[From the U.S. Government Printing Office, www.gpo.gov]
... ......... . .............. ...... .......... .......... .. ... .. .... . .....- ..... ...... .......... . . ............. .................. .. ....... .... ... .......... .. . ........ EL _eArl ... ..... ...... . . . ............... . .. .................. . . . ........ ......... . ........... .. ........... ............ ...................... ............ ................. .... ......- .......... .... ...... @Wetlands Ecology By Pam Mason OBJECTIVES The purpose of this unit is to familiarize yourself Wetlands can be vegetated or nonvegetated. The with the fundamental aspects of wetlands ecology. This plants that vegetate wetlands are called hydrophytes review will touch on the role of wetlands in the ecosys- meaning water loving. Hydrophytes are adapted to life in tem, what makes a wetland, different wetland types and the water. Adaptations can be morphological, reproduc- the plants and animals of the wetland. tive or physiological. Plants that have adapted to grow in Following completion of the module, you will: saline waters are called halophytes meaning salt loving. 1) have a basic understanding of the role of wetlands in Some common terms can be used to describe tidal the ecosystem, 2) be able to recognize wetlands, and 3) wetlands by their relative position in the landscape. The define/describe hydrophyte, primary production, detritus, low marsh is the area of the lowest elevation, often vege- secondary production, food web, zonation. tated by saltmarsh cordgrass or pickerelweed. The high LECTURE NOTES marsh is generally above mean high water and is vege- tated with saltmeadow hay, saltgrass, big cordgrass. The Introduction marsh-upland ecotone, where the wetlands grade into the uplands, is often indicated by the growth of shrubs What is a wetland? A wetland is composed of three like the marsh elder, highwater bush and wax myrtle. fundamental components; Water, plants and soil. The water that makes wetlands wet is referred to as Ecosystem Functions wetland hydrology. Wetland hydrology can come from Primaryproduction and the food web many sources. The water may come from tides or nonti- Green plants use the sun's energy to convert inor- dal sources such as rain, streams or groundwater. Tidal ganic (nonliving) minerals to organic (living) plant tissue. waters can be salt or fresh. In most tidal systems there is This process is know as photosynthesis. As the first- a gradient from saltwater to freshwater progressing up- level of production of organic material the process is stream. In general nontidal wetlands are freshwater, al though some are saltwater. This can occur where there is called primary production. a high amount of dissolved minerals in the water and high Plant species common to wetlands have high levels of primary production. Estimates of the primary produc- evaporation rates increase the concentrations (Chapman, 1960). tion of tidal wetlands are as high as 4-6 tons per acre per Wetland soils are hydric soils. When soils are inun- year. Wetlands productivity rivals or surpasses the most dated with water, anaerobic (without air) conditions usu- -productive farmlands (Tiner, 1984). ally result. Anaerobic conditions lead to chemical Animals that feed directly on plant material are changes in the soils. Hydric soils are generally dark in called primary consumers. However, few animals eat color. the vegetation and most of the plant material becomes de- 2 tritus'. Detritus is partially decomposed plant material. 2. High rates of primary productivity may lead to Depending on the wetland system, some or most, of the high rates of nutrient uptake and subsequent burial material is exported to the estuary. when the plants die. Many types of microorganisms grow on the detri- 3. The nutrients used by the growing wetlands tus. The tiny plants and animals which populate the de- plants are not available to support the development of tritus increase the value of the detritus as food for algal blooms (Mitsch & Gosselink, 1986).. estuarine organisms. The d .etritus is consumed by Fish and Wildlife Habitat many animals, including crabs, fish and shellfish. The A great variety of birds, fish, mammals and inver- consumers digest the microorganisms growing on the tebrates use .wetlands for foraging, shelter, nesting, detritus. However, the detritus is only partially broken- down and passes through mostly undigested. The.detri' spawning and nursery areas. Some species of plants and animals can survive only in wetlands such as salt- tus is repopulated with microorganisms and the process marsh cordgrass a .nd muskrats. However, many other is repeated. species use wetlands for food or refuge but not as a pri- The higher level consumers in the process are mary residence. those that feed on the detrital consumers. Examples of Many species of finfish, including most