824–26 as what became known as the "Meckel-Serres Law" which attempted to provide a link between comparative embryology and a "pattern of unification" in the organic world. In 1961.[3] [8] with genetic change following.g.[10] in a continuous "ordained becoming". that act analogously to the external stimuli in bacteria. Another focus of evo-devo is developmental plasticity. ultimately. suggested that the action of natural selection on promoters responsive to Hox and other "switch" genes may play a major role in evolution. morphogens. a theory central to modern biology. Most biologists working within the modern synthesis assumed that an organism is a straightforward reflection of its component genes. Geoffroy and Grant. rather than being always active. If generation of phenotypes is conditional. The implication that large evolutionary changes in body morphology are associated with changes in gene regulation. or evolution of new. In addition to providing new support for Darwin's assertion that all organisms are descended from a common ancestor. those DNA sequences at which the products of some genes bind to and control the activity of the same or other genes. after which more specialised forms diverged from this shared unity in a branching pattern. rather than the evolution of new genes. with new species appearing by natural birth. the formation of morphological and other phenotypic novelties. fruit flies and human beings) may use the same genes for embryogenesis (e. These discoveries drew biologists' attention to the fact that genes can be selectively turned on and off. and that highly disparate organisms (for example. It was supported by Étienne Geoffroy Saint-Hilaire as part of his ideas of idealism. [15] D'Arcy Thompson postulated that differential growth rates could produce variations in form in his 1917 book On Growth and Form. Hox genes. it served as a backdrop for a renewed interest in the evolution of development after the modern evolutionary synthesis was established (roughly 1936 to 1947).[17] The developmental-genetic toolkit The developmental-genetic toolkit consists of a small fraction of the genes in an organism's genome whose products control its development. transcription factors containing the more broadly distributed homeobox protein-binding DNA motif. proposed that "ontogeny recapitulates phylogeny. why humans. or not expressed at all." that is. . and indeed all vertebrates. Lewis discovered homeotic genes. Thus. Edward B. cell surface receptor proteins.[20] This same gene determines the spot pattern in butterfly wings. as if every evolutionary advance was added as new stage by reducing the duration of the older stages. and pattern of body parts. which in turn form the body plan of the organism. by combinatorial specifying the identity of particular body regions. either by the genes changing their expression pattern or acquiring new functions. These genes are highly conserved among Phyla. even if mouse Pax6/eyeless was expressed in Drosophila. and encode for the production of transcription factors. generating spatial and temporal patterns.[16] and an entire 2005 issue of the Journal of Experimental Zoology Part B: Molecular and Developmental Evolution was devoted to the key evo-devo topics of evolutionary innovation and morphological novelty. in his endeavour to produce a synthesis of Darwin's theory with Lamarckism and Naturphilosophie. A paragon of a toolbox gene is Pax6/eyeless. which controls eye formation in all animals. Among the most important of the toolkit genes are those of the Hox gene cluster. for a natural [13] classification of course includes all ages.[14] This was extended by the more general idea of heterochrony (changes in timing of development) as a mechanism for evolutionary change. in Geoffroy's linear model rather than Darwin's idea of branching evolution. A good example of the first is the enlargement of the beak in Darwin's Large Ground-finch (Geospiza magnirostris). Stephen Jay Gould called this approach to explaining evolution as terminal addition. the development of the embryo of every species (ontogeny) fully repeats the evolutionary development of that species (phylogeny). in the regions where limbs would form in other tetrapods. ” Ernst Haeckel (1866). in which the gene BMP is responsible for the larger beak of this bird. In 2000. The majority of toolkit genes are components of signaling pathways. identity. or complex. cell adhesion proteins. In this case the gene Distal-less is very under-expressed.[18] This means that a big part of the morphological evolution undergone by organisms is a product of variation in the genetic toolkit. rooting the emerging discipline of evo-devo in molecular genetics. a special section of the Proceedings of the National Academy of Sciences (PNAS) was devoted to "evo-devo". Hox genes determine where limbs and other body segments will grow in a developing embryo or larva. relative to the other finches. and secreted morphogens.[12] Haeckel's concept explained. It has been found to produce eyes in mice and Drosophila. function in patterning the body axis. His theory has since been discredited. all of these participate in defining the fate of undifferentiated cells.[19] The loss of legs in snakes and other squamates is another good example of genes changing their expression pattern.[21] which shows that the toolbox genes can change their function. The idea was based on observations of neoteny.Evolutionary developmental biology 292 “ We can see why characters derived from the embryo should be of equal importance with those derived from the adult. for example. have gill slits and tails early in embryonic development. However. Differences in deployment of toolkit genes affect the body plan and the number. He showed the underlying similarities in body plans and how geometric transformations could be used to explain the variations. translocation and duplication of genes. providing a basis for early macroevolutionary changes. The discovery of the homeotic Hox gene family in vertebrates in the 1980s allowed researchers in developmental biology to empirically assess the relative roles of gene duplication and gene regulation with respect to their importance in the evolution of morphological diversity.[23] 293 Development and the origin of novelty Among the more surprising and. there is an apparent paradox: "where we most expect to find variation. we find conservation. Newman and Gerd B. also tend to evolve the same function convergently or in parallel. there is significant variation in limb morphologies amongst salamanders and in differences in segment number in centipedes. or.[26] These researchers argue that the combinatorial nature of transcriptional regulation allows a rich substrate for morphological diversity. as Gerhart and Kirschner have noted.[8] The biologists Stuart A. the occurrence of novel forms within a population does not generally correlate with levels of genetic variation sufficient to account for all morphological diversity. counterintuitive (from a neo-Darwinian viewpoint) results of recent research in evolutionary developmental biology is that the diversity of body plans and morphology in organisms across many phyla are not necessarily reflected in diversity at the level of the sequences of genes. but is controlled by the same genes.[28] . for evo-devo studies. including those of the developmental genetic toolkit and other genes involved in development. at a finer scale. perhaps. including Sean B. even when the respective genetic variation is low. pattern. Classic examples of this are the already mentioned Distal-less gene. which states that morphological evolutionary novelty is generated by regulatory changes in various members of a large set of conserved mechanisms of development and physiology. but rather to alterations in gene regulation. which is responsible for appendage formation in both tetrapods and insects. a lack of change". A major question then. For example. is: If the morphological novelty we observe at the level of different clades is not always reflected in the genome.Evolutionary developmental biology Toolbox genes. Indeed. Epigenetic alterations of gene regulation or phenotype generation that are subsequently consolidated by changes at the gene level constitute another class of mechanisms for evolutionary innovation. where does it come from? Apart from neo-Darwinian mechanisms such as mutation. as well as being highly conserved. the generation of wing patterns in the butterflies Heliconius erato and Heliconius melpomene.[22] The previous supports Kirschner and Gerhardt's theory of Facilitated Variation. but highly divergent regulatory mechanisms for these genes. These butterflies are Müllerian mimics whose coloration pattern arose in different evolutionary events. since variations in the level. novelty may also arise by mutation-driven changes in gene regulation. Carroll of the University of Wisconsin–Madison suggest that "changes in the cis-regulatory systems of genes" are more significant than "changes in gene number or protein function". Epigenetic changes include modification of the genetic material due to methylation and other reversible chemical alteration [27] . has introduced an important new element into evolutionary theory. The finding that much biodiversity is not due to differences in genes. or timing of gene expression may provide more variation for natural selection to act upon than changes in the gene product alone.[25] Diverse organisms may have highly conserved developmental genes. Müller have suggested that organisms early in the history of multicellular life were more susceptible to this second category of epigenetic determination than are modern organisms. as distinct from the functioning of the individual genes in the network. Several biologists. resulting from the interaction and timing of activity of gene networks. Changes in gene regulation are "second-order" effects of genes.[24] Even within a species. as well as nonprogrammed remolding of the organism by physical and other environmental effects due to the inherent plasticity of developmental mechanisms. [16] Goodman CS and Coughlin BS (Eds).1126/science. Callaerts. Proceedings of the National Academy of Sciences 94 (6): 2421–2426.5595. 190-191 [13] Darwin. [8] West-Eberhard. [7] Monod. Joram (1997). 120–128. RA (2004). doi:10. Evolution (http:/ / www. . M-J. (2003). ISBN 978-1-4051-0345-9. doi:10. ISBN 0-674-63940-5. Lidia.Evolutionary developmental biology 294 See also • • • • • • • • • • • • • • • • • • • • • Developmental biology Animal evolution Plant Evolutionary Developmental Biology Baldwin effect Developmental systems theory Evolution of multicellularity Genetic assimilation Ontogeny Ontogeny recapitulates phylogeny List of gene families Important publications in evolutionary developmental biology Evolution and Development Leading journal Body plan Cell signaling Signal transduction Cell signaling networks Transcription factor Enhancer Enhanceosome promoter (biology) Gene regulatory network References [1] Prum. 252–253 [10] Bowler 2003. Massachusetts: Harvard University Press. Stephen Jay (1977).. Changeux. "Special feature: The evolution of evo-devo biology" (http:/ / www. (2000). 337–340 Secord 2003. Desplan. PMID 12411686.4424. (2000).2000.9. London: John Murray.97. pp. 424. Evolution & Development 2: 177–178. Kos. [5] Pichaud. doi:10. PMID 13936070. pp. pnas. A.1046/j. Claude (August 2002). doi:10. JP.1016/S0959-437X(02)00321-0. Brian K. Rina. Journal of Molecular Biology 6: 306–329. full).2421. Walter.1103707. [15] Gould. . blackwellpublishing. Gehring.953. 208 Secord 2003. Franck. Patrick. PMID 12616863.298. pp. (March 2003).O. On the Origin of Species. "Evo-devo or devo-evo—does it matter". Jacob. ISBN 0801413192.1073/pnas.6. pp. 439–430 (http:/ / darwin-online. pp.94. Developmental plasticity and evolution. PMID 15514148. Current opinion in genetics and development 12 (4): 430–434. org. Piatigorsky. [6] Pennisi.. "Squid Pax-6 and eye development" (http:/ / www. ISBN 978-0-19-512235-0. Halder. Science 306 (5697): 828–833. .H. com/ ridley/ ). doi:10. Scientific American 288 (3): 84–93. Stanislav I. Wiley-Blackwell. Mark (2003).x. "Symmetry breaking and the evolution of development". [4] Tomarev. Science 298 (5595): 953–955. doi:10. Proceedings of the National Academy of Sciences 97 (9): 4424–4456. [2] Hall. Georg.00003e.1038/scientificamerican0303-84.1126/science. "EVOLUTIONARY BIOLOGY:Evo-Devo Enthusiasts Get Down to Details". F (1963). PMID 12100888. "Which Came First. E (2002). Ontogeny and Phylogeny. doi:10. 490–491 [12] Bowler 2003. the Feather or the Bird?". [3] Palmer. [14] Ridley. Zinovieva. pp. 53–53.1073/pnas. "Allosteric proteins and cellular control systems". PMID 10781035. Cambridge. org/ content/ 94/ 6/ 2421. org/ cgi/ content/ full/ 97/ 9/ 4424). New York: Oxford University Press. 170. Charles (1859). pnas.. [9] Desmond 1989. doi:10.1525-142x.1016/S0022-2836(63)80091-1. p. 86–88. J. 512 [11] Desmond & Moore 1991. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=457). "Pax genes and eye organogenesis". R. Brush. R. Woo.. C. (1987). Blackwell Publishing. London: Michael Joseph.1038/415315a. doi:10. Marc. • Secord. .1038/20944. nih. P. (2005). A.M. [17] Müller GB and Newman SA (Eds. The MIT Press. Tabin. Maas.O. Brian K. D. [22] Baxter. [26] Carroll.J. Zool.X. • Minelli. and Newman. [23] Gerhart. Nature 399 (6735): 474–479. John. New Delhi. Her. Yale University Press. Blackwell Science. ed (2007). interscience... fcgi?tool=pmcentrez& artid=2581958). "The theory of facilitated variation" (http:/ / www.J. Weatherbee.. Beier. Leo W. Marc (1997). "Convergent Evolution in the Genetic Basis of Mullerian Mimicry in Heliconius Butterflies" (http:/ / www. ISBN 9780300108651. Embryos and Evolution.0701035104. Morphology. H. (2003). doi:10. "Endless forms: the evolution of gene regulation and morphological diversity". . PMID 9006082. [25] Carroll. doi:10. PMID 11797007. Nature 415 (6869): 315–318. Marion (1995). Eva. C. ISBN 978-0-262-12283-2. [18] Xu. medicine. S. Peter J. Genetics 180 (3): 1567–1577. and secret authorship of Vestiges of the natural history of creation. "Developmental basis of limblessness and axial patterning in snakes. Further reading • Buss. fcgi?tool=pmcentrez& artid=1876433). Jennifer K. & Olsen. ISBN 978-0521808514. (1999).. Development 124 (1): 219–231. Chamberlain. ISBN 978-0691084688. Phoenix Giants. Cell 101: 577–80. Victorian sensation: the extraordinary publication. P. reception. ISBN 0-226-14374-0.. Tickle. New York: Oxford University Press. doi:10. Grant. ISBN 978-81-8356-256-0. The politics of evolution: morphology. Gerhart. [21] Beldade. Epigenetic Inheritance and Evolution: The Lamarckian Dimension.Evolutionary developmental biology PMC 18255. P. Stuart A. ISBN 978-0198540632. [19] Abzhanov. Proceedings of the National Academy of Sciences 104 (suppl1): 8582–8589. R. (2000). (2002).1098095. Science 305 (5689): 1462–1465. Part B: Molecular and Developmental Evolution 304B: 485–631. (2003). (2008). doi:10. ffrench-Constant.R. "Bmp4 and Morphological Variation of Beaks in Darwin's Finches".. Norton. Brakefield. India: Discovery Publishing House. Humphray. Adrian J. The Evolution of Individuality. Darwin. Manfred D.107.H.. • Laubichler. and reform in radical London. Gerd B. Cells. ISBN 0-7181-3430-3. Chicago: University of Chicago Press. Evolution: the history of an idea. Sean B. PMID 17494755. Grenier.. Joron. (2004). W. M. • Carroll. I. [24] Gerhart.. nih. C. M. [28] Müller. [20] Cohn. N. (2005).. From DNA to Diversity: Molecular Genetics and the Evolution of Animal Design — Second Edition. Alessandro (2003). Marc (2007). Protas. A.. Oxford. Adrian J. C. John.082982. John (2005). ISBN 0-226-74411-6. Papa. Moore. Kirschner.D. Kirschner.. wiley.. • Kirschner. P. PMC 2581958. The Development of Animal Form: Ontogeny. (1989). Chicago: University of Chicago Press. gov/ articlerender.. "Mouse Eya homologues of the Drosophila eyes absent gene require Pax6 for expression in lens and nasal placode". PMID 18791259. Origination of Organismal Form: Beyond the Gene in Developmental and Evolutionary Biology. pubmedcentral. pubmedcentral.. • Desmond. How the Leopard Changed its Spots. and Maienschein. ISBN 9780691088099.. • Desmond. [27] Jablonka. Morrison. Berkeley: University of California Press. Sean B. com/ cgi-bin/ jissue/ 112149101). Grant. ed (2003). • Goodwin. Long..) (2005). The Plausibility of Life: Resolving Darwin's Dilemma. (1997).1073/pnas.1126/science. Cambridge University Press. Lamb.. Jiggins. ISBN 0-520-23693-9. PMID 15353802.1016/S0092-8674(00)80868-5. MIT Press. R.W. McMillan. "Contribution of Distal-less to quantitative variation in butterfly eyespots". PMC 1876433. doi:10. Sean B. ISBN 9780393060164. Wendy M. Keywords and Concepts in Evolutionary Developmental Biology. "Special issue: Evolutionary Innovation and Morphological Novelty" (http:/ / www3. and Evolution.R.. Scott D. M.R.J.L. • Hall..1534/genetics. . Princeton University Press. S. James A. 295 Sources • Bowler. Jane. ed (2007). gov/ articlerender. From Embryology to Evo-Devo: A History of Developmental Evolution. PMID 10365960. James William (1991). Endless Forms Most Beautiful: The New Science of Evo Devo and the Making of the Animal Kingdom. B. ISBN 978-0865425743. Journal of Exp.".. Brian (1994)..D. ISBN 1405119500.. Evolutionary developmental biology • Orr, H. Allen (2005-10-24). "Turned on: A revolution in the field of evolution?" (http://www.newyorker.com/ critics/books/articles/051024crbo_books1). The New Yorker. Discussion of Carroll, Endless Forms Most Beautiful • Raff, Rudolf A. (1996). The Shape of Life: Genes, Development, and the Evolution of Animal Form. The University of Chicago Press. ISBN 978-0226702667. • Sommer, Ralf J. (2009). "The future of evo–devo: model systems and evolutionary theory". Nature Reviews Genetics 10 (6): 416–422. doi:10.1038/nrg2567. PMID 19369972. 296 External links • Scott F. Gilbert, The morphogenesis of evolutionary developmental biology (http://www.ijdb.ehu.es/ fullaccess/fulltext.03078/ft467.pdf) • Tardigrades (water bears) as evo-devo models, a short video from NPR's Science Friday (http://www.npr.org/ templates/story/story.php?storyId=99800021) Molecular evolution Molecular evolution is the process of evolution at the scale of DNA, RNA, and proteins. Molecular evolution emerged as a scientific field in the 1960s as researchers from molecular biology, evolutionary biology and population genetics sought to understand recent discoveries on the structure and function of nucleic acids and protein. Some of the key topics that spurred development of the field have been the evolution of enzyme function, the use of nucleic acid divergence as a "molecular clock" to study species divergence, and the origin of non-functional or junk DNA. Recent advances in genomics, including whole-genome sequencing, high-throughput protein characterization, and bioinformatics have led to a dramatic increase in studies on the topic. In the 2000s, some of the active topics have been the role of gene duplication in the emergence of novel gene function, the extent of adaptive molecular evolution versus neutral drift, and the identification of molecular changes responsible for various human characteristics especially those pertaining to infection, disease, and cognition. Principles of molecular evolution Mutations Mutations are permanent, transmissible changes to the genetic material (usually DNA or RNA) of a cell. Mutations can be caused by copying errors in the genetic material during cell division and by exposure to radiation, chemicals, or viruses, or can occur deliberately under cellular control during the processes such as meiosis or hypermutation. Mutations are considered the driving force of evolution, where less favorable (or deleterious) mutations are removed from the gene pool by natural selection, while more favorable (or beneficial) ones tend to accumulate. Neutral mutations do not affect the organism's chances of survival in its natural environment and can accumulate over time, which might result in what is known as punctuated equilibrium; the modern interpretation of classic evolutionary theory. Molecular evolution 297 Causes of change in allele frequency There are three known processes that affect the survival of a characteristic; or, more specifically, the frequency of an allele (variant of a gene): • Genetic drift describes changes in gene frequency that cannot be ascribed to selective pressures, but are due instead to events that are unrelated to inherited traits. This is especially important in small mating populations, which simply cannot have enough offspring to maintain the same gene distribution as the parental generation. • Gene flow or Migration: or gene admixture is the only one of the agents that makes populations closer genetically while building larger gene pools. • Selection, in particular natural selection produced by differential mortality and fertility. Differential mortality is the survival rate of individuals before their reproductive age. If they survive, they are then selected further by differential fertility – that is, their total genetic contribution to the next generation. In this way, the alleles that these surviving individuals contribute to the gene pool will increase the frequency of those alleles. Sexual selection, the attraction between mates that results from two genes, one for a feature and the other determining a preference for that feature, is also very important. Molecular study of phylogeny Molecular systematics is a product of the traditional field of systematics and molecular genetics. It is the process of using data on the molecular constitution of biological organisms' DNA, RNA, or both, in order to resolve questions in systematics, i.e. about their correct scientific classification or taxonomy from the point of view of evolutionary biology. Molecular systematics has been made possible by the availability of techniques for DNA sequencing, which allow the determination of the exact sequence of nucleotides or bases in either DNA or RNA. At present it is still a long and expensive process to sequence the entire genome of an organism, and this has been done for only a few species. However, it is quite feasible to determine the sequence of a defined area of a particular chromosome. Typical molecular systematic analyses require the sequencing of around 1000 base pairs. The driving forces of evolution Depending on the relative importance assigned to the various forces of evolution, three perspectives provide evolutionary explanations for molecular evolution.[1] While recognizing the importance of random drift for silent mutations,[2] selectionists hypotheses argue that balancing and positive selection are the driving forces of molecular evolution. Those hypotheses are often based on the broader view called panselectionism, the idea that selection is the only force strong enough to explain evolution, relaying random drift and mutations to minor roles.[1] Neutralists hypotheses emphasize the importance of mutation, purifying selection and random genetic drift.[3] The introduction of the neutral theory by Kimura,[4] quickly followed by King and Jukes' own findings,[5] lead to a fierce debate about the relevance of neodarwinism at the molecular level. The Neutral theory of molecular evolution states that most mutations are deleterious and quickly removed by natural selection, but of the remaining ones, the vast majority are neutral with respect to fitness while the amount of advantageous mutations is vanishingly small. The fate of neutral mutations are governed by genetic drift, and contribute to both nucleotide polymorphism and fixed differences between species.[6] [7] [8] Mutationists hypotheses emphasize random drift and biases in mutation patterns.[9] Sueoka was the first to propose a modern mutationist view. He proposed that the variation in GC content was not the result of positive selection, but a consequence of the GC mutational pressure.[10] Molecular evolution 298 Related fields An important area within the study of molecular evolution is the use of molecular data to determine the correct biological classification of organisms. This is called molecular systematics or molecular phylogenetics. Tools and concepts developed in the study of molecular evolution are now commonly used for comparative genomics and molecular genetics, while the influx of new data from these fields has been spurring advancement in molecular evolution. Key researchers in molecular evolution Some researchers who have made key contributions to the development of the field: • • • • • • • Motoo Kimura — Neutral theory Masatoshi Nei — Adaptive evolution Walter M. Fitch — Phylogenetic reconstruction Walter Gilbert — RNA world Joe Felsenstein — Phylogenetic methods Susumu Ohno — Gene duplication John H. Gillespie — Mathematics of adaptation Journals and societies Journals dedicated to molecular evolution include Molecular Biology and Evolution, Journal of Molecular Evolution, and Molecular Phylogenetics and Evolution. Research in molecular evolution is also published in journals of genetics, molecular biology, genomics, systematics, or evolutionary biology. The Society for Molecular Biology and Evolution [11] publishes the journal "Molecular Biology and Evolution" and holds an annual international meeting. See also • • • • • • History of molecular evolution Chemical evolution Evolution Genetic drift Evolutionary physiology • • • • • Genomic organization Horizontal gene transfer Human evolution Molecular clock Comparative phylogenetics • • • • Neutral theory of molecular evolution Nucleotide diversity Parsimony Population genetics Selection Evolution of dietary antioxidants • E. coli long-term evolution experiment • References [1] [2] [3] [4] [5] [6] [7] [8] [9] Graur, D. and Li, W.-H. (2000). Fundamentals of molecular evolution. Sinauer. Gillespie, J. H (1991). The Causes of Molecular Evolution. Oxford University Press, New York. ISBN 0-19-506883-1. Kimura, M. (1983). The Neutral Theory of Molecular Evolution. Cambridge University Press, Cambridge. ISBN 0-521-23109-4. Kimura, Motoo (1968). "Evolutionary rate at the molecular level" (http:/ / www2. hawaii. edu/ ~khayes/ Journal_Club/ fall2006/ Kimura_1968_Nature. pdf). Nature 217 (5129): 624–626. doi:10.1038/217624a0. PMID 5637732. . King, J.L. and Jukes, T.H. (1969). "Non-Darwinian Evolution" (http:/ / www. blackwellpublishing. com/ ridley/ classictexts/ king. pdf). Science 164 (881): 788–798. doi:10.1126/science.164.3881.788. PMID 5767777. . Nachman M. (2006). "Detecting selection at the molecular level" in: Evolutionary Genetics: concepts and case studies. pp. 103–118. The nearly neutral theory expanded the neutralist perspective, suggesting that several mutations are nearly neutral, which means both random drift and natural selection is relevant to their dynamics. Ohta, T (1992). "The nearly neutral theory of molecular evolution". Annual Review of Ecology and Systematics 23: 263–286. doi:10.1146/annurev.es.23.110192.001403. Nei, M. (2005). "Selectionism and Neutralism in Molecular Evolution" (http:/ / www. pubmedcentral. nih. gov/ articlerender. fcgi?tool=pmcentrez& artid=1513187). Molecular Biology and Evolution 22 (12): 2318–2342. doi:10.1093/molbev/msi242. PMID 16120807. Molecular evolution PMC 1513187. [10] Sueoka, N. (1964). "On the evolution of informational macromolecules". In In: Bryson, V. and Vogel, H.J.. Evolving genes and proteins. Academic Press, New-York. pp. 479–496. [11] http:/ / www. smbe. org 299 Further reading • Li, W.-H. (2006). Molecular Evolution. Sinauer. ISBN 0878934804. • Lynch, M. (2007). The Origins of Genome Architecture. Sinauer. ISBN 0878934847. Human evolution Human evolution, or anthropogenesis, is the origin and evolution of Homo sapiens as a distinct species from other hominids, great apes and placental mammals. The study of human evolution encompasses many scientific disciplines, including physical anthropology, primatology, archaeology, linguistics and genetics.[1] The term "human" in the context of human evolution refers to the genus Homo, but studies of human evolution usually include other hominids, such as the Australopithecines, from which the genus Homo had diverged by about 2.3 to 2.4 million years ago in Africa.[2] [3] Scientists have estimated that humans branched off from their common ancestor with chimpanzees about 5–7 million years ago. Several species and subspecies of Homo evolved and are now extinct. These include Homo erectus, which inhabited Asia, and Homo sapiens neanderthalensis, which inhabited Europe. Archaic Homo sapiens evolved between 400,000 and 250,000 years ago. The dominant view among scientists concerning the origin of anatomically modern humans is the "Out of Africa" or recent African origin hypothesis,[4] [5] [6] [7] which argues that Homo sapiens arose in Africa and migrated out of the continent around 50,000 to 100,000 years ago, replacing populations of Homo erectus in Asia and Homo neanderthalensis in Europe. Scientists supporting the alternative multiregional hypothesis argue that Homo sapiens evolved as geographically separate but interbreeding populations stemming from a worldwide migration of Homo erectus out of Africa nearly 2.5 million years ago. History of ideas The word homo, the name of the biological genus to which humans belong, is Latin for "human". It was chosen originally by Carolus Linnaeus in his classification system. The word "human" is from the Latin humanus, the adjectival form of homo. The Latin "homo" derives from the Indo-European root, dhghem, or "earth".[8] Carolus Linnaeus and other scientists of his time also considered the great apes to be the closest relatives of humans due to morphological and anatomical similarities. The possibility of linking humans with earlier apes by descent only became clear after 1859 with the publication of Charles Darwin's On the Origin of Species. This argued for the idea of the evolution of new species from earlier ones. Darwin's book did not address the question of human evolution, saying only that "Light will be thrown on the origin of man and his history". Human evolution 300 The first debates about the nature of human evolution arose between Thomas Huxley and Richard Owen. Huxley argued for human evolution from apes by illustrating many of the similarities and differences between humans and apes and did so particularly in his 1863 book Evidence as to Man's Place in Nature. However, many of Darwin's early supporters (such as Alfred Russel Wallace and Charles Lyell) did not agree that the origin of the mental capacities and the moral sensibilities of humans could be explained by natural selection. Darwin applied the theory of evolution and sexual selection to humans when he published The Descent of Man in 1871.[9] A major problem was the lack of fossil intermediaries. It was only in the 1920s that such fossils were discovered in Africa. In 1925, Raymond Dart described Australopithecus africanus. Fossil Hominid Evolution Display at The Museum of The type specimen was the Taung Child, an Australopithecine Osteology, Oklahoma City, USA. infant discovered in a cave. The child's remains were a remarkably well-preserved tiny skull and an endocranial cast of the individual's brain. Although the brain was small (410 cm³), its shape was rounded, unlike that of chimpanzees and gorillas, and more like a modern human brain. Also, the specimen showed short canine teeth, and the position of the foramen magnum was evidence of bipedal locomotion. All of these traits convinced Dart that the Taung baby was a bipedal human ancestor, a transitional form between apes and humans. The classification of humans and their relatives has changed considerably over time. The gracile Australopithecines are now thought to be ancestors of the genus Homo, the group to which modern humans belong. Both Australopithecines and Homo sapiens are part of the tribe Hominini. Recent data suggests Australopithecines were a diverse group and that A. africanus may not be a direct ancestor of modern humans. Reclassification of Australopithecines that originally were split into either gracile or robust varieties has put the latter into a family of its own, Paranthropus. Taxonomists place humans, Australopithecines and related species in the same family as other great apes, in the Hominidae. Hominin species distributed through time Note: 1e+06 years = 1 × 106 years = 1 million years ago = 1 Ma Human evolution 301 Before Homo Evolution of the great apes The evolutionary history of the primates can be traced back 65 million years, as one of the oldest of all surviving placental mammal groups. The oldest known primate-like mammal species, the Plesiadapis, come from North America, but they were widespread in Eurasia and Africa during the tropical conditions of the Paleocene and Eocene. Plesiadapis. With the beginning of modern climates, marked by the formation of the first Antarctic ice in the early Oligocene around 30 million years ago. A primate from this time was Notharctus. Fossil evidence found in Germany in the 1980s was determined to be about 16.5 million years old, some 1.5 million years older than similar species from East Africa and challenging the original theory regarding human ancestry originating on the African continent. David Begun[10] says that these primates flourished in Eurasia and that Notharctus. the lineage leading to the African apes and humans— including Dryopithecus—migrated south from Europe or Western Asia into Africa. The surviving tropical population, which is seen most completely in the upper Eocene and lowermost Oligocene fossil beds of the Fayum depression southwest of Cairo, gave rise to all living primates—lemurs of Madagascar, lorises of Southeast Asia, galagos or "bush babies" of Africa, and the anthropoids; platyrrhines or New World monkeys, and catarrhines or Old World monkeys and the great apes and humans. The earliest known catarrhine is Kamoyapithecus from uppermost Oligocene at Eragaleit in the northern Kenya Rift Valley, dated to 24 million years ago. Its ancestry is generally thought to be species related to Aegyptopithecus, Propliopithecus, and Parapithecus from the Fayum, at around 35 million years ago. In 2010, Saadanius was described as a close relative of the last common ancestor of the crown catarrhines, and tentatively dated to 29–28 million years ago, helping to fill an 11-million-year gap in the fossil record.[11] In the early Miocene, about 22 million years ago, the many kinds of arboreally adapted primitive catarrhines from East Africa suggest a long history of prior diversification. Fossils at 20 million years ago include fragments attributed to Victoriapithecus, the earliest Old World Monkey. Among the genera thought to be in the ape lineage leading up to 13 million years ago are Proconsul, Rangwapithecus, Dendropithecus, Limnopithecus, Nacholapithecus, Equatorius, Nyanzapithecus, Afropithecus, Heliopithecus, and Kenyapithecus, all from East Africa. The presence of other generalized non-cercopithecids Reconstructed tailless Proconsul skeleton. of middle Miocene age from sites far distant—Otavipithecus from cave deposits in Namibia, and Pierolapithecus and Dryopithecus from France, Spain and Austria—is evidence of a wide diversity of forms across Africa and the Mediterranean basin during the relatively warm and equable climatic regimes of the early and middle Miocene. The youngest of the Miocene hominoids, Oreopithecus, is from 9 million year old coal beds in Italy. Human evolution Molecular evidence indicates that the lineage of gibbons (family Hylobatidae) became distinct from Great Apes between 18 and 12 million years ago, and that of orangutans (subfamily Ponginae) became distinct from the other Great Apes at about 12 million years; there are no fossils that clearly document the ancestry of gibbons, which may have originated in a so-far-unknown South East Asian hominoid population, but fossil proto-orangutans may be represented by Ramapithecus from India and Griphopithecus from Turkey, dated to around 10 million years ago. 302 Divergence of the human lineage from other Great Apes Species close to the last common ancestor of gorillas, chimpanzees and humans may be represented by Nakalipithecus fossils found in Kenya and Ouranopithecus found in Greece. Molecular evidence suggests that between 8 and 4 million years ago, first the gorillas, and then the chimpanzees (genus Pan) split off from the line leading to the humans; human DNA is approximately 98.4% identical to that of chimpanzees when comparing single nucleotide polymorphisms (see human evolutionary genetics). The fossil record of gorillas and chimpanzees is quite limited. Both poor preservation (rain forest soils tend to be acidic and dissolve bone) and sampling bias probably contribute to this problem. Other hominines likely adapted to the drier environments outside the equatorial belt, along with antelopes, hyenas, dogs, pigs, elephants, and horses. The equatorial belt contracted after about 8 million years ago. Fossils of these hominans - the species in the human lineage following divergence from the chimpanzees - are relatively well known. The earliest are Sahelanthropus tchadensis (7 Ma) and Orrorin tugenensis (6 Ma), followed by: • Ardipithecus (5.5–4.4 Ma), with species Ar. kadabba and Ar. ramidus; • Australopithecus (4–1.8 Ma), with species Au. anamensis, Au. afarensis, Au. africanus, Au. bahrelghazali, Au. garhi, and Au. sediba; • Kenyanthropus (3–2.7 Ma), with species Kenyanthropus platyops; • Paranthropus (3–1.2 Ma), with species P. aethiopicus, P. boisei, and P. robustus; • Homo (2 Ma–present), with species Homo habilis, Homo rudolfensis, Homo ergaster, Homo georgicus, Homo antecessor, Homo cepranensis, Homo erectus, Homo heidelbergensis, Homo rhodesiensis, Homo neanderthalensis, Homo sapiens idaltu, Archaic Homo sapiens, Homo floresiensis. Genus Homo Homo sapiens is the only extant species of its genus, Homo. While some other, extinct, Homo species might have been ancestors of Homo sapiens, many were likely our "cousins", having speciated away from our ancestral line.[12] [13] There is not yet a consensus as to which of these groups should count as separate species and which as subspecies. In some cases this is due to the dearth of fossils, in other cases it is due to the slight differences used to classify species in the Homo genus.[13] The Sahara pump theory (describing an occasionally passable "wet" Sahara Desert) provides an explanation of the early variation in the genus Homo. Based on archaeological and paleontological evidence, it has been possible to infer, to some extent, the ancient dietary practices of various Homo species and to study the role of diet in physical and behavioral evolution within Homo.[14] [15] [16] [17] [18] H. habilis Homo habilis lived from about 2.4 to 1.4 Ma. Homo habilis, the first species of the genus Homo, evolved in South and East Africa in the late Pliocene or early Pleistocene, 2.5–2 Ma, when it diverged from the Australopithecines. Homo habilis had smaller molars and larger brains than the Australopithecines, and made tools from stone and perhaps animal bones. One of the first known hominids, it was nicknamed 'handy man' by its discoverer, Louis Leakey due to its association with stone tools. Some scientists have proposed moving this species out of Homo and into Australopithecus due to the morphology of its skeleton being more adapted to living on trees rather than to moving on two legs like Homo sapiens.[19] Human evolution 303 H. rudolfensis and H. georgicus These are proposed species names for fossils from about 1.9–1.6 Ma, the relation of which with Homo habilis is not yet clear. • Homo rudolfensis refers to a single, incomplete skull from Kenya. Scientists have suggested that this was another Homo habilis, but this has not been confirmed.[20] • Homo georgicus, from Georgia, may be an intermediate form between Homo habilis and Homo erectus,[21] or a sub-species of Homo erectus.[22] H. ergaster and H. erectus The first fossils of Homo erectus were discovered by Dutch physician Eugene Dubois in 1891 on the Indonesian island of Java. He originally gave the material the name Pithecanthropus erectus based on its morphology that he considered to be intermediate between that of humans and apes.[24] Homo erectus (H erectus) lived from about 1.8 Ma to about 70,000 years ago (which would indicate that they were probably wiped out by the Toba catastrophe; however, Homo erectus soloensis and Homo floresiensis survived it). Often the early phase, from 1.8 to 1.25 Ma, is considered to be a separate species, Homo ergaster, or it is seen as a subspecies of Homo erectus, Homo erectus ergaster. In the early Pleistocene, 1.5–1 Ma, in Africa, Asia, and Europe, some populations of Homo habilis are thought to have evolved larger brains and made more One current view of the temporal and geographical distribution of hominid [23] populations. Other interpretations differ mainly in the taxonomy and geographical elaborate stone tools; these differences distribution of hominid species. and others are sufficient for anthropologists to classify them as a new species, Homo erectus. In addition Homo erectus was the first human ancestor to walk truly upright.[25] This was made possible by the evolution of locking knees and a different location of the foramen magnum (the hole in the skull where the spine enters). They may have used fire to cook their meat. A famous example of Homo erectus is Peking Man; others were found in Asia (notably in Indonesia), Africa, and Europe. Many paleoanthropologists now use the term Homo ergaster for the non-Asian forms of this group, and reserve Homo erectus only for those fossils that are found in Asia and meet certain skeletal and dental requirements which differ slightly from H. ergaster. Human evolution 304 H. cepranensis and H. antecessor These are proposed as species that may be intermediate between H. erectus and H. heidelbergensis. • H. antecessor is known from fossils from Spain and England that are dated 1.2 Ma–500 ka.[26] [27] • H. cepranensis refers to a single skull cap from Italy, estimated to be about 800,000 years old.[28] H. heidelbergensis H. heidelbergensis (Heidelberg Man) lived from about 800,000 to about 300,000 years ago. Also proposed as Homo sapiens heidelbergensis or Homo sapiens paleohungaricus.[29] H. rhodesiensis, and the Gawis cranium • H. rhodesiensis, estimated to be 300,000–125,000 years old. Most current experts believe Rhodesian Man to be within the group of Homo heidelbergensis, though other designations such as Archaic Homo sapiens and Homo sapiens rhodesiensis have also been proposed. • In February 2006 a fossil, the Gawis cranium, was found which might possibly be a species intermediate between H. erectus and H. sapiens or one of many evolutionary dead ends. The skull from Gawis, Ethiopia, is believed to be 500,000–250,000 years old. Only summary details are known, and no peer reviewed studies have been released by the finding team. Gawis man's facial features suggest its being either an intermediate species or an example of a "Bodo man" female.[30] H. neanderthalensis H. neanderthalensis lived from 400,000[31] years ago. Also proposed as Homo sapiens neanderthalensis: there is ongoing debate over whether the "Neanderthal Man" was a separate species, Homo neanderthalensis, or a subspecies of H. sapiens.[32] While the debate remains unsettled, evidence from sequencing mitochondrial DNA indicates that no significant gene flow occurred between H. neanderthalensis and H. sapiens, and, therefore, the two were separate species that shared a common ancestor about 660,000 years ago.[33] [34] In 1997, Mark Stoneking stated: "These results [based on mitochondrial DNA extracted from Neanderthal bone] indicate that Neanderthals did not Le Ferrassie Neanderthal skull (cast) contribute mitochondrial DNA to modern humans… Neanderthals are not our ancestors." Subsequent investigation of a second source of Neanderthal DNA supported these findings.[35] However, the 2010 sequencing of the Neanderthal genome indicates that Neanderthals did indeed interbreed with H. sapiens circa 75,000 BC (after H. sapiens moved out from Africa, but before they separated into Europe, the Middle East, and Asia).[36] Nearly all modern humans have 1% to 4% of their DNA derived from Neanderthal DNA.[36] (To appreciate how big of a percentage this is, consider that humans and chimps only differ in 1.5% of their DNA.) This 1-4% bit of DNA is only present in non-African humans.[36] However, supporters of the multiregional hypothesis point to recent studies indicating non-African nuclear DNA heritage dating to one Ma,[37] although the reliability of these studies has been questioned.[38] Competition from Homo sapiens probably contributed to Neanderthal extinction.[39] [40] They could have coexisted in Europe for as long as 10,000 years.[41] Artifacts. However.000 years ago and the second interglacial period in the Middle Pleistocene.[46] Some scientists presently believe that H. shoulder. the DNA of individuals is more alike than usual for most species. A subsequent migration within and out of Africa eventually replaced the earlier dispersed H. or the Denisova hominin.000 BP. Current evidence does not preclude some multiregional evolution or some admixture of the migrant H. The living woman was estimated to be one meter in height. possibly a result of insular dwarfism. with a brain volume of just 380 cm3 (considered small for a chimpanzee and less than a third of the H. This coupled with pathological dwarfism. floresiensis. This is a hotly debated area of paleoanthropology. H.000 years ago to the present.[47] The hypothesis of pathological dwarfism. erectus. sapiens suffering from pathological dwarfism. Denisova hominin In 2008. has been nicknamed hobbit for its small size. which lived from approximately 100. Found in 2003 it has been dated to approximately 18. a team of scientists from the Max Planck Institute for Evolutionary Anthropology in Leipzig. that is. primarily as the result of small groups of people moving into new environmental circumstances. sapiens.[49] . Current research has established that humans are genetically highly homogenous. erectus out of Africa. floresiensis share a common ancestor with modern humans. which may have resulted from their relatively recent evolution or the possibility of a population bottleneck resulting from cataclysmic natural events such as the Toba catastrophe. Between 400.000 years old. archeologists working at the site of Denisova Cave in the Altai Mountains of Siberia uncovered a small bone fragment from the fifth finger of a juvenile hominin. As DNA had survived in the fossil fragment due to the cool climate of the Denisova Cave. around 250. sapiens from H. in addition to internal characteristics such as the ability to breathe more efficiently in high altitudes.[45] H. however.000 years ago. being a concrete example of a recent species of the genus Homo that exhibits derived traits not shared with modern humans.Human evolution 305 H. excavated in the cave at the same level were carbon dated to around 40.000 before present. The direct evidence suggests there was a migration of H. floresiensis is indeed a separate species. including a bracelet. floresiensis H. these features include the form of bones in the wrist. H. This migration and origin theory is usually referred to as the recent single origin or Out of Africa theory. Aside from cranial features.[42] [43] [44] Distinctive genetic characteristics have arisen. dubbed the "X-woman" (referring to the maternal descent of mitochondrial DNA[48] . H. sapiens H. there is an ongoing debate over whether H.000 years ago. because some modern humans who live on Flores. The other major attack on H. forearm. fails to explain additional anatomical features that are unlike those of modern humans (diseased or not) but much like those of ancient members of our genus. sapiens average of 1400 cm3). the trend in skull expansion and the elaboration of stone tool technologies developed. it is argued. floresiensis is intriguing both for its size and its age. The main find was a skeleton believed to be a woman of about 30 years of age. floresiensis is that it was found with tools only associated with H. sapiens. knees. floresiensis was a modern H. is a possible extinct sub-species who lived from about 160. These adapted traits are a very small component of the Homo sapiens genome. the island where the skeleton was found. sapiens idaltu. In other words. then a further speciation of H. but include various characteristics such as skin color and nose form.000 to 12. could indeed create a hobbit-like human. erectus in Africa. providing evidence for a transition from H. sapiens with existing Homo populations. Germany sequenced mtDNA extracted from the fragment. sapiens (the adjective sapiens is Latin for "wise" or "intelligent") have lived from about 250. however. are pygmies. from Ethiopia.[47] This hypothesis is supported in part. and feet. but split from the modern human lineage and followed a distinct evolutionary path. erectus to H. 3–4.6–6.2 – present Zambia Kenya Ethiopia Worldwide 1. rudolfensis H. sapiens sapiens (modern humans) 0.10? – 0.8? 1. cepranensis H.000 2 sites H. The DNA analysis further indicated that this new hominin species was the result of an early migration out of Africa." Pääbo noted that the existence of this distant branch creates a much more complex picture of humankind during the Late Pleistocene.3 ft) 1.300 Very few 1 skull 1921 1972/1986 1997/2003 —/1758 1. sapiens idaltu H.100 (late) 1 skull cap Many 1994/2003 1891/1892 H. China Europe.15 0.[50] Professor Chris Stringer. ergaster 1.7 90 kg (200 lb) ft) 1.[50] Modern humans are known to have overlapped with Neanderthals in Europe for more than 10.9 – 0. Neanderthals. Western Asia 1.6 m (5.35 – 0.2 – 0. floresiensis 0.6 South Africa 2010 H.03 55–70 kg (120–150 lb) (heavily built) 1.000 850 (early) – 1.6 – 0.16 – 0.0 m (3.3 ft) 25 kg (55 lb) 700–850 Many 1975 H.8 600 4 individuals Many 1999/2002 H.900 Many (1829)/1864 H.2 Italy Africa.000 years. remarked: "This new DNA work provides an entirely new way of looking at the still poorly understood evolution of humans in central and eastern Asia. Africa.5 – 0. georgicus 1. but also distinct from the earlier African exodus of Homo erectus. modern humans and the Denisovan hominin may have co-existed together.8 m (5.9 ft) 1.100–1.12 1.9 ft) 60 kg (130 lb) 1.000–1. heidelbergensis 0.9 – 1.4–1. human origins researcher at London's Natural History Museum and one of the leading proponents of the recent single-origin hypothesis.012 400 7 individuals 2003/2004 H.450 1.[48] 306 Comparative table of Homo species Species Lived when (Ma) Lived where Adult height Adult mass Cranial capacity (cm³) Fossil record Discovery / publication of name 1997 H.0–1.2 ft) 1.200–1.0 m (3.4 1.35 Europe.4 Africa 1. India.Human evolution The analysis indicated that modern humans. distinct from the later out-of-Africa migrations associated with Neanderthals and modern humans. antecessor 1.2 ft) (110–220 lb) 1. Eurasia (Java.3 – 1.9 ft) (73–120 lb) 1. neanderthalensis 0.9 m 50–100 kg (4.850 3 craniums Still living .2 ft) 60 kg (130 lb) 510–660 1960/1964 H. habilis 2.9 m (6. and the Denisova hominin last shared a common ancestor around 1 million years ago.8 m (5.9 0.5 m 33–55 kg (3. gautengensis >2 – 0.400 Many 1908 H. Caucasus) Eastern and Southern Africa Indonesia 1. erectus 0.8 Spain 1.3 – 0. China. rhodesiensis H. and the discovery raises the possibility that Neanderthals.75 m (5. Randall Susman used the anatomy of opposable thumbs as the basis for his argument that both the Homo and Paranthropus species were toolmakers. Paleontology has yet to explain the expansion of this organ over millions of years despite being extremely demanding in terms of energy consumption.5 Ma.[52] In 1994. which is one fifth of the energy consumption of a human body. He compared bones and muscles of "A sharp rock". They argue that when most of the Oldowan tools were found in association with human fossils. Homo was always present. A Homo fossil was found near some Oldowan tools.5 to 2. the most basic of human stone tools . which predates the earliest known "Homo" species. suggesting that maybe the Homo species did indeed create and use these tools. but this is debated. The brain of a modern human consumes about 20 watts (400 kilocalories per day).Human evolution 307 Use of tools Using tools has been interpreted as a sign of intelligence. Although there is no direct evidence that points to Paranthropus as the tool makers. Bernard Wood noted that "Paranthropus" coexisted with the early Homo species in the area of the "Oldowan Industrial Complex" over roughly the same span of time. but it is the human species that dominates the areas of making and using more complex tools. and it has been theorized that tool use may have stimulated certain aspects of human evolution—most notably the continued expansion of the human brain.[51] Precisely when early humans started to use tools is difficult to determine. sharp-edged stones) the more difficult it is to decide whether they are natural objects or human artifacts. There is no known evidence that any "Homo" specimens appeared by 2. It should be noted that many species make and use tools. and its age was noted at 2. Researchers have suggested that early hominids were thus under evolutionary pressure to increase their capacity to create and use tools. but not solid evidence. Increased tool use would allow hunting for energy-rich meat products. but Paranthropus was not. one of the greatest human discoveries stone tools" from Ethiopia. because the more primitive these tools are (for example. There is some evidence that the australopithecines (4 Ma) may have used broken bones as tools. the pinnacle of Homo erectus area.3 million years old. The oldest known tools are the "Oldowan Fire. an Oldowan pebble tool. It is surely possible.6 million years old. their anatomy lends to indirect evidence of their capabilities in this An Acheulean hand axe. Most paleoanthropologists agree that the early "Homo" stone working species were indeed responsible for most of the Oldowan tools found. It was discovered that these tools are from 2. and would enable processing more energy-rich plant products. exploration of less hospitable geographical areas. but once that phase started further development was slow. Debate continues as to whether a "revolution" led to modern humans ("the big bang of human consciousness").000–40. Susman defended that modern anatomy of the human thumb is an evolutionary response to the requirements associated with making and handling tools and that both species were indeed toolmakers. anthropologists include specialization of tools. blades. H. finding that humans have 3 muscles that chimps lack. Each phase (H. These Homo species were culturally conservative. or whether the evolution was more gradual.000–300. Humans also have thicker metacarpals with broader heads. Typically.6 Ma. burials with grave gifts). and flattened needles were made.000 years ago).[55] . neanderthalensis) started at a higher level than the previous one. buttons and bone needles show signs of variation among different populations of humans. In this period they also started to make tools out of bone. after about 50. until 50. but after 50. Modern humans started burying their dead. developing sophisticated hunting techniques (such as using trapping pits or driving animals off cliffs). and barter trade networks. H.000–300. Among concrete examples of Modern human behavior.000 years ago is also known as the Acheulean.000 years ago. Modern humans and the "Great Leap Forward" debate Until about 50. making the human hand more successful at precision grasping than the chimpanzee hand.[53] This marks the beginning of the Paleolithic.000 BP. author of The Third Chimpanzee.[53] Finally. organization of living space.000 BP (Before Present) the more refined so-called Levallois technique was developed. or Old Stone Age. by which scrapers. It consisted of a series of consecutive strikes.[52] Stone tools Stone tools are first attested around 2. the Middle Paleolithic (Middle Stone Age. different populations of humans introduced novelty to existing technologies: artifacts such as fish hooks. when H. ergaster (or erectus) made large stone Venus of Willendorf. around 10. ever more refined and specialized flint tools were made by the Neanderthals and the immigrant Cro-Magnons (knives.000 BP. needles. neanderthalensis populations do not vary in their technologies. later "retouched" by additional. The Paleolithic is subdivided into the Lower Paleolithic (Early Stone Age. and engaging in cave painting. more subtle strikes at the sides of the flakes. use of jewellery and images (such as cave drawings). choppers made out of round pebbles that had been split by simple strikes.000 BC modern human culture started to change at a much greater speed. at first quite rough (Early Acheulian). ending around 350. and some anthropologists characterize this as a "Great Leap Forward".[54] As human culture advanced. rituals (for example. its end is taken to be the end of the last Ice Age. an example of Paleolithic art hand-axes out of flint and quartzite. skimmers).000 years ago). and the Upper Paleolithic. H.000 years ago the use of stone tools seems to have progressed stepwise. habilis. habilis in Eastern Africa used so-called pebble tools. when H.Human evolution 308 human and chimpanzee thumbs. something that had not been seen in human cultures prior to 50. ergaster. Jared Diamond. specialized hunting techniques. After 350. The period from 700. making clothing out of hides.000–30. slicers ("racloirs"). Homo erectus evolved more than 1. which support a single migration out of Africa that gave rise to all non-African populations. and by 1.[63] There are differing theories on whether there was a single exodus or several. and then migrating out of Africa and replaced human populations in Eurasia (called the "Out of Africa" Model or the "Complete Replacement" Model). and members of this species populated different parts of Africa in a relatively short time.000 years ago. A multiple dispersal model involves the Southern Dispersal theory. Much of the evidence for the first group's expansion would have been destroyed by the rising sea levels at the end of the Holocene era. Homo habilis. According to the multiregional hypothesis. according to the "Out of Africa" model this is not the first species of hominids: the first species of genus Homo.[64] The multiple dispersal model is contradicted by studies indicating that the populations of Eurasia and the populations of Southeast Asia and Oceania are all descended from the same mitochondrial DNA lineages.000 – 50. near the coastal border of Namibia and Angola. The research also located the origin of modern human migration in south-western Africa. researchers concluded that all were descended from a woman from Africa. Multiregional model Multiregional evolution. evolving worldwide to modern Homo sapiens. Sarah Tishkoff found the San people to express the greatest genetic diversity among the 113 distinct populations sampled. After analysing genealogy trees constructed using 133 types of mtDNA. making them one of 14 "ancestral population clusters".000 years ago. Anthropologists have been divided as to whether current human population evolved as one interconnected population (as postulated by the Multiregional Evolution hypothesis). This group helped to populate Southeast Asia and Oceania.8 Ma. speciated. developed by Chris Stringer and Peter Andrews. explaining the discovery of early human sites in these areas much earlier than those in the Levant. However the presence of archaic admixture in modern humans remains a possibility and has been suggested by some studies. modern H.[59] Evidence from autosomal DNA also supports the Recent African origin. was proposed by Milford H. there was a coastal dispersal of modern humans from the Horn of Africa around 70.[58] Studies of haplogroups in Y-chromosomal DNA and mitochondrial DNA have largely supported a recent African origin. continuous human species. However. a model to account for the pattern of human evolution.[66] . Out of Africa is also supported by the fact that mitochondrial genetic diversity is highest among African populations.5 million years BP to the present day has been within a single.000 years ago and eventually replaced existing hominid species in Europe and Asia. A second wave of humans dispersed across the Sinai peninsula into Asia. population of Homo sapiens sapiens. or evolved only in East Africa.[61] [62] Out of Africa has gained support from research using mitochondrial DNA (mtDNA).5 Ma had spread throughout the Old World. dubbed Mitochondrial Eve.Human evolution 309 Models of human evolution Today. linguistic and archaeological evidence. fossil and genomic data are evidence for worldwide human evolution and contradict the recent speciation postulated by the Recent African origin hypothesis. sapiens evolved in Africa 200. evolved in East Africa at least 2 Ma.[57] Multiregional evolution holds that human evolution from the beginning of the Pleistocene 2.[64] which has gained support in recent years from genetic. undivided by species barrier.[60] Out of Africa According to the Out of Africa model. The fossil evidence was insufficient for Richard Leakey to resolve this debate. In this theory. Wolpoff[56] in 1988. This second group possessed a more sophisticated tool technology and was less dependent on coastal food sources than the original group.[65] The broad study of African genetic diversity headed by Dr. resulting in the bulk of human population for Eurasia. Homo sapiens began migrating from Africa between 70. all humans belong to one. an Australian anatomist and palaeoanthropologist. who first hypothesised the aquatic ape theory of human evolution • Henry McHenry. and its current effects. a British naturalist who documented considerable evidence that species originate through evolutionary change • Richard Dawkins. Haldane. an American anatomist and physical anthropologist whose work has explored the evolution of the vocal tract and speech • Louis Leakey. a Scottish physician and palaeontologist whose work on South Africa led to the discovery and description of the Paranthropus genus of hominins. the evolutionary past that gave rise to it. led to the discovery of Australopithecus africanus • Charles Darwin. Hamilton. an African paleontologist and archaeologist. in South Africa.000 years ago.Human evolution According to the Toba catastrophe theory to which some anthropologists and archeologists subscribe. Scientists postulate one of the reasons for the rapid selection of this genetic variant is the lethality of the disease in non-immune persons. Notable human evolution researchers • Robert Broom. Gregory Cochran and Henry Harpending argue that human evolution has accelerated since and as a result of the development of agriculture and civilisation since some 50. an African archaeologist and naturalist whose work was important in establishing human evolutionary development in Africa • Mary Leakey. a British ethologist. • Sir Alister Hardy. and of "Mrs. Lord Monboddo. Genetics Human evolutionary genetics studies how one human genome differs from the other. a British archaeologist and anthropologist whose discoveries in Africa include the Laetoli footprints • Richard Leakey. medical and forensic implications and applications. Differences between genomes have anthropological.[69] [70] Other reported evolutionary trends in other populations include a lengthening of the reproductive period. evolutionary biologist who has promoted a gene-centered view of evolution • J. with recent findings demonstrating the population which is at risk of the severe debilitating disease kuru has significant over-representation of an immune variant of the prion protein gene G127V versus non-immune alleles.000 Year Explosion. Ples" • James Burnett. B. the supereruption of Lake Toba on Sumatra island in Indonesia roughly 70. and on the evolution of HIV and other human diseases. a British Evolutionary Biologist who expounded a rigorous genetic basis for kin selection. a British zoologist. the origins of bipedality.[71] In their 2009 book The 10.[67] killing most humans then alive and creating a population bottleneck that affected the genetic inheritance of all humans today. an American anthropologist who specializes in studies of human evolution. reduction in cholesterol levels. whose work at Taung. son of Louis and Mary Leakey . and paleoanthropology • Jeffrey Laitman. a British judge most famous today as a founder of modern comparative historical linguistics • Raymond Dart. blood glucose and blood pressure. a British geneticist and evolutionary biologist • William D. Genetic data can provide important insight into human evolution. and that there are consequently substantial genetic differences between different current human populations. S.[68] 310 Recent and current human evolution Natural selection is being observed in contemporary human populations.000 years ago had global consequences. an American paleoanthropologist who is the leading proponent of the multiregional evolution hypothesis. geneticist and statistician. a Swedish biologist specializing in evolutionary genetics • David Pilbeam. leading proponent of the recent single origin hypothesis • Alan Templeton. a paleoanthropologist. an American physical anthropologist and professor of biological anthropology • Chris Stringer. Wolpoff. proponent of the multiregional hypothesis • Philip V. a British naturalist. anthropologist. who independently from Charles Darwin proposed the principles of evolution of animal species • Milford H. • • • Sahelanthropus • Sahelanthropus tchadensis Orrorin • Orrorin tugenensis Ardipithecus • • Ardipithecus kadabba Ardipithecus ramidus • Australopithecus • Australopithecus anamensis • Australopithecus afarensis • Australopithecus bahrelghazali • Australopithecus africanus • Australopithecus garhi • Australopithecus sediba Paranthropus • Paranthropus aethiopicus • Paranthropus boisei • Paranthropus robustus Kenyanthropus • Kenyanthropus platyops • Homo • • • • • • • • • • • • • • Homo habilis Homo rudolfensis Homo ergaster Homo georgicus Homo erectus Homo cepranensis Homo antecessor Homo heidelbergensis Homo rhodesiensis Homo neanderthalensis Homo sapiens idaltu Homo sapiens (Cro-Magnon) Homo sapiens sapiens Homo floresiensis • • See also • • • • • • • • • • • List of human evolution fossils Timeline of human evolution Archaeogenetics Dual inheritance theory Dysgenics Evolutionary anthropology Evolutionary medicine Evolutionary neuroscience Evolution of morality • • • • • • • • • • Evolutionary psychology History of Earth Hominid intelligence Human behavioral ecology Human skeletal changes due to bipedalism Human vestigiality Ida (fossil) Pan prior Physical anthropology Sahara pump theory Sexual selection in human evolution Sociocultural evolution Evolution of human intelligence • Evolutionary origin of religions • . a prominent American paleoanthropologist and expert on Neanderthal biology and human evolution • Alfred Russel Wallace. an Iranian American evolutionary geneticist who developed a statistical method which identifies sections of the genome that have been subject to natural selection 311 Species list This list is in chronological order across the page by genus. Schwartz. researcher and writer on a range of topics involving human and primate evolution. Tobias. • Pardis Sabeti. a South African palaeoanthropologist is one of the world's leading authorities on the evolution of humankind • Erik Trinkaus.Human evolution • Svante Pääbo. • Jeffrey H. sometimes called the "father of biogeography". . [22] Lordkipanidze D. [16] Bogin. archive. scar. Houghton Mifflin Company. doi:10. Al-Masari. Ferring R.H..). which is a serendipitous result from another search. "Human Evolution". Mark F. [11] Zalmout. bartelby. Retrieved 2008-12-29. doi:10. com/ nature/ journal/ v423/ n6941/ full/ 423692a. pbio.1006/jhev. Robertson ML (2007). sciencemag.. Westport. Grine FE. C. 522–543. 265.1007/BF02734246. (1994). p. May). 206. I. Nature 466 (7304): 360–364. 1 (4): 243–53. 32 (1): 17–82. Mass. [13] Bill Bryson (2004). Lordkipanidze D. Also ISBN 0-521-46786-1 (paperback) [3] McHenry. org/ web/ 20031203003838/ http:/ / citd. doi:10.1002/ar. ISBN 0-521-32370-3. . Moniz MA (1997).): e340.2000.1998. Nasser.M. PMID 19334004. [4] "Out of Africa Revisited . CT: Bergin & Garvey. Sciencemag. Nature. M. Robert Martin & David Pilbeam (eds. 2005-05-13.-A. "New Oligocene primate from Saudi Arabia and the divergence of apes and Old World Monkeys".. "A reappraisal of early hominid phylogeny". sex and age categories". (2010). Hammond SL. Hum.308 (5724): 921g . Inc. doi:10. PMID 17439362. de Lumley H. [8] "dhghem" (http:/ / www. plosjournals. "Evolution of early humans". 3–10. pinniped. [7] "Modern Humans . named by him Pithecanthropus erectus: With Remarks on so-called Transitional Forms between Apes and Man" (http:/ / www. [21] Gabounia L. Asa3. "Access : Human evolution: Out of Ethiopia" (http:/ / www.. Subsequent attempts get a targeted search result gave returns without chapter 1). J.0020340. in Reed DL. H. doi:10. The Cambridge Encyclopedia of Human Evolution. Begun DR (2001).A. Discoveries in molecular. (scroll up to view chapter 1 & part of chapter 2.0246.1002/bies. Y.1016/S1631-0683(02)00032-5. "Human nutrition: evolutionary perspectives". html). . Lola. pp. [5] Nature (2003-06-12). J. Hum.M (2009). org/ perlserv/ ?request=slideshow& type=figure& doi=10.1038/nature09094. The Mysterious Biped" (http:/ / books. J.F. This edition published 1981. (2006)...a.pbio. pubmedcentral.1126/science. Georgia". and evolutionary biology 288 (11): 1146–57. [9] Darwin. Phylogeny of Primate Lice from Morphological and Molecular Data (http:/ / biology. PMID 9034954. doi:10.A. . org/ 61/ roots/ IE104. ISBN 0897895169. [17] Barnicot NA (2005.org. Laurence R.: Bergen and Garvey. 36 (1): 115–8.0097. "The evolution of human nutrition" (http:/ / web. Evolution: The First Four Billion Years.Science" (http:/ / www. (1999) The changing face of Genus Homo.1006/jhev. htm). A. Charles (1871. p.. ISBN 9780385660044. Integr Physiol Behav Sci 40 (2): 114–17. Massachusetts: The Belknap Press of Harvard University Press. "'Homo rudolfensis' Alexeev. Comptes Rendus Palevol. Clayton DH PLoS. pp. "Genetic Analysis of Lice Supports Direct Contact between Modern and Archaic Humans". Retrieved 2009-11-23. 40 (1): 17–39. actionbioscience. [18] Leonard WR.1146/annurev. and Selection in Relation to Sex. 27: 311–27. Daniel E. asa3. html). with Introduction by John Tyler Bonner & Robert M. In Romanucci-Ross.061406. com/ books?id=RKHLD9qNs64C). Human Diet: Its Origin and Evolution. 96–142. Annu Rev Nutr. 1371/ journal. "28. A. ActionBioscience. Snodgrass JJ.200800182. Evol. et al.20379. Evol. PMID 9924136. "A fourth hominin skull from Dmanisi. 0020340& id=15540). W. p. In Peter Ungar. PLoS Biol 2. [6] "Origins of Modern Humans: Multiregional or Out of Africa?" (http:/ / www. ca/ ANTAO1/ Projects/ Bogin. [12] Strait DS.nutr. Dubois' Description of Remains recently found in Java. Evol.1371/journal. utoronto. Evol.093659. "On M. M. google. The Anthropology of Medicine: From Culture to Method (3 ed. Sanders. US: Oxford University Press. (2002). & Tancredi. Smith VS. doi:10.Single Origin (Out of Africa) vs Multiregional" (http:/ / www. Barry (1997). Cambridge. ISBN 978-0-674-03175-3. PMID 17031841.S. pp.1006/jhev.0437. nature. Georgia)". gov/ articlerender. In Steve Jones. cellular.308. html).). (11. com/ ?id=6mxZ1hNBHgkC& pg=PA357& lpg=PA357& dq="The+ Calcaneus+ of+ Australopithecus+ afarensis+ and+ its+ implications+ for+ the+ Evolution+ of+ Bipedality"& q="The Calcaneus of Australopithecus afarensis and its implications for the Evolution of Bipedality").. [23] Figure 1. Bioessays 31 (5): 512–25. and the Unknowable. B. PMID 17393680. de Lumley M. ISBN 0897897366. net/ LeonardARN. [15] Peter Ungar & Teaford. 242. google. April/June). doi:10. [19] Wood..5724.).27. The Descent of Man.. Moerman. Random House.1996. Retrieved 2009-11-23. Gunnell.308/ 5724/ 921g). .921g. doi:10. Retrieved 2009-11-23.J. [14] Walker. "Primates from Rudabánya: allocation of specimens to individuals. Princeton. . New Jersey: Princeton University Press. pdf) (PDF). "Effects of brain evolution on human nutrition and metabolism" (http:/ / www. Vekua A. G.Human evolution 312 References Notes [1] Heng HH (May 2009). The American Heritage Dictionary of the English Language (4th ed. . Ali. [24] Turner W (1895). et al. org/ evolution/ johanson. html). doi:10. the Unknown. Hum.B.org. PMID 11139358. nih. org/ cgi/ content/ summary/ sci. "The genome-centric concept: resynthesis of evolutionary theory". The anatomical record. A Short History of Nearly Everything. Cambridge: Cambridge University Press. ISBN 0195183460. 2000. doi:10. 1986-fact or phantom?". "Early Hominin Diets: Overview and Historical Perspectives" (http:/ / books. "Discovery of a new hominid at Dmanisi (Transcaucasia. . Alan (2006). org/ ASA/ education/ origins/ migration. Vekua A. fcgi?tool=pmcentrez& . & Collard. PMID 20631798. South Hadley. 8(6) 195-207 [20] Wood B (1999). . MacLatchy. [10] Kordos L. Evolution of the Human Diet: The Known. Retrieved 2009-11-23. ISBN 0-691-02369-7. [2] Stringer. Rogers AR. Anth. L..M. Part A. In Michael Ruse & Joseph Travis. Al-Mufarreh. (2002). Alla (March 24. Ambrose (1998). Journal of Human Evolution 44: 479. Jared (1992). J. html). Natl. com/ blog/ display/ 57254/ #ixzz0j820ioz1). Discovery News (http:/ / news. 2006). Sutikna T. Fu. A (2003). com). J..2008. .2006. Groves C. uk/ science/ horizon/ 1999/ supervolcanoes_script. Cell 134 (3): 416–426. Donlon D. Biol. Anatoli P. [52] Freeman. Part A. Maclarnon AM. ISBN 0-13-227584-8 pages 786-788 313 . doi:10. discovery. Johannes.1073/pnas. R.. co. J. Chech M. PMID 9230299. [48] Sample. "The Neanderthal taxonomic position: models of intra. PMID 17232143. Wiuf C. R. nih. humans interbred. Journal of Human Evolution 34 (6): 623–651. Jon C.1469-185X.. Herrera. fcgi?tool=pmcentrez& artid=55569). PMID 17031806.1038/nature08976.1038/369645a0.. A. Arsuaga JL. [40] How Neanderthals met a grisly fate: devoured by humans (http:/ / www. nih. Jeffrey M.and inter-specific craniofacial variation". Stone A. and evolutionary biology 288 (11): 1123–45.cell.1016/j. shtml).276. doi:10. gov/ articlerender. [26] Bermúdez de Castro JM. Hum. "Implications of early hominid labyrinthine morphology for evolution of human bipedal locomotion". and differentiation of modern humans". U. Stoneking M. PMID 18368116. doi:10. iu. nature. PLoS Biol. com/ human/ neanderthal-human-interbreed-dna. Ian (March 24. edu/ news/ page/ normal/ 3142. Evol. bbc. Martínez I. Mallegni F. 51 (4): 360–74. "Neanderthals.1016/S0092-8674(00)80310-4.1111/j. Ascenzi A (2001). "No evidence of Neandertal mtDNA contribution to early modern humans" (http:/ / www. "A reanalysis of the ancient mitochondrial DNA sequences recovered from Neandertal bones".x. "Homo floresiensis: microcephalic. ISBN 0060984031. Mosquera M (1997). doi:10. PMC 368159. pygmoid.021. [45] Brown P. May 17. and Differentiation of Modern Humans" (http:/ / www. co. "Evaluating Neanderthal genetics and phylogeny". Pearson Education.00071. "Flores hominid: new species or microcephalic dwarf?". Nature 464 (7290): 894–897. "A cranium for the earliest Europeans: phylogenetic position of the hominid from Ceprano. [30] Indiana University (March 27.A. doi:10. Wood B. Journal of anatomy and physiology 29 (Pt 3): 424–45. PMID 15514638. doi:10. 19 (8): 1359–66. "A new small-bodied hominin from the Late Pleistocene of Flores. Discoveries in molecular.Human evolution artid=1328414). [36] Jennifer Viegas (2010-05-06). "Late Pleistocene human population bottlenecks. Willerslev E (2007). bbc.151259998. Retrieved 2006-11-26. doi:10. or Homo?". K. pubmedcentral. Pääbo. Stanley H. PMID 18692465. 44 (1): 107–32. PMID 15024415. "Neandertal DNA sequences and the origin of modern humans". PMC 55569. Phillips JL. Evol. PMID 17146600. [28] Manzi G.1006/jhev. nih. gov/ articlerender. 2009. PMID 9162001. . [34] Green RE. Australopithecus. pubmedcentral. Zonneveld F (1994). doi:10. PMID 19391204. Langaney A. "Palaeopathological and variant conditions of the Homo heidelbergensis type specimen (Mauer. 3 February 2000 [43] Stanley H. Biological Reviews 84 (2): 245–257. Nature 431 (7012): 1055–61. Viola. Rosas A. BBC News. Science 280 (5368): 1345–47. The Guardian. (2004). et al. Italy" (http:/ / www. PMID 8208290. "Volcanic Winter. [32] Harvati K (2003). "Solving the Brain's Energy Crisis". 2 (3): E57. PMID 16919706. José M.1038/nature02999. [44] Ambrose. "Scientists discover hominid cranium in Ethiopia" (http:/ / newsinfo. PMID 12140248. Wright R (2006). Marín A (2002). uk/ science/ 2009/ may/ 17/ neanderthals-cannibalism-anthropological-sciences-journal). J. PMID 20336068 [50] Katsnelson. Carbonell E. doi:10. PMID 11504953. [46] Argue D. Morwood MJ. The Observer. DNA proves" (http:/ / news. Shunkov. doi:10. Harper Perennial. "To what extent did Neanderthals and modern humans interact?". 98 (17): 10011–6. The Third Chimpanzee: The Evolution and Future of the Human Animal. doi:10.1016/S0047-2484(02)00208-7. Spain: possible ancestor to Neandertals and modern humans". Inc.). [25] Spoor F. "A hominid from the lower Pleistocene of Atapuerca. guardian. J. Retrieved 2010-08-17. Sci. Svante (2010). [41] " DNA identifies new ancient human dubbed 'X-woman' (http:/ / news. Evol. Mol. Evol. The Scientist.5368. Mol.. et al.1998. html).5317. doi:10. BBC2. Cell 90 (1): 19–30.S. "The first hominin of Europe" (http:/ / www. [29] Czarnetzki. uk/ science/ 2010/ mar/ 24/ new-human-species-siberia). Herron. discovery. Pääbo S (1997). Michael V. Eudald. Glenner H.1126/science. Scott. guardian.. [49] Krause. [42] Supervolcanoes (http:/ / www. [27] Carbonell. doi:10. com/ nature/ journal/ v452/ n7186/ full/ nature06815. (2009). volcanic winter.1016/S0047-2484(03)00029-0. PMC 2602844. PMID 12604307. (2008).. Hum. doi:10. (2008-03-27). cellular. Sánchez D. J. uk/ 2/ hi/ 8583254. The anatomical record. Somarelli.1016/j. 2010). et al.0020057.06. co. [37] Gutiérrez G. Dobyns WB (2006). (2004). March 25. Proc. Schmitz RW. Retrieved 2008-03-26. K. Nature 369 (6482): 645–8. Lowery.04.jhevol.1345. Indonesia". Bence. PMID 9634409. 2010. doi:10. (2005). [31] Herrera.1002/ar. pubmedcentral. PMID 9650103. Derevianko. html). "New hominin found via mtDNA" (http:/ / www.1038/nature06815.1371/journal. bradshawfoundation. Good. gov/ articlerender. [33] Krings M.2008. (2007). Bermúdez de Castro et al. doi:10.a. Krainitzki H. "New species of human ancestor found in Siberia" (http:/ / www. [47] Martin RD.280. [38] Hebsgaard MB.0219. 64 (1): 50–60. Science 276 (5317): 1392–5. "The complete mitochondrial DNA genome of an unknown hominin from southern Siberia".1007/s00239-006-0017-y.1392. Retrieved 2006-04-08. fcgi?tool=pmcentrez& artid=368159).20389.1126/science. doi:10. Qiaomei. com/ evolution/ ). stm)". Bradshaw Foundation.pbio. Acad. co. Evolutionary Analysis (4th ed. 2010). the-scientist. Ann (1998). [51] Gibbons.013. Gilbert MT. fcgi?tool=pmcentrez& artid=2602844). . [35] Serre D. Germany)". [39] Diamond. Nature 452 (7186): 465–469. Press release. "A Complete Neandertal Mitochondrial Genome Sequence Determined by High-Throughput Sequencing" (http:/ / www. . PMC 1328414. 1056/NEJMoa0809716. & Ward.. . [67] " The new batch . P.. FH Smith. stm). Retrieved 2009-11-22. A. R. 2003-06-09. Richard (1994). [64] Searching for traces of the Southern Dispersal (http:/ / www.. [59] Jorde LB.. doi:10. D.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3. [71] Byars. The New England journal of medicine 361 (21): 2056–2065. S. J.CO. com/ releases/ 2009/ 11/ 091120091959. com/ news/ 2007/ 07/ 070718-african-origin. • Flinn. doi:10. ISBN 0-521-32370-3.. Geary. [55] Mcbrearty S. S.20157.3125610. . [61] Modern Humans Came Out of Africa. "Evolution in Health and Medicine Sackler Colloquium: Natural selection in a contemporary human population" (http:/ / www. uk/ sn/ prehistoric_life/ human/ human_evolution/ new_batch1. PMID 19923577. PMID 15605391. Science 308 (5724): 1034. Also ISBN 0-521-46786-1 . (1990).. doi:10. org/ cgi/ content/ abstract/ 308/ 5724/ 1034).. MH. not multiple origins" (http:/ / www3. T. nih. [58] Leakey.1126/science. C. Hill. Retrieved 2007-01-05. "Detecting Ancient Admixture and Estimating Demographic Parameters in Multiple Human Populations" (http:/ / www. D. co.1002/(SICI)1521-1878(199802)20:2<126::AID-BIES5>3. Clarke. "Genetic and fossil evidence for the origin of modern humans". PMID 3025745. fcgi?tool=pmcentrez& artid=2868295). Yearbook of Physical Anthropology 31 (59): 49–83.. "The revolution that wasn't: a new interpretation of the origin of modern human behavior". J. Evolution and Human Behavior. Evol. M.2000.1126/science. Bamshad M.1126/science. [65] Macaulay. Science 291 (5509): 1748–53. V. Whitfield. pubmedcentral. com/ journal/ 71008905/ abstract). PMC 2734152.0906199106. (2009). R Eckhardt. Science Masters Series. 1 May 2009. PMID 3125610. "Modern Human Origins" (http:/ / www.1006/jhev. doi:10. 26. by Dr. Hawks J. Science 239 (4845): 1263–8. PMID 19420049. Andrew. D. New York. doi:10. "How Did Humans Evolve? Reflections on the Uniquely Unique Species" (http:// insects. Stoneking M.000 years ago (http:/ / www. Marta Mirazón Lahr. R. Robert Martin. S. Molecular Biology and Evolution 26 (8): 1823.1002/ajpa. ed. Caspari R (2000). Could Offer Insights Into CJD" (http:/ / www. Lohmueller. Ward. Achilli. PMID 10766948. shtml)". PMID 11102266. Nature 325 (6099): 31–6. [70] Mead. sciencedaily. BBC Science & Nature . C. human-evol. D. htm). J. "Definitive" Study Says (http:/ / news. PMC 2868295. & Pilbeam. The Origin of Humankind. and David Pilbeam . H. edu/ ~landc/ html/ cann/ ). and coalitionary arms races: Why humans evolved extraordinary intelligence. O et al. J Radovcic. J. R. NY: Basic Books. and G Clark (1988). uk/ 2/ hi/ science/ nature/ 2975862.ummz. "Paleolithic technology and human evolution". [60] Wall. pubmedcentral. Science Daily (online) (Science News). the results were published in the online edition of the journal Science.". PMID 19858476. 2009)). Shah.edu/pdfs/Alexander1990. "A Novel Protective Prion Protein Variant that Colocalizes with Kuru Exposure. stm). Brooks AS (2000). [57] Wolpoff..1002/ajpa. social competition.1109792. Phys. C.1059487.1093/molbev/msp096. Proceedings of the National Academy of Sciences 107: 1787.2-R. [69] Medical Research Council (UK) ((November 21. doi:10.lsa. J. doi:10. (1994). BBC. Blackburn. nih. doi:10. 87–89. uk/ 2/ hi/ science/ nature/ 8027269. PMID 11249821. PMID 9631658. S. html) [62] Stringer CB. D. et al. doi:10. Steven (1988). [68] "When humans faced extinction" (http:/ / news. wustl.3136545. . Govindaraju. Andrews P (March 1988). pp. gov/ articlerender. . Rengo. C. Poulter. co.2-K. Rogers AR (February 1998).Human evolution [53] Plummer T (2004). doi:10. ISBN 0465053130. ac. University of Michigan Museum of Zoology Special Publication (University of Michigan Museum of Zoology) (1): 1–38. Stearns. Meehan. M. nationalgeographic. PMID 3136545. htm).1038/325031a0.pdf). bbc.pdf)PDF (345 KB) • edited by Steve Jones. "Multiregional. org/ cgi/ pdf_extract/ 241/ 4867/ 772). Ecological dominance. wiley. foreword by Richard Dawkins. Hummerich. sciencemag. [54] Ambrose SH (2001). V.. 39 (5): 453–563. DW Frayer. G.1330310505 • Alexander. Semino. 314 Further reading • Hill.150. (2005). doi:10.1126/science. doi:10. bbc. (2009).umich. Jones. "Mitochondrial DNA and human evolution" (http:/ / artsci. "Single. (2009). Rapid Coastal Settlement of Asia Revealed by Analysis of Complete Mitochondrial Genomes" (http:/ / www. Martin. W. [63] Cann RL.CO. Uphill. H. Bioessays 20 (2): 126–36.. Cambridge: Cambridge University Press. doi:10. The Cambridge Encyclopedia of Human Evolution. et al. Al-Dujaily. co.edu/~gearyd/Flinnetal2005. "Using mitochondrial and nuclear DNA markers to reconstruct human evolution". uk/ Projects/ sdispersal/ sdispersal. . MH. Suppl 39: 118–64. "Flaked stones and old bones: Biological and cultural evolution at the dawn of technology". (http://web. D. K. cam.The evolution of man. Am.. V. Full text. JN Spuhler. [56] Wolpoff. "Brain Disease 'Resistance Gene' Evolves in Papua New Guinea Community. [66] BBC World News "Africa's genetic secrets unlocked" (http:/ / news. fcgi?tool=pmcentrez& artid=2734152). Hum. Am J Phys Anthropol 112 (1): 129–36.0. Plagnol.0435. J. 10-46. E...0.. Anthropol. D.. . gov/ articlerender.. C. "Origin of the hominidae: The record of african large hominoid evolution between 14 my and 4 my". bbc. Campbell. sciencemag. Science 241 (4867): 772–4. Ewbank.1073/pnas. (2005). G Pope. interscience. Wilson AC (1987).missouri. PMID 15890885. V.. Nature 404 (6777): 490.co.si.edu/anthro/humanorigins/faq/encarta/ encarta. presented by Arizona State University's Institute of Human Origins (http://www. Evolution and Human Origins.psu. David R (2001).1006/jhev. Baldini.0495. "Molecular analysis of Neanderthal DNA from the Northern Caucasus".pdf) (PDF). PMID 11681862. doi:10.org/content/free/figures/ch25. C. A..htm) • Smithsonian – The Human Origins Program (http://www.mnh.2001. Ward. Goodwin. PMID 10761915. Kharitonov. D.uk/2/hi/science/nature/6937476.bbc.. et al. Lidén. doi:10.becominghuman.. 315 External links • BBC: The Evolution of Man (http://www. Reeders.org) . S.pnas. Journal of Human Evolution 41 (5): 463. A. Elmar P J. "Molecular evolution of FOXP2.stm) 2007-08-08 Homo habilis and Homo erectus are sister species that overlapped in time.co.—two ancestral ape chromosomes fused to give rise to human chromosome 2. Genetics 88: 9051–9055. a gene involved in speech and language". • BBC: Finds test human origins theory.bbc.html) • J. shtml) • Illustrations from Evolution (textbook) (http://www.htm) • Becoming Human: Paleoanthropology.si.1038/35006625. Begun. "Origin of human chromosome 2: An ancestral telomere-telomere fusion" (http://www. "The oldest Eurasian hominoid". W. • Ovchinnikov. Kerstin. Nature 418: 870.html) • Smithsonian – Homosapiens (http://www. Götherström. T.org/cgi/reprint/88/20/9051. Anders. (http://news. • Heizmann.mnh. Galina P.edu/anthro/humanorigins/ha/sap. • DNA Shows Neandertals Were Not Our Ancestors (http://www. Romanova. Vitaliy M. William (2000). R.edu/ur/NEWS/news/Neandertal. Wells (October 1991).Human evolution • Wolfgang Enard et al. (2002-08-22). IJdo.evolution-textbook.uk/sn/prehistoric_life/human/human_evolution/index. that between the soft human social sciences and the hard natural sciences.Evolutionary psychology 316 Evolutionary psychology Evolutionary psychology (EP) explains psychological traits—such as memory.. and mutually contradictory human disciplines a single. Other adaptations. to cooperate with others. and so on. including mates and relatives. logically integrated research framework for the psychological. mother mammals and their young offspring sometimes struggle over weaning. For example. making it nearly automatic. according to EP." Evolutionary psychology is founded on a computational theory of mind maintaining that our "inner world" is the result of complex neural structures and interactions in the brain. to quote Steven Pinker. Evolved psychological adaptations (such as the ability to learn a language) interact with cultural inputs to produce specific behaviors (e. For example. the specific language learned). as the functional products of natural selection or sexual selection. . or language—as adaptations. and initiate a fear response. such as the heart. which benefits the mother more than the child. Charles Darwin. to identify and prefer healthier mates. and behavioral sciences—a framework that not only incorporates the evolutionary sciences on a full and equal basis. such as fear of spiders and snakes. our brains can perceive a spider. On the Origin of Species Just as human physiology and evolutionary physiology have worked to identify physical adaptations of the body that represent "human physiological nature. have a marked capacity for cooperation under certain conditions as well. fragmentary. but that systematically works out all of the revisions in existing belief and research practice that such a synthesis requires. for example. Psychology will be based on a new foundation. and modularity."[2] In the distant future I see open fields for far more important researches. Anthropologist John Tooby and psychologist Leda Cosmides note: "Evolutionary psychology is the long-forestalled scientific attempt to assemble out of the disjointed. Humans. that humans have inherited special mental capacities for acquiring language. is common in evolutionary biology." the purpose of evolutionary psychology is to identify evolved emotional and cognitive adaptations that represent "human psychological nature. might include the abilities to infer others' emotions.g. Overview Evolutionary psychology (EP) is an approach to the entire discipline that views human nature as a universal set of evolved psychological adaptations to recurring problems in the ancestral environment. Modern evolutionary psychologists argue that much of human behavior is generated by psychological adaptations that evolved to solve recurrent problems in human ancestral environments. evolutionary psychology sees organisms as often in conflict with others of their species. lungs. that is. perception." EP is. social. "not a single theory but a large set of hypotheses" and a term which "has also come to refer to a particular way of applying evolutionary theory to the mind. gene-level selection. Consistent with the theory of natural selection.[1] They hypothesize. Evolutionary psychologists see those behaviors and emotions that are nearly universal. This view is contrary to the idea that human mental faculties are general-purpose learning mechanisms. Evolutionary psychology applies the same thinking to psychology. while inheriting no capacity specifically for reading and writing. compute that it is a potential threat. to discern kin from non-kin. Proponents of EP suggest that it seeks to heal a fundamental division at the very heart of science --. that of the necessary acquirement of each mental power and capacity by gradation. however. and that the fact that human beings are living organisms demands that psychology be understood as a branch of biology. and immune system. Adaptationist thinking about physiological mechanisms. Light will be thrown on the origin of man and his history. as more likely to reflect evolved adaptations. with an emphasis on adaptation. g. cheater detection mechanisms. at the individual level. Examples include language acquisition modules. and others. but there are key differences between them including the emphasis on domain-specific rather than domain-general mechanisms.[3] called psychological adaptations or evolved cognitive mechanisms or cognitive modules designed by the process of natural selection. ontogeny) and • the proximate mechanism (e. EP uses Nikolaas Tinbergen's four categories of questions and explanations of animal behavior. EP is closely linked to sociobiology. law. Why Questions: Sequential vs. archaeology. Most of what is now labeled as sociobiological research is now conducted in the field of behavioral ecology.[5] EP has been applied to the study of many fields. phylogeny) that resulted in the adaptation (functionality). genetics. including economics. The individual-level categories are • the development of the individual (i. as noted in the table below. The term evolutionary psychology was probably coined by American biologist Michael Ghiselin in a 1973 article published in the journal Science. the importance of mismatch theory. aggression. It also draws on behavioral ecology. brain anatomy and hormones). agent detection mechanisms. .e.e. Static Perspective Historical/ Developmental Current Form Explanation of current form in terms of a historical Explanation of the current form of species sequence Proximate How organisms' structures function Ontogeny Developmental explanations for changes in individuals.. the relevance of measures of current fitness. ethology... foraging mechanisms. Evolutionary psychology mostly focuses on the adaptation (functional) category. politics. and psychology rather than behaviour. anthropology. Two categories are at the species level. How vs. two. adaptation) that a behavior serves and • the evolutionary process (i. and zoology.e. biology. from DNA to their current form Phylogeny The history of the evolution of sequential changes in a species over many generations Mechanism Mechanistic explanations for how an organism's structures work Adaptation A species trait that evolved to solve a reproductive or survival problem in the ancestral environment 317 Evolutionary Why organisms evolved the structures (adaptations) they have The species-level categories (often called "ultimate explanations") are • the function (i.Evolutionary psychology EP proposes that the human brain comprises many functional mechanisms. and sex. literature. psychiatry. alliance-tracking mechanisms. Leda Cosmides and John Tooby popularized the term "evolutionary psychology" in their highly influential 1992 book The Adapted Mind: Evolutionary Psychology and The Generation of Culture. incest avoidance mechanisms. intelligence and sex-specific mating preferences. EP has roots in cognitive psychology and evolutionary biology (See also sociobiology).[4] Jerome Barkow.. artificial intelligence. Evolutionary psychology 318 Related disciplines The content of EP has derived from. acknowledged for work on animal behavior behaviour (ethology) emerged with the work of "The Nobel Prize in Physiology or Medicine 1973". Wilson in 1975 as "the systematic study of the biological basis of all social behavior"[8] and in 1978 as "the extension of population biology and evolutionary theory to social organization". notably studies at the molecular level that examine the relationship between genes. coordinated. Selection designed the information processing of many evolved psychological mechanisms to be adaptively influenced by specific classes of information from the environment. Manifest behavior depends on underlying psychological mechanisms. Human psychology consists of a large number of functionally specialized evolved mechanisms. on the one hand. in conjunction with the external and internal inputs that trigger their activation. Establishing some measure of the relative influence of genetics and environment on behavior has been at the core of behavioral genetics and its variants. In the 1970s two major branches developed from ethology. neurotransmitters and behavior. the biological sciences (especially evolutionary theory as it relates to ancient human environments. social and cognitive psychology. 2. 4. with Karl von Frisch. Several behavioural subjects relate to this core Nobel Laureates Nikolaas Tinbergen (left) and Konrad Lorenz (right) who discipline: in the 1930s the study of animal were.[10] Principles Leading practitioners of EP have proposed a foundation of core premises. Nobel Foundation.[9] Secondly. that get combined. Firstly. DIT is a "middle-ground" between much of social science. which views culture as the primary cause of human behavioral variation. . Evolved psychological mechanisms are functionally specialized to solve adaptive problems that recurred for humans over deep evolutionary time. there was behavioral ecology which placed less emphasis on social behavior by focusing on the ecological and evolutionary basis of both animal and human behavior. Pioneers of the subject Leda Cosmides and John Tooby consider the following five principles as critical: . Evolution by selection is the only known causal process capable of creating such complex organic mechanisms. has a slightly different perspective by trying to explain how human behavior is a product of two different and interacting evolutionary processes: genetic evolution and cultural evolution. Dual inheritance theory (DIT). From psychology there are the primary streams of developmental. and integrated with each other to produce manifest behavior. the study of animal social behavior (including humans) generated sociobiology. on the other. the study of paleoanthropology and animal behavior) and. each sensitive to particular forms of contextual input. Dutch biologist Nikolaas Tinbergen and Austrian Retrieved 2007-07-28. the human sciences especially psychology. Those proposed by David Buss state that: 1. 3. Evolutionary biology as an academic discipline emerged with the modern evolutionary synthesis in the 1930s and 1940s. and human sociobiology and evolutionary psychology which view culture as an insignificant by-product of genetic selection. defined by its pre-eminent proponent Edward O. 5. information processing devices housed in the brain. biologists Konrad Lorenz and Karl von Frisch. developed in the late 1970s and early 1980s.[7] although it was not until the 1970s and 1980s that university departments included the term evolutionary biology in their titles. Different neural circuits are specialized for solving different adaptive problems.some traits are passed down from parents to offspring in genes. The insight of Wallace and Darwin was that this "natural selection" was creative .heritable traits vary within a population (now we know that mutation is the source of some of this genetic variation). Consciousness is a small portion of the contents and processes of the mind. metatheoretical framework that integrates the entire field of psychology. a key component of evolutionary theory. The brain is a physical system. It functions as a computer with circuits that have evolved to generate behavior that is appropriate to environmental circumstances 2. 5. and it could explain the broad scale patterns of evolution. involves three main ingredients: • Genetically based inheritance of traits .[11] These researchers suggest that EP is not simply a subdiscipline of psychology but that evolutionary theory can provide a foundational. • Differential survival and reproduction . • Variation . This is the basis of adaptive evolution. it was based on differential survival of variable individuals. Modern skulls house a Stone age mind. survive.[12] Natural selection Natural selection.these traits will vary in how strongly they promote the survival and reproduction of their bearers. . Neural circuits were designed by natural selection to solve problems that human ancestors faced while evolving into Homo sapiens 3. Most problems experienced as easy to solve are very difficult to solve and are driven and supported by very complicated neural circuitry 4. Selection refers to the process by which environmental Darwin's illustrations of beak variation in the finches of the conditions "select" organisms with the appropriate traits to Galápagos Islands.it could lead to new traits and even new species.Evolutionary psychology 1.