2.4 FACTORS CONT3OLLIVG PLANT DEMOGRAPHY<strong>The</strong> distribution <strong>of</strong> plant speciespopulations in any natural situation reflects<strong>the</strong> response <strong>of</strong> individual speciesto soeci f ic environmental parameters. Inwetl and habitats <strong>the</strong>se parameters areespecially varied, primarily due to <strong>the</strong>i nf 1 uence <strong>of</strong> water on habi tat gradients.Siologically-mediated interactions betweenplant species fur<strong>the</strong>r coqplicate <strong>the</strong> perception<strong>of</strong> physical gradients. Much <strong>of</strong><strong>the</strong> information concerning <strong>the</strong> causes forobserved spatial distributions <strong>of</strong> vascularflora in <strong>the</strong>se marshes is anecdotal, althoughenough exists to warrant a generaldiscussion.Inundation<strong>The</strong>re seems to be a general consensusamong researchers investigating plant demographyin <strong>the</strong> tidal freshwater habitatthat <strong>the</strong> frequency and duration <strong>of</strong> floodingis <strong>the</strong> primary factor governing speciesdistributions (Kiviat 1978a; Ooumleleand Silberhorn 1978; Ferren et al. 1981;McCormick and Somes 1932). Despite <strong>the</strong>fact that <strong>the</strong> vast majority <strong>of</strong> plantsoccurring in <strong>the</strong>se marshes must experienceflooding on a daily basis, speciesvary greatly in <strong>the</strong>ir ability to withstandinundation. For some species, extensiveflooding seems to be a physiological requirement for subsistence, whereas foro<strong>the</strong>rs, it can be a detriment to normalgrowth and development. Scul thorpe (1967)notes that nicmerous terres tri a1 pl ants areable to survive long periods ei<strong>the</strong>rcompletelyor partially submerged. It isconceivable that facultative hydrophyteshave evolved in order to avoid competitionor to exploit open niches in habitats suchas <strong>the</strong>se.stunted progeny in deep-wa ter experimentalplots (Yamisaki and Tange 1981). Manyo<strong>the</strong>r researchers working in sal t marsh,mangrove swamp, and freshwater lake environmentshave concl uded that inundationeffectively contributes to segregation <strong>of</strong>pl ant species ~opul a ti ons a1 ong an el evationalgradient (Filandossian and Mc Intosch1950; Aifains 1963; Scul thorpe 1957; Kerwinand Pedigo 1971; Odum 1971). As yet, thisphenomenon has not been quantitativelydetermined for tidal freshwater marshes.However, evidence from o<strong>the</strong>r wetl and si tuationssuggests that tidal freshwaterplant communi ties segregate a1 ong inundationgradients as well.Substrate<strong>The</strong> soil in tidal freshwater marshescan be described as a waterlogged organicmuck with varying amounts <strong>of</strong> sand, silt,and clay (see Section 1.7). Differencesin soil stability, soil moisture retention,and soil nutrient availabil i t,y areall related to <strong>the</strong> physical characteristics<strong>of</strong> a given substrate and may influencespecies distributions directly. <strong>The</strong>spatial heterogeneity <strong>of</strong> substrate characteristicsis not sufficient to explaindistributions or species performance(1Jhigharn and Simpson 1975; Wetzel andPowers 1978). Wetzel and Powers (1978)concluded that substrate characteristicsaffect plant demography only in local izedzones within <strong>the</strong> marsh, and <strong>the</strong>n, <strong>the</strong>sesubtl e differences are largely obscured bymajor environmental gradients acting toproduce species distributions (e.g., elevationand tidal inundation). In ouropinion, this is an area which needsconsiderably more research.Current FlowIn <strong>the</strong> progression from open water Low-gradi ent river courses <strong>of</strong> <strong>the</strong>channels to <strong>the</strong> tnarsh-upland boundary, <strong>the</strong> Atlantic Coastal Plain tend to flow ra<strong>the</strong>rspecies composition <strong>of</strong> vascular flora sluggishly except during extreme storm orchanges noticedbly, even over almost im- flood events. Aside from channel disperceptablevariation in marsh surface charge, however, <strong>the</strong> daily ebb and floodelevations (Hoover 1983). It is known <strong>of</strong> tidal water onto and <strong>of</strong>f <strong>the</strong> marsh surthatcommon and narrow-leaved cattails face can produce significant currentwill segregate along a gradient <strong>of</strong> water velocities. Much <strong>of</strong> this water becomesdepth in nontidal habitats, <strong>the</strong> latter channel ized into dendri tical ly-shapedspecies found in deeper water (Grace and creeks within <strong>the</strong> marsh. <strong>The</strong>se creeksGetzel 1951). Common reed and wild rice deliver ground water from high marsh toa1 SO respond to varying inundation, each low marsh to channel long after <strong>the</strong> tidespecies producing fewer and somewhat has ebbed (Hoover 1983). Concentrated27