commer- higher level consumers are shorebirds, finfish and mam- cial and game fishes use wetlands as habitat. Striped mals including humans. bass, white perch and herrings ai@e just some of the fa- Water Quality miliar fishes which-use wetlands. Crustaceans, like the Wetlands occupy a strategic position in the land- blue crab and. shrimp also use wetlands. Many species scape between uplands and the aquatic environment. of shellfish are found in wetlands or the adjacent shal- Wetlands can intercept upland runoff and filter and trap low waters. Oysters, clams and mussels use wetlands pollutants and sediments before they reach the water- as habitat. way. Several wetlands attributes have the potential to Shoreline Erosion Control affect the flow of chemicals and sediments. Wetlands deter shoreline erosion by reducing wave 1. Wetlands vegetation slows water velocity, caus- ing sediments and sorted chemicals to drop out of the energy and current velocity. Roots and,rhizomes of water column on to the wetland. wetland plants stabilize the s .ubstrate, and the stems and leaves slow the flow of waves and tidal currents. Al- though lacking vegetation, the natural slope of nonvege- Wetlands Program December 1993 tated wetlands provide a buffer to shoreline erosion. College of William and Mary School of Marine Science Wave and tidal curr.ent energy is dissipated as the water Virginia Institute of Marine Science spreads out over the flats. Gloucester Point, Virginia 23062 Flood Protection Dr. Carl Hershner, Program Director Wetlands may slow and temporarily store flood wa- Published by: VIMS Publication Center ters. The capacity for flood water storage is influenced "A publication of the Virginia Department of Environmental by the tidal stage. Portions. of the wetlands already Quality's Coastal Resources Management Program pursuant to flooded by tidal waters may be unable to assimilate ad- National Oceanic and Atmospheric Administration Award ditional flood waters. This is particularly true of low No. NA270ZO312-01. " marsh areas with daily tidal inundation. The absorption "This paper isfunded in part by a grant1cooperative agreementfrom the National Oceanic and of flood water by wetlands and the ability of vegetation S Atmospheric Administration. The views expressed to slow flood waters desynchronizes peak flows mini- herein are those of the author and do not necessarily mizing downstream flows. reflect the views of NOAA or any sub-agencies. Printed on recycled paper. AQ '41 " NW04"111-8 3 Wetland Types high marsh is a mixed community of saltgrass and salt- Generally it is common to group wetlands into cate- meadow hay with saltbush along the upland edge. Oyster- gories according to salinity regimes. Salinity influences catchers, egrets, herons, blue crabs, fiddler crabs and the kinds of plants and animals present in the wetland. finfish are common inhabitants of the saline marsh. Euhaline (Marine). Marine marshes have waters Brackish (Polyhaline and Mesohaline). The with a salinity close or equal to seawater. The vast expan- seawater is gradually diluted by freshwater and the salin- sive marshes behind the barrier islands on the eastern ity ranges from above and below 16 parts per thousand. shore are saline marshes. Dominant vegetative species in- The marshes are found in the lower Chesapeake Bay and clude saltmarsh cordgrass, saltmeadow hay, saltgrass and extending up the major tributaries and creek systems. highwater bush. Zonation of the vegetation is simple and The vegetation is more diverse than the saline mars 'hes. Bulrush, sea lavender and cattails are common in addition easily observed. Tall form saltmarsh cordgrass grows at lower elevations adjacent to the marsh creeks,@ short form to saltmarsh cordgrass, saltmeadow hay and sattgrass. saltmarsh cordgrass occurs in the mid-marsh, and the Saltmarsh cordgrass grows adjacent to the creeks, salt- grass and saltmeadow hay with sea lavender and black 'J i EXTREME HIGH TIDE N ......... . ...... ........ .. ....... .......................................................... UPLAND .MARSH.U LAND MEAN HIGH TIDE BORDER P ER ...... ............................. ............................ IIIG I A 'I PANNE MEAN LOW TIDE POOL .... ..... ....... LOWER HIGHMARS11 LOW MARS11 INTERTI D L ------- FLAT AQUATIC BED Eelgrass Widgeon Grass Smooth Cordgrass Glassworts Sea-Blites Marsh Fleabane Marsh Orach Salt Hay Grass Spike Grass Perennial Salt Marsh Aster Black Grass Hightide Bush Seaside Goldenrod Bulrushes Rose Mallow Common Reed Sea Myrtle Grass-Leaved Goldenrod Switchgrass Marsh Pink Slough Grass Figure 1. Generalized disiribution of vegetation in a salt marsh. Note increased plant diversity along marsh-upland border. (Tiner, 1985, 1987) 4 needlerush occupy the mid-marsh and the upland edge fresh community is much greater than other tidal wet- is marked by highwater bush and marsh'elder. Animals lands. Fifty species per acre is typical. Just a few of found in the brackish marsh include marsh snails, grass- the commorn plants are; arrow arum, pickerelweed, wild hoppers, clapper rails and fiddler crabs. rice, rice cutgrass, cattails, and smartweeds. Zonation Oligohaline. These wetlands form the gradient is less sharply defined, however a general profile may from brackish to tidal freshwater. Located along the be described (Odum, 1984). Arrow arum and pickerel- Bay, niajorSributaries and creeks, increased freshwater weed at the lower elevations adjacent the waterway, input dilutes the seawater to around 3 parts per thou- smartweeds, wild rice, big cordgrass and rice cutgrass sand. Saltmarsh cordgrass may still be found along the at higher elevations, cattails near the upland and wax waters edge, but many other less saline tolerant species myrtle and red maple along the upland edge. A large are also'found. Big cordgrass, sedges and three square - diversity of animals is a consequence of the great vari- grow here. Zonation patterns become less pronounced ety of plants. Amphibians and reptiles are common, as as salinity decreases. Saltmarsh cordgrass is still found are migrating waterfowl, muskrats, river otters, voles, at the lower elevations, big cordgrass, cattails, bulrush raccoons and many other animals. and marsh mallows may all occur at the higher eleva- Tidal Swamps. These wetlands are found at the tions. upper end of tidal influence. The presen ce of trees dif- Tidal Freshwater. The average salinity is 0.5 ppt, ferentiates tidal swamps from tidal marshes., In many or lower, except during drought. These wetlands occur swamps, the tidal swamp grades almost imperceptibly upstream of oligohaline marsh and downstream of non- into a nontidal swamp. Trees in tidal swamps generally Odal wetlands. The vegetative diversity of the tidal grow on hummocks. Trees within tidal range are gener- Figure 2. Food web. y TOP CARNIVORES 'Noma microalgae PS MARY IN CONSUMERS RW PRODUCERS AM-0 - : < bacterl 'k and' e MLW Protozoa -zooplanklon- SECONDARY CONSUMERS V DECOMPOSERS & DETRITUS FEEDERS TI- TERTIARY CONSUMERS SAND 9 MUDFLAT INFAUNA 5 ally smaller and show autumn colors earlier than trees Communities (Types are after the Virginia above tidal influence. Tree species found in tidal swamps Wetlands Cuidelines) include- gums, red maple, green ash and bald cypress. Dragonflies and other insects are common. Amphibians Type 1. Saltmarsh cordgrass community. and reptiles, including spring peepers, water snakes, bull- This community is dominated by Spartina alterni- frogs and turtles, are found in tidal swamps. White-tailed flora. This species is a tall grass reaching 4-6 feet tall deer, red fox and other upland animals wander into the near the waters edge. There are distinct growth habits of swamps. Beavers, which were once gone from the Spartina according to height. There is a tall form, a short Chesapeake Bay region, are once again inhabitants of form and some have proposed an intermediate height tidal and nontidal swamps. form of the species exists. The tall form is found growing adjacent to the marsh creeks and the short form grows fur- AVERAGE ANNUAL SALINITY MARSH TYPE NON-TIDAL FRESHWATER 0-0-0 0-0-0 (DURING LOW 11DAL FLOW LIMIT OF FRESHWATER CONDITIONS) TIDAL INFLUENCE * 0.5 Ppt - - - - - - - - - - - - - - - - - OUGOHALINE I E * 5.0 ppt - - - - - - -- - - - - - - - - S T T MESOHALINE U A < 1&0 Ppt - - - - - - R Y V0, POLYHALINE 1A < 30.0 ppt -------------- ---- J OCEAN EUHALINE (MARINE) Figure 3. The relationshigbetween marsh type and average annual salinity (values are approximate only). Terminology is based on Cowardin et al. (1979). (Odum et al, 1984) 6 ther from the creeks. It has been theorized that the tall with swirls. The vegetation is found growing from form plants receive a "tidal subsidy" from the increased about mean high water to the limit of spring tides. availability of tidal borne nutrients. Primary productivity is around 1-3 tons per acre. Spartina grows from about mean sea level to mean However, tidal flushing of this community is limited to high water which is approximately the-upper two thirds spring and storm tides so much of the vegetative mate- of the tidal range. The location of the community rial remains in the marsh and is not available to the ma- within the tidal zone results in daily flushing by the rine community. tide. The seed heads of the plants provide food for birds The primary productivity of Spartina is very high. and some birds nest in this community. Invertebrates The average production is about 4 tons per acre but like the saltmarsh snail (Melampus) are important food may be as. high as 10 tons per acre. Tidal action trans- for waterfowl, seaside sparrows and rice rats. pofts the detritus making it readily available to the ma- The dense growth habit of the vegetation