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1 Wetland Bryophytes What are What are bryophytes? Why are bryophytes important in wetlands? What are the different groups of bryophytes? Sphagnum Bryophyte preservation What are bryophytes? What are bryophytes? Bryophytes vs. Tracheophytes Waterproof covering – cuticle Have xylem and phloem Vascular system is lignified Flowering plants, ferns, conifers, etc. 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 Bryophytes
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    Wetland Bryophytes

    What are What are bryophytes? Why are bryophytes important in wetlands? What are the different groups of bryophytes? Sphagnum Bryophyte preservation

    What are bryophytes?What are bryophytes?Bryophytes vs. Tracheophytes

    Waterproof covering cuticle Have xylem and phloem Vascular system is lignified Flowering plants, ferns,

    conifers, etc.

    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

    TracheophytesBryophytes

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    Believed to be the first land plants ~ 20,000 species world-wide Bryophyta: Sphagnum and the true

    moss Marchantiophyta: liverworts Anthoceroptophyta: hornworts

    liverwortshornworts

    true moss

    Sphagnum

    The BryophytesThe Bryophytes

    Bryophyte lifecycle: an adaptation to successBryophyte lifecycle: an adaptation to success

    Plants (tracheophytes)Bryophytes

    The advantage of a dominate gametophyteThe 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

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    Global climate modelers are realizing that peatlands have a dramatic effect on global temperatures and water movement.

    The The largelarge scale of bryophytesscale of bryophytes

    Bryophyte influence on the environment Hydrology Nutrient cycling and nitrogen fixation Decomposition Succession Stabilizing soil Biomass production Carbon fixation

    Net primary production (in g m -2 yr -1) for wetland types

    Bryophytes

    TreesVascular plants

    Above ground productivity in fens and bogs in Alberta, Canada

    Categories of Wetland BryophytesCategories of Wetland Bryophytes

    Peat mossesPeat mosses Brown mossesBrown mosses Feather mossesFeather mosses LiverwortsLiverworts Additional Additional

    mossesmosses Ptilium crista-castrensis

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    Feather moss

    Peat moss

    Liverworts

    Brown moss

    Other mosses

    Number of Number of Sphagnum Sphagnum (peat moss) and other (peat moss) and other bryophytes in a bryophytes in a peatlandpeatland from western Canada from western Canada ((VittVitt and and

    BellandBelland 1995)1995)

    53443567

    41282858

    121679

    BogPoor fenModerately rich fen

    Extremely rich fen

    Total

    Other bryophytes

    Sphagnum

    Brown mosses

    Not a taxonomic classification Associated with rich minerotrophic fens Brown, reddish, yellow, and golden Indication of higher pH alkaline conditions Susceptible to N deposition Commonly associated with sedges

    Calliergonella cuspidata

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    Tomenthypnum nitens

    Feather mosses

    Coniferous forest Swamp forests Shaded areas Prefers hummocks and drained sites Hylocomium splendens

    Nitrogen FixationNitrogen Fixation

    http://wslar.epfl.ch/mitchell/edward/Sphagnum.htmCoiled chains of Nostoc are hidden

    in the leaf under light microscopy, but are readily observed as the red cells under ultraviolet-fluorescence microscopy

    Pleurozium schreberi

    Hyaline cells colonized by Cyanobacteria

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    Nitrogen fixation by P. 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

    .5 kg / ha yr previous estimated yearly fixation by all vegetation

    Pleurozium schreberi was recently found to contribute1.5 -2.0 Kg / ha yr fixation

    Implications for forestry management in boreal regions

    Other mosses

    Bryophytes that do not fall into the previous categories

    Polytrichum spp. swamps and fens

    Dicranum spp. hummocks

    LiverwortsLiverworts

    Leaves often lobed Dorsi-ventral organization Two body plans leafy or thallose

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    Liverworts vs. mossLiverworts vs. moss

    1. Lack a protonema2. Have single cell rhizoids3. Have two ranked leaf arrangement

    Mosses are usually 3 or 5 ranked

    Liverworts differ from mosses in three main ways

    Peatmoss a.k.a. Sphagnum

    SphagnumSphagnum factsfacts

    More living carbon in Sphagnum than in any other genus of plants worldwide

    Sphagnum in northern peatlands Covers 1.5 Mkm Stores 150 Gt of carbon

    Ecosystem engineer Resistant to decay Genus contains many species

    ~100 in the northern hemisphere ~ 19 in the Keweenaw peninsula

    2

    2

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    Global distribution of peatlandsGlobal distribution of peatlands

    Structure of Structure of SphagnumSphagnum

    The Head - Capitulum

    Pendent branchesDivergent branches

    Stem

    NO roots or rhizoids !

