Introduction to Biomes

A biome is a large, distinctive complex of plant communities created and maintained by climate. How many biomes are there?

A study published in 1999 concluded that there are 150 different “ecoregions” in North America alone. It is rational to lump these into 8 biomes:

• tundra
• taiga
• temperate deciduous forest
• scrub forest (called chaparral in California)
• grassland
• desert
• tropical rain forest
• temperate rain forest

Figure 29.1 shows the distribution of these 8 biomes around the world. A number of climatic factors interact in the creation and maintenance of a biome. Where precipitation is moderately abundant — 40 inches (about 1 m) or more per year — and distributed fairly evenly throughout the year, the major determinant is temperature. It is not simply a matter of average temperature, but includes such limiting factors as whether it ever freezes or length of the growing season. If there is ample rainfall, we find 4 characteristic biomes as we proceed from the tropics (high temperatures) to the extreme latitudes (low temperatures). In order, they are:

• tropical rain forest or jungle
• temperate deciduous forest
• taiga
• tundra

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 29.1).

1. Tropical Wet Forest

Tropical wet forests are also referred to as tropical rainforests. This biome is found in equatorial regions (Figure 29.1). The vegetation is characterized by plants with broad leaves that fall and are replaced 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 more temperate 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 cm 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 typically 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 29.2). 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 American Amazonian rain forest. 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 example, 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.

2. Savannas

Savannas are grasslands with scattered trees, and they are located in Africa, South America, and northern Australia (Figure 29.1). Savannas are usually 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 29.3). Since fire is an important source of disturbance in this biome, plants have evolved well-developed root systems that allow them to quickly resprout after a fire.

3. Subtropical Deserts

Subtropical deserts exist between 15° and 30° north and south latitude and are centered on the Tropics of Cancer and Capricorn (Figure 29.1). 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). This is largely due to the lack of atmospheric water. 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 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, and then 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 29.4). 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.

4. 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 29.1). 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 29.4, is dominated by shrubs 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.

5. 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 29.1). 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 dense enough to 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.

6. Temperate Forests

Temperate forests are the most common biome in eastern North America, Western Europe, Eastern Asia, Chile, and New Zealand (Figure 29.1). 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 periodically during cold winters. 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 29.6). 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.

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 most 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, which does not develop in tropical rainforests. 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).

7. 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 29.1). 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 (coniferous) 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 above-ground biomass of boreal forests is high because these slow-growing tree species are long-lived and accumulate a large 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 29.7). When conifer needles are dropped, they decompose more slowly than broad leaves; therefore, fewer nutrients are returned to the soil to fuel plant growth.

8. Arctic Tundra

The Arctic tundra lies north of the subarctic boreal forest and is located throughout the Arctic regions of the northern hemisphere (Figure 29.1). The average winter temperature is –34 °C (–29.2 °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 29.8). There is little species diversity, low net primary productivity, and low above-ground 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.

Now that we have covered all eight terrestrial biomes in detail, let’s move onto the second major type of biome – aquatic biomes.

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 and the ability to store much larger amounts of energy than the air) 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 29.9). 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; in fact, 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 29.9). 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 action of tidal ebb and flow. 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 may also be repeatedly struck by waves, and the organisms found there are adapted to withstand damage from their pounding action (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.

The neritic zone (Figure 29.9) extends from the intertidal zone to depths of about 200 m (or 650 ft) at the edge of the continental shelf (the underwater landmass that extends from a continent). Since light can penetrate this depth, photosynthesis can still 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 pelagic or open oceanic zone (Figure 29.9). 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, most of the open ocean lacks a process for bringing the organic nutrients back up to the surface. (Exceptions include major oceanic upwellings within the Humboldt Current along the western coast of South America.) 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 deep-water region beyond the continental shelf (Figure 29.9). The bottom of the benthic realm is composed 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 29.9) 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 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, comprising mostly cnidarians and molluscs, 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 perhaps the best-known and largest reef system in the world—visible from the International Space Station! This massive and ancient reef is 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 29.11). 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 called cnidocytes on their tentacles to capture prey.

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, and planktivores, which consume planktonic organisms such as bacteria, archaea, algae, and protists floating in the pelagic zone.

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 ecosystem benefits, which are aquatic resources for drinking water, crop irrigation, sanitation, and industry. Lakes and ponds are connected with abiotic and biotic factors influencing their 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, as we have seen, 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 larvae and adult 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 the determining factors in the amount of phytoplankton growth that takes place 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 29.12) can become so extensive that they reduce light penetration in water. They may also release toxic byproducts into the water, contaminating any drinking water taken from that source. In addition, 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, resulting in a dead zone. Lake Erie and the Gulf of Mexico represent freshwater and marine habitats where phosphorus control and storm water runoff pose significant environmental challenges.

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 usually clear and free of debris. 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 and other organic material that fall downstream into the river or stream, as well as 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, some species of mayfly (phylum Arthropoda) have 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 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 vegetationconsists 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 29.13). The three shared characteristics among these types—what makes them wetlands—are their hydrology, hydrophytic vegetation, and hydric soils.

Freshwater marshes and swamps are characterized by slow and steady water flow. Bogs, however, develop in depressions where water flow is low or nonexistent. Bogs usually occur in areas where there is a clay bottom with poor percolation of water. (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 used during the decomposition of organic matter is not readily replaced. As the oxygen in the water is depleted, decomposition slows. This leads to a buildup of acids and a lower water pH. The lower pH creates challenges for plants because it limits the available nitrogen. As a result, some bog plants (such as sundews, pitcher plants, and Venus flytraps) capture insects in order to 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.

References

Adapted from Clark, M.A., Douglas, M., and Choi, J. (2018). Biology 2e. OpenStax. Retrieved from https://openstax.org/books/biology-2e/pages/1-introduction

Adapted from Kimball, J. W. (2022) Biology. LibreTexts Biology. Retrieved from https://bio.libretexts.org/Bookshelves/Introductory_and_General_Biology/Book%3A_Biology_(Kimball)/17%3A_Ecology/17.01%3A_Energy_Flow_through_the_Biosphere/17.1C%3A_Biomes