boundary ecosystem

boundary ecosystem


      complex of living organisms in areas where one body of water meets another, e.g., estuaries and lagoons, or where a body of water meets the land, e.g., marshes. The latter are often called wetlands.

      Boundary ecosystems are characterized by the presence of large plants (plant). In the open water of the ocean and large lakes the basic production of living material (primary production) is carried out by microscopic algae ( phytoplankton) floating freely in the water. At the bottom there is not enough light to allow growth of large, attached plants. In boundary ecosystems much of the area is shallow enough for light to reach the bottom and permit large plants to grow. Phytoplankton is also present, but the large plants give the boundary systems their special character.

Boundary systems between waters

Estuaries (estuary)
 Estuaries are places where rivers meet the sea and may be defined as areas where salt water (seawater) is measurably diluted with fresh water. On average, estuaries are biologically more productive than either the adjacent river or the sea because they have a special kind of water circulation that traps plant nutrients and stimulates primary production. Fresh water, being lighter than salt water, tends to form a distinct layer that floats at the surface of the estuary. At the boundary between fresh and salt water, there is a certain amount of mixing caused by the flow of fresh water over salt and by the ebb and flow of tides. Additional mixing may be caused from time to time by strong winds and by internal waves that are propagated along the interface between fresh and salt water. Four types of estuary are recognized according to the degree of mixing: salt wedge estuaries, partially mixed estuaries, vertically homogeneous estuaries, and fjords (Figure 1—>).

 A salt wedge estuary has minimal mixing and the salt water forms a wedge, thickest at the seaward end, tapering to a very thin layer at the landward limit (Figure 1—>). The penetration of this wedge changes with the flow of the river. During flood conditions the wedge will retreat; during low flows it will extend farther upriver. The mouth of the Mississippi River in the United States is a classic example. The mixing at the boundary between fresh and salt water causes the surface layer to entrain salt water and become more saline as it moves toward the sea. To compensate for the entrained salt water there is a slow movement of the salt water up the estuary at depth. Because bottom waters are rich in nutrients (nutrient) derived from decomposing plant and animal remains, this circulation has the effect of pumping nutrients into the estuary and stimulating biological production.

      Organic and inorganic particles carried by rivers tend to flocculate (aggregate into a mass) and sediment out when they encounter salt water. They sink from the freshwater layer to the salt wedge and are carried upstream. When the organic matter decomposes, it adds still more nutrients to the estuary. The inorganic matter settles on the bottom and provides an enriched sediment for flowering plants adapted to salt water. Between the tide marks, mangrove forests flourish in tropical conditions, while salt marshes form in temperate and subarctic conditions. Below low tide, sea grasses form dense beds on muddy substrates. In areas of an estuary where water movement is vigorous enough to remove sediment, leaving a stony or rocky bottom, rooted plants are replaced by seaweeds. These have a special structure known as a holdfast, which attaches itself to any hard surface. Phytoplankton floating freely in the water benefits from the high level of nutrients, especially near the head of the estuary, and grows rapidly. It provides food for the microscopic animals in the water column, the zooplankton. As this community is carried downstream in the surface waters, dead organisms and the fecal pellets of the animals sink toward the bottom and enter the salt wedge to be carried back to the head of the estuary. As they decompose they add still more nutrients to the water.

 In a partially mixed estuary, the vigorous rise and fall of the tide generates strong turbulence and causes partial mixing between the fresh water above and the salt water below (Figure 1—>). Under these conditions the river flow entrains 10 to 20 or more times its own volume of salt water, and the compensatory landward flow of seawater near the bottom is correspondingly increased. The effect of the Earth's rotation (Coriolis effect) is to cause the surface flow to be stronger on the right-hand side facing seaward, or the opposite in the Southern Hemisphere. The bottom flow is stronger on the opposite side of the estuary.

 In a vertically homogeneous estuary the river flow is weak and the tidal flow is strong (Figure 1—>). Consequently, all stratification is broken down and salinity is almost the same from top to bottom at any given place. The salinity is lowest where the river enters the estuary and highest near the sea.

 The fjord-type (fjord) estuary was originally formed by a glacier and has a submerged ridge, or sill, near its mouth, composed of glacial deposits (Figure 1—>). It may be regarded as a partially mixed estuary in which the bottom has been replaced by a basin of undiluted seawater held in place by the sill. When entrainment in river flow causes a strong landward flow at the bottom, water rises over the sill and enters the estuary at intermediate depth, leaving the deep waters undisturbed. Only major intrusions of seawater caused by storms can displace the deep water. Owing to their glacial origin, fjords commonly have steep sides and very little shallow water. Hence, the development of salt marshes or sea-grass beds is minimal, but seaweeds colonize the rocky shores.

