Chapter 20 Ecosystems and the Biosphere PDF

542 20 Ecosystems and the Biosphere William and Mary) are now available and have been used in the construction of experimental oyster reefs. Efforts by Virginia and Delaware to clean and restore the bay have been hampered because much of the pollution entering the bay comes fr...

542 20 Ecosystems and the Biosphere William and Mary) are now available and have been used in the construction of experimental oyster reefs. Efforts by Virginia and Delaware to clean and restore the bay have been hampered because much of the pollution entering the bay comes from other states, which emphasizes the need for interstate cooperation to gain successful restoration. The new, hearty oyster strains have also spawned a new and economically viable industry—oyster aquaculture—which not only supplies oysters for food and profit, but also has the added benefit of cleaning the bay. The Sulfur Cycle Sulfur is an essential element for the macromolecules of living things. As part of the amino acid cysteine, it is involved in the formation of proteins. As shown in Figure 20.16, sulfur cycles between the oceans, land, and atmosphere. Atmospheric sulfur is found in the form of sulfur dioxide (SO2), which enters the atmosphere in three ways: first, from the decomposition of organic molecules; second, from volcanic activity and geothermal vents; and, third, from the burning of fossil fuels by humans. FIGURE 20.16 Sulfur dioxide from the atmosphere becomes available to terrestrial and marine ecosystems when it is dissolved in precipitation as weak sulfurous acid or when it falls directly to Earth as fallout. Weathering of rocks also makes sulfates available to terrestrial ecosystems. Decomposition of living organisms returns sulfates to the ocean, soil, and atmosphere. (credit: modification of work by John M. Evans and Howard Perlman, USGS) On land, sulfur is deposited in four major ways: precipitation, direct fallout from the atmosphere, rock weathering, and geothermal vents (Figure 20.17). Atmospheric sulfur is found in the form of sulfur dioxide (SO2), and as rain falls through the atmosphere, sulfur is dissolved in the form of weak sulfurous acid (H2SO3). Sulfur can also fall directly from the atmosphere in a process called fallout. Also, as sulfur-containing rocks weather, sulfur is released into the soil. These rocks originate from ocean sediments that are moved to land by the geologic uplifting of ocean sediments. Terrestrial ecosystems can then make use of these soil sulfates (SO42-), which enter the food web by being taken up by plant roots. When these plants decompose and die, sulfur is released back into the atmosphere as hydrogen sulfide (H2S) gas. Access for free at openstax.org  20.3 Terrestrial Biomes 543 FIGURE 20.17 At this sulfur vent in Lassen Volcanic National Park in northeastern California, the yellowish sulfur deposits are visible near the mouth of the vent. (credit: “Calbear22”/Wikimedia Commons) Sulfur enters the ocean in runoff from land, from atmospheric fallout, and from underwater geothermal vents. Some ecosystems rely on chemoautotrophs using sulfur as a biological energy source. This sulfur then supports marine ecosystems in the form of sulfates. Human activities have played a major role in altering the balance of the global sulfur cycle. The burning of large quantities of fossil fuels, especially from coal, releases larger amounts of hydrogen sulfide gas into the atmosphere. As rain falls through this gas, it creates the phenomenon known as acid rain, which damages the natural environment by lowering the pH of lakes, thus killing many of the resident plants and animals. Acid rain is corrosive rain caused by rainwater falling to the ground through sulfur dioxide gas, turning it into weak sulfuric acid, which causes damage to aquatic ecosystems. Acid rain also affects the man-made environment through the chemical degradation of buildings. For example, many marble monuments, such as the Lincoln Memorial in Washington, DC, have suffered significant damage from acid rain over the years. These examples show the wide-ranging effects of human activities on our environment and the challenges that remain for our future. 20.3 Terrestrial Biomes LEARNING OBJECTIVES By the end of this section, you will be able to: Identify the two major abiotic factors that determine the type of terrestrial biome in an area Recognize distinguishing characteristics of each of the eight major terrestrial biomes Earth’s biomes can be either terrestrial or aquatic. Terrestrial biomes are based on land, while aquatic biomes include both ocean and freshwater biomes. The eight major terrestrial biomes on Earth are each distinguished by characteristic temperatures and amount of precipitation. Annual totals and fluctuations of precipitation affect the kinds of vegetation and animal life that can exist in broad geographical regions. Temperature variation on a daily and seasonal basis is also important for predicting the geographic distribution of a biome. Since a biome is defined by climate, the same biome can occur in geographically distinct areas with similar climates (Figure 20.18). There are also large areas on Antarctica, Greenland, and in mountain ranges that are covered by permanent glaciers and support very little life. Strictly speaking, these are not considered biomes and in addition to extremes of cold, they are also often deserts with very low precipitation. 