Ecology of Red Maple Swamps in the Glaciated Northeast: A ...

Ecology of Red Maple Swamps in the Glaciated Northeast: A ... Ecology of Red Maple Swamps in the Glaciated Northeast: A ...

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Chapter 4. Abiotic I uences on the Plant Community The structure and floristic composition of red maple swamps are determined by the interplay of a wide variety of environmental factors, including climate and microclimate; abiotic factors such as water regime, soil and water chemistry, and the physical properties of soils; microrelief of the forest floor; biotic factors such as plant competition, disease, insect infestations, and the activities of beavers (Castor canadensis); anthropogenic influences such as logging, grazing, and water level manipulation; and natural catastrophes such as hurricanes and fire. A thorough examination of the role of each of these environmental factors in the ecology of red maple swamps is not possible, simply because most of these topics have not been investigated. Studies of vegetation and environment in northern swamps have identified two key gradients, one related to the position of the water table and the other related to the availability and means of supply of mineral nutrients (Paratley and Fahey 1986). Of the environmental factors that have been studied in red maple swamps, hydrology and nutrient status appear to be most directly responsible for variations in the structure and species composition of the plant community Ultimately, both of these factors are dictated by the wetland's hydrogeologic setting: the physical and chemical composition of the geologic substrate, the size and slope of the drainage basin, and the relative magnitude of the wetland's hydrologic inputs and outputs. Hydrology Research in forested wetlands throughout the United States has shown that hydrology is the primary force controlling the development of these wetlands and their stmctural d floristic attributes (Conner and Day 1976; Gosselink and Turner 1978; Brown et al. 1979; Carter et al. 1979; Harms et al. 1980; Dunn and Stearns 1987a). Hydrology also has been linked to the morphological and chemical properties of wetland soils (Heinselman 1970; Conner and Day 1976; Veneman et al. 1976; Pickering and Veneman 1984), and to the degree of development of surface microrelief (Satterlund 1960; Ehrenfeld and Gulick 1981; Lowry 1984). For these reasons, this chapter emphasizes the central. role of hydrology in shaping the structure and composition of red maple forested wetlands. The influence of water regime on tree growth is addressed in the following chapter. Influence on Community Structure The i~fiuence of hydrology on the structure of red maple swamps is poorly documented in the glaciated Northeast. In floodplain environments, the rate of flow of surface water through wetland forests may restrict woody plant establishment and hasten tree and shrub mortality simply through erosion of soils and mechanical damage to the vegetation itself (Brown et al. 1979; Harms et al. 1980; Huenneke 1982; Ehrenfeld 1986). Brown et al. (1979) found tree density in stillwater wetlands to be more than twice as high as in floodplain wetlands, and they concluded that water movement was a key factor expIaining wetland forest structure in general. Most red maple swamps in the glaciated Northeast are still-water wetlands. Where the swamps occur in streamside locations, either the streams are small and lack true floodplains, or the maple stands are located on the inner floodplain, st some distance from the channel. For these reasons, one might expect the effect of flowing water on eommunity structure to be minimal. Ehrenfeld (1986) found, however, that red maple floodplain forests in the New Jersey Pine Bamens had fewer woody species and lower tree and shrub density and biomass than nonfloodplain red maple swamps. Floodplain forests also had higher h e mortality and lower densities of tree seedlings and saplings. Like Brown et al. (1919), Ehrenfeld concluded that the physical disturbance emsed by flowing water and associated debris in floodplain forests