provides nne community. an effective erosion deterrent, however, the position of Tidal fluctuations allow aquatic animals access to the community at. the higher elevations affords little op- the marsh. Adults of small fish species (killifish, ancho- portunity to guard against shoreline erosion. If the com- vie's) and juveniles of many species (striped bass, spot, monly occurring saltmarsh cordgrass is absent from the croaker) use the marsh for protection and a feeding lower elevations, this community will deter shoreline area. Blue crabs and shrimp also forage and find ref- erosion. The saltmeadow community is often the older uge in the Spartina. The marsh periwinkle, which part of the marsh with deep accumulations of marsh clings to the'stems of the vegetation, and the ribbed peat. Flood waters fro -in high tides and upland runoff marsh peat. The salt- mussel are important as food for aquatic animals as may be absorbed by the deep well as bird and small mammals. meadow acts as a trap, filtering the sediments, nutrients The roots and rhizomes of the plant are used as and toxics carried by the flood waters. food by waterfowl. Some birds use the vegetation as Type Ill. Black Needlerush Community. nesting material including the clapper rail and willet. This community is dominated by black needlerush Muskrats use the plants in the construction of lodges. (Juncus roemerianus). This plant usually grows in pure Raccoons and other small mammals can use the stands with saltmarsh cordgrass, saltmeadow hay and Spartina community as a foraging area. saltgrass occurring at the edges. The'plants have hard The Spartina community is an effective deterrent cylindrical stems tapering to a sharp tip. The'plants ap- to shoreline erosion. The high growing densities, of the pear leafless,as the leaves form a sheath around the vegetation reduce wave. energy. The,peat is very resis- stem. The species blooms from June to August. Jun-' tant to wave energy due to the dense growth of roots cus grows from about mean high water to somewhat be- and rhizomes. The baffling effect of the vegetation low spring tides. The community often grows in slows the flow of tidal waters and upland runoff. As pannes, low areas of the marsh where waters accumu- the water velocity is reduced, sediments are deposited late and evaporation increases salinity. on the marsh. This.property makes the Spartina com- Black needlerush produces about 3-5 tons per year munity effective as a trap for sediments from upland of vegetative material. Due to the rigid natureof the runoff. species, plants decompose slow 'ly and the location of Type 11. Saltmeadow co mmunity. the community in the high marsh limit's the amount of This community is dominated by saltmeadow hay material flushed by the tide. (Spartina patens) and saitgrass (Distichlis spicata).. It It does not appear that any animals feed on Juncus, is common to find saltmarsh cordgrass, marsh elder, however the dense, stiff vegetation provides cover for groundsel tree and other plants growing in this commu- some species. Clapper rails nest in the vegetation. nity. Saltmeadow hay and saltgrass reach about 1-2 The dense system of roots and rhizomes is resistant feet in height. Large expanses appear meadow-like to, erosion. The opportunity to provide shoreline ero- 7 sion protection occurs on some sandy shores and low on the stems and seeds. The meadow vole nests among sand berms. The vegetation is an effective trap for sedi- the stems as does the marsh rice rat, often taking over the ment, although not as effective as the densely matted salt- nest of a marsh wren. meadow community. The absorbent substrate acts to As other species of marsh grass, the dense network buffer coastal flooding. of roots and rhizomes binds the marsh substrate providing Type IV. Saltbush Community. a deterrent to shoreline erosion. Growing at higher eleva- This community is comprised.of groundsel tree or fions in the tidal zone the opportunity to provide shoreline highwater bush (Baccharis halimifolia) and marsh elder erosion control is limited; however, it may be particularly (Ivafrutescens). Saltmeadow hay, and saltgrass are often important.for low salinity wetlands where the lower inter- found growing underneath the bushes. Groundsel tree tidal zone is occupied by species that die back each fall. reaches about 15 feet in height, the leaves are alternately Big cordgrass often is associated with the deep marsh arranged on the branches, and it has conspicuous yellow- peat common at higher elevations in the older portion of white flowers which bloom September through October. the marsh. The deep peat increases the capacity for hold- Marsh elder is 3 to 10 feet high, the leaves are opposite ing flood waters. Where the community is found growing on the branches, and the flowers are small greenish-white adjacent to the uplands, flood buffering may allow for