Community Ecology

In aquatic ecosystems, the availability of light may be limited because sunlight is absorbed by water, plants, suspended particles, and resident microorganisms. Toward the bottom of a lake, pond, or ocean, there is a zone that light cannot reach. Photosynthesis cannot take place there and, as a result, a number of adaptations have evolved that enable living things to survive without light. For instance, aquatic plants have photosynthetic tissue near the surface of the water; for example, think of the broad, floating leaves of a water lily—water lilies cannot survive without light. In environments such as hydrothermal vents, some bacteria extract energy from inorganic chemicals because there is no light for photosynthesis.

The availability of nutrients in aquatic systems is also an important aspect of energy or photosynthesis. Many organisms sink to the bottom of the ocean when they die in the open water; when this occurs, the energy found in that living organism is sequestered for some time unless ocean upwelling occurs. Ocean upwelling is the rising of deep ocean waters that occurs when prevailing winds blow along surface waters near a coastline (Figure 44.9). As the wind pushes ocean waters offshore, water from the bottom of the ocean moves up to replace this water. As a result, the nutrients once contained in dead organisms become available for reuse by other living organisms.

Figure 44.9 Ocean upwelling is an important process that recycles nutrients and energy in the ocean. As wind (green arrows) pushes offshore, it causes water from the ocean bottom (red arrows) to move to the surface, bringing up nutrients from the ocean depths.

In freshwater systems, the recycling of nutrients occurs in response to air temperature changes. The nutrients at the bottom of lakes are recycled twice each year: in the spring and fall turnover. The spring and fall turnover is a seasonal process that recycles nutrients and oxygen from the bottom of a freshwater ecosystem to the top of a body of water (Figure 44.10). These turnovers are caused by the formation of a thermocline: a layer of water with a temperature that is significantly different from that of the surrounding layers. In wintertime, the surface of lakes found in many northern regions is frozen. However, the water under the ice is slightly warmer, and the water at the bottom of the lake is warmer yet at 4 °C to 5 °C (39.2 °F to 41 °F). Water is densest at 4 °C; therefore, the deepest water is also the densest. The deepest water is oxygen poor because the decomposition of organic material at the bottom of the lake uses up available oxygen that cannot be replaced by means of oxygen diffusion into the water due to the surface ice layer.

In springtime, air temperatures increase and surface ice melts. When the temperature of the surface water begins to reach 4 °C, the water becomes heavier and sinks to the bottom. The water at the bottom of the lake is then displaced by the heavier surface water and, thus, rises to the top. As that water rises to the top, the sediments and nutrients from the lake bottom are brought along with it. During the summer months, the lake water stratifies, or forms layers, with the warmest water at the lake surface.

As air temperatures drop in the fall, the temperature of the lake water cools to 4 °C; therefore, this causes fall turnover as the heavy cold water sinks and displaces the water at the bottom. The oxygen-rich water at the surface of the lake then moves to the bottom of the lake, while the nutrients at the bottom of the lake rise to the surface (Figure 44.10). During the winter, the oxygen at the bottom of the lake is used by decomposers and other organisms requiring oxygen, such as fish.

Temperature

Temperature affects the physiology of living things as well as the density and state of water. Temperature exerts an important influence on living things because few living things can survive at temperatures below 0 °C (32 °F) due to metabolic constraints. It is also rare for living things to survive at temperatures exceeding 45 °C (113 °F); this is a reflection of evolutionary response to typical temperatures. Enzymes are most efficient within a narrow and specific range of temperatures; enzyme degradation can occur at higher temperatures. Therefore, organisms either must maintain an internal temperature or they must inhabit an environment that will keep the body within a temperature range that supports metabolism. Some animals have adapted to enable their bodies to survive significant temperature fluctuations, such as seen in hibernation or reptilian torpor. Similarly, some bacteria are adapted to surviving in extremely hot temperatures such as geysers. Such bacteria are examples of extremophiles: organisms that thrive in extreme environments.

Temperature can limit the distribution of living things. Animals faced with temperature fluctuations may respond with adaptations, such as migration, in order to survive. Migration, the movement from one place to another, is an adaptation found in many animals, including many that inhabit seasonally cold climates. Migration solves problems related to temperature, locating food, and finding a mate. In migration, for instance, the Arctic Tern (Sterna paradisaea) makes a 40,000 km (24,000 mi) round trip flight each year between its feeding grounds in the southern hemisphere and its breeding grounds in the Arctic Ocean. Monarch butterflies (Danaus plexippus) live in the eastern United States in the warmer months and migrate to Mexico and the southern United States in the wintertime. Some species of mammals also make migratory forays. Reindeer (Rangifer tarandus) travel about 5,000 km (3,100 mi) each year to find food. Amphibians and reptiles are more limited in their distribution because they lack migratory ability. Not all animals that can migrate do so: migration carries risk and comes at a high energy cost.

Some animals hibernate or estivate to survive hostile temperatures. Hibernation enables animals to survive cold conditions, and estivation allows animals to survive the hostile conditions of a hot, dry climate. Animals that hibernate or estivate enter a state known as torpor: a condition in which their metabolic rate is significantly lowered. This enables the animal to wait until its environment better supports its survival. Some amphibians, such as the wood frog (Rana sylvatica), have an antifreeze-like chemical in their cells, which retains the cells’ integrity and prevents them from bursting.

Water

Water is required by all living things because it is critical for cellular processes. Since terrestrial organisms lose water to the environment by simple diffusion, they have evolved many adaptations to retain water.

• Plants have a number of interesting features on their leaves, such as leaf hairs and a waxy cuticle, that serve to decrease the rate of water loss via transpiration.
• Freshwater organisms are surrounded by water and are constantly in danger of having water rush into their cells because of osmosis. Many adaptations of organisms living in freshwater environments have evolved to ensure that solute concentrations in their bodies remain within appropriate levels. One such adaptation is the excretion of dilute urine.
• Marine organisms are surrounded by water with a higher solute concentration than the organism and, thus, are in danger of losing water to the environment because of osmosis. These organisms have morphological and physiological adaptations to retain water and release solutes into the environment. For example, Marine iguanas (Amblyrhynchus cristatus), sneeze out water vapor that is high in salt in order to maintain solute concentrations within an acceptable range while swimming in the ocean and eating marine plants.