      The high level of plant production in estuaries supports a correspondingly high level of production of invertebrate animals and fish. Estuaries often contain beds of shellfish such as mussels and oysters and large populations of shrimps and crabs. Fish such as plaice and flounders are common. Other species use the estuaries as nursery grounds. Organisms in early stages of development enter the salt wedge at the seaward end and are carried up the estuary by the bottom currents. Juveniles find abundant food as well as protection from predators in the mangrove forests, salt marshes, or sea-grass beds that line the estuary. Later, they may migrate (migration) to the open ocean to continue their growth and development. Other species pass through the estuaries in the course of their migrations. For example, salmon migrate from the sea to the rivers to spawn, while the young fish later migrate back to the sea. Eels migrate in the opposite direction, breeding in the sea but returning to fresh water as juveniles (see marine ecosystem: Patterns and processes influencing the structure of marine assemblages: Migrations of marine organisms (marine ecosystem)).

      Many estuaries are now important sites for aquaculture. There is a long history of mussel culture along the coast of Spain, and Norwegian fjords are much used for salmon culture. In Southeast Asia, artificial ponds are created in mangrove forests and used to culture shrimp. Because estuaries are located at the mouths of rivers, they have been favoured sites for the development of human settlements. This has made them particularly vulnerable to contamination by sewage and industrial effluents. The characteristic circulation that serves to trap natural plant nutrients may also retain high concentrations of pollutants.

Lagoons (lagoon)
      Lagoons are bodies of water partially or completely separated from the open ocean by barriers of sand or coral. In coastal lagoons the barrier most often is formed and reinforced by the movement of sand in alongshore currents. Coral lagoons occupy the space between a coral reef and the shore or within the central basin of a coral atoll. Lagoons are characteristically shallow, and, with an abundance of large plants, they are highly productive.

      The circulation of water in a coastal lagoon is very dependent on the amount of land drainage. A lagoon into which a major river flows is known as an estuarine lagoon and may be regarded as a special kind of estuary. There are, however, many cases in hot arid regions in which lagoons lose more water by evaporation than they receive from land drainage. This causes surface waters to become more dense than seawater and to sink to the bottom. Seawater flows in at the surface to replace that lost by evaporation (vaporization), creating a circulation the reverse of that found in estuaries. If exchange with the open sea is limited, the lagoon may become much more saline than the open sea. Consequently, various species of plants and animals have become adapted to life in high salinities (see biosphere: The organism and the environment: Environmental conditions: Salinity (biosphere)).

Kenneth H. Mann

Boundary systems between water and land
      Wetlands (wetland) generally occur at the interface between terrestrial ecosystems, such as upland forests and grasslands, and aquatic ecosystems, such as rivers, deep lakes, and oceans. Thus, wetlands are neither wholly terrestrial nor wholly aquatic but exhibit characteristics of each. They also depend greatly on both. Wetlands include swamps, bogs, marshes, mires, fens, and other wet ecosystems found throughout the world under many names.

      A wetland usually is defined by its physical, chemical, and biological processes—i.e., its hydrologic conditions, physicochemical environment, and biota. However, the precise definition of “wetland” is somewhat elusive because the ecosystem is an intermediate in a continuous gradient from water to land. The three basic features that distinguish wetlands from aquatic and terrestrial systems—shallow water or saturated (saturation) soil, unique soil conditions in which organic matter decomposes slowly, and vegetation adapted to wet conditions (hydrophytes)—have a considerable range of manifestations. Wetlands range from being permanently to intermittently flooded, and this affects the anaerobic (oxygen-free) conditions and the many types of plants, animals, and microbes that adapted to the conditions of the ecosystem. The current regulatory definition of a wetland in the United States requires that the land be inundated or saturated with a frequency and duration sufficient to support a prevalence of vegetation typically adapted for life in saturated conditions. It also requires the presence of saturated soils or standing water for at least part of the growing season.