the appearing from August to October. The bushes grow at settling and trapping of upland sediment, nutrients and the upper limit of the marsh (the marsh-upland ecotone). toxics. The plants are also effective at trapping larger flot- Primary productivity is low, about 2 tons per year, sam. and little vegetative material reaches the marine environ- Type V1. Cattail Community. ment. The shrubs provide diversity to the grasses and There are two common species of cattails in this com- reeds common to the marsh. Small birds, like the marsh munity, the narrowleaf cattail (Typha angustifolia) and wren, nest among the branches. the broadleaf cattail (Typha latifolia). The plants may be The growth habit of the shrubs does not serve well as found growing with other brackish and freshwater plants a sediment trap, but does effectively trap larger flotsam. including; sedges, bulrushes, smartweeds, arrow arum Often found growing on sand berms in front of small and pickerelweed. Both species grow to heights of 4 to 6 marshes, the bushes do provide some erosion protection. feet, have long strap-like leaves with slightly rounded tips and have the characteristic fruiting head. Both the male Type V. Big Cordgrass Community. and female flowers occur together on one flowering Big cordgrass (Spartina cynosuroides) usually grows in monotypic stands. The grass is- very tall, 6 to 12 spike. In the broadleaf cattail the upper (male) spi@e and feet, and has a distinct feathery fruiting head in the fall. the lower (female) spike are touching, in the narrowleaf The plants are very sturdy and standing dead plants often there is a space between the spikes. persist through the winter. Big cordgrass grows from just Cattails prefer very wet soils and sometimes grow in above mean high water up to thq upland margin. The standing water. They prefer low salinity waters. Look plant is most common in brackish and lower salinity for cattails along the upland margin where groundwater marshes. seepage keeps the soils wet and the salinity low. The Levels of primary productivity range from 3 to 6 tons vegetative material produced by cattails,, about 2 to 4 tons per acre in this community'. Primary productivity rates per acre, is not readily accessible to the marine environ- are similar to saltmarsh cordgrass and higher than most ment. other species growing above mean high water. However, Some species of birds use cattails as habitat. Marsh the position of the community at the higher elevations re- wrens and rice rats build nests attached to the plants. The plant roots are eaten by muskrats. sults in the transport of the vegetative material being lim- ited to high spring, wind and storm tides. The cattail community offers little erosion protection - Various animals use big cordgrass for cover, food or due to the preferred location adjacent to the uplands and building materials. Muskrats use the material for build- the shallow root system of the plants. However, both of ing lodges. Geese eat the rhizomes and other birds feed 8 these characteristics make the community effective as a leaves are shed in the fall, individual stems and seed sediment trap, particularly from upland runoff. -heads often persist through the winter. Phragmites usually grows above mean high tide, Type V11. Arrow Arum - Pickerel Weed Community. . particularly in brackish areas. The plants will also-in- The dominant plants of this community are arrow vade the drier areas of disturbed sites. Primary produc- arum (Peltandra virginica) and pickerelweed (Ponted- tivity rates range from 4 to 6 tons per acre, but the eria cordata). The plants may grow in association with relatively high elevation occupied by the community re- sedges, smartweeds, cattails and pond lily. Both spe- sults in little transport of vegetated material. cies have dark green fleshy leaves and grow in clumps There is little evidence of wildlife use of this com- about 3 feet high. Arrow arum has leaves shaped like munity. The vegetation may provide cover for some birds and small mammals. Where Phragmites invades arrowheads, with a prominent midvein. Arrow arum blooms from May to July, but the flowers. are concealed disturbed sites, there may be a detrimental effect on wildlife use due to the decrease of vegetative species in a sheath called a spathe. Pickerelweed has heart- shaped leaves with parallel v .eins.. The plants bloom which are important to wildlife. from May to October and the showy purple flowers are The sturdy nature of the grass makes for a good borne on flowering spikes up to 4 inches long. shoreline erosion deterrent. Due to the ability to invade Arrow arum and pickerelweed can be found in non- dry, disturbed areas, this species is often found growing on dredge material acting to stabilize the material. tidal freshwater wetlands as well as low salinity and tidal freshwater wetlands. In tidal waters, the commu- Type