MARSH

type of wetland ecosystem characterized by poorly drained mineral soils and by plant life dominated by grasses. The latter characteristic distinguishes a marsh from a swamp (q.v.), whose plant life is dominated by trees. Marshes are common at the mouths of rivers, especially where extensive deltas have been built. The river brings a steady supply of water. The gradient of the river approaches zero at the sea, where flow is sluggish. Because the delta is deposited by sediment settling from the river water, the land that is built will be poorly drained at its driest and will often be underwater. Sediment supplied by the river has often been eroded from the surface soils of the drainage basin and is thus very rich. The combination of water supplied steadily at a low rate over a waterlogged but rich soil creates a perfect environment for marsh grasses. Fibrous-rooted grasses bind the muds together and further hinder water flow, thus encouraging the spread of both the delta and the marsh. Marshes occur in the deltas of most of the world's great rivers. In Europe well-known river-mouth marshes include those of the Camargue in the Rhne Delta, the Guadalquivir in Spain, and the Danube in Romania, all of which are famous as bird sanctuaries. In the Middle East, both the Nile Delta and the delta of the TigrisEuphrates have extensive marshes of historical importance. The marsh dwellers of the Iranian marshes have developed a unique culture adapted to life in the wetlands. Marshes occur in the deltas of the Mekong in Vietnam and the Amazon in Brazil. In the United States, the most extensive delta marshes are those of the Mississippi River. Some low-lying areas with poor drainage at the heads of more extensive drainage patterns contain wetlands. A well-known example is the Pripet Marshes and fens that historically have served as the natural boundary between Poland and Russia. In some places basin-like depressions in the Earth's surface trap waters and make wetlands. Most such areas are drained someplace along their rim by a river that is impeded at that point sufficiently to dam water at times of high flow and create marshes and swamps. The world's two largest rivers, the Amazon and the Congo, fall into this category. Both of the great basins named after these rivers have extensive wetlands. The papyrus marshes of the upper Nile in southern Sudan lie above dams of resistant rocks of the cataracts. The Okavango Marshes east of the Kalahari desert in Botswana are perhaps the best example of marshes formed in an interior, closed basin that has no drainage. Other basins without outlets like that of the Great Salt Lake in Utah have accumulated too much salt for marsh growth. The Florida Everglades constitutes a unique marshswamp combination growing on a limestone base. Because the region is near sea level the water from the abundant rains does not drain but remains on the surface. The Everglades is similar to a huge, shallow, slowly flowing river. The area is an ideal marsh habitat but the Everglades is different from usual marshes. The soils are alkaline because of the limy base, and the water is clear. Some areas, such as the northern Great Plains of the United States, have so many small marshes that they are a characteristic of the landscape. These small marshes formed because the landscape left by the retreat of glacial ice was so irregular and so poorly drained that countless little depressions were filled with water each spring. As snow melted, the depressions supported the growth of temporary marshes, which then dried up during the summer. Larger depressions were occupied by ponds. These gradually became marshy as they filled in with sediment. The number of plant species in marshes is few compared to the numbers that grow on well-watered but not waterlogged land. Grasses, grasslike sedges, and reeds or rushes are of major importance. Wild rice is of some commercial importance, but true rice is undoubtedly by far the most important marsh plant and supplies a major portion of the world's grain. Salt marshes, which are extensive along the east coast of the United States and are also common in the Arctic, northern Europe, Australia, and New Zealand, are formed by seawater flooding and draining, which exposes flat areas of intertidal land. Salt-marsh grasses will not grow on permanently flooded flats; growth is also prevented where the flooded land is subject to strong currents and is therefore unstable. Animals have adapted to the limited supplies of oxygen in salt-marsh water in various ways. Rat-tailed maggots (Tubifera), for example, survive in shallow marshes by means of a telescoping, tail-breathing tube that they extend to the water surface for air. Some larvae of shore flies (Ephydridae) and some nematodes take advantage of the air spaces in plants and obtain oxygen from that source. Many small marsh animals have great resistance to lack of oxygen; for example, many nematodes can live indefinitely in the complete absence of oxygen. This ability is essential for such minute animals that would otherwise be limited in distribution to a thin layer a fraction of an inch deep at the mud surface. Salt-marsh animals living at or in the ground are largely derived from marine ancestors and have a problem in resisting fresh water from rains rather than salt. Some, such as worms, merely hide in the mud until the freshwater has run off the marsh surface. Others, such as fiddler crabs, have developed the ability to control their osmotic concentration in freshwater for periods of up to several days. Insects are the principal land animals found on marshes. Although they can withstand short periods of saltwater immersion, they often avoid saltwater by moving up the plants or flying away. Salt marshes are among the most productive natural systems. Productivities of more than 3,000 grams per square metre per year have been reported for the most productive parts of salt marshes, the tall Spartina alterniflora stands growing along tidal creeks. These values correspond to nearly 30 tons per acre per year and are equal to the highest values that have been achieved in agriculture.