[11] 319 General evolutionary theory Main article: Evolution EP is sometimes seen not simply as a subdiscipline of psychology but as a way in which evolutionary theory can be used as a metatheoretical framework within which to examine the entire field of psychology[11] although a few biologists challenge its basic premises. in the same way it has for biology. these organisms will have such traits more strongly represented in the next generation. conscious experience can mislead individuals to believe their thoughts are simpler than they actually are. which refers to the competition among members of the same sex for mating access to the opposite sex. Inclusive fitness resolved "the problem of altruism" by demonstrating that altruism can evolve via kin selection as expressed in Hamilton's rule: cost < relatedness × benefit In other words. (e. The problem with this was that if one organism in a group incurred any fitness costs on itself for the benefit of others in the group. It is metabolically costly. (i. Thus. Inclusive fitness theory resolved the issue of how "altruism" evolved. and kin selection. However. altruism can evolve as long as the fitness cost of the altruistic act on the part of the actor is less than the degree of genetic relatedness of the recipient times the fitness benefit to that recipient.e. pre-Hamiltonian view was that altruism evolved via group selection: the notion that altruism evolved for the benefit of the group. in recent years group selection has been making a comeback (albeit a controversial one) as multilevel selection. Hamilton in 1964 as a revision to evolutionary theory. is essentially a combination of natural selection. Foundations . in its modern form. the type of selective process that is involved here is what Darwin called "sexual selection". therefore reducing its chances of passing on its altruistic traits. the peacock's tail). (e. the organism that benefited from that altruistic act and only acted on behalf of its own fitness would increase its own chance of survival and/or reproduction. including the "group" level.g. Furthermore. Sexual selection can be divided into two types: • Intersexual selection. which was proposed by William D. sexual selection.g.Evolutionary psychology 320 Sexual selection Many traits that are selected for can actually hinder survival of the organism while increasing its reproductive opportunities. This perspective reflects what is referred to as the gene-centered view of evolution and demonstrates that group selection is a very weak selective force. The dominant. thus increasing its chances of passing on its "selfish" traits. General evolutionary theory." What the peacock's tail does do is attract mates. which refers to the traits that one sex generally prefers in the other sex. which posits that evolution can act not just on the "gene" level but on many levels of functional organization. then that organism would reduce its own ability to survive and/or reproduce. Inclusive fitness Inclusive fitness theory. It refers to the sum of an individual's own reproductive success in addition to the effects the individual's actions have on the reproductive success of their genetic relatives. is essentially inclusive fitness theory. • Intrasexual competition. and essentially a "predator magnet. acted "altruistically"). two stags locking antlers). cumbersome. Consider the classic example of the peacock's tail. emotions of revenge and guilt. etc. this typically results in an "evolutionary stable strategy. in offspring.." or "evolutionary stable equilibrium" -.g. System Level: Kin and Family Problem: How are resources best allocated in mating and/or parenting contexts to maximize inclusive fitness? System Level: Non-kin small group Robert Trivers (1972) Sexually dimorphic adaptations that result in a "battle of the sexes. defect. group. . Peacock's tail.Offspring Conflict / Reproductive Value The two sexes often have conflicting strategies regarding how much to invest in offspring. (Also see Gene-centered view of evolution) Also. the behavior of the social insects with sterile workers (e. doves.Evolutionary psychology 321 System level and problem System Level: Individual Problem: How to survive? System Level: Dyad Problem: How to attract a mate and/or compete with members of one's own sex for access to the opposite sex? System Level: Family & Kin Problem: Gene replication. parental investment. cannot be bettered by alternative strategies. the fur of polar bears or the anus of an octopus).g. timing of reproduction. In a population. Being altruistic toward kin can thus have genetic payoffs.. "mom always liked you best"). sibling Parents allocate more resources to their offspring with higher reproductive value (e.g. parent-offspring conflict.strategies that. offspring may have conflicting interests (e. Parents and rivalry. via shared genes in kin. Reproductive success can thus be indirect." parental favoritism. competitive interactions John Maynard with non-kin? How to Smith (1982) select the best strategy given the strategies being used by competitors? System Level: Non-kin small group Problem: How to maintain mutually beneficial relationships with non-kin in repeated interactions? Robert Trivers (1971) "Tit for Tat" Reciprocity One can play nice with non-kin if a mutually beneficially reciprocal relationship is maintained across multiple social interactions. etc. W. "kin selection") / The evolution of sexual reproduction Selection occurs most robustly at the level of the gene.) Game Theory / Evolutionary Game Theory Organisms adapt. vision. antlers).. Charles Darwin Sexual selection Organisms can evolve physical and mental traits (1859) designed specifically to attract mates (e. and how many offspring to have. Facultative.. fast vs. when to wean. ants). adaptations. Hamilton argued that sexual reproduction evolved primarily as a defense against pathogens (bacteria and viruses) to "shuffle genes" to create greater diversity. Altruism toward kin. etc. or species. skin. John von Neumann and Oskar Problem: Morgenstern How to succeed in (1944). not the individual. and cheating is punished. allocation of resources among offspring. or frequency-dependent. on average. antlers. the organism survive in a particular ecology (for example.. or respond. courtship behavior. Examples: hawks vs. etc. How to help those with whom we share genes survive and reproduce? Author Basic ideas Example adaptations Charles Darwin Natural Selection (or "survival selection") The bodies and minds Bones. coy courtship. Hamilton (1964) Inclusive fitness (or a "gene's eye view" of selection. the Peacock's tail) or to compete with members of one's own sex for access to the opposite sex (e. Strategies are evaluated by the probable payoffs of alternatives.g. to competitors depending on the strategies used by competitors.D. cooperate vs. etc. Parental Investment Theory / Parent . pain (1859) of organisms are made up of evolved adaptations designed to help perception.g. Cheater detection. etc. especially immunological variability. Evolutionary psychology 322 Herbert Gintis (2000. evoked culture. while each offspring may want a little more for themselves. invest more heavily in fewer offspring. During the early 1970s. etc. Generalized Reciprocity (Also called "strong reciprocity"). an offspring may want a little more resources from the parent than the parent is willing to give. racism. depending on the parent's condition.) can replicate and spread from brain to brain. a parent would want to allocate their resources equally amongst their offspring. Thus. One can play nice with non-kin strangers even in single interactions if social rules against cheating are maintained by neutral third parties (e. Problem: How to transfer information across distance and time? Source:[13] Middle-level evolutionary theories Middle-level evolutionary theories are theories that encompass broad domains of functioning. cultural fads. To out-group members: Capacity for xenophobia. ethics and human rights. each of which has a better chance of surviving to adulthood. three very important middle-level evolutionary theories were contributed by Robert Trivers[14] [15] [16] • The theory of parent-offspring conflict rests on the fact that even though a parent and his/her offspring are 50% genetically related. Furthermore. etc. Some possible by-products. Additional middle-level evolutionary theories used in EP include: • The Trivers–Willard hypothesis. Because of that. they are applicable across species." cognitive concepts of justice. rituals. females have a better chance of reproducing at least once in comparison to males. r-selected species.) Richard Memetic Selection / Memetics Genes are not the only replicators Dawkins (1976) subject to evolutionary change. Generalized reciprocity may be a set of adaptations that were designed for small in-group cohesion during times of high inter-tribal warfare with out-groups. (The dark side of generalized reciprocity may be that these adaptations may also underlie aggression toward out-groups. "Memes" (e. In essence. etc. Genes and memes may at times co-evolve ("gene-culture co-evolution").g.. if all things are not equal. They are compatible with general evolutionary theory but not derived from it. and parents in poor condition are predicted to favor investment in daughters. large groups governed by rules and laws Problem: How to maintain mutually beneficial relationships with strangers with whom one may interact only once? System Level: Large groups / culture. However.. a bigger overall "pie" results from group cooperation). or "exaptations. produce many offspring. each of which is unlikely to survive to adulthood. which. a majority group members cooperate by generally adhering to social rules. warfare. and others. tunes. Furthermore. reading. . governments. music. ideas." of language may include writing. genocide. etc. parent-offspring conflict refers to a conflict of adaptive interests between parent and offspring. 2003). parents in good condition are predicted to favor investment in sons. To in-group members: Capacity for generalized altruism. while K-selected species.e.). Today the capacity to be altruistic to in-group strangers may result from a serendipitous generalization (or "mismatch") between ancestral tribal living in small groups and today's large societies that entail many single interactions with anonymous strangers. System Level: Non-kin. • r/K selection theory. Language. mathematics. other individuals. and social interactions create a positive sum game (i. institutions. and many of the same evolutionary principles that apply to genes apply to memes as well. which proposes that parents will invest more in the sex that gives them the greatest reproductive payoff (grandchildren) with increasing or marginal investment. relates to the selection of traits in organisms that allow success in particular environments.g. in ecology. All things being equal. according to the Trivers–Willard hypothesis. they are also 50% genetically different. but males in good condition have a better chance of producing high numbers of offspring than do females in good condition. (in unstable or unpredictable environments). Females are the heavier parental investors in our species. acting like a "good Samaritan.. (in stable or predictable environments). a parent may engage in discriminative investment towards one sex or the other. Because the Pleistocene ended a mere 12. Evolutionary psychologists seek to understand psychological mechanisms by understanding the survival and reproductive functions they might have served over the course of evolutionary history. development. In the environment in which ducks evolved. most human adaptations either newly evolved during the Pleistocene. termed domain-specific. and social relationships. Domain-general mechanisms.[17] Definition The term environment of evolutionary adaptedness was coined by John Bowlby as part of attachment theory. Because the first moving being that a duckling was likely to see was its mother. Evolutionary psychology therefore proposes that the majority of human psychological mechanisms are adapted to reproductive problems frequently encountered in Pleistocene environments. Like other organs and tissues.000 people are killed with guns in the US annually. In novel environments. While philosophers have generally considered human mind to include broad faculties. mating. and should solve important problems of survival and reproduction. cognition has functional structure that has a genetic basis. the EEA is defined as the set of historically recurring selection pressures that formed a given adaptation. kidneys.[18] In broad terms. for example.[19] whereas spiders and snakes kill only a handful. and more easily than an unpointed gun.5 million years ago. whereas guns (and rabbits and flowers) were not. and therefore has evolved by natural selection. There is thus a . the mechanism can malfunction by forming an attachment to a dog or human instead. psychological mechanisms sometimes exhibit "mismatches" to the modern environment.000 years ago. livers. these problems include those of growth. parenting. maintenance. people nonetheless learn to fear spiders and snakes about as easily as they do a pointed gun. Some mechanisms. deal with recurrent adaptive problems over the course of human evolutionary history. appeared between 1. this functional structure should be universally shared amongst a species. Mismatches Because humans are mostly adapted to Pleistocene environments. attachment of ducklings to their mother had great survival value for the ducklings. Environment of evolutionary adaptedness EP argues that to properly understand the functions of the brain. evolutionary psychologists describe EPMs as narrowly evolved to deal with specific issues.Evolutionary psychology 323 Evolved psychological mechanisms At a proximal level. or were maintained by stabilizing selection during the Pleistocene. rabbits or flowers. just like hearts. similar to the attachment patterns of ducks. such as reason and lust. a time that roughly coincides with the start of the Pleistocene 1. on the other hand. such as catching cheaters or choosing mates. as well as those aspects of the environment that were necessary for the proper development and functioning of the adaptation.5 and 2. comprising the genus Homo. or EEA for short.8 million years ago. however. a psychological mechanism that evolved to form an attachment to the first moving being would therefore properly function to form an attachment to the mother. Human EEA Humans.[20] A potential explanation is that spiders and snakes were a threat to human ancestors throughout the Pleistocene. lungs. More specifically. evolutionary psychology is based on the hypothesis that. It refers to the environment to which a particular evolved mechanism is adapted. One example is the fact that although about 10. one must understand the properties of the environment in which the brain evolved. deal with evolutionary novelty. differentiation. That environment is often referred to as the environment of evolutionary adaptedness. and immune systems. public records. individuals within a species. dependent and independent variables. kinship. Therefore. Magazine centerfolds and double cheeseburgers pull instincts intended for an EEA where breast development was a sign of health and fertility in a prospective mate. Waistland[23] she explains junk food as an exaggerated stimulus to cravings for salt.[21] [22] This mismatch also shows up in the phenomena of the supernormal stimulus-. and group living. sugar. women's bodies have adapted to the dangers that the environment may pose to the developing fetus when they eat something.[25] 324 Research methods Evolutionary psychologists use several methods and data sources to test their hypotheses. parenting. data from hunter-gatherer societies.a stimulus that elicits a response more strongly than the stimulus for which it evolved. The term was coined by Nobel Laureate Niko Tinbergen to describe animal behavior. and between the same individuals in different contexts.[29] Areas of research Areas of research in evolutionary psychology can be divided into broad categories of adaptive problems that arise from the broader theory of evolution itself: survival. they hunted together with other men and shared their food. during this time many women experience disgust and even vomiting when eating certain foods which may be toxic to the fetus. Recently. smiling faces and attention-grabbing action. One form of evolutionary adaptiveness can be found in morning sickness in women during their first trimester. They also use more traditional experimental methods involving.Highly acclaimed scientist Luke Gunn describes how 'Over thousands of years."[24] she describes how television is an exaggeration of social cues of laughter. If they did it alone they risked not catching anything at all. In her 2007 book. including archeological records. "Supernormal Stimuli: How Primal Urges Overran Their Evolutionary Purpose.' . self-reports. Also. their lives depended on hunting wild game. methods and tools have been introduced based on fictional scenarios.[28] Evolutionary psychologists also use various sources of data for testing. Vomiting is the body's way of coping with the toxins in the environment and keeping them from reaching the child during this critical period when the vital organs are being formed.[27] and multi-agent computer simulations.[26] mathematical models.Evolutionary psychology mismatch between our evolved fear-learning psychology and the modern environment. They could not risk going about such an arduous task on their own. With men being the providers for the family. The function of this physiological reaction was to protect the fetus. and human products. and fat was a rare and vital nutrient. observational studies. Survival The Hunting Hypothesis might explain the emergence of human coalitions as a psychological mechanism. for example. as well as various comparative methods to test for similarities and differences between: humans and other species. These human coalitions can be seen today. and in her 2010 book. Therefore. mating. males and females. the meat would spoil if they caught a large animal and could not finish it on their own. but Harvard psychologist Deirdre Barrett has pointed out that supernormal stimulation governs the behavior of humans as powerfully as that of animals. and fats. [43] Research by Gangestad et al. sexual selection plays a large role in the direction of human evolution. see also. Creativity may be a proxy for good genes. In essence. financial. This obstacle entails that sexual infidelity would be more aversive than emotional infidelity for a man because investing resources in another man's offspring does not lead to propagation of the man's own genes. This has led some researchers to predict sex differences in such domains as sexual jealousy. Women. which means that a woman would also be more at risk for losing those valued traits in a mate who commits an emotional infidelity. are limited by the fact that they can never be certain of the paternity of their children because they do not bear the offspring themselves. If it were a disease that killed 20 year-olds instead of 70 year-olds this may have been a disease that natural selection could have destroyed ages ago. This particular pattern is predicted because the costs involved in mating for each sex are distinct. This hypothesis allows researchers to examine whether women select mates who have characteristics that indicate high genetic quality during the high fertility phase of their ovulatory cycles. on the other hand. such as Alzheimer disease. Thus. should prefer a mate who can offer some kind of resources (e. what matters most is that individuals live long enough to reproduce and pass on their genes. we suffer the aches. commitment). Natural Selection has not eliminated many harmful conditions and nonadaptive characteristics that appear among older adults.[35] to name a few. Haselton and Miller (2006) showed that highly fertile women prefer creative but poor men as short-term mates. especially under short-term mating conditions. unaided by evolutionary pressures against nonadaptive conditions. the grandparents cared for the young. during the ovulatory phase of a woman's cycle (approximately days 10-15 of a woman's cycle). (2004) indicates that highly fertile women prefer men who display social presence and intrasexual competition.[34] and mate retention. to produce and rear a healthy offspring than a woman who mated with a male with low genetic quality. resources. studies have shown that women's preferences vary across the ovulatory cycle.Evolutionary psychology 325 Mating Given that sexual reproduction is the means by which genes are propagated into future generations.. or would be able to acquire. is of interest to evolutionary psychologists who aim to investigate evolved mechanisms to attract and secure mates. Men.[42] a woman who mated with a male with high genetic quality would have been more likely. And as the benefits of evolutionary selection decrease with age.[44] . these traits may act as cues that would help women predict which men may have. Evolutionary Developmental Psychology In evolutionary theory.[30] Several lines of research have stemmed from this interest. on average. the theory hypothesizes that. the need for culture increases. and infirmities of aging.[36] which makes important predictions about the different strategies men and women will use in the mating domain (see above under "Middle-level evolutionary theories").g. whereas men will not. Much of the research on human mating is based on parental investment theory. Another interesting line of research is that which examines women's mate preferences across the ovulatory cycle. In particular. on average.[37] [38] (however. then. According to Paul Baltes. Human mating. Indeed. pains.[40] [41] The theoretical underpinning of this research is that ancestral women would have evolved mechanisms to select mates with certain traits depending on their hormonal status. For example.)[39] wherein females will react more aversively to emotional infidelity and males will react more aversively to sexual infidelity. such as studies of mate selection[31] [32] [33] mate poaching. So why do humans live so long after reproduction? Many evolutionary psychologists have proposed that living a long life improves the survival of babies because while the parents were out hunting. the benefits granted by evolutionary selection decrease with age. These putative preferences are predicted to be especially apparent for short-term mating domains because a potential male mate would only be offering genes to a potential offspring. it predicts that women will be more selective when choosing mates. social behavior and evolutionary theory in his book Sociobiology:The New Synthesis. generosity would increase the fitness of a group. In addition. and how these mechanisms interact with the developmental and current environmental influences to produce behavior.Evolutionary psychology 326 History 19th century After his seminal work in developing theories of natural selection. He wrote two books. EP attempts to identify underlying psychological adaptations (including emotional. the founder of evolutionary psychology might have been Peter Kropotkin. Psychology will be based on a new foundation. This fact contradicted natural selection which favored the fittest individual. Darwin concluded that while generosity decreased the fitness of individuals. the heated controversies surrounding Sociobiology:The New Synthesis.[47] According to Wilson. motivational and cognitive mechanisms). Darwin pondered why humans and animals were often generous to their group members.The Descent of Man. It was during this period that the modern field of ethology emerged. significantly stigmatized the term "sociobiology".[45] Darwin anticipated evolutionary psychology with this quote from the Origin of Species: In the distant future I see open fields for far more important researches. The specific chapter caused considerable controversy as it reignited the nature versus nurture debate. who argued in his 1902 book Mutual Aid: A Factor of Evolution for the evolutionary benefits of behavioral traits related to mutual aid. p. According to Noam Chomsky. Wilson included a chapter on human behavior. in the 1950s. Konrad Lorenz and Nikolaas Tinbergen were pioneers in developing the theoretical framework for ethology for which they would receive a Nobel prize in 1973. his theories on evolutionary psychology were largely ignored. altruism arose due to competition between groups. E O Wilson built upon the works of Lorenz and Tinbergen by combining studies of animal behavior. -. such as the peacock's tail. and also emphasized that organisms are "adaptation executors" rather than "fitness maximizers" (which can help to explain maladaptive behaviors due to "fitness lags" given novel environmental changes). The Origin of Species. In this case. Evolutionary psychology emerged as a more acceptable term in the 1980s that was not tainted by earlier controversies. that of the necessary acquirement of each mental power and capacity by gradation. [46] Post world war II While Darwin's theories on natural selection gained acceptance in the early part of the 20th century. Only after the second world war. Sociobiology In 1975. E O Wilson argues that the field of evolutionary psychology is essentially the same as sociobiology.[48] [49] .Charles Darwin. 449. Charles Darwin devoted much of his final years to the study of animal emotions and psychology. Darwin felt that acts of generosity decreased the fitness of generous individuals. 1859. rather than focus primarily on overt behavior. He introduced the concepts of sexual selection to explain the presence of animal structures that seemed unrelated to survival. and Selection in Relation to Sex in 1871 and The Expression of the Emotions in Man and Animals in 1872 that dealt with topics related to evolutionary psychology. He also introduced theories concerning group selection and kin selection to explain altruism. did interest increase in the systematic study of animal behavior. [8] Wilson. Cambridge. Evolution and motivation. Tooby J (1997-01-13). CO. 157-171.. In Bernard Campbell. Harvard University Press. "Parent-offspring conflict". However. edu/ catalog/ WILSOR. html''Sociobiology:) Harvard University Pre ss. Lewis. . "Darwin and Evolutionary Psychology: Darwin initiated a radically new way of studying behavior". doi:10. 1978. ucsb. html). 5 [3] evolutionary psychology (http:/ / psychegames. [10] Laland. "The evolution of reciprocal altruism" (http:/ / links.1093/icb/14. Robert L. "Parental investment and sexual selection". Quarterly Review of Biology 46 (1): 35–57. Harvard University Press. Ma. Brown. (2004).. Barkow. Robert L. p. jstor. x. hup. adaptationist approaches to human psychology are contentious. On Human Nature. 2002. Retrieved 2007-07-28. (1974). 2009. pdf). edu/ homepage/ Group/ BussLAB/ pdffiles/ evolutionary_psychology_AP_2010. [12] See for example:Gould. John. M. Edward O. 2004. Sexual selection and the descent of man.179. (March 1971). Oxford [Oxfordshire]: Oxford University Press. PMID 17842154. 2003. Nobel Foundation. edu/ research/ cep/ primer. American Zoologist (The Society for Integrative and Comparative Biology) 14 (1): 249–264. 2005. with critics questioning the scientific nature of evolutionary psychology. Stephen Jay (2002).964. Fleischman. 314. com/ evolutionary-psychology. L. ISBN 0-674-00089-7 p. The Structure of Evolutionary Theory. 1975. 2010 (http:/ / homepage. 287-319. . 2002. (1972). 1871-1971. org/ sici?sici=0033-5770(197103)46:1<35:TEORA>2. 70:1. Easton. Buss. 2005 [2] Tooby & Cosmides 2005. Joseph (eds) Wilson. .4077.[50] [51] Criticisms of the field have also been addressed by scholars. Edward O. Robert L. Center for Evolutionary Psychology. Aldine Transaction (Chicago). Symposium paper presented at the Western Psychological Association Conference. Harvard University Press. html). [11] Cosmides. pp. "Folk economics" Southern Economic Journal. [9] Wilson. and with more minor debates within the field itself. Oxford University Press. Pinker. Phoenix. Kim. Michael & Travis. Goetz. in Rubin.E. The Adapted mind: evolutionary psychology and the generation of culture. Cambridge. The New Synthesis. 2003.. Cambridge. ISBN 0202020053. p. ISBN 0-19-510107-3. [16] Trivers. . Ma. AZ. htm) Psyche Games. Tooby & Cosmides. Sense & Nonsense: Evolutionary Perspectives on Human Behavior. Kevin N. Edward O. [15] Trivers. org/ nobel_prizes/ medicine/ laureates/ 1973/ index. 2007 [4] Ghiselin MT (1973). [13] Mills. utexas.4. (Foreword) Evolution: The First Four Billion Years. [7] Sterelny. Science 179 (4077): 964–968.249. psych.g. Paul H. "Evolutionary Psychology: A Primer" (http:/ / www. Accessed August 22. In Ruse.2-S). 136–179. [5] Tooby. harvard." a variation of the term used in Economics. [14] Trivers.1086/406755. Retrieved 2008-02-16. 978-o674031753. [6] "The Nobel Prize in Physiology or Medicine 1973" (http:/ / nobelprize. Perilloux & Buss. pp. psy. ISBN 0674006135.1126/science. [17] See also "Environment of evolutionary adaptation. Oxford. April. and Gillian R. Ma. 0. Leda (1995).[52] See also • • • • • • • • • • Behavioural genetics Dual inheritance theory Ethnic nepotism Evolutionary developmental psychology Evolutionary educational psychology Evolutionary neuroscience Evolutionary Psychology Research Groups and Centers Gene-centered view of evolution Human behavioral ecology List of evolutionary psychologists • Memetics • Multiple discovery • Outline of psychology Notes [1] Confer. Cosmides. Jerome H.1. e. Durrant & Ellis. (http:/ / www. p.Evolutionary psychology 327 Controversies Applying evolutionary theory to animal behavior is uncontroversial. July 2003. doi:10. doi:10. Boston: Pearson Education.W..08. 63. 137–159.. NY NY: W. utexas. A Topical Approach to Life-Span Development (3rd ed. J. [34] Schmitt. & Buss.1073/pnas. edu/ homepage/ Group/ BussLAB/ pdffiles/ prefs_mate_selection_1986_jpsp.tpb. D. Westen. Parental investment and sexual selection. S. doi:10. 211-215. Retrieved 2008-06-16. D. [27] Omar Tonsi Eldakar. [37] Buss. D. S. edu/ comm/ haselton/ webdocs/ haseltonmiller. Sexual Selection and the Descent of Man. ca/ psychology/ 371g/ Smith2001. pdf) Ethology and Sociobiology. doi:10. now often called evolutionary psychology. J.. (2002).108. (2004). [28] Francisco W. 894-917. (http:/ / www. (http:/ / books. PDF) Psychological Science 3(4). & Semmelroth J. The Adapted Mind: Evolutionary psychology and the generation of culture. [33] Li. Trussell. pdf) (PDF). Likelihood of conception with a single act of intercourse: Providing benchmark rates for assessment of post-coital contraceptives. [36] Trivers. 6. oxy. pdf) Human Nature. R. drawing on theory and data from both biology and the social sciences. pdf) Journal of Personality and Social Psychology. P (2006). . & Barnes. D. J. doi:10. PMC 2383986.. "Selfishness as second-order altruism" (http:/ / www. pp. Psychological Science 13(1). Sex differences in human mate preferences: Evolutionary hypotheses tested in 37 cultures. pdf) 328 . 7–12 [40] Haselton.Evolutionary psychology [18] Symons. Cousins. [43] Miller. (2002) Sexual and romantic jealousy in heterosexual and homosexual adults. From vigilance to violence: Tactics of mate retention in American undergraduates.483. psy. chomsky. M. [31] Buss. M.. M. PMID 16458945. D. uwo.. C. and preparedness: Toward an evolved module of fear and fear learning" (http:/ / instruct. Preferences in human mate selection. R.. Waistland: The R/Evolutionary Science Behind Our Weight and Fitness Crisis (2007) NY. Hammerstein.). "Emotions and actions associated with altruistic helping and punishment" (http:/ / www. Weinberg.. Love and Instinct. 559-570.. B. E and Hammerstein. Women's fertility across the cycle increases the short-term attractiveness of creative intelligence. John (2005).1016/j. Tarik Hadzibeganovic. G. 291-317. Chicago: Aldine-Atherton. [35] Buss. P. Theoretical Population Biology 69 (3): 339. A. The necessities and luxuries of mate preferences: Testing the tradeoffs. A.0712173105. & Baird. ucla. W. F. How the Mind Works. (1999). 50-73.2005. Human mate poaching: Tactics and temptations for infiltrating existing relationships. "Game theory and human evolution: A critique of some recent interpretations of experimental games". 50. pdf) Journal of Personality and Social Psychology. A. (1992).09.62. E. P (2006). David Sloan Wilson. (2009).09. Retrieved 2010-08-15. D. ISBN 0195101073. Psychological Review 108 (3): 483–522. NY: W. 1-49. "Human sociobiology.2005. 203-207. utexas. 386–389 [22] Hagen. utexas. The evolution of desire: Strategies of human mating.1016/j. 251–255 [39] Harris. M. WW Norton & Co. PMID 18448681.1016/j. (http:/ / www. Inc.". Retrieved 2010-08-15. pp. pdf) [20] Ohman.. edu/ homepage/ Group/ BussLAB/ pdffiles/ Vigilance_to_Violence_1988. David Sloan Wilson. and psychology. A. [42] Wilcox. G. New York. M. utexas. M.005. Simpson. (2001). pdf) (PDF). 80. utexas. Norton. "Evolution of ethnocentrism on undirected and directed Barabási-Albert networks" (PDF). T. [46] http:/ / www. PMID 16458945 [26] Omar Tonsi Eldakar. J. .. See especially section "Supernormal Stimuli--Why Birds Are Cuckoo" p. pp. phobias. R. (2006). info/ interviews/ 200401--. Sociobiology (http:/ / books. Proceedings of the National Academy of Sciences of the USA 105 (19): 6982–6986. (2008). EH. google. [38] Buss. Deirdre. D. (http:/ / faculty. (http:/ / homepage. google. [23] Barrett.029. Deirdre. NY: McGraw-Hill. New York. London: Temple Smith. pdf) Journal of Personality and Social Psychology. D. amazon. Evolutionary Psychology: The New Science of the Mind. "Game theory and human evolution: a critique of some recent interpretations of experimental games.2009. SUPERNORMAL STIMULI: How Primal Urges Overran Their Evolutionary Purpose." [48] Controversies in the evolutionary social sciences: a guide for the perplexed (http:/ / instruct. com/ ?id=whG6wOFN-A0C& q=ISBN+ 0674000897& dq=ISBN+ 0674000897). S. 12. uwo. psych. edu/ homepage/ Group/ BussLAB/ pdffiles/ SexDifferencesinJealousy. The Science of Good and Evil (http:/ / books. and Dietrich Stauffer. Campbell (Ed. EO (2000). Donald (1992). "On the use and misuse of Darwinism in the study of human behavior".. Anchor Books: New York. [21] Pinker.. (2006). N. J. In B. Kenrick. Lima. org/ content/ 105/ 19/ 6982. (1994).. & Miller. M. PMID 11488376. Oxford University Press. ISBN 9780805077698. W. full. psy.D. Larsen.. northwestern. 31-51. New York: Basic Books. Henry Holt and Co. Contraception. D. J. cdc. PDF) Behavioral and Brain Sciences. psy.3. ISBN 9780674002357..). D. htm [47] Wilson. (1988). & Christensen. pdf) Psychological Science. Women's preferences for male behavioral displays change across the menstrual cycle. 2010 (http:/ / www. Mineka. Theoretical population biology 69 (3): 339–48.W. (2001). 947-955. com/ s/ ref=nb_sb_ss_c_1_9?url=search-alias=stripbooks& field-keywords=supernormal+ stimuli& sprefix=supernorm) [25] Hagen. W. (http:/ / homepage. (2001). (1989). 15(3). P. Garver-Apgar. J. psy. physiology.1037/0033-295X. & Linsenmeier. 17(1). [30] Wilson. C. M. ISBN 978-0205483389. Norton. psy. doi:10.physa. Dunson. . F. Bailey. ca/ psychology/ 371g/ Ohman2001.S. Physica A: Statistical Mechanics and its Applications 388: 4999–5004. A. [19] CDC pdf (http:/ / www. G.. David (2004). . and Rick O'Gorman. com/ ?id=igN6Q9weoYQC& pg=PA51& lpg=PA51& dq="from+ competition+ between+ groups"). [44] Santrock. Evolutionary Psychology 4: 274–286. "Fears.tpb. has in the last quarter of a century emerged as its own field of study. (1972). edu/ homepage/ Group/ BussLAB/ pdffiles/ SexDifferencesinHuman. . [41] Gangestad. google. 1981.. [29] Buss. doi:10. N. (2000b) The mating mind: How sexual choice shaped the evolution of human nature. P. M. edu/ homepage/ Group/ BussLAB/ pdffiles/ Human_Mate_Poaching_2001. . gov/ nchs/ data/ nvsr/ nvsr54/ nvsr54_10. [32] Buss. . (http:/ / homepage. sscnet. C. edu/ psych/ people/ faculty/ bailey/ Publications/ Li et al. Sex differences in jealousy: Evolution. D. 2002. 9. edu/ clint/ evolution/ articles/ Womenâs Preferences for Male behavioral display change across the menstrual cycle. pnas. epjournal. (1986). [45] Shermer (2004). (http:/ / homepage. Belknap Press of Harvard University Press. net/ filestore/ ep04274286. R. G. com/ books?id=vTsnl4KRLegC& dq=inauthor:Deirdre+ inauthor:Barrett) [24] Barrett.005. pdf+ html) (PDF). (http:/ / homepage.. D. & Ellis. N.ucsb. N. Evolutionary psychology: the new science of the mind. Cambridge. L. Boston: Pearson/A and B. [52] Tooby. New York. ISBN 0-679-76399-6. edu/ research/ cep/ papers/ bussconceptual05.com/nibbs/02/ep. M. ISBN 0-19-516335-4. NJ: Wiley. Evolutionary psychology: A new paradigm for psychological science. ISBN 0-670-03151-8. Gallagher & R. (2006). B. ucsb. 1–33). Mass: MIT Press. "Evolutionary psychology: An emerging integrative perspective within the science and practice of psychology" (http://www. (2005).edu/homepage/Group/BussLAB/pdffiles/ ANewParadigmforPsych. ISBN 0-262-10112-2. com/ books?id=9Ni9ggiew1UC& pg=PA17& dq=sociobiology+ evolutionary+ psychology+ controversy& ei=XqZRSYmJOIzukgSGm-mwBg#PPA17..psy. Retrieved 2009-09-08. . (2002). Human nature after Darwin: a philosophical introduction. google. • Joyce.edu/ research/cep/papers/bussconceptual05. ISBN 0-415-21243-X.wiley. David M. • Santrock. G. Oxford [Oxfordshire]: Oxford University Press. Further reading • Buss.uibk. 10. • Clarke. Jerome H. D. Ullica Christina Olofsdotter (2000). Steven (2002).M1) [50] Alcock.). O. • Buss. • Miller. Handbook of evolutionary psychology. The Topical Approach to Life-Span Development(3rd ed. (1995). 6. Chichester: John Wiley & Sons. (2000). R. Oxford University Press. John W.at/~c720126/ humanethologie/ws/medicus/block1/inhalt. 9. J. M. • Medicus.). (2005).human-nature. New York: Norton. In M. Comprehensive Handbook of Psychology. J. New York. Janet C. Conceptual foundations of evolutionary psychology. Cosmides L (2005). • Richards. & Cosmides. [51] Segerstråle. Cambridge: Belknap Press of Harvard University Press. 329 References • Barkow. (2003). Human Nature Review (2): 17–61. Steven (1997). ISBN 0-674-00089-7. (2005).Y: Viking. Volume Three: Biological Psychology (pp. (1995). pdf) (pdf). The Triumph of Sociobiology. Geoffrey P.J. Full text (http://www.html).Y: Doubleday. New York: Wiley & Sons. John (2001). ISBN 0-393-04535-8. in Buss.psych. Sociobiology: the new synthesis. Psychological Inquiry. ISBN 0-205-37071-3. Full text (http://homepage. New York: Routledge. Robert C. Murray (2004). ISBN 0-19-850505-1. psych. • Wright. Oxford [Oxfordshire]: Oxford University Press. How the mind works. (2004). USA. (2005). Reconstructing reason and representation. Missing the Revolution: Darwinism for Social Scientists. Hoboken. In D. The blank slate: the modern denial of human nature. The Evolution of Morality (Life and Mind: Philosophical Issues in Biology and Psychology). 5–67). Cambridge. ISBN 0-07-322626-2. Nelson (Eds.Evolutionary psychology [49] Evolutionary Psychology By Lance Workman. • Pinker.pdf) . Defenders of the truth : the battle for science in the sociobiology debate and beyond.J. Will Reader (http:/ / books. Edward Raymond (2000).ac.pdf#search='evolutionary psychologypdf') • Kennair. Full text (http://media. L.html). The Handbook of Evolutionary Psychology (pp. New York: Vintage Books. Richard (2006). The moral animal: evolutionary psychology and everyday life. N. 1-30.Y: McGraw Hill. ISBN 0-19-513002-2. Buss (Ed. • Tooby.com/product_data/excerpt/38/04713840/0471384038. Mass: The MIT Press. Garden City. ISBN 0-471-26403-2.PDF) • Durrant. "Evolutionary Theory of Human Sciences" (http://homepage. E.utexas. The mating mind: how sexual choice shaped the evolution of human nature.). • Wilson. "Conceptual foundations of evolutionary psychology" (http:/ / www. M. Evolutionary Psychology. 11. ISBN 0-385-49516-1. pp. (2000). • Pinker. David M. ISBN 0-262-03322-4. social. behavioral biology.hbes.anthro.org/). html) devoted to theoretical advances in the fields of biology and cognition. Evolution and Cognition (http://www.edu/nei/) • The NorthEastern Evolutionary Psychology Society (http://www.org/) • The New England Institute for Cognitive Science and Evolutionary Psychology (http://www. international society dedicated to using evolutionary theory to study human nature • The International Society for Human Ethology (http://evolution. journal of the Human Behavior and Evolution Society (http://www.org/ ) • Human Nature: An Interdisciplinary Biosocial Perspective (http://www.neepsociety.aplsnet. These include the evolutionary. and sociological processes as they interact with human social behavior.ac.htm) page at the Human Behavior and Evolution Society (http://hbes. • Evolutionary Anthropology (http://www3.com/science/journal/10905138). neuroscience. systemsthinker. with an emphasis on the conceptual integration afforded by evolutionary and developmental approaches.com/Hbes/books_c. It focuses primarily on the functional unity in which these factors are continuously and mutually interactive.org/article/Evolutionary_psychology) at Scholarpedia • Evolutionary Psychology (http://www.com).com/cgi-bin/jtoc?ID=38641) • Behavioral and Brain Sciences (http://www. htm) Academic societies • Human Behavior and Evolution Society (http://www.scholarpedia. genetic and ecological knowledge • Society for Evolutionary Analysis in Law (http://www. international and interdisciplinary association concerned with evolutionary. biological.aplsnet.springer.citizendium.at/ishe).politicsandthelifesciences.ac.com) • Politics and the Life Sciences (http://www. see the books (http://hbes.dmoz.com/interests/mind/glabachep/glabachwhatisep.at/publications-a.kli.hbes.epjournal. • Biological Theory: Integrating Development.org/Science/Social_Sciences/Psychology/ Evolutionary_Psychology/) at the Open Directory Project • Evolutionary Psychology page (http://en. artificial intelligence. About 30% of the articles have focused on evolutionary analyses of behavior.sealsite.org/wiki/Evolutionary_psychology) at Citizendium • What Is Evolutionary Psychology? by Clinical Evolutionary Psychologist Dale Glaebach (http://www.Evolutionary psychology • For more readings. .com/) 330 External links • Evolutionary Psychology page (http://www. linguistics and philosophy.wiley.com/social+sciences/ anthropology+and+archaeology/journal/12110) advances the interdisciplinary investigation of the biological.org/) is an interdisciplinary peer-reviewed journal published by the Assoication for Politics and the Life Sciences (http://www.bbsonline.com/).univie.shtml).interscience.psychegames. regional society dedicated to encouraging scholarship and dialogue on the topic of evolutionary psychology Journals • Evolutionary Psychology (http://www. and environmental factors that underlie human behavior. • Evolutionary Psychology-Approaches in Psychology (http://www.une.net/) free access online scientific journal • Evolution and Human Behavior (http://www. cognitive science.org/) interdisciplinary articles in psychology.sciencedirect. promotes ethological perspectives on the study of humans worldwide • The Association for Politics and the Life Sciences (http://www.com/evolutionary-psychology. google.com/videoplay?docid=3554279466299738997) with Steven Pinker by Robert Wright (journalist) discussing evolutionary psychology • Video interview (http://video. contextualizing evolutionary psychology within science.com/index.at/publications-a.Evolutionary psychology • Evolution and Development (http://www3. Free access to Winter 2006 issues (http:// www. emphasizing the conceptual integration afforded by evolutionary and developmental approaches.kli. academics and philosophy .org/toc/biot/1/1) 331 Videos • Brief video clip re what EP is (from the "Evolution" PBS Series) (http://www.youtube. (http://video. Evolutionary & Cultural Psychology (http://www.html) by Steven Pinker about his book The Blank Slate: The Modern Denial of Human Nature • Margaret Mead and Samoa.com/videoplay?docid=-4975549474851602314) with Edward O. nurture debate triggered by Coming of Age in Samoa • Video interview (http://video.interscience. html).com/ watch?v=pEmX8Rim-hs) • TED talk (http://www.com/) • Biological Theory: Integrating Development.php/talks/steven_pinker_chalks_it_up_to_the_blank_slate.google. Wilson by Robert Wright (journalist).ac.wiley.mitpressjournals. Evolution and Cognition (http://www.ted. politics. publishes theoretical advances in the fields of biology and cognition.com/journal/118546131/home) Research relevant to interface of evolutionary and developmental biology • Journal of Social.google.jsecjournal.com/videoplay?docid=-3157675332479529894& q=margaret+mead+and+samoa&total=11&start=0&num=10&so=0&type=search&plindex=0) review of the nature vs. evolution debate or the origins debate) is a recurring cultural.[6] with the politics of creationism primarily focusing on the teaching of creation and evolution in public education. While the controversy has a long history. notably Lamarckism.[5] today it is mainly over what constitutes good science. The scientific establishment dismissed it scornfully and the Church of England reacted with fury. political. palaeontology.[9] The debate is sometimes portrayed as being between science and religion. Scientists and theologians have written eloquently about their awe and wonder at the history of the universe and of life on this planet. Science. versus those who accept evolution. and were harshly repressed. as the National Academy of Sciences states: Today. many religious denominations accept that biological evolution has produced the diversity of living things over billions of years of Earth’s history. life. In England these ideas of continuing change were seen as a threat to the fixed social order.[4] this debate is most prevalent in the United States.[12] . but also the fields of geology. and Creationism[10] History of the controversy The creation-evolution controversy originated from Europe and North America in the late 18th century when discoveries in geology led to various theories of an ancient earth. and virtually nonexistent among those in the relevant fields. and the universe.[2] Though also present in Europe and elsewhere. separation of Church and State. and in 1844 the controversial Vestiges popularised transmutation of species. and theology (particularly how different Christians and Christian denominations interpret the Book of Genesis). Religious denominations that do not accept the occurrence of evolution tend to be those that believe in strictly literal interpretations of religious texts. However. and fossils showing past extinctions prompted early ideas of evolution.[1] The dispute is between those who espouse religious belief and thus support a creationist view. The dispute particularly involves the field of evolutionary biology. humanity. Within the scientific community and academia the level of support for evolution is essentially universal. but many Unitarians. Quakers and Baptists opposed to the privileges of the Established church favoured its ideas of God acting through laws.[8] while support for biblically-literal accounts or other creationist alternatives is very small among scientists. science education (and whether the teaching of the scientific consensus view should be 'balanced' by also teaching fringe theories). nuclear physics and cosmology. thermodynamics. Many have issued statements observing that evolution and the tenets of their faiths are compatible.332 Controversy and social impacts Creation–evolution controversy The creation–evolution controversy (also termed the creation vs.[11] Conditions eased.[7] The debate also focuses on issues such as the definition of science (and of what constitutes scientific research and evidence). as supported by scientific consensus. Evolution. explaining that they see no conflict between their faith in God and the evidence for evolution.[3] and often portrayed as part of the culture wars. —National Academy of Sciences. and theological dispute about the origins of the Earth. free speech. but there was no official resistance to evolution by mainline denominations within the United States of America. less militant theological tradition. as did many nonconformists. In Britain this has been attributed to their minority status leading to a more tolerant. and made it a respectable field of study. whose belief in Christ's return depended on a quasi-literal reading of the Bible. However.[13] There was intense interest in the religious implications of Darwin's book. such as fossils and geological findings. unlike subsequent creationists. However. The Reverend Charles Kingsley openly supported the idea of God working through evolution. Warfield who saw it as a natural law expressing God’s will.[14] In this enterprise they received little aid from conservative Christians in Britain and Europe.[16] Though evolution was never condemned by the church.Creation–evolution controversy 333 Controversies in the age of Darwin Publication of Charles Darwin's On the Origin of Species by Means of Natural Selection in 1859 brought scientific credibility to evolution. The main British Creationist movement in this period was the Evolution Protest Movement. freely granting scientists any time they needed before the Garden of Eden to account for scientific observations. Following the lead of figures such as St. creationist leaders and skeptical scientists were usually willing either to adopt a figurative reading of the first chapter of Genesis.[21] Creationism During the 19th century and up until the mid-20th century. many Christians were opposed to the idea and even some of Darwin's close friends and supporters including Charles Lyell and Asa Gray could not accept some of his ideas."[15] lineage.[19] In the immediate post-Darwinian era. George Jackson Mivart and John Zahm Catholics in the United States were accepting of evolution itself while ambivalent towards natural selection and stressing humanity's divinely imbued soul. in the words of one reviewer. coined the term agnostic to describe his position that God’s existence is unknowable.[22] . but the Church of England's attention was largely diverted by theological controversy over higher criticism set out in Essays and Reviews by liberal Christian authors.[14] Thomas Huxley. or to allow that the six days of creation were not necessarily 24-hour days. development of the eugenics movement led many Catholics to reject evolution. such as B.[20] Evolutionary skeptics. few scientists or clerics rejected the antiquity of the earth or the progressive nature of the fossil record.[14] Around the start of the 20th century some evangelical scholars had ideas accommodating evolution.[14] but evolution was also taken up by ape reflects part of the social controversy over prominent atheists including Edward Aveling and Ludwig Büchner and whether humans and apes share a common criticised. initially the more conservative leaning Catholic leadership in Rome held back but gradually adopted a similar position. as "tantamount to atheism. they were not as concerned about geology.[16] [17] Creationists during this period were largely premillennialists. B. few attached geological significance to the Biblical flood.[20] Likewise. formed in the 1930s. who strongly promoted Darwin's ideas while campaigning to end the dominance of science by the clergy. some of whom expressed support for Darwin.[18] However. Creationism was widely accepted and was considered a foundational truth. and Darwin A satirical 1871 image of Charles Darwin as an also took this position. Section 3 of article 1 is binding alike on the Legislature and the school authorities. The textbook in question was Hunter's Civic Clarence Darrow and William Jennings Bryan chat in Biology (1914)."[26] The Court. while the theory of evolution of man may not be taught in the schools of the State. and following the campaigning of William Jennings Bryan several states introduced legislation prohibiting the teaching of evolution.. 363.. the Court overturned the decision on a technicality (the judge had assessed the minimum $100 fine instead of allowing the jury to assess the fine). —Scopes v. however. such legislation was being considered in 15 states. 289 S. because the Act did not establish one religion as the "State religion.[23] The American Civil Liberties Union offered to defend anyone who wanted to bring a test case against one of these laws.As the law thus stands. The Tennessee Supreme Court's decision held in effect that the Butler Act was constitutional under the state Constitution's Religious Preference Clause. Mencken court during the Scopes trial.. nothing contrary to that theory [such as Creationism] is required to be taught. and that there is no unanimity among the members of any religious establishment as to this subject. L... to any religious establishment or mode of worship. the Court decided that the law was not in violation of the Religious Preference provisions of the Tennessee Constitution (section 3 of article 1). Scopes was convicted. So far we are clear that the Legislature has not crossed these constitutional limitations. the Fundamentalist-Modernist Controversy brought a surge of opposition to the idea of evolution.W. the widespread publicity galvanized proponents of evolution. State. which stated that "that no preference shall ever be given. such as Tennessee. Belief or unbelief in the theory of evolution is no more a characteristic of any religious establishment or mode of worship than is belief or unbelief in the wisdom of the prohibition laws. applying that state Constitutional language. the teaching of evolution remained illegal in Tennessee.Furthermore."[28] As a result of the holding. 1927). It would appear that members of the same churches quite generally disagree as to these things.Protestants. .. It only forbids the teaching of evolution of man from a lower order of animals.. among others.Creation–evolution controversy The Butler Act and the Scopes monkey trial In the aftermath of World War I. So far as we know.. and is commonly referred to as the Scopes Monkey Trial. [the Butler Act] requires the teaching of nothing. . By 1925. by law.. John T. there is no religious establishment or organized body that has in its creed or confession of faith any article denying or affirming such a theory. Catholics. The trial was widely publicized by H. 367 (Tenn.[27] The interpretation of the Establishment clause up to that time was that the government could not establish a particular religion as the State religion.[24] [25] Although it overturned the conviction.[29] [30] 334 . When the case was appealed to the Tennessee Supreme Court....It is not necessary now to determine the exact scope of the Religious Preference clause of the Constitution . Scopes accepted. and Jews are divided among themselves in their beliefs. and he confessed to teaching his Tennessee class evolution in defiance of the Butler Act. and passed in some states. and continued campaigning succeeded in removing evolution from school textbooks throughout the United States. held We are not able to see how the prohibition of teaching the theory that man has descended from a lower order of animals gives preference to any religious establishment or mode of worship. [36] In 1961. the United States Supreme Court invalidated a forty year old Arkansas statute that prohibited the teaching of evolution in the public schools. A Little Rock high school biology teacher. Susan Epperson. Morris and John C. spreading anti-evolutionary ideas at fundamentalist churches.[39] Ultimately.'s The Genesis Flood. Whitcomb Jr. "to use the Bible to support an irrational and an archaic concept of static and undeveloping creation is not only to misunderstand the meaning of the Book of Genesis. flood geology. the CSRC broke up over a divide between sensationalism and a more intellectual approach. The Little Rock Ministerial Association supported Epperson's challenge. in the words of the majority opinion.Creation–evolution controversy Epperson v. that humans lived concurrently with dinosaurs. and several states passed legislative acts requiring that this be given equal time with the teaching of evolution. the first major modern creationist book was published: Henry M.[38] Morris' Creation Science Research Center (CSRC) rushed publication of biology text books that promoted creationism. Morris and Whitcomb argued that creation was literally 6 days long."[31] The Court held that the United States Constitution prohibits a state from requiring. "that teaching and learning must be tailored to the principles or prohibitions of any religious sect or dogma. Following this ruling.[37] On the strength of this. with their own colleges."[32] But the Supreme Court decision also suggested that creationism could be taught in addition to evolution. Waters Daniel v. declaring. Morris became a popular speaker.[41] The flood geologists effectively co-opted "the generic creationist label for their hyperliteralist views".[35] With growth. and conferences. with the Southern Baptist Convention and Lutheran Church . and broadcast media. filed suit charging the law violated the federal constitutional prohibition against establishment of religion as set forth in the Establishment Clause. 335 Creation Science As biologists grew more and more confident in evolution as the central defining principle of biology. schools.Missouri Synod outpacing all other denominations. but to do God and religion a disservice by making both enemies of scientific advancement and academic freedom. Morris and others who supported flood geology adopted the terms scientific creationism and creation science.[42] .[34] American membership in churches favoring increasingly literal interpretations of scripture rose.[33] Daniel v. Arkansas In 1968. colleges. and that God created each 'kind' of life individually. and Morris founded the Institute for Creation Research. and also published other books such as Kelly Segrave's sensational Sons of God Return that dealt with UFOlogy. which was promised to be controlled and operated by scientists.[40] During this time. creationism was stripped of overt biblical references and renamed creation science. and demonology against Morris' objections. publishing houses. Waters was a 1975 legal case in which the United States Court of Appeals for the Sixth Circuit struck down Tennessee's law regarding the teaching of "equal time" of evolution and creationism in public school science classes because it violated the Establishment clause of the US Constitution. these churches became better equipped to promulgate a creationist message. educators.[46] Some see science and religion as being diametrically opposed views which cannot be reconciled. list of scientific societies rejecting intelligent design). Mclean v. In 1987 the Supreme Court of the United States ruled that the Louisiana act was unconstitutional. Arkansas In 1982 another case in Arkansas ruled that the Arkansas "Balanced Treatment for Creation-Science and Evolution-Science Act" was unconstitutional because it violated the establishment clause of the U. More accommodating viewpoints. creationists instead thinking that they had better chances with Edwards v. In the similar case of McLean v. Lower courts ruled that the State's actual purpose was to promote the religious doctrine of creation science. but did require that when evolutionary science was taught. and more than one sees the debate as part of the Christian mandate to evangelize. creation science had to be taught as well. but the State appealed to the Supreme Court. the Neo-Creationist intelligent design movement was formed around the Discovery Institute's Center for Science and Culture.Creation–evolution controversy Court cases McLean v. but is widely rejected as unscientific by the science community — primarily because intelligent design cannot be tested and rejected like scientific hypotheses (see for example. including evidence from Francisco Ayala. . Arkansas (see above) the federal trial court had also decided against creationism. the Louisiana legislature passed a law titled the "Balanced Treatment for Creation-Science and Evolution-Science in Public School Instruction Act". and the scientific community. held by many mainstream churches and many scientists. Aguillard In the early 1980s. old earth creationism or intelligent design) as an alternative. consider science and religion to be separate categories of thought. Arkansas however was not appealed to the federal Circuit Court of Appeals. an argument adopted by the state in support of the act. which ask fundamentally different questions about reality and posit different avenues for investigating it. with the mainstream scientific consensus on the origins and evolution of life challenged by creationist organizations and religious groups who desire to uphold some form of creationism (usually young earth creationism. because the law was specifically intended to advance a particular religion. it stated in its opinion that "teaching a variety of scientific theories about the origins of humankind to school children might be validly done with the clear secular intent of enhancing the effectiveness of science instruction.[43] 336 Intelligent Design In response to Edwards v. Controversy in recent times The controversy continues to this day. Constitution. The act did not require teaching either evolution or creationism as such. Its goal is to restate creationism in terms more likely to be well received by the public. creationism."[45] It has been viewed as a "scientific" approach to creationism by creationists." leaving open the door for a handful of proponents of creation science to evolve their arguments into the iteration of creationism that came to be known as intelligent design. Aguillard. not an undirected process such as natural selection. arguing that the act was about academic freedom for teachers.S.[47] Public opinion in regards to the concepts of evolution. creation science. Later it used a less religious image. Most of these groups are explicitly Christian. Much of the transcript of the case was lost. however. Creationists had lobbied aggressively for the law. and makes the claim that "certain features of the universe and of living things are best explained by an intelligent cause. At the same time. Aguillard. policy makers. then was [44] renamed the Center for Science and Culture. Edwards v. and intelligent The Discovery Institute's Center for the Renewal of Science and Culture used banners based on "The Creation of Adam" from the Sistine Chapel. The Kitzmiller v.[52] and on 13 February 2007. as opposed to God. 4 of the 6 conservative Republicans who approved the Critical Analysis of Evolution classroom standards lost their seats in a primary election. the Board voted 6 to 4 to reject the amended science standards enacted in 2005..[54] This ultimately resulted in the "Dover Trial. because the law was specifically intended to advance a particular religion. The Kansas Evolution Hearings were a series of hearings held in Topeka. and is essentially creationism. the Discovery Institute. the Intelligent Design movement has taken an anti-evolution position which avoids any direct appeal to religion. which went to trial on 26 September 2005 and was decided on 20 December 2005 in favor of the plaintiffs. who charged that a mandate that intelligent design be taught in public school science classrooms was an unconstitutional establishment of religion. though the only paper arguing for it published in a scientific journal was accepted in questionable circumstances and quickly disavowed in the Sternberg peer review controversy. "contrary to typical editorial practices"."[50] Kansas evolution hearings In the push by intelligent design advocates to introduce intelligent design in public school science classrooms. The Kansas State Board of Education eventually adopted the institute's Critical Analysis of Evolution lesson plans over objections of the State Board Science Hearing Committee. and hence religious. Kansas 5 May to 12 May 2005. antecedents". made widely publicised claims that it was a new science. was a "significant departure" from the journal's normal subject area and was published at the former editor's sole discretion. creationists renewed their efforts to introduce creationism into public school science classes. the Discovery Institute. Young Earth creationists often believe that the Universe has a similar age to the Earth's.. so people can understand what the debate is about. Aguillard decision by the Supreme Court of the United States. This effort resulted in intelligent design. Dover decision held that intelligent design was not a subject of legitimate scientific research." Kitzmiller v.000 years. This belief generally has a basis in a literal and inerrant interpretation of the Bible."[53] The Dover Trial Following the Edwards v. literally as described in Genesis. The definition of science was once again limited to "the search for natural explanations for what is observed in the universe.[55] 337 Viewpoints Young Earth creationism Young Earth creationism is the belief that the Earth was created by God within the last 10.[51] On 1 August 2006. within the approximate timeframe of biblical genealogies (detailed for example in the Ussher chronology). More recently. arranged to conduct hearings to review the evidence for evolution in the light of its Critical Analysis of Evolution lesson plans. and that it "cannot uncouple itself from its creationist.[49] President Bush commented endorsing the teaching of Intelligent design alongside evolution "I felt like both sides ought to be properly taught . Dover Area School District. . The moderate Republican and Democrats gaining seats vowed to overturn the 2005 school science standards and adopt those recommended by a State Board Science Hearing Committee that were rejected by the previous board.Creation–evolution controversy design is fluctuating. with the Biological Society of Washington stating that it did not meet the journal's scientific standards. in which the Court held that a Louisiana law requiring that creation science be taught in public schools whenever evolution was taught was unconstitutional. and electioneering on behalf of conservative Republican candidates for the Board. which sought to avoid legal prohibitions by leaving the source of creation to an unnamed and undefined intelligent designer. Scientists argue that Intelligent design does not represent any research program within the mainstream scientific community. Creationist cosmologies are attempts by some creationist thinkers to give the universe an age consistent with the Ussher chronology and other Young-Earth timeframes. the hub of the intelligent design movement.[48] Its leading proponent. specifically. instead of faith being in opposition to biological evolution. Miller. but rather as a literary framework or allegory. Old Earth creationists interpret the Genesis creation narrative in a number of ways. Neo-Creationism Neo-Creationists intentionally distance themselves from other forms of creationism. who is a prominent board member of Kansas Citizens for Science). 24-hour day creation of the Young Earth Creationist view. Neo-Creationism currently exists in the form of the intelligent design movement. Dover Area School District. . but that details of the evolutionary theory are questionable. Notable examples have been biologist Kenneth R. This position does not generally exclude the viewpoint of methodological naturalism. it is therefore well accepted by people of strong theistic (as opposed to deistic) convictions. on the interpretation of the Bible). Their goal is to restate creationism in terms more likely to be well received by the public. Theistic evolution Theistic evolution is the general view that. including. Theistic evolution can synthesize with the day-age interpretation of the Genesis creation myth.g. with specific reference to points raised by intelligent design proponents. Theistic evolutionists have also been active in Citizens Alliances for Science that oppose the introduction of creationism into public school science classes (one example being evangelical Christian geologist Keith B. and hold a range of underlying theological viewpoints (e. but that the creation event of Genesis is not to be taken strictly literally. education policy makers and the scientific community. This group generally believes that the age of the Universe and the age of the Earth are as described by astronomers and geologists. and to bring the debate before the public. It aims to re-frame the debate over the origins of life in non-religious terms and without appeals to scripture. who testified for the plaintiffs in Kitzmiller v. It generally views evolution as a tool used by a creator god. Miller and theologian John Haught. Theistic evolutionists have frequently been prominent in opposing creationism (including intelligent design). consecutive. an organization that has created and maintains a statement signed by American Christian clergy of different denominations rejecting creationism. which has a 'big tent' strategy making it inclusive of many Young Earth Creationists (such as Paul Nelson and Percival Davis). Another example is the Clergy Letter Project. a long standing convention of the scientific method in science. preferring to be known as wholly separate from creationism as a philosophy. who is both the first cause and immanent sustainer/upholder of the universe. evolution. however most adherents consider that the first chapters of Genesis should not be interpreted as a "literal" description. Neo-creationists may be either Young Earth or Old Earth Creationists. some or all classical religious teachings about God and creation are compatible with some or all of modern scientific theory.Creation–evolution controversy 338 Old Earth creationism Old Earth creationism holds that the physical universe was created by God. that each differ from the six. has often been made against the exclusive teaching of evolution.Creation–evolution controversy 339 Naturalistic evolution Naturalistic evolution is the position of acceptance of biological evolution and of metaphysical naturalism (and thus rejection of theism and theistic evolution). Principles such as uniformitarianism. —National Academy of Sciences. and therefore supernatural explanations for such events are outside the realm of science.[60] Definitions Fact: In science. fact The argument that evolution is a theory.[62] The argument is related to a common misconception about the technical meaning of "theory" that is used by scientists. and subjectivity. If the deductions are incorrect.[59] Mainstream proponents accuse the creationists of conflating the two in a form of pseudoscience. Hypothesis: A tentative statement about the natural world leading to deductions that can be tested. a well-substantiated explanation of some aspect of the natural world that can incorporate facts. and what is accepted as a fact today may be modified or even discarded tomorrow. laws. without assuming the existence or non-existence of the supernatural. as well as on avoidance of Baconian idols. In common usage. and unproven assumptions."[63] . "theory" often refers to conjectures. inferences. explanations are limited to those based on observations and experiments that can be substantiated by other scientists. Theory: In science. hypotheses. A prominent proponent of this viewpoint is British evolutionary biologist Richard Dawkins. Occam's Razor or parsimony. an observation that has been repeatedly confirmed and for all practical purposes is accepted as "true. is never final. it remains neutral on subjective subjects such as religion or morality. the original hypothesis can be abandoned or modified. The methodological assumption is that observable events in nature are explained only by natural causes.[56] In countering this claim. —National Academy of Sciences. Law: A descriptive generalization about how some aspect of the natural world behaves under stated circumstances." Truth in science. it becomes more probable that the hypothesis is correct. Science and Creationism[61] Limitations of the scientific endeavor In science. not a fact. which is equated by many creationists with atheism. however. philosophers of science use the term methodological naturalism to refer to the long standing convention in science of the scientific method. Hypotheses can be used to build more complex inferences and explanations. Explanations that cannot be based on empirical evidence are not a part of science. and tested hypotheses. If the deductions are verified. Science and Creationism[61] Theory vs. error. in science.[57] Creationists claim that supernatural explanations should not be excluded and that scientific work is paradigmatically close-minded. "theory" usually means "a plausible or scientifically acceptable general principle or body of principles offered to explain phenomena. However. and the Copernican principle are claimed to be the result of a bias within science toward philosophical naturalism.[58] Because modern science tries to rely on the minimization of a priori assumptions. Arguments relating to the definition and limits of science Critiques such as those based on the distinction between theory and fact are often leveled against unifying concepts within scientific disciplines. Duane Gish. but those that are untestable are not. I have changed my mind about the testability and logical status of the theory of natural selection.[66] In what one sociologist derisively called "Popper-chopping. in Unended Quest. —Stephen Jay Gould.S.. and I am glad to have an opportunity to make a recantation". Theories are structures of ideas that explain and interpret facts. Judge William R." while pointing out it had "scientific character". While many Creationists claim creation is a scientific theory other Creationists have stated that neither creation nor evolution is a scientific theory (and each is equally religious). Evolution as Fact and Theory[64] 340 Falsifiability Philosopher of science Karl R. but apples did not suspend themselves in mid-air. a time before most similarly complex lifeforms had evolved.[68] For example. later in the article is "The theory of natural selection may be so formulated that it is far from tautological."[67] opponents of evolution seized upon Popper's definition to claim evolution was not a science. but it turns out to be not strictly universally true. creationists claimed that a key evolutionary concept. a leading Creationist proponent. paleontologist Stephen Jay Gould wrote: Evolution is a theory.Creation–evolution controversy Exploring this issue. And humans evolved from ape-like ancestors whether they did so by Darwin's proposed mechanism or by some other yet to be discovered. that all life on Earth is descended from a single common ancestor. wrote in a letter to Discover magazine (July 1981): "Stephen Jay Gould states that creationists claim creation is a scientific theory. Popper wrote. Popper wrote admiringly of the value of Darwin's theory."[73] Debate among some scientists and philosophers of science on the applicability of falsifiability in science continues. Facts are the world's data. Popper set out the concept of falsifiability as a way to distinguish science and pseudoscience: Testable theories are scientific. Haldane both pointed out that if fossil rabbits were found in the Precambrian era."[69] Popper responded to news that his conclusions were being used by anti-evolutionary forces by affirming that evolutionary theories regarding the origins of life on earth were scientific because "their hypotheses can in many cases be tested."[70] However. ultimately declaring it "simply not science.[71] In fact. and claimed creationism was an equally valid metaphysical research program. and claimed it never would be. And facts and theories are different things. Facts do not go away when scientists debate rival theories to explain them. not rungs in a hierarchy of increasing certainty. In this case it is not only testable..[72] Only a few years later. a possible framework for testable scientific theories. Arkansas Board of Education.B.[74] However. simple falsifiability tests for common descent have been offered by some scientists: For instance. "that would completely blow evolution out of the water."[75] [76] Falsifiability has also caused problems for creationists: In his 1982 decision McLean v. His conclusion. biologist and prominent critic of creationism Richard Dawkins and J.I still believe that natural selection works in this way as a research programme. It is also a fact."[77] . "I have in the past described the theory as "almost tautological". Einstein's theory of gravitation replaced Newton's. was not mentioned as testable by Popper.[65] However. Popper declared "I have come to the conclusion that Darwinism is not a testable scientific theory but a metaphysical research programme. Overton used falsifiability as one basis for his ruling against the teaching of creation science in the public schools. pending the outcome. Nevertheless. Creation–evolution controversy 341 Conflation of science and religion Appeal to consequences A number of creationists have blurred the boundaries between their disputes over the truth of the underlying facts. fossil records. A theory of universal common descent based on evolutionary principles was proposed by Charles Darwin and is now generally accepted by biologists. macroevolution. Biology Disputes relating to evolutionary biology are central to the controversy between Creationists and the scientific community. the most recent common ancestor of all currently living organisms. Morris. starting with the very beginning. as it focuses on embryology. the vast majority of Creationists reject this theory. with their purported philosophical and moral consequences. is believed to have appeared about 3. of evolution. and explanatory theories. controversial exclusively within the animal kingdom. comparative physiology and comparative biochemistry. typically based on faith. In response to perceived crises in modern science. The aspects of evolutionary biology disputed include common descent (and particularly human evolution from common ancestors with other members of the Great Apes). .g.g. Examples of these arguments include those of prominent creationists such as Ken Ham[78] and Henry M.9 billion years ago. common descent of animals is completely noncontroversial. This type of argument is known as an appeal to consequences. The last universal common ancestor.[79] Disputes relating to science Many creationists strongly oppose certain scientific theories in a number of ways. There. creation science. accusations of bias within the scientific community. comparative anatomy. The scientific community has responded by pointing out that their conversations are frequently misrepresented (e.[82] [83] [84] Evidence of common descent includes evidence from genetics. researchers argued over a variety of topics.[80] and claims that discussions within the scientific community reveal or imply a crisis. any controversy resides in the microbial world. creationists claim to have an alternative. and/or intelligent design. that is. Evolution: what's the real controversy?[81] A group of organisms is said to have common descent if they have a common ancestor. Common descent [The] Discovery [Institute] presents common descent as A phylogenetic tree based on rRNA genes. Michael Behe). and that the creationists' alternatives are generally pseudoscientific. namely the relationship among the three main branches of life. including opposition to specific applications of scientific processes. anatomy. geographical distribution of species. —John Timmer. and the existence of transitional fossils. by quote mining) in order to create the impression of a deeper controversy or crisis. and the fossil record to raise questions about them. With a few exceptions (e. and is a logical fallacy. In the real world of science. and not a scientific theory. but distinctively hominid fossils have been found dating to 3. and are taken as strong evidence of recent common ancestry. and then the chimpanzee (genus Pan) split from the line leading to the humans. Molecular evidence further suggests that between 8 and 4 Ma. and the orangutan lineage (subfamily Ponginae) diverged about 12 Ma. Macroevolution is defined by the scientific community to be evolution that occurs at or above the level of species. fossil proto-orangutans may be represented by Sivapithecus from India and Griphopithecus from Turkey. Creationists also dispute science's interpretation of genetic evidence in the study of human evolution. However. Creationists dispute there is evidence of shared ancestry in the fossil evidence. and that by the early Miocene.[90] However Creationists frequently disagree where the dividing lines would be. 342 . The scientific community considers that there is strong evidence for even such more restrictive definitions. and argue either that these are misassigned ape fossils (e. in the case of Genesis) as an alternative viewpoint to the scientific account. and both arguments have been rejected by the scientific community as pseudoscience.[86] We have no fossil record of this divergence. neither argument has been accepted for publication in a peer-reviewed scientific journal. circa 26-24 Ma. and that scientists are coming to this interpretation only because they have preconceived notions that such shared relationships exist.4 percent identical. Fossil evidence suggests that humans' earliest hominoid ancestors may have split from other primates as early as the late Oligocene. and Homo neanderthalensis. only one distinct human species survives. They argue that it is a "dubious assumption" that genetic similarities between various animals imply a common ancestral relationship.[91] Creation myths (such as the Book of Genesis) frequently posit a first man (Adam.[31] Macroevolution Many creationists now accept the possibilities of microevolution within "kinds" but refuse to accept and have long argued against the possibility of macroevolution. the judge wrote "The overwhelming evidence at trial established that ID is a religious view. a mere re-labeling of creationism. at 6 or 7 Ma (see Toumaï). Creationists also argue that genetic mutations are strong evidence against evolutionary theory because the mutations required for major changes to occur would almost certainly be detrimental. Recent arguments against (such restrictive definitions of) macroevolution include the intelligent design arguments of irreducible complexity and specified complexity. Homo habilis." Biologist Richard Dawkins published a book 'The Greatest Show on Earth: The Evidence for Evolution' giving evidence for evolution and Macroevolution. if vaguer. often relating to the emergence of new body forms or organs. macroevolution can be considered to be a fact.[88] Today. that Java man was a gibbon[89] ) or too similar to modern humans to designate them as distinct or transitional forms. Analysis of fossil evidence and genetic distance are two of the means by which scientists understand this evolutionary history.[85] Evidence from the molecular dating of genetic differences indicates that the gibbon lineage (family Hylobatidae) diverged between 18 and 12 Ma.[87] Comparisons of chimpanzee and human DNA show the two are approximately 98. Further when taken to court in attempt to introduce ID into the classroom.2 Ma (see Lucy) and possibly even earlier. including Homo erectus. but the evidence for this is more complex. first the gorillas. but many earlier species have been found in the fossil record.g.Creation–evolution controversy Human evolution Human evolution is the study of the biological evolution of humans as a distinct species from its common ancestors with other animals. dated to around 10 Ma. Under this definition. While there is no fossil evidence thus far clearly documenting the early ancestry of gibbons. Creationists however tend to apply a more restrictive. as evidenced by observed instances of speciation. the adaptive radiation of many different hominoid forms was well underway. definition of macroevolution. however."[94] The evolution of whales has however been documented in considerable detail. 60. usually traceable in the same geological outcrop. It is plausible. Thus. They believe that somehow a whale must have evolved from an ordinary land-dwelling animal. however. might originally have only been meant for gliding. 343 Geology Many believers in Young Earth Creationism – a position held by the majority of proponents of Flood Geology – accept biblical chronogenealogies (such as the Ussher chronology which in turn is based on the Masoretic version of the Genealogies of Genesis). A common creationist argument is that no fossils are found with partially functional features. To explain these jumps.. Dawkins goes on to say that the reason for this "losing battle" is that many of these critics are theists who "simply don't want to see the truth". In particular. progressing research and discovery managed to fill in several gaps and continues to do so. and would have no hope for survival.[96] [97] They believe that God created the universe approximately 6000 years ago. both of which are central to mainstream geological . but it will never be known in detail. even if it were possible for enough fossils to survive to show a close transitional change. As another example. for example. The theory of punctuated equilibrium developed by Stephen Jay Gould and Niles Eldredge is often mistakenly drawn into the discussion of transitional fossils. This theory. could be accomplished over 60. Critics of evolution often cite this argument as being a convenient way to explain off the lack of 'snapshot' fossils that show crucial steps between species. geology. they only exemplify snapshots of this process. Nowadays. the transition itself can only be illustrated and corroborated by transitional fossils.[95] Furthermore. in the space of six days. often show small jumps in morphology between periods of morphological stability. Alan Haywood stated in Creation and Evolution that "Darwinists rarely mention the whale because it presents them with one of their most insoluble problems. only a very small percentage of all life-forms that ever have existed can be expected to be discovered.000 years is too small a gap to be identified or identifiable in the fossil record. a wing. evolution's critics will never be satisfied. promoters of evolution (such as Richard Dawkins) have pointed out that. These transitions. A land mammal that was in the process of becoming a whale would fall between two stools—it would not be fitted Reconstruction of Ambulocetus natans for life on land or at sea. wings can still have all of these functions. as one of the transitional fossils. The precursor to. and planetary science that give ages in conflict with the young Earth idea. which took to the sea and lost its legs . creationists dispute the reliability of radiometric dating and isochron analysis.000 years. as the discovery of one "missing link" itself creates two more so-called "missing links" on either side of the discovery. trapping flying prey.Creation–evolution controversy Transitional fossils It is commonly stated by critics of evolution that there are no known transitional fossils. that a complex feature with one function can adapt a wholly different function through evolution. but they are also used in active flight. Gould and Eldredge envisaged comparatively long periods of genetic stability separated by periods of rapid evolution. For example the change from a creature the size of a mouse. pertains only to well-documented transitions within taxa or between closely related taxa over a geologically short period of time. to one the size of an elephant.. Much of creation geology is devoted to debunking the dating methods used in anthropology. with Ambulocetus. However. Due to the special circumstances required for preservation of living beings. with a rate of change too small to be noticed over any human lifetime. and/or mating display. described as looking like a three-metre long mammalian crocodile. Although transitional fossils elucidate the evolutionary transition of one life-form to another.[92] [93] This position is based on a misunderstanding of the nature of what represents a transitional feature. [100] The scientific community points to numerous flaws in these experiments. depending on its chemical environment. Apparently inconsistent radiometric dates are often quoted to cast doubt on the utility and accuracy of the method.Creation–evolution controversy theories of the age of the Earth.8 billion years ago. beryllium-7 has been shown to change by up to 1.3 ± 1. a completely different kind of decay can occur.. etc. The recent science of nucleocosmochronology is extending the approaches used for Carbon-14 dating to the dating of astronomical features. . which is in a plasma state where electrons are not bound to atoms. has electrons in normal positions. However. All normal matter. or simply the result of incorrectly evaluating legitimate data. 2. or anything colder than several hundred thousand degrees. 344 Other sciences Cosmology Whilst Young Earth Creationists believe that the Universe was created by the Judeo-Chrisitian God approximately 6000 years ago. the Moon. which they claim demonstrate that 1. meteorites. Wiens. and to the fact that the creationist scientists conducting them were untrained in experimental geochronology. Another case is material inside of stars. Mainstream proponents who get involved in this debate point out that dating methods only rely on the assumptions that the physical laws governing radioactive decay have not been violated since the sample was formed (harking back to Lyell's doctrine of uniformitarianism). Nuclear physics Creationists point to experiments they have performed. They also point out that the "problems" that creationists publicly mentioned can be shown to either not be problems at all. a full critique of the entire parameter-fitting analysis. . are issues with known contamination... ..[99] Old Earth Creationists and some others do not necessarily dispute these figures. In the extremely hot stellar environment.5%. a physicist specialising in isotope dating states: There are only three quite technical instances where a half-life changes. to the fact that their results have not been accepted for publication by any peer-reviewed scientific journal. For example based upon this emerging science.5 billion years of nuclear decay took place over a short period of time. Roger C. which relies on dozens of radionuclei parent and daughter pairs. from which they infer that "billion-fold speed-ups of nuclear decay" have occurred. so the dates of rocks made by electron-capture decays would only be off by at most a few hundredths of a percent.5 billion years. has not been done by creationists hoping to cast doubt on the technique...[98] Young Earth creationists reject these ages on the grounds of what they regard as being tenuous and untestable assumptions in the methodology. 'Bound-state beta decay' occurs when the nucleus emits an electron into a bound electronic state close to the nucleus. so these instances never apply to rocks. .. The consensus of professional scientific organisations worldwide is that no scientific evidence contradicts the age of approximately 4.7 billion years ago. the current scientific consensus is that the Universe as we know it emerged from the Big Bang 13. a core principle underlying nuclear physics generally.. the Galactic thin disk of the Milky Way galaxy is estimated to have been formed 8. Only one technical exception occurs under terrestrial conditions.. and radiometric dating in particular. [H]eavier atoms are even less subject to these minute changes. The artificially-produced isotope. . and these do not affect the dating methods [under discussion]":[103] 1. and this is not for an isotope used for dating.[101] [102] In refutation of young-Earth claims of inconstant decay rates affecting the reliability of radiometric dating. They usually dispute these methods based on uncertainties concerning initial concentrations of individually considered species and the associated measurement uncertainties caused by diffusion of the parent and daughter isotopes. a massive violation of the principle that radioisotope decay rates are constant. such as everything on Earth.. . [111] reply that there is no scientific controversy and that the controversy exists solely in terms of religion and politics."[119] . A Christian Perspective[104] 345 Misrepresentations of science Quote mining As a means to criticise mainstream science. being the overwhelming majority of the scientific community and science education organizations. textbooks must teach intelligent design alongside evolution. dogmatism about evolution" has sapped "America's scientific soul.[105] However. These atomic clocks slow down very slightly (only a second or so per year) as predicted by Einstein's theory of relativity. publishers have a strong incentive to be certified by the board as 'conforming 100% to the state's standards'. that the TalkOrigins Archive has created "The Quote Mine Project" for quick reference to the original context of these quotations. and apparently as students learn more about biology. almost universally these have been shown to be quote mines that do not accurately reflect the evidence for evolution or the mainstream scientific community's opinion of it.. —Roger C. The last case also involves very fast-moving matter. or "politics of science and religion. but appear to acknowledge criticisms similar to those of creationists.. at length. and that the scientific community’s resistance to this approach was bad public relations. Opponents. scientists who ostensibly support the mainstream theories. and question the validity of the fossil record. they find objections to evolution less convincing. . creationists have been known to quote. Wiens. Dover Area School District trial.[108] [110] The American Association for the Advancement of Science and other science and education professional organizations say that Teach the Controversy proponents seek to undermine the teaching of evolution[108] [112] while promoting intelligent design.[106] [107] Many of the same quotes used by creationists have appeared so frequently in Internet discussions due to the availability of cut and paste functions. suggesting that “teaching the controversy” rightly as a separate elective course on philosophy or history of science. Radiometric Dating. or highly out-of-date. by a vote of 13 to 2.. "Because Texas is the second-largest textbook market in the United States. the Texas Board of Education.[117] George Mason University Biology Department introduced a course on the creation/evolution controversy. No rocks in our solar system are going fast enough to make a noticeable change in their dates. said.[113] [114] [115] and to advance an education policy for US public schools that introduces creationist explanations for the origin of life to public-school science curricula.Creation–evolution controversy 3. voted that at least in Texas. "I think the new standards are wonderful ..[118] On March 27. a dentist and chair of the board.[106] Public policy issues Science education Creationists promote that evolution is a theory in crisis[108] [109] with scientists criticizing evolution[110] and claim that fairness and equal time requires educating students about the alleged scientific controversy." would undermine creationists’ criticisms. Don McLeroy. It has been demonstrated by atomic clocks in very fast spacecraft.[116] [117] This viewpoint was supported by the December 2005 ruling in the Kitzmiller v." According to Science magazine. 2009. Institute for Creation Research founder Henry M.[121] This argument usually involves scientists either who were no longer alive when evolution was proposed or whose field of study didn't include evolution. the Modern evolutionary synthesis combines Darwin's Evolution with Mendel's theories of inheritance and genetics. opposed the theory of spontaneous generation with biogenesis. Morris has enumerated scientists such as astronomer and philosopher Galileo.[120] To that end. arguing that by openly debating supporters of supernatural origin explanations (creationism and intelligent design). Issues relating to religion Religion and historical scientists Creationists often argue that Christianity and literal belief in the Bible are either foundationally significant or directly responsible for scientific progress.Creation–evolution controversy 346 Freedom of speech Creationists have claimed that preventing them from teaching Creationism violates their right of Freedom of speech.[122] Many of the scientists in question did some early work on the mechanisms of evolution. which could foster an inaccurate public perception and obscure the factual merits of the debate. mathematician and philosopher Blaise Pascal. for example.g. Aronov) have upheld school districts' and universities' right to restrict teaching to a specified curriculum.[125] For example.[126] While receiving positive responses from creationist observers.. geneticist monk Gregor Mendel.[124] Many historical scientists wrote books explaining how pursuit of science was seen by them as fulfillment of spiritual duty in line with their religious beliefs. Pasteur accepted that some form of evolution had occurred and that the Earth was millions of years old. and Isaac Newton as believers in a biblical creation narrative. it was not until the mid-20th century that evolutionary theories stabilized into the modern synthesis. Shermer concluded "Unless there is a subject that is truly debatable (evolution v. Some of the historical scientists marshalled by creationists were dealing with quite different issues than any are engaged with today: Louis Pasteur. hosted by a variety of institutions. However court cases (such as Webster v. e.[123] The relationship between science and religion was not portrayed in antagonistic terms until the late-19th century. some scientists disagree with this tactic. Forums for the controversy Debates Many creationists and scientists engage in frequent public debates regarding the origin of human life. scientists are lending credibility and unwarranted publicity to creationists. whilst for creationists it is "a spiritual war". New Lenox School District and Bishop v. Even so. mathematician and theoretical physicist James Clerk Maxwell. Some extensions to this creationist argument have included the incorrect suggestions that Einstein's deism was a tacit endorsement of creationism or that Charles Darwin converted on his deathbed and recanted evolutionary theory. Michael Shermer debated creationist Kent Hovind in front of a predominantly creationist audience. with a . However. such professions of faith were not insurance against dogmatic opposition by certain religious people. The argument is generally rejected as specious by those who oppose creationism. and even then there have been many examples of the two being reconcilable for evolutionary scientists. with scientists arguing from "an impregnable fortress of evidence that converges to an unmistakable conclusion"." but rather it was "an emotional drama". in May 2004 Dr. Though biological evolution of some sort had become the primary mode of discussing speciation within science by the late-19th century. creation is not). In Shermer's online reflection while he was explaining that he won the debate with intellectual and scientific evidence he felt it was "not an intellectual exercise. an advocacy some creationists describe as a critique on chemical evolution and abiogenesis. state Citizens Alliances for Science. the National Center for Science Education. as well as a number of radio and television programmes (including the PBS series. Some of those rules are: never say anything positive about your own position because it can be attacked. But in courtrooms they are terrible. have debated Hovind. I don't think I could beat the creationists at debate. explaining: Debate is an art form. it only serves to belittle both the magisterium of science and the magisterium of religion. a non-profit organization dedicated to defending the teaching of evolution in the public schools. because in courtrooms you cannot give speeches. These include the Discovery Institute. It is not about the discovery of truth. just popping into existence one day. This has led some commentators to express a concern at what they see as a highly inaccurate and biased understanding of evolution among the general public. They are good at that. There are certain rules and procedures to debate that really have nothing to do with establishing fact — which [creationists] are very good at. the purveyors of talk-radio evolution rail. The talk-radio version had a packed town hall up in arms at the Why Evolution Is Stupid lecture. horses and humans – a scenario as unlikely as a tornado in a junkyard assembling a 747. —Stephen Jay Gould. Others. Unintelligent Designs on Darwin[130] . It is about the winning of arguments.[129] In the media The controversy has been discussed in numerous newspaper articles. lecture 1985[128] 347 Political lobbying A wide range of organisations.