544 20 Ecosystems and the Biosphere FIGURE 20.18 Each of the world’s eight major biomes is distinguished by characteristic temperatures and amount of precipitation. Polar ice caps and mountains are also shown. Tropical Forest Tropical rainforests are also referred to as tropical wet forests. This biome is found in equatorial regions (Figure 20.18). Tropical rainforests are the most diverse terrestrial biome. This biodiversity is still largely unknown to science and is under extraordinary threat primarily through logging and deforestation for agriculture. Tropical rainforests have also been described as nature’s pharmacy because of the potential for new drugs that is largely hidden in the chemicals produced by the huge diversity of plants, animals, and other organisms. The vegetation is characterized by plants with spreading roots and broad leaves that fall off throughout the year, unlike the trees of deciduous forests that lose their leaves in one season. These forests are “evergreen,” year-round. The temperature and sunlight profiles of tropical rainforests are stable in comparison to that of other terrestrial biomes, with average temperatures ranging from 20oC to 34oC (68oF to 93oF). Month-to-month temperatures are relatively constant in tropical rainforests, in contrast to forests further from the equator. This lack of temperature seasonality leads to year-round plant growth, rather than the seasonal growth seen in other biomes. In contrast to other ecosystems, a more constant daily amount of sunlight (11–12 hours per day) provides more solar radiation, thereby a longer period of time for plant growth. The annual rainfall in tropical rainforests ranges from 250 cm to more than 450 cm (8.2–14.8 ft) with considerable seasonal variation. Tropical rainforests have wet months in which there can be more than 30 cm (11–12 in) of precipitation, as well as dry months in which there are fewer than 10 cm (3.5 in) of rainfall. However, the driest month of a tropical rainforest can still exceed the annual rainfall of some other biomes, such as deserts. Tropical rainforests have high net primary productivity because the annual temperatures and precipitation values support rapid plant growth (Figure 20.19). However, the high rainfall quickly leaches nutrients from the soils of these forests, which are typically low in nutrients. Tropical rainforests are characterized by vertical layering of vegetation and the formation of distinct habitats for animals within each layer. On the forest floor is a sparse layer of plants and decaying plant matter. Above that is an understory of short, shrubby foliage. A layer of trees rises above this understory and is topped by a closed upper canopy—the uppermost overhead layer of branches and leaves. Some additional trees emerge through this closed upper canopy. These layers provide diverse and complex habitats for the variety of plants, animals, and other organisms within the tropical wet forests. Many species of animals use the variety of plants and the complex structure of the tropical wet forests for food and shelter. Some organisms live several meters above ground rarely ever descending to the forest floor. Rainforests are not the only forest biome in the tropics; there are also tropical dry forests, which are characterized by a dry season of varying lengths. These forests commonly experience leaf loss during the dry season to one degree or another. The loss of leaves from taller trees during the dry season opens up the canopy and allows sunlight to the forest floor that allows the growth of thick ground-level brush, which is absent in tropical rainforests. Extensive tropical dry forests occur in Africa (including Madagascar), India, southern Mexico, and South America. Access for free at openstax.org  20.3 Terrestrial Biomes 545 FIGURE 20.19 Species diversity is very high in tropical wet forests, such as these forests of Madre de Dios, Peru, near the Amazon River. (credit: Roosevelt Garcia) Savannas Savannas are grasslands with scattered trees, and they are found in Africa, South America, and northern Australia (Figure 20.18). Savannas are hot, tropical areas with temperatures averaging from 24oC –29oC (75oF –84oF) and an annual rainfall of 51–127 cm (20–50 in). Savannas have an extensive dry season and consequent fires. As a result, scattered in the grasses and forbs (herbaceous flowering plants) that dominate the savanna, there are relatively few trees (Figure 20.20). Since fire is an important source of disturbance in this biome, plants have evolved well- developed root systems that allow them to quickly re-sprout after a fire. FIGURE 20.20 Although savannas are dominated by grasses, small woodlands, such as this one in Mount Archer National Park in Queensland, Australia, may dot the landscape. (credit: "Ethel Aardvark"/Wikimedia Commons) Deserts Subtropical deserts exist between 15o and 30o north and south latitude and are centered on the Tropic of Cancer and the Tropic of Capricorn (Figure 20.18). Deserts are frequently located on the downwind or lee side of mountain ranges, which create a rain shadow after prevailing winds drop their water content on the mountains. This is typical of the North American deserts, such as the Mohave and Sonoran deserts. Deserts in other regions, such as the 546 20 Ecosystems and the Biosphere Sahara Desert in northern Africa or the Namib Desert in southwestern Africa are dry because of the high-pressure, dry air descending at those latitudes. Subtropical deserts are very dry; evaporation typically exceeds precipitation. Subtropical hot deserts can have daytime soil surface temperatures above 60oC (140oF) and nighttime temperatures approaching 0oC (32oF). The temperature drops so far because there is little water vapor in the air to prevent radiative cooling of the land surface. Subtropical deserts are characterized by low annual precipitation of fewer than 30 cm (12 in) with little monthly variation and lack of predictability in rainfall. Some years may receive tiny amounts of rainfall, while