was the most likely reason for differences in cammunity stnxeture heLween floodplain and nonfloodplain sites. Wether stand structure in nonfloodplain red maple? swamps variee with water regime is unclear. %e density and basal area have been shown to h bt,h higher (Ehrenfeld and Gulick 1981; I ~wry 1984) and lower (Ehrenfeld 1986; Paratley and Mley 1986) on wetter sites. Cornpariaon of results of diffc?rent, studies is difficult because tho range of hydrologic conditions examined and tile meanings of "wetter" md "'drier" oftmn vary widely Further, tree density is influenced by tmth stand ago and stand origin (Rraiewa 1983). ?Ille ability of red maple to dominate sites that, mnge widely in wet,nc~s itself suggests that, carlee e~tablisff~d, the trees adapt well fa the prev~ilrtlg tiydrcrlogic regime and th:tt, unusually low dr~-lait,y or basrrl arcsiz can bc expect.ed to occur only where sitp wctxless ~ XCOO~W thi: spi*cies' t,oleranc lcvei. I-ts*lntIvc al_tt~nrlnltce ntld lion~;tu.q of shrubs hrtvt bsoxk iihuwrl 14 iricri?i~s~ wit,h wetlleas in rxonflc~)d~rlai~r red nri~ple swurnys (Ehrcrrfcld and Gtilick 1989; Lowry 1084; Swift et a1. 1984; Paratlemy arrd Fahcy 1986). In lthode Island, Lowry folitrld that both ~lcsxlsity and perce~;?.it,agc cover of strm-labs were greakst ~t ita as with the highest nlcexx water l~vels, but he noted that. these sites rtlsr, hixd tha lowent txcw canopy c30vr;r, the most i,mranr~r~rc~ci xllicrort*licf, nnd the highest ground- WH~&T p11. j\. sf.rong rcfnt,io~;?.i bctwccn wrttsr rcgirne and tbn ~Lructure of troth tliti woody ul~ci~rstory trierce 1981; Tiner 1985) or in oxbows or on floodaxtd the grt>tit~tX vqc~tation layer wm obscrvcd by fwati~y and fi'tklley (1986) in a New York mixed carlifer hwrtiwood swamp. In severely flooded and nmeslterratdy floacic;d areas of the swwnp, woody rzadcrstory cica~.rsitieas were 18,M6 and 10,881 sf~*rsts//kn, ms~rcct ivtaly, while values for seeps and moderatc3Iy dry areas were 7,429 and 8,9:5G stn*mJX~n. The perccntngc. cover of woody seedlings, prijtlirroicts, i\lld blyupl~ytes was found conducted in southern New England and New ta vary e3i&-;?.iifionrli,ly aiuorlg six gwoun(I v~~getatiorl York. asaociatiorxs 13s well. Sedges and rnosscs wcrc mosL abundant in those red maple cornnxunities Hydrologic Variation Among Swamp wikh the highest ~a-rean water levels daring the Cammunitios growing season. 12c*rcentnge cover of woody seed- Damn~an and Kershner (1977) identified soil lings was in a msderabfy wet red maple comnunity th~t received large inflow of nutrientrich surface Water fmm a newby creek during the spring. %search in red maple swmnps in southern New Jersey (Ehrenfeld and Gulick 1981; Ehren- feid 1986) reaffirms the conelusiom drawn in glaciated areas of the Northeast. In two separate studies, shrub density and biomass were much higher in wet hardwood swamps than in dry hardwood swamps. While the biomass of herbs was small to negligible at these sites, its relative eontribution to total biomass was much greater at the wetter sites; herb biomass totaled 195 kg/ha in the wet swamps, but only 53 kfia in the dry swamps. If the influence of hydrology on vegetation structure is to be further elucidated, however, detailed measurements of standard hydrologic parameters over several years will be required. Influence on Floristic Composition The influence of water regime or soil moisture on species composition and distribution in wetland forests has been most clearly demonstrated in floodplain communities. In the bottomland hmdwood forests of the southern United States, which often include a red maple component, plant community composition has been shown to be a function of the timing, frequency, and duration of flooding or of anaerobic soil conditions ('Monk 1%; Brown et al. 1979; Huffman and Forsythe 1981; Conner and Day 1982; Parsons and Ware 1982). A strong relation between species distribution and hydrologic regime has been shown on northeastern floodplains as well. In this region, red maple generally occurs in alluvial basins on the inncr floodplain (Buell and Wiskndahl 1955; plain terraces (Pierce 1981; Holland and Burk 1984; Metzler and Damman 1985) where the forest is less frequently flooded by river waters and the soil is less well drained after floods subside than on the outer floodplain. Information on relationships between water regime and the floristics of nonfloodplain red maple swamps in the glaciated Northeast comes primarily from research n~oisLure regime as a key determinant of floristic variation in western Connecticut forests located over till and gneissic bedrock. They described three red maple swamp communities in that region and suggested that the floristic differences anxong those communities were caused by differ-