[126] Scott says that "Evolution is not on trial in the world of science. but chip away at what appear to be the weaknesses in your opponent's position. are involved in lobbying in an attempt to influence political decisions relating to the teaching of evolution. Evolution and Coral Ridge Ministries' Darwin's Deadly Legacy)." and "the topic of the discussion should not be the scientific legitimacy of evolution" but rather should be on the lack of evidence in creationism. In this version of the theory. reports. scientists supposedly believe that all life is accidental. claimed debates are not the sort of arena to promote science to creationists. on both sides of the controversy. yet scientists embrace his ideas because they want to promote atheism. and numerous national science associations and state Academies of Science. Stephen Jay Gould adopted a similar position. like evolutionary biologist Massimo Pigliucci. designed not to enlighten but to deceive and enrage.Creation–evolution controversy format that is fair. op-eds and letters to the editor. There is the genuine scientific theory and there is the talk-radio pretend version. the National Science Teachers Association. at a number of levels. and have expressed surprise to hear Hovind try "to convince the audience that evolutionists believe humans came from rocks" and at Hovind's assertion that biologists believe humans "evolved from bananas. In a courtroom you have to answer direct questions about the positive status of your belief."[126] (see: scientific method). —Edward Humes."[127] Eugenie Scott of the National Center for Science Education. in a forum that is balanced. Edward Humes states: There are really two theories of evolution. a random crash of molecules that magically produced flowers. The evidence against Darwin is overwhelming. I can tie them. Humans come from monkeys in this theory. there has been some growth in fundamentalist and Pentecostal Christian denominations. The war on the theory of evolution and on its proponents most often originates in forms of religious extremism which are closely allied to extreme right-wing political movements. so the case failed. on Culture.[133] [134] [135] Europe Europeans have often regarded the creation-evolution controversy as an American matter. creationism could become a threat to human rights which are a key concern of the Council of Europe.[141] One of the most acrimonious aspects of the Australian debate was featured on the science television program Quantum. Professor of Geology at Melbourne University.Creation–evolution controversy 348 Outside the United States While the controversy has been prominent in the United States.[142] . about a long-running and ultimately unsuccessful court case by Ian Plimer. and Serbia. Science and Education of the Parliamentary Assembly of the Council of Europe issued a report on the attempt by American-inspired creationists to promote creationism in European schools. the United Kingdom.[140] Public lectures have been given in rented rooms at Universities.[134] [135] [136] [137] [131] [132] On 17 September 2007 the Committee Views on human evolution in various countries. Poland. in recent years the conflict has become an issue in a variety of countries including Germany.. In 2010 the Queensland state government introduced the topic of creationism into school classes within the "ancient history" subject where its origins and nature are discussed as a significant controversy. the Netherlands. in 1980 lobbying was so successful that Queensland allowed the teaching of creationism as science to school children."[138] Australia With declining church attendance. against an ordained minister. they were not made in the course of trade or commerce. Allen Roberts. some advocates of creationism are out to replace democracy by theocracy... Although the court found that Dr Roberts had made false and misleading claims. by visiting American speakers. and speakers with doctorates purchased by mail from Florida sites. Italy.[134] However. It concludes "If we are not careful. Dr. who had claimed that there were remnants of Noah's Ark in eastern Turkey.[139] Under the former Queensland state government of Joh Bjelke-Petersen. it has flared up in other countries as well... p.. the controversy has become more prominent in Islamic countries.religion and theologically inclined philosophy have frequently been very significant factors in the forward movement of science. "This isn't just an American problem. talkorigins.[143] Currently. .[135] See also • • • • • • • • • • • • • • • • • • • • • • • • • • Abiogenesis Allegorical interpretations of Genesis Anti-intellectualism Biogenesis Clergy Letter Project Creation science Creationism Evidence of common descent Evolution and the Roman Catholic Church Evolution Sunday Evolutionary origin of religions Hindu views on evolution History of the creation-evolution controversy Intelligent design Jainism and non-creationism Jewish views on evolution Level of support for evolution List of participants in the creation-evolution controversy Mormonism and evolution Natural theology Objections to evolution Politics of creationism Project Steve Relationship between religion and science Teach the Controversy Theology of creationism and evolution Notes [144] [1] See Hovind 2006. 3–240 [6] See: • Peters & Hewlett 2005. p.Creation–evolution controversy 349 Islamic countries In recent times. pp. Mark Isaak. But in the past 5 years. doi:10. [3] Curry.5918. who organized the meeting".1159. html) . PMID 19251601. "News coverage of the creationism-versus-evolution debate tends to focus on the United States . 26. "Creationist Beliefs Persist in Europe". contrary to the usually held tale of science and religion being always opposed. See also Ruse 1999. political clashes over the issue have also occurred in countries all across Europe." [5] Numbers 1992. pp..[133] Creation science has also been heavily promoted in Turkey and in immigrant communities in Western Europe. [2] An Index to Creationist Claims (http:/ / www. Andrew (27 February 2009).323.Copyright © 2006. 247–263 Chapter titled Modern Culture Wars. org/ indexcc/ index.. primarily by Harun Yahya. 1. [4] Larson 2004. for example..1126/science. . in Egypt evolution is taught in schools but Saudi Arabia and Sudan have both banned the teaching of evolution in schools. Talkorigins Archive." says Dittmar Graf of the Technical University of Dortmund.. Science 323 (5918): 1159.. who writes "One thing that historians delighted in showing is that. and the state Constitution required all fines over $50 to be assessed by a jury. evcforum. umkc. [11] Desmond & Moore 1991. (1793-1864). htm). 21 April 2005 • The Newest Evolution of Creationism (http:/ / www. 14 [19] Numbers 1992. deemed applicable to the states. See generally Incorporation doctrine and • 350 .S. James Dwight Dana (1813-1895). • Kitzmiller vs. Edward Hitchcock. 17 [21] Numbers 1992. p. p. The efforts to enact "Butler Acts" in other jurisdictions were abandoned after the Scopes trial. com/ scopes-trial. htm). the general theory that complex life forms did not evolve but appeared 'abruptly'.000 U. earth and life scientists) who give credence to creation-science. com/ wp-dyn/ articles/ A32444-2005Mar13. law. Kitzmiller v. cfm?id=50-years-ago-scope-trial-witness& offset=6). clergyman and respected Amherst College geologist. Ruling. 14–15 [20] Numbers 1992. Morris (1818-1897). • The Political Design of Intelligent Design (http:/ / cstl-cla. See HistoryNet (http:/ / www. A View from the National Academy of Sciences. • Burns.Creation–evolution controversy • Kitzmiller v. pp. • Numbers 1992. However. . Monday. The jury informed the Court that they "passed" the question of the sentence to the Court. Presbyterian evangelist who published The Other Side of Evolution Its Effects and Fallacy [22] Numbers(2006) p161 [23] Similar legislation was passed in two other states prior to the Scopes trial. at the time of the Scopes decision in the 1920s. Evolution. 16 November 2005. p. page 20 [7] See: Battle on Teaching Evolution Sharpens (http:/ / www. but these views were rarely expressed in books and journals. aaas. 258 "Virtually no secular scientists accepted the doctrines of creation science. Darrow told the jury that they essentially had no choice but to convict Scopes on the evidence. in Oklahoma and Florida. nap. htm). Luther T. com/ article. April. & Standish 1982. Scopes' constitutional defense on establishment grounds rested solely on the state constitution. The New York Times. National Academy of Sciences and Institute of Medicine of the National Academies (2008). Dover page 83 [9] Larson 2004. theologian (1797-1878). Hastings (1833?-1899). Dover Area School District. 18. Thus. 965. Dover Area School District. 2002. H. a move which the judge accepted despite the Constitutional provision establishing a $50 limit on judge-imposed fines." See also Martz & McDaniel 1987. Exceptions are also noted. John William Dawson (1820-1899). pages 7-9. Armstrong (1813-1899). George D. edu/ catalog. p. [26] The Establishment Clause of the First Amendment of the United States Constitution was not. php?NewsID=5262). [25] In his closing argument. org/ spp/ dser/ images_Doser/ Publications/ evol_dialogue_study_guide. 12. a Newsweek article which states "By one count there are some 700 scientists (out of a total of 480. Many or most Christians may have held on to a literal six days of creation. Page A01. Jody Wilgoren. p. and Creationism (http:/ / www. Science. Alexander Patterson. • Politicized Scholars Put Evolution on the Defensive (http:/ / pewforum. pdf) study guide (pdf) [15] See: • Hodge 1874. 14. Washington Post. law. Lerner. Ethics and Religion (http:/ / www. Russell D. The jury was out for a total of nine minutes (including time for egress and ingress). noting that this applies to published or public skeptics. org/ news/ display. NSTA 2007. Enoch Fitch Burr (1818-1907). Renka. Barbara Forrest. Townsend (1838-1922. p. html). L. pp. and he did not hold that against them. Ralph. Peter Slevin. See Court's opinion (http:/ / www. National Academy of Sciences. the observation that evolutionary critics had a relaxed interpretation of Genesis is supported by specifically enumerating: Louis Agassiz (1807-1873). AAAS 2006. p. scientificamerican. 503–505. but that did not deter creation scientists from advancing scientific arguments for their position. Reverend Herbert W. 321–323. edu/ Renka/ Renka_papers/ intell_design. historynet. p. umkc. edu/ faculty/ projects/ ftrials/ scopes/ statcase. pp. 23. such as literal interpretations published by Eleazar Lord (1788-1871) and David Nevins Lord (1792-1880). Arnold Henry Guyot (1807-1884). page 80. php?record_id=11876). semo. Natural History. [24] The statute required a minimum fine of $100. and Pinholster 2006. 14 March 2005. ISBN 0-309-10586-2. also pages 64-90 [8] Myers 2006. • Huxley 1902 [16] Witham 2002 [17] Barbour 1997 [18] Numbers 1992. net/ RefLib/ NaturalHistory_200204_Forrest." [10] Committee on Revising Science and Creationism. [13] van Wyhe 2006 [14] AAAS Evolution Dialogues: Science. IAP 2006. Court's opinion (http:/ / www. 34–35 [12] See": • van Wyhe 2006. 177. html). Charles Hodge. edu/ faculty/ projects/ ftrials/ scopes/ statcase. htm) (describing the Florida and Oklahoma acts and Scientific American Article (http:/ / www. p. washingtonpost. • Desmond & Moore 1991. Hodge 1874. Encyclopedia of religious freedom. pp. [28] The Court stated in its opinion that "England and Scotland maintained State churches as did some of the Colonies.. 255–256: "Fundamentalists no longer merely denounced Darwinism as false. php#questionsAboutIntelligentDesign). pl?navby=CASE& court=US& vol=393& page=97) (U. Dorothy (2000). Arkansas. Retrieved 2007-05-13. pfaw. 252 [37] Larson 2004. pp. org/ pfaw/ general/ default. this was before most of the Bill of Rights had been incorporated and applied to the states. [32] Epperson et al. Evolving Banners at the Discovery Institute. p.Creation–evolution controversy Everson v. Trial and Error: The American Controversy Over Creation and Evolution. Board of Education (seminal U. [30] Timeline and Myths of Scopes Trial (http:/ / www. html) by Constitutional Rights Foundation. Edward J. p. [33] Larson. p. [34] Larson 2004. 5--Peters and Hewlett argue that the atheism of many evolutionary supporters must be removed from the debate [57] Lenski 2000. 7 July 2006. they offered a scientific-sounding alternative of their own. . com/ news/ 2006/ jul/ 07/ many_question_groups_move_elections_nearing/ ) 6News Lawrence. (http:/ / www. p. Conclusion of Ruling. (2003).S. .000 U. php) The Associated Press. • TalkOrigins Archive: Post of the Month: March 2006 (http:/ / www. Wiker 2003.200–208 [38] Larson 2004.(explaining incorporation doctrine relative to First Amendment). Peters & Hewlett 2005. [56] Johnson 1998. com/ articles/ ap/ 2007/ 02/ 13/ america/ NA-GEN-US-Kansas-Evolution-History. pp. earth and life scientists) who give credence to creation-science. biolsocwash. [45] "Top Questions-1." [42] Larson 2004. ISBN 0415941814. pp. [54] The "Evolution" of Creationism (http:/ / www. msnbc. 23. 276. Lawrence Journal-World. May. a Newsweek article which states "By one count there are some 700 scientists (out of a total of 480. 393 U. [44] "NCSE Resource" (http:/ / ncse. pdf) (pdf) A Position Paper from the Center for Inquiry. 3 [51] Some question group's move with elections nearing (http:/ / www. 248. 177. pp. 5–6 [43] Ruling. Cookson. 242. 132 et seq." See also Martz & McDaniel 1987. 251 [36] Larson 2004. 6newslawrence. p. msn. p. 31. aspx?oid=20418& print=yes& units=all) Timeline: how creationism has "evolved". See also Book Review (http:/ / volokh. pp. discovery. xi. pp. org/ csc/ topQuestions. The Creation Controversy: Science or Scripture in Schools. 97 (http:/ / caselaw. New York: iUniverse. [48] Larson 2004. archive. 2002-08-29. pp. p. Supreme Court opinion finally applying the Establishment Clause against states in 1947). com/ scripts/ getcase. Supreme Court 1968-11-12).250. [50] Bumiller 2005. Numbers 1998. People for the American Way. Kitzmiller v. html). Peters & Hewlett 2005. p. see also Dobzhansky 1973 [35] Larson 2004. Conclusions [58] Johnson 1998 • • 351 . ISBN 0195154703.. p. [46] Verderame 2007. The History of Creationism by Lenny Flank. and it was intended by this clause of the Constitution [the Religious Preference Clause] to prevent any such undertaking in Tennessee. [53] Evolution of Kansas science standards continues as Darwin's theories regain prominence (http:/ / www. org/ web/ 20070926214521/ http:/ / www. via the International Herald Tribune." [29] See: s:Kitzmiller v. Incorporation Doctrine Explained (http:/ / www. org/ topics/ law/ scopes/ scopes. net/ uploads/ attachments/ Forrest_Paper. ISBN 0-595-00194-7. shtml) of Professor Larson's Pulitzer Prize winning Summer for the Gods at point 4 ("The constitutional case was largely based on state constitutional law.What is the theory of intelligent design?" (http:/ / www. 258 "Virtually no secular scientists accepted the doctrines of creation science. com/ id/ 14137751/ ) MSNBC.S. v. talkorigins.Simon 2006 [47] Dewey 1994. p. [55] Kitzmiller v. Discovery Institute. Dover Area School District. summarizing Gould. org/ origins/ postmonth/ mar06. org/ id_statement. 2005). antievolution. Understanding the Intelligent Design Creationist Movement: Its True Nature and Goals. Retrieved 2009-04-07. 284–285 [40] Numbers 1992. findlaw. but that did not deter creation scientists from advancing scientific arguments for their position. which they called either 'scientific creationism (as distinct from religious creationism) or 'creation science' (as opposed to evolution science. 254–255. Dover Area School District/2:Context#Page 19 of 139. lp. 04 cv 2688 (December 20. html) at the Wayback Machine (archived September 26. Connecticut (1940 Supreme Court case stating that the Free Exercise Clause of the First Amendment is incorporated into the Fourteenth Amendment and is therefore applicable against the states)." [49] Statement from the Council of the Biological Society of Washington (http:/ / web. 13 February 2007. 2007).. 284–6 [41] Quoting Larson 2004. org/ bill-of-rights-in-action/ bria-7-4-b. Office of Public Policy Barbara Forrest. 2007. [27] The Court accordingly did not address the question of whether the teaching of Creationism in the public schools was unconstitutional.S. 255. com/ creationism/ general/ evolving-banners-at-discovery-institute). and Wiker 2003. 2 August 2006. com/ posts/ 1122414938. Oxford University Press. p.Numbers 1992. iht.S. [52] Evolution’s foes lose ground in Kansas (http:/ / www. 255 [39] Numbers 1992. p. Cantwell v."). crf-usa. html) [31] Nelkin. Taylor & Francis. the general theory that complex life forms did not evolve but appeared 'abruptly'. pp. Dover Area School District pp 7-9. centerforinquiry. Catharine (2003). 7 May 2008 [82] The Discontinuity of Life (http:/ / www. 1999. but to help in restoring the foundation of the gospel in our society. John S. Kurt Wise. asp) (Part II of Relevance of Creation). if true. [75] As quoted by Wallis 2005. provide abundant reason why the reader should reject evolution and accept special creation as his basic world-view. The Chinese Intelligentsia who had been schooled by Orthodox Evangelical Missionaries were thus softened for the advent of Marxism. html). fallacyfiles. org/ faqs/ evolphil/ falsify. as well as animalistic attitudes and behaviour by individuals. A new wave of so-called missionaries from mainline Protestant denominations came teaching evolution and a non-supernatural view of the Bible. US District Court for the Northern District of Georgia (2005). pdf). vi-viii. pbs. It is also worth noting the comment in the book. talkorigins. com/ journals/ science. Fallacy Files [80] Johnson 1993. pp. informationphilosopher. org/ FilesPDFs/ cobb county decision. 16.. 247 Lewin 1982 Numbers(2006) p274 Kofahl 1981. answersingenesis.. 13–37. Selman v. 6 Dawkins quoting Haldane [77] Dorman 1996 [78] "One thing we have come to realise in Creation Science is that the Lord has not just called us to knock down evolution. Discovery Institute [84] Evolution versus the People (http:/ / www.I want to list seventeen summary statements which. but we would also see the seeds of revival sown in a culture which is becoming increasingly more pagan each day. [81] Evolution: what's the real controversy? (http:/ / arstechnica. lehman. 247.. Skeptical Inquirer. TalkOrigins Archive Popper 1976. [74] Ruse 1999. com/ solutions/ philosophers/ popper/ natural_selection_and_the_emergence_of_mind. Congregationalist. November 1983. Center for Science and Culture. 69 where Johnson cites three pages spent in Issac Asimov's New Guide to Science that take creationists to task. p. p. 247 Kofahl 1989 as quoted by Numbers 1992. pp. html). org/ indexcc/ CA/ CA211_1.. Moran 1993 . Talk. p. Tolson 2005. aclu. and Christians need to be awakened to that fact!" Ham. and What Does 'Science' Mean? (http:/ / www. p. Casey Luskin. 2004. Belief in special creation has a salutary influence on mankind. Belief in evolution and animal kinship leads normally to selfishness. p. Methodist. Creation Magazine '6'(2):17. Cited in Appeal to Consequences (http:/ / www. Thomas Samuel Kuhn. since it encourages responsible obedience to the Creator and considerate recognition of those who were created by Him. p. p.org/adconseq. .. which discusses conflicting ideas about science among Karl Popper. pdf). The Remarkable Birth of Planet Earth (Creation-Life Publishers. on page 49 and 50: ‘New philosophies and theologies from the West also helped to erode Chinese confidence in Christianity. "Did Popper refute evolution?" (http:/ / www. Bill Moyers et al. and their disciples. "Natural selection and the emergence of mind" (http:/ / www. Interview with Richard Dawkins [63] Merriam-Webster online dictionary. org/ Outreach_files/ 2004-Did Popper refute evolution?-Skeptical Inquirer. Ken. edu/ execsumm_pdf/ 6024. 32. Answers in Genesis [83] A Primer on the Tree of Life (http:/ / www. Accessed 2006-01-29.m-w. 13. [79] ". www. 1972). org/ articles/ am/ v4/ n1/ discontinuity). 873 Talkorigins summary of Karl Popper attitudes towards evolution (http:/ / www. Morris.. . Creation Evangelism (http:/ / www.. [62] Johnson 1993. and fighting between groups. ‘By Their Blood-Christian Martyrs of the 20th Century’ (Most Media) by James and Marti Helfi. org/ article/ evolution-versus-people/ ). Also see Dawkins 1986 and Dawkins 1995 [76] Wallis 2005. aggressiveness. Origins. Steering Committee on Science and Creationism." PBS. nap. icr. Institute for Creation Research . Science and Creationism: A View from the National Academy of Sciences. Second Edition. Destructive books brought by such teachers further undermined orthodox Christianity. edu/ deanhum/ philosophy/ platofootnote/ PlatoFootnote. org/ now/ transcript/ transcript349_full.com [64] Gould 1981 [65] See: • • [66] [67] [68] [69] [70] [71] [72] Number 1992. pp. We believe that if the churches took up the tool of Creation Evangelism in society. Nobel Intent. Dialectica 32(3/4): 339–355.Creation–evolution controversy [59] Einstein 1930. org/ a/ 10651). discovery. . and Northern Baptist schools were especially hard hit. p. . John Timmer. Massimo Pigliucci (Sept-October 2004). not only would we see a stemming of the tide of humanistic philosophy. Presbyterian. html#dawkins). Morris. Cobb County School District (http:/ / www. Wilkins. p. answersingenesis.html). pdf).. org/ creation/ v6/ i2/ creationII. Henry M. Evolution and Philosophy: Is Evolution Science. National Academy of Sciences.’ Evolution is destroying the Church and society. 1–4 [60] Dawkins 1997 [61] Free Executive Summary (http:/ / www. ars/ 2008/ 05/ 07/ evolution-whats-the-real-controversy). Bertrand Russell came from England preaching atheism and socialism. " Now with Bill Moyers (http:/ / www. talkorigins. html) 352 [73] Karl Popper (1978). 63. 168 and 172 quoted in Kofahl 1981 Unknown sociologist quoted in Numbers 1992. while only spending one half page on evidence of evolution. . ISBN 978-0-309-06406-4.." Henry M.. Jim Foley. p20-21 [105] Dobzhansky 1973 [106] Pieret 2006 Isaak. shocked to find that quote mining is going on in there!" (http:/ / pharyngula. The complete world of human evolution. University of California Press (2007).Creation–evolution controversy [85] Stringer. . "Index to Creationist Claims: Claim CA113" (http:/ / www. Meert. samarium-neodymium dating. Kitzmiller v. Reflections of our past: how human history is revealed in our genes. aaas. talkorigins. Judith Hauck. shtml). org/ indexcc/ CA/ CA113. [104] Radiometric Dating. answersingenesis. asp) [97] The Meaning of the Chronogenealogies of Genesis 5 and 11 (http:/ / www. American Scientific Affiliation. [100] Nuclear Decay: Evidence For A Young World (http:/ / www. Mark Isaak. Alan (1985) Creation and Evolution. pp. pdf) American Association for the Advancement of Science.1051/0004-6361:20047060. Andrews. . org/ news/ releases/ 2006/ pdf/ 0219boardstatement. London: Thames & Hudson. lutetium-hafnium. De Mello. More than 70.I. Quoted in Hooking Leviathan by Its Past (http:/ / www. October 2002. 8 August 2005. Chris. talkorigins. Mike (2007-07-02). php?module=articles& action=view& ID=302). John (2003). archive. 13 November 2000 (updated 6 February 2003). pdf). politics. org/ news/ releases/ 2002/ 1106id2. html). A&A 434: 301–308. org/ programs/ atc/ features/ 2002/ july/ toumai/ index. 19 May 2003 [89] Was Java Man a gibbon? (http:/ / www. talkorigins. edu. pp. Henke. . TalkOrigins website. Arany-prado. Colo: Westview Press.Triangle Books. TalkOrigins website. Mark (2004). rubidium-strontium dating. Dover Area School District. and not in science classes. p298 [96] Biblical chronogenealogies (http:/ / www. [103] Dating methods discussed were potassium-argon dating. com/ article/ dn3744-chimps-are-human-gene-study-implies. html). Interacademy Panel on Global Issues. [99] Del Peloso. org/ faqs/ homs/ compare. org/ tj/ v17/ i3/ chronogenealogies. html) New Scientist.. Retrieved 2007-12-27. Hoagland. File/ Master/ 6/ 150/ Evolution statement. Roger C. org/ cgi/ content/ full/ 354/ 21/ 2277) George J.P. ISBN 0-8133-3958-8. Wiens." But there is no significant controversy within the scientific community about the validity of the theory of evolution. London. asa3. Retrieved 2007-12-27. The Discovery Institute's Dissent From Darwin Petition (http:/ / www. L. [86] Relethford.. org/ archives/ quote_mines/ ) on 2007-12-09. Original version: 17 March 2005. unsw. Peter (2005). L. pp. G. A Christian Perspective (http:/ / www. website. [87] "Toumaï the Human Ancestor: Skull of Oldest Known Hominid Unearthed in Chad" (http:/ / www. pp.E: More Faulty Creation Science from The Institute for Creation Research (http:/ / gondwanaresearch. • 353 . doi:10. E. html). stephenjaygould. 30 April 2003.F. argon-argon dating. Jim Foley. org/ faqs/ homs/ gibbon. The current controversy surrounding the teaching of evolution is not a scientific one. org/ index. and uranium-lead dating.000 Australian scientists and educators condemn teaching of intelligent design in school science classes (http:/ / www. Bert Dodson. has 120. org/ origins/ 07053. au/ news/ 2005/ intelligent. Mahlon B. Talk. J. org/ library/ gould_leviathan. talkorigins. ISBN 0-500-05132-1. Volume 354:2277-25 May 2281 2006 [111] See: 1) List of scientific societies rejecting intelligent design 2) Kitzmiller v.T. org/ faqs/ helium/ zircons. net/ Object. Others insist that teachers have absolute freedom within their classrooms and cannot be disciplined for teaching non-scientific "alternatives" to evolution. 1985. Retrieved 2007-12-26. "The age of the Galactic thin disk from Th/Eu nucleocosmochronology". Statements from Scientific Organizations (http:/ / ncse. htm) [98] IAP Statement on the Teaching of Evolution (http:/ / www. [88] Chimps are human. Stephen Jay Gould [95] Exploring the Way Life Works. 78-90 [93] Life—How Did It Get Here?. Boulder. [92] Scientific Creationism. Kevin R. htm). 57-59 [94] Haywood. NPR: All Things Considered. nejm. The Official Journal of the International Association for Gondwana. [101] Young-Earth Creationist Helium Diffusion "Dates" Fallacies Based on Bad Assumptions and Questionable Data (http:/ / www.. The Panda's Thumb (blog). page 89 [110] "That this controversy is one largely manufactured by the proponents of creationism and intelligent design may not matter. 21 June 2006. Morris. html). org/ archives/ quote_mines/ ). newscientist. PZ (2004-09-11). org/ index/ weblog/ comments/ im_shocked_shocked_to_find_that_quote_mining_is_going_on_in_there). The AAAS. icr. neither scientists nor citizens should be concerned. Gondwana Research.F. npr. ISBN 978-0520249264 [91] Comparison of all skulls (http:/ / www. [102] R. Russell Humphreys. dissentfromdarwin. and firmly rejects intelligent design and denies that there is a legitimate scientific controversy (http:/ / www. org/ ASA/ resources/ Wiens. A number of bills require that students be taught to "critically analyze" evolution or to understand "the controversy. Revision: 24 November 2005. science." Intelligent Judging — Evolution in the Classroom and the Courtroom (http:/ / content. Retrieved 2009-02-21. html). aaas. TalkOrigins website. [108] "Some bills seek to discredit evolution by emphasizing so-called "flaws" in the theory of evolution or "disagreements" within the scientific community. "I'm shocked." AAAS Statement on the Teaching of Evolution (http:/ / www. [90] See disputes over the classification of Neanderthals in The Counter-Creationism Handbook. Pharyngula.000 members. org/ ) has been signed by about 500 scientists. html). Impact. • Dunford. rhenium-osmium dating. html). pandasthumb. and as long as the controversy is taught in classes on current affairs. or religion. interacademies. D. Dover page 83. 1985. New England Journal of Medicine. Henry M. org/ web/ 20071209204412/ http:/ / www. 16 February 2006 (PDF file) [109] Ruling. Da Silva. Number 352. html#page 20). gene study implies (http:/ / www.A. Annas. Archived from the original (http:/ / www.S. (2005). G. com/ media/ voices/ science) on the status intelligent design and other forms of creationism. . • Myers. grisda. pandasthumb. com/ rate. 257. 240..origins. "A new (mis)take on an old paper (and other posts)" (http:/ / web. the largest association of scientists in the U. Watchtower Bible and Tract Society. Richard Dawkins. com/ media/ voices/ science). com/ eskeptic/ 04-05-10. html) [114] Teaching Intelligent Design: What Happened When? (http:/ / www. . Monday. [133] Pitock. xml). Owl Books. 11 July 2006 [116] "has the effect of implicitly bolstering alternative religious theories of origin by suggesting that evolution is a problematic theory even in the field of science. in violation of the Establishment Clause. html) edited by Mark Isaak. Michael.. The American Prospect. com/ content/ 2008/ feb/ 16/ na-clash-over-creationism-is-evolving-in-europes-s). [134] Gregory Katz (2008-02-16). "Why doesn't America believe in evolution?" (http:/ / www. "Evolution myths: It doesn't matter if people don't grasp evolution" (http:/ / www. [113] "ID's home base is the Center for Science and Culture at Seattle's conservative Discovery Institute. Scientism./ [135] Taner Edis. Todd (06 2007). pg. au/ religion/ stories/ s820631. htm) Australian Broadcasting Corporation [140] Carly Hennessy (May 30.Claim CA114. Meyer directs the center. "Then a Miracle Occurs: An Obstreperous Evening with the Insouciant Kent Hovind. [129] Statements from Scientific Organizations (http:/ / ncse. co. 15 May 2006. jhtml?view=DETAILS& grid=& xml=/ earth/ 2007/ 10/ 02/ scihist102. 2005 [123] Index to Creationist Claims . com/ x/ pittsburghtrib/ opinion/ columnists/ guests/ s_493631. Dembski"The clarion call of the intelligent design movement is to "teach the controversy. causes students to doubt its validity without scientific justification. [128] Shermer. Michael (2004-05-10). org/ indexcc/ CA/ CA114. co. . p. Ethics.119. html)."Kitzmiller v.Claim CA114 (http:/ / talkorigins. which would require students to learn how scientists "continue to investigate and critically analyze" aspects of Darwin's theory. Retrieved 2007-02-11. com/ rncse/ 19/ 6/ cloning-creationism-turkey). Dover Area School District. au/ news/ national/ creationism-to-be-taught-in-queensland-classrooms/ story-e6frf7l6-1225873019548). pp. org/ sci_rel. 67–114. 20 April 2006. presents students with a religious alternative masquerading as a scientific theory. Pittsburgh Tribune-Review [131] Michael Le Page (19 April 2008). newscientist. Retrieved 2008-02-17. 354 . heraldsun. Retrieved 2008-02-17. London: The Telegraph. newscientist. telegraph. uk/ 1/ hi/ world/ europe/ 3642460. htm) by William A. former Reagan adviser Bruce Chapman heads the larger institute. [137] Roger Highfield (2007-02-10). RNCSE 19 (6): 30-35. the official Richard Dawkins website. 2004-09-09. stm)." [115] Nick Matzke's analysis shows how teaching the controversy using the Critical Analysis of Evolution model lesson plan is a means of teaching all the intelligent design arguments without using the intelligent design label. the ID crowd has pushed a "teach the controversy" approach to evolution that closely influenced the Ohio State Board of Education's recently proposed science standards. 000-evolution-myths-it-doesnt-matter-if-people-dont-grasp-evolution. 2002): ISBN 0878936599 page 102." Ruling . (Sinauer. Retrieved 2008-02-17. . html) Nick Matzke. Denying Evolution: Creationism. pandasthumb. pittsburghlive. 'Why People Believe Weird Things'. From this perch. and at worst a canard. [132] Jeff Hecht (19 August 2006). richarddawkins. 49 Kitzmiller v.Creation–evolution controversy [112] "In summary. misrepresents its status in the scientific community. p. Reason: In the News. but not ID itself. NCSE [130] Unintelligent designs on Darwin (http:/ / www.Why-I-Wont-Debate-Creationists. religioustolerance. bbc. [127] Massimo Pigliucci. No one here but us Critical Analysis-ists. Associate Professor of Philosophy from George Mason University. Dover Area School District. directs them to consult a creationist text as though it were a science resource. Paperback ed. and Religion (http:/ / www. and the Nature of Science.. .disclaimer. retrieved 2007-04-29 [119] Science. "Clash Over Creationism Is Evolving In Europe's Schools" (http:/ / www2. net/ article. 2010). aaas. org/ spp/ dser/ 02_Events/ Lectures/ 2006/ 02_Lecture_2006_0420. 153. National Center for Science Education. 2002. Young Earth Creationist and Defender of the Faith" (http:/ / www. This tactic is at best disingenuous. tbo. "Creationists rewrite natural history" (http:/ / www. skeptic. 2005 [124] (http:/ / www. 72 [139] Christianity – Pentecostalism (http:/ / www. org/ indexcc/ CA/ CA114_22. . Dover Area School District." There is a very real controversy centering on how properly to account for biological complexity (cf.net. Conclusion. with input from the Christian supply-sider and former American Spectator owner George Gilder (also a Discovery senior fellow).. and instructs students to forgo scientific inquiry in the public school classroom and instead to seek out religious instruction elsewhere. ." . com. and it is a scientific controversy. org/ print/ V13/ 22/ mooney-c. The Panda's Thumb. shtml).Richard-Dawkins). New Scientist 198 (2652): 31. html). com/ article/ mg19826523. html#miracle). "Creationism to be taught in Queensland classrooms" (http:/ / www. Emmett Holman. Kitzmiller v. 2010. The effect of Defendants’ actions in adopting the curriculum change was to impose a religious view of biological origins into the biology course. org/ docs/ dembski/ wd_teachingid0201. . "Cloning Creationism in Turkey" (http:/ / ncse. page 89 [118] AAAS Dialogue on Science. BBC. Edward Humes. html). 3 APRIL 2009 [120] Woods 2005.22 (http:/ / talkorigins. eSkeptic Online. net. abc. . htm) Science and religion: Conflicts & occasional agreements [125] Why I Won't Debate Creationists (http:/ / richarddawkins. arn. [138] New Scientist 10 November 2007. uk/ earth/ main. org/ archives/ 2006/ 07/ no_one_here_but. Herald Sun. the disclaimer singles out the theory of evolution for special treatment. Chapter Five: The Church and Science [121] Morris 1982 [122] Index to Creationist Claims . prospect. New Scientist 191 (2565): 11." Chris Mooney. (http:/ / www. [126] Shermer. The goal of the IDM is not to encourage critical thought. the ongoing events in Kansas). page 134 [117] "ID's backers have sought to avoid the scientific scrutiny which we have now determined that it cannot withstand by advocating that the controversy. 2 December 2002 Survival of the Slickest: How anti-evolutionists are mutating their message (http:/ / www. should be taught in science class.. "Science and Islam". Retrieved July 22. but to foment a revolution which would supplant evolutionary theory with ID. com/ article/ dn9786-why-doesnt-america-believe-in-evolution. html) edited by Mark Isaak. Retrieved 2008-02-17. [136] "Serbia reverses Darwin suspension" (http:/ / news. Discover: 36–45. Associated Press. whether ID is science. cfm?story_id=9036706). 2007-04-19. Kent (2006). The Economist.org/dobzhansky.File/Master/6/150/Evolution statement.net/Object.org/humanist/articles/ dawkins.).2think. "Is Science a Religion?" (http://www. abc. "An Episcopal Trilogy 1887" (http://www. ISBN 978-1417973729. ISBN 0-87975-853-8 • Dobzhansky. Moore. Inc. Interacademy Panel (2006-06-21).)..php?spec=67).talkorigins.pdf). interacademies. retrieved 2007-01-14 • IAP. 1996-01-30. com/2005/08/03/politics/03bush.org/files/19192/19192-8. W. Quantum. in Martin Gardner. (1902). common in academic literature. retrieved . Religion and Science: Historical and Contemporary Issues (first revised ed.. Overton (http://www. Evolution as Fact and Theory (http://www. Ian G. Darwin. Robert E.Reason versus Creationism"." 355 Footnotes References • AAAS. Adrian. Hovind's $250. Ralph. Stephen Jay (1981).gutenberg.One Man's Crusade (http:/ / www. Retrieved 2007-04-25. New York Times Magazine: 1–4. earth. Dr. The Blind Watchmaker. and the universe. and Company. Richard (1995). Great Essays in Science. IAP Statement on the Teaching of Evolution (http://www. Theodosius (1973). London: Michael Joseph. "The Influence of Darwinism on Philosophy". retrieved 2007-01-17 • Hodge. retrieved 2007-01-17 • Huxley. Elisabeth (2005-08-03).W. W. htm). interacademies. this article uses the term "creation myth" in the formal sense. Penguin Group. economist. Armstrong.thehumanist. Statement on the Teaching of Evolution (http://www. Charles (1874).org/faqs/mclean-v-arkansas. Basic Books. au/ quantum/ info/ 97lies. Ian "Telling lies for God. John (1994). aaas. meaning "a religious or supernatural story or explanation that describes the beginnings of humanity. Edward M. What is Darwinism? (http://www.000 Offer (http://www. "Bush Remarks Roil Debate on Teaching of Evolution" (http://www. The Unofficial Stephen Jay Gould Archive. [143] "Evolution and religion: In the beginning" (http:/ / www. Thomas H. retrieved 2007-01-14 • Hovind.drdino. [144] While the term myth is often used colloquially to refer to "a false story". Creation Science Evangelism. Prometheus Books. District Court Judge William R. ISBN 0-465-06990-8 • Dawkins.shtml). . 65. James (1991). W. accessed 2008-02-05. "Religion and Science" (http://www. HarperSanFrancisco.txt).stephenjaygould. River Out of Eden.org/news/releases/2006/pdf/0219boardstatement. Collected Essays Science and Christian Tradition (Kessinger Publishing) V: 126–159. Norton & Company.This article gives a worldwide overview of recent developments on the subject of the controversy. htm). com/ world/ displaystory. Humanist. txt). life. ISBN 0060609389 • Burns.nytimes. Albert (1930-11-09). Lerner.pdf). retrieved 2007-01-30 • Desmond.com/aor/einstein/einsci. Standish. Scribner.. retrieved 2007-01-14 • Bumiller.Creation–evolution controversy [141] Plimer. retrieved 2007-01-30 • Gould. See transcript link for detail. Arkansas Board of Education Decision by U. TalkOrigins Archive Foundation.html). Norton & Company.S. Philip Lee. pp. ISBN 0-7181-3430-3 • Dewey.html?ex=1280721600&en=8bbf73d2f5204260&ei=5088&partner=r).com/articles. 58.org. "Nothing in Biology Makes Sense Except in the Light of Evolution" (http:// www. (Random House) [142] "Telling Lies for God"? . retrieved 2007-01-14 • Transcription of McLean v.gutenberg. American Biology Teacher (National Association of Biology Teachers) 35: 125–129. American Association for the Advancement of Science (2006-02-16) (pdf). net.html).aaas. retrieved 2007-02-03 • Barbour.org/files/15905/15905-8.sacred-texts. Richard (January 1997). The New York Times (2005-08-03). (1997). ISBN 0-393-31570-3 • Dawkins. Richard (1986). World Civilizations Their History and Their Culture (Sixth ed.html). ISBN 0-393-95077-8 • Dawkins. Meacham (1982). retrieved 2007-01-31 • Einstein.net. ASIN B0006AEEMO.org/library/ gould_fact-and-theory. The Evolution Controversy: Who's Fighting with Whom about What? (http://web. Darwinism Comes to America. Stephanie (2006). 224.org/programs/darwin/transcript. "Their Own Version of a Big Bang: Those who believe in creationism – children and adults – are being taught to challenge evolution's tenets in an in-your-face way. Damned Lies and Quote Mines (http://www. Ted. pp. Richard E. "Keeping God out of the Classroom (Washington and bureau reports)". pp.Creation–evolution controversy 2007-01-14 Johnson. "Bible-Believing Scientists of the Past" (http://www. (1998-11-15).com/pharyngula/ 2006/06/ann_coulter_no_evidence_for_ev.shtml). retrieved 2007-09-12 • NSTA. html). (June. archived from the original (http://www. retrieved 2007-01-23 Martz. James (2006). National Science Teachers Association (2007).pdf) on 2006-12-11. P. ISBN 0674023390. New Scientist 87 • Ruse. Science (New Series) 212 (4497) Kofahl. Mystery of Mysteries: Is Evolution a Social Construction.edu/docs/EvolutionBrief2. (2000). Expanded Edition.nsta. "An NSTA Evolution Q&A" (http://www. Law & Education. ISBN 0674193121 • Peters. Inc. Ronald L.. ISBN 0-679-64288-9. Ronald L. ISBN 0-679-40104-0.aaas. InterVarsity Press. (1998). retrieved 2007-01-24 356 • • • • • • • • • Moran. php?module=articles&action=view&ID=185).org/news/releases/2006/0219boardstatement.org/crsq/abstracts/sum26_1.html). Pharyngula (ScienceBlogs). Retrieved 2008-02-01. Lenski.php). Ronald (2006-11-30). TalkOrigins Archive Foundation.actionbioscience.org/faqs/cre-error.org/web/ .plts. "Letter to the Editor". CRS Quarterly (The Creation Research Society) 26 (1). (1992). ISBN 0-8308-1929-0 Johnson. Pacific Lutheran Theological Seminary.) CIX (26): 23–24. The Quote Mine Project Or.archive. retrieved 2007-01-23 • Pinholster. Reason in the Balance: The Case Against Naturalism in Science.org/web/20061211212344/http://www. The Creationists: From Scientific Creationism to Intelligent Design. org/faqs/quotes/mine/project. • Numbers.icr. publicradio. "Popper on Darwinism".edu/docs/ EvolutionBrief2. Edward J. Robert E. Larry. (2004). "The Hierarchy of Conceptual Levels For Scientific Thought And Research" (http://creationresearch. John (2006). 1989).pdf). Hewlett. Harvard University Press." (http://web. retrieved 2007-01-14 • Popper. Harvard University Press.plts. (1993). "Ann Coulter: No Evidence for Evolution?" (http://scienceblogs.org/index. ISBN 0028-9604 Moore. Lies. "Evolution and Wonder — Understanding Charles Darwin" (http://speakingoffaith. IntervaVsity Press. Impact (Institute for Creation Research) 103. Evolution: Fact and Theory (http://www. ISBN 0-674-46706-X • Simon. Harvard University Press. Michael (1999-04-30). retrieved 2007-01-23 • Morris. pp. Unended Quest: An Intellectual Autobiography. Newsweek (Newsweek Inc.html).aspx).Z.org.. ISBN 0-87548-366-6 • Popper.org. Evolution. Evolution is a Fact and a Theory (http://www. (2006-06-18).shtml). 320. Open Court Publishing Co. Henry M.talkorigins. American Public Media. retrieved 2007-01-29 Larson. McDaniel.html). Martinez (2005-12-22) (pdf). Creationists: The Evolution of Scientific Creationism. • Numbers. • Numbers. Knopf. 224. retrieved 2007-01-28 • Pieret. Speaking of Faith (Radio Program). Phillip E. Karl (1980). Ann (1987-06-29).org/ publications/evolution. Darwin on Trial (2nd ed. AAAS Denounces Anti-Evolution Laws as Hundreds of K-12 Teachers Convene for 'Front Line' Event (http://www. 624 pages. Karl (1976). ISBN 0-8308-1324-1 Kofahl.org/evolution/lenski. Ginger (2006-02-19). Alfred A. p.archive. (1982-01-01). TalkOrigins Archive Foundation.talkorigins. Phillip E.). aaas. (22 May 1981). Laurence (1993). retrieved 2007-01-20 • Myers. Robert E. Modern Library. AcitionBioscience. . retrieved 2007-01-21 357 • • • • • • Witham. ISBN 1-56338-349-7. Evolution. retrieved 2007-01-24 Verderame. U. Inc. and the Church (CRISIS Magazine). How the Catholic Church Built Western Civilization. • Gould. 144. ISBN 0231054750. para. and the Drama of Life. (1994). Cambridge. Time Magazine Retrieved on 2007-01-31 Wiker. "From Broadway to Biophilia". (1993).9171.edu/article. Jay (2005-09-05).html). Princeton. Old And New: Or The Theories Of Buffon. (2001).pl?sid=06/02/ 12/1727208&mode=thread) on 2006-09-04.edu/article. ISBN 0801058627. Charles (1996).genom. Erasmus Darwin And Lamarck. Mass: Belknap Press. Grand Rapids. James H. RM: 1994. Where Darwin Meets the Bible: Creationists and Evolutionists in America (kindle ed.htm). Claudia (2005-08-07). Louisville: Westminster John Knox Press. Ernst (1985). retrieved 2007-02-03 Tolson. "Does Science Point to God? Part II: The Christian Critics" (http:// www. Mark A. Genetics and the origin of species. Ronald L. doi:10. sect.com/usnews/culture/articles/050928/ 28religion. • Woods.. ISBN 0345409469. • Haught. ISBN 0-89526-038-7 Published books and other resources • Burian.usnews. • Kutschera U. "The Evolution Wars" (http://www.tamucc.). New York: Ballantine Books.1090909-1. answersingenesis. ISBN 0674364465.org.org. • Henig. Jr. Carl (1997). • Numbers. Crisis Politics. As Compared With That Of Charles Darwin (1911). New York: Columbia University Press. • Sagan. (2005). Kessinger Publishing. Chapter 2. An evolving dialogue: theological and scientific perspectives on evolution.crisismagazine. God. The demon-haunted world: science as a candle in the dark. Dobzhansky. and Inheritance. in Adams. Religion in America (http://www. retrieved 2007-02-07 Wallis. MI: Baker Pub Group. Regnery Publishing. "From Broadway to Biophilia". • Pennock RT (2003). Larry A. Benjamin D.pl?sid=06/02/12/1727208&mode=thread). 9. The Evolution of Theodosius Dobzhansky: essays on his life and thought in Russia and America. The monk in the garden: the lost and found genius of Gregor Mendel. Charles Darwin: gentleman naturalist: A biographical sketch (http://darwin-online. uk/darwin. Culture. The Growth of Biological Thought: Diversity. Oxford University Press.com/time/magazine/article/ 0. ISBN 9780664232856. "The modern theory of biological evolution: an expanded synthesis". • Mayr.: Princeton University Press. Los Angeles Times (2006-02-11). archived from the original (http://philosophy.com/julaug2003/feature1. Niklas KJ (2004). ISBN 019283438X. • Beer. Oxford [Oxfordshire]: Oxford University Press. ISBN 0548641323. doi:10. ISBN 0-691-03479-6. The Twilight of Evolution. LLC.1007/s00114-004-0515-y. the father of genetics. The creationists.00. ISBN 978-0195150452. ISBN 0520083938. News & World Report. PMID 15241603. ISBN 0395977657.asp).html). Naturwissenschaften 91 (6): 255–76. Berkeley: University of California Press. (2010). Making Sense of Evolution: Darwin. John (2006). Theodosius Grigorievich (1982). Robin Marantz (2000). John (2007). • Miller. Dobzhansky on Evolutionary Dynamics: Some Questions about His Russian Background. Pa: Trinity Press International. Creation evangelism: cutting through the excess (http://www. Valley Forge.110400. • Morris. PMID 14527300. Boston: Houghton Mifflin. Samuel (2007). Evolution. retrieved 2007-01-06 van Wyhe. John F.tamucc.. (2002).4.org/ docs2001/0510news.htm).070802. • Butler. "Creationism and intelligent design". "Religion in America: Intelligent Design on Trial".S. Gillian.J. Darwin.1146/annurev. Dr. Annu Rev Genomics Hum Genet 4: 143–63.Creation–evolution controversy 20060904151915/http://philosophy.answersingenesis.time. pp. (July/August 2003). HR (1963). Stephen Jay. Thomas E. The origin of species. N. htm) Creationism as social policy • Isaac Asimov. University Press. State (http://www.data360. Biology and Philosophy 14 (4): 481–504. php/ Main/ RoyalHollowayCollegeDebate)) 2007 . pc. • Maynard Smith. "The status of neo-Darwinism.Creation–evolution controversy • Scott.com/index/ T4220X33T0284444.umkc.com/poll/21814/Evolution-Creationism-Intelligent-Design. (http://www. ISBN 0310240506.html) Evolution versus creationism debates Location and link Year Pro-evolution Pro-creationism Stanford University (http:/ / webcast. Johnson 1996 1997 Kenneth R. ucsd.pdf). Michael Ruse. Grand Rapids. Lee (2004).aspx) • Data by country regarding the percentage of the population that believes in evolution (http://www. Annual Review of Anthropology 26 (1): 263–289. "The Use and Abuse of Sir Karl Popper" (http://www.springerlink.gallup. pbs. rhul.org/ctrl/azimov_creationism. David Berlinski & William F.anthro. Creationism. Edinburgh. to/ apologetics/ p45. bringyou. htm) 1994 Will Provine Phillip E. org. D. The "Threat" of Creationism. Eugenie Scott & Barry Lynn Lewis Wolpert Phillip E. ac. "Antievolution And Creationism In The United States".1146/annurev. and Intelligent Design as of May 2007 (http://www. ed.263.edu/faculty/projects/ftrials/ scopes/statcase. (1999).stephenjaygould.26.aspx?Graph_Group_Id=286) • Opinion of Tennessee Supreme Court in Scopes v.C. The case for a Creator: a journalist investigates scientific evidence that points toward God. doi:10. Waddington. • Hull. E. Retrieved 2008-04-11. Johnson. 358 External links • Gallup public opinion poll in regards to the concepts of Evolution." in "Towards a Theoretical Biology" (C. rm) Nova on-line (http:/ / www. uk/ sites/ debate/ debate_audio. Mich: Zondervan/Willow Creek Resources.1. (1997). edu:8080/ ramgen/ UCSD_TV/ 6287DarSciorPhi. Miller Kenneth R. org/ wgbh/ nova/ odyssey/ debate/ ) Firing Line (http:/ / www. Michael Behe.law. Buckley Steve Fuller University of London (http:/ / www.org/ graph_group.L. • Strobel. html) ( transcript (http:/ / bcseweb. Miller. doi:10.1023/A:1006554919188. Johnson Phillip E.H.. 1969. uk/ index. such as those advocating theistic evolution.S. the evolution of an organism has no "objective" other than increasing the organism's ability to survive and reproduce in its environment. and basic physical laws. It can even refer to metaphysical evolution. however. evidence.[4] Although most religions have accepted the occurrence of evolution. In the context of biology. evolution can refer to any sort of progressive development.[2] The observation of evolutionary processes occurring. The scientific community. does not accept such objections as having any validity. as well as the current theory explaining that evidence. to be more highly evolved or advanced than another. methodology. and include objections to evolution's evidence. and often bears a connotation of gradual improvement: here evolution is understood as a process that results in greater quality or complexity. orthogenesis). have been uncontroversial among mainstream biologists for nearly a century. evolution is genetic changes in populations of organisms over successive generations.[3] Since then. Biologists do not consider any one species. the idea of devolution ("backwards" evolution) is a result of erroneously assuming that evolution is directional or has a specific goal in mind (cf. neocreationism.[12] In colloquial contexts. nearly all criticisms of evolution have come from religious sources. there still exist religious beliefs which reject evolutionary explanations in favor of creationism. the belief that a deity supernaturally created the world largely in its current form.Objections to evolution 359 Objections to evolution Objections to evolution have been raised since evolutionary ideas came to prominence in the 19th century. and its suitability is only defined in relation to this environment. citing detractors' misinterpretations of scientific method. However. Such objections have often centered on undermining evolution's scientific basis. Their arguments against evolution have become widespread. leads to frequent misunderstandings. In reality. centric creation-evolution controversy has been a focal point of recent conflict between religion and science. recent objections to evolution have frequently blurred the distinction between the two. In contrast to earlier objections to evolution that were either strictly scientific or explicitly religious.[11] Defining evolution One of the main sources of confusion and ambiguity in the creation-evolution debate is the definition of evolution itself. this can cause errors such as the claim that modern evolutionary theory says anything about abiogenesis or the Big Bang. and scientific acceptance. rather than from the scientific community. the United Kingdom and Canada. For example. and should therefore be taught in public schools.[6] Modern creationism is characterized by movements such as Creation Science. Certain sources have been criticized for indicating otherwise due to a tendency to evaluate nonhuman organisms according to anthropocentric standards rather than more objective ones. When biological evolution is conflated with other evolutionary processes. when misapplied to biological evolution. such as humans. and Intelligent Design which argue that the idea of life being directly "designed" by a god or intelligence is at least as scientific as evolutionary theory. spiritual evolution. there is widespread belief in creationism in the Muslim world.[1] When Charles Darwin published his 1859 book On the Origin of Species. morality. in India 77% of respondents who had heard of Charles Darwin and knew something about the theory of evolution agreed it was backed by scientific evidence. This common definition. New Zealand.[7] While objections primarily originate from the United States. with the intent of combating the teaching of evolution as fact and opposing the spread of "atheistic materialism". plausibility.[10] However. from evolutionary computation to molecular evolution to sociocultural evolution to stellar and galactic evolution.[5] The resultant U.[8] South Africa and India[9] with smaller followings in Australia. the word has a number of different meanings in different fields.[13] . his theory of evolution by natural selection initially met opposition from alternate scientific theories. but came to be universally accepted by the scientific community. or any of a number of evolutionist philosophies. [13] Creationist sources frequently define evolution according to a colloquial. In the early 20th century. These were opposed on scientific grounds. forming the X Club to widespread acceptance.[18] [19] Kelvin's own views favoured a version of theistic evolution speeded up by divine guidance. as well as meeting political and religious objections. Although the history of life shows an apparent trend towards the evolution of complexity. an estimate strongly disputed by geologists. or stay the same. rather. pangenesis was replaced by .[15] As a result. Darwin's gradualistic account was also opposed by saltationism and catastrophism. decrease.[1] These ideas that natural laws controlled the development of nature and society gained vast popular audiences with George Combe's The Constitution of Man of 1828 and the anonymous Vestiges of the Natural History of Creation of 1844. in particular the transmutation of species theory put forward by Jean-Baptiste Lamarck. These figures were corrected in 1907 when radioactive dating of rocks showed that the Earth was billions of years old.Objections to evolution Evolution also does not require that organisms become more complex. within fifteen to twenty years he convinced most of the scientific community that evolution through common descent was true. there is a question if this appearance of increased complexity is real. but there was continued resistance to his views on the significance of natural selection. This also means that advocates of creationism and evolutionary biologists often simply speak past each other. or if this conclusion comes from neglecting the fact that the majority of life on earth has always consisted of prokaryotes.[14] In this view.[20] The specific hereditary mechanism Darwin provided. organisms' complexity can either increase. Darwin's contemporaries eventually came to accept the transmutation of species Darwin's theory of evolution through common descent gained based upon fossil evidence. Depending on the situation. most notably by Georges Cuvier. meaning. but while most accepted that natural selection was a valid and empirically testable hypothesis. When Charles Darwin published his 1859 book On the Origin of Species. which frequently made greater complexity advantageous. it is a consequence of the specific circumstances of evolution on Earth.[17] although the specific evolutionary mechanism which Darwin provided – natural selection – was actively disputed by alternative theories such as Lamarckism and orthogenesis. and thus naturally selected for. defend it against the church and wealthy amateurs. lacked any supporting evidence. and all three of these trends have been observed in evolution. rather than scientific. Lord Kelvin led scientific opposition to gradualism on the basis of his thermodynamic calculations that the Age of the Earth was between 24 and 400 million years old.[12] [16] 360 History Various evolutionary ideas came to prominence around the start of the nineteenth century.[2] The earliest objections to Darwinian evolution were both scientific and religious. pangenesis. complexity is not a necessary consequence of evolution. his view that it was the primary mechanism of evolution was generally rejected. many attempts to rebut evolution do not address the findings of evolutionary biology (see straw man argument). an argument from design still used by the creationist movement. acceptance of evolution remains low in the Muslim world as prominent figures reject evolution's underpinning philosophy of materialism as unsound to human origins and a denial of Allah. Believers in Biblical infallibility attacked Darwinism as heretical.[8] Further objections by Muslim scholars and writers largely reflect those put forward in the Western world. alleging variously that it is unscientific. Natural theology included a range of ideas and arguments from the outset. In 1996. Arkansas that forbidding the teaching of evolution on religious grounds violated the Establishment Clause. and to question that could lead to excommunication. Although early objections dismissed evolution for contradicting their interpretation of the Bible. This occurred relatively quickly as medieval Madrasah's taught ideas of Al-Jahiz. the encyclical Humani Generis of Pope Pius XII first mentioned evolution directly and officially. Pope John Paul II stated evolution was "more than a hypothesis" and acknowledged the large body of work accumulated in its support. but reiterated that any attempt to give a material explanation of the human soul was "incompatible with the truth about man.[29] Creationists have appealed to democratic principles of fairness. However. broke out in "acrid polemics" and argument about evolution from 1860 to the 1870s – with the turning point possibly marked with the death of Louis Agassiz in 1873 – and by 1880 a form of "Christian evolution" was becoming the consensus. such as genetics. especially in America.[28] Since then creationists have developed more nuanced objections to evolution. this argument was invalidated when the Supreme Court ruled in Epperson v. In 1950. infringes on creationists' religious freedoms or that the acceptance of evolution is a religious stance. a Muslim scholar from the 9th century. The ideas that species change over time through natural processes and that different species share common ancestors seemed to contradict the Genesis account of Creation.[27] This position has been adopted by denominations of Christianity and Judaism in line with modernist theology which views the Bible and Torah as allegorical thus removing the conflict between evolution and religion.[22] Frederick Temple's lectures on Relations between Religion and Science (1884) speaking on how evolution was not "antagonistic" to religion highlighted this trend. This led evolutionary theory to be "both permissible and respectable" by 1876.[24] It allowed for inquiry into humans coming from pre-existing living matter. but to not question Adam and Eve or the creation of the soul. which confirmed Darwin's predictions and refuted the competing theories. The modern synthesis rose to universal acceptance among biologists with the help of new evidence. Sir Henry Chadwick notes a steady acceptance of evolution "among more educated Christians" between 1860 and 1885. as this conflicted with First Vatican Council's (1869–70) finding that everything was created out of nothing by God.[23] Temple's appointment to Archbishop of Canterbury in 1896 showed the broad acceptance of evolution within the church hierarchy. and when Darwin's theory was published. leading to the rise of the modern evolutionary synthesis. ideas of theistic evolution were presented in which evolution is accepted as a secondary cause open to scientific investigation. However. This opposition developed into the creation-evolution controversy involving Christian literalists in the United States objecting to the teaching of evolution in public schools. it would rein in Catholics who proposed that evolution could be reconciled with the Bible. arguing that evolution is controversial.Objections to evolution Mendelian inheritance. while still holding belief in God as a first cause with a non-specified role in guiding evolution and creating humans.[22] For decades Catholicism avoided official refutation of evolution."[25] Muslim reaction covered the gamut with those believing in literal creation from the Qur'an while many educated Muslims subscribed to a version of theistic or guided evolution where the Qur'an reinforced rather than contradicted mainstream science. and that 361 .[26] Regardless of acceptance from major religious hierarchies. The natural theology of the early 19th century was typified by Paley's watchmaker analogy.[21] Protestantism. who proposed concepts similar to natural selection. in the 1920s Christian fundamentalists in the United States developed their literalist arguments against Modernist theology into opposition to the teaching of evolution due to fears that ‘‘Darwinism’’ had led to German militarism and was a threat to religion and morality.[22] In Britain while publication of The Descent of Man by Darwin in 1871 reinvigorated debate from the previous decade. early religious objections to Darwin's theory are still used in opposition to evolution.[8] However. referring to this theory's well-established nature. Under the colloquial definition. in which case scientists would indeed consider evolution "proven". a fact can simply refer to anything for which there is overwhelming evidence. however. tested. an attempt to gradually undermine evolution and ultimately to "reverse the stifling materialist world view and replace it with a science consonant with Christian and theistic convictions". an attempt by the Discovery Institute to promote the teaching of intelligent design in public schools. being controversial. however. evolution may be called a "fact" for the same reason that gravity can: under the scientific definition.[37] Critics state that evolution is not a fact. or controversial. therefore. the theory of evolution can also be called a fact. and refined by scientists. From a scientific standpoint. not natural sciences (where the proper term is "validated"). a fact is a verified empirical observation.[33] [34] or attempting to come up with alternative ideas such as Creationism to debate its findings. Thus. this argument makes the weaker claim that evolution. This in turn forms a major part of the Institute's "wedge strategy". the modern synthesis is constantly debated.[35] Creationists often argue.[38] In science.[45] This appeal to "fairness" and a more democratic.[44] Unlike past creationist arguments which sought to abolish the teaching of evolution altogether. and was endorsed by U. in science a theory is simply an explanation or model of the world that makes testable predictions. it refers to an explanation for the diversity of species and their ancestry. in that the colloquial meaning of proof is simply "compelling evidence".[30] .[30] These objections to evolution culminated in the intelligent design movement in the early 2000s that unsuccessfully attempted to present itself as a scientific alternative to evolution. for example. and no more an indictment of evolution than calling it a "theory". When evolution is used to describe a theory.[42] strict proof is possible only in formal sciences such as logic and mathematics. should be presented alongside other. evolution is widely considered both a theory and a fact by scientists. An example of evolution as theory is the modern synthesis of Darwinian natural selection and Mendelian inheritance. This reflects a misunderstanding of the meaning of theory in a scientific context: whereas in colloquial speech a theory is a conjecture or guess. As with any scientific theory.[36] Status as a theory Critics of evolution frequently assert that evolution is "just a theory". that evolution is unproven.[32] [42] [46] This objection forms the basis of the "Teach the Controversy" campaign. or of characterizing it as a matter of opinion rather than of fact or evidence. in colloquial contexts. evolution is an observable process that occurs whenever a population of organisms genetically changes over time. not just evolution. The distinction is an important one in philosophy of science. Bush.[39] [40] [41] Similar confusion is involved in objections that evolution is "unproven". even though they are purely theoretical. and students should be allowed to evaluate and choose between the options on their own.S. and is in a sense self-fulfilling: it argues that evolution is controversial or contentious. in common usage theories such as "the Earth revolves around the Sun" and "objects fall due to gravity" may be referred to as "facts". non-factual. President George W. "balanced" approach in which conflicting views are given "equal time" appeals to American creationists. For example.[43] Degree of acceptance In 1999 a new major objection to evolution appeared in Kansas. with the intent of emphasizing evolution's unproven nature. so this is trivially true.[31] [32] 362 Scientific acceptance Recent objections to evolutionary theory have focused on its scientific validity. alternative views. The confusion arises. as it relates to the lack of absolute certainty in all empirical claims.Objections to evolution science classrooms should therefore "Teach the Controversy". There is an overwhelming consensus in the scientific community that it remains the only robust model that accounts for the known facts concerning evolution. and that supporters of evolution dogmatically reject alternative suggestions out-of-hand. often based on the creationist belief that evolution has never been observed. it would hold no bearing on the merit of the theory itself. determines what is considered acceptable science.[42] [55] These claims have become more popular in recent years as the neocreationist movement has sought to distance itself from religion. creationists have disputed the level of scientific support for evolution. The Discovery Institute has gathered over 600 scientists since 2001 to sign "A Scientific Dissent From Darwinism" in order to show that there are a number of scientists who dispute what they refer to as "Darwinian evolution". supporters of evolution have argued that no scientist's claims. it is not a science". not popular opinion or fairness. or at least should be taught alongside other views (i. meets the basic scientific standards that would be required to make them scientific alternatives to evolution. it is argued that Darwin "recanted" on his deathbed.[50] Most commonly. Examples of claims made in such arguments are statements that evolution is based on faith.Objections to evolution Scientists and U. including Darwin's. Even if this myth were true. 2010) evolution-supporting scientists named "Steve". The scientific consensus of biologists.[46] [49] A similar objection to evolution is that certain scientific authorities – mainly pre-modern ones – have doubted or rejected evolution. it is a false dilemma to characterize this as evidence for intelligent design. This statement did not profess outright disbelief in evolution. but expressed skepticism as to the ability of "random mutation and natural selection to account for the complexity of life. and Project Steve. and that therefore evolution is a religion. and therefore should not be taught in science classes." Several counter-petitions have been launched in turn.. and it is argued that although evolution is clearly controversial in the public arena. that supporters of evolution revere Darwin as a prophet. but on evidence. creationism).000 signatures in four days. or any other creationist explanation. Creationists have argued for over a century that evolution is "a theory in crisis" that will soon be overturned. a false anecdote originating from the Lady Hope Story. and therefore does not require faith. which gathered over 7. as shown by the aspects of Darwin's theory that have been rejected or revised by scientists over the years. and to reframe the debate from being between science (evolution) and religion (creationism) to being between two equally religious beliefs – or even to argue that evolution is religious while intelligent design is not. it is entirely uncontroversial among experts in the field.[52] The purpose of this criticism is to undermine the higher ground biologists claim in debating creationists.S. . It is also argued that even if evidence against evolution exists.[48] based on objections that it lacks reliable evidence or violates natural laws.[51] These objections are generally rejected as appeals to authority. Religious nature Creationists commonly argue against evolution on the grounds that "evolution is a religion. as have claims that intelligent design.[47] In response. These objections have been rejected by most scientists. 363 Scientific status A common neo-creationist objection to evolution is that evolution does not adhere to normal scientific standards—that it is not genuinely scientific.[56] [57] The claim that evolution relies on faith. is likewise rejected on the grounds that evolution has strong supporting evidence. including A Scientific Support for Darwinism.[52] The arguments for evolution being a religion generally amount to arguments by analogy: it is argued that evolution and religion have one or more things in common. are treated as sacrosanct. These objections often deal with the very nature of evolutionary theory and the scientific method. to form first Neo-Darwinism and later the modern evolutionary synthesis.[53] [54] Those that oppose evolution frequently refer to supporters of evolution as "Evolutionists" or "Darwinists". a tongue-in-cheek petition that has gathered 1.140 (as of August 12.e. It is argued that evolutionary biology does not follow the scientific method.[47] In response. thus giving it more reason to make use of a seemingly anti-religious analogy. courts have rejected this objection on the grounds that science is not based on appeals to popularity. philosopher Karl Popper said that "Darwinism is not a testable scientific theory but a metaphysical research programme". in chapter 6 of On the Origin of Species wrote: "If it could be demonstrated that any complex organ existed. They claim that any observation can be fitted into the evolutionary framework.[62] In contrast. Popper later recanted and offered a more nuanced view of its status: Darwin and an ape. the remedy is to stop the teaching of evolution. which .[59] 364 Unfalsifiability A statement is considered falsifiable if there is an observation or a test that could be made that would demonstrate that the statement is false. and perhaps also in common sense. Morris have claimed that evolution is unfalsifiable. such as the fossil record showing no change over time. Darwin's own most important contribution to the theory of evolution. Creationists such as Henry M. courts have also rejected this objection: Assuming for the purposes of argument. as it were. because any result of an experiment or investigation could be the unpredictable action of an omnipotent deity. they claim that evolution is non-scientific. his theory of natural selection. on the grounds that its methodological naturalism is as unproven. that evolution is a religion or religious tenet.[64] However. U. but by its spiritual or supernatural beliefs. evolution is considered falsifiable by scientists because it can make predictions that.[65] [66] The most direct evidence that evolutionary theory is falsifiable may be the original words of Charles Darwin who. however.[60] [61] However. There are some tests.[58] A related claim is that evolution is atheistic. that evolution is not a religion and that teaching evolution does not violate the Establishment Clause. and thus as "faith-based". we can observe natural selection happening under our very eyes. or observations showing organisms being created supernaturally or spontaneously. were they contradicted by the evidence. even some experimental tests. and that therefore it is impossible to demonstrate that evolution is wrong. Statements that are not falsifiable cannot be examined by scientific investigation since they permit no tests that evaluate their accuracy. Nevertheless. it is argued that "science is an atheistic religion". circa 1874 from Julkaistussa picture source. Several kinds of evidence have been proposed that could falsify evolution. as the supernatural and theistic beliefs of creationism. As such. However. much more so than tests of otherwise comparable theories in physics or chemistry. would falsify evolution. and they accuse creationists of equivocating between the strict definition of religion and its colloquial usage to refer to anything that is enthusiastically or dogmatically engaged in.[54] This argument against evolution is also frequently generalized into a criticism of all science. creationists sometimes merge the two claims and describe evolution as an "atheistic religion" (cf.S. such as the famous phenomenon known as 'industrial melanism'. creationism consists largely of unsubstantiated claims that evolution has been falsified. really severe tests of the theory of natural selection are hard to come by.[60] Creationist explanations involving the direct intervention of the supernatural in the physical world are not falsifiable.[63] In 1976.[60] Many of Darwin's ideas and assertions of fact have been falsified as evolutionary science has developed and has continued to confirm his central concepts. Yet it is clearly established in the case law. confirmation that mutations are prevented from accumulating. Evolutionary supporters point out evolution is neither dogmatic nor based on faith. is difficult to test. not establish another religion in opposition to it. and in some cases. humanism).Objections to evolution In general. the argument that evolution is religious has been rejected on the grounds that religion is not defined by how dogmatic or zealous its adherents are. A related claim. avoids being trivially true."[67] In response to the unfalsifiability criticism of evolutionary theory. At any time. Fecundity A great scientific theory. Kitcher agrees with Popper that “there is surely something right in the idea that a science can succeed only if it can fail. and independent testability of auxiliary hypotheses: Unity “A science should be unified . that can be applied to a wide range of problems” (1982: 47).[77] As Kitcher points out.” which are.. this phrase.[70] The assertion of common descent could also have been disproven with the invention of DNA analysis.Objections to evolution could not possibly have been formed by numerous. The fusion hypothesis was confirmed in 2005 by discovery that human chromosome 2 is homologous with a fusion of two chromosomes that remain separate in other primates. it is argued that evolutionary theory is circular reasoning. people suggested that another planet influenced Uranus’ orbit – and this prediction was indeed eventually confirmed. J. but evolution is required to interpret the evidence.[65] Specifically.B. Abusing Science: The Case Against Creationism. fecundity. If not. in that evidence is interpreted as supporting evolution. the evolution of chimpanzees and humans from a common ancestor predicts a (geologically) recent common ancestor.” observations of Uranus when first discovered in 1781 would have “falsified” Newton’s celestial mechanics. Because a theory presents a new way of looking at the world.. Haldane.[76] In his book. my theory would absolutely break down. DNA analysis has shown that humans and chimpanzees share a large percentage of their DNA (between 95% to 99. Rather.[68] [69] Numerous other potential ways to falsify evolution have also been proposed. However. Numerous transitional fossils have since been found.. it can lead us to ask new questions. also once used.[43] For example. it is often argued that the phrase "survival of the fittest" is a tautology. On the contrary.. incompleteness is the mother of fecundity. the fact that humans have one fewer pair of chromosomes than the great apes offered a testable hypotheses involving the fusion or splitting of chromosomes from a common ancestor. If true. A good theory should be productive. in that fitness is defined as ability to survive and reproduce. in most cases strata are not dated by their fossils. Typically.[42] However. but by their position relative to other strata and by radiometric dating. if one took a strictly Popperian view of “theory. Extra. human evolution has passed several falsifiable tests. successive.4% depending on the measure). unlike simple "survivability". According to Kitcher. inactive telomeres and centromeres remain on human chromosome 2 as a result of the fusion.”[78] But he insists that we view scientific theories as consisting of an “elaborate collection of statements. and most strata were dated before the theory of evolution was formulated. human DNA should be far more similar to chimpanzees and other great apes. good scientific theories must have three features – unity. but then abandoned. and so to embark on new and fruitful lines of inquiry. An example of this is the claim that geological strata are dated through the fossils they hold. opens up new areas of research.[72] Hence. philosopher of science Philip Kitcher specifically addresses the "falsifiability" question by taking into account notable philosophical critiques of Popper by Carl Gustav Hempel and Willard Van Orman Quine that reject his definition of theory as a set of falsifiable statements. a flourishing science is incomplete... and others – what he calls “auxiliary hypotheses.” some of which are not falsifiable. by Popper. slight modifications. when asked what hypothetical evidence could disprove evolution. like Newton’s. or a small family of problem-solving strategies. this definition.. first used by Herbert Spencer in 1864. Additionally. it raises more questions than it can currently answer.[73] [74] [75] Similarly. but that fossils are in turn dated by the strata they are in..[71] Also. is rarely used by biologists.. it should raise new questions and 365 . than to other mammals. replied "fossil rabbits in the Precambrian era". But incompleteness is no vice. Good theories consist of just one problem-solving strategy. then common descent is falsified. fitness is more accurately defined as the state of possessing traits that make survival more likely. is that natural selection is tautological. numerous examples of potential ways to falsify evolution have been proposed.S. inherited. From Kitcher’s point of view. can be addressed by advancing Darwinian histories. it was important to resolve general issues about the presuppositions of Darwinian histories.g. and it was clear that biologists had to tackle questions for which they had.. Reasoning like this can be used to answer a host of biological questions. The first step consists in a description of an ancestral population of organisms. including Popper’s. He is also taking into account the way the life sciences work. Evolutionary theory is unified because so many diverse questions . Like other definitions of theories. Moreover. He also gave a structure to our ignorance. The reasoning proceeds by tracing the modification of the population through subsequent generations. showing how characteristics were selected. The production of new hypotheses is another possible – and equally important – observational consequence. like the observation of irregularities in Uranus’s orbit.” But. A Darwinian history is a piece of reasoning of the following general form.Objections to evolution presume that those questions can be answered without giving up its problem-solving strategies (1982: 47–48). these narratives constantly make claims that are subject to independent check. the evidence for the existence of Neptune is independent of the anomalies in Uranus’s orbit). falsification is only one possible consequence of an observation. independently of the theory it is designed to save” (1982: 46) (e. Darwinian theory not only meets the three conditions for a good scientific theory. related by their common employment of a particular style of historical narrative. Kitcher makes it clear that a good theory includes statements that have (in his terms) “observational consequences. The way in which biology should proceed had been made admirably plain. no answers. and became prevalent. Kitcher’s account of a good theory is based not only on his understanding of how physical sciences work.[79] The same kind of story can be told again and again to answer all sorts of questions about all sorts of living things. as yet. Auxiliary hypothesis that are independently testable “An auxiliary hypothesis ought to be testable independently of the particular problem it is introduced to solve. After Darwin.[80] Darwin not only provided a scheme for unifying the diversity of life..[81] 366 . it is without question an extraordinarily successful theory: The heart of Darwinian evolutionary theory is a family of problem-solving strategies. which by definition cannot be directly observed. considering the former to simply be the latter on a larger scale. as biologists define macroevolution. both microevolution and macroevolution have been observed. from asserting that such fossils are hoaxes or that they belong exclusively to one group or the other. including the evolution of fruit flies. and this. not microevolution:[85] [86] most creationist organizations do not dispute the occurrence of short-term. such as that observed even in dog breeding. provide plausible links between several different groups of organisms. despite popular misconceptions to the contrary.Objections to evolution 367 Evidence Objections to the evidence that evolution occurs tend to be more concrete and specific. mice and bacteria in the laboratory. particularly those using microorganisms. have probably been short in comparison with the periods during which these same species remained without undergoing any change. Evolutionary processes." Darwin appealed to the limited collections then available. only inferred from microevolutionary processes and the traces of macroevolutionary ones. Such studies on experimental evolution. as I believe. in the form of populations changing their genetic composition from generation to generation. is the most obvious and gravest objection which can be urged against my theory. for example. Under the conventional biological definition of evolution. though very long as measured by years. have been directly observed many times.[89] or the recently-discovered Tiktaalik linking fish and limbed amphibians.[83] and of tilapia in the field. Transitional fossils. However. Lack of observation A common claim of creationists is that evolution has never been observed.[82] Challenges to such objections often come down to debates over how evolution is defined (see above)."[92] The number of clear transitional fossils has increased enormously since Darwin's day. Rather. relatively minor evolutionary changes. often involving direct analysis of evolutionary biology's methods and claims. the extreme lengths of time involved. The explanation lies. past macroevolution can be inferred from historical traces. and this problem has been largely resolved with the Transitional species such as the Archaeopteryx have been a fixture of .[83] [84] In response to such examples. to asserting that there should be far more evidence of obvious transitional species. it is a simple matter to observe evolution occurring. have been observed in different scientific contexts. for example. in the extreme imperfection of the geological record. Additionally. creationists specify that the creation-evolution debate for almost 150 years. the modern evolutionary synthesis draws no distinction between macroevolution and microevolution. and different rates of change with some living species differing very little from fossils of the Silurian period. Speciations.[87] Additionally. are now providing important insights into how evolution occurs.[91] Darwin himself found the paucity of transitional species to be one of the greatest weaknesses of his theory: "Why then is not every geological formation and every stratum full of such intermediate links? Geology assuredly does not reveal any such finely graduated organic chain. perhaps. they dispute the occurrence of major evolutionary changes over long periods of time.[43] [88] An example of this is ring species. such as Archaeopteryx linking birds and dinosaurs. they are objecting only to macroevolution.[90] Creationists dispute such examples. In later editions he added "that the periods during which species have been undergoing modification. When such observations contradict a theory's predictions. They thus claim that current evolutionary theory is likely to undergo such a revolution in the future. since species which diverged more recently will be more closely related genetically than species which are more distantly related. scientific claims are just speculation or "story-telling". Jonathan Wells criticizes biology textbooks by alleging that they continue to reproduce such evidence after it has been debunked. and Darwin's finches. that current evidence for evolution is likely to be overturned because past evidence has been. Newton's theory of gravitation was replaced by Einstein's theory of General Relativity when the latter was observed to more precisely predict the orbit of Mercury. it may be revised or discarded if an alternative better explains the observed facts. were not merely errors but frauds. Typically. such as Romanes's 1892 copy of Ernst Haeckel's embryo Ernst Haeckel's 19th-century comparative drawings of embryos. such phylogenetic trees show a hierarchical organization within the tree of life.[98] [99] In fields such as astrophysics or meteorology. and have continued to insist that more dramatic demonstrations of evolution be experimentally produced. these claims have been subsequently refuted. This objection goes further than the less substantial "evolution isn't proven" arguments. peppered moth melanism. creationists have redefined their understanding of what amounts to a "created kind".[103] [104] [105] It has also been claimed that certain former pieces of evidence for evolution which are now considered out-of-date and erroneous. For example. It argues that because no one except God could directly observe events in the distant past. In such fields. major scientific revolutions have overturned theories that were at the time considered near-certain.[93] Creationists counter that even observed speciations and transitional fossils are insufficient evidence for the vast changes summarized by such phrases as "fish to philosophers" or "particles to people". Much of the evidence for evolution has been accused of being fraudulent at various times. claiming that evolution isn't even well-evidenced. It is argued that because scientists have been mistaken and deceived in the past about evidence for various aspects of evolution the current evidence for evolution is likely to also be based on fraud and error.[96] [97] DNA sequences of the genomes of organisms allow an independent test of their predicted relationships. [106] Phylogeny"). used drawings. often attributed incorrectly to [102] to illustrate his Recapitulation theory ("Ontogeny recapitulates Haeckel. Creationists point out that in the past.[104] In response. where direct observation or laboratory experiments are difficult or impossible.[100] 368 Instability of evidence A related objection is that evolution is based on unreliable evidence. the test of falsifiability is satisfied when a theory is used to predict the results of new observations.[95] One version of this objection is "Were you there?".[94] As more and more compelling direct evidence for inter-species and species-to-species evolution has been gathered. or that certain types of evidence are inconsistent and dubious. as predicted by common descent. which predicts a primarily stable fossil record broken up by occasional major speciations.[101] Critics of evolution commonly appeal to past scientific hoaxes such as the Piltdown Man forgery. this is either based on the argument that evolution's evidence is full of frauds and hoaxes.Objections to evolution advent of the theory of punctuated equilibrium. Arguments against evolution's reliability are thus often based on analyzing the history of evolutionary thought or the history of science in general. popularized by Ken Ham. the National Center for Science Education notes that none of the textbooks reviewed by . the scientific method instead relies on observation and logical inference. including Archaeopteryx. on the basis that it is a "theory in crisis" for one reason or another. consistent decay rates. and therefore that an intelligence. The idea that it is simply too implausible for life to have evolved is often encapsulated with a quotation that the "probability of life originating on earth is no greater than the chance that a hurricane sweeping through a scrap-yard would have the luck to assemble a Boeing 747" (a claim attributed to astrophysicist Fred Hoyle and known as Hoyle's fallacy). rather than evolution's gradualistic punctuated equilibrium. God. or rocks acting as closed systems. to have arisen "by chance". Improbability A common objection to evolution is that it is simply too unlikely for life. results. based on the Carbon 14 isotope. in its complexity and apparent "design". These include that there are too many "gaps" in the fossil record.[115] A more extreme version of this argument is that evolution cannot create complex structures. It is argued.[116] Because the theory of evolution is often thought of as the idea that life arose "by chance". generates inconsistent. The basic idea of this argument for a designer is the teleological argument. are seemingly "out of place".[112] which some assert is an "ad-hoc" theory to explain the fossil gaps. such as the principle of uniformitarianism. This view is thus invariably justified with arguments from analogy. design arguments such as William Paley's watchmaker analogy have been popular objections to the [114] theory since Darwin's day. Examination by geologists have found polystrate fossils to be consistent with in situ formation. has been particularly criticized.[113] Plausibility Some of the oldest and most common objections to evolution dispute whether evolution can truly account for all the apparent complexity and order in the natural world. It is argued that the odds of life having arisen without a deliberate intelligence guiding it are so astronomically low that it is unreasonable not to infer an intelligent designer from the natural world. an argument for the existence of God based on the perceived .[111] It is argued that certain features of evolution support creationism's catastrophism (cf. that radiometric dating. such as polystrate fossils. as Haeckel's drawings are shown in a historical context with discussion about why they are wrong.[108] Another form of this objection is that fossil evidence is not reliable. the technique of evaluating a material's age based on the radioactive decay rates of certain isotopes. It is argued that evolution is too unlikely or otherwise lacking to account for various aspects of life.[107] 369 Unreliable or inconsistent evidence Creationists claim that evolution relies on certain types of evidence that do not give reliable information about the past. different radiometric dating methods and techniques have independently confirmed each other's results. Great Flood). Such arguments have been dismissed by scientists on the grounds that independent methods have confirmed the reliability of radiometric dating as a whole. and thus unreliable. and specifically from the diversity of life. and the accurate modern drawings and photos used in the textbooks are misrepresented by Wells. It is argued that radiometric decay relies on a number of unwarranted assumptions. Radiocarbon dating. must at the very least be appealed to for those specific features. or that certain fossils. This is based on a much wider range of claims.Objections to evolution Wells makes the claimed error. for example.[109] [110] that fossil-dating is circular (see evolution is unfalsifiable). additionally. language. Similar objections sometimes conflate the Big Bang with evolution. or necessary precursor to. it is irrational to trust whatever reasoning one relies on to conclude that it is true. arguments against "evolution" are based on the misconception that abiogenesis is a component of. that certain structures demonstrate poor design. and altruism (see altruism in animals). or cannot. which argues that certain natural phenomena are analogical to a watch (in that they are ordered. which asserts that it is irrational to reject a supernatural. Plantinga argues that whereas a God would be expected to create beings with reliable reasoning faculties. emotion. Specifically. emotions. However. instinct. have been well-explained by evolution. photosynthesis. and have them be taught alongside evolution. Although there is broad agreement that certain aspects of life remain unexplained.[117] It is also argued that there is insufficient evidence to make statements about the plausibility or implausibility of abiogenesis.[28] [47] This objection is fundamentally an argument by lack of imagination. Supporters of evolution generally respond by arguing that evolution is not based on "chance". evolution. it is the non-random selection of survival-enhancing genes that drives evolution along an ordered trajectory.[47] [114] It has also been noted that arguments against some form of life arising "by chance" are really objections to nontheistic abiogenesis. homosexuality. rather than supernatural beings. they must have been designed by a "watchmaker"—an intelligent agent. explain a certain aspect of the natural world.[119] [120] 370 Unexplained aspects of the natural world It is frequently argued that a great weakness of evolutionary theory is that it does not. Plantinga claims that evolution cannot account for the rise of reliable reasoning faculties. such as homosexuality.[12] Christian apologist and philosopher Alvin Plantinga. The fact that the results are ordered and seem "designed" is no more evidence for a supernatural intelligence than the appearance of complex natural phenomenon (e. and that the implausibility of life evolving exactly as it did is no more evidence for an intelligence than the implausibility of a deck of cards being shuffled and dealt in a certain random order. or complex. Behe argues that such structures were "purposely arranged by an intelligent agent" (see argument from incredulity). Michael Behe has argued that current evolutionary theory cannot account for certain complex structures. and therefore an alternate. but on predictable chemical interactions: natural processes.[122] Most of these. if conducive to survival. meaning that if evolution is true. A common way of using this as an objection to evolution is by appealing to the 18th-century philosopher William Paley's watchmaker analogy. It is argued that an alternative explanation. among the phenomena that critics variously claim evolution cannot explain are consciousness. Indeed. creationists argue that evolution should be abandoned altogether because of the phenomena it does not explain. On this basis. such as intelligent design. rather than as philosophy or theology. a neocreationist movement seeking to establish certain variants of the design argument as legitimate science. music. supporters of evolution contend that no alternative explanation has been able to adequately explain the biological .Objections to evolution order or purposefulness of the universe. and altruism. hominid intelligence. or purposeful). like a watch. morality.[118] This novel epistemological argument has been criticized similarly to other probabilistic design arguments. hominid intelligence. metamorphosis. making the natural development of reliable cognitive faculties more likely than unreliable ones. or only have preliminary explanations. or argument from incredulity: a certain explanation is seen as being counter-intuitive. This argument forms the core of intelligent design. are the "designer". language. more intuitive explanation is appealed to instead. free will. has formalized and revised the improbability argument as the evolutionary argument against naturalism. can explain the things which evolution cannot. not to evolution. photosynthesis.[121] In addition to complex structures and systems. For example. which means that. a supporter of intelligent design. is more likely to be selected for than irrationality. particularly in microbiology. It has also been argued that rationality. religion. while others remain mysterious. instincts. evolution would be just as likely to lead to unreliable ones.g. Although the process involves some random elements. intelligent creator because the apparent probability of certain faculties evolving is so low. snowflakes). evolution is being redefined to refer to the entire history of the universe. not Evolution" . The earliest feathers might have been a different approach to hairiness among reptiles keeping warm. the Big Bang. then the difficulty of believing that a perfect and complex eye could be formed by natural selection. cellular. if all the intermediary stages between an initial organ and obstacles to a gradual evolution of flagella. Richard Dawkins said on the topic of the evolution of the feather in an interview for the television program The Atheism Tapes: There's got to be a series of advantages all the way in the feather. but rather impossible. Creation of complex structures "Living things have fantastically intricate features-at the anatomical. can hardly be considered real. such as abiogenesis.Objections to evolution origin of these phenomena either. it will be impossible for the later organ to develop by the process of natural selection alone. and molecular levels-that could not function if they were any less complicated or sophisticated.[127] Research has confirmed that the natural evolution of the eye and other intricate organs is entirely feasible.[123] [124] Creationists argue against evolution on the grounds that it cannot explain certain non-evolutionary processes. Creationist arguments have been made such as "What use is half an eye?" and "What use is half a wing?". and the variations be inherited.[92] Similarly.. If you can't think of one. and this objection to evolution has been refined in recent years as the more sophisticated irreducible complexity . however. At this point. and it is argued that if one aspect of the universe is seemingly inexplicable. then that's your problem not natural selection's problem. the eye does vary ever so slightly. each grade being useful to its possessor. the organ it will become are not all improvements upon the original. In such instances. Anticipating early criticisms that the evolution of the eye and other complex organs seemed impossible. or the meaning of life. It's perfectly possible feathers began as fluffy extensions of reptilian scales to act as insulators.. Jonathan Sarfati[126] Modern evolutionary theory posits that all biological systems must have evolved incrementally.[128] [129] Creationist claims have persisted that such complexity evolving without a designer is inconceivable. The only prudent conclusion is that they are products of Intelligent Design.from Refuting Evolution 2. claiming that a major aspect of evolution is not merely unscientific or implausible. and if any variation or modification in the organ be ever useful to an animal under changing conditions of life. through a combination of natural selection and genetic drift. Both Darwin and his early detractors recognized the potential problems that could arise for his theory of natural selection if The bacterial flagellum is often invoked to the lineage of organs and other biological features could not be illustrate the concept of irreducible complexity. that if numerous gradations from a perfect and complex eye to one very imperfect and simple. which is certainly the case. because it contradicts some other law of nature or is constrained in such a way that it cannot produce the biological diversity of the world.