IX. Yellow Pond Lily Community. nity grows from'about mean sea level to about mean Yellow pond lily (Nuphar luteum) grows in fresh- high tide in low salinity and freshwater marshes. This water and may be associated with arrow arum and pick- is the same landscape position occupied by saltmarsh erelweed. The large saucer shaped leaves float on the cordgrass in higher saline waters. water surface at high tide. The plants flower from June About 2 to 4 tons per acre of vegetative material is to September and have a large yellow flower. The produced in this community. The organic material is leaves and flowers of the 2 to 4 foot tall plants are emer- readily available to the marine system. Unlike many gent at low tide, other common welland plants such as the marsh The floating leaves of one plant occupy several grasses, these species decompose quickly and com- square feet so plant densities are generally low. Pri- pletely in the fall. mary productivity approaches one ion per acre. Vegeta- The soft, unconsolidated intertidal muds are suscep- tive material enters the marine environment, but the tible to erosion. Without a dense network of roots and contribution is small compared to other marsh species. rhizomes, the plants can serve to stabilize the shoreline Yellow pond lily is found in river stretches associ- only during the year when the leaves can act to baffle ated with spawning striped bass, white perch and river wave energy. Due to the complete -die back of -the vege- herring. The plants provide attachment sites for algae tation each fall there is -no standing dead material to pro- and aquatic animals. Wading herons, egrets and other tect the shoreline from erosion in the winter. The large waterbirds forage on the small fish and invertebrates leaves of the vegetation can slow tidal waters and among the floating leaves. Finfish also forage in the waves to allow some sediment deposition. yellow pond lily. A resident of freshwater marshes, the Type VI I[I. Reed Grass Community. eastern painted turtle may be found feeding here as This community is dominated by reed grass well. (Phragmites communis). This grass often grows in While lacking stiff stems and leaves, the large pure stands, but may be found growing with saltbush leaves do provide some baffling of wave energy. By re- ducing wave action, the community provides some and switchgrass. Phragmites is a tall stiff grass reach- ing 6 to 10 fe .et in height. The plants bloom in July and shoreline protection and allows fo Ir the settling of some August and have a large, feathery seed head. After the water borne sediments. 9 Type X. Saltwort Community. arrow arum, pickerelweed, smartweeds, wild rice and rice There are three species of vegetation commonly cutgrass are among the species which may occur in a called saltwort found in Virginia. The three species are: freshwater mixed community wetland. Because this com- Salicornia virginica, Salicornia biglovii and Salicornia munity encompasses many vegetative species, each with europa. Saltworts are leafless succulents. with thick a preferred elevation, the community may be found grow- green, edible stems. S. europa and S. biglovii are dif- ing from the subtidal zone to the upper limits of wetlands. fusely branched, fleshy annuals. Salicorma virginica is a Primary productivity of this community depends perennial with woody stems and typically grows in dense greatly on the species present. An average value is 3 to 5 mats, Saltworts grow in saline and higher salinity brack- tons per acre. The vegetative material produced by interti- ish marshes above mean high tide in pannes and sparsely dal species is readily available to the marine environment. vegetated areas. The high diversity of the freshwater mixed wetland Primary productivity is less than 1/2 ton per acre. provides foraging areas, nursery areas and nesting sites There is some evidence that some ducks eat the stems. for a myriad of wildlife species. These wetlands occur Wading birds and other marsh birds may forage where along river and creek stretches where striped bass, shad -small fish and invertebrates get trapped by the receding and river herring spawn. The juvenile fish can find shel- tide in Salicorma pannes. ter among the intertidal vegetation. Wildrice, arrow arum The diminutive habit of the leafless plants, the shal- and smartweeds are consumed by native and migratory low root system and the preferred location of this commu- waterfowl, sora rails, redwinged blackbirds and upland nity offers little erosion- protection, flood buffering or birds. Insects, amphibians and reptiles are more common sediment trapping capabilities. than in the brackish and saline marshes. Type X1. Freshwater Mixed Community. The shoreline erosion deterrent of this community is This community type applies to freshwater wetlands the same as the arrow arum and pickerelweed community