The Polychaete Worms. Definitions and Keys to the Order, Families and Genera

March 17, 2018 | Author: laandeer | Category: Anatomical Terms Of Location, Fixation (Histology), Biology, Earth & Life Sciences, Nature


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THE POLYCHAETE WORMSDEFINITIONS AND KEYS TO THE ORDERS, FAMILIES AND GENERA THE POLYCHAETE WORMS Definitions and Keys to the Orders, Families and Genera By Kristian Fauchald NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY In Conjunction With THE ALLAN HANCOCK FOUNDATION UNIVERSITY OF SOUTHERN CALIFORNIA Science Series 28 February 3, 1977 . TABLE PREFACE ACKNOWLEDGMENTS INTRODUCTION CHARACTERS USED TO DEFINE HIGHER TAXA CLASSIFICATION OF POLYCHAETES ORDERS OF POLYCHAETES KEY TO FAMILIES ORDER ORBINIIDA ORDER CTENODRILIDA ORDER PSAMMODRILIDA ORDER COSSURIDA ORDER SPIONIDA ORDER CAPITELLIDA ORDER OPHELIIDA ORDER PHYLLODOCIDA ORDER AMPHINOMIDA ORDER SPINTHERIDA ORDER EUNICIDA ORDER STERNASPIDA ORDER OWENIIDA ORDER FLABELLIGERIDA ORDER FAUVELIOPSIDA ORDER TEREBELLIDA ORDER SABELLIDA FIVE "ARCHIANNELIDAN" FAMILIES GLOSSARY LITERATURE CITED INDEX OF CONTENTS vii ix 1 2 7 9 9 14 19 20 21 21 31 41 45 100 103 104 114 114 115 117 118 135 152 156 161 180 . . but also by persons willing to put in the hard work needed to fill in the gaps in our knowledge. These have been only partially considered. I apologize to my fellow polychaete workers for introducing a complex superstructure in a group which so far has been remarkably innocent of such frills. practised calmly and slowly.statements are to be made about the biology of the benthic environments.Preface THE STUDY of polychaetes used to be a leisurely occupation. This is hardly the case any longer. This review is an attempt at summarizing current information about the taxonomy and morphology of these animals. All of these developments make it important that the major morphological and anatomical features be well understood. I hope this review will bring forth a spate of investigations. The development of some polychaetes appears to be genetically interesting. The discussion is complex enough without the inclusion of speculations as to how each author would have completed his or her scheme. and some polychaetes even have become famous for their value as test animals in polluted areas. and studies of evolutionary rates and genotypic and phenotypic adaptations in these worms may prove very illuminating to our understanding of the benthic environment. Kristian Fauchald 19 May 1976 Los Angeles . In some areas they are even dominant in biomass. and preferably that the terminology and taxonomic categories be agreed upon by most workers. it became painfully obvious to me how exceedingly poorly known the group is and how few generalizations can be made on the ecology and evolution of the polychaetes. During the reviewing process. the keeping of some of these animals in culture has proved feasible. Studies of marine benthos have demonstrated that these animals may be wholly dominant both in numbers of species and in numbers of specimens. Furthermore. A great number of very sound partial schemes have been suggested from time to time. some note must be taken of the polychaetes. provided that he or she had had the evidence and inclination to do so. by persons wishing to prove me wrong in my phylogenetic speculations. and the presence of these worms hardly ever penetrated the consciousness of any but the small group of invertebrate zoologists and phylogenetlcists interested in annulated creatures. If any . . Pat Hutchings advised me of some errors in the terebellid key. I am also very grateful to Mr. prepared the index and cleaned up my English where needed. Abbott. Some of the ideas presented here were also discussed with the late Dr. Edward Ostermeyer for seeing the paper through. Ms.Acknowledgments THIS REvíEW is a direct outcome of a lecture I gave during a class in polychaete biology at Santa Catalina Marine Biological Laboratory in 1973. Jo Anne Woodcock expertly typed most of the manuscript. Catherine Link. I am also very grateful to Dr. for his support and enthusiasm and for letting me have the time to do the writing on this paper. the others I am responsible for. The good illustrations were made by Ms. My co-teacher at Catalina. I would wish to thank Dr. for prodding me into writing this book in this format and for his constant encouragement and enthusiasm for what grew from a moderately long key to families to a rather more massive offering. Fred Piltz and Bob Smith. Paul C. 1 have also discussed parts of the paper with Ray Emerson. . It has benefited greatly from discussions with all of the students in the class. Natural History Museum of Los Angeles County. Robert J. University of Southern California. Olga Hartman. Dr. was a great help in formulating some of these ideas. for which I am grateful. Lavenberg. Allan Hancock Foundation. for which I am grateful. Bernard C. Tom Kauwiing. but perhaps especially Bill Kennedy was instrumental in pushing me into giving the lecture in the first place. Schroeder. R. whose sound advice was always valued by me. . INTRODUCTION Polychaeta is part of the old. Some hermaphroditic polychaetes are known. however. and the Polychaeta marine worms. If these groups are taken into account. Keys are presented to the families and genera of the Polychaetes. This separation is obviously unsatisfactory and as information about these animals accumulated towards the end of the last century. palps. consistent and practically useful definition of what is meant by a polychaete. The three classes appeared also. tentacular cirri) and the gonadal ducts should be simple. They also should have anterior appendages of various sorts (antennae. coelomate worms in which a secondary loss of segmentation may have taken place. but in which traces of such segmentation at least can be recognized internally. Los Angeles. the only separation that consistently can be made between the oligochaetes/leeches and the polychaetes. All taxa down to the generic level are defined and a phylogenetic sequence suggested. These two groups were considered more advanced than the marine. Members of both the former classes are hermaphroditic and have various complex glands associated with reproduction. dioecious polychaetes.segmented. diffuse concept Verines. Definition of the annelid classes has varied. the Oligochaeta the earthworms and their allies. the earthworms terrestrial and the polychaetes were most common in the marine environment. as they must. The polychaetes are multi-segmented annelids with parapodia. at least grossly. is the presence in the former grouping of hermaphroditic gonads limited to a few segments. It is then difficult to give a good. FAMILIES AND GENERA^ By KRISTIAN FAUCHALD^ ABSTRACT. These definitions work if some of the smaller groups are disregarded. This phylum contains . to be separated ecologically in that the leeches were supposed to be parasitic. University of Southern California. The polychaetes have been defined for the last seventy years as dioecious. only rarely occur in bundles. but a definition along the lines suggested below. LAVENBERG DONALD REISCH ^Allan Hancock Foundation. marine annelids with parapodia bearing numerous setae. . They are dioecious and have simple exit ducts from the 'REVIEW COMMITTEE FOR THIS SCIENCE SERIES ROBERT J. most workers settled on a grouping which associated the leeches and the earthworms with each other in one group opposed to the polychaetes. California. if present. should separate them from the other annelids with reasonable accuracy. more formal definition of the phylum can be found in standard text-books. setae are present in distinct fascicles. everyday recognition of members of the three major classes has never been a problem: The Hirudinea contains the leeches. For the last seventy years or so. A review of the classification of the Class Polychaeta (Annelida) with comments on the characters used to identify the different included taxa has led lo the recognition of seventeen orders. both leeches and earthworms lack parapodia (fleshy unjointed segmental appendages) and the setae. Furthermore.THE POLYCHAETE WORMS DEFINITIONS AND KEYS TO THE ORDERS. a group that perhaps can best be defined as comprising all animals that are longer than wide and non-vertebrate. but these usually have gonads in a large fraction of the total number of segments. such as the sand worm (nereids) and bloodworms (glycerids) used for bait in parts of the world. 90007. Other. the polychaetes have been grouped with the oligochaetes and hirudineans and a few smaller groups into the phylum Annelida. Contribution Number 358 of the Allan Hancock Foundation. A key morphological feature and at the same time one of the most important taxonomic characters of the polychaetes is the setal (chaetal) construction. Any of these features need not be present and none of them is essential for the recognition of an animal as a polychaete. despite their tubicolous habits. B. The separation is based on the development of the anterior end and the life habits of the included species. 24-26). but exactly how this takes place has only partially been clarified. Orrhage 1966). The prostomium usually is distinct and may have or lack appendages. Peristomium. 166190. p. 424-425. Hartmann-Schröder 1971. pp. p. These authors gave no formal definition of any taxon above the family level (except by inference from contained taxa) and it has been difficult to evaluate their schemes. more rarely freshwater and only rarely terrestrial or parasitic in habitat. Dales used the variable structures of the eversible stomodeal region (pharynx) to separate different groups. . Antennae are always sensory. Grube 1850. The sedentaries have short parapodia associated with their tubicolous or burrowing habits and are usually deposit. Storch (1968) and Clark (1969). Hartmann-Schröder 1971. p. non-tubicolous eunicidlike animals. p. Clark 1969. Prostomial appendages include antennae and palps. they give no better solutions to classificatory problems than the old. or have treated the group as if it consisted of about 75 distinct and unrelated families (Hartman 1968. palps always have double roots. Polychaete taxonomists have tended to disregard these attempts and have continued to treat the polychaetes as if the class consisted of two orders (Fauvel 1958. admittedly anificial. Anterior appendages may be absent or a few to many similar appendages may be present. CHARACTERS USED TO DEFINE HIGHER TAXA Major anatomical and morphological features were reviewed during a study of the phylogenesis of the polychaetes (Fauchald 1974a). 281 and tables. These definitions have not changed much over time (cf. These worms are considered freeliving and. All polychaetes with jaws are included in this order. either from the brain or from the circumesophageal ring (Âkesson 1963. ERRANTIA and SEDENTARIA (Audouin and Milne Edwards 1834. Prostomium. 29) or subclasses (Uschakov 1955a). They are usually marine. It is impossible to distinguish any other classes of prostomial appendages either on morphological or anatomical grounds. The formation of structural details in the setae is very well controlled. Antennae are innervated through single roots directly from the brain. pp. The body may be separated into different regions. The material in the setae is a glycoprotein. tentacular cirri. palps may be sensory or may be used as feeding appendages.NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY gonads. antennae. In several families it is more or less fused with the peristomium and the first segments. Furthermore. formed by ectodermal cells that during the development have migrated to a position well below the rest of the ectodermally derived epidermis. 27-29.or filter-feeders. are considered errants since their large jaw-apparatus resembles the jaw-apparatus in other. The presence of either one or both categories of appendages is considered here of great importance. etc. This topic has been treated in considerably greater detail by Clark (1969) and to a lesser extent by Fauchald (1974a). The larval peristomium is the immediate prototrochal region of the trochophora larva. The three most ambitious recent proposais were by Dales (1962). generally. The position and function of the palps furnish important taxonomic characters. A. Polychaetes traditionally are separated into two large orders. The errants are supposed to have a large number of equal body-segments. The degree of fusion is difficult to determine even in an examination of the nervous system so the degree of distinctness of the prostomium is a character that can have no great taxonomic value (see Benham 1894. The separation is otherwise unsatisfactory since neither order can clearly be defined. 1969). separation into two orders. Several attempts have been made to subdivide the polychaetes in a more acceptable manner (Dales Science Series 28 1962. The position of the palps varies from ventral to dorsal. Clark used a variety of different structures to characterize his eight orders. from frontal to occipital.000-1described species of polychaetes separates into two roughly similar groups in terms of numbers of species and genera as well as families. The problem with all proposed schemes is that they are internally inconsistent. pp. Each seta consists of a bundle of filaments laid down by a basal chaetoblast and up to several lateral cells. Fauvel 1923a. The best current review of this topic was made by O'Clair and Cloney (1974) from which most of the above information has been gleaned. thus the onuphids. 1896). The arrangement of the body-wall musculature was used by Storch. Below is given a survey of the findings with an expanded discussion of their taxonomic aspects. 47). The sedentaries are supposed to have a limited number of body segments. should be rapacious in habits. 29). pp. consisting of chitin (a polysaccharide) and a protein cross-linked at the time of formation. The advantage of the system is that the bulk of the 8. The setae are ectodermal derivatives. The anterior appendages are few in number and differentiated into palps. Usually each family has a characteristic kind of pharynx. the presente of branchiae is a specific character. Amphinomida. Generally. teeth and other chitinized structures associated with the anterior end. even if tentacular cirri are absent.g. in other cases the presence may be used as a generic character (Asychis and Branchioasychis). however. In the latter case. The axial pharynx may be followed by a strongly muscular region (nereids. Eversible pharynx. This problem suggests that the structure of the pharynx cannot be used as the single definitory character for higher taxa. but in some families (e. D.). it should. Two different constructions can be recognized (Dales 1962). syllids.). at least in some forms (Altesson 1967). or uniramous with only the neuropodia developed. others have an axial pharynx that is developed symmetrically. maldanids.). nephtyids. in preparation). Polychaete parapodia can be biramous. The presence of notopodia is a very important character at the supra-familial and ordinal levels as is the presence of acicula and setae. C. However. or at least only a few alternative numbers (nereids. but current usage seems ingrained and little would be gained by coining a new term. A ventral plate-muscle pharynx is present in several forms (Eunicea. or at least nearly symmetrically.1976 THE POLYCHAETE WORMS it may persist as the adult peristomium at least in some forms and it appears to be completely pre-segmental In nature. The detailed development of each ramus with the various parapodial lobes and cirri is very important at the generic and specific levels.) or this musculature may be absent (arenicolids. etc. etc. it appears that the detailed structure of the jaws is correlated very precisely to other variable morphological features and to environmental variables (Fauchald and Smith. Spionidae) both plate-muscle and axial pharyngés are present (Orrhage 1966). the notopodia are considered secondarily reduced (Fauchald 1974a). other families tend to have PALP ANTENNA PROSTOMIUM PERISTOMIAL CIRRUS PERISTOMIUM SEGMENT liMticaroHil SETIQER PARAPODIUM SETA SEG MENTAL POSTSEQMENTAL PYGIDIUM ANUS ANAL CIRRUS NOTOPODIUM EUROPODIUM VENTRAL NERVE PARAPODIUM I. FIGURE a constant number. be remembered that the current term may cover two very different structures. The variability of the jaw-apparatus of several members of the super-family Eunicea is presently under investigation. etc. Most polychaetes can everse a part of the anterior digestive tract. Some plate-muscle pharyngés are poorly muscularized and may be difficult to distinguish from weakly developed axial pharyngés so the apparent overlap in distribution of the two kinds of pharyngés may in part be due to definitory problems. It would be valuable to distinguish the two kinds of peristomia. phyllodocids.and neuropodia developed. Parapodia. The presence of branchiae may not even be considered a specific character (Fauchald 1970 on Eunice (Nicidion) cariboea and Palola spp. especially the equipment of jaws. The fused segments may carry parapodial remnants called tentacular cirri. the structure called the peristomium in most taxonomic studies consists of a fusion of this larvally derived structure and one or more true segments. The larvally derived peristomium may carry a single pair of dorsal cirri called peristomial cirri. Diagrams showing the major morphological features of a generalized polychaete. Preliminarily. The presence of branchiae associated with the parapodia is of variable importance. with both note. etc. The number of tentacular cirri vary from one to four pairs in the hesionids. Fused segments are often present. but with generic . are very important characters at the generic and specific levels. however. The detailed structure of the pharynx..). glycerids. the whole posterior end. screening. large animals. even when the water becomes very foul. it is therefore useless to put a bottom sample in the ordinary game-bags used in diving. should be put in a large container of some sort. Most polychaete families are characterized by a complex set of features and cannot be identified by reference to a single structure. I recommend hauls as short as practicable. The process can be speeded up by adding 7% MgCla. algal mats. rubble. F. Shipboard sampling can be done with any of several kinds of a series of gear. Nephridia. SOME USEFUL TECHNIQUES The following comments are necessarily incomplete and are meant only as a first-hand guide to work on polychaetes. If dredges are used. Screening. both in numbers of . since the churning of the material in the dredge will tend to grind up the polychaetes or disturb them enough to make them autotomize appendages and. it is best to remove most or all of these as soon as possible. The character may be of wider usefulness at higher levels. such as crabs or brittle-stars. The most complete. often. such as sands and muds. limbate setae. etc. The main topics covered are collection. As the oxygen concentration in the water decreases. The detailed construction of the setae is important at the specific level. The proposed taxonomic schemes have failed because they did not take this into account. the substrate should be carefully sorted through•plucked to pieces if necessary•and the water screened for animals that left the substrate but did not reach the surface.C. They all were based on two or perhaps three manifestations in one important structure. Collection. all phyllodocids have composite setae.•Polychaetes are found nearly everywhere in the marine environment and thus can be caught with every kind of gear imaginable. or where the material is gently screened. The structure and distribution of nephridia have been used at the subfamilial and generic levels (Hessie 1917 on terebellids). It is especially important to leave the sample in the dark if it contains algae in quantity. Most polychaetes are small and active animals. Thus. the polychaetes will leave the substrate and congregate around the rim of the vessel at the air-water interface. must be screened. Usually. excluding the handling necessary for other kinds of investigations on either live or dead polychaetes. They are soft-bodied animals. but the variation in these features has been too little studied in most families to make the character useful at the present time. seaweed tufts. etc. apparent "intermediate" forms appeared and had to be included in one or another category as an exception or be left outside the proposed system. but not all polychaetes with composite setae are phyllodocids. Numerous kinds of setae have been described (uncini. The samples should be left standing for several hours (2•12 hours) depending on the lighting.•Samples of soft substrates. and put them in separate plastic bags or similar waterproof sample bags for later treatment. It became impossible to find these "hidden" taxa in the system. The samples should be placed in the dark or at least in the shade and should be kept cool. etc. composite spinigers and falcigers. and thus most easily identifiable specimens will be caught with gear that takes chunks of the environment and in situations where the animals are allowed to crawl out. etc. subacicular hooks. but we have not found this to be any great advantage. E. covered completely with sea water and be left standing undisturbed for several hours. recent studies with scanning electron microscope have demonstrated clearly the importance of accurate work on the setae (Thomassin and Rcard 1972). The basic kind of setae in each parapodial ramus is usually a family character. though not necessarily refrigerated below the ambient temperature at the samplmg site. precise observations require close microscopic work to elucidate them and this kind of work may be necessary even in routine identifications. This whole process should be done as quickly as possible. Moreover.B. since polychaetes deteriorate very rapidly after death.). the temperature. pectinate setae. but these undersample the polychaetes badly. so fairly good evidence must be presented to demonstrate that these structures are of less than specific importance. the definition of the taxon would then be left intact.A. quantitative as well as qualitative. sandsamples. Branchiae are of sufficient biological importance. especially the handling necessary to perform identificatory work. since some polychaetes are unable to leave their tubes or burrows and others generally do not approach the surface. They can easily be scooped from the surface with either a small screen (0.5 mm mesh-size) or simply a spoon. but not uniquely so. Setae. Even in S. The importance of accurate examination of the setae still is underestimated by most taxonomists. fixation and preservation and the most common techniques in laboratory handling. the "intermediate" form would be loosely appended to one taxon or another. If the sample contains a large number of motile. After the samples have been treated in this manner. The method will not quantitatively remove polychaetes. and hand-collected specimens tend to be rather poorly treated. the quantity of water in relation to sample size.U. By defining each manifestation precisely. Samples of hard substrates. Quantitative benthic studies now routinely use I mm screens. since most of the worms will escape before the sample can be treated further. collection it is advisable to collect masses of material. rocks.4 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 importance where warranted by evidence. The pan is then flooded with a solution of 7% MgCU in sea water and allowed to stand for about half an hour. and again the fraction of unidentifiable specimens goes up drastically. with as little water as possible. It is useful to have the water prescreened so that pelagic organisms can be avoided. if this is not done. Allow the sample to settle. a certain number of incomplete specimens are to be expected in any treatment. usually 70% isopropyl or ethyl alcohol in distilled water. Standard fixating agent for polychaetes is 10% neutralized formalin in sea water. Once the sediment has been removed from the screens as completely as possible. at least some samples be screened with a double set of screens. the samples must be washed in fresh water once or twice to remove the salt. and the results compared so that the level of inaccuracy engendered by the use of 1. Fixation and Preservation. but we have found that we aie amply rewarded by a much higher than usual fraction of identifiable specimens. Specimens treated as recommended above usually retain most of their appendages. The treatment suggested is rather more elaborate than usual. not least. the samples are ready to be transferred to the preservative. The most commonly used neutralizing agent is borax (Na2B407. The material on the screens is washed down onto a large enamel pan with as little sea water as possible. and the fixative should be changed twice in rapid succession to avoid dilution effects. Deep-water samples that cannot be screened immediately should be refrigerated. both I. The specimens become completely unusable after a short period of time. decant off about one-half of the solution and fill the jar with fresh formalin solution. The net result will be a waste of sampling time and effort and. The sample should fill no more than one-third of the sample jar and the jar should be completely filled with the formalin solution. the sands and gravel will make the use of 0. Thus. for each area studied. Thus. Samples should be left in formalin for at least 24 hours and can be left in the fixative for several weeks. It is much better to split a sample into several jars than to fill one jar completely with the entire sample and get incomplete and imsatisfactory fixation. In terms of the time and effort needed for the different parts of the processing of a single sample. and these are difficult to remove after they have formed.O and 0. However. Before transfer. However. This has to be done for every major sediment type and cannot be guessed at from one sediment type to the next. but each sample should be screened as soon as possible after getting it on deck. and deep-water samples should be treated with water as cold as possible. In Inshore areas. The narcotizing solution can be used several times if so desired. even if sensitivity to formalin becomes a problem.5 mm. Samples should under any circumstances be washed gently with very large quantities of water. but if a complete survey of the fauna is contemplated. this cannot be avoided.1976 THE POLYCHAETE WORMS species and in specimens.5 mm screens rather impractical. DO NOT ATTEMPT TO USE ETHYL ALCOHOL AS A FIXATIVE. to a sample jar. Histological fixatives can also be used on bulk samples. This is because the fraction of small species appears to vary geographically. after 24 hours. loss of time needed for identification of the specimens after the samples have been returned to the laboratory.0 mm screens can be estimated. and a large fraction of complete specimens is usually present. fragile polychaetes if done prior to any sorting. poorly screened samples contain a large number of mangled specimens. we now generally calculate that on the average it takes about 50 minutes of shipboard time to take and process a single shelf sample with a box-core and about three weeks of manhours to adequately treat and interpret this sample in the lab. the objective of the study undertaken must be considered carefully. Then the contents of the pan are poured carefully through a screen and transferred. but 0. setae and other details will become the crystallization sites for salt crystals. The use of 0. One change of preservative is necessary to ensure full strength. the retained material should be chased down to one side of the screen with the help of a gentle stream of water applied to the outside of the screen. Thus. especially with standard narcotizing time as suggested above. If such fixatives are used. the ratio between sample and fixative must be even lower than the one indicated above.75 mm screens does not improve matters greatly. Postfixation with formalin of material originally fixed in alcohol does not work. and such specimens usually cannot be identified. It is of the utmost importance for good results that the screening process be done carefully. and in general such fixation is better on small. technical grade).5 mm screens appear to catch most polychaetes quantitatively. the few minutes of ship time to be gained in preparing the samples poorly will be offset by the need for a larger number of replicate samples to get adequate numbers for mathematical treatments in the lab. The jar should be capped and gently but thoroughly inverted several times to get complete mixing. I would generally recommend that. Do not sort or pick through the material on the screens: non-preserved polychaetes should be handled as little as possible and never moved at all except with the help of a gentle stream of water applied to the outside of screens or elsewhere. . Nothing is gained by hurrying the screening process.•For the last few years we have routinely narcotized the whole sample as retained on the screens. nor from one geographical location to the next. less replication of samples is needed in benthic surveys and this in itself represents a savings in both time and money. Each person will also need two pairs of watchmaker's forceps. a posterior view may be more informative. but not so large as to cover the contents. in certain families and genera. and it is usually much easier to move the specimen than to move the dish. and to store these in jars filled with alcohol. a small scalpel (we use pieces of razor blades glued to applicator sticks with epoxy glue). Generally. and in the genus Magelona . Some workers find it easier to use scissors or scalpel. The sorting and identification of polychaetes is a two-step operation. When polychaetes are being identified. Screw-capped vials almost invariably have a shoulder which makes it difficult to remove specimens and labels when needed. but in special cases the anteriormost or one specific parapodium will have to be removed to study some specific detail of importance in that taxon. This sort of scan is usually all that is needed to get a polychaete to family. and they should end up well below the upper margin of the vial. cotton. a median parapodium from a long series of similar-looking parapodia will be best. Small polychaetes tend to dry out very quickly. This is done because we have found it con- Science Series 28 siderably more efficient to identify the polychaetes family by family in larger numbers. This can be done with the use of two pairs of forceps. Which parapodiimi should be removed depends on which family is being studied. as the minimum. if nothing else. Most specimens preserved as indicated above will be relatively easy to handle with two pairs of forceps. Thus. We use glycerol-alcohol for these mounts and only rarely make permanent mounts. Sorting can be done directly into vials completely filled with alcohol. and these instruments must be used on poorly preserved specimens or on larger animals. as well as with the station number for that particular sample.•The equipment needed to identify polychaetes includes one stereo microscope per person and one compound microscope per two persons. For most medium-sized polychaetes in good condition. The labels should be big enough to stand up in the vial unsupported. a bottle of immersion oil. Polychaetes always have to be treated while completely submerged. and dried out polychaetes are largely unidentifiable. but normally a flat slide is better. for example. Thus. in members of the genus Pista. two needles (we use insect pins glued to applicator sticks with epoxy glue). the parapodium should be mounted with the anterior side facing the observer. and after being transferred to alcohol cannot be left dry for more than a few minutes at a time. If the samples are to be stored as units. and #2 pencils (HB or F works well). After the sample has been sorted to family. since depression slides cannot be used with high-power compound microscopes. fine scissors. The compound microscope must be capable of magnifications up to 1100 times. cover slips. To identify the animals to genus and species. It is imperative to do this immediately rather than having to remember the content and position of each vial later. We prefer to use straight-sided vials. petri dishes of various sizes (preferably with tight-fitting lids). A parapodium mounted in a dorso-ventral position gives no more information than does looking at the whole animal. certain standard observations should always be made. It is thus of very great importance that careful and accurate notes be taken on the numbers of vials for each station. Always find the anterior end and take note of the number and arrangement of anterior appendages. paper. more accurate observations are usually necessary and various dissections must be performed. It is always necessary to remove a parapodium. so a mount showing both notopodium and neuropodium is necessary. so that later the station can be reconstructed accurately on paper. The parapodium should be mounted on a slide. we collect the members of each family in a single jar. high-rag-content paper of sufficient weight (20 pound or higher) and a good pencU. larger ones must be mounted on depression slides. capped with cotton plugs.NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Laboratory Treatment of Samples. and since light from the microscope lamp is apt to evaporate the alcohol it should be refi^shed iiom time to time. Screwcaps normally are not air tight. anterior. Care must be taken that both parapodial rami and associated cirri and branchiae come off. the vials can always be reunited afterwards if desired. a small bottle of glycerol alcohol mixed with one-half glycerol and one-half 70% alcohol. a scalpel or iris scissors. the sample is no longer intact as such. and each vial receives a label with the name of the contained taxon. Before mounting the parapodium. Use good. Identificatory Techniques. Generally. median and posterior thoracic parapodia must be removed. however. A focusabîe microscope lamp is necessary for use with the stereo microscope. but has been distributed taxonomically among families. and the stereo microscopes should have magnifications to about 100 times. The samples must be sorted while completely submerged in alcohol. so the vials will have to be stored within a larger jar regardless. Each sample should first be sorted to family under a stereo microscope. It is much easier to compare specimens from different samples. because a good look at the setae is needed. just pulling off a parapodium with a pair of forceps is the easiest method. and the number of dubious identifications can be decreased drastically by this means. depression slides and flat slides. look at some appropriate parapodial illustrations showing the features to be observed. Scan the body for obvious differences in parapodial structures and for such features as the position and number of branchiae. Especially in the case of the complex jaw structures of the eunicean polychaetes.. the left jaw being mentioned first in each formula. Setal distribution varies from one group of polychaetes to another. The sequence of families otherwise indicates phylogenetic relationships within each order. should not be used at another (lower) level within the same classificatory plan. the structure of the eversible pharynx) may have been used at the family level in one order. parapodial structures are most easily observed at relatively low magnifications under the compound microscope. jaw structures are usually situated anterior in the body. attempt to put immersion oil also between the condensor and the lower side of the slide. This makes it possible to lift a flap of tissue containing the body-wall proper. Be sure to adjust the lighting of the compound microscope every time the magnification is switched. but since the several orders and families are considered the results of a rapid radiation in Pre-Cambrian to Cambrian times (Fauchald 1974a). but at the sub-ordinal or superfamilial levels in other orders. generally. The framework formed by including the anatomical features is more satisfactory in that each order now can be defined to exclude all non-members. this saves wear and tear on the specimens and also saves time in preparation. The jaws in these forms are always observed from the dorsal side and the jaw formulae are given from the posterior to the anterior end. since the position of the different kinds of pharyngeal organs is considered of taxonomic importance. in some cases. separate vial stored within the larger vial with the specimens. if it is possible to remove one single parapodium and get all setal types represented at once. but was only partially successful. We generally make a longitudinal slit on the dorsal side. thus. this sequential arrangement can only poorly represent the phylogenetic pattern. the slit can be made farther back than usual. Theoretically different evidence should be used to define each taxonomic level. . The sequence of families indicates an increasing morphological distance from the ancestral polychaete as this was defined on that occasion. etc. a dissection should avoid cuts into the midlines of the animals. this must be done. in that one feature (e. may be most easily followed at higher magnifications. This may make them difficult to use in practical taxonomic work. they are almost invariably lost if removed. The length and position of the slit will depend on the purpose of the dissection and on the relative position of the eversible pharynx. If the purpose of the dissection is observation of the lining of the pharynx in one of the forms with a long pharynx. Dissection of the anterior end of the polychaete may also be necessary for observation of the structure of the eversible pharynx or jaw structures. Setal structures. are best seen under oil immersion with lOOx objectives. CLASSIFICATION OF POLYCHAETES The scheme proposed below (cfr.. it gives a much more satisfactory picture of the structure and will also.g. Important structures are situated in the midline. to see that the setae are as complete as possible so no extra mounts of setae become necessary. the structure of the eversible pharynx) once used at one taxonomic level. One feature (e. Leaving small parts loose in the vial will invariably lead to their loss. Again. Generally. If available light is insufficient for using the immersion objective. especially the presence of fine hairs along the cutting edge of simple setae. should be removed. it is usually not necessary to remove the lining in nephtyids and phyllodocids. dorsally or ventrally or both.g. so that they are all flattened to a similar degree when the observations are being made. This sort of separation was attempted here. even in forms with a long pharynx. especially on structures on top of a thick preparation. The pharyngeal lining has characteristic structures in members of several families. a lOx ocular is more than adequate. in the long run. and critical lighting is of the utmost importance for a clarification of these structures. save the eyesight of the worker. how many parapodia. also Table 1). Suborders have been recognized only where warranted. Families not included in any intermediate category are listed alphabetically at the end of each order. We have found it most useful to leave the pharynx and the jaws in situ as much as possible. The pharynx may be preserved in the completely retracted position or in various stages of eversión. but critical decisions. it is important to treat them in a similar manner in all specimens. it is wonhwhile to look through the key to be used before deciding which parapodium and. well lateral to the midline. no attempts were made to create intermediate categories in all orders. We usually continue by making transverse slits at both ends of the longitudinal one. It is also worthwhile to scan the body of the animal carefully before deciding on a specific parapodium. Major anatomical as well as morphological features were used to define the orders. It does not help much to use very high oculars. Parts that have been dissected out are best put in a small. is based on phylogenetic ideas presented elsewhere (Fauchald 1974a). but in order to characterize the lining of members of glycerids and goniadids. The fine structure of the glycerid pharyngeal organs cannot be seen except in oil immersion.1976 THE POLYCHAETE WORMS the ninth parapodium must be detailed. It is of great importance that the lining be well oriented in the goniadids. Onuphidae F. TABLE Science Series 28 1. especially in the larger families. Imajima and Hartman (1964). Chaetopteriformia F. Glyceridae F. Sloppiness has been the cause of more errors in the polychaete literature than all other causes combined. Ichthytomidae 12. Lysaretidae F. Antonbruunidae F. Spionidae F. Annpharelidae F. Lopadorhynchidae F. the orders are justifiable scientifically. SABELUDA F. Acrociiridae 6. Chrysopetalidae F. Capitellidae F. PHYLLODOCIDA F. Arabellidae F. O. O. O. O. 1969). Some taxa can be characterized by a single unique structure. the usage of adjectives and adverbs is rather different from common English usage. Nereidiformia F. Euphrosinidae F. The keys are intended as aids in getting a first approximation in identification. Pisionidae F. The generic keys are dependent on the presence of deciduous features. Dorvilleidae F. Under all circumstances. Pholoididae 10. Eulepethidae F. For several reasons. A name erroneously introduced in the literature for any area is in practice indelible. Nerillidae F. O. Cirratuliformia F. Saccocirridae All known taxa to the generic level have been defined. The general approach to keys taken here is that they are tools to make it possible to identify the taxa swiftly and easily and with as little damage to the specimens as possible. Ctenodrilidae F. Pilargiidae F. Sabellariidae F. Sabellongidae F. SPIONIDA SO. Introduction of a genus into the faunal lists should not be attempted based on identifications made through the keys in this paper. etc. Orbiniidae F. Spioniformia F. Poecilochaetidae F. Maldanidae O. For publication purposes. but rather difficult in practical taxonomic work. as in all other keys. AMPHINOMIDA F. Alciopidae F. Amphinomidae F. O. Aphroditacea F.. all keys were made reversible. Phyllodociformia Suborder not F. Phyllodocidae recognized: F. O. to famUy. O. Oweniidae 14. Superfamilies and Families. Pectinariidae F. Goniadidae O. Preferably. Palmyridae F. For that reason one master key to families is proposed. Ksionacea F. O. Poeobiidae 15. Dinophilidae F. Slemaspidae 13. Eunicea F. Trochochaetidae F. Polygordiidae F. one can work backwards into the key to check the identification. OPHELUDA SO. Poiynoidae F. Arenicolidae F. Caobangidae F. identifications made through the key should be checked by using the definitions of genera given. O. Sabellidae F. Identification of orders may be possible only after a detailed anatomical study. Lumbrineridae F. Day (1967) and Banse and Hobson (1974). the original literature must be consulted. Protodríiidae F. Spiiorbidae Families of uncertain afñniíies: F. Chrysopetalacea SF. CTENODRILIDA F. COSSURIDA F. Opheliidae F. STERNASPIDA F. Sphaerodoridae SO. If one has a fairly good idea of what kind of animal one has at hand from an illustration or otherwise. Syllidae Super-family not F. Terebellidae F. Cirratulidae F. O. Aphroditiformia F. Serpulidae F. keys should have been made first to order and within each order. OWENIIDA F. trick wording was avoided. FAUVELIOPGIDA F. Fauvei (1953). Typhloscolecidae F. Pontodoridae F. Once achieved. Chaetopteridae SO. but . Nephtyidae F. SPINTHERIDA F. TEREBELLIDA F. a series of very valuable regional handbooks are available. Iphitimidae SO. Nereidae F. Tomopteridae SF. several families key out at different points. PSAMMODRILIDA F. Most of these keys are short enough to allow scanning of the total key for necessary corrections if such features have been lost. Eunicidae SF. Magetonidae F.NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY 1 Survey of Polychaete Orders. FLABELLIGEREDA F. Older. in addition to complete specimens. Parergodrilidae 3. Lacydoniidae SO. Spimheridae F. Polyodontidae F. Glyceriformia F. Scalibregmidae F. Questidae 2. Hartmann-Schröder 1971). Suborders. Hartman (1968. Cossuridae 5. Flabelligeridae F. However. Such include Uschakov (1955a and 1972). Hesionidae F. as mentioned above. Because of this and because of the general variability of key features within each family. O. Bogueidae 17. Paraontdae F. Sigalionidae no. The absence of this structure in all other taxa at that level has been left unstated to save space. Trichobranchidae F. The family key was intended to discriminate between specimens that had lost deciduous features such as branchiae or antennae. The keys and definitions given are wholly inadequate as a base for description of new genera. Fauveliopsidae 16. lospilidae F. ORBINUDA F. EUNICIDA SF. Heterospionidae SO. this approach was abandoned. Calamyzidae recogaized: F. Aphroditidae 9. Psammodrilidae 4. Histriobdellidae F. O. Apistobranchidae F. The keys were made strictly dichotomous. O. Furthermore. CAPITELLIDA F. For each family. but defining this concept would have been very nearly impossible. 3a (2a). SPIONIDA. followed by taxonomic notes. The approach taken here. These explain new taxonomic combinations and taxa and specific positions taken in this paper. Uschakov 1955a. FLABELLIGERIDA. TEREBELLIDA. In addition. pelagic POEOBIIDAE Body a flattened disc with indistinct segmentation. 2b (la). A glossary contains most of the terms used. SPIONIDA. since interpretation of descriptions frequently is dependent on poorly understood and used terminology. small interstitial forms DINOPHILIDAE Body long and slender. by far the largest with most of the well-known families. External segmentation and setae absent 2 External segmentation and/or setae present 4 Paired antennae on the prostomium 3 Paired antennae absent. lb. Then comes a key to genera and definitions of all contained genera. the highly modified species now included in the orders STERNASPIDA. Fauchald and Belman 1972. the small ectoparasitic spintherids have been assigned to their own order. PSAMMODRILIDA. OWENIIDA. Blake 1975). CAPITELLIDA. Intermediate taxa were employed only where appropriate. especially noting features useful in field identification. The bulk of the recognized orders thus comes from the old collective group (or order) SEDENTARIA. the original research papers must be consulted. tactile hairs along the body. This group was never adequately defined. EUNICIDA. 2a (la). identification features have been illustrated. Clark 1969). A major revision is a long and very laborious process. 1952). However. parapodial development and setal distribution. as stated by Clark (1969). Dales 1962. The literature cited contains references to the original descriptions of all genera listed. FLABELLIGERIDA. it is wholly unsatisfactory to base such revisions on previous descriptions only. interstitial forms PROTODRILIDAE (in part) Body short and saclike. It should be noted that this paper contains little that is wholly new. as much as possible. OPHELIIDA. a definition may be followed by a brief note. AMPHINOMIDA and EUNICIDA. but must be undertaken. STERNASPIDA. except the invalid ones and to major revisions and handbooks. OWENIIDA. and the simple-bodied forms now included in the orders ORBINIIDA. TEREBELLIDA and SABELLIDA. The type-species is named for all genera and an approximate number of species is given. was that major different body constructions were given the rank of order and that modifications on these major body plans were given familial rank. These books can be used for a second approximation to the identifieatory problems. 1927). small line drawings have been added to the glossary to illustrate idiomatic usage. or followed clues indicated by revisions currently under way. FAUVELIOPGIDA. a list of invalid genera has been added. before publication. However. such a revision must be based on materials.1976 THE POLYCHAETE WORMS still indispensable are the handbooks by Fauvel (1923a. PHYLLODOCIDA. in most cases I have followed the most recent major revision of any family. the polychaetes do present an intractable problem of phylogeny for the time being. COSSURIDA. a fact that frequently has obscured close similarities in structure (cfr. COSSURIDA. Major recent reviews are also mentioned at this point. 3b (2a). but this should not be taken to indicate that the orders for that reason are related closely to each other. it will be easier to debate the classification and phylogeny of pclychaetes sensibly. These forms are about as far apart as any other grouping of polychaetes that might be proposed. ORDERS OF POLYCHAETES The following polychaete orders are recognized: ORBINHDA. etc. not on descriptions. It is hoped that this compilation of information may make it simpler to get more complete revisions made for each family where needed. CTENODRILIDA. one would hope that by organizing and defining each known taxon as clearly as possible. judging from differences in tagmatization. Members of the old order ERRANTIA are separated into three orders. because two very disparate subgroups had to be included. They could have been included under the old concept DRILOMORPHA (cfr. where necessary. SPINTHERIDA. Finally. CAPITELLIDA and OPHELÍIDA. CTENODRILIDA. PSAMMODRILIDA. The family key is followed by an order by order review of all the families. SABELLIDA. The review below is separated into several distinct parts. 4a (lb). ectoparasitic SPINTHERIDAE . Ultimately. except those in general usage in invertebrate zoology. Friedrich (1938) and Berkeley and Berkeley (1948. The latter seven orders contain structurally rather simple forms. KEY TO FAMILIES la. AMPHINOMIDA. Illustrations are given for one member of each family. PHYLLOE)OCIDA. prostomium with one to three antennae SIGALIONIDAE Elytrae alternate with dorsal cirri along the whole length of the body. 10b (8b). tube conical. Body not a flattened disc. distally curved. if present. 6a (5a). 16b (15a). notosetae usually distinctly lateral in position.. 20a (15b). 17a (16b). 18a (17b). more than four thoracic setigers SERPULIDAE . sometimes coiled near base. (8a). partially supported by acicula PISIONIDAE (in part) Prostomium not completely retracted between the first parapodia which are otherwise equipped . 7b (6b). (1 la). 18b (17b). notosetae in rosettes . including in part the prostomium. prostomium with seven antennae ONUPHIDAE (part) Specialized setae stout. specialized setae either covering the retractable anterior end or forming an operculum or a series of long protective spines (paleae) 16 Anterior end without exceptionally long.. (8a). if indistinct. (7b). forming a protective cage around the retractable anterior end. skin-papillae few and small. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 10a (8b). specialized setae (NOTE: Short. or absent. 5b (4b). 13b ( 1 lb). anterior end with two to four pairs of branchiae . 19a (18b). thoracal membrane absent 23 Tube irregularly twisted or straight. body with numerous epithelial papillae FLABELLIGERIDAE (part) Specialized anterior setae do not form a protective cage. thoracic membrane present 22 Tube raucoid or homy. 17b (16b). 16a (15a). prostomium without appendages or with numerous tentacles 18 Specialized setae in a transverse row. setae otherwise 17 Specialized setae slender. usually formed of closely fitted sandgrains PECTINARIIDAE Specialized anterior setae either as a fan-shaped group of paleae on either side of the anterior end or as an operculum to the tube 19 Specialized setae form paleae. median antenna. 11a 1 lb 12a 12b (5b). 13a (lib). elytral scars or felt 11 Neuracicula distally hammer-headed EULEPETHIDAE Neuracicula distally pointed 7 Prostomium with a single median antenna. one antenna present PHOLOIDIDAE Spinning glands present. median antenna attached at the anterior margin of the prostomium. attached near the posterior or middle of the prostomium. notosetae in transverse rows CHRYSOPETALIDAE Posterior end covered ventrally by a chitinized shield STERNASPIDAE Posterior end not covered by a shield 14 Prostomium completely retracted between the first parapodia which have three pairs of tentacular cirri. golden or brassy setae that more or less cover the dorsum 12 Notosetae otherwise (may be absent) 13 Prostomium with large facial tubercle and a median antenna. smooth and not distally curved. transformed into a tentacular crown 21 Anterior end not transformed into a tentacular crown (NOTE: Antennae and tentacular cirri may be crowded near the anterior end) 25 Tube calcareous. never harpoon-shaped 8 Neurosetae composite 9 Neurosetae simple 10 All posterior segments with elytrae. 21a (20a). paired lateral and a median antenna present. segmentation usually distinct. 20b (15b). 21b (20a). (11a). anterior end without branchiae SABELLARIIDAE Anterior end. 8a 8b 9a 9b (7b). 5a (4b). 19b (18b). body symmetrical. 15a (14b). AMPHARETIDAE (part) Specialized setae form an operculum. 22a (21a). notosetae usually present POLYNOIDAE Notopodia with expanded. dorsum with felt.10 4b (lb). (5b). often covered with sand-grains. then body clearly longer than wide 5 Dorsura with series of elytrae (scales) or distinct elytral scars present at the dorsal side of notopodial bases in several segments. or notosetae harpoon-shaped or held erect over the dorsum APHRODITIDAE Prostomium with one to three antennae. anterior end not retractable. 14a (13b). 15b (14b). 7a (6b). notosetae absent POLYODONTIDAE Spinning glands absent. dorsum without felt. 15 Anterior end with one or several series of long. 14b (13b). strong hooks may be present) 20 Specialized setae long and chambered. PALMYRIDAE Prostomium without a facial tubercle. often spinous. 6b (5a). felt of matted notosetae may be present 6 Dorsum without elytrae. 33b (32a). if homy. or without appendages 29 Branchiae in a transverse or oblique row or grouped in two groups on either side of the anterior dorsum. (29a). small. short tentacular crown with branching tentacles OWENIIDAE (part) Digestive tract recurved with anus far anteriorly. acicula present only in the prolonged acicular lobes of the second segment (first segment in juveniles). (30b). 28a (27b). peristomium usually with paired palps or tentacular cirri 30 Prostomium without appendages or with a single antenna. (30b). abdominal ones short-handled TRICHOBRANCHIDAE Both thoracic and abdominal uncini short-handled. parasitic in decapod crustaceans IPHITIMIDAE All setae composite. 35b (34a). more rarely bipinnate or lamellate (NOTE: Branchiae are often lost. branched or as numerous filaments. tubes never parchmentlike. 25b (20b). on two-three successive segments. usually digitiform and smooth. thorax with hooks in most setigers SABELLIDAE Setiger 4 with one or a few thick spines. (37b). 28b (27b). parapodia uniramous anteriorly and biramous posterioriy GONIADIDAE Jaws present 37 Jaws absent 40 Each Jaws consisting of a series of denticles in a row DORVILLEIDAE (part) Each Jaw consisting of a single piece 38 Four or five paiiï of jaws LYSARETIDAE A single pair of jaws present 39 Both composite and simple setae present. 26b (25b). uncini with several teeth in one or more transverse rows above the main fang (crested) 28 Thoracic uncini long-handled. (34b). thorax without hooks except in the first setiger CAOBANGIIDAE Digestive tract straight with far posterior or terminal anus. large notosetae furcate. THE POLYCHAETE WORMS ¡i 27b (26a). rarely smooth. 31 Caruncle absent. if homy. peristomium with paired dorsal palps. 35a (34a). otherwise absent TOMOPTERIDAE Setae or acicula or both present in most segments 34 Prostomium long and conical. (36a). (38b). Tube completely coiled. (36a). 26a (25b). 30a 30b 31a 3 lb 32a 32b (29a). (30a). sometimes as ventrolateral pads on the peristomium or fused to the anterior end of the prostomium so that the latter appear cleft. branchiae conspicuous branching tufts AMPHINOMIDAE Palps absent 33 Palps present. (30a). if present. with two pairs of antennae at the tip 35 Prostomium no more than twice as long as wide. usually fanshaped. buccal tentacles retractable. no parapodia fanshaped. usually) number of antennae and tentacular cirri. limited to a few pairs of anterior setigers 27 Anterior end with a limited (10 or fewer pairs. 29b (26b). buccal tentacles non-retractable. 29a (26b). uncini with teeth in one or a few rows AMPHARETIDAE (part) Branchiae. 34b (33b). EUPHROSINIDAE Notosetae in tufts on the notopodial lobes. stalked or sessile. tubes parchmentlike. 23a (21b). 34a (33b). usually annulated. then without annulations 26 Numerous tentacles on the lower side of the prostomium or on the peristomium. body asymmetrical.¡g76 22b (21a). branchiae shorter than setae . others smooth or serrated . if present. never annulated. parapodia either all uniramous or all biramous GLYCERIDAE Eversible pharynx with more than four jaws. but usually free and digitate 49 Setae absent. maximally two pairs of tentacular cirri or without appendages 58 Prostomium continued posteriorly in a caruncle. distinctly annulated CHAETOPTERIDAE Setiger 4 without thick spines (NOTE: Other setigers may have modified setae). or furcate setae only kind of setae present . or. furcate notosetae. (37b). antennae long or short 36 Eversible pharynx with four jaws in a cross. (38b). sometimes with a posterior prolongation in thoracic uncini TEREBELLIDAE Prostomium with at least one pair of antennae. some median parapodia strongly modified. four thoracic setigers present SPIRORBIDAE (Usually considered a sub-family of the SERPULIDAE) Parapodia with uncini in one or a few distinct rows. 24a (23a). tentacular crown with smooth or pennate radióles 24 Small uncini massed in dense fields in the neuropodia only. 24b (23a). 36a 36b 37a 37b 38a 38b 39a 39b (34b). parasitic in fish ICHTHYOTOMIDAE . 25a (20b). 23b (2lb). but scars remain). 32 Notosetae arranged in transverse rows on dorsum. branchiae. 27a (26a). 33 a (32a). if present. if present. inquilines in bivalved moUusks ANTONBRUUNIDAE Four antennae. 51b (50b). NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 42a (41b). 52b (49b). several pairs of appendages along the body 45 All setae composite. (45b). 53b (52b). 49a (32b). two or three pairs of lateral antennae and one median. 53a (52b). (40b). with a pair of antennae at the tip PISIONIDAE (part) Prostomium no longer than wide (NOTE.and peristomium equipped as indicated 85 Paired palps on the first or one of the first postperistoraial segments 60 Paired palps absent 61 . 59b (58a). 56a (55b). if present. small or large forms 47 Five prostomial antennae. 44a (43b). but are associated with the parapodia rather than situated on the dorsum) 43 Dorsal cirri large and folióse PHYLLODOCIDAE Dorsal cirri. 54b (53a). maximally five antennae 53 Palps free ventrolateral projections. pelagic PONTODORIDAE Parapodia not strongly prolonged. 55b (54b). 58a (29b). (44b). NEPHTYIDAE Interramal cirri absent. interstitial forms PROTODREJDAE More than one pair of antennae 4j Eyes larger than the rest of the prostomium. 56b (55b). paragnaths or soft papillae or both on the surface of the everted pharynx or pharynx smooth. pelagic LOPADORHYNCHIDAE At least some setae simple. 54a (53a). 57a (53b). with a few pairs of appendages anteriorly and posteriorly. usually small pigment cups. 50b (49a). parapodia often sub-biramous or uniramous HESIONIDAE Palps ventrolateral pads on the peristomium. ciniform 44 Setae absent. 57b (53b). 59a (58a). 46b (45b). everted pharynx often with a circlet of distal papillae or lappets. tentacular cirri absent DORVILLEIDAE (part) Palps biarticulated. 51a (50b). 43b (42b). with well defined lenses and pigment layers ALCIOPIDAE Eyes. smaller than the prostomium proper. paragnaths or pharyngeal papillae always absent. parasitic in lobsters HISTRIOBDELLIDAE Setae present. but lensed eyes occur 42 Series of large epithelial capsules on the dorsum. 43a (42b). sometimes fused to the prostomium so the latter appears cleft or forming ventrolateral pads on the peristomium 52 Palps multiarticulated. minute interstitial forms NERILLIDAE External ciliation limited mostly to small patches in sensory and respiratory organs. including both pro. parapodia usually biramous NEREIDAE Pharynx usually without jaws. free-living benthic forms 48 Interramal ciiri between the noto. sometimes fused to each other 54 Palps fused to the prostomium so that the latter appears anteriorly cleft 57 Prostomium longer than wide. Fused palps may make it appear longer than it is) 55 Jaws present DORVILLEIDAE (part) Jaws absent 56 Parapodia strongly prolonged.or multiarticulated 50 Palps simple. 49b (32b).¡^ 40a 40b 41 a 41b (38b). 58b (29b). or peristomium with paired palps or tentacular cirri or both pro. 42b (41b). 47a (46b). 44b 45a 45b 46a (43b). (36b). with long supportive acicula.and peristomium without appendages (NOTE: Appendages may be present on some anterior setigers) 59 Prostomium with a single median antenna. (44b). 47b (46b). all setae simple . acicula short.and neuropodia in most forms. One pair of antennae. anterior part of the digestive tract not visibly separated into distinct parts. at least one pair of tentacular cirri 51 Pharynx with paired jaws. neurosetae composite LACYDONIIDAE Palps bi. unarmed PILARGIIDAE Eversible pharynx with four pairs of upper and one pair of lower jaws EUNICIDAE Anterior end. (40b). notosetae simple. benthic or parasitic forms 46 With ciliary bands on the dorsum of each segment in addition to a large ciliary organ on the prostomium. if present. five occipital and two frontal antennae ONUPHIDAE (part) Palps either fused anteriorly to the prostomium or as free ventrolateral projections. 48b (47b).. 50a (49a). 52a (49b). anterior part of the digestive tract with a visible proventriculus in most species SYLLIDAE Eversible pharynx. 55a (54b). 48a (47b). unpaired antenna SPHAERODORIDAE Epithelial capsules absent (NOTE: Etorsal cirri of phyllodocids may be inflated. . segments usually elongated (bamboo-worms) MALDANIDAE Setal distribution otherwise. 73a 73b 74a 74b 75a 75b 76a (70b).. all setae simple. retractable filaments on the abdomen CAPITELLIDAE Slender capillary setae present on anterior. (63b). usually more than twice as long as wide. including in most cases true capillary setae 68 Thorax with series of long dorsal cirri supported by acicula PSAMMODRILIDAE Body usually not clearly regionated. 66a (65b). rostrate long-shafted uncini and spinose or smooth capillaries. 70a (68b). LUMBRINERIDAE Hooded hooks absent. three maxillary carriers ARABELLIDAE Branchiae present on maximally 15-20 segments. long-handled uncini and companion-setae. or found scattered over a large part of the body 77 Setae include anterior spines. 63a (62b). 70b (68b). apart from papillae and reduced parapodia. setal distribution and parapodial shapes grade along the body . 73 Thorax with lateral parapodia. rostrate long-shafted uncini and spinose or smooth capillaries. nearly always articulated 80 . Both composite faicigers and simple setae present ACROCIRRIDAE (part) All setae simple. including the branchial region. 65a (64b). if present. 61a (59b). 76b (73b). 72a (71b). 64b (63a). 79a (78b). sometimes acicular CIRRATULIDAE (part) With a single mid-dorsal palp on one of the first setigers (setigers 3-6 usually) . abdomen with both noto. true capillary setae absent 64 Parapodia usually well developed or at least present as low folds. 62a (61b). (66b). notopodia often reduced in posterior segments and neuropodia may form nearly complete cinctures 72 Slender capillary setae in thorax and sometimes in the first few abdominal segments only.and neuropodia in a long region of the body QUESTIDAE Simple bifid faicigers absent 79 Prostomium an elongated cone. setae otherwise 67 Setae in four fascicles on each segment (biramous condition) CTENODRILIDAE Setae in two fascicles on each segment (uniramous condition) PARERGODRILIDAE Prostomium an oblique plaque. usually bordered by a flange 69 Prostomium pointed. setae smooth and slightly recurved FAUVELIOPSIDAE Without papillae between the rami.and neuropodia in dorsal positions ORBINIIDAE Parapodia lateral in all parts of the body. 71a (70a). starting from one of setigers 4-10 PARAONIDAE (part) Branchiae. segments rarely elongated 78 Setae include simple capillaries and simple bifid faicigers in both noto. 77a (76b). THE POLYCHAETE WORMS 13 69b (68a). 60b (59a). 63b (62b). one pair of maxillary carriers . 62b (61b). 61b (59b). (68a). (74b).. 64a (63a). 65b (64b). 71b (70a). median and sometimes posterior parts of the body. COSSURIDAE Mid-dorsal palp absent 62 With series of long..1976 60a (59a). scars remain) CIRRATULIDAE (part) Branchial filaments and tentacular cirri absent or limited to a few segments 63 Parapodia strongly reduced so that the setae appear sessile on the body-wall. slender branchial filaments and tentacular and dorsal cirri along the body (often lost. setae usually of several different kinds. 77b (76b). (66b). (73a). 78b (77b). 72b (7lb). (70b). if present. (63b).. rounded or blunt 70 Setae include anterior spines. distally curved or straight. (73a). either limited to the extreme anterior end. other appendages absent 65 At least some setae with an internal structure of small chambers FLABELLIGERIDAE (part) No setae chambered 66 With a papilla between the rami of each parapodium. 78a (77b). segments not prolonged SABELLONGIDAE Body separated into two regions with different kinds of setae in a thoracic and abdominal region (NOTE: Regions may sometimes also be definable on parapodial features) 71 Body not separated into regions. 66b 67a 67b 68a 68b 69a (54b). (73b). (74b). bushy or simple filaments ARENICOLIDAE Anterior end with complex jaw-apparatus 74 Jaw-apparatus absent 76 Each jaw consisting of a series of small denticles DORVILLEIDAE Each jaw consisting of a single piece 75 Hooded hooks present in at least some setigers. segments usually elongated (bamboo-worms) MALDANIDAE Setae include bilimbate and spatulate kinds. branchiae non-retractable. branchiae. limited to the anterior end and strongly arborescent. prostomium entire. setae composite 89 Without tentacular cirri. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 84b (83b). setae simple 90 Setal appendages long and slender. pelagic lOSPILIDAE Setal appendages short. All setae simple. may be bluntly conical. as wide as the widest part of the body MAGELX)NIDAE I*rostomium not flattened. The orbiniids have been the subject of several comprehensive studies. branchiae cirriform. easily dehiscent cirri. bifid or furcate setae. simple hooks and sometimes brush-topped. (90b). parapodia either all uniramous or all biramous GLYCERIDAE (part) Eversible pharynx with more than four jaws. including capillary setae and usually acicular spines and serrated or spinose setae. setae limited to lateral tufts 93 All parapodia biramous. (91b). prostomium T-shaped or bifid SCALIBREGMIDAE Prostomium with a median antenna 86 Prostomium without appendages 88 Branchiae present on maximally 15-20 segments first starting from one of setigers 4-10 PARAONIDAE (part) Branchial distribution otherwise or branchiae absent 87 Notopodial cirri flask-shaped in some setigers. (88a). 85b (58b). no peristomial appendages. (79b). except possibly the first one 94 Median parapodia uniramous TROCHOCHAETIDAE Uncini absent. parasitic CALAMYZIDAE With multiple series of small. branchiae. 86a (85a). Prostomium without appendages. abdominal segments prolonged with setae forming complete cinctures around the body HETEROSPIONIDAE Parapodia well developed. plumose setae absent SPIONIDAE (part) With two pairs of tentacular cirri. (82b). (91b). abdominal segments usually not prolonged. FAMILY ORBINIIDAE HARTMAN 1942 Orbiniida with lateral parapodia in a thoracic region and. 80b (79a). plumose setae present POECILOCHAETIDAE Notopodial cirri cirriform or folióse. one or two asetigerous segments present anteriorly. (82b). (88a). A saclike proboscis present. button-shaped projections. if present. (93b). eversible pharynx either an axial sac or a ventrolateral pad. notopodia with acicula. rounded or truncate. never articulated 81 Eversible pharynx with four jaws. pelagic TYPHLOSCOLECIDAE Parapodia low folds or blunt. all setae simple. no additional cephalized segments present. 86b 87a 87b 88a 88b 89a 89b 90a 90b 91a 91b 92a 92b 93a 93b 94a 94b (85a). Pettibone (1957a) and Day (1973).14 79b (78b). (81b). pectinate or absent. (86b). (85b). . (85b). if present. distinctly narrower than the widest part of the body 92 Parapodia inconspicuous. The major generic groupings appear clear and were reviewed by Day (1973) for the subfamily Orbiniinae. cirriform 82 Composite falcigers present in anterior setigers ACROCIRRIDAE (part) All setae simple 83 Neurosetae uncinal in structure BOGUEIDAE Uncini absent 84 AH setae simple capillaries. (83b). The subfamily Protoariciinae has yet to be reviewed in detail. (93b). including Hartman (1957). at least in the neuropodia. (92b). 80a (79a). maximally two asetigerous anterior segments present. dorsal parapodia in an abdominal region. pointed or rounded OPHELIIDAE Furcate and acicular setae usually present. but without setae. Palps absent. notopodia with setae in addition to acicula. Prostomium less than twice as long as wide. (86b). long-shafted uncini in the neutopodia OWENIIDAE (part) Uncini in single rows or absent 91 Prostomium flattened and spatulate. (90b). including capillaries. posterior ones biramous GONIADIDAE Parapodia rounded lobes with large. usually. 81a 81b 82a 82b 83a 83b 84a (79b). 85a (58b). anterior parapodia uniramous. (88b). true branchiae absent APISTOBRANCHIDAE Uncini normally present. Parapodia biramous. branchiae present or absent SPIONIDAE (part) ORDER ORBINIIDA Prostomium without appendages. (92b). (81b). (88b). cirri. 7a (lb). about 22x. diagrammatic. 6b (5b). (D). 5a (4b). KEY TO GENERA la. 7b (lb). Family PARAONIDAE. 3a (2b). Two asetigerous anterior segments A single asetigerous anterior segment Branchiae absent Branchiae present Branchiae present on all but a few anterior segments Branchiae limited to abdominal segments Only crenulated capillaries present Crenulated capillaries and other kinds of setae present Abdominal neurosetae include swan-shaped hooks Swan-shaped hooks absent Abdominal neurosetae all crenulated capillaries. lb. Cirrophorus sp. 9b (8b). 4b (3b). 18x. Quesia caudicirra. 2a (la). 3b (2b). Family QUESTIDAE. 54x. 24x. 5b (4b). prostomium rounded Abdominal neurosetae include acicular hooks. after Hartman 1966.1976 THE POLYCHAETE WORMS 15 FIGURE 2. (E) anterior end of the above. transverse section of the abdomen of the above. 4a (3b). anterior end. (A). 10a (9a). 5x. 6a (5b). prostomium pointed Branchiae absent Branchiae present on at least some segments Prostomium rounded or truncate Prostomium more or less pointed All thoracic parapodia with only slender. Family ORBINHDAE. after Hartman 1969.. 9a (8b). (c). pointed setae Some thoracic neuropodia with setae of another kind Abdominal neuropodia with acicular spines 2 7 Orbiniella 3 Protoariciella 4 Scoloplella 5 Proscoloplos 6 Protoaricia Schroederella Microrbinia 8 Naineris 9 10 11 Berkeleya . (B). Orhinia johnsoni. 8a (7b). 2b (la). anterior end. 8b (7b). posterior seta. Thoracic . never more than four papillae combined. Haploscoloplos Monro 1933. 13b (I lb). PROTOARICIINAE with rounded prostomium and stout branchiae present from setiger 8. Abdominal notosetae crenulated capillaries and sometimes furcate setae. Proscoloplos Day 1954. projecting from the parapodial lobes. Separation between thorax and abdomen indistinct. O. other notosetae camerated capillaries and posterior notosetae curved and serrated acicular spines. Abdominal notopodia with crenulated capillaries and furcate setae. 14a (13b). branchiae first present from any segment from 2-23. Neurosetae similar. ORBINIINAE with conical prostomium. PROTOARICIINAE with rounded prostomium and branchiae limited to abdominal segments. PROTOARICIINAE with rounded or pointed prostomium and without branchiae. Thoracic neurosetae include crenulated capillaries. Thoracic setae all camerated or crenulated capillaries. papillae sometimes also on parapodial postsetal ridges Orbinia Without rows of papillae on the ventrum 14 Median and posterior abdominal neuropodia with thick acicula. . Protoaricia Czerniavsky 1881a. Ai. Orbinia Quatrefages 1865. Phylo Kinberg 1866b. posterior thoracic neuropodia with two papillae and maximally two ventral papillae present. 14b (13b). heavy hooks and heavy spearshaped setae. Thoracic neurosetae include crenulated capillaries. Transition from thorax to abdomen indistinct. Phylo Some thoracic segments with rows of papillae along the ventrum. 2 species. Thoracic neurosetae include hooks (or subuluncini) and crenulated capillaries. minuta Day 1954. Scoloplos cylindrifer Ehlers 1905. cygnochaetus Day 1954. Twelve thoracic segments. ORBINIINAE with rounded or truncate prostomium. Aricia oerstedii Claparède 1864. 12a (1 la). 4 species. ventral papillae absent. Thoracic neurosetae include anterior brush-topped setae. with a combined total of at least five papillae on each segment. 13a (lib). felix Kinberg 1866b. 1 lb (10b). transition from thorax to abdomen abrupt. profunda Hartman 1971. 16 species. only species. ORBINIINAE with pointed prostomium. Maximally two accessory papillae on posterior parapodia. acicular setae and sometimes furcate setae. I2b (11a). Orbiniella Day 1954. Nais quadricuspida Fabricius 1780. Posterior thoracic parapodia with several accessory papillae. Microrbinia Hartman 1965. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Acicular spines absent in abdominal neuropodia Haploscoloplos Thoracic neuropodia of two abruptly different kinds 12 Thoracic neuropodia not abruptly different 13 Anterior three thoracic neuropodia with bristle-tipped setae Califia Posterior thoracic neuropodia with thick. P. First notosetae smooth capillaries. 19 species. C. linea Hartman 1965. numerous ventral papillae present. Furcate and capillary setae present in abdominal notopodia. branchiae and accessory papillae absent. candida Hartman 1957. branchiae not present before setiger 9. only species. crenulated capillaries and in some cases blunt hooks. Posterior thoracic parapodia with several accessory papillae and numerous ventral papillae. branchiae not present before segment 8. P.16 10b (9a). abdominal setae include crenulated capillaries. posterior thoracic neuropodia with two or fewer papillae. branchiae usually present from fifth or sixth segment Scoloplos (Leodamas) Thick projecting acicula absent. 13 species. abdominal neuropodia with crenulated capillaries and acicular spines. abdominal branchiae and all thoracic setae crenulated capillaries. branchiae usually not present before tenth segment (Scoloplos) Generic Definitions Berkeleya Hartman 1971. Aricia cuvierii Audouin and Milne Edwards 1833c. 4 species. hooks and intermediate forms. notosetae crenulated capillaries and in some cases furcate setae. ventral papillae absent. modified spines associated with a glandular pouch . 2 species. Naineris Blainville 1828. ORBINIINAE with pointed prostomium. 1 la (10b). At least five papillae in combined total present on a segment. ORBINIINAE with pointed prostomium and first pair of branchiae on setiger 5-9. Califia Hartman 1957. Setae include crenulated capillaries and one or two swan-shaped hooks in posterior neuropodia. All neurosetae crenulated capillaries. ORBINIINAE with pointed prostomium. 