others receive more. In some cases, the annual rainfall can be as low as 2 cm (0.8 in) in subtropical deserts located in central Australia (“the Outback”) and northern Africa. The low species diversity of this biome is closely related to its low and unpredictable precipitation. Despite the relatively low diversity, desert species exhibit fascinating adaptations to the harshness of their environment. Very dry deserts lack perennial vegetation that lives from one year to the next; instead, many plants are annuals that grow quickly and reproduce when rainfall does occur, then they die. Perennial plants in deserts are characterized by adaptations that conserve water: deep roots, reduced foliage, and water-storing stems (Figure 20.21). Seed plants in the desert produce seeds that can lie dormant for extended periods between rains. Most animal life in subtropical deserts has adapted to a nocturnal life, spending the hot daytime hours beneath the ground. The Namib Desert is the oldest on the planet, and has probably been dry for more than 55 million years. It supports a number of endemic species (species found only there) because of this great age. For example, the unusual gymnosperm Welwitschia mirabilis is the only extant species of an entire order of plants. There are also five species of reptiles considered endemic to the Namib. In addition to subtropical deserts there are cold deserts that experience freezing temperatures during the winter and any precipitation is in the form of snowfall. The largest of these deserts are the Gobi Desert in northern China and southern Mongolia, the Taklimakan Desert in western China, the Turkestan Desert, and the Great Basin Desert of the United States. FIGURE 20.21 Many desert plants have tiny leaves or no leaves at all to reduce water loss. The leaves of ocotillo, shown here in the Chihuahuan Desert in Big Bend National Park, Texas, appear only after rainfall and then are shed. (credit “bare ocotillo”: "Leaflet"/Wikimedia Commons) Chaparral The chaparral is also called scrub forest and is found in California, along the Mediterranean Sea, and along the southern coast of Australia (Figure 20.18). The annual rainfall in this biome ranges from 65 cm to 75 cm (25.6–29.5 in) and the majority of the rain falls in the winter. Summers are very dry and many chaparral plants are dormant during the summertime. The chaparral vegetation is dominated by shrubs and is adapted to periodic fires, with some plants producing seeds that germinate only after a hot fire. The ashes left behind after a fire are rich in nutrients like nitrogen that fertilize the soil and promote plant regrowth. Fire is a natural part of the maintenance of this biome and frequently threatens human habitation in this biome in the U.S. (Figure 20.22). Access for free at openstax.org  20.3 Terrestrial Biomes 547 FIGURE 20.22 The chaparral is dominated by shrubs. (credit: Miguel Vieira) Temperate Grasslands Temperate grasslands are found throughout central North America, where they are also known as prairies, and in Eurasia, where they are known as steppes (Figure 20.18). Temperate grasslands have pronounced annual fluctuations in temperature with hot summers and cold winters. The annual temperature variation produces specific growing seasons for plants. Plant growth is possible when temperatures are warm enough to sustain plant growth, which occurs in the spring, summer, and fall. Annual precipitation ranges from 25.4 cm to 88.9 cm (10–35 in). Temperate grasslands have few trees except for those found growing along rivers or streams. The dominant vegetation tends to consist of grasses. The treeless condition is maintained by low precipitation, frequent fires, and grazing (Figure 20.23). The vegetation is very dense and the soils are fertile because the subsurface of the soil is packed with the roots and rhizomes (underground stems) of these grasses. The roots and rhizomes act to anchor plants into the ground and replenish the organic material (humus) in the soil when they die and decay. FIGURE 20.23 The American bison (Bison bison), more commonly called the buffalo, is a grazing mammal that once populated American prairies in huge numbers. (credit: Jack Dykinga, USDA ARS) Fires, which are a natural disturbance in temperate grasslands, can be ignited by lightning strikes. It also appears that the lightning-caused fire regime in North American grasslands was enhanced by intentional burning by humans. When fire is suppressed in temperate grasslands, the vegetation eventually converts to scrub and dense forests. 548 20 Ecosystems and the Biosphere Often, the restoration or management of temperate grasslands requires the use of controlled burns to suppress the growth of trees and maintain the grasses. Temperate Forests Temperate forests are the most common biome in eastern North America, Western Europe, Eastern Asia, Chile, and New Zealand (Figure 20.18). This biome is found throughout mid-latitude regions. Temperatures range between –30oC and 30oC (–22oF to 86oF) and drop to below freezing on an annual basis. These temperatures mean that temperate forests have defined growing seasons during the spring, summer, and early fall. Precipitation is relatively constant throughout the year and ranges between 75 cm and 150 cm (29.5–59 in). Deciduous trees are the dominant plant in this biome with fewer evergreen conifers. Deciduous trees lose their leaves each fall and remain leafless in the winter. Thus, little photosynthesis occurs during the dormant winter period. Each spring, new leaves appear as temperature increases. Because of the dormant period, the net primary productivity of temperate forests is less than that of tropical rainforests. In addition, temperate forests show far less