 Wetlands are ubiquitous, found on every continent except Antarctica and in every clime from the tropics to the tundra (Figure 2—>). They are estimated to cover 4 to 6 percent of the Earth's land surface (5.3 to 8.6 million square kilometres [2 to 3.3 million square miles]). In humid, cool regions wetlands occur as bogs, fens, and tundra; along temperate, subtropical, and tropical coastlines as salt marshes, mud flats, and mangrove swamps; and in arid regions as inland salt flats, seasonal playas, and vernal pools. They occur along rivers and streams as riparian wetlands, seasonally flooded forests, and backswamps, and they are found in the deltas of the world's great rivers. Rice paddies (paddy), a human-created type of wetland often formed where natural wetlands used to be, are estimated to cover an additional 1.3 to 1.5 million square kilometres around the world. It has been estimated that half the world is fed from these domesticated wetlands.

      While there are many kinds of wetlands, the types can be reduced to two major categories—coastal and inland systems.

William J. Mitsch

Coastal (coast) systems
      Wetlands in coastal areas can be classified into three basic types: mangroves, salt marshes, and freshwater tidal marshes. Other important coastal systems not formally considered wetlands but found at the boundary between land and water are seaweed-based systems, sea-grass beds, and coastal mudflats.

      The fundamental characteristics of shoreline ecosystems are determined by the amount of energy in the water available to move sediments. This energy is supplied by wind-driven currents, tidal currents, and wave action. In high-energy areas the fine sediment is carried away, leaving bedrock, boulders, or cobbles. This creates a prime habitat for seaweeds. As the energy level of water movement progressively lessens, sediments ranging from pebbles to sand, silt, and mud can settle and remain in place. Soft sediments provide a suitable habitat for salt marshes or mangrove forests between tide marks and for sea grasses below the low-tide mark. On a coastline consisting of alternating headlands and embayments, the headlands are most likely to be exposed to strong wave action and to be inhabited by seaweed communities, while the sheltered embayments are more likely to have soft sediments with rooted plant communities. The characteristics of shoreline communities are discussed according to the type of plant production on which they are based.

Mangrove swamps
 Mangrove swamps are found along tropical and subtropical coastlines throughout the world, usually between 25° N and 25° S latitude. The mangrove swamp is an association of halophytic trees, shrubs, and other plants growing in brackish to saline tidal waters of tropical and subtropical coastlines. This coastal forested wetland (called a “mangal” by some researchers) is infamous for its impenetrable maze of woody vegetation, unconsolidated peat, and many adaptations to the double stresses of flooding and salinity. Approximately 68 species of mangrove trees exist in the world. Their uneven distribution is thought to be related to continental drift and possibly to transport by primitive humans. Mangrove swamps are dominant particularly in the Indo-West Pacific region, where they have the greatest diversity of species—30 to 40 species of mangroves, compared with about 10 species in the Americas.

      In the tropics and subtropics the intertidal areas of soft sediment are usually colonized by mangrove trees. Beneath them lies a waterlogged mixture of mud and decaying mangrove leaves that has very little oxygen; an aboveground root system allows the trees to take in air. This network of aerial roots forms a tangled mass that traps sediment but makes a mangrove forest very difficult for large animals (or humans) to penetrate. Small seaweeds and microscopic algae grow on the trunks and roots of the mangroves, and microscopic algae grow on the surface of the mud. This substrate, along with the decaying mangrove leaves, supports a rich and diverse animal community. Crabs and shrimps are often abundant, and clams and snails of many kinds abound. Mudskippers (family Periophthalmidae), which are fish that have developed the capability of leaving the water and moving over the mud surface in pursuit of prey, are found in mangrove systems, as is the mud lobster (Thalassina anomala), which lives in burrows. Because the plankton of adjacent coastal waters is often relatively unproductive, the productivity of the mangrove forests is an important element of the productivity of the whole coastal zone.

Salt marshes (salt marsh)
      Along intertidal shores in middle and high latitudes throughout the world, salt marshes replace the mangrove swamps of tropical and subtropical coastlines. These marshes flourish wherever the accumulation of sediments is equal to or greater than the rate of land subsidence and where there is adequate protection from destructive waves and storms. Dominated by rooted vegetation—primarily salt-tolerant grasses (grass)—that is periodically inundated with the rise and fall of the tide, salt marshes have a complex zonation and structure of plants, animals, and microbes. This biota is tuned to the stresses of salinity fluctuations, alternate drying and submergence, and extreme daily and seasonal temperature variations. Salt marshes are among the most productive ecosystems of the world. A maze of tidal creeks that exhibit fluctuating water levels and carry plankton, fish, and nutrients crisscross the marsh, forming conduits for energy and material exchange with the adjacent estuary. The salt marsh forms an important interface between terrestrial and marine habitats.