18 species. ORBINIINAE with pointed prostomium and branchiae first present from setiger 5-7. B. hooks and subuluncini. Lumbricus armiger 0. Subgenus Scoloplos: Species with first branchiae on setigers 8-10 or later. Branchiae present on a limited number of median setigers in most species. F. FAMILY PARAONIDAE CERRUTI 1909 Body long and slender with lateral parapodia. capensis Day 1963. Thoracic neurosetae include blunt hooks and crenulated capillaries (may be absent). indeterminable Gisela Müller 1858. Prostomium a simple triangular lobe. most neurosetae similar. Subgenus Leodamas: Species with first branchiae on setiger 5-6. these subgenera are here considered distinct genera. Abdominal notosetae include crenulated capillaries. see Orbinia Incertae Sedis Falklandiella Hartman 1967. Strelzov recognized only two genera with prostomial antennae. Alcandra Kinberg 1866b. in addition heavy acicular spines present in posterior neuropodia. 3 species. see Naineris Naidonereis Malmgren 1867. only species. 3b (2b). the latter with a series of subgenera. furcate setae and spines. Müller 1776. with heavy emerging notacicula in posterior setigers. crenulated capillaries and thick smooth spines in posterior segments. Neurosetae include thick hooks with three teeth.F. Schroederella Laubier 1962. 2b (lb). acicular setae present in posterior notopodia. PROTOARICIINAE with pointed prostomium and branchiae present on the abdomen. sec Scoloplos (Leodamas) Anthostoma Schmarda 1861. All setae crenulated capillaries. see Naineris Theostoma Eisig 1914. seeOrbiitia Branchetus Chamberlin 1919c. hooks or otherwise modified setae. Key to Genera la. pauliana Laubier 1962. Thoracic notosctae all are crenulated capillaries. uncinata Hartraann-Schiöder 1962. PROTOARICIINAE with pointed prostomium and branchiae present on all but a few anterior segments (from setiger 3 in type species). Prostomium with a single antenna or antennae absent. Setae present 6om the first segment Aparaonis Setae present from the second segment 2 Modified setae notopodial Cirrophorus Modified setae. 4a (3a). The key given below is based on the important revision by Strelzov (1973). 4b (3a). P. ORBINIINAE with pointed prostomium and branchiae first present from setiger 5 or later. Scoloplella Day 1963. annulata Hartman 1967.1976 THE POLYCHAETE WORMS 77 Invalid Genera neurosetae include crenulated capillaries. usually postbranchial. All setae simple. Resembles the orbiniids in setal structures. Notosetae all slender and camerated. 5a (4b). including capillaries and various. only species. but appears to differ in other features. 3a (2b). error for Naineris Theodisca Müller 1858. two asetigerous segments present. S. S. s&e Protoaricia Venadis Castelnau 1842. seeNaineris Aricia Savigny 1820. but shorter. slender acicular setae with flat teeth. 2 species. without emerging notacicula in posterior setigers. Parapodia biramous with very strongly reduced par^odial lobes. Cirrophorus and Aricidea. some features are taken from Fauchald (1972). PROTOARCIINAE with strongly pointed prostomium and branchiae present on the abdomen. indeterminable Labotas Kinberg 1866b. see Scoloplos (Leodamas) Clytie Grube 1855. Scoloplos Blainville 1828. 38 species. abdominal neurosetae all crenulated capillaries. Protoariciella Hartmann-Schröder 1962. including both ventral and parapodial kinds. Accessory papillae never exceeding four. lb. if present. 2a (lb). neuropodial 3 Prostomium with a median antenna 4 Prostomium without a median antenna 7 Modified setae absent Aedicira Modified setae present 5 Modified setae either pseudocomposite or curved with a subterminal arista on concave side of the shaft Aricidea . see Scoloplos Lacydes Kinberg 1866b. Notosetae include crenulated capillaries and acicular spines in far posterior segments. Neurosetae include crenulated capillaries and acicular hooks. Taxonomic Notes The generic subdivision given above. Median antenna present. Aonides fulgens Levinsen 1884.¡8 5b (4b). Invalid Genus Levinsenia Mesnil 1897. Strelzov (1973) introduced several new taxonomic characters to the ones previously in common usage. such as Paraonides. only species. Nuchal organs peristomial. Nuchal organs prostomial. 4 species. Until the value of Strelzov's system has been tested. modified setae present in the postbranchial notopodial fascicles. may be invalid. A. 8a (7b). More than three prebranchial setigers present. either with numbers of long aristae or bearded . 10 species. three pairs of branchiae on segments 2-4. or tapering abruptly near the middle. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Modified setae otherwise 6 Modified setae not greatly different from the capillary setae. 2 species. Median antenna present. terminal prostomial sense organs absent Sabidius More than three prebranchial setigers. Cirrophorus Ehlers 1908. modified setae either pseudocomposite or recurved with a subterminal arista arising from the concave side of the shaft. Modified setae neuropodial.. median antenna absent. or bifid with several aristae or with a short stiff beard. giving the outer part of the seta the appearance of a long. Modified setae neuropodial. 7 species. 7a (2b). 6a (5b). 13 species. smooth arista. 6b (5b). 10 species. Aricidea Webster 1879b.. Paraonella Strelzov 1973. terminal prostomial sense organs present Tauberia Generic Definitions Acesta Strelzov 1973. Aedicira Hartman 1957. Nuchal organs on the peristomium. with a single preoral ciliary band or prostomial ciliary bands absent 8 Modified setae absent Paraonella Modified setae present 9 Three prebranchial setigers. only species. of prostomial terminal sense organs and the localization of the nuchal organs must now be investigated. Paraonis Cerruti 1909. only species. median antenna absent. Several other genera. fragilis Webster 1879b. Prostomial terminal sense organs present. One asetigerous anterior segment present. Median antenna present. somewhat more abruptly tapering. those in the neuropodial fascicles often shorter than the notopodial ones. Pharynx with a muscular pad. C. but somewhat thicker and more abruptly tapering and with a long arista Allia Modified setae distally strongly curved. 8b (7b). Tauberia Strelzov 1973. median antenna absent. Aricidea catherinae Laubier 1967a. 9b (7b). A. Setae present in the first segment. median antenna absent. branchiatus Ehlers 1908. Paraonis cornatus Hartman 1965. Branchiae and anal cirri present or . The presence of prostomial ciliary bands. Modified setae neuropodial. each strongly curved distally. Aricidea albairossae Pettibone 1957b. Prostomial terminal sense organs absent. All setae simple capillaries.. or bifid. Paradoneis and others used in the traditional sub-division of the family. Paraonides nórdica Strelzov 1968a. Aricidea pacifica Hartman 1944c. Similar characters have been used in other famihes. abyssalis Hartman 1965. and it is anticipated that they will gain acceptance also in the treatment of the paraonids. see Paraonis. prostomium with several transverse bands of cilia Paraonis Nuchal organ on the reduced peristomium. All setae slender capillaries. I hesitate to treat these genera as invalid. Modified setae slightly thicker than the non-modified capillary setae. Modified setae neuropodial. differs markedly from the more traditional pattern summarized by Fauchald (1972). FAMILY QCIESTIDAE HARMAN 1966a Body long and slender. Parapodia biramous with reduced lobes. Sabidius Strelzov 1973. Aparaonis Hartman 1965. Modified setae absent. 9a (8b). 7b (2b). Median antenna present or absent. Prostomium without appendages. Aonides gracilis Tauber 1879. Acesta Nuchal organ on posterior part of prostomium. Three prebranchial setigers. Nuchal organs peristomial. Allia Strelzov 1973. 10 species. Median antenna present. 1976 THE POLYCHAETE WORMS \9 absent. only species. Body short and gniblike. Pharynx a ventral muscular pad. prostomium without appendages. Questids with serrated capillary setae and bidentate hooks. some segments with median dorsal cirri. posterior cirriform branchiae present Setae include capillaries. unpaired dorsal cirri on one or two anterior segments or larval branchiae present RHAPHIDRILINAE 2 . Key to Genera la. Branchiae present on posterior segments. It is quite possible that the family should be considered among the oligochaetes. this generally is considered the key feature separating the oligochaetes from the polychaetes (Fauchald 1974a). only species. caudicirra Hartman 1966a. FAMILY CTENODRILIDAE KENNEL Quesxa Novaquesta (1971). This small family of tiny polychaetes recently has been reorganized and redefined by Hartmann-Schröder FIGURE 3. However. Key to Genera la. bidentate hooks and trifúrcate spines. Stygocapitella subterránea. branchiae and anal cirri absent. 1882 Body short or long. Some anterior segments may be fused and asetigerous. lb. At least one asetigerous anterior segment present. after HanmannSchröder 1971. unpaired dorsal cirri and branchiae absent CTENODRILINAE Ctenodrilus Body long and slender. Setae include serrated capillaries. ORDER CTENODRILIDA Prostomium without appendages. about 30Ox. Setae include capillaries and bidentate hooks. Family CTENODRILIDAE. Family PARERGODRILIDAE. Ctenodrilus serratus. generally grub-shaped polychaetes that tend to turn up in massculture in aquaria more frequently than in the field (especially true for the ctenodrilids). Specimens of Ctenodrilus also have turned up associated with FlabelUderma commensalis at Santa Catalina Island and have the same dark purple pigmentation on sea urchins as these commensals do. all setae simple. Parapodia uni. bidentate hooks and trifúrcate spines. paired lateral branchiae present in some juveniles. 284x. palps absent. with at least 15 segments.or biramous. a feature rarely found among the oligochaetes. (B). Questa Harman 1966a. Parapodial lobes absent. Members of this order are very small. the questids have several different kinds of setae. anal cirri present. with few segments. As remarked by Hobson ( 1970) the questids have the gonads Umited to a few segments. Proboscis a ventral muscular pad. Setae in double fascicles. (A). each with only a few species. branchiae absent Generic Definitions Novaquesta Hobson 1970. trifitrcata Hobson 1970. Questids with serrated capillary setae. modified after Hartman 1944. palps absent. N. lb. it now is divided into two sub-families with three genera in all. parapodial lobes absent. Q. bidentate hooks and trifúrcate spines with the middle tine shorter and slenderer than the other. Parapodia biramous. Pro. Two anterior and two posterior asetigerous segments. Body with maximally 15 segments. branchiae absent Unpaired dorsal cirri absent. Family PSAMMODRILIDAE. lb. palps absent. whiplike tips and short furcate setae with two accessory teeth in the crotch. The family is poorly known and understood. Rkapkidrilus Monticelli 1910. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Unpaired dorsal cirri present. 2 species. Setae include capillaries and genital spines in the adult males.Cossurabrmnea. Invalid Genera Monostylos Vejdovsky 1884. only species. 12. neuro|X)dial tori present in one form. Ctenodrilus monostylos Zeppelin 1883. smooth bilimbate setae with long. otherwise parapodial lobes absent (except for the cirri mentioned above). Key to Genera la. P. with at least 18 segments. subterránea KnöUner 1934. Notopodia in a median region strongly prolonged and supported by acicula.20 2a (lb). eight or nine setigers present. Parapodia. One or two unpaired dorsal cirri present anteriorly. marginally coarsely dentate or smooth. Stygocapitella Knöllner 1934. keideri Reisinger 1925. parapodial lobes absent. Setae thick. grub-shaped worms with at least one anterior asetigerous segment. Uving either interstitially in shallow marine sands (Stygocapitella) or in rotting terrestrial plant material (Parergodrilus). ten or 11 setigers present. FAMILY PSAMMODRILIDAE SWEDMARK Stygocapitella Parergodrilus t 4 1952 FiGimE 4. (B). nemasoma Monticelli 1910. Setae all simple. 5 species. ORDER PSAMMODRILIDA Prostomium without appendages. short bilimbate setae. Prostomium without appendages. Setae include smooth. except . CTENODRILINAE. Setae include capillaries and short. Psammodriloides fauveli after Swedmark 1958. except copulatory hooks in males. Body not translucent. afterFauchald 1972. (A). one achaetous segment present. RHAPHIDRILINAE. 8-9 setigers present. Small. Body long and slender. RHAPHIDRILINAE. with 10-11 setigers. Body long and slender. At least one asetigerous anterior segment present. only species. The parergodrilids are very small worms. setae present in single fascicles. only species. grub-shaped worms. Family CO%S\JRJDk'E. thicker dentate or smooth spines. One anterior and no posterior asetigerous segments. see Zeppelinia Partkenope Schmidt 1857. all setae smooth rods Generic Definitions Parergodrilus Reisinger 1925. Setae all smooth rods. Partkenope cirrata Schmidt 1857. branchiae present Generic Definitions Science Series 28 Zeppelinia Raphidrilus Ctenodrilus Claparède 1863. about 50x. Paired branchiae present in the juvenile stages.5x. one asetigerous segment present. the revision by Hartmann-Schröder (1971) is followed here. all setae simple. Ä. S. palps absent. each with one or a few setae only. 2b (lb).and peristomium without appendages. asetigerous segments absent. with at least IS segments. Zeppelinia Vaillant 1890. set Ctenodrilus FAMILY PARERGODRILIDAE REISINGER 1960 Small. setae limbate and furcate Body translucent. Apistobranchus tullbergi. Body in three regions. The only currently described genus is Cossura Webster and Benedict. 25x. each with several uncini in each neuropodium. proboscis a ventral pad. up to about thirty abdominal setigers. dorsal view of the above. only species. a single median palp present on the dorsum of an anterior setiger (usually between setigers 3 and 6). Parapodia biramous. lb. Pharyngeal apparatus present. ORDER SPIONIDA Prostomium distinct. P. modified after Hartman 1969. only species. (B). dorsal view of the above. A pair or two groups of grooved feeding palps usually present anteriorly. lOx. Two of these differ rather sharply from the others and may deserve separate generic standing. each with a dorsal cirrus supported by an aciculum. head. lateral view. abdominal neuropodial tori present in one form. each with a single uncinus in each neuropodium. Cossurids are common in sand and especially in deep slope and abyssal muds. Parapodia biramous. pharyngeal region and trunk. ventral view. Psammodrilus Swedmark 1952. these small interstitial worms are very isolated among the polychaetes. abdominal uncinigers with several uncini Psammodrilus Dorsal cirri increasing in length from first to fourth segment. with very low parapodial lobes. single uncini present in abdominal neuropodia Psammodriloides Generic Definitions Psammodriloides Swedmark 1958. six thoracic setigers. Body with two distinct regions: head and trunk. (C). Pharynx either axial or as a ventral pad. ORDER COSSURIDA Prostomium without appendages. All setae simple. All setae simple. Key to Genera la. Dorsal cirri decreasing evenly in length from the first to the sixth thoracic segment. (1887) with genotype C. Boccardia proboscidea. As remarked by Swedmark (1958). Parapodia well developed or reduced. . Psantmodritus balanoglossoides Swedmark. Six thoracic setigers with dorsal cirri supported by acicula. 1963 Prostomium without appendages. Paired tentacular cirri sometimes present at the base of the palps. and apparently feed on detritus with the help of the pharynx. either on the prostomium or on an anterior segment. Proboscis a ventral pad.]97() THE POLYCHAETE WORMS 2/ the thoracal dorsal cirri reduced. The palp in this case appears to be sensory. P. Trochochaetus multisetosum. Family APISTOBRANCHIDAE. without appendages or with an occipital antenna. lOx. including bilimbate or hirsute setae in two or more fascicles. (E). They are burrowers. balanogtossoides Swedmark 1952. lOx. a single peristomial asetigerous segment present. fauveli Swedmark 1958. and perhaps may additionally be respiratory in function since it is well-equipped with blood-vessels (Fauchald in preparation). (D). FAMILY COSSURIDAE DAY rata Webster and Benedict (1887) with a total of about fifteen species. lOx. thick spines and capillary setae present in the abdomen of some forms. longocir- FIGURE 5. after Pettibone 1963. Family SPIONIDAE. is unique in that the eversible pharynx is formed from the longitudinal body muscles. Ten abdominal setigers. Eversible pharynx absent. Family TROCHOCHAETIDAE. (A). with reduced parapodial lobes. a single median palp present dorsally on one anterior setiger. after Pettibone 1963. The first form described. those of segments five and six very short. all setae simple. 3a (2a). This is especially true in those genera where the structure of the branchiae is important for species separation. eversible pharynx either a ventral pad or an axial sac. see Apistobranchus Spioniforms with body elongated. Parapodia biramous. hooded or non-hooded hooks. lb. but with palps attached at the postectal margins. Key to Genera la. The apistobranchids are known for one genus. lb. never serrated. 2b (lb). 7b (6b). 6b (5b). or pointed. FAMILY APISTOBRANCHIDAE MESNIL AND CAULLERY 1898 Spioniforms with the anterior end inflated and the rest of the body cylindrical. mainly capillaries. they are. Most commonly found in shelly sands. as some of their spionid relatives. 2a (lb). 4b (3b). Attempts at identification of these taxa on incomplete material should be avoided. parapodial lobes mostly cirriform. 7a (6b).and peristomium. others build permanent tubes in soft substrata and some forms combine the two activities. Species of some genera. Numerous forms are freeliving in sands and muds. 5b (4b). 3b (2a). Apistobranchids are in fact not as rare as the low number of species should indicate. not limited to a tubicolous existence.and peristomium. Body separated into at least two distinct regions. 4a (3b). Some forms are burrowers (Polydora. uncini present Chaetopterifonnia Body rarely separated into distinct regions. Members of the genus Aonides are dominant forms in sandy beaches in temperate and warm areas of the world (Foster 1971). including capillaries and bi. Palps at the postectal comers of the prostomium. All setae simple. All setae simple. Apistobranchus Levinsen 1883. but will feed from loosely constructed burrows. 5a (4b). Each branchia with several (maximally seven) processes in median parapodia Potybranchia Branchia. Prostoraium without appendages. but serrated postsetal lobes present in some setigers. 8a (2b). sands and muds. if present. uncini always absent 2 Palps at the junction of the pro. no segment with remarkably different setae (except large hooks in the first setiger) 8 Setiger 16 modified Morants Either setiger 4 or 5 modified 4 Setiger 4 modified Polydorella Setiger 5 modified 5 Neuropodial uncini distally trifid Tripolydora Neuropodial uncini distally bifid or entire 6 Branchiae first present from setiger 2 Boccardia Branchiae first present posterior to the modified segment 7 Setae in modified segment arranged in a horseshoe-shaped series Pseudopolydora Setae in the modified segment in a straight line or a small patch Polydora Branchiae absent. with frontal horns. 2a (lb). Spionids are very common in all environments.22 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Key to Suborders Science Series 28 la. Prostomium anteriorly blunt. Boccardia et al. the family is speciose and serious errors are easUy introduced into the literature. most frequently perhaps of Prionospio SENSU LAW and of the PolydoralBoccardia complex. with type species Aricia tullbergi Theel 1879. parapodial lobes poorly developed Cirratuliformia Suborder Spioniformia FAMILY SPIONIDAE GRUBE 1850 Palps on the peristomium or at the junction between the pro. parapodial lobes well developed at least in some setigers Spioniformia Palps post-peristomial in origin.) in calcareous substrata or in rock. and frequently most of their branchiae in fixation. 2b (lb). first neuropodia with stout hooks Spiophanes . A total of three species currently are recognized. if care is not taken at all identificatoiy levels. see Apistobranchus Skardaria Wesenberg-Lund 1951. an occipital papilla may be present. 6a (5b).or multidentate. other appendages absent. very likely are present in any kind of benthic marine soft-bottom sample taken anywhere on the globe. single process 2 One setiger with strongly modified setae 3 Setae change gradually along the body. parapodial lobes cirriform or folióse. Invalid Genera Ethocles Webster and Benedict 1887. The spionids usually lose their palps. Parapodia biramous. Nerine oxycephala Sars 1862. 9b (8b). 19b (9b). (22b). 9a (8b). 5 species. Anterior parapodial lobes very large and folióse. A. Four pairs of branchiae from setiger 2. 14a (10b). Neuropodial postsetal lobes smooth. where an additional single pair is present on the second setiger Pygospio Branchiae concentrated near the anterior end only 21 Peristomium forms large lateral wings on either side of the prostomium 22 Peristomium does not form lateral wings on the sides of the prostomium 24 Three pairs of pinnate branchiae present from setiger I Paraprionospio Branchiae otherwise. (22b). (26b). All anterior setae capillaries. first neuropodia without stout hooks 9 Branchiae present on nearly the whole body 10 Branchiae limited to less than half the length of the body 19 Branchiae present from setiger 1 11 Branchiae present from setiger 2 14 Accessory branchiae on some segments Dispio Accessory branchiae absent 12 Posterior notopodia with uncini Marenzelleria Notosetae all capillaries 13 Prostomium anteriorly rounded Spio Prostomium with laterofrontal horns Malacoceros Posterior notopodia with uncini Scolelepis All notosetae capillaries 15 Prostomium with laterofrontal horns 16 Prostomium anteriorly blunt or pointed 17 Branchiae partially fused to the notopodial postsetal lobes Rhynchospio Branchiae completely separated from the notopodial postsetal lobes Mesospio Occipital antenna absent Microspio Occipital antenna present 18 Branchiae partially fused to the notopodial postsetal lobes Pseudomalacoceros Branchiae completely separated from the notopodial postsetal lobes Laonice A single pair of branchiae present Streblospio At least two pairs of branchiae present 20 Branchiae concentrated in a medio-posterior region. 18b (17b). Prionospio sexoculata Augener 1918. 16b (15a). 17a (15b). (21a). only species. neuropodia with uncini from setigers 10-11. 17b ( 15b). 15a ( 14b). present on anterior end only. I5b (14b). (20b). (24b). first three cirriform. 18a (17b). (26b). 19a (9b).]g7(. 16a (15a). Branchiae from setiger 2. fourth pinnate. Prostomium anteriorly pointed. 10a (9a). Notosetae all capillaries (in 36 setigers). 13b (12b). 12b (1 lb). 20b 21a 21b 22a 22b 23a 23b 24a 24b 25a 25b 26a 26b 27a Tib (19b). (24b). peristomium without lateral wings. 10b (9a). Prostomium anteriorly rounded. Two pairs of branchiae on setigers 3 and 4. except in the males. 2 species. (25b). 12a ( 1 lb). Aquilaspio Foster 1971. free from the notopodial postsetal lobes. the last pinnate Apoprionospio Branchiae cirriform or pinnate in another pattern Prionospio free from the notopodial postsetai lobes. (21a). THE POLYCHAETE WORMS 2i At least one pair of branchiae present. posterior parapodia with uncini in both rami. posteriorly uncini in both rami. (21b). 8b (2b). Neuropodial postsetal lobes large and folióse in setiger 2. completely . Apoprionospio Foster 1969. (20b). Prostomium anteriorly blunt. (25b). 20a (19b). Anterior setigers with capillary setae only. day i Foster 1969. Generic Definitions Anaspio Chamberlin 1920. (21b). 14b (10b). A. first present from setiger 2 23 Two to four pairs of branchiae present Aquilaspio At least six pairs of branchiae present Minuspio Prostomium anteriorly pointed Aonides Prostomium anteriorly blunt or with frontal homs 25 Prostomium with frontal homs Scolecolepides Prostomium anteriorly rounded 26 Branchiae two pairs on setigers 3•4 Anaspio Four pairs of branchiae from setiger 2 27 Three first pairs of branchiae cirriform. lib (10a). Chamberlin 1920. 1 la ( 10a). 7 species. boreus. 13a (12b). Aonides Claparède \%(A. neurosetae include capillaries and uncini in posterior setigers. Malacoceros Quatrefages 1843. Prostomium anteriorly blunt or bifid. Mesospio Gravier 1911a. first setiger more or less reduced. 36 species. First four setigers with branchiae. stout setae. in other setigers. Prionospio Malmgren 1867. 15 species. posteriorly uncini in both rami. Notosetae all capillaries. Branchiae present from the second setiger. neuropodia with uncini present from setiger 15. Branchiae fiwm setiger 6. occipital antenna present. Prostomium with frontal horns. Branchial parapodia with large postsetal lobes. but with uncini present from setiger 6. either cirriform or pinnate or both. occipital antenna present. Notosetae all capillaries. Prostomium anteriorly rounded. proliféra Augener 1914. Polydora (Leucodore) polybranchia Haswell 1885. Other setae include notopodial capillaries. fused to the notopodial postsetal lobes. only species. 10 species. Two to four pairs of branchiae from setiger 2. all setae uncmi m both rami. moorei Gravier 1911a. Branchiae present from setiger 2. Fifth setiger modified with strong setae. posterior ones uncini. Nerine cirrata Sars 1850. only species. Ihtjstomium anteriorly blunt. D. each branchia with maximally six to seven processes in median setigers. Capillary setae absent. All anterior setae capillaries. Laonice Malmgren 1867. Branchiae present from setiger 1. Four pairs of branchiae. Curved hooks present in the first notopodia. Setiger 5 with strongly modified. M. Microspio Mesnil 1896. Prostomium anteriorly blunt or bifid. short and tapering. Prostomium anteriorly blunt or pointed. only species. peristomium without lateral wings. all notopodia with capillaries only. Pofydorella Augener 1914. Three pairs of pinnate branchiae present from setiger 1 . Notosetae all capillaries. neuropodia with capillaries. P. neuropodia posteriorly with uncini. in median and posterior parapodia uncini also present. wireni Augener 1913b. posterior end with uncini in both rami. setiger 16 with strongly modified setae. M. posterior parapodia with imcini in both rami. All anterior setae capillaries. All anterior setae capillaries. Pseudomalacoceros Czemiavsky 1881b. P. Marenzelleria Mesnil 1896. stout setae. cornuta Bosc 1802. Anterior neurosetae capillary. All anal cirri similar. neurosetae include capillaries and uncini. Dispio Hartman 1951a. foxi Potts 1928. parapodia of fu^t setiger well developed. Notosetae all capillaries. . posterior setigers with uncini in both rami. all cirriform. Genital pouches in at least some setigers. Branchiae present from setiger 2. at least partially fused to the notopodial postsetal lobes. partially fused to the notopodial postsetal lobes. Branchiae not present before setiger 6. 15 species. accessory branchiae present in some setigers. Boccardia Carazzi 1895. 11 species. peristomium with large lateral wings. sometimes also with simple posterior spines. completely free from the notopodial postsetal lobes. Second segment asetigerous. Prionospio pinnala Ehlers 1901. Branchiae present irom setiger 2. first present from setiger 2. 7 species. Nerinides cántabra Rioja 1918. Prostomium with lateral horns. P. Prostomium anteriorly blunt. Branchia present from setiger 1. Polydora Bosc 1802. Prostomium anteriorly rounded. steenstriipi Makngren 1867. Prostomium anteriorly rounded. duplex Chamberlin 1919a. Branchiae present from setiger 2 to the middle of the body. Prostomium anteriorly rounded. 5 species. peristomium with large lateral wings. M. Anterior setigers with capillary setae in both rami. at least partially fused to the notopodial postsetal lobes. Prostomium with frontal horns. 10 species. 65 species. Prostomium anteriorly blunt. except in setiger 4. Morants Chamberlin 1919a. Branchiae present from setiger 1. Spio mecznikowianus Claparède 1870a. peristomium enfolding sides of prostomium. partially fused to the notopodial postsetal lobes. completely free of the postsetal lobes. all other notosetae capillary. Notosetae all capillaries. Spio vulgaris Johnston 1827. Anterior setae all capillaries in both rami. Minuspio Foster 1971. Neutopodial uncini present fiom setiger 7-10 in most species. Prostomium anteriorly blunt. Fourth setiger modified with large. 5 species. peristomium with large lateral wings. only species. Branchiae from setiger 1 to the end of the body. Poly bronchia Potts 1928. P. Paraprionospio CauUery 1914b. neuropodia posteriorly also with uncini. partially fused to the notopodial postsetal lobes. 2 species. Notosetae all capillaries. Prionospio cirrifera Wiièn 1883.24 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Prostomium anteriorly blunt. Notosetae all capillaries. neurosetae include capillaries and uncini. Branchiae present from setiger 2 to about setiger 40. neurosetae include capillaries and uncini in median and posterior setigers. uncinata Hartman 1951a. spinosa Woodwick 1964. Branchiae absent. partially fiised to the notopodial (Ktstsetal lobes. prostomiimi otherwise without appendages.g. Prostomium with frontal horns. S. Polydora antennata Claparède 1870a./976 THE POLYCHAETE WORMS 25 Pseudopolydora Czerniavsky 1881b. T. posterior setae include capillaries and uncini in both rami. Paired spioniform palps present. Spio Fabricius 1785. Hannerz (1956) and Söderström (1920) reviewed the major taxa. Taxonomic Note The above survey of the genera follows in the main Foster's (1971) survey of the fauna fixsm the Gulf of Mexico. Prostomium anteriorly rounded. From the number of species listed for each genus. rather than saclike pharynx. benedicti Webster 1879a. neurosetae anteriorly all capillaries. The structure of the proboscis may not be a very important feature in this group as noted by Orrhage (1966). Branchiae present fix)m setiger 1. A.F. R. Fifth setiger modified. Tripolydora Woodwick 1964. with a few modified setae only. Nereis filicornis O. Dorsal and ventral cirri and small branchiae jwesent. Anterior setae all capillaries. S. Lumbrkus squamatus O. except in the males where a single pair is present on the second setiger. rarely blunt. Setiger 2 with a dorsal collar between the notopodia. The family is one of the better studied. and best luiderstood families. xt Scolelepis Aonopsis Wagner 1885. neurosetae include capillaries and uncini. Spiophanes Grube 1860. arenincola Hartman 1936b. The total absence of uncmi may not be particularly surprising. Notosetae all capillaries. Scolelepis BlainvUle 1828. Incertae Sedis Aberranta Hartman 1965. Pharynx muscularized. 3 species. until it can be better investigated. The biology of the more common forms also has been rather well studied. rather than use some of them as subgenera. all others with bkamous parapodia. Branchiae present from setiger 1. kroeyeri Grube 1860. only species. Blake 1971). except that I have preferred to list all the taxa at the generic level. larval forms Prostomium conical. occipital antenna present. neurosetae include capillaries and uncini. 4 species. all are fused to the notopodial postsetal lobes. Branchiae concentrated in a short posterior region. According to Hartman (1%5) this differs from the true spionids in that it has a muscular. see Pseudopolydora Chaetosphaera Haecker 1896. First neuropodia with stout hooks. sec Ironice Bilobaria Sveshnikov 1959. at least anteriorly fused with the notopodial postsetal lobes. eyes absent.F. 3 species. S. all other anterior setae capillaries. Branchiae first present from setiger 7. I^stomium usually pointed. All setae simple capillaries. One apodous segment present. partially fused to the notopodial postsetal lobes. 15 species. In addition to Poster's extensive review cited above. A single pair of branchiae on setiger 1. it further differs from the spionids in that it completely lacks hooks of any kind. All notosetae capillaries. Pygospio Claparède 1863. especially the reproductive biology of members of Polydora and Boccardia has been the subject of several studies by Blake (e. neurosetae include capillaries and uncini. Rhynchospio Hartman 1936b. 2 species. Prostomium blunt or with frontal horns. However. 20 species. in that one of the specimens described was ovigerous. only species. indeterminable Aricideopsis Johnson 1901. Prostomium anteriorly rounded. This is not a larval form. posterior parapodia with uncini and capillaries in both rami. posterior neuropodia with uncini. benhami Ehlers 1907. Notosetae all capillaries. enigmática Hartman 1965. Müller 1776. it is considered best to leave the genus as a fnee-standing genus. 16 species. Prostomium with frontal horns. . neurosetae include capillaries and posterior uncini. Invalid Genera Aonis Audouin and Milne Edwards 1833c. Streblospio Webster 1879a. it will be noted that I have been rather more conservative than Foster in retaining species. Branchiae present from setiger 2. limited to the anterior part of the body. Anterior setae all capillaries. Ftestomium anteriorly rounded. neurosetae include capillaries and trifid uncini first present from setiger 9. considering how many spionid genera may lack such hooks in either of the two rami. 11 species. P. Notosetae all capillaries. Scolecolepides Ehlers 1907. Notosetae all capillaries. or over long stretches of the body. Notosetae all capillaries. Branchiae present from setiger 2. Fifth setiger modified with large. Prostomium anteriorly blunt or bifid. thick setae arranged in a horseshoe-shaped pattern. Müller 1806. posteriorly also uncini. ekgans Claperède 1863. Branchiae present fiom setiger 2. Neuropodial jxjstsetal lobes notched at least in posterior setigers. larval forms Carazzia Mesnil 1896. Dr. indetemiinable Pseudonerine Augener 1926. hypothetical Protopolydora Czemiavsky 1881b. Prostomium^flattened and anteriorly ovate or truncate. seePrionospio Diplotis Montagu 1815. see Spio Kinbergella Mclntosh 1909. in Quatrefages 1865. NOMEN NUDUM Spionides Webster and Benedict 1887. separated into two regions. see Polydora Pseudonerine Czemiavsky 1881b. larval fomi Rhynophylla Carrington 1865. with type species M. see Polydora Euspio Mclntosh 1915. Magelonids are common in sandy bottoms. see Polydora Leucodora Johnston 1838. they are rather characteristic with the flattened. see Prionospio Pygophyllum Schmarda 1861. see Pseudomalacoceros Nerinopsis Ehlers 1912. indeterminable Neopygospio Berkeley and Berkeley 1954. fringed setae and straight spines.and peristomium on the ventral side. see Polydora Dipolydora Verrill 1881. larval form Paranerine Czemiavsky 1881. see Scolelepis Nerinides Mesnil 1896. see Laonice Uncinia Quatrefages 1865. indeteraiinabie {IPolydora) Colobranchus Schmarda 1861. papillicornis Müller 1858 and a total of about 35 described species. (A). Setae include capillaries and hooded bi. California. Trochochaeta Levinsen 1884 with type species Disoma multisetosum Örsted 1844 and a total of nine recognized species. Magelona Müller 1858. see Scolelepis Pteriptyches Grube 1872a. diagram of dorsal view. Family MAGELONIDAE. ftostomium a flattened ridge with palps present at the postectal comers. Invalid Genera Maea Johnston 1865. indeterminable Scolecolepis Malmgren 1867. shovel-shaped prostomium often much wider than the rest of the animal. sec Magelona Papillaria Sveshnikov 1959. indeterminable Spionereis Sars 1853. Magelonids build only very flimsy tube-structures and tend to move through the sediment. 25x. Palps at the junction of the pro. off Santa Barbara. set Scolelepis Ctenospio Sars 1867. Magelonids currently are assigned to a single genus. seeAonides Perialia Kinberg 1866. see Polydora Pseudokucodore Czemiavsky 1881b. Magelona sp.26 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Cheimnotus Costa 1861b. without appendages. see Pseudopolydora Nerine Johnston 1838. First parapodia directed forwards along the sides of the prostomium. questionably Boccarrfia Prospio Mesnil 1896. (B). slender bodies. mis-spelling of Scolelepis Spione Örsted 1844. Pettibone (1963) correctly pointed out that the . see Streblospio Heterospio Czemiavsky 1881b. see Polydora Mandarte Kinberg 1866b.or multidentate hooks. 25x. RGURE FAMILY MAGELONIDAE CUNNINGHAM AND RAMAGE 1888 Spioniforms with long. see Prionospio Leipoceras Möbius 1874. Setae capillaries. Meredith L. Jones of the Smithsonian Institution is revising the family. ventral view of the above. The family is recognized for a single genus. which tends to be rather threadlike with very long segments. acicular falcate spines. seeSpio Hekaterobranchus Buchanan 1890. sen Magelona FAMILY TROCHOCHAETIDAE PETTIBONE 1963 Body flattened cylindrical. It is anticipated that more taxa may be recognized when this revision is completed. see Scolelepis 6.. originally applied to these animals by Örsted (1844). mirabilis Laubier and Ramos 1973. Kalaminochaeta and Kaluntmaria both based on pelagic larval forms. Setae in nearly complete cinctures around the abdomen. Santa Barbara. slender bodies. Palps present at the postectal comers. Neuropodial acicular spines are present in most setigers from segment 4.1976 THE POLYCHAETE WORMS 27 name Disoma. but some neurosetae are thickened and gently falcate. slender non-tubicolous spioniforms mainly reported from shallow water in soft substrates. 1956 Spioniforms with long. was used ten years earlier by Ehrenberg for a protozoan. 1941 was described for two genera. Long. P. The family and both genera are here considered invalid. First segment with long setae forming a cephalic cage and second setiger similar to all following segments. First segment is asetigerous and first setiger lacks dorsal and ventral cirri. indeterminable Disoma Örsted 1844. off Loon Point. most simple capillaries. First segment with or without setae. Poecilochaetids with a median antenna. Poecilockaetus johnsoni. see Trochochaeta Pilearia Sveshnikov 1963. Trochochaetids are long. Neuropodial acicular spines present in a few anterior segments only. with lateral sense-organs between FIGURE 7. larval forms Thaumastoma Webster and Benedict 1884. with one or two pairs of tentacular cirri. longissima Ehlers 1874 and for a total of four species. Key to Genera Antenna frontal. Poecilochaetids with a frontal antenna. Prostomium small with either a frontal or a median antenna. filiform branchiae on the thoracic setigers. possibly of a Trochochaeta. Parapodia biramous. 35 m. 25x. Taxonomic Notes Setae are present from the second segment in Elicodasia and from the first in Poecilochaetus. palps attached at the pwstectal comers. see Trochochaeta The family Kalaminochaetidae Nolte. Setae include capillaries. 12 species. They appear to be nowhere truly numerous. see Trochochaeta Disomides Chamberlin 1919c. California. but frequently are reported from areas where quantitative benthic investigations have been undertaken. FAMILY POECILOCHAETIDAE HANNERZ rami. see Trochochaeta Nevaya Mclntosh 1911. only species. FAMILY HETEROSPIONIDAE HARTMAN 1963a Body with short thoracic and greatly prolonged abdominal setigers. plumose and acicular setae. pectinate. Family POECILOCHAETIDAE. fulgoris Claparède in Ehlers 1875. Heterospio Ehlers 1874 with type species H. Dorsal and ventral cirri spindle or bottleshaped . . without nuchal organ. first segment asetigerous Poecilochaetus Elicodasia Generic Definitions Elicodasia Laubier and Ramos 1973. E. but not identifiable. first segment with series of long setae Antenna median. Prostomium blunt without appendages. The family is known for a single genus. Invalid Genera Cherusca Müller 1858. Poecilochaetus Claparède 1875. nuchal organs present. Chaetopterus variopedatus. neuropodia with pectinate uncini. antennae present. Peristomium may have one or two pairs of tentacular cirri. Pectiniform uncini present in posterior setigers. taylori Potts 1914. only species. Mesochaetopterus Potts 1914. Median notopodia all simple. pharynx a simple. Setae include capillary. P. anterior region with uniramous parapodia with lancet-shaped setae. 80x. all notopodia of median region asetigerous. which has stout spines. M. antennae absent. FIGURE 8. neuropodia with pectinate uncini. bilobed and asetigerous. Median notopodia foliaceous.28 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Heterospionids have been reported from widely scattered areas. Tricoelia variopedatus Renier 1804. neuropodia with pectiniform uncini. Uncini present. Median notopodia bilobed Median notopodia never bilobed. some posterior notopodia fused to form dorsal fans. Median region with biramous parapodia. FAMILY CHAETOPTERIDAE MALMGREN 1867 Body with two or three distinct body-regions. S. median neuropodia with uncini. except setiger 4. palps very short Notopodia never fused. mainly in deep shelf and slope sediments. 2a. see Heterospio Suborder Chaetopteriformia Short palps on prostomium. Family CHAETOPTERIDAE. 3b. typicus Sars 1856. neuropodia with pectinate uncini. median and posterior regions with biramous parapodia. palps of varying lengths always present. (A). posterior region with pointed notopodia supported by internal acicula. . 2b. Invalid Genus Longosoma Hartman 1944c. supported by internal acicula. neuropodia with pectinate uncini. They are rather fragile and non-descript and thus easily overlooked if quantitative studies are not undertaken. 3a. Phyllochaetopterus Grube 1863. 12 species. neuropodia with pectinate uncini. posterior region with pointed notopodia with internal acicula. posterior region with pointed notopodia. Spiochaetopterus Sars 1853. posterior region with long. palps large Some median notopodia fused to form dorsal fans. palps large 2 3 Phyllochaetopterus Spiochaetopterus Chaetopterus Mesochaetopterus (la) (la) (lb) (lb) Generic Definitions Chaetopterus Cuvier 1827. 17 species. but no setae. Tharyx moniloceras. modified from various sources. Body with three regions. Median notopodia bilobed and foliaceous. gracilis Grube 1863. 11 species. lb. Body divided in three distinct regions. (B). neuropodia with uncini. Anterior region with uniramous parapodia. limbate setae and modified spines in setiger 4. noneversible tube. natural size. Key to Genera la. asetigerous. Body with three distinct body-regions. antennae absent. the family is poorly known and the several species may be considerably more common than their scattered records indicate. after Hartman. 1969. Family CIRRATULIDAE. Body with three distinct regions. pointed notopodia with a few contained acicula. may be fused A pair of small tentacular cirri present at the base of the large palps Tentacular cirri absent. falcate spines Pseudocirratulus At least some capillary setae present 6 Anterior region without long tentacular structures Raricirrus One or more anterior segments with groups of long tentacular cirri or branchiae 7 Dorsal tentacular cirri first present posterior to the anteriormost branchiae Cirriformia Dorsal tentacular cirri first present from the same segment as the anteriormost branchiae 8 Tentacular cirri on one segment only Cirratulus Tentacular cirri on two or more segments Timareie Generic Definitions Caulleriella Chamberlin 1919c. 16 species. Heterospio catalinensis. 1969. . Setae include capillaries and acicular spines that are usually entire distally. Cirratulus Lamarck 1801. lb. Acrocirris fromifilis. Chaetozone Malmgren 1867. Acrocirris crassifiUs.F. 25 species. see Spiochaetopterus Tricoelia Renier 1804. after Banse. or a group of grooved palps on one or several post-peristomial segments. Müller 1776. 19 species. Paired palps inserted dorsally anterior to. 5b (lb). 2b (la). A pair of long grooved palps attached on the anterior dorsum 2 Two groups of grooved tentacular cirri present 5 All setae slender. (B). I2x. Parapodia reduced. body usually light colored 4 Acicular spines in posterior segments distally entire Chaetozone Acicular spines in posterior segments distally bi. FAMILY CIRRATULIDAE CARUS 1863 FIGURE 9. body usually dark green or brown Dodecaceria Acicular setae not excavate. eversible pharynx a thick ventral pad. 7a (6b). Paired palps inserted dorsally anterior to. setosa Malmgren 1867. 4a (3b). Prostomium without appendages. 4b (3b). tentacular cirri present on one segment only. after Banse. 6b (5b). or at the first setiger.1976 Invalid Genera THE POLYCHAETE WORMS 29 Leptochaetopterus Berkeley 1927. 3b (2b). 6a (5b). Family ACROCIRRIDAE. anteriormost branchiae present from the same segment. see Chaetopterus Suborder Cirratuliformia One pair of grooved palps. (c). Cirratulus bioculatus Keferstein 1862. 1969. see Spiochaetoplerus Mesotrocka Leuckart and Pagenstecher 1855. lateral view. 3a (2b).or multifid Caulleriella All setae acicular. (A). see Mesockaelopterus Ranzanides Chamberlin 1919c. 8a (7b). Body cylindrical. or at the first setiger. curved spines. 2a (la). 8b (7b). Cirratulids without palps. modified from Hartman. about 12x. 7b (6b). Key to Genera 1 a. Family HETEROSPIONIDAE. distally pointed Tharyx At least some setae either curved hooks or spines 3 Acicular setae distally excavate. see Phyllochaetopterus Ranzania Claparède 1870b. about 5x. Prostomium conical or blunt. C. All setae simple including capillaries and curved or excavate hooks. slender filiform or clávate branchiae present on at least some setigers. Setae include capillaries and distally entire. see Mesochaetopterus Telepsavus Costa 1861a. 5a (lb). peristomium fused with at least two segments. Lumbricus cirratus O. 1944c. Setae include capillaries and distally bidentate or multidentate curved spines. see Timarete Promenia Kinberg 1866b. 5 species. see Cirriformia Cirratulispio Mclntosh 1911. neurosetae compound hooded falcigers. Terebella tentaculata Montagu 1808. the genus Flabelligella known from several deep-water locations was transferred to this family (Orensanz 1974b). notosetae segmented and spinose. but sometimes with serrated cutting edges. 2a (lb). Setae include capillaries and curved. tentacular cirri present on one segment only. entire spines or hooks. papillata Hartman 1965. anteriormost branchiae present anterior to the tentacular cirri. 1 b. and even more recently. see Caulleriella and in part Tharyx Labranda Kinberg 1866b. Posterior part of body often flattened and expanded. Setae all gently curved. The TAaryJc-complex is presently rather confused. see Cirratulus FAMILY ACROCIRRIDAE BANSE 1969 Body either slender and elongate. Anterior setigers with capillary setae. epithelium nearly smooth Flabelligella 2 Macrochaeta Acrocirrus Generic Definitions Acrocirrus Grube 1872b. D. Parapodia biramous with small parapodia. The acrocirrids recently were recognized as a separate family.30 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Taxonomic Notes Science Series 28 Cirriformia Hartman 1936a. 22 species. and additional characters will have to be included to clarify the relations between the several species described in this genus. distally entire spines. Setae include capillaries in the notopodia and neuropodia and curved hooks with dentate cutting edges. Prostomium blunt. F. 8 species. Branchiae absent Branchiae present Palpal bases well separated. Timarete Kinberg 1866b. R. indeterminable Heterocirrus Grube 1855. Branchiae present on some anterior segments. Dodecaceria Örsted 1843b. T. paired palps usually present. Acrocirrids with palps originating close together. concharum Örsted 1843b. i&e. mostly capillary and smooth. Caulleriella Monticellirux Laubier 1961a. 8 species. Thorax of 12 segments. then abruptly broad at seliger 9 and tapering from there to the posterior end. Raricirrus Hartman 1961. maculatus Hartman 1961. see Cirratulus Audouinia Quatrefages 1865. Heterocirrus fi-ontifilis Grube I860. 26 species. branchiae present from the same segment as the first tentacular cirri. filiform or clávate. kingstonensis Augener 1922. acutus Webster and Benedict 1887. branchiae present on a few segments. see Cirriformia Mesocirrineris Czerniavsky 1881b. a row of papillae ventral to the neurosetae in all setigers. Several segments usually crowded near the anterior end. Flabelligella Hartman 1965. Pseudocirratulus Augener 1922. Invalid Genera Ambo Chamberiin 1918. tentacular cirri present on at least two anterior setigers. Tharyx Webster and Benedict 1887. a papilla on third segment below the gill. see Dodecaceria Pentacirrus Wesenberg-Lund 1958. cirri and branchiae present on most segments. see Timarete Archidice Kinberg 1866b. Palps absent. separation . 2b (lb). or short and maggot-shaped. Body anteriorly narrow. Acrocirrids without branchiae and palps. questionably Cirratulus or Chaetozone Cirrhineris Blainville 1818. Four pairs of gills. only species. It appears likely that the generic division of the palpate cirratulids will change with future revisions. All setae slender. two achaetous anterior segments present. setiger 13 with a pair of papillae. see Tharyx Naraganseta Leidy 1855. Palps present. posterior neuroopodia with gently curved spines. 16 species. these segments usually asetigerous. Palps present anterior to or at the first stiger. $&e Dodecaceria Heterocirrus Saint-Joseph 1894. Key to Genera la. Palps absent. Cirratulus anchylochaetus Schmarda 1861. P. epithelium densely papillated Palpal bases nearly abutting. Cirratulids without palps. only species. Setae include notopodial capillaries and anteriorly neuropodial capillaries. median and posterior ones also with excavate stout spines. Macrochaeta Grube 1850. ten thoracic segments Genital spines absent. Noiomasms lenuis. Number of thoracal segments variable. Invalid Genus Ledon Webster and Benedict 1887. 4a (lb). 2b (la). usually slender. neuropodia long transverse welts in at least some setigers. after Hartman. Faraiiy CAPlTEhLlDAB. All parapodia biramous. about natural sire. 3b (2b). Parapodia biramous. genera and species cannot be identified without this information. thorax with capillary setae and abdomen with rostrate hooded hooks. 1947. 4b (lb). The characteristic rostrate uncini are small. Prostomium short and truncate. 1862 Body cylindrical. hooded hook of the above.1976 THE POLYCHAETE WORMS i] between thorax and abdomen indistinct. generally. including capillaries and rostrate uncini. Thorax and abdomen defined on internal structures and parapodial development. near marina. 6 species. (C). The anterior thorax of most capitellids is usually strongly areolated and they are usually reddish pink or brown. such as Capitella capitala are considered pollution indicators. 3a (2b). either ten or thirteen thoracic segments present Setigers 7-8 with genital spines. Capitellids are among the most commonly encountered polychaetes. All setae simple.. However. The rostrate uncini look truncate in this sort of observation. 5b (4a). Family ARENICOLIDAE. trying to observe the setae as the light catches them. Usually four pairs of branchiae. without appendages. Three first setigers with capillary setae only At least four setigers with capillary setae only Some thoracic segments with flanged hooks. )i50x. 2a (la). twelve thoracic segments present Flanged hooks absent. FAMILY CAPITELLIDAE GRUBE difficult. 5a (4a). Body-surface densely papillated. modified from several sources. some forms. they are perhaps the most earthwormlike of the common marine polychaetes. epidermis usually densely papillated. in that they are capable of invading areas where disastrous defaunation of natural or man-made causes has taken place (Grassle and Grassle 1974). lb. One or two anterior asetigerous segments. 7x.4rert(fo/fl. ^t& Macrochaeta ORDER CAPITELLIDA Prostomium without appendages. and the capillaries will retain their tapered appearance. In general appearance. 13 thoracic segments Four first setigers with capillary setae only At least five anterior setigers with capillary setae Genital spines present Genital spines absent 2 4 Peresiella 3 Capitomastus 9 Heteromastides 5 8 6 7 . 1 have found it useful to rotate the specimen while counting the segments. Anterior segments uniramous in some species. palps absent. making the determination of the first occurrence of these setae 10. Nais clavicomis Sars 1835. Acrocirrids with bases of palps separated by at least the width of the palps. FIGURE Key to Genera la. (b). (A). 22a (21b). . (23a).and neuropodia. 29b (28b). 27b (23b). 28a (27a). ( 19a). branchiae absent Parheteromastides Six anterior setigers with capillary setae only 11 At least seven anterior setigers with capillary setae 13 Genital spines in setigers 8-9 Capitellides Genital spines absent ¡2 Thorax with 11 segments Neomediomastus Thorax with 12 segments Barantolla Seven anterior setigers with capillary setae only 14 At least nine anterior setigers with capillary setae only 17 Genital spines present 15 Genital spines absent 16 Notopodial cirriform branchiae on some abdominal segments Branchiocapitella Branchiae absent Capitella Thorax with eight segments. branchiae present Heteromastus First setiger with notopodia only. (20b). twelve thoracic segments present Paraleiocapitella Eleven anterior setigers with capillary setae only 24 Twelve or more anterior setigers with capillary setae only 27 At least some capillary setae in the first two abdominal setigers 25 All abdominal segments with hooks only 26 First setiger with neuropodia only. notopodial acicula absent. 17b (13b). 26b (24b). first two abdominal setigers with hooks in the neuropodia and mixed hooks and capillary setae in the notopodia Mastobranchus Abdominal notopodial tori nearly coalesce. I3a (10b). 6b (5a). lia (10a). all thoracic setigers with capillary setae only LeiocapitelUdes Thorax with 12 segments. Pulliella Ten first setigers with capillary setae only 20 At least 11 anterior setigers with capillary setae 23 First setiger with both noto. (17b). NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 21b (20b). 10b (8b). 14b (13a). 13b (10b). notopodial hooks present Notomastus Twelve setigers with capillary setae only 28 Thirteen or more setigers with capillary setae only 30 Anal plaque with imbedded spines present Scyphoproctus Anal end rounded or with anal cirri 29 Thirteenth setiger with both hooks and capillary setae Leiochrus No setiger with both hooks and capillary setae present Leiochrides Thirteen anterior setigers with capillary setae only 31 At least 15 anterior setigers with capillary setae 35 . (17b). notopodia with acicula. 9b (8a). 30a (27b). 16b (14b). first two abdominal setigers with capillary setae in both rami Notodasus First setiger with both noto. 7a (5b). (19a).and neuropodia. 18b 19a 19b 20a 20b 21a ( 17a). 30b (27b). thorax with eleven segments Neonotomastus Last thoracic segment with both hooks and capillary setae. 29a (28b). 23a 23b 24a 24b 25a (19b). 12a (11 b). 26a (24b). thorax with ten segments Pseudoleiocapitella Anterior asetigerous segment absent. 27a (23b). (24a). mixed segments absent Capitellethus At least one segment with mixed hooks and capillary setae 22 Two first abdominal segments with mixed hooks and capillary setae. 22b (21b). 25b (24a). thorax with nine segments . 15a (14a). 7b (5b). 10a (8b). posterior thoracic setigers with hooks Neoheteromastus Nine anterior setigers with capillary setae only 18 Ten or more anterior setigers with capillary setae 19 One anterior asetigerous segment present.32 6a (5a). all abdominal setigers with hooks only. notopodial hooks absent Rashgua Abdominal notopodial tori well separated.and neuropodia Decamastus First setiger with notopodium only 21 All thoracic setigers with capillary setae. 28b (27a). (19b). 12b (lib). . 8a (4b). nine thoracic segments present Capitella (part) Thorax with 11 segments Mediomastus Thorax with 12 segments Parheteromastus Five setigers with capillary setae 9 At least six anterior setigers with capillary setae only 10 First setiger with both noto. ten thoracic segments present Capitomastus 6 Genital spines in setigers 8-9. 8b (4b). I lb (10a). 17a (13b). 15b (14a). 16a (14b). (23a). 18a (17a). 14a (13a). Genital spines in setigers 7-8. 9a (8a). first setiger with notopodia only. CapitelUdes Mesnil 1897b. 10 species. Up to segment 17 or 18 capillary setae only. 2 species. Branchiae absent. Branchiocapitella Fauve I 1932.1976 31a (30a). THE POLYCHAETE WORMS ii 35b (30b). one segment with both capillary setae and hooks present Leiocapitella Thorax with 15 to 18 segments. segments 18 and 19 transitional with notopodial setae and neuropodia with mixed hooks and setae. Lumbricus capitatus Fabricius 1780. 37a (36b). D. Thorax with nine segments.and neuropodia Dasybranchetus First setiger with notopodia only 37 Twenty setigers with capillary setae Lumbricomastus Seventeen or 18 setigers with capillary setae only Anotomastus Generic Definitions Anotomastus Hartman 1947. Capitobranchus Day 1962. next three with hooks. then three segments with mixed hooks and setae in both rami. maximally two transitional segments present 36 First setiger with both noto. segment 8 with mixed setae and hooks in notopodia and hooks in neuropodia. first setiger with notopodia only. BarantoUa Southern 1921. singularis Fauvel 1932. macgregori Day 1962. (33b). 31b (30a). 33a 33b 34a 34b 35a (31b). Dasybranchetus Monro 1931. additional four segments have both setae and hooks in the notopodia Eunotomastus Thoracic count otherwise. Capitella minima Langerhans 1881. (30b). sculpta Southern 1921. 36a (35b). one anterior asetigerous segment present. Branchiae may be present in posterior segments. B. first setiger complete. 3 species. Females: up to segment 4 with capillary setae. then genital spines in segments 8-9. notopodial 34 All notopodial setae capillary Protomastobranchus Abdominal notopodial setae hooks Dasybranchus At least 16 setigers with capillary setae only. 2 species. CapitelUdes dispar Ehlers 1907. capillary setae only. 37b (37b). branchiae. . if present. Thorax with 16 segments. Branchiae absent. asetigerous segment absent. Notopodial cirriform branchiae present. Thorax with 12 segments.and neuropodia 33 Thorax with 14 or 15 segments. one anterior asetigerous segment present. first setiger complete. up to segment 7 with capillary setae only. first setiger complete. only species. one asetigerous segment present. all thoracic setigers with capillary setae. branchiae absent. First seliger with notopodia only 32 First setiger with both noto. Males: up to segment 5 with capillary setae only. First six setigers with capillary setae only. Eunotomastus gordioides Moore 1909. the next three segments with hooks and segments 8-9 with genital spines. Up to segment 7. Dasybranchus Grube 1850. which has genital spines. Neuropodial branchiae present in posterior setigers. 32a (31a). 3 species. asetigerous segment absent. first setiger complete. Thorax with 19 segments. Capitella Blainville 1828. B. (33b). first setiger complete. the next two segments with hooks. first setiger complete. one asetigerous segment present. branchiae neuropodial Capitobranchus Thorax with maximally 15 segments. (31b). C. giardi Mesnil 1897b. All thoracic setigers with capillary setae only. Capitellethus Chamberlin 1919c. Either four first segments with capillary setae only. fauveli Monro 1931. Palmately branched branchiae in posterior segments. Thorax with ten segments. segments 8-9 with genital spines. Thorax with nine segments. then segments 8-9 with genital spines. Thorax with 18 or 19 segments. or first seven segments with capillary setae only. Capitomastus Eisig 1887. segments 8-9 with genital spines. asetigerous segment absent. next four segments mixed with capillary setae in the notopodia and hooks in the neuropodia. only species. Up to segment 15 with capillary setae only. first setiger complete. Thorax with nine segments. Thorax with 11 segments. 32b (31a). except notopodia 8-9. 3 species. asetigerous segment absent. three or four segments with both capillary setae and hooks present Pseudocapitella Thorax with 19 segments. 36b (35b). one asetigerous segment present. only species. 15 species. C. Branchiae not seen. Dasymallus caducus Grube 1846. one asetigerous segment present. Branchiae not seen. first setiger complete. Thorax with 13 segments. branchiae absent. one asetigerous segment present. one asetigerous segment present. M. one asetigerous segment present. approximately 20 thoracic segments present. first setiger complete. capillary setae only. glabra Hartman 1947. 2 species. Branchiae absent. N. Thorax with 13 segments. Branchiae not seen. californiensis Hartman 1944b. Thorax with 12 segments. Branchiae absent. Genus poorly defined. all hooks. Mastobranchus Eisig 1887. first abdominal with notopodial capillary setae and neuropodial hooks. L. first setiger complete. one asetigerous segment present. first seliger complete. segment 14 with mixed setae and hooks in both rami. N. Thorax with 12 segments. 7 species. Thorax with 21 segments. Capitella filiformis Claparède 1864. D. Thorax with 12 segments. lineus Hartman 1960. Leiochrides Augener 1914. Up to segment 4. first setiger complete. first abdominal notopodia with capillary setae. glabrus Fauchald 1972. Up to segment 8 with capillary setae only. one asetigerous segment present. one asetigerous segment present. Notodasus Fauchald 1972. tulearensis Thomassin 1970. all thoracic setigers with capillary setae only. Neoheteromastus Hartman 1960. Branchiae absent. Thorax with eight segments. L. Leiochrus Ehlers 1908. australis Augener 1914. only species. one asetigerous segment present. all thoracic setigers with capillary setae only. Up to segment 6. E. two first abdominal segments with mixed capillary setae and hooks in the notopodia and hooks in the neuropodia. one asetigerous segment present. first setiger with notopodia only. from segment 7. 4 species. First abdominal neuropodia with mixed hooks and capillary setae. magnus Fauchald 1972. Thorax with 12 segments. first setiger with notopodia only. one asetigerous segment present. one asetigerous segment . capillary setae only. Up to segment 13. Notomastus Sars 1850. thereafter. Lumbricomastus Thomassin 1970. analis Hartmann-Schröder 1960a. Mediomastus Hartman 1944b. All thoracic setigers with capillary setae only. Leiocapitella Hartman ¡947. Heteromastus Eisig 1887. Branchiae not seen. Notopodial branchiae present. only species. Thorax with 11 segments. Thorax with 14 or 15 segments. first setiger complete. Thorax with 11 segments. 34 species. Up to segment 14 with capillary setae only. Thorax with 11 segments. segment 5 with mixed capillary setae and hooks in both rami. only species. first setiger with notopodia only. Small notopodial branchial processes present in far posterior setigers. alutaceus Ehlers 1908. first setiger complete. segment 15 with capillary setae in the notopodia and hooks in the neuropodia. 7 species. from segment 6. 2 species. Leiocapitellides Hartmann-Schröder 1960a. one asetigerous segment present. hooks only. L. All thoracic setigers with capillary setae only. N. Mediomastus glabrus Hartman 1960. Thorax with 13 or 14 segments. Neomediomastus Hartman 1969. alt thoracic segments with capillary setae only. Simple or composite notopodial branchiae present. first setiger with neuropodia only. trinchesii Eisig 1887. first setiger with notopodia only. all setae hooks. 2 species. Neonoiomastus Fauchald 1972. L. All thoracic setigers with capillary setae only. one asetigerous segment present. segment 9 with capillary setae in the notopodia and hooks in the neuropodia. L. Notopodial branchiae present. only species. 2 species. only species. Up to segment 5 with capillary setae only. of which 16 have capillary setae only and the next four have mixed setae and hooks in the notopodial fascicles. first setiger complete. Heteromastides Augener 1914. only species. capillary setae only. one asetigerous segment present. gracilis Hartman 1963a. Decamastus Hartman 1963a. second with mixed hooks and capillary setae. N. Eunotomastus Mclntosh 1885. only species. one asetigerous segment present. bifidus Augener 1914. Thorax with 11 segments. Branchiae not seen. grübet Mclntosh 1885. H. second with hooks only. 7 species. Branchiae absent. All thoracic setigers with capillary setae only. Up to segment 7 with capillary setae only. Thorax with 12 segments. Branchial processes on posterior notopodia. then all setigers with hooks. one asetigerous segment present.34 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Thorax with 14 segments. latericeus Sars 1850. first setiger complete. All thoracic setigers and the first two abdominal setigers with capillary setae only. Branchiae absent. M. such as the current one. one asetigerous segment present. Thorax with ten segments. is considered here a synonym of Mediomastus Hartman 1944b. a table has been constructed to indicate in a different manner the relationships between the genera. all with capillary setae. one asetigerous segment present. A review only. first setiger with notopodia only. 2 species. cannot solve the problem of the number of valid capitelHd genera. lack notopodial hooks. probably of the genus Dasybranchus as first suggested by Gallardo (1968). segment 7 with both capillary setae and hooks and neuropodia with hooks only. Paraleiocapitella Thomassin 1970. To make a review somewhat easier. All thoracic setigers with capillary setae only. only species. capillary setae only. which is easily enough seen. Up to segment 4. next three with notopodial capillary setae and neuropodial hooks. first se tiger complete. present. Branchiae absent. Up to segment 15. Branchiae absent. Thorax with 13 segments. first setiger complete. Pseudoleiocapitella Harmelin 1964. first setiger with notopodia only. Thorax with 13 or 14 segments. one asetigerous segment present. fauveli Harmelin 1964. huloti Gallardo 1968. Branchiae absent. multiocuiatus Hartmann-Schröder 1962a. from there on. First setiger complete. Scyphoproctus Gravier 1904. Simple branchiae present. but has been based on the number of segments with capillary setae. Peresiella Harmelin 1968. capillary setae only. hooks. Up to segment 10. Expanded imbedded spines present. mossambica Thomassin 1970. P. All notopodia with limbate capillary setae only. which may be difficult to see. first setiger with notopodia only. P. segments and setigers. clymenoides Harmelin 1968. Protomastobranchus Gallardo 1968. Parheteromastus Monro 1937a. capillary setae only. first setiger complete. the generic identification of capitellids is at best difficult. capillary setae only. only species. P. first setiger complete. up to segment 13 or 14. Thorax with 12 segments. be kept apart. one asetigerous segment present. first setiger complete. Pseudocapitella Fauvel 1913. Branchiae absent. Invalid Genera Ancistria Quatrefages 1865. The key to genera follows the traditional system.]976 THE POLYCHAETE WORMS 35 present. rubrocinaa Wesenberg-Lund 1949. Pulliella Fauvel 1929. It is of the utmost importance that the two concepts. ise Heteromastus Arenia Quartrefages 1865. incerta Fauvel 1913. only species. 2 species. P. Thorax with 11 segments. Thorax with nine segments. P. Parheteromastides Hartmann-Schröder 1962a. only species. and on the distribution of the capillary and hooked setae. tenuis Monto 1937a. P. first setiger with notopodia only. Up to segment 6. which must be based on the variability of all observable characters in a large amount of material. S. Thorax with 12 segments. capillary setae in neuropodia also. Branchiae absent. all other segments with hooks only. Thorax with 12 segments. s^s Dasybranchus Bucherta Rullier 1965a. djiboutensis Gravier one asetigerous segment All thoracic setigers with anal plaque with acicular Taxonomic Notes The generic sub-division of the capitellids is unsatisfactory. questionably//eííromajfujBranchoscolex Schmarda 1861. see Dasybranchus . as first suggested by Hartmann-Schröder (1962a). one asetigerous segment present. Up to segment II with capillary setae only. first two abdominal segments with notopodial capillary setae and neuropodial hooks. The genus Bucherta Rullier (1965a) is considered as the posterior end of a capitellid. Dorsal abdominal tori nearly coalesce. one asetigerous segment present. notopodial acicula present. Branchiae may be present. 1904. armata Fauvel 1929. since both are used in key features of the capitellids. P. rather than on the number of thoracic segments. P. P. The genus Capitita Hartman 1947. As stated above. one asetigerous segment present. segment 12 with notopodial capillary setae and neuropodial hooks. the remainder of thorax with modified. but poorly understood in terms of variability. Up to segment 5. capillary setae only. flanged hooks in at least some segments. se& Noiomasius Areniella Verrill 1874. capillary setae and normal hooks in the others. Thorax with 12 segments. Thorax with 15 to 18 segments. Branchiae absent. only species. only species. 7 species. capillary setae only. Rashgua Wesenberg-Lund 1949. since the former is the more easily observed character. it is based on the number of thoracic segments. all thoracic setigers with capillary setae. one asetigerous segment present. 2 species. compl notop. of setigers with mixed capillaries and rostrate uncini Column 7: First segment complete or incomplete Column 8: EYesence of branchiae and special kinds of setae. No. of thoracic segments present Column 2: Presence or absence of an anterior asetigerous segment Column 3. compl notop. notop. no branchiae no branchiae notacicula près no branchiae no branchiae no branchiae branchiae no branchiae no branchiae no branchiae no branchiae no branchiae small knobs no branchiae ?no branchiae ?no branchiae Hanged setae no branchiae branchiae ± branchiae Tbranchiae no branchiae ± branchiae no notouncini branchiae branchiae ?branchiae no branchiae 7branchiae anal plaque no te-anchiae no notouncini branchiae no la-anchiae no branchiae ?branchiae branchiae neurop. . see Notomastus Isomastus Gravier 191 la. compl compl neuro compl compl compl compl compl compl notop. compl notop. notop. compl notop. 6 7 compl compl compl compl compl compl compl compl compl compl notop. see Capitellethus Dasymallus Grube 1846.36 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Capitellides Ehlers 1907. in Grube 1850. branchiae branchiae 8 + _ + + + + + + + _ 8-9 8-9 8-9 8-9 8-9 8-9 _ 3 _ 2 _ _ _ _ _ _ _ _ - _ 1 _ 2 var. see Capitella Lumbriconais Örsted 1842. Airanged in order of increasing number of thoracic segments present. compl notop. compl compl compl notop. set Notomastus Valla Johnston 1865. see Capitella TABLE 2 Genera of Capitellidae. Column 1: No. notop. stt Capitella Sandanis Kinberg 1867b. + + + + + _ 1 1 1 + + + 7 10 11 11 11 13 3 12 12 12 13-14 13 13 14 15 17-18 14 20 + + + + + _ 2 1 1 + + + + + + + + + + 1 3 1 4 - The genus Eunotomastus is too incompletely known to be included in the table. of setigers with capillary setae Column 6: No. see Dasybranchus Ditrocha Sveshnikov 1959. of thoracic setigers Column 4: Essence of genital spines Genus Leiocapitellides Puiiiella Capitella (1) Capitellides Capitella (2) Branchiocapitella Capitomastus 9 Capitomastus 6 Pseudoleiocapitella Mediomastus Parheteromastides Neomediomastus Capitellethus Decamastus Neonotomastus Peresiella Parheteromastus Heteromastus BarantoUa Neoheteromastus Paraleiocapitella Notomastus Rashgua Mastobranchus Notodasus Heteromastides Leiochrides Scyphoproctus Leiocfirus Protomastobranchus Dasybranchus Leiocapitella Pseudocapitetla Dasybrancketus Anotomastus Capilobranchus Lumbricomastus 1 8 9 9 9 9 9 10 10 10 11 11 11 11 11 11 12 12 12 12 12 12 12 12 12 12 13 13 13 13-14 13-14 14 14-15 15-18 16 18-19 19 21 2 3 7 9 9 9 9 9 9 9 9 10 10 10 10 10 10 11 11 11 11 11 11 11 11 11 11 12 12 12 12-13 13-14 13 13 14-17 15 17-18 18 20 4 5 7 9 4 6 7 7 3 4 9 4 5 6 10 10 10 3 4 5 6 Column 5: No. larval fonns Eisigella Gravier 1901. or at least. All neuropodia approach midventrally. 16 species and subspecies. see Branchiomaldane Pteroscolex Lütken 1864. Branchiae from setiger 7. However. larval forms Telethusae Savigny 1818. Neuropodia approach midventrally in branchial segments. vitwenti Langerhans 1881. one must have both anterior and posterior ends from the same specimen to get them safely iden- . 3b (2b). With the exception of Branchiomaldane. neuropodia elongated tori. It is necessary to have complete specimens to get them even to sub-family. Invalid Genera Chorizobranchus Quatrefages 1865. Arenicola Lamarck 1801. 9 species and subspecies. Prostomium without appendages. Branchiae fix)m setiger 7. 3a (2b). as thick. 1 b. The family has been the object of very intensive studies by Wells and his students. more than one pair of oesophageal sacs present Abarenicola Generic Definitions Abarenicola Wells 1959. Two body-regions. Body slender. Arenicola claparedii Levinsen 1884. see Arenicolides Protocapitella Berkeley and Berkeley 1932. More than one pair of oesophageal sacs. The bamboo-worms are recognized easily in that for once. the distinct branchial region with their strongly tufted branchiae and the habitat. including a prebranchial and branchial region as well as an asetose caudal end. separated into two or three distinct regions. neuropodia elongated tori. Neuropodia widely separated in the branchial region. Wells (1%2) and other papers established the major classificatory criteria.1976 FAMILY ARENICOLIDAE JOHNSTON THE POLYCHAETE WORMS 37 1835 Body cylindrical. anterior modified spines present in several forms. the long segments with the parapodia at one end. including a prebranchial and branchial region. sessile filaments. Arenicolides Mesnil 1898. Three body-regions. cylindrical segments. with a pair of nuchal slits and a median cephalic keel. they are not as easily identified to genus and species. neurosetae rostrate hooks. Prostomium without appendages. or later. as thick filaments arranged with maximally two or three in a tuft Branchiomaldane Body thick. strongly areolated epidermis in most forms. usually no more than three filaments in a group. the arenicolids are not easily confused with any other polychaetes. All neuropodia widely separated. branchiae first present from setiger 18. give the worms the jointed appearance of slender bamboo-shoots. &ee Arenicola Paparía Sveshnikov 1959. including a prebranchial and a branchial region. Branchiae first present fix)m setiger 18 or behind. Branchiae present on some setigers in a median or posterior region. ä. Two body-regions. The very thick. usually truncate at one or both ends. Arenicola ecaudata Johnston 1835. 2b (lb). Branchiae from setiger 12-17 to the posterior end. Key to Genera la. including a prebranchial and branchial region as well as an asetose caudal end. Lumbricus marinus Linnaeus 1758. One pair of oesophageal sacs present. Three body-regions. a single pair of oesophageal sacs present Arenicola Neuropodia of branchial segments well separated. Notopodia short and rounded. only species. makes it easy to recognize the "sand-worms" from all over the world. Notosetae capillary or limbate. Branchiomaldane Langerhans 1881. 2 Asetose caudal end absent Arenicolides Asetose caudal end present 3 Neuropodia of branchial segment approach midventrally. even though easily recognized to family. see Arenicola FAMILY MALDANIDAE MALMGREN 1867 Capitelliform polychaetes with long cylindrical bodies. neurosetae rostrate hooks. 2a (lb). 2 species. the popular name of the members of this family makes sense. most species with long. sec Arenicola Clymenides Claparède 1863. Notopodia bluntly truncate. branchiae first present from a more anterior setiger as thick tufts of very fine filaments . Notosetae smooth or spinose capillaries. 17x. California. flattened cone Lumbriclymenella Pygidium otherwise 6 Pygidium flattened with anus dorsal Notoproctus Pygidium conical with anus terminal Lumbriclymene Cephalic plaque absent. anal plaque present NICHOMACHINAE 8 Both cephalic and anal plaques present 10 Rostrate uncini in all setigers Micromaldane Acicular spines in first three setigers. MALDANINAE: Both anterior and posterior ends with plaques. (B). 7b (Ib). 4b (3b). 2 7 RHODININAE Rhodine Rostrate uncini in single rows. Most frequently the worms are darkly red or orange in life. and are usually quite large animals. anal plaque present. 4a (3b). 3b (2b). 17x. 9a (8b). is that of Verrill (1900). except that the anterior point of the prostomiura may project as a short palpode. 