Inorganic Nutrients and Soil

Inorganic nutrients, such as nitrogen and phosphorus, are important in the distribution and the abundance of living things. Plants obtain these inorganic nutrients from the soil when water moves into the plant through the roots. Therefore, soil structure (particle size of soil components), soil pH, and soil nutrient content play an important role in the distribution of plants. Animals obtain inorganic nutrients from the food they consume. Therefore, animal distributions are related to the distribution of what they eat. In some cases, animals will follow their food resource as it moves through the environment.

Other Aquatic Factors

Some abiotic factors, such as oxygen, are important in aquatic ecosystems as well as terrestrial environments. Terrestrial animals obtain oxygen from the air they breathe. Oxygen availability can be an issue for organisms living at very high elevations, however, where there are fewer molecules of oxygen in the air. In aquatic systems, the concentration of dissolved oxygen is related to water temperature and the speed at which the water moves. Cold water has more dissolved oxygen than warmer water. In addition, salinity, current, and tide can be important abiotic factors in aquatic ecosystems.

Other Terrestrial Factors

Wind can be an important abiotic factor because it influences the rate of evaporation and transpiration. The physical force of wind is also important because it can move soil, water, or other abiotic factors, as well as an ecosystem’s organisms.

Fire is another terrestrial factor that can be an important agent of disturbance in terrestrial ecosystems. Some organisms are adapted to fire and, thus, require the high heat associated with fire to complete a part of their life cycle. For example, the jack pine—a coniferous tree—requires heat from fire for its seed cones to open (Figure 44.11). Through the burning of pine needles, fire adds nitrogen to the soil and limits competition by destroying undergrowth.

Figure 44.11 The mature cones of the jack pine (Pinus banksiana) open only when exposed to high temperatures, such as during a forest fire. A fire is likely to kill most vegetation, so a seedling that germinates after a fire is more likely to receive ample sunlight than one that germinates under normal conditions. (credit: USDA)

Abiotic Factors Influencing Plant Growth

Temperature and moisture are important influences on plant production (primary productivity) and the amount of organic matter available as food (net primary productivity). Net primary productivity is an estimation of all of the organic matter available as food; it is calculated as the total amount of carbon fixed per year minus the amount that is oxidized during cellular respiration. In terrestrial environments, net primary productivity is estimated by measuring the aboveground biomass per unit area, which is the total mass of living plants, excluding roots. This means that a large percentage of plant biomass which exists underground is not included in this measurement. Net primary productivity is an important variable when considering differences in biomes. Very productive biomes have a high level of aboveground biomass.

Annual biomass production is directly related to the abiotic components of the environment. Environments with the greatest amount of biomass have conditions in which photosynthesis, plant growth, and the resulting net primary productivity are optimized. The climate of these areas is warm and wet. Photosynthesis can proceed at a high rate, enzymes can work most efficiently, and stomata can remain open without the risk of excessive transpiration; together, these factors lead to the maximal amount of carbon dioxide (CO2) moving into the plant, resulting in high biomass production. The aboveground biomass produces several important resources for other living things, including habitat and food. Conversely, dry and cold environments have lower photosynthetic rates and therefore less biomass. The animal communities living there will also be affected by the decrease in available food.

44.3 | Terrestrial Biomes

The Earth’s biomes are categorized into two major groups: terrestrial and 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. Comparing the annual totals of precipitation and fluctuations in precipitation from one biome to another provides clues as to the importance of abiotic factors in the distribution of biomes. Temperature variation on a daily and seasonal basis is also important for predicting the geographic distribution of the biome and the vegetation type in the biome. The distribution of these biomes shows that the same biome can occur in geographically distinct areas with similar climates (Figure 44.12).

Tropical Wet Forests

Tropical wet forests are also referred to as tropical rainforests. This biome is found in equatorial regions (Figure 44.12). The vegetation is characterized by plants with broad leaves that fall off throughout the year. Unlike the trees of deciduous forests, the trees in this biome do not have a seasonal loss of leaves associated with variations in temperature and sunlight; these forests are “evergreen” year-round.

The temperature and sunlight profiles of tropical wet forests are very stable in comparison to that of other terrestrial biomes, with the temperatures ranging from 20 °C to 34 °C (68 °F to 93 °F). When one compares the annual temperature variation of tropical wet forests with that of other forest biomes, the lack of seasonal temperature variation in the tropical wet forest becomes apparent. This lack of seasonality leads to year-round plant growth, rather than the seasonal (spring, summer, and fall) growth seen in other biomes. In contrast to other ecosystems, tropical ecosystems do not have long days and short days during the yearly cycle. Instead, a constant daily amount of sunlight (11–12 hrs per day) provides more solar radiation, thereby, a longer period of time for plant growth.

The annual rainfall in tropical wet forests ranges from 125 to 660 cm (50–200 in) with some monthly variation. While sunlight and temperature remain fairly consistent, annual rainfall is highly variable. Tropical wet forests 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 wet forest still exceeds the annual rainfall of some other biomes, such as deserts.

Tropical wet forests have high net primary productivity because the annual temperatures and precipitation values in these areas are ideal for plant growth. Therefore, the extensive biomass present in the tropical wet forest leads to plant communities with very high species diversities (Figure 44.13). Tropical wet forests have more species of trees than any other biome; on average between 100 and 300 species of trees are present in a single hectare (2.5 acres) of South America. One way to visualize this is to compare the distinctive horizontal layers within the tropical wet forest biome. 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, fungi, animals, and other organisms within the tropical wet forests. For instance, epiphytes are plants that grow on other plants, which typically are not harmed. Epiphytes are found throughout tropical wet forest biomes. 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 and have adapted to this arboreal lifestyle.

Figure 44.13 Tropical wet forests, such as these forests of Madre de Dios, Peru, near the Amazon River, have high species diversity. (credit: Roosevelt Garcia)

Savannas

Savannas are grasslands with scattered trees, and they are located in Africa, South America, and northern Australia (Figure 44.12). Savannas are hot, tropical areas with temperatures averaging from 24 °C to 29 °C (75 °F to 84 °F) and an annual rainfall of 10–40 cm (3.9–15.7 in). Savannas have an extensive dry season; for this reason, forest trees do not grow as well as they do in the tropical wet forest (or other forest biomes). As a result, within the grasses and forbs (herbaceous flowering plants) that dominate the savanna, there are relatively few trees (Figure 44.14). 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 44.14 Savannas, like this one in Taita Hills Wildlife Sanctuary in Kenya, are dominated by grasses. (credit: Christopher T. Cooper)

Subtropical Deserts

Subtropical deserts exist between 15 ° and 30 ° north and south latitude and are centered on the Tropics of Cancer and Capricorn (Figure 44.12). This biome is very dry; in some years, evaporation exceeds precipitation. Subtropical hot deserts can have daytime soil surface temperatures above 60 °C (140 °F) and nighttime temperatures approaching 0 °C (32 °F). In cold deserts, temperatures can be as high as 25 °C and can drop below -30 °C (-22 °F). 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. 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 vegetation and low animal diversity of this biome is closely related to this low and unpredictable precipitation. 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. Many other plants in these areas are characterized by having a number of adaptations that conserve water, such as deep roots, reduced foliage, and water-storing stems (Figure 44.15). Seed plants in the desert produce seeds that can be in dormancy for extended periods between rains. Adaptations in desert animals include nocturnal behavior and burrowing.