diversity of tree species than tropical rainforest biomes. The trees of the temperate forests leaf out and shade much of the ground; however, more sunlight reaches the ground in this biome than in tropical rainforests because trees in temperate forests do not grow as tall as the trees in tropical rainforests. The soils of the temperate forests are rich in inorganic and organic nutrients compared to tropical rainforests. This is because of the thick layer of leaf litter on forest floors and reduced leaching of nutrients by rainfall. As this leaf litter decays, nutrients are returned to the soil. The leaf litter also protects soil from erosion, insulates the ground, and provides habitats for invertebrates and their predators (Figure 20.24). FIGURE 20.24 Deciduous trees are the dominant plant in the temperate forest. (credit: Oliver Herold) Boreal Forests The boreal forest, also known as taiga or coniferous forest, is found roughly between 50o and 60o north latitude across most of Canada, Alaska, Russia, and northern Europe (Figure 20.18). Boreal forests are also found above a certain elevation (and below high elevations where trees cannot grow) in mountain ranges throughout the Northern Hemisphere. This biome has cold, dry winters and short, cool, wet summers. The annual precipitation is from 40 cm to 100 cm (15.7–39 in) and usually takes the form of snow; little evaporation occurs because of the cold temperatures. The long and cold winters in the boreal forest have led to the predominance of cold-tolerant cone-bearing plants. These are evergreen coniferous trees like pines, spruce, and fir, which retain their needle-shaped leaves year-round. Evergreen trees can photosynthesize earlier in the spring than deciduous trees because less energy from the Sun is Access for free at openstax.org  20.3 Terrestrial Biomes 549 required to warm a needle-like leaf than a broad leaf. Evergreen trees grow faster than deciduous trees in the boreal forest. In addition, soils in boreal forest regions tend to be acidic with little available nitrogen. Leaves are a nitrogen- rich structure and deciduous trees must produce a new set of these nitrogen-rich structures each year. Therefore, coniferous trees that retain nitrogen-rich needles in a nitrogen limiting environment may have had a competitive advantage over the broad-leafed deciduous trees. The net primary productivity of boreal forests is lower than that of temperate forests and tropical wet forests. The aboveground biomass of boreal forests is high because these slow-growing tree species are long-lived and accumulate standing biomass over time. Species diversity is less than that seen in temperate forests and tropical rainforests. Boreal forests lack the layered forest structure seen in tropical rainforests or, to a lesser degree, temperate forests. The structure of a boreal forest is often only a tree layer and a ground layer. When conifer needles are dropped, they decompose more slowly than broad leaves; therefore, fewer nutrients are returned to the soil to fuel plant growth (Figure 20.25). FIGURE 20.25 The boreal forest (taiga) has low lying plants and conifer trees. (credit: L.B. Brubaker, NOAA) Arctic Tundra The Arctic tundra lies north of the subarctic boreal forests and is located throughout the Arctic regions of the Northern Hemisphere (Figure 20.18). Tundra also exists at elevations above the tree line on mountains. The average winter temperature is –34°C (–29.2°F) and the average summer temperature is 3°C–12°C (37°F –52°F). Plants in the Arctic tundra have a short growing season of approximately 50–60 days. However, during this time, there are almost 24 hours of daylight and plant growth is rapid. The annual precipitation of the Arctic tundra is low (15–25 cm or 6–10 in) with little annual variation in precipitation. And, as in the boreal forests, there is little evaporation because of the cold temperatures. Plants in the Arctic tundra are generally low to the ground and include low shrubs, grasses, lichens, and small flowering plants (Figure 20.26). There is little species diversity, low net primary productivity, and low aboveground biomass. The soils of the Arctic tundra may remain in a perennially frozen state referred to as permafrost. The permafrost makes it impossible for roots to penetrate far into the soil and slows the decay of organic matter, which inhibits the release of nutrients from organic matter. The melting of the permafrost in the brief summer provides water for a burst of productivity while temperatures and long days permit it. During the growing season, the ground of the Arctic tundra can be completely covered with plants or lichens. 550 20 Ecosystems and the Biosphere FIGURE 20.26 Low-growing plants such as shrub willow dominate the tundra landscape during the summer, shown here in the Arctic National Wildlife Refuge. (credit: Arctic National Wildlife Refuge, USFWS) LINK TO LEARNING Watch this Assignment Discovery: Biomes (http://openstax.org/l/biomes)video for an overview of biomes. To explore further, select one of the biomes on the extended playlist: desert, savanna, temperate forest, temperate grassland, tropic, tundra. 