6a (5b). Tómales Bay. thereafter rostrate uncini 9 Anal funnel symmetrically developed Nicomache Both cephalic and anal plaques absent At least anal plaque present Rostrate uncini in double rows. 3a (2b). anus dorsal. The subfamilies may be defined as follows: RHODININAE: Anterior and posterior ends without plaques. The cephalic plaque always has paired nuchal slits. or funnels. often up to 20 cm in length. The anal plaque may have series of marginal anal cirri or be unadorned. It is here strongly suggested that identification of fragments not be undertaken. plaques. posterior segments with encircling collars . posterior segments not collared LUMBRICLYMENINAE 3 More than 20 setigers present Praxillura Nineteen setigers present 4 Setiger 4 with a deep encircling collar Clymenopsis Setiger 4 without collar 5 Pygidium with anus at the end of a simple. Axiothella rubrocincta. NICOMACHINAE: Anterior end without plaque. anus terminal. Another very important study. Family MALDANIDAE. He established the subfamilies and by and large the currently accepted genera. 5b (4b). especially on members of the subfamily Euclumeninae. uncini in single rows. 2b (la). posterior setigers with numerous encircling collars. except of course when one can be sure about the provenance of the fragments. intertidal. 8b (7a). (A). upturned. The major taxonomic revision of the maldanids was done by Arwidsson (1907). uncini in double rows. These can be plain and rounded. ^^^-* ] FIGURE 11. 8a (7a). 7a (lb). Key to Genera la. 2a (la). lb.38 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 tified. EUCLYMENINAE: Anterior and posterior ends with plaques. The subfamilies are defined on the development of the anterior and posterior ends. often with lighter glandular fields and neuropodial tori. LUMBRICLYMENINAE: Anterior and posterior ends without plaques. or they may form flattened discs. with mud-walled tubes. posterior segments without collars. They are all tubicolous. Maldanids are especially common in shelf sediments. posterior end of the above. uncini in single rows. but is otherwise unadorned. 5a (4b). 6b (5b). 12b (lib). MALDANINAE. 12 species. EUCLYMENINAE. 26b (25b). straight and parallel 28 Anal cone projecting well beyond the rim of the anal plaque Praxillella Anal cone low and not projecting beyond the rim of the anal plaque 29 Apart from the long ventral anal cirrus. acicular spines absent. 28b (27b). Axiothea catenata Malmgren 1865. cephalic keel short and low. A. Cephalic rim incised or entire. 27b (26b). 20a (19b). 23a (18b). Asychis Kinberg 1867b.and neurosetae. lib (10a). anal plaque with a few long cirri only Clymenura More than 30 setigers present 24 Eighteen to 20 setigers present 25 Setiger 4 with a deep encircling collar Macroclymenelia Setiger 4 without a collar Gravierella Anterior neuropodia with rostrate uncini Axiothella Anterior neuropodia with acicular spines 26 Anal plaque with two large ventral and several shorter lateral and dorsal anal cirri Proclymene A single large ventral cirrus and several shorter lateral and dorsal anal cirri present 27 Nuchal slits short. Nuchal slits straight. atlanticus Kinberg 1867b. 23b (18b). anal plaque smooth Microclymene All anal cirri similar in length 19 Midventral annal cirrus (rarely two) distinctly longer than all other anal cirri 23 More than 30 setigers present Macroclymene Nineteen or 20 setigers present 20 Acicular spines in the first neuropodia 21 First neurosetae rostrate uncini 22 Setiger 4 with a deep encircling collar Clymenella Setiger 4 without a collar Isocirrus First setiger with notosetae only. Numerous gill filaments on median setigers.and neurosetae. 18b (16b). Axiothella Verrill 1900. Neurosetae absent in first setiger. cephalic rim shallowly notched laterally Maldane Cephalic keel short and low. THE E*OLYCHAETE WORMS i9 Anal funnel asymmetrical. 10a (7b). 19b (18a). 17b (16a). 11a (10a). all other anal cirri similar in length Euclymene Anal cirri of varying lengths Heteroclymene ' Generic Definitions Abyssoclymene Hartman 1967. 16b (15b). MALDANINAE. 29a (28b). . B. Cephalic rim deeply incised laterally. 14a (13b). 28 species. straight and diverging anterioriy Pseudoclymene Nuchal slits long. 3 species. 15a (10b). 10b (7b). 13a (12b). 21b (20a). 12a (1 lb). anal plaque gently crenulated Abyssoclymene First setiger with noto. gently crenulated along margin. 19a (18a).1976 9b (8b). 22b (20b). cephalic keel short and low. Branchiae absent. 24a (23a). anal plaque with numerous cirri Maldanella First setiger with both noto. anal plaque nearly smooth. 27a (26b). 15b (10b) 16a (15b). 25a (23b). 13b (12b). first with notosetae only. 28a (27b). colmani Monro 1939c. 24b (23a). 22a (20b). A. annularis Hartman 1967. Anal plaque with long midventral and shorter lateral and dorsal anal Branchioasychis Monro 1939c. 14b (13b). 29b (28b). Neurosetae absent in first setiger. cephalic rim deeply incised laterally 14 Numerous branchial filaments on median setigers Branchioasychis Branchial filaments absent Asychis Series of vascular caeca cover the surface of the two last setigers Johnstonia Vascular caeca absent 16 Anal plaque marginally smooth or gently crenulated 17 Anal plaque bordered by distinct anal cirri 18 First setiger with notosetae only. 17a (16a). 25b (23b). 21a (20a). Cephalic rim with deep lateral incisions. 18a (16b). 26a (25b). EUCLYMENINAE. with the dorsal side reduced Petaloproctus Anus dorsal MALDANINAE 11 Anus terramal EUCLYMENINAE 15 Rostrate uncini in two or three series in most setigers Sonatsa Rostrate uncini always in single series 12 First three setigers with acicular neuropodial spines Clymaldane First setiger either without neurosetae or with rostrate uncini 13 Cephalic keel long and high. Nineteen setigers. only species. 20b (19b). 18-20 setigers. anal cone strongly projecting. J. Clymene oerstedii Claparède 1863. with large triangular glandular field on setiger 8. Nineteen setigers. Nuchal slits long and straight. 8 species. M. Anal cone symmetrical. EUCLYMENINAE. 2 species. Micromaldane Mesnil 1897a. Lewis nuchal in anevenly Macroclymenella Augener 1926. flattened cone with anus distal. far posterior ones campanulate. Anal plaque with the anal cirri increasing in length from the dorsal side towards the long midventral cirrus. collars absent. acicular spines present. 3 species. EUCLYMENINAE. 27 species. About 20 setigers. EUCLYMENINAE. LUMBRICLYMENINAE. Clymenopsis Verrill 1900. Acicular spines present. Anal plaque with one long midventral and several shorter anal cirri. Lumbriclymene Sars 1872. Anal plaque a flattened disc with few long anal cirri. 5 species. stewartensis Augener 1926. Sabella lumbricalis Fabricius 1780. avicular uncini on all setigers. Setiger 4 with deep encircling collar. anterior neuropodia with acicular spines or strongly reduced uncini. Nineteen setigers. Branchiae absent. acirrata Arwidsson 1907. L. Maldanella Mclntosh 1885. slender cirri. H. Anal plaque with one long midventral and numerous shorter anal cirri of varying lengths. M. Anal plaque with crenulated margin. Acicidar spines absent. . anal cone low. Clymene torquatus Leidy 1855. M. Heteroclymene Arwidsson 1907. Numerous setigers. Maldane Grube 1860. clymenoides Quatrefages 1850b. Neurosetae absent from first setiger. EUCLYMENINAE. Anal end a simple upturned. Acicular spines present in anterior neuropodia. Clymenella Verrill 1873b. Setiger 4 with deep encircling anterior collar. Clymene cingulata Ehlers 1887. multiannulata Fauvel 1919. LUMBRICLYMENINAE. Anal plaque with all cirri of the same length. cylindricauda Sars 1872. C. NICOMACHINAE. acicular spines present. 2 species. Strongly curved. nuchal slits long and straight. anal plaque with all cirri evenly short. EUCLYMENINAE. Acicular spines present terior neuropodia.40 NATURAL H/STORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Clymaldane Mesnil and Fauvel 1939. cephalic keel long and high. antárctica Mclntosh 1885. omithochaeta Mesnil 1897a. nuchal slits long and straight. slits long and straight. MALDANINAE. Clymene planiceps Sars 1872. Anal cone large. Clymene cirrata Ehlers 1887. 2 species. EUCLYMENINAE. Two last setigers covered with series of vascular caeca. nuchal slits long and straight. Acicular spines absent. Nineteen setigers present. Nineteen setigers present. only species. Gravierella Fauvel 1919. nuchal slits strongly curved. First three neuropodia with spines. cephalic rim notched. 2 species. Clymenura Verrill 1900. 3 species. About 20 setigers. EUCLYMENINAE. 18 species. Johnstonia Quatrefages 1850b. Microcfymene Arwidsson 1907. nuchal slits strongly angled. acicular spines absent. Anal plaque smooth-rimmed. Uncini in single rows. nuchal slits long and straight. Anal plaque with all cirri evenly long. Nuchal slits long and straight. glebifex Grube 1860. with circular cross-section. only species. 11 species. Clymene producía 1897. EUCLYMENINAE. robusta Arwidsson 1907. M. G. Nineteen to 20 setigers. Eighteen to 20 setigers. Nineteen setigers. Acicular spines absent. Nineteen setigers. Eighteen to 20 setigers. M. anal cone low. only species. Euclymene Verrill 1900. Keel short and very low. Acicular spines in first neuropodia. nuchal slits long and straight. Anal cone low. Anal plaque with one long median and numerous evenly long anal cirri. More than 30 setigers. LUMBRICLYMENINAE. Nicomache Malmgren 1865. Anal plaque with a long midventral and numerous short. robusta Arwidsson 191 la. nuchal slits strongly curved. EUCLYMENINAE. Macroclymene Verrill 1900. Acicular spines absent. More than 30 setigers. nuchal slits long and straight. Lumbriclymenella Arwidsson 1911a. Isocirrus Arwidsson 1907. 5 species. Acicular spines in the first neuro- podia. 16 species. EUCLYMENINAE. sibogae Mesnil and Fauvel 1939. Acicular spines absent. Acicular spines in anterior neuropodia. EUCLYMENINAE. nuchal slits long and straight. MALDANINAE. Twenty-two setigers. Rostrate uncini in single rows. Anal plaque with numerous long cirri. nuchal slits long and straight. 2 species. L. setiger 4 with deep encircling collar. Cephalic rira shallowly notched. 7 species. palps absent. questionably PrcwiZ/e/fc Leiocephalus Quatrefages 1865. Rostrate hooks absent. 2 species. set Euclymene Chrysothemis Kinberg 1867b. Anal plaque short. see Asychis ORDER OPHELIIDA Prostomium without appendages. Maximally one anterior asetigerous segment present. terricolus Quatrefages 1865. Eighteen to 20 setigers. Nineteen setigers. 9 species. Composite setae have been reported from a single species of the family Scalibregmidae. see Petaloproctus Paraxiothea Webster 1879a. Prostomium without appendages. Rostrate uncini in single rows. Rostrate uncini in two or three series in some setigers. Acicular spines in anterior neuropodia. nuchal slits anteriorly curved. with anus dorsal. first setigers with notosetae only. see Maldane ¡phianissa Kinberg 1867b. S. Acicular spines in anterior and posterior neuropodia. including capillary setae. ornata Verrill 1880. Rhodine Malragren 1865. see Mylitta Nicomachella Levinsen 1884. only species. with a long midventral and several shorter anal cirri. see Clymenura Maldanopsis Verrill 1900. Eighteen to 19 setigers. see Eue lyme ne Leiochone Grube 1868b. Numerous encircling collars on posterior setigers. projecting well beyond the rim of the anal plaque. 11 species. see Praxillella Promaldane Mesnil 1897a. loveni Malmgten 1865. R. see Asychis Clymene Oken 1815. Acicular spines in anterior neuropodia. acicular spines in the first three neuropodia. Anal cone very large. indeterminable Neco Kinberg 1867b. Sonatsa Chamberlin 1919c. LUMBRICLYMENINAE. Parapodia uniramous or (usually) biramous. this form may be incorrectly assigned to family and in some respects resembles members of the Syllidae. oculatus Arwidsson 1907. Anal plaque asymmetrical with the dorsal side strongly reduced. smooth-rimmed. nuchal slits strongly curved. EUCLYMENINAE. the posterior end of the only known species remains unknown. incompletely known Praxilla Malmgren 1865. Acicular spines present in anterior neuropodia. Pseuäoclymene Arwidsson 1907. FAMILY OPHELIIDAE MALMGREN 1867 Body with a limited number of segments. all setae capillary. Uncini in double rows. blunt or conical. prostomium with high. N.]976 THE POLYCHAETE WORMS 4Í Taxonomic Notes NICOMACHINAE. NICOMACHINAE. ses Euclymene Axiothea Malmgren 1865. All neuropodia short and truncate. see. LUMBRICLYMENINAE. P. Anal plaque with two large midventral and several shorter anal cirri. straight and diverging anteriorly. EUCLYMENINAE. Acicular spines in the first neuropodia. Numbers of setigers variable. All setae simple. Anal plaque with subequal anal cirri. cephalic rim notched. Notoproctus Arwidsson 1907. Proclymene Arwidsson 1907. thick and grub-shaped. acicular spines absent. î&t Asychis Mandrocles Kinberg 1867. Nineteen setigers. see Maldane and Asychis Leiochone Saint-Joseph 1894. it resembles closely members of the genus Euchone. Praxillella Verrill 1881. Numbers of setigers unknown. MALDANINAE. Parapodia biramous or uniramous. others are very slender. Invalid Genera Arwidssonia Mclntosh 1914b. meridionalis Chamberlin 1919c. RHODININAE. long cephalic keel. The genus Sonatsa was described originally in its own subfamily. Praxilla praetermissa Malmgren 1865. hypothetical Sabaco Kinberg 1867b. anal cirri all of the same length except the larger midventral one. nuchal slits long and straight. Anal plaque flat. P. nearly smooth and torpedo-shaped. Opheliids have a series of three very distinct bodyshapes. EUCLYMENINAE. Clymene muelleri Sars 1856. Twenty to 29 or more setigers. Some are short. 8 species. . only species. with small button-shaped parapodial lobes. nuchal slits short. The genus Chaponella Rullier 1972 is considered here in the Sabellidae. Praxillura Verrill 1880. Petaloprocius Quatrefages 1865. often with a deep ventral furrow. 20 species. The anterior end closely resembles well-known species of Maldane and the genus is considered here in the subfamily Maldaninae. indeterminable Heteromaldane Ehlers 1908. prostomium large. 6 species. Nineteen setigers. except for its loss of tentacular crown. Axiothella Caesicirrus Arwidsson 1911b. either smooth or marginally dentate. Acicular spines present. hoodlike. see Ctymenella Petaloclymene Augener 1918. indeterminable Clymene Savigny 1818. Clymene quadrilobata Sars 1856. branchiae present. branchiae present. Lateral eyes absent. if present Ophelia Branchiae absent 7 Branchiae present 8 Lateral eyes absent Tachytrypane Lateral eyes present Polyophtkalmus Lateral eyes present Armandia Lateral eyes absent 9 Anal tube short. A. with two internally attached ventral cirri. two internally attached ventral cirri present. all anal cirri of similar length Antiobactrum Anal tube long. 8a (6b). 2b (la). Dindymenides Chamberlin 1919c. limited to the posterior end of body only Ammotrypanella Generic Definitions Ammotrypanella Mclntosh 1879. Body short and grnblike without ventral furrow. about 5x. FIGURE 12. Body fusiform without ventral groove 2 Body cylindrical with at least posterior ventrum deeply grooved 4 Branchiae absent Kesun Branchiae present 3 First setiger appear in front of the mouth. 2a (la). 4b (lb).42 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Finally. 7a (6a). branchiae in posterior region only Euzonus Body not clearly regionated. Ventral groove along the whole body. some setae spinose. 10b (9b). along the whole body . 3 species. Armandia Filippi 1861. all setae smooth Travisia First setiger appear behind the mouth. 5x. with paired long internally attached ventral cirri and shorter dorsal cirri. Ophelina Branchiae always present. 1971. Antiobactrum Chamberlin 1919c. 6a (4b). 18 species. Family SCALIBREGMIDAE. Anal tube short and with all anal cirri of the same length.0. including inflated head. 9a (8b). Ophelina brasiliensis Hansen 1882. 8b (6b). (B). Ventral groove present along the whole body. only species. árctica Mclntosh 1879. Branchiae present. 3a (2b). only species. expanded palpode at the tip of the prostomium. Lateral eyes present. 7b (6a).. Scalibregma inflatum. 9b (8b). 1 b. First setiger appears behind the mouth. . Key to Genera la. Dindymene concinna Kinberg 1866b. 3b (2b). off Anacapa Island. Opheliids are common animals in sandy and muddy bottoms and have been studied extensively by ecologists interested in sandy beaches. 4a (lb). 10a (9b). Ventral groove present along the whole body. some setae spinose Dindymenides Ventral groove present in posterior part of body only 5 Ventral groove present along the whole body 6 Three body-regions. 90 m. inflated anterior part and narrow posterior part. inflated anteriorly and grooved posteriorly. dorsal anal cirri short 10 Branchiae. California. 6b (4b). branchiae from setigers 8-10. if present. some are anteriorly inflated and posteriorly cylindrical or narrow. modified fix)m Hartmann-Schröder. Family OPHELIIDAE. Lateral eyes absent. A. cirrhosa Filippi 1861. branchiae present and limited to the posterior end only. anal tube long and slender. (A). 5b (4a). Anal tube long and narrow.?Mi<i rathkei. 5a (4a). No notopodial lobes modified. especially the latter resembles forms usually associated with the order Phyllodocida. Body short and stocky or long and slender. 32 species. No attempts have been made below to move these genera to other families. Ophelina Örsted 1843b. see Ophelia Omaria Grube 1869b. Body short and grublike without a ventral groove. All setae smooth. E. on most setigers from setigers 8-10. Anal cirri all short and thick. Setae include capillaries and furcate setae and in some cases acicular spines. Anal tube cut away ventrally forming an open hood with marginal anal 1 am presently following Hartmann-Schröder (1971) and others in accepting Ophelina as the valid name for worms often referred to under the generic name -4mmotrypane. usually combined with a strongly wrinkled or areolated epidermis. Some of these forms {Proscalibregma and Scalispinigera) may in fact not be as closely associated with this family as previously indicated. 3b (2b). see Ophelina FAMILY SCALIBREGMIDAE MALMGREN 1867 Travista Johnston 1840. but always with a slender posterior end 4 Two or three anterior asetigerous segments Neolipobranchius One anterior asetigerous segment 3 Branchiae present Polyphysia Branchiae absent Lipobranchius . Prostomium anteriorly bifid or with T-shaped frontal horns. Branchiae and lateral eyes absent. 2a (la). O. Branchiae limited to posterior region. 44 species. Key to Genera la. indeterminable Terpsichore Kinberg 1866b. Lateral eyes absent. see Ophelina Nais Dujardin 1839. see Dindymenides Ladice Kinberg 1866b. Tachytrypane Mclntosh 1879. with small. Branchiae absent. a head consisting of prostomium and one or two setigers. O. 20 species. arcticus Grube 1866. Invalid Genera Aloysina Clapatède 1864. with paired internally attached ventral cirri and shorter dorsal cirri. often anteriorly inflated. Polyophthalmus Quatrefages 1850a. lateral eyes present. lb.1976 THE POLYCHAETE WORMS 43 Taxonomic Notes Euzonus Grube 1866. first setiger appears before the mouth. Last setigers without epipodial pads. see Ophelina Pygophelia Sars 1869. set Polyophthalmus Neomeris Costa 1844. but it is anticipated that the content of this family may change rather drastically in the near future. acuminata Örsted 1843b. Body with three regions. 3 species. anal cylinder furrowed. bicornis Savigny 1818. Hartmann-Schröder (1971) also recognized a series of subfamilies within this family. all characierized by having capillary and furcate setae. an anterior swollen region and a posterior region with ventral groove. Body with ventral furrow along the whole length. jeffreysii Mclmosh 1879. fusus Chamberlin 1919c. Branchiae absent. 3a (2b). see Polyophthalmus Ammotrypane Rathke 1843. Body with ventral furrow along the whole length. see Ophelina Cassandane Kinberg 1866b. see Ophelia Nitetis Kinberg 1866b. posterior setigers with epipodial pads. Ophelia Savigny 1818. maggotlike and inflated 2 Body long. The scalibregmids contained up to a î'ïW years ago a ivêll-circumscribed group of forms. Branchiae rarely absent. 5 species. Branchiae present. Body fusiform with inflated anterior end and ventral furrow posteriorly. Body short and grublike without ventral furrow. There are two distinct groups of genera within the family. Kesm Chamberlin 1919c. Composite setae reported in one species. 10 species. forbesii Johnston 1840. sometimes anteriorly inflated. Last setiger of anterior region with specialized notopodia. see Ophelia Dindymene Kinberg 1866b. Branchiae presently. making a definition of the family more difficult. triangular or button-shaped parapodia. see Ophelina Urosiphon Chamberlin 1919c. Anal cone long. T". T. Nais picta Dujardin 1839. During last ten years several new forms have been added. Anal tube short with small anal cirri. Parapodia biramous. 2b (la). but without papillae. only species. rarely absent. K. but the differences between the groups and the size of the family does not seem to warrant such treatment. Body with ventral furrow along the whole length. on most setigers. Body short. Setae include capillaries and furcate setae. 13a (12b). one asetigerous segment present. 8b (7b). Branchiae present. Body elongated. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Neurosetae composite heterogomph spinigers Neurosetae capillary. 13b (12b). setae include furcate and capillary setae. Body short and maggotlike. 6b (5b). Acicular setae absent. branchiae and dorsal and ventral cirri absent. linea Hartman 1967. 3 species. Dorsal cirri absent. 2 species. dorsal and ventral cirri absent. furcate and capillary setae present. Acicular spines absent. One asetigerous segment present. 1 lb (8b). longiseta Schmarda 1861. 8a (7b). only species. prostomium bilobed. 9a (8a). 10b (9b). 12a (1 la). Polyphysia Quatrefages 1865. Neolipobranchius Hartman and Pauchald 1971. dorsal and ventral cirri absent. Setae include capillaries and furcate setae. . 3 species. Body short and maggothke. 9b (8a). 6a (5b). 1 la (8b). prostomium T-shaped. Branchiae absent. eyes absent. 2 species. prostomium T-shaped. one asetigerous segment present. Body elongated. 7a (6b). Eyes absent. furcate or acicular Furcate setae absent Furcate setae present Median parapodia with long. H. prostomium trapezoidal. acicular spines present in some forms. 7b (6b). Body elongated. anus within a collar-shaped pygidium. P. glabrus Hartman and Fauchald 1971. branchiatus Fauvel 1928b. Acicular spines in up to three anterior setigers. Branchiae. 5b (4b). 12b (lia). Acicular spines. other setae include capillaries and furcate setae. Acicular spines present in up to four anterior setigers. Body short and maggotlike. digitate postsetal lobes Postsetal lobes short and triangular or absent Prostomium with long. 14a (lib). Body elongated. No asetigerous segment present. Lipobranchius Cunningham and Ramage 1888. Parasclerocheilus Fauvel 1928b. ventral cirri present in posterior setigers. Two or three asetigerous anterior segments present. prostomium T-shaped. Scalibregma Rathke 1843. N. prostomium bilobed. eyes present. Lipobranchius intermedius Saint-Joseph 1894. Furcate and capillary setae present. Branchiae absent. Branchiae present. Proscalibregma Hartman 1967. Eumenia glabra Ehlers 1887. cirriform frontal antenna Prostomium T-shaped. Pseudoscalibregma Ashworth 1901. Body elongated. all setae capillary. otherwise with furcate and capillary setae. 10a (9b). 5a (4b). prostomium a conical lobe./*. Body elongated. Kebuita Chamberlin 1919c. 4 species. prostomium T-shaped. S. furcate setae. Eyes absent. 4b (lb). 14b (Ub).44 4a (lb). only species. eyes absent. Branchiae. 2 species. inflatum Rathke 1843. Dorsal and ventral cirri present in posterior setigers. Eyes present. 4 species. Eumenia jeffreysii Mclntosh 1869. dorsal and ventral cirri absent. One asetigerous segment present. without appendages Neither dorsal nor ventral cirri present At least ventral cirri present Acicular setae present Acicular setae absent Eyes present Eyes absent Branchiae present Branchiae absent Acicular spines absent Acicular spines present Eyes present Eyes absent Dorsal cirri absent Dorsal cirri present Science Series 28 Scalispinigera 5 Proscalibregma 6 Scalibregmides 7 Scalibregmella 8 9 11 Asclerocheilus 10 Hyboscolex Kebuita 12 14 Scalibregma 13 Parasclerocheilus Sclerobregma Sclerocheilus Pseudoscalibregma Generic Definitions Asclerocheilus Ashworth 1901. prostomium T-shaped. only species. Eumenia crassa Örsted 1943b. Branchiae. Hyboscolex Schmarda 1861. Qne asetigerous segment. Scalibregma parva Hansen 1878. only species. furcate and capillary setae present. Body elongated. Acicular spines in first setiger. prostomium T-shaped. branchiata Hartman 1965. Scalibregmella Hartman and Fauchald 1971. acicular spines absent. see Scalibregma Oncoscolex Schmarda 1861. is considered here a synonym of Hyboscolex Schmarda as suggested by Day (1967). Most members of this order are modified moderately from the assumed ancestral condition (Fauchald 1974a) in that they have biramous parapodia and only one. see Polyphysia Eusclerocheilus Hartman 1967. simple. small. Invalid Genera Eumenia Orsted 1843b. Dorsal and ventral cirri absent. indicating that this process is going on independently in these taxa. one asetigerous segment present. For that reason. Acicular setae absent. this order contains most members of the old order ERRANTIA. Body elongated. see Polyphysia Eumeniopsis Bidenkap 1895. more commonly associated with hard substrates than with sands and muds. Taxonomic Note The genus Oncoscolex Schmarda 1861. Sclerobregma Hartman 1965. Suborder PhyUodociformia Phyllodocida with at least two pairs of prostomial appendages and at least two pairs of tentacular cirri. when present. They are frequently highly colored in life. first parapodia lateral. prostomium quadrangular with cirriform long antennae and nuchal organs. Eversible pharynx always muscular and cylindrical. and these colors are diagnostic. S. 5 species. Branchiae absent. eyes present. supported by acicula at least in one ramus. FAMILY PHYLLOIXJCIDAE WILLIAMS 1851 PhyUodociformia with long and slender bodies. palps. One asetigerous segment present. notosetae. Similarly. chilensis Hartmann-Schröder 1965. prostomium T-shaped. dorsal cirri absent. A muscular eversible pharynx present. Body elongated. 2 species. it was considered appropriate to erect suborders and superfamilies within the order. Scalibregmides Hartmann-Schröder 1965. Several families show varying levels of cephalization. However. Parapodia with long. furcate setae. ventral cirri present on posterior setigers. One asetigerous segment present. eyes. even if some families are left without specific intermediate designation. It is possible to divide this order into several orders recognizing as such. Apart from the amphinomids and eunicids and associated families. when present. prostomium with four or five antennae. Branchiae present. furcate setae in notopodia only. It is by far the most speciose of the polychaete orders. Parapodia uniraraous (usually) or biramous with the notopodia represented by a short stalk and large. Acicular spines. making such a change undesirable. neurosetae all composite heterogomph spinigers. only species. but jaws often absent. eversible pharynx unarmed. Palps absent. capillary and furcate setae present. but fade very rapidly upon preservation. the Phyllodocida is characterized by the same characters as all the other orders of polychaetes whereas the suborders of Phyllodocida are characterized by a slightly different set of characters. folióse dorsal cirri held erect over the dorsum. if present. All neurosetae composite. dorsal and ventral cirri absent. Eyes present. Scalispinigera Hartman 1967. One asetigerous segment present. one asetigerous segment present. The phyllodocids are common shallow-water polychaetes.¡976 THE POLYCHAETE WORMS 45 ORDER PHYLLODOCIDA Body elongated. Capillary and furcate setae present. Notosetae all capillary. minutus Grube 1863. rarely with acicula or setae. antennata Hartman and Fauchald 1971. oculata Hartman 1%7. prostomium T-shaped. Sclerocheilus Grube 1863. Body elongated. taxa characterized as suborders here. Branchiae present. smooth capillary setae in both rami. Maximally two pairs of jaws present. The phyllodocids produce copious amounts of mucus when . Parapodia distinct in most forms. frontal or frontolateral. furcate and capillary setae present. Branchiae. see Hyboscolex Prostomium usually with at least one pair of antennae. prostomium T-shaped. S. or perhaps a few segments involved in the anterior tagma. only species. S. a change in function of the notopodia from locomotory to defensive appears to have taken place independently in several of the groups. Two to four pairs of tentacular cirri present. see Kebuita Oligobranchus Sars 1846. Eyes absent. see Pseudoscalibregma Gwasitoa Chamberlin 1919c. 5. S. Branchiae absent. Body elongated. prostomium rectangular. slender postsetal lobes in median and posterior setigers. seta of the above. Family ALCIOPIDAE. 112x. 43-70 m. Lopadorhynchus errans. lOx. deep pelagic. Alaska. 1972. (H). FIGURE . (G). Family PHYLLODOCIDAE. Vanadis formosa. pelagic. Family PONTODORIDAE. modified after Uschakov. (F). iOx. 125x. median parapodium of the above. (A).46 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 13. seta of the above. seta from median setiger. Fontodora pelágica. about 50x. 12x. central Pacific Ocean. (D). 19x. lOOx. (C). off central California. (B). anterior end of the above. Family LOPADORHYNCHIDAE. Canoe Bay. Anaitides groenlandica. (E). 5b (4a). and reductions and fusions can be assessed. 7a (4b). over towards the dorsal side. 9a (8b). This latter character may be somewhat troublesome. in most cases it cannot be avoided. The flattening of the ventral cirri is very striking and once seen. and I have found it helpful to use a low light-angle. 11a (7b). 4a (2b). If possible. Distortions in fixation of ordinary tapering ventral tentacular cirri may be misleading. or if setae are present. 8a (7a). and are of great importance. As stated above. dorsal and ventral cirri broadly ovate My slides Second segment without setae. so that strong shadows fall across the specimen. The best way to observe these structures is to hold the specimen with a pair of forceps so that one can look in at the animal in three-quarter dorsal view. and the third segment has dorsally a tentacular cirrus and ventrally a normal parapodium. However. The papillation of the eversible pharynx is another important character and if the pharynx is not everted. This is easily done. 2a (1 b). 6b (5b). 3a (2a). Important identificatory characters and methods of observations: The most important characters in the Phyllodocidae includes the number and arrangement of antennae on the prostomium and the numbers of tentacular cirri and their arrangement on the first few segments (Bergström 1914). Key to Genera la. Notophyllum Five antenna present 12 Four antennae present 22 Ventral tentacular cirri (on segment 2) folióse 13 .¡976 THE POLYCHAETE WORMS 47 disturbed. 10b (9b). 7b (4b). 6a (5b). 8b (7a). Two antennae present Cirrodoce At least four antennae present 2 Two pairs of tentacular cirri on one segment 3 At least three pairs of tentacular cirri on two or three segments 4 Eversible pharynx diffusely papillated or smooth Eteone Eversible pharynx with large lateral papillae in rows Mysta Three pairs of tentacular cirri on two or three segments 5 Four pairs of tentacular cirri on tiiree segments 7 Third segment with dorsal cirri Lugia Third segment without dorsal cirri 6 Second segment with setae. often assymmetrical and flattened. 12a (11a). so the observations have to be rather careful. in that varying fusions obscure the primitive arrangement. In certain groups of species. 3b (2a). It may be necessary to make parapodial preparations and observe these under the compound microsocpe to ascertain the presence or absence of the internal acicula. most easily visible laterally. the ventral tentacular cirri (on the second segment) may be folióse. but may be difficult to do on smaller animals. In this manner it is possible to follow the segmentai furrows. Uschakov (1972) and Banse (1973). 5a (4a). Important recent revisions include HartmannSchröder (1971). contain internal acicula. this character is never again confused with the distortions one can find in preserved material. 4b (2b). lb..e. the stems of the dorsal cirri. dorsal and ventral cirri lanceolate Hesionura Parapodia biramous (with notacicula and sometimes with notosetae) 8 Parapodia uniramous (without notacicula or setae) 11 Ventral tentacular cirrus (on segment 2) foliaceous Hesperophyllum Ventral tentacular cirri digitate or subulate 9 First tentacular segment a complete ring Austrophyllum First tentacular segment dorsally reduced 10 First segment with two pairs of cirri and setae Nipponophyllum First segment with a single pair of cirri. each specimen should be preserved separately. In principle. the next segment usually both a dorsal and a ventral pair. and xnust be relaxed carefully before preservation. 1 lb (7b). 2b (lb). either ventrally or dorsally on most specimens. This whole process is best done under a stereo microscope. lOa (9b). and then rotate the animal slowly from dorsal to lateral positions. the first segment has a single. and the difference essentially is deñned by convention: the parapodia are considered biramous if the notopodia. dissection may be necessary. dorsal pair of tentacular cirri. The blood-vessels in the notopodial rudiments may resemble the internal acicula. discrepancies from this pattern are common. 9b (8b). Biramous and uniramous parapodia in the phyllodocids are exceedingly similar. i. setae first present from the second segment . second segment with setae Pirakia Pharynx with numerous papillae. 22a (lib). 23b (22b). follows in the main the principles first promulgated by Bergström (1914) and followed by most later authors. 25a (23b). 24a (23a). 21a (I9b). 28b (27b). Phyllodocids with five antennae. 17b (16a). 26b (25a).48 12b (1 la). 27 Simple enlarged setae in segments 2-4 Chaetoparia Segments 2-4 with composite spinigers Prochaetoparia First segment with a pair of small dorsolateral papillae in addition to the tentacular cirri Prophyllodoce First segment with tentacular cirri only 28 First to third pair of tentacular cirri and the antenna globose. Papillae in rows on the base of the eversible pharynx. heart-shaped prostomium with nuchal papilla in the crevice between the lobes. Generic Définitions Anaitides Czemiavsky 1882. pharynx smooth Eumida All three tentacular segments free from one another. 13b ( 12a). 20b (19a). 14b (13a). First segment dorsally reduced and partially fused to the second. 27b (25b). 20a (19a). Certain taxa here considered genera often are considered as subgenera. 27a (25b). first segment dorsally reduced Vitiaziphyllum First and second tentacular segment fused. 22b (I lb). Parapodia uniramous. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Ventral tentacular cirri digitate or subulate 16 First segment dorsally reduced 14 All tentacular segments complete rings 15 Second tentacular segment with setae Sige Second tentacular segment without setae Pterocirrus Second tentacular segment with setae. usually narrow and elongated 21 Pharyngeal papillae distally in six rows Bergstroemia Pharyngeal papillae dispersed Clavadoce Pharynx with few papillae. 30a (29b). This creates a problem of setting the relative taxonomic value of the same character in different parts of the family. 28 species. 29b (28b). four pairs of cirri digitate Sphaerodoce All pairs of tentacular cirri and the antennae digitate or subulate 29 Nuchal papilla on a posterior prolongation from the prostomium Paranaitis Prostomiura heart-shaped with the nuchal papilla in a posterior incision 30 Pharyngeal papillae basally in rows Anaitides Pharyngeal papillae basally dispersed Phyllodoce Taxonomic Notes The system indicated above.. 28a (27b). 16a (12b). First two segments fused dorsally 23 Nuchal papillae absent 24 Nuchal papillae present 25 Tentacular cirri flattened Nereiphylla Tentacular cirri digitate or subulate Genetyllis First and second segment fused to each other and to the prostomiura 26 First and second segment fused to each other at least dorsally. 15a (13b). 30b (29b). 26a (25a). 18b (16b). Phyllodocids with four antennae. 29a (28b). It incorporates the main features of the systems organized by Uschakov (1972) and Banse (1973). Phyllodoce groenlandica Örsted 1843a.. Banse 1973). pharynx papillated 19 Reniforra ventral cirri broadly attached transverse to the long axis of the parapodia 20 Ventral cirri narrowly attached. proboscis papillated Steggoa Second tentacular segment without setae. Eulalia charcoti Gravier 191 lb. 18a (16b). 19b (18b). 5 species. 25b (23b). For the sake of consistency. 14a (13a).. pentagonal or . 16b (12b). these characters are here all considered to be of generic value. to the subgeneric level in some cases and to the generic level in others (cf. 15b (13b). 19a (18b). 13a (12a). 17a (16a). all tentacular cirri cylindrical. proboscis smooth Natalia Nuchal epaulettes present 17 Nuchal epaulettes absent 18 First and second tentacular segment free from each other and complete rings Vitiazia First and second tentacular segment fused. 24b (23a). 21b (19b). Austrophyllum Bergström 1914. 23a (22b). but free from the prostomium . second segment without setae Eulalia All three tentacular segments form complete rings Protomystides and Pseudeulalia (these two cannot be separated on current information). Phyllodocids with four antennae. 9 species. Phyllodocids with five antennae of which the median is very small. nilssoni Malmgren 1867. cristate Hartman and Fauchald 1971. Phyllodocids with four antennae. Phyllodocids with four antennae and a nuchal papilla. only species. fragilis Hartmann-Schröder 1958. 3 species. all cirri cylindrical. only species. Hesperophyllum Chamberiin 1919a. Mystides Théel 1879. Eversible pharynx diffusely papillated. Nereiphylla Blainville 1828. First and second segment fused and dorsally reduced. C. G. all tentacular cirri cylindrical. long prostomium and no nuchal papillae. Eversible pharynx diffusely papillated or smooth. short wide prostomium and no nuchal papilla. Eversible pharynx diffusely papillated. Third segment without dorsal cirri. Eversible pharynx with diffuse papillation. Specialized.1976 THE POLYCHAETE WORMS 49 rounded prostomium and without a nuchal papilla. Nereis viridis Linnaeus 1767. two pairs of cylindrical tentacular cirri present. Eteone aurantiaca Schmarda 1861. H. First segment dorsally reduced. which. ventral cirri large and reniform and attached transverse to the long axis of the parapodium. All tentacular segments complete and free from one another. borealis Théel 1879. all tentacular cirri cylindrical. All tentacular cirri cylindrical. pentagonal prostomium and indistinct nuchal papilla. Parapodia uniramous. Clavadoce Hartman 1936c. Phyllodocids with four antennae. . M. Eulalia Savigny 1818. Parapodia blramous. All anterior segments free from one another. Genetyllis Malmgren 1865. Phyllodocids with four antennae. Hesionura Hartmann-Schröder 1858. A pair of auricular lobes attached between prostomium and first parapodia. pentagonal prostomium and no distinct nuchal papilla. Two pairs of tentacular cirri on first segment. 2 species. All anterior segments complete and free from one another. N. Eversible pharynx smooth or with very small papillae. first segment with normal setae. Phyllodocids with five antennae. 8 species. ventral cirrus large and transversely attached. tectum Chamberlin 1919a. all four pairs of tentacular cirri club-shaped and slightly flattened. quadrangular prostomium and no nuchal papilla. Third segment with dorsal cirrus. only species. Eulalia nigrimaculata Moore 1909. Two tentacular segments. Phyllodocids with two antennae. 13 species. all tentacular cirri flattened. in turn. Phyllodocids with five antennae. all cylindrical. quadrangular prostomium and without nuchal organs. 40 species. 50 species. All three tentacular segments free from one another and from the prostomium. Lugia Quatrefages 1865. long prostomium and no nuchal papilla. Cirrodoce Hartman and Fauchald 1971. Phyllodocids with five antennae. Phyllodocids with five antennae. C. is fused to the second. Mysta Malmgren 1865. 11 species. Eteone Savigny 1818. Parapodia uniramous. pentagonal prostomium and without a nuchal papilla. parapwdia uniramous. ventral cirrus reniform. All tentacular segments complete. barbota Malmgren 1865. ventral cirrus foliaceous. all others cylindrical. Parapodia uniramous. short wide prostomium and no nuchal papilla. but free from the other tentacular segments. Nereis flava Fabricius 1780. free from each other and from the prostomium. 6 species. short and wide prostomium and no nuchal papilla. Parapodia uniramous. 3 species. all tentacular cirri cylindrical. but the first dorsally reduced. Eversible pharynx diffusely papillated. Chaetoparia Malmgren 1867. ventral cirri large and reniform and attached transverse to the long axis of the parapodium. Eumida Malmgren 1865. Three pairs of cylindrical tentacular cirri on two segments. only species. pentagonal prostomium and no nuchal papilla. Eulalia sanguínea Örsted 1843b. Third segment without dorsal cirrus. enlarged simple setae in segments 2-4. Parapodia uniramous. M. C. Phyllodocids with five antennae. lutea Malmgren 1865. paretti Blainville 1828. Parapodia blramous. all tentacular cinri cylindrical. all tentacular cirri cylindrical. Eversible pharynx with papillae in six rows distally. First tentacular segment dorsally reduced and fused to the second. Phyllodocids with five antennae. /i. Parapodia uniramous. Eversible pharynx with large lateral papillae in rows. all three pairs of tentacular cirri cylindrical. Three pairs of tentacular cirri on two segments. Phyllodocids with four antennae. Parapodia uniramous. prolonged prostomium and no nuchal papilla. Bergstroemia Banse 1973. First and second tentacular segments are fused and reduced dorsally. triangular or trapezoidal prostomium and a small nuchal papilla. splendida Hartman 1936c. Two tentacular segments with three pairs of tentacular cirri. prostomium fused to the first segment. All tentacular segments are separate and complete rings. All tentacular segments free from one another and from the prostomium. all tentacular cirri cylindrical. Vitiazia Uschakov 1953. Sphaerodoce Bergström 1914. Phyllodocids with four antennae. Notophyllum Örsted 1843b. P. setae first present in the third segment. First and second segments fused and dorsally reduced. all tentacular cirri cylindrical. but with setae. setae present from second segment. 6 species. all others cylindrical. Four pairs of cylindrical tentacular cirri on two segments. Phyllodocids with four antennae and no nuchal papilla. Eversible pharynx closely papillated. parapodia uniramous. All tentacular segments free. Phyllodocids with four antennae and no nuchal papilla. exigua Eliason 1962. All three anterior segments fi-ee from each other. ventral cirri folióse. Phyllodocids with five antennae.50 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Nipponophyllum Imajima and Hartman 1964. all others cylindrical Parapodia uniramous. penîagonal prostomium and no nuchal papilla. all tentacular cirri cylindrical. pentagonal prostomium and no nuchal papilla. 3 species. Eversible pharynx smooth. Phyllodocids with four antennae. dogieli Uschakov 1953. Phillodoce quadraticeps Grube 1878. heart-shaped prostomium with a nuchal papilla between the lobes. all tentacular cirri cylindrical. First and second segment fused. first segment dorsally reduced. Phyllodocids with four antennae. laminosa Savigny 1818. Eversible pharynx with few scattered papillae. broadly transverse prostomium and nuchal epaulettes present. only species. Phyllodocids with five antennae. pentagonal prostomium and no nuchal papilla. Eulalia magalaensis Kinberg 1866b. First segment with a pair of small papillae dorsal to the tentacular cirri. Notophy Hum japonic urn Marenzeller 1879. Sige fusigera Malmgren. Phyllodocids with five antennae. Paranaitis Southern 1914. Eulalia picta Kinberg 1866b. first three pairs of tentacular cirri and the antennae globose. only species. First and second segment fused and dorsally reduced. Phyllodocids with four antennae. Pkyllodoce foliosa Sars 1835. 4 species. Mystides bidentata Langerhans 1880. Pkyllodoce Savigny 1818. P. setae first present in second segment. first segment dorsally reduced. Eversible pharynx with diffuse papillation. Sige Malragren. Pkyllodoce {Eulalia) punctifera Grube 1860. Parapodia biramous. last pair of tentacular cirri (on third segment) digitate. Parapodia uniramous. V. Ventral tentacular cirrus folióse. Genetyllis brevis Ehlers 1901. . Eversible pharynx with rows of lateral papillae. Pterocirrus Claparède 1868. ventral tentacular cirrus folióse. 15 species. setae present from second segment. Steggoa Bergström 1914. Phyllodocids with five antennae. three or four pairs of cylindrical tentacular cirri on three segments. Parapodia uniramous. only species. Prophyllodoce Hartman 1966b. Prostomium with diffuse papillae. first segment dorsally reduced. Parapodia uniramous. only species. broadly transverse prostomium. heart-shaped prostomium with nuchal papilla in the crevice between the lobes. rounded prostomium and a nuchal papilla present. first segment dorsally reduced. all others cylindrical. about five species. Parapodia uniramous. Parapodia uniramous. All three tentacular segments complete rings. Eversible pharynx diffusely papillated. rounded pentagonal prostomium and no nuchal papilla. Phyllodocids with five antennae. only species. all tentacular cirri are cylindrical. Parapodia uniramous. Parapodia uniramous. only species. only species. Phyllodocids with five antennae. All tentacular segments complete rings. 1865. Phyllodocids with four antennae. First segment dorsally reduced and fused to the second. pentagonal prostomium and no nuchal papilla. First and second segment fused to each other and to the prostomium. Protomystides Czemiavsky 1882. fi^ee from one another. Pseudeulalia Eliason 1962. 48 species. Notalia Bergström 1914. Anaitis wahlbergi Malmgren 1865. All tentacular segments free from one another. 13 species. all tentacular cirri cylindrical. all others cylindrical. P. All tentacular segments free from one another. ventral tentacular cirrus folióse. 1865. Parapodia uniramous. but first or first and second dorsally reduced. Parapodia biramous. heart-shaped prostomium with a nuchal papillae between the lobes. a nuchal papilla present. hawaiia Hartman 1966b. Pkyllodoce (Eulalia) macToceros Grube 1860. no setae in first segment. pentagonal prostomium and no nuchal papilla. Pirakia Bergström 1914. with normal composite setae in all setigers. Prochaetoparia Bergström 1914. 10 species. all tentacular cirri cylindrical. Phyllodocids with five antennae. see Eteone Eteonides Hartmann-Schröder 1960a. see Nereiphylla Carobia (Paracarobia) Czemiavsky 1882. or with a single such appendage. spherical eyes. 3b (2b). indeterminable Myriacycium Grube 1880.1976 THE POLYCHAETE WORMS 5/ Phyllodocids with four large and one small median antenna. indeterminable Lepadorhynchus Schmarda 1861. Invalid Genera Carobia Quatrefages 1865. see Polyodontes (family Polyodontidae) Porroa Quatrefages 1865. error for Eumida Eunomia Risso 1826. 4a (lb). see Notophyllum Phyllodoce Ranzani 1817. hypothetical Paraeulalia Czemiavsky 1882. 8a (7b). broadly truncate prostomium and paired nuchal epaulettes. see Eumida (?) Eumenia Quatrefages 1865. see Hesionwa Eulalides Czemiavsky 1882. Vitiaziphyllum Uschakov 1972. indeterminable Mesomystides Czemiavsky 1882. sec Notophyllum Globiodoce Bergström 1914. nearly all setae composite spinigers Alciopa Parapodia without cirriform appendages. Phyllodocids with five antennae. dorsal and ventral cirri foliaceous. 7b (5b). see Notopkyllum Mesoeulalia Czemiavsky 1882. 8b (7b). see Eulalia ana Sige Eteonella Mclntosh 1874. extending well beyond the tip of the parapodia Watelio . see Anahides Carobia (Protocarobia) Czemiavsky 1882. All tentacular segments fully developed and separate from one another. 5a (4b). see Phyllodoce Cirraria Sveshnikov 1959. 6b (5a). slender pelagic organisms. broadly transverse prostoinium and paired nuchal epaulettes. see Eteone Hypocirrus Giard 1913. indeterminable Hypoeulalia Bergströra 1914. see Anaitides Macrophyllum Schmarda 1861. Prostomium with five antennae and a pair of very large. 4b (lb). 6a (5a). all tentacular cirri cylindrical. follows these two authors. see Notophyllum Trachelophyllum Levinsen 1883. nuchalum Uschakov 1972. only species. Parapodia uniramous. 3a (2b). simple or acicular setae present 5 All setae simple g At least some composite setae present 7 Parapodia with a cirriform appendage Krohnia Parapodia without cirriform appendages Alciopina Parapodia with cirriform appendages 8 Parapodia without cirriform appendages Plotohdmis Acicula barely extending beyond the tip of the parapodia Rhynchonerella Aciculum prolonged. mainly known for their very large. 5b (4b). Parapodia uniramous. complex cameratype eyes (Hermans and Eakin 1974). transparent long bodies found exclusively pelagically. see Eulalia Protocarobia Czemiavsky 1882. 2a (la). The key to genera has been rewritten from Uschakov (1972). first segment reduced dorsally. Several anterior segments with mdimentary parapodia 2 Anterior segments with fully developed parapodia 4 All setae simple capillaries Naiades Setae composite spinigers 3 Parapodia distally with a cirriform appendage Vanadis Parapodia distally without a cirriform appendage Torrea Parapodia with two distal cirriform appendages. see My slides Myriana Savigny 1820. The pelagic polychaetes recently were reviewed by Dales and Peter (1972). lapsus calami ioxSphaerodoce Hypereteone Bergström 1914. Key to Genera lalb. 7ä (5b). Three to five pairs of tentacular cirri. see Phyllodoce Pseudonotophyllum Czemiavsky 1882. tentacular cirri cylindrical. The system indicated below. indeterminable Eunotophyllum Czemiavsky 1882. indeterminable Eumidia Verriil 1873b. First and second segment fused. V. Parapodia uniramous. error for Eunomia. see Eulalia Nothis Pruvot 1885. larval forms Eracia Quatrefages 1865. setae simple or composite. see Eulalia Kinbergia Quatrefages 1865. Alciopids are delicate. see Notophyllum FAMILY ALCIOPIDAE EHLERS 1864 Phyllodociforms with slender. 2b (la). Alciopids with rudimentary parapodia on the two first segments. Five pairs of tentacular cirri. Alciopids with parapodia fully developed on anterior segments. Alciopids with parapodia fully developed on all segments. 5 species. N. It is preferable to segregate them in their own family. 2a (la). Dales and Peter (1972) and Uschakov (1972) have given recent reviews. Alciopids with parapodia fully developed on anterior segments. 2 species. see Vanadis Nauphanta Greeff 1876. Alciopids with parapodia fully developed on anterior segments. Members of this family often are considered as truly phyllodocids and are treated as such. Parapodia with a single digitiform cirrus distally. Three pairs of tentacular cirri. Four or five pairs of tentacular cirri present. 2 species. including numerous slender capillaries and several shorter acicular setae. Four pairs of tentacular cirri present. Parapodia without digitiform distal cirrus. Parapodia without digitiform cirri. short-bodied phyllodociform polychaetes. stt Rhynchonerella CaUizonella Apstein 1891. lapsus calami for Vanadis Corynocephalus Levinsen 1885. gracilis Costa 1862. only species. possibly three species. Each parapodium with two digitiform cirri distally. including slender composite setae with long appendages. parapodial cirri long and digitate 2 3 Pelagobia . including capillaries and thicker. Plotohelmis Chamberiin 1919c. A. maximally ten segments. Setae of several kinds. All setae simple capillaries. ste Alciopa Halodora Greeff 1876. A. Three pairs of tentacular cirri. Each parapodium with a digitiform distal cirrus. only species. Parapodia with single digitiform distal cirri. one. formosa Claparède 1870b. Vanadis Claparède 1870b. Alciopids with parapodia fully developed on anterior segments. R. Five pairs of tentacular cirri present. reynaudi Audouin and Milne Edwards 1833b. All setae composite spinigers with long narrow appendages. Setae include numerous composite spinigers. Torrea Callizona Greeff 1876. but a few slender capillaries may be present. Two or three pairs of tentacular cirri present. All setae composite spinigers with long. Three or four pairs of tentacular cirri. cantrainii delle Chiaje 1830. parasitica Claparède and Panceri 1867. Alciopa lepidota Krohn 1845. Callizona gravieri Benham 1929. ste. all setae composite except for modified anterior setae in some forms. Alciopina Claparède and Panceri 1867. Alciopa candida delle Chiaje 1828. V. inclusion in the phyllodocids would make that family very difficult to define. see Torrea Mauita Chamberiin 1919c. slender appendages. slender appendages and a few simple capillaries. Three or four pairs of tentacular cirri. Alciopids with several segments with rudimentary parapodia. 13 species. 3 species. Prostomium with four antennae and a pair of small eyes. Parapodia without digitiform distal cirri. Alciopina Greeffia Mclntosh 1885.52 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Generic Definitions Science Series 28 Alciopa Audouin and Milne Edwards 1833b. Five pairs of tentacular cirri. 2 species. Krohnia Cleta Claparède 1870b. All setae simple. ste. Setae include numerous composite spinigers with long. but the latter considered the lopadorhynchids as part of the phyllodocids. acicula extending well beyond the tip of the parapodia. Watelio Stöp-BowiU 1948. Torrea Quatrefages 1^5. see Alciopa 1868 FAMILY LOPADORHYNCHIDAE CLAPARèDE Exclusively pelagic. Parapodia uniramous. Krohnia Quatrefages 1865. Setae present at the base of the tentacular cirri Setae absent at the base of the tentacular cirri Two pairs of tentacular cirri on one segment. Rhynchonerella Costa 1862. All setae simple. dorsal and ventral cirri cirriform. Key to Genera la. Parapodia without digitiform cirrus. and shorter acicular setae simple or composite. Alciopids with rudimentary parapodia on first three body segments. Rhynchonerella capitata Greeff 1876. see Alciopa Liocapa Costa 1862. see. Each parapodium with a digitiform distal cirrus. Setae of two kinds including simple acicular setae and composite setae with slender pointed appendages. Invalid Genera Asterope Claparède 1870b. Alciopids with parapodia fully developed on the anterior segments. Naiades delle Chiaje 1830. lb. acicular setae in anterior segments and ventrally in other parapodia. All parapodia similar with long setal lobes and very long dorsal and ventral cirri. All setae composite with narrowly oar-shaped. All parapodia similar. curium Reibisch 1895. Lopadorhynchids with three pairs of tentacular cirri on two segments. Prostomium not fused to the first setiger. with small rounded dorsal cirri and long. 2 species. All setae simple. including in some genera finely attenuated felt setae and in others harpoonshaped. ite Maupasia Haiiplanes Reibisch 1895. facial tubercle absent Stout supportive notosetae present. Setae include composite spinigers with slender appendages and sometimes simple setae in the first seliger. only species. Pedinosoma Reibisch 1895. L. lb. Lopadorhynchids with two pairs of tentacular cirri on first segment. eyes small. xt Lopadorhynchus FAMILY PONTODORIDAE BERGSTRöM 1914 Phyllodociform. pelágica Greeff 1879. all setae composite spinigers Generic Definitions Lopadorhynchus Grube 1855. Lopadorhynchids with three pairs of tentacular cirri on two segments. slender. 4 species. Eversible pharynx. see Pontodora Suborder Aphroditiformia Phyllodocida with up to three antennae (may be absent). Pontodora Greeff 1879 and a single species P. simple setae All parapodia similar. It has so far been reported from warm waters in the Atlantic Ocean and from colder waters in the Pacific Ocean (Uschakov 1972). if armed. oar-shaped appendages with dentate cutting margin. elongated uniramous parapodia with long dorsal cirri. Pelagobia Greeff 1879. erect. slender ventral cirri. Maupasia Viguier 1886. setae composite. each marginally smooth or irregularly lobed. caeca Viguier 1886. longicirrata Greeff 1879. Normal setae composite with broad. each conical with short dorsal and ventral cirri. Key to Genera la. P. Invalid Genus Epitoka Treadwell 1943. 2a (la). marginally dentate appendage. but sometimes subdistally spurred 2 4 Heteraphrodita 3 Tricertia Aphrodita . All setae composite spinigers. Dorsum covered with felted notosetae Dorsum without feh. First parapodia directed forwards. slender or thick. Eversible pharynx with a pair of jaws or unarmed. Al! parapodia similar. Notosetae of several kinds. with two or four jaws. Superfamily Aphroditacea Dorsal ciiri alternating with elytrae in at least part of the body. 3b (lb). 3a (2b). Invalid Genera Haiiplanella Treadwell 1943. see Maupasia Hydrophones Claparède 1870b. Two pairs of tentacular cirri present. Prostomium with four antennae. Slender. 3a (lb). see. brevis Grube 1855. long ventral palps. facial tubercle present Neutosetae distally bifid Neurosetae distally entire. Maupasia Halyplanes Reibisch 1893. The family is known for a single genus. FAMILY APHRODITIDAE MALMGREN 1867 Aphroditaceans with dorsoventrally flattened bodies and one antenna present. no setae on tentacular segment. Elytrae alternating with dorsal cirri. sts Lopadorhynchus Mastigethus Chamberlin 1919c. setae absent from tentacular segments. notosetae cylindrical in cross-section. flattened spines or cylindrical spines. parapodial cirri short First two or three parapodia modified and with large. 9 species. 3b (2b). P. Lx)padorhynchids with two pairs of tentacular cirri on the first segment. or with a very few feh setae Stout supportive notosetae absent. Normal segments with conical setal lobes and rounded presetal lobes. M. Two or sometimes three anterior setigers enlarged with markedly modiñed parapodia and simple setae. one pair of tentacular cirri.1976 2b (la). &çs Lopadorhynchus Reibischia Bergström 1914. setae present on tentacular segment. THE POLYCHAETE WORMS 53 Maupasia Lopadorhynchus Pedinosoma Three pairs of tentacular cirri on two segments. 2b (la). pelagic polychaetes. setae present on tentacular segments. small. altoni Pettibone 1966a. Harpoon-shaped notosetae absent. Harpoon-shaped notosetae absent.54 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Seríes 28 FIGURE 14. Long protective notosetae present. notosetae long. other long supportive notosetae present. 28 species. Harpoon-shaped notosetae present. Facial tubercle present. felt absent. 5 species. filicomis Kinberg 1855. Halfmoon Bay. Family APHR0DrnDAE. 2. felt present. araeoceras Haswell 1883. 5b (4b). L. Aphrogenia Kinberg 1855. Facial tubercle present. H.5x. 26 species. marginally serrated Notosetae flattened and marginally serrated Notosetae cylindrical and smooth Laetmonice 5 Aphrogenia 6 Pontogenia Hermionopsis Generic Definitions Aphrodita Linnaeus 1758. Facial tubercle absent. only species. Harpoon-shaped notosetae present Harpoon-shaped notosetae absent Notosetae sabre-shaped and smooth Notosetae. Tricertia Haswell 1883. A. California. Heteraphrodita Pettibone 1966a. H. felt absent. levisetosa Seidler 1923. T. 6b (5b)./4pÄr£>t//ia refulgida. felt present. only species. 6a (5b). Protective notosetae flattened and marginally serrated. felt present. alba Kinberg 1855. 4a (lb).^4. Stout supportive notosetae absent. straight and cylindrical without asperities. flattened and curved. if flattened. felt removed on right side. Laetmonice Kinberg \i55. Harpoon-shaped notosetae absent. . felt poorly developed or absent. Facial tubercle present. Facial tubercle present. only species. felt absent. Hermione chrysocoma Baird 1865. notosetae long. 4b (lb). Neurosetae distally bifid. Facial tubercle present. aculeata Linnaeus 1761. 5a (4b). Hermionopsis Seidler 1923. Facial tubercle present. sabrelike or smooth and capillary. Pontogenia Claparède 1868. only species. tOx. intertidal. generally considered valid. however. 84 m. California. Sunset Bay. Marginally smooth or fringed FIGURE IS. Halosydna brevisetosa. have been excluded from the named genera. elytrae removed on left side. both have been defined on characters usually used in the group. ste Laetmonice Miinesia Quatrefages 1865. stc Laetmonice Hermonia Hartman 1959. Invalid Genera Bartularía Sveshnikov 1959^ NOMENNUDUM Cyanippa Castelnau 1842. usually. Eversible pharynx with four jaws. . st^ Laetmonice Laetmatonice Ex A UCTORE. off Santa Barbara. s&eAphrodita Halithea Savigny 1818. (A). sec Laetmonice Hermione Blainville 1828. Family POLYODONTIDAE. Family POLYNOIDAE.1976 Taxonomic Notes THE POLYCHAETE WORMS 55 The above treatment follows Pettibone (1966a) closely in that several genera (e. two or.g. The validity oiTricertia and Hermionopsis is doubtful. ste Aphrodita FAMILY POLYNOIDAE MALMGREN 1867 Aphroditaceans with dorsoventrally flattened bodies. three antennae present. elytrae removed on the right side. and therefore. s&t Laetmonice Halogenia Horst 1916b. Polyodomes sp-. one. see Laetmonice Letmonicella Roule 1898. Oregon. 15x. (B). have been included. Hermonia Hartman 1959). 1 la (10a). parapodia short 13 Some notocirri with spherical subdistal inflations Andresia Notocirri maximally gently inflated 14 Eyes very large and confluent Robertianella Eyes small Antinoana Fifteen pairs of elytrae 16 Sixteen or more pairs of elytrae 42 More than 50 segments present. (5b). 13b (12b). . and usually distinctly lateral in position. 8b (4a). 4b (3a). Key to Genera 1 a. There are no recent reviews of the whole family. lOa (8b). They differ from Harmothoe SENSV STRICTU. especially members of HARMOTHOINAE. 8a (4a). preserve each scale-wotm separately. Nearly 100 genera of scale-worms presently are recognized and more are being described every year. elytrae cover the whole body.56 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 elytrae alternate with dorsal cirri at least on the anterior end. and especially Marian H. 3a (2a). If possible. but never as felt. 4a (3a). 16-17 segments present Macellicephaloides Prostomium a single unified lobe. care must be taken that scales are maintained with the specimens.or harpoon-setae. recent authors interested in these worms. (2b). parapodia prolonged Herdmanella Twelve to 14 pairs of elytrae present. The best way of doing this. with paired rounded anterior projections. if the specimens are larger. 1 lb ( 10a). 10b (8b). include Hartman and Hartmann-Schröder. 9b (8a). notosetae variously developed. 13a (12b). 17-19 segments Macellicephala Notosetae coarser than neurosetae Bylgides Notosetae much thinner than neurosetae ¡phione First setigers with simple. 15b (10b). rarely large worms. 16b (15a). have been considered as separate genera below. those with strongly ornamented elytrae tend to be free-living. All setae simple. since these furnish one of the best diagnostic characters for species identifications. when preserving scale-worms. (5b). 5a 5b 6a 6b (2b). is either to bulk-relax the whole sample (MS 222. or at most. most genera are well defined and the species are recognizable. 3b (2a). lateral antennae reduced or absent 7a (3b). 9a (8a). only anterior part of body covered by elytrae 17 Less than 45 segments present. 12b (lib). Forms with smooth elytrae are often commensals with other organisms. Several taxa that usually are considered as subgenera under Harmothoe. They are usually medium-sized. 15a (10b). 16a (15a). 7b (3b). Lepidonotus. strongly curved hooks 9 Anterior hooks absent 10 Two segments with anterior hooks Australaugeneria Three segments with anterior hooks Uncopolynoe Fourteen or fewer paks of elytrae 11 Fifteen or more pairs of elytrae 15 Notosetae absent Polynoella Notosetae present 12 Eight or nine pairs of elytrae present. Cervilia 2 3 MACELLICEPHALINAE 5 Median antenna present 4 Median antenna absent IPHIONINAE 7 Lateral antennae attached sub-distally or ventrally on the prostomium (ceratophores below the anterior points of the prostomium which may be produced into cephalic peaks) HARMOTHOINAE 8 Lateral antennae attached distally on the prostomium (ceratophores continuations of the prostomium) LEPIDONOTINAE 51 Fifteen pairs of elytrae Macelloides Maximally 13 pairs of elytrae 6 Prostomium consisting of two inflated lobes. 12a ( I lb). the last 10-15 segments uncovered 23 Lateral antennae inserted posteriorly on the prostomium Lateral antennae inserted anteriorly At least two antennae well developed Median antenna well developed. in exactly the characters that usually are considered valid generic characters elsewhere in the family and cannot be related to Harmothoe by overlap. 2a (lb). but within several of the larger genera (Harmothoe. 14b (13b). Halosydna) problems occur. or MgClj) or. Pettibone of the Smithsonian Institution. 14a (13b). 2b (lb). The polynoids are the most commonly occurring scale-worms. lb. (40b). 27b). Eunoe Neurosetae distally pencillate Austrolaenilla Neurosetae never pencillate 38 Both noto. straight spines Hartmania Neurosetae distally falcate Gattyana Ventral cirri folióse.]<)76 17a ( 16a). (23b). (30b). (33b).. prostomium quadrangular Nemidia Eyes distinct. 33b). 31b). 21b 19b). ventrum usually smooth 28 Neurosetae in part distally trifid Gorekia Neurosetae distally uni. (31b). 15b). (35b). (37b). filamentous tips Antinoella All neurosetae with slender tips. 24b). 29b). without pockets 18 Notosetae absent Heteropolynoe Notosetae present 19 All notosetae thicker than or at least as thick as the neurosetae 20 At least some capillary notosetae present 21 Notosetae held erect over body Hermadion Notosetae short and lateral in position Hemilepidia Eyes reduced or absent. 33a). 19b 18b). neurosetae bidentate. 31a). 20a { 19a). 23a). 17b 16a). (31a). 22b 21b).or bi-dentate 33 Some neurosetae with fine. 21a 19b). (42a). (34b). 25b). (27b). dentate capillaries Melaenis Neuropodial capillary setae absent 40 Some neurosetae deeply cleft with both teeth long and narrow Eucranta Neurosetae with main fangs very much larger than secondary teeth 41 Posterior eight to ten segments not covered by elytrae Lagisca Whole body covered by elytrae Harmothoe Sixteen pairs of elytrae 43 Eighteen or more pairs of elytrae 45 Neurosetae with semilunar pockets Subadyte Neurosetae slender and serrate. (33a). (28b). prostomium rounded 22 Notosetae mainly capillary. [25b). thick and slender Hesperonoe All notosetae thick and similar in shape 36 Eyes absent Intoshella Eyes present . neurosetae unidentate Enipo Notosetae mainly thick and blunt-tipped. Kermadecella All dorsal cirri similar in length. 18a ( 17b). 34b). short dorsal cirri basally inflated . 43b). THE POLYCHAETE WORMS 51 23a 23b 24a 24b 25a 25b 26a 26b 27a 27b 28a 28b 29a 29b 30a 30b 31a 31b 32a 32b 33a 33b 34a 34b 35a 35b 36a 36b 37a 37b 38a 38b 39a 39b 40a 40b 41a 41b 42a 42b 43a 43b 44a 44b 16b). 39b).. semilunar pockets absent 44 Notosetae distinctly serrated Leucia Notosetae with a few spines Scalisetosus . but without filaments Antinoe All neurosetae unidentate 34 At least some neurosetae bidentate 37 Dorsal cirri alternating between short and long. 30b). ventrum papillose Phyllosheila Ventral cirri digitate. 20b 19a). none basally inflated 35 Notosetae of two kinds. notosetae with capillary tips Tenonia Notosetae and most neurosetae coarse 39 Neurosetae include furcate spines with subequal tips and slender. 28b). (15b). (38b). (29b). 40b). 19a ( 18b). (43b).and neurosetae slender. uni. except in far posterior setigers Polynoe Notosetae distinctly slenderer than the neurosetae 24 At least some notosetae as thick as or thicker than the neurosetae 27 Ventral surface with paired folióse appendages Phyllohartmania Ventral surface smooth 25 Most neurosetae bidentate Arcteobia All neurosetae unidentate 26 Neurosetae slender. 23a). 38b). 16b). (39b).. Neurosetae with semilunar pockets Adyte Neurosetae spinose or smooth. 35b). (42a). 18b 17b).or bidentate 29 Neurosetae with semilunar pockets Paradyte Neurosetae spinose or nearly smooth 30 Notosetae with distal tufts of long hairs Barrukia Notosetae without distal tufts of hairs 31 Neurosetae distally with capillary tips 32 Neurosetae distally falcate. 24b). 22a 21b). 23b). 37b). 48b (47b). then neuropodia short with ventral cirri basal 57 Twelve or 13 pairs of elytrae 58 More than 13 pairs of elytrae 66 With branchial filaments on the elytrophores 59 Branchial filaments absent 61 With pseudelytrae on non-elytrophoral. 47b (46a). Eighteen pairs of elytrae Acanthicolepis Nineteen or more pairs of elytrae 46 Notosetae distinctly slenderer than neurosetae 47 Notosetae as thick as. 67b (66a). neuropodium prolonged with the ventral cirrus attached near the middle Frennia Notopodia usually present. or thicker than neurosetae 49 Neurosetae with semilunar pockets Pottsiscalisetosus Semilunar pockets absent 48 Neuropodia with subacicular digitiform lobes Parahotalepidella Neuropodia with large. 66b (57b). 46a (45b). 63a (62b). 60b (59b). (55b).58 45a (42b). 62a (61b). (54b). 50b (49b). 60a (59b). (54b). 69a (68b). 59a (58a). lancet-shaped and tapering and serrated Thormora Notosetae all of one kind. 57b (56b). 69b (68b). alternating segments posteriorly Euphionella Pseudelytrae absent 60 Neurosetae coarsely spinose Chaetacanthus Neurosetae finely plumose Euphione Notopodia absent Drieschia Notopodia present 62 With one very large bidentate hook in the middle of each neuropodial fascicle Sheila All neurosetae similar in thickness. 49b (46b). NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 56b (55b). 71b (70b). 50a (49b). 51b (4b). 65a (64b). if absent. rarely trifid. 45b (42b). none bidentate hooks 63 Neurosetae distally trifúrcate Hermenia Neurosetae distally entire 64 Notosetae of two kinds. 64b (63b). 46b (45b). (52b). 63b (62b). supra-acicular presetal lobes Grubeopolynoe All neurosetae unidentate Polyeunoa At least some neurosetae bidentate 50 Neurosetae with long bare tips beyond the serrated regions Hololepidella Neurosetae with very short bare tips beyond the serrated region Neohololepidella Palps represented by two small tubercles Bouchiria Palps well developed 52 Ventral cirrus heavily fimbriated. 49a (46b). 70b (69a). 57a (56b). ventrum with transverse rows of three papillae on each segment Lepidofimbria Ventral cirrus and ventrum smooth 53 Neurosetae distally bifid with several small accessory teeth in the crotch Lepidogyra Neurosetae unidentate or bifid. 52a (51b). 61b (58b). 64a (63b). 62b (61b). 68a (66b). if bifid. 47a (46a). 61a (58b). 70a (69a). 58a (57a). 65b (64b). 58b (57a). or thicker than the neurosetae 71 All neurosetae unidentate Eulagisca At least some neurosetae bidentate 72 . 67a (66a). 68b (66b). usually slender and serrated 65 Pseudelytrae on non-elytrophoral segments Dilepidonotus Pseudelytrae absent Lepidonotus Fourteen pairs of elytrae 67 Fifteen or more pairs of elytrae 68 Neurosetae bidentate Lucopia Neurosetae entire Podarmus Only anterior half of body covered by elytrae Pseudopolynoe Elytrae on most parts of the body 69 Fifteen pairs of elytrae 70 Sixteen or more pairs of elytrae 74 Notosetae thin Parahalosydna Notosetae as thick as. (52b). 71a (70b). 48a (47b). 59b (58a). then without accessory teeth 54 Prostomium anteriorly produced into two large lobes with small lateral antennae attached distally Allmaniella Prostomium anteriorly tapering into the ceratophores 55 Renal papillae greatly prolonged in some median setigers Bathymoorea Renal papillae of similar length in all setigers 56 Notopodia completely reduced. (53b). 66a (57b). (53b). 51a (4b). 52b 53a 53b 54a 54b 55a 55b 56a (51b). HARMOTHOINAE. only species. Antennal scales at base of lateral antennae. neurosetae numerous 89 Twenty-one or 22 pairs of elytrae. 72b (71b). Acholoe Claparède 1870b. Neurosetae bidentate with slender secondary tooth. marginally serrated with long. up to 30 pairs of elytrae Admetella Setae flattened.]976 72a (71b). 76a (75a). 85b (84b). 74b (69b). curved tips Hyperhalosydna Numerous pairs of elytrae. 84a (83a). Notosetae thicker than neurosetae. 77a (76a). Dorsal cirri and antennae papillated. 82a (81b). 92b (91a). Polynoe asperrima Sars 1861. LEPIDONOTINAE. 83a (82b). neurosetae with long. neurosetae long. neurosetae few Arctonoe Lateral anteimae terminal. 73a (72b). 80a (75b). Neurosetae few.. finely dentate and very abruptly tapering. lateral antennae terminal Haiosydna Only few notosetae present. serrated. with transverse rows of spines. lateral antennae subterminal Alentia Notosetae present 81 Notosetae absent 90 Parapodia distally with series of long papillae Halosydnopsis Parapodia distally without papillae 82 Neurosetae with semilunar pockets Hololepida Neurosetae spinose or smooth 83 Notosetae thinner than most neurosetae 84 Notosetae as thick as. 75 segments. 75a (74b). A. 88b (86b). 2 species. 92a (91a). approx. 73b (72b). 93b (91b). flattened. pre. unidentate with serrated subdistal areas. 90b (80b). finely Acanthicolepis Mclntosh 1900. 89a (88b). 79b (76b). 75b (74b). straight or curved tips Lepidasthenia Ventral cirri with two-three knobs Perolepis Ventral cirri smooth 91 Antennal scales present 92 Antennal scales absent 93 Setae flattened. 23 pairs of elytrae Bathyadmetella Neurosetae stout. 93a (91b). 86b (83b). 89b (88b). Notosetae thinner than neurosetae. with whorls of teeth. 78a (77b). 74a (69b). 85a (84b). 80b (75b). Numerous segments and elytrae. neurosetae with short. llh (76a). 88a (86b). Cirriferous segments with T-shaped papillae. 91b (90b). Nereis squamosa delle Chiaje 1828. . Notosetae absent. Benhamipolynoe Neurosetae slender. 87b (86a). Admetella Mclntosh 1885. 91a (90b). 87a (86a). longipedata Mclntosh 1885. 82b (81b). 78b (77b). . slender tips. falcate. 40 segments.and postsetal neuropodial lobes of similar length . THE POLYCHAETE WORMS 59 Dorsal cirri of two kinds. short and expanded or long and gently inflated Malmgreniella All dorsal cirri long and slightly inflated subdistally 73 Notosetae nearly smooth Malmgrenia Notosetae densely serrated Paralepidonotus With 16 pairs of elytrae Arctonoella At least 17 pairs of elytrae 75 Seventeen to 20 pairs of elytrae 76 At least 21 pairs of etytrae 80 Notosetae absent 77 Notosetae present 79 Dorsum with a median pustule on each segment Weheria Dorsal pustules absent 78 Neurosetae with capillary tips Drieschella Neurosetae with falcate tips Alentiana Notosetae numerous. presetal lobes distinctly longer than the postsetal ones in the neuropodia Telolepidasthenia Generic Definitions LEPIDONOTINAE. 81b (80a). 79a (76b). 86a (83b). Up to 30 pairs of elytrae. or thicker than the neurosetae 86 Superior neurosetae slender and spinose. 81a (80a). 84b (83a). Eighteen pairs of elytrae. 76b (75a). inferior ones acicular Pseudohalosynda All neurosetae stout 85 At least some neurosetae bidentate Halosydnella All neurosetae unidentate Acholoe All neurosetae unidentate 87 At least some neurosetae bidentate 88 Ventrum with large segmentally arranged lamellae Gastrolepidia Ventrum smooth Lepidastheniella Lateral antennae subterminal. 83b (82b). 90a (80b). 3 species. HARMOTHOWAE. thick and falcate. Antinoana Hartman and Fauchald 1971. only species. with the distal end pencillate. Neurosetae unidentate. Lateral antennae subterminal. with a few coarse teeth on the cutting edge. serrated Some notocirri with large. only species. Neurosetae distally with slender. only species. Antinoe sarsi Kinberg in Malmgren 1865. 8 species. filamentous tips. Polynoe gelatinosa Sars 1835. Allmaniella Mclntosh 1885. Alentiana Hartman 1942. LEPIDONOTINAE. Seventeen to 20 pairs of elytrae. LEPIDONOTINAE. Barrukia Bergström 1916. Notosetae thicker than neurosetae with transverse rows of teeth. Fifteen pairs of elytrae. fusca Hartman and Fauchald 1971. spherical ampullae subdistally. 2 species. Fifteen to 16 pairs of elytrae. 35 segments. Bathyadmetella Pettibone 1967. numerous segments. only species. Neurosetae few. Upper neurosetae sharp-tipped. Lateral antennae subterminal. Polynoe rutilans Grube 1878. 