      The most common site for a salt marsh, after estuaries and lagoons, is on the sheltered side of a sand or shingle spit. Alongshore currents deposit coarser material on beaches but carry the fine material until it reaches the quieter water behind the barrier. As plants colonize the area, they slow down the flow of water and cause even more silt to accumulate. The Atlantic coast of North America has over 600,000 hectares (2,300 square miles) of salt marshes dominated by the marsh grass Spartina.

 On the European side of the North Atlantic the flora includes other important components such as the sea pink (Armeria), sea lavender (Limonium; see photograph—>), and sea plantain (Plantago maritima). In the course of history large areas of salt marsh in Europe have been used for grazing cattle and sheep, and these areas subsequently have been dominated by the grasses Puccinella and Festuca. Early colonists in North America often erected dikes around the marshes to keep out the sea; the reclaimed land was used for agriculture in much the same way that it had been in Holland and Belgium.

      Only a very small proportion of salt marsh vegetation is eaten directly by animals. The remainder dies, decays, and becomes suspended as fine particles (detritus) in the water. It was thought at one time that the export of this detritus on every ebbing tide supplied large amounts of nutritious food material to the animals in nearby estuarine or coastal waters. Detailed field studies have failed to support this view, and it is now thought that most of the production of salt marsh plants is decomposed by bacteria and fungi and that the plant nutrients are recycled within the marsh. Salt marshes are important habitats for oysters, shrimps, crabs, flatfish, and mullet. They also support large numbers of birds that stop over in the course of migration.

Freshwater tidal marshes
      This category includes freshwater marshes close enough to coasts to experience significant tides but far enough upriver in the estuary to be beyond the reach of oceanic salt water. This set of circumstances usually occurs where fresh river water runs to the coast and where the morphology of the coast amplifies the tide as it moves inland. Freshwater tidal marshes are interesting because they receive the same “tidal subsidy” as coastal salt marshes but without the stress of salinity. They act in many ways like salt marshes, but the biota reflect the increased diversity made possible by the reduction of the salt stress found in salt marshes. Plant diversity is high, and more birds use these marshes than any other marsh type. In most parts of the world, the location of freshwater tidal marshes corresponds to sites determined by humans as optimal for habitation and eventual development of cities—i.e., those areas that provide a reliable source of fresh water as well as a physical connection to the sea for ships. Thus freshwater tidal marshes are among the wetland types that have been most altered or destroyed by urban development around the world. Examples of the impact human development has had on wetlands are found in Chesapeake Bay and the lower Delaware River in the eastern United States.

Seaweed-based (seaweed) systems
 In seaweed-based systems seaweeds vary in size from giant kelps 40 metres (130 feet) or more in length, through the common rockweeds that are 1 or 2 metres long, to species that are so small as to be barely visible. They are algae and differ from flowering plants in having a holdfast instead of roots, a stipe instead of a stem, and a blade or thallus instead of leaves (see algae). They depend on water movement to continuously provide nutrients, which they take up through the surface of the blade. kelp is a general term for large brown algae of the order Laminariales (Figure 3—>). They live predominantly just below low-tide mark and form dense beds reminiscent of underwater forests. They absorb a great deal of wave action, helping to defend shorelines against storms.

      The giant kelps that occur along the Pacific coast of the United States and South America have been studied extensively because they are harvested for the extraction of alginates and other substances used in food processing. Typically growing in about 10 metres of water, they have large holdfasts from which several stipes originate. A young stipe grows as much as 45 centimetres (18 inches) per day, reaches the surface of the water, and then trails downstream. A large number of relatively small blades grow from the stipe and form a surface canopy with which they intercept light and nutrients. Giant kelp beds are home to a rich variety of invertebrates and fish, and, in many regions, to the sea otter (Enhydra lutis). Sea otters were once abundant around the North Pacific rim from Japan to California, but their range was greatly reduced by hunting. They have recently been reintroduced and populations are growing in many parts of British Columbia in Canada and Washington, Oregon, and California in the United States. Sometimes the sea urchin Strongylocentrotus becomes extremely abundant; in the course of feeding on the stipes of the kelps it may destroy kelp beds over large areas. The sea otter is a predator of sea urchins, and where it is abundant it has been shown to control sea urchin numbers. Abalone, a favourite food of sea otters as well as humans, are often abundant in kelp beds.