20.4 Aquatic and Marine Biomes LEARNING OBJECTIVES By the end of this section, you will be able to: Describe the effects of abiotic factors on the composition of plant and animal communities in aquatic biomes Compare the characteristics of the ocean zones Summarize the characteristics of standing water and flowing water in freshwater biomes Like terrestrial biomes, aquatic biomes are influenced by abiotic factors. In the case of aquatic biomes the abiotic factors include light, temperature, flow regime, and dissolved solids. The aquatic medium—water— has different physical and chemical properties than air. Even if the water in a pond or other body of water is perfectly clear (there are no suspended particles), water, on its own, absorbs light. As one descends deep enough into a body of water, eventually there will be a depth at which the sunlight cannot reach. While there are some abiotic and biotic factors in a terrestrial ecosystem that shade light (like fog, dust, or insect swarms), these are not usually permanent features of the environment. The importance of light in aquatic biomes is central to the communities of organisms found in both freshwater and marine ecosystems because it controls productivity through photosynthesis. In addition to light, solar radiation warms bodies of water and many exhibit distinct layers of water at differing temperatures. The water temperature affects the organisms’ rates of growth and the amount of dissolved oxygen available for respiration. The movement of water is also important in many aquatic biomes. In rivers, the organisms must obviously be adapted to the constant movement of the water around them, but even in larger bodies of water such as the oceans, regular currents and tides impact availability of nutrients, food resources, and the presence of the water itself. Finally, all natural water contains dissolved solids, or salts. Fresh water contains low levels of such dissolved substances because the water is rapidly recycled through evaporation and precipitation. The oceans have a relatively constant high salt content. Aquatic habitats at the interface of marine and freshwater ecosystems have complex and variable salt environments that range between freshwater and marine levels. These are known as brackish water environments. Lakes located in closed drainage basins concentrate salt in their waters and can have Access for free at openstax.org  20.4 Aquatic and Marine Biomes 551 extremely high salt content that only a few and highly specialized species are able to inhabit. Marine Biomes The ocean is a continuous body of salt water that is relatively uniform in chemical composition. It is a weak solution of mineral salts and decayed biological matter. Within the ocean, coral reefs are a second type of marine biome. Estuaries, coastal areas where salt water and fresh water mix, form a third unique marine biome. The ocean is categorized by several zones (Figure 20.28). All of the ocean’s open water is referred to as the pelagic realm (or zone). The benthic realm (or zone) extends along the ocean bottom from the shoreline to the deepest parts of the ocean floor. From the surface to the bottom or the limit to which photosynthesis occurs is the photic zone (approximately 200 m or 650 ft). At depths greater than 200 m, light cannot penetrate; thus, this is referred to as the aphotic zone. The majority of the ocean is aphotic and lacks sufficient light for photosynthesis. The deepest part of the ocean, the Challenger Deep (in the Mariana Trench, located in the western Pacific Ocean), is about 11,000 m (about 6.8 mi) deep. To give some perspective on the depth of this trench, the ocean is, on average, 4267 m or 14,000 ft deep. Ocean The physical diversity of the ocean has a significant influence on the diversity of organisms that live within it. The ocean is categorized into different zones based on how far light reaches into the water. Each zone has a distinct group of species adapted to the biotic and abiotic conditions particular to that zone. The intertidal zone (Figure 20.28) is the oceanic region that is closest to land. With each tidal cycle, the intertidal zone alternates between being inundated with water and left high and dry. Generally, most people think of this portion of the ocean as a sandy beach. In some cases, the intertidal zone is indeed a sandy beach, but it can also be rocky, muddy, or dense with tangled roots in mangrove forests. The intertidal zone is an extremely variable environment because of tides. Organisms may be exposed to air at low tide and are underwater during high tide. Therefore, living things that thrive in the intertidal zone are often adapted to being dry for long periods of time. The shore of the intertidal zone is also repeatedly struck by waves and the organisms found there are adapted to withstand damage from the pounding action of the waves (Figure 20.27). The exoskeletons of shoreline crustaceans (such as the shore crab, Carcinus maenas) are tough and protect them from desiccation (drying out) and wave damage. Another consequence of the pounding waves is that few algae and plants establish themselves in constantly moving sand or mud. FIGURE 20.27 Sea stars, sea urchins, and mussel shells are often found in the intertidal zone, shown here in Kachemak Bay, Alaska. (credit: NOAA) The neritic zone (Figure 20.28) extends from the margin of the intertidal zone to depths of about 200 m (or 650 ft) at the edge of the continental shelf. When the water is relatively clear, photosynthesis can occur in the neritic zone. 552 20 Ecosystems and the Biosphere The water contains silt and is well-oxygenated, low in pressure, and stable in temperature. These factors all contribute to the neritic zone having the highest productivity and biodiversity of the ocean. Phytoplankton, including photosynthetic bacteria and larger species of algae, are responsible for the bulk of this primary productivity. Zooplankton, protists, small fishes, and shrimp feed on the producers and are the primary food source for most of the world’s fisheries. The majority of these fisheries exist within the neritic zone. Beyond the neritic zone is the open ocean area known as the oceanic zone (Figure 20.28). Within the oceanic zone there is thermal stratification. Abundant phytoplankton and zooplankton support populations