Figure 44.15 To reduce water loss, many desert plants have tiny leaves or no leaves at all. The leaves of ocotillo (Fouquieria splendens), shown here in the Sonora Desert near Gila Bend, Arizona, appear only after rainfall, and then are shed.

Chaparral

The chaparral is also called the scrub forest and is found in California, along the Mediterranean Sea, and along the southern coast of Australia (Figure 44.12). 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, shown in Figure 44.16, is dominated by shrubs and is adapted to periodic fires, with some plants producing seeds that only germinate 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.

Figure 44.16 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; they are also in Eurasia, where they are known as steppes (Figure 44.12). 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 and when ample water is available, which occurs in the spring, summer, and fall. During much of the winter, temperatures are low, and water, which is stored in the form of ice, is not available for plant growth.

Annual precipitation ranges from 25 cm to 75 cm (9.8–29.5 in). Because of relatively lower annual precipitation in temperate grasslands, there are few trees except for those found growing along rivers or streams. The dominant vegetation tends to consist of grasses and some prairies sustain populations of grazing animals Figure 44.17. 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 44.17 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 Agricultural Research Service)

Fires, mainly caused by lightning, are a natural disturbance in temperate grasslands. When fire is suppressed in temperate grasslands, the vegetation eventually converts to scrub and dense forests. 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 44.12). This biome is found throughout mid-latitude regions. Temperatures range between -30 °C and 30 °C (-22 °F to 86 °F) 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).

Because of the moderate annual rainfall and temperatures, deciduous trees are the dominant plant in this biome (Figure 44.18). Deciduous trees lose their leaves each fall and remain leafless in the winter. Thus, no photosynthesis occurs in the deciduous trees during the dormant winter period. Each spring, new leaves appear as the temperature increases. Because of the dormant period, the net primary productivity of temperate forests is less than that of tropical wet forests. In addition, temperate forests show less diversity of tree species than tropical wet forest biomes.

Figure 44.18 Deciduous trees are the dominant plant in the temperate forest. (credit: Oliver Herold)

The trees of the temperate forests leaf out and shade much of the ground; however, this biome is more open than tropical wet forests because trees in the temperate forests do not grow as tall as the trees in tropical wet forests. The soils of the temperate forests are rich in inorganic and organic nutrients. This is due to the thick layer of leaf litter on forest floors. 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 (such as the pill bug or roly-poly, Armadillidium vulgare) and their predators, such as the red-backed salamander (Plethodon cinereus).

Boreal Forests

The boreal forest, also known as taiga or coniferous forest, is found south of the Arctic Circle and across most of Canada, Alaska, Russia, and northern Europe (Figure 44.12). 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 required to warm a needle-like leaf than a broad leaf. This benefits evergreen trees, which 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 may have 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. Plant species diversity is less than that seen in temperate forests and tropical wet forests. Boreal forests lack the pronounced elements of the layered forest structure seen in tropical wet forests. The structure of a boreal forest is often only a tree layer and a ground layer (Figure 44.19). 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 44.19 The boreal forest (taiga) has low lying plants and conifer trees. (credit: L.B. Brubaker)

Arctic Tundra

The Arctic tundra lies north of the subarctic boreal forest and is located throughout the Arctic regions of the northern hemisphere (Figure 44.12). The average winter temperature is -34 °C (-34 °F) and the average summer temperature is from 3 °C to 12 °C (37 °F–52 °F). Plants in the arctic tundra have a very short growing season of approximately 10–12 weeks. However, during this time, there are almost 24 hours of daylight and plant growth is rapid. The annual precipitation of the Arctic tundra is very low with little annual variation in precipitation. And, as in the boreal forests, there is little evaporation due to the cold temperatures.

Plants in the Arctic tundra are generally low to the ground (Figure 44.20). 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 deep into the soil and slows the decay of organic matter, which inhibits the release of nutrients from organic matter. During the growing season, the ground of the Arctic tundra can be completely covered with plants or lichens.

Figure 44.20 Low-growing plants such as shrub willow dominate the tundra landscape, shown here in the Arctic National Wildlife Refuge. (credit: USFWS Arctic National Wildlife Refuge)

Watch this Assignment Discovery: Biomes video (http://openstaxcollege.org/l/biomes) 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.

44.4 | Aquatic Biomes

Abiotic Factors Influencing Aquatic Biomes

Like terrestrial biomes, aquatic biomes are influenced by a series of abiotic factors. The aquatic medium—water— has different physical and chemical properties than air, however. 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 into a deep body of water, there will eventually be a depth which the sunlight cannot reach. While there are some abiotic and biotic factors in a terrestrial ecosystem that might obscure light (like fog, dust, or insect swarms), usually these are not 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. In freshwater systems, stratification due to differences in density is perhaps the most critical abiotic factor and is related to the energy aspects of light. The thermal properties of water (rates of heating and cooling) are significant to the function of marine systems and have major impacts on global climate and weather patterns. Marine systems are also influenced by large-scale physical water movements, such as currents; these are less important in most freshwater lakes.

The ocean is categorized by several areas or zones (Figure 44.21). 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. Within the pelagic realm is the photic zone, which is the portion of the ocean that light can penetrate (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. These realms and zones are relevant to freshwater lakes as well.

Marine Biomes

The ocean is the largest marine biome. It 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 kind of marine biome. Estuaries, coastal areas where salt water and fresh water mix, form a third unique marine biome.

