Wetland Bryophytes• • • • • What are bryophytes? Why are bryophytes important in wetlands? What are the different groups of bryophytes? Sphagnum Bryophyte preservation What are bryophytes? Bryophytes vs. Tracheophytes Bryophytes • Typically small • Do not have xylem and phloem • Non-lignified • No roots! • External transport • Thrive in hostile and barren habitats • Biochemical diversity • Gametophyte is dominant Tracheophytes • • • • Waterproof covering – cuticle Have xylem and phloem Vascular system is lignified Flowering plants, ferns, conifers, etc. 1 000 species world-wide • Bryophyta: Sphagnum and the “true moss” • Marchantiophyta: liverworts • Anthoceroptophyta: hornworts true moss Sphagnum liverworts hornworts Bryophyte lifecycle: an adaptation to success Bryophytes Plants (tracheophytes) The advantage of a dominate gametophyte • • • • • • Bryophytes are immobile Have different life cycle strategies from other plants Immediate fitness – If change is favorable Immediate selection – If change is unfavorable Thus bryophytes can express new genes in the generation they first occur Diploid plants mask deleterious genes 2 .The Bryophytes • Believed to be the first land plants • ~ 20. The “large” large” scale of bryophytes • Bryophyte influence on the environment – Hydrology – Nutrient cycling and nitrogen fixation – Decomposition – Succession – Stabilizing soil – Biomass production – Carbon fixation Global climate modelers are realizing that peatlands have a dramatic effect on global temperatures and water movement . Net primary production (in g m -2 yr -1) for wetland types Trees Vascular plants Bryophytes Above ground productivity in fens and bogs in Alberta. Canada Categories of Wetland Bryophytes • • • • • Peat mosses Brown mosses Feather mosses Liverworts Additional mosses Ptilium crista-castrensis 3 . Feather moss Brown moss Peat moss Liverworts “Other mosses” Number of Sphagnum (peat moss) and other bryophytes in a peatland from western Canada (Vitt and Belland 1995) Extremely rich fen Moderately rich fen Poor fen Bog Sphagnum 9 7 16 12 Other bryophytes 58 28 28 41 Total 67 35 44 53 “Brown mosses” Calliergonella cuspidata • • • • • • Not a taxonomic classification Associated with rich minerotrophic fens Brown. reddish. and golden Indication of higher pH – alkaline conditions Susceptible to N deposition Commonly associated with sedges 4 . yellow. Tomenthypnum nitens Feather mosses • • • • Coniferous forest Swamp forests Shaded areas Prefers hummocks and drained sites Hylocomium splendens Nitrogen Fixation Hyaline cells colonized by Cyanobacteria Coiled chains of Nostoc are hidden in the leaf under light microscopy.ch/mit chell/edward/Sphagnu m.epfl.htm Pleurozium schreberi 5 . but are readily observed as the red cells under ultraviolet-fluorescence microscopy http://wslar. 5 -2.5 kg / ha yr – previous estimated yearly fixation by all vegetation Pleurozium schreberi was recently found to contribute1. scherberi • • • • • • • • • Biological nitrogen fixation Boreal forest nitrogen supplies? Feather mosses contribute to 60 -80% of ground cover in boreal forest Pleurozium schreberi – common worldwide Boreal bryophytes provide greater primary productivity then overstory vegetation 3 kg / ha yr – Organic carbon accumulation in boreal forest . hummocks Liverworts • Leaves often lobed • Dorsi-ventral organization • Two body plans leafy or thallose 6 . swamps and fens • Dicranum spp.0 Kg / ha yr fixation Implications for forestry management in boreal regions Other mosses Bryophytes that do not fall into the previous categories • Polytrichum spp.Nitrogen fixation by P. Sphagnum Sphagnum facts 2 Sphagnum than in • More living carbon in any other genus of plants worldwide • Sphagnum in northern peatlands – Covers 1.5 Mkm2 – Stores 150 Gt of carbon • Ecosystem engineer • Resistant to decay • Genus contains many species – ~100 in the northern hemisphere – ~ 19 in the Keweenaw peninsula 7 . 3. 2. Lack a protonema Have single cell rhizoids Have two ranked leaf arrangement Mosses are usually 3 or 5 ranked Peatmoss a.a. moss Liverworts differ from mosses in three main ways 1.k.Liverworts vs. Capitulum Pendent branches Divergent branches (fascicles) Stem NO roots or rhizoids ! Sphagnum cell structure Water adaptations 25x Weight in water One cell in twenty is alive Sphagnum leaf Pores Photosynthetic cells Hyline cells Fibirals http://kentsimmons.Global distribution of peatlands Structure of Sphagnum The Head .uwinnipeg.jpg 8 .ca/2152/sphagnum3. 