of fish and whales. Nutrients are scarce and this is a relatively less productive part of the marine biome. When photosynthetic organisms and the organisms that feed on them die, their bodies fall to the bottom of the ocean where they remain; the open ocean lacks a process for bringing the organic nutrients back up to the surface. Beneath the pelagic zone is the benthic realm, the deepwater region beyond the continental shelf (Figure 20.28). The bottom of the benthic realm is comprised of sand, silt, and dead organisms. Temperature decreases as water depth increases. This is a nutrient-rich portion of the ocean because of the dead organisms that fall from the upper layers of the ocean. Because of this high level of nutrients, a diversity of fungi, sponges, sea anemones, marine worms, sea stars, fishes, and bacteria exists. The deepest part of the ocean is the abyssal zone, which is at depths of 4000 m or greater. The abyssal zone (Figure 20.28) is very cold and has very high pressure, very low or no oxygen content, and high nutrient content as the dead and decomposing material that drifts down from the layers above. There are a variety of invertebrates and fishes found in this zone, but the abyssal zone does not have photosynthetic organisms. Chemosynthetic bacteria use the hydrogen sulfide and other minerals emitted from deep hydrothermal vents. These chemosynthetic bacteria use the hydrogen sulfide as an energy source and serve as the base of the food chain found around the vents. VISUAL CONNECTION FIGURE 20.28 The ocean is divided into different zones based on water depth, distance from the shoreline, and light penetration. In which of the following regions would you expect to find photosynthetic organisms? a. The aphotic zone, the neritic zone, the oceanic zone, and the benthic realm. b. The photic zone, the intertidal zone, the neritic zone, and the oceanic zone. c. The photic zone, the abyssal zone, the neritic zone, and the oceanic zone. d. The pelagic realm, the aphotic zone, the neritic zone, and the oceanic zone. Coral Reefs Coral reefs are ocean ridges formed by marine invertebrates living in warm shallow waters within the photic zone of the ocean. They are found within 30˚ north and south of the equator. The Great Barrier Reef is a well-known reef Access for free at openstax.org  20.4 Aquatic and Marine Biomes 553 system located several miles off the northeastern coast of Australia. Other coral reefs are fringing islands, which are directly adjacent to land, or atolls, which are circular reefs surrounding a former island that is now underwater. The coral-forming colonies of organisms (members of phylum Cnidaria) secrete a calcium carbonate skeleton. These calcium-rich skeletons slowly accumulate, thus forming the underwater reef (Figure 20.29). Corals found in shallower waters (at a depth of approximately 60 m or about 200 ft) have a mutualistic relationship with photosynthetic unicellular protists. The relationship provides corals with the majority of the nutrition and the energy they require. The waters in which these corals live are nutritionally poor and, without this mutualism, it would not be possible for large corals to grow because there are few planktonic organisms for them to feed on. Some corals living in deeper and colder water do not have a mutualistic relationship with protists; these corals must obtain their energy exclusively by feeding on plankton using stinging cells on their tentacles. LINK TO LEARNING In this National Oceanic and Atmospheric Administration (NOAA) video (http://openstax.org/l/coral_organisms), marine ecologist Dr. Peter Etnoyer discusses his research on coral organisms. Coral reefs are one of the most diverse biomes. It is estimated that more than 4000 fish species inhabit coral reefs. These fishes can feed on coral, the cryptofauna (invertebrates found within the calcium carbonate structures of the coral reefs), or the seaweed and algae that are associated with the coral. These species include predators, herbivores, or planktivores. Predators are animal species that hunt and are carnivores or “flesh eaters.” Herbivores eat plant material, and planktivores eat plankton. FIGURE 20.29 Coral reefs are formed by the calcium carbonate skeletons of coral organisms, which are marine invertebrates in the phylum Cnidaria. (credit: Terry Hughes) 554 20 Ecosystems and the Biosphere EVOLUTION CONNECTION Global Decline of Coral Reefs It takes a long time to build a coral reef. The animals that create coral reefs do so over thousands of years, continuing to slowly deposit the calcium carbonate that forms their characteristic ocean homes. Bathed in warm tropical waters, the coral animals and their symbiotic protist partners evolved to survive at the upper limit of ocean water temperature. Together, climate change and human activity pose dual threats to the long-term survival of the world’s coral reefs. The main cause of killing of coral reefs is warmer-than-usual surface water. As global warming raises ocean temperatures, coral reefs are suffering. The excessive warmth causes the coral organisms to expel their endosymbiotic, food-producing protists, resulting in a phenomenon known as bleaching. The colors of corals are a result of the particular protist endosymbiont, and when the protists leave, the corals lose their color and turn white, hence the term “bleaching.” Rising levels of atmospheric carbon dioxide further threaten the corals in other ways; as carbon dioxide dissolves in ocean waters, it lowers pH, thus increasing ocean acidity. As acidity increases, it interferes with the calcification that normally occurs as coral animals build their calcium carbonate homes. When a coral reef begins to die, species diversity plummets