with a heterogeneous mix of vegetation and no single spe- 'because these species grow at the lower elevations adja- cies covers more than 50 percent of the site. Bulrushes, cent to the waterways. However, due to-the presence of grasses, sedges, rushes and a peaty substrate at the higher Figure 4. Marsh to uplands transition in a typicalfreshwater marsh. BIDENS (seasonal) N N-TIVAL DEPRESSION GREENBRIER BUTTON BUSH SEDGES SMARTWEED RED MAPLE TOUCH-ME-NOT MARSH MALLOW ARROW ARUM POISON IVY ROYALFER TEARTHUMA PICKEREL WEED SWEET PEPPERBUSH- CATTAIL 810 CORDGRASS ARROWHEAD SWAMPROSE- IRONWEED .11- 1 - YELLOW POND LILY SOUARESTEM SPIKEFIUS SEDGES MARSH FERN WILD RICE 10 elevations, the freshwater mixed community is effec- As this community is most often bordered by salt- tive as a sediment trap and assimilator of flood waters marsh cordgrass, the community provides effective (Figure 4). shoreline erosion control. Likewise, the attributes of Type X11. Brackish Water Mixed Community. the commonly, occurring saltmeadow community spe- This community is comprised of various brackish cies, saltmeadow hay and saltgrass, provide effective wetlands species. No single species cover more than sediment trapping and flood buffering capabilities 50 percent of the site. This diverse vegetative commu-' (Figure 5). nity may include saltmarsh cordgrass, saltmeadow hay, Type X111. Intertidal Beach Community. saltgrass, saltbushes, threesquares, big coidgrass, com- The intertidal beach is nonvegetated. The domi- mon reed and cattails: This community may extend nant animal species are burrowing invertebrates. Ani- from the lower limit of saltmarsh cordgrass, about mals that live in the substrate are called infauna. mean sea level, to the upland margin. Wandering species and nonmobile species attached to Primary productivity of the brackish mixed com- hard substrates are called epifauna. There is some vafi- munity ranges from 3 to 5 tons per acre. The detritus of ability between species commonly found on bay the intertidal species is available to the marine food beaches and those most often associated with ocean web. beaches. On the ocean beach, mole crabs, donax clams As is the case with the freshwater mixed wetlands, and haustorid amphipods are common. Amphipods are a wide diversity of vegetation provides a variety of also common on bay beaches as are oligochaete worms wildlife food and shelter. Many species of waterfowl and sand fleas. These animals occur in large numbers; feed on the seed heads. Shorebirds and wading birds in the summer there may be as many as 5000 individu- forage for plentiful invertebrates. Fiddler crabs and als per square meter- marsh crabs are common. Raccoons forage for crabs Lacking the macrophytes, large plants, of the vege- and shellfish. Small mammals such as the meadow tated wetlands, primary production is relatively low. vole and rice rat, feed on the vegetation and inverte- The primary producers of the nonvegetated wetlands brates. Foxes are attracted by the small mammals, are microscopic plants including algae and phytoplank- marsh birds and invertebrates. Figure 5. Marsh to uplands transition in a typical saltwater marsh. ---------- TRANSITION ZONE ----------- SALTMARSH BULRUSH ALOM FRIF"ATER STREAMETS A... PINE GROUNDSEL TRE MARSH ELDE BLACK NEEDLERUSH GREENBRIER SEA OXEYE RA SALTMEADOW HA SPARTINA ALTERNIFLORA RED CEDAR FOXTAIL GRASS SALTMEAD HAY Short form WAX MYRTLE GREENBRIER SALT GRASS Tall form GOLDENROD ton. The production of animals, or secondary production, population densities are generally higher than sand or is very high. mud flats and range from 5300 to 8300 individuals per The large numbers of invertebrates are important square meter. food for shorebirds. At high tide, finfish and blue crabs Primary production is similar to sand flats. How- also forage for food. The areas above high tide are used ever, the smaller particle size and high organic matter con- as nesting sites for terns, plovers and skimmers. tent, means greater microbial activity. Bacteria and The low slope of the beach makes it a natural wave fungi, fundamental to the breakdown of detritus, are dissipator. The dissipation of wave energy reduces the found in higher numbers in sediments containing small potential for shoreline erosion. The beach interacts with grain size particles. Nutrient cycling activity is greater dunes, offshore bars and adjacent shoreline in the distribu- than on the sand flat. tion of sand. Shorebirds and wading birds feed on the invertebrate Type XIV. Sand Flat Community. populations of the sand/mud flat. Waterfowl are attracted Sand flats are populated by various surface dwelling by the mollusks. Blue crabs and fish use the area for for- and burrowing animals. Sandworms, bloodworms, am- aging and shelter. This community often borders a vege- phipods and clams are all found on the sand flat. Animal tated