Ocean

The physical diversity of the ocean is a significant influence on plants, animals, and other organisms. 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, which is the zone between high and low tide, is the oceanic region that is closest to land (Figure 44.21). 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 or muddy. The intertidal zone is an extremely variable environment because of tides. Organisms are exposed to air and sunlight at low tide and are underwater most of the time, especially during high tide. Therefore, living things that thrive in the intertidal zone are 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 44.22). 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 the constantly moving rocks, sand, or mud.

Figure 44.22 Sea urchins, mussel shells, and starfish are often found in the intertidal zone, shown here in Kachemak Bay, Alaska. (credit: NOAA)

The neritic zone (Figure 44.21) extends from the intertidal zone to depths of about 200 m (or 650 ft) at the edge of the continental shelf. Since light can penetrate this depth, photosynthesis can occur in the neritic zone. The water here contains silt and is well-oxygenated, low in pressure, and stable in temperature. Phytoplankton and floating Sargassum (a type of free-floating marine seaweed) provide a habitat for some sea life found in the neritic zone. Zooplankton, protists, small fishes, and shrimp are found in the neritic zone and are the base of the food chain for most of the world’s fisheries.

Beyond the neritic zone is the open ocean area known as the oceanic zone (Figure 44.21). Within the oceanic zone there is thermal stratification where warm and cold waters mix because of ocean currents. Abundant plankton serve as the base of the food chain for larger animals such as whales and dolphins. Nutrients are scarce and this is a relatively less productive part of the marine biome. When photosynthetic organisms and the protists and animals that feed on them die, their bodies fall to the bottom of the ocean where they remain; unlike freshwater lakes, the open ocean lacks a process for bringing the organic nutrients back up to the surface. The majority of organisms in the aphotic zone include sea cucumbers (phylum Echinodermata) and other organisms that survive on the nutrients contained in the dead bodies of organisms in the photic zone.

Beneath the pelagic zone is the benthic realm, the deepwater region beyond the continental shelf (Figure 44.21). The bottom of the benthic realm is comprised of sand, silt, and dead organisms. Temperature decreases, remaining above freezing, 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 exist.

The deepest part of the ocean is the abyssal zone, which is at depths of 4000 m or greater. The abyssal zone (Figure 44.21) is very cold and has very high pressure, high oxygen content, and low nutrient content. There are a variety of invertebrates and fishes found in this zone, but the abyssal zone does not have plants because of the lack of light. Hydrothermal vents are found primarily in the abyssal zone; chemosynthetic bacteria utilize the hydrogen sulfide and other minerals emitted from the vents. These chemosynthetic bacteria use the hydrogen sulfide as an energy source and serve as the base of the food chain found in the abyssal 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 system located several miles off the northeastern coast of Australia. Other coral reef systems are fringing islands, which are directly adjacent to land, or atolls, which are circular reef systems surrounding a former landmass that is now underwater. The coral organisms (members of phylum Cnidaria) are colonies of saltwater polyps that secrete a calcium carbonate skeleton. These calcium-rich skeletons slowly accumulate, forming the underwater reef (Figure 44.23). Corals found in shallower waters (at a depth of approximately 60 m or about 200 ft) have a mutualistic relationship with photosynthetic unicellular algae. 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. Some corals living in deeper and colder water do not have a mutualistic relationship with algae; these corals attain energy and nutrients using stinging cells on their tentacles to capture prey.

Watch this National Oceanic and Atmospheric Administration (NOAA) video (http://openstaxcollege.org/l/marine_biology) to see marine ecologist Dr. Peter Etnoyer discusses his research on coral organisms.

It is estimated that more than 4,000 fish species inhabit coral reefs. These fishes can feed on coral, the cryptofauna (invertebrates found within the calcium carbonate substrate of the coral reefs), or the seaweed and algae that are associated with the coral. In addition, some fish species inhabit the boundaries of a coral reef; 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 44.23 Coral reefs are formed by the calcium carbonate skeletons of coral organisms, which are marine invertebrates in the phylum Cnidaria. (credit: Terry Hughes)

Estuaries: Where the Ocean Meets Fresh Water

Estuaries are biomes that occur where a source of fresh water, such as a river, meets the ocean. Therefore, both fresh water and salt water are found in the same vicinity; mixing results in a diluted (brackish) saltwater. Estuaries form protected areas where many of the young offspring of crustaceans, mollusks, and fish begin their lives. Salinity is a very 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.

The short-term and rapid variation in salinity due to the mixing of fresh water and salt water is a difficult 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 the salinity resulting from saltwater on their roots or from sea spray. In some halophytes, filters in the roots remove the salt from the water that the plant absorbs. Other plants are able to pump oxygen into their roots. 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 and ponds (standing water) as well as rivers and streams (flowing water). They also include wetlands, which will be discussed later. Humans rely on freshwater biomes to provide aquatic resources for drinking water, crop irrigation, sanitation, 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. In the summer, thermal stratification of lakes and ponds occurs when the upper layer of water is warmed by the sun and does not mix with deeper, cooler water. Light can penetrate within the photic zone of the lake or pond. Phytoplankton (algae and cyanobacteria) are found here and carry out photosynthesis, providing 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 phosphorus are important limiting nutrients in lakes and ponds. Because of this, they are determining factors in the amount of phytoplankton growth in lakes and ponds. When there is a large input of nitrogen and phosphorus (from sewage and runoff from fertilized lawns and farms, for example), the growth of algae skyrockets, resulting in a large accumulation of algae called an algal bloom. Algal blooms (Figure 44.24) 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, and resulting dead zones are found across the globe. Lake Erie and the Gulf of Mexico represent freshwater and marine habitats where phosphorus control and storm water runoff pose significant environmental challenges.

Figure 44.24 The uncontrolled growth of algae in this lake has resulted in an algal bloom. (credit: Jeremy Nettleton)

Rivers and Streams

Rivers and streams are continuously moving bodies of water that carry large amounts of water from the source, or headwater, to 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.

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 than at any other place along the length of the river or stream. Because of this, the current is often faster here than at any other point of the river or stream.

The fast-moving water results in minimal silt accumulation at the bottom of the river or stream; 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. An additional input of energy can come from leaves or other organic material that falls into the river or stream from 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. Plants and animals have adapted to this fast-moving water. For instance, leeches (phylum Annelida) have elongated bodies and suckers on both ends. These suckers attach to the substrate, keeping the leech anchored in place. 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 and the current slows. This slow-moving water, caused by the gradient decrease and the volume increase as tributaries unite, has more sedimentation. 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. Worms (phylum Annelida) and insects (phylum Arthropoda) can be found burrowing into the mud. The higher order predator vertebrates (phylum Chordata) include waterfowl, frogs, and fishes. These predators must find food in these slow moving, sometimes murky, waters and, unlike the trout in the waters at the source, these vertebrates may not be able to use vision as their primary sense to find food. Instead, they are more likely to use taste or chemical cues to find prey.