36 segments. B. Neurosetae slender and bifid. A. Neurosetae thin. with slender. only species. HARMOTHOINAE. Eighteen pairs of elytrae. Arctonoella Buzhinskaya 1967. A. nearly capillary setae. slender. Neuropodia prolonged. A. Neurosetae unidentate or bidentate. Sixteen pairs of elytrae. 26-27 segments. commando Pettibone 1967. ampullifera Prenant 1924: only species. Notosetae slenderer than neurosetae. Polynoe vittata Grube 1855. Arcteobia Annenkova 1937. HARMOTHOINAE. 4 species. 40-43 segments. Neuropodia prolonged. Thirteen to 14 pairs of elytrae 32-33 segments. approximately 40 segments. Notosetae absent. bidentate and vaguely serrated. Twelve to 13 pairs of elytrae. smooth or faintly serrated. 43 segments. Lateral antennae subterminal. LEPIDONOTINAE. often pilose. Notosetae thicker than neurosetae. otherwise blunt and serrated. A. HARMOTHOINAE. Notosetae thick and finely serrated. neurosetae of two kinds. Andresia Prenant 1924.or bidentate with coarse serrations. Notosetae thicker than neurosetae. serrated or smooth. Notosetae thinner than neurosetae.60 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 marginally serrated with smooth tips. LEPIDONOTINAE. Fifteen pairs of elytrae. Antinoella Augener 1928b. 58 segments. smooth with few spines. Notosetae few. HARMOTHOINAE. Notosetae thicker than neurosetae. 7 species. 6 species. long posterior region without scales. finely dentate and distally very abruptly tapering. but less than 30 pairs of elytrae. Fifteen pairs of elytrae. Austrolaenilla Bergström 1916. 2 species. 3 species. uni. others bidentate. HARMOTHOINAE. distally capillary. Twenty-three pairs of elytrae. Fifteen pairs of elytrae. usually bifid. 36-39 segments. Adyte Saint-Joseph 1899.or bidentate. neurosetae . Gattyana cristata Willey 1902. Neurosetae unidentate. Notosetae absent. HARMOTHOINAE. More than 15. Prostomium anteriorly produced into two large lobes with the small lateral antennae attached distally. recurved uni. thick. LEPIDONOTINAE. Notosetae short and slender. Polynoe aurantiaca Verrill 1885. Forty or more pairs of elytrae. 5 species. Neurosetae with slender tips. A. 35-36 segments. Alenda Malmgren 1865. microps Kinberg 1855. HARMOTHOINAE. vaguely serrated. Notopodia reduced with a few. Neurosetae unidentate. Hertnadion assimile Mclntosh 1876. flattened setae. fine hairs. only species. upper slender and smooth. Notosetae distally with tuft of long. Fifteen pairs of elytrae. LEPIDONOTINAE. Australaugeneria Pettibone 1969d. Notosetae at least as thick as neurosetae. Presetal neuropodial lobes longer than postsetal ones. Notosetae thicker than neurosetae coarsely serrated. Neurosetae with semilunar pockets and faintly bifid tips. Antinoe Kinberg 1855. approximately 40 segments. otherwise uni. antárctica Bergström 1916. with blunt or capillary tips. Arctonoe Chamberlin 1920. Bathymoorea Pettibone 1967. 41 segments. LEPIDONOTINAE. Fifteen pairs of elytrae. Neurosetae varied: as large golden hooks in the second and third segment. 2 species. Notosetae thicker than neuro- setae with rows of spines. Eupolynoe anticostiensis Mclntosh 1874. lower thicker. setubalensis Mclntosh 1885. serrations faint. Numbers of elytrae not known. often spurred and subdistally inflated. Harmothoe sinagawaensis Izuka 1912. but not Glamentously prolonged. HARMOTHOINAE. Neurosetae unidentate. transversely serrated. blunt. Polynoe Trenotuberculata Moore 1910.or bidentate. notopodia reduced. Dilepidonotus Hartman 1967. Iphione magnifica Grube 1875. unidentate and laterally covered with fine hairs. Euphione Mclntosh 1885. but a few bidentates also present.1976 THE POLYCHAETE WORMS 61 thicker. Fifteen pairs of elytrae. 36^0 segments. only species. D. only species. 37 segments. Notosetae absent. F. 6 species. Notosetae capillary. Enipo Malmgien 1865. Fifteen pairs of elytrae. Neurosetae slender and tapering to capillary tips. Numbers of pairs of elytrae and segments not known. neurosetae unidentate and spinose. Notosetae and neurosetae similar in thickness. neurosetae thicker. neurosetae unidentate and serrated. Notosetae slender. E. Prostomium round. Twelve pairs of elytrae. pseudelytrae present. shortbodied. E. Notosetae slender and capillary. 47 segments. Two antennae. D. 26 segments. neurosetae all unidentate. 6 species. Notosetae mainly capillaries. only species. Twelve to 13 paire of elytrae. Bylgides Chamberlin 1919c. 2 species. HARMOTHOINAE. smooth capillaries. with more or less well-marked spinose region. Some neurosetae distally split with both parts about equally long and thick. neurosetae thicker. Notosetae thicker than neurosetae with rows of spines. neurosetae unidentate and thicker. unidentate and smooth. Dorsum of cirral segments crested. Bylgia elegans Theél 1879. neuropodia strongly prolonged with ventral cirri attached near the middle. LEPIDONOTINAE. numerous segments. Neurosetae thick. Fifteen pairs of elytrae. lower thicker and slightly serrated. Frennia Viguier 1912. dubia Viguier 1912.£. Notopodia absent. 40 species. Notosetae capillary. Notosetae coarser than neurosetae. More than 21 pairs of elytrae. pectinate. LEPIDONOTINAE. Notosetae thicker than neurosetae with rows of teeth. Lepidasthenia antipathicola Benham 1927. Sub-family unknown. Benhamipolynoe Pettibone 1970d. Notosetae fine and capillary. mainly unidentate. 3 species. approximately 40 segments. neurosetae slender and tapering to capillary tips. Anteiuiae and dorsal cirri strongly inflated subdistally. HARMOTHOINAE. E. Notopodia completely reduced. All setae slender. Drieschella Augener and Pettibone in Pettibone 1970d. Notosetae slender and silky capillaries. a few thicker blunt setae present. facial tubercle absent. LEPIDONOTINAE. Fifteen pairs of elytrae. Eunoe Malmgren 1865. Three antennae. Ventrum with large lamellae. Euphionella Monro 1936. only species. only species. Notosetae absent. but otherwise similar setae ventrally. LEPIDONOTINAE. only species. neurosetae thicker. posterior half of body not covered by elytrae. 3 species. More than 17 pairs of elytrae. falcate with weakly marked serrations. Elytrophores with branchiae. long. Presetal lobes longer than postsetal ones. D. E. Physalidonotus lobulatus Seidler 1922. kinbergi Malmgren I86S. LEPIDONOTINAE. elisabethae Mclntosh 1885. clavigera Schmarda 1861. 26 setigers. smooth and pointed. Renal papillae greatly prolonged in some median setigers. Bouchiria Wesenberg-Lund 1949. LEPIDONOTINAE. 4 species. numerous. Neurosetae thicker than notosetae. Eulagisca Mclntosh 1885. IPHIONINAE. LEPIDONOTINAE. LEPIDONOTINAE. acicular lobes not projecting. japónica Frickinger 1916. bidentate and marginally serrated. with slender tips. LEPIDONOTINAE. Elytrophores with branchiae. Pseudelytrae present. Twenty pairs of small elytrae. facial tubercle present. LEPIDONOTDJAE. only species. other neurosetae slender and unidentate. upper neurosetae capillary. Palps reduced to two small tubercles. Twelve pairs of elytrae. 25 segments. HARMOTHOINAE. Cervilia Frickinger 1916. oerstedi Malmgren 1865. Elytrophores with branchiae. eyes conspicuous. vesiculosa Wesenberg-Lund 1949. Chaetacanthus Seidler 1924. . Twelve pairs of elytrae. 3 species. C. neurosetae include slender capillary setae with long slender spines and thicker more acicular. pelágica Michaelsen 1892. thicker than the neurosetae. corrientis Mclntosh 1885. maculata Augener and Pettibone in Pettibone 1970d. LEPIDONOTINAE. 26-27 segments. Drieschia Michaelsen 1892. B. G. lateral antennae attached posteriorly on the prostomium. Nimibers of pairs of elytrae and segments not known. falklandicus Hartman 1967. Notosetae absent. Lateral antennae subterminal. 4 species. Eucranta Malmgren 1865. Gastrolepidia Schmarda 1861. notosetae blunt and serrated. Fifteen pairs of elytrae. Numerous stalked papillae present on the parapodia. Twelve pairs of elytrae. villosa Malmgren 1865. but usually also some unidentate in inferior positions. 3 species. Nuchal flap present. HARMOTHOINAE. short. of two kinds.62 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Gattyana Mclntosh 1900. thick serrated and blunt and slender. nearly smooth with blunt tips. pointed and serrated. Hermenia Grube 1856. Malmgrenia crassicirris Willey 1902. H. lanceolate and bidentate with transverse rows of pockets. neurosetae thicker. Neurosetae in part distally trifid. Polynoe ascidioides Mclntosh 1885. Hololepidella Willey 1905. HARMOTHOINAE. LEPIDONOTINAE. magalhaensi Kinberg 1855. Notosetae ñner than neurosetae. Grubeopolynoe Pettibone 1969b. 58 segments. Fifteen pairs of elytrae. Notosetae thicker than neurosetae and strongly serrated. only species. of two kinds. small. approximately 40 segments. LEPIDONOTINAE. 50 segments. Eight to 9 pairs of elytrae. short and blunt and slender and tapering. HARMOTHOINAE. Most notosetae capillary with fine dentition. LEPIDONOTINAE. Halosydnella Hartman 1938. Fifteen pairs of elytrae. vaguely bidentate or unidentate. numerous segments. HARMOTHOINAE. 36-38 segments. nordgaardi Bidenkap 1907. 4 species. only species. Herdmanella Darboux 1899. Polynoe tuta Grube 1855. HARMOTHOINAE. Notosetae held erect over body. curved . pointed tips. Twelve pairs of elytrae.or bidentate. approximately 40 segments.or unidentate. 14 species. Notosetae at least as thick as neurosetae. Notosetae in part at least as thick as neurosetae. Notosetae few in numbers.or bidentate. Notosetae few. more than 50 segments. slender and serrated. 4 species. 14 species. Notosetae as thick as neurosetae. with rows of spines. Eighteen pairs of elytrae. Neurosetae tapering to sharp. Notosetae thicker than neurosetae. body covered. HARMOTHOINAE. HARMOTHOESTAE. serrated. only species. Neurosetae all of one kind. 40 segments. Neurosetae distally bi. 5 species. H. Fifty or more pairs of elytrae. 120 species. distally unidentate. Fifteen pairs of elytrae. 38-40 segments. HARMOTHOINAE. 2 species. Distal end of parapodia with series of large papillae. parapodia prolonged. of two kinds. H. HARMOTHOINAE. with coarse serrations. neurosetae distally trifúrcate. 55 or mote segments. Halosydnopsis Uschakov and Wu 1959. moorei Pettibone 1955. serrated. Notosetae absent. capillary. Fifteen pairs of elytrae. neurosetae less so. Twenty-six or more pairs of elytrae. LEPIDONOTINAE. Notosetae slenderer than neurosetae. Hemilepidia Schmarda 1861. Hermadion Kinberg 1855. H. patagónica Kinberg 1855. and serrated. 45 segments. serrated and pointed. magna Moore 1905.//. Neurosetae thicker than notosetae. Harmothoe Kinberg 1855. Neurosetae all unidentate. Twenty-seven pairs of elytrae. usually slender superior and thick in inferior positions. commensalis Willey 1905. Numerous pairs of elytrae. Notosetae much fmer than neurosetae. Hyperhalosydna Augener 1922. Hesperonoe Chamberlin 1919. tapering to sharp tips. Notosetae few. Notosetae fmer than neurosetae. tuberculata Schmarda 1861. Neurosetae uni. Aphrodita cirrhosa Pallas 1766. Gorekia Bergström 1916. 3 species. not overlapping. only species. serrated. Fifteen pairs of elytrae. Halosydna Kinberg 1855. both conspicuously long and unidentate. LEPIDONOTINAE. HARMOTHOINAE. Notosetae spinose. Halosydna australis Kinberg 1855. numerous segments. Neurosetae distally uni. H. LEPIDONOTINAE. Halosydna pilosa Horst 1917. posterior part of body not covered by elytrae. serrated. Neurosetae at least in part bidentate. Neuropodia with long supraacicular postsetal lobe. spinosa Kinberg 1855. Heteropolynoe Bidenkap 1907. Hartmania Pettibone 1955. Lepidonotus striatus Kinberg 1855. finely serrated. only species. Fifteen pairs of elytrae. 9 species. H. verruculosa Grube 1856. Notosetae fmer than neurosetae. 15-17 segments. Fifteen pairs of elytrae. Numbers of pairs of elytrae not known. uni. vaguely serrated.or bidentate. Twenty-one pairs of elytrae. all neurosetae unidentate and marginally serrated. finely serrated. Harmothoe complánala Johnson 1901. not falcate. Neurosetae thick. subdistally serrated. slender in superior and coarse in inferior positions. H. HARMOTHOINAE. Twenty-one to 22 pairs of elytrae. Ä. 37 segments. neurosetae nearly smooth and unidentate. 2 species. Neurosetae distally bidentate or entire. Notosetae shorter and thicker than the neurosetae. 5 species. Hololepida Moore 1905. covering the body. Posterior 8-10 segments not covered by elytrae. antárctica Uschakov 1957. HARMOTHOINAE. Polynoe nivea Sars 1863. Lateral antennae subterminal. M. LEPIDONOTINAE. Macelloides Uschakov 1957. Fifteen pairs of elytrae. Numbers of pairs of elytrae and segments not known. M. shortbodied. Fourteen pairs of elytrae. only species. Twelve pairs of elytrae. Eight to 13 pairs of elytrae. L. dicirra Hartman 1967. LEPIDONOTINAE. 30 segments. only species. neurosetae bidentate and serrated. Ventrum with transverse row of three papillae on each segment. only species. serrated. LEPIDONOTINAE. M. Notosetae few or absent. most are thicker than notosetae except inferior ones in each fascicle. unidentate and serrated. 4 species. notosetae smooth-tipped. Lateral antennae terminal. M. IPHIONINAE. oculata Hartman 1967. more distinctly serrated. the short ones basally inflated. neurosetae smooth and distally entire. Notosetae thinner than neurosetae. Dorsal cirri alternating long and short. only species. Numerous pairs of elytrae and segments. Lepidogyra Hartman 1967. M. Dorsal cirri strongly inflated. Notosetae coarser than neurosetae and serrated. nearly smooth. Macellicephaloides Uschakov 1955 b. Dorsal cirri of two kinds. LEPIDONOTINAE. notosetae falcate. 16-17 segments. weakly serrated. Notosetae few. usually paddle-shaped. Iphione Kinberg 1855. notosetae capillary. 41-56 segments. Lepidastheniella Monro 1924. L. Malmgrenia Mclntosh 1874. Neurosetae with rows of coarse teeth. 3 species. neurosetae uni. Fifteen pairs of elytrae.and neurosetae similar in thickness. magnicirra Pillai 1965. Lepidonotus Leach 1816.or bidentate with very small secondary teeth. slender. neurosetae spinose. HARMOTHOINAE. L. MACELLICEPHALINAE. all tapering with whorls of spines. HARMOTHOINAE. usually blunt. Fifteen pairs of elytrae. MACELLICEPHALINAE. M. rarely bidentate. 65 species. only species.or bidentate. 24 species. approximately 40 segments. LEPIDONOTINAE. Noto. Sixteen pairs of elytrae. Aphrodita clava Montagu 1808. Eyes absent. Numbers of pairs of elytrae and segments not known. 26 segments. only species. Noto. Notosetae aosent. curved tips. short body. only species. only species. Up to 90 pairs of elytrae. Leucia Malmgren 1867. usually unidentate. Fifteen pairs of elytrae. Neurosetae spinose and distally entire. 27 segments. 36-41 segments.and neurosetae similar in thickness. distally unidentate. Polynoe (Langerhansia) euplectellae Mclntosh 1885. . Notosetae with dense rows of teeth. Polynoe rarispina Sars 1861. distally bind with a series of accessory teeth in the crotch between the two major teeth. ringed with spines. Macellicephala Mclntosh 1885. slender secondary tooth. LEPIDONOTDslAE. Lepidasthenia Malmgren 1867. Notosetae absent. Noto. Lateral antennae terminal. Fifteen pairs of elytrae. 18 species. uni. Maximally 9 pairs of elytrae. Neurosetae bidentate with long. Kermadecella Darboux 1899. Neurosetae distally inflated. neurosetae serrated and distally entire. Neurosetae with transverse rows of spines. 9 species. Notosetae finer than neurosetae. neurosetae unidentate. Neurosetae long. Neurosetae long. marginally dentate. all neurosetae serrated. Polynoe magnipalpa Mclntosh 1885. grandicirra Uschakov 1955b. Noto. ventral cirrus heavily fimlniated. approximately 50 segments. Notosetae nearly smooth.and neurosetae similar in thickness. Notosetae absent. Notosetae absent. if present then with marginal teeth. Polynoe muricata Savigny 1818.1976 THE POLYCHAETE WORMS 63 and blunt.and neurosetae about equally thick. Lepidofimbria Hartman 1967. Prostomium with two very strongly inflated lobes. Maimgreniella Hartman 1967. only species. HARMOTHOINAE. MACELLICEPHALINAE. Lucopia Pillai 1965. long and slender and short and expanded. lovent Malmgren 1865. Thirteen pairs of elytrae. Polynoe comma Thomson 1902. neurosetae numerous. Polynoe elegans Grube 1840. Intoshella Darboux 1899. Notosetae thicker than neurosetae. LEPIDONOTINAE. Melaenis Malmgren 1865. Notosetae coarser than neurosetae. alba Hartman 1967. neurosetae at least in part bidentate. LEPIDONOTESIAE. 17-29 segments. 37 species. 3 species. Neurosetae long. Neurosetae bidentate with long. mirabilis Mclntosh 1885. Lagisca Malmgren 1865. Fifteen pairs of elytrae. whiteavesi Mclntosh 1874. but a few thick spines present. Polynoe ampullifera Grube 1878. Polynoella Mclntosh 1885. regularis Ehlers 1908. taylori Pettibone 1961.64 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 HARMOTHOINAE. less than 50 segments. only species. Neurosetae unidentate and serrated along both edges. Fifteen pairs of elytrae. subdistally expanded and dentate. 30 segments. Fifteen pairs of elytrae. Twenty-eight or more pairs of elytrae. Ventral surface with paired folióse appendages on each segment. Paradyte Pettibone 1969a. Pseudohalosydna Fauvel 1913. Nineteen to 30 pairs of elytrae. . nearly smooth. Notosetae absent. only species. Notosetae absent. Notosetae finer than neurosetae. HARMOTHOINAE. but a few capillary setae present. thick and straight or slender and capillary. serrated. less than 40 segments. 3 species. Fifty or more pairs of elytrae. Numerous pairs of elytrae and segments. nearly smooth. Nearly all neimisetae bidentate. neurosetae all distally entire. except usually one or two unidentate in each of the posterior setigers. Twelve pairs of elytrae. A'. Fourteen pairs of elytrae. neurosetae mainly unidentate. spinose. Fifteen pairs of elytrae. Notosetae thicker than neurosetae. pointed and distally bifid. LEPIEXDNOTINAE. HARMOTHOINAE. Neurosetae at least in part bidentate. slender. Pottsiscalisetosus Pettibone 1969a. 4 species. sibogae Horst 1915a. Notosetae mainly thick and blunt-tipped. serrated. Neurosetae unidentate. 2 species. but still thick. levisetosa Mclntosh 1885. ventral surface papiUated. Fifteen pairs of elytrae. Neurosetae very thick. spinose and with bifid tips. Notosetae absent. LEPIDONOTINAE.and postsetal lobes of similar length. P. Polynoe crinoidicola Potts 1910. distally bidentate or entire. Notosetae slender and spinose with capillary tips. rosea Fauvel 1913. At least 20 pairs of elytrae. P. slender. pre. HARMOTHOINAE. HARMOTHOINAE. Notosetae thinner than neurosetae. murrayi Pettibone 1969b. Neurosetae of two kinds: numerous slender dentate with capillary tips. Notosetae thicker than the neurosetae. 2 species. HARMOTHOINAE. Perolepis Ehlers 1908. 17 species. only species. tapered to blunt tips. with semilunar pockets and entire tips. 4 species. unidentate and falcate. posterior part of body without elytrae. Notosetae spinose and capillary. Fifteen pairs of elytrae. with blunt tips. HARMOTHOINAE. few in number. Neurosetae smooth and distally bidentate. P. Podarmus Chamberlin 1919c. neurosetae distally bidentate. Ventral cirri folióse. HARMOTHOINAE. P. Fifteen pairs of elytrae. Notosetae thicker than neurosetae. slightly hooked. nmnerous segments. only species. densely serrated. Neuropodia long. P. Paralepidonotus Horst 1915a. Scatisetosus praelongus Marenzeller 1902. 3 species. Subacicular ones thicker. Numerous pairs of elytrae and segments. 9 species. laevis Mclntosh 1885. Neurosetae with very short bare tip beyond a dense spinose region. HARMOTHOINAE. Neurosetae long. HARMOTHOINAE. wg/fyi Pettibone 1961. eyes missing or strongly reduced. Neuropodia with subacicular digitiform process. of two kinds. faintly serrated. P. 38 segments. Nemidia Malmgren 1865. P. LEPIDONOTINAE. most neurosetae coarser than the notosetae. Fifteen pairs of elytrae. sabrelike with entire or slightly notched tips. HARMOTHOINAE. Neurosetae of two kinds: supracicular ones with semilunar pockets. posterior part of body not covered by elytrae. posterior part of body not covered by elytrae. smooth or faintly structured. P. distally spinose and spinigerous. Prostomium quadrangular. few furcates with subequal blunt tips. Parahalosydna Horst 1915a. neurosetae similar in thickness. Hololepidella greeffii Augener 1918. shortbodied. Notosetae thicker than neurosetae. Ventral cirri with three knobs. but a few bidentate setae present. Notosetae mainly capillaries. numerous segments. pica Chamberlin 1919c. numerous segments. distally entire and more or less falcate. P. N. LEPIDONOTINAE. Neurosetae with semilunar pockets. 40 segments. 39-41 segments. Phyllosheila Pettibone 1961. Parahololepidella Pettibone 1969b. 25-26 segments. Polynoe Savigny 1818. only species. cirrophores enlarged dorsally. Notosetae thicker than neurosetae. scolopendrina Savigny 1818. torelli Malmgren 1865. Notosetae slenderer than neurosetae. 2 species. Phyllohartmania Pettibone 1961. Fourteen pairs of elytrae. entire and very faintly spinose. LEPIDONOTDJAE. Polyeunoa Mclntosh 1885. only species. Notosetae thicker than neurosetae. Neohololepidella Pettibone 1969b. 1976 THE POLYCHAETE WORMS 65 neurosetae of two kinds; superior ones slender and spiralled spinose; inferior ones acicular and spinose. Pseudopolynoe Day 1962, Polynoe inhacae Day 1951; only species. LEPIDONOTINAE. Fifteen to 17 pairs of elytrae; posterior half of body not covered by elytrae. Notosetae finer than neurosetae, serrated; neurosetae bi- or unidentate, serrated. Robertianella Mclntosh 1885, R. synophthalma Mclntosh 1885; only species. HARMOTHOINAE. Thirteen pairs of elytrae; approximately 30 segments. Noto- and neurosetae of similar thickness; notosetae blunt, nearly smooth; neurosetae distally minutely notched, nearly smooth. Eyes very large and nearly confluent on the prostomium. Scalisetosus Mclntosh 1885, S. ceramensis Mclntosh 1885; 8 species. HARMOTHOINAE. Sixteen pairs of elytrae; 40 segments. Notosetae much coarser than neurosetae, with a few spines only. Neurosetae slender, distally entire and spinose. Both noto- and neuropodia with long lobes. Sheila Monro 1930, S. bathypelagica Monro 1930; only species. LEPIDONOTINAE. Thirteen pairs of elytrae; 29 segments. Notosetae slender, dentate capillaries; neurosetae of several kinds: numerous superior dentate capillaries; most of the others coarse, dentate and entire; one very large bidentate hook in the middle of each fascicle. Subadyte Pettibone 1969a, Polynoe pellucida Ehlers 1864; 3 species. HARMOTHOINAE. Fifteen to 16 pairs of elytrae; approximately 40 segments. Notosetae similar in thickness to the neurosetae; with spinose pockets and slightly notched tips. Neurosetae with semilunar pockets, spinose and distally bidentate. Telolepidasthenia Augener and Pettibone in Pettibone 1970d, T. lobetobiensis Augener and Pettibone in Pettibone 1970d; only species. LEPIDONOTINAE. More than 21 pairs of elytrae and more than 50 segments. Notosetae absent; neurosetae slender with spinose regions and entire tips. Presetal lobes longer than postsetal ones. Tenonia Nichols 1969, T. kitsapensis Nichols 1969; only species. HARMOTHOINAE. Fifteen pairs of elytrae; approximately 40 segments. All setae similar in thickness; all slender; notosetae capillary, neurosetae in part bidentate. Thormora Baird 1865, T.jukesii Baird 1865; 5 species. LEPIDONOTINAE. Twelve pairs of elytrae; short- bodied. Notosetae of two kinds; short and lancetshaped, and longer, serrated and tapering. Neurosetae serrated and unidentate. Uncopolynoe Hartmann-Schröder 1960a, £/. comllicola Hartmann-Schröder 1960a; only species. HARMOTHOINAE. Numbers of elytrae not known, approximately 44 segments. Notopodia absent; first three neuropodia with strongly curved hooks; others with uni- or bidentate setae with serrated subdistal areas. Weberia Horst 1915b, W. pustulata Horst 1915b; 3 species. LEPIDONOTINAE. Eighteen pairs of elytrae; shortbodied. Notosetae absent; neurosetae unidentate and curved. Ventral cirri absent except in second segment; dorsum with a pustule on each segment. Invalid Genera Agnodice Mclntosh 1885, see Lagisca Bathynoe Ditlevsen 1917, see Weberia Bylgia Theél 1879, see Bylgides Chaetosphaera Haecker 1898, larvae of several genera Dasylepis Malmgren 1867, see Acanthicolepis Eumolpe Oken 1807, see Lepidonotus Eupolynoe Mclntosh 1874, see Eucrunw Evarne, Malmgren 1865, see Harmothoe Evarnella Chamberlin 1919c, see Harmothoe Gastroceralella Darboux 1899, ^ee Thormora Halosydnoides Seidler 1924, see Arctonoe Harmopsides Chamberlin 1919c, see Lepidasthenia Iphionella Mclntosh 1885, seelphione Laenilla Malmgren 1865, see Harmothoe Langerhansia Mclntosh 1885, see Intoshella Lepidametria Webster 1879b, see Lepidasthenia Nectochaeta Marenzeller 1892, in part Lepidasthenia, also generally polynoid larvae and juveniles Norepa Baird 1865, seelphione Nychia Malmgren 1865, see Gattyana Oligolepis Levinsen 1887, see Macellicephala Paranychia Czemiavsky 1882, questionably la^iico Parapolynoe Czemiavsky 1882, see Polynoe Parmensis Malmgren 1867, see Harmothoe Pkysalidonotus Ehlers 1905, see Euphione Plotolepis Chamberlin 1919c, see Drieschia Quetieria Viguier 1911, juvenile form Tricosmochaeta Morgera 1918, see Harmothoe FAMILY POLYODONTIDAE BUCHANAN 1894 Aphroditaceans with dorsoventrally flattened bodies. Two or three antennae present, median antenna when 66 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 present, attached dorsally or posteriorly on the prostomium (antennae may be absent). Eversible pharynx with four jaws. Marginally fringed or smooth elytrae alternate with dorsal cirri in all setigers. All setae simple. Spinning glands present. The polyodontids are large, solid-bodied scaleworms, usually covered with thick, loosely constructed tubes consisting of thin threads filled with clay or sand particles. The most important recent revision was made by Strelzov (1968b). Key to Genera la. lb. 2a (la). 2b (la). 3a (2a). 3b (2a). 4a (2b). 4b (2b). 5a (lb). 5b (lb). Ommatophores present Ommatophores absent Two antennae present Three antennae present Ommatophores fused in the midhne Ommatophores separated from each other Superior neurosetae long and tapering, more or less hirsute Superior neurosetae short and brush-topped Three antennae present Antennae absent 2 5 3 4 Neopanthatis Eupolyodontes Polyodontes Panthalis Eupanthalis Resno Generic Definitions Eupanthalis Mclntosh 1876, E. kinbergi Mclntosh 1876; 8 species. Ommatophores absent, three antennae present; median antenna attached dorso-posteriorly. Setae include slender, pilose tapering capillaries; bluntly acicular, aristate ones with pilose shafts, and side-shaped, dentate ones. Eupolyodontes Buchanan 1894, E. cornishii Buchanan 1894; 5 species. Ommatophores present and separated from each other. Two frontal antennae present; small nuchal papilla present. Setae include bipinnate pencillate setae; serrulate setae and smooth, acicular spines. Neopanthalis Strelzov 1968b, N. pelamida Strelzov 1968b; only species. Ommatophores present but fused in the midline. Lateral antennae short and inserted dorsally on the ommatophores. Panthalis Kinberg 1855, P. oerstedii Kinberg 1855; 12 species. Ommatophores present and separated; three antennae present. Setae include short brush-topped setae; thick, blunt acicular setae with rows of teeth subdistally and dentate sicle-shaped setae. Polyodontes Renier in Audouin and Milne Edwards 1832, Phyllodoce maxillosa Ranzani 1817; 12 species. Ommatophores present and separated; three antennae present. Setae include long, tapering hirsute ones; stout acicular setae with or without aristae, usually hirsute or dentate; and dentate sickle-shaped setae. Restio Moore 1903, R. aenus Moore 1903; only species. Ommatophores and antennae absent. Setae include long, slender ones with awnlike spines, thick, colorless and tapering, slightly curved with pilose middle part, acicular in appearance, thick, with broad lance-shaped ends and transverse rows of fine hairs. Taxonomic Note Pseudeupanthalis Fauvel (1957), described in this family, appears to be synonymous with Sthenelanella in the family Sigalionidae. Invalid Genera Acoetes Audouin and Milne Edwards 1832, see Polyodontes Euarche Ehlers 1887, sec Eupanthalis Eupompe Kinberg 1855, see Polyodontes Lepidio Savigny in Audouin and Milne Edwards 1832, see Panthalis. Pholoididae NEW NAME Aphroditaceans with flattened bodies. One median antenna present. Four jaws present. Fringed elytrae alternate with dorsal cirri in all setigers. Each elytron with concentric rings. Neurosetae composite, notosetae simple. The family consists of one genus, Pholoides Pruvot 1895 with genotype P. dorsipapillata Pruvot 1895. The family-name is to replace Peisidicidae Hartman 1976 THE POLYCHAETE WORMS 67 FIGURE 16. (A), Family PHOLOIDIDAE, Phoioides áspera. Mugu Submarine Canyon, Caiifomia, about 100 m, 50x; (B), elytron of the above, 50x; (C), Family EULEPETHIDAE, Grubeuiepisfimbriata, Consag Rock, Golfo de California, elytron, 11 x; (D), diagram of tiie anterior end of the above, 8x; (E), median parapodium of the above, 25x; (F), anterior end of the above, first two elytrophores removed, lOx. 68 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 and Fauchald 1971, based on the genus Peisidice Johnson 1897. There appears to be no reason to maintain a separation between the two genera; the older name has priority, and the family name should be based on this generic name. Four species are presently considered valid. FAMILY EULEPETHIDAE CHAMBERLIN 1919C Invalid Genera Parapholoe Hartmann-Schröder 1965, see Phohides Peisidice Johnson 1897, seePhoioides Aphroditaceans with flattened bodies. Two antennae, four jaws present. Elytrae alternate with dorsal cirri in anterior setigers, present on all posterior setigers; elytrae marginally notched or with flattened marginal lappets. All setae simple. Neuracicula distally hammerheaded. The euiepethids (formerly pareulepids SENSU Hartman) were recently the subject of a monographic study by Pettibone (1969e). The present survey follows hers closely in most matters. Key to Genera la. lb. 2a 2b 3a 3b (la). (la). (lb). (lb). Elytrae 12 pairs, increasing in length progressively, followed by posterior pseudelytrae 2 Elytrae more than 12 pairs; the first 12 pairs increasing in length posteriorly; the more posterior pairs smaller , 3 Elytrae with lateral border notched Pareulepis Elytrae with flattened lateral marginal lappets Grubeulepis Elytrae with lateral borders notched Eulepethus Elytrae with flattened lateral marginal lappets Mexieulepis Taxonomic Note The pseudelytrae referred to in the key are sessile in contrast to the true elytrae, which are equipped with distinct elytrophores. They are in the same general position as the elytrae and appear to have similar functions. Generic Definitions Eulepethus Chamberlin 1919c, Eulepis hamifera Grube 1875; only species. Body with 60-70 segments. Elytrae with marginal notches on segments 2,4,5,7 . . . 21,24 and from segment 28 small elytrae with entire margins on every segment. Grubeulepis Pettibone 1969e, Eulepis fimbriata Treadwell 1901; 7 species. Body with about 38 segments. Elytrae with flattened marginal lappets including in all twelve pairs, on segments 2,4,5,7 . . . 21,24; pseudelytrae begin on segments 26-29. Mexieulepis Rioja 1961, Ai. elongam Rioja 1961; 2 species. Body with about 50 segments. Elytrae with flattened marginal lappets, including numerous pairs, on segments 2,4,5,7 . . . 21,24; smaller elytrae on every segment starting on segments 27 or 28. Pareulepis Darboux 1899, Eulepis wyvillei Mclntosh 1855; 2 species. Body with about 37 segments. Twelve pairs of elytrae on segments 2,4,5,7 . . . 21,24; all with lateral border notched. Pseudelytrae begin on segment 28. Invalid Genus Eulepis Grube 1875, see Eulepethus FAMILY SIGALIONIDAE MALMGREN 1867 Aphroditaceans with quadrangular or flattened bodies. One to three antennae present; four jaws present. Marginally fringed elytrae alternate with dorsal cirri on anterior setigers and are present on all posterior setigers. Neurosetae composite, notosetae simple. Sigalionids are common in soft bottoms; they tend to be long-bodied and the scales are usually rather closely appended to the bodies. This, combined with the well-developed notopodial fascicles of setae, tend to give them a rather quadrangular appearance. The sigalionids are presently under revision by Pettibone, who has already published a series of important monographs (Pettibone 1969c, 1970a, 1970b, 1970e, 1971b, 1971c). The key given below must be considered provisional. Key to Genera la. lb. One median antenna present At least two antennae present 2 3 1976 THE POLYCHAETE WORMS 69 FIGURE 17. Family SIGALIONIDAE, Sthenolepis japónica. Bay of Nha Trang, Viet Nam, 50 m, 25x. 2a 2b 3a 3b 4a 4b 5a 5b 6a 6b 7a 7b 8a 8b 9a (la). (la). (lb). (lb). (3b). (3b). (4a). (4a). (4b). (4b). (6a). (6a). (7a). (7a). (7b). 9b (7b). 10a 10b 1 la lib 12a 12b 13a 13b 14a 14b (6b). (6b). (10b). (10b). (1 lb). (lib). (12b). ( 12b). ( 13b). (13b). Composite neurosetae with long, slender, articulated appendages Mayella Composite neurosetae short, smooth unidentate falcigers Pholoe Two antennae present Sigalion Three antennae present 4 Lateral antennae on the prostomium proper; all antennae with short ceratophores or short and inconspicuous 5 Lateral antennae fused to first setiger; median antenna with large median ceratophore 6 Eyes large, antennae with ceratophores Euthalenessa Eyes small, antennae without ceratophores Thalenessa Median ceratophore without auricles 7 Median ceratophore with auricles 10 Dorsal cirri absent on setiger 3; elytrae not sand-incrusted 8 Dorsal cirri present on setiger 3; elytrae sand-incrusted 9 Neurosetae composite spinigers with short, canaliculated appendages Leanira Neurosetae composite, unidentate falcigers with marginally serrated appendages Pareupholoe Elytrae small, leaving the middle of the dorsum uncovered; neurosetae with short bidentate appendages Eupkoloe Elytrae covering the dorsum, neurosetae with uni- or bidentate appendages of varying lengths Psammolyce Auricles large, auricles also present on the tentacular segment Horstileanira Auricles small, absent from tentacular segment 11 All neurosetae composite unidentate falcigers with relatively short, straight appendages . . Sthenelanella At least some neurosetae otherwise (bidentate, or with articulated, canaliculated or spinigerous appendages) 12 Ventral cirri covered medially by long papillae Willeysthenelais Ventral cirri smooth 13 Parapodial lobes and sty lodes covered with fine fimbriae Fimbriosthenelais Parapodial lobes and stylodes smooth 14 Long dorsal cirri on setiger 3 Neoleanira Dorsal cirri absent from setiger 3 15 70 15a 15b 16a 16b (14b). (14b). (15b). (15b). NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY At least some neuropodial falcigers present All neurosetae spinigerous Dorsal tubercle present on setiger 3 Dorsal tubercle absent fron) setiger 3 Science Series 28 Sthenelais 16 Sthenolepis Ehlersileanira Generic Definitions Ehlersileanira Pettibone 1970c, Sthenelais incisa Grube 1877; 2 species. Three antennae; auricles present. Dorsal cirrus on setiger 3 absent. All neurosetae spinigerous, with canaliculated appendages. Eupholoe Mclntosh 1885, E. philippinensis Mclntosh 1885; only species. Three antennae; auricles absent. Dorsal cirrus on setiger 3 present. Elytrae small, leaving middle of body uncovered. Neurosetae with short, bidentate appendages. Encrusted with sand, Euthalenessa Darboux 1899, Thalenessa digitata Mclntosh 1885; 8 species. Three small antennae, all on the prostomium, ceratophores present. Two pairs of large eyes. Neuropodia with large foUose lobes anteriorly. Neurosetae slender, composite with multiarticulated distally bidentate appendages. Fimbriosthenelais Pettibone 1971b, Sthenelais longipinnis Grube 1870a; 2 species. Three antennae; auricles present. No dorsal cirrus on setiger 3. All parapodial lobes and stylodes fimbriated. Neurosetae either simple and spinose or composite with distally bidentate, short, slender and articulated appendages. Horstileanira Pettibone 1970a, H. vanderspoeii Pettibone 1970a; 2 species. Three antennae, large auricles, auricles present also on the tentacular segment. Setiger 3 with prominent dorsal tubercle. Neurosetae simple and spinose and composite spinigers with long, canaliculated appendages; a few may be bidentate. Leanira Kinberg 1855, L. quatrefagesi Kinberg 1855; 11 species. Three small antennae; auricles absent. No dorsal cirrus on setiger 3. Neurosetae composite spinigers with relatively short, canaliculated appendages, Mayella Hartmann-Schröder 1959, M. articulata Hartmann-Schröder 1959; only species. One antenna, auricles absent. Neurosetae composite spinigers with articulated appendages. Neoleanira Pettibone 1970a, Sigalion tetragonum Örsted 1845; 2 species. Three antennae; auricles present. Long dorsal cirri on setiger 3. Neurosetae composite spinigers with canaliculated, relatively short appendages. Pareupholoe Hartmann-Schröder 1962a, P. fimbriatus Hartmann-Schröder 1962a; only species. Three antennae; auricles absent. No dorsal cirri on setiger 3. Neurosetae composite, unidentate falcigers with serrated cutting edge. Pholoe Johnston 1839, Aphrodita minuta Fabricius 1780; 10 species. One antenna; auricles absent, Neurosetae composite falcigers with short, unidentate appendages. Notosetae in part geniculate and strongly dentate. Psammolyce Kinberg 1855, P. flava Kinberg 1855; 26 species. Three antennae, auricles absent. Third setiger with long dorsal cirri. Neurosetae composite falcigers with uni- or bidentate appendages of varying lengths. Elytrae cover body, sand-incrusted. Sigalion Audouin and Milne Edwards 1832, S. mathildae Audouin and Milne Edwards 1832; 12 species. Two lateral antennae; auricles absent. Neurosetae simple and spinose or composite with appendages of varying lengths, multiarticulated and bidentate. Sthenelais Kinberg 1855, S. helenae Kinberg 1855; 45 species. Three antennae, auricles present. No dorsal cirrus on setiger 3'. Neurosetae include simple spinose and two kinds of falcigers, stout with short, bidentate appendages, and slenderer falcigers with articulated appendages. Sthenelanella Moore 1910, S. uniformis Moore 1910; 4 species. Three antennae; auricles present. No dorsal cirri on setiger 3, All neurosetae with short, unidentate appendages. Sthenolepis Willey 1905, Leanira japónica Mclntosh 1885; 20 species. Three antennae; auricles present. With dorsal tubercles on setiger 3. Neurosetae all composite spinigers with canaliculated appendages, Thalenessa Baird 1868, Sigalion edwardsi Kinberg 1855; 8 species. Three small antennae, all on the prostomium, ceratophores absent. Two pairs of small eyes. Neurosetae composite falcigers and a few simple spinose setae. Willeysthenelais Pettibone 1971b, Sthenelais diplocirrus Grube 1875; only species. off Santa Catalina Island. . 50 m. shallow subtidal. held erect over the dorsum or covering the back as tiles on a roof. notosetae removed on four first segments on the right side and on the first segment on the left. Eniwetok. 50x. California. short appendages or more slender with articulated appendages. see Sthenelanella Superfamily Chrysopetalacea Elytrae absent. (C). (E). Neurosetae simple spinose or composite falcigers with thick. Family CHRYSOPETALIDAE. Neurosetae composite. whole animal. lOx. see Psammolyce Pseudeupanthalis Fauvel 1957. Notosetae in transverse rows. see Psammolyce Polylepis Grube 1878. notoseta (palea) of the above. Ventral cirri covered medially with long papillae. Chrysopetalum occidentale. 75x. FIGURE 18. see Sthenelais Euleanira Horst 1916a. notoseta (palea) of the above. see Sthenelais and Leanira Haswellia Darboux 1899. 50x. notosetae flattened and expanded paleae covering the dorsum. anterior end of the above. Chrysopetalids and palmyrids are very similar. the two families are often considered synonymous. Marshall Islands. (A). see Sthenelanella Eusigalion Augener 1918. see Psammolyce Pelogenia Schmarda 1861. see Thalenessa Eusthenelais Mclntosh 1876. 200x. Prostomium not fused to the first setiger. Palmyra aurífera. see Euthalenessa Lepidopleurus Claparède 1868. auricles present.1976 THE POLYCHAETE WORMS 71 Three antennae. No dorsal cirri on setiger 3. Invalid Genera Conconia Schmarda 1861. Family PALMYRIDAE. Day 1967). usually strongly flattened. in which case the name Palmyridae applies (cf. (D). Three antennae. setae of four first notopodia removed on the right side. (B). FAMILY CHRYSOPETALIDAE EHLERS 1864 Chrysopetalaceans with short or long bodies. dorsal paleae of two abruptly different kinds Paleanotus First segment with paired similar cirri. 5 species. neurosetae composite falcigers with blades of varying lengths. D. not covered by paleae. Fisione oerstedi. 19. see Dysponetus FAMILY PALMYRIDAE KINBERG Superfamily Pisionacea Elytrae absent. FIGURE . 2b (lb). P. Body with up to 300 segments. usually sandy or generally hard substrates in warm water and appear to be most common in the western Pacific Ocean.. body long. flattened bodies. 6 species. myriolepis Schmarda 1861. completely covered by paleae.. Four jaws present. Body with few segments. Body with about 40 segments. first segment with paired. shallow water. broad and narrow. 9 species. 4 species.72 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Key lo Genera Science Séries 28 la. Invalid Genera Heteropale Johnson 1897. Palmyra debilis Grube 1855. Caruncle present. Laubier (1967b) and Stecher (1968) recently have reviewed the family in terms of the generic sub-division. Peru. Paleanotus Schmarda 1861. dorsal and ventral cirri usually clávate. dorsal paleae of one kind only Chrysopetalum Generic Definitions Bhawania Schmarda 1861. lb. similar ventral cirri. The key below is after Laubier (1967b). prostomium deeply imbedded in the first segment. Setae composite and simple. The family is limited to shallow water. B. Caruncle absent. Palmyra Savigny 1818 with genotype/*. FAMILY PISIONIDAE SOUTHERN 1914 Pisionaceans with maximally two pairs of antennae on the prostomium. 3b (2b). The palmyrids are known for one genus. First segment with two pairs of tentacular cirri. aurífera Savigny 1818 and possibly for one or two more species. Notosetae cylindrical and erect over the dorsum. chrysolepis Schmarda 1861. Paleae of two kinds. body short 3 First segment with asymmetrical ventral cirri. completely covered by paleae. 2a (lb). Independencia Bay. 3a (2b). 25x. Family PISIONIDAE. not covering dorsum Dysponetus Notosetae flattened paleae 2 Prostomium without a conspicuous caruncle. prostomium retractile under a fold from the first setigers. Pisionura Hartman and Fauchald (1971) does not belong to this family (HartmannSchiöder 1975). 1858 Chrysopetalaceans with short. ses Paleanotus Psectra Grube 1868a. Caruncle may be present. Caruncle absent. completely covered by paleae. Dysponetus Levinsen 1879. Chrysopetalum Ehlers 1864. First segment with strongly asymmetrical ventral cirri. Siewing (1953). abruptly differing in shape. Notosetae in rosettes on the notopodial tubercles. parapodia sub-biramous or uniramous. Paleae of one or two kinds. Paleae of one kind only. most described forms are considered synonyms of P. Body with about 40 segments. pygmaeus Levinsen 1879. see Bhawania Taphus Webster and Benedict 1887. Notosetae cylindrical rather than flattened. Bhawania Prostomium with a conspicuous caruncle overlapping the peristomium. Neurosetae composite. consisting of many segments . One antenna present. or projecting freely between the first segments. aurífera or indeterminable. generally hesionids should be handled as scale-wotms: each specimen preserved separately. 2a (ib). Dorsal cirri long and slender. Ib. They tend to be fragile and fragment easily upon collection. lb. first segment without parapodia and setae. 6b (5a). the generic sub-division suggested by the key below is very tentative. a large number of additional genera may be expected described within a few years. 6a (5a).1976 THE POLYCHAETE WORMS 7i Key to Genera la. palps may be absent or have from one to three articles. frequently biarticulated. maximally three antennae Four or more pairs of tentacular cirri Three pairs of tentacular cirri Five or more pairs of tentacular cirri Pour pairs of tentacular cirri Five pairs of tentacular cirri Six or more pairs of tentacular cirri Seven or more pairs of tentacular cirri Six pairs of tentacular cirri Seven pairs of tentacular cirri Eight pairs of tentacular cirri Parapodia uniramous Parapodia biramous Hesiosyllis 2 3 7 4 8 Friedericielta 5 6 12 Periboea 22 Orseis Alikunhia . simple. Pisionella Hartman 1939. but notopodia always reduced compared to the neuropodia. 4a (3a). Pisionids with palps. if present. see Pisionidens Praegeria Southern 1914. 4b (3a). Parapodia uniramous or biramous. A. 2b (lb). First parapodia lateral. all segments with parapodia and setae. Proboscis with four jaws. 5b (4b). 7b (2b). Two or three antennae (antennae rarely absent). Fisione Grube 1857. all anterior segments with parapodia and setae. Key to Genera la. proboscis unarmed No apodous and asetigerous segment present. only species. dorsoventrally flattened worms. 2b (lb). FAMILY HESIONIDAE SARS 1862 Relatively short-bodied. Pisionella indica Aiyar and Alikunhi 1940. 7a (2b). notosetae. Pisionids with two pairs of similar cephalic appendages. F. Two to eight pairs of tentacular cirri present. Neurosetae composite. hancocki Hartman 1939. see Pisionidens Pisionella Aiyar and Alikunhi 1940. 2a (lb). 3a (2a). 3b (2b). Two pairs of tentacular cirri. see Fisione Suborder Nereidiformia Phyllodocida with at least one pair of antennae. all anterior segments with parapodia and setae. Invalid Genera Fauveliella Tebble 1953. Pisionids with palps and two pairs of tentacular cirri present. at least one pair of tentacular cirri. Hesionids are common animals in hard substrates and in shallow water. 5a (4b). oerstedii Grube 1857. Median unpaired antennae present Median unpaired antennae absent With two pairs of sinnilar cephalic appendages With three pairs of cephalic appendages of different structure First segment asetigerous and apodous. proboscis with four jaws Pisionella 2 Pisionidens 3 Anoplopisione Fisione Generic Definitions Anoplopisione Laubier 1967b. 3b (2a). only species. five antennae At least three pairs of tentacular cirri. only species. Pisionids with palps and two pairs of tentacular cirri present. minuta Laubier 1967b. they are more rarely found in deep water. palps short and usually distally blunt. Pisionidens Aiyar and Alikunhi 1943. The hesionids are one of the least known families of polychaetes. Proboscis unarmed. Proboscis with four jaws. Eversible pharynx. 11 species. Jaws may be present. P. if armed. two pairs of tentacular cirri and a median unpaired antenna present. 3a (2b). Proboscis with four jaws. with one pair of lateral jaws and sometimes with accessory denticles. outer harbor. 10b (8b). (B). Baja California. intertidal. diagrammatic oudine from die dorsal side. intertidal. Puerto Rico. Family CALAMYZIDAE. Family SYLLIDAE. Los Angeles. 9a (8a). 50x. 1 Calamyzas sp. Hesione interlexia. (F).74 8a (3b). 9b (8a). California. Family PILARGIIDAE. Family HESIONIDAE. (A). 50x. . median parapodium of the above. (D). TyposylHs armillaris. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Parapodia distinctly uniramous Parapodia sub-biramous or biramous First setiger with large hooks First setiger without hooks Antennae absent Antennae present Tentacular cirri on three segments (1-2-1). lOx. 1 la (10b). 8b (3b). 50x. Sigambra bassi. 50 m. anterior end of the above. 10a (8b). median paiapodium. 15x. (C). (E). El Descanso. dorsal cirri articulated Science Series 28 9 10 Stmwela Hesiocaeca Bonuania 11 Hesionella FIGURE 20. 50x.. Friedericiella Laubier 1967c. 27a (22b). 20b (19b). Dalhousiella Mclntosh 1901. 25b (24b). carpenteri Mclntosh 1901. 15b (14b). 12a (5b). 14b (13b). Eversible pharynx distally papillated. 16a (15b). 29a (28b). only species. 14a (13b). 29b (28b). Anophthalmus erythraeus Alikunhi 1949. Eversible pharynx distally smooth. THE POLYCHAETE WORMS 75 Hesionides 13 17 Syllidia 14 "Syllidia" 15 Parasyllidea 16 Neopodarke Micropodarke 18 21 Microphthalmus 19 Heteropodarke 20 Ophiodromus "Podarke" Parahesione Nereimyra 23 27 24 26 Amphiduros 25 Leocratides Leocrates Gyptis Podarkeopsis Hesiospina 28 Hesione 29 Wesenbergia 30 Dalhousiella Kefersteinia Tentacular cirri on two segments (2-2). jaws absent. . Amphiduros Hartman 1958. 30b (29b). 15a (14b). Two antennae. Median antenna attached medially. B. 11?L (6b). 5 species. 24a (23b). 16b ( 15b). 3 species. Parapodia biramous. palps and three pairs of tentacular cirri present. Pillai 1965. 21a (17b). 13b (12a). 17b (12b). Bonuania Pillai 1965. 26b (23b). only species. Parapodia uniramous. dorsal cirri smooth All parapodia uniramous At least some parapodia sub-biramous or biramous First three segments dorsally reduced Maximally first segment dorsally reduced Tentacular cirri on four segments (first reduced) so that they appear as 3-2-1 Tentacular cirri on three segments (2-2-2) Pharynx distally with a circlet of fine fimbriae Pharynx with either eleven or twenty-one distal papillae Pharynx with 21 distal papillae Pharynx with eleven distal papillae Three antennae present Two antennae present Median antenna attached medially or posteriorly on the prostomium Median antenna attached frontally Palpi simple Palpi biarticulated Setae present from the second segment Setae present from the fourth segment Dorsal cirri smooth Dorsal cirri articulated Three antennae Two antennae Median antenna attached medially or posteriorly on the prostomium Median antenna attached frontally Eversible pharynx distally fimbriated Eversible pharynx distally papillated Parapodia uniramous Parapodia biramous Setae present from the second segment Setae present from the fifth (or fourth) segment Notopodia with falcate spines Parapodia uniramous Palps absent Palps present Setae present from the fourth segment Setae present from the third segment Eversible pharynx smooth Eversible pharynx distally fimbriated Generic Definitions Alikunhia Hartman 1959. first setae in third segment. Eversible pharynx distally fimbriated. 28b (27b). 20a (19b). 19a (18b). 4 species. Three antennae. jaws absent. D. Antennae absent. Amphidromus setosus Hessle 1925. parva. 27b (22b). jaws absent. 25a (24b). 23b (22a). 17a (12b). 21b (17b). 12b (5b). 23a (22a). biarticulated palps and eight pairs of tentacular cirri present. 26a (23b). 18b (17a). 24b (23b). Parapodia biramous.1976 1 lb (10b). Parapodia biramous. biarticulated palps and eight pairs of tentacular cirri present (2-2-2-2). 13a (12a). 19b (18b). biarticulated palps and four pairs of tentacular cirri present. 22b (6b). 30a (29b). 18a (17a). 28a (27b). Three antennae. Hesioneila pacifica Friedrich 1956. jaws absent. biarticulated palps and six pairs of tentacular cirri present. Parapodia biramous. H. Median antenna attached posteriorly. biarticulated palps and six pairs of tentacular cirri present. jaws present. Hesiospina Imajima and Hartman 1964. Three antennae. only species. Kefersteinia similis Kessle 1925. jaws absent. jaws absent. jaws absent. . 7 species. Kefersteinia dubia Kessle 1925. Heteropodarke Hartmann-Schröder 1962a. 3 species. H. O. Nereimyra Blainville 1828. Nereis flexuosa delle Chiaje 1825. Parapodia biramous. Leocrates Kinberg 1866b. first setae in fourth segment. N. Two antennae. Parapodia biramous. Parapodia uniramous. biarticulated palps and eight pairs of tentacular cirri present. jaws absent. Eversible pharynx distally smooth. only species. jaws absent. Parapodia sub-biramous. simple palps and three pairs of tentacular cirri present. Eversible pharynx with distal circlet of fimbriae. only species. H. 13 species. Parapodia sub-biramous to biramous. Three antennae. jaws absent. first setae in third segment. jaws absent. simple palps and six pairs of tentacular cirri present. Eversible pharynx with distal circlet of papillae. Three antennae. palps absent. Three antennae. smooth palps and six pairs of tentacular cirri present. Parapodia sub-biramous.76 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Three antennae. jaws absent. bermudensis Hartman 1965. H. Notopodial falcate spines present. Micropodarke Okuda 1938. Eversible pharynx with 40 distal papillae. Two antennae. smooth palps and two pairs of tentacular cirri present. 5 species. Three antennae. only species. palps absent. Three antennae. Three antennae. Nereis punctata O. Median antenna attached posteriorly. Parapodia biramous. mccullochae Hartman 1939b. Eversible pharynx distally with many fine fimbriae. L. Eversible pharynx with ten distal papillae. only species. Five antennae (four frontal. Two antennae. Microphthalmus Mecznikow 1865. Hesionella Hartman 1939b. eight pairs of tentacular cirri present. Parapodia uniramous. Eversible pharynx with ten distal papillae and two longer cirri. Two antennae. Eversible pharynx with 21-27 distal papillae. Parapodia biramous. Two antennae. enigmática Wesenberg-Lund 1950. Eversible pharynx with 21 distal papillae. Eversible pharynx with jaws. only species. Eversible pharynx with jaws. Parapodia uniramous. Hesionides Friedrich 1937. Parapodia uniramous. setae from fourth segment. Neopodarke Hartman 1965. Gyptis Marion and Bobretzky 1875. Three antennae. Leocratides Ehlers 1908. Parapodia uniramous. Hesione Savigny 1818. arenaria Friedrich 1937. pulla Ehlers 1864. jaws and teeth present. Orseis Ehlers 1864. Hesiocaeca Hartman 1965. Median antenna attached posteriorly. simple palps and five pairs of tentacular cirri present. M. woodsholea Hartman 1965. chinensis Kinberg 1866b. filamentosa Ehlers 1908. only species. Two antennae. first setae in second segment. Three antennae. Eversible pharynx with circlet of papillae distally. Hesiosyllis Wesenberg-Lund 1950. Müller 1776. setae first in third segment. Parapodia uniramous. first setae in fourth segment. propinqua Marion and Bobretzky 1875. L. sczelkowii Mecznikow 1865. first setae in second segment. first setae in fourth segment. jaws absent. splendida Savigny 1818. biarticulated palps and six pairs of tentacular cirri present. Parapodia sub-biramous. biarticulated palps and four pairs of tentacular cirri present. 11 species. H. biarticulated palps and eight pairs of tentacular cirri present. only species. biarticulated palps and six pairs of tentacular cirri present. Kefersteinia Quatrefages 1865. biarticulated palps and eight pairs of tentacular cirri present. biarticulated palps and eight pairs of tentacular cirri present. Two antennae. Median antenna attached posteriorly. Ophiodromus Sars 1862. H. only species. one dorsal). 16 species. biarticulated palps and four pairs of tentacular cirri present. four pairs of tentacular cirri present. Eversible pharynx with a few distal papillae. Parapodia uniramous. Psamathe cirrata Keferstein 1862. Parapodia sub-biramous. Eversible pharynx with 11 papillae distally. biarticulated palps and eight pairs of tentacular cirri present. G. first setae in third segment. 11 species. heteromorpha Hartmann-Schröder 1962a. first setae in second segment. Parapodia biramous.F. 12 species. Parapodia uniramous. Wesenbergia Hartman 1955. Podarke Ex AUCTORE. confused. only species. unidentifiable Schmardiella Czemiavsky 1882. biarticulated palps and eight pairs of tentacular cirri present. . biarticulated palps and eight pairs of tentacular cirri present. 2 species. unidentifiable Stephania Claparède 1870b. Two antennae. but notopodia always reduced. The same is true of the concept Syllidia as it has been used in the literature. Pilargiids have been the subject of two recent revisions on the generic level. Notosetae may be absent. possibly five species. only species. longocirmta Ehlers 1864. indeterminable Dalhousia Mclntosh 1885. see Ophiodromus Castalia Savigny 1820. biarticulated palps and six pairs of tentacular cirri present. Eversible pharynx with jaws. seeNereimyra Cirrosyllis Schmarda 1861. only species. The key is wholly dependent on correct identification of the numbers of pairs of tentacular cirri. Parapodia uniramous. differs slightly from Ophiodromus. fu-st setae in fourth segment. Two antennae. P. jaws absent. Pilargiids are never really numerous. Two antennae. possibly about 12 species. ? jaws absent. Parapodia biramous. Eversible pharynx with a distal circlet of fimbriae. the only exceptions are the large. Most hesionids have dehiscent antennae and tentacular cirri so this may be the only means of getting the numbers of anterior appendages correctly identified. and Emerson and Fauchald (1971). Two or three antennae present (rarely absent). Two antennae. Invalid Genera Anophthalmus Alikunhi 1949. Parasyllidea Pettibone 1961. the concept as usually used. Eversible pharynx with a circlet of papillae. Periboea Ehlers 1864. Hesionella problemática Wesenberg-Lund 1950. Syllidia Quatrefages 1865. first setae on fourth visible segment (fifth segment). Parapodia biramous. Three antennae. but are present in most areas in moderate numbers. and have to be handled very carefully in collections. so that each tentacular cirrus will cast a shadow. first setae in fourth segment. Pearson (1970). jaws absent. Hesionids in general are rather fragile. and with the light coming in under a low angle. 5 species. biarticulated palps and six pairs of tentacular cirri present. Podarke luteola Webster 1880. first setae in third segment. Two antennae. jaws absent. humesi Pettibone 1961. first setae from fourth segment. Parapodia biramous. only species. Three antennae. compared to the version originally proposed. first setae in third segment. Parapodia biramous. see Ophiodromus Oxydromus Grube 1855. Setae simple. see Ophiodromus Onhodromus Ehlers 1908. SeeNereimyra Mania Quatrefages 1865. However. only species. Eversible pharynx with circlet of papillae. two pairs of tentacular cirri (rarely absent). S. biarticulated palps and six pairs of tentacular cirri present. Eversible pharynx with distal circlet of fimbriae. first setae in third segment. noodti HartmannSchröder 1959. armata Quatrefages 1865. jaws absent. Parapodia uniramous. confused. Uniramous parapodia. triarticulated long palps and seven pairs of tentacular cirri present. First to third segment dorsally reduced. seeGyptis Pseudosyllidia Czemiavsky 1882. Podarkeopsis Laubier 1961b. and rather solid-bodied members of the genus Hesione. Palps simple or biarticulated. Struwela Hartmann-Schröder 1959. Pettibone (1966b) added to the generic descriptions and revised several species. Two pairs of antennae. first setae in second segment. and has been included in the key and in the definitions. Parapodia uniramous. Parapodia sub-biramous. galangaui Laubier 1961b. Proboscis unarmed.1976 THE POLYCHAETE WORMS 77 Taxonomic Notes Parahesione Pettibone 1956. seeLeöcra/ej FAMILY PILARCDDAE SAINT-JOSEPH 1899 Nereidiforms with ribbon-shaped or cylindrical bodies. The genus Podarke usually is considered a synonym of Ophiodromus. simple palps and six pairs of tentacular cirri present. First setiger with large hooks. four pairs of tentacular cirri present. notosetae sometimes as a thick spine or hook. P. see Leocra/es Halimede Rathke 1843. First segment reduced. Two antennae. biarticulated palps and six pairs of tentacular cirri present. see Ophiodromus Telamone Claparède 1868. Eversible pharynx with 16-22 distal papillae. P. they tend to be associated with shelf depths and moderately coarse to mixed sediments. Syllidia Ex AUCTORE. see Alikunhia Anoplonereis Giard 1882. This is perhaps best observed on the lateral side of the animal. first setae in second segment. see Hesione Tyrrhena Claparède 1868. S. biarticulated palps. P. simplex Chamberlin 1919c. Otopsis Ditlevsen 1917. 4a (3b). verrucosa Saint-Joseph 1899.78 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Key to Genera Science Series 28 la. 5b (4b). P. but not articulated. A. biarticulate palps and two pairs of tentacular cirri present. Emergent notopodial spines present. A. Emergent notopodial spines present. parapodia distinct. palps present. straight spines 6 Peristomium dorsally entire Ancistargis Peristomium dorsally incised 4 Dorsal and ventral cirri reduced or absent. 6 species. body subcylindrical Cabira Dorsal and ventral cirri well developed. Emergent notopodial hooks present. Parandalia Emerson and Pauchald 1971. 9a (8b). 5a (4b). with two antennae. antennae Emd tentacular cirri absent. annandalei as reported by . 3a (2a). annandalei Fauvel 1932. body dorso-ventrally flattened . three antennae and two pairs of tentacular cirri present. C. biarticulate palps present Pilargis Prostomium with three antennae. Cabira Webster 1879b. parapodia poorly developed. parapodia well developed. incerta Webster 1879b. 8b (Ib). O. groenlandica McIntosh 1879. biarticulate palps and two pairs of tentacular cirri present. 2a (la). Emergent spines absent. Peristomium dorsally entire. Body cylindrical. 7a (6b). 7 species. Body flattened. L. 5. 6b (2b). Peristomium dorsally incised. Emergent notopodial hooks present. Sigambra Müller 1858. aberrans Monro 1936. 7b (6b). with three antennae. longipes Ditlevsen 1917. and has been considered an INCESTAE SEDIS. only species. only sjwcies. grubii Müller 1858. Emergent notopodial hooks present. Dorsal and ventral cirri reduced. 9b (8b). Body cylindrical. 3 species. 5 Antennae shorter than palps. but not set off from body 7 Tentacular cirri present. Antennae shorter than palps. 6 species. Taxonomic Note Talehsapia Fauvel 1932. 6a (2b). 7 species. 1970. L. Body cylindrical. Loandalia Monro 1936. parapodia sharply set off from body Synelmis Prostomial antennae absent. 4b (3b). has been considered a member of the family. ocularis Emerson and Pauchald 1971. two pairs of tentacular cirri present. Body cylindrical with biarticulate palps. Body flattened. palps without palpostyles Otopsis Generic Definitions Ancistargis Jones 1961. Emergent notopodial hooks present. Litocorsa Pearson. with three antennae. Ancistrosyllis Mclntosh 1879. Antennae shorter than palps. Emergent spines absent. 2b (la). papillosus Jones 1961. Antennae longer than palps. two pairs of tentacular cirri. biarticulate palps. palps present Parandalia Prostomial antennae and tentacular cirri absent Loandalia Prostomial antennae and tentacular cirri present 9 Prostomium with two antennae. two pairs of tentacular cirri. Body cylindrical without antennae and palps. lb. S. Emergent notopodial spines present. 8a (Ib). biarticulate palps and two pairs of tentacular cirri. integument papillated Ancistrosyllis Antennae longer than palps. 4 species. Pilargis Saint-Joseph 1899. biarticulate palps and two pairs of tentacular cirri present. it cannot be considered a pilargiid. Peristomium dorsally incised. with three antennae. 11 species. Body flattened. Body flattened. parapodia reduced. As noted by Emerson and Pauchald (1971). 2 species. stremma Pearson 1970. T. two antennae. biarticulate palps. three antennae. biarticulate palps present. integument smooth Sigambra Prostomial antennae present. 3b (2a). palps absent Litocorsa Tentacular cirri absent. Body flattened. Emergent spines absent. with genotype T. Notopodia with stout emergent hooks or spines 2 Notopodia without emergent hooks or spines 8 Notopodia with recurved emergent hooks 3 Notopodia with stout. Synelmis Chamberí in 1919c. antennae and tentacular cirri absent. . 3a (2b). occipital flap absent 4 Dorsal cirri cylindrical 5 Dorsal cirri clávate or folióse 6 Bayonet-setae thick-shafted. 5b (4a). Similarly. see Pilargis FAMILY SYLLIDAE GRUBE 1850 Small to medium-sized nereidiform polychaetes with usually. and tend to be most numerous on hard substrates. palps fused for at least half their lengths . Revision of the California fauna is under way and may be expected within a few years (Piltz. they are considered here as unidentifiable Parandalia spp. or pigmented specimens will have to be dissected. 7a (6b). Hartmann-Schröder has also concentrated considerable attention on the syllids. 8b (7b). In certain genera {Trypanosyllis. set Synelmis Harpochaeta Korschelt 1893. occipital flap present Alluaudella At least some setae composite. lib (9a). 12b (1 lb). Ventral cirri absent AUTOLYTINAE 2 Ventral cirri present 9 Dorsal cirri on first setiger only Procerastea Dorsal cirri on most setigers 3 All setae simple. 6b (4b). the latter sometimes fused to each other. more than . Larger specimens. Three antennae and simple palps present. 4b (3b). Autolytus and others). 5a (4a). Two pairs of tentacular cirri. making the identification of syllids a time-consuming occupation. as is amply demonstrated in the key below. however. s&e. Key to Genera la. lb. setae simple or composite. rather than flattened nuchal epaulettes Autosyllis Nuchal organs flattened epaulettes 8 Antennae and dorsal cirri clávate Umbellisyllis Antennae cylindrical. EXOGONINAE 11 Larger forms. Invalid Genera Glyphohesione Friedrich 1951. Gidholm 1962) and more are expected. dorsal cirri smooth or irregularly wrinkled EUSYLLINAE 23 Palpi free to base or absent. slender bodies (sometimes dorsoventrally flattened). in preparation). even at the generic level. Exogoninae. the structure of the trepan. is of great importance. only fused basally 10 Palps fused at base. indeterminable Kynephorus Ehlers 1920. also is well represented in abyssal depths. see Synelmis Phronia Webster 1879b.. Ancistrosyllis Hermundura Müller 1858. The examination of the trepan can be done through the body-wall. in a series of papiers revised the Japanese syllids. dorsal cirri articulated SYLLINAE 43 Antennae. if fused at all. ciliary bands present on every segment Autolytus Eversible pharynx unarmed Phyllosyllis Eversible pharynx with teeth 7 With projecting nuchal lobes. 6a (4b). 8a (7b).5 mm across. The pharynx is only rarely eversed in preserved material. if the specimen is very small and unpigmented.g. palps. Recent monographs include Imajima (1966a-d 1967) who. tentacular and dorsal cirri absent Exogonella At least tentacular cirri present. 1 la (9a). 9b (lb). 2b (la). close examination of the structure of the setae is also necessary. dorsal cirri usually conspicuous. 10a (9b). Examination of the eversible pharynx cannot be dispensed with in this family. dorsal cirri flattened Myrianida Small forms (usually less than 8 mm). median and posterior region should be mounted on a slide for setal examination under a compound microscope.7976 THE POLYCHAETE WORMS 79 Fauvel (1935) and Mesnil and Fauvel (1939) differ from the species as originally described. 7b (6b). 12a (lib). 9a (lb). Eversible pharynx armed with a single tooth or a circlet of smaller teeth or unarmed. Pro ventricle present in nearly all forms. usually also antennae and dorsal cirri 12 Two pairs of tentacular cirri present 13 One pair of tentacular cirri present 17 . personal communication). the denticles along the cutting edge of the eversible pharynx. 10b (9b). The syllids are very common shallow-water forms. Parapodia uniramous. and the whole specimens should be scanned for modified setae of any kind. 3b (2b). 2a (la). Gidholm has published a series of papers on the subfamily Autolytinae (e. (Olga Hartman. segmental cUiary bands absent Proceraea Bayonet-setae thin-shafted. 4a (3b). one sub-family. Parapodia from anterior. 15b (14b). 21a (20a). 31b (30b). dorsal cirri well developed if sometimes short Dorsal cirri long and slender Dorsal cirri short. 34a (33a). 20a (18b). 18a (17b). 27a (26b). 28b (27b). 41a (24b). 36a (33b). 23b (10a). 28a (27b). 21b (20a). palps twice as long as prostomium Parapodia short and conical. 30a (24a). 37b (36b). 19a ( 18a). 13b ( 12a). 36b (33b). 38a (37b). 29a (28b). 35b (34b). 22a (2üb). 33b (32b). 37a (36b). straight Dorsal cirri long and filiform Dorsal cirri globular Body with dorsal globular papillae in addition to the dorsal cirri Body without dorsal globular papillae One antenna present. 39a (38a). 17a (12b). 26b (25b). 22b (20b). 32a (31b). 27b (26b). 24b (23b). 40b (38b). palps maximally as long as prostomium Eversible pharynx with a series of very small teeth Science Series 28 Exogonita 14 Campesyllis 15 Brania 16 Eurysyllis Plakosyllis Spermosyllis 18 19 20 Anguillosyllis Braniella 21 22 Exogonoides Pseudexogone Sphaerosyllis Exogone 25 24 30 41 Rhopaiosyllis 26 Pharyngeovalvata 27 Fauvelia 28 Atelesyllis 29 Streptosyllis SylUdes Eudontosyllis 31 Nudisyllis 32 Irmula 33 34 36 Lamellisyltis 35 Parapionosyllis Petitia Clavisyliis 37 Opisthodonta 38 39 40 Synsyllis Eusyllis Dioplosyllis PionosylUs Parautolytus . 32b (31b). 29b (28b). 17b (12b). 35a (34b). dorsal cirri rudimentary Tentacular cirri present. 30b (24a). 19b (18a). 33a (32b). 24a (23b). 14b (13b). contractile dorsal lobes Parapodia without dorsal lobes Eversible pharynx unarmed Eversible pharynx with a single anterior tooth Ventral cirri fused to lower edge of parapodia Ventrial cirri free from parapodia Dorsal cirri pyriform (flask-shaped) Dorsal cirri papilliform or ovoid Eversible pharynx unarmed Eversible pharynx either with a single tooth or with a chrlet of smaller teeth Eversible pharynx with a single large tooth Eversible pharynx with a series of teeth Dorsum covered with small papillae Dorsum without papillae Pharynx with an internal valve Pharynx without an internal valve Tentacular cirri absent. 31a (30b). thick simple setae Median body-region with composite falcigers Parapodia long. 18b (17b). 26a (25b). 40a (38b). 16b (15b). dorsal cirri rudimentary Three antennae present. 38b (37b). dorsal cirri well developed Antennae and tentacular cirri very short Antennae and tentacular cirri not noticeably short Enlarged knobbed acicula present in anterior parapodia Enlarged knobbed acicula absent Notacicula present Notacicula absent Antennae and dorsal cirri absent Antennae and dorsal cirri present Three first segments fused with six pairs of tentacular cirri Maximally two pairs of tentacular cirri on one segment One pair of tentacular cirri present Two pairs of tentacular cirri present Dorsal cirri flattened Dorsal cirri cylindrical Composite setae spinigerous Composite setae falcigerous Tentacular and dorsal cirri very large. 25a (23a). ovoid or pyriform Parapodia with erect. 14a (13b). ovoid and inflated Tentacular and dorsal cirri more or less cylindrical Middorsal tooth situated posterior in the eversible pharynx Middorsal tooth situated anterior in the eversible pharynx Anterior margin of eversible pharynx denticulated Anterior margin of eversible pharynx smooth Median body-region with furcate. 39b (38a). NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Antennae absent Three antennae present Eversible pharynx non-muscular and sinuous Eversible pharynx at least partially muscular. 16a (15b). 15a (14b). 34b (33a). 23a (10a). 25b (23a).-so 13a (12a). 20b (18b). 50a (45a). (10b). first dorsal cirri longer than the rest. Three antennae. 46b (44b). 47a (46b). large occipital flap present. Amblyosyllis Grube 1857. 52b (5Ib). dorsal cirri present on second segment 54 Parapodia with pseudocomposite and simple setae in addition to the composite ones Syllis Parapodia maximally with two simple setae in addition to the composite ones 55 Both composite spinigers and falcigers present Ehlersia Only composite falcigers present Typosyllis Generic Definitions EUSYLLINAE. 51a (50b). AUTOLYTINAE.7976 41b 42a 42b 43a 43b 44a 44b 45a (24b). setae all simple. two pairs of tentacular cirri. palps separated to the base. (43b). 53b (52b). Three antennae. A. Three long antennae. Alluaudella Gravier 1905c. dorsal cirri absent from second segment Paratyposyllis Tentacular and dorsal cirri with at least five articles. A. SYLLINAE. Palps completely fused. Eversible pharynx with six or seven bi. eversible pharynx unarmed. Parapodia with long contractile dorsal lobes. (10b). Each segment with ciliated band. first dorsal cirri short. Autosyllis Imajima and Hartman 1964. (41b). 46a (44b). digitate distal lobes Branchiosyllis Parapodia without distal lobes 48 Paired. Dorsal cirri clávate or folióse. Paired long nuchal epaulettes present. A. sometimes with an additional larger single tooth as well 49 Eversible pharynx with a single tooth 50 Dorsum covered with small papillae Xenosyllis Dorsum smooth 47 Parapodia with long. nuchal epaulettes absent Odontosyllis Palps absent. only species. 49a (45b). with few segments. Palps free to base. Eversible pharynx with trepan with varying number of teeth. rubrofasciata Pruvot 1930. a trepan or both 45 Eversible pharynx unarmed 46 Eversible pharynx with a trepan of several teeth. oculata Ehlers 1887. with numerous segments. (44a). posteriorly directed nuchal lappets present Parapterosyllis Nuchal lappets absent Pseudosyllides Palpi as long as prostomium. two pairs of tentacular cirri. Müller 1788. rhomheata Grube 1857. long palps separated to the base. 5 species. large nuchai epaulettes present Amblyosyllis Body longer. 54b (53b). one pair of tentacular cirri. 8 species. Dorsal cirri emerge well above base of parapodia. 55b (54b). Eversible pharynx unarmed. Autolyms Grube 1850. bulbously fusiform Parasphaerosyllis Dorsal cirri may alternate between long and short. first segment without parapodia and setae. 49b (45b). EXOGONINAE. two pairs of tentacular cirri. 50b (45a). usually with tentacular cirri 44 Eversible pharynx with a single tooth. Long slender forms with three minute antennae. 55a (54b). Atelesyllis Pruvot i 930. nuchal projecting lobes present. Eversible pharynx unarmed. AUTOLYTINAE. 54a (53b). two pairs of tentacular cirri. japónica Imajima and Hartman 1964. Three antennae. body cylindrical Geminosyllis Palpi small and conical. 51b (50b). 48a (47b). (41b). but all are of similar thickness 53 Tentacular and dorsal cirri with very few articles. 47b (46b). Eversible pharynx unarmed. 2 species. Three short antennae and occipital flap present.tri. capensis Day 1963. . (43b). 48b (47b). Bayonet-setae thin-shafted. 110 species. A.E. 52a (51b). Antennae and tentacular cirri very short. Dorsal cirri wrinkled. body flattened Trypanosyliis Middorsal tooth attached posterioriy OpisthosylUs Middorsal tooth attached anteriorly 51 Setae simple. B.or pentacuspid teeth. Eversible pharynx with a limited number of large teeth 42 Body short. Branchiosyllis Ehlers 1887. first segment with parapodia and setae Haplosyllides Palps present. Parapodia with long digitate distal lobes attached pre. Anguillosyllis Day 1963. EUSYLLINAE. madagascariensis Gravier 1905c. AUTOLYTINAE.and postsetally. A. 53a (52b). only species. THE POLYCHAETE WORMS 8] 45b (44a). distally furcate or subdistally bossed Haplosyltis At least some setae composite 52 Dorsal cirri in posterior region alternately long and slender or large. Nuchal epaulettes present. Nereis proliféra O. only species. ventral cirri short. two pairs of tentacular cirri present. Haplosyllis Langerhans 1879. SYLLINAE. ventral cirri as long as the setal lobes. margin denticulated. D. tentacular cirri and dorsal cirri also globular. SYLLINAE. Three antennae and two pairs of tentacular cirri. Doisai cirri longer than. only species. nuchal ridges present or absent. dorsal cirri long and filiform. E. Antennae absent. ventral cirri fused to par^Kidia. EXOGONINAE. Exogone Örsted 1845. ovoid and inflated. slender and smooth. Setae composite falcigers. all cirri articulated and slender. Clavisyllis Knox 1957. Exogone pusilla Dujardin 1851. EXOGONINAE. Composite spinigers. Eversible pharynx smooth-rimmed with single dorsal tooth. alternata Knox 1957. two pairs of tentacular cirri. All antennae and cirri slender and articulated. Exogonita Hartman and Fauchald 1971. Three ovoid antennae. all cirri smooth. Palpi as long as prostomium. ovate antennae. minor Chamberlin 1919a. Eversible pharynx unarmed. Two pairs of tentacular cirri. Body short and flattened. Ehlersia Quatrefages 1865. floridana Augener 1922. 21 species. 