water movement may (1) impair <strong>the</strong> abil i t y<strong>of</strong> seeds, seedlings, or adult plants togrow and develop and (2) may confine <strong>the</strong>dispersion <strong>of</strong> particular seeds to portions<strong>of</strong> <strong>the</strong> marsh wqth little or no water flow.Several studies provide evidence tosupport <strong>the</strong>se ideas. Whigham et al.(1979) noted that arrow-arum seed1 ings,which develop uni formly throughout mostsections <strong>of</strong> <strong>the</strong> marsh, were absent fromstreambafik areas. Higher rates <strong>of</strong> watermovetilent a1 ong <strong>the</strong> s treambank apparentlyprevented seeds from establ ishing <strong>the</strong>mselves,since seeds collected from <strong>the</strong>sesariie areas were found to be physiologicallycapable <strong>of</strong> germination. In contrast,pickerelweed is known to prefer streambank1 oca tions, taklng advantage <strong>of</strong> greatersoil surface temperatures on <strong>the</strong> exposedmud as we1 1 as reduced competitive pressuresin this area (Garbisch and Coleman1978).Sal i nitv<strong>The</strong> variahil ity and complexity <strong>of</strong>wetland plant communities increases withdecreasing salinity. Anderson et a1 .(1968), studying a 25-mJle stretch <strong>of</strong> <strong>the</strong>Patuxent River estuary in Maryland, il lustratedthis fact by quantifying plant species'diversity at sltes with differentsalinfty regimes (Table 5). Ry definition,<strong>the</strong> tidal freshwater habitat shouldnot encounter average water sal ini tiesgreater than 0.5 opt. However, thisboundary between tidal fresh and 01 i goha-1 ine waters has been seen to mi grate considerabledistances over <strong>the</strong> course <strong>of</strong> ayear in responseperiods. <strong>Marshes</strong>to drought and floodwhich intermittentlycome into contact with elevated watersalinities may harbor slightly lessdiverse plant co~notun.! ties dominated byfacultative halophytes (see Section 2.2).<strong>Freshwater</strong> species which appear to dropout <strong>of</strong> <strong>the</strong> plant communities in <strong>the</strong>seareas include spatterdock, sweetfl ap,blueflag, various sedges, and giant cutgrass,Physiological Capabil i tylAnaerobic ToxinsPreliminary evidence suggests thattidal freshwater marsh soils are not asreduced as some salt marsh substrates, atleast in <strong>the</strong> surface horizon (see Section1.7). Presumably, <strong>the</strong> extent <strong>of</strong> oxygendeficiency is not homogeneous over <strong>the</strong>entire marsh pr<strong>of</strong>ile, being 1 ess intensein those areas which drain regularly wi<strong>the</strong>ach tidal cycle. Never<strong>the</strong>less, soi 1s andassociated microbial populations shift<strong>the</strong>ir predominant metabol ic pathways underanoxic conditions, affecting both inorganicand organic soil constituents --- thiscan have important consequences for wetlandplant life.<strong>The</strong> bioavailabil i ty <strong>of</strong> most nutrientsand toxins responds to <strong>the</strong> oxidationreductionconditions <strong>of</strong> wetland soils(Gambrel 1 and Patrick 1978). Increasedlevels <strong>of</strong> soluble iron and managanese insome reduced soils are reported to be toxicto plants (Armstrong 1975), and fur<strong>the</strong>rmore,may facil i tate <strong>the</strong> formation <strong>of</strong>inorganic-oxide 1 ayers around roots whichpotentially impedes <strong>the</strong> transport <strong>of</strong> nutrientsfrorn soil to plant (Armstrong andBoatman 1967; Howel er 1973). Extremelyreduced soil s with appreciable organiccarbon mity develop toxic sulfide compounds.Many wetland plants have adapted to<strong>the</strong>se extreme conditions , devel oping means<strong>of</strong> metabol izing anaerobical ly and excludingtoxins froin roots. <strong>The</strong> provision <strong>of</strong>air-space or aerenchy~iiatous tissue is onemechanism enabling plants to transportat~nospheric gas to anoxic rhizospheres.<strong>The</strong> functioning <strong>of</strong> this pressurized, flowthroughsystem has been documented indetail for spatterdock (Dacey 1980), aspecies which typically thirves in <strong>the</strong>most waterlogged portions <strong>of</strong> <strong>the</strong> marsh.O<strong>the</strong>r emergent macrophytes which possessaerenchynatous tissue include arrow-arum,pickerelweed, and even certain grasses andsedges. Most <strong>of</strong> <strong>the</strong>se species will befound in tile lower intertidal zones intidal freshwater marshes.CompetitionFrom an ecological point <strong>of</strong> view, <strong>the</strong>diverse flora indigenous to tidal freshwaterwetlands would seem to have a highpotential for speci es-speci es i nteractions.A conspicuous feature <strong>of</strong> manyplant communities that is <strong>of</strong>ten consideredevidence <strong>of</strong> competitive displacement is<strong>the</strong> segregation <strong>of</strong> species along a habitatgradient. A1 though species segregations