as animals lose food and shelter. Coral reefs are also economically important tourist destinations, so the decline of coral reefs poses a serious threat to coastal economies. Human population growth has damaged corals in other ways, too. As human coastal populations increase, the runoff of sediment and agricultural chemicals has increased, causing some of the once-clear tropical waters to become cloudy. At the same time, overfishing of popular fish species has allowed the predator species that eat corals to go unchecked. Although a rise in global temperatures of 1°C–2°C (a conservative scientific projection) in the coming decades may not seem large, it is very significant to this biome. When change occurs rapidly, species can become extinct before evolution leads to newly adapted species. Many scientists believe that global warming, with its rapid (in terms of evolutionary time) and inexorable increases in temperature, is tipping the balance beyond the point at which many of the world’s coral reefs can recover. Estuaries: Where the Ocean Meets Fresh Water Estuaries are biomes that occur where a river, a source of fresh water, meets the ocean. Therefore, both fresh water and salt water are found in the same vicinity; mixing results in a diluted (brackish) salt water. Estuaries form protected areas where many of the offspring of crustaceans, mollusks, and fish begin their lives. Salinity is an important factor that influences the organisms and the adaptations of the organisms found in estuaries. The salinity of estuaries varies and is based on the rate of flow of its freshwater sources. Once or twice a day, high tides bring salt water into the estuary. Low tides occurring at the same frequency reverse the current of salt water (Figure 20.30). Access for free at openstax.org  20.4 Aquatic and Marine Biomes 555 FIGURE 20.30 As estuary is where fresh water and salt water meet, such as the mouth of the Klamath River in California, shown here. (credit: U.S. Army Corps of Engineers) The daily mixing of fresh water and salt water is a physiological challenge for the plants and animals that inhabit estuaries. Many estuarine plant species are halophytes, plants that can tolerate salty conditions. Halophytic plants are adapted to deal with salt water spray and salt water on their roots. In some halophytes, filters in the roots remove the salt from the water that the plant absorbs. Animals, such as mussels and clams (phylum Mollusca), have developed behavioral adaptations that expend a lot of energy to function in this rapidly changing environment. When these animals are exposed to low salinity, they stop feeding, close their shells, and switch from aerobic respiration (in which they use gills) to anaerobic respiration (a process that does not require oxygen). When high tide returns to the estuary, the salinity and oxygen content of the water increases, and these animals open their shells, begin feeding, and return to aerobic respiration. Freshwater Biomes Freshwater biomes include lakes, ponds, and wetlands (standing water) as well as rivers and streams (flowing water). Humans rely on freshwater biomes to provide aquatic resources for drinking water, crop irrigation, sanitation, recreation, and industry. These various roles and human benefits are referred to as ecosystem services. Lakes and ponds are found in terrestrial landscapes and are therefore connected with abiotic and biotic factors influencing these terrestrial biomes. Lakes and Ponds Lakes and ponds can range in area from a few square meters to thousands of square kilometers. Temperature is an important abiotic factor affecting living things found in lakes and ponds. During the summer in temperate regions, thermal stratification of deep lakes occurs when the upper layer of water is warmed by the Sun and does not mix with deeper, cooler water. The process produces a sharp transition between the warm water above and cold water beneath. The two layers do not mix until cooling temperatures and winds break down the stratification and the water in the lake mixes from top to bottom. During the period of stratification, most of the productivity occurs in the warm, well-illuminated, upper layer, while dead organisms slowly rain down into the cold, dark layer below where decomposing bacteria and cold-adapted species such as lake trout exist. Like the ocean, lakes and ponds have a photic layer in which photosynthesis can occur. Phytoplankton (algae and cyanobacteria) are found here and provide the base of the food web of lakes and ponds. Zooplankton, such as rotifers and small crustaceans, consume these phytoplankton. At the bottom of lakes and ponds, bacteria in the aphotic zone break down dead organisms that sink to the bottom. Nitrogen and particularly phosphorus are important limiting nutrients in lakes and ponds. Therefore, they are determining factors in the amount of phytoplankton growth in lakes and ponds. When there is a large input of nitrogen and phosphorus (e.g., from sewage and runoff from fertilized lawns and farms), the growth of algae skyrockets, resulting in a large accumulation of algae called an algal bloom. Algal blooms (Figure 20.31) can become so extensive that they reduce light penetration in water. As a result, the lake or pond becomes aphotic and photosynthetic plants cannot survive. When the algae die and decompose, severe oxygen depletion of the water occurs. Fishes and other organisms that require oxygen are then more likely to die. 556 20 Ecosystems and the Biosphere FIGURE 20.31 The uncontrolled growth of algae in this waterway has resulted in an algal bloom. Rivers and Streams Rivers and the narrower streams that feed into the rivers are continuously moving bodies of water that carry water from