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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of the year, the forest floor commonly consists of deep hollows and convex mounds; in swamps that lack surface water entirely or that are flooded only temporarily, nnicrorelief is not as well developed (Ehrenfeld and GuXick 1981). Lowry (1984) took spot elevations at over 700 points in each of six red maple swamps and six Atlantic white cedar swamps in southern %ode Island and determined that microrelief was more highly developed in the cedar swamps, which had significantly higher mean water levels as well. He dso confumed that the extent of microrelief in the red maple swamps was related to water level. Considering all points more than 20 cm above the average level of the depressions to be mounded, he calculated that nearly 75% of the variation in the amount of mounded ground among the six swamps could be explained by differences in the 7-year mean water levels among the sites. Figure 4.5 illustrates pronounced microrelief in a seasonally flooded red maple swamp. How active a role vegetation plays in the development of microrelief is unclear. Initially, the dis- tribution of trees and shrubs in a swamp is determined by the relative wetness of various possible germination sites on the forest floor. Once they are established, those trees that have the ability to develop a compact, elevated root system clearly stand a greater chance of surviving the effects of prolonged high water levels. Root system development thus may increase mound size. Significantly, radial growth of red mqle trees in any given year appears to be directly related to the deviation of that year's average water level from the long-term average. Lowry (1984) demonstrated that, in Rhode Island swamps, growth was greatest in years when water levels were closest to the 7-year mean. This finding suggests that in each swamp there may be an optimal distance, dependingupon water regime and soil characteristics, between the elevation of the average water level and the depth of tree roots. Whether the role of vegetation in microrelief development is active or passive, variation in surface elevation within a swamp maximizes the opportunity for any tree to achieve that optimum position and to maximize its growth. Fig. 4.5. Mound-and-pool microrelief in a seasonally flooded red maple swamp. Swamps with particularly high water levels, such as this one, generally have high mounds and little vegetation growing in the pools. The measuring stick is graduated in ZQ-cm incremenk; the water averages 15-25 cm in depth. The photograph was taken in mid-April.

Influence on Swamp Vegetation Floristic Composition Through its influence on soil aeration (Huenneke 1982; Paratley and Fahey 1986), nutrient availability (Ehrenfeld and Gulick 1981; Paratley and Fahey 1986), and relative litter accumulation (Little 1950; Malecki et al. 1983; Paratley and Fahey 1986), microrelief creates a variety of microhabitats and thus has a major effect on species composition and distribution of swamp flora. Beatty's (1984) research in a sugar maple-Arnerican beech upland forest in eastern New York showed that microrelief may cause local variations in soil acidity and soil temperature as well. Pronounced microrelief allows species with widely differing soil moisture requirements or tolerances to coexist in a limited area in red maple swamps (Bergman 1920; Sampson 1930; Thompson et al. 1968; Huenneke 1982; Paratley and Fahey 1986). Wile mosses, liverworts, and hydrophilic herbs thrive in seasonally flooded or saturated depressions and at the bases of mounds, species unable to tolerate prolonged saturation grsw higher up on the mounds (Niering 1953; Thompson et al. 1968; Paratley and Fahey 1986). Figure 4.6 shows the influence of microrelief on plant distribution in a Rhode Island swamp. Faratley and Fahey (1986) found plant species richness to be positively correlated with microrelief; in fact, they cited high microsite heterogeneity as one of the factors most responsible for the unusually high species richness observed in their central New York study area. Under a given water regime, certain species of plants tend to occur either primarily on mounds or primarily in depressions. However, the microsite preferences of some species may change depending on mound height or on the relative wetness of the depressions. In a detailed analysis of the relation between species distribution and microrelief in a New York swamp with organic soils, Paratley and Fahey (1986) found that five ground-layer plants-including spotted touchme-not, marsh marigold, mosses of the genus Mnium, sensitive fern, and northern bugleweed-