      The characteristic kelps of the North Atlantic are species of Laminaria that grow in dense beds but extend only one or two metres above the bottom. A characteristic inhabitant of these kelp beds is the Atlantic lobster, Homarus americanus, which includes sea urchins in its diet. In the 1970s in Nova Scotia, Can., there was a major outbreak of destructive grazing by sea urchins. This outbreak was accompanied by a sharp decline in lobster populations, suggesting that when lobsters are scarce sea urchin numbers proliferate. However, the question of whether lobsters control sea urchin numbers is still undecided. In the Southern Hemisphere Macrocystis and Laminaria also occur, but the giant kelp Lessonia is important in South America, as is Ecklonia in South Africa and Australia.

      Rockweed (Fucus) is a general term for the familiar brown seaweeds of the order Fucales, which grow between high- and low-tide marks (the intertidal zone (littoral zone)) on rocky shores. In the Northern Hemisphere Fucus and Ascophyllum are common genera. The latter may be recognized by possession of small air-filled bladders on the fronds. It usually grows in more sheltered locations than Fucus. The intertidal zone is of interest because of the zonation of organisms that occurs there. It is in many ways an ideal laboratory in which to study the factors controlling the population size of seaweeds and invertebrates. A high proportion of the animals and algae in this zone are firmly attached to the rocks in order to withstand the force of waves breaking on the shore. Attached fauna include barnacles (barnacle), limpets (limpet), periwinkles (periwinkle), and mussels (mussel). Barnacles are crustaceans that are attached to rocks along their backs, with upward-pointing legs that are surrounded by a row of protective hard plates. Limpets are mollusks that live under a very strong conical shell and cling to the rock by an adhesive “foot.” Barnacles filter fine particles of seaweed and plankton from the water, while limpets graze on the very small algae growing on the rock surface. Periwinkles are marine snails with hard shells that find shelter among the rockweeds on which they browse. Clamlike mussels are able to anchor themselves firmly to the rocks by means of strong threads; they feed by filtering water. Characteristic predators of these animals are large snails known as whelks, as well as crabs and starfishes. Several kinds of fish enter the rockweed zone at high tide and feed on the invertebrates.

      Zonation of seaweeds and animals in the intertidal zone results partly from adaptation to a gradient of physical conditions and partly from competitive interactions between the organisms. The upper part of the intertidal zone is exposed to the air for a longer period and thus is at greater risk of drying out, baking, freezing, or being exposed to rainwater. Algal (algae) zonation occurs according to the ability of a species to tolerate these environmental factors, and this in turn influences the type of animal that will inhabit each zone of seaweed. The reverse effect also operates, because by their feeding activity, grazers exclude some seaweeds from zones to which they are otherwise suited.

      At the next level in the food web (that of consumers), predators such as starfish control the abundance of grazing animals. In classic experiments on the coast of Washington state, the ecologist Robert Paine demonstrated that removal of the starfish Pisaster ochraceus from a section of shoreline caused the community to change from one containing 30 species to one totally dominated by the mussel Mytilus californianus. Mussels in this location have the ability to outcompete all other organisms for space on the rocks. Only when the mussel population is controlled by the starfish is a diverse community able to develop. Since these pioneering studies were carried out, many comparable effects have been demonstrated elsewhere. For example, in some places, barnacles are competitive dominants, but their abundance is controlled by limpets and whelks.

Sea-grass beds
      Sea-grass beds are found just below low-tide mark in all latitudes. In north temperate waters Zostera is the most common genus, while in tropical climates Thalassia, known as turtle grass, is an important element. As with marsh grasses, it seems that most of the plant material produced is decomposed by fungi and bacteria while the nutrients are recycled. The sea-grass beds slow the flow of water, causing deposition of silt in which worms and clams may burrow. The plants present a large surface area on which small algae grow, providing a nutritious source of food for browsing animals. The sea-grass beds also shelter many small organisms from their predators, and various species of fish lay their eggs close to sea-grass beds so that the young fish can take advantage of this shelter. Manatees (manatee) and dugongs (dugong), often known as sea cows, are marine mammals that specialize in feeding on sea grasses. This was once a diverse and abundant group, but there are now only three species of manatee (genus Trichecus) and one dugong species (Dugong dugon). The manatees inhabit the eastern and western shores of the Atlantic, while dugongs are found from East Africa to Southeast Asia and Australia. They reach two to three metres in length and feed by ploughing along the bottom, ingesting rhizomes, stems, and leaves of sea grass. Dugongs in northern Australia can occur in herds of 100 to 200 and need very large areas of sea-grass beds to support them. Green turtles (Chelonia midas), which compete with dugongs for sea grass as food, occur throughout the tropics and are much more abundant than dugongs. In the area of the Great Barrier Reef, nesting colonies of green turtles have been observed that contain between 11,000 and 12,000 individuals.