[125] 371 Implausibility This class of objections is more radical than the above. can be shown to exist. though insuperable by our imagination. objections leave the arena of evolutionary biology and become general scientific or philosophical disputes. the entire body of scientific theories must be baseless. if further. accounted for by merely gradual. Darwin noted that: [R]eason tells me.. step-by-step changes over successive Careful analysis shows that there are no major generations.. Behe's primary examples of irreducible complexity has been cellular and biochemical in nature. and that evolution without an intelligent agent cannot account for the generation of information that would be required to produce specified complexity. objection of creationists to evolution is that evolutionary mechanisms such as mutation cannot generate new information. such as the evolution of nylon-eating bacteria.[142] However.[130] "In fact. makes frequent appeals to concepts from information theory in its objections to evolution and affirmations of the Genesis account of Creation. as mutating organisms either reproduce or fail.[133] [134] The idea that seemingly irreducibly complex systems cannot evolve has been refuted through evolutionary mechanisms. If that happened. Dembski has argued that life demonstrates specified complexity. grow it for ten thousand generations. or any other unintelligent process."-Michael Behe[131] In the years since Behe proposed irreducible complexity. which developed new enzymes to efficiently digest a material that never existed before the modern era. new information is regularly generated in evolution. and see if a flagellum—or any equally complex system—was produced.[143] ."[138] However. The flip side of this claim is that the flagellum can’t be produced by natural selection acting on random mutation. such as exaptation (the adaptation of organs for entirely new functions)[135] and the use of "scaffolding". Creationists such as William A. Here is a thought experiment that makes the point clear. a scientist could go into the laboratory. these claims have been widely rejected by the scientific community. which are initially necessary features of a system that later degenerate when they are no longer required. and Lee Spetner have attempted to use information theory to dispute evolution. It is not created. but rather gathered from the environment through research—by trial and error. He has argued that the components of systems such as the blood clotting cascade. formulated by biochemist Michael Behe. my claims would be neatly disproven. and the minority of mutations which are beneficial or harmful are often situational. and strongly supportive of Genesis creation. Whereas past arguments of this nature generally relied on macroscopic organs. when an organism is considered together with the environment it evolved in. Dramatic examples of entirely new. and therefore that they could not have evolved naturally from less complex or complete systems. a mutation that is harmful in one environment may be helpful in another. "[I]t should be clear that a rigorous application of the science of information is devastating to materialistic philosophy in the guise of evolution. and increasingly common. my argument for intelligent design is open to direct experimental rebuttal. whenever a novel mutation or gene duplication arises. place a bacterial species lacking a flagellum under some selective pressure (for mobility.[141] A related argument against evolution is that most mutations are harmful. such as an improved understanding of the evolution of flagella. for example. say). the immune system.Objections to evolution argument of the intelligent design movement. there is no need to account for the creation of information. The Christian apologetics site Answers in Genesis. In Darwin’s Black Box (Behe 1996) I claimed that the bacterial flagellum was irreducibly complex and so required deliberate intelligent design.[133] [136] [137] 372 Creation of information Another new. Dembski. and the bacterial flagellum are so complex and interdependent that they could not have evolved from simpler systems.[28] Irreducible complexity is the idea that certain biological systems cannot be broken down into their constituent parts and remain functional.[139] [140] In fact. unique traits arising through mutation have been observed in recent years. To falsify such a claim. in reality the vast majority of mutations are neutral. Werner Gitt.[132] have already undermined these arguments. new developments and advances in biology. The information in the genome forms a record of how it was possible to survive in a particular environment. Potential evolutionary pathways have been provided for all of the systems Behe used as examples of irreducible complexity. . because the enormous increase in entropy due to the Sun and Earth radiating into space dwarfs the local decrease in entropy caused by the existence and evolution of self-organizing life. Though the law applies to all systems. an ideal isolated system's entropy (a measure of the dispersal of energy in a physical system so that it is not available to do mechanical work) will tend to increase or stay the same. In other words. The Sun-Earth-space system does not violate the second law.Objections to evolution 373 Violation of the second law of thermodynamics Another objection is that evolution violates the second law of thermodynamics. Water freezes into ice and fertilised eggs turn into babies. are open systems. but [we do] not invoke divine intervention to explain the process [. mathematician Jason Rosenhouse stated: The fact is that natural forces routinely lead to decreases in entropy. or disorder. in the case of a closed one it states. approaching a maximum value at equilibrium". Creationists argue that evolution violates this physical law by requiring a decrease in entropy. this argument can be analyzed quantitatively.[144] However. in contrast. the Earth absorbs energy from the Sun and emits energy back into space. and radiate and absorb heat..[148] . over time. who concluded: "Quantitative estimates of the entropy involved in biological evolution demonstrate that there is no conflict between evolution and the second law of thermodynamics. which do not exchange matter or energy with their surroundings. Organisms. Similarly.[146] [147] This was done by physicist Daniel Styer. Plants use sunlight to convert carbon dioxide and water into sugar and oxygen.] thermodynamics offers nothing to dampen our confidence in Darwinism."[146] In a published letter to the editor of The Mathematical Intelligencer titled "How anti-evolutionists abuse mathematics". this claim is based on a manifestation of the law only applicable to isolated systems. "the entropy of an isolated system not in equilibrium will tend to increase over time. not decrease. as they constantly exchange energy and matter with their environment: for example animals eat food and excrete waste.[39] [145] Since the second law of thermodynamics has a precise mathematical definition. Evolutionary theory stands at the base of moral relativism and the rejection of traditional morality". Assertions of ape ancestry in the 1844 publication Vestiges of Creation gained much public acceptance but were scornfully attacked by establishment scientists. homosexuality and a host of other issues are really debates about the origin—and thus the meaning—of human life.[154] [155] Creationist Ken Ham likens evolution to a horde of termites. teen pregnancies. some people feel that humans should be considered separate from. such as Catholicism. however. Jr. President of Southern Baptist Theological Seminary in Louisville. and events.. abortion. behaviors. abortion. "Taking Issue". homosexuality. The mediaeval concept of a great chain of being set out a static hierarchy in which humans are "above" animals. but below angels and God. and they argue against evolution on the basis that it contradicts their literal interpretation of creation myths about separate "kinds". have reconciled their beliefs with evolution through theistic evolution. animals. environmentalism. It is argued that the teaching of evolution degrades values. and fosters irreligion or atheism. Ham suggests that "evolutionary termites" are responsible for pornography. most religions. Albert Mohler. These may be considered appeals to consequences (a form of logical fallacy). including bad beliefs. and/or superior to. accompanied by a satirical disclaimer claiming a fait accompli for its [149] inclusion Huxley's book Man's Place in Nature was the first devoted to human evolution and an early example of comparative biology.[150] Traditionalists still object to the idea that diversity in life. English biologist Thomas Henry Huxley showed that they were wrong and overcame their opposition to Darwin's ideas. but instead shows that animals behave in different ways. "I'm not saying that evolution is . undermines morals. Anatomists claimed that humans had unique physical features. Humans as animals Even though biology has long shown that humans are animals.. in National Public Radio's forum. 2005. homosexual behavior and lawlessness. Kentucky. As evolutionary thought developed it was feared that evolution implied that there was no real separation between man and brute.. arose through natural processes without a need for supernatural intervention. Social effects Some Creationists claim that perceived social ills like crime..[153] R. that "Debates over education. However. weakening society's foundation.Objections to evolution 374 Moral implications Other common objections to evolution allege that evolution leads to objectionable results. including human beings. and teaches that humans behave like humans. immorality. He also writes. and though Darwin avoided the subject when announcing his theory in On the Origin of Species in 1859. as the potential ramifications of belief in evolutionary theory have nothing to do with its objective empirical reality. Evolutionary common descent does not imply that human beings should behave like other animals. wrote August 8. In Why Won't They Listen?.[151] [152] 1871 caricature of Charles Darwin as an ape. the issue was quickly raised. and that it undermined human social hierarchy. wars and genocide are caused by a belief in evolution. "[159] [160] [161] Discovery Institute fellow Richard Weikart has made similar claims. these strong nations could be said to have "survived" in the struggle for dominance. Morris. and what has it brought us? Whether Darwin intended it or not. and Darwin and evolutionary theory cannot explain Hitler’s genocidal madness. and our world". and some argue that the opposite seems to be the case. D. avoiding lies. no Darwin. Kennedy also states that. fidelity. typically European. including belief in creationism and disbelief in evolution. was part of Plato's and Aristotle's philosophies. unwed mothers. Paul found that religious beliefs. The Anti-Defamation League describes such claims as outrageous misuse of the the Holocaust and its imagery. and as trivializing "the many complex factors that led to the mass extermination of European Jewry.[173] Strong. moral codes and the rule of law. unwed births. World War II. Careful analyses of the creationist charges that evolution has led to moral relativism and the Holocaust yields the conclusion that these charges appear to be highly suspect. the more common image subscribed to by creationists." particularly in the nineteenth century. divorce and child abuse. Hitler did not need Darwin or evolution to devise his heinous plan to exterminate the Jewish people. are positively correlated with social ills like crime. "We have had 150 years of the theory of Darwinian evolution."[156] Former Texas Republican Representative Tom DeLay claimed that the Columbine school shootings were caused by the teaching of evolution. with the exception of Christian missionaries.[82] Kent Hovind's son Eric Hovind has now taken over the family business while his father is in prison. premarital sex. and the accompanying pamphlet with the same title. millions of deaths. Europeans.Objections to evolution the cause of abortion or school violence.[173] With this attitude. the destruction of those deemed inferior. claims that evolution was part of a pagan religion that emerged after the Tower of Babel. both sects which reject evolution and embrace creationism.[158] Rev. and their attitudes. the devaluing of human life. World War I. will also change. DeLay is quoted as stating that "Our school systems teach the children that they are nothing but glorified apes who are evolutionized [sic] out of some primordial soup.[164] The claim was central to Ben Stein's film Expelled: No Intelligence Allowed promoting intelligent design creationism. Darwin’s Deadly Legacy.[162] [163] as have other creationists. Stalin's war crimes. James Kennedy's documentary. James Kennedy of The Center for Reclaiming America for Christ and Coral Ridge Ministries claims that Darwin was responsible for Adolf Hitler's atrocities. and as such. engineering professor and founder of the Creation Research Society and the Institute of Creation Research. the Vietnam War and Pol Pot's Cambodian killing fields on evolution.[167] The Barna Group surveys find that Christians and non-Christians in the US have similar divorce rates.[173] 375 . the exploitation of "lesser breeds without the law" by "superior races. A study published by the author and illustrator Gregory S. and was responsible for everything from war to pornography to the breakup of the nuclear family. as well as the increase in crime.[169] He goes on to suggest that evolution gives more support to the notion of an omnipotent creator. then the way people think. increasing hopelessness."[161] [165] Kent Hovind of Creation Science Evangelism blames communism. and claims that evolution is responsible for tattoos. no Hitler. nations successfully expanded their empires. anti-Semitism existed long before Darwin ever wrote a word. seldom adopted the customs and languages of local people under their empires. What I'm saying is that the more a culture abandons God's word as the absolute authority. socialism. Kennedy states that "To put it simply. and the more a culture accepts an evolutionary philosophy. the Holocaust. and other social ills.[168] Michael Shermer argued in Scientific American in October 2006 that evolution supports concepts like family values. rather than a tinkerer with limitations based on a human model. and the highest divorce rates in the US are among Baptists and Pentecostals." In his efforts to expose the "harmful effects that evolution is still having on our nation.[171] [172] Evolution has been used to justify Social Darwinism.[166] Supporters of evolution dismiss such criticisms as counterfactual."[157] Henry M. racism.[170] Such analyses conclude that the origins of the Holocaust are more likely to be found in historical Christian anti-semitism than in evolution. our children. Moreover. sexually transmitted diseases (STDs). body piercing. In D. Also.[183] While about 55% of scientists surveyed were atheists. and one a member of the Russian Orthodox Church and the author of a book on religion and science. and Theodosius Dobzhansky — one was a devout Anglican who preached sermons and published articles in church magazines.[179] Molleen Matsumura of the National Center for Science Education found that "of Americans in the twelve largest Christian denominations. "Can we seriously expect non-Christians to develop a respect for Christianity if we insist on teaching the brand of science that creationism brings with it?"[176] However. Presbyterian Church (USA). one an apparent atheist. These churches include the United Methodist Church.6% belong to churches that support evolution education". Sewall Wright. National Baptist Convention of America.[180] A poll in 2000 done for People for the American Way found that 70% of the American public felt that evolution was compatible with a belief in God. and believe in immortality.[174] It is commonly claimed that all proponents of evolutionary theory are "materialistic atheists". Very similar results were reported from a 1997 Gallup survey of the American public and scientists. one a practicing Unitarian. Evangelical Lutheran Church in America. the Roman Catholic Church.[177] H. S. about 40 percent believe in both evolution and an active deity (theistic evolution). notes that: Of the five founding fathers of twentieth-century evolutionary biology — Ronald Fisher. B. professor of biology at University of Rochester. however. the Episcopal Church. J. Allen Orr.Objections to evolution 376 Atheism Another charge leveled at evolutionary theory by creationists is that belief in evolution is either tantamount to atheism. Philosopher Robert Pennock makes the comparison that evolution is no more atheistic than plumbing.[175] On the other hand. Young asks. Only 48% of the people polled could choose the correct definition of evolution from a list. African Methodist Episcopal Church. or nonreligious theists. National Baptist Convention USA. a wide range of religions have reconciled a belief in a supernatural being with evolution. Ernst Mayr. Haldane. and others.[182] This is similar to the results reported for surveys of the general American public.[184] Group [184] Belief in Young Earth Creationism 44% 5% Belief in God-guided evolution 39% 40% Belief in evolution without God guiding the process 10% 55% American public American scientists . agnostics. 89.[181] One poll reported in the journal Nature showed that among American scientists (across various disciplines). about 40 percent of the scientists polled believe in a God that answers prayers. evolution neither requires nor rules out the existence of a supernatural being. atheism is far from universal among scientists who support evolution. or conducive to atheism. or among the general public that supports evolution. one a dabbler in Eastern mysticism. Davis Young argues that Creation Science itself is harmful to Christianity because its bad science will turn more away than it recruits.[178] In addition. Evolution: The History of an Idea. 900-worldwide-creationism-shotgun-stunner-and-more.Objections to evolution 377 See also • Faith and rationality Notes [1] Johnston. ca/ ~johnstoi/ darwin/ title. com/ article/ mg20327153. cfm?id=is-the-human-race-evolvin). [6] Sarfati. Laurence (1993). ewtn. 2006. "Charles Darwin: gentleman naturalist: A biographical sketch" (http:/ / darwin-online. html) (html). Retrieved 2010-05-24. [20] Bowler. html). talkorigins. html) (HTML).). Peter J. Completely Revised and Expanded. Shotgun stunner. ISBN 0-393-30154-0. GSA Today 17 (1): 4–9. pp. com/ about-us#who_we_are) (html). html) [25] John Paul II. Retrieved 2010-05-24. . Retrieved 2010-06-14. P. "What is Evolution?" (http:/ / www. uk/ darwin. aaas. p. talkorigins. John (2002-7). org/ cgi-bin/ templatefiller/ ?type=& id=) (html). . PJ (2003). [14] Carroll SB (2001). The eclipse of Darwinism: anti-Darwinian evolution theories in the decades around 1900 (paperback ed. htm#LECTURE_IV) Lecture IV of Eight Lectures Preached Before the University of Oxford in 1884 [24] Pius XII. F.1130/GSAT01701A. org/ files/ 17194/ 17194-h/ 17194-h. p. 10. "Section Three: The Origins of Evolutionary Theory" (http:/ / records. Liberal Studies Department. historian Ronald Numbers traces the religious motivations and scientific pretensions. [11] "Opinions on evolution from ten countries | NCSE" (http:/ / ncse. Retrieved 2007-03-24. James R. gutenberg. [8] Abdul Majid (2002). scientificamerican. Whitcomb. New Scientist. Ian C. (January 2007). . National Center for Science Education. [9] "Worldwide creationism. Scientists Confront Creationism. asp). com/ timeline/ November. . html). . Jonathan & Matthews. Harry Rimmer. html) [18] England. [16] Doolan. Nature 409 (6823): 1102–9. ISBN 978-0520236936. answersingenesis. Retrieved 2010-05-24. "Oh! My aching wisdom teeth!" (http:/ / www. 1998. "The Muslim Responses To Evolution" (http:/ / diberri. vatican. Retrieved 2007-07-25. And Still We Evolve. talkorigins. ISBN 0-8018-4391-X. [13] "Ask the experts:Biology-Is the human race evolving or devolving?" (http:/ / www. B. org/ indexcc/ CA/ CA212. (1907). com/ library/ PAPALDOC/ JP961022. org/ creation/ v18/ i3/ wisdom_teeth. Message to the Pontifical Academy of Sciences on Evolution (http:/ / www. American Association for the Advancement of Science. 624 pages. interacademies. . Expanded Edition. Cambridge University Press. University of Cambridge. Retrieved 2010-06-07. org/ news/ releases/ 2006/ pdf/ 0219boardstatement. The Complete Work of Charles Darwin Online. Numbers. Pg 8. va/ holy_father/ pius_xii/ encyclicals/ documents/ hf_p-xii_enc_12081950_humani-generis_en... . Harvard University Press. Henry M. Retrieved 2010-05-24. . "On the ultimate disintegration products of the radio-active elements. The TalkOrigins Archive (http:/ / www. PMID 11234024. ISBN 0674023390. Retrieved 2007-03-24. Argument: Creationism is religion. . "John Perry's neglected critique of Kelvin's age for the Earth: A missed opportunity in geodynamics". org/ get-answers/ topic/ religion). com/ article. "Chance and necessity: the evolution of morphological complexity and diversity". University of California Press. .1038/35059227. Malaspina University College. Molnar. Johnson and the Intelligent design movement. [15] "CA212: Definition of evolution" (http:/ / www. The Post-Darwinian Controversies: A Study of the Protestant Struggle to Come to Terms with Darwin in Great Britain and America. [22] Moore. Answers in Genesis. newscientist. viu. The Creationists: From Scientific Creationism to Intelligent Design. American Journal of Science 23: 77–88. net/ Object. Third Edition. [23] The Relations Between Religion and Science by Frederick Temple (http:/ / www. answersingenesis. Baltimore: Johns Hopkins University Press. File/ Master/ 6/ 150/ Evolution statement. aboutdarwin. dyndns.. [5] Godfrey. P.1. [21] Bowler. org. talkorigins. [7] "Statement on the Teaching of Evolution" (http:/ / www. and more" (http:/ / www. Interacademy Panel [4] In his comprehensive treatise on Creationism. [17] AboutDarwin. [19] Boltwood. com/ news/ 2009/ 07/ opinions-evolution-from-ten-countries-004885) (html). of prominent creationists from George Frederick Wright through George McCready Price. Inc. doi:10. pdf) (PDF). Morris and his Institute for Creation Research (and lesser figures) to Phillip E.org.. org/ faqs/ evolution-definition. W. Righter. [10] "About Us" (http:/ / creation. Answers in Genesis. HTM) [26] Adnan Oktar (1999). . B. John C. com/ evolution_specialpreface. htm). The Creationists. Retrieved 2007-07-25. not science (http:/ / www. [12] Moran.. [2] van Wyhe. Robert (1996). Scientific American. Retrieved 2007-03-20.com (http:/ / www. . doi:10. 23–24. . Laurie R. (1983). Michael (2000). "The Evolution Deceit" (http:/ / www. Part II. . pdf). [3] IAP Statement on the Teaching of Evolution (http:/ / www. (1999). Norton & Company (1984). php) (html). org). harunyahya. 2006). Ronald (November 30. Retrieved 2007-03-24. encyclical Humani Generis (http:/ / www. The disintegration products of uranium". Islamic Research Foundation International. ISBN 9780521285179. W. (1981). "Teach the controversy" (http:/ / www. [53] Dembski. actionbioscience. org).Objections to evolution [27] "Darwin Correspondence Project – Darwin and design: historical essay" (http:/ / www. National Science Teachers Association Press (http:/ / www. org/ library/ gould_fact-and-theory. Induction and Decision Theory). Evan Ratliff. Retrieved 2010-05-04. org/ indexcc/ CA/ CA040. html) . ISBN 0393017168. ISBN 978-0521678674. Creationism: An Introduction. [32] "National Science Teachers Association Disappointed About Intelligent Design Comments Made by President Bush" (http:/ / www. . Matzke NJ (May 2007). Sci. html). "Biological design in science classrooms" (http:/ / www. David N. Douglas (2004). org/ docs/ meyer/ sm_teachthecontroversy. html)". Mark (2004). creationsafaris. talkorigins. Master Books. and the scientific method" (http:/ / www. Laurence (1993). org/ docs/ dembski/ wd_isidtestable. [51] "Was Darwin a Christian? Did he believe in God? Did he recant evolutionism when he died? . M (2004). [39] Isaak. Acad. . Retrieved 2010-05-01. Retrieved 2007-03-24. creationists. publicopinionpros. org/ features/ 2006/ aug/ bishop3. asp). talkorigins. Retrieved 2007-03-24. The Lie: Evolution (http:/ / www.Supp. ISBN 0-89051-158-6.1073/pnas. htm). Retrieved 2010-05-23. Retrieved 2007-03-24. 1255 (http:/ / www. talkorigins. [40] Gould. discovery.0701505104. Cambridge University Press. talkorigins. . October 2004. Retrieved 2007-03-24. [57] Kutschera U. asp). Wired magazine [45] Meyer. Chapter 2. com/ wired/ archive/ 12. [50] "World's Greatest Creation Scientists from Y1K to Y2K" (http:/ / www. answersingenesis. org/ indexcc/ CA/ CA611. The TalkOrigins Archive (http:/ / www. talkorigins. org/ faqs/ mclean-v-arkansas. . [47] Scott. WA (2006). [54] Morris. talkorigins. html). Mark (2001-09-24). National Poll Challenge Darwinism" (http:/ / www. org/ about/ pressroom. doi:10. Retrieved 2007-03-24. . org/ creation/ v21/ i4/ design. September 2001. [36] Ham. The TalkOrigins Archive (http:/ / www. entouch. darwinproject. talkorigins. [58] McLean v Arkansas Board of Education. Retrieved 2008-10-27. K (1987). talkorigins. Claim CA040: Equal time" (http:/ / www. Ken (2010). SC (2002). or a Law?" (http:/ / mall. Retrieved 2010-05-01. actionbioscience. . "Evolution: Fact and Theory" (http:/ / www. htm). Retrieved 2007-10-30. [49] Morton. Discovery Institute. pdf) (PDF). [48] "Blown away by design" (http:/ / www. The TalkOrigins Archive (http:/ / www. Proc. [43] Theobald. Hen's Teeth and Horse's Toes (http:/ / www. html). . Norton & Company. org/ evolutionisreligion. University of California Press. . ISBN 978-0890510025. ISBN 0520246500. html). [34] Edwards. christiananswers. "Evolution Is Religion—Not Science" (http:/ / www. [37] Moran. org/ faqs/ comdesc/ sciproof. Niklas KJ (June 2004). Retrieved 2007-03-24. scientific evidence. wired. Scientific Creationism. crisismagazine. . The Design Inference: Eliminating Chance through Small Probabilities (Cambridge Studies in Probability. "Five Major Misconceptions about Evolution" (http:/ / www. Mark (2003). "Separation of Christianity and State" (http:/ / www. htm). Discovery Institute. 529 F. html). GR (2002). com/ julaug2003/ feature1. org/ cgi/ pmidlookup?view=long& pmid=17494747). com/ press/ pressRelease_100Scientists. net/ C/ cs/ theory. com/ wgcs_4.org (http:/ / www. Retrieved 2010-06-16. org). "Does Science Point to God? Part II: The Christian Critics" (http:/ / www. Harris Interactive. . Retrieved 2007-03-24. [55] Wiker. . crisismagazine. [30] A copy of the Discovery Institutes Wedge Strategy document can be found here: "Wedge Strategy" (http:/ / www. [41] Lenski. BD (2003). [35] Dembski. RE (2000). org/ ). Retrieved 2007-03-25. . pnas. arn. uk/ content/ view/ 110/ 104/ ). W. [31] "Polls Apart on Human Origins" (http:/ / www. Retrieved 2007-03-24. "Index to Creationist Claims. antievolution. . htm). 1999. talkorigins. net/ dmd/ moreandmore. EC (2004). 253–262. HM (1985). " Index to Creationist Claims. [56] Isaak. Naturwissenschaften 91 (6): 255–76. org/ pdf/ imp/ imp-332. htm). ActionBioscience. org/ faqs/ faq-misconceptions. . org/ evolution/ lenski. Retrieved 2008-09-02. stephenjaygould. "100 Scientists. "Evolution is a Fact and a Theory" (http:/ / www.1007/s00114-004-0515-y. Impact: Vital Articles on Science/Creation 332.S. Menton. [29] "Evolution is Religion" (http:/ / www. "Is Intelligent Design Testable?" (http:/ / www. 104 Suppl 1: 8669–76. nsta. W. PMID 17494747. Retrieved 2010-05-04. . com/ ). U. pp. . html?pg=2). pdf) (pdf). htm) (html). . 378 . turnpike. Evolution vs. original "100 Scientists" advertisement. HM (2001). arn. Retrieved 2007-10-30. [46] Isaak. . Crisis Magazine (http:/ / www. Retrieved 2007-03-24. pdf) (PDF). The TalkOrigins Archive (http:/ / www. Natl. icr. . org). aspx?id=50794). [44] The Crusade Against Evolution (http:/ / www. 10/ evolution. Retrieved 2007-03-24. net/ q-aig/ darwin. [28] Scott EC. nsta. PMC 1876445. Claim CA611: Evolution Sacrosanct? (http:/ / www. Missouri Association for Creation. .ChristianAnswers. php) (php). "Is Evolution a Theory. See Evolution is Religion. "The modern theory of biological evolution: an expanded synthesis". Retrieved 2007-03-24. org/ evolutionism-is-a-religion. Cincinnati Enquirer. . html). answersingenesis. William (2001). html). [33] "A Scientific Dissent from Darwinism" (http:/ / www. html). org).A. a Fact. 2005. org). org/ articles/ au/ separation-of-christianity-and-state). org/ faqs/ evolution-fact. [42] Morris.Net" (http:/ / www. "29+ Evidences for Macroevolution: Scientific "Proof". . Master Books. html). org/ features/ wedge. . SJ (1994).. talkorigins. creationists. "The Imminent Demise of Evolution: The Longest Running Falsehood in Creationism" (http:/ / home. TalkOrigins Archive. Retrieved 2010-05-23. doi:10. PMID 15241603. [52] Ham. ac. . norc. org/ articleFiles/ PDFs/ 100ScientistsAd. reviewevolution. . [38] Dr. The TalkOrigins Archive (http:/ / www. 45. html). Retrieved 2008-05-10. Retrieved 2010-03-07. The TalkOrigins Archive (http:/ / www. pp. "Evolution and Philosophy: Is Evolution Science. org). p. html).00. org/ indexcc/ CA/ CA211. editor. doi:10.1120331. "Myth: Science is a Religion for Atheists that Requires Faith" (http:/ / atheism. org/ resource/ evol07. Henry M. org/ faqs/ evolphil/ tautology. . "Evolution and Philosophy: A Good Tautology is Hard to Find" (http:/ / www. [61] Scientific Creationism. Retrieved 2007-03-24. talkorigins. Retrieved 2007-03-24. . 189. Rev. doi:10.1746-8361. talkorigins. html#part5). The TalkOrigins Archive (http:/ / www. Genet. [92] Darwin. [75] See Survival of the fittest for a more thorough discussion.V. J (2006). Dialectica 32 (32): 339. newscientist. [77] Hempel. and What Does 'Science' Mean?" (http:/ / www. Retrieved 2007-03-25. [69] Wallis.1038/nrg1088.tb01321. "The pectoral fin of Tiktaalik roseae and the origin of the tetrapod limb". [70] "Human Chromosome 2" (http:/ / www. genconnect/ ). [73] Wilkins.1038/nature07892. USA: Creation Science Evangelism. [84] Elena SF. talkorigins.G. html). M (2003). [88] Wilkins. talkorigins.x. talkorigins. doi:10. html). talkorigins. html). uk/ content/ frameset?viewtype=text& itemID=F385& pageseq=391) 379 . Blackwell Publishing Limited. 359–360 (http:/ / darwin-online. Abusing science: the case against creationism. "Evolution experiments with microorganisms: the dynamics and genetic bases of adaptation". com/ od/ atheismscienceevolution/ a/ ScienceFaith. The Dangers of Evolution. talkorigins. "The Evolution Wars" (http:/ / www. org). Brockhurst MA. Craig Maclean R. [63] "Expelled Exposed: Why Expelled Flunks » Science & Religion" (http:/ / www. Jenkins FA (April 2006). "Chimps are human. org). expelledexposed. 2006. Retrieved 2007-03-25. The TalkOrigins Archive (http:/ / www. Open Court. Morris. org/ faqs/ faq-transitional. Retrieved 2007-03-24. talkorigins. (2006). ISBN 978-1405103459. A (2006). Cambridge: Harvard University Press [78] Kitcher. Lenski RE (June 2003).1090909.1126/science. talkorigins. "Transitional Vertebrate Fossils FAQ" (http:/ / www. TalkOrigins. .1111/j. Arkansas. org/ faqs/ macroevolution. Philosophy and History" (http:/ / www. [60] TalkOrigins Claim CA211 (http:/ / www. K (1985). . org/ FAQ. org/ Home/ Area/ faq/ dont_use. Kansas evolution hearings [87] Boxhorn. . Pohl B. 52 [81] Kitcher 1982 p. p. [66] Misquoted Scientists Respond (http:/ / ncse. Daeschler EB. Retrieved 2008-05-10.com. teachersdomain. [71] Jeff Hecht (2003-05-19). "Observed Instances of Speciation" (http:/ / www. Nature 457 (7231): 824–9. com/ cej/ 2/ 4/ misquoted-scientists-respond) National Center for Science Education 1981 by John R. or the Preservation of Favoured Races in the Struggle for Life. C (2005-08-07). [85] Questions frequently asked about the TBSEF (http:/ / tbsef. Quine. .1978. html). evo. Peters DS (December 2005). p. Unended Quest: An Intellectual Autobiography. 50 [80] Kitcher 1982 p. life. C. Cole quoting Popper: "I have changed my mind about the testability and logical status of the theory of natural selection. The TalkOrigins Archive (http:/ / www. sci. Retrieved 2007-03-24.O 1952 “Two Dogmas of Empiricism” reprinted in From a Logical Point of View. org). [89] Mayr G. [72] Jim Foley. com/ index. . PMID 19212400. and I am glad to have the opportunity to make a recantation. Science (journal) 310 (5753): 1483–6. Answers in Genesis. talkorigins. Cambridge. org/ faqs/ kansas/ kangaroo10. The TalkOrigins Archive (http:/ / www. html). . . Index to Creationist Claims. com/ time/ archive/ preview/ 0. "Fossil Hominids: The Evidence for Human Evolution" (http:/ / www. Nat. On the Origin of Species by Means of Natural Selection. [76] MacRae. 4 (6): 457–69. PMID 16598250. JS (1997). about. Texans for Better Science Education Foundation [86] Kansas Evolution Hearings Part 10 (http:/ / www. org). John Murray. .Objections to evolution [59] Cline. talkorigins. about. "A well-preserved Archaeopteryx specimen with theropod features". [79] Kitcher 1982 p. Mark Isaak. Evolution. Mass: MIT Press. C (1859). W. A (1998). Joseph (1995). ISBN 0-262-61037-X. . PMID 16322455. [DVD]. [90] Shubin NH. 32. "Natural selection and the emergence of mind". Philip (1982). Retrieved 2007-03-24. 4th edition of 1866. 1974 Master Books. asp). [74] "Arguments we think creationists should NOT use" (http:/ / www. Third Edition. ISBN 978-0087583436. html). doi:10. JS (1997). NewScientist. org/ faqs/ evolphil/ falsify. answersingenesis. "The Beagle in a bottle". PMID 12776215. p." [67] Darwin. Retrieved on 2008-04-20. 52-53 [82] Kent Hovind. Retrieved 2007-03-24. [65] Popper. 2002–2008 WGBH Educational Foundation. org/ faqs/ faq-speciation. Charles (1859). National Center for Science Education. [64] Popper. talkorigins. htm). talkorigins. Retrieved 2008-12-05. [83] Buckling A.1038/nature04637. uk/ content/ frameset?viewtype=text& itemID=F373& pageseq=298). . time. Retrieved 2007-03-24. htm#Is TBSEF against teaching evolution?). talkorigins. "Radiometric dating and the geological time scale: Circular reasoning or reliable tools" (http:/ / www. "Macroevolution: Its Definition. org). org. Nature 440 (7085): 764–71. org. org/ faqs/ homs/ ). com/ article/ dn3744-chimps-are-human-gene-study-implies. pp. gene study implies" (http:/ / www. 2006. [68] Ridley. 280–313 (http:/ / darwin-online. . talkorigins. org/ faqs/ dating. 6-7 [62] Wilkins. doi:10. html). php/ the-truth/ science-religion). K (1978). [91] Hunt (1997). On the Origin of Species. Colegrave N (February 2009). Time Magazine.10987. Glencoe: the Free Press. 1951 “Problems and Changes in the Empiricist Criterion of Meaning” in Aspects of Scientific Explanation. Science 276 (5310): 227–32. html) (HTML). com/ creationism/ analysis/ icon-4-haeckels-embryos). . The TalkOrigins Archive (http:/ / www. htm). html). org). html). org/ indexcc/ CC/ CC200. org/ docs/ johnson/ pjdogma1. talkorigins. "Variation. M (2005). Retrieved 2009-12-05. National Center for Science Education. org/ faqs/ archaeopteryx/ forgery. . Damned Lies. . org/ docs/ behe/ mb_dm11496. . and Probability of Abiogenesis Calculations (http:/ / www. Retrieved 2007-03-24. doi:10. D (1995). . doi:10. . talkorigins. The TalkOrigins Archive (http:/ / www. The TalkOrigins Archive (http:/ / www. . . Inc. Icons of Evolution. talkorigins. [108] Isaak. The TalkOrigins Archive (http:/ / www. The TalkOrigins Archive (http:/ / www. "Index to Creationist Claims. M (2004). November 23. "Haeckel’s ABC of evolution and development. . talkorigins. Retrieved 2008-07-13. p. The TalkOrigins Archive (http:/ / www. B. Retrieved 2007-03-24. [96] Ham. 2006. "On Archaeopteryx. Retrieved 2007-03-24. html) (PDF).227. . talkorigins. [105] "Icons of Evolution FAQs" (http:/ / www. Answers in Genesis. Retrieved 2007-03-24. html). . Brinkmann H. "Missing links still missing!" (http:/ / www. [107] "Icon 4 — Haeckel's Embryos" (http:/ / ncse. The TalkOrigins Archive (http:/ / www. . M (2004). org/ indexcc/ CA/ CA110. [104] Wells. 516 [103] Nedin.0001. P (1990). info/ gallery/ gtext3." (http:/ / www. html). Retrieved 2008-12-17. Oxford University Press. [102] Richardson and Keuck. C (1991). MJ (1996-10-29). CC201: Phyletic gradualism" (http:/ / www. org/ indexcc/ CC/ CC201. org/ indexcc/ CC/ CC363. Retrieved 2010-06-07. talkorigins. [99] Delsuc F. talkorigins. org/ origins/ postmonth/ feb98. [100] Einstein. talkorigins. [119] Fitelson. .. "Plantinga's Probability Arguments Against Evolutionary Naturalism" (http:/ / fitelson. talkorigins. "Evolution and Chance" (http:/ / www. org). 2006. CC340: Out-of-place fossils" (http:/ / www. Retrieved 2007-03-24. talkorigins. org/ faqs/ wells/ ). Retrieved 2007-03-24. org). . [98] Huelsenbeck JP. . "Index to Creationist Claims. The TalkOrigins Archive (http:/ / www. talkorigins. "Index to Creationist Claims. talkorigins. J (2002). M (2004). Retrieved 2007-03-24. org).+ information+ and+ the+ created+ kind& as_publication=Journal+ of+ Creation& as_ylo=1991& as_yhi=1991& btnG=Search)). talkorigins. [110] Isaak.001. Retrieved 2007-03-24. Statistics.1038/nrg1603.1093/0195078640. . 380 . [114] Wilkins. M (2004).276. org/ faqs/ chance/ chance. htm). Claim CD010: Radiometric Dating" (http:/ / www. . html). talkorigins. M (2004). [111] Isaak. information and the created kind" (http:/ / creation. C (1997). Claim CC363: Requirements for fossilization" (http:/ / www. . org/ docs2004/ 0806faithful. talkorigins. "The Foundation of the General Theory of Relativity" (http:/ / www. [94] "A faithful man takes on faith-based teaching" (http:/ / www. talkorigins. Ken (1989). co. uk/ scholar?hl=en& lr=& q=author:Wieland+ intitle:Variation. [124] "CB400: Evolution of consciousness" (http:/ / www. [118] Plantinga. html). [101] Isaak. "Index to Creationist Claims. "Phylogenetic methods come of age: testing hypotheses in an evolutionary context"." (http:/ / www. html). org/ indexcc/ CI/ CI100. "Phylogenomics and the reconstruction of the tree of life". The TalkOrigins Archive (http:/ / www. CC200. M (2005). Answers in Genesis. Retrieved 2008-07-13. Nat. com/ variation-information-and-the-created-kind) ( – Scholar search (http:/ / scholar. html). alberteinstein. org). [113] Isaak. . "Cheating with chance" (http:/ / creation.5310. Retrieved 2007-03-24. html) [117] "CI100: Intelligent Design" (http:/ / www. M (2004). html). [115] Battern. Annalen der Physik 49: 769–822. [116] Lies. doi:10. WR (1998). Regnery Publishing. Journal of Creation 5 (1): 42–47. pdf) (PDF). org). Retrieved 2009-12-05. . Retrieved 2008-07-13. com/ cheating-with-chance). . [122] Johnson. "Evolution as dogma: The establishment of naturalism" (http:/ / www. Claim CA120: Mind's fallibility" (http:/ / www. Retrieved 2007-03-24. google.Objections to evolution [93] Elsberry. talkorigins. Retrieved 2007-03-24. talkorigins. The TalkOrigins Archive (http:/ / www. 6 (5): 361–75. org/ indexcc/ CB/ CB400. [106] "CB701: Haeckel's embryo pictures. Retrieved 2006-09-03. talkorigins. CC200: Transitional fossils" (http:/ / www. The TalkOrigins Archive (http:/ / www. "Were You There?" (http:/ / www. org). 25. Albert (1916). Retrieved 2007-03-24. org). org/ indexcc/ CA/ CA221. talkorigins. php?module=articles& action=view& ID=670). arn. Genet. org/ indexcc/ CC/ CC200_1. "Darwin under the microscope" (http:/ / www. [120] Isaak. org/ indexcc/ CD/ CD010. talkorigins. The TalkOrigins Archive (http:/ / www. Institute for Creation Research. talkorigins. PMID 15861208. A (1993). html). M (2004). org/ faqs/ abioprob/ abioprob. Retrieved 2007-03-24.1126/science. talkorigins." p. . arn. [123] "CB401: Inconceivable instinct" (http:/ / www. talkorigins. org/ indexcc/ CB/ CB401. Retrieved 2008-07-13. Retrieved 2007-03-24. . Rev. . org). talkorigins. "Index to Creationist Claims. org/ index. Claim CA110: Evolution will soon be widely rejected. "Index to Creationist Claims. html). org). Philippe H (May 2005). Rannala B (April 1997). The TalkOrigins Archive (http:/ / www.org. [97] Isaak. [112] Isaak. . talkorigins. org). org/ plant.1: Transitional fossil abundance" (http:/ / www. talkorigins. [121] Behe. Retrieved 2007-03-24. and Forgery" (http:/ / www. html). Warrant and Proper Function. Astronomers. talkorigins. org). Retrieved 2009-12-06. talkorigins. ISBN 978-0895262004. [109] Isaak. [95] Wieland. talkorigins. J (1997). org). org/ indexcc/ CB/ CB701. New York Times. talkorigins. E (1997). html). Sober. ISBN 0-19-507864-0. html). org/ indexcc/ CC/ CC340. org/ indexcc/ CA/ CA120. asp). "Index to Creationist Claims. . icr. "Index to Creationist Claims. answersingenesis. Claim CA221: Were you there?" (http:/ / www. html). talkorigins. "Index to Creationist Claims. PMID 9092465. html). PNAS 104 (17): 7116–7121. The TalkOrigins Archive (http:/ / www. H (1982). math. org/ indexcc/ CB/ CB921_2. uk/ Genesis/ results of believing evolution. [130] Behe. p. . lwbc. and references therein [134] Ussery. [137] Sanz. co. talkorigins. [156] Ham. org/ docs/ behe/ mb_philosophicalobjectionsresponse.0700266104.. [154] Mohler. The Troubled Waters of Evolution.1038/ng1482. . uk/ content/ frameset?viewtype=image& itemID=CUL-DAR141. edu/ ~rosenhjd/ sewell. "Evolution and Information: The Nylon Bug" (http:/ / www. . The TalkOrigins Archive (http:/ / www. The TalkOrigins Archive (http:/ / www. talkorigins. Bunn. co. past and future" (http:/ / www. PE/ 0510007). [132] Liu. talkorigins. M (2006). doi:10. M (2004). D. Baldwin. 2003 Refuting Evolution 2. org). Kevin Knight. . asp). Mr." American Journal of Physics. Vol. Journal of Heredity 96 (3): 171–184. (2007). [151] Churches urged to challenge Intelligent Design (http:/ / www. I told him most clearly and positively to draw me a life-like portrait of that profound philosopher. ISBN 978-0743290319. [142] "CB101: Most mutations harmful?" (http:/ / www. "Truth Cannot Contradict Truth". theTalkOrigins Archive (http:/ / www. nih. org. talkorigins. "New Perspectives on Eye Development and the Evolution of Eyes and Photoreceptors". "Darwin's Black Box: Irreducible Complexity or Irreproducible Irreducibility?" (http:/ / www. [139] Musgrave. [144] Lambert. shtml) [152] Pope John Paul II. Ochman.2: Half a wing" (http:/ / www.newadvent. Retrieved 2007-03-24. [138] Gitt. nmsr. com/ loom/ archives/ 2005/ 02/ 15/ eyes_part_one_opening_up_the_russian_doll. talkorigins. . co.C.htm> [153] Morris. Claim CB200: Irreducible complexity" (http:/ / www. et al. [141] Bergstrom. [133] Isaak. October 2009. Vol. Retrieved 2007-03-24. "Claim CA009: Evolution teaches that people are animals. org/ home/ area/ WWTL/ index.J.D. "Index to Creationist Claims. answersingenesis. Retrieved 2007-03-24. org/ faqs/ behe/ review. entropysite. A. . . I. npr. org/ tj/ v10/ i2/ information. 11. org). (2005). cbs. org). talkorigins. html). [143] Harter. [149] A venerable Orang-utang (http:/ / darwin-online." – The Hornet. html). Gaidukov. . "Does Life On Earth Violate the Second Law of Thermodynamics?" (http:/ / physics. [126] Jonathan Sarfati. [136] Robison. talkorigins. doi:10. uk/ index. com/ cracked_crutch. K (2002). talkorigins. Living Word Bible Church (http:/ / www. "A biochemist's response to "The biochemical challenge to evolution"" (http:/ / www. Retrieved 2010-01-02.. 1031 [147] Emory F. "The fitness value of information" (http:/ / arxiv. S. W. Roodveldt. lwbc. [155] "The Result of Believing Evolution" (http:/ / www. org). . Retrieved 2010-05-30.1021/ed079p187. php?storyId=4760816). talkorigins. C. Gould. "Index to Creationist Claims. Retrieved 2007-03-24. Teorema 28: 173–188. Tawfik. html) (HTML). Retrieved 2007-03-24. htm). ISBN 978-0890510872. (2009). talkorigins. (2005). dtu. Styer. PMID 15653558. . Khersonsky. Lachmann. arn. gov/ articlerender. Retrieved 2007-03-26. [135] Aharoni. org). 10." American Journal of Physics.. M (2005). Retrieved 2007-03-24. Retrieved 2007-09-22. Retrieved 2007-03-24. The TalkOrigins Archive (http:/ / www. RN (2006).. edu/ ~roerter/ EvolutionEntropy. "Disorder — A Cracked Crutch For Supporting Entropy Discussions" (http:/ / www. htm). "Stepwise formation of the bacterial flagellar system" (http:/ / www.. talkorigins. . The Mathematical Intelligencer 23 (4): 3–8. Answers in Genesis. html). org/ nylon. "The 'evolvability' of promiscuous protein functions". doi:10.S.1073/pnas. No. "Information Theory and Creationism" (http:/ / www. MJ (1996). Retrieved 2007-03-24. talkorigins. . dk/ staff/ dave/ Behe. p. J (2001). org/ indexcc/ CE/ CE440. Retrieved 2007-03-24.. html) (html). talkorigins. Retrieved 2007-03-24. F (2002). The Loom: A blog about life. 77. "The Origins of Life: An Evangelical Baptist View" (http:/ / www. 76. answersingenesis. Information. R.1093/jhered/esi027. ekklesia. htm). Journal of Chemical Education 79: 187–192. Retrieved 2007-03-24.Objections to evolution [125] Isaak. We should not be surprised when people who are taught evolution start behaving like animals" (http:/ / www. [128] Gehring. "How Anti-Evolutionists Abuse Mathematics" (http:/ / www. . [145] Oerter. ed. Bios 70: 40–45. pdf) (PDF). [140] Thomas. org/ faqs/ mutations. Part One: Opening Up the Russian Doll. from the collection of Darwin Online [150] Isaak. Retrieved 2007-03-24. npr. html). Responses of biochemistry to intelligent design".org/library/docs_jp02tc. Nature Genetics 37 (1): 73–76. (1999). science and biology (http:/ / www. 922 [148] Rosenhouse. "Entropy and evolution. Ph. The TalkOrigins Archive (http:/ / www.org. ISBN 0890513783. "Evolution and the Second Lqw of Thermodynamics. 381 . talkorigins. M (2004). htm).. doi:10.org. PMID 17438286. 2009-Feb-15 <http://www. Michael (2000-07-31). htm). "Philosophical Objections to Intelligent Design: Response to Critics" (http:/ / www. org/ pdf/ q-bio. PMID 15568024. NPR (http:/ / www. html). html). . PMC 1852327. R (1999). org/ indexcc/ CB/ CB200. New Advent. jmu. pubmedcentral. RA (2005). 5& pageseq=1) "I have to apologize once more for the wild flights of my incorrigible artist. [131] Behe. . D.. . org). talkorigins. No. K (1996). . org). gmu. corante. org/ indexcc/ CA/ CA009. Why Won't They Listen? A Radical New Approach to Evangelism (http:/ / www. fcgi?tool=pmcentrez& artid=1852327). . H. Darwin's Black Box: The Biochemical Challenge to Evolution. asp). 1871. November 2008. Free Press. Retrieved 2010-06-07. Claim CE440: The origin of it all" (http:/ / www. R. [146] Daniel F. CT. Darwin. D. "Darwin's golden flame. (2005). "Are Mutations Harmful?" (http:/ / www. org/ indexcc/ CB/ CB101. Werner (1996). uk/ content/ news_syndication/ article_060220creationism. Master Books. php).Y. New Mexicans for Science and Reason. org/ faqs/ information/ infotheory. org/ templates/ story/ story. Master Books ISBN 0-89051-387-2 [127] "CB921. [129] "Eyes. L. V. . html). O. talkorigins. Master Books.M. org/ archive/ 8/ 252006b. Politics. . Social Darwinists vigorously advocated the acquisition of empires. To these elitists. 32. 2006. com/ books?id=kKGgoNo4un0C& pg=PA653& lpg=PA653). CEN Technical Journal. The Case for a Creator. com/ 200605/ 20060507_1. some nations were deemed more fit than others for the competition. A. "Darwin on the Right: Why Christians and conservatives should accept evolution" (http:/ / www. and Racism in Germany. barna. J (2006). Everyday Life: Says outlook on Genesis account affects every aspect of life (http:/ / www. In the popular mind. Jonathan Sarfati. Retrieved 2007–03–25. agapepress. Right Wing Watch. adl. org/ 2006/ 08/ kennedy_evoluti. December 15. html). org). Hopelessness in World" (http:/ / www. adl. Retrieved 2007-03-24. [162] Weikart. The Barna Group (http:/ / www. Retrieved 2007-03-24. GS (2005). . Retrieved 2004-03-24. Hector Avalos. C (1999-09-06). answersingenesis. Even among Europeans. had wide currency in the Western states. 382 [165] "Anti-Evolution Film Misappropriates the Holocaust" (http:/ / www. 1999.date=Feb+ 06. pqarchiver. evolutionnews. The Holocaust and evolution (http:/ / creation. org/ indexcc/ CA/ CA006_1. an attitude that sparked their competitive enthusiasm. Scientific American. Usually. com/ globe/ search/ stories/ reprints/ darwin100199. darwintohitler. [160] Martin. created 2001-4-29. Dakota Voice. Evolution News and Views. Jerry Bergman. ISBN 0-89051-291-4. [168] "Born Again Christians Just As Likely to Divorce As Are Non-Christians" (http:/ / www. creighton. . In the nineteenth century. From Darwin to Hitler: Evolutionary Ethics. Creation 22(1):4. adl. org/ barna-update/ article/ 5-barna-update/ 194-born-again-christians-just-as-likely-to-divorce-as-are-non-christians). rightwingwatch. 2008 [171] Talkorigins Claim CA006. Agape Press (http:/ / headlines. Smith. Retrieved 2007-04-26. Jonathan Witt. Index to Creationist Claims. cfm?chanID=sa006& articleID=00068F43-E189-150E-A18983414B7F0000& colID=13). html?FMT=ABS& amp. cfm). Palgrave Macmillan. org/ PresRele/ HolNa_52/ 5277_52. org/ 2006/ 12/ from_darwin_to_hitler_a_straig. barna. Retrieved 2009-12-07. except for missionaries. The paper was criticized by Moreno-Riaño. html#more). "TV Producer Defends Documentary Exposing Darwin-Hitler Link" (http:/ / headlines. Eugenics. and Paul does not produce any speculations about the cause of these correlations. Talkreason [173] "Western Civilization: Ideas. 2006. and Society" (http:/ / books. HM (1989). The Long War Against God: The History and Impact of the Creation/Evolution Conflict. html). . google. in contrast to the seventeenth and eighteenth centuries. dakotavoice. 2004. htm). edu/ History/ Faculty/ Weikart/ FromDarwintoHitler. From Darwin to Hitler: Evolutionary Ethics. Missionaries were the first to meet and learn about many peoples and were the first to develop writing for those without a written language. Retrieved 2009-12-07. org/ religious_freedom/ resource_kit/ intelligent_design. htm). For example: • • • Darwinism and the Nazi race Holocaust (http:/ / www. . 5/7/2006 [167] Paul. This language of race and conflict. . those that were successful at expanding industry and empire-would survive and that others would not. "Intelligent Design: It's Not Science – Religious Freedom Resources" (http:/ / www. com/ the-holocaust-and-evolution). They had little sense that other cultures and other people had merit or deserved respect. Lee (2004). Michael Ruse. edu/ JRS/ 2005/ 2005-11. [169] Shermer. "In my quest to determine if contemporary science points toward or away from God. Discovery Institute. org/ articles/ Genocide. [158] Morris." [174] Strobel. asp). Retrieved 2009-12-07. 2009. org/ tj/ v13/ i2/ nazi. Anti-Defamation League. Christian missionaries were ardently opposed to slavery. [161] "ADL Blasts Christian Supremacist TV Special & Book Blaming Darwin For Hitler" (http:/ / www. html) because "[Paul's] methodological problems do not allow for any conclusive statement to be advanced regarding the various hypotheses Paul seeks to demonstrate or falsify. saying that strong nations-by definition. December 1999. modified 2005-7-1. I knew I had to first examine the claims of evolution in order to conclude once and for all whether . talkorigins. com/ tallahassee/ access/ 1715211751. rarely adopted the customs or learned the languages of local people. [159] "Kennedy: Evolution to Blame for Death. 13(2):101–111. August 22.+ 2008). 2004. com/ ). Parker. edu/ JRS/ 2006/ 2006-1. © 2006 [172] Creationists for Genocide (http:/ / www.1: Adolf Hitler exploited the racist ideas of Darwinism to justify genocide (http:/ / www. asp). html). org/ PresRele/ HolNa_52/ 4877_52." Of course. the concepts of evolution justified the exploitation of "lesser breeds without the law" by superior races. Social Darwinists thought their own nation the best. agapepress. . p. . all white men were more fit than non-whites to prevail in the struggle for dominance. Retrieved 2007-03-24.Objections to evolution [157] Raymo. html). Talkorigins. htm). boston. Retrieved 2007-03-24. sciam. htm). My View. Palgrave MacMillan. and Racism in Germany (http:/ / web. Richard Weikart. . correlation does not imply causality. M (2006). Boston Globe. "Cross-National Correlations of Quantifiable Societal Health with Popular Religiosity and Secularism in the Prosperous Democracies: A First Look" (http:/ / moses. [163] From Darwin to Hitler: A Pathway to Horror (Updated) (http:/ / www. of superior and inferior people. April 29. "Darwin's Dangerous De-evolution" (http:/ / www. Discovery Institute) and is amply repeated in creationist literature. and Mach in a published article in the same journal (http:/ / moses. Zondervan. Eugenics. 2008. Many westerners believed that it was their duty as Christians to set an example and to educate others. . Mark Isaak. org). com/ article. February 6. asp). ISBN 978-1403972019. talkreason. R (2004). [164] This creationist claim that is part of a Discovery Institute campaign ( New book by Discovery Institute Fellow shows influence of Darwinian principles on Hitler's Nazi regime (http:/ / www. Anti-Defamation League. Baker Book House. [166] Creationist Links Origins to Faith. Bob Ellis. Journal of Religion & Society 7. Europeans. Anti-Defamation League. creighton. [170] Darwin and Hitler: a not-very-intelligent link (http:/ / pqasb. Tallahassee Democrat. csustan. ISBN 0310241448. Houghton Mifflin Harcourt Publishing Company. "The most extreme ideological expression of nationalism and imperialism was Social Darwinism. . BA (1995). . com/ viewer?a=v& q=cache:UDwdl6q3IgwJ:67. Ph. ISBN 075460912X. . Connecticut: Greenwood Press. Leslie Carlin (2004). "15 answers to creationist nonsense" (http://www. Retrieved 2007-03-24. "The Church of Darwin" (http:/ / www. doi:10. 2002. Artisan Publishers. Wichita Eagle. Tower of Babel: The Evidence Against the New Creationism (http:/ / ncse. The Counter-Creationism Handbook. D (1988). pdf+ evolutionandcreationisminpubliceducation. • Isaak. Retrieved 2007-03-24. org/ ev_publi. "Scientists are still keeping the faith". Allen Orr. org/ docs/ johnson/ chofdarwin. com/ rncse/ 17/ 6/ many-scientists-see-gods-hand-evolution). 17A. arn. emporia." [175] Johnson. com/ archive/ 2005/ 05/ 30/ 050530fa_fact?currentPage=1). ISBN 978-0262661652. "Public beliefs about evolution and creation" (http:/ / www. [180] Schrock. [177] Pennock. . "Many scientists see God's hand in evolution" (http:/ / ncse. Am. pdf& hl=en& gl=ca& pid=bl& srcid=ADGEESj-27vCwtszKiottg8mjbOKovsUvtJqRXXlLit-R3VitD5tlX4OfMPkfEEzNwQjvUSioQSua6y3LN1SR72877bW3dc6wFzFlG2pFdWl-21CVU68FW sig=AHIEtbQjqyUehD_pbin4wiuZlHKkn--pTA) (PDF). 287 (1): 78–85. L (1997). PMID 12085506. EJ.pdf) (PDF). L (1997). Reports of the National Center for Science Education 17 (6): 33. [183] Witham.1038/scientificamerican0702-78. 238. ISBN 093466627X. pp. . Retrieved 2007-03-24. [184] Robinson. Master Books ISBN 0-89051-387-2 • Coleman. MIT Press. 330. 2002. RT (2000). Christianity and the Age of the Earth. htm). com/ store/ title/ tower-babel). [178] Devolution: Why intelligent design isn’t (http:/ / www. ISBN 031333305X. [182] Larson. [176] Young. Retrieved 2007-03-24. Burlington. Witham. H. religioustolerance. Wall Street Journal. [181] "Evolution and Creationism In Public Education: An In-depth Reading Of Public Opinion" (http:/ / docs. May 30. JR (2005-05-17). Sci. . Retrieved 2010-05-23. then the atheist conclusions I reached as a student might still be valid. • Rennie J (July 2002). htm).Objections to evolution 383 Darwinism creates a reasonable foundation for atheism.D. Cambridge: The MIT Press • Jonathan Sarfati. Vermont: Ashgate. National Center for Science Education. p. com/ media/ voices/ religion). Simon. The Cultures of Creationism: Antievolution in English-speaking Countries. edu/ biosci/ schrock/ docs/ Eagle-25.1038/386435a0. . That's because if the materialism of Darwinian evolution is a fact. Nature 386: 435–436. 2003 Refuting Evolution 2. newyorker. Annals of Science. Westport. 195. Further reading • Philip Kitcher 1982 Abusing Science: The Case Against Creationism. Evolution Not in Conflict" (http:/ / www. [179] "Statements from Religious Organizations" (http:/ / ncse. Retrieved 2007-03-24. Mark (2005). "Christianity. 2005. google. The New Yorker. . Phillip (1999). 59/ multimedia/ pdf/ Reports/ evolutionandcreationisminpubliceducation. 192. Retrieved 2007-03-24.swarthmore.edu/NatSci/cpurrin1/ textbookdisclaimers/wackononsense. People for the American Way. doi:10. pdf) (PDF). p. creationism in America has contested scientific theories.[23] After the legal judgment of the case Daniel v.[19] [20] [21] [22] History The history of creationism is part of the history of religions. Aguillard (1987) ruled that creation science similarly contravened the constitution.[5] [6] which derive from natural observations of the universe and life. or to a concept about the origin of the soul.[10] [11] [12] [13] while different members of individual faiths vary in their acceptance of scientific findings. and complexity of life on Earth. diversity. creationists often reject the conclusions of the research[15] or its underlying scientific theories[16] or its methodology. which was subsequently claimed to be a new scientific theory. Dover (2005) ruling concluded that intelligent design is not science and contravenes the constitutional restriction on teaching religion in public school science classes. In the 1920s the term became particularly associated with Christian fundamentalist movements that insisted on a literalist interpretation of the Genesis creation narrative and likewise opposed the idea of human evolution. the term "anti-evolutionists" became more common. then in 1929 in the United States the term "creationism" first became specifically associated with Christian fundamentalist opposition to human evolution and belief in a young Earth. in particular much of evolution. Waters (1975) ruled that teaching creationism in public schools contravened the Establishment Clause of the First Amendment. For the movement in Spanish literature. As the creation–evolution controversy developed. the formation of the solar system and the origin of the universe. all references to "creation" in a draft school textbook were changed to refer to intelligent design.[7] [9] Other religions have different deity-led creation myths. see Creacionismo.[3] At this time those holding that species had been separately created were generally called "advocates of creation" but they were occasionally called "creationists" in private correspondence between Charles Darwin and his friends.[18] The most notable disputes concern the evolution of living organisms.[24] . the Earth. and the universe are the creation of a supernatural being. the term is more commonly used to refer to religiously motivated rejection of certain biological processes. though the term itself is modern. although evolution accounts for the nature of the biosphere.Creationism 384 Creationism "Creationism" can also refer to creation myths.[8] Strict creationists of the Christian faith usually base their belief on a literal reading of the Genesis creation narrative. Strict creationists[7] believe that evolution cannot adequately account for the history. such as that of evolution. various views developed which aimed to reconcile science with the Genesis creation narrative. as an explanation accounting for the history. Creationism is the religious belief[1] that humanity. These groups succeeded in getting teaching of evolution banned in United States public schools.[14] When mainstream scientific research produces theoretical conclusions which contradict a strict creationist interpretation of scripture. then from the mid-1960s the young Earth creationists promoted the teaching of "scientific creationism" using "Flood geology" in public school science classes as support for a purely literal reading of Genesis. evolution itself is cosmologically attributable to a Creator deity. evolutionary creationists maintain that. However. When the court case Edwards v. the idea of common descent. the content was stripped of overt biblical references and renamed creation science. life. The Kitzmiller v.[4] Since the 1920s.[17] The rejection of scientific findings has sparked political and theological controversy. In contrast to the strict creationists.[2] As science developed from the 18th century onwards. though its usage was contested by other groups who believed in various concepts of creation. and complexity of life on earth. diversity.[2] Two offshoots of creationism—creation science and intelligent design—have been characterized as pseudoscience by the mainstream scientific community. the geological history of the Earth. and Maimonides described the story of Eve and the serpent as "most absurd in its literal sense. with the spiritual and allegorical interpretations of Genesis often being seen as more important than the literal. James Ussher. asserted that there are "waters above the heavens".[29] In 1650 the Archbishop of Armagh. though a German peasant listening to a translation would have different perceptions from a Jew familiar with early Jewish language and culture. but did not take the chronology or the days as ‘literal’. but as an allegory it contains wonderful wisdom. In particular. and obstacles be placed to their believing. only in such measure as to be ready to abandon it if it be proved with certainty to be false. he noted that Moses said the world was created in six days. Saint Basil in the fourth century while literal in many ways. The first century Jewish writer Philo admired the literal narrative of passages concerning the Patriarchs. asserted the need to hold the truth of Scripture without wavering while cautioning "that since Holy Scripture can be explained in a multiplicity of senses. like Augustine. published the Ussher chronology based on Bible history giving a date for Creation of 4004 BC. with Martin Luther advocating the idea that creation took six literal days about 6000 years ago. For example. but God through Moses described them in figurative or allegorical language. one should not adhere to a particular explanation.Creationism 385 Judaism and early and medieval Christianity The Genesis creation narrative appears in the Jewish Torah. contradicting the contemporary understanding of nature.[28] Natural theology developed the study of nature with the expectation of finding evidence supporting Christianity.[27] In the thirteenth century Thomas Aquinas. Similarly. being immeasurable and indivisible."[25] Natural theology From 1517 the Protestant Reformation brought a new emphasis on lay literacy. and Luther still had to refer to allegorical understandings such as the meaning of the serpent.[25] [26] To a large extent. and a new belief developed that each of these biological species had been individually created by God. Augustine emphasized that the text was difficult to understand and should be reinterpreted as new knowledge became available. but also shows creation occurring simultaneously. Christians should not make absurd dogmatic interpretations of scripture which contradict what people know from physical evidence.[25] Augustine of Hippo in The Literal Meaning of Genesis was insistent that Genesis describes the creation of physical things. but the development of modern geology in the 18th and . but in other passages viewed the literal interpretation as being for those unable to see an underlying deeper meaning. and his Baconian method introduced the empirical approach which became central to modern science. and claiming that "Moses wrote that uneducated men might have clear accounts of creation". John Calvin also rejected instantaneous creation. Genesis was about real events. lest holy Scripture be exposed to the ridicule of unbelievers. Thus Origen believed that the physical world is ‘literally’ a creation of God. For him. This was generally accepted. In 1605 Francis Bacon emphasized that the works of God in nature teach us how to interpret the word of God in the Bible. with days not meant literally. light was the illumination of angels rather than visible light. described creation as instantaneous and timeless.[25] Discoveries of new lands brought knowledge of a huge diversity of life. and early Jewish teachers believed that the inspired biblical text contains layers of meaning. the early Christian Church Fathers read creation history as an allegory. and followed Philo's ideas of time beginning with an instantaneous creation. but without taking a purely literal view of the texts. but did not consider this as a length of time as "we must think of God as doing all things simultaneously" and the six days were mentioned because of a need for order and according with a perfect number. with the days of creation being categories for didactic reasons. and spiritual light was just as literal as physical light. and fully agrees with the real facts". and numerous attempts were made to reconcile new knowledge with the biblical Deluge myth and story of Noah's Ark. a logical framework which has nothing to do with time. The tradition of such writers as Abraham ibn Ezra consistently rejected overly literal understandings of Genesis. Christian orthodoxy rejected the second century Gnostic belief that Genesis was purely allegorical. but criticised those who. In Evolution Vs.[42] Some theists took the general view that. Asa Gray argued that evolution is the secondary effect.[32] Theistic evolution Through the 19th century the term creationism most commonly referred to direct creation of individual souls. of the first cause. Eventually it was realised that supernatural intervention could not be a scientific explanation. but this was found to be untenable[29] and by 1850 all geologists and most Evangelical Christians had adopted various forms of old Earth creationism. it is therefore well accepted by people of strong theistic (as opposed to deistic) convictions. but their attention quickly turned to the much greater uproar about Essays and Reviews by liberal Anglican theologians. which he thought ridiculous. mostly amongst anatomists.[38] In America. some or all classical religious teachings about Christian God and creation are compatible with some or all of modern scientific theory.[34] [35] Darwin's view at the time was of God creating life through the laws of nature. This book re-examined the Bible and cast doubt on a literal interpretation. Natural Selection is not inconsistent with Natural Theology. This acceptance was resisted by conservative evangelicals in the Church of England.[33] When On the Origin of Species was published.[43] It generally views evolution as a tool used by God.[39] and published a pamphlet defending the book in theistic terms. though he later regretted using the term rather than calling it an unknown process. Following the publication of Vestiges there was interest in ideas of Creation by divine law. Creationism. the liberal theologian Baden Powell argued that this illustrated the Creator's power better than the idea of miraculous creation.[23] At this time those holding that species had been separately created were generally called "advocates of creation". and fears of republican revolutions such as the American Revolution and French Revolution led to a harsh repression of such evolutionary ideas.[31] and was also used in a response by Charles Lyell. but they were occasionally called "creationists" in private correspondence between Charles Darwin and his friends. or modus operandi.Creationism 19th centuries found geological strata and fossil sequences indicating an ancient Earth.[4] The term appears in letters Darwin wrote between 1856 and 1863. Catastrophism was favoured in England as supporting the Biblical flood. Charles Darwin's development of his theory of natural selection at this time was kept closely secret. In 1859 Darwin's On the Origin of Species provided that evidence from an authoritative and respected source. In particular. and the anonymous publication of Vestiges of Creation in 1844 aroused wide public interest with support from Quakers and Unitarians. including specifically evolution.[25] Britain at that time was enmeshed in the Napoleonic Wars. design.[25] 386 Growing evidence for evolution From around the start of the nineteenth century. however most adherents . ideas such as Jean-Baptiste Lamarck's concept of transmutation of species had gained a small number of supporters in Paris and Edinburgh. it is also known as "evolutionary creation". often involving special creation of human beings. which introduced into the controversy "the higher criticism" begun by Erasmus centuries earlier. which challenged the divine hierarchy justifying the monarchy. instead of faith being in opposition to biological evolution.[30] By 1875 most American naturalists supported ideas of theistic evolution. while continuing to firmly reject evolution. the cleric Charles Kingsley wrote of evolution as "just as noble a conception of Deity".[34] [40] [41] Theistic evolution became a popular compromise. and St. Repression eased. which emphasized the need for solidly backed science. Eugenie Scott and Niles Eldredge state that it is in fact a type of evolution. but was strongly criticised by the scientific community. who is both the first cause and immanent sustainer/upholder of the universe.[36] [37] and the book makes several references to "creation". in contrast to traducianism. and gradually convinced scientists that evolution occurs. George Jackson Mivart was among those accepting evolution but attacking Darwin's naturalistic mechanism. and naturalistic mechanisms such as neo-Lamarckism were favoured as being more compatible with purpose than natural selection. Theistic evolution can synthesize with the day-age interpretation of the Genesis creation narrative. [4] Several U.[44] In one form or another.. It is rooted in a belief that everything depends upon God."[46] Roman Catholic schools teach evolution without controversy on the basis that scientific knowledge does not extend beyond the physical. while Old Earth creationism accepts geological findings and other methods of dating the earth and believes that these findings do not contradict the Genesis myth. By the start of the twentieth century. but rather as a literary framework or allegory. 387 Re-emergence in the United States In the United States. as upheld in the Scopes Trial. They typically believe the days in Genesis Chapter 1 are 24 hours in length. After World War I. naturalistic explanations. theistic evolution is the view of creation taught at the majority of mainline Protestant seminaries[45] For Catholics. (Vatican's chief astronomer between 1978 and 2006): . Evolution was omitted entirely from school textbooks in much of the United States until the 1960s. there are other religious people who support creationism. human evolution is not a matter of religious teaching.[49] including atheists. renewed efforts to introduce teaching creationism in American public schools in the form of flood geology.Creationism consider that the first chapters of Genesis should not be interpreted as a "literal" description. but reject evolution. From a theistic viewpoint. In the creation-evolution controversy its proponents generally take the "evolutionist" side.[47] Theistic evolution can be described as "creationism" in holding that divine intervention brought about the origin of life or that divine Laws govern formation of species.S. Since then. In the United States the term started to become associated with Christian fundamentalist opposition to human evolution and belief in a young Earth in 1929. popular belief that German aggression resulted from a Darwinian doctrine of "survival of the fittest" inspired William Jennings Bryan to campaign against the teaching of Darwinian ideas of human . which is neither precluded nor required by the sources of faith. Judaic-Christian faith is radically creationist. life forms developed in biological evolution. Alternately. creationism has come to mean some fundamentalistic. evolution was widely accepted and was beginning to be taught in U.[4] Such beliefs include Young Earth creationism. many modern philosophers of science. and are so self-sufficient that the complexity of the entire physical universe evolved from fundamental particles in processes such as stellar evolution. public schools. but in terms of allegorical interpretations of Genesis. some religious communities have refused to accept. and intelligent design have been consistently held to contravene the constitutional separation of Church and State by a succession of legal judgments. and tried instead to counter them. and must stand or fall on its own scientific merits. George Coyne. or better. though many creationists (in the strict sense) would deny that the position is creationism at all. as theistic evolutionists have accepted.[50] refer to the long standing convention in the scientific method that observable events in nature should be explained by natural causes. The Catechism of the Catholic Church comments positively on the theory of evolution.. Evolution and the Roman Catholic Church are not in conflict. but by the 1980s it had been co-opted by proponents of creation science and flood geology. The term theistic evolution has been coined to refer to beliefs in creationism which are more compatible with the scientific view of evolution and the age of the Earth. scientific interpretation of Genesis. the development of life-forms and the appearance of man. literal. states passed laws against the teaching of evolution in public schools. stating that scientific studies "have splendidly enriched our knowledge of the age and dimensions of the cosmos.in America. creation science. the proponents of theistic evolution reject the implication taken by some atheists that this gives credence to ontological materialism. This sentiment was expressed by Fr. with the distinction that it does not assume the actual existence or non-existence of the supernatural. all is a gift from God. but in a totally different sense. and in the same way the origin of life by natural causes has resulted from these laws.[48] While supporting the methodological naturalism inherent in modern science. and scientific truth and religious truth cannot be in conflict.[24] The meaning of the term creationism was contested. In fact. the underlying laws of nature were designed by God for a purpose. proponents of which believe that the earth is thousands rather than billions of years old.S. the Fundamentalist-Modernist Controversy led to an upsurge of fundamentalist religious fervor in which schools were prevented from teaching evolution through state laws such as Tennessee’s 1925 Butler Act.[56] but in Kitzmiller v. Arkansas judgment ruled that state laws prohibiting the teaching of evolution violate the Establishment Clause of the First Amendment of the U. Knowledge & Truth that evolution did occur but only through God being the One who brings it about.[54] The 1968 Epperson v.[59] Rather than wholly adopting the theory of natural selection. and defining creation science as positing the “creation of the universe.[25] and by 1965 the term "scientific creationism" had gained currency. Rationality.[58] Ahmadis interpret scripture from the Quran to support the concept of macroevolution and give precedence to scientific theories. and unable to dissociate itself from its creationist roots.” as well as explaining the earth’s geology by "the occurrence of a worldwide flood". mandating that "creation science" be given equal time in public schools with evolution.[54] This was ruled unconstitutional at McLean v. unlike more orthodox Muslims. the percentage of people in the USA who accept the idea of evolution declined from 45% in 1985. Ahmadis believe that mankind has gradually evolved from different species.[24] In 1981 the state of Arkansas passed a law. Morris brought the Seventh-day Adventist biblically literal flood geology of George McCready Price to a wider audience. Creationism became associated in common usage with opposition to evolution. a new group identifying themselves as creationists promoted a "Creation science" which omitted explicit biblical references. Dover the court found intelligent design to be essentially religious. It does not occur itself. Ahmadiyya Muslim Community The Ahmadiyya Movement is perhaps the only denomination in Islam that actively promotes evolutionary theory. and indeed claimed that "creationism" only referred to young Earth creationism with flood geology. to 40% in 2005. and in drafts of the creation science school textbook Of Pandas and People all references to creation or creationism were changed to refer to intelligent design.[57] 388 Movements Creationist movements exist among peoples with various religions perspectives such as Judaism. Louisiana introduced similar legislation that year. according to the Ahmadiyya Muslim Community. Christianity and Islam.[60] Mirza Tahir Ahmad. Hinduism. as part of the ruling that teaching intelligent design in public school science classes was unconstitutional. energy.[54] Undaunted.[24] The Genesis Flood by the Baptist engineer Henry M.[23] In the 1920s. an attempt to repeal the Butler Act failed.[53] In 1961 in the United States.Creationism evolution. Ahmadis promote the idea of a "guided evolution". Waters ruled that a state law requiring biology textbooks discussing "origins or creation of man and his world" to give equal treatment to creation as per Book of Genesis was unconstitutional. Arkansas in January 1982 as the creationists' methods were not scientific but took the literal wording of the Book of Genesis and attempted to find scientific support for it.[55] and when in 1975 Daniel v. Ahmadis regard Adam as being the first Prophet of God – as opposed to him being the first man on Earth. Fourth Caliph of the Ahmadiyya Muslim Community has stated in his magnum opus Revelation. viewing each stage of the evolutionary process as having been selectively woven by God. A series of judgments and appeals led to the 1987 Supreme Court ruling in Edwards v. and life from nothing. Aguillard that it too violated the Establishment Clause of the First Amendment.[52] Proponents of the intelligent design movement organised widespread campaigning to considerable effect. They officially denied any links to creation or to religion. popularizing a novel idea of Young Earth creationism. Constitution which prohibits state aid to religion.S. . Furthermore.[52] "Creation science" could no longer be taught in public schools. Act 590.[52] However.[51] [52] and by getting evolution removed from biology textbooks nationwide. is quite compatible with a degree of uncertainty or latitude about how precisely that unfolds in creative time.[72] It appears that both Philo and Augustine felt uncomfortable with the idea of a seven day creation because it detracted from the notion of God's omnipotence. in the United States. Bahá'í theology regards the Earth as ancient. However. ".[73] The historic Christian literal interpretation of creation requires the harmonization of the two creation stories. who argue that evolution is one of the principles through which God created living beings. Bahá'ís hold."[66] Leaders of the Anglican[67] and Roman Catholic[68] [69] churches have made statements in favor of evolutionary theory. Political partisanship affecting religious belief may be a factor because whilst political partisanship in the U.S. both Jews and Christians had been considering the idea of the creation history as an allegory (instead of a historical description) long before the development of Darwin's theory of evolution. and is capable of abstract thought and of spiritual development.[74] [75] They sometimes seek to ensure that their belief is taught in science classes. as have scholars such as John Polkinghorne. mainly in American schools (see Young Earth Creationism. Members of Protestant (70%).[70] and the French Jesuit priest and geologist Pierre Teilhard de Chardin saw evolution as confirmation of his Christian beliefs.[62] Humanity.[63] such as Anglicans[64] and Lutherans. albeit important and full of symbolic meaning. Bahá'í literature maintains that humanity is distinct from other parts of creation on Earth . Few Jews today (17%) are likely to accept the Biblical literal interpretation of the creation and favor an alternative explanation. humanity has gone through numerous physical changes and adaptations in time. or in any set amount of time.for most of the history of Christianity (and I think this is fair enough) an awareness that a belief that everything depends on the creative act of God. for example). Bahá'í theology holds that humanity is a species essence . and do not take a literal view of the Genesis creation narrative. the cultural environment. Philo wrote that it would be a mistake to think that creation happened in six days. is highly correlated with fundamentalist thinking this is not so in Europe. as a biological species..[62] The Bahá'í faith regards evolution (as a progress of physical form) and the act of divine creation as related processes or even as the same process viewed in different contexts. Genesis 1:1-2:3 and Genesis 2:4-25. Asa Gray and Charles Kingsley who were enthusiastic supporters of Darwin's theories upon their publication.[65] consider that there is no conflict between the spiritual meaning of creation and the science of evolution. for there to be a consistent interpretation. However evangelical Christians have continued to believe the literal claims of Genesis. however. Another example is that of Liberal theology. not providing any creation models.that only mankind has a soul.[57] Most contemporary Christian leaders and scholars from mainstream churches. Bahá'ís regards the biblical account of creation as symbolic. Mormon (76%) and Jehovah's Witnesses (90%) denominations are those most likely to reject the evolutionary interpretation of the origins of life. Rowan Williams.[71] Augustine argued that everything in the universe was created by God at the same moment in time (and not in six days as a literal reading of Genesis would seem to require). Earlier supporters of evolutionary theory include Frederick Temple. Two notable examples are the first century Jewish neoplatonic philosopher Philo of Alexandria and Saint Augustine of the late fourth century who was also a former neoplatonist.[62] Judaism and Christianity Many Christians around the world today accept evolution as the most likely explanation for the origins of species. In fact.an essential reality and part of God's eternal creation. . an inversion has happened. However. despite condemnation from Church authorities for his more speculative theories. and comparison to non-Jewish "cosmologies" of that age. but instead focusing on the symbolism in beliefs of the time of authoring Genesis. According to the Archbishop of Canterbury.[61] Far from accepting the idea of a Young Earth..Creationism 389 Bahá'í Faith The Bahá'í Faith holds the harmony of religion and science as a fundamental principle. has changed in physical form over time. The United States is the exception where belief in religious fundamentalism is much more likely to affect attitudes towards evolution than it is for believers elsewhere. Scientifically accepted age. From the days of the early Christian Church Fathers there were allegorical interpretations of Genesis as well as literal aspects. Answers in Genesis (AIG) Ministries based in both the US and United Kingdom. Creation Ministries International promotes Young Earth views in Australia. Divine intervention at some point in the past. most people labeled "creationists" are those who object to specific parts of science for religious reasons. has opened a Creation Museum to promote Young Earth Creationism.000 years old. Reshaped by global flood. A few assign a much older age to the Universe than to Earth. . No single common Scientifically accepted age.g.Creationism Opponents reject the claim that the literalistic Biblical view meets the criteria required to be considered scientific. within the approximate time frame of biblical genealogies (detailed for example in the Ussher chronology). The Christian organizations Institute for Creation Research (ICR) and the Creation Research Society (CRS) both promote Young Earth Creationism in the USA. Less than 10. global flood. No Scientifically global flood.[25] Types of Biblical creationism Several attempts have been made to categorize the different types of creationism. literally as described in Genesis creation narrative. Directly created by God. Some adherents claim the existence of Earth is the result of divine intervention Scientifically accepted age Theistic evolution Scientifically accepted age. Another organization with similar views. Macroevolution does not occur. as evidenced by what intelligent-design creationists call "irreducible complexity" Evolution from single common ancestor. and that this appearance is what gives the geological findings and other methods of dating the earth and the universe their much longer timelines. 390 Comparison of major creationist views Humanity Biological species Earth Age of Universe Young Earth creationism Gap creationism Progressive creationism Intelligent design Directly created by God.000 years old. Creationist cosmologies are attempts by some creationist thinkers to give the universe an age consistent with the Ussher chronology and other Young-Earth time frames. accepted age. however many (if not most) people who believe in a divine act of creation do not categorically reject those parts of science. Proponents hold various beliefs. Scientifically accepted age. Directly created by God. Macroevolution does not occur. Most Young Earth creationists believe that the Universe has a similar age as the Earth. South Africa and New Zealand. ancestor. so that the world appears to be much older than it is. Other Young-Earth creationists believe that the Earth and the universe were created with the appearance of age. Young Earth creationism Young Earth creationism is the belief that the Earth was created by God within the last ten thousand years.[76] [77] [78] Creationism covers a spectrum of beliefs which have been categorized into the broad types listed below. Reshaped by global flood. accepted age. Directly created by God. Canada. and create a "taxonomy" of creationists. Behe accepts evolution from primates Evolution from primates. As a matter of popular belief and characterizations by the media. Directly created by God Direct creation + evolution. Among Catholics. e. the Kolbe Center for the Study of Creation promotes similar ideas. Many religious sects teach that God created the Cosmos. Less than 10. No Scientifically (based on primate anatomy). Omphalos hypothesis The Omphalos hypothesis argues that in order for the world to be functional. Creation science Creation science is the attempt to present scientific evidence interpreted with Genesis axioms that supports the claims of creationism. holds that life was recently created on a pre-existing old Earth. These views became popular with publications of Hebrew Lexicons such as the Strong's Concordance. and placed it in the center of the universe. 391 ." (Original act of creation." This allows an indefinite "gap" of time to be inserted after the original creation of the universe. planets and everything else in the universe revolve around it. but that details of modern evolutionary theory are questionable. explanations for the fossil record as a record of the destruction of the global flood recorded in Book of Genesis (see flood geology). It is considered that the words formless and void in fact denote waste and ruin. and so on. rock strata. The Sun.Creationism Modern geocentrism Modern geocentrism holds that God recently created a spherical world. Old Earth creationism Old Earth creationism holds that the physical universe was created by God. Old-Earth creationism itself comes in at least three types: Gap creationism Gap creationism. and darkness was upon the face of the deep. but that the creation event of Genesis is not to be taken strictly literally. trees with growth rings. which may also be associated with Lucifer's rebellion. and explanations for the present diversity as a result of pre-designed genetic variability and partially due to the rapid degradation of the perfect genomes God placed in "created kinds" or "Baramin" (see creation biology) due to mutations.000 years are generally ascribed to this "world that then was". Some gap theorists expand the basic theory by proposing a "primordial creation" of biological life within the "gap" of time. Various claims of creation scientists include such ideas as creationist cosmologies which accommodate a universe on the order of thousands of years old. attacks on the science of radiometric dating through a technical argument about radiohalos. and void. also called "Restitution creationism". taking into account the original Hebrew and other places these words are used in the Old Testament." Thus. who have extended the argument to light that appears to originate in far-off stars and galaxies (see Starlight problem). Genesis 1:1-2 is consequently translated: "In the beginning God created the heavens and the earth. This theory relies on a particular interpretation of Genesis 1:1-2.[80] Discoveries of fossils and archaeological ruins older than 10. This group generally believes that the age of the Universe and the age of the Earth are as described by astronomers and geologists. while maintaining a literal interpretation of the biblical text. the six days of creation (verse 3 onwards) start sometime after the Earth was "without form and void. And the Spirit of God moved upon the face of the waters. This is thought to be "the world that then was" mentioned in 2 Peter 3:3-7. and Bible commentaries such as the Scofield Reference Bible and the Companion Bible. but prior to creation according to the Genesis creation narrative(when present biological species and humanity were created). Gap theorists can therefore agree with the scientific consensus regarding the age of the Earth and universe.[79] The idea has seen some revival in the twentieth century by some modern creationists. God must have created a mature Earth with mountains and canyons. therefore no evidence that we can see of the presumed age of the earth and universe can be taken as reliable.) "And the earth was without form. if it were to be 392 .S. public schools. in the context of Genesis 1. where curricula are under the control of State governments rather than local school boards. or billions of years of human time). founded by Hugh Ross. Its goal is to restate creationism in terms more likely to be well received by the public. usually made together with a tacit admission of supernaturalism. particularly where they point towards supernatural elements. and the Discovery Institute has run a series of campaigns to change school curricula." is at the root of many of contemporary society's ills including social unrest and family breakdown. and to bring the debate before the public. One of its principal claims is that ostensibly objective orthodox science is actually a dogmatically atheistic religion. Neo-Creationism Neo-Creationists intentionally distance themselves from other forms of creationism. Neo-Creationists argue that the scientific method excludes certain explanations of phenomena. or in a dogmatically literal interpretation of the Bible. can be properly interpreted as "age. and an open and often hostile opposition to what they term "Darwinism". as an "atheistic enterprise.[52] [83] It is widely accepted in the scientific and academic communities that intelligent design is a form of creationism. This view accepts most of modern physical science including the age of the earth.[24] In Australia. They argue that this effectively excludes any possible religious insight from contributing to a scientific understanding of the universe. Unlike their philosophical forebears. Day-Age creationism is not so much a creationist theory as a hermeneutic option which may be combined with theories such as progressive creationism. with various ideas as to how the process operated—though it is generally taken that God directly intervened in the natural order at key moments in Earth/life's history. education policy makers and the scientific community. there was a public outcry when the notion of ID being taught in science classes was raised by the Federal Education Minister Brendan Nelson. but rather much longer periods (for instance." Some adherents claim we are still living in the seventh age ("seventh day"). the minister quickly conceded that the correct forum for ID. This theory often states that the Hebrew word "yôm". Neo-Creationists largely do not believe in many of the traditional cornerstones of creationism such as a young Earth. Intelligent design Intelligent design (ID) is the claim that "certain features of the universe and of living things are best explained by an intelligent cause.[86] [87] [88] ID originated as a re-branding of creation science in an attempt to get round a series of court decisions ruling out the teaching of creationism in U. Progressive creationism can be held in conjunction with hermeneutic approaches to Genesis chapter 1 such as the day-age theory or framework/metaphoric/poetic views. Common to all forms of Neo-Creationism is a rejection of naturalism. Progressive creationism Progressive creationism holds that species have changed or evolved in a process continuously guided by God. The most recognized form of Neo-Creationism in the United States is the Intelligent Design movement. promote this theory. preferring to be known as wholly separate from creationism as a philosophy. Neo-Creationists also argue that science.[84] [77] [78] [85] and some have even begun referring to it as "intelligent design creationism". but rejects much of modern evolutionary biology or looks to it for evidence that evolution by natural selection alone is incorrect.Creationism Day-Age creationism Day-Age creationism states that the "six days" of Book of Genesis are not ordinary twenty-four-hour days. Strictly speaking. each "day" could be the equivalent of millions.[81] All of its leading proponents are associated with the Discovery Institute.[82] a think tank whose Wedge strategy aims to replace the scientific method with "a science consonant with Christian and theistic convictions" which accepts supernatural explanations. Organizations such as Reasons to Believe. It aims to re-frame the debate over the origins of life in non-religious terms and without appeals to scripture. not an undirected process such as natural selection". which generally is meant to refer to evolution. and it is We (Allah) Who are steadily expanding it. Most Muslims generally accept the scientific positions on the age of the earth. Similar to Christian creationism. Khalid Anees. president of the Islamic Society of Britain.[92] Islam Islamic creationism is the belief that the universe (including humanity) was directly created by God as explained in the Qur'an. a promise (binding on Us). However. made points including the following:[93] There is no contradiction between what is revealed in the Koran and natural selection and survival of the fittest. not evolution. Arkansas. and by the application of the Lemon test. which holds that mainstream scientific analysis of the origin of the universe is supported by the Qur'an. They said: We come. that creates a legal hurdle to teaching Intelligent Design in public school districts in other Federal court jurisdictions. and thus religious.[89] In the United States."[Qur'an 41:11] "And it is We (Allah) Who have constructed the sky (space) with might. teaching of Intelligent Design in public schools has been decisively ruled by a Federal District court to be in violation of the Establishment Clause of the First Amendment to the United States Constitution.Creationism taught. at a conference called 'Creationism: Science and Faith in Schools'. where fewer than 25% of people believe in evolution. then We (Allah) clove them asunder and We (Allah) created every living thing out of the water. It usually views Genesis as a corrupted version of God's message. however unlike Ahmadi Muslims. do not agree that one species can develop from another. based on previous Supreme Court decisions in Edwards v. antecedents. obedient. This sets a persuasive precedent. there is concern regarding the perceived conflicts between the Qur'an and the main points of evolutionary theory. surely We will bring it about. willingly or loth. Some Muslims believe in evolutionary creationism. The main location for this has been in Turkey. Islam also has its own school of Evolutionary creationism/Theistic evolutionism.[95] There are several verses in the Qur'an which some modern writers have interpreted as being compatible with the expansion of the universe. especially among Liberal movements within Islam. Dover Area School District. as We (Allah) originated the first creation. Will they not then believe?"[Qur'an 21:30] "Then turned He to the sky (space) when it was smoke.[91] The International Society for Krishna Consciousness (ISKCON) includes several members who actively oppose "Darwinism" and the modern evolutionary synthesis. In Kitzmiller v. Aguillard and Epperson v. some Muslims.[94] But there is also a growing movement of Islamic creationism. and said unto it and unto the earth: Come both of you. namely the Vedas. and hence cannot be taught as an alternative to Evolution in public school science classrooms under the jurisdiction of that court.". is in religious or philosophy classes.[99] . Muslims believe that humans have been inserted on earth on a much later date from the creation of earth. such as Adnan Oktar.[52] [90] 393 Hinduism Some Hindus find support for. the court found that intelligent design is not science and "cannot uncouple itself from its creationist. or foreshadowing of evolutionary ideas in scriptures."[Qur'an 51:47] "On the day when We (Allah) will roll up the sky (space) like the rolling up of the scroll for writings. Big Bang and Big Crunch theories:[96] [97] [98] "Do not the Unbelievers see that the skies (space) and the earth were joined together. the age of the universe. (so) We (Allah) shall reproduce it. The creation myths in the Qur'an are vaguer and allow for a wider range of interpretations similar to those in other Abrahamic religions."[Qur'an 21:104] Many Islam majority countries have expelled science textbooks containing human evolution. and strict creationist views are much less common in other developed countries.Creationism 394 Judaism Judaism has a continuum of views about creation. who expounds that there was a Neanderthal-like species with which Adam mated (he did this long before Neanderthals had even been discovered scientifically). although Conservative Judaism does not have an official view on the subject. our own epistemological limits are to blame for any apparent irreconcilable point. Prevalence Most vocal strict creationists are from the United States. of years old. from Nachmanides. Many kabbalists accepted the teachings of the Sefer Temunah. his close student Yitzhak of Akko. Japan and Europe showed that public acceptance of evolution is most prevalent in Iceland. does generally embrace science and therefore finds it a "challenge to traditional Jewish theology. a survey of the United States. Forty-two percent believe in a "wholly scientific" explanation for the origins of life. According to a study published in Science. and the RADBAZ. including many Orthodox Jewish groups. so too can they believe that the world was created in its "adult" state. Other interesting parallels are derived. Just as they believe God created man and trees and the light on its way from the stars in their adult state. Turkey. This belief has been advanced by Rabbi Dr."[107] A 2009 Nielsen poll showed that almost a quarter of Australians believe "the biblical account of human origins" over the Darwinian account.3 billion years ago were in line with the 19th century scientific theories. the origin of life and the role of evolution in the formation of species. with the understanding that there are. To these groups. The major Jewish denominations. Denmark and Sweden at 80% of the population. former philosophy professor at Johns Hopkins University. according to Rabbi Aryeh Kaplan. of the Australian population now believe that Earth is thousands. They point out the fact that the even root word for "world" in the Hebrew language — ( םלועOlam) — means hidden — םלענ (Neh-Eh-Lahm). rather than billions. and can be. including the Ramban.