the source or headwater to the mouth at a lake or ocean. The largest rivers include the Nile River in Africa, the Amazon River in South America, and the Mississippi River in North America (Figure 20.32). FIGURE 20.32 Rivers range from (a) narrow and shallow to (b) wide and slow moving. (credit a: modification of work by Cory Zanker; credit b: modification of work by David DeHetre) Abiotic features of rivers and streams vary along the length of the river or stream. Streams begin at a point of origin referred to as source water. The source water is usually cold, low in nutrients, and clear. The channel (the width of the river or stream) is narrower here than at any other place along the length of the river or stream. Headwater streams are of necessity at a higher elevation than the mouth of the river and often originate in regions with steep grades leading to higher flow rates than lower elevation stretches of the river. Faster-moving water and the short distance from its origin results in minimal silt levels in headwater streams; therefore, the water is clear. Photosynthesis here is mostly attributed to algae that are growing on rocks; the swift current inhibits the growth of phytoplankton. Photosynthesis may be further reduced by tree cover reaching over the narrow stream. This shading also keeps temperatures lower. An additional input of energy can come from leaves or other organic material that falls into a river or stream from the trees and other plants that border the water. When the leaves decompose, the organic material and nutrients in the leaves are returned to the water. The leaves also support a food chain of invertebrates that eat them and are in turn eaten by predatory invertebrates and fish. Plants and animals have adapted to this fast-moving water. For instance, some species of mayfly (phylum Arthropoda) have Access for free at openstax.org  20.4 Aquatic and Marine Biomes 557 flattened bodies and legs with modified claws to help them cling to the underside of submerged rocks. This body form reduces drag and allows these species to benefit from the high oxygen concentrations in fast-moving currents without being dislodged. Freshwater trout species (phylum Chordata) are an important predator in these fast- moving rivers and streams. As the river or stream flows away from the source, the width of the channel gradually widens, the current slows, and the temperature characteristically increases. The increasing width results from the increased volume of water from more and more tributaries. Gradients are typically lower farther along the river, which accounts for the slowing flow. With increasing volume can come increased silt, and as the flow rate slows, the silt may settle, thus increasing the deposition of sediment. Phytoplankton can also be suspended in slow-moving water. Therefore, the water will not be as clear as it is near the source. The water is also warmer as a result of longer exposure to sunlight and the absence of tree cover over wider expanses between banks. Worms (phylum Annelida) and insects (phylum Arthropoda) can be found burrowing into the mud. Predatory vertebrates (phylum Chordata) include waterfowl, frogs, and fishes. In heavily silt-laden rivers, these predators must find food in the murky waters, and, unlike the trout in the clear waters at the source, these vertebrates cannot use vision as their primary sense to find food. Instead, they are more likely to use taste or chemical cues to find prey. When a river reaches the ocean or a large lake, the water typically slows dramatically and any silt in the river water will settle. Rivers with high silt content discharging into oceans with minimal currents and wave action will build deltas, low-elevation areas of sand and mud, as the silt settles onto the ocean bottom. Rivers with low silt content or in areas where ocean currents or wave action are high create estuarine areas where the fresh water and salt water mix. Wetlands Wetlands are environments in which the soil is either permanently or periodically saturated with water. Wetlands are different from lakes and ponds because wetlands exhibit a near continuous cover of emergent vegetation. Emergent vegetation consists of wetland plants that are rooted in the soil but have portions of leaves, stems, and flowers extending above the water’s surface. There are several types of wetlands including marshes, swamps, bogs, mudflats, and salt marshes (Figure 20.33). FIGURE 20.33 Located in southern Florida, Everglades National Park is vast array of wetland environments, including sawgrass marshes, cypress swamps, and estuarine mangrove forests. Here, a great egret walks among cypress trees. (credit: NPS) Freshwater marshes and swamps are characterized by slow and steady water flow. Bogs develop in depressions where water flow is low or nonexistent. Bogs usually occur in areas where there is a clay bottom with poor percolation. Percolation is the movement of water through the pores in the soil or rocks. The water found in a bog is stagnant and oxygen depleted because the oxygen that is used during the decomposition of organic matter is not replaced. As the oxygen in the water is depleted, decomposition slows. This leads to organic acids and other acids 558 20 Ecosystems and the Biosphere building up and lowering the pH of the water. At a lower pH, nitrogen becomes unavailable to plants. This creates a challenge for plants because nitrogen is an important limiting resource. Some types of bog plants (such as sundews, pitcher plants, and Venus flytraps) capture insects and extract the nitrogen from their bodies. Bogs have low net primary productivity because the water found in bogs has low levels of nitrogen and oxygen. Access for free at openstax.org