Beaches (beach) and mudflats
      Large areas of coastal habitat have sediments that are too unstable to support communities of large plants. They often have populations of microscopic algae growing at the surface, and they receive particles of decomposing organic matter derived from nearby seaweed or sea-grass beds. A beach near the high-tide level may be so unstable that few animals are able to live in it, but a little farther out to sea the mudflats or sand flats support a rich community of burrowing animals such as polychaete worms, clams, and burrowing shrimps. Many of the worms ingest the sediment and derive nourishment from the organic matter contained in it. Others have tubes that reach to the surface so that they can filter food particles from the water when they are covered by the tide. Clams usually feed in the same way. Crustaceans, starfish, and various kinds of finfish, especially flatfish, move over the mudflats at high tide in search of prey. Mudflats and sand flats are important feeding grounds for wading birds (shorebird) such as sandpipers (sandpiper), oystercatchers (oystercatcher), and plovers (plover). In temperate climates such birds may remain year-round, but many hundreds of thousands of birds make seasonal migrations between high-latitude summer habitats and low-latitude wintering grounds. Large flocks rely on intertidal flats for feeding along the way. For example, it has been shown that about 70,000 semipalmated sandpipers stop on the mudflats of the upper Bay of Fundy, in eastern Canada, in July and August of each year. Feeding predominantly on the burrowing amphipod shrimp Corophium volutator, each bird takes 10,000 to 20,000 shrimps and accumulates 13 to 18 grams (0.46 to 0.63 ounce) of fat, comprising one-third to one-half of the body weight, before taking off on a nonstop journey to the Lesser Antilles or the north coast of South America. At one time there was a plan to build a dam for tidal power that would have permanently flooded these tidal flats, and this would have been a disastrous loss of habitat for these migratory birds.

Kenneth H. Mann William J. Mitsch

Inland systems
Freshwater marshes
      The wetlands in this diverse group are unified primarily by the fact that they are all nontidal freshwater systems dominated by grasses (grass), sedges (Cyperaceae), and other freshwater hydrophytes. However, they differ in their geologic origins and their driving hydrologic forces, and they vary in size from small pothole marshes less than a hectare in size to the immense saw grass monocultures of the Florida Everglades. Vegetation is dominated by graminoids and sedges such as the tall reeds Typha (cattails) and Phragmites, the grasses Panicum and Cladium, the sedges Cyperus and Carex, and floating aquatic plants such as Nymphaea and Nelumbo in temperate regions and Eichhornia crassipes in tropical and subtropical climes. Some inland marshes, such as the prairie glacial marshes of North America, follow a 5- to 20-year cycle of drought. During this period the marsh dries out and exposes large areas of mudflat upon which dense seedling stands germinate. When the rains return, flooding drowns the annual seedlings while allowing the perennials to spread rapidly and vigorously. Deterioration of the marsh follows and is sometimes associated with concentrated muskrat activity. The cycle then repeats.

      The substrate of inland marshes has a higher pH and a greater availability of minerals than does the substrate of bogs. Freshwater marshes are often very productive ecosystems, and most of that productivity is routed through detrital pathways. Herbivory can be important, particularly by muskrats and geese, and consumers can have very significant effects on ecosystem development.

Bogs (bog) and fens (fen)
      Bogs and fens belong to a major class of wetlands called peatlands, moors, or mires, which occur throughout much of the boreal zone of the world. Bogs and fens are distributed in cold temperate climates, mostly in the Northern Hemisphere. There, ample precipitation and high humidity from maritime influences, combined with low evapotranspiration, lead to moisture accumulation. Bogs are acid peat deposits that generally have a high water table (the upper surface of groundwater) but no significant inflow or outflow of streams. Because of their low pH, they support acidophilic (acid-loving) vegetation, particularly mosses. Fens are open wetland systems that generally receive some drainage from surrounding mineral soils and are often covered by grasses, sedges, or reeds. Extensive areas of bogs and fens occur in Finland, eastern Europe, western Siberia, Alaska, Canada (especially Labrador), and the north-central United States. Canada has approximately 1.3 million square kilometres of peatlands, making it the largest resource for peat in the world. In the United States, bogs and fens usually develop in basins scoured out by the Pleistocene glaciers and are clustered primarily around the Great Lakes region and in Maine.