[101] [102] [103] [104] In 1838 rabbi Israel Lipschitz again restated that the calculations of the ancient kabbalists that Creation occurred about 15. the notion that science and the Bible should even be reconciled through traditional scientific means is questioned. relatively old Kabbalistic sources from well before the scientifically apparent age of the universe was first determined are in close concord with modern scientific estimates of the age of the universe. among other sources. They point to various discrepancies between what is expected and what actually is to demonstrate that things are not always as they appear. but rather as a symbolic or open-ended work. science is as true as the Torah and if there seems to be a problem. Many Conservative Rabbis follow theistic evolution. no physical ways to verify it. Dovid Gottlieb. an early kabbalistic work attributed to the 1st century Tanna Nehunya ben ha-Kanah. Australian Young Earth [105] [106] Creationists claimed that "five percent Views on human evolution in various countries. while 32 percent believe in an evolutionary process ."[100] Reform Judaism does not take the Torah as a literal text. and based on Sefer Temunah.[57] Australia According to a PBS documentary on evolution. accept evolutionary creationism or theistic evolution. Also. Conservative Judaism however. For Orthodox Jews who seek to reconcile discrepancies between science and the Bible. denounced evolution as "one of many lies" taught in Polish schools."[120] Poland saw a major controversy over creationism in 2006 when the deputy education minister.Creationism "guided by God". head of the worldwide Anglican Communion. In the United Kingdom. 2007. 17% opted for intelligent design. prime minister Silvio Berlusconi wanted to retire evolution from schools in the middle level. strict creationism is less well accepted.[115] In response."[109] Europe In Europe. In countries with a Roman Catholic majority. Minister of Education Roman Giertych. Member of the European Parliament Maciej Giertych. Čolić's deputy made the statement. the Archbishop of Canterbury Rowan Williams. which runs three government-funded 13 to 19 schools in the north of England (out of several thousand in the country) teaches that creationism and evolution are equally valid "faith positions". the Parliamentary Assembly of the Council of Europe has released a draft report entitled The dangers of creationism in education on June 8. this deals with it as a historical belief and addresses hostility towards evolution rather than promoting it as an alternative to naturalistic evolution. Mirosław Orzechowski.[114] There continues to be scattered and possibly mounting efforts on the part of religious fundamentalists throughout Europe to introduce creationism into public education.[117] Serbia suspended the teaching of evolution for one week in September 2004. has opposed the teaching of evolution and has claimed that dinosaurs and humans co-existed.[119] Čolić resigned after the government said that she had caused "problems that had started to reflect on the work of the entire government. 48% selected evolutionary theory.[113] However. after one week of massive protests. teachers and opposition parties" says the BBC report. In Italy.[110] A 2006 poll on the "origin and development of life" asked participants to choose between three different perspectives on the origin of life: 22% chose creationism. he reversed his opinion. papal acceptance of evolution as worthy of study has essentially ended debate on the matter for many people. only allowing schools to reintroduce evolution into the curriculum if they also taught creationism. under education minister Ljiljana Čolić. and the rest did not know.[118] "After a deluge of protest from scientists. "I have come here to confirm Charles Darwin is still alive" and announced that the decision was reversed. "as long as most scientists in our country say that it is the right theory. though regular opinion polls are not available.[116] reinforced by a further proposal of banning it in schools dated October 4. 2007. has stated that the theory of evolution would continue to be taught in Polish schools." Giertych's father. One exam board (OCR) also specifically mentions and deals with creationism in its biology syllabus.[121] .[108] 395 Canada A 2008 Canadian poll revealed that "58 percent accept evolution. His superior. Mainstream scientific accounts are expressed as fact. in the United Kingdom the Emmanuel Schools Foundation (previously the Vardy Foundation). views the idea of teaching creationism in schools as a mistake.000 years.[111] [112] Exceptionally. Most people accept that evolution is the most widely accepted scientific theory as taught in most schools. while 22 percent think that God created humans in their present form within the last 10. but God had no part in this process".[122] about 45% of North Americans believe that "God created human beings pretty much in their present form at one time within the last 10. Besides the United States the study also compared data from 32 European countries.[128] According to a study published in Science. and whether teaching creationism in science classes conflicts with the separation of church and state. of those with postgraduate degrees.[57] Education controversies In the United States.[123] and 14% believe that "human beings have developed over millions of years from less advanced forms of life. and most North Americans had heard of evolution." Another 37% believe that "human beings have developed over millions of years from less advanced forms of life. but many did not fully understand the basics of the theory.000 years or so. the controversy comes in the form of whether advocates of the Intelligent Design movement who wish to "Teach the Controversy" in science classes have conflated science with religion. creationism has become centered in the political controversy over creation and evolution in public education. Georgia [125] In 1987.Creationism 396 United States According to a 2001 Gallup poll.[122] Belief in creationism is inversely correlated to education.S.000 U.[124] Anti-evolution car in Athens. Newsweek reported: "By one count there are some 700 scientists with respectable academic credentials (out of a total of 480. earth and life scientists) who give credence to creation-science. but God guided this process". the general theory that complex life forms did not evolve but appeared 'abruptly. 74% accept evolution. They found that most North Americans were not familiar with Creationism. the number of adults who reject evolution declined from 48% to 39% and the number of people who were unsure increased from 7% to 21%. The only country where acceptance of evolution was lower than in the United States was Turkey (25%). People for the American Way polled 1500 North Americans about the teaching of evolution and creationism in November and December 1999. Turkey. between 1985 and 2005 the number of adult North Americans who accept evolution declined from 45% to 40%. Currently.'"[126] [127] A 2000 poll for People for the American Way found 70% of the United States public felt that evolution was compatible with a belief in God. The main findings were: .[90] The Truth fish. and Japan. one of the many creationist responses to the Darwin fish. O God of Israel. God had to limit or "empty" Himself. is an "endeavor designed to demonstrate that religion and science can be compatible. a theology of the cross requires that Christians accept a methodological naturalism. even death on a cross. for the crucifixion to occur. is direct evidence of God's act of creation (Murphy quotes Phillip Johnson's claim that he is speaking "of a God who acted openly and left his fingerprints on all the evidence. Political creationists are opposed by many individuals and organizations who have made detailed critiques and given testimony in various court cases that the alternatives to scientific reasoning offered by creationists are opposed by the consensus of the scientific community. Have this mind among yourselves. religious explanations can be discussed in another class Creationism can be discussed in science class as a 'belief. taking the form of a servant. Murphy argues that this view of God is incompatible with the Christian understanding of God as "the one revealed in the cross and resurrection of Jesus. On the contrary." The basis of this theology is Isaiah 45:15. In March 2006.[132] Other Christians have expressed qualms about teaching creationism. God limits divine action in the world to be in accord with rational laws God has chosen. Several religious organizations. Archbishop of Canterbury Rowan Williams." In the article "Intelligent Design as a Theological Problem". which has collected more than 13. George Murphy argues against the view that life on Earth.' not a scientific theory Creationism and evolution should be taught as 'scientific theories' in science class Only Creationism should be taught Teach both evolution and Creationism." He also said: "My . among them the Catholic Church.[129] [130] Christian criticism Many Christians disagree with the teaching of creationism. as if the Bible were a theory like other theories. in Philippians 2:5-8. but unsure how to do so No opinion In such political contexts. but emptied himself.000 signatures. For Murphy. Just as the son of God limited himself by taking human form and dying on the cross. Murphy concludes that. did not count equality with God a thing to be grasped. "Truly. though he was in the form of God. stated his discomfort about teaching creationism. while recognizing that such acceptance does not require one to accept a metaphysical naturalism.[131] The Clergy Letter Project. who. the Savior. the leader of the world's Anglicans. And being found in human form he humbled himself and became obedient unto death. creationists argue that their particular religiously based origin belief is superior to those of other belief systems. being born in the likeness of men." Murphy observes that the execution of a Jewish carpenter by Roman authorities is in and of itself an ordinary event and did not require Divine action. which is yours in Christ Jesus. which proposes that nature is all that there is. in particular those made through secular or scientific rationale. meaning that one cannot invoke God to explain natural phenomena. thou art a God who hidest thyself. It was for this reason that Paul wrote. in all its forms. This enables us to understand the world on its own terms. but it also means that natural processes hide God from scientific observation.Creationism 397 Americans believe that: Public schools should teach evolution only [128] 20% 17% 29% 13% 16% 4% 1% Only evolution should be taught in science classes. hold that their faith is not in conflict with the scientific consensus regarding evolution. saying that creationism was "a kind of category mistake."). [140] reject the non-overlapping magisteria and argue that. Early science incorporated elements of these beliefs. creationism is based on literal interpretations of the narratives of particular religious texts. many creationist beliefs can be framed as testable predictions about phenomena such as the age of the Earth. as well as other "worldviews that focus on speculation regarding the origins of life represent another important and relevant form of human inquiry that is appropriately studied in literature or social sciences courses. since creationist beliefs are not supported by empirical evidence. empirical evidence and testable explanations of natural phenomena. they.[136] [137] Some scientists. Other scientists. distributions and relationships of living organisms found on it.[138] consider science and religion to be two compatible and complementary fields. Such study. "a Christ-Focused Creation Ministry" Creation Research Society Intelligent design • • Access Research Network Center for Science and Culture Evolution • • TalkOrigins Archive National Center for Science Education Old Earth Creationism • • Reasons to Believe led by Hugh Ross [144] Answers In Creation led by Greg Neyman . with authorities in distinct areas of human experience. By contrast.[135] However. Irrespective of this diversity in viewpoints. so-called non-overlapping magisteria. a group promoting Young-Earth Creationism Creation Ministries International. the American branch of the Anglican Communion. the scientific consensus is that any attempt to teach creationism as science should be rejected. but favour verifiable scientific explanations of natural phenomena over those of creationist beliefs. such as supernatural intervention. who believe that ultimate origins and meaning are addressed by religion. the American Academy of Religion issued Guidelines for Teaching About Religion in K‐12 Public Schools in the United States which included guidance that creation science or intelligent design should not be taught in science classes.[141] [142] [143] Organizations Young Earth Creationism • • • • Answers in Genesis." However. and these cannot be confirmed or disproved by scientists. however. its geological history and the origins."[134] 398 Scientific criticism Science is a system of knowledge based on observation. an organisation promoting biblical creation Institute for Creation Research.[133] In April 2010.[139] This view is also held by many theologians." The views of the Episcopal Church. in disproving literal interpretations of creationists. on teaching creationism are also the same as Williams. such as Richard Dawkins. must include a diversity of worldviews representing a variety of religious and philosophical perspectives and must avoid privileging one view as more legitimate than others.Creationism worry is creationism can end up reducing the doctrine of creation rather than enhancing it. but as science developed these beliefs were gradually falsified and were replaced with understandings based on accumulated and reproducible evidence. the scientific method also undermines religious texts as a source of truth. as "Creation science and intelligent design represent worldviews that fall outside of the realm of science that is defined as (and limited to) a method of inquiry based on gathering observable and measurable evidence subject to specific principles of reasoning. such as Stephen Jay Gould. Some creationist beliefs involve purported forces that lie outside of nature. [12] Dundes. 2008. [16] Isaak. University of California Press. the Scopes Trial of 1925 brought creationism and evolution into the adversarial environment of the American justice system. stephenjaygould. html). Mark (2004). [6] Evolution's status as a "theory" has played a prominent role in the creation–evolution controversy. Laurie Patton and Wendy Doniger. "Creationism History: Topic Index" (http:/ / www. com/ media/ voices/ science). that "evolution is a fact and a theory". html). "CA230: Interpretation and observation" (http:/ / www. MD: Rowman & Littlefield. Mark (2005). html). html). Index to Creationist Claims. Ed.. Original work published 1993. net/ history/ ordcreat-frame. Retrieved 07 April. . drawing upon the vernacular conception of "theory" as "part of a hierarchy of confidence running downhill from fact to theory to hypothesis to guess". co. google.Defending the Teaching of Evolution in Public Schools. "theories are structures of ideas that explain and interpret facts". "CH100. Charlottesville: University Press of Virginia [pp. see Evolution of complexity. . highereducation). [5] For example. TalkOrigins Archive. "CA215: Practical uses of evolution. Numbers. Mark (2004). Retrieved 2009–08-20. 147)." (http:/ / www. org/ indexcc/ CA/ CA230. 21 February). [3] Ronald L. e. Retrieved 12 April 2010. Counterbalance Meta-Library. TalkOrigins Archive. google. talkorigins. "Binary Opposition in Myth: The Propp/Levi-Strauss Debate in Retrospect". . Alan (Winter. "Madness in Method Plus a Plea for Projective Inversion in Myth". Scott (with forward by Niles Eldredge) (2004). p. [18] "Statements from Scientific and Scholarly Organizations" (http:/ / ncse.. Myth & method (http:/ / books. talkorigins. National Center for Science Education. [9] Ronald L. html). com/ creationism). com/ books?id=OgsTmeRHpeUC& printsec=frontcover& source=gbs_navlinks_s#v=onepage& q& f=false). Evolution vs. html). In scientific terminology. org/ faqs/ evolution-fact. . Creationism: An Introduction (http:/ / books. Retrieved 2007-08-15. 2010. In contrast. Berkley & Los Angeles. they often mean to characterize evolution as an "imperfect fact". talkorigins. Creationism History. Moran L (2002). therefore. html). California: University of California Press. counterbalance. Retrieved 2010-08-11. "Madness in method plus a plea for projective inversion in myth". Corey MA (1993). . ac. Numbers. talkorigins. "Making sense of the 'coincidences'". + creationism& q). Creationism. org/ indexcc/ CA/ CA215.Creationism 399 See also • • • • • • • Abiogenesis Biogenesis Biblical inerrancy Cosmological argument Flying Spaghetti Monster Teleological argument Watchmaker analogy Notes on terminology [1] Eugenie C. Counterbalance Meta-Library. "Antievolutionists and Creationists" (http:/ / www. The Guardian. Counterbalance Meta-Library. Isaak. Western Folklore (56): 39–50. [15] Isaak. . 114. Index to Creationist Claims. Mark (2005). Retrieved 2009–08-20. [10] While the term myth is often used colloquially to refer to "a false story". asp?year=& id=4298) . com/ books?id=M8wYPYrKnHgC& printsec=frontcover#v=onepage& q& f=false) (pp. 1997). org/ indexcc/ CA/ CA301. html). and served as a catalyst for the wider creation–evolution controversy. uk/ world/ 2006/ feb/ 21/ religion. "Academics fight rise of creationism at universities" (http:/ / www. . Evolution as fact and theory (http:/ / www. net/ history/ anticreat-frame. Index to Creationist Claims. royalsoc. Retrieved 2009–08-20. ISBN 0-520-24650-0. In MA Corey. org/ library/ gould_fact-and-theory. TalkOrigins Archive. Index to Creationist Claims. Myth and Method. see Irreducible complexity. [7] Campbell D (2006. 1996. 147-162]. 3-4. For a creationist perspective. when strict creationists refer to evolution as a theory. org/ indexcc/ CH/ CH100_1. Introduction. Giberson & Yerxa (2002). Retrieved 2008-08-28. 157-174). The trial was well-publicized. Charlottesville: University of Virginia Press. It is understood." (http:/ / ncse. "The ‘Ordinary’ View of Creation" (http:/ / www. guardian. Retrieved 2009-06-22. [13] Dundes. Alan (1984). Alan (1996). Connecticut: Greenwood Press. talkorigins. this article uses the term in the academic meaning of "a sacred narrative explaining how the world and mankind came to be in their present form" (Dundes A. com/ ?id=03b_a0monNYC& printsec=frontcover& dq=evolution+ vs. google. [17] Isaak. Sacred Narrative: Readings in the Theory of Myth. Retrieved 2009-06-22.g. In LL Patton & W Doniger (Eds. God and the new cosmology: The anthropic design argument (http:/ / books. . html). . "CA301: Science and naturalism" (http:/ / www. TalkOrigins Archive. counterbalance. [11] Dundes. Retrieved 16 June 2010 Also: Westport. counterbalance. [4] Ronald L. . Gould SJ (May 1981).. [19] Royal Society statement on evolution. Alan Dunes.1: Science in light of Scripture" (http:/ / www. Retrieved 12 April 2010. [14] See. ISBN 0313321221 [2] "NCSE : National Center for Science Education . pp. Evolution is a fact and a theory (http:/ / www. p. uk/ news.). Retrieved 2009–08-20. [8] For the biological understanding of complexity. net/ history/ intro-frame. Lanham. Numbers. Ed. creationism and intelligent design (http:/ / www. ac. 15 March 1863" (http:/ / www. Atlantic Monthly (Darwin Correspondence Project . . htm). . Asa (1860). 5 (July 1856)" (http:/ / www. C.Creationism [20] National Association of Biology Teachers Statement on Teaching Evolution (http:/ / www. File/ Master/ 6/ 150/ Evolution statement. com/ yonge/ book2. Retrieved 2010-08-11. C. Schafersman. D. . and Evolution (http:/ / ncse. . London: Michael Joseph. uk/ entry-1919). [35] "Darwin Correspondence Project . ISBN 0-7181-3430-3. html). 2007. darwinproject. html) by Steven D. asp). [30] Desmond. translated from the Greek by Charles Duke Yonge: Philo: Allegorical Interpretation. uk/ darwinletters/ calendar/ entry-2534. php?id=18504) [49] The Tower of Babel (http:/ / www. interacademies. com/ yonge/ index-2. Numbers. 31 May (1863)" (http:/ / www. Science. "Evolution and the Magisterium" (http:/ / www. Retrieved 2007-06-30. The Oxford Handbook of Philosophy of Religion. Young (1988). catholic. Niles Eldredge. Marston. ISBN 1854244418. net/ Object. org/ query?id=1257023127596318) 2009-10-31. Eugenie Scott. 139 [34] Darwin and design: historical essay (http:/ / www.Genesis Through History" (http:/ / www. aaas. Talk Reason [25] Forster. [36] Quammen 2006. org/ docs/ odesign/ od182/ ntse182. Moore. TalkOrigins Archive. Dr Paul (2001). org.. oxfordscholarship. Retrieved 2009-04-11. C. pdf) Joint statement issued by the national science academies of 67 countries. p62-63 [44] Albrecht Moritz (October 31. 202–208 [43] Evolution Vs. . asa3. com/ gp/ product/ product-description/ 026216180X) by Robert T. 207 (http:/ / darwin-online. American Public Media. the world's largest general scientific society: 2006 Statement on the Teaching of Evolution (http:/ / www. [26] The Works of Philo Judaeus (http:/ / www. . "Evolution and Wonder . darwinproject. . org/ national/ national_story. I (http:/ / www. aaas. . [48] Text of talk by Vatican Observatory director on ‘Science Does Not Need God. Adrian. ac. Report on "Naturalism. findarticles. 1995) by Davis A. uk/ content/ view/ 110/ 104/ ). shtml). Darwin Correspondence Project. uk/ entry-4041). uk/ entry-4196). adapted from The Biblical Flood: A Case Study of the Church's Response to Extrabiblical Evidence (Eerdmans. talkreason. darwinproject. to Gray. Roger. pdf)PDF (44. asp) [21] IAP Statement on the Teaching of Evolution (http:/ / www. com/ evolution/ education/ science-religion-evolution) by Eugenie Scott (accessed at 2007-07-09). James (1991). publicradio. [42] Bowler 2003. utexas. edu/ blogs/ archives/ bleiter/ 001072. uk/ content/ frameset?viewtype=text& itemID=F1577& pageseq=9). . "Letter 4196 — Darwin. Reason Science and Faith. Creationism. html) by 400 . 27 [40] Miles 2001 [41] Gray. C. Retrieved 2008-11-22. Jimmy (January 2004). Theism and the Scientific Enterprise" conference (http:/ / www. Retrieved 2007-08-15. Perspectives on Science and Christian Faith. Charles to Darwin. R. Asa. William Coleman: Microsoft Encarta Online Encyclopedia 2007. [33] Bowler 2003. This Rock. [28] Moore. ivycottage. [39] Dewey 1994. html). 11: God. Retrieved 2008-09-17.Understanding Charles Darwin" (http:/ / speakingoffaith. org/ articles/ HistoryID. . html) [27] Davis A. Retrieved 2008-08-18. com/ oso/ public/ content/ philosophy/ 0195138090/ acprof-0195138090-chapter-12.. [46] Akin. 18 November 1859" (http:/ / www. [47] Jeff Severns Guntzel (2005). pp. "The Origin of Life" (http:/ / www. freeinquiry. org/ faqs/ abioprob/ originoflife. talkorigins. R. Retrieved 2009-04-11. nabt. uk/ content/ view/ 84/ 69/ ). R.. arn. [31] "Letter 1919 — Darwin. Chester. Retrieved 2010-08-11. org/ news/ releases/ 2006/ 0219boardstatement. p. The Leiter Reports (http:/ / webapp. Retrieved 2007-06-30. Retrieved 2010-08-11. org/ programs/ darwin/ transcript. to Hooker. html#intro). to/ apologetics/ p82. . England: Monarch Books. [24] Creationism/ID. "Chapter 7 . shtml) [23] " Creationism (http:/ / encarta. bringyou. Speaking of Faith (Radio Program). catholic. aspx?id=6826). including the United Kingdom's Royal Society (PDF file) [22] From the American Association for the Advancement of Science. Naturalism is an Essential Part of Science and Critical Inquiry (http:/ / www. Pennock. com/ naturalism. Darwin Correspondence Project. Darwin Correspondence Project. Darwin. com/ p/ articles/ mi_m1141/ is_21_41/ ai_n13592804/ print). msn. ac. org/ ASA/ PSCF/ 1988/ PSCF3-88Young. National Catholic Reporter. com/ text_761580511___0/ Creationism. Retrieved 2007-06-27. org/ group/ group. earlychristianwritings. ac. J. Or Does It? A Catholic Scientist Looks at Evolution’ Catholic Online (http:/ / www. Religion. darwinproject. html).Letter 2534 — Kingsley.8 KB). org/ news/ releases/ 2006/ pdf/ 0219boardstatement. Archived (http:/ / www. R. ac. html)" Contributed By: Ronald L. [32] "Letter 4041 — Lyell. and Naturalism (http:/ / www. amazon. p. Young. . [29] History of the Collapse of "Flood Geology" and a Young Earth (http:/ / www. darwinproject. cfm) By Lenny Flank. Charles to Darwin. Penguin Group. James. earlychristianwritings. org/ sub/ position_statements/ evolution. ac. AAAS Denounces Anti-Evolution Laws (http:/ / www. webcitation. Darwin Correspondence Project. htm). Retrieved 2007-08-15. 2006). darwinproject. p. p. 119 [37] Moore 2006 [38] Barlow 1963. "Natural Selection is not inconsistent with Natural Theology" (http:/ / www. A Short Legal History (http:/ / www. html). [45] Science. "National Catholic Reporter: Catholic schools steer clear of anti-evolution bias" (http:/ / www. com/ thisrock/ 2004/ 0401bt.Essay: Natural selection & natural theology). "The contemporary relevance of Augustine" (http:/ / www. com/ yonge/ book2. evolutionnews. 53. html). Statement on Intelligent Design (http:/ / www. html). Schick (1965). n. King James Version" (http:/ / www. org/ library/ articles/ Guided_evolution_and_punctuated_equilibrium-20081104MN. com/ media/ voices/ lutheran-world-federation). Gary L. Utke. Emeritus Professor of Philosophy at the Simon Fraser University (New Zealand). [54] McLean v. 17. [51] s:Kitzmiller v. html). 90–5. biblegateway. [55] Edwards v. Chapter 2. html) [59] http:/ / textonly. . html) by Michael Martin (philosopher) [50] Butterflies and wheels article (http:/ / www.1126746. Keven. Science 313 (5788): 765–6. "The Lutheran is the magazine of the Evangelical Lutheran Church in America" [66] Archbishop of Canterbury. May. Von Kitzing. Decision January 5. n. org/ origins/ postmonth/ mar06. com/ articleprint. Patrick Russell (October 2006). itl-usa.Every Wind of Doctrine (http:/ / textonly. Ross. "Lutheran World Federation" (http:/ / ncse. pdf) [74] Wayne Jackson. Creationism's Propaganda Assault on Deep Time and Evolution. co. Scott EC. [75] "The Creation Myths: Internal Difficulties" (http:/ / www. Omphalos: An Attempt to Untie the Geological Knot. thelutheran. org/ files/ nagt/ jge/ abstracts/ Ross_v53n3p319. Marcus R. cfm?article_id=6093). v. n. infidels. org/ 2005/ 12/ post_6. [77] Wise. org/ ASA/ topics/ Bible-Science/ PSCF3-88Young. uk/ 2006/ 03/ 21/ archbishop_backs_evolution/ ) [68] What Catholics Think of Evolution? They don't not believe in it (http:/ / slate. doi:10. July 12. pp. htm). p. Bradley. biology. Evolution and Bahá'í belief: ʻAbduʾl-Bahá's response to nineteenth-century Darwinism. Retrieved 2010-05-17. butterfliesandwheels. com/ creationism/ general/ creationevolution-continuum). etc. Public acceptance of evolution". General Convention (2006)" (http:/ / ncse. 401 . pp. [64] "Episcopal Church. com/ ?id=egDAfpkONRsC). "'God allows the universe to create itself and evolve'" (http:/ / www. com/ id/ 2122506/ ).1126/science. Tuesday 21 March 2006 (http:/ / www. org/ faqs/ edwards-v-aguillard. 2007. [53] TalkOrigins Archive: Post of the Month: March 2006 (http:/ / www. 19. talkorigins. v. p. Kalimat Press. Slate Magazine. Journal of Geoscience Education. . ISBN 9781890688080. 319-323 [79] Gosse. com/ ?id=EGbZMLCmE1IC& lpg=PA90& pg=PA90#v=onepage& q). html) [56] Evolution News & Views: Dover Judge Regurgitates Mythological History of Intelligent Design (http:/ / www. (http:/ / www. [78] Who Believes What? Clearing up Confusion over Intelligent Design and Young-Earth Creationism (http:/ / nagt. pdf [61] Matthews. The Register. National Center for Science Education. PMID 16902112. Justifying Methodological Naturalism (http:/ / www. 30-35. 3. .1:42-45 (3/1988)). Office of Public Policy Barbara Forrest. Volume 12 of Studies in the Bábí and Bahá'í religions (http:/ / books. The History of Creationism by Lenny Flank. . 2005 4:43 PM. Retrieved 2007-05-23. 23-25. org/ pdf/ report2religious-landscape-study-chapter-2. "Science communication. 4. google. Retrieved 2006-12-31. Van Voorst. Aguillard (http:/ / www. D. org/ article/ article_buy. [65] Edwin A. com/ media/ voices/ episcopal-church-general-convention-2006). The Lutheran. [63] "Denominational Views" (http:/ / ncse. ISBN 9781931847162. Draper. 6. apologeticspress. org/ ahmadi/ ahmadi13. J. [58] ITL: Jesus and the Indian Messiah . edu/ ID. Retrieved 2010-05-17. html). May. earlychristianwritings. The Challenge of Baha'u'llah (http:/ / books. . National Center for Science Education. google. org/ faqs/ mclean-v-arkansas. Henry Philip. com/ paulntobin/ creationint. talkorigins. 1999. . [76] The Creation/Evolution Continuum (http:/ / ncse. . php?num=158) by Raymond D. National Center for Science Education. com/ religion/ denominational-views). Arkansas Board of Education (http:/ / www. . Discovery Institute. geocities. 2005. US Baha'i Publishing Trust. eds (2001). Dover Area School District/2:Context#Page 19 of 139 [52] Understanding the Intelligent Design Creationist Movement: Its True Nature and Goals. uiowa. Chris Williams. Eugenie Scott. com/ embor/ journal/ v6/ n12/ full/ 7400589. 1982. Young. he is a Primate. in The Encyclopedia of the Lutheran Church. talkorigins. Keelin McDonell. [69] See also the article Evolution and the Roman Catholic Church. org/ library/ modern/ michael_martin/ naturalism. alislam. The American Scientific Affiliation [73] Pugh Forum findings Page 95 (http:/ / religions. html [60] http:/ / www. (2005). org/ issue46/ 46pigliucci. Retrieved 2007-07-01 [57] Miller JD. 2005. nature. Eberhard. org/ 354) [67] Archbishop of Canterbury backs evolution: Well. Philosophy Now: The Alleged Fallacies of Evolutionary Theory (http:/ / www. html) by Massimo Pigliucci. html). "Are There Two Creation Accounts in Genesis?" (http:/ / www. html) [72] ASA3.U. v. net/ uploads/ attachments/ Forrest_Paper. 49. [70] see eg John Polkinghorne's Science and Theology pp6-7 [71] The Works of Philo Judaeus. "The Contemporary Relevance of Augustine's View of Creation" (From: Perspectives on Science and Christian Faith 40. July/August. p. Posted by Jonathan Witt on December 20.org (http:/ / www. Science and fundamentalism (http:/ / www. Retrieved 2007-05-23. centerforinquiry. translated by Charles Duke Yonge (http:/ / www. 2008. philosophynow. London [80] "The Holy Bible. Retrieved 2010-05-17. [62] Brown. Okamoto S (August 2006). Davis A. html). October 17. itl-usa. Retrieved 2010-05-17. archbishopofcanterbury. pdf). com/ passage/ ?book_id=68& chapter=3& version=9).. asa3. org/ articles/ 2194). 1857. 16-17. theregister. 2001. Allen R. Transcript of interview with the Guardian (http:/ / www. .. Explainer. NCSE Reports. org/ ahmadi/ ahmadi13. "Evolution".Creationism Paul R. Journal of Geoscience Education. pewforum. 117. pdf)PDF (413 KB) A Position Paper from the Center for Inquiry. 1. [83] Wedge Strategy (http:/ / www.org (http:/ / www. October 21. Office of Public Policy (http:/ / www. Nathan. com/ articles/ 70big_bang. [86] Forrest. pdf). html). ID is not a scientific theory. eds. harunyahya. com/ article/ dn9786-why-doesnt-america-believe-in-evolution. html). Beyond this there is considerable variability. usc. html). Forrest. Volume 19.R.. 40(2). and Gross. php?fellow=36) [96] Harun Yahya. org/ spp/ sfrl/ per/ per26. Cyril. The Big Bang Echoes through the Map of the Galaxy (http:/ / www.L. Cambridge. pbs. 402 . and Science (http:/ / www.00. 4. suggesting that doing so is merely a rhetorical ploy to "rally the troops". New Scientist 198 (2652): 31.C.. Barbara Forrest. v. smh. Jewish Virtual Library [101] Aviezer. Greenwood Press. com/ content/ muslim-cleric-challenges-darwin-we-are-descendants-human-beings-jews-brothers-apes-and-pigs) [100] Judaism and Evolution (http:/ / www. R. Hardcover. but a creationist pseudoscience. Genesis. 16-17. p. php#questionsAboutIntelligentDesign). Barbara (May. ISBN 0-88125-328-6 [102] Carmell. Pennock (editor). 2005. and "Scientific Creationism". [82] Kitzmiller v. au/ national/ god-is-still-tops-but-angels-rate-well-20091218-l5v9... 2001." Trojan Horse or Legitimate Science: Deconstructing the Debate over Intelligent Design (http:/ / www. org/ features/ wedge.com (http:/ / news. 1998. newscientist. org/ csc/ topQuestions. where the latter is understood to be supernatural. Evolution and the Wedge of Intelligent Design: The Trojan Horse Strategy. 440 p. metanexus. uk/ news/ science/ creationist-offers-prize-for-fossil-proof-of-evolution-945289. 000-evolution-myths-it-doesnt-matter-if-people-dont-grasp-evolution. hcs. The Qur'an. 1995). co. Retrieved 2007-05-17. Inc. p. "The Quran and Modern Science". . E. Expanded Edition. Dover Area School District Ruling [85] The Creationists: From Scientific Creationism to Intelligent Design. American Association for the Advancement of Science. The Rabbinical Assembly [105] Michael Le Page (19 April 2008). uk/ conferences/ story/ 0. Issue 1. "Evolution myths: It doesn't matter if people don't grasp evolution" (http:/ / www. Sydney Morning Herald. (2) Am I (and the many others who see Dembski's movement in the same way) misrepresenting their position? The basic notion of creationism is the rejection of biological evolution in favor of special creation. Deborah Smith Science Editor. well. Robert T. com/ creationism/ general/ creationevolution-continuum). 2005 [91] Moorty. . MIT Press. Public Broadcasting Service [108] Jacqueline Maley (2009-12-19). antievolution. ISBN 0-87306-174-8 [103] Schroeder. Eugenie Scott.. Retrieved 2008-07-18. ISBN 0674023390. P. sunnysideatheism. ISBN 0195157427 [87] "Dembski chides me for never using the term "intelligent design" without conjoining it to "creationism. [106] Jeff Hecht (19 August 2006)." Professional Ethics Report (http:/ / www.C. harvard. pdf).". iskcon. aspx). 2001.1117752. Robert T. 825 p. Evolution. Gerald L. . Retrieved 2007-05-13.S. Westport. com/ article/ mg19826523. 1999. . 1990. 2004. Toby (29 September 2008). Oxford. Evolution vs. Theological. J. edu/ ~hsr/ fall2005/ mu.. com.T. p. html). Washington..com (http:/ / www. University of Southern California. "Creationist offers prize for fossil proof of evolution" (http:/ / www. dated December 20. Harvard University Press. org/ faqs/ dover/ day6pm. net/ uploads/ attachments/ intelligent-design. guardian. org/ jsource/ Judaism/ jewsevolution. Pennock. 645-667 of Intelligent Design Creationism and Its Critics: Philosophical. The Bible the Qur'an and Science. html).Creationism [81] "Top Questions-1. Cambridge. "Creationism: Science and Faith in Schools" (http:/ / education. The Age. html).Narayana (May 18–21. Abd-Allah. Fall 2005. Harvard Science Review.C. edu/ dept/ MSA/ quran/ scislam. aaas.. 1999. 224 p. Discovery Institute. Dover Testimony (http:/ / www. jewishvirtuallibrary. html) [97] Maurice Bucaille (1990). 19. "Creationists are repackaging their message as the pseudoscience of intelligent design theory. [94] Green.Krishnamurti: A Case Study" (http:/ / www.R. html). God is still tops but angels rate well (http:/ / www. David (2004-01-07). ISBN 8171011322. "Science and spirituality: Any Points of Contact? The Teachings of U. html). 1999. In the Beginning: Biblical Creation and Science. November 30. B. Vol. talkorigins. 2003. Numbers. Conservative Judaism. A Position Paper from the Center for Inquiry. MIT Press. 1976.G. com/ user/ jct/ science. org/ wgbh/ evolution/ religion/ revolution/ 1980. Tower of Babel: Evidence Against the New Creationism. newscientist. [92] ISKCON. . and Scientific Perspectives. pdf). Pennock. Retrieved 2007-08-22. Challenge: Torah Views on Science New York: Association of Orthodox Jewish Scientists/Feldheim Publishers. Scott. Guardian (London). London: The Independent. html). Retrieved 2008-12-26. templeton-cambridge. . NCSE Reports. 23-25. Massachusetts. . New Scientist 191 (2565): 11. David Mu. from Wizards of ID: Reply to Dembski (http:/ / www.. "Why doesn't America believe in evolution?" (http:/ / www. Krishnamurti Centennial Conference. net/ magazine/ ArticleDetail/ tabid/ 68/ id/ 2645/ Default. The Science of God: The Convergence of Scientific and Biblical Wisdom Broadway Books. Ktav. org/ fellows/ showfellow. Science. Oxford University Press. Conclusion of Kitzmiller v. [95] Templeton-Cambridge. 2006. discovery. [84] "for most members of the mainstream scientific community.20-27. [107] Evolution Revolution (http:/ / www. Winter 1987/1988." He implies (though never explicitly asserts) that he and others in his movement are not creationists and that it is incorrect to discuss them in such terms. pdf). [88] The Creation/Evolution Continuum (http:/ / ncse. independent.: Center for Inquiry. theage. centerforinquiry. Tigay. com. ISBN 0-7679-0303-X [104] Jeffrey H. n. ISBN 0520246500 [89] Intelligent design not science: experts (http:/ / www.. 296p.What is the theory of intelligent design?" (http:/ / www. Aryeh and Domb. au/ news/ national/ intelligent-design-not-science-experts/ 2005/ 10/ 20/ 1129775902661. ISBN 0262661241.2007) (PDF). Knowledge. 2005. Understanding the Intelligent Design Creationist Movement: Its True Nature and Goals. Cambridge.. Creationism: An Introduction. D. Ronald L. com/ node/ 1592/ 2008-12-27/ expelled_no_intelligence_allowed) [93] Papineau. July/August. [90] Full text of Judge Jones' ruling. [98] A. Discovery Institute. co. [99] Sunnysideatheism. html). html). . de/ dw/ article/ 0." (http:/ / www. reproducible and therefore can be checked independently by others. [140] Dawkins. telegraph. guardian.00. pfaw.org. Committee on Culture. coe. 11297. Natural History 106 (3): 16–22. uk/ 2/ hi/ europe/ 3663196. The Economist. . religioustolerance. 3 May 2004 [115] In the beginning: The debate over creation and evolution. April 19th 2007. int/ Main. org/ ev_publi. htm) [128] Evolution and Creationism In Public Education: An In-depth Reading Of Public Opinion (http:/ / media. August 8. gov/ nihrecord/ 07_28_2006/ story03." (http:/ / www. Tuesday March 21. 10–12. Natural causes are. org/ library/ gould_noma. co.9 percent of scientists accept evolution" Finding the Evolution in Medicine (http:/ / www. [126] "Keeping God Out of the Classroom". nap. 23. kva. Richard (2006). 11297. com/ media/ voices/ religion) [132] Murphy. religious affairs correspondent.. htm) From: religioustolerance. June 29. stm). 26 January 2006 [112] "BBC Survey On The Origins Of Life" (http:/ / www. uk/ schools/ story/ 0. Rapporteur: Mr Guy LENGAGNE. org/ indexcc/ index. . 2002. [118] "Darwin is off the curriculum for Serbian schools" (http:/ / www. asp?Link=/ documents/ workingdocs/ doc07/ edoc11297. . and Society" (http:/ / www. "Intelligent Design as a Theological Problem. Evolution. [135] Committee on Revising Science and Creationism: A View from the National Academy of Sciences.. explanations must be based on naturally occurring phenomena. html). Socialist Group. co. and Technology [133] The Guardian. com/ world/ displaystory.00. edu/ catalog/ 11876. [138] Gould. co. dw-world. com/ news/ 2008/ 08/ polling-creationism-canada-001375). aspx) [123] Substantial Numbers of Americans Continue to Doubt Evolution as Explanation for Origin of Humans (http:/ / www. nih. Archbishop: Stop teaching creationism. Newsweek. ISBN 0-5930-5548-9. "Evolution Beliefs. gallup. 2007. scientists have no way of either confirming or disproving those explanations. Parliamentary Assemble Council of Europe. Doc. Douglas J. [116] The dangers of creationism in education (http:/ / assembly. org/ pdf/ creationism/ creationism-poll. 18 December 2006 [122] Majority of Americans Doubt Theory of Evolution (http:/ / www. J. unl. [139] Gould. Retrieved 2008-08-10. . 2006. 5. in principle. int/ main. National Center for Science Education.Creationism [109] "Polling creationism in Canada" (http:/ / ncse. Rocks of Ages: Science and Religion in the Fullness of Life. html). Science. 403 . Williams backs science over Bible (http:/ / www. Retrieved 2008-08-12. Washington. Retrieved on November 11. Retrieved 2008-10-27. talkorigins. Committee on Culture. 2007. D. Retrieved 2008-08-12.. stm). Warsaw Business Journal. religioustolerance. 1987. html) See transcript of Guardian interview for primary source [134] "American Academy of Religion on teaching creationism" (http:/ / ncse. American Association for the Advancement of Science. 9 September 2004 [119] "Serbia reverses Darwin suspension" (http:/ / news. html). .. Creationism.00. July 23. Retrieved 2007-03-20. [110] Archbishop: stop teaching creationism-Williams backs science over Bible (http:/ / education. June 8. Retrieved 2010-08-09.Resolution 1580 (http:/ / assembly. [130] "99. France. pl/ ?command=article& id=35336& type=wbj)". (http:/ / www. co." [136] "An Index to Creationist Claims" (http:/ / www. The God Delusion.: National Academies Press. "Nonoverlapping Magisteria" (http:/ / www. p. Deutsche Welle. org/ news/ releases/ 2006/ pdf/ 0219boardstatement. org/ ev_publi. se/ arkiv/ FutuymaCh22final. economist.. aaas. Stephen Bates. pdf) (PDF). 2007. MOSCOW AND ROME. BBC News. Doc. stm)."Public beliefs about evolution and creation". BBC News.1728235. edu/ rhames/ courses/ current/ creation/ evol-poll. uk/ schools/ story/ 0. Socialist Group. com/ poll/ 14107/ Third-Americans-Say-Evidence-Has-Supported-Darwins-Evolution-Theory. html). html) (free pdf download ed. Stephen Jay (1997). guardian.C. htm).).1188423. uk/ news/ 1471367/ Darwin-is-off-the-curriculum-for-Serbian-schools. (2002). France. telegraph. bbc. ISTANBUL. gallup. co. Science and Education. Science and Education. stephenjaygould. html) [114] Italy Keeps Darwin in its Classrooms (http:/ / www. [117] The dangers of creationism in education . National Academy of Sciences and Institute of Medicine of the National Academies (2008). uk/ religion/ Story/ 0. uk/ 2/ hi/ science/ nature/ 4648598.org (http:/ / www. 2006. 30 January 2006 [113] Exam board brings creationism into science class (http:/ / education.2144. "In science. George L. bbc. asp?link=/ Documents/ AdoptedText/ ta07/ ERES1580. and Creationism (http:/ / www. [127] US poll results . If explanations are based on purported forces that are outside of nature. aspx) Gallup Organization. wbj. pp. 2010. [125] "Public beliefs about evolution and creation. [111] "Britons unconvinced on evolution" (http:/ / news. BBC News. uk/ 2/ hi/ europe/ 3642460. com/ video/ 27838/ Evolution-Beliefs. html). ISBN 0-309-10586-2. Rapporteur: Mr Guy LENGAGNE. "Evolutionary Science. Ipsos MORI for BBC Horizon. htm) National Institutes of Health [131] National Center for Science Education: Statements from Religious Organizations (http:/ / ncse. htm) [124] Frank Newport. coe. htm). ipsos-mori.1735731. bbc. 2007. religioustolerance. 9 September 2004 [120] "'Anti-Darwin' Serb minister quits" (http:/ / news. com/ news/ 2010/ 07/ american-academy-religion-teaching-creationism-005712). Transworld Publishers. "Evolution" (2005). National Center for Science Education. once most conspicuous in America.. is fast going global (http:/ / www. 16 September 2004 [121] " And finally. The Guardian.1735730. biologi.co. Science. co. 2008. Evolution and religion. June 11.uk. aspx?oItemId=262). co.00. . com/ researchpublications/ researcharchive/ poll. S. pdf) (PDF). guardian. New York: Ballantine Books. p. October 4. pdf)PDF (481 KB) [129] "Statement on the Teaching of Evolution" (http:/ / www. Parliamentary Assemble Council of Europe. [137] Futuyma. cfm?story_id=9036706).." in Covalence: the Bulletin of the Evangelical Lutheran Church in America Alliance for Faith. American Public Media. • Darwin. Retrieved 2009-06-10. Nora. • Quammen.Creationism [141] "Royal Society statement on evolution. ed. ISBN 0-520-23693-9. PZ (2006-02-15). • Forster. London: Michael Joseph. "Ann Coulter: No Evidence for Evolution?" (http:/ / scienceblogs. John (1994).html)] 1809–1882. .html). php). James A. England: Monarch Books. Reason Science and Faith (Ivy Cottage: E-Books ed. In Martin Gardner. and Reform in Radical London.ivycottage. Prometheus Books. Retrieved 2009-03-24. Perspectives on Science and Christian Faith 53: pp. uk/ news.org/programs/darwin/transcript. • Hayward. James L. Darwin. Marston. Sara Joan (2001). Chicago: University of Chicago Press. royalsoc. Darwin.org.Understanding Charles Darwin (http://speakingoffaith. (2000).). (2003). • Desmond. pp.). "10 Significant Court Decisions Regarding Evolution/Creationism" (http:/ / ncse. • Dewey. Louise (2007-07-31). 253. David (2006). Edited and with Appendix and Notes by his Granddaughter Nora Barlow]. answersincreation. Barlow. Great Essays in Science. "Charles Darwin and Asa Gray Discuss Teleology and Design" (http://www. (1998). Molleen. Mead. Retrieved 2008-11-22. aspx?id=6826). 2006-04-11. Evolution: The History of an Idea (3rd ed. Victorian Sensation: The Extraordinary Publication. 196–201. Reception. publicradio. . [144] Answers In Creation (http:/ / www. The Royal Society. Penguin Group. [143] Myers. University of California Press. uk/EditorialIntroductions/Freeman_LifeandLettersandAutobiography. ISBN 0-393-05981-2. Chester. With the Original Omissions Restored. Roger. ISBN 1-85424-441-8. • Bowler. James (1991). Charles (1958). James (2006).org. Chicago: University of Chicago Press. com/ pharyngula/ 2006/ 06/ ann_coulter_no_evidence_for_ev.org/ ASA/PSCF/2001/PSCF9-01Miles. London: Collins. Adrian (1989). 7): pp. ISBN 0-8108-3386-7. • Miles. [142] Matsumura. Pharyngula (ScienceBlogs).org/group/group. Moore. ac. The Reluctant Mr. National Center for Science Education. ISBN 0-87975-853-8. 201–278. "The Influence of Darwinism on Philosophy". "Genesis Through History" (http://www. and Secret Authorship of Vestiges of the Natural History of Creation. Retrieved 2007-09-12. Bulletin of the British Museum (Natural History) Historical Series 2 (No. . . creationism and intelligent design" (http:/ / www. ISBN 0-7181-3430-3. Retrieved 2008-11-22. Speaking of Faith (Radio Program).uk/content/frameset?viewtype=text& itemID=F1577&pageseq=1). Adrian. • Secord. • Moore. [[The Autobiography of Charles Darwin (http://darwin-online. com/ taking-action/ ten-major-court-cases-evolution-creationism). ISBN 0-226-74411-6. asp?year=& id=4298).shtml). Dr Paul (1999).asa3. The Politics of Evolution: Morphology. org) 404 Notes References • "Darwin's Ornithological Notes" (http://darwin-online. Retrieved 2009-01-09. ISBN 0-226-14374-0. Scarecrow Press/Salem Press. • Desmond. The Creation/Evolution Controversy: an annotated bibliography. Retrieved 2008-11-04. 1963. Medicine. New York: Atlas Books. Retrieved 2007-04-23. Evolution and Wonder . Peter J. org/) This site features many videos bringing a Creationist perspective.edu/entries/creationism/) by Michael Ruse • HowStuffWorks: How creationism works (http://www. 2008 • Numbers. Larson and Larry Witham Leading scientists still reject God in Nature. 2005. 1999 • Aryeh Kaplan.themilitant.stephenjaygould. • Edward J. 10. Creationism & ID Timeline (http://www. htm) Focuses on major historical and recent events in the scientific and political debate • Evolution and Creationism (http://images. 313.com/creationism. No. 2000. Vol. A Dictionary of Creation Myths. OUP. Vol.sciam.stanford.talkorigins.htm) • Harun Yahya (http://www. • Anderson.html) by Isaac Asimov.at/20051012/Evolution-and-Creationism. NY.talkorigins. Ballantine Books.com/2005/6935/693551. 66–73 External links • The Edinburgh Creation Group (http://edinburghcreationgroup. Bernhard W. ISBN 0674023390. 1993 • Stuart Kauffman Reinventing the Sacred.howstuffworks. Bernhard W. NJ. • Workers have stake in defending science (http://www. Primack and Nancy Ellen Abrams In a Beginning.org/RESOURCES/EVOLUTION/timeline.html) from talk. Ronald (November 30.php) Examines whether Biblical creation and neo-darwinistic evolution can be reconciled.derstandard.cfm?id=15-answers-to-creationist) from Scientific American . • Stephen Jay Gould Rocks of Ages: Science and Religion in the fullness of life. 394. Tikkun. ISBN 978-0195102758.. Harvard University Press. in association with the Association of Orthodox Jewish Scientists. p. Creation Versus Chaos: The Reinterpretation of Mythical Symbolism in the Bible (ISBN 1-59752-042-X) • Ian Barbour When Science Meets Religion.com/creationism/general/creationevolution-continuum) by Eugenie Scott. Muslim creationist web site • Darwinism Refuted (http://www. Resurrection. Ktav. A Guide for Museum Docents • What is creationism? (http://www. No. Expanded Edition.org/creationevolutionboth.org/faqs/mom/groves.com) based on Harun Yahya's writings • Can Creation and Evolution Both Be True? | epologetics (http://epologetics.com).. • Evolution.org (http://www. 1997. Harper SanFrancisco.: Quantum Cosmology and Kabbalah. • Joel R. The Creationists: From Scientific Creationism to Intelligent Design. and the Age of the Universe: A Kabbalistic View.origins • 15 Answers to Creationist Nonsense (http://www.html) from talk. David (1996). • Creationism (Stanford Encyclopedia of Philosophy) (http://plato.org/ctrl/azimov_creationism.darwinismrefuted.freethought-web.html) a materialist statement on creationism by The Militant.origins • The Creation/Evolution Continuum (http://ncse. pp. 6691 (1998). Online at Freethought-web.org/faqs/wic. 1. pdf)PDF (204 KB).com/article. Harper SanFrancisco • Ian Barbour Religion and Science: Historical and Contemporary Issues.allviewpoints. 2006).org/ctrl/news/file002. • Armies of the Night (http://www. Immortality. (editor) Creation in the Old Testament (ISBN 0-8006-1768-1) • Anderson.harunyahya.html) • Creationism: The Hindu View (http://www.Creationism 405 Further reading • Adams Leeming. [6] [7] Intelligent design was developed by a group of American creationists who revised their argument in the creation–evolution controversy to circumvent court rulings that prohibit the teaching of creationism as science. District Judge John E. Intelligent design is the proposition that "certain features of the universe and of living things are best explained by an intelligent cause. Constitution. a politically conservative think tank[4] [5] —believe the designer to be the God of Christianity.[12] [13] [14] [15] and indeed is pseudoscience. not an undirected process such as natural selection.S. they seek to fundamentally redefine science to include supernatural explanations.[21] ."[1] [2] It is a form of creationism and a contemporary adaptation of the traditional teleological argument for the existence of God. antecedents". leading to the 2005 Kitzmiller v. Dover Area School District trial. together with its Center for Science and Culture.[1] In so doing. that it "cannot uncouple itself from its creationist.[11] The overwhelming consensus in the scientific community is that intelligent design is not science. which barred the teaching of "creation science" in public schools as breaching the separation of church and state.[3] Its leading proponents—all of whom are associated with the Discovery Institute. Aguillard ruling. Jones III ruled that intelligent design is not science.S. and thus religious.[8] [9] [10] Proponents argue that intelligent design is a scientific theory. see Teleological argument.[8] The first significant published use of intelligent design was in Of Pandas and People.Intelligent design 406 Intelligent design For the philosophical "argument from design". where U. but one which deliberately avoids specifying the nature or identity of the designer. a 1989 textbook intended for high-school biology classes. planned and funded the "intelligent design movement".[20] [4] They advocated inclusion of intelligent design in public school curricula.[16] [17] [18] Intelligent design originated in response to the United States Supreme Court Edwards v. and that the school district's promotion of it therefore violated the Establishment Clause of the First Amendment to the U.[19] From the mid-1990s. intelligent design proponents were supported by the Discovery Institute which. and must have originated with an intelligent agent. Past examples have included the eye and the feathered wing.[29] Intelligent design in the late 20th and early 21st century is a development of natural theology that seeks to change the basis of science and undermine evolutionary theory. blood clotting. In correspondence about the question with Asa Gray. used the watchmaker analogy. is an ancient one. is the result of chance.Intelligent design 407 History Origin of the concept Whether the order and complexity of nature indicates purposeful design has been the subject of debate since the Greeks.[23] [24] In De Natura Deorum (On the Nature of the Gods. who consider modern science and the theory of evolution to be compatible with the concept of a supernatural designer. Darwin wrote that "I cannot honestly go as far as you do about Design. in December 1988. in his Summa Theologiae. and bacterial flagella. Plato posited a good and wise "demiurge" as the creator and first cause of the cosmos in his Timaeus.[33] At the Sources of Information Content in DNA conference in 1988 he said that his intelligent cause view was compatible with both metaphysical naturalism and supernaturalism. see irreducible complexity. a chemist and creationist. "Sources of Information Content in DNA. as we see it. though the essential argument remains the same: complex systems imply a designer.[31] In March 1986 a review by Meyer used information theory to suggest that messages transmitted by DNA in the cell show "specified complexity" specified by intelligence. Thaxton. for the movement." instead of creationism." which attracted creationists such as Stephen C."[28] Though he had studied Paley's work while at university. Similar reasoning postulating a divine designer is embraced today by many believers in theistic evolution. Aquinas. 45 BCE) Cicero wrote that "the divine power is to be found in a principle of reason which pervades the whole of nature. Aristotle developed the idea of an "Unmoved Mover". Thaxton held a conference in 1988. The teleological argument. current examples are typically biochemical: protein functions. Thaxton decided to use the term "intelligent design.[6] [7] [35] [36] [37] Whether this lack of . Paley's argument from design in Natural Theology (1802).[22] In his Metaphysics. ultimately led to Charles Darwin's On the Origin of Species (1859).[32] In November of that year Thaxton described his reasoning as a more sophisticated form of Paley's argument from design. the study of nature A marble bust based on a portrait ca. Forrest writes that. In the 4th century BCE. I cannot think that the world. Meyer. Some well-known forms of it were expressed in the 13th century by Thomas Aquinas and in the 19th century by William Paley. used the concept of design in his "fifth proof" for God's existence. which held in some form by Plato and Aristotle."[25] This line of reasoning has come to be known as the teleological argument for the existence of God. passion for collecting fossils and other biological specimens. or "argument from design". 370 as way of understanding "the mind of God".[27] and such arguments led to the development of what was called natural theology. This movement fueled the BC of Plato. the examples held up as evidence of design changed.[34] Intelligent design avoids identifying or naming the agent of creation—it merely states that one (or more) must exist—but leaders of the movement have said the designer is the Christian God.[26] In the early 19th century. Philosopher Barbara Forrest writes that the intelligent design movement began in 1984 with the publication by Jon A. & yet I cannot look at each separate thing as the result of Design. by the end of his life he came to regard it as useless for scientific development. I am conscious that I am in an utterly hopeless muddle.[30] As evolutionary theory expanded to explain more phenomena. Buell's the Foundation for Thought and Ethics of The Mystery of Life's Origin by Charles B. Therefore.[46] Of Pandas and People was the first modern intelligent design book. . the argument runs: The world exhibits teleological order (design. For the cause of this cause we have sought in vain among the physical forces which surround us. until we are at last compelled to rest upon an independent volition. Schiller: "It will not be possible to rule out the supposition that the process of evolution may be guided by an intelligent design". Rethinking Schools magazine characterizes it as a "creationist treatise dressed up to look like a legitimate [38] discussion of science". has been a matter of great debate between supporters and critics of intelligent design. which render evolution possible—in heredity and in adaptability. A derivative of the phrase appears in the Macmillan Encyclopedia of Philosophy (1967) in the article titled. The Kitzmiller v.[39] in an 1850 book by Patrick Edward Dove. for these properties are the cause and not the effect of evolution. and. or just a posture taken to avoid alienating those who would separate religion from the teaching of science. adaptation).[44] The phrases "intelligent design" and "intelligently designed" were used in a 1979 philosophy book Chance or Design? by James Horigan[45] and the phrase "intelligent design" was used in a 1982 speech by Sir Fred Hoyle in his promotion of panspermia."[42] The phrase can be found again in Humanism. "Teleological argument for the existence of God": "Stated most succinctly. a 1903 book by one of the founders of classical pragmatism.S. above all.Intelligent design specificity about the designer's identity in public discussions is a genuine feature of the concept. it was produced by an intelligent designer". F.[41] The Paleyite botanist George James Allman used the phrase in an address to the 1873 annual meeting of the British Association for the Advancement of Science: "No physical hypothesis founded on any indisputable fact has yet explained the origin of the primordial protoplasm.[43] Robert Nozick (1974) wrote: "Consider now complicated patterns which one would have thought would arise only through intelligent design". 408 Origin of the term The phrase "intelligent design" can be found in an 1847 issue of Scientific American.C.[40] and in an 1861 letter from Charles Darwin. of its marvellous properties. Dover Area School District court ruling held the latter to be the case. a far-seeing intelligent design. full-color illustrations. in the case of Edwards v. and thought "That's just what I need. and was the first book to make frequent use of the phrases "intelligent design.[50] Of Pandas and People was published in 1989. such as "creationism" and "creation science".[49] Stephen C.Intelligent design 409 The modern use of the words "intelligent design". and later recalled that "the term came up". and chapter titles such as 'Homology' and 'Genetics and Macroevolution'".[52] It was the first place where the phrase "intelligent design" appeared in its present use. as stated both by its publisher Jon Buell. Meyer was at the conference. A Discovery Institute report says that Charles Thaxton.[48] In drafts of the book over one hundred uses of the root word "creation".[34] and in December decided to use the label "intelligent design" for his new creationist movement. and it was described by an ACLU lawyer as a political tool aimed at students who did not "know science or understand the controversy over evolution and creationism". began after the Supreme Court of the United States.[53] [54] and by William A. [sic][47] In June 1988 Thaxton held a conference titled "Sources of Information Content in DNA" in Tacoma. were changed. editor of Of Pandas and People. and was actively promoted by creationists for public school use. in one instance. saying it was a "creationist treatise" packaged to look like a high quality science textbook.[51] "Intelligent design” was the most prominent of around fifteen new terms it introduced as a new lexicon of creationist terminology to oppose evolution without using religious language." and "design theory". thus representing the beginning of the modern "intelligent design" movement. "cdesign proponentsists". with a "glossy cover. as a term intended to describe a field of inquiry. it's a good engineering term". Washington. had picked the phrase up from a NASA scientist.[55] The book presented all of the basic arguments of intelligent design proponents before any research had been done to support these arguments. with numerous "professionally prepared charts and illustrations appear to show how concrete scientific evidence supports the existence of the unnamed 'designer'".[38] . Dembski in his expert witness report. Philosopher of science Michael Ruse believes the contents were "worthless and dishonest".[51] Rethinking Schools magazine has criticized the book. ruled that creationism is unconstitutional in public school science curricula. to "intelligent Use of the terms "creationism" versus "intelligent design" in sequential drafts of [47] [19] the book Of Pandas and People design"." "design proponents. Aguillard (1987). almost without exception. while "creationists" was changed to "design proponents" or. [51] Behe defines it as "a single system which is composed of several well-matched interacting parts that contribute to the basic function. He provides the following examples: "A single letter of the alphabet is specified without being complex. This is sometimes called the "scaffolding objection" by an analogy with scaffolding.Intelligent design 410 Integral concepts Irreducible complexity The term "irreducible complexity" was introduced by biochemist Michael Behe in his 1996 book Darwin's Black Box. the spring and the hammer—all of which must be in place for the Darwin's Black Box. which can support an "irreducibly complex" building until it is complete and able to stand on its own. philosopher. and theologian William Dembski. in the Dover trial. in his 1996 book. because the selectable function is present only when all parts are assembled. though he had already described the concept in his contributions to the 1993 revised edition of Of Pandas and People. especially the "patterns" of molecular sequences in functional biological molecules such as DNA. Dembski states that when something exhibits specified complexity (i. they argue. and the adaptive immune system. Furthermore. popularised by Michael Behe. one can infer that it was produced by an intelligent cause (i. the blood clotting cascade.[64] He states that details of living things can be similarly characterized. Removal of any one piece destroys the function of the mousetrap.[57] [58] Critics point out that the irreducible complexity argument assumes that the necessary parts of a system have always been necessary and therefore could not have been added sequentially. simultaneously). evolution often proceeds by altering preexisting parts or by removing them from a system.. cilia. coli.[56] Behe uses the analogy of a mousetrap to illustrate this concept. and that his "argument against Darwinism does not add up to a logical proof". and must have originated with an intelligent agent.[32] The intelligent design concept of "specified complexity" was developed in the 1990s by mathematician.e. Intelligent design advocates assert that natural selection could not create irreducibly complex systems. mousetrap to work.e. .. A long sentence of random letters is complex without being specified. the catch. A Shakespearean sonnet is both complex and specified".[63] Specified complexity In 1986 the creationist chemist Charles Thaxton used the term "specified complexity" from information theory when claiming that messages transmitted by DNA in the cell were specified by intelligence. Behe argued that irreducibly complex biological mechanisms include the bacterial flagellum of E.[59] [60] They argue that something which is at first merely advantageous can later become necessary as other components change. is both complex and "specified". that it was designed) rather than being the result of natural processes. wherein the removal of any one of the parts causes the system to effectively cease functioning".[62] Irreducible complexity has remained a popular argument among advocates of intelligent design.[61] Behe has acknowledged using "sloppy prose". A The concept of irreducible complexity was mousetrap consists of several interacting pieces—the base. rather than by adding them. the court held that "Professor Behe's claim for irreducible complexity has been refuted in peer-reviewed research papers and has been rejected by the scientific community at large". [81] This. such as galaxies. oxygen. proponents argue. Stenger and other critics say both intelligent design and the weak form of the anthropic principle are essentially a tautology.[74] Fine-tuned Universe Intelligent design proponents have also occasionally appealed to broader teleological arguments outside of biology. living things could not be born and grow.[76] Victor J.[82] The second law applies to closed systems only. an intelligent designer of life was needed to ensure that the requisite features were present to achieve that particular outcome. . They argue that this procedure is flawed as a model for scientific inference because the asymmetric way it treats the different possible explanations renders it prone William Dembski proposed the [65] concept of specified complexity. If Granville's argument were valid.[66] [67] [68] The conceptual soundness of Dembski's specified complexity/CSI argument has been widely discredited by the scientific and mathematical communities. most notably an argument based on the fine-tuning of universal constants that make matter and life possible and which are argued not to be solely attributable to chance. Life as we know it might not exist if things were different. Some scientists argue that even when taken as mere speculation. finally defaulting to design. as this also would be a decrease in entropy. and that it thus violates the second law of thermodynamics and so supports intelligent design. electromagnetism. argues in The God Delusion that allowing for an intelligent designer to account for unlikely complexity only postpones the problem. These include the values of fundamental physical constants. as such a designer would need to be at least as complex. these arguments are poorly supported by existing evidence.[80] Proponent Granville Sewell argues that the evolution of complex forms of life represents a decrease of entropy.[77] [78] [79] The claim of the improbability of a life-supporting universe has also been criticized as an argument by lack of imagination for assuming no other forms of life are possible. sunlight) whose production requires an offsetting net increase in entropy.[72] Richard Dawkins. is a misapplication of thermodynamic principles. and gravity between fundamental particles. so the real question becomes whether or not CSI actually exists in nature. making it impossible for many chemical elements and features of the Universe. aeronautic and automotive systems which are considered problems too complex for human "intelligent designers". these arguments amount to the claim that life is able to exist because the Universe is able to support life.[75] Thus. A number of critics also suggest that many of the stated variables appear to be interconnected and that calculations made by mathematicians and physicists suggest that the emergence of a universe similar to ours is quite probable. Neither evolution nor the growth of living things violates the second law of thermodynamics because living things are not closed systems—they have external energy sources (e. as well as the ratios of masses of such particles. Scientists have generally responded that this argument cannot be tested and is therefore not science but metaphysics. John Wilkins and Wesley Elsberry characterize Dembski's "explanatory filter" as eliminative. but a different sort of life might exist in its place. to making false conclusions. because it eliminates explanations sequentially: first regularity. Critics say that this renders the argument a tautology: complex specified information cannot occur naturally because Dembski has defined it thus. the universe would be dramatically different. in his view. Intelligent design proponent and Center for Science and Culture fellow Guillermo Gonzalez argues that if any of these values were even slightly different. food.[69] [70] [71] Specified complexity has yet to be shown to have wide applications in other fields as Dembski asserts.Intelligent design 411 Dembski defines complex specified information (CSI) as anything with a less than 1 in 10150 chance of occurring by (natural) chance. the relative strength of nuclear forces. to form. another critic of intelligent design.[73] Other scientists have argued that evolution through selection is better able to explain the observed complexity. then chance.g. however. as is evident from the use of selective evolution to design certain electronic. [68] Richard Dawkins sees the assertion that the designer does not need to be explained. for some as-yet undetectable practical purpose.[90] Movement The intelligent design movement is a direct outgrowth of the creationism of the 1980s. Later it used a less religious image. then was [91] renamed the Center for Science and Culture. For example. The new question raised by the explanation is as problematic as the question which the explanation purports to answer". The Discovery Institute's intelligent . established in 1996 as the creationist wing of the Discovery Institute to promote a religious agenda[99] calling for broad social..S. The authoritative description of intelligent design. Dembski. Dembski concludes that "no intelligent agent who is strictly physical could have presided over the origin of the universe or the origin of life". measurable evidence. for example.[85] Asserting the need for a designer of complexity also raises the question "What designed the designer?"[86] Intelligent design proponents say that the question is irrelevant to or outside the scope of intelligent design. to the exclusion of all other religions. in Darwin's Black Box. speculates that an alien culture could fulfill these requirements. Acknowledging the paradox. amphibians. the designer is often implicitly hypothesized to have intervened in a way that only a god could intervene..[6] [7] [35] Beyond the debate over whether intelligent design is scientific. Odd designs could. "have been placed there by the designer . and freshwater fish.[84] The leading proponents have made statements to their supporters that they believe the designer to be the Christian God. not as a contribution to knowledge.[88] [89] In the absence of observable. so such questions cannot be answered definitively. Although they do not state that God is the designer. along with a U. or for some unguessable reason". The movement is headquartered in the Center for Science and Culture (CSC). for artistic reasons.[10] The scientific and academic communities. mammals. but from evolution.[10] [96] [97] [98] The Discovery Institute's Center for the Renewal of Science and Culture used banners based on "The Creation of Adam" from the Sistine Chapel. Coyne responds that in light of the evidence.[85] Previously.Intelligent design 412 Intelligent designer Intelligent design arguments are formulated in secular terms and intentionally avoid identifying the intelligent agent (or agents) they posit. academic and political changes. irrespective of its status in the world of science. "either life resulted not from intelligent design.[83] however. Jerry Coyne asks why a designer would "give us a pathway for making vitamin C. the very question "What designed the designer?" leads to an infinite regression from which intelligent design proponents can only escape by resorting to religious creationism or logical contradiction. despite the suitability of such islands for these species". or the intelligent designer is a cosmic prankster who designed everything to make it look as though it had evolved". in The Design Inference. Behe had argued that we are simply incapable of understanding the designer's motives. Invoking an unexplained being to explain the origin of other beings (ourselves) is little more than question-begging. but then destroy it by disabling one of its enzymes" (see pseudogene) and why he or she would not "stock oceanic islands with reptiles.[87] Richard Wein counters that the unanswered questions an explanation creates "must be balanced against the improvements in our understanding which the explanation provides. a number of critics argue that existing evidence makes the design hypothesis appear unlikely. Coyne also points to the fact that "the flora and fauna on those islands resemble that of the nearest mainland. view intelligent design as either a form of creationism or as a direct descendant that is closely intertwined with traditional creationism. federal court.[17] [18] [92] [93] [94] [95] and several authors explicitly refer to it as "intelligent design creationism". even when the environments are very different" as evidence that species were not placed there by a designer. to show off. explicitly states that the Universe displays features of having been designed. but as a thought-terminating cliché. "[99] Barbara Forrest.[104] Johnson emphasizes that "the first thing that has to be done is to get the Bible out of the discussion".[99] Phillip E. We intend these to encourage and equip believers with new scientific evidences that support the faith... in my personal experience. which suffocates the human spirit.[108] Both Johnson and Dembski cite the Bible's Gospel of John as the foundation of intelligent design. to be sure. describes this as being due to the Discovery Institute's obfuscating its agenda as a matter of policy. systematic agenda for promoting not only intelligent design creationism.] Not only does intelligent design rid us of this ideology (materialism). they state that intelligent design has its foundation in the Bible. although efforts have been made in other countries to promote intelligent design. Christians. as well as to 'popularize' our ideas in the broader culture. Leaders of the movement say intelligent design exposes the limitations of scientific orthodoxy and of the secular philosophy of naturalism. and Stephen C.[35] [100] .[36] [100] All leading intelligent design proponents are fellows or staff of the Discovery Institute and its Center for Science and Culture. while Jonathan Wells is a member of the Unification Church. We will do this primarily through apologetics seminars. The overall goal of the movement is to "defeat [the] materialist world view" represented by the theory of evolution in favor of "a science consonant with Christian and theistic convictions". evangelistic orientation: "Alongside a focus on influential opinion-makers. She has written that the movement's "activities betray an aggressive. "ID is part of God's general revelation [.[106] In this work Dembski lists a god or an "alien life force" as two possible options for the identity of the designer. we also seek to build up a popular base of support among our natural constituency. even if its practitioners don't have a clue about him. the institute affirms its Christian. But the conceptual soundness of the theory can in the end only be located in Christ. I've found that it opens the path for people to come to Christ". Johnson explicitly calls for intelligent design proponents to obfuscate their religious motivations so as to avoid having intelligent design identified "as just another way of packaging the Christian evangelical message".[103] the majority of principal intelligent design advocates are publicly religious Christians who have stated that in their view the designer proposed in intelligent design is the Christian conception of God.[101] Nearly all intelligent design concepts and the associated movement are the products of the Discovery Institute. but. namely.[100] Recognizing the need for support. "after we have separated materialist prejudice from scientific fact [. In statements directed at the general public. Johnson has stated that cultivating ambiguity by employing secular language in arguments that are carefully crafted to avoid overtones of theistic creationism is a necessary first step for ultimately reintroducing the Christian concept of God as the designer. an expert who has written extensively on the movement. and Michael Behe is a Roman Catholic. Phillip E. in his book Intelligent Design: The Bridge Between Science and Theology. which guides the movement and follows its wedge strategy while conducting its Teach the Controversy campaign and their other related programs. Intelligent design proponents allege that science should not be limited to naturalism and should not demand the adoption of a naturalistic philosophy that dismisses out-of-hand any explanation which contains a supernatural cause. when addressing conservative Christian supporters. Phillip E.[105] The strategy of deliberately disguising the religious intent of intelligent design has been described by William Dembski in The Design Inference."[107] Dembski also stated.. but the religious world-view that undergirds it". William Dembski. Johnson stated that the goal of intelligent design is to cast creationism as a scientific concept.. Meyer are evangelical Protestants. however. be pursued without recourse to Christ.] only then can 'biblical issues' be discussed". they say intelligent design is not religious. The pragmatics of a scientific theory can. Leading intelligent design proponents have made conflicting statements regarding intelligent design. Johnson. Stuart Burgess. Dembski states that "Christ is indispensable to any scientific theory.[102] 413 Religion and leading proponents Although arguments for intelligent design are formulated in secular terms and intentionally avoid positing the identity of the designer.Intelligent design design campaigns have been staged primarily in the United States. in 1999. Charles Thaxton. that ID. "left the Bible out of it" and thereby unwittingly aided and abetted the modern rejection of the Bible. Henry M. She lists connections of (current and former) Discovery Institute Fellows Phillip Johnson.[118] A series of Gallup polls in the United States from 1982 through 2008 on "Evolution.” According to Morris: “The evidence of intelligent design… must be either followed by or accompanied by a sound presentation of true Biblical creationism if it is to be meaningful and lasting."[113] Polls Several surveys were conducted prior to the December 2005 decision in Kitzmiller v.[115] [116] [117] A May 2005 survey of nearly 1500 physicians in the United States conducted by the Louis Finkelstein Institute and HCD Research showed that 63% of the physicians agreed more with evolution than with intelligent design. The polls also noted answers to a series of more detailed questions.[109] She writes that the leading proponents of intelligent design are closely allied with the ultra-conservative Christian Reconstructionism movement. and it will fail today. Introducing a biblically based.Intelligent design Barbara Forrest contends such statements reveal that leading proponents see intelligent design as essentially religious in nature. support for "God created human beings in pretty much their present form at one time within the last 10. at best. 10% of adults in the United States viewed human beings as "so complex that they required a powerful force or intelligent being to help create them". such as having a very low response rate (248 out of 16. In 2002 he wrote: "Winning the argument for design without identifying the designer yields. "even if well-meaning and effectively articulated. not merely a scientific concept that has implications with which their personal religious beliefs happen to coincide. will not work! It has often been tried in the past and has failed. Wieland explained that "AiG's major 'strategy' is to boldly. Creationism. a sketchy origins model. two of the most prominent Young Earth creationism organizations in the world have attempted to distinguish their views from intelligent design. believes that the efforts of intelligent design proponents to divorce the concept from Biblical Christianity make its hypothesis too vague."[111] Likewise. a leading figure in the Reconstructionist movement. a single leap into the origins fray. and the extent of the funding provided the Institute by Howard Ahmanson Jr.[119] . call the church back to its Biblical foundations… [so] we neither count ourselves a part of this movement nor campaign against it. but God had no part in the process" varied from 9% to 14%. and containing leading questions. Jonathan Wells and Francis Beckwith to leading Christian Reconstructionist organizations. According to a 2005 Harris poll. but God guided the process" of between 35% and 40%. a proponent of Old Earth creationism. Such a model makes little if any positive impact on the community of scientists and other scholars… The time is right for a direct approach. Hugh Ross of Reasons to Believe. Richard Weikart. Dover. but humbly. Carl Wieland of Answers in Genesis (AiG) criticized design advocates who. being conducted on behalf of an organization with an expressed interest in the outcome of the poll.[114] Although Zogby polls commissioned by the Discovery Institute show more support. which sought to determine the level of support for intelligent design among certain groups.000)."[112] In 2002.. Michael Behe. scientifically verifiable creation model represents such a leap. Intelligent Design" found support for "human beings have developed over millions of years from less advanced formed of life. The reason it won't work is because it is not the Biblical method. and support for "human beings have developed over millions of years from less advanced formed of life. though well-intentioned.[110] 414 Reaction from other creationist groups Not all creationist organizations have embraced the intelligent design movement.000 years or so" varied from 43% to 47%. Morris of the Institute for Creation Research (ICR) wrote. these polls suffer from considerable flaws. and present intelligent design in the language of science as though it were a scientific hypothesis. In response to Bush’s statement and the pending federal trial. being asked to leave the theater. Michelangelo’s God points at the image of a chimpanzee contemplating the caption which read "The push to teach "intelligent design" raises a question: Does God have a place in science class?". [120] [121] Creating and teaching the controversy The intelligent design movement states that there is a debate among scientists about whether life evolved. and they view their work in the promotion of intelligent design as a way to return religion to a central role in education and other public spheres. among a significant proportion of the general public in the United States the major concern is whether conventional evolutionary biology is compatible with belief in God and in the Bible. finding that God and intelligent design were both distinct from the material that should be covered in a science class. and how this issue is taught in schools.[103] [105] However. Intelligent design proponents seek to change this fundamental basis of science[132] by eliminating "methodological naturalism" from science[133] and replacing it with what the leader of the intelligent design movement. Rather than pointing at Adam. of their beliefs in Intelligent Design. and cultural leaders that schools should "Teach the Controversy". which means belief in a transcendent. Dover trial in late 2005 and after President George W. Time magazine ran an eight-page cover story on the Evolution Wars in which they examined the issue of teaching intelligent design in the classroom.[131] In the Kitzmiller v. nonnatural dimension of reality inhabited by a transcendent.[129] [130] The cover of the magazine featured a parody of The Creation of Adam from the Sistine Chapel. Johnson.[122] But in fact.[134] Some have called this approach "methodological supernaturalism". politicians. One of the film's first screenings resulted in Paul "PZ" Myers. saying that teaching only evolution unfairly discriminates against those holding creationist beliefs.[136] Proponents say evidence exists in the forms of irreducible complexity and specified complexity that cannot be explained by natural processes. allows for the possibility of religious belief. though others note that intelligent design serves as an effective proxy for the religious beliefs of prominent intelligent design proponents in their efforts to advance their religious point of view within society. seeking to convince the public. an interviewee in the film. the court ruled that intelligent design was a religious and creationist position.[137] . nonnatural deity. The movement stresses the importance of recognizing the existence of this supposed debate. also has helped finance some religious films such as The Passion of the Christ. Many intelligent design followers believe that "Scientism" is itself a religion that promotes secularism and materialism in an attempt to erase theism from public life.[135] Intelligent design proponents argue that naturalistic explanations fail to explain certain phenomena and that supernatural explanations provide a very simple and intuitive explanation for the origins of life and the universe. Phillip E. There have also been allegations from some interviewees that interviews were recorded many times in order to get the exact phrasing required by the producer. Dover case. the film insinuates. Premise Media.[9] Empirical science uses the scientific method to create a posteriori knowledge based on observation and repeated testing of hypotheses and theories. the scientific consensus is that life evolved.[128] The public controversy was given widespread media coverage in the United States. This documentary. Teaching both. spends much time focusing on professors who have been asked to leave or have left numerous institutions because. there is no such controversy in the scientific community. they argue. Bush expressed support for the idea of teaching intelligent design alongside evolution in August 2005. hosted by Ben Stein. which they argue leaves no room for the possibility of God.[1] They also hold that religious neutrality requires the teaching of both evolution and intelligent design in schools. Some allege that this larger debate is often the subtext for arguments made over intelligent design. particularly during the Kitzmiller v. calls "theistic realism".[126] [127] Advocates of intelligent design seek to keep God and the Bible out of the discussion.[123] [124] [125] Intelligent design is widely viewed as a stalking horse for its proponents' campaign against what they say is the materialist foundation of science.Intelligent design 415 Film The film Expelled: No Intelligence Allowed sparked further controversy in 2008. without causing the state to actually promote such beliefs. The production company. but it is even more scientific than evolution.[106] has been compared to the a priori claim that aliens helped the ancient Egyptians build the pyramids. From a strictly empirical standpoint. physics or geology.Intelligent design [138] [139] 416 Intelligent design has not presented a credible scientific case and is an attempt to teach religion in public schools. they have also been critical of its refusal to identify the designer.[149] [150] [151] and have pointed to previous failures of the same argument.[140] If the argument to give "equal time for all theories" were actually practiced.[143] [144] In both cases.[141] [142] Intelligent design proponent Michael Behe concedes "You can't prove intelligent design by experiment". which is limited to dealing only with the natural world[145] — a position described by the term theistic evolution. and have often said that not only is their own position scientific.[156] For a theory to qualify as . Supporters of intelligent design have also reached out to other faith groups with similar accounts of creation with the hope that the broader coalition will have greater influence in supporting science education that does not contradict their religious views.[128] Intelligent design proponents cannot legitimately infer that an intelligent designer is behind the part of the process that is not understood scientifically. there would be no logical limit to the number of mutually incompatible supernatural "theories" regarding the origins and diversity of life to be taught in the public school system. and it violates the principle of parsimony.rejected by science because they do not fit the facts. wondering why no one claims that God's hand should be taught in other secular classes. Slifkin also asserts that the intelligent design movement is inordinately concerned with portraying God as "in control" when it comes to things that cannot be easily explained by science. but not in control in respect to things which can be explained by scientific theory.[154] Defining science The scientific method is a body of techniques for investigating phenomena and acquiring new knowledge of the natural world without assuming the existence or nonexistence of the supernatural. While creationist organizations have welcomed intelligent design's support against naturalism. states that intelligent design is not falsifiable because "[d]efenders of ID always have a way out". such as history. and that they want a redefinition of science as a revived natural theology or natural philosophy to allow "non-naturalistic theories such as intelligent design". such as the astronomer George Coyne and the biologist Ken Miller. Slifkin criticizes intelligent design's advocacy of teaching their perspective in biology classes. observable or falsifiable.[146] As well as pointing out that intelligent design is not science.000 clergy who signed the Clergy Letter Project. Prominent scientists who strongly express religious faith. they also reject it for various philosophical and theological reasons.[136] Many religious bodies have responded by expressing support for evolution. and having failed religion because they think too little of God. since they have not shown that anything supernatural has occurred. Intelligent design proponents believe that this can be equated to materialist metaphysical naturalism. which in the philosophy of science is about how and where to draw the lines around science. do not truly understand it.[152] Rabbi Natan Slifkin directly criticized the advocates of intelligent design as presenting a perspective of God that is dangerous to religion. The Roman Catholic church has stated that religious faith is fully compatible with science. Philosopher of biology Elliott Sober. substituting public support for scientific research. for example.[153] Those who promote it as parallel to religion. have been at the forefront of opposition to intelligent design. an approach sometimes called methodological naturalism. including intelligent design parodies such as the Flying Spaghetti Monster "theory".[155] This presents a demarcation problem. one may list what is known about Egyptian construction techniques. intelligent design does not provide a mechanism for discriminating among them.[147] [148] The arguments of intelligent design have been directly challenged by the over 10. the effect of this outside intelligence is not repeatable. he asserts. which advocate William Dembski has said could alternately be an "alien" life force.[153] Kenneth Miller expressed a view similar to Slifkin's: "[T]he struggles of the Intelligent Design movement are best understood as clamorous and disappointing double failures . but one must admit ignorance about exactly how the Egyptians built the pyramids. The inference that an intelligent designer created life on Earth. • There should be a known rate of error that can be used in evaluating the results. of these criteria. the philosopher Thomas Nagel argues that intelligent design is very different from creation science. the less scientific it is. we have addressed the seminal question of whether ID is science. not a disagreement between science and something else. observable and ultimately testable data and which require explanations to be based on empirical evidence. and ideally all. The fewer criteria are met.[162] is not falsifiable.[161] is not scientifically useful. he argues that it is a scientific disagreement. and other claims of supernatural intervention in the origin of life or of species are not science because they are not testable by the methods of science.[157] [158] [159] it is expected to be: • • • • • • • • Consistent Parsimonious (sparing in its proposed entities or explanations. In Kitzmiller v.[169] The U.[176] Intelligent design proponents believe that their writings are rejected for not conforming to purely naturalistic. • The methods should preferably be published in a peer-reviewed journal. Against this. Whatever the merits of the positions. see Occam's Razor) Useful (describes and explains observed phenomena. Dover Area School District.[163] is not empirically testable. repeated experiments Correctable and dynamic (modified in the light of observations that do not support it) Progressive (refines previous theories) Provisional or tentative (is open to experimental checking. by appealing to a supernatural agent. The Daubert Standard governs which evidence can be considered scientific in United States federal courts and most state courts. and moreover that ID cannot uncouple itself from its creationist. We have concluded that it is not. in that it does not depend on distortion of evidence. hypothesis or conjecture to be considered scientific. intelligent design. using these criteria and others mentioned above.[173] [174] Peer review The failure to follow the procedures of scientific discourse and the failure to submit work to the scientific community that withstands scrutiny have weighed against intelligent design being accepted as valid science.S. antecedents".[160] violates the principle of parsimony.[171] [17] [18] Others in the scientific community have concurred. which limit its inquiries to empirical.. National Science Teachers Association and the American Association for the Advancement of Science have termed it pseudoscience. non-supernatural mechanisms rather than because their .[172] and some have called it junk science. It depends only on the idea that the hypothesis of a designer makes sense.[165] [166] [167] Critics also say that the intelligent design doctrine does not meet the Daubert Standard. • The methods should be generally accepted within the relevant scientific community. Typical objections to defining intelligent design as science are that it lacks consistency. Behe and other intelligent design proponents say bias by the scientific community is to blame for the failure of their research to be published. provisional or progressive. Dembski.Intelligent design scientific. and if it meets only a few or none at all. and thus religious. it must meet most.[175] The intelligent design movement has not published a properly peer-reviewed article in a scientific journal. National Academy of Sciences has stated that "creationism. Its four criteria are: • The theoretical underpinnings of the methods must yield testable predictions by means of which the theory could be falsified. Judge Jones ruled that ". then it cannot be treated as scientific in any meaningful sense of the word. dynamic.S. directly conflicts with the principles of science. and can be used predictively) Empirically testable and falsifiable (see Falsifiability) Based on multiple observations.."[170] The U. often in the form of controlled.[164] and is not correctable. and does not assert certainty) 417 For any theory.[168] the criteria for scientific evidence mandated by the US Supreme Court.[175] Intelligent design. or on the assumption that it is immune to empirical evidence. [189] As summarized by the judge.[177] Michael Shermer has rebutted the claim. Harper Jr. Critics and advocates debate over whether intelligent design produces new research and has legitimately attempted to publish this research.[192] 418 . and faulty logic".[186] which mentioned neither irreducible complexity nor intelligent design and which Behe admitted did not rule out known evolutionary mechanisms."[183] In a 2001 interview. Charles L. Why? Because they are doing science. reject this claim. largely members of the scientific community. said: "From the point of view of rigor and intellectual seriousness. Some scientists describe this claim as a conspiracy theory.[175] The Discovery Institute insists that a number of intelligent design articles have been published in peer-reviewed journals. the judge wrote: "A final indicator of how ID has failed to demonstrate scientific warrant is the complete absence of peer-reviewed publications supporting the theory"."[178] The issue that supernatural explanations do not conform to the scientific method became a sticking point for intelligent design proponents in the 1990s. foundation vice-president. Meyer. says that it asked intelligent design proponents to submit proposals for actual research.[188] In sworn testimony. Dembski has written that "perhaps the best reason [to be skeptical of his ideas] is that intelligent design has yet to establish itself as a thriving scientific research program.[185] There. but none were ever submitted. Behe conceded that there are no peer-reviewed articles supporting his claims of intelligent design or irreducible complexity.[180] Written by the Discovery Institute's Center for Science & Culture Director Stephen C.Intelligent design research is not up to "journal standards".[181] The article was a literature review. intelligent design proponents cited just one paper. inappropriate mathematical modeling. however. but rather culled quotations and claims from other papers to argue that the Cambrian explosion could not have happened by natural processes. the judge found that intelligent design features no scientific research or testing. because it was so outside the normal subject matter (see Sternberg peer review controversy[182] ). Behe said: "There are no peer reviewed articles by anyone advocating for intelligent design supported by pertinent experiments or calculations which provide detailed rigorous accounts of how intelligent design of any biological system occurred"." He noted that scientists such as Joan Roughgarden and Lynn Margulis have challenged certain Darwinist theories and offered explanations of their own and despite this they "have not been persecuted.[184] In the Dover trial.[179] The only article published in a peer-reviewed scientific journal that made a case for intelligent design was quickly withdrawn by the publisher for having circumvented the journal's peer-review standards.[187] Michael Lynch called the conclusions of the article "an artifact of unwarranted biological assumptions. not religion. For instance. stating that no established scientific journal has yet published an intelligent design article.. Rather. fired or even expelled. and is addressed in the wedge strategy as an aspect of science that must be challenged before intelligent design can be accepted by the broader scientific community.[191] consisting entirely of intelligent design supporters. which means that it did not present any new research. a former funder of the Discovery Institute and a major supporter of projects seeking to reconcile science and religion. it appeared in the peer-reviewed journal Proceedings of the Biological Society of Washington in August 2004. shunned.[190] including in its list the two articles mentioned above. noting "Anyone who thinks that scientists do not question Darwinism has never been to an evolutionary conference. on simulation modeling of evolution by Behe and Snoke. the intelligent design people don't come out very well in our world of scientific review". Dembski said that he stopped submitting to peer-reviewed journals because of their slow time-to-print and that he makes more money from publishing books. In his ruling. the Templeton Foundation. Critics. The choice of venue for this article was also considered problematic. intelligent design proponents have set up their own journals with peer review that lacks impartiality and rigor. and that the merit of their articles is overlooked. [196] This claim is similar in type to an assumption of Cartesian dualism that posits a strict separation between "mind" and the material Universe. we assume intelligent cause. selection and the survival of the fittest to solve diverse mathematical and scientific problems that are usually not solvable using conventional methods. if a computer program can access randomness as a function. However. in studies of artificial intelligence. or intentions (which scientists do know when searching for the results of human intelligence). and that "we don't know yet" is a more appropriate response than invoking a cause outside science. appeal to a designing intelligence is indispensable". They contend most scientists would reply that the unexplained is not unexplainable.