Wetlands

Wetlands are environments in which the soil is either permanently or periodically saturated with water. Wetlands are different from lakes because wetlands are shallow bodies of water whereas lakes vary in depth. 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 44.25). The three shared characteristics among these types—what makes them wetlands—are their hydrology, hydrophytic vegetation, and hydric soils.

Figure 44.25 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 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.

44.5 | Climate and the Effects of Global Climate Change

All biomes are universally affected by global conditions, such as climate, that ultimately shape each biome’s environment. Scientists who study climate have noted a series of marked changes that have gradually become increasingly evident during the last sixty years. Global climate change is the term used to describe altered global weather patterns, including a worldwide increase in temperature, due largely to rising levels of atmospheric carbon dioxide.

Climate and Weather

A common misconception about global climate change is that a specific weather event occurring in a particular region (for example, a very cool week in June in central Indiana) is evidence of global climate change. However, a cold week in June is a weather-related event and not a climate-related one. These misconceptions often arise because of confusion over the terms climate and weather.

Climate refers to the long-term, predictable atmospheric conditions of a specific area. The climate of a biome is characterized by having consistent temperature and annual rainfall ranges. Climate does not address the amount of rain that fell on one particular day in a biome or the colder-than-average temperatures that occurred on one day. In contrast, weather refers to the conditions of the atmosphere during a short period of time. Weather forecasts are usually made for 48-hour cycles. Long-range weather forecasts are available but can be unreliable.

To better understand the difference between climate and weather, imagine that you are planning an outdoor event in northern Wisconsin. You would be thinking about climate when you plan the event in the summer rather than the winter because you have long-term knowledge that any given Saturday in the months of May to August would be a better choice for an outdoor event in Wisconsin than any given Saturday in January. However, you cannot determine the specific day that the event should be held on because it is difficult to accurately predict the weather on a specific day. Climate can be considered “average” weather.

Global Climate Change

Climate change can be understood by approaching three areas of study:

• current and past global climate change
• causes of past and present-day global climate change
• ancient and current results of climate change

It is helpful to keep these three different aspects of climate change clearly separated when consuming media reports about global climate change. It is common for reports and discussions about global climate change to confuse the data showing that Earth’s climate is changing with the factors that drive this climate change.

Evidence for Global Climate Change

Since scientists cannot go back in time to directly measure climatic variables, such as average temperature and precipitation, they must instead indirectly measure temperature. To do this, scientists rely on historical evidence of Earth’s past climate.

Antarctic ice cores are a key example of such evidence. These ice cores are samples of polar ice obtained by means of drills that reach thousands of meters into ice sheets or high mountain glaciers. Viewing the ice cores is like traveling backwards through time; the deeper the sample, the earlier the time period. Trapped within the ice are bubbles of air and other biological evidence that can reveal temperature and carbon dioxide data. Antarctic ice cores have been collected and analyzed to indirectly estimate the temperature of the Earth over the past 400,000 years (Figure 44.26a). The 0 °C on this graph refers to the long-term average. Temperatures that are greater than 0 °C exceed Earth’s long-term average temperature. Conversely, temperatures that are less than 0 °C are less than Earth’s average temperature. This figure shows that there have been periodic cycles of increasing and decreasing temperature.

Before the late 1800s, the Earth has been as much as 9 °C cooler and about 3 °C warmer. Note that the graph in Figure 44.26b shows that the atmospheric concentration of carbon dioxide has also risen and fallen in periodic cycles; note the relationship between carbon dioxide concentration and temperature. Figure 44.26b shows that carbon dioxide levels in the atmosphere have historically cycled between 180 and 300 parts per million (ppm) by volume.

Figure 44.26 Ice at the Russian Vostok station in East Antarctica was laid down over the course 420,000 years and reached a depth of over 3,000 m. By measuring the amount of CO2 trapped in the ice, scientists have determined past atmospheric CO2 concentrations. Temperatures relative to modern day were determined from the amount of deuterium (an isotope of hydrogen) present.

Figure 44.26a does not show the last 2,000 years with enough detail to compare the changes of Earth’s temperature during the last 400,000 years with the temperature change that has occurred in the more recent past. Two significant temperature anomalies, or irregularities, have occurred in the last 2000 years. These are the Medieval Climate Anomaly (or the Medieval Warm Period) and the Little Ice Age. A third temperature anomaly aligns with the Industrial Era. The Medieval Climate Anomaly occurred between 900 and 1300 AD. During this time period, many climate scientists think that slightly warmer weather conditions prevailed in many parts of the world; the higher-than-average temperature changes varied between 0.10 °C and 0.20 °C above the norm. Although 0.10 °C does not seem large enough to produce any noticeable change, it did free seas of ice. Because of this warming, the Vikings were able to colonize Greenland.

The Little Ice Age was a cold period that occurred between 1550 AD and 1850 AD. During this time, a slight cooling of a little less than 1 °C was observed in North America, Europe, and possibly other areas of the Earth. This 1 °C change in global temperature is a seemingly small deviation in temperature (as was observed during the Medieval Climate Anomaly); however, it also resulted in noticeable changes. Historical accounts reveal a time of exceptionally harsh winters with much snow and frost.

The Industrial Revolution, which began around 1750, was characterized by changes in much of human society. Advances in agriculture increased the food supply, which improved the standard of living for people in Europe and the United States. New technologies were invented and provided jobs and cheaper goods. These new technologies were powered using fossil fuels, especially coal. The Industrial Revolution starting in the early nineteenth century ushered in the beginning of the Industrial Era. When a fossil fuel is burned, carbon dioxide is released. With the beginning of the Industrial Era, atmospheric carbon dioxide began to rise (Figure 44.27).

Figure 44.27 The atmospheric concentration of CO₂ has risen steadily since the beginning of industrialization.