marsh which may increase the role of both densities range from 300 to 3000 individuals per square 'communities as wildlife habitat. The marsh provides meter. habitat variety and organic material to support nutrient cy- Most of the animals complete their life cycle in one cling and food webs. or two years. High rates of production and predation re- Shoreline erosion control is similar to sand flats. sults in a rapid turnover of individuals. High predation Type XVI. Mud Flat Community. acts as a control on the size of the animals, in general indi- Mud flat animal populations are dominated by viduals are small. spionid- worms, mud snails, razor clams and bloodworms. Primary productivity by microscopic plants ranges These animals are mostly surface dwelling or shallow bur- from 100 to 200 grams per square meter. Primary con- rowers. The instable, anoxic mud limits the presence of sumers graze on the small plants incorporating the vegeta- deep burrowers and deters species other than surface detri- tive material into the food web. This community has a tal feeders. Animal -populations are highly variable. Gen- reduced role in nutrient cycling due to the low percentage erally lower than sand/mud flats but higher than sand of organics and the large particle size of the sand. flats, densities range from 50 to 5000 animals per square . Sand flats are used as foraging areas, by shorebirds, meter.* finfish and blue crabs. Juvenile blue crabs and finfish Of the nonvegetated wetlands, mud flats probably seek shelter in the shallow waters where larger aquatic have the highest primary productivity. Mud flats interact animals cannot go. Some sand flats support high shellfish with adjacent vegetated areas in nutrient cycling. The populations. marshes provide organic material to the animal popula- Like the intertidal beach, the low slope of this co in- tions of the mud flats which breakdown and transform the munity acts as a natural dissipator of wave energy reduc- organic material. ing the potential for shoreline erosion. Foraging wildlife are attracted by the invertebrate Type XV. Sand/Mud Mixed Flat Community. populations. Waterfowl, wading birds, finfish and blue crabs all feed on the mud flats. Hard clams, soft clams, mud snails, parchment worms and polychaetes are commonly associated with in- Mud flats occur frequently in quiescent areas and tertidal flats composed of sand and mud. The animals-are have a limited opportunity to control shoreline erosion. - surface dwellers, burrowers and some build permanent However, the gradual slope of the mud flat acts like the tubes. Variability in particle size allows some overlap of other nonvegetated wetlands in the dissipation of wave en- species common to sand flats and mud flats. As a result, ergy- NOAA COAST ly 12 862 P Type XVII..Intertidal Oyster Reef Community. The nooks and crannies of the reef offer niches for Oysters are the dominant- animal, but there is great manyaquatic animals. Small finfish find shelter among diversity in the attached and associated organisms. the shells, larger fish forage on the reef. Blue crabs Hard clams, mud crabs, sand w'orms, barnacles and feed on the reefs extensively. Oystercatchers and other sponges are.associated with oyster reefs. When oyster birds also forage on the reefs. reefs are managed by man, the oysters are most com- The rigid structure of the permanently attached mon. In a natural state, fouling organisms are d 'omi- shells are resistant to erosion and may dissipate wave nant. Populations are very variable, but are generally energy. greater than flat communities. Bibliography Chapman, V.J. 1960. Salt marshes and salt deserts of Stickney, R.R. 1984. Estuarine ecology of the southeast- the world. lnter@cience Publishers, Inc. New York, em United States and Gulf of Mexico. Texas A&M New York. University Press. College Station, Tx. 310pp. Long, S.P. and C.F. Mason. 1983. Saltmarsh ecology. Tiner, R.W. 1985. Wetlands of New Jersey. U.S. Fish Chapman and Hall. New York, New York. and Wildlife Service, National Wetlands Inventory, Mitsch, W.J. and J.G. Gosselink. 1986. Wetlands. Van Newton Comer, MA. 117 pp. Nostrand Reinhold. New York, New York. Tiner, R.W. 1987. Mid-Atlantic wetlands: a disappear- Odum, W.E., T.J. Smith Jr., J:K. Hoover and C.C. ing natural treasure. U.S. Fish and Wildlife Serv- McIvor. 1984. The ecology of tidal freshwater ice. Newton Comer, MA. 28pp. marshes of the United States east coast: a commu- Virginia Wetlands Guidelines. Virginia Institute of Ma- nity profile. U.S. Fish Wildl. Serv. FWS/OBS- rine Science and Virginia Marine Resources Com- 83/17. 177pp. mission, VMRC, Newport News, VA. 57 pp.. Silberhom, G.M. 1982. Common plants of the Mid- White, C.P. 1989. Chesapeake Bay: a field guide. Tide- Atlantic coast: a field guide. Johns Hopkins Univer- water Publishers. Centreville, MD.'212pp. sity Press. Baltimore, MD. 256pp.