    (fascicles)

    Water adaptationsWater adaptations

    http://kentsimmons.uwinnipeg.ca/2152/sphagnum3.jpg

    SphagnumSphagnum cell structurecell structure

    25x Weight in water

    PoresHyline cells

    Photosynthetic cells

    Fibirals

    Sphagnum leafOne cell in twenty is alive

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    Chemical properties of Chemical properties of SphagnumSphagnumCoping with a harsh environmentCoping with a harsh environment

    Often grows in low nutrient areas Creates cation sites Contains secondary compounds

    Phenolics Can translocate metabolites

    COOH

    +

    COOH+

    COOH+COOH +CO

    OH

    +Ca++

    CationCation exchangeexchange

    H+

    H+

    pH

    Active substances are uronic acids-galacturonic acid and 5KMA

    The uronic acids sometimes referred to as sphagnanHigh pH - phenolics such as sphagnum acid are active

    Microtopography of SphagnumpH

    3.1

    3.2

    3.5

    4.0

    capillifolium

    magellanicum

    recurvum

    cuspidatum

    fuscum2.9

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    Differences in Sphagnum species in a hummock hollow complex in northern Michigan. (redrawn from Crum1992)

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    Desiccation

    Mosses are among the most desiccation-tolerant of all plants. They can be completely dried at the cellular level and recover completely. When re-wet after months or years of being dry, they begin photosynthesis within 5 minutes!

    http://ucjeps.herb.berkeley.edu/bryolab/pww/

    Known durations of desiccation survival in

    bryophyte plants.

    Sphagnum fuscum 2-4 dSphagnum papillosum 2-4 dSphagnum balticum 2-4 dSphagnum cuspidatum 2-4 dSphagnum magellanicum 2-4 dSphagnum magellanicum 14 dSphagnum fallax 14 dFontinalis flaccida 3 mosBarbula torquata 18 mosOxymitra 4 yrsRiccia canescens 7 yrsGrimmia laevigata 10 yrsSyntrichia ruralis 14 yrs 3Tortula muralis protonema 14 yrsRiccia macrocarpa 23 yrs

    Cold tolerance

    Increase sugar as natural antifreeze

    Species of Species of SphagnumSphagnum in the Keweenaw peninsulain the Keweenaw peninsula

    ~119 species of Sphagnum worldwide 19 species of Sphagnum in the

    Keweenaw Divided into sections based on:

    Morphology Growing conditions

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    http://plants.usda.gov/gallery/large/spsq70_004_lhp.jpg

    Sphagnum squarrosum

    Medium to robust Squarrose leaves

    widely spreading Large terminal

    capitulum Prefers depression Shade and moist

    areas Thuja swamps

    http://plants.usda.gov/gallery/large/spsq70_006_lhp.jpg

    Sphagnum squarrosum

    http://web.odu.edu/webroot/instr/sci/plant.nsf/files/10378.jpg/$FILE/10378.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. magellanicum Poor fens and acid lakes

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    Sphagnum papillosum

    sphagnum magellanicum

    http://plants.usda.gov/gallery/large/spma70_006_lhp.jpg

    Sphagnum magellanicum

    http://plants.usda.gov/gallery/large/spma70_004_lhp.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 Common and widespread

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    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

    http://www.bd.lst.se/publishedObjects/10001510/bollvitmossa.jpg

    Sphagnum Sphagnum wulfianumwulfianum

    Robust growthClover-like capitulumGreen to yellowBrittle stemCommonGrows almost exclusively in Thujaswamps

    Has a loose, shaggy, top-heavy appearance

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    Sphagnum girgensohnii

    http://plants.usda.gov/gallery/pubs/spgi70_001_php.jpg

    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://www.nps.gov/olym/crypto/M_SPGI.jpg

    Sphagnum girgensohnii

    Vulnerability of bryophytesVulnerability 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

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    Threats to BryophytesThreats to Bryophytes

    Deforestation Forest cultivation Urbanization Land reclamation Road construction Wetland drainage Invasive species

    Peatland lossPeatland loss

    Agriculture 250,000 km2

    Forestry 150,000 km2

    Peat harvesting 50,000 km2

    Does not include tropical peatlands

    65000Total125000Total121300Estonia131300Iceland391900Lithuania81905Norway852000Netherlands22000Finland

    121370Sarawak53000Sweden11410Canada505000Ukraine131930China707620Poland715760West Malaysia409631Belarus1617100USA8512000Germany

    1837200Indonesia1270400Russia

    % countrys peatland

    Agriculture% countrys peatland

    AgricultureEurope

    Peatland drained or altered for agriculturePeatland drained or altered for agriculture

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    Bryophyte harvesting not including Bryophyte harvesting not including peatlandspeatlands

    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?Sustainable?

    Can take several decades for regeneration Accumulation rate 22.4g/m /yr Commercial harvest rate of at least 21 years

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    Web sitesWeb sites

    Dr. Janice Glimes Online Bryology book http://www.bryoecol.mtu.edu/

    International Association of Bryologist http://www.bryology.org/index.html

    American Bryological and Lichenological Society (ABLS) http://www.unomaha.edu/~abls/

    Moss images http://plants.usda.gov/

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