[195] As a means of criticism. have been used to mathematically demonstrate that randomness and selection can be used to "evolve" complex. where either evolution or design is the proper explanation. Michael Behe's demands for ever more detailed explanations of the historical evolution of molecular systems seem to assume a false dichotomy. artificial intelligence need not be bound to an inflexible system of rules. for the most part.Intelligent design 419 Intelligence as an observable quality The phrase intelligent design makes use of an assumption of the quality of an observable intelligence.[198] In the argument from ignorance. they say. highly adapted structures that are not explicitly designed by a programmer. Intelligence derived from randomness is essentially indistinguishable from the "innate" intelligence associated with biological organisms. disputed Dembski's comparison of SETI and intelligent design. for example. relies on the assumption that intelligence is readily apparent as a fundamental and basic property of complex systems. and any perceived failure of evolution becomes a victory for design. asserts that "in special sciences ranging from forensics to archaeology to SETI (the Search for Extraterrestrial Intelligence). Scott and Branch say that intelligent design is an argument from ignorance because it relies on a lack of knowledge for its conclusion: lacking a natural explanation for certain specific aspects of evolution.[197] Arguments from ignorance Eugenie Scott. Rather. are proposing both searching for a designer without knowing anything about that designer's abilities. certain skeptics have pointed to a challenge of intelligent design derived from the study of artificial intelligence. William Dembski. The characteristics of intelligence are assumed by intelligent design proponents to be observable without specifying what the criteria for the measurement of intelligence should be. that intelligence is irreducible to natural processes". while there is an implicit assumption that supposed "intelligence" or creativity of a computer program is determined by the capabilities given to it by the computer programmer. specifically that "no preprogrammed device can be truly intelligent. undermining the key elements that make it possible as legitimate science. Dembski. Cognitive science continues to investigate the nature of intelligence along these lines of inquiry. parameters. a subfield of machine learning (itself a subfield of artificial intelligence). a lack of evidence for one view is erroneously argued to constitute proof of the correctness of another view. creative. Intelligent design proponents. a researcher with the SETI Institute.[198] Particularly. The criticism is a counter to intelligent design claims about what makes a design intelligent. Seth Shostak. saying that intelligent design advocates base their inference of design on complexity—the argument being that some biological systems are too complex to have been made by natural processes—while SETI researchers are looking primarily for artificiality. and poses a challenge to the intelligent design conception that intelligence itself necessarily requires a designer.[193] How this appeal is made and what this implies as to the definition of intelligence are topics left largely unaddressed.[194] Critics say that the design detection methods proposed by intelligent design proponents are radically different from conventional design detection. has argued that many points raised by intelligent design proponents are arguments from ignorance. and adaptive intelligence. Evolutionary algorithms use the Darwinian metaphor of random mutation. along with Glenn Branch and other critics. has written that "Intelligence leaves behind a characteristic signature". instead. as well as denying the very distinction between natural/artificial design that allows scientists to compare complex designed artifacts against the background of the sorts of complexity found in nature. a concept that has no scientific consensus definition. Scott and Branch also . The intelligent design community. this effectively allows for a flexible. Evolutionary algorithms. said: . and that the school board policy thus violated the Establishment Clause of the First Amendment to the United States Constitution.[202] The suit was tried in a bench trial from September 26. saying in his decision that: "Those who disagree with our holding will likely mark it as the product of an activist judge. A key feature of this type of argument is that it merely answers outstanding questions with explanations (often supernatural) that are unverifiable and ultimately themselves subject to unanswerable questions. and at worst a canard. On December 20. The eight Dover school board members who voted for the intelligent design requirement were all defeated in a November 8. and teachers of the Dover Area School District deserved better than to be dragged into this legal maelstrom.[201] Eleven parents of students in Dover. If so. John G. Rather. Michael Behe. should be taught in science class. and barring intelligent design from being taught in Pennsylvania's Middle District public school science classrooms.Intelligent design contend that the supposedly novel contributions proposed by intelligent design proponents have not served as the basis for any productive scientific research. sued the Dover Area School District over a statement that the school board required be read aloud in ninth-grade science classes when evolution was taught." Reaction Judge Jones himself anticipated that his ruling would be criticized. Intelligent design has also been characterized as a "god of the gaps" argument. 2005 to November 4. Americans United for Separation of Church and State (AU) and Pepper Hamilton LLP. Dover Area School District was the first direct challenge brought in the United States federal courts against a public school district that required the presentation of intelligent design as an alternative to evolution. Associate Director of the Center for Science and Culture at Discovery Institute. The students.[199] A god-of-the-gaps argument is the theological version of an argument from ignorance. 2005 before Judge John E.[199] which has the following form: • There is a gap in scientific knowledge. The National Center for Science Education (NCSE) acted as consultants for the plaintiffs. Barbara Forrest and John Haught served as expert witnesses for the prosecution. This tactic is at best disingenuous. West. this case came to us as the result of the activism of an ill-informed faction on a school board. ruling that the Dover mandate was unconstitutional. The plaintiffs were represented by the American Civil Liberties Union (ACLU). 2005 election by challengers who opposed the teaching of intelligent design in a science class. Judge Jones made the following condemnation of the Teach the Controversy strategy: "Moreover. The breathtaking inanity of the Board's decision is evident when considered against the factual backdrop which has now been fully revealed through this trial. The plaintiffs successfully argued that intelligent design is a form of creationism. aided by a national public interest law firm eager to find a constitutional test case on ID.[200] 420 Kitzmiller trial Kitzmiller v. 2005 Judge Jones issued his 139-page findings of fact and decision."[204] As Jones had predicted. Robert Pennock.[203] In his finding of facts. Kevin Padian. Jones III. parents. and the current school board president stated that the board does not intend to appeal the ruling. ID’s backers have sought to avoid the scientific scrutiny which we have now determined that it cannot withstand by advocating that the controversy. • The gap is filled with acts of God (or intelligent designer) and therefore proves the existence of God (or intelligent designer). they will have erred as this is manifestly not an activist Court. The defendants were represented by the Thomas More Law Center. Pennsylvania. Steve Fuller and Scott Minnich served as expert witnesses for the defense. but not ID itself. who in combination drove the Board to adopt an imprudent and ultimately unconstitutional policy. with its resulting utter waste of monetary and personal resources. Ken Miller. Brian Alters. it described intelligent design as "anti-science" and involving "blatant scientific fraud" and "intellectual deception" that "blurs the nature. argued that intelligent design is a valid scientific theory. the American Academy of Religion issued Guidelines for Teaching About Religion in K‐12 Public Schools in the United States which included guidance that creation science or intelligent design should not be taught in science classes. they. the Council of Europe's Parliamentary Assembly approved a resolution stating that schools should "resist presentation of creationist ideas in any discipline other than religion". The dangers of creationism in education. is not based on facts.[210] In the first. including "intelligent design" which it described as "the latest.[211] while in the third. In the Spring of 2007 the University of Montana Law review published three articles. objectives and limits of science" and links it and other forms of creationism to denialism. He has conflated Discovery Institute's position with that of the Dover school board. West and Casey Luskin. but to "warn against certain tendencies to pass off a belief as science".Intelligent design "The Dover decision is an attempt by an activist federal judge to stop the spread of a scientific idea and even to prevent criticism of Darwinian evolution through government-imposed censorship rather than open debate. however.[219] [220] The DfES subsequently stated that "Intelligent . "presented in a more subtle way". as well as other "worldviews that focus on speculation regarding the origins of life represent another important and relevant form of human inquiry that is appropriately studied in literature or social sciences courses. the decision has been examined in a search for flaws and conclusions. and many "faith schools" that teach the ethos of particular denominations. fear of a similar lawsuit has resulted in other school boards abandoning intelligent design "teach the controversy" proposals. as "Creation science and intelligent design represent worldviews that fall outside of the realm of science that is defined as (and limited to) a method of inquiry based on gathering observable and measurable evidence subject to specific principles of reasoning. must include a diversity of worldviews representing a variety of religious and philosophical perspectives and must avoid privileging one view as more legitimate than others. such as 'intelligent design'.[217] and claimed they were being used by 59 schools. The resolution emphasises that the aim of the report is not to question or to fight a belief."[205] Newspapers have noted with interest that the judge is "a Republican and a churchgoer". which states "Creationism in any of its forms. arguing that the decision was extremely well reasoned and spells the death knell for the intelligent design efforts to introduce creationism in public schools.[10] In April 2010. On October 4. DeWolf et al. public education includes Religious Education as a compulsory subject. 2007. and are not specified in the science curriculum" (part of the National Curriculum which does not apply to independent schools or to Education in Scotland)."[213] 421 Status outside the United States Europe In June 2007 the Council of Europe's "Committee on Culture. John G.[206] [207] [208] [209] Subsequently. Science and Education" issued a report. When it was revealed that a group called Truth in Science had distributed DVDs produced by the Discovery Institute affiliate Illustra Media[216] featuring Discovery Institute fellows making the case for design in nature." However. does not use any scientific reasoning and its contents are pathetically inadequate for science classes. Such study. more refined version of creationism". and that the Kitzmiller decision will have no effect at all on the development and adoption of intelligent design as an alternative to standard evolutionary theory. partly by intelligent design supporters aiming to avoid future defeats in court.[212] However. David K. and he totally misrepresents intelligent design and the motivations of the scientists who research it."[214] In describing the dangers posed to education by teaching creationism.[130] In the second Peter Irons responded. answer the points made by Irons. DeWolf.[218] the Department for Education and Skills (DfES) stated that "Neither creationism nor intelligent design are taught as a subject in schools. all of the Discovery Institute. the Jones court should not have addressed the question of whether it was a scientific theory.[215] In the United Kingdom. and it won't work. could provide the opportunity to explain or explore why they are not considered to be scientific theories and. public meetings promoting intelligent design were sponsored by the local government. as part of a syllabus set by a local Standing Advisory Council on Religious Education. and is not accepted by the science community as a whole. 2007. It states that "Intelligent design lies wholly outside of science".[227] Plans by Dutch Education Minister Maria van der Hoeven to "stimulate an academic debate" on the subject in 2005 caused a severe public backlash. a notable Muslim in Canada. it is not included in the science curriculum". intelligent design and other theories of origin". in Belgium the President of the Flemish Catholic Educational Board (VSKO) Mieke Van Hecke declared that: "Catholic scientists already accepted the theory of evolution for a long time and that intelligent design and creationism doesn't belong in Flemish Catholic schools."[230] 422 Relation to Islam Creationism has strong political support in many Islamic countries. staunch atheist and opponent of intelligent design".[231] In general.[232] Ideas similar to intelligent design have been considered respected intellectual options among Muslims. history or citizenship may deal with creationism and intelligent design in their lessons".[221] In 2006 the Qualifications and Curriculum Authority produced a Religious Education model unit in which pupils can learn about religious and nonreligious views about creationism.[233] .[228] After the 2007 elections she was succeeded by Ronald Plasterk. signed the Scientific Dissent list of the Discovery Institute. why evolution is considered to be a scientific theory". in Northern Ireland the Democratic Unionist Party claims that the revised curriculum provides an opportunity for alternative theories to be taught.[222] [223] On June 25. some use was made of intelligent design antievolution resources. "questions about creationism and intelligent design which arise in science lessons. It's not the tasks of the politics to introduce new ideas. but left the way open for it to be explored in religious education in relation to different beliefs. Muzaffar Iqbal. "Teachers of subjects such as RE.[231] and David Berlinski of the Discovery Institute was the keynote speaker at a meeting in May 2007. Muslim creationists have partnered with the Institute for Creation Research for ideas and materials which they adapted to their own theological positions.[226] In Lisburn the DUP has arranged that the City Council will write to post primary schools asking what their plans are to develop teaching material in relation to "creation.[13] The British Centre for Science Education lobbying group has the goal of "countering creationism within the UK" and has been involved in government lobbying in the UK in this regard. 2007. described as a "molecular geneticist. for example. Similarly. and has sought assurances that pupils will not lose marks if they give creationist or intelligent design answers to science questions.[225] However.[229] As a reaction on this situation in the Netherlands. that's task and goal of science.[224] Detailed government "Creationism teaching guidance" for schools in England was published on September 18. or explanations. However. in the right context. intelligent design and evolution by natural selection. as a result of media coverage.Intelligent design design is not a recognised scientific theory. and in Turkey many intelligent design books have been translated. has no underpinning scientific principles. and antievolutionary views are mainstream among academic theologians and scientists. though teachers would be expected to answer pupils' questions within the standard framework of established scientific theories. therefore. In Istanbul in 2007. the UK Government responded to an e-Petition by saying that creationism and intelligent design should not be taught as science. Though it should not be taught as science. p. the Christian God". Ruling p. In: Wilgoren. aaas. html). Politicized Scholars Put Evolution on the Defensive (http:/ / www. org/ faqs/ dover/ day6pm. New York Times. discovery. is in religious or philosophy classes. Dover Area School District Trial transcript: Day 6 (October 5). Ronald L. Dover Area School District. . Amasino. com/ bnpndxtr/ download/ HorsesMouth-BP007. The Designer of intelligent design is. citizenlink. A selection of writings and quotes of intelligent design supporters demonstrating this identification of the Christian God with the intelligent designer are found in the pdf Horse's Mouth (http:/ / home. org/ spp/ dser/ 03_Areas/ evolution/ issues/ peerreview. Intelligent Design (http:/ / www. And are almost all of the individuals who are involved with the intelligent design movement associated with the Discovery Institute? A. PM Session. 2006. Has the Discovery Institute been a leader in the intelligent design movement? A. discovery. .1172/JCI28449. Elliot Sober. php#questionsAboutIntelligentDesign) [cited 2007-05-13]. [3] The Creationists. Science & Religion Guide. Retrieved 2007-07-20. php?command=download& id=602) (PDF). raised the notion of intelligent design being taught in science classes. Kitzmiller v. • Intelligent Design network. Dover Area School District trial. php?command=view& program=DI Main Page . Who is behind the ID movement? (http:/ / www. Expanded Edition. aclu. • "The engine behind the ID movement is the Discovery Institute". org/ scripts/ viewDB/ filesDB-download. • . [2] Intelligent Design and Evolution Awareness Center. 26. 2008 at the Wayback Machine. discovery. org/ religion/ schools/ 16371res20050916. "CitizenLink: Friday Five: William A. Harvard University Press. (PDF) by Brian Poindexter. . org/ stuff/ contentmgr/ files/ 393410a2d36e9b96329c2faff7e2a4df/ miscdocs/ intelligentdesigntheoryinanutshell. When the former Australian Federal Education Minister. 04 cv 2688 (December 20. Attie. Devon Williams (December 14. TalkOrigins Archive. org/ csc/ topQuestions. Science & Theology News. Retrieved 2007-12-15.News& id=2745). Top Questions-1. The Boston Globe. Dembski" (http:/ / www. ultimately. shtml). kc. Primer: Intelligent Design Theory in a Nutshell (http:/ / www. The Evolution of George Gilder. 2005. pdf) [PDF]." Barbara Forrest. jci. kc. org/ ). [6] "the writings of leading ID proponents reveal that the designer postulated by their argument is the God of Christianity". (http:/ / www. Thomas Powell and Michael M. 2007 [cited 2007-05-13]. rr. doi:10.New York Times. testifying in the Kitzmiller v. September 16. Retrieved 2007-07-20. J. "Defending science education against intelligent design: a call to action" (http:/ / www. org/ scripts/ viewDB/ index. Q. Yes... pdf) [PDF].. November 2005. dated 2003. pdf) June 27. rr.What is the theory of intelligent design? (http:/ / www. Part 1. 2007. stated "I believe God created the world for a purpose. ideacenter.[234] See also • • • • • • Abiogenesis Cosmological argument Creation science Flock of Dodos Flying Spaghetti Monster • • • • • Intelligent falling List of works on intelligent design Neo-Creationism Old Earth creationism Santorum Amendment Watchmaker analogy Argument from poor design • Notes [1] Discovery Institute. 2005).. Kitzmiller v. the Discovery Institute's Center for Science and Culture. 2005 [cited 2007-07-19]. [7] William A. Dembski. July 27. American Association for the Advancement of Science. Alan D. JP. ISBN 0674023390. The Author And Tech-Sector Guru Has A New Cause To Create Controversy With: Intelligent Design (http:/ / www. com/ bnpndxtr/ download/ HorsesMouth-BP007. org/ web/ 20080627021627/ http:/ / home. intelligentdesignnetwork. Numbers. when asked in an interview whether his research concluded that God is the Intelligent Designer. cfm). 2005 [cited 2007-07-19]. 2007). • American Civil Liberties Union. Focus on the Family. the public outcry caused the minister to quickly concede that the correct forum for intelligent design. All of the leaders are. org/ content/ A000006139. archive. A publication of the American Society for Clinical Investigation. [5] "Science and Policy: Intelligent Design and Peer Review" (http:/ / www. if it were to be taught. Terese Berceau. August 21. [4] "Q. pdf) Archived (http:/ / web. Beth Cox4. 373. 2004 [cited 2007-05-13]. edu/ course/ te/ 407/ FS05Sec3/ te408/ files/ Politicized Scholars Put Evolution on the Defensive . yes. html). 379–380. Cox (2006). . Journal of Clinical Investigation 116:1134–1138. Brendan Nelson.Intelligent design 423 Australia The status of intelligent design in Australia is somewhat similar to that in the UK (see: Education in Australia). msu. • "Who's Who of Intelligent Design Proponents" (http:/ / www. talkorigins. Retrieved 2007-07-19. org/ articles/ view/ 28449). • Kahn. 2005 [cited 2007-07-19]. 2005 [cited 2007-07-20]. "The Discovery Institute is the ideological and strategic backbone behind the eruption of skirmishes over science in school districts and state capitals across the country". Richard M. aspx?id=50794). shtml)." Kitzmiller v. who framed the argument as a syllogism: Wherever complex design exists.S. harvard.. Attempts to establish an idea of the 'specified complexity' needed for intelligent design are surrounded by complex mathematics. 2005). 04 cv 2688 (December 20. According to The New York Times "There is no credible scientific challenge to the theory of evolution as an explanation for the complexity and diversity of life on earth". Intelligent design lies wholly outside of science. teachernet. org/ docs/ johnson/ ratzsch. hcs. Office of Public Policy." "This argument for the existence of God was advanced early in the 19th century by Reverend Paley" (the teleological argument) "The only apparent difference between the argument made by Paley and the argument for ID. com/ articles/ ap/ 2007/ 02/ 13/ america/ NA-GEN-US-Kansas-Evolution-History. The AAAS. Meyer and Paul A. "Starting a Conversation about Evolution" (http:/ / www. discovery. Understanding the Intelligent Design Creationist Movement: Its True Nature and Goals. discovery. Schools and Families.000 science teachers and administrators National Science Teachers Association (August 3. the idea seems to be essentially a modern version of the old idea of the "God-of-the-gaps". See also "Evolution of Kansas science standards continues as Darwin's theories regain prominence" (http:/ / www. ncbi. ." [17] David Mu. 04 cv 2688 (December 20. Furthermore they assert that this claim is scientifically testable and should therefore be taught in science lessons. nlm." [18] American Association for the Advancement of Science.7401131 (http:/ / dx. Dover Area School District. 2005).19(1). 2007.embor. archive. and firmly rejects ID (http:/ / www. the immune system and blood clotting mechanisms." [14] Nature Methods Editorial. gov. cfm?id=11890). 2005). which encompasses ID. p. com/ 2007/ 09/ 27/ science/ 27expelled. htm).4(12):983. 04 cv 2688 (December 20. findlaw." [19] Kitzmiller v. nytimes... . . Press release. 1038/ nmeth1207-983). . Dover Area School District. Nelson (May 1. a professional association of 55. Creationism teaching guidance. "We stand with the nation's leading scientific organizations and scientists.8(12):1107–1109. nsta. Retrieved 2007-05-20. 3) The Discovery Institute's A Scientific Dissent From Darwinism petition begun in 2001 has been signed by "over 700 scientists" as of August 20. gov/ pmc/ articles/ PMC2267227/ ). php?command=view& id=1685). com/ media/ voices/ science) on the status intelligent design and other forms of creationism. would also include astrology. including Dr. 1996). org/ scripts/ viewDB/ index. au/ news/ 2005/ intelligent.S.. [15] Mark Greener. Professional Ethics Report (http:/ / www. pdf) [PDF].000 Australian scientists and educators condemn teaching of intelligent design in school science classes (http:/ / web. Document bank" (http:/ / www. Retrieved 2007-10-01. doi. 24. [13] "Teachernet. org/ news/ releases/ 2002/ 1106id2.. centerforinquiry.. ID is not a scientific theory. html?_r=2& oref=slogin& oref=slogin).1038/sj. org/ about/ pressroom. . nih. • • Phillip E. com/ scripts/ getcase. but is rather an old religious argument for the existence of God. 1996). arn. aaas. PMID 18059309. unsw. 2005). A book review. citing Edwards v. 2006. Dover Area School District. iht. Despite this. Cordelia (September 27. org/ 10. "CSC – Getting Rid of the Unfair Rules (http:/ / www. 31 – 33. aaas. Sometimes examples are quoted that are said to require an 'intelligent designer'. 2005). However. Dean. 1038/ sj. 32 ff. Phillip Johnson Files.Intelligent design [8] Kitzmiller v. He traced this argument back to at least Thomas Aquinas in the 13th century. Harvard Science Review. 578 (http:/ / caselaw. 2001. "National Science Teachers Association Disappointed About Intelligent Design Comments Made by President Bush" (http:/ / www. the largest association of scientists in the U. "For most members of the mainstream scientific community. nature is complex. Ignatius Press. . net/ uploads/ attachments/ intelligent-design. Lead defense expert Professor Behe admitted that his broadened definition of science. Johnson (August 31. An intelligently designed response. pl?navby=CASE& court=US& vol=482& page=578) . edu.C. php). . Kitzmiller v. the president's top science advisor. but a creationist pseudoscience". [16] National Science Teachers Association. Fall 2005. lp. Lack of a satisfactory scientific explanation of some phenomena (a 'gap' in scientific knowledge) is claimed to be evidence of an intelligent designer. in stating that intelligent design is not science. Origins & Design"]. org/ spp/ sfrl/ per/ per26. many of these have subsequently been shown to have a scientific explanation. doi:10. php?command=view& id=1780). Which arguments and evidence counter pseudoscience? (http:/ / www. Meyer (December 1. New York Times. there must have been a designer. 482 U. D. EMBO Reports. A four day A Scientific Support for Darwinism petition gained 7733 signatories from scientists opposing ID. 2007).It is simply not fair to present pseudoscience to students in the science classroom. International Herald Tribune. September 18. org/ scripts/ viewDB/ index. [9] "ID is not a new scientific argument. "Creationists are repackaging their message as the pseudoscience of intelligent design theory.1038/nmeth1207-983 (http:/ / dx. doi:10. Ruling. Nat. uk/ docbank/ index. (http:/ / www. Retrieved 2007-09-28. has 120. 424 • • • [12] See: 1) List of scientific societies explicitly rejecting intelligent design 2) Kitzmiller v. Trojan Horse or Legitimate Science: Deconstructing the Debate over Intelligent Design (http:/ / www. 2007. "Scientists Feel Miscast in Film on Life's Origin" (http:/ / www. UK Department for Children. therefore nature must have had an intelligent designer. Methods. 2005). Dover Area School District. 2002). [10] Washington. Context pg. org/ 10. 2007 May [cited 2007-08-06]. Retrieved 2007-05-20.: Center for Inquiry. Dover Area School District. 04 cv 2688 (December 20. Aguillard. February 13. pdf) [PDF]. 04 cv 2688 (December 20. science. Access Research Network. Whether ID Is Science Kitzmiller v. doi. as expressed by defense expert witnesses Behe and Minnich. 7401131). 2007. edu/ ~hsr/ fall2005/ mu. org/ web/ 20060115091707/ http:/ / www. "The intelligent design movement claims there are aspects of the natural world that are so intricate and fit for purpose that they cannot have evolved but must have been created by an 'intelligent designer'. Dover page 83. [11] Stephen C.. 2007. "The Scientific Status of Intelligent Design: The Methodological Equivalence of Naturalistic and Non-Naturalistic Origins Theories" (http:/ / www. pp. Stephen C. Retrieved 2007-05-20. Taking on creationism. html) List of statements from scientific professional organizations (http:/ / ncse.000 members. embor. . pdf) [PDF]. . is that ID's 'official position' does not acknowledge that the designer is God. for example. John Marburger. More than 70.. ) Leading intelligent design proponent William Dembski (2001) argues the opposing view in Is Intelligent Design a form of natural theology? (http:/ / www. 1986 March [cited 2007-10-10]. 001. "Aristotle's analysis of phenomenon and change.D. p. htm) . org/ web/ 20070926220422/ http:/ / www.g. [35] Dembski: "Intelligent design is just the Logos theology of John's Gospel restated in the idiom of information theory. Let's Be Intelligent About Darwin (http:/ / www. htm) [31] Forrest. 2006 [cited 2007-07-27]. 31. [27] William Paley. org/ web/ 20070608233455/ http:/ / www.uk. html). org/ web/ 20070808152431/ http:/ / www. leaderu. Meyer. leaderu. 001. Thaxton. pdf) [PDF]. uk/ AS+ Subjects/ Philosophyofreligion/ fiveways. Touchstone magazine. revised July 1988 and May 1991 [cited 2007-10-06]. Johnson (http:/ / web. Media Backgrounder: Intelligent Design Article Sparks Controversy (http:/ / www. php?command=view& id=2190)." ( Darwin." Johnson 1996. org. php).. arn. R. ca/ news/ social-issues/ 2004/ 03. Dover Area School District. It's about religion and philosophy.. 1993. August 10." Touchstone Magazine. Steinmetz (2005) "The Debate on Intelligent Design" in The Christian Century. thelatinlibrary. Access Research Network. archive. 2005 [cited 2007-07-22]. Ph. origins. ac. edu/ course/ te/ 407/ FS05Sec3/ te408/ files/ Politicized Scholars Put Evolution on the Defensive . design theory demonstrates that Christians can sit in the supernaturalist’s “chair” even in their professional lives.." Johnson 2000. org/ web/ 20070609033219/ http:/ / www. While there are distinctive differences between Paley's argument and those used today by intelligent design theorists and creationists. com/ documents/ 2001. Meyer: "I think the designer is God . ISBN 0812690397. html). Book I. html) Archived (http:/ / web. and numerous other anatomical features throughout the natural world are presented as arguments for God's presence and concern.New York Times. discovery. 1986 [cited 2007-10-10]. the publisher's editorial description of the 2006 paperback printing of William Paley (1803) Natural Theology" : "William Paley's classic brings depth to the history of intelligent design arguments. We Are Not Alone (http:/ / www. Berkeley's Radical (http:/ / touchstonemag. *Richard Hooker (1996) "Aristotle" (http:/ / wsu. [26] Thomas Aquinas." *Monte Ransome Johnson (2005) Aristotle on teleology. The Philosophy of Religion: An Historical Introduction. August 21. php?id=15-05-037-i). html) June 8. "So the question is: "How to win?" That's when I began to develop what you now see full-fledged in the "wedge" strategy: "Stick with the most important thing"—the mechanism and the building up of information. the marketing of Intelligent Design (http:/ / www. Christian Leadership Ministries. is fundamentally teleological. html). com/ story/ 2006/ 3/ 11/ 8448/ 52824) [32] Stephen C. February 1. Letter 2998 — Darwin. Metaphysics Bk. London. Design and the Origin of Life (http:/ / www.org. [25] Linda Trinkaus Zagzebski (2006).. htm) ( archive link (http:/ / web. [22] Stanford Encyclopedia of Philosophy. Open Court Publishing. 140. christianity. ID_as_nat_theol. Nightline ABC News. Christianity. e. 2004. Latin Library (http:/ / www. with Ted Koppel. That means concentrating on.. Barbara. Issue4: July/August. 27–31. Summa Theologiae " Thomas Aquinas' 'Five Ways' (http:/ / web. org/ docs/ meyer/ sm_notalone. dailykos. Volume 12. Asa. 2002 June. [23] Aristotle. christianity. World Magazine. 03. Twelfth Edition. Phrase the argument in such a way that you can get it heard in the secular academy and in a way that tends to unify the religious dissenters.. ca/ news/ social-issues/ 2004/ 03. 04 cv 2688 (December 20. seeing the cosmos through the lens of a comprehensive • • • • 425 . October 25. com/ archives/ article. 36–37. edu/ entries/ plato-timaeus/ ). Know Your Creationists: Know Your Allies (http:/ / www. 1988. com/ docs/ issues/ 15. 2005. com/ offices/ thaxton/ docs/ inpursuit. org/ articles/ thaxton_dnadesign. [28] Darwin Correspondence Project. com/ pjohnson/ world2. edu/ ~dee/ GREECE/ ARIST. touchstonemag.g. (December 27." Johnson 2004. [37] Stephen C. Clarendon Press. C. September 7. The contrivance of the eye. [33] Charles B. archive. 2005. "This isn't really. e. archive. James M. Get the Bible and the Book of Genesis out of the debate because you do not want to raise the so-called Bible-science dichotomy. before the academic world and into the schools. and never has been a debate about science. David C. com/ archives/ issue. htm))" in faithnet. 2005). to/ apologetics/ p90. uk/ content/ frameset?itemID=A142& viewtype=text& pageseq=1). faithnet. Nancy Pearcey: "By contrast. and the sociology of knowledge. htm). com/ 2006-02-01/ news/ discovery-s-creation. stanford. rationality. shtml). De Natura Deorum. com/ cicero/ nd. Jodi Wilgoren. DNA. Politicized Scholars Put Evolution on the Defensive (http:/ / www. Seattle Weekly. php?id=49) [36] Phillip Johnson: "Our strategy has been to change the subject a bit so that we can get the issue of Intelligent Design. which people are always trying to do. org. faithnet. Natural Theology: or. designinference. uk/ AS+ Subjects/ Philosophyofreligion/ fiveways. Thaxton. msu. Plato's Timaeus (http:/ / plato. 2007). bringyou. November 13–16. Evolutionary epistemology. org/ scripts/ viewDB/ index. Kushiner. it remains a fascinating glimpse of the nineteenth-century's debate over the roles of religion and science". html) at the Wayback Machine (archived June 9. 26 Nov (1860) (http:/ / www. pp. [34] Charles B. the ear. darwinproject. June 23–26.ca. [30] See. 2005). archive. 2007 at the Wayback Machine. to Gray. In Pursuit of Intelligent Causes: Some Historical Background (http:/ / www. Evidences of the Existence and Attributes of the Deity (http:/ / darwin-online. 1809. Berkeley's Radical An Interview with Phillip E. • Discovery's Creation (http:/ / seattleweekly. 1999 (http:/ / touchstonemag. which really means the reality of God. 12 [24] See also. Cicero. uk/ entry-2998) [cited 2010-08-11]. then. "Do you need a Creator to do the creating. or can nature do it on its own?" and refusing to get sidetracked onto other issues. [29] Gerard Radnitzky. Witnesses For The Prosecution (http:/ / www.Intelligent design [20] Discovery Institute.. org. HTM). [21] Kitzmiller v. Conclusion of Ruling. 5docs/ 15-5pg40. 2004 [cited 2007-09-24]. org/ content/ 104/ suppl. edu/ cgi/ t/ text/ pageviewer-idx?c=scia.'" [47] Nick Matzke. nih.didno=scia0002-48. [51] Nick Matzke. cited 2009-11-18]. library. html). R. Retrieved 2009-06-02. Dembski. nlm. 2006 [cited 2007-10-05]. . 2007 [cited 2009-06-02]. org/ 10. Bridgham et al. May 23. org/ ctrl/ archive/ design/ aulie_of-pandas. [41] Charles Darwin. org/ 2005/ 12/ post_6.1126/science. The true origin of "intelligent design" (http:/ / pandasthumb.1123348 (http:/ / dx.. html?position=& ei=5090& en=f2de0d764cc7e0e8& ex=1282276800& adxnnl=1& partner=rssuserland& emc=rss& pagewanted=print& adxnnlx=1132902202-gyP0H4EZfG7IeNHPMWlcBw). LC 08031381 "Intelligence-Intelligent Design". org/ 10. "I guess ID really was "Creationism's Trojan Horse" after all" (http:/ / www. Expert_Report_Dembski. [46] Nicholas Timmins. to Herschel. June 29. Jon Buell (October 13. Anarchy. and Utopia. Horigan. com/ sir-fred-hoyle-and-the-origins-of-id/ ). html)" [61] For example. html) [58] The Collapse of "Irreducible Complexity" Kenneth R.0701505104 (http:/ / dx. dtu. New York City. col A.idno=scia0002-48. 1979. ( Total Truth (http:/ / web. NCSE Resource (http:/ / ncse. ISBN 0028949900. Discovery Institute. January 13.. [52] Richard P. Evolution according to Hoyle: Survivors of disaster in an earlier world (http:/ / telicthoughts. pdf) [PDF]. cornell. ncseweb. aspx) [57] Irreducible complexity of these examples is disputed. PMC 1876445 (http:/ / www. London. 1974. ncseweb.0. equipping us to defend it in the public arena". 2006. 1123348).seq=00383. State. This article draws from the following exchange of letters in which Behe admits to sloppy prose and non-logical proof: Discovery Institute. com/ archive/ 2005/ 05/ 30/ 050530fa_fact). Chance or Design?. apologetics. pnas. London: Macmillan Publishing Company. 1998 [cited 2007-10-05]. 23 May 1861 (http:/ / www. The Panda's Thumb. 1850. December 20. Aulie. org/ wp/ ?p=80) 2007-01-14. [50] William Safire. 2005). C. shtml). Evolution of Hormone-Receptor Complexity by Molecular Exploitation. J. Miller Brown University (http:/ / www. 1873:10. org/ archives/ 2006/ 01/ ken_miller_webc. Missing Link discovered! (http:/ / www2. html). National Center for Science Education. ncbi. ektopos. darwinproject. 2006 [cited 2009-11-18]. Winter 1997/98 [cited 2009-02-08].. NCSE Resource -. March 11. html). org/ web/ 20061028204903/ http:/ / rightreason. Intelligent Design steps boldly into the scientific arena to build a case based on empirical data. html)" [60] David Ussery. pp.1073/pnas. Scott. gov/ pmc/ articles/ PMC1876445/ ). F. Johnstone & Hunter. 1/ 8669. Patrick Edward. 2005 [cited 2007-10-05]. cbs. designinference. Matzke (2006): The Story of the Pandas Drafts (http:/ / ncse. Proceedings of the National Academy of Sciences.9. com/ archives/ 2005/ 09/ why_sciencetype. The Times. " A Biochemist's Response to 'The Biochemical Challenge to Evolution' (http:/ / www. March 29. McDonald's " reducibly complex mousetrap (http:/ / udel. 1982:22. com/ documents/ 2005. The theory of human progression. (http:/ / www. html). 2005 [cited 2009-06-02]. May 30. pp. Rethinking Schools Online (http:/ / www. uk/ darwinletters/ calendar/ entry-3154. org/ archives/ 2007/ 08/ the-true-origin. dailykos. millerandlevine. ac. Crossway Books. Bridgham et al. html). PMID 17494747. see Kitzmiller. [43] William P. archive.104(Suppl 1):8669–8676. Letter 3154—Darwin. 1073/ pnas. [44] Robert Nozick. *Nick Matzke. and natural probability of a reign of justice. On Language: Neo-Creo (http:/ / www. Encyclopedia of Philosophy. 2005. The Free Press. org/ cgi/ content/ abstract/ 312/ 5770/ 97) showed that gradual evolutionary mechanisms can produce complex protein-protein interaction systems from simpler precursors. com/ creationism/ analysis/ critique-pandas-people). discovery. Michael (1997): Molecular Machines: Experimental Support for the Design Inference (http:/ / www. edu/ ~mcdonald/ mousetrap. com/ story/ 2006/ 3/ 11/ 8448/ 52824). The New Yorker. [62] Devolution (http:/ / www. org/ wp/ ?p=80). 19. evolutionnews. archive. 76–78. W. [40] Dove. full).view=image. [53] Eugenie C. "F. php?program=CRSC Responses& command=view& 426 .node=scia0002-48:1) [cited 2010-01-21]. Philosophical Library. [45] James E. 2007 [cited 2010-01-21]. 0701505104). [39] Nick Matzke. newyorker. doi:10.312(5770):97–101. 1861. [55] William A. html). Journals: Scientific American (1846 . com/ rncse/ 26/ 1-2/ design-trial). and Ken Miller Webcast (http:/ / www. Alston. PMID 16601189.. It restores Christianity to its status as genuine knowledge. doi. August 14. Hoyle stated in a 1982 speech: '. [42] The British Association. pandasthumb. ISBN 0465097200. August 21. May 15. Science. com/ 2005/ 08/ 21/ magazine/ 21ONLANGUAGE. dk/ staff/ dave/ Behe.. html) [59] John H. It takes Christianity out of the ineffectual realm of value and stakes out a cognitive claim in the realm of objective truth. In: Paul Edwards. [49] DarkSyde. 1967. Biological design in science classrooms (http:/ / www. [56] Behe. doi. Daily Kos: Know Your Creationists: Know Your Allies (http:/ / www. org/ archives/ 2005/ 10/ i_guess_id_real. USA: Basic Books. 1126/ science. 2005. rethinkingschools. The Panda's Thumb. ISBN 1581344589. p. National Center for Science Education. A Reader's Guide to Of Pandas and People (http:/ / www. Has Darwin met his match? Letters—An exchange over ID (http:/ / www. com/ km/ evol/ design2/ article. National Center for Science Education. pandasthumb. sciencemag.one arrives at the conclusion that biomaterials with their amazing measure or order must be the outcome of intelligent design. 2006 [ archived (http:/ / web. org/ scripts/ viewDB/ index. [48] Jonathan Witt. National Association of Biology Teachers.Intelligent design biblical worldview. 204-205) [38] Leon Lynn. doi:10. org/ MolecularMachines/ tabid/ 99/ Default. org/ web/ 20070114121029/ http:/ / www2. [54] Nick Matzke. org/ archive/ 12_02/ panda. 09. nytimes. Collier Macmillan Publishers.1869) (http:/ / digital. Expert Witness Report: The Scientific Status of Intelligent Design (http:/ / www. September 20. stephenjaygould. 2004. Evolution News & Views: Dover Judge Regurgitates Mythological History of Intelligent Design (http:/ / www.rgn=full text.cc=scia. Elliott Sober. [83] Dembski. September 27. discovery. [68] Richard Wein. p. ISBN 0618680004. talkorigins. (PDF file) [78] Victor J. The God Delusion. 000-evolutionary-algorithms-now-surpass-human-designers. July 28. ISBN 0-89526-065-4. v. [79] Joseph Silk. Dover Area School District. [64] Dembski. colorado. edu/ cluster/ pdf/ coyne/ New_Republic_ID. Gerald Feinberg and Robert Shapiro. html) [cited 2007-07-17]. org/ intelligentdesignwatch/ designer. Photo by Wesley R. leaderu. August 22–29.. html). org/ articles/ newmath. 2005. Christopher Stephens. Elsberry. New Scientist. March 26.g. org/ contentmgr/ showdetails. The American Spectator. e. 2005 [cited 2007-02-16]. Washington. colorado. Information Theory and Creationism (http:/ / www.. org/ archives/ 2005/ 12/ 28/ evolutions-thermodynamic-failu). math. org/ design/ faqs/ nfl/ ). html). Biology and Philosophy. org/ web/ 20060618235930/ http:/ / pondside.Intelligent design amp. e. taken at lecture given at University of California at Berkeley." [84] LeaderU. University of Colorado. Creationism's Propaganda Assault on Deep Time and Evolution. 2001. org/ archives/ 2004/ 08/ privileged_plan_4. 220–221.. com/ article/ mg19526146. org/ faqs/ thermo/ entropy. pdf) [PDF]. Discovery Institute. edu/ philosophy/ vstenger/ Cosmo/ FineTune. Christianity postulates the religious answer to this question that the designer is God who by definition is eternally existent and has no origin. Nature. uchicago.16:711–724. The Blind Watchmaker: Why the Evidence of Evolution Reveals a Universe without Design. Dembski. 2004. 49.. pdf) 2006-06-18]. [76] The Panda's Thumb. 2005 [ archived (http:/ / web. (Also available from the Discovery Institute (http:/ / www. jmu. ARN. Rev Max (July–August 2006). this question is essentially irrelevant to intelligent design theory. html). 2005. talkdesign. lhup. wisc. php/ id/ 1147) [88] Jason Rosenhouse. [85] The Case Against Intelligent Design (http:/ / pondside. edu/ sober/ dembski. [73] Richard Dawkins. [67] Some of Dembski's responses to assertions of specified complexity being a tautology can be found at William A. p. html) [77] Is The Universe Fine-Tuned For Us? (http:/ / www. 2003 [cited 2006-11-30]. p. Committee for the Scientific Investigation of Claims of the Paranormal. December 28. Dembski 'displaces Darwinism' mathematically -. 2007. 2002. In: Huchingson. [90] See. Thus. talkorigins. "The Incredibly Strange Story of Intelligent Design". arn. 2005). 2006.. Intelligent Design: The Glass is Empty (http:/ / www. pdf) [PDF]. today. discovery.. [70] Mark Perakh.. Our place in the Multiverse (http:/ / www. [89] Richard Dawkins. [80] See. 47 [65] William Dembski. csicop. 1998. Wilkins. 2002 [cited 2007-10-07]. 2001. org/ scripts/ viewDB/ index. Simanek. [71] Jason Rosenhouse. cfm). "Another way to detect design" (http:/ / www. html). [91] NCSE Resource (http:/ / ncse. . "The theory of Intelligent Design holds that certain features of the universe and of living things are best explained by an intelligent cause. aspx). which merely seeks to detect if an object was designed . A Puddlian Fable. nature. How Not to Detect Design: A review of William A.23(4):3–8. Stenger. php?command=view& id=3122)) [82] TalkOrigins Archive. [66] Branden Fitelson. archive. Joseph Manson. edu/ cluster/ pdf/ coyne/ New_Republic_ID. August 29. Stenger. DC: Regnery Publ. newscientist. There is no logical philosophical impossibility with this being the case (akin to Aristotle's 'unmoved mover') as a religious answer to the origin of the designer. The Act of Creation: Bridging Transcendence and Immanence (http:/ / www. edu/ ~rosenhjd/ sewell. Dover Area School District Ruling [93] Wise.or does he? (http:/ / www. talkorigins." FAQ: Who designed the designer? FAQ: Who designed the designer? (http:/ / www. Disorder and Life (http:/ / www. p. edu/ portal/ ut/ 050927voices_pseudoscience. [75] Guillermo Gonzalez. Fall 2001. Questions About Intelligent Design (http:/ / www. The Privileged Planet: How Our Place in the Cosmos is Designed for Discovery. htm).. [74] Evolutionary algorithms now surpass human designers (http:/ / www. Religion and the Natural Sciences. [86] Donald E. [72] John S.g..233(8/9):21–33. University of Colorado. pdf#search="Fine tuned universe") Victor J. The Anthropic Principle (http:/ / www. org/ csc/ topQuestions. [81] Evolution's Thermodynamic Failure (http:/ / spectator. Journal of Geoscience Education. com/ creationism/ general/ evolving-banners-at-discovery-institute). The Mathematical Intelligencer. September 14. org/ faqs/ information/ dembski. html). htm). p. org/ docs/ dembski/ wd_anotherwaytodetectdesign. 141. pandasthumb. D. Wesley R. ideacenter. edu/ philosophy/ vstenger/ Cosmo/ ant_encyc. 1. php#questionsAboutIntelligentDesign). 2006/03/17. Intelligent design theory cannot address the identity or origin of the designer—it is a philosophical / religious question that lies outside the domain of scientific inquiry. edu/ ~dsimanek/ philosop/ empty. How Anti-Evolutionists Abuse Mathematics (http:/ / www. [87] IDEA "One need not fully understand the origin or identity of the designer to determine that an object was designed. Elsberry. 427 . n. 2005.U. review of The Privileged Planet (http:/ / www. 64. talkreason. The Advantages of Theft over Toil: The Design Inference and Arguing from Ignorance (http:/ / www. Dembski's The Design Inference—Eliminating Chance Through Small Probabilities (http:/ / philosophy. 04 cv 2688 (December 20. Intelligent Design. 1993. com/ offices/ dembski/ docs/ bd-the_ac. 2006. [63] Kitzmiller v. ucla. [92] Conclusion of Kitzmiller v. pdf) [PDF]. Cambridge University Press. Boston: Houghton Mifflin. New Dawn Magazine (97). not an undirected process such as natural selection. Entropy. org/ cs/ theft_over_toil).id=1406).443(7108). The New Republic. Who Designed the Designer? (http:/ / www.. [69] Rich Baldwin. pdf) [PDF]. Intelligent design is pseudoscience (http:/ / www. com/ nature/ journal/ v443/ n7108/ full/ 443145a. uchicago. Not a Free Lunch But a Box of Chocolates: A critique of William Dembski's book No Free Lunch (http:/ / www. 30–35. In: Discovery Institute. asp) [101] Discovery Institute fellows and staff (http:/ / www. Pennock. That stands on its own. 428 . [104] Phillip Johnson. moral character and purposes of this intelligence lie beyond the competence of science and must be left to religion and philosophy". Evolution vs. com/ documents/ 2005. And the materialist scientists are deluding themselves. php). (2) Am I (and the many others who see Dembski's movement in the same way) misrepresenting their position? The basic notion of creationism is the rejection of biological evolution in favor of special creation. Dembski has defined intelligent design. The Bible had that right.. Theological. Wedge Document (http:/ / www.the nature. The Design Inference. org/ docs/ johnson/ le_wedge. . ISBN 0262661241. 1999. org/ library/ modern/ barbara_forrest/ wedge. and continued in Reason in the Balance and Defeating Darwinism by Opening Minds. The very beginning of this strategy. while relatively small. [105] Phillip Johnson.] Now the way that I see the logic of our movement going is like this. [107] Dembski. They involve the supernatural". 2005. The Wedge (http:/ / www. [. org/ scripts/ viewDB/ filesDB-download.. where the latter is understood to be supernatural. Journal of Geoscience Education. As symptoms..19(4):16–17." p.This is not to say that the biblical issues are unimportant. Intelligent Design Creationism and Its Critics: Philosophical. 'Keeping the Darwinists Honest' an interview with Phillip Johnson. purpose. 1998. metanexus. pdf) [PDF]. 645–667. which is devoted to scholarship and writing that furthers this program of questioning the materialistic basis of science.C.. 2004. Massachusetts: MIT Press.the first thing that has to be done is to get the Bible out of the discussion. discovery. [98] Eugenie Scott. 2005. the next question that occurs to you is. And both of those are basically religious. I'm also talking about the definition of intelligent design by Dr. and to replace it with a science consonant with Christian and theistic convictions". pdf) Discovery Institute. Tower of Babel: Evidence Against the New Creationism. B. Reply_to_Henry_Morris.R.. archive.. ISBN 0195157427. the point is rather that the time to address them will be after we have separated materialist prejudice from scientific fact". Design theory promises to reverse the stifling dominance of the materialist worldview. discovery. May. [108] William Dembski.. infidels. It's falsified by all of the evidence and the logic is terrible. The first thing you understand is that the Darwinian theory isn't true.. When you realize that. 1999. [102] Barbara Forrest. arn. 2001. 210... Connecticut: Greenwood Press. Ross. Numbers. [97] Robert T.53(3):319–323. [103] ". 2001. antievolution. Michael Behe's highly successful Darwin's Black Box followed Johnson's work. Massachusetts: MIT Press. Cambridge. does involve a supernatural creator. Truth Sheet # 09-05 Does intelligent design postulate a "supernatural creator? (http:/ / www. Scott. Cambridge. designinference. we must cut it off at its source. htm). org/ files/ nagt/ jge/ abstracts/ Ross_v53n3p319. And I am objecting to it as they have defined it. which has come to be called the theory of intelligent design (ID). htm) [106] William Dembski. Dover Area School District trial transcript. 1999.Intelligent design [94] Marcus R.. Pennock. In: Robert T. Intelligent Design: The Bridge Between Science and Theology. and Gross. NCSE Reports. Evolution and the Wedge of Intelligent Design: The Trojan Horse Strategy. and the Wedge strategy stops working when we are seen as just another way of packaging the Christian evangelical message. Day 6 (October 5) "What I am talking about is the essence of intelligent design. and Scientific Perspectives (http:/ / www. Wizards of ID: Reply to Dembski. Cambridge.. org/ specialdocs/ evolutiondebate. org/ features/ wedge. Dembski as the Logos theology of John's Gospel. org/ csc/ fellows. ISBN 0674023390.. as it is understood by the proponents that we are discussing today. org/ fellows/ ).. net/ magazine/ ArticleDetail/ tabid/ 68/ id/ 2645/ Default. (PDF file) [100] "I have built an intellectual movement in the universities and churches that we call The Wedge. In the beginning was intelligence. where might you get the truth? [. [96] Forrest. broadening the wedge with a positive scientific alternative to materialistic scientific theories. Massachusetts: Harvard University Press. How the Evolution Debate Can Be Won (http:/ / web. The Wedge at Work: Intelligent Design Creationism and Its Critics (http:/ / www. 2006. 1999. [. well. [95] Ronald L. "Center for Science and Culture fellows and staff" (http:/ / www.. Eugenie C.] Intelligent design.intelligent design does not address metaphysical and religious questions such as the nature or identity of the designer. [109] Barbara Forrest. This is precisely our strategy. That source is scientific materialism.. The Creationists: From Scientific Creationism to Intelligent Design. Oxford: Oxford University Press. 2003. aspx). ISBN 0520246500. Expanded Edition. The Creation/Evolution Continuum (http:/ / ncse. can split the trunk when applied at its weakest points.. If we view the predominant materialistic science as a giant tree. He implies (though never explicitly asserts) that he and others in his movement are not creationists and that it is incorrect to discuss them in such terms. and as Dr. coralridge. as Professor Johnson has defined intelligent design. 1999. Beyond this there is considerable variability. "Intelligent Design is an intellectual movement. "Dembski chides me for never using the term "intelligent design" without conjoining it to "creationism". p. Reclaiming America for Christ Conference. Pennock. com/ creationism/ general/ creationevolution-continuum). suggesting that doing so is merely a rhetorical ploy to "rally the troops".. php?command=download& id=565) [cited 2007-07-19]. Touchstone: A Journal of Mere Christianity. those consequences are certainly worth treating. 02. Westport. our strategy is intended to function as a 'wedge' that. Robert T. and the essence of it is theistic realism as defined by Professor Johnson. ". P. ." Johnson 1999.. and I hope our work opens up for them some doors that have been closed"." and ". Intelligent Design's Contribution to the Debate Over Evolution: A Reply to Henry Morris (http:/ / www. Who Believes What? Clearing up Confusion over Intelligent Design and Young-Earth Creationism (http:/ / nagt. Creationism: An Introduction..' was Phillip Johnson's critique of Darwinism begun in 1991 in Darwinism on Trial. Kitzmiller v. discovery. In the beginning was the word. 1999. and wisdom. and that is my objection.. [.] I start with John 1:1. html).] The evangelists do what they do very well. [99] "The social consequences of materialism have been devastating. However. the 'thin edge of the wedge. November 30. Now that stands on its own quite apart from what their motives are. we are convinced that in order to defeat materialism. 23–25. Expert Testimony.. org/ web/ 20071107005414/ http:/ / www.. We are building on this momentum. au/ news/ world/ intelligent-design-a-trojan-horse-says-creationist/ 2005/ 11/ 26/ 1132966007431. August 11.1177/0963662506060588 (http:/ / dx. 2007]. asp). Polling for ID (http:/ / www. com/ harris_poll/ index. July 6. doi. Time Magazine. asp) [ archived (http:/ / web. reasons. org/ web/ 20030816135718/ http:/ / blogs. Joint statement issued by the national science academies of 67 countries. [115] New Mexicans for Science and Reason. archive. htm) [cited 2007-07-13]. "Evolution. • 429 . Design is not enough! (http:/ / www. net/ Object. cited 2007-02-16]. time. Adults Believe Human Beings Were Created by God (http:/ / www. [122] Shaw. com/ 0001092/ 2003/ 07/ 30. Retrieved 2008-10-17. doi:10. [124] The Interacademy Panel on International Issues. California: Institute for Creation Research. 2003 [ archived (http:/ / web. cited 2007-07-13]. nwsource. Answers in Genesis. [126] Intelligent design a Trojan horse.9171. umt. August 7. doi. Retrieved 2008-10-17. File/ Master/ 6/ 150/ Evolution statement. University of Montana Law Review.68(1). imdb. pdf) [PDF].00. Linda. net/ uploads/ attachments/ intelligent-design. edu/ mlr/ Discovery Institute Article. 2002 [cited April 25. org/ web/ 20080327041611/ http:/ / csicop. Nearly Two-thirds of U. 2006. • Television wildlife programming as a source of popular scientific information: a case study of evolution. html). hcdi. org/ sub/ position_statements/ evolution. pdf) [PDF].1207/s15328415jmr0501_3 (http:/ / dx. aaas. [117] David Harris. nmsr. [123] National Association of Biology Teachers. 2007 May. AiG's views on the Intelligent Design movement (http:/ / www." (http:/ / www. 2005 [cited 2007-07-13]. cfm?ID=93). Salon. 2002 [cited April 25. [116] Committee for Skeptical Inquiry. 2005-08-10 [cited 2009-02-16]. "Statement on the Teaching of Evolution" (http:/ / www. pdf) (PDF). 1177/ 0963662506060588). com. org/ docs2005/ 0810time. au. com/ html/ localnews/ 2002225932_design31m. shtml). org/ site/ PageServer?pagename=cs_2005_02_special). pdf) [PDF]. • Answers in Genesis. html) 2003-08-16. salon. answersingenesis. com/ news/ 2005/ 08/ evolution-wars-time-00696). Retrieved 2007-10-08. Understanding the Intelligent Design Creationist Movement: Its True Nature and Goals. More than intelligent design (http:/ / www. Pasadena. Intelligent Design Will Survive Kitzmiller v. . html) [ archived (http:/ / web. 2005 February [cited 2007-07-29]. org/ 10. [121] New Anti-Evolution Film Stirs Controversy (http:/ / www. Press release. March 31. shtml#more_than_id). nabt. org/ news/ releases/ 2006/ 0219boardstatement. org/ docs2002/ 0830_IDM. Dover (http:/ / www. [120] Expelled. gallup. Press release. [119] Gallup. September 11. 2005 [cited 2009-11-18]. David K. Intelligent Design: Creationism's Trojan Horse (http:/ / www. org/ web/ 20060927160040/ http:/ / www. 18% of the physicians believed that God created humans exactly as they appear today. csicop. html). Ross. Office of Public Policy. November 27. livescience. California: Reasons to Believe. 2006). [111] H. A Position Paper (http:/ / www. June 21. harrisinteractive. salon. 2007. archive. asp) 2006-09-27]. [113] Carl Wieland. centerforinquiry. 1207/ s15328415jmr0501_3). org/ sub/ position_statements/ evolution. org/ news/ releases/ 2006/ pdf/ 0219boardstatement. html). Journal of Media and Religion. Public Understanding of Science. • American Association for the Advancement of Science (February 19. [114] Harris Interactive. [112] Henry M. Santee.com. No Intelligence Allowed (2008) (http:/ / www. May 4.Intelligent design [110] Center for Inquiry. doi:10.1090909. 2006 [cited 2008-10-17]. aspx). says creationist (http:/ / www. The poll also found that "an overwhelming majority of Jewish doctors (83%) and half of Catholic doctors (51%) believe that intelligent design is simply "a religiously inspired pseudo-science rather than a legitimate scientific speculation". "The evolution wars" in Time (http:/ / ncse. Does Seattle group "teach controversy" or contribute to it? (http:/ / seattletimes. IAP Statement on the Teaching of Evolution (http:/ / www. Another 42% believed that God initiated and guided an evolutionary process that has led to current human beings. The poll also found that "more than half of Protestant doctors (63%) believe that intelligent design is a "legitimate scientific speculation". org/ article/ 859/ 17/ ). org/ 10. interacademies. creationism. [125] From the world's largest general scientific society: American Association for the Advancement of Science (February 16. aaas. Press release. . 'Intelligent Design'-ers launch new assault on curriculum using lies and deception (http:/ / blogs. [130] DeWolf. [128] The Evolution Wars (http:/ / www. intelligent design. • Journalism and the Debate Over Origins: Newspaper Coverage of Intelligent Design. 2005 [cited 2007-07-23]. [129] National Center for Science Education. archive. 2007]. com/ strangenews/ 080404-expelled-movie. org/ resources/ fff/ 2002issue10/ index. com/ 0001092/ 2003/ 07/ 30. org/ doubtandabout/ polling/ ) 2008-03-27. com/ time/ magazine/ article/ 0. asp).15:131–152. answersingenesis. [127] Americans United for the Separation of Church and State. Retrieved 24 August 2010.S. 2007]. net/ News/ PressRelease. 2005. "AAAS Denounces Anti-Evolution Laws" (http:/ / www. Sandia National Laboratories says that the Intelligent Design Network (IDNet-NM/Zogby) "Lab Poll" is BOGUS! (http:/ / www. icr. org/ doubtandabout/ polling/ ). . Morris. NABT's Statement on Teaching Evolution (http:/ / www. Time for evolution wars (http:/ / www. asp?PID=581). 2006. Sydney Morning Herald. com/ title/ tt1091617/ ). com/ poll/ 21814/ evolution-creationism-intelligent-design. smh. 2005 [cited 2007-07-29]. 1999 [cited April 25. org/ id-poll.5(1):49–61. 2006). [118] According to the poll. including the United Kingdom's Royal Society. "Majority of Physicians Give the Nod to Evolution Over Intelligent Design" (http:/ / www. html). Seattle Times. nabt. which really means the reality of God. is it Christian? (http:/ / www. 2007. ca/ news/ social-issues/ 2004/ 03. Johnson quoted]: We are taking an intuition most people have and making it a scientific and academic enterprise . p. [143] Franklin & Marshall College.3(2):7–29. Science & Theology News. html). [141] What is wrong with intelligent design? (http:/ / philosophy. Intelligent Design: William A. org/ resources/ fff/ 2002issue10/ index. Press release. • The Evolution Wars: Who Is Fighting with Whom about What?. discovery. infidels. christianity. [153] Natan Slifkin (2006). murray/ Providence. 2006 [cited 2007-07-21]. html). 2001 [cited 2007-07-22]. 2007 [cited 2007-07-23]. edu/ michael. org/ web/ 20080114094157/ http:/ / www. doi:10. 001. Intelligent Design as a Theological Problem (http:/ / puffin. php). 1996 [cited 2007-07-23]. [145] Bring You To. 1996 [cited 2008-10-18]. Philo. [148] Creighton University. particularly in biology. Dallas Christian Leadership. com/ creationism/ general/ creationevolution-continuum). html) [cited 2007-07-23].1086/511656 (http:/ / dx. fandm. [151] Harun Yahya International (2007). edu/ NRCSE/ NRCSEPosReID. answersingenesis. [132] Methodological Naturalism and Philosophical Naturalism: Clarifying the Connection (http:/ / www. Vienna (http:/ / www.].11(28):18. 1995. In: Robert B. Johnson quoted]: Our strategy has been to change the subject a bit so that we can get the issue of Intelligent Design. Phillip (http:/ / www." [135] See. christianity. Johnson. org/ 10. "[Phillip E. ca/ news/ social-issues/ 2004/ 03. org/ capabilities_2004/ press02. es/ pdfs/ DarwSciOrPhil. html). What is the position of the NRCSE on the teaching of intelligent design [ID] as an alternative to neo-Darwinian evolution in Nebraska schools? (http:/ / puffin. • The Politically Incorrect Guide to Darwinism and Intelligent Design (http:/ / books. February 4. S. More Than Intelligent Design (http:/ / www." [146] National Center for Science Education. 2000 [cited 2009-11-18]. 2004. ISBN 0800662180. August 30. Catechetical Lecture at St. 2007 [cited 2007-07-23]. [154] Miller. doi.. 2005 [cited 2007-07-23]. org/ articles/ wow/ is-idm-christian). Intelligent design: is it intelligent. • 430 . arn.] [134] Phillip E. 2000 [cited 2007-07-27]. [150] Reasons to Believe. [144] Creighton University. 1992 March [cited 2007-07-23]. to/ apologetics/ p91. Kenneth. com/ pjohnson/ world2. "Purpose and design in the natural world. We are removing the most important cultural roadblock to accepting the role of God as creator. 2006 March [cited 2007-07-17]. php?programs=CSCstories& command=view& id=613). August 31. This means that we affirm that God is objectively real as Creator. 2003 [ archived (http:/ / web. Quarterly Review of Biology. Darwinism: Science or Philosophy (http:/ / ebd10. archive. [has] no difficulty [. Stewart. Science. edu/ NRCSE/ IDTHG. Cambridge University Press.. InterVarsity Press. 1997 February [ archived (http:/ / web. answersingenesis. November 30. 'mere creation'—as the defining concept of our [the ID] movement. htm) [Reprint]. Let's Be Intelligent About Darwin (http:/ / www. html) 2007-06-08. [147] National Council of Churches. creighton. 2006. edu/ cola/ depts/ philosophy/ faculty/ koons/ ntse/ papers/ Vuletic. org/ a/ 2920). edu/ cola/ depts/ philosophy/ faculty/ koons/ ntse/ papers/ Vuletic. com/ new_releases/ news/ intelligent_design. 2005 [cited 2007-07-22]. google. AiG's views on the Intelligent Design Movement (http:/ / www. Debating Design. templeton. Austin. com/ releases/ 2007/ 02/ 070222155420. Stephan's Cathedral. October 2. org/ docs2002/ 0830_IDM. p. harunyahya. pdf) [PDF]. [142] Science Daily. discovery. Law and Education. Regnery Publishing. Templeton Foundation. and the Teaching of Evolution in Public School Science Classes (http:/ / www. 001. google. pdf) [PDF]. reasons.] with the theory of evolution [within] the borders of scientific theory.. Minneapolis. . Forthcoming [cited 2007-07-23]. [136] Los Angeles Times. March 25." [139] The Foundation for Thought and Ethics. com/ ?id=MjKkFG8qVjcC& pg=PA44& vq=evolution+ wars& dq=intelligent+ design+ michael+ ruse+ william+ dembski+ 2007). and the C. The Flagellum Unspun. December 7. for instance: University of Texas. html) [Reprint]." [137] Witnesses For The Prosecution (http:/ / www. Calgary Herald. htm). org/ docs/ johnson/ ratzsch. World Magazine. cited 2007-07-23]. "The "Intelligent Design" Distraction" (http:/ / www. The Challenge of Creation (New York: Yashar Books) 288 ff. creighton. [149] Answers in Genesis. wisc. utexas. [138] The Evangelical Fellowship of Canada. shtml#more_than_id) [cited 2007-07-21]. [152] Answers in Genesis. org/ web/ 20070608233455/ http:/ / www. archive. MN: Fortress Press. ISBN 0830819290. html) 2008-01-14. asp). Retrieved 2007-07-20. Lewis Fellowship. [140] Intelligent design's long march to nowhere (http:/ / www. "My colleagues and I speak of 'theistic realism'—or sometimes.. 273. edu/ sober/ what's wrong with id qrb 2007. org/ library/ modern/ barbara_forrest/ naturalism. ncccusa. Enlisting Science to Find the Fingerprints of a Creator (http:/ / www. Access Research Network.82(1):3–6. ebd. February 23. Dembski & Michael Ruse in Dialogue (http:/ / books. What Is Wrong With Intelligent Design? (http:/ / www. ISBN 1596980133. sciencedaily.Intelligent design Does God have a place in class?: Intelligent design ignites great debate (http:/ / www. [133] Reason in the Balance: The Case Against Naturalism in Science. org/ pdfs/ evolutionbrochurefinal. 2002 [cited 2007-07-21]. html) [Reprint by Leadership U. leaderu. The Creation/Evolution Continuum (http:/ / ncse. bringyou. htm). 2005-08-25 [cited 2009-02-16]. 95. "[Phillip E. cited 2007-07-27].[Johnson positions himself as a "theistic realist" against "methodological naturalism". December 5. Methodological Naturalism and the Supernatural (http:/ / www. Religion. pdf) [PDF]. and that the reality of God is tangibly recorded in evidence accessible to science. 1086/ 511656). csic. pdf) [PDF]. January 10. 2002 [cited 2007-07-20]. before the academic world and into the schools. Natural Providence (or Design Trouble) (http:/ / edisk. utexas. org/ scripts/ viewDB/ index. html). Starting a Conversation about Evolution: Johnson. com/ ?id=H8yn0iaRRfoC& pg=PA1& vq=evolution+ wars& dq=politically+ incorrect+ guide+ to+ intelligent+ design). edu/ ~joelv/ teaching/ 167/ nagel 08 . [167] Intelligent Design is not Science: Scientists and teachers speak out (http:/ / web. org/ index/ science/ comments/ creationism_and_the_daubert_test/ ). [168] PZ Myers. since its existence is typically asserted without sufficient conditions to allow a falsifying observation. John Marburger. Pennock. The ruling discusses central aspects of expectations in the scientific community that a scientific theory be testable. The designer being beyond the realm of the observable. 2005. See. com/ articles/ Skeptic_paper.g. intelligent design cannot be sustained by any further explanation. com/ pdf/ 2005/ 09/ 15/ nobel_letter. Lehigh University. The Elie Wiesel Foundation for Humanity. 1999. 04 cv 2688 (December 20. . p. designer. Branden Fitelson. Kitzmiller v. edu.public education and intelligent design. Adding entities (an intelligent agent. p. nsta.. e. 196–197. The October 2005 statement. Mark Perakh. 2005.. pdf) (PDF). cfm). [159] Kitzmiller v. University of New South Wales. "We stand with the nation's leading scientific organizations and scientists." [163] The designer is not falsifiable. Retrieved 2007-07-19. archive. pdf) [PDF]. and parsimony. doi:10.000 science teachers and administrators National Science Teachers Association (August 3. How Not to Detect Design–Critical Notice: William A.. e. au/ news/ 2005/ intelligent. January 11. stanford. 1038/ 416250a). e. See. Creationism and the Daubert test? (http:/ / pharyngula. The idea of the progressive growth of scientific ideas is required to explain previous data and any previously unexplainable data. [170] National Academy of Sciences. org/ scripts/ viewDB/ index. science. in stating that intelligent design is not science. Nature. By asserting a conclusion that cannot be accounted for scientifically. claims about its existence can be neither supported nor undermined by observation.416(6878):250.. testability. Dover Area School District. archive. pdf) 2007-07-24. December 1. unsw. p. or "Occam's razor". and Scientific Perspectives. Chapter 8 discusses parsimony (Occam's razor) [158] Research Methods in Psychology. 2005. 2005). correctible. php?isbn=0309064066& page=25). lehigh. 22 and p. it cannot be tested as scientific theory because its central conclusion is based on belief in the intervention of a supernatural agent". Chapter 2. [169] Nagel. 2005). a designer) to the equation is not strictly necessary to explain events. [161] Intelligent design fails to pass Occam's razor. including the principles of falsifiability. because it fails to qualify on every count as a scientific theory". nap. Science and Creationism: A View from the National Academy of Sciences (http:/ / www. [160] See. the brief explanation in Kitzmiller v. Ruling. naturalism. cv 2688 (December 20.Intelligent design [155] Center for Science and Culture. Once it is claimed that a conclusion that need not be accounted for has been established. 2005. Talk Reason. Dover Area School District. and provisional. [156] University of California at San Diego. and say that it is being used as a Trojan Horse to introduce the teaching of creationism into schools" 431 . lacks supporting data and terminates further thought. ucsd. a professional association of 55. Theological. Department of Biological Sciences. See... "But many scientists regard ‘intelligent design’ as pseudoscience. p. aspx?id=50794).. but when presented as a scientific hypothesis. Thoughts on Evolution and Intelligent Design (http:/ / www. Discusses the scientific method.It is simply not fair to present pseudoscience to students in the science classroom. cv 2688 (December 20. by a coalition representing more than 70. Vol. "Q: Why couldn't intelligent design also be a scientific theory? A: The idea of intelligent design might or might not be true. 2002. doi. making intelligent design and the argument from design analytic a posteriori arguments. . Chapters 5–8. 77. p. 2005)." [172] Evolution critics seek role for unseen hand in education. 2005).. Wadsworth Publishing. [166] "Nobel Laureates Initiative" (http:/ / media. no. org/ web/ 20060614003243/ www.. The Dream World of William Dembski's Creationism (http:/ / talkreason.org. edu/ faculty/ wuthrich/ teaching/ 2007_145/ Lecture02. "National Science Teachers Association Disappointed About Intelligent Design Comments Made by President Bush" (http:/ / www.1038/416250a (http:/ / dx. e. The September 2005 statement by 38 Nobel laureates stated that: "Intelligent design is fundamentally unscientific. pdf). 36.g. php?command=view& id=1780).000 Australian scientists and science teachers said: "intelligent design is not science" and called on "all schools not to teach Intelligent Design (ID) as science. 2002 [cited 2007-07-19]. Thus intelligent design is not a provisional assessment of data which can change when new information is discovered. See. html). 2005 October [cited 2009-01-09]. the intelligent agent. ISBN 0521017084. 54–65. org/ about/ pressroom. Discovery Institute. there is simply no possibility of future correction. e. discovery. [162] See. 2007 [ archived (http:/ / web. 2008. htm). PMID 11907537. [171] National Science Teachers Association. 2001. testability. Dembski The Design Inference. it is not useful because it is based on weak assumptions.g. ljworld. edu/ openbook. . September 9. 66. the president's top science advisor. progressive. Kitzmiller v. dynamic self-correcting of hypotheses. Pharyngula. Philosophy and Public Affairs. 2. dynamic. based upon multiple observations. Ruling. et al. Discusses principles of induction. and objections raised to those who accept intelligent design make little headway. edu/ ~inbios/ schneider/ evolution. Dover Area School District. Thomas. 22 and p. [157] Scientific Method in Practice. org/ 10.. In: Robert T.g. 4: whether ID is science. Cambridge UP. Demarcating science vis-à-vis pseudoscience (http:/ / philosophyfaculty. ISBN 0534609767. Dover Area School District. 597–616. ucsd. [165] Intelligent design professes to offer an answer that does not need to be defined or explained. Press release. 8th ed. 66. and multiple observations. May 21. The Scientific Status of Intelligent Design: The Methodological Equivalence of Naturalistic and Non-Naturalistic Origins Theories (http:/ / www. pp. 2005. "Public Education and Intelligent Design" (http:/ / www. progressive development of theory. MIT Press. 2003. including Dr. 2005). cv 2688 (December 20. deduction and probability related to the expectation of consistency.g. org/ web/ 20070724203349/ http:/ / philosophyfaculty. [164] That intelligent design is not empirically testable stems from the fact that it violates a basic premise of science. cited 2007-07-19]. the designer. e.g. edu/ faculty/ wuthrich/ teaching/ 2007_145/ Lecture02. Intelligent Design Creationism and Its Critics: Philosophical. " • Robert T. [178] Skeptic: eSkeptic: Thursday. The President and the teaching of evolution (http:/ / johnhawks. September 2005 [cited 16 March 2009]. October 2004 [cited 16 March 2009]. Index to Creationist Claims (http:/ / www. Devolution—Why intelligent design isn't (http:/ / www. talkorigins. do not propose a mechanism. PMID 16670753. [183] Design Inference Website. com/ eskeptic/ 08-04-17.041171805 (http:/ / dx. net/ weblog/ topics/ creation/ bush_intelligent_design_2005. [. needs. the signals actually sought by today's SETI searches are not complex. while the other has a more overtly philosophical orientation. weaknesses. February 25. 4: whether ID is science [188] Simple evolutionary pathways to complex proteins (http:/ / www. Protein Science. 2005). New York Times. html). 1110/ ps. gov/ pmc/ articles/ PMC2253472/ ). org/ scripts/ viewDB/ index. 1172/ JCI28449). php?command=view& id=2640& program=CSC . [194] Space. Peer-Reviewed. Whether ID is Science. one has languished for want of material and quietly ceased publication. 2007 July [cited 2007-07-17]. [189] Kitzmiller v. nlm. doi:10. as the ID advocates assume. p.C. com/ documents/ 2001. it would be on the basis of artificiality" [195] "For human artifacts. Washington University Law Quarterly. SETI and Intelligent Design (http:/ / www. gov/ pmc/ articles/ PMC1451210/ ). World Magazine. and the designer. 03. PMC 1451210 (http:/ / www. Defending science education against intelligent design: a call to action (http:/ / www. nih. com/ ?id=kHeQhdNQvrUC& pg=PA210& lpg=PA210& dq=intelligent+ design+ junk-science). Dover Area School District. org/ faqs/ dover/ day12am. cv 2688 (December 20. htm). ID_as_nat_theol. edu/ inprint/ 83-1/ p 1 Brauer Forrest Gey book pages.13(10):2651–64. 2006. 210 ff. nlm. we know the designer's identity.83(1). html). php) Discovery Institute. discovery. human. October 19.1172/JCI28449 (http:/ / dx. doi. wustl. skeptic. [184] Darwinism Under Attack (http:/ / chronicle. org/ web/ 20070926214521/ http:/ / www. and significant omissions in fact and argument. biolsocwash. "Biologists aren't alarmed by intelligent design's arrival in Dover and elsewhere because they have all sworn allegiance to atheistic materialism. htm). and other Scientific Publications Supporting the Theory of Intelligent Design (Annotated) (http:/ / www. nih. google. org/ 10. Journal of Clinical Investigation. 04802904). doi. talkorigins. archive. org/ 10. Peer-Edited. December 4. S.Intelligent design [173] Attie. 2005 December [cited 2007-07-18]." [192] TalkOrigins Archive. as well as many other attributes including the designer's abilities. org/ scripts/ viewDB/ filesDB-download. 2004 [cited 2007-05-10].14(9):2217–2225. Allen Orr. ISBN 9780312352417.1110/ps. Proceedings of the Biological Society of Washington. discovery. With ID. org/ indexcc/ CI/ CI001_4. 04 cv 2688 (December 20. D. org/ evolution/ nhmag. New Yorker. com/ 2005/ 12/ 04/ weekinreview/ 04good. 2001 [cited 2007-07-19].117(2):213–239. com/ archive/ 2005/ 05/ 30/ 050530fa_fact). The origin of biological information and the higher taxonomic categories (http:/ / www. Detecting Design in the Natural Sciences (http:/ / www.. html). [191] Is It Science Yet?: Intelligent Design Creationism and the Constitution (http:/ / lawreview. December 21. Macmillan. worldmag. html#part1) [179] Intelligent Design Might Be Meeting Its Maker (http:/ / www. Kitzmiller v. and the mechanism of design. ncbi. AM session Kitzmiller Testimony. 2001 [cited 2008-12-10]. [181] Meyer. nlm. com/ free/ v48/ i17/ 17a00801. nih. pdf) [PDF]. com/ searchforlife/ seti_intelligentdesign_051201. org/ csc/ freeSpeechEvolCampMain. However. 4. "ID leaders know the benefits of submitting their work to independent review and have established at least two purportedly "peer-reviewed" journals for ID articles. Dover Area School District. The purpose of peer review is to expose errors. 2008 (http:/ / www. doi:10. 2006. ncbi. [185] Ruling. designinference. Pennock Tower of Babel: The Evidence Against the New Creationism. doi. That purpose is not served if the reviewers are uncritical" [193] Natural History Magazine. 2007). proponents assert that they refuse to propose hypotheses on the designer's identity. ncbi. H. the reviewers are themselves ardent supporters of intelligent design. html) at the Wayback Machine (archived September 26. html) [190] Discovery Institute. p. PMID 16131652. 2005. org/ 10. html?ex=1291352400& en=feb5138e425b9001& ei=5088& partner=rssnyt& emc=rss). Both journals employ a weak standard of "peer review" that amounts to no more than vetting by the editorial board or society fellows. "With some of the claims for peer review. org/ id_statement. Dover Area School District 4: whether ID is science [186] Simulating evolution by gene duplication of protein features that require multiple amino acid residues (http:/ / www3.1110/ps. org/ articles/ view/ 28449). html). Dover Area School District. 2005 [cited 2007-07-18]. wiley. [187] Kitzmiller v. Junk science (http:/ / www. and desires. Protein Science.. actionbioscience. 2005 August [cited 2007-07-19]. they're alarmed because intelligent design is junk science. "In fact. 2002 April [cited 2007-07-18]. nytimes.. Center for Science and Culture.Science). [174] Junk Science (http:/ / books. PMID 15340163. 2006. jci. php?command=download& id=549).. PMC 2286568 (http:/ / www. [182] The Sternberg peer review controversy and several similar academic disputes are the subject of the 2008 documentary "Expelled: No Intelligence Allowed". 2005). [175] Kitzmiller v. April 17th. com/ cgi-bin/ fulltext/ 121602290/ HTMLSTART).] If SETI were to announce that we're not alone because it had detected a signal.. • 432 . notably Campbell and Meyer (2003) and the e-journal PCID. com/ articles/ 11553).Scientific Research and Scholarship .116(5):1134–1138. 2006. • Mark Bergin. gov/ pmc/ articles/ PMC2286568/ ). 041171805). 2005 May. discovery. 2005 [cited 2007-07-19]. 1110/ ps. as we have experience based upon empirical evidence that humans can make such things. doi:10.com. Behe (http:/ / www. 87 [176] Free Speech on Evolution Campaign Main Page (http:/ / www. space. interscience. Is Intelligent Design a Form of Natural Theology? (http:/ / www. A.04802904 (http:/ / dx. newyorker. [180] Statement from the Council of the Biological Society of Washington (http:/ / web. [177] John Hawks Weblog. 780fc9a. csicop.Intelligent design he/she/it/they. bbc. [206] Judge rules against 'intelligent design' (http:/ / www. West.'. [212] DeWolf. [203] Judge Rules Against 'Intelligent Design' (http:/ / pewforum. Guardian. uk/ science/ 2006/ nov/ 27/ controversiesinscience. 2006 [cited 2009-11-18].14:203–220. guardian. org/ news/ display. co. September 29. pdf) [PDF]. pp. umt. 2005 [cited 2007-09-03]. April 10. defense expert Professor Minnich agreed that in the case of human artifacts and objects. 1971. html). com/ news/ 2007/ 10/ council-europe-approves-resolution-against-creationism-001200). org/ web/ 20011018142820/ http:/ / www. 2007-10-04 [cited 2007-10-05]. July 15. 2005 [cited 2007-09-03]. com/ news/ 2010/ 07/ american-academy-religion-teaching-creationism-005712). com/ id/ 10545387/ ). MSNBC. gov/ kitzmiller/ 04cv2688-111. archive. coe. Dover Area School District. [197] Intelligent Design and Evolution Awareness Center. html) 2001-10-18. org/ si/ 2001-03/ intelligent-design. org/ contentmgr/ showdetails. org/ scripts/ viewDB/ index.68(1). 2006 [cited 2007-07-13]. NCSE Resource—Council of Europe approves resolution against creationism (http:/ / ncse. (London) November 27. December 21. htm) [cited 2007-08-03]. Dover Intelligent Design Decision Criticized as a Futile Attempt to Censor Science Education (http:/ / www. Links to both the article and a response by the Discovery Institute (our partners in the production of Unlocking the Mystery of Life and The Privileged Planet)" [217] Center for Science and Culture. December 20. [216] Illustra Media. "Intelligent Design" Not Accepted by Most Scientists (http:/ / ncse. et al. htm#0611021004183). org/ 2005/ 12/ dover_intelligent_design_decis. html). skeptic. Peter. December 20.Video and Curriculum . php?command=view& id=2116& program=CSC . php?NewsID=5945).Multimedia)]]. php/ id/ 1136). msnbc. we know the designer and its attributes and we have a baseline for human design that does not exist for design of biological systems. cited 2007-07-17]. In addition. 2004 [cited 2007-07-13]. Associated Press.68(1). 2006 [cited 2007-07-13]. [207] Godless: The Church of Liberalism (http:/ / www. [215] National Center for Science Education. 3). Luskin. publications. University of Montana Law Review. [199] Del Ratzsch (2005) "Teleological Arguments for God's Existence". 2005). In that vein. 2006 [cited 2007-09-03]. 2005 [ archived (http:/ / web. int/ Main. but we do not know any of those attributes for the designer of biological life. php?NewsID=5945) 2007-09-28. pdf) [PDF]. November 1. illustramedia. com/ creationism/ general/ intelligent-design-not-accepted-by-most-scientists).. Journal of the Evangelical Theological Society. [214] Council of Europe. v. uk/ pa/ cm200506/ cmhansrd/ cm061101/ text/ 61101w0010. July 23. umt. Dallas Morning News. 2007 [cited 2008-10-16]. 2010 [cited 2010-08-09]. csicop. [220] Parliament of the United Kingdom. com/ ID01WiredMagPage. Casey. com/ sharedcontent/ dws/ dn/ latestnews/ stories/ 122105dnnatidesign. 2007 [cited 2007-07-14]. pdf) [PDF]. 2002 September [cited 2009-11-18]. April 27. (http:/ / www. edu/ entries/ teleological-arguments/ #4. in Stanford Encyclopedia of Philosophy. html). Reuters. com/ creationism/ general/ discovery-institute-tries-to-swift-boat-judge-jones). ideacenter. Discovery Institute tries to "swift-boat" Judge Jones (http:/ / ncse. et al. pdf) . April 27. 2008 [202] Tammy Kitzmiller. Dover Area School District. Section 4. October 4. The dangers of creationism in education (http:/ / assembly. 2005 [cited 2008-10-17]. [218] Revealed: rise of creationism in UK schools (http:/ / www. Professor Behe agreed that for the design of human artifacts. [208] National Center for Science Education. edu/ mlr/ Editors' Note. BBC News. David K. University of Montana Law Review. Disaster In Dover: The Trials (And Tribulations) Of Intelligent Design (http:/ / www. we know the identity and capacities of the human designer. Primer: Intelligent Design Theory in a Nutshell (http:/ / www. National Center for Science Education. 137 – 138. com/ eskeptic/ 06-09-21. [[Unlocking the Mystery of Life (http:/ / www. evolutionnews. htm) [cited 2007-07-13]. American Academy of Religion on teaching creationism (http:/ / ncse. • 433 . January 4. University of Montana Law Review. [204] Kitzmiller v.3. 2006 [cited 2008-10-17]. com/ article/ scienceNewsMolt/ idUKL0417855220071004). Council of Europe firmly opposes creationism in school (http:/ / uk. reuters. archive. The “Intelligent Design” (ID) Movement (http:/ / plato. discovery. co. [205] Discovery Institute. December 20. org/ si/ 2001-03/ intelligent-design. religion). [200] See.68(1). [209] Intelligent design policy struck down (http:/ / www. 2007 [cited 2009-11-18]. 81 [196] Skeptical Inquirer Magazine. asp?link=/ Documents/ WorkingDocs/ Doc07/ EDOC11297. Rebuttal to Irons (http:/ / www. org/ web/ 20070928055938/ http:/ / pewforum. 04 cv 2688 (December 20. The Washington Post. [210] Articles: Editor's Note: Intelligent Design Articles (http:/ / www. parliament. edu/ mlr/ Irons Response. Written Answers (http:/ / www. Dover (http:/ / ncse. 2007 [cited 2008-10-16]. September 21. 2007 [cited 2008-10-16].—Kitzmiller v. org/ news/ display. Darwin in Mind: Intelligent Design Meets Artificial Intelligence (http:/ / www. Dover Area School District. dallasnews. [219] 'Design' attack on school science (http:/ / news. stanford. [198] National Center for Science Education. edu/ mlr/ DI Rebuttal. [201] Intelligent Design on Trial: Kitzmiller v. Professor Behe's only response to these seemingly insurmountable points of disanalogy was that the inference still works in science fiction movies". 2001 March/April [ archived (http:/ / web. WIRED Magazine response (http:/ / www. stm). cv 2688 (December 20. umt. October 17th. uk/ 1/ hi/ education/ 5392096. msn. pamd. has never been seen. [213] National Center for Science Education. uscourts. John G. p. Discovery Institute. com/ creationism/ legal/ intelligent-design-trial-kitzmiller-v-dover).. "It's also important that you read a well developed rebuttal to Wired's misleading accusations. html). for instance: Man Come Of Age: Bonhoeffer's Response To The God-Of-The-Gaps. 2005). cited 2007-09-03]. [211] Irons. com.ca/books?id=raqJjM9LkeAC& printsec=frontcover#v=onepage&q&f=false) at Google Books • Humes E. [232] National Center for Science Education.nl. How can we answer questions about creation and origins? (http:/ / www. [230] De Morgen. uk/ pa/ ld200607/ ldhansrd/ text/ 61218w0006.1126/science. New Brunswick. September 27. [222] NCSE. Cloning Creationism in Turkey (http:/ / ncse. Preview (http://books. org/ cgi/ content/ summary/ 308/ 5727/ 1394b). 2007. sciencemag. 2007 [cited 2008-10-17]. ISBN 0-06-088548-3. lisburntoday. 2005 [cited 2007-07-13]. [225] Lengagne. Dembski (http://www. 3233193. ISBN 0-19-515742-7. nl/ news/ archives/ print/ 001501. Monkey Girl: Evolution. The Battle over the Meaning of Everything: Evolution. php). [226] Henry.Y: ECCO. Forrest B. com/ rncse/ 19/ 6/ cloning-creationism-turkey) [cited 2009-11-18]. Martin. ethz. Taner. Science. 1126/ science. 2005 [231] Edis. June 8. 11297. 434 References Further reading • Gross PR.J: Rutgers University Press. New York. May 23. com/ news/ 2007/ 02/ guidance-creationism-british-teachers-001170). Intelligent Design. and a School Board in Dover. Oxford [Oxfordshire]: Oxford University Press. dutchnews. htm). Why Intelligent Design Fails: A Scientific Critique of the New Creationism. uk/ Page12021). 21 June 2007 [cited 2007-07-13]. org/ 10. according to Lisburn folk (http:/ / www. gov. [227] Dup Call For Schools To Teach Creation Passed By Council—Lisburn Today (http:/ / www. Education. Lesley-Anne. Religion. Tussle of Biblical proportions over creationism in Ulster classrooms (http:/ / www. 2007 [cited 2007-10-01]. doi:10. html). * Viewpoint: The world. 2008 January [cited 2008-02-23]. Preview (http://books. London: Her Majesty's Stationery Office. ISN Security Watch. 2007. uk/ news/ Creation-Motion-Passed-By-Council. Young M. 2007 [cited 2009-11-18]. DutchNews. PA. 2006 [cited 2007-10-01].google. co. The History of Science Society : The Society (http:/ / www. September 25. au/ news/ national/ intelligent-design-not-science-experts/ 2005/ 10/ 20/ 1129775902661.1394b (http:/ / dx. Guy. PMID 15933170.308. [223] Qualifications and Curriculum Authority for England. p. coe. Schools: Intelligent Design (http:/ / www. org/ publications/ Newsletter2008/ NewsletterJanuary2008Creationism. uk/ opinion/ viewpoint-the-world-according-to-lisburn-folk-13479536. Doc. pdf) [PDF]. June 3. jp). March 12. uk/ news/ education/ tussle-of-biblical-proportions-over-creationism-in-ulster-classrooms-13479246. uk/ libraryAssets/ media/ qca-06-2728_y9_science_religion_master. int/ Main. doi. 2007 [cited 2007-10-01]. and the Battle for America's Soul. N. smh. • Slack G. qca.epetition reply (http:/ / www. [228] Enserink.google. html). isn. co. hssonline. 5727. • Edis T. 2004. NCSE Resource—Guidance on creationism for British teachers (http:/ / ncse. 2007 [cited 2008-05-31].arn. ISBN 978-0-8050-8306-4. Belfast Telegraph.Intelligent design [221] Parliament of the United Kingdom. Creationism's Trojan Horse: the Wedge of Intelligent Design. [233] Jones.308(5727):1394b.google. 2007. 2006. belfasttelegraph. assembly. number10. belfasttelegraph. htm). Ulster Star. History of Science Society. N. October 21. September 26. Belfast Telegraph. ISBN 0-8135-3872-6.org/) • Design Inference: The website of William A.ca/ books?id=hYLKdtlVeQgC&printsec=frontcover#v=onepage&q&f=false) at Google Books • Why Darwin Matters: The Case Against Intelligent Design. February 13. [224] Prime Minister's Office. Evolution Politics: Is Holland Becoming the Kansas of Europe? (http:/ / www. Council of Europe. Sydney Morning Herald. [229] Cabinet ministers announced (update 2) (http:/ / www.5727. [234] Intelligent design not science: experts (http:/ / www. 308. The dangers of creationism in education (http:/ / www. Turkey's survival of the fittest (http:/ / www. Preview (http://books. NoCreSciEd . 2006 [cited 2007-07-13]. San Francisco: Jossey-Bass. asp?link=/ Documents/ WorkingDocs/ Doc07/ EDOC11297. 2005.com/) .designinference. co. September 26. org. publications.ca/ books?id=1kpx8HrQ08cC&printsec=frontcover#v=onepage&q&f=false) at Google Books ID perspectives • Access Research Network (http://www. New York: Henry Holt. Dorian L. 1394b). 2007 [cited 2008-10-17]. html). Newsletter. December 18. 2008 [cited 2008-03-13]. parliament. ch/ isn/ Current-Affairs/ Security-Watch/ Detail/ ?id=54183& lng=en). ISBN 0-7879-8786-7. html). org/news/ releases/2002/1106id2.pbs.origins) • 139 page in-depth analysis of intelligent design.129. (Seattle Weekly) • Intelligent Design vs. and Design (ISCID) (http://www.Natural-Knowledge-and-Natural-Design.Richard-Dawkins) http://web.pitt.utm.(PBS) • Discovery's Creation (http://www.edu/d/ design. irreducible complexity.edu/forumy/2005/12/ kitzmiller-intelligent-ruling-on.edu/openbook.org/intelligentdesignwatch/differences.com/) William Dembski's blog Non-ID perspectives • Scientific American .evolutionnews.nap.iscid.discovery.wisc.com/) A multiple contributor weblog by Discovery Institute fellows. cfm?id=15-answers-to-creationist) • ACLU site on Intelligent Design (http://www.pdf) Elliott Sober.com/darwinanddesign.com/2006-02-01/news/discovery-s-creation/) An overview of the origin of the intelligent design movement.org/) • Uncommon Descent (http://www.org/) (Archive of the UseNet discussion group talk.washingtonpost.uncommondescent.html) • The Design Argument (http://philosophy. PA Intelligent Design trial in 2005.archive. org/web/20080307082606/http://richarddawkins.com/wp-dyn/content/article/2005/12/16/ AR2005121601559.uscourts. • Short discussion on Natural Knowledge and Natural Design as a contrast to Intelligent Design (http:// richarddawkins.php).naturalhistorymag. Center for Science and Culture (http://www.pdf) by the Kitzmiller v. Evolution (http://www.gov/kitzmiller/kitzmiller_342.Intelligent design • Discovery Institute.aclu.law.com/article. • ID The Future (http://www.tvw.talkorigins.aaas.Richard-Dawkins by Richard Dawkins Media articles • Judgment Day: Intelligent Design on Trial (http://www.org/) (Hub of the intelligent design movement) • EvolutionNews.html) What would "intelligent design" science classes look like? All we have to do is look inside some 19th century textbooks.net/article.org (http://www.129.org/MediaPlayer/Archived/WME.Natural-Knowledge-and-Natural-Design.archive. (The Washington Post) 435 .sciam.org/religion/intelligentdesign/index.com/creationism/ general/what-is-intelligent-design-creationism) • Resolution from the American Association for the Advancement of Science (http://www.15 Answers to Creationist Questions (http://www.edu/sober/design argument 11 2004.org/wgbh/nova/id/) A PBS-NOVA documentary on the Dover.shtml) • Science and Creationism: A View from the National Academy of Sciences (http://www. • International Society for Complexity.com/darwinanddesign.org/ web/20070620122151/http://www. php?isbn=0309064066) Second Edition (1999) • Talk Origins Archive (http://www.pamd.iep.html) • Intelligent Design? (http://www. Information.htm) • National Center for Science Education What Is Intelligent Design Creationism? (http://ncse. Dover Area School District judge • Kitzmiller: An Intelligent Ruling on 'Intelligent Design' (http://jurist. and the book Of Pandas and People (http://www.net/ article.org/) Discovery Institute website tracking media coverage of intelligent design.html) http://web.csicop.seattleweekly.naturalhistorymag. 2004.idthefuture. cfm?EVNum=2006040103&TYPE=V) debate between paleontologist Peter Ward and Stephen Meyer co-founder of the Discovery Institute • Intelligent Design Deja Vu (http://www. JURIST • ID and Creationism (http://www.html special feature in the Natural History Magazine • Internet Encyclopedia of Philosophy: Design Arguments for the Existence of God (http://www. [2] Darwin himself addressed eugenic concepts and denounced them as "evil" in his book. though it was never universally accepted by either. explanations." Later advocates of this theory would suggest radical and often coercive social measures to attempt to "correct" this imbalance. . Some have vigorously opposed acceptance of the scientific explanation due to its perceived religious implications (e. Weikart claims that Darwinism's impact on ethics .Charles Darwin. Thomas Huxley spent much time demonstrating through a series of thought experiments that it would not only be immoral. and at no point in Nazi literature is Charles Darwin or the scientific theory of evolution mentioned.justicetalking. without deterioration in the noblest part of our nature. 1871 Neo-creationist Polemics single out "Darwinism" as the cause of many.8599. it could only be for a contingent benefit. it greatly fell out of favor with public and scientific opinion.html) (Christian Science Monitor) • Devolution (http://www. of modern society's ills. The surgeon may harden himself whilst performing an operation.nejm.com/2005/1221/p01s01-ussc. for he knows that he is acting for the good of his patient. it has had a profound impact on human societies.org/templates/story/story.00.org/cgi/content/full/ 354/21/2277) (New England Journal of Medicine) 436 Social effect of evolutionary theory The social effects of evolutionary thought have been considerable. sometimes very widely held. In the controversial book From Darwin to Hitler by Richard Weikart [3]. which holds that "the survival of the fittest" (a phrase coined in 1851 by Herbert Spencer.[1] Stephen Jay Gould and others have argued that social Darwinism is based on misconceptions of evolutionary theory.html) (TIME) • Intelligent Design: Scientific Inquiry or Religious Indoctrination? (http://web. which claimed that human civilization was subverting natural selection by allowing the "less fit" to survive and "out-breed" the "more fit.php?storyId=5014428) (NPR) • Darwin Victorious (http://www. and many ethicists regard it as a case of the is-ought problem.org/web/ 20070928143835/http://www. Because the theory of evolution includes an explanation of humanity's origins.time.asp) (Columbia Journalism Review) • Banned in biology class: intelligent design (http://www. an idea that preceded the publication of The Origin of Species. if not all. This has led to a vigorous conflict between creation and evolution in public education.newyorker. 6 years before Darwin published his theory of evolution) explains and justifies differences in wealth and success among societies and people. After the atrocities of the Holocaust became linked with eugenics. Francis Galton. but impossible. known as eugenics. primarily in the United States.com/archive/2005/05/30/050530fa_fact) (The New Yorker) • The Evolution Debate (http://www.com/time/health/article/0.org/viewprogram. with an overwhelming present evil.csmonitor.asp?progID=506) (Justice Talking) • Intelligent Judging—Evolution in the Classroom and the Courtroom (http://content.npr.1142672.com/pages/science/sciencespecial2/) (The New York Times) • Debating Evolution in the Classroom (http://www.Intelligent design • How the media have covered ID (http://cjrarchives. such as social Darwinism. popular in the 19th century. Nor could we check our sympathy. even at the urging of hard reason.nytimes. As the scientific explanation of life's diversity has developed. Evolution and ethics The theory of evolution by natural selection has also been adopted as a foundation for various ethical and social systems. The Descent of Man.archive. The Descent of Man.org/issues/2005/5/mooney. A similar interpretation was one created by Darwin's cousin. but if we were intentionally to neglect the weak and helpless. its implied rejection of the special creation of humans described in the Bible).g. it has often displaced alternative. Darwin further argued that the most "sympathetic" societies would consequently be the most "successful. and racial extermination. Hitler often used Christian beliefs like. such men are separated from him by great differences in appearance or habits.. abortion. claiming that environment dictates our moral beliefs.[4] Some argue that six million of the people killed during the Holocaust were killed because of their religion (Judaism) not their race. which has been very active in promoting intelligent design as an alternative to evolution in public school science classrooms. 1997 at the Wayback Machine. in fact. seems to arise incidentally from our sympathies becoming more tender and more widely diffused. until they are extended to all sentient beings. and eventually becomes incorporated in public opinion. the most detailed and comprehensive of the two sections devoted to it. .. ." . and small tribes are united into larger communities. This "state of art" is not permanent: It is instead the replacement of natural selection by artificial selection through the human energy expended in maintaining it. "materialism spawned a virulent strain of utopianism. experience unfortunately shows us how long it is. may be maintained in a "state of art". Sam Harris argues that Nazism was largely a continuation of Christian anti-Semitism. it spreads through instruction and example to the young. but by constant diligence of the gardener. the moral arguments only come in later in the essay. Thinking they could engineer the perfect society through the application of scientific knowledge. the simplest reason would tell each individual that he ought to extend his social instincts and sympathies to all the members of the same nation." to justify his anti-Semitism. As soon as this virtue is honored and practiced by some few men." Archived [6] June 8. but also in euthanasia.Social effect of evolutionary theory and morality played a key role not only in the rise of eugenics. Many proponents of animal rights hold that if animals and humans are of the same nature. This virtue. This point being once reached. It should be noted that Huxley is here attempting to disprove the science behind Social Darwinism. there is only an artificial barrier to prevent his sympathies extending to the men of all nations and races. Jim Walker has compiled a list of 129 quotes from Mein Kampf in which Hitler described himself as a Christian or mentioned God or Jesus or a biblical passage. piece by piece. Charles Darwin. Many critics contend this. indeed. Materialists denied the existence of objective standards binding on all cultures." or any reason with an obvious link to the mechanism of Darwinian evolution.. "Jews killed Jesus. one of the noblest with which man is endowed. If. indeed. Consider a garden. all ultimately embraced by the Nazis. before we look at them as our fellow-creatures.