Forested swamps (swamp)
      The term “swamp” usually refers to a wetlands system dominated by trees or other woody vegetation. A wide variety of such systems are found throughout the world. In the tropics vast swamps (also called riparian systems; see below) are found along the great rivers, by which they are often inundated for many months. In temperate regions forested swamps can be dominated by trees that tolerate permanent to semipermanent flooding such as the bald cypress (Taxodium) or swamp tupelo (Nyssa) in the southern United States or the alder (Alnus) or maple (Acer) in more temperate climes.

Riparian systems
      Riparian systems occur along rivers and streams that periodically crest their channel confines, causing flooding. They are also in evidence in places in which a meandering channel creates new sites for plant life to take root and grow. The soils and amount of moisture they contain are influenced by the adjacent stream or river. These systems are distinguished by their linear form and by large fluxes of energy and materials delivered by upstream systems. In arid regions riparian systems can exist along or in ephemeral streams and on the floodplains of perennial streams. In most nonarid regions riparian zones usually develop first along the region of the stream where water flow is constant—i.e., the point at which sufficient groundwater enters the channel to sustain flow through dry periods. Riparian ecosystems exist as broad, alluvial valleys several tens of kilometres wide, as in the Amazon Basin in South America and in Bangladesh, or they can be narrow strips of vegetation along the bank of a stream, as is often seen in the arid western United States. The riparian zone is valuable to animals as a refuge, as an abundant source of water, and as a corridor for migration. This is particularly true in arid regions, where riparian zones may support the only significant vegetation in many kilometres.

Functions and values of wetlands
      Wetland functions are physical, chemical, and biological processes or attributes that are vital to the integrity of the wetland system. Because wetlands are often transition zones (ecotones (ecotone)) between uplands and deepwater aquatic systems, many processes that take place in them have a global impact: they can affect the export of organic materials or serve as a sink for inorganic nutrients. This intermediary position is also responsible for the biodiversity often encountered in these regions, as wetlands “borrow” species from nearby aquatic and terrestrial systems. Wetlands play a major role in the biosphere by providing habitats for a great abundance and richness of floral and faunal species; they are also the last havens for many rare and endangered species. Some wetlands are considered among the Earth's most productive ecosystems. The wetland's function as a site of biodiversity is also valuable to humans, who rely on these ecosystems for commercial and sport fishing, hunting, and recreational uses. The capacity of wetlands to absorb a great amount of water also benefits developed areas. A wetland system can protect shorelines, cleanse polluted waters, prevent floods, and recharge groundwater aquifers, earning wetlands the sobriquet “the kidneys of the landscape.”

Wetland management
      Among ecosystems that are easiest to destroy permanently are these boundary ecosystems. Wetlands can be isolated from their hydrologic source and essentially destroyed if drainage areas are altered or impoundments are built. Major cities in the United States such as Chicago and Washington, D.C., stand on sites that were, in part, wetlands. The amount of wetlands lost worldwide is almost impossible to determine. However, it is known that in the lower 48 states of the United States, a relatively newly developed region of the world, more than half of the original wetlands have been lost, primarily to agricultural development.

      Humans have been utilizing wetlands for centuries. Early civilizations, such as the ancient Babylonians, the Egyptians, and the Aztec, developed unique systems of water delivery that involved wetlands. Among the peoples currently living in proximity to wetlands (known as “wetlanders”) whose culture is linked to these systems are the Camarguais of southern France, the Cajuns of Louisiana, and the Maʿdan, or Marsh Arabs, of southern Iraq; after hundreds of years, all still live in harmony with wetlands. Countless plant and animal products are harvested from wetlands in countries such as China. A thriving modern industry continues to depend on the harvest of cranberries from bogs in the United States. The Russians and the Irish have mined their peatlands for several centuries as a source of energy. Many countries throughout South and Southeast Asia, East Africa, and Central and South America depend on mangrove wetlands for timber, food, and tannin. For centuries salt marshes in northern Europe and the British Isles, and later in New England, have been used to graze animals and raise crops of hay. Thatch roofs and fences have been built from materials retrieved from these areas. Reeds from the wetlands of Romania, Iraq, Japan, and China are used for similar purposes. Techniques to produce fish in systems integrated into rice paddies or shallow ponds were developed several thousand years ago in China and Southeast Asia; crayfish harvesting is still practiced in the wetlands of Louisiana and the Philippines.