Current and Past Drivers of Global Climate Change

Since it is not possible to go back in time to directly observe and measure climate, scientists use indirect evidence to determine the drivers, or factors, that may be responsible for climate change. The indirect evidence includes data collected using ice cores, boreholes (a narrow shaft bored into the ground), tree rings, glacier lengths, pollen remains, and ocean sediments. The data shows a correlation between the timing of temperature changes and drivers of climate change: before the Industrial Era (pre-1780), there were three drivers of climate change that were not related to human activity or atmospheric gases. The first of these is the Milankovitch cycles. The Milankovitch cycles describe the effects of slight changes in the Earth’s orbit on Earth’s climate. The length of the Milankovitch cycles ranges between 19,000 and 100,000 years. In other words, one could expect to see some predictable changes in the Earth’s climate associated with changes in the Earth’s orbit at a minimum of every 19,000 years.

The variation in the sun’s intensity is the second natural factor responsible for climate change. Solar intensity is the amount of solar power or energy the sun emits in a given amount of time. There is a direct relationship between solar intensity and temperature. As solar intensity increases (or decreases), the Earth’s temperature correspondingly increases (or decreases). Changes in solar intensity have been proposed as one of several possible explanations for the Little Ice Age.

Finally, volcanic eruptions are a third natural driver of climate change. Volcanic eruptions can last a few days, but the solids and gases released during an eruption can influence the climate over a period of a few years, causing short-term climate changes. The gases and solids released by volcanic eruptions can include carbon dioxide, water vapor, sulfur dioxide, hydrogen sulfide, hydrogen, and carbon monoxide. Generally, volcanic eruptions cool the climate. This occurred in 1783 when volcanos in Iceland erupted and caused the release of large volumes of sulfuric oxide. This led to haze-effect cooling, a global phenomenon that occurs when dust, ash, or other suspended particles block out sunlight and trigger lower global temperatures as a result; haze-effect cooling usually extends for one or more years. In Europe and North America, haze-effect cooling produced some of the lowest average winter temperatures on record in 1783 and 1784.

Greenhouse gases are probably the most significant drivers of the climate. When heat energy from the sun strikes the Earth, gases known as greenhouse gases trap the heat in the atmosphere, as do the glass panes of a greenhouse keep heat from escaping. The greenhouse gases that affect Earth include carbon dioxide, methane, water vapor, nitrous oxide, and ozone. Approximately half of the radiation from the sun passes through these gases in the atmosphere and strikes the Earth. This radiation is converted into thermal radiation on the Earth’s surface, and then a portion of that energy is re-radiated back into the atmosphere. Greenhouse gases, however, reflect much of the thermal energy back to the Earth’s surface. The more greenhouse gases there are in the atmosphere, the more thermal energy is reflected back to the Earth’s surface. Greenhouse gases absorb and emit radiation and are an important factor in the greenhouse effect: the warming of Earth due to carbon dioxide and other greenhouse gases in the atmosphere.

Evidence supports the relationship between atmospheric concentrations of carbon dioxide and temperature: as carbon dioxide rises, global temperature rises. Since 1950, the concentration of atmospheric carbon dioxide has increased from about 280 ppm to 382 ppm in 2006. In 2011, the atmospheric carbon dioxide concentration was 392 ppm. However, the planet would not be inhabitable by current life forms if water vapor did not produce its drastic greenhouse warming effect.

Scientists look at patterns in data and try to explain differences or deviations from these patterns. The atmospheric carbon dioxide data reveal a historical pattern of carbon dioxide increasing and decreasing, cycling between a low of 180 ppm and a high of 300 ppm. Scientists have concluded that it took around 50,000 years for the atmospheric carbon dioxide level to

increase from its low minimum concentration to its higher maximum concentration. However, starting recently, atmospheric carbon dioxide concentrations have increased beyond the historical maximum of 300 ppm. The current increases in atmospheric carbon dioxide have happened very quickly—in a matter of hundreds of years rather than thousands of years. What is the reason for this difference in the rate of change and the amount of increase in carbon dioxide? A key factor that must be recognized when comparing the historical data and the current data is the presence of modern human society; no other driver of climate change has yielded changes in atmospheric carbon dioxide levels at this rate or to this magnitude.

Human activity releases carbon dioxide and methane, two of the most important greenhouse gases, into the atmosphere in several ways. The primary mechanism that releases carbon dioxide is the burning of fossil fuels, such as gasoline, coal, and natural gas (Figure 44.28). Deforestation, cement manufacture, animal agriculture, the clearing of land, and the burning of forests are other human activities that release carbon dioxide. Methane (CH4) is produced when bacteria break down organic matter under anaerobic conditions. Anaerobic conditions can happen when organic matter is trapped underwater (such as in rice paddies) or in the intestines of herbivores. Methane can also be released from natural gas fields and the decomposition that occurs in landfills. Another source of methane is the melting of clathrates. Clathrates are frozen chunks of ice and methane found at the bottom of the ocean. When water warms, these chunks of ice melt and methane is released. As the ocean’s water temperature increases, the rate at which clathrates melt is increasing, releasing even more methane. This leads to increased levels of methane in the atmosphere, which further accelerates the rate of global warming. This is an example of the positive feedback loop that is leading to the rapid rate of increase of global temperatures.

Figure 44.28 The burning of fossil fuels in industry and by vehicles releases carbon dioxide and other greenhouse gases into the atmosphere. (credit: “Pöllö”/Wikimedia Commons)

Documented Results of Climate Change: Past and Present

Scientists have geological evidence of the consequences of long-ago climate change. Modern-day phenomena such as retreating glaciers and melting polar ice cause a continual rise in sea level. Meanwhile, changes in climate can negatively affect organisms.

Geological Climate Change

Global warming has been associated with at least one planet-wide extinction event during the geological past. The Permian extinction event occurred about 251 million years ago toward the end of the roughly 50-million-year-long geological time span known as the Permian period. This geologic time period was one of the three warmest periods in Earth’s geologic history. Scientists estimate that approximately 70 percent of the terrestrial plant and animal species and 84 percent of marine species became extinct, vanishing forever near the end of the Permian period. Organisms that had adapted to wet and warm climatic conditions, such as annual rainfall of 300–400 cm (118–157 in) and 20 °C–30 °C (68 °F–86 °F) in the tropical wet forest, may not have been able to survive the Permian climate change.

Watch this NASA video (http://openstaxcollege.org/l/climate_plants) to discover the mixed effects of global warming on plant growth. While scientists found that warmer temperatures in the 1980s and 1990s caused an increase in plant productivity, this advantage has since been counteracted by more frequent droughts.