0 cuspidatum Differences in Sphagnum species in a hummock – hollow complex in northern Michigan.1 capillifolium 3.Chemical properties of Sphagnum Coping with a harsh environment • Often grows in low nutrient areas • Creates cation sites • Contains secondary compounds – Phenolics • Can translocate metabolites Cation exchange H+ + OH CO OOH+ C O CO COOH+ CO OH + pH Ca++ + H H+ Active substances are uronic acids -galacturonic acid and 5KMA The uronic acids sometimes referred to as sphagnan High pH .2 magellanicum 24” recurvum 3. (redrawn from Crum1992) 9 .phenolics such as sphagnum acid are active Microtopography of Sphagnum pH 2.9 fuscum 3.5 4. When re-wet after months or years of being dry.berkeley.herb. They can be completely dried at the cellular level and recover completely.edu/bryolab/pww/ Cold tolerance Increase sugar as natural antifreeze Species of Sphagnum in the Keweenaw peninsula • ~119 species of Sphagnum worldwide • 19 species of Sphagnum in the Keweenaw • Divided into sections based on: – Morphology – Growing conditions 10 . they begin photosynthesis within 5 minutes! •http://ucjeps. Sphagnum fuscum 2-4 d Sphagnum papillosum 2-4 d Sphagnum balticum 2-4 d Sphagnum cuspidatum 2-4 d Sphagnum magellanicum 2-4 d Sphagnum magellanicum 14 d Sphagnum fallax 14 d Fontinalis flaccida 3 mos Barbula torquata 18 mos Oxymitra 4 yrs Riccia canescens 7 yrs Grimmia laevigata 10 yrs Syntrichia ruralis 14 yrs 3 Tortula muralis protonema 14 yrs Riccia macrocarpa 23 yrs • Mosses are among the most desiccation-tolerant of all plants.Desiccation • • Known durations of desiccation survival in bryophyte plants. jpg Sphagnum papillosum • • • • • • • • Golden brown lawns Plump and swollen Short stubby spreading branches Brown stems Wet and acidic habitat Forms lawns and low hummocks Followed by S.jpg 11 .gov/gallery/large/spsq70_004_lhp.jpg/$FILE/10378.odu.gov/gallery/large/spsq70_006_lhp.usda.usda.edu/webroot/instr/sci/plant.nsf/files/10378.jpg Sphagnum squarrosum http://plants. magellanicum Poor fens and acid lakes http://web.Sphagnum squarrosum • Medium to robust • Squarrose leaves – widely spreading • Large terminal capitulum • Prefers depression • Shade and moist areas • Thuja swamps http://plants. jpg Sphagnum magellanicum • • • • • • • Robust and large Grows in partial shade and in the open Red produced as sun response Initiates hummocks Poor fen to bog Moist to dry sites http://plants.usda.jpg Common and widespread 12 .Sphagnum papillosum Sphagnum magellanicum sphagnum magellanicum http://plants.gov/gallery/large/spma70_004_lhp.usda.gov/gallery/large/spma70_006_lhp. bd.lst. shaggy.Sphagnum magellanicum Sphagnum cuspidatum • • • • • • Medium to slender Grows in wet depressions Can grow submerged Long branches and leaves White to yellow Looks like a wet cat – wet matted masses Sphagnum wulfianum •Robust growth •Clover-like capitulum •Green to yellow •Brittle stem •Common •Grows almost exclusively in Thuja swamps •Has a loose.se/publishedObjects/10001510/bollvitmossa. top-heavy appearance http://www.jpg 13 . nps.Sphagnum girgensohnii • • • • • • Vibrant green Large with flat top capitulum When dry white streaks visible on leaves Thuja swamps Long graceful spreading branches Capitulum appears 5-rowed (star shaped) http://plants.usda.jpg Vulnerability of bryophytes • • • • • • Sensitive drought Have a low growth rate Very sensitive to pollution Lack of “image” Are not large and charismatic Lack of understanding of how they contribute to ecosystem functioning 14 .gov/gallery/pubs/spgi70_001_php.jpg Sphagnum girgensohnii http://www.gov/olym/crypto/M_SPGI. Threats to Bryophytes • • • • • • • Deforestation Forest cultivation Urbanization Land reclamation Road construction Wetland drainage Invasive species Peatland loss • • • • Agriculture 250.000 km2 Does not include tropical peatlands Peatland drained or altered for agriculture Europe Agriculture % country’s peatland Russia 70400 12 Germany 12000 85 Belarus 9631 40 Poland 7620 Ukraine Agriculture % country’s peatland Indonesia 37200 18 USA 17100 16 West Malaysia 5760 71 70 China 1930 13 5000 50 Canada 1410 1 Sweden 3000 5 Sarawak 1370 12 Finland 2000 2 Netherlands 2000 85 Norway 1905 8 Lithuania 1900 39 Iceland 1300 13 Estonia 1300 12 Total 125000 Total 65000 15 .000 km2 Peat harvesting 50.000 km2 Forestry 150. org/index.bryology.edu/ • International Association of Bryologist – http://www.gov/ 16 .edu/~abls/ • Moss images – http://plants.mtu.4g/m /yr • Commercial harvest rate of at least 21 years Web sites • Dr.bryoecol.unomaha.Bryophyte harvesting not including peatlands • A nontimber forest product • Commercial importance increasing • Harvested – – – – Soil conditioner Cultivation medium Packing material Floral decorative • 6 million dollar industry in the US • With illegal harvest it is estimated as a $15 million industry in the Northwest and Appalachian region Sustainable? 2 • Can take several decades for regeneration • Accumulation rate 22.html • American Bryological and Lichenological Society (ABLS) – http://www.usda. Janice Glime’s Online Bryology book – http://www. 17 .