[5] Of course. spent much of his book Evolution and Ethics debunking Social Darwinism. as such. though personally unknown to him. a product of natural selection and a trait beneficial to social animals (including humans). 1871 437 Thomas Huxley: Evolution and Ethics Thomas Huxley. Without constant upkeep. and whose "Wedge document" and former mission statement expand on the theme: "The cultural consequences of this triumph of materialism were devastating." He also stated that our sympathy should be extended to "all sentient beings": As man advances in civilization.Charles Darwin. even the very walls surrounding it crumbling in sufficient time. Research for Weikart's book was funded by the Discovery Institute. The Descent of Man.. The notion that humans share ancestors with other animals has also affected how some people view the relationship between humans and other species. . then rights cannot be distinct to humans. all indications are that Hilter was raised a catholic but rejected his beliefs and certainly did not live them out. "Darwin's Bulldog". infanticide. In his book The End of Faith. "strength. it would return to the "state of nature". considered "sympathy" to be one of the most important moral virtues — and that it was. materialist reformers advocated coercive government programs that falsely promised to create heaven on earth. The following is a summary of his arguments in the "Prolegomena". Nature uses unrestricted breeding to let hundreds compete for the natural resources that would only support one. they introduce English grain and fruit trees. protects from frost and drought—and.are all weeded out and destroyed. for example. the chances are that the old state of nature will have the best of it. It is thus no less natural than. they find themselves in the midst of a state of nature. However. Can we then apply this to humans? Let's see how far we can take the analogy with respect to colonization: Suppose a shipload of English colonists sent to form a settlement. thenceforward. The characteristic of the former is the elimination of that struggle. English dogs. Those to replace them are chosen with a view to his ideal of the colony. of the English animals and plants some will be extirpated by their indigenous rivals. to conquer or be vanquished. The common plants. a competitor in the struggle for existence. cattle. variation. requiring not just strength. Not only is the state of nature hostile to the state of art of the garden. in such a country as Tasmania was in the middle of the last century.Social effect of evolutionary theory This artificial selection is. They clear away the native vegetation. just as a gardener tries to create through his selection his ideal garden. On landing. finally. a gardener restricts multiplication. within the old state of nature. as yet we lack an organized gardener. and some form of selective pressure. stupid. even though the "histological process" may remove many aspects of the "struggle for existence" that is a key part of the natural laws that apply to biology. in fact. the common birds and quadrupeds. in watchful antagonism to the old regime. and take measures to defend themselves from the re-immigration of either. are as totally distinct as the men from anything to be seen on the side of the globe from which they come. widely different from that left behind them in everything but the most general physical conditions. that is natural laws. Their farms and pastures represent a garden on a great scale. gives each plant sufficient space and nourishment. to ensure that no struggle for existence between the colonists interferes with the 438 . Let us imagine an idealized one: an administrative authority of intelligence and foresight as much greater than men as men are to their livestock. sheep." However. The characteristic feature of the latter is the intense and unceasing competition of the struggle for existence.men. if they are slothful. In their place. but "flexibility and good fortune. Considered as a whole. however. they set up a new Flora and Fauna and a new variety of mankind. others will pass into the feral state and themselves become components of the state of nature. attempts to modify the conditions to benefit the forms that most nearly approach the result he desires. through selective breeding. The struggle for existence is not actually required for improvement: only heritability. The unwanted native species . and uses frost and drought to kill off the weak and unlucky. On the other hand. but the principle of the horticultural process. The presence of humans may change the dynamic. The tendency of the cosmic process is to bring about the adjustment of the forms of plant life to the current conditions. should he find them wanting. part of natural selection: It is the action upon a set of species by the human species by way of the human species expending energy through evolved intelligence on its choice of selection. but in a perfectly natural way. and English men. However. a predator expending energy through evolved instinct on preferentially hunting a certain prey species. if the work of the colonists be carried out energetically and with intelligent combination of all their forces. extirpate or drive out the animal population. from its preferred plant species by substituting human work for work done by the species itself. And. and themselves the gardeners who have to keep it up. The native savage will destroy the immigrant civilized man. or plants . Hence. so far as may be necessary. or if they waste their energies in contests with one another. though the gardener's actions may have circumvented natural selection. In a few decades. by the removal of the conditions which give rise to it. the colony is a composite unit introduced into the old state of nature. it is part of the "cosmic process". in every other way. animals. Under the conditions supposed. the tendency of the horticultural process is the adjustment of the conditions to the needs of the forms of plant life which the gardener desires to raise. and. he can still improve the species. The colonists proceed to put an end to this state of things over as large an area as they desire to occupy. is antithetic to that of the cosmic process. all other traces of the settlement will have vanished. by which the latter is created and maintained. and careless. horses. there is no doubt of the result. bonds of such a singular character. but for the means of existence. "With every step of this progress in civilization. a considerable proportion appear to me to be based upon the notion that human society is competent to furnish. housing. in consequence of the competition. if it be one. carefully matched. the weak or deformed in body or in mind. and. with the least chance of success. is to be endowed with the preternatural intelligence. we do not have an idealized administrator: Of the more thoroughgoing of the multitudinous attempts to apply the principles of cosmic evolution. many do not show themselves before the practical difficulties of life stimulate manhood to full exertion. required for the purpose of carrying out the principle of improvement by selection. or the fierce struggle for existence must recommence and destroy that peace. as equally sure to be stupid. If they were endowed with desires. However." However. nor flowers: the organization of human society is kept together by .. And by that time the mischief is done. many will consider the preternatural ruthlessness. by selection directed towards an ideal. an administrator of the kind I have imagined. with the somewhat drastic thoroughness upon which the success of the method depends. the administrator would have to face the tendency to the reintroduction of the cosmic struggle into his artificial fabric. A despotic government. The fact that "saviors of society" take to that trade is evidence enough that they have none to spare. If the administrator is guided purely by scientific considerations. not merely for the commodities. and the excess of infants born. whether individual or collective. problems arise: "as soon as the colonists began to multiply. But intelligence is another affair. though this might create a paradise where every aspect of nature works to support its colonists. it seems probable (indeed the belief is not without support in history) that a collective despotism. more and more. their lives would be conditioned by a state of art. When the colony reached the limit of possible expansion. as certain to be serviceable members of the polity. The "points" of a good or of a bad citizen are really far harder to discern than those of a puppy or a short-horn calf. and with what. on the well-known grounds of the aphorism which denies both body and soul to corporations. and selection is nullified. or vicious. the surplus population must be disposed of somehow.Social effect of evolutionary theory struggle against nature. he provides them with sufficient food. each could desire to perform none but those offices for which its organization specially fits it. with a view to the progeny best adapted to the purposes of the administrator. which have appeared of late years. would be capable of more thorough work in this direction than any single tyrant. that the attempt to perfect society after his fashion would run serious risk of loosening them. the colonists would become more and more independent of the state of nature. The pigeons. I am afraid." And so we have reached Social Darwinism. industry. the infirm aged. However. or the breeder destroys undesirable cattle. those who should be kept." just as a "gardener pulls up defective and superfluous plants. And such as they possess is generally sold to the capitalists of physical force on whose resources they depend. Experience certainly does not justify us in limiting the ruthlessness of individual "saviors of society". or what are supposed to be such. would be permitted to perpetuate their kind. in short. from its own resources. which is the fundamental condition of the maintenance of the state of art against the state of nature. if he had before him a hundred boys and girls under fourteen. and those who should be chloroformed. idle. Only the strong and the healthy. the administrator would have to avail himself of the courage. it is proper it should do. in view of the good of the whole. In order to attain his ends. I doubt whether even the keenest judge of character. In other words. has ever achieved.. However. and which. and co-operative intelligence of the settlers. and it is plain that the interest of the community would be best served by increasing the proportion of persons who possess such qualities. has had time to multiply. and so on. humans are not cattle. Among mankind. are to be their own Sir John Sebright. a mob got to believe in its own divine right by demagogic missionaries. to social and political problems. on the 439 . The evil stock. he would work to restrict the population by removing "the hopelessly diseased. "The members of the society are each organically predestined to the performance of one particular class of functions only. could pick out. They do not even correspond to social insects such as bees: With bees. puffed up with the same illusion. and diminishing that of persons devoid of them. indirectly. familial and social bonds. they risk eternal punishment.Social effect of evolutionary theory contrary. and the superfluous. when it would have been only too easy to qualify for a place among the "unfit. with one requirement: the punishment of wrongdoers being necessary for the continuation of society. without the least reference to the welfare of the society into which they are born". there is no such predestination to a sharply defined place in the social organism. and moral. must rank medicine among the black arts and count the physician a mischievous preserver of the unfit. of the bonds which hold society together. through our military and industrial wars with other nations. apart from other reasons. with all their enormous differences in natural endowment. Moreover. one must be very "fit. and the delicacy of their sensations. the creation of morality. and experience tells us how much that is worth. in one's life. I sometimes wonder whether people. the cool and daring subtlety to which the great financier owes his fortune. including: • Historical evidences against: Consider the vast changes of society between the Tudor and the Victorian eras. who justify that conduct on the ground that it has the sanction of the cosmic process. men agree in one thing. or to the hulks. whose whole lives. It strikes me that men who are accustomed to contemplate the active or passive extirpation of the weak. However much men may differ in the quality of their intellects. nor any restraint on the conduct of men. it is fairly probable that the children of a "failure" will receive from their other parent just that little modification of character which makes all the difference. and. however. under unfavorable conditions. therefore. may very easily. intellectual. are an education in the noble art of suppressing natural affection and sympathy. But. without them. the innate qualities. Surely. to do nothing but that which it pleases them to do. remains the same. if they are consistent. lead their possessors to the gallows. checked only by sympathy. the intensity of their passions. self-restraint must not be taken so far that wrongdoers may act unrestrained: Without the protection of society against them. physical. there is no conscience. Moreover. but they must reckon with the certainty that other people will be masters of the earth. in short. are not likely to have any large stock of these commodities left." • Whether some qualities are virtues or vices depends on circumstance: "The benevolence and open-handed generosity which adorn a rich man. while they hold back from those which are opposed to the sympathies of their associates. except the calculation of self-interest. may make a pauper of a poor one. nor can play. the balancing of certain present gratifications against doubtful future pains. "the followers of the "golden rule" may indulge in hopes of heaven. and fear of the judgment of ones fellow man. the unfortunate. we see firm believers in the hell of the theologians commit acts by which. and increases the importance of the pleasures and pains derived from sympathy. who. In short. Every day. it cannot be said that one is fitted by his organization to be an agricultural laborer and nothing else. human nature. after the fashion of the horticulturist and the breeder. as evidenced by their writing. on whose matrimonial undertakings the principles of the stud have the chief influence. "Every forward step of social progress brings men into closer relations with their fellows. Since morality is what keeps the desire for selfishness in check. Huxley finishes with a series of short. or perhaps two. and is the only way of ensuring the progress of the race. as they believe when cool. "In my belief. of our nation have remained substantially the same for the last four or five centuries."[7] Huxley sums up this section of his argument against Social Darwinism: I have further shown cause for the belief that direct selection. and another to be a landowner and nothing else." indeed."" 440 . the energy and courage to which the successful soldier owes his rise. If the struggle for existence has affected us to any serious extent (and I doubt it) it has been. it may be the destruction. neither has played. ever dispassionately consider their own history. who talk so freely about extirpating the unfit. it is necessary to the propagation of society. further evidences against Social Darwinism. because I do not see how such selection could be practiced without a serious weakening. any important part in the evolution of society. not to know of an occasion. and that is their innate desire to enjoy the pleasures and to escape the pains of life. evolution was accepted by some religious groups such as the Unitarian church and the liberal Anglican theologians who went on to publish Essays and Reviews. Today. in their view. Marx even sent Darwin a copy of his book Das Kapital. as did most scientists in the English scientific establishment. there has been a strong religious backlash to the teaching of evolution theory. Many important political figures on the left have never publicized their views on biology. Pope John Paul II updated the Church's position. many religions have formally or informally synthesized the scientific and religious viewpoints. This view is commonly referred to as creationism.. They have expressed concerns about the effects of the teaching of evolution on society and their faith (see Creation-evolution controversy). my predecessor Pius XII has already affirmed that there is no conflict between evolution and the doctrine of the faith regarding man and his vocation. overemphasize the role of competition and ignore elements of co-operation in nature such as symbiosis. more than a half-century after the appearance of that encyclical. Karl Marx's work was based on a material view of the world that showed natural causes and effects for all aspects of human society and economy. following a series of discoveries in different scholarly disciplines. Marxists are the exception. were also Christians who saw no incompatibility between their experimental and theoretical confirmations of evolution and their faith. where God provides a divine spark that ignited the process of evolution and (or). beginning in 1950 with Pope Pius XII's encyclical Humani Generis. However.. Dobzhansky) whose work confirmed Darwin's theory. address to the Pontifical Academy of Science. He recognized that Darwin's work provided a similar material explanation for all of nature.Social effect of evolutionary theory Evolution and Ethics [8] at Project Gutenberg 441 Evolution and religion See also: History of evolutionary thought Before Darwin's argument and presentation of the evidence for evolution. and so their opinions of evolutionary theory are unknown. though Darwin never wrote back to him. In fact it is remarkable that this theory has had progressively greater influence on the spirit of researchers. thus supporting Marx's worldview. From the 1920s to the present in the US. Western religions generally discounted or condemned any claims that diversity of life is the result of an evolutionary process."In his encyclical Humani Generis. To some extent. notably Robert Edmund Grant. "The Church does not forbid that. Some religions have adopted a theistic evolution viewpoint. in as far as it inquires into the origin of the human body as coming from pre-existent and living matter. Karl Marx..research and discussions." [9] In an October 22. which to some means species. on the part of men experienced in both fields. some new findings lead us toward the recognition of evolution as more than an hypothesis. Evolution and the Roman Catholic Church The Roman Catholic Church. Literal or authoritative interpretations of Scripture hold that a supreme being directly created humans and other animals as separate "Created kinds". where God has guided evolution in one way or another. In response to the wide scientific acceptance of the theory of evolution. 1996. Friedrich Engels and Vladimir Lenin supported Darwin's evolutionary theory. particularly by conservative evangelicals." [10] Evolutionary theory and the political left The majority of those on the left do not oppose Darwinism per se. . but are critical of interpretations of evolutionary theory that. Several important 20th century scientists (Fisher. take place with regard to the doctrine of evolution.. recognizing that Evolution is "more than a hypothesis" . took up a neutral position with regard to evolution. as well as by many scientists in France and Scotland and some in England. For example. Richard Dawkins' book. except for Christian missionaries. 442 Evolution in relation to Social Darwinism and Imperialism Evolution has often been linked to Social Darwinism. In his book.[12] During the age of New Imperialism. which is further linked with nationalism and imperialism. seldom adopted the customs and lanugages of local people under their empires. Dawkins also made a documentary of the same name. on the other hand.[11] According to the documentary. which caused agricultural problems.[12] Being critics of Darwinism. survive. these strong nations would survive in the struggle for dominance. has a chapter.believed that evolutionary theory conflicted with their economic and social ideals. Dawkins explains that when an animal sacrifices itself or uses its resources for the survival of other members of the same species. 'teleology' in natural science is not only dealt a mortal blow but its rational meaning is empirically explained. were the very first individuals to meet new peoples and develop writing systems for local inhabitants' languages that lacked one. because there is a 50% chance of a particular gene being present in its offspring. "Darwin’s work is most important and suits my purpose in that it provides a basis in natural science for the historical class struggle. This led to the practice of Lysenkoism. if a mother dies to save three of its pups.[12] Christian missionaries.the idea that an organism can pass on characteristics that it acquired during its lifetime to its offspring. one and a half copies (on average) of its genes will survive. for the first time. strong nations were composed of white people who successful at expanding their empires. ." Most later Marxists agreed with this view.Social effect of evolutionary theory In 1861 Karl Marx wrote to his friend Ferdinand Lassalle. the concepts of evolution justifed the exploitation of "lesser breeds without the law" by "superior races. some authors such as Peter Singer (in his book. On the contemporary moderate left. anarcho-communist Peter Kropotkin argued that co-operation and mutual aid are as important in the evolution of the species as competition and mutual strife. Europeans. which attempts to explain the role of altruism and cooperation in evolution and how social animals not only cannot survive without such traits. its genes. they came to support Lamarckism instead . Mutual Aid: A Factor of Evolution. and as such. it is here that. The Selfish Gene.[12] With this attitude. As a result.. "Nice guys finish first". but how evolution will create them. ardently opposed slavery and provided an education and religious instruction to the new peoples they interacted with since they felt that this was their duty as Christians."[12] To elitists. but some .particularly those in the early Soviet Union . present on the other animals. if not more so.. Dawkins added that chapter as a way of overcoming modern day misinterpretations of the concept of "survival of the fittest". A Darwinian Left) support Darwinism but reach different political and economic lessons than more conservative observers.[12] See also • • • • • • • Social evolutionism Neo-Creationism Garden of Eden Hypergamy Natural philosophy Freethought Age of the Earth . Despite all shortcomings. org/ etext/ 2940) [2] The fallacious nature of [[reductio ad Hitlerum (http:/ / www. youtube. com/ hitler. in contrast to the seventeenth and eighteenth centuries. Many westerners believed that it was their duty as Christians to set an example and to educate others." as well as Adam Smith [8] http:/ / www. htm) [5] Religion and Hitler . Even among Europeans. saying that strong nations-by definition. Social Darwinists thought their own nation the best. the concepts of evolution justifed the exploitation of "lesser breeds without the law" by superior races. org/ crsc/ aboutcrsc.) for not only having "laid the foundations but built up much of the superstructure of a true theory of the Evolution of the intellectual and moral faculties and for having realized that the creation of civilization will in itself create ethics. Missionaries were the first to meet and learn about many peoples and were the first to develop writing for those without a written language. rarely adopted the customs or learned the languages of local people. vol. archive. Social Darwinists vigourously advocated the acquistion of empires. org/ etext/ 2940 [9] http:/ / www. In the nineteenth centruy. html [7] Huxley credits Hartley (Observations on Man (1749). com/ books?id=kKGgoNo4un0C& pg=PA653& lpg=PA653& dq=social+ darwinism+ missionaries+ india& source=bl& ots=Y_LLqh3b93& sig=zQsURzU455ZsXRa-SOQJIzIzP8w& hl=en& ei=7TuhTLfJM8aNnQfF5bSiAw& sa=X& oi=book_result& ct=result& resnum=4& ved=0CCYQ6AEwAw#v=onepage& q=social darwinism missionaries india& f=false). google. discovery. if not in an evolutionary context. htm) [6] http:/ / web. va/ holy_father/ pius_xii/ encyclicals/ documents/ hf_p-xii_enc_12081950_humani-generis_en. some nations were deemed more fit than others for the competition. had wide currency in the Western states. nobeliefs. all white men were more fit than nonwhites to prevail in the struggle for dominance. net/ articles/ darwin_nazism. Usually. Houghton Mifflin Harcourt Publishing Company. gutenberg. "The most extreme ideological expression of nationalism and imperialism was Social Darwinism. They had little sense that other cultures and other people had merit or deserved respect. This language of raece and conflict.a history (http:/ / www. Politics. htm)] arguments by anti-evolutionists. org/ web/ 19970608130849/ http:/ / www. of superior and inferior people. html [10] http:/ / www. gutenberg. 281. To these elitists.Social effect of evolutionary theory 443 References [1] Eugenics. and Society" (http:/ / books." . nobeliefs. com/ library/ PAPALDOC/ JP961022. ewtn. darwintohitler. HTM [11] Nice guys finish first (http:/ / www.] [3] http:/ / www. com/ [4] Webpage containing 129 Mein Kampf quotes relating to his Christianity (http:/ / www. com/ watch?v=FzeCn02l_Rw) [12] "Western Civilization: Ideas. In the popular mind. except for missionaries. those that were successful at expanding industry and empire-would survive and that others would not. Retrieved 2007–03–25. vatican. Christian missionaries were ardently opposed to slavery. Europeans. rationalrevolution. ii p. an attitude that sparked their competitive enthusiasm. . com/ hitlerchristian. etc (http:/ / www. Ludachris78641. Jacek Kendysz. Adi. Baristarim. Ashenai. Encephalon. Merzul. B. Ems2.Spudeman. Gdo01. Andrew c. Agathman. Sashag. Abuzin.. Fieldday-sunday. Lcarscad. Hari Seldon. Hubcapwiki. Mann jess. Dpr. DGG. Tomandlu. FlyingToaster. Bdaay. InShaneee. KingTT. Bendzh. Boomcoach.. Vsmith. Raeky. Bull Market. Can't sleep. Badreligion. Enviroboy. BigGoose2006. Inthebeginning. Howaboutthat. DocJohnny. Fvw. Hrafn. Headbomb. Delta Tango. CUSENZA Mario. John Stumbles. Iamnotanorange. Kaal. Danerz34. BlueFireIce. Gniniv. Hippietrail. Lobe lob. Driski555. 4 Gnon. JimWae. Kelly Martin. Evercat. SandyGeorgia. Badanedwa. UncleBubba. Jitterro. KillerChihuahua. Lightdarkness. Emw. Alexf. Ec5618. Dr.. Hob Gadling. Axel147. Jos Jio. Juamh. Bender235. Bollyx. Dial lean. Leptictidium. DCDuring. Depolarizer. Landroo. Emmett. Arkatox. Amcaja. John. Ekologkonsult. Dafergu3. Dante Alighieri. Ag545. NeilN. Biglonstud. Algebraist. Bodnotbod. GraemeL. Darwindis. Katalaveno. DopefishJustin. Jim62sch. Fxmastermind. Cosmic Latte. Lars Washington. Ear Sid. Boom. Iridescent. Foggg. Gilgameshfuel. LA2. Agrabiec4. Demi. 489thCorsica. Jokerst44. Aecis.xxx. Lukasz Lukomski. Joel. Giuegg. Grooy. Hbeop. Galwhaa. NuttyProSci-Fi3000. Afasmit. Binabik80. Jordan117. Cpl Syx. Apostlealex. Eenu. Dash92. Asemoasyourmom. JeremyA. Android79. Alpha166. KayEss. Drooling Sheep. 123heyho. Edivorce. Dr. FisherQueen. GoonerDP. Lazza 99.delanoy. Bkell. Burtonsarpa. JeffStickney. Evil saltine. Ediblehuman. CommonsDelinker. Dfarrar. Flimg. Z10x. Macdonald-ross. Duja. Harald Khan. Dangee63. Laurence Boyce. Pcu123456789. Kteey. Hanksname. Ilya. Kinst. J. Hriow. Evolutioniswrong. Gscshoyru. Bart133. Bkl3x. Crazytail2.129. 9 noon. Elmerglue. Sjö. Daelin. Dustimagic. B89smith. The Nut. Manuel Trujillo Berges. 162. Bornhj. K. JoanneB. Bio-queen. Dontlookatme. Fuzheado. Woudloper. PhDP. LonelyMarble. Intranetusa. Eve oft. Crohnie. Dinaey. Jak Tac. Laik G. Bogdangiusca. John Wilkins. Base tonne. Liz de Quebec.13. Tstrobaugh. 6 Four.illuminatus. Dressagebea. Da Gingerbread Man. Hmpxrii. Aaarrrggh. BlackMaria. Edwardmayhem. Dsevans93. Gabbe.O. Borisblue. David. AzureCitizen. Avicennasis. Feeeshboy. ChadThomson. Iankeir. Jamesofur. Aarnu.org/w/index. Cwilldagangsta. Titanium Dragon. Franz lino. Fubar Obfusco. Laegg. AH9.aston. Khalid Mahmood. Will Beback. Glen. Leeg 7. Fang 23. J. Gmlk. Dekisugi. Aaron Schulz. Bobby Mason. Lakle. Magister Mathematicae. Aartp. 1nic. Doalla 8. Lee Daniel Crocker. Logicus. Dachshundboy25. MAXimum Xtreme. Lscminn. MisterScott99. Fic-in. ManaUser.xxx. Hollywoodgenes. Ascánder. Cougarkid. AdamRetchless. 3 Lane. Matthead. Kandar. Kungfuadam. Leibniz. Craigmac41. KMFDM Fan. Bevo. Kdbuffalo. JoergenB. Joka1991. Gits (Neo). IanCheesman. F-402. Notmymall. Felizdenovo.delanoy. Delphinian. Bassbreaker. Draco 2k. DanielCD.26. Dar book. Doc Tropics. Kuru. Demong. Barny Fife.108. Jimfbleak. A Softer Answer.126. IzzyReal. Dg10050. Lord Voldemort. Raphite. Jpowell. DSG2. Eel eat. Geneb1955. Dabs. SU Linguist. Ggbroad. LightningPower. AlanHarmony. Sarefo. Lancer873. Fox1. B. Special-T. Dveim.. Ashmoo. Liquid2ice. Antandrus.lot. Cybercobra. Teapotgeorge. Lif 91. Deor. Harryboyles. Bauxzaux. Gorillasapiens. BillySharps. Crazytail3. David Merrill. RucasHost. FCYTravis. Haikon. Harehawk. Aainsqatsi. Conspiracyfactory. Ali. Kondspi. Hdt83. Baegis. Ginsengbomb. Arm. Johnuniq. Gracenotes. Betbs. C. MER-C. Flowerpotman.xxx. Ale jrb. Alexwebb2. David Z. Miranda. Daycd.. Jurto. CuteWombat. Dmerrill. Michael Devore. Richard001. J4V4. Lremo. Hipporoo. ClarenceCM3. Malvaro. Calamine Lotion. Knight of BAAWA. Koavf. DLH. Animum. Joyous!. H2g2bob. Nishkid64. Aunt Entropy. DreamGuy. Jacob1207. Benandorsqueaks. GOC-in-C. Pslide. Jimbobsween. Grutter. Jrtayloriv. Bluegrass32333233. why are there still apes?. Benhocking. Joannamasel. JustSomeKid. Karatenerd. Davud363000. Az1568. GSlicer. Wegotthe. Jrmccall. Barnaby dawson. Gog. Chairboy. Furrypig. Wknight94. David D. Clayoquot. The Thin Man Who Never Leaves. EncycloPetey. Spencer. Knuv moor. Briguy. Alan Peakall. Clarityfiend. clown will eat me. Abi Don. EncycloPetey. Levi P. Peter coxhead. Amaltheus. Hrimfaxi. Andrewa. Brainstormer1980.wikipedia. MONGO. MPF. Boffey. Fastfission. Cen Upp. Sander Säde. Eaglizard. Maxis ftw. Manph. Juffe. Igiffin. John Fader. JohnnyCalifornia.wikipedia. BlytheG. Axeman89. Korinkami. Jefffire. Drpickem. Donhoraldo. David Shankbone. Rror. Alex. AVengel. Cold Sandwhich. Basil45. Homestarmy. Banes. Ande B. GVP Webmaster. Fingers80. Dcooh. Doulos Christos. Chun-hian. Fagstein. Coppit. Joshurtree. CapitalR. Andrewlp1991. Alienus. Creationlaw. Aldaron. 65. CalebNoble. Frazzydee. GK. 40 tune. CommonJoe. ArielGold. Dwmyers. Eriathwen. Cards44izzy. Daa89563.xxx. Lime Ore. Krukouski. VVtam. Natalie Erin. Dave souza. Graham87. Fm.175. Filll. Charlesdrakew. Skittle. Hubrid Noxx. Ancheta Wis. MasterVageSha. CMacMillan. Fubar Obfusco. Deltabeignet. ESkog. Hu. Besidesamiracle. Apokryltaros. Aks818guy. Luna Santin. Falcon8765. Flaiw. Andriesb. Coopercmu. Black Kite. Fen Fin. The undertow. NawlinWiki. A8UDI. Lfh. Icairns. Achoo5000. Kljenni. Aldenrw. Lyotchyk. Brazucs. Enochlau. Igor233. Eu. CarpeScientia. BlueNight. Boothy443. Gurch. ConfuciusOrnis. Paul Carpenter. AdultSwim. Malcolm rowe. Khukri. Danielkueh. Avb. Fbartolom. Equendil. Lickarice. Caltechdoc. 4 trin. Armchair info guy. Frenchgeek. Alfio. Causa sui. Dancingspring. Five part. Borgx. AxelBoldt. Chaojoker. MZMcBride. Doggie Bark. IP69. Another sockpuppet of Outoftuneviolin. Benzocane. Herakles01. Dubc0724. Zapvet. Abce2.Monniaux. FT2. PDH. No Guru.:Ajvol:. Fvasconcellos. JoshuaZ. BradBeattie. Davidhorman. Joelr31. Gabbe. Leandrod. Lhasanimir. Bucketsofg. Konjikiashisogijizo.a.scheme.xxx. K. Sting au. ASDFGHJKL. Furry great. Donner meat123. Bharatveer. Aurush kazemini. Danny. Garik. Ixfd64. Mal hen. Professor marginalia. Sintaku. Chrislk02. Cinik. Fang 23. Finn up. Laughing tyre. Jnothman. Breathe ing. Hannes Hirzel. Andrevan. Seb az86556. 24. JWSchmidt. Erik9. Hadal. Dave souza. Maartenvdbent. Lst27. Guoed. Angelo. MSGJ. MECU. Darthgriz98. Apokryltaros. 2357. Bbatsell. Duffman. FeloniousMonk. CommonsDelinker. G4rfunkel. Bryan Derksen. Joe1978. Athenean. Caasiopia68. Joshjoshjosh1. HisSpaceResearch. Bcasterline. John. Googleaseerch. J. Jania902. Kanfood. MSTCrow. Kilda. CambridgeBayWeather. AzaToth. Ardric47. Artichoker. Jaberwocky6669. Lexor. Surendre91. ArcticFrog. Foofighter20x. Betacommand. Random Replicator. Chameleon. Tktktk. Gadfium. HaeB. Jedwina. JimmyButler. Arjun01. FeemisDragon. EP8841. Headbomb. Binarypascal. Download. Bookandcoffee. 8r13n. Madeleine Price Ball. Bwatb. ConservativeChristian. Kay of Kintyre. Wassupwestcoast. Elkman. Awadewit. Awon. G-123. Gzkn. Hadrian89. Golden two. JHunterJ. Copper canned. Methcub. Jan van Male. MKoltnow. Agg56tt. Adambiswanger1. Elembis.. 119. Busy ironing. Jclerman. Geni. Arch dude. Dinaayoub. Unhinged1122. Justbob. Rambam rashi. Ajgisme. IanCheesman. Jdhunt. Icthus123. Karmosin. Craigy144. HenryLi. KeloheDeb. Jjb123. Gfoley4. Thingg. Adamsiepel. Gooago. Ghaor. Caulde. Jagged 85. Breed3011. FlavrSavr. AlphaEta. NeoNerd. Burmiester. Conversion script. Macaddct1984. AlanD. Alphachimp. Fred Hsu.94. Jmeppley. IronChris. Linecircle. 168. Kukini. Loom yellow. H2O.68. Fredbauder. Firsfron. Gnixon. GeoMor. ForbiddenWord. Danielparker. Livingboner. Gregbard. Ed g2s. Adenosine. El C. AnthonyQBachler. Jesus Loves You. Masterpiece2000. Joelsmades6. Mexeno1. ClockworkSoul. Amaltheus. Andrewrost3241981. Jketola. Irishguy.php?oldid=392593427 Contributors: 100110100. Durin.89. DLR. Isac. Cretog8. Andrewpmk. Haukurth. Chaos. Here4thefood. Fang Aili. Bevo. ScientistDragonmouse. CanisRufus. Kajisol. Kazuba. JESUS LOVS YOU. DJ Clayworth. GD. JohnOwens. Kirbytime. MarcoTolo. Magician that makes things white. John Callender. Chicken Soup. Carcharoth. Leithp. FieryPhoenix. Lakinekaki.226. Heimstern. Back23. Rosesflowers8. Inn par. TimVickers. LonelyMarble. Jamyskis. Jdfoote. DragonFlySpirit. Grandmastergrawp. Chanting Fox. Koavf. Acalamari. Dashmast3r. Narayanese. Kniknight. GetAgrippa. Cyp. Jeremypyle. KnightLago. Dw4372. R. Koeplinger. HansHermans. Gleng. Jason Potter. Crab or. Snalwibma. Lights. Electric free. Favonian. FlyingOrca. Thethornofarose. Arker. Aitias. David Fuchs. Gzornenplatz. Cyr egi. Chrislk02. ComputerPlace. Jlrobertson. Cynicism addict. Heyits5. Cosmic Latte. Justforasecond. Hairchrm. Fr8we. Daniel5127. Edwy. A bit iffy. Hard Sin. Fott leigh. Clicketyclack. AlienHook. Ljbhebjbb. JForget. Brighterorange. Artichoker. Butros. Gwernol. JNeal. Jenequa. GSlicer. Billare. Adrian. GeometryGirl. Johnuniq. 210. 0nlyth3truth. Azcolvin429. Bobo192. ABoerma. Paste. Hillock65. Abtract. Dispenser. Icseaturtles. BlueGoose. PervyPirate. John254. Bubba73.. Apostrophe. Kate. Kaisershatner. Celador. Herk1955. KnowledgeOfSelf. Jlefler. Christian List. Alan Liefting. Hanoachide. Fred Bauder. Psychodolly. Joe11. LifeScience. Mimihitam. Fraslet. Myrridias. David Schaich. JinJian. MHall. BatteryIncluded. Johnstone. Aunt Entropy. Jeffrey O. Dark Shikari. Chas zzz brown. Rich Farmbrough. Danimoth. Wat-up-guys. Hawknel. Dawn Bard. FrummerThanThou. Kotjze. Adz71. Danielrcote. Frymaster. Josh Grosse. TableManners. Esseh. Kugamazog. Vanished user. PlattF. Aznmastermind5. Jsjacobs. AuburnPilot. Algorithms. GeorgeLouis. Kablammo. Kaboomywhackwhack. Betterusername. Eath Br. Graft. Hmains. Lawn 42. Flubbit. DKqwerty. 16@r. Galoubet. Instinct. 99DBSIMLR. Candorwien. FreplySpang. Diza. Jesus Is Love. Athf1234. Everyking. Thefellswooper. GeneralPatton. James. Jimaltieri. Ly4321. ConfuciusOrnis. Hut 8. Lethe. 100110100. Harvestdancer. Illuminatiscott. Gurvindert. J. Christopher Parham. Grandpa2390. Smith. Butwhatdoiknow. Woohookitty. Jpk. GN. Cheshire Boy. Adashiel. Missbmd. ArthurWeasley. The Last Melon. Octavian history. Camembert.Mestel. Kdeo. Lostcaesar. Acroterion. Philanthropic1989. JF Mephisto. JoshuaZ. Adriansrfr. KimvdLinde. Javascap. IVAN3MAN. Infophile. Azra99. Ebush. EdJohnston.Gilmore. God Emperor. Chris55. JeremyA. Oreo Priest. Dyac L. Fixtgear. Eubulides. Hensa. Chris the speller. Fredrik. Croft. Avenue. Ian Dalziel. Geddone. Mahmudmasri. JedRothwell. Giftlite. Magicman710. Connelly. Shinglor. Dcoetzee. Lisasilzaz. Cynical. Joymmart. A455bcd9. Bth. DanTheMan2.. Jedi of redwall. Autonova. Macdonwald. Leranedo. Keno. BrokenSegue. Ballista. Glomove. Coffee2theorems. Farsight001. CJLL Wright. GoEThe. Jrockley. Canterbury Tail. Boo wall. Eloquence. Elijya. Blainster. Kinhull. Gogo Dodo. Amarilloarmadillo. Jackrm. FeloniousMonk. Astroview120mm. Hadrian89.stefan. KC Panchal. Mchavez. Jgardner7777. Ali. RatKingSmurf. Aggn. Dipset1991. Davril2020. Laorv. I. Shadowsoftheriots. FF2010. Geothermal. Bowslayer. LaMenta3. IVAN3MAN. BlankVerse. Mike Searson. Centrx. Jok2000. KsprayDad. Jhouser24. Debresser. Aude. Doll ten.Fred. Byrll. Boe syl. Iota. Gilawson. ElkeK. Loom91. Dina. Ju66l3r. Daeoque. Frank A. Twooars. Discospinster. Joke137. Ethancole117. M. Knowledge Seeker. BG.54. Jossi. Canadian-Bacon. Five edges. Fisher. Flockmeal. HJ Mitchell. Harej. Cyde. Jvbishop. LexCorp. Andrew Lancaster. Thompsma. BirgitteSB. Illspirit. Billystut. Faradayplank. Igilli. Ggbroad. Blanchardb. Duncharris. Gcior. Goatasaur. Debresser. DerHexer. Itsmine. Ligulem.org/w/index. Lang. Delta Tango. Johnleemk. Broalka. Kim Bruning. Cometstyles.14. Kwion. Gmaxwell. 12 Cove. Clayoquot. Art LaPella. Killdevil. AndrewWTaylor. Silly rabbit.250. Gnfnrf. Epbr123. Andromeda321. Mgiganteus1. Genius523. Axxel Sel. Chardon. Seegoon. Barbary lion. John D. Leon. GromXXVII. Joe. . Davodd. Filll. CharlesGillingham. Kylu. Christopherlin. Gringer. Kmweber. Hackwrench. Delirium.. Beezhive. BlytheG. Ratphobia. Enormousdude. Fen zero. CryptoDerk. Aircorn. Orangemarlin. Flo98. Arnoutf. GodsWarrior. Ciar. Silence. Ec5618. Dougp59. AshLin. Mindmatrix. Calaschysm. Astrobayes. Ajf99. Brent0270. Fgleb. Bogey97. Amused Usher. Frosty0814snowman. Deviator13. Delarge3. Sfvace. Ettrig. Keet Onn. . 7 Fej. Anilocra. Boring bus. Altenmann. Writtenright. Kev 63. Hdhcst95. Hrafn. DavidL (usurped). Alias Flood. Drmies. Gaius Cornelius. GrahamColm. Emw... Bill37212. BrownHairedGirl. Gdavidp. Agentmoose. Ettrig. Halaqah. Freestyle10evan. Bakasuprman. AaronFX. It da. Elamere. Christpower79. 216.Article Sources and Contributors 444 Article Sources and Contributors Introduction to evolution Source: http://en.50. MBCF.87. Jiejunkong. LFaraone. AC+79 3888. Hillock65. 6 y go. Donnellyj. Ezra Wax. Fred Hsu. David D. Axa4975. Dyoes. Iamme06. AlexiusHoratius. Emc2. Ginkgo100. Africangenesis. Izztek. DDek. GalaazV. Dreary101. Dlohcierekim. JayHenry. GloriousLeader. Leave Power Behind. Guettarda. Dan ald. Jack Walsh. NuclearWarfare. Wizardboy777. GHe. Lycanthrope. 10. Caesura. Lenin13. Fourier. Deeptrivia. Vanished User 0001. Tikiwont. Lowellian. JimmyButler. JosephPayseur.103. CardinalDan. Lailablossom. JiFish. Dullhunk. Farsight001. Chunky Rice. Friedgreenkillertomatoes. WAS 4. Freakofnurture. JackH. Coffee2theorems. Butwhatdoiknow. Country050. Randomblue. Hojimachong. Cmayy. Ballin789. Dancingspring. Bwhack. Gortu. Asbestos. Eisnel. Bam2014. Hordaland. Edgar181. Broabey.5. DEFRT. ForestDim. Goethean. Academic Challenger. Heron. Goxxen. Lapabc. Ahoerstemeier. Kyorosuke. CheeseDreams. Isaac. Grunt. Andymarczak. Insearchoftruth. Curps. Kungming2. Armchair info guy. Iwanttofitin. Jecar. Acsparkman. Edward. . Drilnoth. Backslash Forwardslash. Colin Keigher. Jon234567890. Haham hanuka. Greeneto. Jumacdon. Anthony. BenB4. Etxrge. AmiDaniel. Dracontes. Blue401. Brian0918. Audacity. Edgarde. Abrech. ScienceApologist. Die4christ. Alphazeta33. Amitch.یناریا 248 anonymous edits Evolution Source: http://en. Digitalme. Cutsman. Gustafson. Deor. Eviom. Malo. Shyam. Drpsarge. Greenrd. Foxxmuuldr. Jumbo Whales. La goutte de pluie. CerealKiller. Haemo. Crazytales. Faradn. Cybercobra. AndrewTJ31.php?oldid=393126466 Contributors: !!!niloivenutfotuO. Jedi Davideus. Dongless. LeperColony. Adambro. Chris Melton. Gregbard. Dhomm. Digresser. Christian41691. Fastman99. Nneonneo. Ed Poor. CanDo. Clean Sweep. Ryantremblay. Compute14. Ian Pitchford. Imag Qu. J Di. El-Spectre. Dysmorodrepanis. Crust. Joyous!. Benhocking. Comder. Skier Dude. Cinnamon. Bow bowl. Lithfo. ElTyrant. Ignignot. Adraeus. Kurykh. J0m1eisler. Gregcaletta. Abdullais4u. JYolkowski. AvicAWB. Levineps. FisherQueen. Cassan. Tuxedo junction. Ck lostsword. Jaknouse. Gimmetrow. ChicXulub. Kosebamse. Aron-Ra. GregorB. Garylhewitt. DVD R W. Ikanreed. Beltho. AlphaEta. Dionyseus. Grsz11. If humans came from apes. Lou Sander. Dweller. Ikh. Arof. Gary Cziko. Barras. David Kernow. Circeus. Cloud silver. Irpen. Kelsey Francis. BrianG5. Jiy. Neptunerover. Loom peat.tan. Guettarda. Rjwilmsi. Malik Shabazz. David. HXJ. Cherhillsnow. DeadEyeArrow. Iamdadawg1. Dbtfz. Kangaru99. Bikeable. DrKiernan. Paulr. Rls. Striver. Sjö. Montgomery '39. Noivb. Snalwibma. Sickopath. RexNL. Sophie. PatrickA. Tannin. Papapryor. Eheneden. Tgees. Sam Spade. Slrubenstein. Rickert. Philcha. Unflappable. The Anome. SuperLexicon. Possum. Wavesmikey. Oking83. Nbound. Octopus-Hands. IslaySolomon. Hmains. Elfred. Soap. Nnp. Dave souza. Skywriteing. JSF16. Outoftunebassguitar. Raphus Cucculatus. Melono V. Ed Poor. Ra2007. Scott Noyes. Worship Him!. Q. Numerousfalx. Rjoebrandon. Pablo-flores. Oboeboy. Wdraa. NuttyProSci-Fi3000. Sobored1. Wiki alf. Toothturtle. Cst17. Master Jay. Nondescript. Teapotgeorge. Pschemp. Raymond. Robert Stevens. Pizzaghost. Ranveig. Varith. Samsara. Myoe83. D. WAvegetarian. Tom Schmal. JohnCD. Aprock. Slrubenstein. Workaphobia. Owendude1210. GeneCallahan. Robertbowerman. Zappernapper. Blue Dinosaur Jr.php?oldid=392830825 Contributors: A little insignificant. TheSlowLife. Persian Poet Gal. Trabucogold. Yosef 52. SWAdair. Thomas Veil. Picus viridis. Armchair info guy. MarSch. Jaketola. Mattpickman. Stirling Newberry. SpookyMulder.org/w/index. Dbachmann. Zashaw. Crystallina. Hut 8. Mastahcheeph. Rjwilmsi. WLU. TxMCJ. Qcoo. Excirial. Eaglegordon. Raz1el. Sommers. Marc210. Edison. Sushilover2000. Raymondofrish. Silly rabbit. WillowW. Salvio giuliano. RJHall. Kingdon. Boeingbenn. Sophia. SixPurpleFish. Outoftuneviolin 4. NuclearWarfare. Sundar. Questioning. Moondyne. Aunt Entropy. Mickchaaya. Spotty11222. Peruvianllama. Sikatriz. Prezbo. OutoftunevioIa. Nefariousski. Phlegm Rooster. Nimur. Shadowin. Viskonsas. Wing Nut. Spark. Timir2. Martin Hogbin. WebEdHC. Ork rule1.. Zx-man. Strreeh. Guettarda. Mor. Xiner. FlagSteward. Pgan002. Gail. Yorek.tk. GregorB. Xiahou. Mxn. LilHelpa. TheNightRyder. Pbuee. Nick. WAS 4. NoSeptember. Phaup.hoyland. PWhittle. Glloq. Aitias. Skittle. Sfvace. Naddy. StaticGull. Wacxi. SkyWalker. Anarchangel. Uriah923. Titoxd. Schaefer. MichaelBillington. Vind r. Vox Rationis.hoyland. Randwicked. Ultimus. Keilana. Nukeh. Thomas Arelatensis. Alan Liefting. Tstrobaugh. Shglien. No1lakersfan. Smalljim. Pbarnes. Ac02111993. Mdotley. Rappadizydizzy. 5 albert square. Rd232. Xaosflux. Kozuch. Unlocked. Raeky. Rokfaith. Ricardo sandoval. Spotfixer. Robert1947. O. Savant13. Mkrose. Tznkai. Tide rolls. Leivick. Sally1309. Nihiltres. Billion. SamuelGrauer.org/w/index. Vanderdecken.. Satyrium. Scfencer. Snigbrook. Mildly Mad. Hans Adler. Shicoco. Philanthropic1989. Titanium Dragon. Squiddy. Cremepuff222. Calvin 1998. Wilson44691. Yuodd. Warpsmith. Prom ten. No Guru. Semantics. TheAlphaWolf. Monterey Bay. Svartkell. ScienceApologist. The Nut. SupaStarGirl. VictorAu. Michael2. Gjmulder. RoyBoy. Tobyk777. Philipholden. MattDal. Skaterbøy. IRP. Randywombat. Nowa. Scipain. Jeff G. Zbvhs. StradivariusTV. Paleoderek. Methcub. This user has left wikipedia. CooPs89. Thiessen. Sharifneuro.wikipedia. Onecatowner. Mikemoral. O keyes. Iridescent. Woudloper. Zarru. NewEnglandYankee. Widictionaryia. Max David. Jomasecu. EncycloPetey. WBardwin. Outoftuneviolin's son. Mattunseal. Jarry1250. RJFJR. Matt poo watt. Muijz. Wildfirejmj. Nivix. Tony Sidaway. Waldow. Stephenb. Phantomsteve. RandomXYZb. Nathanielsmithisaloser. Thadius856. Tiberius47. Pbarnes. West store. Causa sui. Utcursch. Quux. Rudjek. Darth Panda. Waterspyder. RS1900. Omicronpersei8. Orangemarlin. Niaga Niloivenutfotuo. Teapotgeorge. Michael Devore. Tuxedo junction. RetiredUser2. Janus01. Gareth E Kegg. VolatileChemical. SallyForth123. Tntnnbltn. Mirv. Yar owe. Marcperkel. Editor2020. AlphaEta. Mourn. Truth0r. Truth777. Pilotguy. Another Stickler. Sting au. Montgomery '39. Those Debate Studs. Andrew c. Sikkema. Oblivious. Rammadd. Elmerfadd. Rossnixon. Scen heal. Phaldo. SparrowsWing. BenFrantzDale. Zappernapper. Shark96z. W00tboy. SodiumHydroxide. Pce3@ij. Nakon. Linkthewindow. Bkell. Thompsma. Tail. Weyes. Mike Rosoft. NawlinWiki. Quizkajer. RexNL. Ripepette. Paxsimius. Sander Säde. Thinradred9. Johnuniq. Hrafn. Maz haz. HiDrNick. TimVickers. Tailpig. Tbarron. Johnuniq. Minna Sora no Shita. Postlewaight. Random Replicator. Parrikk. Visionholder. Portillo. Nvav. VVtam. Raeky. ThinkBlue. Rich Farmbrough. Wassupwestcoast. Deconstructhis. Risker (Anne Criske) is old & has gray hair & wears glasses. Mike Young. Sifaka. BellaKazza. NuclearWarfare. Veratien. Kittiekatsu. Just plain Bill. Raybechard. Wolfkeeper. Qomee. Ptaat. Spangineer. Tschwenn. Samuraipizzaguy. Wimt. Atl braves. Matěj Grabovský. Techyactor15. The Duke of Waltham. Mindmatrix. Suzanne Elsasser. Ajstov. Mary Calm. Tsusurfanami. SFC9394. Uesonne. Blind designer. Walkiped.. WeddingCrashError. Woohookitty. Yasmine shenn. R0M4NC1NG H3LL. Vituperex. ArielGold. Z. Moulder. Micahbode. Andrewlp1991. OwenX. Redrocket. Mike6271. Marshman. Jchthys. Ritchy. Northfox.Fred. Scorpionman. Willtron. Unclebulgaria. Red Act. Swaggart. Gniniv. Plommespiser. Thevenerablez. Ncaroe. NawlinWiki. Ryulong. Ulric1313. Realitygenerationx. Meson man. RyanParis. O. Julesd. Yamakiri. Nickptar. Ezra Wax. Sluzzelin. Urco. Wool sixty. St3p5. R Lowry. Mr Stephen. Pigsonthewing. Wikipeterproject. Tile join. Outoftuneviolin. SchfiftyThree. Taco325i. Vsmith. Declan Clam. Wikidea. Auraclan. Teenwriter. Reywas92. Pjotr Morgen. TedE. Texture. Wennj. LeCour. Qtoktok. Mausy5043. Yandman. Nwbeeson. Slicky. Thue. VI. Vssun. ProfessorRight. Sbarry12. Paw yanne. Wafulz. Nutella002. Bdoom. Operation V. Kal Sir. WadeSimMiser. Zazaban. DavidLaurenson. Upsidown. Solaricon. Verwoerd. TBadger. Plaster of.. Newtman.php?oldid=391449486 Contributors: 10outof10die. Tmol42. Wapondaponda. Monkey4160. Arbeo. Outoftuneviolin 3. Gregrutz. SDas. Smith609. Gabbe. AkselGerner. Salva31. Postdlf. Ollyoxenfree. TeaDrinker. Red went. TimVickers. Diucón. Pluthra. Shot info. WatchingYouLikeAHawk. Ystram. ScienceApologist. Pine fame. Butwhatdoiknow.-). Seven rawt. Staxringold. Sushant gupta. MuZemike. Skid slid. Robert Merkel. Nwbeeson. Maver1ck. Mcy jerry. Sagi Nahor. Gnangarra. Skamel85. Vquirk. Endgame69. Aksi great. ElijahOmega. Pippo2001. SmilesALot. Kku. Pollinator. StoatBringer. Matekm. May Ced. Recentchanges. Smooth O. Margareta. Shadowjams. Graham87. Tim@. Stuthomas4. Timwi. J04n. Nightscream. Alan Liefting. Tuatara. Syvanen. Nurg. Ms2ger. Reuqr. Peter coxhead. Opabinia regalis. Unhinged1122. Yarovit. Artoannila. Ocatecir. Spencer. Vduer. Venn 87. Traumerei. Th1rt3en. Uogl. Rmky87. Ensign beedrill. Zoe. Dougweller. Aunt Entropy. Yowuza. Staticmemory. UTvolfan711. Moreschi. Ate98. Vipinhari.C. Twilh.. Wild Deuce.. Someone else. Rusty Cashman. SpK. Dawn Bard. Sheep81. Sleepnomore. Toyokuni3. Mwanner. Number 04. Webbrg. The pupils president. WLU. NativeForeigner. Wikididact. Rnt20. Steht. Wisdom89. Matthewcieplak. Swatjester. Nol888. UtherSRG. Robin Johnson. Fama Clamosa. Sean. Teflon Don. Tune. Sparrows point. Stephenchou0722. Nrcprm2026. Mkmori. Anas Emad. Rursus. Orangemarlin. Evrimfeyyaz. Rcuub.net. Swpb. NewEnglandYankee. Headbomb. Neurolysis. Paul August. Larsen. OmerSelam. MarcoTolo. Sonikkua. Mgs 90. MisfitToys. U mene. Sancassania. You wouldnt dare block me. Thesquire. Steinsky. Variable. Nicolae Coman. Ymous. MarcIam. PDH. Nath Hebb. JSpung. Navme. Otterathome. McSly. Alansohn. Rreagan007. Leptictidium. Tatarize. Samboy. Colinclarksmith.Article Sources and Contributors Maork. StaticGull. Payp two. Jag149. Joshuajohnlee. Tfietkau. Skizzik. Sfvace. Thorht. N6. Nicholasink. Mikael Häggström. Michael Johnson. Teh one who blocks me is an evolutionist. Oasrocks. 16. Tezero. Sengkang. Soom zo. Poopmaster1234567. Theoldhenk. Wmahan. Noisy. Roland Deschain. Sampi. GVnayR. Politoed666. Oliver Lineham. MarsRover. PieCam. ElationAviation. Tawker. Scientizzle. Warrush. Rgamble. Narayanese. Beardnomore. TonyClarke. Yqbd. Arthur Rubin. Escape Orbit. MartinHarper. Mimihitam. Tothebarricades. Weightofair.. SuperTycoon. Two teem. RDBrown. Maximus Rex. Smith609. RDF. Ppe42. Urod. Nate slayer0. Slashme. Zundark. Wyher. Dawn Bard. SMesser. Rhynchosaur. Vary. THobern. Sionus. Pixel . Naoh. Noodleman. Su huynh. Onorem. Mark Renier. Zeromegamanx. Sweetpoet.E. Wikiskimmer. Michael Devore. Siroxo. Scientizzle. Ugur Basak. Superdudefreak. Craigmac41. Docu. Mike Jones. Proopnarine. Tlame. Saucepan. MarineHebert. Hryhorash. Wikid77. Siddiq ahmed. Shauri. Rosencrantz1. Verybigfish86. Ndteegarden. Wetman. Mjharrison. RobertG. Shambalala. O18.N. Srivange. Outoftuneviola. 362 anonymous edits Evolutionary history of life Source: http://en. Table pot. Spotty11222. Petri Krohn. Sushant gupta. Timothy Usher. Ucanlookitup. Wegge 9. DarkLink. Paul Drye. Rockin dead. Fatapatate. Shlomi Hillel. Rossman123. Philopp. Daniel J. Snalwibma. NickBush24. Screen five. Shyamal. Mygerardromance. Hqb. SJP. Icairns. Trishm. Methcub. Ooogage. Snakey Jake. Schrandit. Salanus. Tygrrr. Plantsurfer. Matusz. Refusedalways. Raymond arritt. SandyGeorgia. VirtualEye. Aks818guy. Zebspace. Pmj. When nine. MrFish. Thuom. 2608 anonymous edits Evolution as theory and fact Source: http://en. Discospinster. Simpsons contributor. Wknight94. Yiwb.250. Wildie. Petri Krohn. Whoistheroach. Tweetbird. Phaedriel. Rror. Vanished User 0001. Parallel or Together?. NorCal764. Saros136. Olav L. Skiddum. Bluerasberry. Sopher99. Tycho. The Rogue Penguin. NorCal764. Pharos. Memestream. Rebroad. RJN. SpeakEasier. Glass Sword. Uberjivy. Treisijs. Mcapplbee. Tpduden. Mister Five. Sting au. PseudoSudo. Glane23. Kerotan. Orangemarlin. Ragesoss. The Rambling Man. Redioz. MirDoc. Wwoods. TomShmells247. Munita Prasad. Rusty Cashman. Poiuyt Man. 81 anonymous edits 445 . NBeale. Smelialichu. Vain vase. Jesusforever. Rolf Schmidt. Twas Now. Outoftunebassoon. Wlodzimierz. Ptk. Shanel. Ravenswood Media. Tim1988. Tynews2001. Nexon123. TheSun. River flowing. Jeff G. Valluvan. Tariqabjotu. Pierpontpaul2351. Woohookitty. Lars Washington. Raz grime. NAZISMISNTCOOL. Roadsoap. Tintero. REF4. Tteil. Sting-fr. Radred. Type how. Wap. Spliffy. Sleeppointer. Montana's Defender. Pathoschild. Silly rabbit. GodMadeTheUniverse. Misteror. JoshuaZ. Rugby471. PervyPirate. Richard Arthur Norton (1958. Nikai. Siim. Spellcast. Vsmith. Richwil. Superior IQ Genius. Looie496. Mirlen. Dan8080. YellowMonkey. Stevertigo. Mr. Vasile. Chowbok. Spitfire. Squeakytoad. Sully. Sebastiano venturi. Rdsmith4. Mikearion. Edgar181. The sunder king. Schnurrbart. Standonbible. Zachorious. Osbus. Sillygrin. N Shar. Yerpo. Vegetator. Piotrus. Ruud Koot. Ocaasi. Outoftuneviolin and his sockpuppets. Noctibus. Chrisjj. Where. Peace237. Teh tennisman. Ynist. J. FullMetal Patch. TigerShark. Outoftuneviolin 9. Tomandlu. Sadi Carnot. Outoftuneclarinet. Nishauncom. Cabe6403. Sitearm. Remiel. OFVWT. Ryguasu. The Thing That Should Not Be. Yamamoto Ichiro. Wuob. Peter. Savidan. Mlaat. Wafulz. Mcmillin24. Tabletop. Tigers4657. Gabbe. Dan Gluck. Earthmichael. Rcorcs. Nihiltres. Trevor MacInnis. Separa. Dcljr. Noleander. Lradrama.O. Spotfixer. Tylertreso. Rankiri. Sted Go. Vanished user. Wawmaw. Shoaler. Minasbeede.O. Poise too. Tresiden. Neohart. Rock nj. Temtem. Zero g. Oontoo. Ocmec. Sango123. Nowimnthing. Mindmatrix. MartinSpamer. Ytrottier. Stevenmitchell. Iridescent. TableManners. Mat8989. Mildly Mad. Outoftuneguitar. Diamondslide. PhDP. Xeatc. Michael Johnson. Stone Oak Drive. Mojosam. Vicenarian. Thebestlaidplans. Terjen. VirtualDelight. Sratt.. Sjsitler92. Matnkat. Ohnoitsjamie. Gnevin. Ykse. Onlyme31. Ciphergoth. Materialscientist. Plumbago. FeloniousMonk. Snowmanradio. SomedatapacketS. Artichoker. StudyAndBeWise. Van der Hoorn. Knucmo2. Pleuu. Just plain Bill. Randomblue. Sviev. Rich333. Slowking Man. Tom Allen. SpNeo. Twci. Zachary. Marskell. Mattinbgn. Valich. Trabucogold. Cybercobra. Wtfmate. Sander Säde. Vicki Rosenzweig. Pendragonneopets. Z10x. PiCo. Wilke. KillerChihuahua. Ricimer. Pharaoh of the Wizards. Messenger777. Samsara noadmin. SpaceTycoon. Pengo. Vanished user. RK. Maxozo. Wikipedia Administration. Xiner. Realknowledge. Michael. Radman622. Neutrality. Barryruth. Shanata. RainbowOfLight. Quietmarc. Noleander. MykReeve. PTHS. UberScienceNerd. Rintrah. Rab Eyye. DMacks. Walor. JRtheDude. Ched Davis. Thetorpedodog. Yournoangei. Brougham96. Rhobite. Zyasu. Netesq. Tijmz. Vanished User 0001. Smartse. Tiptophiphop. Psychohistorian. Tsackton. Modulatum. Yaqix. OGGVOB. Misza13. Xted. Elassint. Pkpyro8. NeuroBells123. Steel. Wteaw. Wayward. NellieBly. Welsh Crazy Frog. IanCheesman. TomTheHand. Pasado. Swazamee.. Mpallen. Zyyx. Sifaka. Shayne T. Smeira. KimvdLinde. Vlmastra. UBeR. Vsast. RG2. NCartmell. Nostalgiphile. Thomas H. Wesley. The Obfuscator. Quadzilla99. Mseas. Coffee2theorems. Mindspillage. Timwi. Shantavira. ZeWrestler.wikipedia. MrMorgan. Rory096. Nixeagle. Scruffy1. Welsh. Prolog. TheTrueSora. Faustnh. Midnightcomm. Wpaa. Songwriter17. Raiph. OutOfTuneViolin. Unschool.5. NeilN. Rfl. Wolfrock. Pond sew. Textangel. Wfward. XJamRastafire. Nsk92. RoyBoy. NameThatWorks. NerdyScienceDude. C. ZimZalaBim. Maqsarian. Jennavecia. Lightmouse. Rednblu. Pretzelpaws. Narayanese. Astatine-210. StuTheSheep. Richard001.Teavee. Tie Bon. Meepster. Mnptl. Vlipvlop. Lord of the Pit. Sarrh. Nicklott. Panoptical. Piano non troppo. John. Thingg. Nova77. J. Temmor. Quadraxis. Schrandit. Tjfulopp. Felix Folio Secundus. Antiuser. Popefauvexxiii. NatureA16. Hpa. Sepia officinalis. Wildnox. SpuriousQ. Sfnhltb. Mindmatrix. The Placebo Effect. Theuser. Theodolite. Sarg. StealthCopyEditor. Mikker. Raul654.I. Inniverse. Mild Bill Hiccup. Moon&Nature. Mixmoney. Zzuuzz. WhyBeNormal. Mgegh. Oidia. Shanes. Headbomb. Tstrobaugh. Tooth Fairy. Mporch. Tysto. Sugarcaddy. Zsinj. Axel147. Tuxedo junction. Tmopkisn. Tar concr. RE. Sciolus. Samgoody777. Reilly. Jazzed. Rebecca. Winhunter. Plaonui. Zarniwoot. PierreAbbat. Tom.). Red lorry. Wdavies. Benclewett. Mishuletz. MarcoTolo. Rich Farmbrough. Senortypant.. Wavelength. SixPurpleFish. Whoutz. Sawahlstrom. Tony1. II MusLiM HyBRiD II. Johannordholm. Peyre. Methcub. Tarinth. Giovanni33. Pydan. Dock312. PrivateRyan.. Staam. NuclearWarfare. Wellwicked94. Pftaylor. Marr Plv. Martycota. Neural. StN. Tarcieri. Taffboyz. Nacre 10. Prodego. Wiki Raja. Ryankhart. Wildie. Six moss. MikeJ9919. Polsmeth. Pinethicket. Blanchardb. Mikker. Wik. Regebro. Seb951. Vewar. RxS. Rjwilmsi. Shed 8*6. Wisdom89. Tyro55555. Wisco. Sandgem Addict. Trevor Andersen. Walton One. Oxymoron83. Volnav. PAK Man. Zaharous. Chick Bowen. Qymme. ZA. Phe enn. Pyroclastic. Saganaki-.R. GSlicer. Ray vivid. Silicon-28. Nihiltres. TimVickers. Ramdrake. Mav. Yday. Gregkaye. Rveek. Nishkid64. Pseiir. StuartH. Farsight001. Versageek. P4k. Azcolvin429. Critter2482. Nick123. Muza47. PrometheusX303. BK4ME. Kevmin. Mivmirre. Philcha. Silence. Wi-king. Theforester. Splash. UncleBubba. Sam. Micahmn. Filll. KnowledgeOfSelf. Sean. Mausy5043. Pgk. Michaelas10. Courcelles. Olorin28. Charlesdrakew. Mgiganteus1. Wasell. Winterspan. TheIncredibleEdibleOompaLoompa. Mattbr. Rubisco. AlphaEta. Tesseran. Strum enge. Mann jess. Scwlong. Sin-man. Nate Slayer0.I. PopiethePopester.delanoy. Ten tinted. Rorrenig. PhilipO. Ydonne 2.L. Vsmith. Mooselover801. Stevenwmccrary58. VegaDark. Uuger. Woody.k. Saltforkgunman. Sir48. Mayumi. Stephen e nelson. Michael Hardy. Itisnotme. Tommy2010. Pyiid. Stfg. Triqqi. Non believer evolution. Tocir. AdjustShift. Sdornan. Ploum. Taw. Plow Col. PhDP. Tixity. NiLoIvEnUtFoTuO. RDBrown. Fred Hsu. Outoftunetrumpet. Neptunerover. Rjwilmsi. Premeditated Chaos. SusanLesch. Mikenorton. Salmanmdkhan. Hello32020. Mchavez. McBeardo. Nethgirb. Vuri. Myrridias. Richard D. Phsource. Supercrazy617. Moss oul. Aunt Entropy. Outriggr. DerHexer. Ttiotsw. Diego pmc. Supertask. Abbenm. John Abbe. Jelammers. Human.php?oldid=392950288 Contributors: 1 use. Tstrobaugh. La Pianista. Nableezy. Johnbod. Dweir. MSchnitzler2000. Kosmocentric. Plinkit. Bobby D. PhDP. Voyagerfan5761. Ano-User.wikipedia. Frunobulax. FanCollector. Alba. Scythia. Silence. Andrea105. Suruena. Kaldari. Addshore. S h i v a (Visnu). Pmanderson. Richerman. Taffboyz. Dolphin51. VetteDude. Jkbell. Entropix. Gregbard. Angela. Armchair info guy. Kennethjackson. Meggar. David Gerard. Athaler. Sam Hocevar. Lexor. MPF. Katieh5584. Smaines. Marknesbitt. CanisRufus. Axl.Verite. Hyacinth. BaseballDetective. Dragons flight. Lexor. Aranel. Fred Hsu. Tutmosis. Armchair info guy. Pingveno. Steinsky. Petter Bøckman. Rreagan007. Stephen Morley. Stephan Schulz. Makeemlighter. Igoldste. Cygnis insignis. Skirby17. Bogdangiusca. AdultSwim. Allinthebrain. Dr. JureCuhalev. Mikenorton. KimvdLinde. TimVickers. Pince Nez. Dabomb87. Twanderson. Pepperosie03. Andycjp. Ethan Mitchell. Nihiltres. Jaknouse. Lucyintheskywithdada. Neelix. Manuel Anastácio. Johnstone. Steinsky. Philippe. Mtg400. Bendzh. Atlant. Dolphonia. Doulos Christos. GRBerry. Binadot. Johnuniq. JeffersonianView. Mark Carden. GSlicer. KyNephi. Coffee2theorems. Friginator. Bngrybt. Orangemarlin. Bongwarrior. Pgk. Auréola. Jooler. Johann Wolfgang. Keraunos. Flexijane. GCarty. RobertG. Sean. JoshuaZ. CapitalR. Tomruen. JohnnyB256. Csernica. Unimaginative Username. Millosh. TimothyRias. Epbr123. Reportsporusskie. Vsmith. Mild Bill Hiccup. Craftyminion. Rabidfoxes. Raymondwinn. Mindmatrix. MAXimum Xtreme. Anlace. BD2412. Wilson44691. Tex. LittleDan. Rajeevvp. George Hernandez.php?oldid=391503292 Contributors: 16@r. RobertG. Francis Schonken. HalfShadow. WAS 4. Silverxxx. Ceramufary. Gilliam. LonelyMarble. Infochanger904. Rennie84. KirbyManiac. Shaw. SmilesALot. Extremophile. Twas Now. Dabomb87. Extremophile. Coelacan. Akersmc. Duffman. Old Moonraker. Tinz. RoyBoy. Suslindisambiguator. Rysz. PDH. Pollinator. J0m1eisler. Mencial.wikipedia. Duncharris. Dojarca. Qmwne235. Mercury. Rdsmith4. clown will eat me. CSVR4. The wub. CCooke. Brion VIBBER. Esobocinski. Blehfu. Jarry1250. Blanchardb. User A1. Timwi. AlphaEta. Robert Stevens. Charles Matthews. Bobblewik. Cam. LilHelpa. Diedo1982. Allinthebrain. Halaster. Piotrus. BrenMan 94. Snowolf. SlimVirgin. CreaoTnismKickR. Legoland. Pennywisdom2099. ArthurWeasley. Zappernapper. Eras-mus. Moondyne. CeoMaiy. Lordaraq. Alansohn. StudyAndBeWise. Dodo bird. Jon186. Aderksen. CommonsDelinker. Dysmorodrepanis. Bryan Derksen. Natalie Erin. Coricus. HereToHelp. J. Aunt Entropy. Dave souza. Brogers0436. Tulostoma. Mangostar. Nemo bis. Elfguy. Duncharris. Snalwibma. Hu12. Richard Arthur Norton (1958.hoyland. Crum375. Aziz1005. Fred Hsu. Onevalefan. Arde Madrid. Funhistory. Shirik. Xmahahdu.39. Bensaccount. Guest9999. Digitata. Wai Wai. Galoubet. Eleassar777. Mrh30. Gary Cziko. JohnWoolsey.org/w/index. Afernand74. Joel7687. Aksi great. SheffieldSteel. Kozuch. Armchair info guy. KnowledgeOfSelf. Marek69. Finell. Bobo192. Henrygb. RK. Anton Kos. Jmc112693. Garion96. Daniel Brockman. Ksanyi. Rich Farmbrough. Fanatix. Tuckerj1976. Luk. DLH. Redtigerxyz. RyanCross. Dannyno. Sanfranman59. Veda784. Ec5618. Chris the speller. Bschopman. Wayland. Bcasterline. Stemonitis. ArcaneMachine. Malleus Fatuorum. Quiddity. Ejosse1. Cyde. Bobo192. TutterMouse. Faro0485. Diadem. Rjwilmsi. Ancheta Wis. Aa77zz. Michael Johnson. Heron. Arbeiter.250. Rling. Bsadowski1. PhilosopherGeneral. Rusty Cashman. Isolation booth. Purnajitphukon. NuclearWarfare. Art LaPella. Smallweed. Samuelscat. Murderbike. StudyAndBeWise.org/w/index. Boreas74. Hibernian. Joe hill. Sonett72. Fastfission. Achmed123456789. Marskell. Gilliam. Huckleboy. Leonardorejorge. Alba. 124 anonymous edits Lamarckism Source: http://en. Henry Flower. Johnuniq. Katalaveno. Zafiroblue05. Jullz. Bleent. Leptictidium. Michael Johnson. Aljullu. Alansohn. TypoDotOrg. Sverdrup. Evertype. JForget. OrangUtanUK. Jsc83. Nwbeeson. Huysman. Andres. Marcus MacGregor. WLU. Mindmatrix. Momabdishu. D-Notice. Artichoker. Yerpo. Vera Cruz. Magneticstockbrokingpetdetective. Macdonald-ross. Antandrus. Theprivateer83. Rich Farmbrough. Tim!. Zamphuor. Ottre. Imc. Spencer. Eurosong. Heron. Tmol42. Octavian history.delanoy. Cesare Barbone. Dudesleeper. FT2. A little insignificant. Horned1. Bosmon. Cachan0001. Astor. Rachel1. 609 anonymous edits Darwinism Source: http://en. Gimmetrow. Brighterorange. ConfuciusOrnis. Cmdrjameson. Stone. 213. Memestream. RussGrim. LittleHow. Jrmccall. Rajender Sethi. Maxis ftw. Akamad. Art LaPella. GameKeeper. Anlace. Hadrian89. Bomac. Dethme0w. Freakofnurture. Oblivious. VacuousPoet. William M. Ryan. Technopat. Alethe. Euroster. Rossami. Philly jawn. Bolivian Unicyclist. Eric Forste. Bevo. Transisto. That Guy. Fetchcomms. Hairhorn. RJHall. Josh Grosse. CenozoicEra. TeleComNasSprVen. Sabik. Xagent86. Gogo Dodo. Acc3ss. Johnuniq. Hereforhomework2. Wheely Guy. Martial75. Jebus989. Redheylin. Streona. Headbomb. Xabian40409. OpenToppedBus. Baegis. CSWarren. Vitovino. Dyed Purple. Iapetus. White Shadows. Dichro. Katharina Simon. Matterfoot. Leptictidium. Dawn Bard. RJHall. Kingturtle. Gaius Cornelius. Samsara. Artichoker. Knight1993. RevRagnarok. Workster. One-dimensional Tangent. Imarealscientest. Pbarnes. Elemesh. Cinik. Ano-User. Marcus334. Shadowjams. Persian Poet Gal. JHunterJ. Richard001. JeffW. A Raider Like Indiana. Pee Tern. Gwern. GN. Damneinstien. Dbachmann. Nakon. JoshuaZ. The Anome. BenSittler. William Avery. Versus22. Lauranrg. Howardsr. Shanel. Memestream.250. IanCheesman. Eastlaw. Merzul. Lord Patrick. Magafuzula. Memestream. TechBear. Macdonald-ross.php?oldid=391895866 Contributors: AS. Michael Hardy. Ankit jn. Halaqah. Null Nihils. Agathman. TimVickers. WAS 4. Morning277. Snigbrook. NuclearWarfare. Colonies Chris. Tide rolls. Novangelis. Rick Norwood. Black Kite. Ragesoss. 22 anonymous edits On the Origin of Species Source: http://en. Nivix. Roentgenium111. Spotty11222. Mais oui!. Vanished User 0001. Arch dude. Fieldday-sunday. MeisterPL. Dumarest. Jsethedwards. Csmwiki. Graham87. Sillygrin. Shoujun. Brian0918. Kuru. Johnuniq. Antandrus. A8UDI. A. AS. Xuehxolotl. WolfmanSF. Miss Madeline. Rusty Cashman. Eric Kvaalen. Dave L. Drmaik. Descendall. SMC.wikipedia. Anadverb. Marek69. RoundSparrow. Gadfium. Brennfalcon. Doubting thomas. Hephaestos. Mateuszica.kenny. Trilobitealive.wikipedia. Eisnel. Androstachys. Jayc. NameIsRon. Cmdrjameson. Koavf. Johnuniq. Esdraelon. OpenToppedBus. Dawn Bard. Wimt. Oldag07. Piledhigheranddeeper. Svetovid. Manning Bartlett. Heliogabalus1. Gyrofrog. Thibbs. Cmayy. Myles325a. Rich Farmbrough. Andres. RedWolf. Fastfission. Smith609. Tide rolls. Marc Mongenet. Headbomb. Lauranrg. Closedmouth. Maurreen. Anonywiki. Bahudhara. Jeargle. Macdonald-ross. Rusty Cashman. Bhoke81. Livingrm. BD2412. SnowFire. Vsmith. Onco p53. Rds865. Bigbird31416. Sjdunn9. Headbomb. Allen3. Knowledge Seeker. Mm40. CharlesGillingham. Saikiri. Infophile. Thunderfun. WojPob. Gil Gamesh. Fastfission. Alienus. Plumbago. PeterisP. Diza. Dysepsion. 446 . Ebyabe. Edgar181. Ruffmonster. From That Show!. Fvasconcellos. Alexrio1013. M Alan Kazlev. Peter Karlsen. Donald. Jsonitsac. Bergsten. Twinxor. DoubleBlue. Parsa. G716. François-Dominique. Mattisse. Slrubenstein. Onnerfors. OlEnglish. Graham87. Orangemarlin. Viriditas. Nihiltres. Aldaron. Vrenator. Conversion script. Omegatron. Phi*n!x. Bryan Derksen. Fang 23. Icairns. JH-man. Riapress. Faithlessthewonderboy. Ngio. Tpbradbury.php?oldid=392624508 Contributors: 1or2. Vintermann. Filll. Cgingold. Pamdhiga. EOBeav. Marshman. Edhubbard. Sander Säde.wikipedia. Curps. Headbomb. Stbalbach. DrKiernan. Knowledge Seeker. Kyleain. Tbhotch. Alba. Husond. AxelBoldt. FuNRtux6. Nedlum. Marshman. Szquirrel. RichardKennaway. Josiah Rowe. Orangemarlin. The C of E. Hmains. Spencer. Garcsera82. Dekimasu.vilbig. HaeB. Ronhjones. DrKiernan. Zoicon5. Srnec. Arkuat. John. H. Ncmvocalist. HumbleGod. Antrophica. LilHelpa. Cephal-odd. Cyan. Thoroughbred Phoenix. Lupo. Jcmo. Seegoon. Brunnock. Alyeska. Pulsar06. Fang 23. Piledhigheranddeeper.php?oldid=392998003 Contributors: ***Ria777. Krclathrate. Evolauxia. Mitteldorf. Ollie senter. Writtenright. Abosaleh911. H. User99. GoEThe. Thatguyflint. Ballista.welsh. Markjoseph125. Amatulic. Barto. Grimhelm. Hillsboro. Tbjablin. Filelakeshoe. Andrewlp1991. Golem Voice. Lightmouse. Tycho. Kekm23. Steinsky. Johnbibby. Tannin. Fuzheado. Jeff3000. Simnicol. Magnani.v2. Allventon. Fred Hsu. HarryHenryGebel. Tony1. Kluv0008. Zachlipton.76. Hugin&Munin. JTBurman. Quuxplusone.Article Sources and Contributors Timeline of evolution Source: http://en.delanoy. Shadowjams. SusanLesch.40.Kurtz. Charlesdrakew. John. Pengo. Theultimatum. Vremya. StN. Samak47. Merovingian. Bender235. Vicenarian. Superborsuk. Antandrus. Skater. Thesoxlost. Nascar1996. Gogo Dodo. Mr. Animum. Paul from Michigan. Bcasterline. Wells. John Lynch. Ashmoo. Duncharris. Magnus Manske. DancingPhilosopher. Dave souza. Robofish. AdamantlyMike. Fred Bradstadt. AC+79 3888. Barnaby dawson. Batmanand. Alan Canon. Adam Zivner. Olivier. Blainster. Athenean. Spammer34567.org/w/index.3. Chocolateboy. Tevildo. Barticus88. Philcha. Rlinfinity. Simon12. VolatileChemical. Greeneto. Gene Nygaard. THEN WHO WAS PHONE?. Neverquick. Civil Engineer III. Leroy Ussery. Duncharris. Rst20xx. BenB4. Gomm.org/w/index. Chris. Surv1v4l1st. KevinBrowning. Leandrod. Jrmccall. Tuxedo junction. Akasbrian. Mav. Wiki Raja. Wikididact. Fences and windows. Ciphergoth. Dendodge. Martinssaz. Evolve1976. Brooke021. Uncle Milty. CMacMillan. Neonblak. KWMec. Philip Trueman. JaGa. Vmaldia. Edivorce. Havermayer. Francis Hoar. Vcrs. Quiddity. Spencer. Headbomb. Jag19. Mattisse. Ellen Kearns. William Avery. Zef. LordAmeth. Erich168. Linas. Maelnuneb. R:128. Reywas92. Brufiki. Dcljr. Ccarroll. Mchavez. Selkem. Lexor. Jappalang. Reedy. Justtheinformation. Auriam. Boreas74. Causa sui. Theresa knott. Proxima Centauri. Abdul Muhib. Mikker. AnonMoos. PDH. Gniniv.urs-o. Yuh66h.org/w/index. Mercury. Bania. Jaraalbe. F345. Join Tile. Classicstruggle2. Anna Lincoln. Neutrality. Z10x. TheAlphaWolf. Rich Farmbrough. Philmcgrove. Vclaw. Tomisti. Odycee. Goalie1002002. Eaglizard. Minia23. Varlaam. Jatkins. MickWest. Dbabbitt. Ballista. LeonardM. Isnow. Llehctimj. Big iron.sanat. Jusdafax. Philip Stevens. Wordwright. Twang. Vanished user 03. RyanGerbil10. Richard001. JorisvS. Edward. Leptictidium. Ling.253. Madeleine Price Ball. Blades95. Pauli133. J. Aristotle2600. Safay. Extremophile. S. DSatz. Edital. Satori Son. Tiptoety. Gaggle. Barticus88. DJ Clayworth. Fred114. Michael Devore. Enchanter. Autodidactyl. Discospinster. Keilana. GoingBatty. Hadal. Rsrikanth05. Salleman. Nn123645. Kevinalewis. Trialsanderrors. Xiner. Brichard37. Download. Richard001. Pinethicket. Kvn8907. Gabbe. Egmontaz. Rjwilmsi. Ms2ger. Random account 47. MayerG. Karvok. Royboycrashfan. AdiJapan. Stappsclass.Nut. Circeus. Syncategoremata. Dave souza. Ragesoss. Pharaoh of the Wizards. Discospinster. McSly. Wavelength. AlphaEta. Easterbradford. Sin-man. Dreish. Tmol42. Mrfunnyd. HGB. Vatsan34. Nosson77. Emil Teofanov. MaxSem. Havermayer. Sweetness46. Ams80. Qertis. Alan Liefting. RainbowOfLight. Ed Poor. Raul654. Martin-vogel. Ettrig.php?oldid=382880304 Contributors: Adzyaye. Guettarda. GSlicer. Orphan Wiki. Chris55. Burner0718. Syzygos. SpuriousQ. Urhixidur. Beyond My Ken. Jrockley. Grim23. Shrimp wong. Filll. Armchair info guy. Butterscotch. Trevor Andersen. CardinalDan. Jacob1207. Astatine-210. LGagnon. Kleenex11. Jamesdowallen. GoonerDP. OldakQuill. CJLL Wright. Talkstosocks. John D. J. Sonicblade128. Nneonneo. Lauranrg. Opabinia regalis. Immunize. Connolley. Fred Bradstadt.php?oldid=385164271 Contributors: 3tcetera. Dysmorodrepanis. SchfiftyThree. DavidCBeck. DanielCD. Ronhjones. Rednblu. Emw. WolfmanSF. Gdr. Bender235. Igiffin. Robert K S. Extremophile. Anwegmann. Gadfium. Rusty Cashman. Fubar Obfusco. Tim Q. Hugo999. NancyHeise. Valich. Rusty Cashman. Akamad. Koebie. Lquilter. Stefan. Gakrivas.xxx. Iwanttofitin. Sala427. Commander Keane. KimvdLinde. Jefffire. DarwinianMan. MasterOfHisOwnDomain. Stiepan Pietrov. Leibniz. NatureA16. Coyoty. Vanished User 0001. Ante Aikio. Shyamal. Ewlyahoocom. StuffOfInterest. UtherSRG. 213 anonymous edits Saltationism Source: http://en. Cohesion. Matthew Yeager. Alansohn. Kimiko. Lilac Soul. Hippalus. Phthoggos. AxelBoldt. J. CSWarren. StN. Klarno. Berton. Armchair info guy. Peripitus. Zundark. RJHall. Template namespace initialisation script. Xaliqen. Slarre. Lacatosias. Hrafn. Mmcannis. Justinfh2Point0. Codemonkey87.org/w/index. Gdr. Lexor. KnowledgeRequire. Peruvianllama. Enrique Cordero. Taco325i. Bomac. Philip Trueman. GSlicer. Taffboyz. Sietse Snel. EchetusXe. Wereon. Kompar. Sleeppointer. The Thing That Should Not Be. Interrelation. DirkvdM. ParlerVousWiki. DARTH SIDIOUS 2. Someone else. Deus Ex. Chris55. SMesser. Yvori. Martial75. Ealdgyth. R. Malljaja. DeLarge. Dissembly. Petri Krohn. Juliancolton. Discospinster. SU Linguist. Wikid77. Harryboyles. IronGargoyle. Hitman59.Bastedo. Skizzik. Lisatwo. Jamesmarkhetterley. PoochieR. AlphaEta. James A. Dreaded Walrus. Kansan. Williamb. Narayanese. Samsara. Icairns. Steinsky. Abecedare. Eternian4ever. Aidan Elliott-McCrea. Everyking. Rich Farmbrough. Vegetator. Tbjablin. Vero. Dgri. Svetovid. Alexei Kouprianov. Pwner594. Mainefan1.wikipedia. Jagged 85. JohnCD. Kaimiddleton. Magda wojtyra. Александър. Anlace. Specter01010. Alansohn. Teapotgeorge. Aitias. R Lowry. Jack Merridew. Juliancolton. Meco. Timwi. Calor. John Vandenberg. Mr Chuckles. Woohookitty. SirGrant. Rune. Johnor. Tony360X. Ipooponyouforyou. Ashmoo. Hephaestos. VBGFscJUn3. Tonyfaull. Trueblood.org/w/index. GrahamP.wikipedia. Mervyn. Portillo. Sandahl. 15 anonymous edits Orthogenesis Source: http://en. Zoe. Kbdank71. BigBurkey. Vanished user. Bhadani. Gregbard. Yamamoto Ichiro. Malick78. Soulkeeper. Peter b. Quadpus. Aliekens. Drumbeatsofeden. The Thing That Should Not Be. Bongoo. IstvanWolf. Maximus Rex. Andkore.0. Vsmith. Emmanuelm. Alansohn. Garygromet. Bryant. FvdP. Conversion script. Samiam1955. PiCo. Jonas Mur. Smeggysmeg. Enviroboy. Albie34423. Koavf. Brian Crawford. David Cheater. Escape Orbit. Tuganax. Adrian 1111. CommonsDelinker. Voyevoda. GrahamColm. Popsracer. Logicus. Tuxedo junction. Jay ryann. Robert1947. Plumbago. Vasiľ. Bcasterline. TeleComNasSprVen. Mgerb. Kablammo. Cyberix. Dave souza. JakeVortex. Opie. Rlevse. JoeshB. Hadal. Timwi. Can't sleep. Croft. Courcelles. John37309. Benjaburns.). Gdr. LovesMacs. SandyGeorgia. Kapow. SchnitzelMannGreek. Leonardorejorge. Lazylaces. Soulkeeper. 838 anonymous edits History of evolutionary thought Source: http://en. Mattbr. Waggers. CeoMaj. Igiffin. Stirling Newberry. DharmaDreamer. Moulder. PSWG1920. Kdbuffalo. ConfuciusOrnis. AxelBoldt. JLaTondre. Deb. Categorer. Yelyos. Ttiotsw. That0neguy01. Mladifilozof. Henrygb. ForestDim. LFaraone. Malten. Slrubenstein. Tijfo098. Sbeath. Rumping. Antiuser. Noisy. Shrimp wong. RHaworth.k. 437 anonymous edits The Genetical Theory of Natural Selection Source: http://en. Keno. Nuttycoconut. Alfalfa9.org/w/index. Lizard1959. 1exec1. JamesHilt62. Grantus4504. Vachung 82. Andrew c. Gabbe. ThePointblank.wikipedia. Wikieditor06. J. Tuxedo junction. Agathman. Deeplogic. RScavetta.250. R. Johnuniq. Vivian. Cement123. Myanw. Minisimon. Mouse555. Skarl the Drummer. Irishguy. El C. Terence. Mattisse. O. Szquirrel.org/w/index. Irish Souffle. Cat Whisperer. WAS 4.php?oldid=392814660 Contributors: 10outof10die.muller. Maniadis. Arminius. Gregbard. Sam Medany. K. Wetman. Reedy. Felinasmith. Northfox. Azeleas=awsome. Hmains. Kbp001. Simeon24601. Andreybigb. Tgearty. Gamesta3. SteveBaker. Jweiss11.0. Henrionline. Benhocking. PDH. Ratphobia. Dr. Ksyrie. Portillo. Jfdwolff. Kleuske. NewEnglandYankee. Escape Orbit. Echoray. Traxs7. Alexpavlov12345. Djlayton4. Omicronpersei8. Txomin. Smith609. Warofdreams. Johnuniq. FeloniousMonk. Duncharris. Radagast83. Gimme danger. Dave souza. Valich. Murtasa. Samsara. Syncategoremata. Andersrask1977. Cyde. Palnatoke. Woohookitty. GromXXVII. Yankeefan292. Carrionluggage. Hrafn. Infraredeclipse. Emw. Hut 8. CzarB. clown will eat me. Olianov. Edward.org/w/index. Aianrnoens. Writtenonsand. Jonverve. Rico12121212. Des. VIDEOGAMES. Prohlep. Ragesoss. Edgar181. The Anome. Spizzer2. Odd nature. Silverfish. Jusdafax. Enviroboy. Geoff. David Gerard. Icairns. Taneli HUUSKONEN. Johnuniq. 18 anonymous edits Neo-Darwinism Source: http://en. Betacommand. Agathman. Dan Polansky. Orangemarlin. Mailer diablo. Gog. FisherQueen. Manco Capac. Richhoncho. Hmains. Jrockley. Daniel5127. Donarreiskoffer. MAURY. Blotchhead. Mensfortis. Yerpo. JeremyA.m.org/w/index. Rich Farmbrough. Agathman. Dripping Flame. Nicolae Coman. Ed Poor. Happysailor. Lunakeet. Gimmetrow. Kjkolb. Martial75. AmericanKido. B00P. Gloriamarie. Mboverload. J'raxis. Richard001. Qurat-ul-ain Basit. Proyash. Fabricebaro. Azcolvin429. Headbomb. Senortypant. Luna Santin. Mild Bill Hiccup.Article Sources and Contributors Audacity. Tiddly Tom. Barak Sh. Headbomb. Azcolvin429. 84user.wikipedia. MetsFan76. TimothyHorrigan. Jredmond. TimVickers. Techman224. Template namespace initialisation script.V. Beboots. Senortypant. Kzollman. Flarity. Rjwilmsi. Jondw. Digitwoman. Uzzo2. Sharkface217. Mcy jerry. Kingdon. WLU. Sghussey. John D. Woudloper. Ocsadanielsw@hotmail. Mlewan. Ettrig. ValenShephard. Headbomb. Arkarind. Reinyday. Firsfron. Zeamays. Myahoo. StudyAndBeWise. Calebrw. Philosophistry. Rob Hooft. Savana-ona-rolla. TheKMan. Sraoof18.10. Ezra Wax. Marquez. DeadEyeArrow. Ec5618. Mbc362. Jstanley01. Cerealkiller13. JForget. GlassFET. Jengod. Macdonald-ross. Guettarda. Joriki. Earlypsychosis. Brian0918. Curtis Clark. Nuttycoconut. Man with two legs. Security`57. Michael Johnson. Pgk. J. GoEThe. Azcolvin429. Harley peters. Meggar. Capricorn42. Wavelength. Ashnard. R Lee E. Gtrmp. B. Dbachmann. ElijahOmega. Orangemarlin. Bryan Derksen. Liftarn. DGtal. Aunt Entropy. SEWilco. Peripitus. MayerG. Daniel J. Filll. Garik. Silly rabbit. Bkkbrad. Massimo74. Kaiba. Gnomon Kelemen. WadeSimMiser. Z10x. Fationia. Robert Stevens. Phantomsteve. SimonD. Samsara. AresAndEnyo. Corvi42. Isaac Dupree. Bobianite. Hunter. Inwit. Heironymous Rowe. AxiomOfFaith. Kilmer-san. Thunderfrog07. Nads240721. Koveras. ErikHaugen. TedE. Rafikgl. Ucucha. Ec5618. Keilana. Orangemarlin. JavaTenor. Keesiewonder. Job tungol. Jooler. Ixfd64. Davril2020. Davril2020. Obli. Neverquick. Alan Liefting. Gary King. Tony Fox. Thomas Arelatensis. Coelacan. Menchi. Dawn Bard. SWAdair. BenAveling. Old Moonraker. T-rex. Conversion script.org/w/index. Александър. Camw.org/w/index. Opabinia regalis. GraemeL. Zhou Yu. Richard001. Madhero88. Human. Gaius Cornelius. Reconsider the static. Vicenarian. Calabraxthis. GSlicer. RandomP. PoolPartay. Petiatil. Strictscrutiny. Rock nj. Sexy Back 16. Persian Poet Gal. MTDinoHunter. Onevalefan. I am not a dog. Lindosland. GuyChaham. Apeloverage. Rgamble. Wheely Guy. Pkeck. Wayne Olajuwon. RoyBoy. Danielrcote. GeneralBelly. Nwbeeson. Lexor. TimVickers. Bcasterline. Trimp. Shoemaker's Holiday. Rory096.khot. John Lynch. SheffieldSteel. Mrmariokartguy. ChrisDuben. Skullslayer15. Snalwibma. 22 anonymous edits Modern evolutionary synthesis Source: http://en. Piano non troppo. Cribbswh. Ed Poor. The Thing That Should Not Be. Digitwoman. Jpbowen. Dingar. Magister Mathematicae. Mitch Ames. Tide rolls. Mwills011. Hkhenson. Robert Stevens. Mchavez. Sylentkorean. Timo Honkasalo. 477 anonymous edits Fitness Source: http://en. JoshuaZ.php?oldid=393073936 Contributors: 2D. Waldow. Glen. Snarius. 2D. Melaen. OhanaUnited. Lexor. Rogue-pilot. Shyamal. Tom. DiLLuS. EPM. ForestDim. Epbr123. Twirligig. Jojhutton. BirgitteSB. Artichoker. Adrian J. Hertz1888. CanisRufus. Odd nature. Equendil. Bobo192. Cavrdg. TimVickers. Lexor. Spencer. Jesus954. Vanished user. Kjells. Allstarecho. Cyde. Resmar48. Jagged 85. Roland Deschain.php?oldid=366488537 Contributors: Alan Liefting. Vsmith. Conversion script. Waldow. Stiepan Pietrov. Angr. Drbug. Graft. Rambam rashi. Pterodactyler. Margareta. Belovedfreak. Dionyseus.thomson. Joannamasel. TehBrandon. PDH. DoubleBlue. Godisnotmocked. Filll. Jaxl. Portillo. Dionyseus. DoktorDec. Mav. Mindmatrix. Erikhansson1. Kjkolb. LossIsNotMore. Jhbadger. WikiLaurent. Kzollman. Lantios. Titoxd. Gaius Cornelius. Thelivingbrian. Oxymoron83. Nadiatalent. Bairh.250. Brockert. Pinethicket. Gregbard. WAS 4. Fastfission. Bueller 007. HKT. Guettarda. Derek Andrews. Jdvelasc. Furor1. Hazzeryoda. Ldavies. Alienus. Mengela. Superborsuk. Ryanmcg2006. Senortypant. Nakon. Yuanchosaan. Iamsnorlax. Rico191919. The wub. Ospalh. DonSiano. Dysepsion. I do not exist. Hult041956. Horselover Frost. Malo. N5iln. Delirium. Nectarflowed. Tamfang. Frank101. KimvdLinde. Earl Andrew. Shot-caller8. ChXu. Akendall. VoteFair. R Lowry. MathEconMajor. Syp. SomeStranger. Hu12. Chowbok. Rhetth. Can't sleep. Authorized User. RexNL. Vimescarrot. Vcolin. Samspeedy1985. Tommy2010. Lexor. Snalwibma. TedE. Duncharris. MER-C. King of Hearts. Jarhed. Gabbe. Azcolvin429. Movses. Dominus. Wimt. Carbuncle. 5 albert square. Pavel Vozenilek. HiEv. Sandahl. Woohookitty. Tobby72. Tempodivalse. Hut 8. CardinalDan. SpuriousQ. Lowellian. Justforasecond. Fama Clamosa. Dtheobald. Grim23. JH-man. Kzollman. ShelfSkewed. Kripkenstein.delanoy. Christian Skeptic. 104 anonymous edits Evidence of common descent Source: http://en. Mswake. VVtam. Hibernian. Smalljim. Morenooso. Heron. MarcoTolo. Gloriamarie. ClamDip. Thatguyflint. Nunh-huh. Aznmastermind5. Jefffire. Masterofsuspense. Lee Daniel Crocker. JFFCakes. Ed Poor.. AnonMoos. Julianonions. Dave souza. Husond. JaGa. Mccajor. Dekimasu. Richard001. Slrubenstein. Hqb.php?oldid=347178552 Contributors: Adoniscik. Salvio giuliano. Dhochron. Gogo Dodo. Harland1. Nivix. Headbomb. Snowmanradio. Hrafn. GregAsche. Jebus989. Andre Engels. Quietmarc. Richard001. Guettarda. Sango123. Nectarflowed. Hello32020. Carradee. Sammalin. Knowledge Seeker. Winterschlaefer. Bender235. Casull. Lopoh. Bobblehead.alsayegh. Peteraandrews. Giftlite.J. Joe Decker.delanoy. Canada Jack. Digwuren. Orangemarlin. Richard001. Lexor. Livingrm. Minimac. DrThompson. Bonerface. Shamrogorr. Menchi. DabMachine. Borgx. BG. Sir Vicious. Leivick. Tannin. Dinofan135. Anthony. Jon186. Aldado. J heisenberg. Fracker. Meestaplu. Pyschedelicfunk. Minhtung91. Fubar Obfusco. Courcelles.delanoy. Rjensen. Timwi. ConfuciusOrnis. DavidL (usurped). Lijealso. Kukini. Eeekster. Decltype. Jason Potter. Dawn Bard. Aecis. Bobo192. Sander Säde. Lordandmaker. Diannaa. Hadrian89. Smigs. HTBrooks. JHunterJ. AnakngAraw. RadicalBender. Jogers. Tameeria.. Apokryltaros. Excirial. Cenarium. Brighterorange.wikipedia. BlaiseFEgan. Falcon8765. Ian. KimvdLinde. JoshuaZ. JoanneB. Maximus Rex. Filll. Reinis. Christian Skeptic. I am not a dog.org/w/index. JRother. Dougweller. Kukini. KillerChihuahua. Livingrm. Mouse8128. TDogg310. Firsfron. Richard David Ramsey. Springnuts. Tail. RxS. J. Tautologist. Haggiaomega. Qxz. Join Tile. Fastfission. Jon Ascton. BlueGoose. Jni. Briggsalex. LedgendGamer. Nihiltres. MartinHarper. Hydrogen Iodide. Lightmouse. Mygerardromance. Headbomb. Cornopeanus. Froonyhoob. Deli nk.php?oldid=392613323 Contributors: 168. Fences and windows. Tomtheman5. DannyMuse. Duncharris. Brianrein.php?oldid=384873781 Contributors: 168. Michael Johnson. Jarhead483. Brightmoon. Rusty Cashman. Robert Stevens.wikipedia. TonyBallioni.wikipedia. ConfuciusOrnis. DancingPenguin. Chris5858. Teh roflmaoer. FaerieInGrey. Fred. Joxernolan. Iridescent. Doc Tropics. IRP. Aunt Entropy. Myanw. La hapalo. Freedomlinux. Octavian history. Andycjp. Richard. Jackie moons. Dj Capricorn. Naddy. Hibernian. Ec5618. Ground Zero. Conversion script. DRosenbach. Teenageactivist. Erik Corry. DLH. PinkPanthress. The Anome. Ano-User. WTGDMan1986. Imjustmatthew. Patrick0Moran. The Man in Question. PMDrive1061. Harburg. Chad625.php?oldid=390608842 Contributors: Abrahami. Iridescent. Vasiľ. JoaoRicardo. Draicone.250. The Merciful. Beland.U. Rusty Cashman. Filemon. James089. Ljhliesl. Wiki alf. Jefffire. Fiskeharrison. T. Loren. Nakon. The Rambling Man. Shyamal. Rich Farmbrough.delanoy. Logicus. Opabinia regalis. Knowledge Seeker. Rjwilmsi. Backmask. Aunt Entropy. YouWillBeAssimilated. Graham87. DLH. JoeSmack. Connormah. Doczilla. Doc Tropics. Pmanderson.wilton. Nibuod. Matthew Yeager. Nahallac Silverwinds. Headbomb. Extremophile. Onevalefan. Sp. Steinsky. DARTH SIDIOUS 2. Andonic. Ottre. Shadowjams. Likeminas. Cheddarbob23. Opie. Obli. JaGa. Sven Heinicke. Neo-Jay. Dziewa. Bobo192. Cornellrockey. GreatWhiteNortherner. Mrl. Splanxna. Ollyoxenfree. Balzi. Tom harrison. I'm the Cavalry. Dfinch. Zappernapper. Smartse. I am not a dog. Kraziur.. Amwyll Rwden. Hugin&Munin. FeloniousMonk. Poeloq. John Vandenberg. Red Act. Ambystoma. Fences and windows. Princeofexcess. ForestDim. Fratrep. Aunt Entropy. TimShell. Wlodzimierz. Ceyockey. Melanie or mely. Hrafn. Frenkmelk. Ivankb. Ground Zero. Pgold. Finell. Nicwright. Silicon-28. Rumping. Pdcook. Bongwarrior. SmurfLover1982. Randwicked. RA0808. Karan. Jockcampbell. Archaeopteryx. MartinSpamer. JoshuaZ. Some jerk on the Internet. Pjvpjv. Leptictidium.. TedE. Shadowjams. Mr. Keegscee.com. Run!. M. AlphaEta. Horselover Frost. Bob f it. Aalexandros. VityUvieu. Vsmith. Neutrality. Yamamoto Ichiro. Tiddly Tom. Voyagerfan5761. PAK Man. Persian Poet Gal. Gnomon Kelemen. Bebo liebe. NellieBly. Lexor. Razorflame. YanaGator. Alansohn. Armchair info guy. Karol Langner. Sander123. MichaelSH. Dina. Anarkisto. The Ogre. GSlicer. Paul August. Vsmith. Oldekop. ScientistDragonmouse. Haemo. Benjaminmyklebust.. Auntof6. Afaz. Kim Bruning. Vietbio. Dawn Bard. Bibliophile20. Jedi6. JH-man. Asrghasrhiojadrhr. Samsara. Pbarnes. Stephenb. Gaius Cornelius. Johnuniq. Andonic. J. Dave souza. Radiocar. Kafziel. Marknen. Fr78. LilHelpa. Webridge. Tyrenius. Andrea105. FeloniousMonk. Blove maple. Rhobite. J Di. Alex. Silly rabbit. DRosenbach. Nerdygeek101. Builder w. Demfranchize. BrainUser. Northfox. Uncle Dick. William Avery. The Transhumanist. Can't sleep. Sdlr. Ungtss. Kuru. Mynameisvirgil. The morgawr. Eequor. GSlicer. Rednblu. Grafen. Alec . JWSchmidt. Livingrm. Philip Trueman. Darrenhusted. SallyForth123. Sceptre. RHaworth. EdJohnston. Susurrus. Jebus989. E-citizen. Spammer34567.delanoy. Headbomb. Dave souza. Johan1298. Bloodofox. Memestream. Tedernst. Bheest. JmCor.v2. Tstrobaugh. Plumbago. TableManners. Transcendence. Burntsauce. Andrewpmk. Vipinhari. Bubbamagic87. Hrafn. Andy Marchbanks. BlytheG. RoyBoy. Vsmith. Missbay67. Mindmatrix. Jim62sch. DO11. Silence. EarthRise33. Lexor. Rich Farmbrough. Dave souza. CaptainAmerica. D.wikipedia. NamfFohyr. Wafulz. Hesperian. Lotje. Spitfire. Alexs.. SpuriousQ. Xaosflux. Dude112233. TimVickers. Sam Hocevar. Dfarrar. Io Katai. David Shear. Acroterion. JustSomeKid. Ramdrake. Dreadstar. Discospinster. Dbiel. J. Condem. Bless sins. Omicronpersei8. Akerans. Vanished user. J. Aquishix. G4rfunkel. Professer elling. Madigan531. Kevinmon. Trusilver. Armchair info guy.. Shawnc. N328KF. Chase me ladies. Hairy Dude. 122 anonymous edits Heredity Source: http://en. Shenme. Ed Poor. Orion 5. Duncharris. Mietchen. Frap. Catgut. Acather96. PaulRuttenfromBrunei. RDBrown. Nbound. Missvain. Fernando S. Nikai. Bowlhover. Squirt2711. Wik. Sslevine. Debresser. Duncharris. Peak. Antandrus. Cureden. TestPilot.e. Daughter of Mímir. DerHexer. Granf. Tommy2010. Cuaxdon. MaseratiFerarri. Ilyushka88. Qertis. Roland Deschain. Teapotgeorge.. Ohconfucius. Gombo. Phantomsteve. Fbartolom. Duncharris.wikipedia. Croft. Snowolf. Rjwilmsi. Will Beback. LittleOldMe. Shadikka. Andres. Haham hanuka. ArthurWeasley.5. Kilonum. Vanished User 0001. PhilipO.5. Encyclopedia77. Vsmith. Spa toss. Vanished user. Fastfission. Discospinster. Stevenmitchell. Dsbrod. Burris. Jorgesalgueiro. Filll. Killiondude. 447 . XxCrazySnoopxX. Ourai. Jusdafax. Emperorbma. Gzhanstong. Lang. Sommers. Gurch. Tuxedo junction. Herbee. Yamamoto Ichiro. TheDJ. Altenmann. Memestream. Mausy5043. Levineps. Pinotgris. Charles Matthews. Eall Ân Ûle. LDHan. CapitalR. Jefffire. Gtrmp. Gaius Cornelius. NawlinWiki. Petri Krohn. Antandrus. Duncharris.decal. Pariah. Smiggle16. WAS 4. Nick. Maurog. Vanished User 0001. Northfox. Join Tile. Gökhan. SchnitzelMannGreek. Synchronism. Viriditas. Stemonitis. Obey. Snalwibma. Psbsub. DavidWBrooks. Lindert. BenB4. Jimmy Pitt. Mr Stephen. clown will eat me. Uriobolski. Littlealien182. Poor Yorick. Apparition11. Blacklake. CanisRufus. Edward Ockham. Tide rolls. TimVickers. Rotational. Deglr6328. Bcasterline. TangentCube. Mark Richards.K. Ewe nik. TedE. Armchair info guy. Crystallina. Glenmin. La goutte de pluie. BW52. K. Mkamensek. 94 anonymous edits Common descent Source: http://en. Shiritai. KittyKAY4. Smoove Z. Vegetator. John Quiggin. King of Hearts. Soulkeeper. Michael Hardy. Rcrosby. Brian0918. Tim Long. Fubar Obfusco. Insanity Incarnate. Snalwibma. Evolutionsucks. Finngall. Bongwarrior. Daecon. Johnuniq. Mpvdm. Dawn Bard. GusChiggins21. Andrewpmk. TomS TDotO. Rusty Cashman. KillerChihuahua. Mastad. The Matrix Prime. Bhawthorne. Axeloide. Boris Živ. Fangfufu. Mattb112885. Gary King. A. Makemi. Van helsing. Mboverload. Narayanese. The Nut. DVD R W. Epbr123. CWY2190. Grafen. Nihiltres. Mchavez. Nascar1996. Oleksii0. Dr Dima. Axel147. Cvata. Forluvoft. Yamamoto Ichiro. Drmies. Dysmorodrepanis. Writtenallover. Jcobb. Mintleaf. Deville. DabMachine. CryptoDerk. Loren1994. BrokenSegue. Bogey97. Voyajer. Salamurai. Odiem00n. Samsara. Joie de Vivre. Lewa. UtherSRG. Kodemage. Memills. Waldow. Mrsssgt. MacMed. Magnus Manske. Malevious. Jamesooders. Hydrogen Iodide. Andres. Radon210. 7. Scepia. TomTheHand. ChrisCork. Stemonitis. Porqin. Gsmgm. Phantomsteve. Wayne Olajuwon. Bueller 007. Wedian. Eatme20005. Dan100. Kazikameuk.250. JoshuaZ. BananaFiend. Rror. Melsaran. Adriaan. Andre Engels. Smartse. Fanatix. Dodgydobson. TheOtherJesse. Hrafn. Giftlite. Gr8niladri. Atubeileh. 478jjjz. Metzenberg. Majorly. Katalaveno. Leandro Palacios. Openstrings. Delldot. Freakoverdose. LookingGlass. Nomer0591. Iridium ionizer.wikipedia. Korou. Ches88. Danski14. Waggers.. Tuxedo junction. Emw. ArielGold. Davidhorman. Miquonranger03. Seahorseruler. Taw. Aircorn. Michael Devore. StripeyBadger. Bioephemera. Jakirfirozkamal. Sionus. Nethgirb. Electric goat. The Anome.. FayssalF. Ryanjunk. Oxymoron83. Isilanes. Phlyght. Pinethicket. Wolfrock. Izehar. Gaius Cornelius. MAXimum Xtreme. Kyoko. Ravenswood Media. PleaseStand. Mav. Nhandler. Michael Johnson. Livajo. Khatru2. Pilotbob. Sithu. Alikingofkings. Haetmachine. Ktfreese. Rich Farmbrough.org/w/index. WAS 4. Sander123. Ketiltrout. Iridescent. Barsonian20. Quintote. Sasata. Cantus. Bryan Derksen. Therucks. CanadianLinuxUser. AnonMoos. Eric-Wester. Jerry Zhang. Jason. Bevo. Ben1002. Decltype. IanCheesman. AxelBoldt. Michael Hardy. Ioeth. Duncharris. Tide rolls. Steinsky. Conversion script. Luuva. Tktktk.wikipedia. Fubar Obfusco. Zerokitsune. Ooikahkooi0507. Pixeltoo. Gilgamesh he. Stevenj. Richard001. 65 anonymous edits Mutation Source: http://en. Duckling78. Hobartimus. Bobamnertiopsis. Dr d12. Tohd8BohaithuGh1. Meepmoop. YDaniel7. Bhadani. Ajgorhoe. Dnvrfantj. Citizen Premier. Guitarist0820. Minimac. Oxwil. FeloniousMonk.Rafe.. Gioto. 15lsoucy. Aled 12345. Cimex. Dancey2.delanoy. Wickey-nl. Yamamoto Ichiro. Aranel. Duncharris. Aunt Entropy. TFOWR. Jeff G. Kcordina. Guillom. Ucucha. 21655. Furrykef. Gcm. 65.lifeguard. Timwi. Jefffire. TestPilot. Cww. WhatTheBlack. Cyde. Oneiros. Ettrig. Jerzy. Camw. CJLL Wright. Plop. JoanneB. Zahid Abdassabur. NickSeigal. ImpalerBugz.96. Atulsnischal. LestatdeLioncourt. EverSince. Dingopup. Saganaki-. Anastacie. Turkishbob. Joelmills. Magnus Manske. Alienus. Myanw. Ukexpat. Grcaldwell. Lordlosss2. Etxrge. Arjun01. Ndteegarden. Jeffmcneill. Kipholbeck. MECU. Habbilow. Deryck Chan. Bwangia. Keilana. Captain-tucker. Genetics411. JaGa. Fedir. TimVickers. Oda Mari. Thingg. Nikola Smolenski. JForget. Quadell. MBisanz. Gabbe. Axel Driken. Friginator. Nadsozinc. GK. ClockworkTroll. Kupirijo. Logan. Cremepuff222. Johnuniq. Dysmorodrepanis. Hordaland. Elvin fimel.27. Lid6. GSlicer. John. Ed Poor. Dave souza. Kungfuadam.josh. Johnstone. Rgamble.com. Doczilla. KathrynLybarger. Freedom to share. Steel. Bar fly high. Ex nihil. Snigbrook. PDH. Jagged 85. Bradshej. Lova Falk. NeilN. Shenme. Ghost2. Sverdrup.. Pb0823. J. Eperotao. Samsara. Gwernol. Gtstricky. Deskana. HJ Mitchell. Dysmorodrepanis. Icek. Joycierules. Gabbe. Instinct. Mike. Jonahib. UkPaolo. EPM. KimvdLinde. LeaveSleaves. Reedy. NuclearWarfare. Jmjanzen. Jamie Lokier. Jon Cates. CodeCat. Leithp. TheresaWilson. Hadal. Abstraktn. AdjustShift. IMatthew. Tuxedo junction. InBalance. Deathgod1995. SomeStranger. GraemeL. Harland1. Pgan002. Biskot. Ivan Bajlo.Article Sources and Contributors TheChrisD. Cfsenel. Ngsmart. Markhurd. Kaarel. ParisianBlade. Giftlite. Whosasking. Sintaku. SallyForth123. AnonMoos. Daarznieks. Timwi. David D. Enviroboy. WaysToEscape. Alai. Paps 70. Aliekens. WikipedianMarlith. Eloquence. Shell Kinney. D6. Naohiro19 revertvandal. Bobo192. Hoppity25. Tmol42. Amaltheus. DARTH SIDIOUS 2. Boing! said Zebedee. SimonSayz. Hadal. Josh Parris. ScaldingHotSoup. Macy.. Fanatix. Frankie816. Jiy. Enfresdezh. MiRroar. Steven Zhang. Matdrodes. JohnRobertMartin. Husond. SixteenBitJorge. Graham87. ZuluPapa5. Hylobius. Graham87. Graft. Jonny1q2w3e4r5t6y7u8i9op. Luna Santin. Hede2000. Jakeisbaked. Juliancolton. Bencherlite. Kungfuadam.132. Marj Tiefert. Juicy fisheye. Dendodge. Academic Challenger. Big iron.wikipedia. Ronhjones. Aadamescu. TUF-KAT. JWB. Gadfium. Kosnapyo. Isilanes. Chaosof99. JaGa. NYKevin. Satyrium. MONGO. Abdullais4u.. Russell. Poechalkdust. Bender235. Reconsider the static. Jinxed. Shadowjams. Giftlite. LovesMacs. Julian Mendez. Delmonte. Showgun45. Backslash Forwardslash. Gracenotes. Template namespace initialisation script. FinnWiki. Electrified mocha chinchilla. Benhocking. Thewall2. Eliyak. Barefootmatt. Flubajub boy. Ettrig. HTBrooks. Jkoeslag. Aitias. Graft. Joyous!. Marek69. Tellyaddict. Gleng. Josh Grosse. Doc Tropics. Abdullais4u. Mokotillon. Fungusnunchuck. Dxballer08. Zerothis. Shanes. Kzollman. Sushant gupta. Diego pmc. PierreAbbat. Mac mckinlay. Mouse555. Uncle Dick. NickSeigal. Professor marginalia. Methcub.. Marek69. Forluvoft. Redwrathe. Cool3. Snek01. Protez. Jennavecia. ITxT. R'n'B. Johnuniq. Jwissick. Hornlitz. Geologyguy. Alex314058. Anypodetos. Bobkenshaw. LAX. Mlpkr. GoldenTorc. Nixeagle. Brian Kendig. Lingwitt. Someone else. Amorymeltzer. Forluvoft. DMacks. Immunize. Lavateraguy. Rogerd. Nectarflowed. EdwinHJ. KimvdLinde. Joejoeftw. Virginia Dutch. Thedude212. Kevin aylward. WAS 4. Epbr123. Jobin104. Piperh. DMacks. Culverin. Robert Stevens. MeSoDark. Freerow@gmail. Malcolm Farmer. YassineMrabet. Sir Nicholas de Mimsy-Porpington. Courcelles. Doc glasgow. Extremophile. Seb951. Brian0918. 4v4l0n42. Stevertigo. Bobo192. Lexor. Whittrocks. Alasdair. Kubra. Lexor. Closedmouth. Andonic. Bomac. Danthedude. Xyzzyplugh. Dmerrill. Filll. Methcub. Trek-a-Tolk-a-Lewisite. Zsingaya. Capeo. Ceranthor. GLaDOS. SineCurve. Mc4th. Rjwilmsi. 597 anonymous edits Genetic drift Source: http://en. Chris55. CanbekEsen. Poethical. Gclassballer. Az1568. Cohesion. Dreadstar. Watchdogb. Pharaoh of the Wizards. Kalv372. S3000. Woohookitty. Miquonranger03. Abc518. Andre Engels. Salvio giuliano. Ansell. Notjimsteele. Kbh3rd. Ninly. Rahul sig. Reinyday. Rabo3. Angusmclellan. ForestDim. D. RJC. Dureo. Mashford. Jackelfive. Flewis. Lazylaces. Rgoodermote. Tombomp. Palnatoke. Nakon. Aaron Bowen. Wimt. Pip2andahalf. Rrburke.Miller. Joannamasel. Sikatriz. Mx740. Enlil Ninlil. YechezkelZilber. Harland1. Kim Bruning. Jackol. Zurrr. Irixman. Gabbe. Antandrus. Akeldamma. Mike6271. Dmitri Lytov. Habj. Ajoust. John Reaves. Michael Johnson. Mikael Häggström. WikiDisambiguation. Andonic. DerHexer. Roland Deschain. TheIncredibleEdibleOompaLoompa. Magister Mathematicae. Cannaya. Guettarda. Master of Puppets. DJ Clayworth. RoyBoy. EuPhyte. Tanthalas39. JellyWarriorAllele. İnfoCan. Tempodivalse. Dinosaurdarrell. Samsara. Philip Trueman. Sander123. Savant13. 5 albert square. Golden. Headbomb. Agreene175. MC10. JWSchmidt. Slrubenstein. Garik. Kahkooi. Calliopejen1. Wimt. The High Fin Sperm Whale. PhDP. Bwhack. Tom harrison. Graevemoore. Bcasterline. Dawn Bard. Vsmith. L Kensington. I try this account. Calmypal. King of Hearts. Abdullais4u. Sancassania. JosephCCampana. Mentifisto. Pete. Equendil. Zhou Yu. LFaraone. Cp111.xxx. Rieser. Rang3rofskil. Borgx. SchnitzelMannGreek. WAS 4. Skysmith. Balohmann. Royboycrashfan. Jackelfive. Talking image. Peternewell. Jusdafax. Lupo. DanielCD. Susan118. Alansohn. Wiki alf. Ifiber. 12345ryan12345. Hadrians. Piano non troppo. Vanished user. Dweje. Modulatum. Karajade. Edsova. Delldot. Femto. Muntuwandi. AdamXtreme18. Ixfd64.php?oldid=392416091 Contributors: 0zymandias. Mouseomatic. Lawrence Cohen. Plumpurple. Mkpg. Calvin 1998. Can't sleep. Cpu111. Malcolm Farmer. TedE. Mav.. Courcelles. Polinizador. CanbekEsen. Deville. Template namespace initialisation script. WikiHigor. IRP. Duncharris. Yath. Bongwarrior. Standardfact. Rustygin. ErgoSum88. SMC89. Jonathanstray. Nk. Gaius Cornelius. Eisnel. RDBrown. BorisTM. Nikai. Apers0n. Mboverload. Ec5618. Eeekster. Omegatron. Irishguy. Steinsky. Amorymeltzer.wikipedia. Allstarecho. Gawaxay. Svick. TheIncredibleEdibleOompaLoompa. RexNL. Artichoker. DARTH SIDIOUS 2. Circeus. Trichodinamitra. Waggers. Terrace4. Ettrig. Haetmachine. Harry R. Mav. Zephyris. Edwy. CWii. Duncharris. Werdan7. DeadEyeArrow. Egmontaz. Johnuniq.:Ajvol:.org/w/index. Annasweden. Z10x. Turnstep. Mihoshi. Usernodunno. Ronhjones. Conversion script. Dunn8. Closedmouth. Addshore. A8UDI. Javascap. RexNL. Shanel. Hanmi74. Nmerriam. DianaGaleM. Abergabe. Aericanwizard. Gilliam. Keegan. Vipinhari. ElectricRay. Abdullais4u. Bongwarrior. HalfShadow. BiT. Munita Prasad. GD. Arnold90. Copypaste45. AC+79 3888. Michael Hardy. Exponent. 168. Bluerasberry. Archanamiya. Gimme danger. Ian Dunster. CWY2190. Sheitan. Lavateraguy. CharlotteWebb. clown will eat me. Atulsnischal. Gimboid13. Ahoerstemeier. Adenosine. Liontooth. Archanamiya. Crazy Boris with a red beard. WikiDao. JerryFriedman. Bakerccm. Pdorrell. Gimme danger. Violetriga. Omc. Davril2020.org/w/index. Dlohcierekim's sock. Galygal. Capricorn42. Jojhutton. Gniniv. Tio1. DoktorDec. Aunt Entropy. Gurch. Sango123. DeadEyeArrow. Dncarley. Tiddly Tom.php?oldid=391795037 Contributors: 10outof10die.delanoy. Tameeria. Gusgould. HexaChord. Ocaasi. Yamamoto Ichiro.org/w/index. I am not a dog. Jcurious. NuclearWarfare. Clappingsimon. Goethean. TimVickers. Arjun01. PizzaBox.org/w/index. Archie Paulson. Dysmorodrepanis. Venmon. Alex tudor. Mann jess. GetAgrippa. Ahoerstemeier. 2468Aaron. Alansohn. Planettelex23. JSpudeman. Michael Hardy. MCTales. Hannes Röst. Xezbeth. Chrislk02. BiT. Ciar.. Lindenburg. ParticleMan.php?oldid=390815129 Contributors: 0612. Jisily. Hobartimus. Mercenario97. LindsayH. Trusilver.delanoy. Wouterstomp. Faustnh.php?oldid=393098533 Contributors: -Ril-. Lee Daniel Crocker. Uncle Dick. Jebus989. Darwinek. Hogyn Lleol. Tide rolls. Big Bird. WAS 4. Mintleaf. Favonian. Alan Liefting. Quizkajer. Marcosantezana. ArchStanton69. Agathman. 351 anonymous edits Adaptation Source: http://en. JzG. ConfuciusOrnis. Mauler90. Dhollm. Chewie. ClockworkSoul. Macdonald-ross. InfoCan. Arakunem. Bendzh. Muad. On the other side. Wavelength. Defender of torch.wikipedia. Excirial. Gogo Dodo. Kubigula. Sugarfish. LeoNomis. BananaFiend. LazyBoi633. Hatch68. Vicenarian. Eagleal. Enviroboy. Sleigh. Amcguinn. The Claw. Gurch. Amit man. Radagast83. 197 anonymous edits Gene flow Source: http://en. Kyknos. Hut 6. Suisun. Misza13. Tannin. Johnuniq. Discospinster. Cremepuff222. Res2216firestar. Wikibofh. Guettarda.Win. Flowerpotman. Jwing79. Arcadian. Dante mars. GrahamN. Gwernol. GSlicer. Ragesoss. Imarockstarfoo. Hut 8. Sk741. Swastikde. Harryboyles.php?oldid=390717271 Contributors: 10outof10die. Huji. Abdullais4u. Vangoghhasamachette. LeadSongDog. Kpjas. Mc4th. Sceptre. Donarreiskoffer. Teerickson. Carifio24. GSlicer. Alansohn. Madeleine Price Ball. Somnathroy. Bomac. Stardust8212.Hurd. Zandperl. MKS. Bobo192. Pyrrhus16. Leek Francis. Derek Ross. Agathman. Kingjalis3. KnowledgeOfSelf. EWS23. DerekLaw. Tothebarricades. Volfy. Versus22. Cal 1234. FCSundae. Christinebenson58. DrMontik. Lotje. Romanm. 10outof10die. Standonbible. Dysepsion. Batmanand. J.. Plumbago. FF2010. Sewings. Hede2000. Timo Honkasalo. Gimme danger. Engelmann15. Martial75. CanadianLinuxUser. John Vandenberg. Mccready.tk. Vespristiano. Novangelis. Danimoth. Ettrig. Ahoerstemeier. Shirik. Fireyair. WikiDao. Vanished user. Rich Farmbrough. Leonard^Bloom. Jaxl. KillerChihuahua. Maedin. Cooner7777777. Khoikhoi. Borgx. Zargulon. Foant. Frank Lofaro Jr. Z10x. 15mypic. DVD R W. Specs112. Gimme danger. Kuru. Chain funds. Thue. Jheald. Soccerdu82. Nabarry. . Noclevername. Dmanning. Jebba. Tide rolls. Genenome. Dcooper. Lexor. Ychennay.250. Pakaran. Credema. Tgr. Synapomorphy. Dycedarg. The Thing That Should Not Be. Kazkaskazkasako. Derek Ross. Azcolvin429. Bongwarrior. Wmahan. 10outof10die. Bill37212. FCYTravis. MigueldelosSantos. Bayerischermann. Join Tile. Extremophile. Megan1967. C. Tonicthebrown. Chopchopwhitey. Lightmouse. Fox. Vsmith. K. HIGA. Ceyockey. Kdawson66. Jungenbergs. The Thing That Should Not Be. Yuanchosaan. Bigger digger. Garcsera82. John Lynch. Zundark. Atulsnischal. Addaick. Fl. Aitias. Jusdafax. Antiuser. Millahnna. Gunnar Mikalsen Kvifte. KnightofZion. Courcelles. Lradrama. Roke. Curehd.. Shaocc.250. Shogun. Richard001. Leptictidium. Guettarda. Brianga. Amcbride.. Lexor. GoodScienceForYou. Guettarda. Pengo. Steinsky. Christomaniac76. DoctorDNA. Katalaveno. EPM. Solenoozerec. Wtmitchell. Fastfission. Jeff G. LifeScience.. HiDrNick. Gaius Cornelius. Risk one. Guyzero. Netnuevo.250. FT2. Aunt Entropy. Mortartion. TimVickers. Eleassar. Bensaccount. Dysmorodrepanis. Dardasavta. VK35. Letatcestmoi94. Pigman. Emw. Ocon. Livajo. Childhoodsend. Chazza 94. ESkog. Peachiezworld. Avillia. Atlas1. Wilhelm meis. Knowledge Seeker. Triona. DerHexer. ZeroJanvier. Sdarwin.5. Calvinloa. PAR. Gail. Johnpseudo. Leafyplant. Giordanobruno. Ian Dunster. Andrea105. Faradayplank. Mindmatrix. Tycho. Joannamasel. Alansohn. Unint. Phenz. Mxn. 448 . Zzyzx11. Technopilgrim. Ideogram. Cgingold. Jessisamaknows. Thomas Arelatensis. Bomac. Indosauros. FF2010. Tupeliano. Andres. Discospinster. Seb951. Mentifisto. Smith. 9mw. Ehrenkater. Maedin. Carlroller. Numbo3. Keilana. Brentt. Dwheeler. Aitias. Elliskev. Jiddisch. Doc Tropics. J. Pengo. Octopus-Hands. Jcorry10. Narayanese. Rjwilmsi. GeneralAtrocity. Azcolvin429. Stephenpratt. DGG. Tommy2010. Limbo socrates. Allangmiller. Jason7825. Jrockley. Jeff G. CSWarren. J04n. Egmontaz. 