Present Climate Change

A number of global events have occurred that may be attributed to climate change during our lifetimes. Glacier National Park in Montana is undergoing the retreat of many of its glaciers, a phenomenon known as glacier recession. In 1850, the area contained approximately 150 glaciers. By 2010, however, the park contained only about 24 glaciers greater than 25 acres in size. One of these glaciers is the Grinnell Glacier (Figure 44.29) at Mount Gould. Between 1966 and 2005, the size of Grinnell Glacier shrank by 40 percent. Similarly, the mass of the ice sheets in Greenland and the Antarctic is decreasing:

Greenland lost 150–250 km³ of ice per year between 2002 and 2006. In addition, the size and thickness of the Arctic sea ice is decreasing.

Figure 44.29 The effect of global warming can be seen in the continuing retreat of Grinnel Glacier. The mean annual temperature in the park has increased 1.33 °C since 1900. The loss of a glacier results in the loss of summer meltwaters, sharply reducing seasonal water supplies and severely affecting local ecosystems. (credit: modification of work by USGS)

This loss of ice is leading to increases in the global sea level. On average, the sea is rising at a rate of 1.8 mm per year. However, between 1993 and 2010 the rate of sea level increase ranged between 2.9 and 3.4 mm per year. A variety of factors affect the volume of water in the ocean, including the temperature of the water (the density of water is related to its temperature) and the amount of water found in rivers, lakes, glaciers, polar ice caps, and sea ice. As glaciers and polar ice caps melt, there is a significant contribution of liquid water that was previously frozen.

In addition to some abiotic conditions changing in response to climate change, many organisms are also being affected by the changes in temperature. Temperature and precipitation play key roles in determining the geographic distribution and phenology of plants and animals. (Phenology is the study of the effects of climatic conditions on the timing of periodic lifecycle events, such as flowering in plants or migration in birds.) Researchers have shown that 385 plant species in Great Britain are flowering 4.5 days sooner than was recorded earlier during the previous 40 years. In addition, insect-pollinated species were more likely to flower earlier than wind-pollinated species. The impact of changes in flowering date would be mitigated if the insect pollinators emerged earlier. This mismatched timing of plants and pollinators could result in injurious ecosystem effects because, for continued survival, insect-pollinated plants must flower when their pollinators are present.

KEY TERMS

nonliving components of the environment

total mass of aboveground living plants per area

deepest part of the ocean at depths of 4000 m or greater

rapid increase of algae in an aquatic system

part of the ocean where no light penetrates

(also, benthic zone) part of the ocean that extends along the ocean bottom from the shoreline to the deepest

parts of the ocean floor

study of the geographic distribution of living things and the abiotic factors that affect their distribution

ecological community of plants, animals, and other organisms that is adapted to a characteristic set of

environmental conditions

living components of the environment

branches and foliage of trees that form a layer of overhead coverage in a forest

width of a river or stream from one bank to the other bank

frozen chunks of ice and methane found at the bottom of the ocean

long-term, predictable atmospheric conditions present in a specific area

individuals that are members of the same species

ocean ridges formed by marine invertebrates living in warm, shallow waters within the photic zone

invertebrates found within the calcium carbonate substrate of coral reefs

study of interaction between living things and their environment

human benefits and services provided by natural ecosystems

wetland plants that are rooted in the soil but have portions of leaves, stems, and flowers extending

above the water’s surface

species found only in a specific geographic area that is usually restricted in size

biomes where a source of fresh water, such as a river, meets the ocean

seasonal process that recycles nutrients and oxygen from the bottom of a freshwater ecosystem

to the top

altered global weather patterns, including a worldwide increase in temperature, due largely to

rising levels of atmospheric carbon dioxide

warming of Earth due to carbon dioxide and other greenhouse gases in the atmosphere

atmospheric gases such as carbon dioxide and methane that absorb and emit radiation, thus trapping

heat in Earth’s atmosphere

effect of the gases and solids from a volcanic eruption on global climate

individuals that are members of different species

part of the ocean that is closest to land; parts extend above the water at low tide

Chapter 44 | Ecology and the Biosphere 1313

Milankovitch cycles

neritic zone

net primary productivity

ocean upwelling

oceanic zone

pelagic realm

permafrost

photic zone

planktivore

predator

Sargassum

solar intensity

source water

thermocline

weather

cyclic changes in the Earth’s orbit that may affect climate

part of the ocean that extends from low tide to the edge of the continental shelf

measurement of the energy accumulation within an ecosystem, calculated as the total amount

of carbon fixed per year minus the amount that is oxidized during cellular respiration

rising of deep ocean waters that occurs when prevailing winds blow along surface waters near a

coastline

part of the ocean that begins offshore where the water measures 200 m deep or deeper

(also, pelagic zone) open ocean waters that are not close to the bottom or near the shore

perennially frozen portion of the Arctic tundra soil

portion of the ocean that light can penetrate

animal species that eats plankton

animal species that hunt and are carnivores or “flesh eaters”

type of free-floating marine seaweed

amount of solar power energy the sun emits in a given amount of time

point of origin of a river or stream

layer of water with a temperature that is significantly different from that of the surrounding layers

conditions of the atmosphere during a short period of time

CHAPTER SUMMARY

44.1 The Scope of Ecology

Ecology is the study of the interactions of living things with their environment. Ecologists ask questions across four levels

of biological organization—organismal, population, community, and ecosystem. At the organismal level, ecologists study

individual organisms and how they interact with their environments. At the population and community levels, ecologists

explore, respectively, how a population of organisms changes over time and the ways in which that population interacts

with other species in the community. Ecologists studying an ecosystem examine the living species (the biotic components)

of the ecosystem as well as the nonliving portions (the abiotic components), such as air, water, and soil, of the

environment.

44.2 Biogeography

Biogeography is the study of the geographic distribution of living things and the abiotic factors that affect their

distribution. Endemic species are species that are naturally found only in a specific geographic area. The distribution of

living things is influenced by several environmental factors that are, in part, controlled by the latitude or elevation at which

an organism is found. Ocean upwelling and spring and fall turnovers are important processes regulating the distribution of

nutrients and other abiotic factors important in aquatic ecosystems. Energy sources, temperature, water, inorganic

nutrients, and soil are factors limiting the distribution of living things in terrestrial systems. Net primary productivity is a

measure of the amount of biomass produced by a biome.

44.3 Terrestrial Biomes

The Earth has terrestrial biomes and aquatic biomes. Aquatic biomes include both freshwater and marine environments.