<strong>of</strong> <strong>the</strong> year, <strong>the</strong> forest floor commonly consists <strong>of</strong><br />

deep hollows and convex mounds; <strong>in</strong> swamps that<br />

lack surface water entirely or that are flooded only<br />

temporarily, nnicrorelief is not as well developed<br />

(Ehrenfeld and GuXick 1981). Lowry (1984) took<br />

spot elevations at over 700 po<strong>in</strong>ts <strong>in</strong> each <strong>of</strong> six red<br />

maple swamps and six Atlantic white cedar<br />

swamps <strong>in</strong> sou<strong>the</strong>rn %ode Island and determ<strong>in</strong>ed<br />

that microrelief was more highly developed <strong>in</strong> <strong>the</strong><br />

cedar swamps, which had significantly higher<br />

mean water levels as well. He dso confumed that<br />

<strong>the</strong> extent <strong>of</strong> microrelief <strong>in</strong> <strong>the</strong> red maple swamps<br />

was related to water level. Consider<strong>in</strong>g all po<strong>in</strong>ts<br />

more than 20 cm above <strong>the</strong> average level <strong>of</strong> <strong>the</strong><br />

depressions to be mounded, he calculated that<br />

nearly 75% <strong>of</strong> <strong>the</strong> variation <strong>in</strong> <strong>the</strong> amount <strong>of</strong><br />

mounded ground among <strong>the</strong> six swamps could be<br />

expla<strong>in</strong>ed by differences <strong>in</strong> <strong>the</strong> 7-year mean water<br />

levels among <strong>the</strong> sites. Figure 4.5 illustrates pronounced<br />

microrelief <strong>in</strong> a seasonally flooded red<br />

maple swamp.<br />

How active a role vegetation plays <strong>in</strong> <strong>the</strong> development<br />

<strong>of</strong> microrelief is unclear. Initially, <strong>the</strong> dis-<br />

tribution <strong>of</strong> trees and shrubs <strong>in</strong> a swamp is determ<strong>in</strong>ed<br />

by <strong>the</strong> relative wetness <strong>of</strong> various possible<br />

germ<strong>in</strong>ation sites on <strong>the</strong> forest floor. Once <strong>the</strong>y are<br />

established, those trees that have <strong>the</strong> ability to<br />

develop a compact, elevated root system clearly<br />

stand a greater chance <strong>of</strong> surviv<strong>in</strong>g <strong>the</strong> effects <strong>of</strong><br />

prolonged high water levels. Root system development<br />

thus may <strong>in</strong>crease mound size. Significantly,<br />

radial growth <strong>of</strong> red mqle trees <strong>in</strong> any given year<br />

appears to be directly related to <strong>the</strong> deviation <strong>of</strong><br />

that year's average water level from <strong>the</strong> long-term<br />

average. Lowry (1984) demonstrated that, <strong>in</strong><br />

Rhode Island swamps, growth was greatest <strong>in</strong><br />

years when water levels were closest to <strong>the</strong> 7-year<br />

mean. This f<strong>in</strong>d<strong>in</strong>g suggests that <strong>in</strong> each swamp<br />

<strong>the</strong>re may be an optimal distance, depend<strong>in</strong>gupon<br />

water regime and soil characteristics, between <strong>the</strong><br />

elevation <strong>of</strong> <strong>the</strong> average water level and <strong>the</strong> depth<br />

<strong>of</strong> tree roots. Whe<strong>the</strong>r <strong>the</strong> role <strong>of</strong> vegetation <strong>in</strong><br />

microrelief development is active or passive, variation<br />

<strong>in</strong> surface elevation with<strong>in</strong> a swamp maximizes<br />

<strong>the</strong> opportunity for any tree to achieve that<br />

optimum position and to maximize its growth.<br />

Fig. 4.5. Mound-and-pool microrelief <strong>in</strong> a seasonally flooded red maple swamp. <strong>Swamps</strong> with<br />

particularly high water levels, such as this one, generally have high mounds and little vegetation<br />

grow<strong>in</strong>g <strong>in</strong> <strong>the</strong> pools. The measur<strong>in</strong>g stick is graduated <strong>in</strong> ZQ-cm <strong>in</strong>cremenk; <strong>the</strong> water averages<br />

15-25 cm <strong>in</strong> depth. The photograph was taken <strong>in</strong> mid-April.

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