- Page 2 and 3: LibraryNational Wetlands Restarch C
- Page 4 and 5: PREFACEThis report is part of a ser
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- Page 36 and 37: able physiognonies in this species
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other rails) gather to feed on the
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the effects of seabirds on nutrient
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the various soecies of mammals in t
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h?ve not been directly studied. Stu
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CHAPTER 9- VALUES, ALTERATIONS, AND
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estuary to the rlext dependinq upon
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Section 3.3). Simpson et al. (1981)
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Tab1 e 24. Hypothetical comparisons
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Table 24.Concluded.Characteristics
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REFERENCESAdams, D.A. 1963. Factors
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J.E. Clark eds. Wetland functions a
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Sci. Rep. 40.4. Nat. Resour. Inst.U
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University Press, Ithaca, N.Y.345 p
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fish fauna of Tivoli Bays. Bard Col
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Patrick, eds. Two studies of Tinicu
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Sci . 16:77-78.Penney, J.T. 1950, D
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Smith, B.A. 1971. The fishes of fou
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R.L. Simpson, eds. Freshwater wet1
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APPENDIX APlants of the Tidal Fresh
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PontederiaceaePontederia cordataZos
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Ona graceaeJossiaeare~en~LudwuriaDa
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APPENDIX BFISH OF TIDAL FRESHWATERS
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Amiidae -bowfins&Labrvf inElopidae
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Geographic Salinity Relativerange r
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NameCatostomidae -suckersGeographic
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Name GeographicrangeSalinityrangeRe
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NameGeographicrangeSalinity Relativ
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Centrarchidae -sunfishesName Geogra
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Percidae -perchesEtheostomafusiform
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Name Geographic Salinity Relativera
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Reference Numbers Key1. Adams 19702
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l m L o .Arl- >4J 3o m uc, mEU mmm
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Family / SpeciesRegionStatusFood ha
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- fIn-f W--7.40 -> 0171- Lw m mUtu
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APPENDIX D:Avifauna of tidal freshw
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Hooded merganser(LoDbodvtes-)Common
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FamilySpeciThreskiornithidae - ibis
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Family / SpeciesRegionSeasonStatusH
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Cathartidae - vulturesRegionDIURNAL
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CULLS, TERNS, KINGFISHERS, AND CROW
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ARBOREAL BIRDSFamily / Species Regi
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Family / Species Region Season Stat
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Family / Species Region Season Stat
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Family / Species Region Seasonnorth
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Family / SpeciesMarsh wren(Long-bil
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drQFamily / SpeciesFringilidae - fi
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APPENDIX E:Mammals of tidal freshwa
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Family / SpeciesRegionStatusFood ha
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