15 species. Dorsal and ventral cirri shorter than setal lobes. Three antennae and two pairs of tentacular cirri. Eurysyllis Ehlers 1864. EXOGONINAE. martinensis Gravier 1900. Three antennae. two pairs of tentacular cirri. naidina Örstcd 1845. Eversible pharynx with middorsal tooth. Antennae absent. B. EUSYLLINAE. E>orsal cirri long. Three antennae. EXOGONINAE. Three antennae. ventral cirri folióse. H. occipital flap present. SYLLINAE. . EUSYLLINAE. E. one pair of ovoid tentacular cirri. all anterior appendages articulated (except palps). two pairs of tentacular cirri. Exogonoides Day 1963. Tentacular and dorsal cirri large. Eversible pharynx with a single tooth. Eversible pharynx with a trepan with 10 teeth and a middorsal tooth present. EXOGONINAE. Setae simple. Fauvelia Gravier 1900. Eversible pharynx with a single tooth. Three short. Antennae. dorsal cirri ovoid or papilliform. EUSYLLINAE. Dioplosyllis Gidholm 1962. EUSYLLINAE. 27 species. Campesyllis Chamberlin 1919a. Eversible pharynx with trepan of ten teeth and in addition a single large tooth. Dorsum covered with rows of globular papillae. tentacular cirri absent. tentacular and dorsal cirri absent. E. Trypanosyllis (Trypanedenta) ohma Imajima and Hartman 1964. palps absent. one pair of tentacular cirri. Eversible pharynx with a single tooth. 2 species. Geminosyllis Imajima 1966c. pupa Hartman 1965. cirrosa Gidholm 1962. Three antennae. Setae include composite spinigers and falcigers and in posterior setigers one or two simple setae per fascicle. aciculata Knox 1960. eversible pharynx unarmed. Body subcylindrical. oculata Hartman and Fauchald 1971. two pairs of tentacular cirri. brunnea Hartman 1961. Notacicula present. EXOGONINAE. dorsal cirri long. smooth-riramed or with a few teeth. Three antennae. C. E. dorsal cirri rudimentary. Braniella Hartman 1965. prominent nuchal epaulettes present. Exogonella Hartman 1961. only species. three antennae. 2 species. C. blomstrandi Malmgren 1867. parapodia long. Palps very large and Ungulate. SYLLINAE. Eversible pharynx sinuous and nonmuscular. 40 species. Eversible pharynx with middorsal tooth. only species. F. Three short antennae. only species. Eversible pharynx with middorsal tooth. two pairs of tentacular cirri. Eudontosyllis Knox 1960. antennata Day 1963. Occipital flap may be present. 3 species. only species. E. one pair of tentacular cirri. Eversible pharynx with middorsal tooth and denticulated margin. EXOGONINAE. tuberculata Ehlers 1864. only species. 10 species. only species. E. Three globular antennae. Eversible pharynx with single tooth. Tentacular and dorsal cirri articulated. EUSYLLINAE. Haplosyllides Augener 1922. Eversible pharynx with anterior dorsal tooth. First segment with parapodia and setae. EXOGONINAE. E. Syllis sexoculata Ehlers 1864. only species. Syllis spongicola Grube 1855.82 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Brania Quatrefages 1866. 3 species. Dorsal cirri ovoid. Eusyllis Malmgren 1867. distally furcate or with a subdistal boss or knob. Eversible pharynx unarmed. 1976 THE POLYCHAETE WORMS 83 Irmula Ehlers 1913, /. spissipes Ehlers 1913; only species. EUSYLLINAE. Three antennae; six pairs of tentacular cirri on three fused segments. All cirri smooth. Eversible pharynx with anterior single tooth. Lamellisyllis Day 1960, L. comans Day 1960; only species. EUSYLLINAE. Flattened, small form. Three foliaceous antennae; one pair of tentacular cirri. Eversible pharynx with single tooth. Dorsal cirri flattened; ventral cirri cylindrical. Myrianida Milne Edwards 1845, Nereis pinnigera Montagu 1808; 8 species. AUTOLYTINAE. Three antennae; two pairs of tentacular cirri. All dorsal cirri flattened; antennae cylindrical. Eversible pharynx with trepan with varying numbers of teeth. Nudisyllis Knox and Cameron 1970, N. tinihekia Knox and Cameron 1970; only species. EUSYILLINAE. Antennae and dorsal cirri absent. Eversible pharynx with a single tooth; margin smoothrimmed. Odontosyllis Claparède 1863, Syllis fidgwans Audouin and Milne Edwards 1833a; 35 species, EUSYLLINAE. Three antennae; two pairs of tentacular ciiri. Occipital flap usually present. Eversible pharynx with a series (less than 20) curved teeth. Opisthodonta Langerhans 1879, O. morena Langerhans 1879; 2 species. EUSYLLE^AE. Three antennae; two pairs of tentacular cirri; all antennae and cirri smooth. Eversible pharynx with a single large, posteriorly located tooth. Some anterior parapodia with very thick acicula. Opisthosyllis Langerhans 1879, O. brimnea Langerhans 1879; 10 species. SYLLINAE. Three antennae; two pairs of tentacular cirri; occipital flap may be present. All antennae and cirri articulated. Eversible pharynx with posteriorly attached mid-dorsal tooth; anterior margin smooth. Parapionosyllis Fauvel 1923, Pionosyllis gestarts Pierantoni 1903; 9 species. EUSYLLINAE. Three antennae; one pair of tentacular cirri. Eversible pharynx with a single tooth. Composite setae spinigerous. Parapterosyllis Hartmann-Schröder 1960a, P. sexoculata Hartmann-Schröder 1960a; 2 species. SYLLINAE. Three antennae and two pairs of tentacular cirri. Piostomium with paired posteriorly directed nuchal lappets. All appendages articulated. Eversible pharynx unarmed. ParasphaerosylUs Monro 1937b, P. indica Monro 1937b; 4 species. SYLLINAE. Three antennae; two pairs of tentacular cini. Dorsal cirri anteriorly all slender and moniliform; posteriorly alternating between slender and large, bulbously fusiform cirri. Eversible pharynx with middorsal tooth. Paratyposyllis Hartmann-Schroder 1962b, P. paurocirrata Hartmann-Schröder 1962; 2 species. SYLLINAE. Three antennae; two pairs of tentacidar cirri; eversible pharynx with single tooth. All composite setae falcigers; one or two simple setae present in each of the posterior setigers. All antennae and cirri with less than five articles; second segment without dorsal cirri. Parautolytus Ehlers 1900, P. fasciatus Ehlers 1900; 2 species. EUSYLLINAE. Three antennae; two pairs of tentacular cirri. Antennae and cirri smooth. Eversible pharynx fmely denticulated, large tooth absent. Petitia Slewing 1955, P. amphophthalma Slewing 1955; only species. EUSYLLINAE. Three antennae; one pair of tentacular cirri. Palpi biarticulate in adults. Eversible pharynx with a single tooth. Composite setae falcigerous. Pharyngeovalvata Day 1951, P. natalensis Day 1951; only species. EUSYLLINAE. Three antennae; two pairs of tentacular cirri. Occipital flap present. Pharynx with valve; teeth absent. Phyllosyllis Ehlers 1897, P. albida Ehlers 1897, only species. AUTOLYTINAE. Three antennae; first segment setose, with two large, folióse cirri. Eversible pharynx without teeth. Dorsal cirri folióse. Pionosyllis Malmgren 1867, P. compacta Malmgren 1867; 31 species. EUSYLLINAE. Three antennae; two pairs of tentacular cirri. Tentacular and dorsal cirri smooth and cyhndrical. Eversible pharynx with single tooth; anterior margin smooth. Plakosyllis Hartmann-Schröder 1956, P, brevipes Hartmann-Schröder 1956; 2 species. EXOGONINAE. Body short, three globular antennae; tentacular cirri and dorsal cirri also globular. Eversible pharynx with a trepan with ten teeth; a single tooth also present. Dorsal globular papillae absent. Proceraea Ehlers 1864, P. picta Ehlers 1864; 7 species. AUTOLYTINAE. Three antennae; two pairs of tentacular cirri. Palps small and ventrally located, Eversible 84 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 pharynx with trepan with numerous teeth. Epaulettes present. Bayonet-setae thick-shafted. Ciliary, bands absent from the setigers. Procerastea Langerhans 1884, P. nematodes Langerhans 1884; 3 species. AUTOLYTINAE. Three antennae, two pairs of tentacular cirri. Dorsal cirri on first setiger only. Eversible pharynx with trepan with numerous teeth. Pseudexogone Augener 1922, P. backstromi Augener 1922; only species. EXOGONINAE. Three anteimae; one pair of tentacular cirri. Eversible pharynx unarmed. Dorsal cirri present on second segment. Pseudosyllides Augener 1927b, P. curacaoensis Augener 1927b; only species. SYLLINAE. Three antennae; two pairs of tentacular cirri; antennae and cirri strongly articulated. Eversible pharynx unarmed with a smooth margin. Rhopalosyllis Augener 1913a, R. hamuUfera Augener 1913a; only species. EUSYLLINAE. Three antennae and two pairs of tentacular cirri. Antennae and cirri smooth. Dorsum covered with long papillae. Eversible pharynx unarmed. Spermosyllis Claparède 1864, S. torulosa Claparède 1864; 3 species. EXOGONINAE. One antenna and one pair of tentacular cirri. Eversible pharynx with a single tooth. Dorsal cirri rudimentary, ventral cirri absent. Sphaerosyllis Claparède 1863, S. hystrix Claparède 1863; 28 species. EXOGONINAE. Three antennae; one pair of tentacular cirri. Dorsal cirri pyriform (flask-shap)ed), absent on second segment. Body with adhesive papillae. Strepiosyllis Webster and Benedict 1884, S. arenae Webster and Benedict 1884; 7 species. EUSYLLINAE. Three antennae and two pairs of tentacular cirri. Eversible pharynx unarmed. Large knobbed acicula present in anterior setigers. Syllides Örsted 1845, 5. longocirrata Örsted 1845; 13 species. EUSYLLINAE. Three antennae and two pairs of tentacular cirri. Eversible pharynx unarmed; distal margin of pharynx smooth. No enlarged setae or acicula. Syllis Savigny 1818, S. monilaris Savigny 1818; 45 species. SYLLINAE. Three antennae; two pairs of tentacular cirri, all articulated. Eversible pharynx with a single tooth. Pseudocomposite and simple setae present in addition to the composite setae in all parts of the body. Synsyllis Verrill 1900, S. longigularis Verrill 1900; 2 species. EUSYLLINAE. Three antennae and two pairs of tentacular cirri. Eversible pharynx with mid-dorsal tooth; margin denticulated. Middle part of body with large, distally furcate simple setae. Trypanosyllis Claparède 1864, Syllis zebra Grube 1860; 26 species. SYLLINAE. Body flattened with numerous short segments. Three antennae and two pairs of tentacular cirri; all articulated. Trepan with several teeth; a single tooth may be present. Typosyllis Langerhans \S79, Syllis krohnii Ehlers 1864; 89 species. SYLLINAE. Three antennae and two pairs of tentacular cirri; all articulated. Eversible pharynx with a single tooth. Setae include uni- or bidentate falcigers in addition to one or two simple setae in each of the posterior setigers. Umbellisyllis Sars 1869, U. fasciata Sars 1869; 2 species. AUTOLYTINAE. Three clávate antennae; two pairs of tentacular cirri. Nuchal organs foliaceous. Dorsal cirri clávate. Xenosyllis Marion and Bobretzky 1875, Syllis scabra Ehlers 1864; 2 species. SYLLINAE. Three short, thick antennae; two pairs of tentacular cirri. Tentacular cirri and dorsal cirri with few moniliform or collared articles. Dorsum covered with small papillae. Eversible pharynx unarmed, Taxonomic Notes The subfamilies have been accepted strictly as defined above; as a consequence several of the genera have beeil moved from one subfamily to another. This is not considered to be of any great importance: the differences between the subfamilies, especially between the EUSYLLINAE and SYLLINAE appear to be of more practical than scientific value. The genus Irmula was originally described in the Hesionidae. It was moved to Syllidae by Day (1967). It has a very isolated position in the family due to the presence of three modified anterior segments with six pairs of tentacular cirri. However, the structure of pharynx, parapodia and setae is typically syllid, so it appears best to retain it in the Syllidae. Hesiosyllis Wesenberg-Lund 1950, described as intermediary between the syllids and the hesionids, is treated here among the latter, in that the structure of the pharynx, setae, parapodia and tentacular cirri appear to resemble members ofthat family much more than it resembles any member of the Syllidae. J 1976 Invalid Genera THE POLYCHAETE WORMS 85 Amytis Savigny 1818, &et Proceraea Anoplosyllis Claparède 1868, see Syllides Aporosyllis Quatrefages 1865, seeSyllis Autolytides Malaquin 1893, seeAutolytus Brachysyllis Imajima and Hartman 1964, see Dioplosyllis Chaetosyllis Malmgren 1867, seeEhlersia Cirrosyllis Schmarda 1861, see Amblyosyllis and Autolytus Claparedia Quatrefages 1865, see Eusyllis (?) Crithida Gosse 1855, seeAutolytus Cystonereis Köllikerm Koch 1846, seeExogone Desmosyllis Verrill 1900, see Eusyllis Diploceraea Grube 1850, seeAutolytus Doyeria Quatrefages in Milne Edwards 1848, NOMEN NUDUM Eucerastes Ehlers 1864, seeMyrianida Eurymedusa Kinberg 1866b, see Trypanosyllis Exotokas Ehlers 1864, seeExogone Gattiola Johnston 1863, see Amblyosyllis Gnathosyllis Schmarda 1861, seeSyllis Gossia Quatrefages 1865, seeExogone (?) Grubea Quatrefages 1865, seeBrania Grubeosyllis Verrill 1900, see Brania Hemisyllis Verrill 1900, see Haplosyllis Hesperalia Chamberiin 1919a, lOdontosyllis Heterosyllis Claparède 1863, indeterminable loda Johnston 1840, seeSyllis ¡sosyllis Ehlers 1864, see Typosyllis Lalage Müller 1858, NOMEU NUDVM {see Syllis) Langerhansia Czemiavsky 1882, see Ehlersia Laomedora Kinberg 1866b, indeterminable Lapithas Kinberg 1866b, indeterminable Lophosyllis Sars 1867, indeterminable Lycastis Savigny 1818, see Typosyllis Microsyllis Claparède 1863, questionably fjíogo/ie Monocerina Costa 1861a, indeterminable Nereisyllis Blainville 1828, seeSyllis Nicotia Costa 1864, see Amblyosyllis Oophylax Ehlers 1864, see Exogone Paedophylax Claparède 1868, seeExogone Pagenstecheria Quatrefages 1865, see Typosyllis Parasitosyilis Potts \9\2, NOMENNVDVM Parexogone Mesnil and Caullery 1916, seeExogone Periboea Kinberg 1866b, indeterminable Photocharis Ehrenberg 1835, indeterminable Platysyltis Grube 1878, indeterminable Podonereis Blainville 1818, seeAutolytus Polybostrichus Örsted 1843a, see Proceraea Polymastus Claparède 1864, see Eurysyltis Polynice Savigny in Grube 1850, questionably Autolytus Procome Ehlers 1864, see Odontosyllis Protogrubea Czemiavsky 1881a, seeBrania PseudosyHides Czemiavsky 1882, set Amblyosyllis Pterautolytus Ehlers 1907, seeAutolytus Pterosyllis Claparède 1863, see Amblyosyllis Sacconereis Müller 1853, seeAutolytus Salvatoria Mclntosh 1885, seeBrania Sckmardia Quatrefages 1865, Siee Exogone Stephanosyllis Claparède 1864, see Proceraea Syllia Quatrefages 1865, seeExogone Sylline Grube 1860, seeAutolytus Sylline Claparède 1864, seeExogone Tetraglene Grube 1863, see Trypanosyllis Thoe Kinberg 1866b, see Typosyllis Thylaciphorus Quatrefages 1865, see Amblyosyllis Trichosyllis Schmarda 1861, seeSyllis Virchowia Langerhans 1879, see Umbellisyllis Xenosyllides Perejaslavzeva in Jakubova 1930, questionably Umbellisyllis FAMILY CALAMYZIDAE HARTMANN-SCHRôDER 1971 Body short, anteriorly and posteriorly rounded Prostomiura small and without appendages with a suck ing mouth on the ventral side. All cirri (tentacular cirri, dorsal and ventral cirri) short and digitate. Anal cirri absent. Setae composite. Eversible pharynx with stylet shaped sucking tube, otherwise without specializations The only known genus and species, Calamyzas amphictenicola Arwidsson 1932 is parasitic on the ampharetid polychaete,/Impfcicie/i gunneri (Sars 1835) from Sweden. A review can be found in HartmannSchröder 1971. FAMILY NEREIDAE JOHNSTON 1845 Elongated, multi-segmented nereidiform polychaetes. Two, rarely one, antennae; palps biarticulated. Two or four pairs of tentacular cirri. Eversible pharynx with a pair of jaws and often accessory denticles or papillae. Parapodia nearly always biramous, usually with complex flattened lobes and cirri. Setae composite or simple, spinigerous or falcigerous. The nereids are common forms in all depths, and penetrate fteshwater and to a very limited extent, even terrestrial environments (Pflugfelder 1933). Some nereids are easy to maintain under laboratory conditions and have been used extensively for experimental research. The most popular of the nereids thus used, is Hediste diversicolor, referred to in the experimental literature incorrectly as Nereis diversicolor in most cases; in other cases, more correctly as Neanthes diversicolor. Some critical problems are associated with the recognition of species within the family, in that all species dorsal view. Boiler Bay. Key to Genera la. 2b (lb). (B). 3a (2a). diagrams of the pharyngeal areas of nereids. California. (F). all setae composite spinigers Cheilonereis 2 3 4 Dendronereis FIGURE 21. vexillosa from Dillon Beach. median parapodium of Ai. Aíereíj vexUlosa. 2a (1 b). 25x. lb. in ventral and dorsal views. 50x. defined on non-morphological features. It is suspected that in fact a series of widespread species (Nereis pelágica. lOx. (E). Peristomium forms a large ventral collar Peristomium not ventrally enlarged Some notopodia with pectinate branchiae Branchiae absent Branchiae arise from the dorsal cirrus. Platynereis dumerilii and others) may turn out to be species-complexes. median parapodium of the above. (C) and (D). Neanthes virens. . (A). Family NEREIDAE. It is thus of importance that the provenance of experimental organisms be stated clearly in all pubUcations.86 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 cannot be recognized on strictly morphological characters (Smith 1958). Oregon. tntertidal. lOx. venu-al view of the above. 12a (1 lb). paragnaths present 26 Paragnaths present on one pharyngeal ring only 27 Paragnaths present on both pharyngeal rings 30 Paragnaths present on the maxillary ring only 28 Paragnaths present on the oral ring only 29 Paragnaths in eight groups. apodous segment present 17 Dorsal cirri attached basally on the notopodial superior lobes 18 Dorsal cirri attached distally on the notopodial superior lobes 19 Notopodial homogomph falcigers present in posterior setigers Rullierinereis Notopodial homogomph falcigers absent Nicon Superior notopodial lobes long and straplike. apodous segment absent Micronereides Four pairs of tentacular cirri. anterior apodous segment present Cryptonereis A single median antenna present 8 Two antennae present 9 Paragnaths present on the maxillary ring. 21b (20a). parapodia uniramous Dawbinia Notocirri of parapodia 5-7 broadly elytraeform Kainonereis Notocirri of parapodia 5-7 cylindrical and cirriform 10 Notopodia strongly reduced or absent 11 Median and posterior notopodia well developed. in addition to the jaws 15 Eversible pharynx with jaws. dorsal cirri distally attached .]ÇJ6 3b (2a). 8b (7a). 18a (17a). 15a (14a). 23a (22b). 19a (17b). but otherwise smooth 16 Eversible pharynx with soft papillae only 20 Eversible pharynx with at least some paragnaths 25 Two pairs of tentacular cirri. 27a (26a). 24a (23b). 14a (10b). 26b (25b). 18b (17a). with lobes and setae 14 Notosetae present Namanereis Notosetae absent 12 Tentacular cirri articulated Lycastoides Tentacular cirri smooth 13 Notacicula present. 10b (9b). 26a (25b). 16b (14b). 5b (4b). 28b (27a). antennae and cirri well developed Namalycastis Notacicula absent. 9b (7b). 6a (5 a). all conical Ceratonereis Notopodial homogomph falcigers present in posterior setigers Eunereis Notopodial homogomph falcigers absent Websterinereis . 7b (5b). 20b (15a). 27b (26a). 9a (7b). 5a (4b). 20a (15a). 29b (27b). 11a (10a). 15b (14a). parapodia biramous (except the first two) Unanereis Paragnaths absent. some composite falcigers present Dendronereides Anterior ventnim with transverse fleshy ridges Australonereis Anterior ventrum smooth * Antennae absent ^ At least one antenna present ^ Two pairs of tentacular cirri present.. 16a (14b). 4b (2b). 7a (5b). 4a (2b). 22b (20b). 1 lb ( 10a). anterior apodous segment absent Micronereis Three pairs of tentacular cirri present. 17a (16b). posterior dorsal cirri long and whiplike Gymnonereis Accessory dorsal cirri absent. dorsal cirri basally attached Kinberginereis Eversible pharynx with both papillae and paragnaths Leonnates Papillae absent. 29a (27b). posterior dorsal cirri not whiplike 24 Pharyngeal papillae on both rings. 10a (9b). 13a (12b). antennae and cirri reduced Lycastopsis Eversible pharynx with either papillae or paragnaths or both. 25a (15b). THE POLYCHAETE WORMS 87 Branchiae arise from the notopodial lobes. Tylorrhynchus Pharyngeal papillae on oral ring only. 23b (22b). 24b (23b). inferior neuropodial lobe absent Steninonereis Superior notopodial lobes large and folióse. 8a (7a). 22a (20b). 25b (15b). 19b (17b). inferior neuropodial lobes present Leptonereis Pharyngeal papillae at least in part in tufts 21 Pharyngeal papillae solitary 22 All setae homogomph spinigers Tylonereis Setae include also neuropodial homogomph falcigers in posterior setigers Laeonereis Ventral cirri double at least in some setigers Ceratocephale All ventral cirri simple 23 Accessory dorsal cirri on some anterior setigers. inferior neuropodial lobes present. inferior neuropodial lobes absent. 6b (5a).. all rod-shaped Solomononereis Paragnaths in patches and bands. 21a (20a). 12b ( 1 lb). 17b (16b). 28a (27a). 14b (10b). 13b (12b). malaitae Gibbs 1971. Eversible pharynx smooth or with soft papillae. Four pairs of tentacular cirri. 3 species. C. 53 species. Four pairs of tentacular cirri. parapodia biramous with capillary setae. 7 species. 32b (31b). if present. 31a (30a). Notosetae include homogomph spinigers and blunt simple falcigers. Notosetae homogomph spinigers. Nereis cyclurus Harrington 1897. Branchiae present as subdivisions of the dorsal cirri. neurosetae heterogomph spinigers and falcigers. only species. Branchiae present as a subdivision of the notopodial superior lobes. parapodia biramous. Three pairs of tentacular cirri. 36a (35b). arborifera Peters 1854. Eversible pharynx with soft papillae on both rings. D. inferior neuropodial lobe may be present. C. usually on most. Eversible pharynx without paragnaths or papillae. Eversible pharynx with conical paragnaths on both rings. biramous parapodia. Frontal antennae absent. With fleshy transverse ridges across anterior ventrum. biramous parapodia. parapodia uniramous. Superior notopodial lobes large and folióse with dorsal cirrus attached medially. neurosetae homo. Cryptonereis Gibbs 1971. Four pairs of tentacular cirri. Dendronereis Peters 1854. Neuropodia in anterior setigers multifid. composite falcigers present or absent 34 Notopodial homogomph falcigers present in posterior setigers Nereis Notopodial homogomph falcigers absent Neanthes Paragnaths include pectinate bars and usually small patches of cones. only species. Eversible pharynx with conical paragnaths on both rings. Cheilonereis Benham 1916. biramous parapodia. transverse smooth bars absent Platynereis Paragnaths include transverse smooth bars. Eversible pharynx without paragnaths or papillae. 35a (30b). pectinate bars absent Perinereis Generic Definitions Australonereis Hartman 1954. Peristomium greatly expanded ventrally. 33b (32b). Ceraiocephale Malmgren 1867. Eunereis Malmgren 1867. D. heteropoda Southem 1921. Cirronereis Kinberg 1866a. Neuropodial inferior lobes absent. Four pairs of tentacular cirri. Neurosetae homogomph spinigers and heterogomph falcigers. 35b (30b). 34b (33b). C. 2 species. A single median antenna present. Nereis longissima Johnston 1840.and heterogomph falcigers. Dendronereides Southern 1921. Ventral cirri double on at lest some setigers. Notosetae homogomph spinigers. gracilis Kinberg 1866a. Notosetae homogomph spinigers. neurosetae homo. 36b (35b). 33a (32b). oral ring bare. in notopodial positions 33 With blunt simple falcigers in notopodia Cirronereis Simple falcigers absent. Four pairs of tentacular cirri. D. pectinate bars usually present Pseudonereis Superior notopodial lobes not expanded in any setigers. 30b (26b). . parapodia biramous. Four pairs of tentacular cirri. Dawbinia Benham 1950. Eversible pharynx with soft papillae on the maxillary ring. at maturity. only species. 35 All setae homogomph spinigers Nectoneanthes At least some falcigers present 32 Middle and posterior neuropodia with single homogomph falcigers Hediste Homogomph falcigers. patches of cones and sometimes pectinate bars 36 Superior notopodial lobes greatly expanded in posterior setigers. only sf)ecies.88 30a (26b). loveni Malmgren 1867. 32a (31b). Two pairs of tentacular cirri. mirabilis Kinberg 1866a. 2 species. Four pairs of tentacular cirri. 31b (30a). C. 34a (33b).and heterogomph spinigers and heterogomph falcigers. biramous parapodia. Setae include homogomph and heterogomph spinigers and heterogomph falcigers. neurosetae homoand heterogomph spinigers and heterogomph falcigers. uniramous parapodia.. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 All paragnaths conical 31 Cones and in addition either transverse or pectinate paragnaths or both present on the pharynx . All setae homogomph spinigers. Nereis (Leonnates) ehlersi Augener 1913a. biramous parapodia. neurosetae homogomph spinigers and heterogomph falcigers. II species. Ceratonereis Kinberg 1866a. Eversible pharynx with soft papillae on both rings. Dorsal cirri attached basally to the superior notopodial lobe. Neurosetae heterogomph spinigers and falcigers. Notosetae include homogomph spinigers and falcigers. Eversible pharynx with paragnaths on the maxillary ring only. aucklandica Benham 1950. only species. Two pairs of tentacular cirri present. All setae homogomph spinigers. Tentacular cirri jointed. Nereis culvert Webster 1879a. neurosetae heterogomph falcigers with coarsely serrated blades. alata Chamberlin 1919c. neurosetae heterogomph spinigers and falcigers. Antennae bifid at the tips. Kinberginereis Pettibone 1971a. Antennae absent.1976 THE POLYCHAETE WORMS m Eversible pharynx with paragnaths on oral ring only. 2 species. only species. Superior notopodial lobes large and folióse in posterior setigers. variegata Claparède 1863. Eversible pharynx without papillae or paragnaths. parapodia biramous. the latter with long appendages. Notopodial superior lobes prolonged in posterior setigers. Four pairs of tentacular cirri. eyes absent. parapodia sub-biramous or uniramous. neurosetae homogomph and heterogomph spinigers. parapodia biramous. the latter in posterior setigers. Leptonereis Kinberg 1866a. Two pairs of tentacular cirri present. Eversible pharynx smooth or with soft papillae. neurosetae homo. only species. Four pairs of tentacular cirri. Four pairs of tentacular cirri. Laeonereis Hartman 1945. Neurosetae homoand heterogomph spinigers. Four pairs of tentacular cirri. Four pairs of tentacular cirri. L. Four pairs of tentacular cirri. Neanthes Kinberg 1866a. Inferior neuropodial lobe present. Notosetae homogomph spinigers and falcigers. Notosetae homogomph spinigers. M. L. Neurosetae heterogomph falcigers and spinigers. Three pairs of tentacular cirri present. All setae homogomph spinigers. parapodia biramous. Micronereides Day 1963. Notosetae homogomph spinigers. Kainonereis Chamberlin 1919c. parapodia biramous. parapodia biramous. Four pairs of tentacular cirri present. Lycastis quadraticeps Blanchard 1849.and heterogomph spinigers and heterogomph falcigers. Apodous segment absent. Eversible pharynx with conical paragnaths on both rings. 5 species. Inferior neuropodial lobe present. L. Paranereis abiuma Müller in Grube 1871. vaalii Kinberg 1866a. Three or four pairs of tentacular cirri present. parapodia biramous. parapodia biramous. Eversible pharynx without papillae or paragnaths.and heterogomph spinigers and heterogomph falcigers. N. Nereis (Alitta) oxypoda Marenzeller 1879. neurosetae homo. Nereis diversicolor O. 2 species. Lycastopsis Augener 1922. Eversible pharynx without papillae or paragnaths. parapodia uniramous. laevis Kinberg 1866a.F. indicus Kinberg 1866a. 18 species. Accessory dorsal cirri present on some anterior segments. parapodia biramous. Eversible pharynx without papillae or paragnaths. only species. Four pairs of tentacular cirri. Eversible pharynx with papillae on the oral ring only. Antennae and cirri reduced. L. parapodia biramous with notopodia strongly reduced. Four pairs of tentacular cirri. 2 species. Namanereis Chamberlin 1919c. only species. posterior dorsal cirri long and whiplike. Namalycastis Hartman 1959. broad elytraeform dorsal cirri on parapodia 5-7. capensis Day 1963. Neurosetae heterogomph spinigers and falcigers. Eversible pharynx with conical paragnaths on both rings. K. Four pairs of tentacular cirri. Notosetae homogomph spinigers. Gymnorhynchus sibogae Horst 1918. 6 species. Neurosetae include heterogomph spinigers and falcigers. 50 species. Nectoneanthes Imajima 1972. Neuropodia with a single setal lobe only. heterogomph falcigers. Gymnonereis Horst 1919a. alticola Johnson 1903. All setae homogomph or slightly hemigomph spinigers. parapodia . Four pairs of tentacular cirri present. M. Apodous segment absent. 2 species. Eversible pharynx with papillae on the oral ring and paragnaths on the maxillary ring. parapodia biramous. Notosetae homogomph spinigers. Eversible pharynx with conical paragnaths on both rings. beumeri Augener 1922. Eversible pharynx without papillae or paragnaths. Eversible pharynx without papillae and paragnaths. Notosetae homogomph spinigers. 10 species. Lycastoides Johnson 1903. Micronereis Claparède 1863. dorsal cirri basal. parapodia uniramous. Four pairs of tentacular cirri present. Leonnates Kinberg 1866a. Inferior neuropodial lobes present. Hediste Malmgren 1867. Nereis (Leptonereis) inermis Hoagland 1920. Müller 1776. neurosetae heterogomph spinigers and falcigers. Eversible pharynx with soft papillae on the oral ring only. Eversible pharynx with tufts of papillae on both rings and large solitary papillae on area VI. parapodia biramous. neurosetae homogomph spinigers and falcigers. Notosetae homogomph spinigers. 6 species. parapodia biramous. A single homogomph falciger present in median and posterior neuropodia. Eversible pharynx without paragnaths or papillae. Notosetae usually absent. P. 2 species.and heterogomph spinigers and heterogomph falcigers. novaehollandiae Kinberg 1866a. Eversible pharynx with eight groups of rod-shaped paragnaths on the maxillary ring. parapodia biramous. Unanereis Day 1962. Eversible pharynx without papillae and paragnaths. Sotomononereis Gibbs 1971. Notosetae homogomph spinigers. only species. Eversible pharynx with paragnaths on both rings. but the character has the distinct drawback that it is dependent on interpretation of shapes. only species. including cones. pictus Kinberg 1866a. Four pairs of tentacular cirri. Platynereis Kinberg 1866a. Four pairs of tentacular cirri. 7 species. neurosetae hemi. Notosetae homogomph spinigers and falcigers. A^.and heterogomph spinigers and heterogomph falcigers. parapodia biramous. neurosetae homoand heterogomph spinigers. transverse smooth bars and pectinate bars. Eversible pharynx with paragnaths on the oral ring only. rnaranensis Gibbs 1971. magalhaensis Kinberg 1866a. including cones. Notosetae homogomph spinigers. parapodia biramous. Inferior neuropodial lobes present. Eversible pharynx with soft papillae on the maxillary ring only. long and straplUce in posterior setigers. Inferior neuropodial lobes absent. Four pairs of tentacular cirri. Pettibone (1971a) introduced characters of the parapodial lobes (called ligules by Pettibone) as major features in the generic identification. neurosetae homo. and heterogomph falcigers. parapodia biramous. Websterinereis Pettibone 1971a. Perinereis Kinberg 1866a. heterogomph spinigers and heterogomph falcigers. parapodia biramous. neurosetae homo. Eversible pharynx with conical paragnaths on both rings. with dorsal cirrus inserted between the superior and the median lobes. U. neurosetae homoand heterogomph spinigers and heterogomph falcigers.and heterogomph spinigers and heterogomph falcigers. Eversible pharynx with conical and transverse paragnaths on both rings. Notosetae homogomph spinigers. Nereis tridentata Webster 1880. Taxonomic Notes The generic subdivision of the nereids has been based mainly on the pharyngeal structures and the presence of specific kinds of setae in the parapodial rami. four pairs of tentacular cirri.and heterogomph spinigers and heterogomph falcigers. parapodia biramous. Eversible pharynx without papillae or paragnaths. Four pairs of tentacular cirri. 134 species. bogoyawlenskyi Fauvel 1911. Four pairs of tentacular cirri present. macgregori Day 1962. only species. T. 5 species. parapodia biramous. Notosetae hemigomph spinigers.and heterogomph spinigers and heterogomph falcigers. gallapagensis Kinberg 1866a.90 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 biramous. Leptonereis zebra Rullier 1963. Both characters are subject to some variation within each genus. Eversible pharynx with paragnaths on both rings.and heterogomph spinigers and heterogomph falcigers. which is noto- . Notosetae homogomph spinigers and falcigers. parapodia biramous. pelágica Linnaeus 1758. 60 species. Rullierinereis Pettibone 1971a. Four pairs of tentacular cirri. neurosetae homo. and pectinate bars. Nicon Kinberg 1866a. neurosetae include homo. Eversible pharynx without papillae or paragnaths. Superior notopodial lobes. and especially the pharyngeal structures require acciu'ate dissection of the anterior end. Four pairs of tentacular cirri. martini Wesenberg-Lund 1958. Four pairs of tentacular cirri present. Nereis Linnaeus 1758. Notosetae include homogomph spinigers and falcigers. 5. Tylonereis Fauvel 1911. neurosetae homo. neurosetae homogomph spinigers and falcigers. Expanded superior notopodial lobes in median and posterior seliger.and heterogomph spinigers and heterogomph falcigers. only species. Eversible pharynx with conical paragnaths on the maxillary ring. this may be valid. A single median antenna present. neurosetae hemi-. All setae homogomph spinigers. Four pairs of tentacular cirri. parapodia biramous. Steninonereis Wesenberg-Lund 1958. 2 species. Notosetae include homogomph spinigers and falcigers. Notosetac include homogomph spinigers and falcigers. parapodia biramous. neurosetae include homo. Notosetae include homogomph spinigers and falcigers. the latter in posterior setigers. Notosetae homogomph spinigers. parapodia biramous. parapodia biramous. P. the latter in median and posterior setigers. Tylorrhynchus Grube 1869a. Eversible pharynx with soft papillae on both rings. Four pairs of tentacular cirri. Four pairs of tentacular cirri. the latter sometimes fused to form simple falcigers. oral ring bare. 5. Inferior neuropodial lobes present. All setae homogomph spinigers. Pseudonereis Kinberg 1866a. Nereis heterochaeta Quatrefages 1865. N. 15 species. P. 20 species. NOMEN NUDUM Stratonice Malmgren 1867. see Perinereis Heminereis Quatrefages 1865. At the tip of the pharynx are four short. indeterminable Heteronereis Örsted 1843a. see Platynereis Nereilepas Blainville 1828. see Pseudonereis Meganereis Blainville 1818. s&e Platynereis Lepidonereis Blainville 1818. notosetae. seeNeanthes Arete Kinberg 1866a. which they will evert when disturbed. lb. see Pseudonereis Nectonereis Verrill 1873b. NOMEN NUDUM Alitta Kinberg in Malmgren 1865. slender bodies and conical prostomia. 2a (lb). see Nereis Protolycoris Hatschek 1893. FAMILY GLYCERIDAE GRUBE 1850 Glyceriform polychaetes with long. see Hediste Phyllonereis Hansen 1882. Parapodia either all biramous or all uniramous. see Perinereis Paranereis Kinberg 1866a. in all depths. aileron of jaws with lateral wing E*rostomium short. see Perinereis Tetratrocha Sveshnikov 1959. see Perinereis Branchionereis Blainville 1818. Eversible pharynx unarmed. see Platynereis FAMILY Antonbruunidae NEW NAME Nereidiform polychaetes with cylindrical bodies. s&e Platynereis Leontis Malmgren 1867. see Gymnonereis Hedyle Malmgren 1867. indeterminable Uncinereis Chamberlin 1919c. indeterminable Naumachius Kinberg 1866a. indeterminable Lipephila Malmgren 1867. seeCeratocephate Chinonereis Chamberlin 1924. larvae. cylindrical eversible pharynx. indeterminable Gnatholycastis Ehlers 1920. The glycerids are long. Key to Genera la. indeterminable Praxithea Malmgren 1867. see Namanereis Lycastis Audouin and Milne Edwards 1833b. see Tylorrhynchus arroceros Claparède 1863. see Pseudonereis Phaetusa Castelnau 1842. with four jaws or with a circlet of jaws. reflecting the more conservative approach taken to lumping at the specific level in this study. 2b (lb). with small males. Parapodia uniramous throughout Parapodia biramous throughout Prostomium long. slender polychaetes with numerous segments. simple. this is unfortunate. Eversible pharynx with four jaws in a cross. All setae simple. but the numbers of species indicated for each genus is higher than as given by Pettibone. usually black jaws. inquilines in bivalves. indeterminable Cirronereis Blainville 1818. Parapodia biramous. They appear to be mainly carnivores. they are rarely present in large numbers. see Pseudonereis Pisenoe Kinberg 1866a. reflects the added insights of Pettibone at the generic level. with about three rings. see Neanthes Nicomedes Kinberg 1866a. Suborder Glyceriformia Phyllodocida with two pairs of antennae. Perhaps their most noticeable feature is the very long. indeterminable Nossis Kinberg 1866a. NOMEN NUDUM Chaunorhynchus Chamberlin 1919c. viridis living in the mantle cavity of the bivalve Lucina fosteri Hartman and Boss. Invalid Genera Aceronereis Blainville 1818. see Nereis ¡phinereis Malmgren 1865. Antonbruunia Hartman and Boss 1965 with genotype ^4. Glycerids are present. Sexual dimorphism present. palps and tentacular cirri absent. three antennae and simple palps. First parapodia lateral. Pettibone in the same paper lumped series of species based on overlapping ranges in different characters. no species named Tkoosa Kinberg 1866a. when present. see Ncanalycastis Lycastoides Jakubova 1930. ^ee Perinereis Lycastella Feuerbom 1932. see Nereis Masligonereis Schmarda 1861. Proboscis either unarmed. indeterminable Paralycastis Ehlers 1920. Two pairs of tentacular cirri present. see Nereis Typklonereis Hansen 1878. with more than three rings. This family is known for a single genus. see Namanereis Lycoris Savigny 1818. but notopodia reduced to dorsal cirri supported by internal acicula. see Platynereis Podonereis Blainville 1826. aileron of jaws a simple rod Hemipodus 2 Glycera Glycerella . and very difficult to quantify. Neurosetae composite. The generic key given above.1976 THE POLYCHAETE WORMS 91 riously dependent on the experience of the observer. since the variability within any single population of these animals has never been examined and quantified in detail. see Perinereis Gymnorhynchus Horst 1918. mainly in soft sandy or muddy substrates. essentially in two major genera. Bathyglycinde). since the characteristic ridges may be very difficult to see and depend on the full resolution of the mictoscope. the whole body must be scanned for the presence or absence of these structures. FIGURE Invalid Genera Euglycera Verrill 1881. neurosetae composite spinigers. (B). Key to Genera la. Pharyngeal organs of tnany kinds. The pharyngeal organs are considerably larger than in the glycerids. off Santa Barbara. in addition to parapodial lobes and branchiae. Specific identification depends on study of the pharyngeal organs. The goniadids often are considered as part of the glycerids.64x. but because of their structure. 22. with at least three. eversible pharynx with a circlet of smaller and larger jaw-pieces. and frequently are retracted in parts of the body. Glycera amerkam. notosetae simple. unicornis Savigny 1818. (A) Family GLYCERIDAE. Aileron of jaws rodlike. 2 species. 55 species. 1866b Glyceriforms with long and slender bodies. with seven to ten vague annuli. aileron of jaws rod-shaped. seit Hemipodus Proboscidea Blainville 1825. lb. Branchiae may be retractable. Hemipodus Quatrefagcs 1865. 1950. after Hartman. Pharyngeal organs elongate oval or filamentous. usually five to seven annuli. Phaiyngeal organs long and slender. posteriorly biramous. Phaiyngeal organs can be characterized only by very close work under compound microscopes. all setae composite spinigers. They resemble the glycerids in that their most remarkable structure is the eversible pharynx. very long. covered with pharyngeal organs. median parapodium. Glycera capitata. has been stable for the last 50 years and not much change is anticipated. Parapodia uniramous. Prostomiiun is conical. Taxonomic Notes The generic subdivision. Prostomium short. see Glycera Hemipodia Kinberg 1866b. usually slender and crowned with a series of teeth. Identification of species. It is of great importance that the light be very accurately adjusted. but the fwo groups differ sufficiently that a recognition at the family level is warranted. posterior view. Neurosetae composite. Glycera rosea Blainville in Quatrefages 1865. not because of their small sizes. posterior ones biramous Eversible pharynx with organs of many kinds Progoniada 2 3 . Parapodia anteriorly uniramous. see Glycera Hamiglycera Ehlers 1908.92 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Prostomium long. rarely all uniramous. and are partly sclerotinized in some genera {Glycinde. see Glycera Rhynchobolus Claparède 1868. see Glycera Telake Chamberlin 1919c. intertidal. and the number of species may be subject to considerable adjustment. Aileron of jaw with lateral wing. 2a (lb). 15 species. Glycerella Arwidsson 1899. especially in certain groups of Glycera is quite difficult. Hemipodus magellanicus Mclntosh 1885. capillary or acicular. Notosetae simple. neurosetae composite spinigers. Prostomium long. All parapodia uniramous Anterior parapodia unh-amous. with maximally four annuli. see Glycera FAMILY GONIADIDAE KINBERG Generic Definitions Glycera Savigny 1818. lOx. Notosetae capillaries. G. posterior ones both spinigers and falcigers Anterior setae all falcigers. 5b (4a). B. only species. Eone gracilis Verrill 1873b. Goniadella Hartman 1950. 3b (2a). mainly of one kind. posterior neurosetae both falcigers and spinigers. Bookhoutia Mohammad 1973. 7a (6b). posterior view. 1950. (C). mexicana Fauchald 1972. emérita Audouin and Milne Edwards 1833b. Notosetae acicular. aciculata Hartmann-Schröder 1960a. pharyngeal organs small. neurosetae composite spiningers. Chevrons absent. (A). anterior neurosetae all falcigers. median parapodium. 4b (2b). G. pharyngeal organs of one kind. 6a (4b). All neurosetae composite falcigers. oligognatha Mohammad 1973. B. 20 species. Chevrons present. Posterior segments biramous. 1950.5x. 31x. Posterior segments biramous. Goniada Audouin and Milne Edwards 1833b. 8b (7b). 12. G. Glycinde armígera. Chevrons absent. Notosetae capillaries. THE POLYCHAETE WORMS 93 4 Bathyglycinde Glycinde 5 6 Goniada Goniadella Ophioglycera 7 Goniadides 8 Bookhoutia Goniadopsis Eversible pharynx with organs of one or a few kinds Notosetae slender capillaries Notosetae knobbed or falcate hooded hooks Eversible pharynx with chevrons Eversible pharynx without chevrons Neuropodia with spmigers only Neuropodia with both spinigers and falcigers Neuropodia with spinigers only At least some falcigers present Notopodia reduced to a rounded knob with a few thick acicular setae and a cirrus Notopodia well developed Anterior setae all falcigers. Notosetae knobbed or falcate hooded hooks. after Hartman. 25x. Notosetae acicular spines. Goniadopsis Fauvel 1928a. Chevrons present. Neuropodia in uniramous part of body with composite falcigers.¡g76 2b (lb). Goniada brurmea. neurosetae composite spinigers. Posterior segments biramous. 3 species. Glycinde Müller 1858. G. Posterior segments biramous. 4a (2b). 5a (4a). after Hartman. 34 species. anterior end with pharynx half everted. neurosetae composite spinigers. 2 species. agnesiae Fauvel 1928a. (B). pharyngeal organs large and of several kinds. Goniadides Hartmann-Schröder 1960a. multidens Müller 1858. anterior end of the above. pharyngeal organs large and of several kinds. FIGURE . G. Notopodia reduced to short rounded lobes with a cirrus and one or two coarse acicular setae. 2 species. 3 species. Family GONIADIDAE. spinigers. 3a (2a). 7b (6b). Posterior segments biramous. Chevrons absent. 6b (4b). Chevrons absent. Notosetae acicular or capillary. Composite spinigers and falcigers present on all neuropodia. pharyngeal organs small and of one kind. 8a (7b). Posterior segments biramous. posterior ones all spinigers Generic Definitions Bathyglycinde Fauchald 1972. those in biramous part. Posterior segments biramous. ¿5> 23. Chevrons absent. Neurosetae include composite falcigers and spinigers in all parapodia. prostomium trapezoidal in outline. in most species at least two kinds are distinctly chitinized projections. Paralacydonia sp. regularis Hartman 1965. 25x. Antennae short and biarticulated. They traditionally have been considered as an appendix to the phyllodociform polychaetes. 2b (lb). but the parapodia and setal structures. San Clemente Basin. Family LACYDONIIDAE. mainly because of the lack of proboscideal armament. Invalid Genera Eone Malmgren 1866. caeca Rullier 1965b. miranda Marion and Bobretzky 1875. only first parapodia uniramous. 2 species. first twcethree parapodia uniramous. California.and neuropodia are developed equally. (A). notosetae simple. all others biramous. Tentacular cirri present Tentacular cirri absent Antennae short and biarticulated. In the other genera. Antennae long and slender. . 1914 Glyceriform polychaetes with short and slender bodies. SOx. median parapodium of the above. The lacydoniids recently were reviewed by Uschakov (1972). pharyngeal organs of one kind and short. paradoxa Fauvel 1913. 3 species. eversible pharynx unarmed. First parapodia uni. Paralacydonia Fauvel 1913. only species. 6 species. 24. gigantea Verrill 1885.94 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Ophioglycera Verrill 1885. 1500 m. tentacular cirri absent. 4 species. first setiger biramous Lacydonia 2 Paralacydonia Pseudolacydonia Generic Definitions Lacydonia Marion and Bobtetzky 1875.. all neurosetae composite spinigers. Posterior segments biramous. Progoniada Hartman 1965.or biramous. In Glycinde and Bathyglycinde at least three distinct kinds of organs are present. (B). 2a (lb). The following five families do not appear to be closely related to any other I%yllodocida nor are they obviously related to each other. see Glycinde Epicaste Kinberg 1866b. Notosetae slender and acicular. most species have only one kind of pharyngeal organ. ally them more closely with the glycerids than with any other group of polychaetes. FIGURE Key to Genera la. O. but in some cases two kinds may be identifiable. L. Pseudolacydonia Rullier 1965b. first setiger uniramous Antennae long and smooth. They are listed in alphabetical order. Chevrons absent. as well as the structure of the prostomium. The only difficult point in the key above appears in dichotomy 2 on the question of a few or many kinds of pharyngeal organs. Neurosetae composite. All segments uniramous. P. all parapodia birjimous. not articulated. the present treatment follows his closely. tentacular cirri absent. Antennae short and smooth. Taxonomic Notes The genera appear well established in the Goniadidae. P. see Glycinde Lacharis Kinberg 1866b. see Goniada FAMILY LACYDONIIDAE BERGSTRöM The lacydoniids somewhat resembles the nephtyids. in that both noto. P. tentacular cirri present. Chevrons present. indeterminable Leonnatus Kinberg 1866b. lb. but recognizable parapodia and setae. the presence and structure of the anterior appendages (sometimes the lack of such appendages). makes it difficult to ally these forms with any other polychaetes. Phalacrophorus pictus borealis. 50x. in that they have one pair of antennae and one pair of palps. Prostomium without antennae. This connection seems based on adaptive convergencies to the pelagic environment. from which they differ sharply in the development of the parapodia. 3b (2b). a reduction in the number and importance of the setae and the lightly built. Family ISOPILIDAE. They are not related to the nephtyids. 1914 Pelagic. The nephtyids and sphaerodorids either are considered related to the glyceriforms or placed in the vicinity of the nereids. Eversible pharynx present. but appear to differ rather sharply from members of this family in most of the characters usually considered at the familial level. it appears for the time being best to leave the nephtyids as a free-standing fiimily within the order Phyllodocida. Key to Genera la. all setae composite. 1972. but has not been assigned to any suborder or super-familial group. appear to have an extensive and varied degree of cephalization (Fauchald 1974b) and appear isolated in the Phyllodocida. modified after Uschakov. In all five cases. 3a (2b). and insist that all families must be allied with a suborder. Rather than forcing the issue. small. slender Phyllodocida. The sphaerodorids. Eversible pharynx with a parr of curved jaws Eversible pharynx unarmed Up to 11 anterior segments with reduced parapodia Parapodia fully developed from segment 5 Dorsal and ventral cirri present. in that even the first seliger carries small. The nephtyids are extremely poorly cephalized compared to most other polychaetes. FIGURE 25. The same can be said for the tomopterids and the typhloscolecids (Uschakov 1972). armed in some cases. FAMILY IOSPILIDAE BERGSTRôM The iospilids oñen are considered allied with the phyllodocids. but small on segments 2-3 Dorsal and ventral cirri absent on segments 2-3 Phalacrophorus 2 ¡ospilopsis 3 Paraiospilus lospilus . 2b (lb). For these reasons. 2a (lb). The iospilids differ from the phyllodociform families in the presence of palps and the lack of antennae. most of which are related to the Phyllodocidae. lospilidae.7976 THE POLYCHAETE WORMS 95 The three families with pelagic members. Tomopteridae and Typhloscolecidae. The family was revised by Dales and Peter (1972) and Uschakov (1972). without any commitment as to further affiliation. according to the innervation pattern whereas the phyllodocids have at least two pairs of antennae and true palps are absent. usually are considered with the bulk of the other pelagic polychaetes. two short palps present. such as a frequently folióse condition of the parapodial lobes. 1 b. The lack of cephalization places the nephtyids close to the phyllodocids. It is here considered a member of the order Phyllodocida. it here is considered best to leave all five as separate entities. often translucent bodies in these forms. Parapodia uniramous. however. and rather more gently posteriorly. affinis Viguier 1911. 70 m. pelagic. P. 3 species. Up to U anterior segments with rudimentary parapodia. 75x. jaws absent. dorsal and ventral cirri present. straightsided bodies abruptly tapering anteriorly. Interramal. parapodium of the above. (B). Nephtys catiforniensis. off Santa Catalina Island. lospilus Viguier 1886. One pair of antennae and one pair of very short palps present. Maximally four anterior segments with reduced parapodia. off central California. 15x. Family NEPHTYIDAE. and are most common in sandy and muddy substrates. (A). All setae simple. Family SPHAERODORIDAE. 2 species. The nepthyids may superficially be confused with the sigalionids in that both groups have long. off Santa Catalina Island. only species. Family TOMOPTERIDAE. dorsal and ventral cirri absent on segments 2 and 3. respiratory cirri present in nearly all foims. P. (E). FAMILY NEPHTYIDAE GRUBE lospilopsis Augener 1922. phalacroides Viguier 1886. /. /.and postsetal lobes.96 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Generic Definitions Science Series 28 Maximally ten anterior segments with reduced parapodia. Nephtyids are present at all depths. Eversible pharynx with paired jaws. 15x. antillensis Augener 1922. and of course the scales present in the sigalionids. FIGURE 26. A pair of lateral jaws present. but rudimentary on segments 2 and 3. California. 1850 Phyllodocida with long. lOx. Maximally four anterior segments with reduced parapodia. the nephtyids lack obvious. (C). (D). median parapodium of the above. only species. 5x. Tomopteris sp. Parapodia biramous with both rami well developed with complex pre. Sphaerodoropsis sphaerulifer. jaws absent. slender bodies with quadrangular cross-sections. Paraiospilus Viguier 1911. long appendages anteriorly.. pictus Greeff 1879. Phalacrophorus Greeff 1879. . jaws absent. However. 50 m. recurved or involute Interranial cirri recurved Interramal cirri involute Eversible pharynx with subterminal and terminal papillae Eversible pharynx without subterminal and terminal papillae Micronephtkys 2 Nephtys 3 Agiaophamus Inermonephtys Generic Definitions Agiaophamus Kinberg 1866b. Interramal cirri involute. lyratus Kinberg 1866b. and cannot be confused with any other group of polychaetes. Ib. Nephtys (Agiaophamus) inermis Ehlers 1887. except perhaps Sphaerosyllis (Syllidae). see Agiaophamus Aonis Savigny 1822. acicula distally hooked. acicula in most forms with a distal cap. larval forms. Interramal cirri involute. The first start of the interramal cirri is seen best from the lateral side. Identification to species is not difficult. which tend to be better treated than shallow-water ones. terminal papillae long Levidorum 2 3 6 Sphaerephesia 4 . Eversible pharynx with 14 (rarely 16) rows of papillae. second third and last third). Eversible phiirynx with 22 rows of papillae. however. 45 species. acicula distally hooked in most species. Invalid Genera Aglaopheme Kinberg 1866b. Both dorsal and ventral surfaces smooth At least two rows of dorsal macrotubercles Macrotubercles with terminal papillae Macrotubercles distally rounded Macrotubercles in four rows. Uniramous parapodia with simple or composite setae. Eversible pharynx without papillae. Nephthys minuta Theél 1879. Interramal cirri reduced or absent. set Nephtys FAMILY SPHAERODORIDAE MALMGREN 1867 Small Phyllodocida with short and thick or long. but not capped. since the shape of the parapodia changes along the length of the body. Dorsum with two to many rows of large spherical tubercles. fust third. 2a (lb). the dorsal cirri. To overcome the problem of varying body-length. Two to six antennae and one pair of tentacular cirri present. Inermonephtys Fauchald 1968a. Eversible pharynx unarmed. which has similar rows of tubercles. and the corresponding adults as Nephtys. these tend to have small. 3b (2b). 2b (lb). hombergii Savigny 1818. 3a (2a). Portelia Quatrefages 1865. Nephtys Cuvier in Audouin and Milne Edwards 1833b. Key to Genera la. no species named. see Nephtys Pellucidaria Sveshnikov 1959. They turn out to be quite frequent in deepwater samples. A.1976 THE POLYCHAETE WORMS 97 Key to Genera la. are quite different. Recent revisions include Hartman (1950) and Fauchald (1968a). Interramai cirri rudimentary or absent Interramal cirri well developed. lb. Eversible pharynx with 14 rows of papillae. N. but care must be taken that appropriate segments are compared with each other (or with illustrations). 2b (lb). acicula blunt-tipped. see Nephtys Diplobranchus Quatrefages 1865. Fauchald (1974b) reviewed the group. The setal structures. in the latter case. 50 species. they are short and grub-shaped or more slender. Micronephtkys Friedrich 1937. vermiform. 3b (2a). 2a (lb). Taxonomic Notes Identification of nephtyids to genus can be tricky on small specimens. It is probably only a question of time before they are found in relatively large numbers also in shallow water samples. 3a (2b). Interramal cirri recurved. Characteristically.e. 3 species. barely emerging interramal cirrus can be seen on its ventral side. The sphaerodorids have been overlooked in most collections. and a probe must be used to lift the notopodial cirrus so that the small. nearly straight interramal cirri and one could identify these as Micronephtkys. relatively slender bodies. 5 species. terminal papillae very short Macrotubercles in two rows. comparisons should be made on distinct fractions of the bodies (i. along the dorsum. Invalid Genera Bebryce Johnston 1844. Ephesiella Chamberlin 1919c. Ephesiopsis Hartman and Fauchald 1971. Setae composite. A large recurved hook present in the first seliger in most species. (4b). 20 species.98 4a 4b 5a 5b 6a 6b 7a 7b 8a 8b (3b). Stalked macrotubercles without terminal papillae. Anterior end with a median and two or three pairs of lateral antennae. 3 species. L. A number of genera have been described. E. Sphaerodorum claparedii Greeff 1866. (6a). which has simple recurved hooks Ephesiopsis Macrotubercles stalked 7 Macrotubercles sessile 8 Median antenna as long as. the lateral antennae Clavodorum Median antenna shorter than the lateral antennae Sphaerodoridium All setae composite Sphaerodoropsis All setae simple Commensodorum Generic Definitions Sphaerodoridium Liitzen 1961. setae semicomposite or simple. 1848 Pelagic. scotiarum Hartman 1967. Ephesia gracilis Rathke 1843. anterior appendages absent. Anterior end with a median and two pairs of lateral antennae. only species. Levidorum Hartman 1967. (6b). flattened Phyllodocida. Setae composite. see Sphaerodorum Hypephesia Perrier 1897. Setae simple. All tubercles and papillae absent. folióse rami. questionably Sphaerodorum FAMILY TOMOPTERIDAE GRUBE Clavodorum Hartman and Fauchald 1971. setae in other setigers both simple and composite. Stalked macrotubercles in six or eight rows. or longer than. Anterior end with a short median and two pairs of longer lateral antennae. Sphaerephesia Fauchald 1972. Eversible pharynx unarmed. Commensodorum Fauchald 1974b. Parapodia biramous with elongated bases and short. 5 species. Sessile macrotubercles in four rows. Sphaerodoridium commensalis Liitzen 1961 . atlanticum Hartman and Fauchald 1971. First segment well developed in juveniles and reduced in adults. Four rows of sessile macrotubercles with short terminal papillae. The tomopterids have been referred to the phyllodociform group of polychaeles. guayanae Hartman and Fauchald 1971. (2b). Anterior end with long median antenna and two pairs of lateral antennae. Two rows of macro. all other setae composite. Second segment in adults with a pair of long tentacular cirri supported by long acicula. s&t Ephesiella Pollicita Johnston 1845. all anterior appendages short. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Ail setae simple Sphaerodorum At least some setae composite 5 All setae composite apart from the recurved hooks in the first seliger Epkesiella Both composite and simple setae in all setigers. (3b). Prostomium fused with the first two segments.and two rows of microtubercles on the dorsum. .and two rows of microtubercles on the dorsum. longiseta Fauchald 1972. macrotubercles without terminal papillae. Anterior end with a median and two pairs of lateral antennae. but it appears best to consider the family as consisting of only two genera. (4b). Parapodia with two or three acicula. Sphaerodoropsis Hartman and Fauchald 1971. Anterior end with a median and one or two pairs of lateral antennae. ste Sphaerodorum Thysanoplea Schmidt 1857. Recent revisions include Dales and Peter (1972) and Uschakov (1972). Sphaerodorum Örsted 1843b. only species. apart from the first. but differs rather sharply from these in the structure of the anterior end (Uschakov 1972) and appears best considered as a free-standing family in the order Phyllodocia. C. Sphaerodorum abyssorum Hansen 1878. only species. s&t Sphaerodorum Ephesia Rathke 1843. setae absent. A large recurved hook in the first seliger. Glands of different kinds in the parapodia. including in most species large recmved hooks in the first seliger. 9 species. (6a). Two rows of macro. Setae composite. Anterior end with a median and two or three pairs of lateral antennae. (6b). Setae composite. the sessile macrotubercles with terminal papillae. Four rows of sessile macrotubercles with terminal papillae. only species. Four or more rows of macrotubercles without terminal papillae. two antennae. Sphaerodorum sphaerulifer Moore 1911. S. transparent. (2b). Setae simple. Anterior end blunt. the sessile macrotubercles with terminal papillae. Anterior end with a median and two pairs of lateral antennae. macrotubercles without terminal papillae. 5 species. ¡976 THE POLYCHAETE WORMS 99 Key to Genera la. lb. Tentacular cirri of second segment much longer than the body; parapodial fins restricted to the far distal part of each ramus Enapteris Tentacular cirri as long as, or barely longer than the body; parapodial fins surround the distal part of the rami Tomopieris Generic Definitions Enapteris Rosa 1908, Tomopteris euchaeta Chun 1887; only species. Parapodial rami with folióse lobes along the distal margin only. Tentacular cirri much longer than body. Tomopteris Eschscholtz 1825, T. onisciformis Eschscholtz 1825; 40 species. Parapodia rami with folióse lobes surrounding most of the rami. Tentacular cirri shorter than, or about as long as the body. Invalid Genera Briaraea Quoy and Gaimard 1827, questionably Tomopteris Escholtzia Quatrefages 1865, see Tomopteris Johnstonella Gosse 1853, see Tomopteris FAMILY TYPHLOSCOLECIDAE ULJANIN 1878 Pelagic, transparent, fusiform or torpedo-shaped Phyllodocida. Prostomium without appendages; large folióse nuchal lobes present. Two pairs of tentacular cirri. Eversible pharynx unarmed, but with retortorgan. Parapodia uniramous, with large folióse dorsal and ventral cirri; setae simple. The typhloscolecids are poorly studied in terms of their biology and even basic biological information is missing. Recent reviews are given by Uschakov (1972) and Dales and Peter (1972). 27. (A), Family TYPHLOSCOLECIDAE, Travisiopsis lobifera, after Uschakov, 1972, about 12.5x; (B), Travisiopsis levinseni, anterior end, cirri on left side removed, after Uschakov, 1972, 25x. FIGURE Key to Genera la. lb. 2a (lb). 2b (lb). Prostomium with ciliated ridges dorsally and ventrally Prostomium without ciliated ridges Median prostomial papilla on the dorsal side present Prostomium without median dorsal papilla Generic Definitions Sagitella Wagner 1872, S. kowalewskii Wagner 1872; only species. Prostomium without ciliated ridges; median antenna absent. Travisiopsis Levinsen 1885, T. lobifera Levinsen 1885; 6 species. Prostomium without ciliated ridges; median antenna present (may be indistinct). Typhloscolex Busch 1851, T. muelleri Busch 1851; 6 species. Prostomium with dorsal and ventral ciliated ridges; median antenna present. Typhloscolex 2 Travisiopsis Sagitella JOO NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Taxonomic Notes Science Series 28 The above definitions are after Uschakov (1972: 220-225); there appears to be differences in the structure of the nuchal organs in this family, and this may bt of value as a generic character in a future revision. Invalid Genera Acicularia Langerhans 1878, see Sagitella Nuchubranchiata Treadwell 1928, see Travisiopsis Plotobia Chamberlin 1919c, see Travisiopsis ORDER AMPHINOMIDA Prostomium distinct, a caruncle present and at least one antenna. Pharynx with a muscular, rasplike eversible ventral pad; jaws absent. Parapodia distinct with branching branchiae on at least some setigers. The family Spintheridae usually is associated with the Amphinomidae (Fauvel 1923a; Hartmann-Schröder 1971). The external resemblance of the ectoparasitic spintherids to the euphrosinids first were remarked upon by Sars (1850:210) and next by Johnston (1865: 127-128). Augener (1913a:87), while recognizing the overall external similarity to the short-bodied amphinomids, found the differences sufficient to erect a new family for the single genus. Mantón (1967:10 and 21) remarked upon the unique construction of the eversible pharynx in Spinther compared to all other polychaetes. The genus also differs sharply from the order Amphinomida in general and the erection of a separate order to contain this family seems warranted. The new order, Spintherida, is listed below (p. 103). FAMILY AMPHINOMIDAE SAVIGNY 1818 Amphinomida with either elongate or ovate and flattened bodies. One to five antennae present; palps present. Noto- and neurosetae in tufts, notosetae protective spinous setae, at least in part. Branchiae in branching tufts. The amphinomids are common in shallow water. They generally are referred to as fire-worms, since the spines can lead to general discomfort and infections, if they break off within the inflicted cuts. The amphinomids generally are highly colored shallow-water animals, with very distinct color patterns (cf. Fauvel 1953), but are also present, in less showy editions, in deeper water. Key to Genera la. lb. 2a (lb). 2b (lb). 3a (2a). 3b (2a). 4a (3b). 4b (3b). 5 a (4a). 5b (4a). 6a (5b). 6b (5b). 7a (4b). 7b (4b). 8a (7b). 8b (7b). 9a (8b). 9b (8b). 10a (2b). 10b (2b). 1 la (10a). 1 lb ( 10a). 12a ( 1 la). 12b (lia). 13a (lib). 13b ( 1 lb). Caruncle completely absent; neurosetae simple hooks Hipponoa Caruncle present, usually well developed (may be difficult to discern in some species); neurosetae otherwise 2 Body ovate or fusiform 3 Body elongated with parallel sides and usually abruptly tapering anteriorly and posteriorly 10 Branchiae present on all setigers Branchamphinome Some setigers (anterior or posterior) without branchiae 4 One dorsal cirrus per notopodium 5 Two dorsal cirri per notopodium 7 Caruncle with three parallel longitudinal ridges Benthoscolex Caruncle long and folded with indistinct lateral folds 6 Eyes absent, fu-st pair of branchiae larger than the following ones Bathychloeia Eyes present, first branchiae not larger than the following ones Chloeia Caruncle with high central ridge and two wide flattened lateral folds Notopygos Lateral lobes of caruncle small and hidden under the central ridge, or absent 8 Caruncle wedge-shaped, without crest and folds Sangiria Caruncle with crest and folds 9 Caruncle high, loosely plaited and rugose Chloenopsis Caruncle low, narrowly plaited with a crenulated plate Parachloeia Caruncle small and inconspicuous, stretching through maximally three segments 11 Caruncle large and conspicuous, stretching through at least three segments 14 Branchiae present on all segments from the second or third 12 Branchiae limited to the anterior part of the body 13 Caruncle broadly triangular or cordate Amphinome Caruncle narrow and elongated Pareurythoe First segment with large, anteriorly directed hooks Paramphinome First segment without hooks Linopherus 1976 THE POLYCHAETE WORMS mi FIGURE 28. (A), Family AMPHINOMIDAE, Eurythoe complanata, San Felipe, Golfo de California, intertidal, lOx; (B), median pajapodium of the above, lOx; (C), Family EUPHROSINIDAE, Euphrosine borealis. Murchinson Sound, Greenland, 100 m, lOx; (D), Euphrosine sp., off Point Loma, California, median parapodium, 19x; (E), Family SPINTHERIDAE, Spinlher sp., Point Barrow, Alaska, dredged, 5x. narrowly plaited with crenulated plates. 9 species. with a few filaments only. Chloenea Kinberg. 12 species. present on some anterior setigers only. only species. caruncle limited to the two-three first segments. eyes present. 2 species. Dorsal cirri single. eyes present. reported absent in some forms. if present. only species. Hermodice Kinberg 1857. present on most of the body. Body long. 3 species. Dorsal cirri single. Body long. Hipponoa Audouin and Milne Edwards 1830. Body long. eyes present. ( 14b). marmorata Horst 1912. Body ovate. 1867c is considered a synonym of Chloeia. P. Dorsal cirri single. Bipinnate branchiae from segment 5. eyes present or absent. Aphrodita carunculata Pallas 1766. caruncle low. eyes present. 20 species. eyes present. broadly triangular or cordate. Parachloeia Horst 1912. eyes present. Pareurythoe Gustafson 1930. Branchiae pennate. caruncle about as wide as long. Benthoscolex Horst 1912. caruncle large. N. elongated. First setiger with anteriorly directed hooks. first pair larger than the others. Linopherus Quatrefages 1865. Body long. eyes present. Dorsal cirri single. Chloenea Mclntosh. Chloenea atlántica Mclntosh 1885. Dorsal cirri double. especially strongly developed on posterior setigers. sinuous. 14b 15a 15b 16a 16b (10b). bushy. Notosetae large. Body short and ovate. caruncle short. Dendritically branched branchiae. loosely plaited and rugose. Notopygos Grube 1855. sibogae Horst 1910. 14 species. caruncle small and inconspicuous. eyes present. Branchiae poorly developed. without a distinct median ridge. Small forms. bushy in appearance. (14b). Body ovate. Dendritically branched branchiae present on all setigers. Chloeia Savigny 1818. Dendritically branched branchiae from setiger 6. 4 species. caruncle small. caruncle absent. japónica Gustafson 1930. eyes present or absent. . 6 species. Branchiae dendritically branched. caruncle large. (15b).102 14a (10b). Dorsal cirri single. only species. Branchiae pennate. Eurythoe Kinberg 1857. B. Dorsal cirri single. caruncle with three digitate posteriorly directed lobes. smooth median ridge covering the narrow folded lateral folds. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Caruncle longer than wide with a large smooth. Branchiae dendritically branched. Body ovate. H. Body ovate. with several transverse folds. eyes present. 10 species. Dorsal cirri single. Y-shaped or elongated. sinuous median ridge nearly covering the narrow lateral parts Eurythoe Caruncle about as long as wide or wider. caruncle long with a thick. capensis Kinberg 1857. coeca Horst 1912. caruncle with three longitudinal parallel ridges. Branchiae dendritically branched. ( 15b). Hipponoe Jeffreysii Mclntosh 1868. Branchiae tufted. limited to anterior setigers. E. smooth narrow ridge 16 Furcate setae absent Pherecardia Furcate setae present Pherecardites Generic Definitions Amphinome Bruguière 1789. Eyes absent. with a few deep transverse folds Hermodice Caruncle with a distinct. median ridge. P. Branchamphinome Hartman 1967. Aphrodita flava Pallas 1766. 19 species. caruncle with high central ridge and large flattened lateral lobes. Branchiae dendritically branched. gaudichaudi Audouin and Milne Edwards 1830. Dorsal cirri double. Branchiae tufted. Dorsal cirri single. Amphinome incaruncutata Peters 1854. 1885 differs generically from Kinberg's genus and needs a new name. 1885. Dorsal cirri single. Bathychloeia Horst 1910. Dorsal cirri single. Small species. sinuous. Paramphinome Sars 1869. eyes absent. folded with lateral folds. crinita Grube 1855. with indistinct lateral folds. only species. Body short and ovate. much longer than wide. Chloenopsis new name. Chloenopsis is here proposed as the generic name for the species originally named Chloenea atlántica Mclntosh. retractile curved hooks. 5. narrow 15 Caruncle without distinct median ridge. but elongated bodies in most forms. Aphrodita rostrata Pallas 1776. Etendritically branched branchiae on all segments from the second or third. Body ovate. but long-bodied. Dorsal cirri double. B. antárctica Hartman 1967. caruncle high. Body short and ovate. Notosetae bifurcate. Invalid Genera Amphibranchus Kinberg 1867c. . 2 species. Short-bodied forms with an elongated caruncle. Dorsal cirri single. see Chloeia Chloochaeta Kinberg 1867c. as noted by Fauvel (1913) to combine characters of the euphrosinids with those of the palmyrids. see Amphinome Blenda Kinberg 1867c. As indicated by Orrhage on several occasions. Dorsal cirri single. Short-bodied forms with short prostomium. see Hipponoa Pleione Savigny 1818. larval forms Veleda Castelnau 1842. branching branchiae in rows between the notosetae. which would ally the genus closely to Palmyra rather than to the amphinomidlike forms with their ventral plate-muscle pharynx. indeterminable Eucarunculatus Malaquin and Dehorne 1907. see Chloeia Colonianella Kinberg 1867c. with cylindrical shafts Notosetae flattened. 4 species. caruncle with three longitudinal lobes. Parapodia transverse ridges. see Euryihoe Chloenea Kinberg 1867c. but the euphrosinids make up a distinct. caruncle with median axis and lateral lamellae directed posteriorly. Branchiae bushy. see Amphinome Lirione Kinberg 1867c. structure of the pharynx may not be an overwhelmingly strong character. Neuropodia with composite. 2 species. palps absent. Notosetae flattened smooth paieae. Invalid Genus Lophonota Costa 1841. and it appears best to await further study of these forms to decide the question. Prostomium with a median antenna. Palmyreuphrosyne Fauvel 1913.One pair of antennae. Body long. indeterminable Zothea Risso 1826. Sangiria Horst 1911. Setae include capillaries. strongly curved hooks. eyes absent. Bushy branchiae present from the first seliger. parva Horst 1912. In the meantime. myrtosa Savigny 1818. eyes present. Notopodia represented by membranous ridges supported by simple or furcate setae. smooth and cylindrical. Pherecardites Horst 1912. compact group of forms. indeterminable FAMILY EUPHROSINIDAE WILLIAMS 1851 Amphinomida with short and thick bodies. see Linopherus Rastraría Haecker 1898. Branchiae with few filaments. see Chloeia Thesmia Kinberg 1867c. and furcate setae. possibly Amphinome Pseudeurythoe Fauvel 1932. see Amphinome Didymobranchus Schmarda 1861. caruncle with a narrow smooth ridge bordered by wide lateral lobes with deep parallel folds on both sides. Pharynx retractable and cylindrical (resembles a turbe Harlan pharynx). only species. Fauvel indicates that the eversible pharynx of Palmyreuphrosyne should be ventral. smooth paieae Euphrosine Palmyreuphrosyne Generic Definitions Euphrosine Savigny 1818. paradoxa Fauvel 1913. see Notopygos Lycaretus Kinberg 1867c. Key to Genera la. Branchiae in transverse rows in the dorsum of each segment.1976 THE POLYCHAETE WORMS 103 Pherecardia Horst 1886. S. Dorsal cirri double. Body long. larval forms Strategis Kinberg 1867c. The two families are closely related. 40 species. eyes present. caruncle wedge-shaped without crest and folds. Palmyreuphrosyne is maintained where it was placed by its original describer. Euphrosinids often are considered in the amphinomids SENSV LATU. Body ovate. Hermodice striata Kinberg 1857. 1 b. notosetae in transverse rows on the dorsum. ^e Chloeia Thetisella Baird 1870. Taxonomic Note Palmyreuphrosyne appears. and it appears best to treat them separately. see Pherecardia Lenora Grube 1878. E. Pectinate branchiae in three dorsal groups per segment. hysirix Horst 1911. Ectoparasitic on sponges. P. see Hermodice Asloegia Kinberg 1867c. seeEuryihoe Metamphinome Treadwell 1940. P. see Euphrosine ORDER SPINTHERIDA Body ovate. Fauvel (1913). Neurosetae in tufts. Furcate neurosetae present. strongly muscular and with at least one pair of jaws. Maxilla I smooth and curved. 2b (la). 25x. pectinate setae. 6a (5b). 5a (4a). 3b (2b). (C).4 Two modified anterior setigers. Nothria elegans. all dorsal cirri digitate Diopatra . Most onuphids are tubicolous. see Spinther Oniscosoma Sars 1850. Paranorthia More than two modified anterior setigers 3 Modified parapodia with strongly curved. off Santa Barbara. others are sessile. grapple-hook shaped setae Rhamphobrachium Modified parapodia with composite falcigers Americonuphis Branchiae in part spiralled 5 Branchiae pectinate. FAMILY ONUPEDDAE KINBERG 1865 Eunicea with two frontal and five occipital antennae. 3a (2b). (B). 5b (4a).or bidentate hooks and capillary setae . 29. or only first parapodium longer than the following ones . About 12 species presently are recognized. 35x. oniscoides Johnston 1845. limbate setae and subacicular hooks. Eversible pharynx ventrolateral. 15x. often supported by internal acicula. with strongly developed neuropodia and reduced notopodia. (A). Spinther Johnston 1845. Family ONUPHIDAE. 1972). third carrier absent. (D). Supetfamily Eunicea Two to five pairs of lateral jaws (maxillae) and usually one pair of lower jaws (mandibles). with or without appendages.104 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Family Spintheridae Science Series 28 As the order. FIGURE Key to Genera la. 4b (lb). 20 m. simple or absent 7 Tentacular cirri absent Epidiopatra Tentacular cirri present 6 Frontal antennae short and conical. maxillae of the above. but may be able to leave their tubes in emergencies (Schäfer 1972). see Spinther ORDER EUNICIDA Prostomium distinct.. Onuphids tend to be common at all depths. others may actively hunt for debris. next to the lumbrinerids. 2a (la). the family of Eunicea best represented in deep water. Recent revisions were made by Fauchald (1968b. Setae include composite and pseudocomposite hooks and spinigers. some of them carry the tube around (Hyalinoecia). Notopodia represented by the base of the branchiae and the dorsal cirri. S. these with uni. lb. Invalid Genera Cryptonota Stimpson 1854. third parapodium of the above. with the genotype. Parapodia distinct. the family is known for a single genus. median parapodium of the above. Two or more anterior setigers with prolonged parapodia and modified setae 2 Anterior parapodia not prolonged. 