was <strong>the</strong> most likely reason for differences <strong>in</strong> cammunity<br />

stnxeture heLween floodpla<strong>in</strong> and nonfloodpla<strong>in</strong><br />

sites.<br />

We<strong>the</strong>r stand structure <strong>in</strong> nonfloodpla<strong>in</strong> red<br />

maple? swamps variee with water regime is unclear.<br />

%e density and basal area have been<br />

shown to h bt,h higher (Ehrenfeld and Gulick<br />

1981; I ~wry 1984) and lower (Ehrenfeld 1986;<br />

Paratley and Mley 1986) on wetter sites. Cornpariaon<br />

<strong>of</strong> results <strong>of</strong> diffc?rent, studies is difficult<br />

because tho range <strong>of</strong> hydrologic conditions exam<strong>in</strong>ed<br />

and tile mean<strong>in</strong>gs <strong>of</strong> "wetter" md "'drier"<br />

<strong>of</strong>tmn vary widely Fur<strong>the</strong>r, tree density is <strong>in</strong>fluenced<br />

by tmth stand ago and stand orig<strong>in</strong> (Rraiewa<br />

1983). ?Ille ability <strong>of</strong> red maple to dom<strong>in</strong>ate sites<br />

that, mnge widely <strong>in</strong> wet,nc~s itself suggests that,<br />

carlee e~tablisff~d, <strong>the</strong> trees adapt well fa <strong>the</strong> prev~ilrtlg<br />

tiydrcrlogic regime and th:tt, unusually low<br />

dr~-lait,y or basrrl arcsiz can bc expect.ed to occur only<br />

where sitp wctxless ~ XCOO~W thi: spi*cies' t,oleranc<br />

lcvei.<br />

I-ts*lntIvc al_tt~nrlnltce ntld lion~;tu.q <strong>of</strong> shrubs<br />

hrtvt bsoxk iihuwrl 14 iricri?i~s~ wit,h wetlleas <strong>in</strong><br />

rxonflc~)d~rlai~r red nri~ple swurnys (Ehrcrrfcld and<br />

Gtilick 1989; Lowry 1084; Swift et a1. 1984; Paratlemy<br />

arrd Fahcy 1986). In lthode Island, Lowry<br />

folitrld that both ~lcsxlsity and perce~;?.it,agc cover <strong>of</strong><br />

strm-labs were greakst ~t ita as with <strong>the</strong> highest<br />

nlcexx water l~vels, but he noted that. <strong>the</strong>se sites<br />

rtlsr, hixd tha lowent txcw canopy c30vr;r, <strong>the</strong> most<br />

i,mranr~r~rc~ci xllicrort*licf, nnd <strong>the</strong> highest ground-<br />

WH~&T p11. j\. sf.rong rcfnt,io~;?.i bctwccn wrttsr rcgirne<br />

and tbn ~Lructure <strong>of</strong> troth tliti woody ul~ci~rstory trierce 1981; T<strong>in</strong>er 1985) or <strong>in</strong> oxbows or on floodaxtd<br />

<strong>the</strong> grt>tit~tX vqc~tation layer wm obscrvcd by<br />

fwati~y and fi'tklley (1986) <strong>in</strong> a New York mixed<br />

carlifer hwrtiwood swamp. In severely flooded and<br />

nmeslterratdy floacic;d areas <strong>of</strong> <strong>the</strong> swwnp, woody<br />

rzadcrstory cica~.rsitieas were 18,M6 and 10,881<br />

sf~*rsts//kn, ms~rcct ivtaly, while values for seeps and<br />

moderatc3Iy dry areas were 7,429 and<br />

8,9:5G stn*mJX~n. The perccntngc. cover <strong>of</strong> woody<br />

seedl<strong>in</strong>gs, prijtlirroicts, i\lld blyupl~ytes was found<br />

conducted <strong>in</strong> sou<strong>the</strong>rn New England and New<br />

ta vary e3i&-;?.iifionrli,ly aiuorlg six gwoun(I v~~getatiorl York.<br />

asaociatiorxs 13s well. Sedges and rnosscs wcrc<br />

mosL abundant <strong>in</strong> those red maple cornnxunities Hydrologic Variation Among Swamp<br />