      Recognition of the importance of wetlands is growing, with the result that many are being protected by local and national policies (particularly in the United States) as well as by international projects. Examples of these efforts include the Ramsar Convention, which is an international agreement set up for the protection of habitat for migratory waterfowl and other avian life, and the North American Waterfowl Management Plan. Wetland recognition and protection is becoming one of the most important facets of global natural resource protection.

Wetland ecology
      Combining the attributes of both aquatic and terrestrial ecosystems, but falling outside each category, wetlands inhabit a space betwixt and between the disciplines of terrestrial and aquatic ecology. Consequently, their unique properties are not adequately addressed by present ecological paradigms. With their unique characteristics of standing water or waterlogged soils, anoxic conditions, and plant and animal adaptations, wetlands serve as testing grounds for “universal” ecological theories and principles such as succession and energy flow, concepts developed primarily with aquatic or terrestrial ecosystems in mind (see biosphere: The organism and the environment: Resources of the biosphere: The flow of energy (biosphere) and community ecology: Patterns of community structure: Ecological succession (community ecology)). These boundary ecosystems also provide an excellent laboratory for the study of principles related to transition zones, ecological interfaces, and ecotones. In order for wetlands to be protected or restored in the best possible manner, a multidisciplinary approach to their study is required.

William J. Mitsch

Additional Reading
Donald S. McLusky, The Estuarine Ecosystem, 2nd ed. (1989), is a concise account of the subject at the college level. John W. Day, Jr., et al., Estuarine Ecology (1989), deals with physical aspects, plants, animals, organic detritus, and human impacts, including some information on lagoons. K.H. Mann, Ecology of Coastal Waters: A Systems Approach (1982), discusses estuaries as well as sea grass, marsh grass, mangrove, seaweed, and mudflat communities. S.P. Long and C.F. Mason, Saltmarsh Ecology (1983), treats such topics as the formation, flora, fauna, physiography, and conservation of salt marshes. J.r. Lewis, The Ecology of Rocky Shores (1964), is still a standard work on organisms and their relationship to the environment. Roger N. Brehaut, Ecology of Rocky Shores (1982), is a concise, nontechnical text with suggestions for further reading. A.C. Brown and A. McLachlan, Ecology of Sandy Shores (1990), discusses sandy beaches worldwide. John R. Clark, Coastal Ecosystems: Ecological Considerations for Management of the Coastal Zone (1974), and Coastal Ecosystem Management: A Technical Manual for the Conservation of Coastal Zone Resources (1977, reprinted 1983), discuss the ecology of marine boundary ecosystems and the problems of management.Kenneth H. Mann Patrick Dugan (ed.), Wetlands in Danger: A World Conservation Atlas (1993), summarizes all the world's major wetlands and wetland types. William J. Mitsch and James G. Gosselink, Wetlands, 2nd ed. (1993), describes seven major types of wetlands and the principles common to all wetlands. William A. Niering, Wetlands (1985), an illustrated text, details the habitats' features and characteristics. William J. Mitsch (ed.), Global Wetlands: Old World and New (1994), covers topics such as the biogeochemistry, modeling, and ecological engineering of wetlands, as well as wildlife management and river delta management. Regional accounts can be found in Canada Committee on Ecological (Biophysical) Land Classification, National Wetlands Working Group, Wetlands of Canada (1988), including an extensive bibliography; A.J. McComb and P.S. Lake, Australian Wetlands (1990); and Bates Littlehales and William A. Niering, Wetlands of North America (1991), both heavily illustrated with photographs. Edward Maltby, Waterlogged Wealth: Why Waste the World's Wet Places? (1986), describes the functions of wetlands and the degree to which they are threatened around the world. Jon A. Kusler, William J. Mitsch, and Joseph S. Larson, “Wetlands,” Scientific American, 270(1):64–70 (January 1994), summarizes the structure and function of wetlands, emphasizing the importance of a fluctuating water level on ecosystem function. Dennis Whigham, Dagmar Dykyjová, and Slavomil Hejný (eds.), Wetlands of the World: Inventory, Ecology, and Management (1993– ), is a scholarly treatment. Max Finlayson and Michael Moser (eds.), Wetlands (1991), deals with all inland waters of the world from a conservation perspective.William J. Mitsch

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