4a (lb). California. AU species appear to be scavenging and feed on both plant and animal debris floating past their tube openings. Maxillary carriers short. and are. 25x. Ancirri with cirri Paradiopatra Ehlers 1887. E. 10b (9b). dorsal cirri digitate. dorsal cirri digitate. 8a (7a). Hyalinoecia Malmgren 1867. Frontal antennae short and conical. 1818 Eunicea with fi-om one to five occipital antennae. 60 species. Maxilla I . 5 species. dorsal cirri digitate. Branchiae spiralled. tentacular cirri absent. 2 species. dorsal cirri folióse in some anterior setigers Branchiae present or absent. Branchiae absent. The separation is retained here for practical reasons. see Hyalinoecia Northia Johnston 1865. tentacular cirri present. 20 species. grapple-hooklike setae. Diopatra magna drews 1891. agassizi Ehlers 1887. Anterior setigers and setae not modified. species of both genera resemble each other closely. fragosa Ehlers 1887. Anterior setigers and setae not modified. Invalid Genera Leptoecia Chamberlin 1919c. Branchiae simple and straplike or absent. Frontal antennae short and conical. tentacular cirri present. Epidiopatra Augener 1918. tentacular cirri present. only species. Anterior setigers and setae not modified. Anterior setigers and setae not modified. Onuphis Audouin and Milne Edwards 1833b. Five or more anterior setigers modified composite falcigers. Taxonomic Notes Nothria often is considered a synonym of Onuphis. Branchiae present. dorsal cirri digitate. Anterior setigers and setae not modified. Rhamphobrachium Ehlers 1887. simple and straplike in most species. 7b (4b). gracilis Ehlers 1887. all dorsal cirri digitate Generic Definitions Americonuphis Fauchald 1973. Anterior setigers and setae not modified. P. branchiae spiralled. 10 species. Frontal antennae long and slender. Branchiae pectinate. a final decision of this question will have to await the study of larger materials than is presently available. R. dorsal cirri folióse in some setigers. Paranorthia Moore 1903. P. P. Frontal antennae short and conical. D. Müller 1788. 9a (7b). O. Frontal antennae short and conical. tentacular present. Branchiae absent. Frontal antennae short and conical. dorsal cirri digitate. dorsal cirri digitate. at least some dorsal cirri folióse Tentacular cirri absent Tentacular cirri present Branchiae present Branchiae absent Branchiae in part pectinate Branchiae simple or absent Branchiae absent. Nothria Malmgren 1867. Anterior setigers and setae not modified. Anterior setigers and setae not modified. 10a (9b). Two modified anterior setigers present with uni' and bidentate hooks and capillary setae. 5 species. Frontal antennae short and conical. This is probably correct. tentacular cirri absent. Nereis tubicola O. 8b (7a). see Nothria Trapodia Baird 1870. Paronuphis Ehlers 1887. Branchiae simple and straplike or bifid. hupferiana Augener 1918. dorsal cirri digitate. Branchiae pectinate. Frontal antennae short and conical. tentacular cirri present. see Paronuphis Nereitube Blainville 1828. Two or more anterior setigers modified with strongly curved. eremita Audouin and Milne Edwards 1833b. dorsal cirri folióse in some setigers. Branchiae pectinate. dorsal digitate. 2 species. 40 species. 9b (7b). 40 species. see Onuphis FAMILY EUNICIDAE SAVIGNY Diopatra Audouin and Milne Edwards 1833b. Branchiae spiralled. Frontal antennae short and concial. 5 species. Onuphis conchylega Sars 1835.F. tentacular cirri present. Diopatra phyllocirrus Schmarda 1861. tentacular cirri absent. Frontal antennae short and conical. brevicornuta Moore 1903. tentacular cirri present. tentacular cirri present. amboinensis Audouin and Milne Edwards 1833b. Frontal antennae short and conical. third carrier absent. Maxillary carriers short. dorsal cirri digitate. THE POLYCHAETE WORMS 105 Heptaceras 8 9 Hyalinoecia Paronuphis Onuphis 10 Paradiopatra Nothria Frontal antennae long and slender. 7a (4b). Heptaceras Ehlers 1868.7976 6b (5b). One occipital antenna. Family EUNICIDAE. California. pectinate setae. pectinate setae. sometimes supported by internal acicula. lb. Setae include composite falcigers and spinigers. Notopodia represented by branchiae and dorsal cirri. 6a (lb). Tube-building is known for some species. Three occipital antennae. 2b (la). Most species are associated with hard substrates and thus with shallow water (Fauchald 1969. lOOx. limbate setae. The eunicids are among the largest of polychaetes. aculeaia Wesenberg-Lund 1949. A^. tentacular cirri and branchiae absent. pectinate setae. 4b (3b). Point Loma. (B). Setae include limbate setae. 3a (2a). Nematonereis Schmarda 1861. Eunice siciliensis Grube 1840. ninetta Audouin and Milne Edwards 1833a. composite spinigers and subacicular hooks. 4 species. Generally. branchiae present. Setae include limbate and pectinate setae. but some may be scavengers or live on large detrital particles. pectinate setae. composite falcigers and subacicular hooks. intertidal. subacicular hook of the above. 25x. Lysidice Savigny 1818. Marphysa Quatrefages 1865. Five occipital antennae. (C). 50 species. 1970). 170 species. Five occipital antennae. composite spinigers and falcigers and subacicular hooks. 10 species. 5a (2b). 2 species. L. Major revisions include Hartman (1944a) and Fauchald (1970). 3b (2a). tentacular cirri and branchiae absent.106 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 smooth and curved. Eunice antennata. lOx. Setae include limbate setae. composite spinigers and subacicular hooks. Setae include limbate setae. Euniphysa Wesenberg-Lund 1949. 5b (2b). pectinate setae and subacicular hooks. Five occipital antennae. branchiae present. Palola Gray 1847. Five occipital antennae present One to three occipital antennae present Tentacular cirri present Tentacular cirri absent Subacicular hooks absent Subacicular hooks present Composite setae falcigers Composite setae spinigers Branchiae present Branchiae absent One occipital antenna present Three occipital antennae present Generic Definitions Eunice Cuvier 1817 (NOMEN CONSERVANDUM) Nereis aphroditois Pallas 1788. 30. only species. tentacular cirri absent. (A). branchiae present. Setae include limbate setae. the eunicids are considered carnivores. E. some Eunice aphroditois have been reported as long as two meters. 4a (3b). FIGURE . Key to Genera la. tentacular cirri present. 2 6 3 5 Palola 4 Eunice Euniphysa Marphysa Paramarphysa Nematonereis Lysidice . 2a (la). median parapodium of the above. tentacular cirri present. Nereis sanguínea Montagu 1815. composite falcigers and subacicular hooks. others are burrowing into limestone or other calcium carbonate substrates. unicornia Schmarda 1861. 6b (lb). 2b (lb). indeterminable Pseudopalolo Friedländer in Woodworth 1907. The latter is a large genus. see Eunice Lithognatha Stewart 1811. tentacular cirri and branchiae present. but also in the deep scoopshape of the mandibles. see Palola Lycidice Williams 1851. indeterminable Leodice Savigny 1818. it is of the utmost importance that both paths in a key be explored. error foT Lysidice Macduffia Mclntosh 1885. An important character in most keys to species concerns the length of the posterior parapodial lobes in relation to the length of the anterior ones. buirowing forms in sand or mud or between algal hold-fasts and plantroots. tentacular cirri and branchiae absent. Maxillary carriers short. is that the posterior lobes have to be distinctly longer than the anterior ones.and peristomium. ^& Eunice Taxonomic Notes Palola frequently is considered a subgenus of. error for Aphelothrix. Notopodia absent or represented by small. third carrier absent. but care must be taken in noting the exact location of the parapodium examined in relation to the total length of the animal. but also occur in numbers in deep-water areas. set Eunice Eunice Rafinesque 1815. that no assumption is made about the structure of the posterior lobes.¡976 THE POLYCHAETE WORMS 107 Five occipital antennae. lb. Length and shape of the parapodial lobes also are valuable characters. With a fragmented specimen this may be impossible to observe. Heteromarphysa Verrill 1900. Setae include limbate setae. in such cases. Subacicular hooks and pectinate setae are absent. Most dependable appears to be the structure of the anterior setigers. FAMILY LUMBRINERIDAE MALMGREN 1867 Invalid Genera Amphelothrix Chamberlin 1919c. Members of Palola differ consistently and uniquely from the rest of the family not only in the characters listed above. Paramarphysa and Marphysa are very similar and can be separated only as indicated in the key. see Marphysa Mayeria Verrill 1900. set Marphysa Nereidice Blainviile 1828. ste Marphysa Nausicaa Kinberg 1865. the structure of the jaw•apparatus and the detailed structure of the hooks. This often is difficult since the lumbrinerids fragment readily. composite falcigers and subacicular hooks. see Nematonereis Eriphyle Kinberg 1865. see Marphysa Amphiro Kinberg 1865. see Eunice Eunicea without prostomial appendages. They are among the most common polychaetes in sandy and muddy bottoms in shelf-depths. 2a (lb). see Marphysa Amphiron Chamberlin 1919c. sometimes with internal acicula (but see Kuwaita below). appears to be of value. Paramarphysa Ehlers 1887. P. 4 species. the separation is maintained here provisionally. Five occipital antennae. or synonymous with Eunice. Marphysa and Palola Palpiglossus Wagner 1885. error for Amphiro. Nauphanta Kinberg 1865. not least since it appears related to the ecology of the contained species. It thus appears valuable to retain the distinction. Setae include limbate and pectinate setae. see Lysidice Tibiana Lamarck 1816.fide Fauvel 1918:338). Key to Gertera la. usually stated as "prolonged posterior lobes. but sometimes with one to three nuchal papillae emerging from a pocket between the pro. see Marphysa ^p/ie/oi/irix Chamberlin 1919c. Identification of lumbrinerids is dependent on very accurate work and is not altogether easy. in other words. simple and composite hooks. 3b (2b)." The key feature in this character. 3a (2b). button-shaped projections. Setae include limbate and pectinate setae and composite falcigers. Parapodia with distinct dorsal cirri Dorsal cirri absent Pharyngeal apparatus absent Pharyngeal apparatus present Branchiae absent Branchiae present Kuwaita 2 Ophiwicola 3 Lumbrineris Ninoe . NOMEN NUDUM {Eunice Cuvier 1817 is the valid name for the main genus in this imvAy. longula Ehlers 1887. sesLysidice Nereidonta Blainviile 1828. see Eunice. Maxilla I smooth and curved. and anything that can be done to subdivide it. Most lumbrinerids are free-living. see Marphysa Blainvillea Quatrefages 1865. 52x. California. Family LUMBRINERIDAE. (J). FIGURE . Lumbrineris californiensis. (B). Family ARABELLIDAE. (D). Oenone fulgida. 385x. from Loxorhynchus grandis. posterior simple hooks of the above. 30 m.W8 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 31. Arabella tricolor. 26x. (A). 26x. 53x. (F). 95x. 100th parapodium of the above. Santa Catalina Island. (E). (C). Bahia Magdalena. third parapodium of the above. (I). 30 m. anterior end of the above. Dillon Beach. (H). Baja California. fifth parapodium of the above. California. California. anterior composite hooded hook. Family LYSARETDDAE. fifteenth parapodium of the above. 385x. intertidal. Family IPHITIMIDAE. 16x. (G). 16x. Iphitime loxorhynchi. 53x. off Point Finnin. but experience has taught me that surveys based on descriptions from the literature are considerably less valuable than ones based on actual specimens. Setae include simple and composite falcigers. The change in concept of the previously described Augeneria and Ninoe indicated in the alternative key above appears unfortunate and is not recommended for future adoption. Pilger (1972) and Fauchald (1970) reviewed the family. 4a (3b). cynips Ludwig 1905. subtriangular and not expanded posteriorly Lumbrinerides Digitiform branchiae emerging from the pwstsetai lobes on a number of anterior setigers. dorsal cirri present and elongate in posterior segments. their feeding modes are unknown. maxilla IV greatly expanded with a central thin area Augeneria Paraf)odia with or without composite setae. Maxillae different Lumbrineris Generic Definitions Kuwaita Mohammad 1973. One pair of maxillary carriers fused to maxilla I. or III and IV usually with denticulated cutting edge Ninoe If branchial structures present. error for Zygolobus. 2b (la). gesae Orensanz 1974a as type species respectively. maxillae III and IV either smooth or with a few large teeth 4 Anterior parapodia with composite hooded hooks. Both genera may be valid. O. see Lumbrineris Unciniseta Bidenkap 1907. Notopodia represented by simple or branched branchiae. maxillary carriers robust. see Lumbrineris Eranno Kinberg 1865. doederleini Marenzeller 1902. dorsal cirri and branchiae absent. magna Mohammad 1973. i^t Lumbrineris Zygolobus Grube 1863. It is now time to study the variability of the characters we are using to define the many species. see Lumbrineris Scoletoma Blainville 1828. distally multidentate (anterior parapodia may have incompletely formed hooks) 3 Maxilla IV with a series of articulated spines. Invalid Genera Aotearia Benham 1927. chilensis Kinberg 1865. and five species are known. The genus Kuwaita is isolated in the family and may belong to a different. it is possible that they parasitize the crustacean host-. Lumbrineris Blainville 1828. 158 species. Iphitime Marenzeller 1902. 2a (la). branchiae present. only species. Jaw apparatus present. if present. Hooded hooks distally bidentate 2 Hooded hooks. see Lumbrineris Zygophyllus Grube 1863.J976 THE POLYCHAETE WORMS 109 Key to Genera Viàe Orensanz (1974a) la. maxillary carriers large and posteriorly expanded Lumbrineriopsis Maxilla IV without denticles or articulated spines. third carrier is absent. latreilli Audouin and Milne Edwards 1834. which is smooth and curved. dorsal cirri absent. mandibles posteriorly bifid. Lumbrinerids without nuchal papillae. only species. 28 species. mandibles posteriorly entire. mucronata (Ehlers 1908) and i. Lumbrinerids with three nuchal papillae. set Lumbrineris Lumbriconereis Grube 1840. 3b (lb). Lumbrinerids without or with one to three nuchal papillae. 3a (lb). N. Ninoe Kinberg 1865. L. lb. . Admittedly the genus Lumbrineris as presently accepted. but little damage seems to be present on the branchial tissue of the host. maxilla II with three large teeth. All Iphitimidae are inquilines in branchial chambers of crustaceans. see Lumbrineris 1970 FAMILY IPHITIMIDAE FAUCHALD Eunicea with a single pair of frontal antennae. A single genus. perhaps undescribed family. contains a large number of rather different fonns. K. Lumbrinerids without nuchal papillae. Ophiuricola Ludwig 1905. 4b (3b). see Lumbrineris (but see above) Cenogenus Chamberlin 1919c. maxillae IV. Jaw apparatus present. they are in a different position. dorsal cirri or jaw apparatus. Taxonomic Notes Orensanz (1974a) described the two genera Lumbrineriopsis and Lumbrinerides with L. so that we may use knowledge of this variability when we attempt to describe more genera in this family. Jaw apparatus present. branchiae absent. see Lumbrineris Augeneria Mon'o 1930. with genotype /. Fauchald 1970. Nereis iricolor Montagu 1804. 2a (la). Parapodia without acicular spines. Taxonomic Notes The generic sub-division is presently inconsistent in that in the group of arabellids with emergent acicular spines. &ee Iphitime Enonella Stimpson 1854. unpaired carrier present. Maxillae I proximally dentate. Maxilla I smooth or basally dentate. Maxillary carriers long and narrow. lb. O. chilensis Schmarda 1861. Drilognathus Day 1960. Parapodia without acicular spines. or may stay parasitic throughout life (Wirén 1886). Arabellids with two pairs of minute maxillae and the carriers fused for part of their length. Biborin Chamberlin 1919a. L. bonelliae Spengel 1882. 3b (2b). thick. mandibles fused into horseshoe-shaped piece. 2 species. Oligognathus Spengel 1882. 20 species. Arabellids with five pairs of maxillae and mandibles present. N. Acicular spines absent. Arabellids are never tubicolous. Arabellids with two or three pairs of maxillae present. Arabellids with the maxillary apparatus reduced to a single rod. only species. Acicular spines present. B. 2 species. notopodia absent. Acicular spines absent. Pettibone 1957c. maxilla I distally falcate. mandibles present. only species. H. parasiticus Saint-Joseph 1888. Arabellids without maxillary apparatus.no NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Invalid Genera Science Series 28 Coelobranchus Izuka 1912. Notocirrus Schmarda 1861. 5b (4b). mandibles present. Lais acutus Kinberg 1865. 5a (4b). 4a (lb). Acicular spines absent. ecbola Chamberlin 1919a. Major revisions: Hartman 1944a. Haematocleptes Wirén 1886. 2b (la). a third carrier present. Notopsilus Ehlers 1868. Acicular spines present Acicular spines absent Maxillary apparatus represented by a single rod Four or five pairs of jaws present Maxilla I distally falcate Maxilla I distally dentate Maxillary apparatus absent Maxillary apparatus present. Arabellids with five pairs of maxillae and mandibles present. Setae include limbate setae and in a number of cases. Key to Genera la. Arabellids with four or five pairs of maxillae. 6b (5b). but differ sharply from the latter in that they have long narrow maxillary carriers and have three. they may be parasitic in other animals. 6a (5b). 7b (6b). D. mandibles usually present. 4b (lb). Parapodia with acicular spines. indeterminable FAMILY ARABELLIDAE HARTMAN 1944a Eunicea without prostomial appendages. Lumbriconereis filum Claparède 1868. Acicular spines absent. Arabellids with four or five pairs of maxillae. 10 species. Arabellids with the maxillary apparatus consisting of two pairs of small plates. Labrorostratus Saint-Joseph 1888. 3a (2b). but also in echiurans. mandibles present and separate. They may be parasitic as juveniles and become free-living as adults (Pettibone 1957c). Parapodia with acicular spines. but often reduced Five pairs of maxillae present Two or three pairs of maxillae present Mandibles fused and horseshoe-shaped Mandibles two triangular plates Maxillary carriers anteriorly bilobed and fused along most of their lengths Maxillary carriers anteriorly rounded and fused along their whole length 2 4 Drilognathus 3 Drilonereis Notocirrus Biborin 5 Arabella 6 Oligognathus 7 Labrorostratus Haematocleptes Generic Definitions Arabella Grube 1850. lerebellides Wirén 1886. forms with maxilla I distally falcate are con- . 40 species. Arabellids resemble lumbrinerids closely in overall body-construction. Drilonereis Claparède 1870b. rather than two carriers. Maxillae I dentate to the tip. 7a (6b). only species. emergent spines. usually in other polychaetes. capensis Day 1960. only species. Halla parthenopeia also has been reported from cosmotropical areas and may be another species-complex. indeterminable Plioceras Quatrefages 1865. Aglaura fulgida Savigny 1818. Proximal end of maxilla I is dentate./ sidered generically distinct from forms with maxilla I distally dentate.¡976 THE POLYCHAETE WORMS . 3 species. three antennae and the distal end of maxilla I falcate. Crossland (1924) investigated some of the variabiHty of the jaw-apparatus. a third carrier is present. or to fuse the well-known genera Notocirrus and Drilonereis to ensure generic conformity within the family. but the large dorsal cirri are distinct. iridescens Knox and Green 1972. 2a (lb). brasiliensis Kinberg 1865. Lysaretids with two distinct peristomial rings. Nereis parthenopeia delle Chiaje 1828. Lysaretidae is a small family of rather large. Setae include simple and composite hooks. indeterminable Aglaura Savigny 1818. Lysarete Kinberg 1865. . Dorvilleids are mainly small polychaetes. Lysaretids with two distinct peristomial rings. see Notopsilus Laranda Kinberg 1865. bidentate hooded setae. The general appearance of the worms is that of a lumbrinerid. see Drilonereis Aracoda Schmarda 1861. Proximal end of maxilla I smooth. Lysaretids with two distinct peristomial rings. Key to Genera 1 a. considered most common in shallow water. Invalid Genera Aenone Risso ¡n Quatrefages 1865. ^ee Arabella Labidognathus Caullery 1914a. L. mandibles present. see Oenone Cirrobranchia Ehlers 1868. see Oenone Andromache Kinberg 1865. T. 4 species. &ee Notocirrus FAMILY LYSARETIDAE KINBERG 1865 Eunicea with one or three occipital antennae. Invalid Genera Arabes Ehlers 1920. Tainokia Knox and Green 1972. which has been defined in the generic definitions. 3a (2b). mainly tropical shallow-water polychaetes. three antennae and the distal end of maxilla I divided into two large fangs. it is not clear whether the best procedure would be to revise Arabella to separate the two sets of forms. maxilla I proximally smooth or dentate. see Halla Danymene Kinberg 1865. This genus has been largely disregarded. Oenone Savigny 1818. see Halla FAMILY DORVILLEIDAE CHAMBERLIN 1919C Eunicea with two pairs of antennae. but recently recovered in increasing numbers from deeper water (Jumars 1974). see Oenone Aglaurides Ehlers 1868. but did not have large enough materials to confirm or reject the presence of distinct sub-groups. . one antenna and distally falcate maxilla I. but with setae and acicula in most forms. Notopodia represented by large dorsal cirri supported by acicula. distally falcate maxilla I. Taxonomic Notes The number of species involved in the circumtropical complex referred to as Oenone fulgida has not been determined. distal end falcate Halla Proximal end of maxilla I smooth or nearly smooth. The proximal end of maxilla I is dentate. each distal end with two large fangs . three antennae and usually. One distinct peristomial segment present Oenone Two distinct peristomial segments present 2 One short antenna present Tainokia Three antennae present 3 Proximal end of maxilla I dentate. Imjwrtant revisions include Fauchald (1970) and Knox and Green (1972). see Arabella. as is the jaw-apparatus.. Proximal end of maxilla I dentate. Maxillary carriers long and narrow. 3b (2b). but not considered in the key. 2 species. indeterminable Larymna Kinberg 1865. Notopodia reduced. lb. Setae include limbate setae and in some genera. Maxillae consist of one or two series of small jaw-pieces and paired carriers. indeterminable Maclovia Grube 1871b. The corresponding separation should split the gtnus Arabella so that the forms with maxilla 1 dentate should go to the genus Nowpsilus. Lysaretids with one distinct peristomial ring. Lysarete Generic Definitions Halla Costa 1844. only species. Notocirrus and Lumbrineris Cenothrix Chamberlin 1919c. sec Drilonereis Lais Kinberg 1867. furcate and limbate setae. see Arabella Pterothrix Chamberlin 1919c. 2b (lb). 3 species. Dorvilleids with very long. Key to Genera 1 a. Dorvillea Parfitt 1866. Only simple. 2a (la). Ophryotrocha Claparède and Mecznikow 1869. Dorsal cirri small or absent. 4b (3a). Dorsal cirri absent. Dorvilleids with both antennae and palps well developed and of the same length. 3a (lb). O. 7a (6b). Either one or both pairs may be reduced. 11 species. large. Protodorvillea Pettibone 1961. Carriers and two rows of denticles present. 6a (5b). lb. carriers fused to the basal plates. Newport. California. Setae include capillaries and heterogomph falcigers. Setae include capillaries and composite heterogomph falcigers and in the first seliger. only species.112 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 One pair of the antennae is referred to as palps. 5a (3b). 50x. Setae of two kinds. Both pairs of carriers and the four basal plates of the jaws fused into a single structure. cirriform antennae and well-developed palps. furcate setae and composite heterogomph falcigers. Carriers and four rows of denticles always present. FIGURE 32. 4a (3a). cropion Jumars 1974. Setae include capillaries and heterogomph falcigers. only species. Staurocephalus rubrovittatus Grube 1855. Dorvilleids with both antennae and palps well developed and of the same length. Staurocephalus kefersteini Mclntosh 1869. they differ in structure from the other pair. Jumars (1974) has revised the generic classification of the family. Dorsal cirri absent. Dorvilleids with antennae and palps absent. Dorsal cirri with acicula. acicular setae present. 2b (la). A. capillaries and composite heterogomph falcigers. One pair of carriers and two rows of denticles present. . shallow subtidal. 7b (6b). carriers may be variously fused to the basal plates. never with acicula. Setae include capillaries. Four rows of denticles present. The jaw-apparatus may also be reduced in some forms. only species. Dorsal cirri without acicula. 3b (lb). curved composite hooks. 5b (3b). mutabiliseta Jumars 1974. Meiodorvillea Jumars 1974. Notacicula present Notacicula absent Furcate setae present Furcate setae absent Furcate or geniculate setae present Furcate or geniculate setae absent Palps well developed Second pair of antennae reduced or absent Only simple acicular setae present Both capillary and compound setae present Setae of first seliger markedly different from the others Setae of first seliger similar to others First antennae long and cirriform First antennae reduced and papilliform 2 3 Schistomeringos Dorvillea 4 5 Protodorvillea Meiodorvillea Parophryotrocha 6 Exallopus 7 Apophryotrocha Ophryotrocha Generic Definitions Apophryotrocha Jumars 1974. Protodorvillea minuta Hartman 1965. Exallopus Jumars 1974. Dorvillea articúlala. may be absent. Dorvilleids with antennae and palps reduced to papilliform projections. Dorvilleids with small clávate antennae. in that they often are articulated. 6b (5b). Parophryotrocha Hartmann-Schröder 1971. and small palps. Ophryotrocha isochaeta Eliason 1962. Dorsal cirri without acicula. puerilis Claparède and Mecznikow 1869. 5 species. Four rows of denticles present. 15 species. Family DORVlLLEmAE. carriers and basal plates fused into one piece. which may be absent. Carriers and basal plates fused into a characteristic ice-tong shape. E. FIGURE . respectively. Dorsal cirri with acicula. see Dorvillea and Schistomeringos Telonereis Verrill 1900. furcate setae and composite heterogomph falcigers. Siernaspis scutata. Invalid Genera Anisoceras Grube 1856. 10 species. antennae small or absent. a varying 33. which makes them resemble members of the Eunicida more closely than they resemble members of any other jawed family. combined from several sources. and parasitic on decapod crustaceans and fishes. (C). They are both very small. Carriers and four rows of denticles present. 95x. 23 m. the above with the anterior end everted. Woods Hole. (A). see Dorvillea Stauroceps Verrill 1900. ventral view. furcate setae and composite heterogomph setae. unrelated families of the order Eunicida. see Schistomeringos Staurocephalus Grube 1855. Histriobdella homari. Nereis rudolphii delle Chiaje 1828. Four rows of denticles present. Family STERNASPIDAE. Setae include capillaries. Dorsal cirri without acicula. Setae include capillaries. Massachusetts. from Homarus americanus. Dorvilleids with well-developed antennae and palps of approximately the same length. see Dorvillea Stauronereis Verrill 1900. One pair of anterior and one pair of posterior appendages always present. see Dorvillea and indeterminable Eleonopsis Esmark 1878. 5x. one pair of lower jaws and a single lateral jaw. about 30x. (B). Family HISTRIOBDELLIDAE. Thanks to these two surveys. anterior end inverted. they have been modified so that the only characters they have in common with other members of the order. off Santa Catalina Island.1976 THE POLYCHAETE WORMS U3 Dorvilleids with well-developed palps. Probably in response to this habit. is the structure of the jaw-apparatus. see Ophryotrocha Prionognathus Keferstein 1862. (D). carriers sometimes fused with basal plates. see Dorvillea The following two families are considered here free-standing. lOx. in terms of numbers of species. the matter seems to have been adequately clarified. Taxonomic Notes The genera of this family have been confused to a considerable extent. as noted by Pettibone (1961) and Jumars (1974). FAMILY HISTRIOBDELLIDAE VAILLANT 1890 Eunicida with five antennae. Family ICHTHYOTOMIDAE. Schistomeringos Jumars 1974. Ichthyotomus sanguinarias. carriers may be fused with basal plates. Stemaspis Schreiberius Otto 1821. with type species. the illustration given here has been redrawn from that illustration. chitinized. Histriobdellids are known as parasites in the branchial chambers of crustaceans. Setae absent. but do not appear to be common in abyssal depths. They have turned out to be quite frequently reported from moderate depths on the continental slopes. lb. Histriobdella FAMILY IcHTHYOTOMtDAE EISIG 1906 Eunicida with one antenna. but are perhaps most usually found in about 100-200 m depth. Parasitic on replant decapod crustaceans. combined with their habitat makes it likely that they are considerably more common than the few scattered records indicate. Renier 1804. prostomium without appendages. lb. proboscis a muscular pad. The shape of the rather characteristic small hooks has been well demonstrated by Thomassin and Picard (1972) with help of scanning electron microscope. Key to Genera la. 3a (lb). Notopodia with acicula. Stratiodrilus Has well 1900. FAMILY OWENIIDAE RIOJA The genera are defined as indicated in the key. The family is known for one genus. those associated with the shield. sanguinarius Eisig 1906 from the Gulf of Naples. and they are often capable of moving around with the tube. tubicolous. Invalid Genera Eckinorhynchus Jfivst. They are common in sandy and muddy substrates in all depths. tubicolous animals. The sternaspids are among the most easily recognized polychaetes with the usually dark yellow or reddish chitinized shield. Histriobdella van Beneden 1858. It is parasitic on the fins of eels. see Stemaspis Thalassema Ranzani 1817. can be inverted with the first three setigers. Sternaspids are only rarely found in large numbers. ventrally enth-e deeply bilobed with paired palps 2 3 Galathowenia Owenia Myriowenia . They are very small. 2a (la). both from fresh-water and marine environments. prostomium sometimes produced in lobes or as a folded membrane. H. see Stemaspis ORDER OWENIIDA Prostomium fused to the anterior segments. see. Invalid Genus Histriodrilus Foettinger 1884. Eversible pharynx axial. falcate spines. tasmanicus Has well 1900. Parasitic on fishes. The oweniids are characteristically rather small. ventrally deeply incised forming a tentacular crown. homari van Beneden 1858. 4 species. the tubes are usually short. those in the first three setigers thick. They are burrowers in the sand and mud. only species. /. and that. slender capillanes. With a single pair of lateral appendages With at least two pairs of lateral appendages Histriobdella Stratiodritus Renier 1807. mineral-impregnated shield. neurosetae very small bi. Key to Genera la. Fauvel (1958) gives a good description and illustration of the form. ORDER STERNASPIDA Posterior ventrum covered by a stiff. 5. The total number of currently recognized species is about ten.or tridentate hooks in dense fields. Eckinorhynchus scutatus Body cylindrical with long anterior segments and short posterior ones. Prostomium Prostomium Prostomium Prostomium Prostomium anteriorly produced into a low collar or tentacular crown rounded or bilobed forming a low collar. one pair of lateral jaws. Notosetae capillary.114 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 number of lateral appendages also present. neurosetae composite. Ichthyotomus is known for a single species. 2b (la). FAMILY STERNASPIDAE CARUS 1917 1863 Short-bodied polychaetes with indistinct segmentation. but otherwise asetigerous. Neuropodial hooks in dense fields. see. Each genus is known in just a few species. Stemaspis Otto 1821. All setae simple. i976 3b (lb). Fauveliopsis brevis. sec Myriochele ORDER FLABELLIGERIDA Anterior pharynx with ventrolateral muscular pad or unreinforced. larval forms Ops Carrington 1865. set Owenia Psammocollus Grube 1868. Only one anterior setiger with notosetae only. 12x. M.5x. Prostomium produced anteriorly into a low tentacular crown with flattened lobate projections. about 5x. Pro. Anterior end retractable within a sheath formed by the first setigers. Notosetae cross-barred or smooth. 1971. 4b (3b). Three first setigers without neurosetae. M. First two or three setigers with notosetae only. redrawn and simplified after Robbins. O. after Hartman and Fauchald. Invalid Genera Ammochares Grube 1847. Prostomium anteriorly rounded. Owenia delle Chiaje 1841. Foeobius meseres. First three setigers without neurosetae. Myriowenia Hartman 1960. Family POEOBIIDAE. 2 species. most forms with epidermis covered by papillae. First three setigers without neurosetae. Eyes present. Family FLABELLIGERIDAE. 11 species. Prostomium a slender ridge with palps at the posterolateral sides. Family OWENIIDAE. (D). FIGURE Body cylindrical or fusiform. after Hartman. Myriochele Malmgren 1867. Family FAUVELIOPSIDAE. see Owenia Mitraría Müller 1851. heeri Malmgren 1867. 1965. and slender. palps absent First two or three setigers with notosetae only Only first setiger with notosetae only Generic Definitions Galathowenia Kirkegaard 1959. only species. (A). Pherusa inflata. Prostomium deeply bilobed with a pair of grooved palps attached anteriorly. 4a (3b). Prostomium anteriorly rounded. Peristomium with an expanded dorsal membrane c^irrying branchiae. Owenia collaris. FAMILY FLABELLIGERIDAE SAINT-JOSEPH 1894 34. 12 species. neurosetae either similar or more thickened . Prostomium anteriorly produced into a low collar that ventrally is deeply incised with one lobe overlapping the other. only species. G. 5x. Myrioglobula Hartman 1967. 1969. antárctica Hartman 1967. californiensis Hartman 1960.and peristomia retractable between the first three setigers. combined from several sources. 12. (C). THE POLYCHAETE WORMS 115 4 Myriochele Myrioglobula Prostomium rounded. (B). M. africana Kirkegaard 1959. fusiformis delle Chiaje 1841. 4a (2b). Body short and fusiform. C. distally acicular. which secretes the mucus. All setae capillaries. Flabeiligera commensalis Moore 1909. B. Dissection of the retractable anterior end is necessary for safe identification of the several similar genera. 5b (4b). 10a (9a). Key to Genera la.. has its separate cover of particles. rounded or triangular 7 Branchial membrane club-shaped. 7b (5b). 12b (lib). Body short and fusiform. papillota Hartman 1967. Body anteriorly inflated. The number and structure of the branchiae and the structure of the branchial membrane are important identificatory characters. with branchial filaments attached distally on all sides . 8a (7a). 3a (2a). but in most each individual papilla. only species. 9b (7b). 7a (5b). cephalic cage present. 10b (9a). Oral tube present. Neurosetae slightly thicker than notosetae. 9a (7b). with the branchial filaments attached on all sides distally. 8b (7a). Body anteriorly inflated. 7 species. Neurosetae simple hooks. Bradabyssa Hartman 1967. branchial membrane short and folded. but usually distinctly cross-barred 4 Body encased in a smooth continuous mucus sheath Flabeiligera Body with individual papillae covered with mucus and impregnated with debris Flabelliderma Branchiae absent Bradabyssa Branchiae present 5 Branchial membrane long. 5a (4b). I2a (11 b). in some forms this mucus forms a complete smooth encasing for the animal {Flabeiligera). 4b (2b). Branchial membrane triangular with numerous branchial filaments. Branchiae absent. Body anteriorly inflated. . numerous pairs of branchiae Brada Body covered by a thick mucus sheath Buskiella Body not covered by a mucus sheath. Flabelliderma Hartman 1969. mucus sheath covering the whole body. 2a (lb). 13a (12b). Four pairs of branchiae of two kinds on a short branchial membrane. 21 species. lib (9b). 1 la (9b). longisetosa Haswell 1892. Neurosetae acicular. flattened and nearly disc-shaped llyphagus Cephalic cage poorly developed or absent 10 Cephalic cage well developed 11 Neurosetae distinctly thicker than notosetae.116 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 spines or composite with falcate. Cephalic cage present. 6b (5a). abyssorum Mclntosh 1885. lb. only species. only species. Neurosetae simple hooks. Coppingeria Haswell 1892. tonguelike. Trophonia glauca Malmgren 1867. cephalic cage present in some forms. B. 13b (I2b). Buskiella Mclntosh 1885. Branchial membrane long. Coppingeria Branchial membrane flattened. sometimes doubled 6 Branchial membrane short. often sand-incrusted 12 A long oral tube present Therochaetella Oral tube absent 13 Notosetae serrated and plumose Pantoithrix Notosetae cross-barred capillaries Pherusa Generic Definitions Brada Stimpson 1854. with branchial filaments attached on one side only Piromis All setae capillary 8 At least some neurosetae acicular or falcigerous 9 Body anteriorly inflated with tapering posterior end Diplocirrus Body. short. The flabelligerids characteristically are heavily impregnated with sand or mud in a matrix of mucus. Diplocirrus Haase 1915. cephalic cage absent. B. cephalic cage absent. 6a (5a). Body with a distinct incision just posterior to the setigers carrying the cage-forming setae Therochaeta Body without distinct incisions 2 Neurosetae composite or pseudocomposite 3 Neurosetae entirely simple. Numerous branchial filaments on a short branchial membrane. 2b (lb). parapodia very prominent. four pairs of branchiae Trophoniella Neurosetae only slightly thicker than notosetae. granosa Stimpson 1854. cylindrical and slender. 3b (2a). 2 species. unidentate or bidentate appendages. T. see Flabelligera Trophonia Cuvier 1830. Neurosetae bi. All . Pherusa chilensis Schmarda 1861. P. neurosetae bidentate hooks. ORDER FAUVELIOPSIDA Pro. cephalic cage present. Further information may demonstrate that they are related to other groups of polychaetes.F. only species. FAMILY FAUVELIOPSIDAE HARTMAN 1971 Small. retractile branchiae and palps present. body papillae covered with thick mucus and encrusted with debris.e Flabelligera Siphostoma Otto 1821. Müller 1776. T. se. sec Pherusa Lophocephalus Costa 1841. indeterminable FAMILY POEOBIIDAE HEATH 1930 Body saclike without external segmentation or setae. Body anteriorly inflated. Pelagic. deep-water polychaetes are incompletely known. Stylarioides coUarifer Ehlers 1887. 10 species. it has been reported only once. see Pherusa Pycnoderma Grube 1877. smooth-bodied polychaetes without anterior appendages. see Pherusa Tecturella Stimpson 1854. flattened and tongueshaped. Two distinct septa only polychaete characters. 43 species. Trophoniella Caullery 1944. terwillei Betrem 1924. cephalic cage present. Anterior end inflated. Proboscis a ventral muscular pad. and because they have chlorocruorin as one of their pigments (Robbins 1965). Amphitrite plumosa O. usually anteriorly directed papillae. Flabelligera Sars 1829. see Pherusa Zorus Webster and Benedict 1887. Aristenia Savigny in Quatrefages 1865. Neurosetae mostly uni. /. All setigers biramous with simple limbate setae and a small rounded papilla between the rami. only species. Anterior end inflated. Pantoithrix Chamberlin 1919c. Poeobius meseres Heath 1930. chilensis Hartman 1967. only species.or bidentate hooks. Incertae Sedis Enigma Betrem 1924. This form resembles the poeobiids in several respects. either single or double.and peristomium without appendages. Body anteriorly inflated. Anterior end fused to the rest of the body. The shape of the prostomium also is markedly different. Six pairs of branchiae. F. Most neurosetae uni. see Diplocirrus Semiodera Chamberlin 1919c. Cephalic cage present. Body anteriorly inflated. Four pairs of branchiae on a short branchial membrane. avicularia Cauilery 1944. Ilyphagus Chamberiin 1919c. The poeobiids are considered related to the flabelligerids because the anterior end is retractable. 18 species. Therochaeta Chamberlin 1919c. Numerous pairs of branchiae on a short branchial membrane. 6 species. see Pherusa Chloraema Dujardin 1839a. Composite hooks present on some anterior setigers in most species. with numerous branchial filaments. bythincola Chamberlin 1919c. indeterminable Balanochaeta Chamberlin 1919c. flattened and nearly disc-shaped. cephalic cage very poorly developed or absent. arenosus Kinberg 1867b. pelagic. The family contains a single genus and species.or unidentate hooks. Notosetae serrated and plumose capillaries. E. ses Piromis Saphobranchia Chamberlin 1919c. Four pairs of branchiae on a short branchial membrane. covered with large papillae and mud. Long narrow oral tube present. see Piromis Siphonostoma Rathke 1843. Therochaetella Hartman 1967. First post-incisional setiger with series of enlarged. cephalic cage present. but unfortunately the original material has been lost (Hartman 1967). Branchial membrane prolonged. cephalic cage present. Anterior end inflated. Piromis Kinberg 1867b. cephalic cage present. These small. Body stout. a separate order appears justified for the time being. affinis Sars 1829. Pharynx with a ventral muscular pad. Pherusa Oken 1807. Cephalic cage present. Neurosetae distally bifid and falcate. 3 species. see Flabelligera Stylarioides delle Chiaje 1841. Body cylindrical. one septum observed. the retractable anterior end and the characteristic setae. Parapodia and setae absent. All setae capillaries. covered completely by smooth mucus sheath. 11 species. Either four or many pairs of branchiae on a short branchial membrane. A distinct incision present behind the last setiger that carries the cage setae. pseudocomposite or composite hooks in the neuropodia. otherwise simple neuropodial hooks.1976 THE POLYCHAETE WORMS m Invalid Genera Body cylindrical. containing a circlet of eversible tentacles. see Flabelligera Flemingia Johnston 1846. They were grouped formerly with the flabelligerids from which they differ in that they lack the papillar investments. Pseudocomposite hooks present in most neuropodia. Cephalic cage present in some forms.or bidentate hooks.or sand-incrusted in most forms. setae absent. L. Flabeliigella was shown to belong to the Acrocirridae by Orensanz (1974b) and are cited under that family above. as well as for Flabeliigella Hartman 1965. ORDER TEREBELLDDA Prostomium without appendages. Setae of first setiger forming an operculum with setae in one to three rows. 4b (3a). Peristomium with five tentacular cirri. with eight described species. 3b (lb). consequently. 4a (3a). Parapodial lobes reduced. 2a (la). Incertae Sedis Bruunilla Hartman 1971. operculimi flattened cone Phragmatopoma Middle opercular paleae not concealing the inner ones. 6b (3b). armatus Kinberg 1867b. /. 5a (4b). The sabellariids are important as reef-builders in certain areas (Schäfer 1972). Flota Hartman 1967. At least one pair of branchiae present (rarely absent). natalensis Hartman 1971. accessory setae absent. Paired short palps lateral to the mouth as flat pads. slender smooth acicular rods and thicker and cross-barred acicular setae. opercular peduncles fused. only species. hooks present dorsal and proximal to the paleae. opercular peduncles long and separated. I b. Body with less than ten segments. with embedded acicula. most of them are present in shallow water only. indicus Kinberg 1867b. only species. Prominent lateral parapodia present. Thorax with two rudimentary segments and three or four parathoracic setigers. FAMILY SABELLARIIDAE JOHNSTON 1865 Tubicolous polychaetes with the body in three regions. short and flattened. 6a (3b). Three parathoracic setigers. Pharynx muscular and eversibje. B. 2b (la). 5b (4b). Dorsal and ventral cirri present on most segments. peristomium with a series of feeding appendages. Body with papillae. The family was erected by Hartman (1971) for Fauveliopsis Mclntosh. With a single row of paleae in the operculum 2 With at least two rows of paleae in the operculum 3 The prolonged opercular peduncles free from one another Phalacrostemma Opercular peduncles short and fused Monorchos Two rows of paleae 4 Three rows of paleae 6 Hooks absent Gunnarea A pair of large hooks dorsal and proximal to the opercular paleae 5 Three parathoracic setigers Idanthyrsus Four parathoracic setigers Lygdamis Middle opercular paleae cover the inner ones. 3a (lb). operculum open and bristly Sabellaria Generic Definitions Gunnarea Johansson 1927. posterior region an asetigerous anal tube. Two rows of paleae. but members of two genera (Phalacrostemma and Monorchos) appear to be most common in slope depths. Four parathoracic setigers. F. Two rows of paleae.¡18 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 setigers biramous with simple limbate setae and a small rounded papilla between the rami. Hermella capensis Schmarda 1861. both rami distally strongly pointed. all biramous. Flota Hartman 1967 and Bruunilla Hartman 1971. opercular peduncles fused. 12 species. All of them are tubebuilders and tend to build on firm substrates. challengeriae Mclntosh 1922. Idanthyrsus Kinberg 1867b. hooks present dorsal and proximal to the paleae. Prostomium small and triangular attached ventral to the large peristomium. hooks and accessory setae absent on operculum. Flota and Bruunilla differ sharply from Fauveliopsis and are characterized here as free-standing genera without obvious familial affiliations. Two rows of paleae. especially on the parapodia. The whole family is then reduced to the type-genus. Three parathoracic setigers. Fauveliopsis. Encased in a thick mucus sheath. Prostomium a simple conical lobe with a pair of tri- lobed processes. Key to Genera la. 5 species. ftostomium a narrow ridge fused laterally to the first setiger. pharynx with a ventral muscular pad. only species. Median region with notopodial pectinate uncini and ventral capillary setae. pelagic. Parapodia biramous with setae of two kinds. Lygdamis Kinberg 1867 b. with type F. flabelligera Hartman 1967. . scoop-shaped and open dorsally. accessory setae absent. Sabellaria Savigny 1818. hooks and accessory setae absent. natural size. Amphictene capensis. P. (B). Four parathoracic setigers. opercular peduncles fused. Middle opercular paleae covers the inner ones. 2x. 28 species. (A). caudate Morch 1863. (D). Sabella alveolata Linnaeus 1767. accessory setae absent. hooks present dorsal and proximal to the paleae and two rows of accessory setae between the paleal rows. 5 species. A single row of paleae.1976 THE POLYCHAETE WORMS 119 c^\ ?n OA J FIGURE 35. tube pattern of Phragmatopoma sp. M. after Day. 1969. Three rows of paleae. pectinariid-tube. 2 species. 8 species. A single row of paleae. after Hartman. . Phragmatopoma califomica. Phalacrostemma Marenzeller 1895. 3x. cidariophilwn Marenzeller 1895. Monorchos Tread well 1926. Three parathoracic setigers. opercular peduncles short and fused. 5x. Phragmatopoma Mörch 1863. (C). opercular peduncles long and separated. hooks present dorsal and proximal to the paleae. Family SABELLARIIDAE. 1967. philippinensis Treadwell 1926. P. Three parathoracic setigers. Family PECTINARllDAE. operculum conical. strong setae (paleae) forming a comb used in digging. scaphe distinctly set off from abdomen Twelve uncinigers. pusilla Malmgren 1866. The family often is recognized for only two genera. 13 uncinigers present. 14 uncinigers. Petta and Pectinaria. see Pectinaria . cephalic veil at least partly fused to the operculum Thirteen uncinigers. Scaphe distinctly set off from abdomen. Cephalic veil free from operculum. Pectinaria Savigny 1818. The generic sub-division appears reasonably stable. Invalid Genera Centrocorone Grube 1850. they usually are acicular in appearance and may be present between the rows of paleae (Monorchos) or proximal to the paleae proper (Sabellaria). accessory setae sometimes present. Scaphe distinctly set off from the abdomen. 4a (3b). 8 species. 4b (3b). include setae associated with the operculum. pectiniform uncini and scaphal spines. 13 uncinigers present. The family is under current revision by Dr. marginally smooth. see ¡daitíhyrsus Tetreres Caullery 1913. Tubes short. see Sabellaria Cryptopomatus Gravier 1908. tusk-shaped. Uncini with major teeth in double rows. Petta Malmgren 1866. marginally cirrate. Müller 1776. P. Scaphe distinctly set off from abdomen. Uncini with major teeth in double rows. tapering. more or less tusk-shaped. opercular peduncles short and fused. 8 species. Uncini with major teeth in two or more rows. Prostomium reduced. 4 species. Cephalic veil free from operculum. 3a (2b). see Lygdamis Hermella Savigny 1818. lb. L. Nereis cylindraria bélgica Pallas 1766. hooks absent. scaphe not distinctly separated fiwm abdomen Cephalic veil marginally cirrate. Cephalic veil marginally smooth. abdomen and a posterior scaphe. 2b (lb). Cephalic veil free from operculum. see Sabellaria Pallasia Quatrefages 1848.P.¡20 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Three rows of paleae. 12 uncinigers present. coppery or brassy paleal setae and the gently curved. Scaphe distinctly set off from abdomen. Cistenides Malmgren 1866. Lagis Malmgren 1866. 13 uncinigers. Uncini with major teeth in a single row. Amphitrite auricoma O. in which case the other listed genera are considered subgenera of Pectinaria. see Idantkyrsus Pallasina Annenkova 1925. see Sabellaria Chrysodon Okenin Quatrefages 1865. indeterminable Cistena Leach 1816. koreni Malmgren 1866. opercular rim marginally cirrate. opercular rim marginally smooth. 3b (2b). marginally cirrate. built of relatively large particles. marginally cirrate. Key to Genera la. David Kirtley and the numbers of species assigned to each genus may be changed drastically. Sabella granúlala Linnaeus 1767. Middle opercular paleae pointed distad. operculum open and generally rather bristly in appearance. Other setae include short capillaries. marginally cirrate. opercular rim marginally smooth. Cephalic veil at least partly fused to operculum. but usually very fragile tubes. Three parathoracic setigers. Scaphe not distinctly set off firom abdomen. Opercular rim marginally cirrate. First seliger with expanded. 8 species. Cephalic veil free from operculum. The pectinariids are among the most characteristic polychaetes with their strongly golden. including thorax. 18 species. cephalic veil completely free from the operculum Opercular rim cirrate Opercular rim smooth Uncini with major teeth in a single row Uncini with major teeth in two rows Petta 2 Lagis 3 Amphictene 4 Cistenides Pectinaria Generic Definitions Amphictene Savigny 1818. Invalid Genera Ariapithes Kinberg 1867b. Opercular rim marginally smooth. 2a (lb). see Idanthyrsus Eupallasia Augener 1927a. Taxonomic Notes The "accessory setae" referred to in the above definitions. Uncini with major teeth in a single row. see Lygdamis FAMILY PECTINARIIDAE QUATREFAGES 1865 Body separated into three regions. 13a (12b). larval forms Scalis Grube 1846. 4b (3a). 3b (2a). 4a (3a). The major classification was reviewed by Day (1964) who reduced greatly the number of monotypic genera. anterior region with biramous parapodia. 14 thoracic uncinigers Melinna Four pairs of branchiae. Thoracic uncini with major teeth in one or a few rows. 10a (lb). it is not clear that the characters Day used to identify his genera are any more precise than those he disregarded. 6b (2b). 12b (10b). Two to four pairs of smooth or pinnate branchiae present. indeterminable THE POLYCHAETE WORMS 111 FAMILY AMPHARETIDAE MALMGREN 1867 Body with two regions. The ampharetids have turned out to be very common in deep water and a whole mass of previously unrecognized genera have been reported from deep water over the last ten years. paleae present Ecamphicteis Maximally 12 thoracic uncinigers. The uncini often are flattened plates in the ampharetids and nearly always distinctly crested in the terebellids. 3a (2a). all buccal tentacles similar Melinantipoda Maximally 14 thoracic uncinigers present. MELINNINAE 2 First severed neuropodia without acicular setae. It has been included in the definitions. 6a (2b). until the current deep-water material has been worked up in detail. abdominal uncini similar. lib (lOa). lOb (lb). one or two buccal tentacles very large 9 Segment 6 with a distinct dorsal crest between the notopodia Melinnexis A dorsal crest absent Amelinna At least three pairs of branchiae 11 Two pairs of branchiae 12 Three pairs of branchiae present 15 Four pairs of branchiae 35 Fourteen thoracic uncinigers. posterior region with well-developed neuropodia. The ampharetids resemble the terebellids. lb. 12a (lOb). 1 la (lOa). 7b (6a). the latter do not. Addition- ally. capillary notosetae on setigers 3-4 Isolda Three pairs of branchiae. never the masses of arborescent branchiae or the numerous sessile filaments present in the terebellids. it appears best to await further fusions of genera. first uncini in a short row similar to all others Egamella Twelve uncinigers. 12 thoracic uncinigers Moyanas A single pair of nuchal hooks. 13b (12b). one smooth. 9a (8b). 5b (3b). 9b (8b). all neurosetae uncini AMPHARETINAE 10 Nuchal hooks present 3 Nuchal hooks absent 6 Branchiae smooth 4 Branchiae in part papillose or pennate 5 Two pairs of nuchal hooks. other neuropodia with uncini . 2a ( I a). two smooth and two papillose. notosetae absent on setigers 3-4 Irana Ten thoracic uncinigers present 7 At least 12 thoracic uncinigers present 8 Three pairs of branchiae Melinnopsides Four pairs of branchiae Melinnopsis Sixteen thoracic uncinigers present.i976 Labiaria Sveshnikov 1959. The present review retains most of the genera Day fused. but more frequently crested. Prostomium simple or complex with lateral folds and glandular ridges. two papillose. the main feature used to separate the two families is behavioral in that the former will withdraw the buccal tentacles completely within their mouth. 14a (13b). 8b (6b). 2b (la). Since a large number of additional taxa are now being described. neuropodial uncini present both in thorax and abdomen. 5a (3b). The genus Oeorpata is not clearly separable from Isolda according to Day (1964) and is incompletely known. 7a (6a). Notopodia! capillary setae present in thorax. paleae absent 13 Nine uncinigers. 8a (6b). rarely crested. the ampharetids usually have a few pairs of simple branchiae. notopodia reduced or absent. but not in the key. Key to Genera la. first uncini in a long row 14 Prostomium anterioriy pointed Auchenoplax First several neuropodia with fine acicular setae. Nuchal hooks and anterior acicular setae present in some forms. modified after Fauchald. Boguella ornata. 1972. Family AMPHARETIDAE. 12. modified from live sketch.5x. about 23x. 3x. Terebellides straemi. (A). combined from several sources. Neoamphitrile. Family TRICHOBRANCHIDAE. (B). redrawn after Hartman and Fauchald. Family TEREBELLIDAE. (C). FIGURE . about 5x.122 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 36. oeai johrtstoni. Amphicteis scaphobranchiata. Family BOGUEIDAE. 1971. (D). 38a (37a). 30a (29b). 16a (15a). if present. setae present on segment 3. 41b (40a). 30b (29b). 36b (35a). 17b (16b). THE POLYCHAETE WORMS 123 Prostomium anteriorly truncate Melinnoides All branchial pairs arranged in a distinctly segmental fashion 16 Only last pair of branchiae clearly associated with a specific segment 18 Eleven thoracic uncinigers. 22a (21b). 29b (28b). 15b (1 la). 24a (23b). if present. anterior in position. 38b (37a). tentacular membrane small 17 First setae on segment 2. as the first notosetae in the body) 36 Paleae absent (i. 21a (20b). 33a (31b). 34b (33b). 11 thoracic uncinigers Amythasides Buccal tentacles. 19b ( 18a).1976 14b (13b). 39b (37b). 32b (31a). Neosamytha Dorsal crest on either segment 6 or 7 Melirmampharete Ten thoracic uncinigers Muggoides At least 11 thoracic uncinigers 26 Eleven thoracic uncinigers Glyphanostomum At least 12 thoracic uncinigers 27 Twelve thoracic uncinigers 28 At least 13 thoracic uncinigers 31 At least one pair of branchiae basally flanged Samythella All branchiae cylindrical 29 Buccal tentacles papillose Neosabellides Buccal tentacles smooth 30 Oral membrane greatly prolonged Pabits Oral membrane short Eusamytha Thirteen thoracic uncinigers 32 Fourteen thoracic uncinigers 33 Notopodia of setiger 13 elevated with hirsute notosetae Sosanella Notopodia of setiger 13 at the same level as all others.e. ten thoracic uncinigers Melinnaia Flange or crest. 15a (11a). 26b (25b). 28a (27a). 34a (33b). 31a (27b). 41a (40a). 24b (23b). notosetae limbate or capillary Ampharete . 26a (25b). 22b (21b). 21b (20b). at least 11 thoracic uncinigers 21 Few stout buccal tentacles present. 35b (lib). 16b (15a). 25b (18b). 27b (26b). 40a (39a). 20a (19b). 40b (39a). 18b (15b). setae absent on segment 3 or setae already present from segment 2) 52 Flanged branchiae present on four successive segments Phyllampharete Branchiae flanged. hirsute setae absent Alkmaria Oral membrane smooth and folded Amythas Oral membrane tentaculate 34 Glandular ridges on prostomium Samythopsis Glandular ridges absent Samytha Paleae present (i. 19a (18a). 15 thoracic setigers Eusamythella Oral membrane short. buccal tentacles papillose Pterampharete Both branchiae and buccal tentacles smooth Sabellides Twelve thoracic uncinigers present 40 At least 13 thoracic uncinigers present 44 Buccal tentacles papillose 41 Buccal tentacles smooth 42 Setiger 11 with notopodia elevated and notosetae distally hirsute Anobothrella Setiger 11 with notopodia at the same level as all other setigers. 23a (22a). 39a (37b). 36a (35a). 32a (31a). 17a (16b). only one pair clearly associated with a segment 37 Eleven thoracic uncinigers 38 At least 12 thoracic uncinigers 39 Branchiae pennate.e. tentacular membrane large Ampharana At least 14 thoracic uncinigers. 27a (26b). 28b (27a). 31b (27b). 35a (1 lb). numerous. 37b (36b). lamellate. 18a (15b). 20b (19b). 14 thoracic uncinigers Neopaiwa First setae in segment 3. pennate or cylindrical. 33b (31b). 12 thoracic uncinigers 22 Dorsal crest present on one anterior setiger 23 Dorsal crest absent Eclysippe Oral membrane broadly folded. 29a (28b). 25a (18b). last thoracic notopodia elevated Mugga Ten or more thoracic uncinigers 20 Flange present between last thoracal notopodia. 37a (36b). 23b (22a). 15 thoracic uncinigers Weddellia Paleae present 19 Paleae absent 25 Nine thoracic uncinigers. 14 thoracic setigers 24 Dorsal crest on segment 4 . 42b (40b). 53b (52b). 46b (45b). 52b (35b). 43a (42b). 53a (52b). 11 thoracic uncinigers. 27 species.124 Ala. 2 species. Four pairs of smooth branchiae. (55a). (58a). Eleven thoracic uncinigers. Glandular ridges absent. 52a (35b). 50b (49a). A. (58b). one or two large oral tentacles present in addition to numerous smaller ones. Twelve or 13 thoracic uncinigers. Amphicteis acutifrons Grube 1860. no paleae. Three pairs of smooth branchiae on three successive segments. anus surrounded by a circle of papillae Phyllocomus Three pairs of lamellate and one pair of cylindrical branchiae. AMPHARETE>ÍAE. rominji Horst 1919b. Ampharete Malmgren 1866. auricula Malmgren 1866. Two pairs of anal cirri. others smooth Pterolysippe All branchiae smooth 46 Glandular ridge on prostomium Hypania Glandular ridge absent Lysippe Two of the four pairs of branchiae lamellate 48 All four pairs of branchiae cylindrical 49 Paleae large. Glandular ridges present. Four pairs of smooth branchiae. A. 54a (53a). Three pairs of smooth branchiae. no notopodial rudiments in abdomen. anus with two pairs of anal cirri . Paramage All anterior notopodia with setae 57 One pair of anal cirri Grubianella Two pairs of anal cirri Amage Twelve thoracic uncinigers 59 Fourteen thoracic uncinigers 60 Buccal tentacles papillose. (40b). . 50a (49a). 54b (53a). Paleae absent. AMPHARETENAE. 45a (44a). (55b).. abdominal notopodial rudiments absent Pkyllamphicieis Paleae small. AMPHARETINAE. Amelinna Hartman 1969.. Schistocomus Eleven thoracic uncinigers 56 At least 12 thoracic uncinigers 58 First two notopodia (segments 4 and 5) asetigerous . 44b (39b). (55b). 47b (44b). notosetae absent in segments 3 and 4. A. notopodial cirri present Paramphicteis Buccal tentacles smooth. 14 species. (58a). 46a (45b). antárctica Hartman 1967. Ampharana Hartman 1967. all notosetae capillary Asabellides Buccal tentacles smooth. 43b (42b). notopodial cirri absent Amphisamytha Generic Definitions MELINNINAE. (53b). 45 Fourteen thoracic uncinigers 47 One pair of branchiae pennate. 49b (47b). Alkmaria Horst 1919b. 51a (49b). 48b (47a). Amage Malmgren 1866. only species.. (56b). NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Seliger 14 with elevated notopodia and hirsute notosetae Sosane Setiger 10 or 11 with elevated notopodia and tapering or hirsute notosetae 43 Notopodial rudiments present in abdomen Anobothrus Notopodia! rudiments absent from abdomen Sosanides Thirteen thoracic uncinigers . Oral membrane large and folded with buccal tentacles in two lateral groups. 48a (47a). but small capillary setae present in segment 3. (55a). 13 thoracic uncinigers. A. (58b). 47a (44b). 45b (44a). 44a (39b). no paleae. abdominal notopodial rudiments present. last thoracic notosetae modified Sosanopsis Buccal tentacles papillose. abyssalis Hartman 1969. 49a (47b). 55a 55b 56a 56b 57a 57b 58a 58b 59a 59b 60a 60b (53b). abdominal notopodial rudiments present Lysippides Glandular ridges on prostomium 50 Glandular ridges absent 51 Abdominal notopodial rudiments present Amphicieis Abdominal notopodial rudiments absent Parhypania Abdominal notopodial rudiments present Paiwa Abdominal notopodial rudiments absent Hypaniola Branchiae arranged in oblique series directly associated with distinct segments Mexamage Only the last pair of branchiae clearly associated with a segment 53 At least three pairs of branchiae lamellate 54 All branchiae cylindrical 55 All four pairs of branchiae lamellate. (56b). only species. 51b (49b). Nuchal hooks absent. Eusamytha Mclntosh 1885. E. AMPKARETINAE. A. Glandular ridges absent. Ampharete gracias Malmgren 1866. branchial membrane high and laterally free from branchial bases. Four pairs of smooth branchiae. AMPHARETINAE. AMPHARETINAE. Auchenoplax Ehlers 1887. A. paleae present. Eleven thoracic uncinigers. 2 species. Four pairs of smooth branchiae. glandular ridges absent. Four pairs of smooth branchiae. Four pairs of smooth branchiae. Four pairs of papillose branchiae. glandular ridges absent. 12 thoracic uncinigers. Amythasides Eliason 1955. E. Fourteen thoracic uncinigers. only species. Twelve thoracic uncinigers. Three pairs of smooth or grooved branchiae. 30 species. Anobothrus Levinsen 1884. first row of uncini very long and ventrally located. Twelve thoracic uncinigers. Twelve uncinigers. Four pairs of smooth branchiae. Glandular ridges absent. Glandular ridges absent. nine uncinigers present. 14 thoracic uncinigers. oral membrane folded and smooth. AMPHARETINAE. abdominal notopodial rudiments absent. Four pairs of smooth branchiae. Abdominal notopodial rudiments present. rarely folióse. Twelve thoracic uncinigers. Ecamphicteis Fauchald 1972. paleae absent. paleae absent. Buccal tentacles papillose. paleae absent. Anobothrella Hartman 1967. dorsal crest across segment 5. Samytha pallescens Théel 1879. Egamella Fauchald 1972. pacifica Mclntosh 1885. Grubianella Mclntosh 1885. only species. only species. paleae present. . only species. crinita Ehlers 1887. paleae present. Fourteen thoracic uncinigers. Notosetae ab- sent in segments 3 and 4. Amphicteis invalida Grube I860. paleae absent. Amphicteis Grube 1850. Glandular ridges absent. Two pairs of smooth branchiae. Eclysippe Eliason 1955. notosetae absent in segment 4. Three pairs of smooth branchiae. Glandular ridges present. Sabellides sibirica Wirén 1883. Abdominal notopodial rudiments absent. Thirteen thoracic uncinigers. 8 species. AMPHARETINAE. E. Glandular ridges present. Abdominal notopodial rudiments absent. Amphicteis (lAryandes) kowalewskii Grimm 1877. Twelve thoracic uncinigers. paleae present. abdominal notopodia] rudiments present. Anobothrus antárctica Monro 1939a. quadribranchiata Fauchald 1972. abdominal notopodial rudiments absent. Paleae absent. only species. AMPHARETINAE. Three pairs of smooth branchiae (Day 1964: four pairs of branchiae in type species). AMPKARETINAE. Glandular ridges absent. Glandular ridges absent. usually cylindrical. Paleae present. Eleven thoracic uncinigers. paleae present. only species. AMPKARETINAE. Two pairs of branchiae on two successive segments. Seliger 11 with notopodia elevated and hirsute notosetae. Prostomium anterioriy sharply pointed. AMPHARETE^iAE. Asabellides Annenkova 1929. Amphisamytha Kessle 1917. 2 species. Eusamythella Hartman 1971. AMPHARETINAE. Hypania Ostroumouw 1897. Notosetae absent in segment 4. macroglossus Eliason 1955. only species. only species. paleae present. Three pairs of smooth branchiae. Buccal tentacles papillose. 2 species. AMPHARETINAE. Buccal tentacles papillose. Three pairs of branchiae. antárctica Mclntosh 1885. japónica Kessle 1917. Twelve thoracic uncinigers. AMPKARETINAE. 3 species. abdoininal notopodial rudiments present. Glandular ridges and abdominal notopodial rudiments present. Oral membrane broad and folded. 14 uncinigers. Setiger 10 or 11 with notopodia elevated and modified notosetae. abdominal notopodial rudiments present. Eleven thoracic uncinigers. Abdominal notopodial rudiments present. only species. AMPHARETINAE. Two pairs of smooth branchiae on first two segments. Amythas Benham 1912. Segment 3 with small capillary setae. Lysippe vanelli Fauvel 1936. paleae present. Clávate notopodial abdominal rudiments present. A. Glandular ridges absent. Glyphanostomum Levinsen 1884. one pair of anal cirri. only species. G. elongata Fauchald 1972. AMPHARETINAE. Twelve thoracic uncinigers. AMPHARETINAE. Three pairs of smooth branchiae. Amphitrite gunneri Sars 1835. Glandular ridges and abdominal notopodial rudiments absent. paleae present. Buccal tentacles few and large. Four pairs of branchiae.1976 THE POLVCHAETE WORMS 125 AMPHARETINAE. paleae absent. Eusamytha sexdentata Hartman 1967. Glandular ridges and abdominal notopodial rudiments absent. membranifera Benham 1921. Hypaniola Annenkova 1927. only species. AMPKARETINAE. A. Glandular ridges absent. Prostomium a small quadrangular lobe. Twelve thoracic uncinigers. Thirteen thoracic uncinigers. dorsal crest on segment 6 present. AMPHARETINAE. Mugga Eliason 1955. pulchella Müller 1858. Glandular ridges and abdominal notopodial rudiments absent. abdominal notopodial rudiments absent. notopodia on segments 2 and 3. paleae present. Nuchal hooks absent. Four pairs of smooth branchiae. Melinnopsides Day 1964. explorons Chamberlin 1919c. Ten thoracic uncinigers. segment 6 without neurosetae. Moyanus Chamberlin 1919c. M. Abdominal notopodial rudiments present. ridge across dorsum on segment 4. M. wahrbergi Eliason 1955. nearly always smooth. Buccal tentacles all similar. MELINNINAE. only species. first notosetae on segment 5. small paleae present. Three pairs of smooth branchiae. atlántica Mclntosh 1885.¡26 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 AMPHARETINAE. Four pairs of smooth branchiae. Mexamage Fauchald 1972. corrugata Fauchald 1972. first notosetae in segment 5. small paleae present. M. MELINNINAE. Ten thoracic uncinigers. Four pairs of smooth branchiae. Last thoracic notopodia elevated with modified notosetae. AMPHARETINAE. 5 species. MELINNINAE. Four pairs of branchiae. paleae present. dorsal crest on segment 6. only species. paleae absent. Glandular ridges absent. Three pairs of branchiae. 14 uncinigers. Three pairs of branchiae. Twelve thoracic uncinigers. Nuchal hooks present. only species. only species. Melinnopsis Mclntosh 1885. antárctica Hartman 1967. Neopaiwa Hartman and Fauchald 1971. AMPHARETINAE. Four pairs of smooth branchiae. only species. but not setae. Melinnata Hartman 1965. flange across dorsum between last thoracic notopodia. other two pennate. All buccal tentacles similar. heterobranchiata Wesenberg-Lund 1949. Capillary notosetae first present in segment 7. one smooth. Dorsal crest on segment 6. Melinnexis Annenkova 1931. First notosetae in segment 5. Melinnopsis capensis Day 1955. only species. cinctus Hartman 1965. Abdominal notopodial rudiments present. MELINNINAE. Glandular ridges absent. Melinna Malmgren 1866. Dorsal ridge either on segment 6 or 7. Ten thoracic uncinigers. Nuchal hooks on both segments 4 and 5. only species. AMPHARETINAE. Isolda Müller 1858. two flanged. M. small paleae present. One very large and numerous small buccal tentacles present. nuchal hooks and dorsal crest absent. Lysippides Hessle 1917. Four pairs of branchiae on four successive segments. /. first notosetae in segment 5. Thirteen or 14 thoracic uncinigers. Prostomium prolonged with oral lobe suspended below it. Glandular ridges absent. First notosetae in segment 5. last thoracic notopodia elevated with modified notosetae. AMPHARETINAE. Twelve or 13 thoracic uncinigers. Melinnampharete Annenkova 1937. /. M. Four pairs of smooth branchiae. Two pairs of smooth branchiae. Three pairs of smooth branchiae. M. . MELINNINAE. abdominal notopodial rudiments absent. Glandular ridges absent. Three pairs of smooth branchiae. nuchal hooks absent. Twelve thoracic uncinigers. Lysippe Malmgren 1866. nuchal hooks absent. 8 species. labiata Malmgren 1866. nelsoni Benham 1927. Three pairs of smooth branchiae. N. 4 species. only species. MELINNINAE. Four pairs of branchiae. Nuchal hooks on segment 4. only species. AMPHARETINAE. Dorsal crest on segment 6. Capillary notosetae present in segments 5 and 6. M. cirrata Hartman and Fauchald 1971. 2 species. AMPHARETINAE. americana Hartman 1965. Ai. Sabellides cristata Sars 1851. two cylindrical. All buccal tentacles similar. Dorsal crest on segment 6. no fused anterior segments. Dorsal crest absent. Four pairs of branchiae. MELINNINAE. Nuchal hooks present. paleae absent. Nine thoracic uncinigers. Ten thoracic uncinigers. M. Melinnoides Benham 1927. 16 thoracic uncinigers. Muggoides Hartman 1965. two smooth and two pennate. eoa Annenkova 1937. Ten thoracal uncinigers. Four pairs of smooth branchiae. only species. paleae present. Fourteen thoracic uncinigers. Irana Wesenberg-Lund 1949. only species. Melinantipoda Hartman 1967. 12 thoracic uncinigers. MELINNINAE. Glandular ridges absent. árctica Annenkova 1931. Paleae absent. 26 species. M. AMPHARETINAE. L. first row of uncini long and ventrally displaced. dorsal crest on segment 6. Fourteen thoracic uncinigers. only species. Amphicteis fragilis Wollebaek 1912. Four pairs of smooth branchiae. Phyllampharete Hartman and Fauchald 1971. only species. N. Three pairs of smooth branchiae. Sabellides Milne Edwards in Malmgren 1966. Glandular ridges absent. Twelve thoracic uncinigers. AMPHARETINAE. only species. luderitzi Augener 1918. only species. Segment 4 without notosetae. Oeorpata Kinberg 1867b. only species. Thirteen thoracic uncinigers. notopodial rudiments present in abdomen. . Glandular ridges absent. Glandular ridges absent. Glandular ridges absent. collaribranchis Augener 1918. notosetae present from segment 2. Buccal tentacles papillose. Three pairs of smooth branchiae. Notosetae absent in segment 4. AMPHARETINAE. Eleven thoracic uncinigers. 6 species. Fourteen thoracic uncinigers. Three pairs of branchiae on three successive segments starting on segment 2. only species. paleae absent. paleae absent. Glandular ridges and abdominal notopodial rudiments absent. Phyllamphicteis Augener 1918. Glandular ridges and abdominal notopodial rudiments absent. Four pairs of pennate branchiae. only species. Glandular ridges absent. abdominal notopodial rudiments present. AMPHARETINAE. only species. 2 species. Eleven thoracic uncinigers. P. Samytha Malmgren 1866. Samythella Verrill 1873a. 8 species. paleae absent. Twelve thoracic uncinigers. P. paleae present. Neosamytha Hartman 1967. notopodial lobes present on segments 4 and 5. P. Eleven thoracic uncinigers. Paleae absent. elongata Verrill 1873a. abdominal notopodial rudiments present. Three pairs of smooth branchiae. Amphicteis brevispinus Grube 1860. small paleae present. only species. small abdominal notopodial rudiments present. crocea Grube 1877. Buccal tentacles papillose. Glandular ridges present. first notosetae on segment 6. Sabellides angustifolia Grube 1878. Pterolysippe Augener 1918. Pterampharete Augener 1918. small paleae reported in some species. AMPHARETINAE. paleae present. three smooth and one pennate. P. Glandular ridges absent. AMPHARETINAE. longicirrata Hartman and Fauchald 1971. Glandular ridges absent. AMPHARETINAE. Three pair? of smooth branchiae. AMPHARETINAE. two smooth and two lamellate. 2 sjiecies. Four pairs of smooth branchiae. Glandular ridges and abdominal notopodial rudiments absent. Four pairs of smooth branchiae. Four pairs of smooth branchiae. Twelve thoracic uncinigers. Paramphicteis Caullery 1944. paleae absent. Glandular ridges and abdominal notopodial rudiments absent. deroderus Chamberlin 1919c. AMPHARETINAE. abdominal notopodial rudiments well developed. small paleae present. Fourteen thoracic uncinigers present. Eleven thoracic uncinigers. paleae present. Twelve thoracic uncinigers. Paleae absent. paleal development absent. Dorsal ridge across segment 4. madurensis Caullery 1944. Pabits Chamberlin 1919c. AMPHARETINAE. Capillary notosetae present from segment 2. Glandular ridges absent. Four pairs of branchiae. Buccal tentacles papillose. armata Kinberg 1867. Sabe lia octocirrata Sars 1835. abdominal notopodial rudiments present. Fourteen thoracic uncinigers. Four pairs of flanged branchiae. Fourteen thoracic uncinigers. elongata Kessle 1917. Fourteen thoracic uncinigers. first notosetae present on segment 5. P. only species. Glandular ridges present. abyssi Chamberlin 1919c. not clearly separable from Isolda (cf. Anus surrounded by a circle of papillae.1976 THE POLYCHAETE WORMS m AMPHARETINAE. AMPHARETINAE. P. N. Paleae present. Neosabellides Hesslc 1917. Paramage Caullery 1944. small abdominal notopodial rudiments present. Sabellides sexcirrata Sars 1856. Four pairs of branchiae. AMPHARETINAE. AMPHARETINAE. Parhypania Annenkova 1928. S. P. AMPHARETINAE. Phyllocomus Grube 1877. Paiwa Chamberlin 1919c. O. Four pairs of flanged branchiae on four successive segments starting on segment 2. Fourteen thoracic uncinigers. Incompletely known. Prostomium anteriorly incised. AMPHARETINAE. Oral lobe prolonged. Day 1964). abdominal notopodial rudiments very small. bipennata Augener 1918. 7 species. MELINNINAE. only species. Glandular ridges and abdominal notopodial rudiments absent. only species. paleae absent. Four pairs of smooth branchiae. abdominal notopodial rudiments present. gracilis Hartman 1967. P. Twelve thoracic uncinigers.. 5a (3a). Piostomium a simple fold. AMPHARETINAE. Glandular ridges absent.128 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 AMPHARETINAE. abdominal notopodial rudiments present. 12 thoracic uncinigers. abdominal noto- podial rudiments present. 3 species.. Glandular ridges absent. Circle of anal papillae. Glandular ridges absent. 17 thoracic setigers Artacama Proboscis grooved. AMPHARETINAE. paleae absent. The usually numerous buccal tentacles cannot be fully retracted into the mouth. see Melinnopsides Microsamytha Augener 1928a. Four pairs of smooth branchiae. 2b (lb). Twelve thoracic uncinigers. only species. Invalid Genera Aryandes Kinberg 1867b. Glandular ridges absent. wireni Hessle 1917. 2a (lb). Weddellia Hartman 1967. 15 thoracic setigers Artacamella First notosetae on first postbranchial segment Pseudampharete . when present. Sosanella Hartman 1965. AMPHARETINAE. ses Sabellides Melinnides We sen berg-Lund 1950. lb. 4a (la). Fourteen thoracic uncinigers. 5. Samythopsis Mclntosh 1885. S. AMPHARETINAE. only species.. Four pairs of smooth branchiae. paleae absent. Paleae absent. AMPHARETINAE. 