168. Standinguptoit. Ency456. Code Burner. Lothar von Richthofen. Townmouse. UncleBubba. Buickid. Bheest. James086. ZxqamF. Hydrogen Iodide. Cossey mo. Shirulashem. MIT Trekkie. Ppgardne. Pigman. Flashinfinity. Cohesion. PhDP. RainbowOfLight. CYD. Klgroom. Dlohcierekim. JohnOwens. Cyde. Micha L. Hornlitz. Hensa. Mike2vil. Oliver Lineham. Joannamasel. GirasoleDE. Etxrge. Finalius. Gcrossan. Drew R. Larrybarry. CreationScientist. Gog. CardinalDan. Dockingman. Daycd. Eptalon. Mandarax. Nunh-huh. 838 anonymous edits Natural selection Source: http://en. Template namespace initialisation script. JHunterJ. Samsara. Lexor. CanbekEsen. Rat at WikiFur. RedWolf. Epbr123.5. Tuganax. Alchemes. Viridae. Chris 73. Daleep123. RexNL. Dr d12. Ganny. David D. Corpx. Johnuniq. Dantecubed. Gracenotes. Razorflame. Crazynas. Nezzadar. Ed Poor. Myanw. Donama. Lowellian. Thebestlaidplans. Leuko. Da monster under your bed. Discospinster. Toastybread. ChrisTheBrown. Semperf. Meaghan. Sharonlees. Natalie Erin. Ncfuzzy101. Daggermouth. Mjb. Lab-oratory. Seven of Nine. Nut-meg. Firebat08. Farquaadhnchmn. FireBrandon. Donarreiskoffer. Oxymoron83. Natalie Erin. Skysmith. Kingdon. Graminophile. VirtualDelight. Joseph Solis in Australia. Amaltheus.wikipedia. Titoxd. JWSchmidt. ElfQrin. Kralizec!. WereSpielChequers. J. Owenman. Arnold90. Capitaljay. Zhou Yu. Wimt. Pexego. SEJohnston. Strobilomyces. Sobreira. Zro. Casito. Walkingwithyourwhiskey. Espoo. Roland Deschain. Raul654. Chanting Fox. Vsmith. Knownot. HJJHolm.. Poo HEAD JESS. Oblivious.. Darth Ag. Kosigrim. Dj Capricorn. Sitearm. Wotnow. Tellyaddict. Dave souza. Bcasterline.191. Alan Liefting. Lauranrg.wikipedia. Dinoguy2. Furrykef. Aranae. Bryant. Anlace. DianaGaleM. ABF. WAS 4. Banno. Benpayne2007. Moosepuggle. Sjö. Nsaa. Onevalefan. Mr. C. Template namespace initialisation script. Ragesoss. Chris Roy. Auno3. Tualha. Bruno Dantas. HalfShadow. Ed Poor. MER-C. Gabbe. Pharaoh of the Wizards. Alarichall. GoEThe. Orangemarlin.php?oldid=391298611 Contributors: 10outof10die. NickCT. Slysplace. Pinethicket. Vishnava. Radon210. Can't sleep. BlytheG. RexNL. Guettarda. SpankDatAzz. Sadalmelik. Jfraatz. XCalPab.g. CinchBug. Dyanega. 172. Fastily. Graft. Müslimix. A8UDI. Jan Pospíšil. Brazucs. Kku. Wobble. Ettrig. Time9. Ecorahul. Springbok26. Anthere. Bcasterline. No Guru. Poethical. Bigtimepeace. MarcoTolo. Egern. WadeSimMiser. TheAlphaWolf. Raymondwinn. Pete. Neutrality. Sinbad68101. Pekaje.mace. Eugene van der Pijll. Rossami.. Valich. Woudloper. Diagonalfish. Psarj. Dysepsion. Autonova. EdJohnston. Wickey-nl. Leet3lite. Recordinghistory. Template namespace initialisation script. Ulluxo. Robert Stevens. Bikeable. Mendaliv. Mav. Fama Clamosa. Spiritia. Dkabban-GMU. Albedo. Uncle Dick. Bmeirose. Wayward. Omegatron. Jaknouse. Arfn24. Lindosland. Nomoskedasticity.org/w/index. Olivier. Shyamal. PrestonH. Sfmammamia. Michael Johnson. Buckley. Themfromspace. CDN99.wikipedia. C. Plumpurple. Fratrep. Plantsurfer. True Pagan Warrior. Dbabbitt. Chewie. Atulsnischal. BenB4. Razorflame. ESkog. Mike Dillon. Mindmatrix. Nurg. RoyBoy. MrDolomite. OLH06. Palica. Johnuniq. 04carrolld. 168. 2004-12-29T22:45Z. Sacquebout. Snowmanradio. CharlotteWebb. Lapaz. Beland. Canadian-Bacon.org/w/index. Mario1952. Fui in terra aliena. Jasmaunder.xxx. Alan Liefting. Ryulong. ConfuciusOrnis. Brya. 130. Enzymes-GMU. Aviv007. Mike Dillon. Florentino floro. Boreal99. Jeffpw. Wmahan. CHW100. DavidLevinson. BadDoggie. Nicke L. Adam Retchless. 2over0. Just Another Dan. Rich Farmbrough. Gurch. Nmg20.153. SilkTork. Dexterhaven.Fred. TotoBaggins. Junyor. Zfr. Arctic. Ajsh. Tremello22. Cladist. Afasmit. Sietse Snel. Marooned Morlock. Boredzo. Ecobion. Andres. Hu12. Ugen64. APH. Shyamal. BirdValiant. Fatapatate. Sikatriz. 4444hhhh. Buffison. Anburnett. Visionholder. Bubbha. Kosigrim. Slrubenstein. Sedulus.gaze. Debresser. Animum. Victor falk. Île flottante. RhiannonAmelie. Samashton. Extra999. Slon02. Carlaude. GoEThe.org/w/index. Spencer. MichaelBillington. CKCortez. Ahimsa xrqhd. Antipastor. Rmhermen. Noleander. Aymatth2. Andersmusician.wikipedia. Scbomber. Neapoli. Courcelles. Blechnic. Geoff. Tktktk. Silly rabbit. Antoni Barau.wikipedia. Nishkid64. Sandman. Quizkajer. Vsmith. Kevin Saff. LoganFrost. BG. Opabinia regalis. Gaurav. Rillian. Mr. RK. Berton. Conversion script. Aeon1006. Marshman. Inferno. AnonMoos. Castjean.. Jerainseltran. Mikespedia. The sock that should not be. Vercillo. Samuell. Brian0918. PierreAbbat. Tito-. Peak. Bobo192. Shanes. AlienHook. Aua. Lexor.. KimvdLinde. Emaraite. Mark t young. EncycloPetey. Nlu.. BlaiseFEgan. Neutrality. Bornhj. Noodleman. Benthehutt. Woland37. Narayanese. Airplaneman. NatureA16. Valich. Entnahme. BD2412. Triwbe. Mario1952. Sundar. Memestream. Recognizance. Standonbible. Al-Zaidi. Paalexan.Article Sources and Contributors Moe Epsilon. Brya. Ragesoss. Vsmith. Olianov..hoyland. RoyBoy. Breandandalton. Dmerrill. Ke4roh. Ospalh. Iridescent. Redian. Bdb484. Gdvorsky. Narayanese. Shazalusose. Tarquin. Persian Poet Gal. Ecphora. TorreFernando. LOL. TimVickers. Arakunem. Babylonian Armor. BerndH. Romanm. Jamiejoseph. Bassbonerocks. Gaius Cornelius. Spamsara. Will Beback Auto. TutterMouse. Agathman. Sid mishra94. Alex543211. BobKawanaka. ChrisCork. WillJeck. Johnuniq. Hrmaham. Whatiguana. Syp. Wordbuilder. The Anome. Kiwiboy1221. 200. Robert Stevens. Quietmarc. Okedem. Kencf0618.. Gilliam. Sentausa.wikipedia. Cmporter. Shao. Natisto. Aunt Entropy. Seqsea. Julian Mendez. Rigadoun. Dr. 332 anonymous edits Phylogenetics Source: http://en. Captinhippie. Petri Krohn. Attilios. Thue.Ent. NickelShoe. M stone. Fbartolom. Vaughan Pratt. VeryVerily. Trevor MacInnis. Themfromspace. Roland Deschain. Slrubenstein.Fred. PseudoSudo. Mgiganteus1. Basher018. Antelan. AxelBoldt. Aunt Entropy. Consist. Richard001. Samsara. Wafulz. PDH. Bluefrog67. Pjvpjv. Boston6788. Levineps. Patrick0Moran. Gadfium. Mindmatrix. LilHelpa. Ballista. JackH. Jrockley. Carlosp420. Popnose. WillowW. PhJ. Bruno Dantas. MrDolomite. Steven Argue. Brandon. Petter Bøckman. Calaschysm. Pen-pen. Sheep81. Uncle Milty. Ryulong. Alexius08. Wzhao553. JerrySteal. LossIsNotMore.bit. Unyoyega. Susurrus. Wmgetz. Thatguyflint. Snalwibma. Sirkad. Earrnz. West. Demi. Bookandcoffee. Alteripse.org/w/index. Thorwald. J. Bueller 007. Nakon. Fastfission. Pschemp. Hurmata. AshLin. Lexor.. Verson. Naddy. Vsmith. I'm 449 .96. Ahoerstemeier. Carlosp420. Onco p53. Gdr.j. 100110100. Conversion script. Kembangraps. Rich Farmbrough. Rdsmith4. UtherSRG. Fedor. Stephenchou0722. Tslocum. Drphilharmonic. Template namespace initialisation script.org/w/index. PhDP. User6985. Camw. Conversion script. David Gerard. Extremophile. Marknau. Juliancolton. Josiahtimy. Beland. I dream of horses. 1180 anonymous edits Speciation Source: http://en. Rintrah. Styrofoam1994. Halidecyphon. Veryhuman. Hamamelis. Igodard. Bosmon. Seglea. Wikipeditor. AndriyK. Yerpo. Armchair info guy. Ilmari Karonen. Bomac. Template namespace initialisation script. Erianna. Seglea.php?oldid=393080483 Contributors: !1029qpwoalskzmxn. SgtDrini. Gioto. Snigbrook. Insanity Incarnate. Metamagician3000. TimVickers. Sebastiano venturi. Welsh. Slrubenstein. Rossnixon. Robert1947. TEPutnam. Addshore. Brianlucas. ThisIsAce. Dr d12. Tyw7. Rjwilmsi. Antandrus. Snek01. Ggriffit1. The Anome.muller. Michael Hardy. GraphicArtist1. Oxymoron83. Discospinster. Omnipedian. Richard001. Polly. DéRahier. Dj Capricorn. Cromwellt. 220 anonymous edits Cladogram Source: http://en. Duncharris. PDH. Paul Richter. Calair. MAH!. Colonies Chris. Pinethicket. Juan448. Snowmanradio. NawlinWiki. Yousalame23. Mercury543210. SpuriousQ.. Ernestfax. Tameeria. Etxrge. Dave souza. Tristanf12. NCC-8765. CardinalDan. Mdhowe. Professor marginalia. JohnCD. Wikichickenlol. Alastair Haines. 8 anonymous edits Molecular phylogenetics Source: http://en. Hadrianheugh. TongueSpeaker. Patstuart. Debresser. Gunnar Mikalsen Kvifte. Johnuniq. Jknabe. Calypso. Netsnipe.wikipedia. Tom Schmal. Wheely Guy. Townmouse. Opabinia regalis. Knnwill. Touchstone42. ScienceApologist. Evolve17. Cfortunato. Mygerardromance. Neptune5000. Gruzd. 168. R. M. Moshe Constantine Hassan Al-Silverburg. CanisRufus. Tuxedo junction. Tobias Bergemann. PeterMSimons. O. Waldow. Shanel. Bryan Derksen. Radiocar. Samsara.org/w/index. Bluecurtis. Ragesoss. Ksbrown. BananaFiend. Polinizador. William R. Ombudsman. Tony1. Vanished User 0001. Takometer. Cybercobra. TheProject. Bobby D. Tony Sidaway. The Thing That Should Not Be. Grouchy Chris. Lexor. Gongoozler123. Jacek FH. OnBeyondZebrax.. Chase me ladies. Bender235. Leptictidium. Rjwilmsi. Sluzzelin. TheCatalyst31. Rich Farmbrough. D I Williamson. BSquared04. Consist. Sluzzelin. MrDarwin. SandyGeorgia. SidP. Dmitri Lytov. Imagine-GMU. Agathman. Dysmorodrepanis. TUF-KAT. Radagast83. Lexor. Mat8989. Hkim43. RUL3R. Sander Säde. Schnolle. PAvdK. MrSomeone. Skydog100. Sam Clark. Tide rolls. Dysmorodrepanis.Eff. Restre419. Katefan0. Corpx. The Thing That Should Not Be. Blue98. Trabucogold. Ultramarine. Amorymeltzer. Touchstone42. ConcernedVancouverite. ZjarriRrethues. Peter coxhead. Utcursch. Samw. Gcm. Anthon. Gauravm1312. Stirling Newberry. Lord of Penguins.122. 999powell. JoshuaZ. Susan118. Armchair info guy. EdJohnston. TedE. Pallab1234. Grendelkhan.250. RedAlphaRock. Syncategoremata. Cthompson. EdJohnston. Ummhihello. Mariachi2324. GetAgrippa. Avb. Beland. Diannaa. Vanished user. Fastily. ForestDim. Nrcprm2026. Josh Grosse. Stephen C. From That Show!. SilkTork. Tofof. The Thing That Should Not Be. Dragon Helm. Frankenpuppy. Regancy42. Pasquale. Matthias. Oking83. Snoyes. BlackPh0enix. Duncharris. Bookuser. Swpb. Faustnh. Naddy.xxx..Bastedo. JoJan. Striker2712. PDH. Alex. Clicketyclack. Marj Tiefert. Littlealien182. BD2412. Shoemaker's Holiday. Belizefan. Nielses. KiloByte. RandomStringOfCharacters. Qxz. Scilit. Zvika. Xaosflux. Mav. NoahElhardt. Ulric1313. Paul Hjul. M1ss1ontomars2k4. Wintran. AdamRetchless. Julesd. Littenberg. Northfox. AxelBoldt. Royalguard11. DianaGaleM. SqueakBox. SimonGreenhill. Treisijs. Mdsam2. Seglea. Lightmouse. Berton. Rmarquezp. Martinwguy. Toddst1. Gogo Dodo. Ppe42. Jojhutton. AdultSwim. BaNkR17. Johnuniq. JRR Trollkien. Pawyilee. Borgx. Pigsonthewing. Dpr. Mzsabusayeed. RickK. Naj-GMU. Woohookitty. ParlorGames. Damian Yerrick. JForget. Genypholy. Huson. Wykis. Sandstein. Dysmorodrepanis. Fredrik. Bobblewik. Ian mccarthy. LilyCylinder. Willking1979. Template namespace initialisation script. Andycjp. Karmosin. Johnuniq. Ak1255. Heron. SimonGreenhill. Themusicgod1. Who then was a gentleman?. Andre Engels. WikiDao. Richard001. Bussech. CardinalDan. Noleander. Anabus. JerrySteal.eyes. Shanes. Nixeagle. Thesoxlost. Safay. Dcljr. Greeneto. Rasmuss. Vanished User 1004. Hughcharlesparker. Tide rolls. Vanished User 0001. Vanished user. Theuser. Afronathan. Borgx. Ann arbor street. Wgmccallum. Zbvhs. Darth Panda. Sadi Carnot. RJO. Katalaveno. Tjunier. Twaz. Kosigrim. Nareek. Craigy144. Good Olfactory. Quizkajer. Ph. Pizza1512. MB83. Christian Kreibich. Stemonitis. Visionholder. Dante Alighieri. Stemonitis. Sabik. Bigturtle. Mav. Magnus Manske. Carlson. Benbest. NielsHietberg. Carlosp420. Bomac. Lexor. Philcha. Jmeppley. Michael93555. Pinkgirl941222. Unyoyega. Imasleepviking. Blackout. Biol433. Carolmooredc. Abdullais4u. Zsinj. Aunt Entropy. GeoMor. Beyazid. Shenme. Timwi. Samsara. Eroston. Scndlbrkrttmc. Slack---line. Sidhekin. John of Reading. Tea Tzu. Nudve. Thomas Arelatensis. MrOllie. Brentt.andrew. Debivort. Tmol42. Eyallow. RGTraynor. Gimme danger. Nakon. Philip Trueman. Slowking Man. Headbomb. Shyamal. Bwsmith1. Vary. Fanghong. ArglebargleIV. Hadal. Neelix. Alansohn. APH. Opie. Bryan Derksen.gnome. Alpha Quadrant (alt). Branko. Samsara. Steinsky. Arkuat. ABF.wikipedia.org/w/index. Taffboyz. Azcolvin429. Lexor. Mcw1139. Nonsuch. Mokele. Whatiguana. Calliopejen1. Chandrax. Pharaoh of the Wizards. BenFrantzDale. Bobo192. Antiuser. StephenWeber. Lee Daniel Crocker. Yuekling. Sjschen. Nikola Smolenski. Outriggr. MichaelSH. TestPilot. Chameleon. Nurg. 59 anonymous edits Molecular evolution Source: http://en. Stfg. That Guy.xxx. Kevin Forsyth. SJP. Sholto Maud. Dayewalker. Arnon Chaffin. RexNL. Aidan Elliott-McCrea. Purnajitphukon. Bfinn. Agathman. Aranae. Stevegiacomelli.php?oldid=392458047 Contributors: -Ril-. Nakon. BryanEkers. Invertzoo. clown will eat me. ZeroJanvier. Benhocking. Chakazul. Andycjp. B jonas. Wdanwatts. Scibah (new account). Deltabeignet. Culmination. Circeus. Karada. K. Samir. DMG413. Akriasas. WikiRat1. Yamamoto Ichiro. Woohookitty. Sjö. UtherSRG. Predheini. Tide rolls. Buddho. Ewen. 54 anonymous edits Human evolution Source: http://en. Mensfortis. Aldenrw. IanCheesman. Athf1234. Snek01. Inquam. Koavf. SomeStranger.188.Hurd. Squids and Chips. Santa Sangre. 140 anonymous edits Cladistics Source: http://en. Aranae. MrStalker. Cyde. RedHillian. Modify. HorsePunchKid. Manitobamountie. Miaers. BiologyForksWA. TimVickers. Ppe42. Arny. Harry R. Surachit. BlastOButter42. Snailwalker. Theresa knott. NerdyScienceDude. JoJan. Ian Pitchford. The Braz. Michael Hardy. Quailman. RJaguar3.php?oldid=377909149 Contributors: 168. Shyamal. Lakmiseiru. Atif. Smmurphy. Sergeantbreinholt. LeaveSleaves. Badams5115. Bhawthorne.php?oldid=387836487 Contributors: ArkinAardvark. A robustus. AshLin. Zvika. F. Steinsky. LittleHow. 168. GerardM. Toytown Mafia. Bonadea.org/w/index. Atl braves. Drphilharmonic. Betacommand. Amaltheus. Rjwilmsi. Peter morrell. Wickey-nl. Duncan. Adrian. Eugene van der Pijll. Rjwilmsi. Samsara. Wilhelm meis. Nate1481. Smith609. Anthonyjameswood. Nbarth.php?oldid=392634694 Contributors: 10outof10die. Rjanag. Staffwaterboy. BrettAllen. StormBlade. GSlicer. Wykymania. Consists. Drdaveng. Anaxial. Joriki. clown will eat me. Timwi. Cohesion. Ainlina. EJF. Richard Arthur Norton (1958. Gabbe. Themfromspace. Ligulem. Separa. JerryFriedman. Yamamoto Ichiro. WinContro. Ballinchad15. Beland. Notreallydavid. Vanished User 0001. Bobo192. Tmkeesey. Verbist. Ish ishwar. Juliokhat. Huji. Tommy Kronkvist. KillerChihuahua. Chamaeleon. BarretBonden. Swithrow2546. OGoncho. Neutrality. PierreAbbat. BrianGV. Thankyousir8. Thingg. Can't sleep. Nandreeson. SteinbDJ. Caltas. RoRo. Buritdd. Niteowlneils. Wavesmikey.delanoy. A. Gökhan. EEye. Quiddity. Herbm. Zundark. GreatWhiteNortherner. 27 anonymous edits Evolutionary developmental biology Source: http://en. Aranae. TomGreen. Noctibus. RG2. Biophys. Dawn Bard. Raven in Orbit. Bendzh. Sean.t2. Roosme. Stevertigo. Redsunrising15. Zephae. Ameliorate!. Pepper. JonHarder. Euyyn. CzarNick. Alamowit. Sango123. StN. Slakr. Pathoschild. Belthil. MarkGallagher. AubreyEllenShomo. YK Times.php?oldid=390075235 Contributors: AS. Thehelpfulone. Bongwarrior. 9eyedeel. Rickproser. Moreschi. Yue44003. Syp. Who. Alan Peakall. The Merciful. Dysmorodrepanis. Eratosignis. Blue Tie. Azzors. Adoniscik. N4nojohn. Samsara. Seb951. UncleBubba. Stefan Kruithof. Cephal-odd. Ravenswood Media. Justanotherdomino. Rusty Cashman. Captain-tucker. Smith609. Agathman. Atarr.). Rjwilmsi. CalumH93. Betterusername. Boing! said Zebedee. Frostus. JackWasey. Acaeton. LittleHow. Michael Hardy. Shyamal.php?oldid=392407939 Contributors: 10outof10die. Ioverka. Figma. Bz75. Ancheta Wis.94. Infomuse. Slrubenstein. Samsara. Cholling. Pschemp. Kazvorpal. Emw. Ormy. BMF81. Grimey109. Nethgirb. Diego. DHBoggs. Rjbonacolta. ParisianBlade. Fubar Obfusco. Ljhliesl. Koavf. Roke. Blingbling1000. Yuzhong. Deus Ex. MAXimum Xtreme. Der Falke. Don231194. Drc79. Universal1300. Karn. Narayanese. GRider. Ashmoo. Chrislk02. Kuru. MK8. R Lowry. Rror. CuteWombat. Armchair info guy. Juliancolton. Aquishix. Kooblikon. Azcolvin429. Saul Greenberg. Philip Trueman. JoshuaZ. Chanting Fox. Snoyes.Smoothie. Ajedi32. Ctbolt. PerryTachett. Rramir16. Efnar. FreplySpang. Robofish. Danielrcote. Andonic. Klingoncowboy4. Joegoodfriend. Exxxs. Denny. Quantumor. Kazvorpal. Funhistory. The Storm Surfer. Grigri. Livajo. Brothejr. 474 . Picaroon. Wiki4ata. Maxis ftw. Eisnel. Norris. Light in Water. RexNL. Koavf. Quintote. Arkuat. Frowanda. Weredozen. CardinalDan. King of the Dancehall. TheNewPhobia. Mjk2357. Tempodivalse. Ishootudie. CDN99.:. Levil.5. Zoz. Lynn Wilbur. Ed Poor. Josue arnold. Fationia. Kevs.wikipedia. Kaylabelle. Bjoh249. Ec5618. Temtem. Aalit4. Clicketyclack. FreplySpang. Hu. Moumine. Kevmus. JH-man. KateH. Twas Now. Davril2020. Joyous!. Johnkarp. Hairhorn. Keganizcool. Sylandi. Lauriec. Ed Hagen. Beaven. Knowledge Seeker. Skysmith. Grayt2. Dmyersturnbull. Bryan Derksen. Mzajac. Feinbergdavid. Manscher. Sonjaaa. Thingg. Gt2003. Nivix. JFreeman. Darklilac. Edward Z. JoanneB. Hawke666. JForget. JNW. Discospinster. Billion. FlyingOrca. Possum. Dimionix. Master Scott Hall. Josuechan. BBrihem. NawlinWiki. Ming the Merciless. Gaius Cornelius. Filll. Mindmatrix. Davril2020. Dhochron. Jason Potter. Andonic. Headbomb. DopefishJustin. Dieall. Cptmurdok. Smoove Z. DRTllbrg. TriNotch. OwenX. Robert Stevens. JonHarder. JohnWittle. Richard001. The Rambling Man. Dogposter. John Vandenberg. Mike Christie. Engleaugen666. Duae Quartunciae. Gary D. Wap. Svidrillion. Alan Peakall. La Pianista. Danoconnell. Crusio. Geologyguy. Darth Panda. Rob117. Wangi. TestPilot. It Is Me Here. BOARshevik. Emptymountains. Davidzuccaro. Mikker. Leafyplant. Raeky. Suffusion of Yellow. Lord Pistachio. Discospinster. KimvdLinde. Fastfission. Kpjas. Hello32020. FishoFish. RedHouse18. Humanevol. Schrandit. Leinad-Z. The Anome. Gimmetrow. Mikker. Kuimov. Iss246. Ems57fcva. Sean D Martin. Slrubenstein. Ndteegarden. Yath. Nowimnthing. Aschwa5. Hrafn. Goethean. Hypershock. Kariteh. BigGuyandaGuitar. Laurascudder. Park3r. Harriv. Daweirdo911. Mlaffs. User2004. Will Beback. Mordgier. Ucucha. McSly. Luokehao. Romanm. Slowking Man. Dysepsion. Lincspoacher. Gaia1CB3. Cbruno. JavaTenor. DopefishJustin. SidP.250. Hurmata. Cookiehead. Mjsandiego2003. Hut 8. Tony1. Waggers. Mudkipperz. Diego Vara. The ed17. Jtneill. Defyn. GusChiggins21. Shalom Yechiel. Proma96. MBisanz. Jackollie.Lutherson. C56C. QuizzicalBee. Nightscream. Fvw. IVAN3MAN. Wiooiw. Gilliam. Viriditas. Tomandlu. Calvin 1998. Editor2020. Ashlux. Hans van Deukeren. Senecasd. DoktorDec. Tobby72. Landroo. Midgley. The PIPE. Crust. DDek. Jiffles1. RainbowOfLight. Dijxtra. DerHexer. Nectarflowed. Jlujan69. Superweapons. Waywethin. Leadwind. CommonsDelinker. The stuart. Kestrel. Jefffire. AlexBadea. Jaguar. Stimpster. Rhofro. LeaveSleaves. Fang 23. TheThomas. Little Mountain 5. Snuffkin. Vanished user. George100. Scwlong. Ftjrwrites. Lord Patrick. Mietchen. SimonP. Dave souza. Jtle515. Everyking. Restepc. ImprobabilityDrive. Ballista. Jhanley. Enochlau. Wiml. Hmrox. Arthur B. Gnaussor. Falcon8765. Lemmey. Jossi. Chrissmith22. Hans Adler. Quadell.. Bmcirillo. Sander Säde. Jeeny. Wine Guy. Methcub. Lifeartist. Tresiden. Gladysvale. Cheryl dennison. Brainbuster. Opelio. Filemon. DVD R W. J. N5iln. Chris55. Johan Elisson. ColinJF. Lala357. Fred Hsu. Signa727. Brienno. Froes. Suki77. Silly rabbit. Pasquale. Pproctor. Lando Calrissian. Xook1kai Choa6aur. Edivorce. CLC Editorial. GSlicer. David85. Zaxas.W. RedWolf. BTfromLA. Ohnoitsjamie. Geoff archibald. Mr. Sir Nicholas de Mimsy-Porpington. DARTH SIDIOUS 2. Vina. Rhynchosaur. Magioladitis. AmanUwellCant. S h i v a (Visnu). Mkilly. Mattisse. Fruit. Ruedi33a. GregorB. Pigman. Tiptoety. Martin Breheny. ChrisCork. Jemaura. FilmFan69. DJ Clayworth. Izehar. Alephx. Euchiasmus. FreeThinker09. Charles Matthews. Pioneer42. Tomwsulcer. Lee Daniel Crocker. Moeron. Gary King. Nareek. Ihcoyc. Dougweller. Dr. Ghelae. RyanHoliday. Noah Salzman. Igiffin. SDJ. Suntag. Crohnie. Tubby. Cmdrjameson. Phirosiberia. James pic. Delldot. Florian Blaschke. Eu. Hollyvic. IlSoge. Pinethicket. Allstar86. Frédérick Lacasse. Figma. Gwernol. Jamesontai. Timwi. Inforazer. Millosh. Johanmathiesen. ScienceApologist. Corington. J.php?oldid=392083460 Contributors: -Ril-. Hans-Friedrich Tamke. Torrubirubi. Randi-Gordan-did-it. Guettarda. Shoaler. Jebus989. Gabbe. Raven in Orbit. Aceofelves. Joy.welsh. Fermion.O. IsaacGS. VladimirGolovin. Mongreilf. Picus viridis. JFreeman. Verwoerd. R Lowry. Duncan. Uncle Dick. Melaen. Stegosaur. Gregzeng. JohnAlbertRigali. Johnuniq. ChadThomson. Taw. Bahhasg. Ludovika26. Michael Johnson. DFS454. DanMS. Knowledge Seeker. Rosiestep. Jpgordon. Psykhosis.malshe. Numerousfalx. Mikeeg555. Sardanaphalus. AlistairMcMillan. Elven Dragon Rider. Cold Sandwhich. RJHall. Archivar7. HrvojeSimic. Tjnelso1. GSlicer.dreams. Alansohn. Malo. Peak. Spencer. Kyknos. Peyre. Gimme danger. InShaneee. Bongwarrior. Jeff3000. Wavelength. Habj. Enormousdude. Sam Hocevar. Brisvegas. Mboverload. Margareta. LionKimbro. Gensanders. Rjwilmsi. Rjwilmsi. Steinbach. Sietse Snel. Headbomb. Altenmann. Ilse@. Kris the Kangaroo. Netesq. Maldek. M3taphysical. Loquacious conundrum.france. Kukini. Spliffy. The. Brian0918. Bkkbrad. Stevenmitchell. Ben davison. Treisijs. Malcolm. Weebs. Epsilon60198. FuelWagon. A vivid dreamer. Glloq. Gregchapman. Elassint. Fasten. Graham87.tk. Fredbauder. Lottamiata. Mange01. Arru. FeloniousMonk. Krovisser. Chris 73. Plushpuffin. SpuriousQ. Sturgeonz1. John D. Camo99. F. Greeneto. Florkle. Vsmith. Lord Patrick. Aunt Entropy. MBisanz. Fryed-peach. Laug. Yang. Betterworld. Meaghan. LuisFagundes. Gdr.com. Andrewlp1991. Findaknow. Yath. Drbreznjev. SwiftlyTilt. BryanSWiley. Fences and windows.delanoy. The Thing That Should Not Be. Hmwith. Wtmitchell. Pilif12p. Daycd. Lexor. Mindwarz. FernoKlump. Paul Gard. Amire80. Jim62sch. Skarebo. Janthony. Hmains. La Fuente. Irmgard. Gnixon. Mr 450 . Cedders. Hecd2. Ka Faraq Gatri. Zelmerszoetrop. Kkrueger. Husond. Lightmouse. EChronicle. Kuzaar. Iffykid. Insomniac3. Graft. Cmyers. Favonian. DreamGuy. Maelnuneb. Sander123. Bkell. Fbc215. Kbdank71. Fenice. Joriki.Cardenas. Waltpohl. AllanLee. BarryCull. Iridescent. Colin012. Stefan. GreatWhiteNortherner. IMWeazel.. Abhishikt. Sebesta. RingtailedFox. Postdlf. GordonUS. Hobartimus. Karol Langner. Cheesemancheeseman. Trevor MacInnis. JCarriker. Adoniscik. Intelligentsium. Kurykh. Wikipe-tan. Slakr. SpookyMulder. Edward. Clicketyclack. Hpoliko1221. Kurykh. Hut 8. ABlake. Hvn0413. Jason Potter. Duncharris. Ed Poor.delanoy. VonRichthofen. Tyc20. Shadowjams. Mcferran. Pairadox. LittleHow. H2g2bob. Gbkorol. Ja 62. Grutter. Qatter. Lemonflash. Mexeno1. Joe Decker. F13nd. Jose Icaza. Esanchez7587. WLU. Onorem. CanisRufus. Gabbe. Akuchling. Ladlergo. Cj67. Avb. BBiiis08. Rileyhegan. Spotty11222. Dissembly. Formeruser-81. Limulus. Icseaturtles.Article Sources and Contributors the Cavalry. Emptymountains. GoodCommonsense. BrokenSegue. Kotiwalo.ca. J. Duncan. ForestDim. ElijahOmega. EPM. Jamesontai. Igiffin. Mac Davis. BlankVerse. Katefan0. Corrigann. Huysman. Wmahan. Salvio giuliano. Frogbounce. Reedy. Jkorz. Epf. Vividpresentwakefulness. Delmonte. The Letter J. Epbr123. TheaterMarine. Jdhunt. Lightmouse. Oblivious. Ligulem. RK. ScotchMB. HalJor. Topbanana. Tectonicura. Daniel Case. Elemesh. Kuru. Jpgordon. Ckatz. Ibis3. Wiki alf. Geoff. AlphaEta. Alan Liefting. Andy Marchbanks. Semmler. Gaia Octavia Agrippa. Snek01. Hisaac. Illspirit. Icairns. Memills. Duncharris. Matt Crypto. MindstormsKid. Lingo pen. Vctull. Hrafn. Gilliam. Warbola. Yonghokim. Jmcc150. Wingnut99. FloNight. Ekimiheart. Claudelepoisson.stefan. Clayheart. Balabiot. Eiler7. MistyMorn. Pixelface. Emijrp. Joed6829. El C. Sommers. Shambalala. Joseph Solis in Australia. Tarad09. Drur93. Karl Dickman. Kingpin13. Visionholder. Fred Hsu. Nunocordeiro. KYPark. Dbachmann. Baegis. Shawnhath. Revoprod. Washburnmav. Joscquin. Hairy Dude. Eterminator. Whatiguana. Pstanton. Rich Farmbrough. Saforrest. PrestonH. Leon. TimVickers. DarkKunai. RexNL. Lorenzo Braschi. Ombudsman. Matau. Zackp. Keitei. Mixie2me. Jidliqz. Petter Bøckman. Dpr.لیقع فشاکanonymous edits Creation–evolution controversy Source: http://en. Smalljim. Olegwiki. Yoga sawant. Phenylalanine. Vaughan. IDD55. Cheesegoduk. UtherSRG. David Pro. Piano non troppo. JorisvS. Kendroche.france. Orangemarlin. Clasqm. Fui in terra aliena. Stars4change. GreyCat. SunCreator. Chaca chik. Bart133.php?oldid=389004714 Contributors: 2004-12-29T22:45Z. Garik. Snigbrook. Das Murphs. Grick. AlexCatlin. Luk. Gail.. Dresdenia. DCDuring. Ronabop. Chet Ubetcha. Read-write-services. Ucgaweh. Tommy2010. DocWatson42. PedEye1. L Trezise. KimDabelsteinPetersen. John Abbe. Muntuwandi. Floris V. Orthologist. HiDrNick. Grundle2600. Iknowshower. John of Reading. Conversion script. Guettarda. Jamesooders. Cybercobra. Wyndynera. Karada. Anticipation of a New Lover's Arrival. Cquan. Cesar Tort. Hermanjvr. John. NatureA16. Nbarth. Edgerunner76. Catgut. Parl2001. Hrafn. Minority2005. Konman72. Dreamyshade. Tyfoo1011805. Heron. Mpbx3003. Duncharris. Robin klein. TedE. Danjelka. Lova Falk. The Anome. Moverton. SimonP. Michaelas10. Terry Longbaugh. GoldenXuniversity. Dustinasby. Saucepan. Thiago90ap. Osubuckeyeguy. Ian Pitchford. Mjresin. Merope. Eric-Wester. Tide rolls. Pgdudda. Jwanders. Scorpios. Donaldpugh. Marc Verhaegen. Hrvoje Simic. P. Cavrdg. CNicol. Hires an editor. Ickydog. Access Denied. ProfesserElvinLonghair. Gilgameshfuel. Gidonb. Whispering. Kaiwen1. Mortene. Arbeiter. Penrithguy. Fisher. Chris Rodgers. Moverton. Ybbor. JBFrenchhorn. Maikstrik. Grouf. Karatenerd. PDH. Pete. Sklmsta. Universal Life.org/w/index. Cheese. Jeremyloveshistory. Derelk. Kelly Martin. Dr Zen. J.. Kasreyn. Ikanreed. NerdyScienceDude. Liface. Hu12. Leopoldhausen. Monaca. Eldrebreath. Zyxw. Maximus Rex. WAS 4. Reinyday. Cookiemobsta. Indian Chronicles. Vox Rationis. Smalljim. Jor. Parsa. Beland. Mister Matthew. Bonzsu. Magister Mathematicae. Kazkaskazkasako. Grenavitar. Hackwrench. Ernham. Graft. Pound. Leadwind. Mangostar. Jfdwolff. ForestDim. Evercat. Salim. Vancj. AndreNel. WLU. Dawn Bard. Kzzl. A. WikiDao. Mshecket. W Pete Welch. Rmallott1. Lorenzo Braschi. Jayantanth. Graham87. Kingturtle. Doc Tropics. Factster. Maristoddard. . Frosty0814snowman. CheeseDreams. Jasmine c. JimVC3. Delphicoracle. Urthogie. Mann jess. Decltype. The wub. Concerning. Helon. Insanity Incarnate. Silverleaftree. BD2412. Londers. HumusSuepiens. Circeus. Ptresnan. Darrelljon. LizardJr8. Das Baz. Coffee2theorems. BlueNight. Cst17. Rodeosmurf. Tom Schmal. Clicketyclack. Samuell. Mgiganteus1. Joymmart. Kevin j. Williamb. Akadruid. Josephjordania. Charles Matthews. Octopus-Hands. Kabads. Michael Johnson. DougsTech. Kljenni. Epastore. Fastfission. Jpalme. Dinesh. NHJG. Nickshanks. Tycho. Sautiller. Theresa knott. Tikiwont. Davril2020. PierreAbbat. Kosigrim. Paxsimius. Dawn Bard. Phil1988. Marcus MacGregor. Erianna. Rodhullandemu. Gökhan.wikipedia. Jillysteve. Drvenik. Emc2. Dooky. Monkeyblue. JuJube. Bachrach44. Johnstone. Cyde. Samsara. Promit. Digthepast. Gonzonoir. Josh Grosse. PhilKnight. Fabartus. Trilobitealive. Cashtastrophy. Ronk01. Peproject. Falcon9x5. Marek69. DivineAlpha. Lindosland. Dmarquard. Stephenb. Neurolysis. Midgrid. Ian Pitchford. Kubigula. Natalie Erin. Darkchaos556. Icrywhen ipoo. Pjacobi. Allinthebrain. LarissaCP78. Wapondaponda. Fæ. Jonny-mt. Jagz. NawlinWiki. Evolve37830. Wes!. Ubiq. Nakon. Enlil Ninlil. Agathman. 5 albert square. G worroll. Spettro9. Nihil novi. Wikidas. Daniel. MER-C. Midnightblueowl. Scootermcknight. Gary King. Arbor. LA2. Barnaby dawson. Mr Stephen. Monado. Xezbeth. Vlad24601.E. BrainMagMo. Orgone. D6. David Andel. Manning Bartlett. John Broughton. Frowanda. Thebrid. Afaprof01. Antandrus. Dbrodbeck. Igoruha. Wmspriggs2008. Uavfun. Jockocampbell. Useight. Eamonnca1. DrFrankencelery. Nova77. Wafulz. Samsara. Conversion script. Johnuniq. Schaefer. Wet dog fur. Sonett72. Ems2. Thorell. La goutte de pluie.org/w/index. Cmart1. Jæs. Wikidudeman. Deus Ex Machina. Metta Bubble. MilitaryTarget. IRP. Cyrius. RayAYang. Fama Clamosa. Betacommand. Islandsage. Flightx52. Mstroeck. All Hallow's Wraith. Croft. Ttony21. The Thing That Should Not Be. Terence. Goethean. Z10x. Liftarn. Stefan Kruithof. Persian Poet Gal. Amh1219. Mattisse. Alansohn. Parrot. Awon. Mjpieters. K. EverSince. Rcberwick. RoyBoy. Sadi Carnot. Mjk2357. GoEThe. Mapetite526. Couloir007. Rune. Wavesmikey. Deglr6328. Yakushima. ThreeOfCups. Romanski. Alienus. Hadal. Gzkn. WikipedianProlific. Rlevse. Rkurzban. Achilton. Jorvik. Micov. SunCreator. RyanCross. Closedmouth. Glenn. Danski14. Ryulong. ABF. Kww. Cgingold. Nohat. BKalesti. Eliz81. Gurch. Zaparojdik. Jade Knight. Dancingspring. Gregconquest. Bmeacham. Bodinagamin. Joe Kress. Excirial. Phanerozoic. Josh Parris. MZMcBride. Jonathan Hall. Jmcw37. Breslauer. Rarichter. Feyre. Da monster under your bed. Foobaz. MFNickster. Jordgubbe. Aquillion. Surv1v4l1st. Nipisiquit. Luigifan. Otolemur crassicaudatus. JayJasper.delanoy. Noisy. Michelet. Anclation. Washburnmav. Stevenmitchell. Lexicon. Jjshapiro. Derekt75. Sarah. Dionyseus. JamieA. Irrevenant. NellieBly. Quena@sympatico. Siim.. Martin451. John Wilkins. Karukera. HiLo48. Tuvyah. Rednblu. Radagast83. Scilit. Iddo56. Jglyon. Robin S. Senortypant. A8UDI. Fledgeling. Zapvet. Veda784. Islescape. Mildly Mad. Jamalj03. MrWotUp. Maniadis. Wotnow. Craptree. Olivierd. Wigren. CalebNoble. Enric Naval. Jlujan69. Jackaranga. Kajekai. Jguk. Missdipsy. Sethwegner. Supersexyspacemonkey. Khamsin. ConfuciusOrnis. Tothebarricades. Zhou Yu. Realm of Shadows. Masonrose. Markunator. Mindmatrix. Vytal. KillerChihuahua. Bensaccount. Guslacerda. Teapotgeorge. Kusunose. Master Jay. Mazca. The sock that should not be. Correctia. Totorotroll. Gberkeleywannabe. Dmorg632. Granitethighs. Eleanor J Miller. Diego Grez. Oleg Alexandrov. Ga13be. NBeale.5. Waldow. Dbrodbeck. Feinoha. Gogo Dodo. Wabakimi. Adoniscik. Darkhorse82. Mayumashu. Minimac's Clone. Pauli133. Krator. Jason Potter. KuroiShiroi. Excalibur. Fieldday-sunday. Coffee. Mirv. Twinsday. Ideogram. Philippe. Cat Cubed. Hob Gadling. Dillypickle. Ramdrake. Eleanor J Miller. EvilFlyingMonkey. Giraffedata. Lskil09. David H Braun (1964). Michaelbusch. Wapondaponda. Chris goulet. Keahapana. Rich Farmbrough. TheKMan. Haemo. Dan. Arbustoo. Spammer34567. Mav. Yojasor. Zarius. DanMS. Mateuszica. Beetstra. LilDice. Greensburger. Vindicated vigilanty. MarcoTolo. Karn. 1559 anonymous edits Evolutionary psychology Source: http://en. Shambalala. Hanjabba. Clintville. Omicronpersei8. Doc Tropics. Larry_Sanger. Richfife. Dave souza. Tmh. Bluerasberry. Wayne Olajuwon.balaa. Greenwraith. Ozean. Muntuwandi. Cybercobra. Kingpin13. Kotuku33. Maunus.Hurd. ConservativeChristian. Vanished User 0001. Allinthebrain. Evoluent. Wolfrock. Lighthead. Big iron. Mkemper331. Ablewisuk. Homestarmy. Netalarm. EPM. Royalbroil. Yeohac. Shotgunlee. Headlikeawhole. Jlrobertson. Lenoxus. Rebent. Nuno Tavares. GoodSamaritan. Dupz. Rholton. LexCorp. Moonriddengirl. Nburden. Sexybomber. Kemet. Jcw69. Mentifisto. Smith609. Josh Parris. Noclevername. Steinsky. Wangi. TeleComNasSprVen. Chriskl. Shawnc. Wik. Mygerardromance. Mayalld. Pharos. Alai. Warfwar3. Adhib. Jersyko. Jossi. Cometstyles. Fleacircus. Gazpacho. SiobhanHansa. Jrockley. Non believer evolution. Hannes Hirzel. DrunkDriver333. Michael Hardy. 128. Retired username. Steinsky. Ashmoo. Hikarites. Aunt Entropy.Winsatlife. Hajatvrc. Gmlk. Markunator. Gail. Scherzbold3000. CheeseDreams. Ospalh. Theneokid. Thethornofarose. Vanished User 0001. Fuzheado. Favonian. Lemmey. ProGloriaDei. Zashaw. Fleacircus. JSF16. Rnt20. Capricorn42. Delirium. Schwnj. Leroy7. AuxBuissonets. Makeemlighter. BarrackStubbs. A. Madchester. SheffieldSteel. Fastnet. Jan. Kim Bruning. Simonmaltz. IZAK. Nickdc. JVersteeg. Imaglang. Hrafn. AJR. Honest cretan. MZMcBride. Bloopa1. PS2pcGAMER.96. Liken to a glove. Chowbok.xxx.HG. Piano non troppo. Ollj. Dr U. Lawsinium. JGrusd. Midnightblueowl. Gniniv. Marokwitz. Sunny256. JH-man. Novalis. Batman144. Grantmidnight. Jim Ellis. Limideen. PMLawrence. Joffan. Kyrisch. Croft. KHM03. JForget. TedE. 206pm.132. 65. Kedi the tramp. Dantaeos. Antandrus. DanielJosphXhan. Andre Engels. Cirt. Vaceituno. Cromwellt. Hcaudill. John D. Crazynas. Setanta747 (locked). Amd628. CanadianLinuxUser. Dreamyshade. Vassyana. Jpgordon. Tevildo. Tohd8BohaithuGh1.193. Thomas Arelatensis. Adriancurrier. Fama Clamosa. Ahoerstemeier. Gaius Cornelius. Vestbors. Braninja. Lawrencekhoo. Chrisjj. Roland Deschain. EMC125. Matturn. Shivi617. Standonbible. Jim Apple. CeoMaj. Hzcummi. AbstractEpiphany. Func. Jay-Sebastos. Aaker. Nefariousski. Khym Chanur. Daniel J. Az1568. Ianhowlett. Finlay McWalter. Keahapana. Art LaPella. D6. Logicalthinker33. RaiseYourGaze. StudyAndBeWise. Mindmatrix. N5iln. Th1rt3en.lee. Larry_Sanger. HiLo48. Ohnoitsjamie. LeeHunter. Mann jess. Hadal. Rusty Cashman. Sir. Chris3145. Hilighter555. Bucko2007. GregorB. HamSquidLllamaHam. Andrewa. Collinhasaids. Prussian725. Metamagician3000. Wdanwatts. Richard001. Phocks. Devicus. Bob frasier. Sophia. Mambaman. MARussellPESE. Chris 73. Ramdrake. Roadcreature. Calton. Sango123. Texture. David. Lskil09. Emw. Artichoker. Richard Keatinge. Caltas. Dreaded Walrus. Galoubet. TimVickers. NewEnglandYankee. Yellow Ant. Sever. JoshuaZ. MarritzN. Vestbors. Antihelios. GLGermann. Falcon8765. Lights. Fox. Slrubenstein. AdjustShift.McBratney. Gracer-t-racer. Jlujan69. Halbwolf. Melchoir. Wikipediatoperfection. Keepyouhonest. Lee J Haywood. Crispmuncher. ArcticFrog. Pallab1234. UBeR. Firefly322. Chaser. God Emperor. Rothery. HairyNevus. Pctopgs. Yanksox. Kww. Day21 07. Jiang. Wasted Sapience. Ccrashh. Pollinator. Brutaldeluxe. Martin451. Adriancurrier. Kdbuffalo. Rairai707. Titanium Dragon. Bobo192. Black Kite. Z1perlster. Mbhiii. Scientizzle. Pdelongchamp. Dave souza. Sochwa. AdamRetchless. K. WJBscribe. Frad70. Jusdafax. Cassivs. Nightscream. Epsilon60198.kt. OurGodLives. Crathen Spiroth. Cyrus Andiron. Gimme danger. DARTH SIDIOUS 2. Endomion. SonoftheMorning. Rd232. Saddhiyama. Cyde. Tmol42. Lightdarkness. Bradeos Graphon. WLU. Thegreyanomaly. Mastahcheeph. Dacxjo. Uncle Scrooge. Ethereal. Mike0001. GCarty. Eugenides. Circeus. Mildly Mad. Ref ward. Nick R. Robert Stevens. Ztobor. Maury Markowitz. JonHarder. RoyBoy. Mannafredo. Terjen. Deadbarnacle. Lord Yaar. Thebestlaidplans. El Cubano. Conroyous. Stormie. Dbachmann. Roux-HG. Temtem. ADude. Rich Farmbrough. Octopus-Hands. Jose Ramos. Grahamec. Bbatsell. Arb. Bmancini42. Biggreenjelly. Kookoo114. Dwacwise. Samsara.5. JonHarder. MER-C. Frasierwise. Czeese. Saganaki-. Kevin B12. Egern. 100110100. Leranedo. Kmarinas86. Az7997. Jamesooders. DLH. The Transhumanist. Ermeyers. Henrik46. Matej Vela. Davril2020. MarkBrownlee. Justinbrown. Richard Keatinge. Suttkus. Dpr. UncleBubba. Lethe. Gurch. RexNL. 92random. Shyhiloguy31. Caper13. JEmfinger. JamesG. Pcu123456789. Douglasers. Raeky.132. Charles Matthews. Macaddct1984. Agathman. Dancingspring. Darwinftw. Mholland. ConfuciusOrnis. UberCryxic. Schneelocke. Ragesoss. Emijrp. Velvet-Glove. Lord Emsworth. Rehumanist. Henrygb. Kairos. Addihockey10. Bueller 007. Anon vict. Jebba. Diego. TimVickers. Fecc. Vclaw. Appostle. Esurnir. Fbc215.191. Dcoetzee. Jok2000. Kwub. Maximusveritas. Goobergunch. LexCorp. Khaosworks. Professor Chaos. ThatGuamGuy. Jayjg. Cielomobile. Astrale01. JamieS93. Javit. Silly rabbit. HeikoEvermann. Krawi. Valaggar. Lmtm. SlimVirgin. IW. PEAR. Cutewithoutthet. Eugene van der Pijll. Leafyplant. Keihar. Geoffrey. Erdal Ronahi. Mindmatrix. Aunt Entropy. Plantsurfer. Como. Gregbard. Durin. American Eagle. Northwestgnome. Portillo. Miguel. Raul654. YVNP. 203. DerHexer. LibLord. Jelle564. Andy Johnston. ThaddeusB. Wereon. Johnnysmitthy. Parhodes. FridayFourthMay2007. Old-copy-editor. David Gerard. Arthena. Yuckfoo. Alex. Erik Lumberjack. Kungming2. Salsb. Algont. GoldenTorc. Luk. Kurieeto. Gerfinch. Thedewi. Goodone121. Frankchris2. Kbh3rd. Torchiest. FreezBee. Bobblewik. Gnixon. Geschichte des Bambiteletubbies. Jeff G. GB fan. Dougweller. Hburg. 129. Masterthief104. Garganthaclopes. AlphaEta. Merzul. Sasquatch. Bonkerzbanks. Homestarmy. Sjö. Meowmix4194.php?oldid=392192847 Contributors: 2over0. Haiduc. Millahnna. Gareth McCaughan. Odd nature. Headbomb. Xiong Chiamiov. Zoolanderhotept. Conscious. Habfellow. Komodon. Figma. X-factor. Causa sui. Lulzworth. Its devie whoa. Make No Name. Avicennasis. Kungfuadam. Brian Kendig. Dysprosia. Kenstal. Avb. Tritium6. RxS. Aktonin. Flowerpotman.Chesswas. Noah Salzman. Ian Pitchford. Malvenue. Arthur3030. Ligulem. Wizzywiz. Christopher Parham. Teapotgeorge. Curtmack. Inflow. Ordinary Person. Metamagician3000. Nefariousski. Ashnard. Tony Sidaway. Adraeus. Professor marginalia. John Callender. Dancingspring. Fæ.Mestel. Rominandreu. Heqwm. Legendary aa. Dawn Bard. T1980.org/w/index. DinoDude65. Vsmith. God ofcoffee. Aidje. Jnestorius. Lukeyboymcr.250.. Piddle. Big iron. Anarchangel. HIScouter. Hipocrite. Shinmawa. RadioFan2 (usurped). Vanished User 0001. Ghostintheshell. Andreworkney.68. Davewho2. Appleeater5. Hermione1980. Dharris6877. Gzkn. Fradulent Ideas. Charlesdrakew. KrakatoaKatie. Michael Johnson. Zzz111zzz. JohnOwens. Can't sleep. Kittiekatsu. Ida Shaw. Davepin. Snalwibma. Guettarda. P4k. Malerin. Wurzeller. Apokryltaros. Grice. Hammer1980. Jim62sch. Fubulover023. Shagudiga. Ronhjones. Svanslyck. Argyrios Saccopoulos. Scientizzle. Zenlax. Connolley. Nantoz. FestivalOfSouls. Kgasso. Coelacan. Diannaa. Itsnotaboutme. Sci fi writer. Kutless fan1. Closedmouth. Glane23. Janejellyroll. Irate. The Thing That Should Not Be. FeloniousMonk. CoolMike. Andycjp. Aaron Bowen. Tylerdmace. 213. Domingo Portales. Agathman. Oashi. CDV. Ben Standeven. Dastryaize. Australopithecus.164. Jzick33. Methcub. Ezra Wax. MartinHarper. Methcub. Heron. Sean. Tavengen. Dabbler. JeanandJane. JohnWoolsey. Taintedwatcher. Nneonneo. Seraphimblade. Ra2007. Kenmiller7883. Tommy2010. Gekritzl. Philjohnston. Hauskalainen. LoolTool. Sagartan. Boogeryspok. Mildly Mad. Mike Rosoft. Ricagambeda. CruftCutter. VirtualDelight. Mel Etitis. RoyBoy. GirasoleDE. PinchasC. Andrewrp. 6024kingedward1. MyNameIsNotBob. Orangemarlin. Voortle. DMacks. Spellmaster. Heimstern. Beardnomore. AVPsagaman. Hannes Röst. Woland37. IndulgentReader. Hughdbrown. JoshuaZ. Jhemdal. Bryant. Rlsheehan. Rjwilmsi. GoodDamon. ScienceApologist. Durac. Kaisershatner. Epbr123.landis. Ed Poor. FergusM1970. KRS. Meelar. Lacerda. Harmil. Barwick. Oashi. Rjwilmsi.39. Courcelles. Raymond arritt. Hanacy. Stephen B Streater. Summer Song. Ziounclesi. Invidus. Darth Panda. Hob Gadling. Doc Tropics. Cheeseypooofs. Cardsharque. GSlicer. Rick Norwood. Orangemarlin. Home Computer. Duncharris. Admiral Rupert. ArglebargleIV. DKqwerty. Nova77. Craigjmoss. Gabbe. Demonsoul5. JBlanchard. Axfy15e. Johnuniq. Bfigura's puppy. Utahraptor40. GeeJo. MichaelGensheimer. Agamemnon2. JoanneB. Filll. GraemeL. WAS 4. Islamanswering. Amcaja. Nakon. NathanLee. Sarah Waggoner.xxx. clown will eat me. Jlrobertson. JackH. Fusionbomb. FreplySpang. TimDanaher. Peyre. RedAlphaRock. Kansan. Xkoalax. Clawson. Danielrcote.J.org/w/index. Badger Drink. Natural Cut. Sfvace. 14ledzeppelin. Fubar Obfusco. Crazycomputers. Jemcalex. Kman543210. Zzzxcvb. Pschemp. Rickyrab. ChadThomson. Manta7. Kpjas. Neural.Kamenicek. Gyrus. Loodog. Polocrunch. Melvalevis. Leinad-Z. Thethornofarose. CyberAnth. Endomorphic. Jonny34567. Conversion script. Stefan Kruithof. Remiel. Jjjsixsix. Savant13. Jomasecu.hoyland. Seicer. Rareram. MONGO. Itsabouttime. Nowa. Dejvid. Alai. Karn. Benanhalt. Lear's Fool. Fred Hsu. TheMadBaron. Nondistinguished. PyroGamer. Shniken. Escape Orbit. Colin012. Wknight94. Brian0918. David Levy. Elembis. Artichoker. C56C. Aude. Mikker. Ashenai. Adashiel. Raeky. Leandrod. Laughing Joker. SheffieldSteel. ConfuciusOrnis. Markb. Polly. Mgiganteus1. Coocoo200234. Armchair info guy. Another Stickler. Stevertigo. Hunnieyes. Alan Liefting. Bigroger27509. Pmanderson. Bryan Derksen. Icairns. DragonflySixtyseven. MHess. Dugganjim.xxx. Bobby D. Malcolm Farmer. Kelovy. TShilo12. Walkiped. Zillakilla. Rednblu. SammyJames.Article Sources and Contributors Christopher. Zashaw. Jnine8.xxx. Joker828. GSlicer. Cookiehead. Jeff3000.250. Localh77. Ra2007. Chipmunkdavis. Fordmadoxfraud. Excirial. Curps. Builtfoaprice. Godardesque. Galadrien. SimonP. Didactohedron. Jparenti. Hmrox. Iridescent. A Quest For Knowledge. Xytor. Bonalaw. Jason. Jorend. Draco 2k. Nightkey. VorpalBlade. Maraman25. Silence. CO. Goldom. Guettarda. Gregkaye. Dawn Bard. J. Annoyed with fanboys. DeadEyeArrow. Teapotgeorge. BBiiis08. Ziggurat.s. M. Imgi12. Exert. Illnab1024. HalfShadow. KillerChihuahua. Edward. Gdo01. Cosmic Latte. Filll. Knowledge Seeker. Juicedboi. Art LaPella. Bensaccount. Maedin. Chrislk02. NaturalBornKiller. ManningBartlett. Northfox. Dominick. Alan_d. Sin-man. Duefiori. 200. Borb. Cryptichaos. SlimVirgin. Irishguy.. Kubigula. ZUDIN23. CinchBug. Deor. Falphin.delanoy. Abtract. Jj137. Nightkey. Gonzo fan2007. MarkSutton. Rossnixon. CatTruck. Bongwarrior. Thingg. Intelligentsium. Wisdom89. Wysprgr2005. Alan Liefting. Fbartolom.. J. Mdandrea. Haney cody. Carlaude. ClairSamoht. Janet1983. Sam Hocevar. Ucucha. Josh Keen. H. Zondor. D24m0p. Darthvader023. Mifter. Ukvilly. Frahod. Rayment. Netanel h. DoubleBlue. EALacey. Titanium Dragon. Kentfowl. Aparker62. Hrafn. Zaslav. Elm-39. SelfQ. Cardigan3000. Jtocci. Ian13.Nut.xxx. Galygal. Nigholith. Wickifrank. FilmFan69. Goethean. Caasiopia68. Jonathan. SparrowsWing. Jamht1972. Paste. Bigfun. ScottMHoward. Avs5221. H2g2bob. Johnuniq. Lightmouse.rosenheim. Hroðulf. Flowerpotman. CommonsDelinker.xxx. Ajraddatz. Cherlin. Xiner. DanielCD. Rray. CoastTOcoast533. Hazza85. CptHowdy. DigitalEnthusiast. Northfox. DaveTheRed. Antwon Bob. Orphan Wiki. Rdrozd. Scorpionman. Tesseran. Silversmith. Fubar Obfusco. 2toise. Athiest123. Htrbabaseball. Daverocks. Jacob Haller. Caster23. Vanished user. Ed Poor. Ultimus. Avengerx. Grondemar. Bassplr19. Jim62sch. Finalius. Kingoomieiii. IreverentReverend. Awayforawhile. CodeWeasel. Tobyw87. Robert Stevens. Enormousdude. Tisthammerw. Fingerz. Atwarwiththem. John D. Ec5618. Routinerob!. DCDuring. Gamescoper. SouthernNights. 62. Jelson25. Nowimnthing. Cybermonsters. Evil Monkey. WLU. Stijak. Cardsplayer4life. Hektor. 65. Shadowwolf1917. Dekisugi. BD2412.Sdraulig. LexCorp. Fairandbalanced. Rtc. Fang 23. Alansohn. Tsumetai. SkepticBanner. Joe Kress. Flex. Leptictidium. Liliuokalani. Johnleemk. Buckelz. Lkjgnhrkjghnz. Ksyrie.t. DJ Clayworth. Lulu71339. Jabal al Lawz. Esrogs. Simpsons contributor. Magister Mathematicae. Graham87. Wavelength. Discospinster. TheIncredibleEdibleOompaLoompa. Frank G Anderson. GDonato. Cedars. Rursus. James086. Justice is Needed. Farsight001. GeorgeOrr.xxx. Scott McNay. RookZERO. Panoptical. Manning Bartlett. Ground Zero. Anetode. Cosmic Latte. Vicenarian. Sikatriz. Jointquest. 1rachelkanter. Hans-Friedrich Tamke. Wikiblastfromthewikipast. Rawkfist525. Baegis. CrimsonLine. Pbarnes. La Pianista. CRouleau. Sad mouse. Harryboyles.php?oldid=392955936 Contributors: -Midorihana-. Tmol42. Escape Artist Swyer. Cbrown1023. Novachek. Goatasaur. Azcolvin429. Bhadani. Grayshi. MegamanXplosion. Kobinks. DVD R W. Exucmember. KillerChihuahua. Allstarecho. Rebroad. Wiki Raja. HarryAlffa. Minimac. Apologistdb. Rfl. StudyAndBeWise. Yeff. Rich Farmbrough. Jrbfrank. Kosebamse. Trödel. Handface. Jdavidb. Robin S. Christian *Yendi* Severin. Llort. IronGargoyle. Lacrimosus. LeaveSleaves. Twas Now. ChemoEmarles. Bluemoose.wikipedia. Bokiva. Adriansrfr. Dacer. Seeleschneider. Michael Glass. Canadian-Bacon. Andy Marchbanks. Ministry of 451 . Fredrik. Keimzelle. Eldrebreath. Emperorbma. Barnaby dawson. Narayanese. Rtc. Sxoa. Mathewsyriac.delanoy. Silly rabbit. Jefffire. Floydfan821. Salva31. Slrubenstein. Ungtss. Atradaemion. Erik the Red 2. David hogan. AndrewWTaylor. Bartledan. Spark. Cunado19. Plumbago.wikipedia. Amorymeltzer. Dpol. Richard001. Keilana. Icthus123. Bento00. Nemadude. Ian a wright. Dbabbitt. Bibliomaniac15. Columcille. Chris55. PhilipO. 979 anonymous edits Objections to evolution Source: http://en. Kainer2000.75. Frowanda. Gregbard. Wdyoung. Kzollman. Acroterion. BadgerBadger. Quadzilla99. Sander Säde. E946. XerKibard. Michael Hardy. Whoutz. Deeptrivia. Grant Gussie. KeyStroke. FeloniousMonk. Soulkeeper. Juggerboyy. ScienceApologist. Anupam. Cxz111. Factversusfiction. fiend. Mav. Peterdjones. Edwardsvaguilard. Raul654. Paul August. Ab762. JNeal. Jabberwocky03. SWoods. Mdd4696. Hdt83. Keio. Gniw. Chanting Fox. Assmar975. Poohneat. Eloquence. Raprat0. Pasado. Uberhobo. BYOB2795. Nescio. Tatarize. Kww.J. Philip J. Duileasc. SixPurpleFish. Therucks. E. DarkFalls. Inter. TLVR2004. TableManners. 5-HT8. Sandahl. Cmdrjameson.109. Jessica Simmonds. Leszek Jańczuk. Ivan Bajlo. Hassandoodle. Samsara. Crogers79. J-beda.87. Francs2000. AdultSwim. Fbartolom. Jecar. Phobos042. MattieTK. Zalgo. Johnny8899. Jkelly. Im not a vanal. 343 anonymous edits Creationism Source: http://en. Diberri. Tedmullin. Dungodung. Croft. Iamblichus. KnowledgeOfSelf. Brharris. Trishm. Iamthebordestpersononearth. Headbomb. IronCrow. Luna Santin. Dejongm. Gzuckier. J. Beeblebrox. Colin MacLaurin. The Mad Echidna. Mighty Antar. InfrequentHelper. Javierito92. Gary D. Dude1818. Malc82. PhilKnight. KSchutte. Brilliand. EamonnPKeane. Leivick. BoomerAB. AlexiusHoratius. Angela. Paul Foxworthy. Jasballz. Mark oo00. Zappernapper.158. Bob A. Sonicsuns. Elliskev. I'm the Cavalry.253. Koavf. Rhobite. Brandon5485. Empiric. Rettetast. Geogeogeo. CAF51. DanielDemaret. Tashman7. John254. 144. Finetooth. Lexor. PhilVaz. Agarner. Goal2004. Ferengi. That. Farsight001. Chunky Rice. Marriex. Fieldday-sunday. Bronks. Dougweller. Bogey97. Khehreb. MiloDC.188. Jaxl. Tawker. Chase me ladies. FireBrandon. Jigglyfidders. Anjouli. Ilkali. Matthew Yeager. Charlesdarwin. Hut 8. StuffOfInterest. Allinthebrain. Jacobolus. DavidCary. KimvdLinde. FF2010. Soulkeeper. Gscshoyru. Dave souza. Gerrit. Firsfron. Tide rolls. MEGslayer. Tony Sidaway. Lemoranjelo. RichardMathews. TrulyGroup. Smalljim. Armchair info guy. Nv8200p. Markhadman. Administer. Aaarrrggh. RIPSAW1986. Scgtrp. TokyoJapan. Gaius Cornelius. Cpl Syx. Catbar. Pinethicket. Gcbirzan. Yath. Kingpin13. IanCheesman. Apologian. R. Junjk. William M. Drmies. Hojimachong. Jchthys. Eob. GusChiggins21. Acalamari. Pigman. Creationist Zoo. Aveytare. Jimfbleak. Yorick8080. Zahid Abdassabur. Quasipalm. Gabbe. AxelBoldt. Kbolino. Tardiz1. Lee Daniel Crocker. Ling. Arjun01. Nof20. Stephenchou0722. Marknen. Gogo Dodo. Ben D. Dzubint. Casper2k3.xxx. Itsovabliss. Hephaestos. ESkog. Hello578. Doggtown. Low German. DVdm. Vernon39. Charlesdrakew. Kirbytime. Wingspantt. Coffee. Dmerrill. Foxyrockets. Mermaid from the Baltic Sea. Nunh-huh. Dougp59. Bobianite. Somepers99. Barak Sh. Jossi. Fang Aili. Dana boomer. KimvdLinde. Ramdrake. Mawest217. Plumbago. Sidonuke. Metzenberg. 24. TreasuryTag. Aleitheiophile. IslandGyrl. DarwinII. Mann jess.tayua. Kcarson. Robert Stevens. Edgar181. Jaxl. Xhaoz. BrokenSegue. LenW. Dachshund. Nakon. WAvegetarian. Sidewinder102. Unsubnl. Trishm. Antonrojo. Mikael Häggström. Rickyrab. Izehar. 24. Pleasantville. MONGO. Ascánder. Philippe. Dabomb87. Knowledge Seeker. Royboycrashfan. Albert Einstien's ghost. David D. Kazrak. Reggin500. Pgreenfinch. Space Cadet. Aristotle1990. Reinis. Zaslav.88. Rursus. Ewald. Vicki Rosenzweig. Encuesta española. Elektrik Shoos. Missvain. CJGB. MEGOP. Wetman. Raul654. Pspeck. Shoaler. Docdoley. Myanw. M2K 2. Blake-. GregAsche. FF2010. Crunchy Frog. Ummel. Ladlergo. Lee Daniel Crocker. Alexf.68. Avertist.. Avirab. XeZaR. Macguysoft. The Anome. Jpittman. KarlFrei. 213. Misonzeng. DWM. Jprw. LittleOldMe. Seabhcan.7. Bensaccount. JDoorjam. Ragesoss. MonkeyPhantom. Grutter. Bloodshedder. Temtem. Abu badali. Pdn. Trc. Swift as an Eagle. Brian Kendig. Krashlandon. Mushroom. Vsmith. Sposer. Spy5295. Bryan Derksen. Fox.E. Gareth Owen. VVtam. Heqwm. Bduke. Tmarse. DrKiernan. Rossnixon. Puddleglum Marshwiggle. Robert Merkel. Petri Krohn. RookZERO. Snemmers. Harrisbradley. RandomP. Vanished User 0001. Wimt. Duae Quartunciae. Agathman. Mark Musante. Ganggreen. Saksjn. Malo. Uncle G. Vistronic. Markaci. Paulbraveheart. Paul Magnussen. Hektor. Spintrian. MarkGallagher. Gabbe. FuelWagon. A ghost. Roflcpta. Eeshking. Postmortemjapan. Harro5. Cenarium. Duncharris. Yrusoad JDST. Michael Devore. JVC. Mmxx. Woohookitty. Frankdracman. Harry491. ScienceApologist. Flcelloguy. Life. TennisInMn. Str1977. Amazins490. Anyeverybody. Arion 3x3. Crism. Sjö. Shoeofdeath.85. Hiccupman. Raspor. Krich. Billjefferys. AndrewTJ31. Xnuala. Shambalala. Bubba73. Jason Potter. OmSKIZNIT. Pfranson. Wndl42. Gr1st. Sugarcaddy. Amarkov. Panpaniscus. Fvasconcellos. Ellebliss.lt. Atfyfe. Vunecal212. Omegatron. Igiffin. D6. Wdanwatts. Patrick0Moran. Sever. Funnyhat.xxx. Butros. Wnissen. Mav. JTBurman. Nsaa. M. Greg. Nawsum526. Bryant. Jmcc150. Serg!o. Hut 8. Caltas. Djewett. BTfromLA. Spirile. Doldrums. Monkeybait. Tomchiukc. Transity. Alweth. Saints17. Saccerzd. Clawson. Indopug. 4theChildren. Sifaka. Ace-o-aces. MickWest. Conversion script. Andrewlp1991. A Man In Black. Aunt Entropy. R'n'B. RxS. Marskell. Alienus. Phenz. Shorty2010. Scythia. Stevertigo. Instinct. DLH. LaszloWalrus. Methcub. Thermometredupochard. Wdavies. Ref ward. Mortene.128. Robert1947. Asbestos. Tacoboy42. Liontherock. 2552 anonymous edits Intelligent design Source: http://en. Jumbo Snails. Fubar Obfusco. Brian0918. Joe1978. Schweiwikist. Ambuj. GregLindahl. John Broughton.PioT. TDaySpawn. Dina. Spitfire. Snickersnee. Mirlen. BraneJ. Bongwarrior. Wizardofdos. Chubs353. Poorsod. Asderoff. Mildly Mad. CSWarren. Authr. Black Kite. DavidCary. RiverBissonnette. Frankpettit. Primalchaos. Zondor. The Mad Echidna. RadioFan. WolfieInu. Jointquest. Tisimefilms. Macarion. Charlesdrakew. Big iron. L Kensington. 217. CesarB. Possum. Mathfan. Cheater. Alansohn. Causa sui. Johnleemk. IMSoP. Guettarda. Daniel Sabo. Towne. Wassermann. Schneelocke. Malcolm Farmer. Andyjsmith. Rama. Abhishikt. Davion. Stephenb. Wikieditor06. Wolfkeeper. Runwiththewind. Gwernol. Pharaoh of the Wizards. SteveWilhelm. Astroceltica. CruftCutter. Jpeob. Baitzph. Gaohoyt. Alex bartho. William Avery. Scottfr. Christian de Quincey. Ifnord. Trevor MacInnis. Friday. Jandalhandler. Nezzadar. Pokk1. Kornbelt. Kusername. Pir. HassourZain.5. Hackwrench. Rei. Tb. Samgb. Sjakkalle. Jc4901. Freyr.xxx. Askari Mark. TheresaWilson. SchfiftyThree. Tablizer. Endomion.. Ohnoitsjamie. Treescharm0987. BenAveling. Khukri. Woodburn. Chaser. KnightLago. The Transhumanist. Jesse Viviano. Kevin B12. Son of darwin. Aragorn2. 129. Leibniz. MER-C. Vinarm. Leadfeather89. Jimfbleak. DavidSJ. Tritium6. Sushiflinger. Yamamoto Ichiro. TheFBH. MAD. Kww. Feedbackanxiety. Syvanen. Woland37. Flowers of Happiness. Texture. RucasHost.121. Karuna8. Richard001. RJaguar3. Supadawg. Working for Him. Mr. Jeff G. Magister Mathematicae. Ghewgill. Robwingfield. Moogwrench. Smadmead. Ninahexan. Eequor. Jok2000. B. Chris 73. Hal Cross. Blanchardb. John of Reading. Sannse. Dragonnas. Danski14. WordyGirl90. Psy guy. Premeditated Chaos. Lang. Matthu999. Mrtrey99. Karada. Ten*10^23. TheTrueSora. FilmFan69. Fang 23. Destoo. Kaldari. Leremyus. Connolley. Jfg284. Lightdarkness. Zundark. Yqbd. Taffboyz. Rjwilmsi.. Helbrax. Allgoodnamesalreadytaken. FQuist. Neurolysis. RedMC. Peaceworld111. Gilliam. Emanresu21. Tommy2010. Spotty11222. BG66. Laurence358. Lightmouse. Bhumburg. NZUlysses. Wenteng. NeilN. David Kernow. WikiLaurent. Gdo01. Unschool. FeanorStar7. Gadfium. Zeeshanhasan. Barbara Shack. PanzerHier. Sfvace. Tevanor. Cynicism addict. Mikker. Born2cycle. Wdflake. Exiled from GROGGS. JeremyA. Geni. Raelx. Art LaPella. Clarinetplayer. Sct72. Frostlion. Laurinaaaaa. CptKirk. Amcaja. Sahilm. Quietmarc. Property. John. Jin. Dacoutts. Jules. WikipedianMarlith. Wildnox. BBiiis08. Scherzbold3000. ThaddeusB. 130. WLU. Bloodwater. James A Whitson. Val42. NawlinWiki. Tlogmer. Richard David Ramsey. DannyMuse. Elmer Clark. Non-dropframe. MikeWren. Pretzelpaws. TechnoFaye. MisfitToys. Mooveeguy. L33tminion. 序名三. PeaceNT. Hob Gadling. Madman. Husond. Chris Melton. CptHowdy. Mallocks. Dlempa. Sopher99. Kazvorpal. Mailer diablo. Dancingspring. Fred Bauder. JokersWild1. Luokehao. Netesq. Winchelsea. Laughing Joker. Damiens. Zandperl. Evercat. Erasmocbc. Rednblu. Wtmitchell. Ck lostsword. Darwinian. Logophile. Samwb123. 65. Therucks. The bellman. Yoggysot. Zeeks. StudyAndBeWise. Grammarfreak. Aua. Cavalorn. Nefariousski. Svick. Hillbrand. Nufy8. Irishsgb. Betacommand. SQGibbon. Lefty. Rainer Wasserfuhr. GraemeL. Ollie Liddycoat. Rholton. Afterall. Bluetd. Jim62sch. Zio tom78. Jokestress. Wik. Uzimakiman2011. Anonymous Dissident. Tawker. PiCo. Coldbourne. Vanda1zew1kipedia. F. Rlsheehan. Timwi. Birdman1. Oliver202. Chad A. Tosayit. Jujutacular. Markbruder. Leeborkman.wikipedia. Web-Crawling Stickler. KnowledgeOfSelf. El Cubano. Mjb. Morning277. Filll. Sfatman1. Tothebarricades. Measure. Angr. Thewaya. Peyre. Picturef1. JzG. Danielrcote. MattShepherd. Mainstreetmark. Havermayer. Perdy80. Isotope23. H3llkn0wz. Gene Ward Smith. Viewfinder. Lilwik. Andrew Lancaster. Sleepyorangejuice. Jagged 85. Stfeeney. Ward20. Lawgor76. Jpgordon. Satori Son. FCYTravis. SiobhanHansa. RK. Honza97. Esprit15d. TML. Bondegezou. Bratsche. We66er. Lacrimosus. Shadow Puppet. Safeguarded. ChrisBradley. Krovisser. Raverguy2k7. Zzyzx11. Marshill. Bueller 007. PhDP. Gnixon. Fragtion. Lookie Louis. JWB. Tsumetai. Dave10115. THEN WHO WAS PHONE?. Rothery. Ig0774. Tannin. Node ue. Hazratio. CottonGrass. Vishnava. Versus22.144.191. K. Mindmatrix. Mego'brien. Axl. Tisthammerw. Silly rabbit. Gaius Cornelius. JoshuaZ. John Fader. Enzyme. NameIsRon. OwenX. The obakeneko. DRosenbach. Srushe. Michael Hardy. Ivar Y. Vogne87. Sandahl. GangofOne. Rosemaryamey. Scwlong. B9 hummingbird hovering. Diego pmc. NoPetrol. BMF81. Hayabusa future. Graft. Monado. Rossami. Wild Deuce. BernardSumption. Yandman. RaspK FOG. Hinotori. Peak. Green Giant. Nate slayer0. Puppppt. Naddy. FlyingOrca. Davidpatrick. Shadowjams. Fernando S. PatGallacher. Stradkid27. Goethean. Ixfd64. Ashmodai. Fish and karate. Jwbaumann. Sosmart101. Ec5618. Gogo Dodo. Izanbardprince. Dr Zen. Tide rolls. Tschel. Simon J Kissane. Scientizzle. M1rtyn. El C. TheologyJohn. Mad Scientist. QuackGuru. TheHerbalGerbil. Malcolmchurch. Izuko. NigelCunningham. ChrisDuben. UberCryxic. Keith-h. Gparker. Acalamari. Vicki Virago. Stickstickleyisafag. Ian Pitchford. Minna Sora no Shita. Twilsonb. McKay. Tresiden.xxx. Spark. Avengerx. FloNight. Mister Magotchi. Handface. Doc Tropics. Egern. Monedula. Wknight94. Oblivious. JustSomeKid. Ilikerad. Kasreyn. Antandrus. Aitias. Erkin2008. Jakes18. Jiohdi. Ciphergoth. Inkypaws. Ilkali. Eotm. AzaToth. Cpritchett42. Useight. Fairandbalanced. Adashiel. A1r. Pkgx. Jtneill. CWii. ArrrghBob.29. COGDEN. TeaDrinker. PhilKnight. Lambiam. Baegis. Nivix. Cybercobra. DGG. J. Fred J. Minna Sora no Shita. Liberty. Leithp.xxx. Steel. ST47. Turkeyphant. Itsmejudith. Jgarth. Andrew c. Nomilkforsanta. Rtc. Zero0000. Pachavi. Wikiposter0123. NickBush24.org/w/index. Computerjoe. Matilda. 198. Randy6767. DavidSaff. Bobo192. MegaMind. Everyking. LinkinPark. Charlesdarwin. Truth777. Pbarnes. Rdsmith4. CJLindsay. Johnor. ChadThomson. DJ Clayworth. SuaveArt. Thampran.php?oldid=393127480 Contributors: -Ril-. Silence. MisterSheik.xxx. Joe Kress. Johnuniq. Shanel. Bbatsell. Waggers. Paine Ellsworth. Reddi.Article Sources and Contributors Truth. Titanium Dragon. CliffC. Roger4911. Martin Hinks. Grover cleveland. Eloquence. FrozenUmbrella. Gurch. Praefectorian. Guest9999. AlexiusHoratius. Aminorex. Iwalters. Postdlf. Nutella002. Wikimk. Littlealien182. Eugeneacurry. Harrygorilla. Guy Hatton. Philip J. Benna. Icairns. Ghepeu. Speed. Simpletoremember. Tydude40. Stormwriter. Monk Bretton. RedWordSmith. GusChiggins21. Triops. Novalis. Brightflash.225. Irishtimes. Ashenai. Hgilbert.93. Dbergan. Profg. DropDeadGorgias. OllieFury. Mahjongg. Slrubenstein. Illspirit. Ikanreed. Showgun45. Orangemarlin. Cosmic Latte. DickClarkMises. Geckofuel1979. Swmeyer. AndyBoySouthPas. PhilHibbs. Justin Eiler. GreatWhiteNortherner. Farsight001. X612NT13. Fences and windows. Cla68. Mkweise. Magioladitis. Demiurge. Salva31. Oliver Pereira. SuperMoonMan. Tartsonawire. Shirahadasha. Linas. Wavelength. Davewild. Pkeck. Matthew Gibson. Trek-a-Tolk-a-Lewisite. Shem. Spriteman8. QuantumEleven. Drumguy8800. Mr-largo. Will-mansfield. Derek Ross. Joshbuddy. Stargate70. RoyBoy. Snalwibma. RexNL. Bill37212. Mirage GSM. Kalense. WalkthrougheRr. 172.2kx. 200. Thethornofarose. Zarel. Hu12. Harris7. WAS 4. ClydeOnline. Glen. DocJohnny. Traverlaw. Mholland. Jtkiefer. Bowslayer. Michael Johnson. A bit iffy.xxx. Mwazzap. SpaceFlight89. Reywas92. Brighterorange. Rick Norwood. ^demon. Teapotgeorge. Ashmoo. Anthonydent. Bevo. Benjaminevans82. Silsor. Kingpin13. 19. WikiDao. Neverquick. Petercksun. Artichoker. Carlcarlcarl. Vilerage. Quasipalm. Melly42. Dodo bird. Mglg. NBeale. Netalarm. IreverentReverend. EncephalonSeven. Adamzad. Stephensuleeman. Jaranda. Plommespiser. Berniejenkins411. FreplySpang. Cortonin. Asmetcalf. Wiki alf. Kevinccc. DKqwerty. Zelmerszoetrop.xxx. Can't sleep. StuartH. Necrogoth. Phlebas. Intranetusa. Tomlillis. Tad Lincoln. Harrydalehuffman. Dbcraft. Lithfo. Gnrlotto. Davril2020. RainbowOfLight. Pablo-flores. Triona. BD2412. ESkog. Momabdishu. Aldado. Hiberniantears. Alai. Michaelas10. Micov. Piano non troppo. GSlicer. Jed 20012. Obaidz96. FeloniousMonk. Atyy. Nutstoyoutoo. EdH. Thaurisil. Cthe. BlueMoonlet. Wotnow. King of Hearts. Oxymoron83. Dysmorodrepanis. Mel Etitis. Nasaman58. The Squicks. Ludwigs2. Ed Poor. LeadSongDog. OverlordQ. Titoxd. Kyle Barbour. Avb. JaGa. Sango123. Mentifisto. Mk270. Rich Farmbrough. Mohitkhullar. Anynobody. Jwissick. Merzul. Hubcapwiki. Counterfact. Malc82. Gimmetrow. Falcorian. Honore Fabri. Pollinator. Cognita.m. Hdhcst95. Falphin. Hans Adler. Prolog. Johnstone. DHN. LeeHunter. CloseEncounters.rf. Oxyman42. Peter Fleet. LexCorp. Dr.172. Samboy. Cuberhobo. Serenacw. Voltageman78. Hannes Karnoefel. Paul Murray. PatrickFisher. Micah Fitch.188. Mark Foskey. Millahnna. Loodog. Quux. Markbassett. Diego001. SheffieldSteel. Lordvolton. Sp. RobertG. ConfuciusOrnis. Kelly Martin.53. Digitalme. Kirill Lokshin. Joker828. Długosz. Yidisheryid. Jimput. Doc glasgow. Thiseye. Chuckgaff. Ohsimone.253.5. Pcpcpc. Mike. Puddhe. Canar. Hans Joseph Solbrig. BradC. Barneyboo. Christian List. Studerby. Johann Wolfgang. 213. Schoolbox. AnalystsAreUs. Avicennasis. Jefffire. DragonflySixtyseven. Stewacide. Welzen. Mange01. Kpearce. Musdan77. The wub. SH.O. Hadal. RickK. Rayment. Jager3333. Darthdavid. Sam. 128. David Merrill. RDF. Nick-in-South-Africa. Monkdom. Mkmcconn. Jiy. Trinirven. TheIncredibleEdibleOompaLoompa. Blackca. AVengel. TheDJ. Donarreiskoffer. Rjanag.Saxena. StephenBuxton. Abecedare. Tentacleguy. Peter G Werner. Curps. Dougp59. Furrypig. FayssalF. Dendlai. Wfward. Piotrus. SquidSix. GinaDana. Advanet. Thulesius. Elwood00. Jonzim.delanoy. Adriansrfr. GregorB. JackH.87. Rfl. AnteaterZot. ILovePlankton. Noloop. InShaneee. Vrenator. Yorick8080.168. Bbagot. Vanished user 03. CBM. Easter Monkey. MvH. Polonium. JosephCCampana. Bunchofgrapes. Populus. Bento00. Mjdellar. Philc 0780. Jamesgibbon. Phoenix-forgotten. Kozuch. IbizaHedonist. Stern. RyanParis. Christopher Parham. Hiflyer. Wapcaplet. Polsmeth. Luna Santin. I-10. Makaristos. Irishguy. Ungtss. Algae. Dark Shikari.J. Edwin Cross. Reach Out to the Truth. Hrafn. Mazin07. Debresser. Kosebamse. Fredrik. 452 . The Thing That Should Not Be. AndrewWTaylor. Stephan Schulz. BalthCat. Bumhoolery. 4twenty42o. Steinsky. DESiegel. H2g2bob. DerHexer. Dawn Bard. Shawnhath. Brusegadi. Wildstarlights. Hannes Röst. K-UNIT. Pilotguy. Toddst1. Avistor. Koavf. Mayalld. Majorly. Kf4bdy. RJN. Rndm85. Morwen. Haxwell. Williamb. Tizio.tk. Markburton. E tac. Crocodilicus. AmiDaniel. Bookandcoffee. Gscshoyru. Smkolins. RekishiEJ. Armchair info guy. Chinfo. Beny1991. Philip Trueman. Gould363. Pinethicket. Johntex. Snigbrook. DonnEdwards. Anthonyhcole. Inthebeginning. Aun'va. Steve Dufour. Treygdor. Kop. Infophile. PinchasC. MartinHarper. Jdvelasc. Dyslexic agnostic. William M. NewEnglandYankee. Supd00dlol. R. Andrewpmk. Matthew Stannard. Bkell. Bearian. Championdante. Nescio. Shoujoubill.28. Jezmck. Mehandas. KillerChihuahua. LilHelpa. Totnesmartin. Benandorsqueaks. Itayabla. PCHS-NJROTC. Sintau. Jose Ramos. Proverbs2221. Gary D. McGrupp10799. RadioFan2 (usurped). Grafen. Th1rt3en. Snoyes. Ligulem. Drernie. Eisnel. Albanaco. Skeetch. Onorem. Az1568. Midnight Gardener. Mourn. Axlq. AvicAWB. Lordsong. Wareware. Bnaur. Onebyone. Swimmer1207. Project2501a. Baley. Gwern. White Cat. SimonP. Edit1243.32. UberScienceNerd. Dave souza. Aecis. Alaffin. Genisock2. Heimstern. R. Dan100. Natl1. Discospinster. Berserkerz Crit. Tohd8BohaithuGh1. Tslocum. Shashamula.xxx. PVCCLearning. Chris the speller. Ndenison. Telop.loutsenko. Kdouglaslee. Stu42. CatLover. Gregbard. Dragons flight. IronCrow. Staffwaterboy. Bragr. Atlant. Mrkwcz. Nnp. Simulcra. Weyoun6. Cyde.101. CSTAR. Anton Gutsunaev. Mr.39. Snowbird. Trevorgoodchild. Rollingstoned413. J. Besidesamiracle. Darrenhusted. Amatulic. Yintan. Sriram sh. Yoninah. FestivalOfSouls. FireWeed. Dalinian. Newpost7. Cool3. SparrowsWing. Apollo Crua. Bobby D. Central2. Gilgamesh. Daniel. Saxifrage. Ansell. General Nolledge. Hillbillyhatfield. Yousaf465. Old-copy-editor. Tonicthebrown. RememberingLife.xxx. Cpcjr. Qwyrxian. Aoi. Tony Fox.250. Eperotao. RavShimon.xxx. The only expert. AySz88. Kbdank71. GrahamN. Chensiyuan. Cspalletta. 14. Tbones1111. Zamphuor. Svetovid. Trusilver. clown will eat me. Ashley Y. Robibrad. Nhishands4ever. Ntr11023. Seba5618.192. CuteWombat. J Milburn. AlexKarpman. Paper45tee. Pwnage8. Draco 2k. Cmdrjameson. Til Eulenspiegel. Agateller. GavinTing. Fsiler. Hypnopomp. Dp76764. Asdasdaasdasdasdasaadsasdasdasd. Shawshankhank. IZAK. Dmerrill. Canderson7. ElinorD. N5iln.164. Bobisbob. Danqaz. ImaPatriot. Foggg. Africangenesis. Exucmember. Andycjp. TehPhil. Brainbuster.. Magnus Manske. Mo0. LukeSurl. Edwy. LarRan. Terjen. Ymous. Margareta. TigerShark. AuburnPilot. Humps. Rlove. Tapir Terrific.xxx. Liftarn. Vanished user. Casliber. Lexor. Thatdoesntmakesense. Delldot. Bigmantonyd. Taw. Omegatron. Piotrus. Mojoe1185. Treisijs. Peter Menke. Tony Avedon. TedE. Vanished user. Swatjester. Trevor MacInnis. RK. The Jacobin. Pjrm. Scientizzle. W8IMP. Truthandi. Robuddha. Robert Stevens. Schama. Cerebellum. Taurrandir. NorCal764. Xyzzy n. Baley. Umpiresafe. Sj. Simpsons contributor. RxS. Ronabop. Trilemma. Pcarbonn. Steve Smith. Vapour. Octavabasso. Nowimnthing. Anupam. Rpspeck. Skyraider7. Smartse. Trilobite. Patstuart. Titoxd. Retired username. Dave souza. Xiahou. Nburden. Xianxxx. OldakQuill. Samsara. Rotem Dan.org. Sciguy47. Subversive. PromX1. Womentime. Splash. ScienceApologist. Ukegi. Ragesoss. Sillygrin. Warpsmith. Myles325a. Wyorunner. Steinsky. Wikipediarules2221. Shahab. Syzygy. WLU. Singletoned. Oliphaunt. NuclearWinner. Spewin. William2233. Splintercellguy. Theo Clark. Ruslik0. Wesley R. Triwbe. Snoyes. Charles Matthews. Moxy. Nethgirb. Robben salter. Ottava Rima. Oxyman42. Monkeyspangler. WTucker. Tomandlu. Nikodemos. Molas. Topbanana. Robert Merkel. Webpendragon. RickReinckens. SallyForth123. Rcarlberg. Parable1991. TableManners. Nishidani. Steven Kippel. Ravenswood. Richard Arthur Norton (1958. Nick Thorne. Wikignizable. Vicki Rosenzweig. Rc brooks. RainbowOfLight. Odd nature. Oz1sej. Mswake. Myork. Wikisk8r. Zscout370. Seidenstud. Rrburke. Tom Ketchum. The tooth. Wafulz. Nikola Smolenski. Unyoyega. Palthrow. Tom Cod.php?oldid=391190096 Contributors: 10outof10die. Polsmeth.org/w/index. Wren337. Mloclam901. Search4Lancer. Timwi. Will231. Ryguasu. WilsonjrWikipedia. TigerShark. Rtc. Petri Krohn. Erik Daniel P. Nigholith. Princeofexcess. R. UberScienceNerd. Saksjn. Umarekawaru. Seglea. Reidgeo. Noisy. Year 2144. Zginder. TimVickers. Profg. Cmdrjameson. Savidan. Spondoolicks. Tesseran. Rdsmith4. Sxeptomaniac. Netesq. Sjö. Node ue. PeR. NJM. Nyenyec. TeamZissou. Quietmarc. Parallel or Together?. Elsberry. Thialfi. Tsinoyboi. Mlm42. Raul654. Tfarrell. Shanel. Sonseeker007. Scetoaux. TheIncredibleEdibleOompaLoompa. Omphaloscope. Shark118. Nearfar. Rbones. THobern. RoyBoy. Vanished user. Uogl. Tschel. Pgan002. Psychohistorian. Teterty. 2391 anonymous edits Social effect of evolutionary theory Source: http://en. Gimme danger. Laurence Boyce. PhilKnight. Smack. Swarm. Munnin. Lacarnean. Reject. Wookipedia. Scope creep. Triops. Peripitus. Soundguy99. Rory096. Peedrag. Sdkl'. Thivierr. PatrickA. Northfox. Septegram. Piotrus. R. Teacherbrock. Pgk. Www. Swmeyer. Plumbago. UnicornTapestry. Wdanwatts. TUF-KAT. Wdavies. Rictus. Reginmund. Skye. Newman. Wasell. Nydas. Hillman. Deepraine. Thanissaro. Rufus843. Sunray. MuncherOfSpleens. Peruvianllama. Parhamr. TSP. Nufy8. Rd232.. The wub. Naddy. Darklilac. Ott. Smitty1276. Quokkapox. Redrocket. PhDP. Reinis. Mr. ZayZayEM. Tom. TheAlphaWolf. Nakon. RussBlau. SJP. Cosmic Latte. Radiant chains. Rjcflyer@aol. Mtdeeley. Stirling Newberry. Philip J. Zleitzen. Teapotgeorge. SMesser. Staxringold. Ted Hewlett. Talkingtoaj. Politepunk. Sovietbill9989. Vanished User 0001. Pasado. Nurban. RucasHost. Stetsonharry. Spliffy. Stokakrishna. Tomlillis. Teckor. Rakwiki. Portillo. PiCo. Selfishjeans. Vinson L Watkins. Mm40. Johnuniq. Scienceman123. Peter M Dodge. StoatBringer. Tryptofish. RedMC. Serababba. Ospalh. Simpletoremember. Pastafarian. Nick Number.tian. NuclearWarfare. Vsmith. Pudor. 46 anonymous edits 453 . Snowded. Voln. NigelCunningham. Wknight94. Scott G. Nick123. Markjoseph125. Vanished User 4517. Popeofdope420. Slrubenstein. Wisdom89. MrPeabody. TheFBH. Sjschen. Moriori. Schneelocke. Ohnoitsjamie. Nv8200p. Xstreeem. Rootxploit. Wshun. Stevertigo. Richard001. Rickyrab. Wassupwestcoast. Tretten. Sesharoo2. Sannse. Summeree. LilHelpa. Videmus Omnia. Rothery. Mr. Zzxc. MosheZadka. The Squicks. N4nojohn. Out for Truth. Steel. Tyreth. Tarinth. Leinad-Z. Tisthammerw. Nsoonhui. Pgecaj. Squady. Poor Yorick. Wereon. TeaDrinker. Tanthalas39. YellowMonkey. Pachydermballet. . Reedy. Rumping. Spotfixer. RobertG. Rich Farmbrough. SpeedyGonsales. Apokryltaros. Unbehagen. Yopienso. StephenFerg. ConfuciusOrnis. RexNL. Nothingbutmeat. Samuell. Yobol. Rousseau. Tmol42. Stevemidgley. TheDoctor10. Nstott. Wmarkham. Sam Hocevar. Scoopczar. Omicronpersei8. Pigman. Rbj. Unitvm. Netcody. RyanParis. PlantEvolBioGuy. Twinsday. SonOfNothing. Shadowin. Samsara. Northfox. Rhobite. VorpalBlade. Toby D. Wildemoi. Nina Gerlach. Orderinchaos. Quizkajer. Whoistheroach. Sophos II. OwenX. Sophia. WikiDao. Rossnixon. The Rev of Bru. VFHwebdev. Punctilius. Refreshments. Waninge. Spangineer. SpookyMulder. WJBscribe. Petral-1. Persian Poet Gal. Petesmiles. Norwikian. Orangemarlin. Seeking Objectivity in Science. JoshuaZ. Murfquake82. Wikitumnus. Tablizer. Wiki alf. TechBear. Natural Cut. Wee Jimmy. Ukkeli. Steveperry112.250.jpfo. Nightbolt. PL290. SpacemanAfrica. Ec5618. Mr Adequate. Selket. Simoes. Paul August. Royboycrashfan. Fastfission. Rahulb64. Pete. Wikiwikibangbang. Raspor. Pollinator. Pitcdbf. Spellcast. Theyeti. TheIncredibleEdibleOompaLoompa. Rednblu. Ta bu shi da yu. Tom harrison. Ossworks. Professor marginalia. Rlsheehan. Semperf. Wayward. NickelShoe. Tevildo. Mysteryboi4u. Shoemaker's Holiday. Naja Haje. Tannin. Neptunius. Tznkai. Molerat. Pediainsight. Timl2k4. Mo0. Notinasnaid. Whatjesusdoforyou. Slicky. Nnp. Suruena. Wikidudeman. Voulouza. J Milburn. Wasabe3543. Tony Fox. Trysha. Mulp. Slicing. Reywas92. Ed Poor. Ulisse0. Thv. NawlinWiki. TheGreatWhiteBuffalo. Shadoom1. Steinsky. Cyde. Sabre23t. Rracecarr. Tixity. Wimt.wikipedia. Nuttycoconut.Article Sources and Contributors Mkmcconn. Morphh. Silly rabbit. Veinor. Agamemnon2. Wesley. Royalguard11. Yidisheryid. Sam Korn. Y0u. Vague Rant. Plazak. Plunge. TreveX. Nil Einne. OmerT. Headbomb. Tony Sidaway. Yath. AdamRetchless. Patstuart. Tezh. KillerChihuahua. Stephenb. SheffieldSteel. Wavelength. We66er. Airminded. Struct. Variable. Tassedethe. Vranak. FeloniousMonk. Pilotguy. Wildspell. SCEhardt.). Paul Stansifer. Paul A. Peter. Sparcusmarcus. Stillnotelf. WAS 4. Smadmead. Wikroach. Wisco. Privatemusings. Sp. The Crying Orc. SparrowsWing. Ramdrake. Nik. Steve Dufour. Zeke pbuh. Ciroa. Alan Liefting. SlimVirgin. Rayment. Silverback. Orthologist. Wing Nut. Stuckerj. Zandperl. SandyGeorgia. R6144. Sardanaphalus. Silence. Patton123. Shikexue2. One-dimensional Tangent. Fubar Obfusco. Purplefeltangel. Overix. Zzuuzz. Ultramarine. Norklut. Zaslav. Rholton. Sad mouse. The Anome. Unknown. Twelvethirteen. XerKibard. Wolfrock. Rolinator. Rothacisms. Ttiotsw. Smith609. RebelChrysanthemum. Zelmerszoetrop. Scott MacDonald.. Vanished user 03. Filll. Duncharris. Sander Säde. Roger4911. Portillo. Yserarau. Petral. Wikipediatoperfection. ZeroOne. WeezerGeek182. Rnt20. Trishm. Tony1. Lexor. Mrt23498. Olivier. Schapel. Pyrospirit. Lefty. Stevertigo. Xerxesnine. Psy guy. Zsinj. Petrafan007. White Wolf. Nivix. Pithecusson. TheRationalistArbiter. Ungtss. Paddyman1989. Jacob1207.houston. Rdrabik. Prkastoria. Rje. Osculate. Venske. Robbrown. Murilo. Nealparr. Modemac. fiend. Rriss. Millosh. Timhenderson. Grutter. Donarreiskoffer. Nescio. Vecrumba. Aunt Entropy. RoyBoy. Phaedriel. Tfpeditor. Shirahadasha. Mqduck. SDY. Xabian40409. UninvitedCompany. FD Johnston.gnome. Billion. Roland Deschain. Whoutz. Beach. Monkeyman. Sceptre. Paul rembrandt. ZS. Yumyumpuppies. PhilipO. Snalwibma. Zargulon. Osbus. Mr Christopher.com. Ukexpat. Rmky87. Stabbycat. Uncle Dick. Black Falcon. Arctic. Morwen. Vesal. Tired time. Stradivarius. Yqbd. Rjwilmsi. PeterGrecian. Random account 47. Whirlicane. Yamamoto Ichiro. Watgap. Nondistinguished. jpg License: Public Domain Contributors: DragonFire1024.org/w/index. Wst Image:Phanerozoic biodiversity blank 01.wikipedia.org/w/index. G.wikipedia.org/w/index.org/w/index.f.betularia.png Source: http://en.svg License: GNU Free Documentation License Contributors: User:Elembis File:Selection Types Chart.wikipedia.php?title=File:Acanthodes_BW.wikipedia.0 Contributors: yaaaay File:Mutation and selection diagram.org/w/index.wikipedia.png License: GNU Free Documentation License Contributors: User:Mstroeck Image:IMG013biglittledogFX wb.png License: Public Domain Contributors: User:Philcha Image:Opabinia BW2. Philcha Image:Acanthostega BW. Aainsqatsi file:Biston.org/w/index.wikipedia.jpg Source: http://en.org/w/index.wikipedia.php?title=File:Phanerozoic_biodiversity_blank_01.jpg License: Public Domain Contributors: Ernst Haeckel File:Sanzio 01 Plato Aristotle.php?title=File:Glyptodon-Armadillo.org/w/index.php?title=File:Collapsed_tree_labels_simplified.org/w/index.php?title=File:Square-townsend-fledermaus.php?title=File:Lion_waiting_in_Nambia.wikipedia.org/w/index.jpg Source: http://en.php?title=File:BelonBirdSkel.org/w/index.org/w/index.jpg License: Creative Commons Attribution 2.jpg License: Creative Commons Attribution 3.php?title=File:Age-of-Man-wiki.php?title=File:DNA_Overview.wikipedia.5 Contributors: Kilom691.php?title=File:Charles_Darwin_by_G.php?title=File:Biston.5 Contributors: Photo by and ©2002 Dustin M.org/w/index.org/w/index. Woudloper.php?title=File:Whale_skeleton.org/w/index.wikipedia.0 Contributors: User:ArthurWeasley Image:Acanthodes BW.org/w/index.dallorto.org/w/index._Richmond.wikipedia.wikipedia.jpg License: GNU Free Documentation License Contributors: User:ToB Image:Aulonocara jacobfreibergi RB3.php?title=File:Acanthostega_BW.jpg Source: http://en.php?title=File:Tyrannosaurus_BW.7209.wikipedia. Pengo.png Source: http://en.jpg License: Creative Commons Attribution 3.jpg License: GNU Free Documentation License Contributors: Paul Harrison Image:Slime mold solves maze.org/w/index. MonteChristof.jpg License: Public Domain Contributors: Flominator.php?title=File:Slime_mold_solves_maze.wikipedia.svg License: Public Domain Contributors: User:Ilmari Karonen File:Darwin's finches.0 Contributors: User:ArthurWeasley File:Lichen.jpg Source: http://en.wikipedia. Licenses and Contributors Image:Tyrannosaurus BW.png License: Public Domain Contributors: User:Philcha Image:Horodyskia per Fedonkin 2003.jpg License: GNU Free Documentation License Contributors: Original uploader was Velho at en.org/w/index.jpg Source: http://en.wikipedia. Geoffrey.php?title=File:Sanzio_01_Plato_Aristotle.jpg Source: http://en.wikipedia.Image Sources.jpg License: Public Domain Contributors: Original uploader was Kosigrim at en.5 Contributors: Kilom691. 3 anonymous edits file:Haeckel Orchidae.jpg Source: http://en.Feb.jpg License: Public Domain Contributors: User:Wilson44691 File:Speciation modes edit.jpg License: Public Domain Contributors: L C Miall .jpg Source: http://en. Ragesoss Image:Glyptodon-Armadillo.png Source: http://en.org/w/index.wikipedia.0 Contributors: w:en:Kathy ChapmanKathy Chapman online Image:Richard dawkins.org/w/index.php?title=File:Liposome.php?title=File:Eopraptor_sketch5.jpg License: Creative Commons Attribution-Sharealike 2.jpg Source: http://en.png License: unknown Contributors: File:Tree of life by Haeckel.org/w/index.org/w/index. Wutsje. 5 anonymous edits File:Great Chain of Being 2.php?title=File:Thamnophis_sirtalis_sirtalis_Wooster.jpg Source: http://en.wikipedia. Blurpeace.svg Source: http://en.wikipedia. Patrick.jpg License: unknown Contributors: User:Sadi Carnot Image:Aquila spinogaster (African Hawk Eagle). TomCatX.php?title=File:Horodyskia_per_Fedonkin_2003. Kilom691.Sep.wikipedia.jpg Source: http://en.jpg License: Creative Commons Attribution 3.org/w/index.php?title=File:Speciation_modes_edit.jpg License: Creative Commons Attribution 2.0 Contributors: User:Azcolvin429 File:Allele-frequency.wikipedia.jpg License: Public Domain Contributors: Unknown.jpg License: GNU Free Documentation License Contributors: User:Radomil Image:James Hutton.wikipedia.php?title=File:ADN_static.org/w/index.wikipedia. Millifolium Image:Cooksonia pertoni.svg Source: http://en.php?title=File:Gene-duplication.jpg License: Public Domain Contributors: BerndH.org/w/index.php?title=File:Darwin_ape.jpg Source: http://en.wikipedia.wikipedia.wikipedia.php?title=File:Mutation_and_selection_diagram.wikipedia.wikipedia.jpg Source: http://en.wikipedia.svg License: Public Domain Contributors: User:K.landis. Kevmin.php?title=File:James_Hutton. Kentin.jpg Source: http://en.org/w/index.wikipedia.jpg Source: http://en.org/w/index.wikipedia File:Darwin ape.org/w/index.png Source: http://en.7200.3.betularia.jpg License: Public Domain Contributors: User:Dave souza File:ADN static.wikipedia file:Charles_Darwin_by_G.php?title=File:Extinction_intensity.wikipedia.org/w/index.wikipedia.php?title=File:Aquila_spinogaster_(African_Hawk_Eagle).org/w/index.php?title=File:Biston.wikipedia.org/w/index.svg Source: http://en.png License: Public Domain Contributors: Didacus Valades (Diego Valades) File:BelonBirdSkel. Tomisti.php?title=File:Stromatolites_in_Sharkbay.php?title=File:Selection_Types_Chart.5 Contributors: User:ArthurWeasley Image:Termite cathedral mounds in a bushfire blackened tropical savanna.org/w/index.php?title=File:Tree_of_life_by_Haeckel. Smith609.png Source: http://en.jpeg Source: http://en. Svdmolen.png Source: http://en.1804 .png License: Public Domain Contributors: Original uploader was TimVickers at en.png Source: http://en.org/w/index.org/w/index.jpg Source: http://en. Ies.wikipedia Image:Collapsed_tree_labels_simplified.jpg License: Creative Commons Attribution 2.jpg Source: http://en.png Source: http://en.png License: Public Domain Contributors: Original uploader was TimVickers at en. Tohma.org/w/index. Ramsey (Kralizec!) Image:Extinction intensity. Orchi.jpg License: Public Domain Contributors: Beria. Dysmorodrepanis.wikipedia.php?title=File:Termite_cathedral_mounds_in_a_bushfire_blackened_tropical_savanna.php?title=File:Great_Chain_of_Being_2.5 Contributors: User:ArthurWeasley Image:Age-of-Man-wiki.wikipedia.php?title=File:Charles_Darwin_aged_51_crop.org/w/index.org/w/index.png Source: http://en.php?title=File:Darwin's_finches.svg License: GNU Free Documentation License Contributors: Beland.png License: GNU Free Documentation License Contributors: Original uploader was Esurnir at en.jpg Source: http://en. Marco Cristo.jpg License: Creative Commons Attribution-Sharealike 2.png License: Public Domain Contributors: User:Brian0918 File:Gene-duplication.php?title=File:Opabinia_BW2.png License: Public Domain Contributors: User:Philcha Image:Stromatolites in Sharkbay.png Source: http://en.php?title=File:Ape_skeletons.jpg License: GNU Free Documentation License Contributors: Farbenfreude.org/w/index.jpg License: Creative Commons Attribution-Sharealike 2.org/w/index.jpg Source: http://en. Dragons flight. Olei file:Biston. The Hornet is no longer in publication and it is very likely for a 20-year-old artist in 1871 to have died before 1939 Image:Liposome.jpg Source: http://en.wikipedia.wikipedia.carbonaria.org/w/index.jpg License: Creative Commons Attribution 2.org/w/index.org/w/index.7200.php?title=File:Lichen.png License: Creative Commons Attribution-Sharealike 3.wikipedia.wikipedia.php?title=File:Gilboa.wikipedia. Jacobolus.png License: Creative Commons Attribution 3.jpg Source: http://en.org/w/index.jpg Source: http://en.f.png License: GNU Free Documentation License Contributors: User:Philcha Image:Eopraptor sketch5.wikipedia. Olei File:Lion waiting in Nambia.org/w/index.php?title=File:Palais_de_la_Decouverte_Tyrannosaurus_rex_p1050042.php?title=File:IMG013biglittledogFX_wb.jpg License: Creative Commons Attribution-Sharealike 2.org/w/index.0 Contributors: Matti Á File:Charles Darwin aged 51 crop.wikipedia.org/w/index.png Source: http://en.jpg Source: http://en.png Source: http://en.jpg License: GNU Free Documentation License Contributors: User:David.svg Source: http://en.wikipedia.wikipedia.wikipedia.wikipedia.0 Contributors: Steve Jurvetson Image:Square-townsend-fledermaus.jpg Source: http://en. 3 anonymous edits Image:Stephen Jay Gould (by Kathy Chapman).Monniaux File:Ape skeletons.jpg Source: http://en.jpg Source: http://en.php?title=File:Cooksonia_pertoni.1881) File:Palais de la Decouverte Tyrannosaurus rex p1050042.carbonaria.jpg Source: http://en. Licenses and Contributors 454 Image Sources.wikipedia Image:DNA Overview. Mattes.betularia. Bibi Saint-Pol.0 Contributors: User:Smith609 Image:Gilboa.wikipedia.wikipedia File:Whale skeleton._Richmond.jpeg License: Public Domain Contributors: John Gould (14.7209.php?title=File:Aulonocara_jacobfreibergi_RB3.png Source: http://en.jpg Source: http://en.jpg License: Public Domain Contributors: ArséniureDeGallium.php?title=File:Stephen_Jay_Gould_(by_Kathy_Chapman).png Source: http://en.jpg Source: http://en.png License: Public Domain Contributors: Meyers Konversionlexikon File:Thamnophis sirtalis sirtalis Wooster.php?title=File:Richard_dawkins.php?title=File:Haeckel_Orchidae.betularia.php?title=File:Allele-frequency.org/w/index. Juiced lemon.wikipedia file:Saurischian & Ornithischian pelvis bone modification w.dymaxion map.jpg License: Public Domain Contributors: Richard Owen File:Darwins first tree.png License: GNU Free Documentation License Contributors: User:Azcolvin429.5 Contributors: user:Aka Image:Asaphiscuswheelerii.org/w/index.org/w/index.php?title=File:Owen_geologic_timescale.wikipedia.wikipedia.jpg License: Public Domain Contributors: User:Alexei Kouprianov File:Huxley .wikipedia.org/w/index.png Source: http://en.Sep.org/w/index.php?title=File:Structure_of_Evolutionary_Biology.wikipedia. Licenses and Contributors File:Owen geologic timescale.wikipedia.php?title=File:Jean-baptiste_lamarck2.org/w/index.org/w/index. Jappalang.jpg License: Public Domain Contributors: Jappalang File:Image-Rhea Darwinii1.jpg Source: http://en. 2 anonymous edits File:Asa Gray (1867).jpg Source: http://en.php?title=File:Evolution_pl.wikipedia.selected species of Dinosauria.png Source: http://en.php?title=File:Route_of_vas_deferens_from_testis_to_the_penis.php?title=File:Camelid_migration_&_evolution_DymaxionMap_01.org/w/index.org/w/index.org/w/index. Glenn.php?title=File:GiraffaRecurrEn. Inductiveload.png License: Public Domain Contributors: User:Azcolvin429.org/w/index.svg License: Public Domain Contributors: Original uploader was User:TimVickers.org/w/index.svg Source: http://en.wikipedia.wikipedia.org/w/index.1804 .wikipedia.php?title=File:Titanothere_Osborn.php?title=File:CollapsedtreeLabels-simplified.php?title=File:Charles_Darwin_by_Julia_Margaret_Cameron_2.wikipedia.wikipedia Later version(s) were uploaded by Obli at en.jpg Source: http://en.3. Shadygrove2007.jpg Source: http://en.php?title=File:Darwin's_finches. Haplochromis.wikipedia.png Source: http://en.png License: Public Domain Contributors: Original uploader was TimVickers at en. Kevmin.png License: Public Domain Contributors: Original uploader was TimVickers at en.wikipedia.svg License: unknown Contributors: User:Sting Image:Brassica oleracea0.jpg Source: http://en.jpg License: Creative Commons Attribution-Sharealike 2.jpg Source: http://en.wikipedia.wikipedia.png Source: http://en.php?title=File:Charles_Darwin_seated.php?title=File:Pedigree_of_man_(Haeckel_1874).png Source: http://en.wikipedia.jpg License: GNU Free Documentation License Contributors: Albasmalko.wikipedia. User:Mario modesto file:Evolution insect mouthparts color.wikipedia Image:Jean-baptiste lamarck2.Feb.org/w/index.jpeg License: Public Domain Contributors: John Gould (14. Kevmin.png License: Creative Commons Attribution-Sharealike 3.wikipedia.wikipedia.org/w/index.jpg Source: http://en. Spawn Man.org/w/index.jpg Source: http://en. DaB. Tagishsimon.png License: GNU Free Documentation License Contributors: User:Azcolvin429 Image:Resin with insect (aka).php?title=File:Darwins_first_tree.php?title=File:Brassica_oleracea0.php?title=File:Saurischian_&_Ornithischian_pelvis_bone_modification_w-_selected_species_of_Dinosauria. Fastfission.org/w/index. Ben Tillman.png Source: http://en.org/w/index.org/w/index.wikipedia.jpg License: Creative Commons Attribution-Sharealike 2.wikipedia.php?title=File:Armadillo.jpg Source: http://en.wikipedia.php?title=File:Marsupial_biogeography_present_day_-_dymaxion_map.wikipedia.php?title=File:Darwin_divergence. file:Camelid migration & evolution DymaxionMap 01.wikipedia.org/w/index. MPF Image:Darwin's finches.wikipedia.jpg Source: http://en.jpg Source: http://en.jpg License: Public Domain Contributors: Aleator.org/w/index.jpg Source: http://en.wikipedia File:Titanothere Osborn.wikipedia.wikipedia.jpg License: Public Domain Contributors: Unknown. User:Ecelan File:GiraffaRecurrEn.jpg License: GNU Free Documentation License Contributors: User:Roepers 455 .jpg License: Creative Commons Attribution 2.jpg License: Public Domain Contributors: Conrad Martens (1801 .png License: GNU Free Documentation License Contributors: User:Azcolvin429.Mans Place in Nature.jpg License: Public Domain Contributors: Robert Chambers File:Vertebrate archetype. 3 anonymous edits Image:Cynognathus BW.png Source: http://en.png License: unknown Contributors: Original uploader was Mcy jerry at en.org/w/index.jpg License: Public Domain Contributors: John Gould File:Darwin divergence. Winterkind File:Origin of Species title page.org/w/index.php?title=File:Asa_Gray_(1867).png License: Creative Commons Attribution-Sharealike 3.jpg License: Public Domain Contributors: User:Electric goat File:CollapsedtreeLabels-simplified.jpg Source: http://en.php?title=File:Glyptodon-1.org/w/index.png License: Public Domain Contributors: Charles Darwin and Francis Darwin File:Alfred Russel Wallace 1862 .jpg License: Public Domain Contributors: User:Petter Bøckman file:Marsupial biogeography present day .png Source: http://en.jpg License: Public Domain Contributors: Hartmann Image:Charles Darwin by Julia Margaret Cameron 2.php?title=File:Rev_Baden_Powell.jpg Source: http://en.org/w/index.wikipedia.jpg License: GNU Free Documentation License Contributors: DanielCD.png License: GNU Free Documentation License Contributors: Original uploader was Mcy jerry at en.org/w/index.jpg License: GNU Free Documentation License Contributors: Ies.org/w/index.jpg Source: http://en.wikipedia.svg Source: http://en.php?title=File:Resin_with_insect_(aka).php?title=File:Huxley_-_Mans_Place_in_Nature.org/w/index.jpg License: GNU Free Documentation License Contributors: User:Valérie75 Image:Giraffes.php?title=File:HMS_Beagle_by_Conrad_Martens. The Hornet is no longer in publication and it is very likely for a 20-year-old artist in 1871 to have died before 1939 Image:Structure of Evolutionary Biology.wikipedia.org/w/index.jpg Source: http://en.png License: Public Domain Contributors: Original uploader was TimVickers at en.Elephant jaw.5 Contributors: User:ArthurWeasley Image:Horseevolution.org/w/index.wikipedia.wikipedia File:Huxley .org/w/index.png Source: http://en.php?title=File:Giraffes.1881) Image:HMS Beagle by Conrad Martens. Pfctdayelise.jpg Source: http://en.jpg Source: http://en.org/w/index.Project Gutenberg eText 15997.php?title=File:Alfred_Russel_Wallace_1862_-_Project_Gutenberg_eText_15997. Pengo.png Source: http://en.jpg License: Public Domain Contributors: Boivie.org/w/index.php?title=File:Sexlinked_inheritance_white.org/w/index. Tolanor file:Armadillo.org/w/index.png Source: http://en.php?title=File:Asaphiscuswheelerii.org/w/index.php?title=File:Origin_of_Species_title_page.php?title=File:Huxley_-_Mans_Place_in_Nature.php?title=File:Darwin_tree.org/w/index.php?title=File:Pangaea_Glossopteris.0 Contributors: User:Azcolvin429 File:Phylogenetic tree.png Source: http://en. Jappalang.php?title=File:Drosophila_melanogaster_-_side_(aka).php?title=File:Image-Rhea_Darwinii1.wikipedia.wikipedia.0 Contributors: User:Azcolvin429 Image:Drosophila melanogaster .png Source: http://en..wikipedia.wikipedia. Original uploader was User A1 at en.wikipedia. LadyofHats.wikipedia.jpg Source: http://en.php?title=File:Arabidopsis_thaliana.wikipedia.org/w/index.png License: Public Domain Contributors: Original uploader was Evercat at en.wikipedia.wikipedia File:Vestiges dev diag. Aristeas.jpg License: Public Domain Contributors: Charles Darwin File:Marsh Huxley horse.org/w/index.jpg License: Public Domain Contributors: User:Davepape Image:Darwin ape.php?title=File:Marsh_Huxley_horse.php?title=File:Charles_Darwin_-_Pigeon_skulls.jpg Source: http://en.jpg Source: http://en.wikipedia. Common Good. 5 anonymous edits File:Charles Darwin seated.png Source: http://en.svg Source: http://en.org/w/index.wikipedia.jpg Source: http://en.jpg: w:Georges CuvierGeorges Cuvier (1769–1832) File:Darwin tree.php?title=File:Darwin_ape.org/w/index.png Source: http://en. Saperaud file:Glyptodon-1.Image Sources.png Source: http://en.php?title=File:Vertebrate_archetype.Pigeon skulls.wikipedia.php?title=File:Evolution_insect_mouthparts_color. Nordelch.php?title=File:Georges_Cuvier_-_Elephant_jaw.jpg Source: http://en.side (aka).jpg Source: http://en.jpg License: Public Domain Contributors: w:Henry MaullHenry Maull (1829–1914) and w:John FoxJohn Fox (1832–1907) (Maull & Fox) UNIQ-ref-0-7337d7d7159c63fb-QINU File:Georges Cuvier .0 Contributors: User:Drbug File:Route of vas deferens from testis to the penis.wikipedia.wikipedia.jpg Source: http://en.org/w/index.org/w/index.png License: Public Domain Contributors: Cuvier_elephant_jaw. Gveret Tered.jpeg Source: http://en.php?title=File:Vestiges_dev_diag.jpg License: unknown Contributors: Henry Fairfield Osborn File:Sexlinked inheritance white. Ragesoss.org/w/index.svg License: Creative Commons Attribution-Sharealike 2. Hunadam.wikipedia. Pengo.org/w/index.org/w/index.org/w/index. SVG conversion by User:User_A1.wikipedia.5 Contributors: user:Aka Image:Arabidopsis thaliana.php?title=File:Horseevolution.org/w/index.wikipedia.wikipedia.jpg License: Public Domain Contributors: w:Benjamin Waterhouse HawkinsBenjamin Waterhouse Hawkins (1807–94) File:Pedigree of man (Haeckel 1874).jpg Source: http://en._Pengo.wikipedia file:Evolution pl.Mans Place in Nature.jpg License: unknown Contributors: Deadstar.jpg Source: http://en.php?title=File:Chromosome2_merge.png License: Public Domain Contributors: Charles Darwin File:Charles Darwin . Alex6122.jpg License: Public Domain Contributors: E Haeckel (1834–1919) File:Rev Baden Powell.php?title=File:Phylogenetic_tree.wikipedia.php?title=File:Cynognathus_BW. Lamiot. User:Mcy jerry file:Pangaea Glossopteris.21 August 1878) Image:Chromosome2 merge.png License: Public Domain Contributors: Grook Da Oger. php?title=File:Proconsul_skeleton_reconstitution_(University_of_Zurich).org/w/index. Dixi. Duesentrieb.jpg License: Creative Commons Attribution 2.jpg License: Public Domain Contributors: Diwas.org/w/index.wikipedia.org/w/index.php?title=File:Charles_Darwin_aged_51. Dirk Hünniger.5 Contributors: Image Credit: (A.wikipedia.wikipedia.JPG Source: http://en.svg Source: http://en.org/w/index.org/w/index.php?title=File:Views_on_Evolution.wikipedia. Visviva Image:Speciation modes.php?title=File:Jean-Baptiste_Lamarck.php?title=File:Polydactyly_01_Lhand_AP.jpg Source: http://en.org/w/index.jpg License: Creative Commons Attribution 2. User:Pdeitiker File:Identical cladograms.php?title=File:Drosophila_speciation_experiment. Maksim. Mattes. subsequently altered by .jpg Source: http://en.org/w/index.jpg Source: http://en.jpg License: Public Domain Contributors: ComputerHotline.svg License: Public Domain Contributors: YassineMrabet File:Notable mutations.svg Source: http://en.svg License: Public Domain Contributors: User:Mikael Häggström Image:Life cycle of a sexually reproducing organism.jpg Source: http://en.jpg License: Attribution Contributors: User:MatthiasKabel Image:Lorenz and Tinbergen1.svg Source: http://en.wikipedia.org/w/index.wikipedia. Ryz. Maksim.svg Source: http://en.0 Contributors: User:Professor marginalia File:Population bottleneck.0 Contributors: Claire Houck from New York City.org/w/index.org/w/index.svg Source: http://en.wikipedia.php?title=File:Canto_tallado_2-Guelmim-Es_Semara.org/w/index.php?title=File:Simple_cladistics.png Source: http://en. Quadell Image:Jean-Baptiste Lamarck.php?title=File:Clade_types.svg License: unknown Contributors: User:Surachit File:Tree of life SVG.php?title=File:Random_genetic_drift_chart.jpg Source: http://en.5 Contributors: User:Alexei Kouprianov Image:Cladistics-Apomorphy.jpg License: GNU Free Documentation License Contributors: Ayacop.5 Contributors: User:120 File:Canto tallado 2-Guelmim-Es Semara. .php?title=File:Tragopogon_porrifolius_flower.org/w/index.jpg License: Creative Commons Attribution 3. (B and D) by Tom Ings Image:Pfau imponierend.svg Source: http://en.org/w/index.php?title=File:Notharctus_tenebrosus_AMNH.org/w/index.svg License: unknown Contributors: User:Pbroks13 456 .0 Contributors: User:Professor marginalia File:Random genetic drift chart.org/w/index.jpg Source: http://en.php?title=File:Venus_of_Willendorf_frontview_retouched_2.jpg Source: http://en.php?title=File:Antibiotic_resistance.wikipedia. USA File:Proconsul skeleton reconstitution (University of Zurich). E.svg License: Public Domain Contributors: User:LadyofHats File:Cladogram-example1.org/w/index.5 Contributors: User:Mateuszica File:Notharctus tenebrosus AMNH.jpg License: Creative Commons Zero Contributors: User:Sklmsta File:PlesiadapisNewZICA.org/w/index.org/w/index. and F) by Rob Knell.wikipedia. Guérin Nicolas.php?title=File:Founder_effect_with_drift.php?title=File:Portulaca_grandiflora_mutant1.org/w/index.wikipedia.jpg Source: http://en.org/w/index. Bstlee. Kurpfalzbilder.php?title=File:Polyploidization. Quadell. User:Earrnz Image:Fossil hominids.php?title=File:Ambulocetus_BW.gif License: Creative Commons Attribution-Sharealike 3.php?title=File:The_Creation_of_Adam.jpg License: unknown Contributors: Unknown File:The Creation of Adam.wikipedia.0 Contributors: User:JerryFriedman Image:Benzopyrene DNA adduct 1JDG. Ragesoss.svg Source: http://en.wikipedia. Nard the Bard. ComputerHotline.php?title=File:E1b1a_ancestry. MikePhobos.org/w/index.php?title=File:Small_bonfire.png License: Creative Commons Attribution 3.wikipedia.svg Source: http://en.jpg Source: http://en. Sailko.php?title=File:Population_bottleneck.jpg License: Public Domain Contributors: Barosaul.org/w/index. Sandpiper.jpg Source: http://en.jpg Source: http://en.org/w/index.jpg Source: http://en.org/w/index.php?title=File:Biface_(France).svg License: Public Domain Contributors: User:BenB4.png License: GNU Free Documentation License Contributors: Benjah-bmm27.5 Contributors: ComputerHotline.wikipedia.png Source: http://en. PxMa. Hammond SL.wikipedia.svg License: Public Domain Contributors: User:Ilmari Karonen File:Drosophila speciation experiment.php?title=File:Phylogenetic-Groups. 5 anonymous edits Image:Gasterosteus aculeatus. and .wikipedia.png License: Public Domain Contributors: User:Luis_Fernández_García File:Clade types.svg Source: http://en.png License: Creative Commons Attribution-Sharealike 2.wikipedia.wikipedia.wikipedia. Clayton DH File:Ferrassie skull. Wst.wikipedia.org/w/index. PFHLai.png Source: http://en.wikipedia.jpg License: Creative Commons Attribution-Sharealike 3.php?title=File:Ferrassie_skull. 1 anonymous edits File:Ambulocetus BW. Infrogmation.svg License: Public Domain Contributors: User:Wykis Image:Pavo cristatus albino001xx. David Levy.wikipedia. Quadell.org/w/index.wikipedia.jpg License: unknown Contributors: Bob Burkhardt.org/w/index.dallorto.php?title=File:PlesiadapisNewZICA.svg Source: http://en.wikipedia.jpg Source: http://en.wikipedia.wikipedia.org/w/index. User:Ilmari Karonen File:Phylogenetic-Groups. Überraschungsbilder. Fastfission.gif Source: http://en. Image:Charles Darwin aged 51.JPG License: GNU Free Documentation License Contributors: user:Guérin Nicolas Image:Humanevolutionchart.wikipedia.wikipedia.php?title=File:Gasterosteus_aculeatus.svg Source: http://en. Licenses and Contributors Image:Tragopogon porrifolius flower.wikipedia.php?title=File:Notable_mutations.org/w/index.jpg License: GNU Free Documentation License Contributors: User:Professor marginalia Image:Portulaca grandiflora mutant1.php?title=File:Wasp_mimicry.org/w/index.0 Contributors: User:Faustnh Image:Simple cladistics.Image Sources.svg Source: http://en.jpg Source: http://en.JPG License: Creative Commons Attribution-Sharealike 3.jpg Source: http://en.wikipedia.php?title=File:Haeckel_arbol_bn.org/w/index.wikipedia.wikipedia.php?title=File:Scopes_trial.wikipedia.svg License: Creative Commons Attribution-Sharealike 2.png Source: http://en.org/w/index.wikipedia.php?title=File:Tree_of_life_SVG.jpg License: GNU Free Documentation License Contributors: .jpg License: Public Domain Contributors: User:Duesentrieb File:Plumpollen0060.org/w/index.svg License: GNU Free Documentation License Contributors: User:Adam Cuerden.svg License: Creative Commons Attribution-Sharealike 2. Ghouston.de.wikipedia File:Haeckel arbol bn.org/w/index.0 Contributors: User:ArthurWeasley File:Views on Evolution.php?title=File:Bee_PD_foto_explained1.org/w/index.5 Contributors: User:Surachit File:E1b1a ancestry.php?title=File:Pavo_cristatus_albino001xx.jpg Source: http://en. Satyrium.php?title=File:Random_sampling_genetic_drift.wikipedia.org/w/index.wikipedia.php?title=File:Carnica_bee_on_sedum_telephium.org/w/index.jpg License: GNU Free Documentation License Contributors: Aphaia. Xhienne.5 Contributors: User:Ilmari Karonen Image:Polyploidization.jpg License: Public Domain Contributors: User:Locutus Borg File:Small bonfire.php?title=File:Life_cycle_of_a_sexually_reproducing_organism.php?title=File:Chromosomes_mutations-en.jpg License: Creative Commons Attribution 2.png License: Public Domain Contributors: User:Muntuwandi.org/w/index.svg License: Public Domain Contributors: Original uploader was TotoBaggins at en.org/w/index.wikipedia.org/w/index.wikipedia. G.php?title=File:Humanevolutionchart.php?title=File:Identical_cladograms.php?title=File:Fossil_hominids.jpg Source: http://en.org/w/index.svg License: Public Domain Contributors: User:Wykis Image:Antibiotic resistance.png License: Creative Commons Attribution-Sharealike 3.svg Source: http://en. 1 anonymous edits File:Carnica bee on sedum telephium.jpg License: Creative Commons Attribution-Sharealike 3.jpg License: GNU Free Documentation License Contributors: user:Dixi Image:Polydactyly 01 Lhand AP.jpg License: Creative Commons Attribution-Sharealike 2. 1 anonymous edits File:Chromosomes mutations-en.jpg Source: http://en.jpg Source: http://en.org/w/index.php?title=File:Cladogram-example1.svg Source: http://en.php?title=File:Plumpollen0060.wikipedia.png License: unknown Contributors: Reed DL. Siebrand.org/w/index.wikipedia. User:Fastfission.php?title=File:Benzopyrene_DNA_adduct_1JDG. Smith VS.php?title=File:Cladistics-Apomorphy.0 Contributors: User:Kenneth Hawes File:Biface (France).php?title=File:Lorenz_and_Tinbergen1. C.0 Contributors: User:Professor marginalia File:Founder effect with drift.jpg License: GNU Free Documentation License Contributors: Max Planck Gesellschaft File:Scopes trial.0 Contributors: User:DocteurCosmos File:Venus of Willendorf frontview retouched 2.org/w/index.org/w/index.wikipedia.wikipedia.php?title=File:Pfau_imponierend.jpg Source: http://en. Rogers AR.php?title=File:Speciation_modes.wikipedia.wikipedia.jpg Source: http://en.JPG Source: http://en.wikipedia.png Source: http://en.jpg Source: http://en.wikipedia.png Source: http://en.svg License: Creative Commons Attribution-Sharealike 2.org/w/index.jpg Source: http://en.png Source: http://en.org/w/index.org/w/index. Satyrium File:Bee PD foto explained1.wikipedia.jpg Source: http://en.wikipedia.jpg License: GNU Free Documentation License Contributors: User:BS Thurner Hof File:Random sampling genetic drift. Quadell Image:Wasp mimicry.jpg License: Creative Commons Attribution 3. wikipedia. Videmus Omnia File:Pandas text analysis.org/w/index. J Milburn. Skier Dude.rf. 1 anonymous edits Image:Archaeopteryx-model.wikipedia.svg Source: http://en.Image Sources. OSX. Kevmin. K. Richard001 File:Truth fish.jpg License: Creative Commons Attribution 2. J.org/w/index.org/w/index.php?title=File:Pandas_and_ppl.png Source: http://en. Melesse.wikipedia.php?title=File:Plato_Silanion_Musei_Capitolini_MC1377. Duae Quartunciae.org/w/index.jpg License: Public Domain Contributors: Derek Ross.jpg Source: http://en. FeloniousMonk.5 Contributors: User:Jastrow Image:Pandas and ppl. Licenses and Contributors Image:Charles Darwin 1880.jpg Source: http://en. Vmenkov.php?title=File:Flagellum_base_diagram_en. G.php?title=File:WilliamPaley.jpg Source: http://en.org/w/index.jpg Source: http://en. Peripitus.-). PxMa.jpg Source: http://en. Hrafn.wikipedia. PFHLai. 1 anonymous edits 457 . Karppinen.jpg License: Public Domain Contributors: Original uploader was Sir Paul at en.png License: Public Domain Contributors: User:TimVickers Image:MichaelBehe. Duncharris.Mans Place in Nature.jpg Source: http://en.wikipedia.jpg License: Public Domain Contributors: Romanes.dallorto.org/w/index. Dinoguy2.jpg License: Creative Commons Attribution 2.wikipedia. Damiens. SnowFire. Dudo.php?title=File:Archaeopteryx-model. Rooivalk. Odd nature.org/w/index.jpg License: Creative Commons Attribution-Sharealike 2.php?title=File:MichaelBehe.wikipedia Image:The Creation of Adam.wikipedia.php?title=File:Haeckel_drawings. Image:WilliamPaley.org/w/index.jpg License: unknown Contributors: Amatulic.jpg Source: http://en. Sanbec.wikipedia. TomCatX.php?title=File:Truth_fish. 2 anonymous edits Image:Haeckel drawings.jpg License: Public Domain Contributors: Barosaul.php?title=File:The_Creation_of_Adam.jpg License: Public Domain Contributors: Original uploader was Ungtss at en.wikipedia.wikipedia.org/w/index.wikipedia File:Plato_Silanion_Musei_Capitolini_MC1377. Vanished user 03.wikipedia.php?title=File:Pandas_text_analysis.jpg Source: http://en. G. Juiced lemon.php?title=File:Creationist_car. Mattes.php?title=File:Charles_Darwin_1880. Pixel .wikipedia File:Flagellum base diagram en.php?title=File:Caricatura_de_Darwin.org/w/index.jpg License: GNU Free Documentation License Contributors: Archaeopteryxdino. Preto(m). Wolfmann. Lobo.org/w/index.jpg License: Public Domain Contributors: w:Benjamin Waterhouse HawkinsBenjamin Waterhouse Hawkins (1807–94) Image:Creationist car.php?title=File:Dembski_head_shot.php?title=File:Huxley_-_Mans_Place_in_Nature. Para.jpg Source: http://en. Mu.jpg License: GNU Free Documentation License Contributors: Campus Photo • Bryan Matluk Image:Dembski head shot.jpg Source: http://en.5 Contributors: Original uploader was Kenosis at en. Nard the Bard.org/w/index. Juiced lemon.jpg Source: http://en.wikipedia.org/w/index.jpg Source: http://en. David Levy. Sailko.jpg Source: http://en.wikipedia.wikipedia. John.org/w/index.org/w/index.jpg License: Public Domain Contributors: w:Elliott & FryElliott & Fry Image:Caricatura de Darwin.wikipedia.0 Contributors: Dodo.svg License: Public Domain Contributors: User:LadyofHats Image:Huxley . License 458 License Creative Commons Attribution-Share Alike 3. 0/ . org/ licenses/ by-sa/ 3.0 Unported http:/ / creativecommons.