There are eight major terrestrial biomes: tropical wet forests, savannas, subtropical deserts, chaparral, temperate

grasslands, temperate forests, boreal forests, and Arctic tundra. The same biome can occur in different geographic

locations with similar climates. Temperature and precipitation, and variations in both, are key abiotic factors that shape the

composition of animal and plant communities in terrestrial biomes. Some biomes, such as temperate grasslands and

temperate forests, have distinct seasons, with cold weather and hot weather alternating throughout the year. In warm, moist

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This OpenStax book is available for free at http://cnx.org/content/col11448/1.10

biomes, such as the tropical wet forest, net primary productivity is high, as warm temperatures, abundant water, and a

year-round growing season fuel plant growth. Other biomes, such as deserts and tundra, have low primary productivity due

to extreme temperatures and a shortage of available water.

44.4 Aquatic Biomes

Aquatic ecosystems include both saltwater and freshwater biomes. The abiotic factors important for the structuring of

aquatic ecosystems can be different than those seen in terrestrial systems. Sunlight is a driving force behind the structure

of forests and also is an important factor in bodies of water, especially those that are very deep, because of the role of

photosynthesis in sustaining certain organisms. Density and temperature shape the structure of aquatic systems. Oceans

may be thought of as consisting of different zones based on water depth and distance from the shoreline and light

penetrance. Different kinds of organisms are adapted to the conditions found in each zone. Coral reefs are unique marine

ecosystems that are home to a wide variety of species. Estuaries are found where rivers meet the ocean; their shallow

waters provide nourishment and shelter for young crustaceans, mollusks, fishes, and many other species. Freshwater

biomes include lakes, ponds, rivers, streams, and wetlands. Bogs are an interesting type of wetland characterized by

standing water, lower pH, and a lack of nitrogen.

44.5 Climate and the Effects of Global Climate Change

The Earth has gone through periodic cycles of increases and decreases in temperature. During the past 2000 years, the

Medieval Climate Anomaly was a warmer period, while the Little Ice Age was unusually cool. Both of these irregularities

can be explained by natural causes of changes in climate, and, although the temperature changes were small, they had

significant effects. Natural drivers of climate change include Milankovitch cycles, changes in solar activity, and volcanic

eruptions. None of these factors, however, leads to rapid increases in global temperature or sustained increases in carbon

dioxide. The burning of fossil fuels is an important source of greenhouse gases, which plays a major role in the greenhouse

effect. Long ago, global warming resulted in the Permian extinction: a large-scale extinction event that is documented in

the fossil record. Currently, modern-day climate change is associated with the increased melting of glaciers and polar ice

sheets, resulting in a gradual increase in sea level. Plants and animals can also be affected by global climate change when

the timing of seasonal events, such as flowering or pollination, is affected by global warming.

ART CONNECTION QUESTIONS

1. Figure 44.10 How might turnover in tropical lakes

differ from turnover in lakes that exist in temperate

regions?

2. Figure 44.12 Which of the following statements about

biomes is false?

a. Chaparral is dominated by shrubs.

b. Savannas and temperate grasslands are

dominated by grasses.

c. Boreal forests are dominated by deciduous trees.

d. Lichens are common in the arctic tundra.

3. Figure 44.21 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

REVIEW QUESTIONS

4. Which of the following is a biotic factor?

a. wind

b. disease-causing microbe

c. temperature

d. soil particle size

5. The study of nutrient cycling though the environment is

an example of which of the following?

a. organismal ecology

b. population ecology

c. community ecology

d. ecosystem ecology

6. Understory plants in a temperate forest have adaptations

to capture limited ________.

a. water

b. nutrients

c. heat

d. sunlight

7. An ecologist hiking up a mountain may notice different

biomes along the way due to changes in all of the

following except:

a. elevation

b. rainfall

c. latitude

d. temperature

Chapter 44 | Ecology and the Biosphere 1315

8. Which of the following biomes is characterized by

abundant water resources?

a. deserts

b. boreal forests

c. savannas

d. tropical wet forests

9. Which of the following biomes is characterized by short

growing seasons?

a. deserts

b. tropical wet forests

c. Arctic tundras

d. savannas

10. Where would you expect to find the most

photosynthesis in an ocean biome?

a. aphotic zone

b. abyssal zone

c. benthic realm

d. intertidal zone

11. A key feature of estuaries is:

a. low light conditions and high productivity

b. salt water and fresh water

c. frequent algal blooms

d. little or no vegetation

12. Which of the following is an example of a weather

event?

a. The hurricane season lasts from June 1 through

November 30.

b. The amount of atmospheric CO2 has steadily

increased during the last century.

c. A windstorm blew down trees in the Boundary

Waters Canoe Area in Minnesota on July 4,

1999.

d. Deserts are generally dry ecosystems having

very little rainfall.

13. Which of the following natural forces is responsible

for the release of carbon dioxide and other atmospheric

gases?

a. the Milankovitch cycles

b. volcanoes

c. solar intensity

d. burning of fossil fuels

CRITICAL THINKING QUESTIONS

14. Ecologists often collaborate with other researchers

interested in ecological questions. Describe the levels of

ecology that would be easier for collaboration because of

the similarities of questions asked. What levels of ecology

might be more difficult for collaboration?

15. The population is an important unit in ecology as well

as other biological sciences. How is a population defined,

and what are the strengths and weaknesses of this

definition? Are there some species that at certain times or

places are not in populations?

16. Compare and contrast ocean upwelling and spring and

fall turnovers.

17. Many endemic species are found in areas that are

geographically isolated. Suggest a plausible scientific

explanation for why this is so.

18. The extremely low precipitation of subtropical desert

biomes might lead one to expect fire to be a major

disturbance factor; however, fire is more common in the

temperate grassland biome than in the subtropic desert

biome. Why is this?

19. In what ways are the subtropical desert and the arctic

tundra similar?

20. Scientists have discovered the bodies of humans and

other living things buried in bogs for hundreds of years,

but not yet decomposed. Suggest a possible biological

explanation for why such bodies are so well-preserved.

21. Describe the conditions and challenges facing

organisms living in the intertidal zone.

22. Compare and contrast how natural- and humaninduced

processes have influenced global climate change.

23. Predict possible consequences if carbon emissions

from fossil fuels continue to rise.

Adapted from:

OpenStax, Biology. OpenStax. May 20, 2013. <http://cnx.org/content/col11448/latest/>

“Download for free at http://cnx.org/content/col11448/latest/.”