4b (la). Thirteen thoracic uncinigers. Terebellids are among the most common shallowwater polychaetes and are found in all environments. Three pairs of smooth or flanged branchiae. Sosanopsis Kessle 1917. Segment 3 with capillary setae. Other forms may stretch the buccal tentacles into the water and capture particles from the water. Four pairs of smooth branchiae. S. include one to three pairs on the first segments. Uncini usually with a large main fang and a crest of smaller teeth. apalea Hartman 1965.. abdominal notopodial rudiments absent. glandularis Hartmann-Schröder 1965. They usually are grooved and used in selective deposit-feeding on the surface. 14 Thoracic uncini in single rows in all setigers 3 Branchiae present THELEPINAE 5 Branchiae absent POLYCIRRINAE 10 I>roboscis papillated. Abdominal notopodial rudiments absent. S. Three pairs of smooth branchiae. hiltoni Chamberlin 1919a. profimda Hartman 1967. Three pairs of smooth branchiae. paleae present. Fifteen thoracic uncinigers. AMPHTTRITINAE . S. Setiger 11 with modified notosetae. Twelve thoracic uncinigers. Glandular ridges and abdominal notopodial rudiments present. Glandular ridges and abdominal notopodial rudiments absent. W. abdominal parapodia with long dorsal cirri. Third from last thoracic notopodia elevated and with modified setae. but without paleal modifications. sec Amphicteis Eusamytha Hartman 1967. AMPHARETINAE. sulcata Malmgren 1866. only species. Glandular ridges absent. indeterminable FAMILY TEREBELLIDAE MALMGREN 1867 Body in two regions. associated distinctly with separate segments. AMPHARETINAE. Sosane Malmgren 1866. one smooth and three lamellate. set Ampharete Crossostoma Gosse 1855. Schistocomus Chamberlin 1919a. 2 species. abdominal notopodial rudiments present. grübet Mclntosh 1885. see Alkmaria Pseudosabellides Berkeley and Berkeley 1943. 3b (2b). paleae absent. only species. Two pairs of anal cirri. indeterminable Branchiosabella Claparède 1863. see Eusamythella Heterobranchus Wagner 1885. Branchiae. Notosetae of last thoracic setiger modiñed. anterior region with biramous parapodia and posterior region with neuropodia only.. Three pairs of smooth branchiae on three successive segments. paleae present. Twelve thoracic uncinigers. Four pairs of branchiae. Key to Genera la. 3a (2b). 5 species. see Asabellides Rytocephalus Quatrefages 1866. see Melinnexis Annenkova. 1931 Melinnopsis Day 1955. Sosanides Hartmann-Schröder 1965. Notopodia in setiger 13 (third from last thoracic notopodium) elevated with hirsute setae. paleae absent. A large scoop-shaped or conical proboscis present ARTACAMINAE 4 Proboscis absent 2 Thoracic uncini in double or alternating rows in at least some setigers . S. 24b (23b). 28b (24b). 33a (32b). but never directly associated with the branchiae 24 Uncini from third thoracic setiger Neoleprea Uncini from second thoracic setiger 25 Sixteen or 17 thoracic setigers 26 Eighteen or more thoracic setigers 28 A single pair of branchiae. 8a (7a). 25a (24b). 32b (31a). uncini from segment 5 33 Branchiae smooth. 25b (24b). 7b (6b). 8b (7a). 17a (16a). 20b (19b). all notosetae denticulated 20 Fifteen thoracic setigers. 17b (16a). some notosetae smooth Proclea Neurosetae fiom setiger 2. 24a (23b). 26b (25a). 6a (5b). 16 thoracic setigers Spiroverma Two or three pairs of branchiae. 15b ( 14a). 13a (1 lb). THE POLYCHAETE WORMS 129 First notosetae on one of the branchial segments 6 First notosetae on the first branchial segment Streblosoma First notosetae on the second branchial segment 7 Lateral lappets present on one or a few anterior segments 8 Lateral lappets absent 9 Uncini from the fifth segment Euthelepus Uncini from the first abdominal segment Telothelepus Uncini from seliger 9 Parathelepus Uncini from setiger 2 Tkelepus Setae completely absent Hauchiella At least some setae present 11 Neurosetae absent 12 Neurosetae present 13 Thoracic notopodia vascularized and in part furcate or branched Enoplobranchus Thoracic notopodia may be vascularized. 27a (26b). 13b (lib). 2 and 5 Polymniella Branchiae on segments 3. uncini from segment 6 Opisthopista Notosetae from segment 4. 1\2L (15b). branchiae sessile filaments Amphitrite Branchiae attached on segments 2-4 29 Branchiae attached on another combination of segments 30 Branchiae arbwrescent from the base Terebella Branchiae distinctly stalked Amphitritides Branchiae on segments 1. 10a (3b). 31b (22b). 7a (6b). 18a (17b). 12b (1 la). 30b (28b). 17 thoracic setigers Laphania Lateral lappets present on one or a few anterior segments. 16 thoracic setigers 18 Third segment with transverse ridge across dorsum Leaena Third segment without transverse ridge Stschapovella Neurosetae from setiger 3. 32a (31a). 12a (1 la). 10b (3b). 9a (7b). 29a (28a). 9b (7b). 16b ( 15a).7976 5b (3a). ampharetinlike Eupistella . 19b (16b). 28a (24b). 20a (19b). but never furcate or branched Lysilla Neurosetae short-handled uncini Polycirrus Neurosetae long-handled spines Amaeana Branchiae absent 15 Branchiae present 22 Seventeen or fewer thoracic setigers 16 More than 20 thoracic setigers 21 All notosetae smooth-tipped 17 At least some notosetae with denticulated tips 19 Lateral lappets absent. 26a (25a). II b (10b). 27b (26b). 17 thoracic setigers 27 Lateral lappets present. 23b (22a). 31a (22b). distally smooth-tipped Baffinia Some notosetae subdistally inflated. 7 and 13 Terebellobranchia Anterior uncini long-handled 32 All uncini short-handled 35 Notosetae from segment 5. 14a (2a). 14b (2a). notosetae finely denticulated Lanassa Fourteen thoracic sefigers. 29b (28a). 21b (15b). 22b (14b). 19a (16b). I la ( 10b). 18b (17b). 16a (15a). 15a (14a). branchiae stalked and branched Neoamphitrite Lateral lappet absent. 23a (22a). 6b (5b). 22a (14b). 30a (28b). some notosetae distinctly pectinate Phisidia All notosetae tapering. distally denticulated Spinosphaera Notosetae with marginally serrated tips 23 Notosetae with smooth tips 31 Three anterior segments with large lateral lappets to which the branchiae are attached . Colymmatops Lateral lappets present or absent. 2 species. Three pairs of smooth. Lateral lappets absent. 8 species. notosetae from segmem 3. 44a (36b). AMPHITRITINAE. 46b (45b). 48b (47b). Müller \11\. Grooved. 35b (31b).130 33b (32b). 36b (35b). Eyes rarely present. Notosetae distally smooth. Uncini absent. thorax with 15 to 40 setigers Nicolea Three pairs of branchiae. Sevemeen thoracic seügers. 3 species. . 39b (38b). 45b (44b). uncini face to face in posterior thoracic setigers. 43a (42b). distally serrated. A. 47a (46b). each uncinus long-shafted. nephridial papillae on segment 3 and fi-om segment 6. Lateral lappets absent. Müller 1771. 38a (37b). 17 species. P^P'"««« proboscis attached ventrally on the peris"""*""'• Artacamella Hartman 1955. branchiae short-shafted or sessile 38 Eighteen thoracic setigers Melinella Seventeen thoracic setigers 39 First four segments flattened with large lateral lappets forming an anterior plaque. 36a (35b). 44b (36b). 34b (33b). cirrata O. Terebella gracilis Grube 1860. Sixteen thoracic setigers. stalked branchiae from segment 2. tube opening unadorned Eupofymnia Large lateral lappets on segments 2 and 4. Terebella flexuosa Grube 1860. 13 to 25 thoracic setigers. branchiae very long-shafted Axionice Seventeen or 18 thoracic setigers. Artacama Malmgren 1866. A.5 species POLYCIRRINAE. branched branchiae. Amphitritides Augener 1922. Two pairs of branching. Polycirrus trilobatus Sars 1863. 39a (38b). ampharetinlike branchiae on segments 2-4. two or three pairs of branchiae 40 Branchiae tufts of sessile filaments Thelepides Branchiae branching off one or a few stems 41 Buccal lateral lappets form a crest across the dorsum Paralanice No dorsal crest across the buccal segment 42 Uncini with teeth in a single row Loimia Uncini with teeth in two or more rows 43 Small lateral lappets on segments 2 and 3. Fifteen tho•cic setigers. 42b (41b). 41a (4üb). 42a (41b). 37b (36a). 38b (37b).F. 40a (39b). tube opening with fan-shaped frills Lanice Thirteen thoracic setigers Ramex At least 15 thoracic setigers 45 First notosetae on first branchial segment Naneva First notosetae on last branchial segment or from one of the first postbranchial segments 46 Uncini present from first setiger Reieterebeila Uncini present from second setiger 47 Uncini in an open circle in posterior thoracal setigers Terebellanice Uncini arranged back to back in posterior thoracal setigers 48 Two pairs of branchiae. Notosetae from fourth segment.F. 47b (46b). all uncini short-handled. Three pairs of sessile branchiae from segment 2. Serrated notosetae present from segment 4. Three pairs of filamentous ''^^'«^Wae on segments 2-4. notosetae distally smooth. lateral lappets present. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Branchiae branched 34 Lateral lappets at least on segments 2 and 4 Pista Lateral lappets limited to segment 3 Lanicides Ventrum with large anteriorly opening glandular folds on segments 3-5 Paraxionice Ventrum with glandular scutes or smooth 36 Lateral lappets present on one or a few anterior segments 37 Lateral lappets inconspicuous or absent 44 Sixteen thoracic setigers. ARTACAMINAE. Ten to 13 thoracic segments. 34a (33b). AMPHITRITINAE. Tentacular lobe trifoil-shaped.A. 41b (40b). 43b (42b). lateral lappets varying in size. abdominal neurosetae long-shafted spines. 40b (39b). thorax with 17 setigers Scionides Generic Definitions Amaeana Hartman 1959. 45a (44b). 48a (47b). 37a (36a). boat-shaped proboscis attached ventrally on the peristomium. Lateral lappets absent. Axionice Malmgren 1866. hancocki Hartman 1955. one pair of branchiae Scionella First four segments not modified into a flattened plaque. 46a (45b). AMPHITRITINAE. uncini in all thoracic setigers. Amphitrite O. Two or three pairs of longshafted. proboscidea Malmgren ^^^^' ^ ^P^*^'**ARTACAMINAE. 35a (31b). Branchiae on segments 2-4. Euthelepus Mclntosh 1885. Branchiae absent. Smooth-tipped notosetae from segment 4. AMPHITRITINAE. POLYCIRRINAE. Lateral lappets present. 7 species. notosetae spinose capillaries. lateral lappets present. notosetae from segment 3. only species. E. 12 species. Notósetae present from third segment to the end of the body (more than 70 segments). Amphitrite affinis Malmgren 1866. only species. Seventeen thoracic setigers. AMPHITRITINAE. AMPHITRITINAE. Smooth-tipped notosetae present from segment 4. Lateral lappets present. Three pairs of branched branchiae. Uncini with short handles. ampharetinlike branchiae. granulatus Peters 1854. long-shafted uncini present from segment 5. notosetae distally smooth. 8 species. only species. Eupista darwini McIntosh 1885. Lateral lappets present. POLYCIRRINAE. Terebella medusa Savigny 1818. Three first segments with large lateral lappets to which are attached branchiae. 4 species. Leaena abyssorum McIntosh 1885.setigers with distally serrated notosetae. Lateral lappets present. AMPHITRITINAE. with usually one seta much longer than the others. vascularized and in part furcate or branched. Three pairs of branched branchiae. Tube with branched fine-meshed fan attached to opening. some anterior uncini with prolonged shafts. 10 species. nordenskioldi Malmgren 1866. L. Lanicides Hessle 1917. One pair of branched branchiae. Amphitrite nesidensis delle Chiaje 1828. A^. then biserial and finally uniserial in last 30-35 setigers. THELEPINAE. Neurosetae completely absent. third segment with transverse ridge across dorsum. multisetosa Wesenherg-Lund 1950. L. Enoplobranckus Webster 1879. denticulated tips. Seventeen thoracic setigers. AMPHITRITINAE. Branchiae absent. Sixteen thoracic setigers. AMPHITRITINAE. Six to 12 thoracic segments. Notopodial lobes prolonged. AMPHITRITINAE. Hauchiella Levinsen 1893. Three pairs of branching branchiae. Two pairs of smooth. AMPHURITINAE. resembles Proclea in setal structures. Naneva Chamberlin 1919a. Notosetae distally serrated. notosetae distally smooth. Three pairs of branched branchiae. Two pairs of dendritically branched branchiae from segment 2. Lateral lappets absent. present on 20 segments. 18 thoracic setigers. Lateral lappets present. Twentyseven thoracic setigers. Eupistella Chamberlin 1919c. B. Lateral lappets sometimes present. setubalensis Mclntosh 1885. Branchiae absent. 3 species. uncini present from second setiger. Seventeen thoracic setigers. smooth-tipped notosetae present from first branchial segment. lateral lappets present. Two pairs of branched branchiae. 16 species. Uncini absent. all with uncini. POLYCIRRINAE. only species. AMPHITRITINAE. Branchiae absent. Branchiae absent. . Thorax of ten segments. Fifteen thoracic setigers. 10 species. Lateral lappets absent. héspera Chamberlin 1919a. 17 thoracic setigers. Thirteen or 14 thoracic setigers. sometimes as single filaments only. crested. lateral lappets on segments 2-3. Notosetae from second branchial segment. L. Cotymmatops Peters 1854. Uncini with all teeth in a single row (pectinate). 6 species. Bathya Saint-Joseph 1894. boecki Malmgren 1866. Loimia Malmgren 1866. notosetae smooth-tipped. 3 species. Terebella abranchiata Sars 1865. Incompletely described. Seventeen thoracic. only species. usually about 70 segments in all. Neoamphitrite Hessle 1917. Capillary setae distally smooth. Uncini first present from first postbranchial segment. only species. Terebella (Phyzelia) bilobata Grube 1877. Uncini present from setiger 7. Nereis conchilega Pallas 1766. AMPHITRITINAE. capillary distally smooth. AMPHITRITINAE. Melinetla Mclntosh 1914. AMPHITRITINAE. C. uncini back to back in posterior thoracic setigers. Laphania Malmgren 1866. Lysilla Malmgren 1866.}976 THE POLYCHAETE WORMS m Baffinia Wesenberg-Lund 1950. Incompletely described. Eupolymnia Verrill 1900. loveni Malmgren 1866. AMPHITRITINAE. AMPHITRITINAE. All setae absent. A/. Seventeen thoracic setigers. Chaetobranchus sanguínea Verrill 1873b. Lanassa Malmgren 1866. Lanice Malmgren 1866. Smooth-tipped notosetae present from segment 4. Nephridial papillae present from segment 3. only species. Leaena Malmgren 1866. II species. Polycirrus tribullata Mclntosh 1869. Uncini present in double-rows in most thoracic segments. macduffi Mclntosh 1914. Smooth-tipped notosetae from segment 4. Notosetae present from segment 4. uniserial in first eight uncinigers. large lateral buccal lappets connected across dorsum with a crest. californiensis Hartman 1944b. First notosetae in segment 5 and first uncini in segment 6. P. Two pairs of branched branchiae. notosetae distally smooth. Three pairs of branched branchiae. 15-40 thoracic setigers. One pair of branchiae on segment 4. notosetae distally smooth. POLYCIRRINAE. Notosetae from segment 4. Notosetae first present from segment 3. aurantiaca Verrill 1900. Four first segments flattened dorsoventrally with very large longitudinally oriented lateral lappets. Paralanice Caullery 1944. 39 species. Branchiae absent. Sixteen thoracic setigers. 2 species. AMPHITRmNAE. 4 species. Sixteen thoracic setigers. pacifica Kessle 1917. notosetae first present on segment 4. uncini from segment 2. lateral lappets present on at least segments 2 and 4. THELEPINAE. Pista Malmgren 1866. Uncini arranged back to back in posterior thoracic segments. lateral lappets absent. notosetae from segment 4. Nicolea Malmgren 1866. AMPHITRmNAE. Twentytwo or more thoracic segments. THELEPINAE. Notosetae distally denticulate. Smooth-tipped notosetae from segment 4. Pseudatnpharete Hartmann-Schröder 1960b. R. Notosetae first present from first postbranchial segment. with pectinate appearance. the shorter coarse. Three pairs of branched branchiae on segments 1. notosetae present from segment 3. only species. Sixteen thoracic setigers. S. AMPHITRmNAE. Amphitrite cristata O. Scionella Moore 1903. Thorax with a variable number of setigers. Two pairs of sessile branchial filaments on large bosses on first and second segment. Phisidia Saint-Joseph 1894. P. only species. timorensis Caullery 1944. Scionides Chamberlin 1919b. tentaculata Hartmann-Schröder 1960b. Uncini present from first setiger. longer finely so. some distally denticulated. P. AMPHITRmNAE.F. AMPHITRITINAE. Leaena oculata Langeriians 1880. thorax with 13 setigers. Ramex Hartman 1944b. uncini present from setiger 2. Lateral lappets present. Uncini arranged back to back in posterior thoracic segments. AMPHITRITINAE. Parathelepus Caullery 1915.132 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Neoleprea Kessle 1917. AMPHITRITINAE. lateral lappets absent. Branchiae absent. only species. Notosetae distally smooth or serrated. Three pairs of branchiae. O. lateral lappets inconspicuous. Three pairs of branched branchiae. Paraxionice Fauchald 1972. seventeen thoracic setigers. Proclea Saint-Joseph 1894. Notosetae from segment 1. Terebella zostericola Örstcd 1844. lateral lappets large. Polycirrus Grube 1850. Uncini from setiger 3. on segments 2 and 4 at least. 22 species. Thelepides collaris Southem 1914. only species. AMPHITRITINAE. 3 species. P. notosetae distally smooth. Müller 1776. marginally dentate. 5 species. Two pairs of branched branchiae. Branchiae absent. Spinosphaera Kessle 1917. Seventeen thoracic setigers. 14 thoracic setigers. One pair of branched branchiae on second segment. 2 species. Uncini from setiger 2. . only species. only species. Notosetae from second branchial segment. Two pairs of stalked branched branchiae. Twenty-three or more thoracic setigers. 3 species. Seventeen-40 thoracic setigers. notosetae from segment 4. AMPHITRITINAE. P. Nephridial papillae present on segments 3-9. S. notosetae distally either dentate or smooth. AMPHITRITINAE. only species. Uncini first present from segments 7-18. some smooth. AMPHITRITINAE. distally smooth. 2 and 5. Three pairs of branched branchiae. smooth-tipped capillaries from segment 4. notosetae distally smooth. lateral lappets also on segments 2 and 3. long-handled uncini present in anterior setigers. Lateral lappets absent. Ventral part of segments 3-5 covered by a large glandular apparatus that opens anteriorly on segment 3 in two trumpetshaped openings. Reteterebella Hartman 1963b. AMPHITRmNAE. Fifteen to 24 thoracic setigers. queenslandia Hartman 1963b. Opisthopista Caullery 1944. japónica Moore 1903. One pair of branched branchiae with double bases on segments 3 and 4. artifex Fauchald 1972. Polymniella Verrill 1900. uncini from setiger 9. Seventeen thoracic setigers. AMPHITRITINAE. lateral lappets absent. Leprea streptochaeta Ehlers 1897. R. Two or three pairs of branched branchiae. only species. Smooth-tipped notosetae present from segment 4. AMPHITRmNAE. medusa Grube 1850. Anterior uncini long-shafted. 40 species. uncini from setiger 7. forming a large oblique plaque at the anterior end. Leaena graffii Langerhans 1880. Terebella reticulata Ehlers 1887. sibogae Caullery 1944. T. uncini from segment 5. Branchiae absent. However. see Enoplobranckus Cyaxares Kinberg 1867b. Body strongly spiralled. Smooth-tipped notosetae from segment 4. Two or three pairs of branchiae from segment 2. Invalid Genera Amaea Malmgren 1866. Notosetae finely capillary rather than limbate. in that these genera resemble branchiate members of the AMPHITRITINAE very closely in setal structures as well as in the structure of the anterior end. Fifteen thoracic setigers. notosetae marginally serrate. AMPHITRITINAE. only species. indeterminable Anisocirrus Gravier 1905a. T. see Pista Ehlersiella Mclntosh 1885. T. no revision was attempted on this occasion. Terebellanice Hartmann-Schröder 1962b. The genus Pseudothelepus Augener 1918 is considered here a synonym of Streblosoma. The genus Pseudampharete has been included among the THELEPINAE since it is branchiate and has the uncini in single rows in all thoracic segments. capensis Day 1955. Two or three pairs of branched branchiae. THELEPINAE. number of setigers. Terebella Linnaeus 1767. Lateral lappets absent. THELEPINAE. AMPHITRITINAE. NOMEN NUDUM Chaetobranchus Verrill 1873b. see Polycirrus Athelepus Chamberhn 1919c. see Polycirrus Dejoces Kinberg 1867b. Thelepus Leuckart 1849. laeviseta Hartmann-Schröder 1962b. T. tatjanae Levenstein 1957. Stschapovella Levenstein 1957. koehleri Gravier 1911a. see Polycirrus Apneuma Quatrefages 1866. 32 species. 3 species. Notosetae smoothtipped. Three pairs of filiform branchiae. 20 species. notosetae from segment 4. see Polycirrus Eugrymaea Verrill 19(X). Smooth-tipped notosetae fi-om segment 3. The treatment here reflects this view. AMPHITRITINAE. It belongs in the abranchiate gorup of AMPHITRITINAE. only species. this latter feature may also be present among members of the AMPHITRITINAE. uncini present on abdomen. S. lateral lappets absent. lapidaria Linnaeus 1767. All nephridia free from one another. only species. Neuiosetae absent on thorax. One pair of sessile branchiae. as suggested by Day (1967). lateral lappets present. Uncini in an open circle on posterior thoracic segments. The genus Batkya Saint-Joseph 1894. filiform branchiae on seg ments 2-4. Sessile. see Eupistella Euscione Chamberlin 1919c. AMPHITRITINAE. T. indeterminable Dendrophora Grube 1870a. 28 species. distally serrated. see Thelepus . seeAxionice Grymaea Malmgren 1866. Grymaea bairdi Malmgren 1866. ononokomachii Uchida 1968. Amphitrite cincinmta Fabricius 1780. usually large. natalensis Day 1951. 17 thoracic setigers present. the longer ones with hispid swellings subdistally. see Streblosoma Heterophenacia Quatrefages 1866. This change appears unfortunate. The placement must be considered temporary. THELEPINAE. Three pairs of sessile branchiae on segments 2-4 (may be absent). Sixteen thoracic setigers. Lateral lappets present. ste Eupolymnia Amphytrite Renier 1804. Two pairs of branchiae from third segment. AMPHITRITINAE. Spiroverma Uchida 1968. and is very poorly known. ss&Amaeana Amphiro Montagu 1808. It further resembles members of this subfamily in that the branchiae are sessile filaments. Thorax with variable. Telotheiepus Day 1955. Uncini from segment 5. se& Amphitrite Amphitritoides Costa 1862. more than 19 thoracic setigers present. Taxonomic Notes Day (1967) altered the definition of the sub-families to include the abranchiate members of AMPHITRITINAE among the POLYCIRRINAE. Tentacular lobe very large and frilly. listed above in the definitions has not been included in the key. Uncini from segment 5. Terebellobranchia Day 1951. Notosetae from first branchial segment (segment 2). Thelepides Gravier 1911a. Three pairs of branched branchiae on segments 3. lateral lappets present. only species. Sixteen thoracic setigers.1976 THE POLYCHAETE WORMS m Notosetae denticulate. Some of the genera are difficult to separate from related forms. Distally serrated notosetae present from segment 4. 2 species. see Polycirrus Aphlebina Claparède 1864. indeterminable Ereutho Malmgren 1866. see Streblosoma Eupista Mclntosh 1885. see Polycirrus Dendrobranchus Wagner 1885. smooth-tipped notosetae present from second branchial segment (segment 3). S. AMPHITRITINAE. Notosetae from second branchial segment (segment 3). lateral lappets absent. Uncini face to face in posterior thoracic segments. 7 and 13. each with maximally eight filaments on segment 2. Streblosoma Sars 1872. 2a (la). see Parathelepus Thelepodopsis Sars 1872. Octobranchus Marion and Bobretzky 1875. Notosetae capillary. seeLanice FAMILY TRICHOBRANCHIDAE MALMGREN 1866 Body separated into two regions. see Terebella ¡dalia Quatrefages 1866. The trichobranchids are related closely to the terebellids and frequently are included among them as a subfamily (Day 1967). 5b (3b). Two pairs of smooth brapchiae on the second and third segment (presetal branchiae). anterior region with biramous parapodia. Novobranchus Berkeley and Berkeley 1954. 6a (5b). &ee Pista Laphaniella Malm 1874. each with pectinate branchial lamellae. see Euthelepus Schmardanella Mclntosh 1885. 2b (la). One species. The large folded prostomium also gives them a characteristic appearance. 4 species. see Thelepus Odysseus Kinberg 1867b. Maximally four pairs of branchiae present on anterior setigers. first uncini on setiger 4. Four pairs of branchiae on segments 2-5. see Thelepus Neottis Malmgren 1866. Terebella lingulata Grube 1863. see Pista Pallonia Costa 1862. in that they tend to be slenderer and more muscular than the terebellids. 3b (lb). each lobe with numerous flat branchial lamellae Two or three pairs of branchiae present Four pairs of branchiae present . 6b (5b). see Pofycirrus Uncinochaeta Quatrefages 1866. . pacificus Berkeley and Berkeley 1954. see Pista Solowetia Ssolowiew 1899. Terebellides Sars 1835. see Eupolymnia Protothelepus Verrill 1900. Members of the family appear to be most common in cold-water soft bottoms and may be represented best in deep water. seeNicolea Thelephusa Verrill 1871. F. 4b (3a). seeAxionice Scionopsis Verrill 1873b. Sixteen thoracic setigers. stc Terebella Leucariste Malmgren 1866. Sixteen thoracic setigers. Two pairs of branchiae. last pair a frilled rosette. first setiger is segment 3. vanhoeffeni Ehlers 1913. only species. Prostomium a large hood over the mouth. see Eupolymnia Phenacia Quatrefages 1866. abdomen with densely crested uncini. seeNicolea Pofymnia Malmgren 1867. see Terebella Heteroterebella Quatrefages 1866. A. Filibranchus Malm 1874. 3a (lb). uncini present on nine last thoracic setigers and on abdomen. se^ Pofycirrus Lumara Stimpson 1854. digitate projection Branchia stalked with four lobes. lb. posterior region with neuropodia only. T. N. first three pairs lanceolate. stroemi Sais 1835. roseus Malm 1874. Four pairs of branchiae on first setiger. They differ rather strikingly in body-proportions. 15 thoracic setigers Each branchia with pectinate branchial lamellae Each branchia simple and digitate or rosette-shaped All branchiae simple and tapering Fourth pair of branchiae sessile rosettes Generic Definitions 2 3 Unobranchus Terebellides 4 5 Filibranchus Trichobranchus Ampharetides 6 Octobranchus Novobranchus Ampharetides Ehlers 1913. A single middorsaUy attached branchia present At least a pair of branchiae present Branchia a single tapering. Four pairs of similar lanceolate branchiae. indeterminabie Venusia Johnston 1865. 4a (3a). Terebellides stroemi. only species. all with uncini. 17 thoracic setigers Three pairs of branchiae. sttLanassa Phyzelia Schmarda 1861. 14 species. see Thelepus Wartelia Giard 1878. see Terebella Scione Malmgren 1866. indetenninable Otanes Kinberg 1867b. only species.134 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Heterophyselia Quatrefages 1866. Fifteen thoracic setigers. see Thelepus Tkelepides Southern 1914. Anterior neuropodia with long-shafted hooks. setLanassa LepreaMa\mgren 1866. has been reported from worldwide areas. Key to Genera la. see Thelepus Torquea Leidy 1855. uncini present ftom setiger 4. 5a (3b). seeProclea Thelepella Chamberlin 1919c. see Thelepus Pherea Saint-Joseph 1894. Seventeen thoracic setigers. an important member of soft-bottom environments. and is apparently nearly everywhere. attempts have been made to avoid the more confusing part of the terminology. uncini present from seliger 6. ornato Hartman and Fauchald 1971. cylindrical and tapering posteriorly. 5 species. such as species of Fabricia and allied genera. Three pairs of simple. The shafts of these Z-shaped uncini may be short or long. fused with the peristomium which usually forms a large tentacular crown. By avicular uncini are meant uncini that are essentially Z-shaped (sometimes called swan-shaped) with the small crested head at the top of the Z. One of the key features in the group lies in the structure of the thoracic uncini. are important. Parapodia biramous with long slender. all thoracic setigers with uncini. these also have been called pennoned setae or pick-ax setae. They occur in an anterior row.or short-handled. only species. but never calcareous. in part in double rows.1976 THE POLYCHAETE WORMS /J5 Invalid Genera Eighteen thoracic setigers. and the presence of small. Uncini crested or with teeth in several rows. prostomium rounded without appendages. first setae on segment 3. T. made of varying material. One anterior asetigerous segment. see Terebellides Corephorus Grube 1846. Boguella Hartman and Fauchald 1971. often nearly straight shaft. often maroon or red-colored tentacular crowns. first setae on segment 6. only species. Twelve thoracic setigers. Single dorsal branchia with four lamellate branchial lobes. by which is meant that the crested head. and the shaft sharply bent. are capable of moving around. have setae resembling the terebellid uncini. U. four first setigers without neurosetae. Most of the larger forms are associated with shallow water. in front of the uncini they accompany and usually are small and deeply imbedded in the epidermal tissues. smaller species are common in deep sea collections. but a complete avoidance of this terminology has not been possible. but lack completely anterior appendages. The neutral term companion seta is preferred here. The major characteristics used to identify the sabellids. Two anterior asetigerous segments. B. first seliger is segment 3. Fifteen thoracic setigers. These may be acicular. especially the larger ones. in part spinose notosetae and short-handled. with one large tooth and several smaller ones. Tube present in most species. Most of the forms. The structure of the tentacular crown such as the number of radioli. . lb. The bogueids. Trichobranchus Malmgren 1866. than in microscopic preparation under the compound microscope. three first setigers without neurosetae. long. avicular crested uncini in the neuropodia. Key to Genera la. enigmática Hartman 1948. Sabellids characteristically have nearly smooth appearing bodies. glacialis Malmgren 1866. Vnobranchus Hartman 1965. They are more easily seen by the reflection they give off under a stereo microscope. In the key below. FAMILY SABELLIDAE MALMGREN 1867 Body cylindrical with a thorax of few setigers and abdomen with few to many. lanceolate branchiae. large lanceolate branchia present mid-dorsally on segment 2. external appendages called sty lodes. see Terebellides FAMILY BOGUEIDAE HARTMAN AND FAUCH ALD 1971 Body slender and cylindrical with few segments. Aponobranchus Gravier 1905a. include the presence or absence of companion setae to the neuropodial uncini in the thorax. with large. ß. the smaller forms. setae include thoracic notopodial limbate and geniculate kinds and neuropodial uncini. only species. A single. known for two genera. setal positions reversed in the abdomen. is supported by a gently curved. uncini first present on setiger 4. ORDER SABELLIDA Prostomium reduced. are strictly sessile and never leave their tubes. abyssalis Hartman 1965. They are known from a few locations in the western Atlantic Ocean only. First three setigers without neurosetae First four setigers without neurosetae Boguea Boguella Generic Definitions Boguea Hartman 1945. 2b (lb). about I7x. (B). Caobangia abbotti. right lateral view of the above. (A). Famüy SABELLONGIDAE. Sabellonga disjoncta. (D). but always strongly bent shafts (avicular). 50 m. gently curved shafts (acicular) companion setae always absent FABRICINAE 3 Thoracic uncini with short or long. off Santa Catalina Island. 18x.136 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 FIGURE 37. discrete tori 2 Thoracic uncini with long. companion setae present in some forms SABELLINAE 15 Abdomen with two or three setigers 4 Abdomen with four or more setigers 9 Abdominal uncini form nearly complete cinctures around the body . 1969. posterior end of the above. 1974. modified after Hartman. Family SABELLIDAE. about 17x. 33x.. Key to Genera lalb. 3b (2a). (C). after Jones. anterior end in right ventrolateral view. 3a (2a). 33x. Chone sp. 2a (lb). Family CAOBANGI•)AE. MYXICOLINAE Myxicola Abdominal uncini in short. (E). with fused segments or greatly expanded pygidium 10 Posterior end not modified 11 Seven last setigers covered dorsaily by the large. 29b (28b). 26a (2lb). 20b (18b). 15a (2b). two pairs of radioli Mamymkia Posterior end distinctly modified. 23a (22b). 6a (5b). 25a (23b). spatulate setae present Eudistylia Large subdistal compound eyes on a few dorsal radioli Megalomma Eyes. three pairs of radioli Augeneriella Ventral vascularized filaments smooth. 28b (27b). external stylodes very small Pseudobranchiomma Eight thoracic setigers. 21b (15b). 28a (27b). 22b (21a). 18b (16b). Spirographis Both parts of the tentacular crown similar in size. Radióles partially united by a membrane Monroika Radióles free to the base 5 With an extensive lower lip. 9a (3b).. if present. 19a (18a). 27a (26b). 8b (7b). 14a (13b). spatulate setae absent . ring-shaped pygidium Euchonella Last several setigers flanged laterally to form a spoon-shaped dorsal cavity Euchone Radióles united by a membrane Chone Radióles free to the base '• 12 Abdominal uncini with a large main fang and a crest of smaller teeth Jasmineira Abdominal uncini with major teeth in two or more rows 13 More than ten pairs of radioli in the tentacular crown Dialychone Maximally five pairs of radioli in the tentacular crown 14 Radióles externally distinctly ridged Oriopsis Radióles externally rounded Desdemom Thoracic neuropodial companion setae absent 16 Thoracic neuropodial companion setae present 21 Radioli with external stylodes 17 Radióles without external stylodes 18 Four thoracic setigers. 18a (16b). 19b (18a). 23b (22b). 5b (4b). 26b (21b). 1 lb (9b). 8a (7b). 16b (15a). smaller and scattered over most of the radioli 27 Collar reduced or absent. 27b (26b). 7b (6b). 14b (13b). THE POLYCHAETE WORMS ^•Í7 N. external stylodes well developed Branchiomma Thoracic spatulate notosetae present 19 Thoracic spatulate notosetae absent 20 Thoracic uncini with gently curved. 20a (18b). transversely striated setae Bispira Lancet-shaped notopodial setae absent 23 Collar two-lobed 24 Collar four-lobed 25 Anterior ventrura irregularly rugose Distylidia Anterior ventrum smooth. 30a (29b). 12a (1 lb). 30b (29b). prostomium anteriorly rounded 6 Abdominal uncini short-handled Fabriciola Abdominal uncini long-handled 1 Without ventral vascularized filaments Fabricia With ventral vascularized filaments in the tentacular crown 8 Ventral vascularized filaments branched. 21a (15b). 15b (2b). 22a (21a). 6b (5b). triangular ventral projection absent 28 Notosetae of the collar-segment in a long row Hypsicomus Notosetae of the collar-segment in a tuft 29 Radioli dichotomously branching Schiiobranchia Radioli not divided 30 Spatulate thoracic notosetae absent Sabella Spatulate thoracic notosetae present 31 Thoracic uncini at least in part nearly straight and long-handled 32 . 29a {28b).]Ç76 4a (3a). 24a (23a). 17a (16a). 25b (23b). 10a (9a). 13a (12b). radioli free to the base Sabellastarte Tentacular crown with radioli arranged in one or two spirals 22 Tentacular crown with radioli in two semicircles 26 Posterior thoracic notopodia with lancet-shaped. 11 a (9b). 24b (23a). 4b (3a). 16a (15a). long handles Fabrisabella Thoracic uncini with sharply bent handles Laonome Collar reduced. 31a (30b). 5a (4b). 17b (16a). 10b (9a). curved up under the weakly bilobed prostomium Pseudofabricia Without a distinct lower lip. 9b (3b). apart from the glandular fields Demonax One half of tentacular crown very much larger than the other. 7a (6b). 13b ( 12b). radioli basally united by a web Euratella Collar well developed. 12b (1 lb). a triangular ventral projection at base of tentacular crown present Amphigiena Collar well developed. Twelve pairs of radioli. Twelve or more pairs of radioli. Sabella analis Kröyer 1856. Abdomen with several setigers. Pygidium expanded to a large dorsal horse-shoe covering the last seven setigers. 8 species. Collar well developed. Bispira Kröyer 1856. Demonax Kinberg 1867b. FABRICINAE. thoracic neuropodial companion setae absent. external stylodes. (31a). webbing and eyes present. 2 species. Amphicora mediterránea Ley dig 1851. 3 species. . only species. webbing absent. Approximately eight thoracic setigers present. 30 species. Distylidia Hartman l%\. Radióles in semi-circles. triangular ventral projection present at base of tentacular crown. Numbers of radióles variable. Branchiomma Kölliker 1858. radiolar backs rounded. Three pairs of radioli. but sharply bent. FABRICINAE. Euchone Malmgren 1866. Abdominal uncini avicular. quadrangular base and crested tip. SABELLINAE. magna Fauchald 1972. E. crested. abdominal uncini long-handled. FABRICINAE. Both thoracic and abdominal uncini long-handled. Radióles in spirals. Collar bilobed. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Thoracic uncini sharply bent.or four-lobed. Collar well developed. eyes and webbing absent. Collar bilobed. Collar well developed. D. Amphitrite bombyx Dalyell 1853. late thoracic notopodia with iancent-shaped. ventrally higher. FABRICINAE. Abdominal uncini avicular. More than five. Abdomen with four to 12 setigers. Thoracic spatulate setae absent. (31b). external stylodes and webbing absent. external stylodes absent. Number of thoracic segments variable. Abdomen with numerous setigers. FABRICINAE. sometimes in semicircles. Radioli in spiral. leucaspis Kinberg 1867b. transversely striated setae. infundibuliformis Kröyer 1856. Radióles spiralled. Collar well developed on both sides. Spatulate thoracic notosetae present. only species. quadrangular bases and teeth in two rows along one side. Sabella vancouveri Kinberg 1867b. ornata Banse 1957. Collar two. Spatulate thoracic noto setae absent. eyes present. SABELLINAE. Radioli in semi-circles. webbing and eyes absent. Abdominal uncini with short. SABELLINAE. SABELLINAE. Dialychone Claparède 1870a. acústica Claparède 1870a. Laonome salmacidis Claparède 1870a. 21 species. only species. quadrangular base and teeth in two or more rows along one side. Abdominal uncini avicular. (31a). Amphitrite volutacornis Montagu 1804. usually many pairs of radioli present. stylodes and webbing absent. Forked ventral vascularized filament present. eyes present. Abdominal uncini with short. Euchonella Fauchald 1972. Eight thoracic and three abdominal setigers. Eudistylia Bush 1904a. highest ventrally. SABELLINAE. Three pairs of radioli. Collar divided dorsally and unhed ventrally. webbing and external stylodes absent. Last abdominal setigers flanged laterally to form a deep spoon-shaped cavity. but very sharply and distinctly bent and distally crested. distinct webbing present between the radioU. Abdomen with several setigers. webbing absent. Collar absent. webbing and external stylodes absent. Desdemona Banse 1957. eyes and external stylodes present. eyes present. abdominal uncini strongly bent. Augeneriella Banse 1957. long. 3 species. webbing present.Disrylia rugosa Moore 1904b. 26 sf)ecies. SABELLINAE. Eight thoracic segments Chone Kröyer 1856. hummelincki Banse 1958. Spatulate notosetae and neuropodial companion setae present in thorax. Euratella Chamberlin 1919c.^. Radicles spiralled. Both thoracic and abdominal uncini with long handles. Abdominal uncini with short. D. 4 species. FABRICINAE. eyes present. collar dorsally low. Collar reduced. Radióles sometimes spiralled. Abdomen with numerous setigers. webbing absent. (3Ib). Collar usually well developed. D. SABELLINAE. 3 species. Anterior ventrum irregularly rugose.or short-handled Abdominal uncini long-handied and nearly straight Abdominal uncini short-handled and strongly bent Thoracic uncini long-handied Thoracic uncini short-handled Science Series 28 33 Potaspina Potamethus Panousea Potamilla Generic Definitions Ampkiglena Claparède 1864. 2 species. branchial heart present.¡38 31b 32a 32b 33a 33b (30b). C. Collar fourlobed. 2 species. Abdominal uncini avicular. eyes and external stylodes absent. abdominal uncini with short. only species. in nearly complete cinctures around the posterior part of the body. Myxicola Koch in Renier 1847. straight or gently curved row. Collar two. Radióles in semi-circles. Amphitrite vesiculosa Montagu 1815. eyes present. 30 species. thoracic neuropodial companion setae absent. Monroika Hartman \')5\h. Megalomma Johansson 1926. SABELLINAE. Radióles in semi-circles. Fabricia Blainville 1828. abdominal uncini long-handled. webbing and eyes absent. J. SABELLINAE. nearly straight handles. webbing. united ventrally. Radióles in semi-circles. neuropodial companion setae present. abdominal uncini short-stemmed. webbing and eyes absent. Abdominal uncini avicular. 22 species. SABELLINAE. Both thoracic and abdominal uncini avicular. Both thoracic and abdominal uncini with short. M. abdomen with three setigers. Thoracic uncini short-handled. Abdomen with long-handled uncini. Collar divided dorsally and fused ventrally. Vascularized filaments absent. Abdomen with four or more setigers. Potamilla malmgreni Hansen 1878. except ventrally where it is produced into a pair of triangular lobes. dorsally incised. Radióles in semi-circles. Collar very low. eyes usually present. 6 species. Companion setae present. FABRICINAE. vasculosa Hartman 1969. Spatulate thoracic notosetae present. Jasmineira Langerhans 1880. only species. 13 species. but deeply and widely separated dorsally. abdominal uncini long-handled. compound eyes present subdistally on a few dorsal radioli. external stylodes and webbing absent. external stylodes and webbing absent. SABELLINAE. SABELLINAE. Three abdominal setigers. in part united by a web. Three pairs of radióles and branchial hearts present. Thoracic uncini long-shafted. Three pairs of radioli and branchial hearts present. 5 species. Radióles in semi-circles. Anterior thoracic neuropodia with avicular uncini. Two abdominal setigers. FABRICINAE. pacifica Hartman 1969. SABELLINAE. Laonome Malmgren 1866. external stylodes and eyes absent. 18 species. webbed. Collar well developed. Eight or more pairs of radioli. Tube mucoid. Radióles in semi-circles. kroeyeri Malmgren 1866. only species. external stylodes and webbing absent. quadrangular handles and teeth in several rows along one margin. flattened base and avicular head. Two pairs of radioli. Abdominal uncini avicular.or four-lobed. Panousea Rullier and Amoureux 1970. ventrally in addition a pair of palplike. eyes. Radióles in semi-circles. 4 species. Manayunkia Leidy 1859. Three to five pairs of radióles. eyes and external stylodes absent. 12 species. Numerous pairs of radióles present. Collar well developed. FABRICINAE. Radióles in semi-circles. Fabrkiola Friedrich 1939. caudata Langerhans 1880. Radicles in semi-circles. with short. Sabella phaeotaenia Schmarda 1861. Six pairs of radióles.1976 THE POLYCHAETE WORMS li9 Spamlate thoracic notosetae and neuropodial thoracic companion setae absent. Fabricia armandi Claparède 1864. Potamethus Chamberlin 1919c. Thoracic uncini with long. Collar ventrally high. FABRICINAE. Collar bilobed. several abdominal setigers present. 10 species. P. Collar well developed. Collar two. Very large. L. strongly bent handles. Terebella inßndibulum Renier 1804. Potamilla Malmgren 1866. Amphicora sabella Ehrenberg 1837. high and bifid ventrally. Collar four-lobed. Hypsicomus Grube 1870b. abdominal ones strongly bent. Potaspina Hartman 1969. Collar high. radióles externally flanged. Abdominal uncini avicular. otherwise absent. eyes. Webbing. FABRICINAE. Thoracic uncini minute and long-handled. SABELLINAE. and external stylodes absent. Sabella neglecta Sars 1850. only species. Three abdominal setigers. abdomen with numerous setigers. Abdominal uncini short-handled. Spatulate thoracic notosetae present. Fabrisabella Hartman 1969. Spatulate thoracic notosetae present. vascularized smooth filaments. eyes present. Spatulate thoracic notosetae present. FABRICINAE. external stylodes and webbing absent. Collar distinct dorsally. Manayunkia africana Monro 1939b. MYXICOLINAE. F. Collar low dorsally. Manayunkia pacifica Annenkova 1934. first notosetae in a long. last thoracic neuropodia with . speciosa Leidy 1859. Thoracic uncini long-handled and gently curved.or four-lobed. thoracic neuropodial companion setae absent. africana Rullier and Amoureux 1970. P. 10 species. strongly webbed. Spatulate thoracic notosetae present. Webbing absent. Oriopsis Caullery and Mesnil 1896. FABRICINAE. external stylodes and webbing absent. Collar often very low dorsally and united ventrally. 35 species. Jones (1974a) revised the original material of Monroika and pointed out that the character used to separate this genus from all other FABRICIIN genera is incorrect. see Hypsicomus Arippasa Johnston 1865. see Oriopsis Oriades Chamberlin 1919c. Spatulate thoracic notosetae absent. Three abdominal setigers. asiática). with one half very much larger than the other. seeBispira Dybowscella Nusbaum 1901. only species. webbing absent. see Branchiomma Distylia Quatrefages 1866. see Potamethus Gymnosoma Quatrefages 1866. see Potamilla Branchiomma Claparède 1870a. Eurato sanctijosephi Gravier 1906. Sabella Linnaeus 1767. see Fabriciola Garjaiowella Dybowskii 1929. Radióles in semi-circles. Abdominal uncini with short base and several rows of teeth. Schizobranchia Bush 1904a. SABELLINAE. questionably Fabricia Parachonia Kinberg 1867b. Radióles spiralled. irrespective of the structure of the uncini. in that members of the FABRICINAE have gently curved. Invalid Genera Amphicora Ehrenberg 1837. acicular. It differs from the latter in that it has short limbate setae in both the first setigers. Members of this genus will key out with Manayunkia above. see Manayunkia Gorbunovia Annenkova 1952. eyes absent. Taxonomic Notes The sub-families as accepted here are based on the structure of the neuropodial uncini of the thorax. see Euchone Dasycharte Sars 1862. Pseudofabricia Cantone 1972. emersoni Jones 1962. The genus Pseudopotamilla Bush is considered here a junior synonym of Potamilla and the genus Trichosobranchella Dybowski 1929. companion setae absent. Spatulate setae and companion setae present. This genus could not be included in the key. avicular uncini. all sabellids with thoracic neuropodial companion setae have been included in the SABELLINAE. see Demonax Potamis Ehlers 1887. is referred to Manayunkiu. Spatulate and companion setae absent. Radicles in semi-circles. Radióles spiralled. Sabellastarte Savigny 1818. 4 species. see Myxicola Aspeira Bush 1904a. see Chone Parasabella Bush 1904a. external stylodes and webbing absent. see Oriopsis Othonia Johnston 1835. distally bifid anterior end and an extended rounded. S. SABELLINAE. eyes present. see Myxicola Leptochone Claparède 1870b. seeMegalomma Chaponella Rullier 1972. S.140 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 thick. see Chone Metachone Bush 1904a. see Hypsicomus Fabriciella Zenkevitch 1935. see Myxicola Eurato Saint-Joseph 1894. external stylodes present. see Potamilla Sabellina Dujardin 1839a. Collar well developed. see Hypsicomus Laonomedes Chamberlin 1919c. Spirographis Viviani 1805. since it was based on dried material. SABELLINAE. webbing and external stylodes absent. indeterminable . companion setae present. Eyes present. and members of SABELLINAE have strongly bent. Monroika has two rather than three abdominal setigers. However. FABRICINAE. Jones (1974a) also described a new genus. see Manayunkia Eriographis Grube 1850. see Fabricia Amphicorina Quatrefages 1866. Collar four-lobed. see Hypsicomus Oria Quatrefages 1866. long-handled uncini. Furthermore. Collar four-lobed. seeBispira Notaulax Tauber 1879. widely separated dorsally. P. It differs from other genera in this complex in that it has pilose setae in the last three thoracic setigers. see Myxicola Megachone Johnson 1901. Manayunkia has such setae only in the first setiger and they are much longer. see Hypsicomus Pseudopotamilla Bush i904a. penicillus Linnaeus 1767. spallanzani Viviani 1805. only species. indeterminable Sabina Williams 1851. acicular spines. Pseudobranchiomma Jones 1962. All uncini avicular. Stylodes and webbing absent. dichotomously divided. external stylodes and webbing absent. see Potamethus Protulides Webster 1884. Collar four-lobed. Abdominal uncini small. eyes present. see Hypsicomus Hypsicomatopsis Augener 1922. Thoracic uncini long-handled. only one part spiralled. 2 species. and no information was available about the structure of the tentacular crown. Brandtika (genotype: B. see Chone Metalaonome Bush 1904a. insignis Bush 1904a. see Oriopsis Oridia Rioja 1917. Thoracic uncini short-handled. but small. lower lip. SABELLINAE. aberrans Cantone 1972. see Oriopsis Anamobaea Kröyer 1856. see Myxicola Haplobranchus Bourne 1883. see Branchiomma Dasychonopsis Bush 1904a. eyes present. see Potamilla Leiobranchus Quatrefages 1850. P. 5. Radióles in semi-circles. Eight radióles on a greatly prolonged. SABELLINAE. see Manayunkia Hypsicomatides Augener 1922. it does not have the webbed tentacular crown claimed by Hartman. 8 species. Four thoracic setigers. but bent. which will key out in the same complex. Setae include long-handled uncini with companion setae and giant falcate spines. The family consists of a single genus. 6b (3b).¡976 THE POLYCHAETE WORMS 141 Sitophaga Gistel 1848. Sabellonga disjuncta Hartman 1969 from northern Baja California. Key to Genera la. 10a (8b). lb. 5b (4b). 8a (7a). Otherwise. and may. 7a (lb). Tube calcareous. 4b (3a). The older taxonomic groupings placed emphasis on the structure of the operculum and on the overall body-construction (development of the thoracic membrane and collar. The serpulids and the closely allied spirorbids represent a sub-specialty of their own within the polychaetes. etc. Caobangia Giard 1893. 6a (3b). or that such has ever been present. with type-species C. Digestive tract U-shaped with the anus opening dorsally and far anteriorly on the body. billeti Giard 1893 and a total of seven known species.). nearly straight capillaries Protis Operculum carried on a branchial radióle 8 Operculum carried on a modified stalk 12 Five thoracic setigers present 9 Six or more thoracic setigers present 10 Collar setae slender and limbate. 1969 The family is known for a single genus and species. The family has been reviewed on a couple of occasions. see Manayunkia FAMILY CAOBANGIIDAE JONES 1974b Small. some authors (especially Zibrowius) have placed more emphasis on the detailed structure of the uncini and setae. Body separated into two regions. there is no trace of the loss of a tentacular crown. The specimen resembles a sabellid that has lost its tentacular crown. see Fabricia Trichosobranchella Dybowski 1929. 3b (2b). Most of the forms burrow in the shell of the host. but the demonstration of a series of forms spread over a larger geographical region. 4a (3a). the setal equipment is largely what one would expect in a member of the SABELLINAE. More recently. presence or absence of setae in specified segments. Saint-Joseph (1894) published an extensive review and Southward (1963) gave a key to all known genera. a thorax with a thoracic membrane (absent in rare instances) and dorsal capillary or limbate setae and an abdomen with ventral setae and dorsal uncini. 9a (8a). be difficult to use. This fainily was recently proposed by Jones (1974b) for a series of small polychaetes that live in close association with molluscs in freshwater in Southeast Asia. The presence of the giant falcate spines in far posterior setigers also is characteristic. the genus has been considered among sabellid enigmaticae. 2b (la). The key below represents a compromise. 3a (2b). However. made the recognition of a new family necessary. 2a (la). as such. 7b (lb). Peristomium forms a bevelled collar around the prostomium (tentacular crown absent). two kinds of avicular hooks present in a posterior region. FAMILY SERPULIDAE JOHNSTON 1865 FAMILY SABELLONGIDAE HARTMAN Body cylindrical with few thoracic and many abdominal segments. 5a (4b). short-bodied sabelüform polychaetes with three pairs of radioli in a tentacular crown. The generic sub-divisions within the family are nevertheless debatable. First setiger with neurosetal palmate hooks. 8b (7a). 9b (8a). Up to this time. limbate or capillary 3 Slender capillary or limbate collar setae present 4 Collar setae modified 6 Thoracic membrane absent Salmacinopsis Thoracic membrane present 5 Thoracic membrane reaches setiger 5 Subprotula Thoracic membrane reaches setiger 7 Salmacina Abdominal setae geniculate Salmacina Abdominal setae slender. Operculum absent 2 Operculum present 7 Abdominal setae trumpet-shaped Pseudoserpula Abdominal setae marginally dentate and geniculate. thorax and abdomen not separated by a long asetose region Dipomatus Collar setae simple and tapered limbate Apomatus . remainder of the thoracic setigers without hooks. long asetose region present between thorax and abdomen Josephella Collar setae basally dentate. One radióle often transformed into an operculum. 32a (31b). 21a (20b).. 17b (13a). 14b (13a). Filogranula Operculum a depressed cone. 29a (28a). 20b (19a). 21b (20b). with basal pilose boss and distal spines Spirobranchus Thoracic neuropodia equidistant in all setigers. 33b (26b). 16b (15b). collar setae bayonet-shaped Temporaria Uncini with anterior peg pointed Omphalopomopsis Uncini with anterior peg gouge-shaped (hollowed out from beneath) Pomatostegus Collar setae absent 27 Collar setae present 33 Abdominal setae absent. I2b (7b). 29b (28a). 12a (7b). 30b (28b). either capillary or with a smooth double-boss basally and smooth tapering tips Crucígera Collar setae absent 18 Collar setae present 19 Operculum with two projections Olga Operculum without projections Pomatoleios Collar setae limbate or capillary 20 Collar setae with two separate limbations (bayonet-type). 27a (26a). 25a (23b). operculum funnel-shaped Scierostyla (in part) Opercular stalk not calcified. 33a (26b). 26a (13b). 34b (33b). uncini usually present 28 At least some abdominal setae geniculate 29 All abdominal setae slender capillaries 30 Uncini with numerous teeth Placostegus Uncini with seven to nine teeth Marifugia Abdominal uncini absent Bonhourella Abdominal uncini present 31 Tube free. 32b (31b).. immovable 34 Abdominal setae slender capillaries Schiiocraspedon Abdominal setae trumpet-shaped or geniculate 35 Abdominal setae trumpet-shaped 36 Abdominal setae geniculate 39 Operculum a simple funnel 37 . 25b (23b). 20a (19a). 14a (13a). 31b (30b). 31a (30b). collar setae all of one kind. 28b (27b). 36a (35a).142 lOb (8b). 17a (13a). bordered by two eccentrically placed thorns connected by a low ridge Pseudopomatoceros At least anterior abdominal setae trumpet-shaped 24 Anterior abdominal setae geniculate 25 Operculum conical. 28a (27b). operculum spherical Dasynema Opercular spines movable Galeolaria Opercular spines. 13a ( 12b). 1 lb (lOb). 15b (14a). if present. seven thoracic setigers present 16 Thoracic neuropodia widely separated anteriorly and approaching posteriorly. 35a (34b). abdominal uncini present Rhodopsis Abdominal setae present. 26b (13b). 16a (I5b). 19a (17b). with a basal boss or setose 23 Anterior abdominal setae stout. acute spines Paumotella Anterior abdominal setae trumpet-shaped or geniculate 21 Opercular cap black and chitinous Crosslandiella Opercular cap calcareous 22 Opercular cap flat. 19b (17b). 22a (21b). collar setae basally pilose Conopomatus Operculum with slanting distal plate. 35b (34b). collar setae of two kinds. 24a (23a). I la (10b). NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Collar setae with a few coarse teeth basally and a Hmbate denticulate blade distally II Operculum a shallowly depressed cone on a spherical swelling of the opercular stalk . 30a (28b). 22b (21b). stalk without the subdistal swelling Filograna Opercular stalk flattened and ribbonlike Mesavermilia Opercular stalk oval or circular in cross-section (sometimes with wings or spines) 13 Opercular stalk with wings or spines 14 Opercular stalk without wings or spines 26 Opercular stalk with spines 15 Opercular stalk with wings (sometimes produced into points distally) 17 Opercular stalk with spines on one side only. 24b (23a). 23b ( 19b). 18a (17a). 27b (26a). 13b (I2b). 18b (17a). with or without spines Pomatoceros Operculum distally excavated. six thoracic setigers present Spirodiscus Opercular stalk with four spines in a cross. 34a (33b). 15a (14a). 23a (19b). tusk-shaped and smooth Ditrupa Tube at least partly attached 32 Opercular stalk calcified. 43a (41b). Collar setae simple limbates. 8 species. 38a (36b). (52b). 45b (40b). abdomjnal setae with only a short geniculate tip Sclerostyla (in part) Opercular stalk not calcified. nearly radial in structure Neopomatus Operculum with a single series of long. 37b (36a). dewae Straughan 1976a. 48a (45b). (51a). abdominal setae capillary. 52a 52b 53a 53b 54a 54b (51a). Uncini with teeth in several rows in both thorax and abdomen. Mercierella Collar setae with teeth in two rows Sphaeropomatus Collar setae with teeth in a single row 44 Operculum distally rounded Ficopomatus Operculum distally excavate Mercierellopsis At least some collar setae with a basal dentate or hirsute region 46 All collar setae limbate 48 Simple limbate collar setae present. 46a (45a). calcareous plate. 50a (49b). (53b).. (52b). 44b (43b). insignis Gravier 1905b. C. 38b (36b). 41b (40a). 47b (46b). sometimes. operculum conical without wings or spines on the stalk. Calcareopomatus Straughan 1967a. only species. Hydroides norvegica groenlandica Mörch 1863. 42a (41a). 45a (40b).) absent 40 Side setae present 49 Collar setae with few coarse teeth in one or two marginal rows 41 Collar setae at least in part limbate 45 Operculum distally ornamented 42 Operculum distally smooth. 42b (41a). Collar . rounded or excavate 43 Operculum with concentric series of teeth. opercular stalk distinct. 40a (39a). 43b (41 b). operculum flat or lightly convex. Vermiliopsis Thoracic uncini with teeth in a single row. 47a (46b). 51a (50a).. only species. in addition to some with a basal group of spines Omphalopoma All collar setae with basal denticulated or hirsute region 47 Six thoracic setigers present Hyalopomatus Seven thoracic setigers present Pseudochitinopoma Opercular stalk calcified. Seven thoracic setigers.. 37a (36a). 48b (45b). glandular hood-shaped distal part Olgaharmania Operculum with series of distal spines and.. smaller hoods Hydroides Side setae {Apomatus-ssX?is. ampulliferus Philippi 1844. side setae present. strong teeth. anterior peg very long. Collar setae absent. abdominal uncini absent. globular operculum on a radióle. anterior peg furcate. making it bilaterally symmetrical . side setae present. operculum with a globular base and a flat. 44a (43b). Six thoracic setigers. A. 49b (39b). B. 2 species. 39b (35b). Collar setae small and limbate. 51b (50a). Thoracic uncini with anterior peg bifid. 38 Collar setae basalty minutely hirsute Paraserpula Collar setae basally dentate Serpula Operculum with a large. SERPULINAE. 40b (39a). Seven thoracic setigers. Seven thoracic setigers. THE POLYCHAETE WORMS ¡43 Operculum with a basal funnel and in addition various spines or hoods forming a distal part ./97(5 36b (35a). abdominal setae with about one-half of the exposed setae in the geniculate tip Neovermilia Opercular stalk annulated Calcareopomatus Opercular stalk not annulated 50 Collar setae limbate 51 Collar setae with a basal boss in addition to the limbate distal portion 52 Both thoracic and abdominal uncini with all teeth in a single row. Chitinopoma Levinsen 1884. 46b (45a). SERPULINAE. 49a (39b). Bonhourella Gravier 1905b. stalk annulated. 41a (40a). anterior peg entire . Thoracic uncini with anterior peg gougeshaped. (53b). SERPULINAE. 39a (35b). SERPULINAE. abdominal uncini with teeth in several rows Pseudovermilia Both thoracic and abdominal uncini with teeth in several rows Filogranula (in part) Thoracic uncini with teeth in a single row 53 Operculum distally rounded Chitinopomoides Operculum distally excavate 54 Opercular stalk with three distal bulbs Janita Opercular stalk without swellings Chitinopoma Generic Définitions Apomaius Philippi 1844. 50b (49b). Seven thoracic setigers. operculum with black. only species. SERPULINAE. Collar setae slender . wilsoni Benham 1927. C. FICOPOMATINAE. Janita Saint-Joseph 1894. SERPULINAE. SERPULINAE. other thoracic setae limbate. all thoracic setae limbate. pilose at base. stalk smooth. Collar setae present. only species. two operculae with branchial filaments on peduncle. SERPULINAE. 5 species. Collar setae notched with limbate expansion at the base. Abdominal setae geniculate. anterior peg furcate. stalk without spines or wings. rounded.144 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 setae of fin and blade construction. operculum spherical. C. Uncini with teeth in several rows. SERPULINAE. side setae absent. abdominal setae trumpet-shaped. Conopomatus Pillari 1960. Six to 12 thoracic setigers. Abdominal setae with only the distalmost tip geniculate. only species. soft or chitinous. operculum pear-shaped. only four radicles present. Serpula fimbriata delle Chiaje 1828. operculum bladder-shaped. 85 species. Seven thoracic setigers. SERPULINAE. F. Josephella Caullery and Mesnil 1896. Seven thoracic setigers. operculum infundibular with three large distal bulbs on the stalk. distal funnel present. stalk without spines or wings. abdominal setae capillaries. Crucígera Benedict 1887. Collar setae present and very short and slender. Collar setae limbate. norvegica Gunnerus 1768. uncini with numerous teeth in several rows. H. FILOGRANINAE. distal end of stalk with four large spines forming a cross. Collar setae absent. side setae present. Uncini with few teeth. or a second. Crosslandiella Monro 1933. only species. 2 species. FILOGRANINAE. only species. Side setae absent. abdominal ones with teeth in several rows. abdomen with geniculate setae. others serrated limbates. Six thoracic setigers. acuiconus Pillai 1960. Uncini with few teeth in a single row. opercular plate chitinous with a single large spine. Hyalopomatus Marenzeller 1878. side setae present. rounded operculum on one of the radióles. gracilis Langerhans 1884. C. Galeolaria Savigny 1818. operculum conical. 2 species. opercular stalk smooth.F. operculum with simple funnel. Seven thoracic setigers. SERPULINAE. implexa Berkeley 1828. D. F. SERPULINAE. Dipomatus Ehlers 1913. operculum at end of normal radióle. tusk-shaped. multispinosa Monro 1933. side setae present. Abdominal setae geniculate with a dentate. Müller 1776. stalk either frondose or smooth. H. Filogranula Langerhans 1884. C. abdominal setae geniculate with long triangular blade. side setae present. SERPULINAE. abdominal setae geniculate with triangular blade. spherical base. marenzelleri Caullery and Mesnil 1896. SERPULINAE. Dentalium arietinum O. Collar setae of fin and blade construction. Some collar setae with dentate boss. thoracic setae limbate and capillaries. Thoracic uncini with teeth in a single row. 8 species. abdominal ones with teeth in several rows. Collar setae limbate. side setae present. abdominal ones capillaries. chitinous plate. 2 species. Uncini with numerous teeth in several rows. Chitinopomoides Benham 1927. anterior peg very large. Collar setae lancet-shaped and tapering. Hydroides Gunnerus 1768. Collar-setae basally dentate. Seven thoracic setigers. operculum with calcareous plate surmounted by movable spines. 5 species. Seven thoracic setigers. Ficopomatus Southern 1921. distally with a crown of spines or smaller hoods. Seven thoracic setigers. side setae present. stalk smooth. abdominal setae trumpet-shaped. side setae absent. macrodon Southern I92I. abdominal setae trumpet-shaped. each distally funnel-shaped and marginally dentate. Serpula chrysogyrus Grube 1878. Seven thoracic setigers. Dasynema Saint-Joseph 1894. Five thoracic setigers. claparedii Marenzeller 1878. Five thoracic setigers. Thoracic uncini with teeth in a single row. J. side setae absent. opercular stalk winged. SERPULINAE. operculum basally with a marginally dentate funnel. operculum a small cone on a small. Uncini with numerous teeth in several rows. Collar setae basally dentate. serpulides Ehlers 1913. G. Side setae and limbate setae present in thorax. 2 species. F. surmounted by a spinose column. websteri Benedict 1887. 4 species. Seven thoracic setigers. Ditrupa Berkeley 1835. slender tip. Collar setae with large boss below a limbate zone. Collar setae present. caespitosa Savigny 1818. Tube free. stalk winged. abdominal setae geniculate. 3 species. stalk winged. operculum an inverted cone with a chitinous plate. Seven thoracic setigers. Filograna Oken 1815. operculum rounded. Collar setae and side setae absent. SERPULINAE. Six thoracic setigers. Thoracic uncini with teeth in a single row. distally cup-shaped. uncini with about twenty teeth. operculum bladder-shaped. Side setae present. anterior abdominal setae stout acute spines. abdominal ones with teeth in several rows. stalk without spines or wings. Seven thoracic setigers. 2 species. stalk smooth. only species. partially in two rows in abdomen. M. FICOPOMATINAE. Collar setae marginally dentate. Thoracic uncini with teeth in a single row. operculum funnel-shaped. Side setae present. thoracic setae slender and straight. uncini with teeth in one row in thorax. only species. Uncini with few teeth (7-12). nearly radial in structure. Six thoracic setigers. Marifugia Absalon and Hrabe 1930. SERPULINAE. Paumotella Chamberlin 1919c. SERPULINAE. prietoi Rioja 1945. operculuni conical and distally smooth. umbilicata Mörch 1863. operculum unknown. collar setae limbate. 4 species. Merciereliopsis Rioja 1945. Seven thoracic setigers. stalk smooth. FICOPOMATINAE. Six or seven thoracic setigers. Uncini with few teeth. Abdominal setae geniculate. Omphalopoma Morch 1863. Operculum distally smooth. operculum with a central tooth and paired lateral horns. Membranopsis Bush 1910. 18 species. Collar setae of two kinds. Collar setae dentate. Seven thoracic setigers present. stalk smooth. Seven thoracic setigers. elegantissima Jones 1962. SERPULINAE. side setae absent. FICOPOMATINAE. M. takemoana Chamberlin 1919c. only species. Collar setae absent. stalk flattened. abdominal setae geniculate. abdominal setae trumpet-shaped. Olga Jones 1962. Seven thoracic setigers. only species. Thoracic membrane fused over the back at ninth seliger. abdominal setae geniculate with a triangular blade. . Collar setae basally with paired bosses. enigmática Fauvel 1923b. stalk smooth. only species. slender capillaries and limbate with a pilose subdistal boss. marginally dentate. abdominal setae geniculate with widely triangular dentate distal plates. posterior ones fine capillaries. uschakovi Pillai 1960. Collar setae limbate. P. Collar setae and side setae absent. 2 species. operculum distally oblique with chitinous plate covered by series of curved spines. SERPULINAE. SERPULINAE. Paraserpula Southward 1963. Serputa tridentala Fabricius 1780. Collar setae dentate. anterior peg deeply bifid. side setae absent. Hydroides glandiferum Rioja 1941a. Nine thoracic setigers. abdominal ones with teeth in several rows. operculum smooth. N. Seven thoracic setigers. Collar setae absent. Collar setae bayonet-shaped and finely pilose. anterior peg very large. Uncini with numerous teeth. side setae present. O. inconspicua Bush 1910. Seven thoracic setigers. abdominal ones with teeth in one or several rows. side setae absent. side setae absent. Thoracic uncini with few teeth in a single row. stalk smooth. maximally nine teeth present in thoracic uncini. Vermilia multicristata Philippi 1844. cavatica Absalon and Hrabe 1930. Olgaharmania Rioja 1941b. AÍ. operculum conical and distally flattened. Placostegus Philippi 1844. finely dentate capillaries. Seven thoracic setigers. side setae absent. Omphalopoma langerhansii Marenzeller 1884.1976 THE POLYCHAETE WORMS 145 limbate. stalk smooth. P. Omphalopomopsis Saint-Joseph 1894. Uncini similar to those m Protula. 2 species. only species. thoracic setae limbate. SERPULINAE. Seven thoracic setigers. Seven thoracic setigers. anterior peg furcate. SERPULINAE. simple limbates and fin and blade setae. operculum cup-shaped. side setae absent. Thoracic uncini with teeth in a single row. Collar setae of two kinds. Neopomatus Pillai 1960. side setae absent. operculum with series of concentric rows of teeth. Neovermilia Day 1961. planorbis Southward 1963. N. side setae absent. SERPULINAE. Collar setae limbate. operculum spherical or slightly cup-shaped. SERPULINAE. anterior peg entire. uncini with double rows of teeth. abdominal setae genicdate or capillaries. stalk winged. anterior peg undercut. Uncini with teeth in a single row. M. anterior peg gouge-shaped. only species. anterior peg large. sometimes ornamented. only species. stalk with wings. ribbon-shaped. margin entire. 2 species. SERPULINAE. opercular stalk with wings. Uncini with numerous teeth. operculum with basal corona of spines surmounted by very large. SERPULINAE. capensis Day 1961. Thoracic uncini with teeth in a single row. O. glandular chitinous hood equipped on one side with four large spines. Collar setae limbate. Mercierelia Fauvel 1923b. Metavermilia Bush 1904a. of)erculum funnel-shaped with chitinous distal plate. Uncini with numerous very fine teeth. Collar setae simple limbates. operculum absent. Five to nine thoracic setigers. operculum on a modified radióle. incrustans Claparède 1870a. blunt. Thoracic membrane absent. Schizocraspedon Bush 1904a. stalk with broad wings. SERPULINAE. FILOGRANINAE. Uncini with teeth in several rows. P. Pseudochitinopoma Zibrowius 1969. Seven thoracic setigers. operculum distally cup-shaped with two eccentrically placed thorns connected by a low ridge. abdominal setae with flat crown and a long spike. Uncini with numerous teeth. abdominal setae nearly capillary. 23 species. Thoracic uncini with teeth in a single row. 3 species. abdominal ones with teeth in several rows. Spirohranchus occidenlalis Mclntosh 1885. 13 species. thoracic setae limbate. anterior peg large. Seven thoracic setigers. shaft smooth. P. Pseudopomatoceros Holly 1936. operculum funnel-shaped. stalk winged. abdominal setae geniculate. simplex Ehlers 1887. Pseudoserpula Straughan 1967b. SERPULINAE. collar setae present or absent. only species. Seven thoracic setigers. Collar setae absent. Seven thoracic setigers. Collar setae and side setae absent. if present finely limbate. abdominal setae trumpet-shaped with one long spine at one side. SERPULINAE. Thoracic uncini with teeth in a single row. Seven thoracic setigers. with very short geniculate tips. setosa Bush 1910. SERPULINAE. stalk without wings. SERPULINAE. SERPULINAE. Seven thoracic setigers. Uncini with teeth in a single row. only species. Protis Ehlers 1887. Salmacinopsis Bush 1910. Protula Risso 1826. Uncini with numerous teeth. operculum a flat calcareous plate. opercular stalk smooth. Uncini with few teeth. Thoracic uncini with teeth . Pomatostegus Schmarda 1861. side setae present. operculum absent. Collar setae small and limbate. other thoracic setae capillary and limbate with a few serrate setae in posterior thoracic setigers. R. 8 species. 2 species. SERPULINAE. SERPULINAE. FILOGRANINAE. Side setae present. Side setae absent. side setae present. SERPULINAE. Salrrmcina Claparède 1870a. S. Abdominal setae capillary. anterior peg furcate. Serpula triquetra Linnaeus 1767. Pseudovermilia Bush 1907b. anterior peg very long. side setae present. P. ctenaclis Mörch 1863. side setae absent. side setae absent. abdominal setae geniculate and serrated. macrosoma Schmarda 1861. Side setae absent. abdominal setae geniculate with narrow blade. abdominal ones with teeth in several rows. Collar setae fine capillaries. slender processes. SERPULINAE. abdominal setae absent. operculum absent. stalk winged. only species. Seven thoracic setigers. rugosa Straughan 1967b. 5. Serpula tubularia Montagu 1803 in Mclntosh 1923. superior thoracic setae geniculate with conspicuous spines at the base. Rhodopsis Bush 1904a. Uncini with about ten teeth in both thorax and abdomen. Sclerostyla Mörch 1863. operculum covered distally with chitinous plate with spines arranged in a rosette. operculum with chitinous plate surmounted by a column with several discs. 2 species. abdominal setae nearly capillary. Nine thoracic setigers. Collar setae limbate. Seven thoracic setigers. stalk winged. P. with or without spines. Seven thoracic setigers. Serpula furcifera Grube 1878. Pomatoleios Pixell i913b. one inside the other. operculum bladder-shaped. distally with a chitinous cover consisting of two concentric funnels. Collar setae of fin and blade construction with basal dentate fm well separated from the blade. operculuin absent. anterior peg furcate. Collar setae with subdistal pilose boss.146 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Pomatoceros Philippi 1844. abdominal setae trumpet-shaped. Pomatoceropsis roxasi Holly 1935. crosslandi Pixell 1913b. Seven thoracic setigers. 8 species. abdominal setae trumpet-shaped. 5 species. SERPULINAE. SERPULINAE. Collar setae notched with limbate subdistal and distal parts. 3 species. Uncini with teeth in several rows. operculum absent. calcified. thoracic setae limbate. operculum distally ornamented. anterior peg furcate. both marginally split with long. Thoracic uncini with teeth in a single row. pusillus Bush 1904a. Collar setae limbate. abdominal setae geniculate with triangular blades. abdominal uncini with teeth in several rows. Seven thoracic setigers. stalk smooth. Abdominal setae geniculate. anterior peg large and blunt. Hyalopomatopsis occidentatis Bush 1904a. Collar setae capillary. Collar setae with fin and blade construction and little separation between the basal fin and the distal blade. operculum distally flat. only species. side setae absent. anterior peg simple. Seven thoracic se tigers. Uncini with teeth in a single row. side setae present. S. Collar setae present. only species. indeterminable Placostegopsis Saint-Joseph 1894. Uncini with about 15 teeth. SERPULINAE. Body asymmetrical and tube coiled in a spiral. vermicularis Linnaeus 1767. SERPULINAE. see Vermiliopsis Piratesa Templeton 1835. see Pseudopomatoceros Proplacostegus Bush 1904a. tube calcareous. Collar and side setae present. opercular stalk smooth. only species. Vermilia poiytrema Philippi 1844. Collar setae spinose with teeth in two rows. included in the generic definitions. Seven thoracic setigers. SERPULINAE. see Hydroides Eupomatus Philippi 1844. FILOGRANINAE. Sphaeropomatus Tread well 1934. Thoracic uncini with at least ten teeth. other setae slender capillaries. operculum rounded or flattened. operculum soft and vesicular. Seven thoracic setigers. see Hyalopomatus Microserpula Dons 1931.YCHAETE WORMS 147 Taxonomic Notes in a single row. Abdominal setae trumpet-shaped. see Chitinopoma Paravermilia Bush 1907b. S.\97f> THE Poi. see Hydroides Pomatoceropsis Gravier 1905b. abdominal uncini with teeth in a single row. It may be of some value to recognize the difference between Hydroides SENSU STRICTU-dnd Eupomatus at the sub-generic level. miamiensis Treadwell 1934. anterior peg very large. Abdominal setae trumpet-shaped. has been omitted from the key. 17 species. Thoracic uncini with teeth in a single row. Seven thoracic setigers. Thoracic uncini with teeth in a single row. Collar setae bayonet-shaped and pilose. 30 species. see Hydroides Filigrana Mörch 1863. Collar setae flattened and lancet-shaped. Collar setae capillaries. side setae absent. Uncini with numerous teeth. Seven thoracic setigers. Abdominal setae trumpetshaped. Seven thoracic setigers. S. stalk with dorsal spines. Invalid Genera Apomatopsis Saint-Joseph 1894. indeterminable Spiramella Blainville in Quatrefages 1866. see Protula Vermilia Savigny 1818. One radióle transformed into an operculum. see Spirobranchus Zopyrus Kinberg 1867b. longiseta Bush 1910. separated by a notch. Knight-Jones and Vine has clarified the generic sub-division of the group considerably and is largely followed here (references to articles can be found be- . see Hyalopomatus Eucarphus Mörch 1863. Temporaria Straughan 1967b. Serpula Linnaeus 1767. four or five segments. as is customarily done by most specialists today. stalk smooth. see Placostegus Psymobranchus Philippi 1844. see Spirobranchus Pomatoceropsis Holly 1935. Recent reviews by BaileyBrock. operculum absent. a thorax with a thoracic membrane. see Placostegus Podioceros Quatrefages 1866. prior to that of course. Thoracic neuropodia are separated widely anteriorly and approaching in late thoracic setigers. some forms have limbate collar setae. s^t Apomatus Cystopomatus Gravier 1911a. •e Filograna Glossopsis Bush 1904a. see Filograna Filopora Fleming 1825. since the structure of the operculum remains unknown. Collar setae either limbate or with a dentate boss. The spirorbids are rather difficult to work with since identification requires precise and accurate work with compound microscopes. stalk winged. abdominal ones with teeth in several rows. Seven thoracic setigers. only species. Spirodiscus Fauvel 1909. The genus Hydroides has been defined here to include Eupomatus Philippi 1844. opercular stalk winged. The genus Membranopsis Bush 1910. anterior peg large. S. FICOPOMATINAE. see Serpula 1970 FAMILY SPIRORBIDAE PILLAI Body separated into two regions. operculum with slanting calcareous plate. Thoracic membrane to fifth setiger only. S. 35 species. operculum funnel-shaped on a small ampulla. Six thoracic setigers. grimaidii Fauvel 1909. dorsal setae and ventral uncini and an abdomen with ventral setae and dorsal uncini. 