wikh <strong>the</strong> highest ~a-rean water levels dar<strong>in</strong>g <strong>the</strong> Cammunitios<br />

grow<strong>in</strong>g season. 12c*rcentnge cover <strong>of</strong> woody seed- Damn~an and Kershner (1977) identified soil<br />

l<strong>in</strong>gs was <strong>in</strong> a msderabfy wet red maple<br />

comnunity th~t received large <strong>in</strong>flow <strong>of</strong> nutrientrich<br />

surface Water fmm a newby creek dur<strong>in</strong>g <strong>the</strong><br />

spr<strong>in</strong>g.<br />

%search <strong>in</strong> red maple swmnps <strong>in</strong> sou<strong>the</strong>rn<br />

New Jersey (Ehrenfeld and Gulick 1981; Ehren-<br />

feid 1986) reaffirms <strong>the</strong> conelusiom drawn <strong>in</strong><br />

glaciated areas <strong>of</strong> <strong>the</strong> Nor<strong>the</strong>ast. In two separate<br />

studies, shrub density and biomass were much<br />

higher <strong>in</strong> wet hardwood swamps than <strong>in</strong> dry hardwood<br />

swamps. While <strong>the</strong> biomass <strong>of</strong> herbs was<br />

small to negligible at <strong>the</strong>se sites, its relative eontribution<br />

to total biomass was much greater at <strong>the</strong><br />

wetter sites; herb biomass totaled 195 kg/ha <strong>in</strong> <strong>the</strong><br />

wet swamps, but only 53 kfia <strong>in</strong> <strong>the</strong> dry swamps.<br />

If <strong>the</strong> <strong>in</strong>fluence <strong>of</strong> hydrology on vegetation structure<br />

is to be fur<strong>the</strong>r elucidated, however, detailed<br />

measurements <strong>of</strong> standard hydrologic parameters<br />

over several years will be required.<br />

Influence on Floristic Composition<br />

The <strong>in</strong>fluence <strong>of</strong> water regime or soil moisture<br />

on species composition and distribution <strong>in</strong> wetland<br />

forests has been most clearly demonstrated<br />

<strong>in</strong> floodpla<strong>in</strong> communities. In <strong>the</strong> bottomland<br />

hmdwood forests <strong>of</strong> <strong>the</strong> sou<strong>the</strong>rn United States,<br />

which <strong>of</strong>ten <strong>in</strong>clude a red maple component, plant<br />

community composition has been shown to be a<br />

function <strong>of</strong> <strong>the</strong> tim<strong>in</strong>g, frequency, and duration <strong>of</strong><br />

flood<strong>in</strong>g or <strong>of</strong> anaerobic soil conditions ('Monk<br />

1%; Brown et al. 1979; Huffman and Forsy<strong>the</strong><br />

1981; Conner and Day 1982; Parsons and Ware<br />

1982). A strong relation between species distribution<br />

and hydrologic regime has been shown on<br />

nor<strong>the</strong>astern floodpla<strong>in</strong>s as well. In this region,<br />

red maple generally occurs <strong>in</strong> alluvial bas<strong>in</strong>s on<br />

<strong>the</strong> <strong>in</strong>ncr floodpla<strong>in</strong> (Buell and Wiskndahl 1955;<br />

pla<strong>in</strong> terraces (Pierce 1981; Holland and Burk<br />

1984; Metzler and Damman 1985) where <strong>the</strong> forest<br />

is less frequently flooded by river waters and<br />

<strong>the</strong> soil is less well dra<strong>in</strong>ed after floods subside<br />

than on <strong>the</strong> outer floodpla<strong>in</strong>. Information on relationships<br />

between water regime and <strong>the</strong> floristics<br />

<strong>of</strong> nonfloodpla<strong>in</strong> red maple swamps <strong>in</strong> <strong>the</strong> glaciated<br />

Nor<strong>the</strong>ast comes primarily from research<br />

n~oisLure regime as a key determ<strong>in</strong>ant <strong>of</strong> floristic<br />

variation <strong>in</strong> western Connecticut forests located<br />

over till and gneissic bedrock. They described<br />

three red maple swamp communities <strong>in</strong> that region<br />

and suggested that <strong>the</strong> floristic differences<br />

anxong those communities were caused by differ-

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