2 species. operculum with calcareous distal plate. Collar setae bayonet-shaped with both regions limbate. 2 species. stalk without wings. Spirobranchus Blainville \&IS. thorax with setigerous rudiments of three. anterior peg furcate. see Pomatoceros Polyphragma Quatrefages 1866. anterior peg entire. side setae absent. collar tri-lobed. Serpula gigantea Pallas 1766. SERPULINAE. the specimens must be dissected out from their tubes as completely as possible. usually ornamented. Serpula irifundibulum Linnaeus 1758. operculum funnel-shaped with crenulated margin. see Serpula Hyalopomatopsis Saint-Joseph 1894. Uncini with few teeth. SERPULINAE. Vermiliopsis Saint-Joseph 1894. see Protula Siliquaria Savigny 1818. Subprotula Bush 1910. side setae absent. see Hydroides Helena Castelnau 1842. (C). tube. FIGURE . lb. 8a (7b). Key to Genera la. 4a (3a). (A). Laeospira sp. The family is most frequently considered a subfamily under the serpulids. 5x. Pillai (1970) has been followed. 15x. Family SPIRORBIDAE. 9b (8b). natural size. Family SERPULÎDAE. blunt anterior peg Simplicaria Thoracic uncini with broadly indented anterior peg Eulaeospira Dextrally coiled 11 Sinistrally coiled 12 Collar setae of fin and blade construction Spirorbella gf**^ 38. 10a (5b).148 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 low). 4b (3a). 5a (lb). simplified after Day 1964. (B). With three or more pairs of thoracic tori 2 With two pairs of thoracic tori 5 With four or more pairs of thoracic tori or at least rudiments of a fourth pair present on the concave side 3 With three pairs of thoracic tori 17 With at least four complete thoracic tori present 4 With remnants of a fourth thoracic torus present 21 Four pairs of thoracic tori present Anomalorbis Approximately seven pairs of thoracic tori present Neomicrorbis Side setae absent 6 Side setae present 10 Margins of collar fused to form dorsal tunnel Dexiospira Margins of collar not fused 7 Dextrally coiled Pillaiospira Sinistrally coiled 8 Collar setae limbate Leodora Collar setae of fin and blade construction 9 Thoracic uncini with simple. 3b (2a). 7a (6b). 6b (5a). 10b (5b). Where the above-mentioned authors have yet to treat a genus. 9a (8b). 3a (2a). 1969. 8b (7b). Serpula vermicularis. 2a (la). 7b (6b). after Hartman. 6a (5a). Protolaeospira patagónica. 2b (la). 1 la (10a). (D). 5b (lb).. after Day 1967. tube. Spirobranchus spinosus. about 15x. I believe that this difference in approach is relatively unimportant for the time being. (22a). 19b (18b). 21b 22a 22b 23a 23b 24a 24b 25a 25 b (3b). 14b (13b). (21b). 20a (17b). (22b). (22a). Genus Ampiaría Anomalorbis Capeospira Circeis Eulaeospira Helicosiphon Janua subgenus Genus Paralaeospira Pileolaria subgenus Duplicarla Pileolaria Protoleodora Simplicaria Dextralia Protolaeospira Bushiella Romanchella Spirorbella Spirorbis Dexiospira Fauveldora Janua Leodora PüUúospira Prodexiospira Protolaeospira subgenus Romanchella subgenus Spirorbis subgenus Metalaeospira Neomicrorbis Paradexiospira . most of the tube uncoiled Helicosiphon Three thoracic tori and four fascicles of setae on the convex side. (24b). 12a (10b). (24b). as indicated in Table 3. 12b ( 10b). THE POLYCHAETE WORMS 149 Collar setae simply limbate Janua Collar setae simply limbate 13 Collar setae of fin and blade construction 15 Collar fused dorsally to form tunnel Romanchella Collar not fused dorsally 14 Incubation of larvae in opercutar chamber. four thoracic tori and five fascicles of setae on the concave side Ampiaría Fourth setiger represented by a rudiment of the torus on the concave side 22 Sinistrally coiled 23 Dextrally coiled 24 Side setae present Protolaeospira Side setae absent Capeospira Collar setae simply limbate Prodexiospira Collar setae of fin and blade construction 25 Thoracic uncini with furcate anterior peg Dextralia Thoracic uncini with bluntly rounded anterior peg Paradexiospira generic level. abdominal setae with blades as those of the collar setae Fauveldora Incubation in the tube. The key given above suggests that all named taxa are at the same level. (21b). Vine (1972) and Knight-Jones (1973) used several of the contained taxa at the sub- TABLE 3 Alphabetic list of genera and sub-genera of Spirorbidae as suggested by Vine (1972) and Knight-Jones (1973). (22b). 18a (17a).¡97(. 19a (18b). 16b (15b). 20b (17b). 15a (12b). 15b (12b). 18b (17a). lib (lOa). abdominal setae very small with short blades Buskiella Thoracic uncini with several rows of teeth Spirorbis Thoracic uncini with a single row of teeth 16 Two or more opercular plates on top of each other on the opere ulum Duplicaría A single opercular plate present Pileolaria Side setae present 18 Side setae absent 20 Collar setae of fin and blade construction Paralaeospira Collar setae simply limbate 19 Incubation of larvae in opercular chamber Protoleodora Larvae attached to stalk in faecal groove Metalaeospira Tube tightly coiled along its whole length Circeis Tube coiled only basally. 13a (12a). 16a (15b). 17b (2b). 14a (13b). 17a (2b). 21a (3b). 13b (12a). two pairs of thoracic tori present. Probably dextrally coiling. biscoeensis Gravier 1907. Margin of collar unfused. Abdominal tori distinctly larger on concave side. Sinistraliy coiled. Incubation in groove along posterior abdomen. Abdominal setae with blades as big as those in the collar seate. Collar setae of fin and blade construction. Dextrally coiled. three thoracic tori and five fascicles of setae on the convex side. abdomen not strongly asymmetrical. Sinistraliy coiled. Incubation in faecal groove attached by stalk. side setae in third fascicle present. Bushiella Knight-Jones 1973. Operculum with two or more plates stacked on top of each other. Dextralia Knight-Jones 1973. Thoracic uncini with blunt anterior peg. 23 species. Metalaeospira Pillai 1970. Thoracic uncini with blunt anterior peg. side setae in third and fourth thoracic fascicle. Side setae present. Sinistraliy coiled. Thoracic uncini with bilobed . Uncini with teeth in several rows. incubation in operculum. Abdominal setae with vestigial brushtop. Incubation in the operculum. 7 species. side setae absent. Abdomen somewhat asymmetrical. Collar setae limbate. Sinistraliy coiling. only species. abdominal setae with blades as big as those in the collar setae. two pairs of thoracic tori present. spiculosa KnightJones 1973. 3 species. Sinistraliy coiled. 2 species. Thoracic uncini with blunt anterior peg. Collar setae limbate. manuatus Vine 1972. 4 species. Dexiospira Caullery and Mesnil 1897. Uncini with teeth in several rows. Both thoracic and abdominal uncini more numerous on concave side. Margins of collar fused to form dorsal tunnel. Uncini with teeth in several rows. two pairs of thoracic tori present. collar setae limbate. side setae absent. not crossstriated. collar setae limbate. Janua Saint-Joseph 1894. Helicosiphon Gravier 1907. Eulaeospira Pillai 1970. incubation in the operculum. Sinistraliy coiled. Collar setae limbate. Anterior abdominal tori divided into two parts. Fin and blade collar setae without cross-striations. Spirorbis pagenstecheri Quatrefages 1865. anterior peg bifid or multifid. Incubation in faecal groove with attachment stalk for the embryos. Anomalorbis Vine 1972. Pronounced abdominal asymmetry. three rows of tori on the concave side and traces of a fourth on the convex side. thorax with three complete and a fourth rudimentary setiger. Spirorbis evolutus Bush 1904b. thoracic uncini with several rows of teeth. Serpula corrúgala Montagu 1803. or almost uncoiled. Spirorbis (Laeospira) koehleri Caullery and Mesnil 1897. Incubation in the operculum. Dextrally coiled. Incubation in faecal groove on attachment stalk. Spirorbis (Paralaeospira) adeonella Day 1963. less than one fourth the size of the side setae. Incubation in the operculum. collar setae limbate. Capeospira Pillai 1970. Sinistraliy coiling. Spirorbis (Laeospira) orientalis Pillai 1960. Duplicaría Vine 1972. Serpula spirillum Linnaeus 1758. A. Incubation in the tube. A. 8 species. Side setae absent. two pairs of thoracic tori present. Usually dextrally coiled. Collar setae of fm and blade construction. side setae absent. three thoracic setigers on concave side. Thoracic uncini with blunt Amplaria Knight-Jones 1973. only species. collar setae limbate. four complete thoracic setigers and remnants of a fifth. side setae in third and fourth fascicle. abdominal setae genie ulate. H. four thoracic tori and four fascicles of setae on the concave side. only species. Spirorbis laevis Quatrefages 1865. side setae absent. side setae in third fascicle present. Collar setae of fin and blade construction. side setae present. Jatiua (Fauveldora) kayi Knight-Jones 1972. numerous. Abdominal setae with blades as big as those in the collar setae. collar setae of fin and blade construction. side setae absent. Circeis Saint-Joseph 1894. three thoracic setigers present. abdominal setae with blades as long as those in the collar seta. incubation in the operculum. Side setae absent. Collar setae with limbate blades. three thoracic tori on the concave side. Dextrally coiled. only species. Side setae present in third fascicle. two pairs of thoracic tori present. incubation probably in tube.¡50 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Generic Definitions Science Series 28 anterior peg. not dentate. Sinistraliy coiled. Leodora Saint-Joseph 1894. Incubation in the tube. only species. Fauveldora Knight-Jones 1972. Uncini with teeth in several rows. 3 species. Sinistraliy coiling. Spirorbis pixelli Harris 1969. two pairs of thoracic tori. two pairs of thoracic tori present. collar setae limbate. Spirorbis (Paradexiospira) falklandica Pixell 1913a. only species. without crossstriations. 4 species. see Janua Neodexiospira Pillai 1970. Incubation in the tube. perrieri Caullery and Mesnil 1897. Incubation in operculum. Thoracic uncini with broad anterior pegs. two pairs of thoracic tori present. blades usually coarsely dentate. side setae absent. side setae in third fascicle. indeterminable Laeospira Caullery and Mesnil 1897. Collar setae of fin and blade construction. three complete and one fourth incomplete thoracic seliger. Spirorbis asperatus Bush 1904b. attachment stalk absent. even if the character should turn out to be useless in certain instances. Incubation in the tube. Serpula vitrea Fabricius 1780. 1973. 7 species. Thoracic uncini with broad anterior pegs. see Dexiospira Heterodisca Fleming 1825. Serputa crenatostriatus Goldfuss 1833. Collar setae limbate. three thoracic tori on the concave side. Spirorbis Daudin 1800. Dextraliy coiled. Dextraliy colled. Sinistrally coiled. side setae present in the third fascicle. Prodexiospira Pillai 1970. Incubation in the operculum. Incubation in faecal groove. Incubation in operculum. Simplicaria Knight-Jones. 48 species. set Spirorbis Marsipospira Bailey 1969. the coiling may vary from sinistral to dextral. Pileolaria Claparède 1870a. Dextraliy coiled. Sinistrally coiled. only species. only species. Spirorbella Chamberiin 1919. see Circeis Dexiorbis Chamberiin 1919c. attachment stalk absent. two pairs of thoracic tori present. Protoleodora Pillai 1970. 3 species. Side setae absent. cross-striated. abdominal setae with blades as long as those in the collar setae. thorax with two pairs of tori. Sinistrally coiled. Incubation in faecal groove. gouge-shaped pegs. Spirorbis (Paralaeospira) aggregata Caullery and Mesnil 1897. R. collar setae of fin and blade construction. 20 species. side setae present in the third fascicle. two pairs of thoracic tori. Side setae in third fascicle.1976 THE POLYCHAETE WORMS ¡51 anterior peg. Sinistrally coiled. Margin of collar fused dorsally. Paralaeospira Caullery and Mesnil 1897. see. two pairs of thoracic tori present. attachment stalk present. see Protolaeospira Mera Saint-Joseph 1894. questionably Spirorbis Coretus Baster 1762. P. It is thus of the utmost importance that great care be taken in using the key and definitions given above and that all the important features be checked carefully. two pairs of thoracic tori. It is quite possible that the coiling will turn out to be of great value in most of the genera. only species. side setae present in posterior thoracic setigers from the third. collar setae of fin and blade construction without cross-striations. collar setae limbate. side setae in third and fourth fascicle. Thoracic uncini with blunt. 8 species. Incubation in faecal groove. cross-striated. 3 species. Sinistrally coiled. militaris Claparède 1870a. three pairs of tori and in addition a fourth incomplete present. Sinistrally or dextraliy coiled. thoracic uncini with blunt anterior pegs. Dextraliy coiled. Incubation in operculum. Sinistrally coiled. attachment stalk absent. Sinistrally coiled. Incubation in the tube. Protolaeospira Pixell 1912. collar setae limbate. 3 species. side setae present. collar setae of fm and blade construction. Incubation in faecal groove. Spirorbis (Laeospira) pseudomiliiaris Thiriot-Quievreux. Romanchella Caullery and Mesnil 1897. Thoracic and abdominal uncini more numerous on the concave side. Side setae present. 4 species. Incubation in tube. Pillaiospira Knight-Jones 1973. Spirorbis violaceus Levinsen 1884. approximately seven thoracic setigers. Collar setae of fin and blade construction. Thoracic uncini with bilobed anterior pegs. three complete and a fourth incomplete seliger present. side setae present in the third fascicle. 1965. Neomicrorbis Roverto 1904. Collar setae of fin and blade construction. Dexiospira . collar setae limbate. attachment stalk present. Janua (Pillaospira) trifurcata Knight-Jones 1973. Spirorbis borealis Daudin 1800. blades usually coarsely dentate. few in number. ambilateralis Pixell 1912. attachment stalk absent. P. 3 species. three (two?) setigers in the thorax. Paradexiospira Caullery and Mesnil 1897. collar setae of fin and blade construction. Invalid Genera Charybs Montfort in Mörch 1863. side setae present in the third fascicle. collar setae of fin and blade construction. Taxonomic Notes Vine (1972) pointed out that the coiling of the spirorbids may not be a character of particularly high value and that even within one species. Spirorbis (Dexiospira) marioni Caullery and Mesnil 1897. prostomium trilobed. Pharyngeal apparatus well developed. 3a (2b). D. usually segraentally arranged. Pharyngeal apparatus well developed. prostomium trilobed Body short. Some forms distinctly sexually dimorphic. punicus Westheide 1971. FAMILY NERILLIDAE LEVINSEN 1883 Interstitial small forms with few segments. Body long and slender. They are characterized. only species. very fine cuticular hairs covering the body. pharyngeal apparatus absent. Ciliary bands on ventral side present. I find it difficult to associate the five families. ciliary tracts present. 8 species. Nerilla Meganerilla . heideri Remane 1925. Short-bodied. Slender. 3b (2b). without adhesive glands. usually distincdy segmented polychaetes. see Paradexiospira Stoa Serres 1855. without appendages. Parapodial cirri usually present. 2b (lb). Diurodrilus Remane 1925. and. I find it difficult to retain the concept of the Archiannelida (Fauchald 1974a) and presently FAMILY DINOPHILIDAE REMANE Very small. 2a (lb). inflated and indistinctly segmented. prostomium rounded Pygidium with two adhesive glands. Body short. 5 species. eyes absent. Key to Genera la. nuchal organs and up to three antennae. 2a (la). Prostomium with paired palps. Key to Genera la.}52 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 Pixellia Pillai 1970. T. and the features in which they resemble each other are exactly those that have been considered to be of selective advantage for the mesopsammon (Swedmark 1964. Body with nine segments including the post-peristomial pharyngeal segment Body with less than nine segments Composite setae present Simple capillary setae present Three antennae present Maximally one antenna present 2 4 Mesonerilla 3 . without adhesive glands. as pointed out by Hermans (1969). 3 species. to any known polychaete family. some forms with adhesive glands posteriorly. 1 b. pygidium rounded or conical. D. The main changes involve terminology to fit the terms used in this paper. A. eyes usually present Apharyngtus 2 Trilobodrilus 3 Diurodrilus Dinophilus Generic Definitions Apharyngtus Westheide 1971. and have for that reason listed them below. The key to genera and definitions given below have been modified after Jouin (1971). vorticoides Schmidt 1848. 1932 The following five families usually are considered as belonging to a separate order. separately or as a unit. Trilobodrilus Remane 1925. more or less inflated. Ventral eversible pharynx with four jaws or unarmed. in addition to the regularly arranged ciliary bands. Muscular pharynx absent Muscular pharynx present Body slender and elongated. Dinophilus Schmidt 1848. distinctly segmented bodies. see Proiolaeospira Sinistrella Chamberlin 1919c. lb. Uniramous parapodia with capillary setae or composite setae in a single fascicle. Pygidium with paired adhesive glands ending distally on conical projections of various shapes. eyes absent Pygidium conical. minimus Remane 1925. 3b (2b). in alphabetical order. In agreement with Jouin (1971) and Westheide (1971). Archiannelida. setae and parapodia. Hermans 1969). The family recently was reviewed by Jouin (1971) and by Westheide (1971). Like Jouin and Westheide. I do not think the five families are particularly closely related to each other. 2b (la). Pharyngeal apparatus present and well developed. indistinctly segment dinophilids. by features adaptive to life in the mesopsammon. 3a (2b). indeterminable FIVE "ARCHIANNELIDAN" FAMILIES consider them free-standing families of the Polychaeta. see Leodora Spirorbides Chamberlin 1919c. pygidium rounded or conical. Family NERILLIDAE. posterior end of the above. 1971. Protodrilus hatscheki. from Jouin. about 50x. about 15x. 1971. about 25x. after Jouin. . posterior end of the above. (A). Family POLYGORDIIDAE. Diurodhlus ankeli.7976 THE POLYCHAETE WORMS 153 FIGURE 39. about 15x. (C). Polygordius neapotitanus. 1971. about 25x. (D). posterior end of the above. Family DINOPHILIDAE. (F). Family PROTODRILIDAE. (H). Mesonerilta sp. 1971. about 15x. after Jouin. 1971. Family SACCOCIRRIDAE. (E). about 15x. (B). after Jouin. after Jouin. about 25x. Saccocirrus polycercus. (G).. A'. Segmentation indistinct anteriorly. more distinct posteriorly. Body with eight segments including the pharyngeal segment. (4b). All setae simple capillaries. two antennae present. intermediate segments with composite setae. Prostomium with two solid antennae and nuchal slits. (5a). only species. gracile Remane 1925. Body with seven segments including the pharyngeal segment. two palps present. palps present. Single slender capillary setae in each of the last 10-12 seg- . luederitzi Remane 1949. Chaetogordius Polygordius Prostomium conical with paired. Nerillidopsis Jouin 1967. only species. Pharyngeal segment with setae. may be absent. Both simple and composite setae present All setae either composite or simple All setae composite All setae simple and capillary Pharyngeal segment asetigerous. NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Eight segments. Antennae absent. (4b). Two large palps and maximally one antenna present. antennata Schmidt 1848. All setae composite spinigers. Setae composite. 5 species. M. marine Composite setae. C. All setae simple capillaries. Anterior and posterior setae simple capillaries. (4a). Mud-dwelling forms. antennae absent. Body with nine segments including the pharyngeal segment. canaliculatus Moore 1904a. including the post-peristomial pharyngeal segment Seven segments present . Setae simple capillaries. M. Mesonerilla Remane 1949. parapodial cirri absent. parapodial cirri present. P. Two palps and three antennae present. palps reduced to triangular lateral comers on the prostomium. palps present. (4a). 7 species. N. Body with eight segments including the pharyngeal segment. Key to Genera la. only species. hyalina Jouin 1967. Psammoriedlia Kirsteuer 1966. Body with nine segments including the pharyngeal segment. palps reduced Setae simple and capillary. Body with eight segments including the pharyngeal segment. except on last segments. Three antennae and two palps present. Antennae and parapodial cirri absent. P. Body with seven segments including the pharyngeal segment. limicola Jouin and Swedmark 1965. palpifoliaceus Ax 1954. Antennae and parapodial cirri absent. FAMILY POLYGORDIIDAE CZERNIAVSKY 1881a Slender. T. freshwater Pharyngeal segment with setae. Antennae absent. T. including the pharyngeal segment. Paranerilla Jouin and Swedmark 1965. All setae composite. Nerillidium Remane 1925. maximally two antennae present. 6 species. swedmarki Boaden 1961. only species. Troglochaetus Delachaux 1921. setae absent in most forms. Freshwater and caves.i54 4a 4b 5a 5b 6a 6b 7a 7b 8a 8b (lb). N. All setae simple capillaries. parapodial cirri present except on last segment Science Series 28 5 g Nerillidopsis 6 Thaiassochaetus 7 Troglochaetus Nerillidium Paranerilla Psammoriedlia Generic Definitions Meganerilla Boaden 1961. Nerilla Schmidt 1848. (5a). Body with eight segments. Thaiassochaetus Ax 1954. only species. mainly interstitial forms. segmentation and ciliation poorly developed. lb. Eyes and parapodia absent. pharyngeal segment asetigerous. beranecki Delachaux 1921. (6b). Eversible or noneversible muscular pharynx ventral. All setae simple capillaries. 2 species. slightly articulated antennae and large nuchal organs. only species. (lb). (6b). ruperti Kirsteuer 1966. Body with nine segments including the pharyngeal segment. Setae present in posterior segments Setae absent Generic Definitions Chaetogordius Moore 1904a. Prostoitiium rounded or conical with paired. The family is known for a single genus. lb. mirabilis Czerniavsky 1881a. papillocercus Bobretzky 1872 and about 12 species. parapodia always absent. chiselshaped setae. The family has been reviewed recently by Jouin (1966). antennae solid. . eyes present. 15 species. 20 species.7976 THE POLYCHAETE WORMS FAMILY PROTODRILIDAE CZERNIAVSKY 155 1881a ments. but may be absent. Setae absent. Polygordius Schneider 1868. Proboscis a ventral muscular pad. Protodrilus chaetifer Remane 1926. antennae hollow. Interstitial slender forms. Segmentation and parapodia poorly developed. 2 species. S. hollow or solid antennae. with simple. Setae or segmentally arranged adhesive organs present Setae and segmentally arranged adhesive organs absent Protodriloides Protodrilus 1881a Generic Definitions Protodriloides Jouin 1966. Jouin (1971) gave a key to species. Pharynx muscular. P. with genotype. lacteus Schneider 1868. Setae or segmentally arranged adhesive glands present. Parapodia uniramous. Segmentation usually indistinct along the whole body. Salivary glands and lateral organs absent. P. Prostomium with paired. FAMILY SACCOCIRRIDAE CZERNIAVSKY Interstitial slender forms. but not eversible. present in most forms. all others with a single seta on either side. eyes present. Key to Genera la. usually smooth antennae and large nuchal organs. Protodrilus Czerniavsky 1881a. setae absent in most known species. Saccocirrus Bobretzky 1872. Setae and segmentally arranged adhesive glands absent. Posterior end of body usually slightly wider than the rest of the body. Prostomium with paired hollow antennae. only one segment with paired setae on either side. Salivary glands and lateral organs present. apodous•without parapodia (66). bifurcate•with two prongs (23). denticulate•with small teeth (40). cirrus (1)•sensory projection. cephalic cage•long protective setae enclosing and protecting the head. abdomen•posterior part of the body. cirriform•slender and cylindrical (41). ceratostyle•distal joint of an antenna (29). tentacles and palps (13). deciduous•liable to fall off (like leaves). accompanying. buccal•pertaining to the mouth. innervated from the first part of the brain (12. Generally. compound (or composite) setae•^jointed setae (37). homologous with the dorsal cirrostyles (43). elytron (a)•dorsal scales found in the aphroditoids. cirrophore•basal joint of a cirrus (35). slender tapering setae. clávate•club-shaped (32). lateral or frontal surface of the prostomium. antenna (e)•sensory projection arising from the dorsal. branchia (e)•any extension of the body wall with a loop of the vascular system or which is wellequipped with capillary blood-vessels. aristate•about setae: simple setae with smooth shaft and a tuft of hairs or a single spine distally (17). bipinnate•formed like a feather with a main stem and two rows of side branches (26). used about antennae. caruncle•posterior sensory organ projecting from the prostomium sometimes over the first several segments (28). asetigerous•without setae (66). simple setae in rows. chevron•V-shaped chitinized jaw piece at the base of the eversible pharynx in some goniadids (31). Number in parenthesis refer to the specific example of this feature illustrated in the appended illustrations. avicular•beaked (shaped like a bird's head) (20). . ceratophore•basal joini of an antenna (30).38. digitiform•finger-shaped (42).78. bidentate•with two teeth (21). bilimbate•simple setae with two wings or flattened margins (24). biarticulate•with two joints. cirrostyle•distal joint of a cirrus (34). cephalic veil•hoodlike membrane between the paleae and the buccal tentacles in pectinariids. companion setae•small. Acicular setae are thick. biramous•with two branches. capillary (ies)•in polychaete literature used either as an adjective in the combination capillary seta or as a noun characterizing long. anal cirrus (i)•one or more elongated projections from the pygidium (11).and neuropodia present (6). from the superior part of the notopodiura (dorsal cirrus) or from the inferior part of the neuropodium (ventral cirrus) (36. etc. cephalic rim•flange encircling the head (25). usually slender and cylindrical. cephalon•head. aileron•accessory jaw plate in the glycerids (9). a gill (27). such as distal and proximal. these terms can be found in a good dictionary. sometimes followed by a tail (2).61). or alternating with larger setae.80). usually used about parapodia having both noto. usually hooks of some kind. The glossary contains the terms characteristically used in systematic polychaete literature and explanation of idiosyncratic usage where terms are used in a different sense from the usual lexicographic one. auricular•ear-shaped (19).156 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY GLOSSARY Science Series 28 The glossary does not contain terms in general use in zoology or in biology in general. behind the thorax. anterior and posterior. projecting setae (5). bifid•split in two (22). arborescent•branching (like a tree) (18). aclculiun (a)•stout supportive setae found internally in each parapodial ramus where these project from the body. dentate•with teeth (39). . homogomph articulation•with articulation distinctly and symmetrically at right angles to the long axis of the shaft (54). usually blunt (used about setae) (45). blunt. foUaceous•leaflike (47). falciger•distally blunt and curved setae (45). limbate•simple seta with a flattened margin or wing (56). imbricated•overlapping (like tiles). smaller hooks arranged in single or double rows are often referred to as uncini. hook•general term used about stout-shafted. hooded setae•setae distally covered by a delicate chitinous envelope or guard (51). harpoon seta•stout pointed seta with recurved barbs near the tip (50). geniculated•bent (like a knee) (49). Diagram of selected terms defined in the Glossary #1-18.1976 THE POLYCHAETE WORMS 157 elytrophore•aphroditoid cirrophore carrying an elytron (44). FIGURE 40. epitoke•modified reproductive stage or specimen. falcate•distally curved. filifonn•slender and threadlike (46). ligule•finger-shaped major process on a parapodium (55). facial tubercle•projecting ridge or lobe on the upper lip of certain polychaetes (especially scaleworms). often swarming. used mainly about flattened kinds. often distally curved and dentate setae. but also more generally about all kinds of major parapodial processes (8). fusiform•cigar. liemigomph articulation•with asymmetrical articulation nearly at right angles to the long axis of the shaft (52).or spindle-shaped (48). inferior•the more ventral (of two or more structures). heterogomph articulation•with articulation clearly oblique to the long axis of the shaft (53). lobe•major parapodial process. Diagram of selected terms defined in the Glossary #19-60.158 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 •52 -53 -54 FIGURE 41. . ^^^ notosetae•setae of a notopodium. ocular peduncle•projecting structure supporting the eyes. moniliform•beaded or beadlike (57).|¿. FIGURE paragnath•chitinous denticle in the pharyngeal cavity of nereids (63). with a long basal rod supporting the uncinus metastomium•rarely used terin covering the segmented body between the prostomium and pygidium. operculum (a)•literally a lid. usually flattened setae. but including neither. onunatophore•see ocular peduncle. neurosetae•setae of the neuropodium. occipital•pertaining to the posterodorsal part of the prostomium (64). pits or grooves. especially in the polyodontids. neuropodiuin (a)•ventral branch (ramus) of a parapodium (7). ^. abruptly tapered (58). see Sabellariidae and Serpulidae. muitiarticulated•with several joints (60). ocular•pertaining to the eyes. sometimes as paired epaulettes stretching posterolaterally from the prostomium.]976 THE POLVCHAETE WORMS 159 long-handled•used about uncini. palmate•resembling the fronds of a palm (67). notopodium (a)•dorsal branch (ramus) of a parapodium (3). palp (s)•sensory or feeding structures innervated from the posterior part of the brain or from the circumesophagal nerve ring. nuchal•pertaining to the neck. . posterodorsal and used in feeding in the spioniform worms (13). muero•with a sharp point or tip. 42.^ ^. anteroventral and sensory in the Phyllodocida. used about a structure some tubicolous worms use as stoppers for their tubes when the occupant is retracted. used about sensory organs found on the posterodorsal side of the head and variously developed as paired or single processes. Diagram of selected terms defined in the Glossary #61-83.^. natatory•swimming. palea (e)•strong or broad. often eversible. terminal or primary one. is absent (79). especially syllids (81). posterior to the head(l).65). plumose•resembling down. used about parapodial lobes or ligules (4). setter•segment carrying setae.'V c. always modified for feeding purposes. penicillate•brushlike (70). pectinate•combiike. the first being the apical. a pre-segmental part of the body anterior to the mouth. S-shaped setae with a distinct beak. in the latter case considered homologous with the dorsal and ventral cirri of normai. fine threads may be present. lumpy (72). secondary tooth•the second of two teeth. spiniger•seta that tapers to a fine point. in practice including also segments fused to this structure. found posterior to the pharynx. pharynx (ges)•anterior part of the digestive tract. often with antennae and eyes (14. without a long rod-shaped support. subulate•awl-shaped. tridentate•with three teeth (82). most frequently the notopodium. used about parapodia in which one ramus. with series of projections arranged like the teeth of a comb (68). Uncini usually are arranged in rows transverse to the long axis of the animal (83). presetal•anterior to the setae. prostomium (a)•anteriormost. pygidium (a)•post-segmental terminal part of the body carrying the anus (10).160 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28 parapodium (a)•segmentaily arranged projections carrying setae. rugose•roughened. often with a platelike base. proventricle•muscularized region of the anterior digestive tract in syllids.64). trepan•chitinized. most frequently used about composite setae (33). usually blunt-tipped. p--sicephalic parapodia (15. with a main stem and lateral branches. Çl^-W-^^-^ VxiXv) C^^A v^^ ^^V-^ ^^^^-^ . strictly including only the region around the tnouth. radicle•one of the main tentacles in the tentacular crown of sabellids and serpulids. uneiniger•segment carrying uncini. apart from the prostomium and pygidium set off internally or externally by septa or otherwise from the preceding and following parts. hairy (69). spatulate•blade-shaped. stylode•small. postsetal•posterior to the setae. uniramous•with one branch only. used about parapodial lobes or hgules (8). fingerlike projection associated with a parapodium. peristomium (a)•first distinct post-prostomial region. seta•secretion from the parapodia forming the armature of these structures. segment•any part of the body. usually small and distinctly longer than wide (73). reniform•kidney-shaped (76). uncinus (i)•general term covering sharply dentate. or small. forming the posterior part of the recognizable head (16. pinnate•featherlike.66). sub-biramous•pertaining to parapodia in which the neuropodia are well developed and the notopodia reduced (77). simple seta•unjointed seta. ramose•branched. scaphe•flattened caudal appendage of pectinariids. deeply imbedded setae. short-handled•used about uncini. anteriorly toothed part of the eversible pharynx of some worms. enclosing at least the anterior part of the brain. spinous pocket•enlarged. thorax•anterior region of the body. foot (16). pocketlike serration along the margin of the setae of some scale-worms (74). superior•the more dorsal (of two or more structures). tentacular cirrus•sensory projections arising either from the peristomium or from cephalized segments. \(A. unidentate•distally entire (45). see also bipinnate (71). sometimes with a muero (59). sometimes also for burrowing (62). tapering to a fine point (75). Leningrad 30:13-20. 1913a. 2. Classißcation des Annelides. Paris (1)20:156-159. Zool Anz. et description de celles qui habitent les côtes de la France. Aarb. Micro. Indian Mus. 81:157-294. Report upon the Annehda Polychaeta of Beaufort. Ergänzung zu den Polychaeten von Spitzbergen. Hamburg wiss. Akad. and R. Kurze Übersicht der Polychaeten der Litoralzone der Bering-Insel (Kommador-Inseln). 290 pp. Zool. 1922. K. Mus. Genoots. Annélides. 1949. Bergens Mus. Fauna Árctica. 1834. Medd. 1928a. Nauk. HRABE. Vidensk. [The polychaete fauna of the northern part of the Japan Sea]. 1922:38-63. The comparative morphology and embryology of the head in scale worms (Aphroditidae. Sedentaria. Quar. Jahrb. Classißcation des Annélides. Nat. 1891. I. 41:253-273. AND H. 1963. dansk naturh. . North Carolina. Polychaeten von Franz Josephland II. Ergebnisse der Hamburger südwest-australiscben Forschungsreise 1905. Ann. 1830. 1925. spec. 42:89-107. Polychaeten von Neuseeland II. 4(5):65-304. 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Sitzber. 1924. 37:495-502. SÔDERSTROM. Morph. 56:254-264. R. [Polychaetous annelids of the family Paraonidae Cerruti. Ann. Mar. Description de Spirorbis (Laeospira) pseudomilitaris n.J. Morphologische Untersuchungen am "Kopf" der Pisioniden (Fisione puzae nov. Some new and little-known serpulid polychaetes from the continental slope. Les Annélides polychètes des Mers de la Nouvelle-Zemble. Akad. Polychaeta). fishery steamer ALBATROSS. Neapel 3:15-52. Contributions to the zoology of the Philippine Archipelago and adjacent regions. 1909 (Polychaeta. 1968a.. 1922. (In Russian). 39:1-42. B. naturf. Ann. Fish. THéEL. Atlantic Deep-sea Exped. [On types of polychaete larvae (Polychaeta)]. Nat. A. Arch. 1855. SPENGEL. J. 1965. Ark. 1835.. Ass. Nauk SSSR 17:74-95. Freunde Berlin 1922:86-81. Irish Acad. The interstitial fauna of marin sand. THIRIOT-QUIEVREUX. 1879. 1963. J. Ser. 1955. n. C. C. Mus. Svensk. Stat. eine schmarotzende Eunicee. St. H. 1953. SsoLOWiEW. 1923. 150:298-313. Zool. Univ.. or the region about the mouth of the Bay of Fundy. Zur Kenntnis der Turbellaria rhabdocoela und einiger anderer Würmer des Mittelmeeres. Sys.G.. Leipzig 1868: 51-60. Über Bau und Entwicklung von Polygordius. 1-91. 1879.S. Gymnocopa og Polychaeta. polychete Spirorbinae. 15:201-261. Wien 23:347-366. New Zealand Inst.M. Note préliminaire sur un polychèle sédentaire aberrant.M. Paris. Akad. U. gen. sp. 286 pp. [Types of reproduction and development of polychaetous worms in connection with their geographical distribution]. Some Serpulidae (Annelida: Polychaeta) from Heron Island. et de sa larve. Nauk SSSR.gen. Étude de la microstructure des soies de polychètes Capitellidae et Oweniidae au microscope électronique a balayage: un critère systématique précis. 16(3):3-75.. Soc. n. SOUTHERN. Smithsonian Contr. 1952. Biol. STôP-BOWITZ. Hist. Zool. SEIDLER. Errantia) de Golfe du Tonkin de la Mer de Chine Méridionale. V. Studien über die Polychaetenfamilie Spionidae. Beiträge zur Kenntnis der Polynoiden IV.L. 6:1-67. The polychaetous annelids of Porto Rico. Polychaeta from the MICHAEL SARS North Atlantic deep-sea expedition 1910. TREADWELL.S. Stat. SERRES. Wiss. On reproductive pattern as a specific characteristic among nereid polychaetes. Sedentaria)]. gen. Proc. Akad. Mag. series 10:71-101. U. Stockholm (2) 4:159-162. THOMASSIN. Rep. STORCH. (In Russian). Marine Serpulidae (Annelida: Polychaeta) of eastern Queensland and New South Wales. Sei. D. 1968. 1972. Doklady Akad. 1901. Annulata Dánica. Oligochaeta. Vie et Milieu 6:413^25. Sei. sp. Descriptions of a few invertebrated animals obtained at the Isle of France. Psammodritus halanoglossoides n. Trav. Sedentaria) of the Barentz Sea. J.] (In Russian). Nauk SSSR Moskva 150(6):1393-1396. Biol. AND C. Handl. 20:181-210. 1968b. Gesells. Zool. P. 1882.. Balanoglossi. (In Russian). 179-200. 1958. SiEwiNO. Mar. Endoume. Ann. Berlin 89A(11):1-217. V. Bull. S. Sei. 1881:699-747. Nemertina and Annelida of the Bermudas. 1. Polychaeta. Arts Sei. . Acad. Rep. Invest. AND B. Danish INGOLF-Exped. 17 pp. _. Arch. estest.P. Polychaetous annelids from Shakotan (Hokkaido) The collection in 1967. Porifera. Ann. 1950. Rep. 1878. Lombriciens. Leipzig. Paris. Adv. Histoire naturelle des Annelés marins et d'eau douce. Nauk SSSR Leningrad 13: 207-209. 1940. Mus. 24:338-341. Proc. Marine fauna of Eastport. L. Mus. F. with a survey and a bibliography of fresh and brackish water polychaetes. Soc.E. Polyzoa. VicuiER. U. 1972. (In Chinese and Russian). Maine. 1879b. Stud. The genera of Arenicolidae (Polychaeta). The Zoology of Iceland 2(19): 1-182. with an account of the physical characters of the region. 1890. Albany Inst. 8:424-448. 1884. Nouvelles études sur le plancton de la Baie d'Alger. Études sur les animaux inférieurs de la Baie d'Alger. [Nouveau groupe d'Annelidcs]. 1968. ViviANl. Fisheries Commission. Conn. Hokkaido Univ. quatordecim luseentiam animalculorum novis speciebus illustrata. Polychaetes of the Iranian Gulf. 19:595-612.S. Annelida Chaetopoda of the Virginian coast. J. Acad. 1885:707-755. 1965 by Israel Program Scientific Translating. H. 1871-72:295-778. 1911. J. [Polychaetous annelids of the family Aphroditidae from the Kurilo-Kamshatska Trench]. 10:595-671. WELLS. Trans. 1874. Rep. Wash. Amer. 1880. Explorations of Casco Bay by the U. BENEDICT. 1880. -. E. Térétulariens et Planariens. Results of recent dredging expeditions on the coast of New England. Additions to the Turbellaria. UcHlDA. 1885. 1871.Y. Akad. C. gen. Nouvelles études sur le plancton de la Baie d'Alger. 1879 und 1882. Hist. with revisions of some New England genera and species. 1934. VINE. Ocean. with annotated list of the species hitherto recorded. Rep. (6) Zool. VAILLANT. Acad. Studies on the fauna of Curacao and other Caribbean Islands. Nat. New Haven (3) 5:1-16. N. Conn. Iran 4:247^00. Comm. zool. [New species of polychaetous annelids of the family Phyllodocidae (Polychaeta)]. B. ein aberranter Archiannelide aus dem Mesopsammal der tunesischen Mittelmeerküste. (In Russian). Jerusalem). Prague. Rep. Akad. Danish Sei. 9:202-269. Zool.. J. Amer. I. Linn. 1884. Arch. A. Nat. Nauk SSSR. Anthozoa. 1873b. a new genus and species of serpulid polychaete. Jerusalem).178 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY 1928. Sinica 1(4): 1^0. Annelida Chaetopoda of New Jersey. 166 pp. Fish.Y. Petersburg 3: 344-347. Ann. chiefly from brackish water. N. Die wirbellosen des Weissen Meeres. zool. Paris (2) 4:347-442. 1089:1-4. N. Arts Sei. The Annelida Chaetopoda from EVovincetown and WeUfleet Mass. Mus. [Polychaetous annelids of the Far Eastern Seas of the USSR]. Amer. Science Series 28 1881. 1887. WESTHEIDE. Proc. exp. 1:1-171. Comm. Nat. Zool. Trans. Novität. System und Morphologie des Oligochaeten. 1886. Echinodermata. Inst. 1958. 1912. Annelida. Ann. Suriname 17:1-41. Inst. 1955b. Trudy Zool. 22:340-395. Phosphorescencia maris. Nemertina. U. 102. Mass. WESENBERG-LUND. Ass. (translated 1974 by the Israel Program for Scientific Translation. U. Sei. W. in 1873. Sei. Paris (9) 15:85-99. 1971. Trudy Inst. 1972. Historical sketch. Notices of recent additions to the marine Invertebrata of the northeastern coast of America. Polychaetous annelids from the ARCTURUS océanographie expedition. (In Russian). Comm. Wagner. 3(2):340-768. Bdellomorphs.S. Arch. Lesser Antillean polychaetes. Mikrofauna des . Cruise VII CARNEGIE 1928-1929:30-59. AND J. Essex Inst. Sei. 4(14): 1-92. Bull. 1872.S. nov. (In Russian). UscHAKOV. Tunicata. Trois Aphroditiens pélagiques. Genoa.J. Gephyrea. [The polychaetous annelids of the families Phyllodocidae and Aphroditidae from the Yellow Sea]. Notice of recent additions to the marine Invertebrata of the northeastern coast of America. Fish. VEJDOVSKY. Res. Nematoda. Polychaeta.E. G. Mollusca. Apharyngtus punicus nov. P. Proc. Fac. 1951. Akad. Hist. Biological results of the last cruise of the CARNEGIE. 32: 101-128. exp. H. Keys to the Fauna of the SSSR 56:1-433 (translated. 1873a. 39:128-159 (reprint of the 1879 paper with added plates). 1805. New England Annelida. 3:2-6.. [Polychaeta 1. WAGNER. Pt. spec. WEBSTER. D.S. Akad. U. Nat. Sur le genre Sagitella N. Zoologische Forschungen an der Küste des Solowetzkischen Meerbusens in dem Sommer-monaten der Jahre 1877. J.L. UUANiN. Maine. with descriptions of new genera and species and critical remarks on others. The Annelida Chaetopoda from Eastport. St. The Annelida Chaetopoda of New Jersey. Roret. Paris 7:1-32. Inst. with description of new genera and species and critical remarks on others. 1879a.E. Salem. Report upon the invertebrate animals of Vineyard Sound and the adjacent waters. Fauna of the SSSR. gen. Fish. Nauk SSSR 12: 311-321. London Zool. VERRILL. 1953. 1959. London 133:301-314. Nauk SSSR. Polychaetous annelids. 4:285-324. Pt. Hirudiniens. Sphaeropomalus miamiensis. 1955a.s.. Polychaetes of the suborder Phyllodociformia of the Polar Basin and the northwestern part of the Pacific]. Zoológica 8:449-489. State Mus. 1900.S. Recherches sur les Annélides pélagiques. Nat. Wu. Sei. Sei. Engelmann. n. (In Russian). N. 1949. Nat. Soc. Oceanol. Paris (9) 13:187-267. Trudy Obsch. U. 1885.Y. A new genus and two new species of polychaetous annelid from Texas and one new species from the Philippine Islands. Sei. P. [943. Spirorbinae (Polychaeta: Serpulidae) from the Red Sea including descriptions of a new genus and four new species. 1959. 51:1-16. 271 pp. Natuurwetensch. 2: 165-205. Proc. State Mus. gen. Trans. Ann. (In Russian). ZENKEWITSCH. H. 1851. Majoru. ZEPPELIN. Zool. Assoc. 1883. WooDWORTH. Anz. Akad. 4:243-324. 11(12):3-10. Vetensk. 1902. Mainz. W. A. The palolo worm. London 21:159-272. Comp. Wiss. VEGA-Exped. T. K. Ceylon Pearl Oyster Fish. Zool. Adv. ZiBROwius. WiRÉN. Chaetopoda frân Sibiriska Ishafvet och Berings Haf insamlade under VEGA-Expeditionen 187879. Ammocharidae. WiLLEY A. at Ceylon in 1902. Polychaeta) from Eniwetok. Review of some little known genera of Serpulidae (Annelida: Polychaeta). WoLLEBAEK. .A. 39:615-652. Kl. 1883. 1911 (2): 1-144. Zool. nouvelle annélide parasite de la famille des Euniciens. Eunice viridis (Gray). Akad. Svensk. Kí. Brit. Pac. Accepted for publication June 30. Über den Bau und die Teilungsvorgänge des Ctenodrilus monostylos n. [971 (6). Rep. Zool. and Bikini Atolls. Nordeuropaeiske Annulata Polychaeta 1935. 109:195-203. Akad. WILLIAMS. Nat. Bull. Amphictenidae.233-249. Vetensk. I. Report on the collections of natural history made in the Antarctic regions during the voyage of the SOUTHERN CROSS. 1964. Report on the British Annelida.M. Nat.) an der Murmankiiste. 12:262-283. 18:146-159. wiss. 2:383-428. Polychaeta. Mus. Marshall Islands. Suppl. 1912.H. A. Zeits. Smithsonian Contr. 1969. Kristiania. Hand!. sp. 1886. WooDWicK. Vidensk. lagtt. Haematocleptes terebellidis. Polydora and related genera (Annelida. Sei. polaris n. Rep. Ampharetidae. Über das Vorkommen der Brackwasser-poiychaete Manayunkia (M. M. 112:1-22. L. VON. Math. Sei. Report on the Polychaeta collected by Professor Herdman. 1975. K. 1907.1976 THE POLYCHAETE WORMS 179 Meeresbodens. sp. _. Terebellidae og Serpulidae. Mat. Skr. 1905. Harvard 51:1-21. 124 Americonuphis. 100. 107 Blenda. 124. 60 Alluaudella. 79. 48. 48 Autolytides. 53. 102 Bergstroemia. 81 Anisoceras. 17. 123. 25 Arippasa. 85 Autolytus. 58. 37 Areniella. 60 Auslralonereis. 140 Amphicorina. 133 Aphrodita. 125 Amythasides. 129. 87. 48 Anamobaea. 91 Ariapithes. 30 Admetella. 59. 129. 130 Axiothea. 59 Adyte. 18 Aricideopsis. 125 Anomaiorbis. 52 Asychis. 103 Amphicora. 60 Bathynoe. 73 Anoplosyllis. 123. 78 Andresia. 59 Acicularia. 107. 75 Alitta. 17. 32. 13. 123. 97 Aglaopheme. 91 Alkmaria.180 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY INDEX Page numbers set in bold face type represent citations of illustrations. 79. 37 Arenicolides. 130 Amage. 60 Alücunhia. 107 Amphibranchus. 60 Austrophyllum. 98 Benhamipolynoe. 10. 49 Beikeleya. 37 Aberranta. 39 Atelesyllis. 14. 60 Antinoe. 22 Apistobranchus. 112 Apoprionospio. 129. 101 Amphiro. 25 Boguea. 78 Ancistria. 77 Anoplopisione. 91 Aonides. 59. 149. 108. 18 Acholoe. 100. 134 Ampheiothrix. 117 Artnandia. 124. 125 Amytis. 92. 57. 16 Bhawania. 35 Ancistrosyllis. 103 Aspe ira. 59. 43 Amaea. 133 Bathyadmetella. 8. 60 Antiobactrum. 105 Ammochares. 85 Anaitides. 12. 131. 23 Aporosyllis. 88 Austrolaenilla. 93. 15. 23 Arabella.52 Alciopidae. 56. 125 Amphictene. 121. 17. 42 Artacama. 120 Amphiduros. 133 Amaeana. 125 Asclerocheilus. 115 Ammotiypane. 51. 109 Augeneriella. 8. 22 Apneuma. 58. 121. 111 AnguillosylUs. 124 Amblyosyllis. 91 Antonbruunidae. 60 Aretonoe. 111 Archidice. 29. 133 Auchenoplax. 110-111 Arabes. 30 Arcteobia. 111 ArabelUdae. 41 Axiothella. 135 Apophryotrocha. 17 Alciopa. 25 Aotearia. 8. 104. 81 AutosylUs. 8. 8. 72 Biborin. 17. 80. 8. 107 Aphlebina. 56. 123. 91 Acesta. 138 Australaugeneria. 77 Anoplonereis. 11. 80. 60 Arenia. 111 Aracoda. 54 Apistobranchidae. 54 Aphrogenia. 110 Bilobaria. 37 Science Series 28 Abatenicola. 51-52 Alciopina. 25 Abyssoclymene. 124. 55 Barrukia. 143 Aponobranchus. 133 Apomatopsis. 152 Aphelothrix. 42 Ampharana. 150 Amythas. 23 Ancistargis. 137. 133 Amphytrite. 42 Antonbruunia. 79. 133 Ampiaría 149. 109 Aparaonis. 75 Amphiglena. 22 Aricidea. 17 Antinoana. 124 Ampharetidae. 18 Apharyngtus. 35 Aicnicola. 130 Arwidssonia. 97 Aglaura. 12. 33. 81 Athelepus. 39 B Baffmia. 59. 25. 11. 125 Anobothrus. 123. 8. 8. 140 Amphicteis. 58. 124 Allia. 133 Anobothrella. 85 Aquilaspio. 65 Alcandra. 140 Asterope. 138 Blainvillea. 59 Aceionereis. 66 AcfociiTidae. 122 Ampharetides. 121128. 60 Aedicira. 140 Aristenia. 141. 10. 60 Andromache. 60 Arctonoella. 33. 22. 150 Anophthalmus. 65 Bebryce. 130 Amphitritoides. 128 Asabellides. 60 Alentiana. 73. 46. 81 Ambo. 59. 41 Aryandes. 18 Allmaniella. 56. 30 Amelinna. 124 Ampharete. 24. 36 Anthostoma. 18 Aenone. 107 Amphisamytha. 137. 60 Bathya. 25 Bispira. 35 Arete. 57. 103 Boccaidia. 110. 125 Audouinia. 60 Bathychloeia. 54 Aphroditidae. 97 Aonopsis. 22. 8. 51. 57. 61 Benthoscolex. 39 Acanthicolepis. 128. 137. 59. 57. 85 Anotomastus. 130 Artacamella. 14. 36 Barbularia. Amphitritides. 43 Ammotrypanella. 128. 81 Axionice. 133 Amphiron. 57. 13. 120 Aricia. 130 Arenicolidae. 57. 111 Aglaundes. 23 Aonis. 129. 44 Asloegia. 100 Acoetes. 117 BarantoUa. 147 Apomatus. 100. 52 Alentia. 125 Amphitrite. 13. 124. 111 Aglaophamus. 113 Anisocirrus. 102 Bathyglycinde. 135 . 53-55. 94 Bathymoorea. 123. 102 Amphinomidae. 30-31 Acrocirrus. 30 Augeneria. 140 Anaspio. 31. 81 Aloysina. 131 Balanochaeta. 148. 100-103. 60 Antinoella. 111 Agnodice. 21. 47. 138 Amphinome. 59. 140. 116 Bradabyssa. 149. 40 Clymenura. 26 Dipolydora. 13. 102 Chloochaeta. 27 Chinonereis. 80. 93 Bouchiria. 144 Diurodrilus. 137. 40 Clytie. 120 Ceratocephale. 151 Chaunorhynchus. 71. 149. 30 Dorvillea. 102 Chloenea. 152 Diopatra. 61 THE POLYCHAETE WORMS 18] Cryptonota. 152. 87. 36 Dasychone. 36 Carazzta. 71 Conopomatus. 91 CheUonereis. 138 Ditrocha. 30 Cirratulus. 104. 87. 18 Cirrosyllis. 65 Chaetosyllis. 13. 32. 140 Branchionereis. 143 Califia. 143 Bonuania. 42 Dinophilidae. 137. 49 Chaetopteridae. 88 Decamastus. 150 Ciiraria. 10. 32. 88. 91 Chitinopoma. 33. 33. 82 Briaraea. 91 Cirrophorus. 128 Cnicigera. 17 Coelobranchus. 33. 49 Clavisyllis. 111 Duplicaria. 24 Distylia. 116. 131 Commensodonim. 8. 88 Cheironotus. 8. 91 Branchiosabella. 87. 78 Caesicimis. 136. 86. 61 Brachysyllis. 111 Centrocorone. 26 Cherusca. 37 Chrysodon. 33. 33. 33. 120 Clapaiedia. 25 Carobia.31. 150 Buskiella. 116. 23. 33. 39. 100. 128 Branchiosyilis. 47. 29. 133 Chaetogordius. 55 Cyaxares. 30 Ciirobranchia. 14. 85 Chaetozone. 151 Dexiospira. 141 Capeospira. 85 Dexiorbis. 88 Dendronereis. 141. 58. 98 Conconia. 8. 142. 33. 17 Branchioasychis. 97 Diploceraea. 36 Capitita. 80. 88 . 74. 31. 102. 13. 29. 39. 140 Brania. 8. 150 Dialychone. 33. 27 Disomides. 154 Chaetoparia. 85 Diplocirrus. 61 Dindymene. 133 Dendronereides. 150 Cabira. 41 Circeis. 116 Branchamphinome. 36 Ditrupa.31-36 Capitellides. 34. 135 Bonhourelia. 117 Diplotis. 100. 140 Distylidia. 100. 21 Crithida. 20 Ctenospio. 109 Cenothrix. 19-20 Ctenodrilus. 141 Caobangiidae. 87. 99 Bruunilla. 52 Campesyliis. 138 Dendrobranchus. 26 Colonianella. 138 Desmosyllis. 35 Brandiika. 77. 32. 102 Branchetus. 80. 65 Dasymallus. 87. 137. 138 Chorizobranchus. 151 Corynocephalus. 29 Chaponella. 135 Boguella. 72 Chrysothemis. 56. 36 Branchiomaldane. 120 Chiysopetalidae. 85 Crosslandiella. 61 Chaetacanthus. 81 Branchoscolex. 40 Clymene. 36 Dasybranchus. 25. 142. 144 Crossostoma. 52 Callizonella. 85 Clavadoce. 35 Capitobranchus. 148. 82 Clavodorum. 149. 82 Caobangia. 140 Dasychonopsis. 144 Disoma. 143 Chitinopomoides. 52 Cossura. 8. 32. 61 Drilognathus. 150 Capitella. 85 Drieschella. 82 Branielia. 41. 152 Dodecaceria. 13. 19. 144 Chloeia. 110 Colobranchus. 56. 65 Bylgides. 36 Dasynema. 28. 40 Clymenides. 43 Castalia. 110. 29 Cirrhineris. 28.7976 Bogueidae. 29-30 Cirratulispio. 142. 122. 41 Calamyzas. 27 Dispio. 85 Cystopomatus. 147 D Dalhousia. 8. 9. 150 Dextralia. 36 Capiteilidae. 20. 103 Colymmatops. 85 Calamyzidae. 76 Danymene. 104 Cryptopomatus. 143. 17. 13. 85 Cistena. 14. 120 CtenodrUidae. 117 Chone. 33. 133 Desdemona. 80. 110 Drilonereis. 111 Dasybranchetus. 48. 43 Dindymenides. 26. 36 Capltomastus. 48. 19. 29 Cenogenus. 133 Cystonereis. 80. 153 Dinophilus. 37 Ciymenopsis. 103 Chloenopsis. 21 Cossuridae. 26 Dipomatus. 77 CauUeriella. 118 Bucheita. 32. 138. 38. 35 Bushiella. 137. 8. 116 Bylgia. 142. 26 Cyanippa. 105 Dioplosyllis. 36 Dejoces. 71-72 Chrysopetalum. 85 Calcareopomatus. 8. 61 Drieschia. 36 Capitellethus. 8. 75. 32. 30 Cimformia. 58. 149. 58. 103 Dilepidonotus. 39. 12. 88 Ceraionereis. 51 Cirratulidae. 52 Clymaldane. 77 Dalhousiella. 74. 8. 112 Dorvilleidae. 137. 140 Dasylepis. 103 Chloraema. 112 Doyeria. 41 Ciymenella. 11. 82 Diplobranchus. 88 Cervilia. 142. 37 Branchiomma. 144 Dawbinia. 85 Brada. 135 Coretus. 8. 144 Coppingeria. 88 Dendrophora. 58. II. 142. 86. 75 Bookhoutia. 111 Ciiroceros. 39 Branchiocapitella. 98 Cleta. 28-29 Chaetoptenis. 144 Cryptonereis. 120 Cistenides. 49 Cirronereis. 111113. 140 Chaiybs. 16 Callizona. 129. 137. 138 Didymobranchus. 133 Demonax. 91 Cirrodoce. 117 Corephorus. 51 Cassandane. 11. 28 Chaetosphaera. 149. 31. 61 Chaetobranchus. 13. 140 Eaiymedusa. 82 Ehlersiella. 55 Halosydna. 45 Gymnonereis. 125 Euscione. 133 Euscleiocheilus. 137.91 Glycerella. 85 Grubeopolynoe. 73. 61 Eudistylia. 138 Eurato. 125. 51 Eteonides. 48. 93 Goniadella. 65 Evamella. 53 Eracia. 39. 93 Goibunovia. 85 Frennia. 107 Escholtzia. 79. 92 Eugrymaea. 62 Geminosyllis. 112 Exogone. 14. 103 Eupista. 92 Haplobranchus. 89 Gymnorhynchus. 150 Fauvelia. 40 Ettcranta. 121. 147 Eucaninculatus. 111 Halodora. 51 Eamolpe. 142. 76 H Haematocleptes. 71 EusylUs. 19. 30. 85 Gattyana. 144 Garjaiowella. 99 Enigma. 59. 120 Eupanthalis. 85 Gnibeulepis. 8. 16 Haplosyllides. 70 Flabellidemia. 59. 71 Eulepethidae. 85 Eurysyllis. 51 Glossopsis. 147 Fimbriosthenelais. 80. 115. 137. 58. 80. 61 Gattiola. 34. 104. 129. 139 FaUclandiella. 125 Giymaea. 49 Eteonella. 149. 82 . 143. 85 Gravierella. 93 Goniadidae. 52 Giubea. 102 Eusamytha. 87. 39. 118 Galathowenia. 47. 45. 57. 66 Euratella. 138 Euclymene. 109 Ereutho. 131 Eupolynoe. 8. 133 Eulaeospira. 125 Glyphohesione. 106. 55 Halla. 36 Elicodasia. 66 Eucaiphus. 58. 10. 139 Fabriciella. 11. 124. 82 Euthalenessa. 133 Cunnarea. 8. 80. 88 Eunice. 85 Euchone. 142. 144 Filopora. 48. 144 Filibranchus. 131 Eupolymnia. 75 Science Series 28 Ecamphicteis. 118 Ficopomatus. 68 Evimenia. 91. 148. 66 Eupomatus. 140 Eriphyle. 125 Echinoiiiyachus. 123. 69. 94 Ephesia. 82 Eurythoe. 133 Eupistella. 92 (T13) Glycinde. 45 Eumida. 140 Haploscoloplos. 49 Eumidia. 117118 Fauveliopsis. 94 Glyphanostomum. II.182 Dybowscella. 49 Gisela. 137. 105-107. 58. 8. 137. 123. 137. 8. 72 E NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Eimotomastus. 70 Euthelepus. S3 Haliplanes. 94 Epidiopatra. 82 Euglycera. 75. 70 Bisigella. 62 Halosydnoides. 147 Eupompe. 66 Euphione. 82 Exogonella. 103 Euphiosinidae. 17 Fauveldora. 10. 140 Gastroceratella. 131 Eone. 129. 40 Greeffia. 140 Fabriciola. 107 Eunicidae. 42. 13. 80. 51 Eranno. 116. 138 Eudontosyllis. 51 Fabricia. 91 Gymnosoma. 116. 62 Grubeosyllis. 128 Emamythella. 99 Eteone. 147 Glycera. 62 Gossia. 137. 118 Gwasitoa. 45 Eusigalion. 57. 140 Gorekia. 125 Ehlersia. 92 Glyceridae. 65 Gastrolepldia. 57. 51 Eupallasia. 133 Ehlersileanira. 80. 58. 92-94. 103 Eucerastes. 82 Fauveliella. 101. 91. 106 Eunoe. 61 Euphionella. 27 Enapteris. 68 Eulepethus. S3 Halithea. 138 Euchonella. 73 Fauveliopsidae. 69. 140 Dysponetus. 65 Exallopus. 68 Eulepis. 77 Haliplanella. 80. 57. 65 Halosydnopsis. 59. 114 Eclysippe. II. 115. 110 Enoplobranchus. 130. 61 Eulalia. 48. 118 Flabelligera. 137. 53 Hamiglycera. 131 Euzonus. 49 Eulalides. 81. 133 Eriographis. 117 Flota. 92. 8. 41. 68 Grubianella. 82 Exotokas. 51 Euleanira. 98 Ephesiella. 139 FalTisabella. 65 Eupolyodontes. 117 Flabelligeridae. 58. 82 Exogonoides. 110 Halimede. 70 Euphrosine. 143. 14. 85 Goniada. 113 Ethocles. 134 Filigrana. 121. 91 Gnathosyllis. 106 Euniphysa. 17 Globiodoce. 62 Halosydnella. 61 Friedericiella. 82 Genetyllis. 123. 93 Goniadopsis. 125 Egamella. 93. 82 Exogonita. 140 Gyptis. 61 Eupholoe. 150 Eulagisca. 52 Halogenia. 67. 12. 65 Eunereis. 147 Filograna. 61 Enonella. 80. 98 Ephesiopsis. 62 Halyplanes. 117 Enipo. 91-92. 93 (T13) Goniadides. 71 Euspio. 105 Epitoka. 61 Eunomia. 56. 144 Filogranula. 115 Galeolaria. 142. 57. 123. 79 Gnatholycastis. 81. 51 Eteonopsis. 115-117 Felmingia. 149. 87. 81. 117 Flabelligella. 69. 114. 51 Eumeniopsis. 98 Epicaste. 22 Euarche. 43 Evarne. 26 Eusthenelais. 8. 33. 129. 36 Eunotophyllum. 59. 121. 85. 79 Hesiocaeca. 114 Ichthyotomus. 57. 40 Josephella. 62 Heterospio. 140 Leptoecia. 51 Hyperhalosydna. 54. 129. 131 Lanicides. «5. 91 Lepidonotus. 126 Irmula. 85 Lamellisyllis. 125 Hypephesia.1976 Hapiosyllis. 36 Leipoceras. 76 Hesionella. 53. 34. 26. 85 Laranda. 11. 52 Kuwaiia. 32. 41 Heteromarphysa. 110 Lacharis. 131 Langerhansia. 42. 17 Labranda. 109 Iphitimidae. 39. 79 Hartmania. 126. 76 Hesiosyllis. 55 Laetmonioe. 85 Lanice. 131 Leanira. 59. 34. 72 Heterophenacia. 131 Laomedora. 137. 143. 58. 89 Kalaminochaeta. 87. 51 Hypoeulalia. 75. 27 Kebuita. 55 Hermionopsis. 102 Histriobdella. 121 Labidognathus. 111 Leaena. 54 Heterobranchus. 144 THE POLYCHAETE WORMS 755 Labotas. 65 Lepidasthenia. 147 Helicosiphon. 59. 8. 134 Hipponoa. 57. 104. 82 Hannopsides. 89 Leonnatus. 74 Hesionides. 105 Hyalopomatopsis. 75. 95-96 lospilopsis. 41 Iphinereis. 130. 137. Heteroterebella. 27 Heterospionidae. 89 Krohnia. 63 Lepidogyra. 80. 24 Laonome. 57. 80. 121. 36 Heteromastus. 40 Heterodisca. 73. 58. 39. 76 Leodice. 26 Helena. 63 Lepidopleunis. 124. 91 Hemipodia. 57. 151 Laetmatonice. 91 Iphione. 114 Idalia. 49 Hesiospina. 124. 127 Isoma^tus. 32. 8. 62 Hespeniphyllum. 97 Ilyphagus. 117 Intoshella. 74. 63 Hyboscolex. 103 Leocrates. 141. 113114 Histriodrilus. 100. 70 Hyalinoecia. 134 Leucia. 128 Heterocimis. 36 Isosyllis. 95. 34. 89 Hedyle. 62 Horstileanira. 65 Iphitime. 63 Kesun. 92 Hemipodus. 120 Hermenia. 44 Hydroides. 118 Inermonephtys. 66 Lepidoñmbria. 76 Leocratides. 71 Lepraea. 57. 56. 12. 31 Leiobranchus. 109-110 Irana. 44 Kefersteinia. 105 Leptoneieis. 23. 111 Laiymna. 14. 54 Lagis. 134 Heteropodarke. 105 Herdmanetla. 75. 56. 65 Laeonereis. 85 Janita. 62 Haipochaeta. 76 Hesionura. 147 Hyalopomatus. 57. 91 Hekaieiobranchus. 62 Heimella. 41 Leiochone.92 Hemisyllis. 29 Leptochone. 114 Hololepida. 30 Labrorostratus. 71 Hauchiella. 63 Lepidonereis. 27 Kalaminochaetidae. 76. 63 Lepidastheniella. 139 Laonomedes. 14. 36 Leiocephalus. 57. 94 Lacydoniidae. 73-77. 134 Leptochaetoptenu. 116. 85 Heptaceias. 62 Heimadion. 63 Lais. 40 Isolda. 150 Hemilepidia. 43 Laenilla. 63 Iphionella. 79 Labiaria. 108. 94 Ladice. 75. 62 Hermione. 65. 62 Haswellia. 140 L^iocapitella. 99 Johnstonia. 125 Hypaniola. 63 Lepidia. 85 Hesperonoe. 30 Hetenxlymene. 85 Laonice. 113. 134 Idanthyrsus. 56. 59. 143. 75. 91 HeteiDpale. 87. 76 Hesione. 17 Lacydonia. 12. 149. 129. 26 Lenora. 63 loda. 33. 133 Heterophyselia. 150 Leoimates. 148. 143. 94 Leontis. 27-28. 91 Lepadorhynchus. 75. 120 Lagisca. 140 Laphania. 81. 11. 137. 140 Hypsicomatopsis. 96 Iphianissa. 74. 95. 113. 76 Heteropolynoe. 83. 58. 131 Hediste. 144 Janua. 27 Kalununaria. 55 Hermundura. 8. 144. 88. 8. 151 Heteiomatdane. 76 Hesionidae. 47. 32. 149. 55 Heimodice. 57. 75. 34. 53 Hypania. 26 Kinbergia. 31. 8. 149. 84 IsociiTus. 65 Harmothoe. 107. 114 Histriobdellidae. 39. 62 Heminereis. 34. 129. 87. 51 Lepidametria. 55 Leucariste. 57. 94 Lacydes. 36 Heteronereis. 59. 129. 62 Hypociirus. 107 Heteromastides. 147 Hydrophane«. 57. 8. 58. 36 Leiochrus. 32. 94. 95. 56. 140 Hypsicomus. 70 Ledon. 111 Lalage. 139 I Ichthyotomidae. 84 Hesperalia. 85 lospilidae. 150 Jasmineira. 47. 91. 96 lospilus. 8. 83 Lanassa. 98 Hypereteone. 89 Letmonicella. 69. 130. 51. 69. 87. 89 Laeospira. 41 Leiochrides. 49 Heteraphrodita. 102 Hermonia. 12. 139 Johnstonella. 144 Kainonereis. 34. 51 Kinberginereis. 131 Laphaniella. 107 Leodora. 51 Hypsicomatides. 29 Heterosyllis. 109 Kynephorus. HI . 62 Hololepidella. 58. 76 Kermadecella. 36 Leiocapitellides. 43 Kinbergella. 134 Lapithas. 87. 152. 108. 8. 63 Macelloides. 22. 154 Meiodotviltea. 70 Mayeria. 39. 51 Myrianida. 40 Maldanidae. 126 Melinella. 145 Nephtyidae. 12. 59. 52 Lipephila. 77 Marenzelleria. 97 Micronereides. 16 Nais. 107 Neieilepas. 23. 109 Lycaretus. 111 Lysaretidae. 91 Mastobranchus. 91 Lygdamis. 139 Meganereis. 107 Nereidonta. 59. 123. 78 Loimia. 66 Neopodarke. 91 Lipobranchius. 49 Mystides. 136. 140 Metamphinome. 55 Minuspio. 87. 126 Melinnides. 65 Nectoneanthes. 129. 126 Mexieulepis. 12. 145 Mercierellopsis. 115 Myrioglobula. 103 Lophosyllis. 126 Neopanthalis. 13. 24 Moyanus. 151 Neoheteromastus. 126 Melinnexis. 39. 12. 24 Mitrana. 88. 75. 126 Mugga. 51. 111 Macrochaeta. 34. 108 Lumbrinendes. 123. 52. 128 Melinnoides. 91 Nauphanta. 57. 47. 121. 43. 26 Malacoceros. 23. 51 Maea. 154 . 36 Neopaiwa. 152. 85 Lucopia. 126 Melinnopsides. 40 Micronephthys. 75. 89 Neco. 131 Lysippe. 106 Lysilla. 36 Lumbriconereis. 102 Liocapa. 126 Melinnopsis. 137. 149. 24 Microsyllis. 39. 40 Lumbricomastus. 15. 86. 85 MUnesia. 36 Mauita. 70 Neoleprea. 34. 37-41. 16 Microsamytha. 87. 107 Litocorsa. 26 Mageiona. 47. 109 Lumbrineris. 123.184 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Mandane. 28 Mesocirrineris. 121. 86. 41 Nectochaeta. 145 Mesochaetopterus. 85-91. 151 Mastigethus. 63 Lugia. 107 Macellicephala. 87. 58. 89 Microphthalmus. 32. 130. 57. 64 Neoamphitrite. 38. 126. 89. 148. 107-109. 134 Lumbriclymene. 69. 29 Metachone. 56. 137. 129. 53 Mayella. 32. 121. 143.38 Maldanopsis. 52 Naidonereis. 145 Neopygospio. 97. 140 Naiades. 96. 23. 49 Myxicola. 44 Lirione. 76 Microrbinia. 56. 48. 43 Naraalycastis. 85 Monorchos. 88. 123. 95. 123. 139. 143. 69. 126 Muggoides. 103 Metavermilia. 86 Nereidice. 152. 8. 26 Mandrocles. 20 Monroika. 139. 127 Neottis. 89 Lycidice. 40 Micromaldane. 88. 32. 23. 106. 124. 51 Mesonerilla. 51 Myriochele. 53 Lophocephalus. 32. 145 Mexamage. 126 Melinnampharete. 26 Neosabellides. 75. 126 Myriana. 8. 118. 76 Neopomatus. 8. 40 Macrophyllum. 49 Nereis. 128 Microserpula. US Monocerina. 46. 119. 132 Neolipobranchius. 17 Nainereis. 139 N Science Series 28 Leucodora. 89 Namanereis. 52 Maupasia. 142. 107 Marsipospira. 126 M Macduffia. 143. 117 Lophonota. 91 Lycastoides. 97 Nereidae. 147 Mera. 142. 26 (TIO) Magelonidae. 87. 5253 Lopadorhynchus. 47. 150 Metalaonome. 30. 68 Microclynaene. 63 Melinantipoda. 106. 26 Levidorum. 8. 100. 130. 24 Maldane. 49 Lumara. 91 Nematonereis. 34. 78 Loandalia. 58. 75. 63 Macellicephaloides. 124. 147 Microspio. 130. 31 Macroclymene. 39. 107 Nausicaa. 115 Myriowenia. 89 Naneva. 121. 34. 36 Lumbriconais. 105 Nerilla. 134 Neovermilia. 41 Malmgrenia. 123. 131 Naraganseta. 13. 41 Mania. 28 Lopadorhynchidae. 151 Mercierella. 103 Lithognatha. 107 Lycoris. 96-97 (TI3) Nephtys. 85. 34. 89 Nectonereis. 79. 30 Naumachius. 24 Mesotrocha. 43. 145 Marphysa. 32. 103 Lycastella. 143. 149. 126 Melinnata. 151 Neonotomastus. 63 Maclovia. 56. 85 Nereitube. 33. 107. 140 Metalaeospira. 40 Lumbriclymenella. 129. 140 Megalomma. 87. 154 Mesospio. 109 Lumbrineridae. 76 Micropodarke. 11. 137. 112 Melaenis. 63 Manayunkia. 8. 118 Lysarete. 36 Neohololepidella. 114. 24 Marifugia. 34. 131 Melinna. 128 Membranopsis. 91 Meganerilla. 87. 38. 23. 36 Neomeris. 115 Mysta. 120 Monostylos. 36 Megachone. 107 Neanthes. 121. 53 Mastigonereis. 91 Nereimyra. 109 Lumbrineriopsis. 39. 63 Malmgreniella. 91 Lycastopsis. 131 Neodexiospira. 90 Nereisyllis. 91 Lycastis. 83 Myriacyclum. 145. 131 Longosoma. 123. 51 Mesomystides. 140 Monticellina. 98 Levinsenia. 15. 64 Neoleanira. 8. 18 Linophenis. 89 Micronereis. 14. 124. 107 Mediomastus. 30 Morants. 38. 107 Nemidia. 40 Macroclymenella. 26. 111 Lysidice. 123. 43 Neomicrorbis. 44 Neomediomastus. 76 Nereiphylla. 127 Neosamytha. 126 Lysippides. 30 Mesoeulalia. 40 Maldanella. 121. 75. 114. 140 Otopsis. 51 Nothria. 83 Phenacia. 73. 77 Ophioglycera. 85 Pagenstecheria. 145 Omphalopomopsis. 149. 127 Paleanotus. 8. 16 THE POLYCHAETE WORMS 1&5 Parapolynoe. 8. 129. 143. 53 Peisidice. 51 Paiahalosydna. 38. 102. 105 Oophylax. 68 Parapionosyllis. 38. 127 Phyllodoce. 14. 109 Nipponophyllum. 15. 124. 77 Parahololepidella. 50 Notaulax. 124. 83 Nychia. 127 Paramphinome.W6 Nerillidae. 80. 80. 39. 26 Nevaya. 107. 28 Phyllocomus. 10. 100 Nudisyllis. 139 Pantoithrix. 23. 140 Oriades. 83 Parautolytus. 127 Olga. 80. 36 Paralepidonotus. 81. 110. 8. 91 Notalia. 151 Paradiopatra. 17-lS Paraonides. 111 Notopygos. 147 Paraxionice. 72 Pallasia. 48. 132 Nicomache. 143. 72 Palmy reuphrosyne. 127 Paramarphysa. 152-154. 66. 66 Pholoididae. 77. 100. 143. 110. 43 Norepa. 134 Palmyra. 87. 70 Pholoides. 102 Parexogone. 114-115. 120 Pectinariidae. 253 Neríllidíum. 48. 79 Phyllampharete. 70 Pareurythoe. 41 Nicomedes. 90 Nicotia. 102 Paranaitis. 154 Paranerine. 58. 44 Paraserpula. 154 Nerillidopsis. 104. 32. 115 Oweniidae. 119. 124. 18 Paradyte. 19. 104. 95. 140 Parascierocheüus. 142. 130. 85 Ninoe. 105 Parophryotrocha. 130. 129. 130. 85 Perinereis. 119 Pharyngeovalvata. 132 Paiatyposyllis. 34. 134 Odontosyllis. 50. 59. 57. 35. 71. 41 Petaloproctus. 68 Peisidicidae. 83 Odysseus. 124. 145 Pectinaria. 103 Pherecardites. 85 Paiwa. 75. 107 Paramphicteis. 8. 77 Parathelepus. 104-105 Onuphis. 67 Photocharis. 118. 129. 118. 64 Paraeulalia. 42. 43 Ophiodromus. 102 Novaquesta. 68 Pareupholoe. 30 Peresiella. 81. 79 Paranereis. 107. 45 Otigognathus. 83 Ops. 18 Parapholoe. 20 Pareulepis. 85 Parheteromastides. 36 Perialla. 36 Noiophyllum. 31. 95. 115 Oxydromus. 117 Papillaria. 18 Paraonidae. 35. 102. 83 Parasabella. 91 Paramage. 36 Parheteromastus. 8. 132 Opisthosyllis. 12. 14. 91 Nicon. 91 Paranerilla. 145 Olgahannania. 137. 81. 35. 26 Parachloeia. 78 Owenia. 11. 65 Paronuphis. 65 Paraprionospio. 103 Palmyridae. 71. 41 Parergodrilidae. 77 Pabits. 18 Paraonis. 65 Omaria. 105 Paranychia. 132 Paraxiothea. 145 Oncoscolex. 47. 8. 123. 13. 140 Oriopsis. 14. 142. 145 Parasphaerosyllis. 66 Pelagobia. 124. 112 Parthenope. 77 Otanes. 20 Paumotella. 35. 8. 48. 97 Pelogenia. 117 Phisidia. 64 Paralycastis. 94 Ophiuricola. 96 Paralacydonia. 85 Phragmatopoma. 111 Oeorpata. 116.20 Parergodrilus. 26 Paranorthia. 102 Paracbonia. 51 Orbiniella. 145 Oligobranchus. 65 Paraonella. 50 Nitetis. 10. 80. 78. 134 Pherea. 64 Parahesione. 138. 96 Phalacrostemma. 26 Nerinopsis. 24 Parapterosyllis. 26 Nerinides. 140 Paradexiospira. 85 Ophelia. 106. 50 Notoproctus. 105 Nossis. 105 Paradoneis. 47. 139 Orseis. 19 Novobranchus. 32. 41^3. 27 Nicolea. 103 Pherasa. 13. 69. 116. 42 Ophelina. 127 Phyllamphicteis. 76 Orthodromus. 52. 120 Pallonia. 127 Phyllochaetopterus. 123. 73. 36 Notomastus. 83 Opisthopista. 34. 134 Nuchubranchiata. 104 Onuphidae. 64 Petaloclymene. 134 Oenone. 140 Oridia. 32. 111 Notodasus. 83 Petta. 8. 106. 43 Opheliidae. IS Oria. 26 Periboea. 42. 80. 91 Phalacrophorus. 132 Paraleiocapitella. 154 Nerine. 119 Phronia. 8. 45 Oniscosoma. 15. 88. 124.72 Palola. 127 Paedophylax. 32. 100. 41 Petitia. 85 Parasyllidea. 32. tOO. 50 Parandalia. 71 Pentacirrus. 53 Pellucidaria. 140 Nothis. 75. 41 Notopsilus. 132 Pholpe. 16 Orbiniidae. 149. 64 Paraiospilus. 12. 112 Opisthodonta. 110 Oligolepis. 109 Ophryotrocha. 43 Omphalopoma. 142. 120. 107 Palpiglossus. 134 Pherecardia. 40 Nicomachella. 8. 120-121 Peduiosoma. 127 Patmensis. 58. 14-17. 76. 66 Panousea. 105 Notocirrus. 107 Panthalis. 134 Othonia. 69. 115 Orbinia. 65 O Octobranchus. 93. 10. 59. 100. 83 Parasitosyllis. 120 Pallasina. 94 Paralaeospira. 83 Paraveimilia. 10. 121 Phaetusa. 80. 83 . 81. 65 Northia. 13. 151 Paralanice. 90 Perolepis. 36 Parhypania. 81. 41 Rhodopsis. 53 Pontogenia. 57. 140 Pseudohalosydna. 155 E>rotodrilus. 146 Pomatoleios. 13. 65-66 Polyophthalmus. 85. 16 Protoariciella. 25 Questa. 111 Plotobia. 132. 79. 139. 85 Pterothrix. 146 Pseudosyllides. 54 Poitoa. 18-19 Quetieria. 12.9. 130. (TÍO) Pixellia. 80. 91 Protomastobranchus. 22. 73 Pisionura. 23. 100 Plotohelmis. 46 (T-13) Phyllohattmania. 142. 127 Pteroscolex. 145 Placostegopsis. 142. 65 Pseudopomatoceros. 65 Ramex. 25 Rhynophylla. 39. 133 Pseudovefmilia. 51 Protodorviliea. 65 Romanchella. 48. 36 Pseudochitinopoma. 64 Phyllonereis. 66 Poiyodontidae. IS. 85 Polymnia. 26 Pseudomalacoceros. 84. 149. 147 Plakosyllis. 14. 141 Pseudosabellides. 10. 117 PoUicita. 83 Procerastea. 112 Protcxlrilidae. 151 Protoleodora. 107 Pseudopolydora. 74. 85 Polynoe. 10. 124. 64 PhyllosylHs. 35. 41 Procome. 43. 53 Pontodoridae. 128 Pseudoscalibregma. 27. 72-73 E^sionidens. 27 Püeolaria. 50 Protopolydora. 33. 153. 55. 92. 43 Polyphragma. 147 Podonereis. 22. 65 Phyzelia. 115 Psammodrilidae. 12. 24 Polycimis. 20-21 Psamraodriloides. 25 Pseudopolynoe. 26 PterociiTus. 140 Potaspina. 84 Rhynchobolus. 38. 45-51. 132 Ranzania. 23. 83 Platyneteis. 8. 41 Praxillura. 15. 151 Ehotolycoris. 8. 103 Pseudexogone. 117 Poeobius. 80. 51 Portelia. 23. 103 . 130. 154-155 Polygordius. 140 Potamis. 52 Plotolepis. 133 Pseudeulalia. 155 Protodriloides. 44 Pomatoceropsis. 8. 22. 73 Pisionella. 8. 48. 111 Pulliella. 8. 154 Psectra. 24 Proboscidea. 35. 151 Progoniada. 139 Potamilla. 139 Pottsiscalisetosus. 27 Poecilochaetus. 35. 21 Psammolyce. 146 Pseudopotamilla. 36 Puparia. 50 Proplacostegus. 149. 91 Phyllosheila. 36 Protomystides. 50 Pterolysippe. 148. 26 Protis. 117 I^gophelia. 70 Psammoriedlia. 117 Pisenoe. 80. 66 Reteterebella. 10. 23. 27 Poeobiidae. 143. 113 Prionospio. 88. 46. 84 Prochaetoparia. 52 Rhynchospio. 32. 141. 85 Protolaeospira. 77 Podarmus. 98 Polybostrichus. 140 Science Series 28 Phyllodocidae. 15. 73 Praxilla. 147 Piromis. 154. 132. 30 Prophyllodoce. 84 Pseudobranchiomma. 50 Pseudeupanthalis. 73 Pisionidae. 8. 22. 19 Questidae. 16 EVospio. 36 Reibischia. 72. 138. 42. 56. 9. 26 Pseudonotophyllum. 58. 146 Psymobranchus. 37 Pycnoderma. 71 Pseudeurythoe. 12. 86. 29 Raphidrilus. 26 Robertianella. 64 Polynoidae. 24. 64 Podioceros. 77 Podarkeopsis. 140 Psammocollus. 134 Protula. 149. 15. 141. 88. 39. 59. 33. 90 Platysyllis. 43. 8. 30 Pseudoclymene. 155 Polylepis. 147 Pseudocapitella. 12. 24 Polyeunoa. 115. 149. 91 Pisione. 57. 44 Pseudoserpula. 129. 8. 66. 127 Pterautolytus. 147 Pterampharete. 41 Pseudofabricia. 83 Pirakia. 91 Prionognathus. 30 Rashgua. 20. 51. 129. 48. 85 Pleione. 55-65 Poiyodontes. 132 Polynice. 138. 146 Rhopalosyllis. 50 Proclea. 151 Rostrada. 146 Pomatostegus. 151 PUlaiospira. 37 Protocarobia. 151 Pionosyllis. 50 Piratesa. 143. 41 Praxithea. 51. 8. 85 Pteriptyches. 22. 58. 48. 146 Pseudocirratulus. 90 Pseudonerine. 35. 64 Praegeria. 128. 142. 20 RariciiTUS. 75. 8. 149. 130. 69. 43 Pygophyllum. 155 Ftetogrubea. 41 PraxUlella. 146 Pontodora. 39. 58. 75. 37 Pterosyllis. 91 Poecilochaetidae. 64 Polygoidiidae. 149. 104. 41 Promenia. 105 Rhodine. 72 Pista. 36 Pseudoleucodore. 80. 13. 153. 92 Rhynchonerella. 123. 79. 129. 83 Phylo. 26 F^gospio. 116. 132 Polydora. 146 Protulides. 8. 29. 152 Placostegus. 56. 147 I^scalibregma. 81. 78 Pilearia. 29. 72 Pseudampharete. 32. 55. 25 Polydorella. 44 Proscoloplos. 140. 53 Restio. 85 Polybranchia. 8. 79. 77 Pseudothelepus. 137. 97 Potamethus. 26 Protothelepus. 24 Pseudonereis. 85 Pseudosyllidia. 65 Podarke. 21 Psammodrilus.]86 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Pomatoceros. 16 Physalidonotus. 142. 58. 35. 138. 71 Polymastus. 142. 48. 103 Plioceras. 94 Promaldane. 64 Polynoella. 17 Protocapitella. 132 Rhamphobrachium. 64 Pseudolacydonia. 134 Polymniella. 92 Proceraea. 94 Pseudoleiocapitella. 51 Pseudopalolo. 137. 134 PUargiidae. 85 Prodexiospiia. 132 Proclymene. 29 Ranzanides. 142. 32. 57. 48. 12. 147 Poiyphysia. 146 Protoaricia. 38. 77-79 Pilargis. 117 Sttophaga. 148 Spirorbides. 43 Travisiopsis. 18 Tecturella. 149. 78 Sige. 133 Stygocapitella. 113 Stauroceps. 142. 127 Samythopsis. 124. 85 Sagiteila. 26 Spiophanes. 147-152. 77 Stephanosyllis. 9.1976 RuUierinereis. 30 Thaumastoma. 128 Sosanopsis. 109 Scoloplella. 84 Sphaerephesia. 12. 90 Rytocephalus. 147 Sphaerosyllis. 95. 117 Telake. 128 Schistomeringos. 45 Scalibregmidae. 25 Spiramella. IS Sabina. 8. 141 Skardaria. 81. 45 Scalis. 57. 27 Thelepella. 137. 44. 113 Schizobranchia. 96. 140 Spirorbella. 43 Tetraglene. 120 Thalassema. 133 Staurocephalus. 25 Spiochaetopterus. 98 Sphaerodoropsis. 92 Telamone. 100. 147 Syllia. 98 Sphaerodorum. 84 Sabaco. 79-85 Syllides. 133. 36 Sangiria. 85 Schmardiella. 124. 65 Terebella. 129. 134 Thelephusa. 21. 80. 128 THE POLYCHAETE WORMS 187 Subprotula. 14. 104 Spintheridae. 77 Stschapoveila. 146 Salvatoria. 134 Scionelia. 123. 35. 149. 147 Spirodiscus. 50 Siliquaria. 69. 147 Simplicaria. 140 Sabellonga. 44. 149. 8. 114 Stemaspis. 65 Scalispinigera. 85 Syllis. 43 Tainokia. 119. 147 Spirographis. 87. 74. 36 Semiodera. 103 Stratiodrilus. 84. 133 Streblospio. 113 Steggoa. 17 Theostoma. 68-71. 111 Talehsapia. 8. 45 Sclerostyta. 85 Tetratrocha. 128 Sandanis. 91 Thormora. 103 Saphobranchia. 133 Terpsichore. 8. 10. 123. 98 Sphaeropomalus. 132 Spinther. 32. 133 Teiebellidae. 43. 148. 17 Therochaeta. 141 Sabellongidae. 42. 103 Thoe. 112. 141. 69. 17 Scoloplos (Scoloplos) 16. 44. 8. 8. 142. 70 Tharyx. 74. 127 Sabellina. 142. 12. 124. 151 Spirorbidae. 124. 152 Siphonostoma. 51. 8. 44 Scaiibregmella. 90 Stephania. 65 Thylaciphorus. 85 Thysanoplea. 100. 25 Streptosyllis. 48. 129. 105 Travisia. 8. 114 Schroederella. 65 Tachy try pane. 99 Torquea. 129. 42. 113 Stauronereis. 143. 51 Trapodia. 69. 155 Saccocirrus. 128-134 Terebellides. 130. 77 Sylline. 77 Schreiberius. 152 Strategis. 134 Terebellobranchia. 119 Sabellastarte. 96. 37 Telolepidasthenia. 141. 140 Schizocraspedon. 129. 103 Thetisella. 107 TimaiBte. 116. 30 Tomopteridae. 117 Thesmia. 141. 146 Schmardanella. 129. 58. 48. 17 Scione. 98 Sphaerodoce. 97. 117 Scalibregma. 70 Sigalionidae. 122. 23. 43^5 Scalibregmides. 39. 95. 148. 41 Sabella. 44. 26 Spionereis. 70 Sthenelanella. 121 Scalisetosus. 117 Subadyte. 8. 25 Scoletoma. 137. 136 Sabellides. 87. 50 Steninonereis. 65 Telonereis. 80. 23. 117 Siphostoma. 146 Salmacinopsis. 28 Spione. 8. 52 Trachelophyllum. 91 Tetreres. 80. 147 Tenonia. 123. 78 Taphus. 130. 13. 11. 98-99 Tomopteris. 101. 127 Samythella. 26 Scoleiepis. 136. 8. 116. 29. 142. 114 Sthenelais. 84 Synelmis. 129. 54 . 134 Sonatsa. 152 Spirorbis. 77 Teiepsavus. 142. 10. 117 Serpula. 134 Thelepus. 70 Sthenolepis. 141 Sabidius. 15. 41 Sosane. 133 Teiebellanice. 57. 57. 151 Spirovenna. 117 Therochaetella. 17 Scyphoproctus. 90 Solowetia. 129. 129. 151 Sinistrella. 154 Thalenessa. 78 Synsyllis. 85 Stemaspidae. 137. 22. 65 Sigalion. 134 Thelepides. 23. 113 Telothelepus. 10. 148 Sheila. 8. 80. 137. 140 Saccocirridae. 140 Sabellidae. 48. 143. 141-147. 147 Spirobranchus. 59. 100. 72 Tauberia. 133 Temporaria. 80. 134 Sclerobregma. 153. 99 Tricertia. 69 Sigambra. 114 Thalassochaetus. 132 Scionopsis. 87. 10. 44. 114 Stratonice. 23. 128 Sosanella. 123. 98 Tibiana. 134 Schmardia. 134 Thelepodopsis. 99 Salmacina. 130. 8. 8. 11. 17 Scoloplos (Leodamas) 16. 14. 84 Struwela. 133 Theodisca. 146 Scolecolepides. 85 Syllidae. 155 Sacconereis. 22-26 Spionides. 53. 26 Spionidae. 42. 123. 9798 Sphaerodoridium. 11. 128 Spermosyllis. 140 Sabellaria. 29 Telethusae. 91 Sireblosoma. 84 Syllidia. 85 Samytha. 85 Thoosa. 11. 15. 130. 45 Schistocomus. 104 Spio. 58. II. 25 Scolecolepis. 8. 50 Sphaerodoridae. 132 Scionides. 118. 113. 128 Sosanides. 45 Sclerocheilus. 120 Sabellariidae. 118-120. 29. 135-141. 20 Stylarioides. 134 Torrea. 97 Spinosphaera. 22 Solomononereis. 147 Serpulidae. 75. 70 Stoa. 143. 99-100 Typhloscolex. 17 Venusia. 90 Tylorrhynchus. 11. 26 Unciniseta. 143. 134 Watelio. 14. 81. 65 Trilobodrilus. 91 Uncinia. 84 Tylonereis. 22. 123. 14. 85 Xenosyllis. 8. 147 Venniliopsis. 140. 26 Trochochaetidae. 69. 59. 90 Uncinerels. 70 ^ . 134135 Trichobranchus. 85 Tricoelia. 141 Trichosyllis. 81. 52 Veleda. 25 Trochochaeta. 84 Tyrrhena. 51 W Wartelia. 87. 8. 103 Zygolobus. 109 Zygophyllus. 81. 109 Science Series 28 Trichobranchidae. 84 ^ Zeppelinia. 134. 51 52 Webena. 56.188 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY U Umbellisyllis. 134. 134 Uncopolynoe. 109 Uncinochaeta. 103 Venadis. 65 ^^ÏT'T^i' o« Weddellia. 79. 147 Zorus. 99. 84 Unanereis. 77 Vitiaziphyllum. 122. 51. 48. 87. 117 Ttypanosyllis. 152 Tripolydora. 50 Xenosyilides. 117 Trophoniella. 134 Vennilia. 116. 147 Virchowia. 135 Trichosobranchella. 135 Urosiphon. 21. 90 Typhlonereis. 8. 20 Zopyrus. 117 Zothea. 12S Wesenbergia 75 77 Wüleysthenelais. 79. 26-27 Troglochaetus. 43 V Valla. 36 Vanadis. 95. 87. 154 Trophonia. 91 Typhloseolecidae. 65 Unobranchus. 29 Tricosmoehaeta. 85 Vitiazia. 48. 99 Typosyllis. 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