Global Ecology

CHAPTER OUTLINE

Chapter

32 3

Life on the Land Rainforests Savannas and Deciduous Tropical Forests Deserts Grasslands Temperate Deciduous Forests Temperate Mixed and Coniferous Forests Mediterranean Scrub The Northernmost ForestsTaiga and Boreal Forest Arctic Tundra

magine driving across the United States from New York City to Los Angeles. As an astute botanist you are able to separate the patches of natural vegetation you see from agricultural elds, heavily grazed pastures, and frequently mowed roadsides. What you observe is a gradual change in this natural vegetation. Leaving New York City, you rst encounter deciduous forest, then pass through grasslands where trees are mostly conned to the margins of rivers, and eventually you enter the deserts of the arid southwest with much bare ground, scattered shrubs, and cacti. Along the way, you also go up and down mountains, perhaps driving from desert up into forests and grasslands, and then back down to desert. Crossing the coastal mountains into southern California takes you into landscapes dominated by dense shrubs, most of them species that you have not seen any-

where else. Such a trip makes it clear that the natural ecosystems of the eastern United States are dramatically different from those of the west. You would also observe that when the topography is reasonably uniform, you tend to be in one kind of vegetation for very long stretches. The major regional ecosystems you encounter on your trip are called biomes. Biomes are the major subunits of the biosphere and are characterized by a more or less homogeneous type of plant cover. The plants and animals that occur in particular biomes have characteristic growth forms and other adaptations that have evolved in relation to specic climates. It is because of these common growth forms that we can recognize biomes, such as grasslands or deserts, wherever they occur, even though the individual species of a biome in one region are often completely different from those of the same biome in another region, having achieved similar characteristics as a result of parallel evolution. Biomes are shaped by climateprimarily temperature and precipitationand so similar biomes are found worldwide in regions of similar climate (Figure 321). Temperate deciduous forest, for example, is found in North America, Europe, and Asia, and tropical rainforest is found in Central and South America, Africa, Asia, and northernmost Australia. Although it is useful to identify the major biomes, it is also important to keep in mind that climatic, geologic, and other factors often change gradually over space and time. Thus, on your cross-country drive, you would have noticed that biomes do not shift abruptly from one to the next. As with other classication systems, opinions differ about the best way to

(a)

(b)

321 Mediterranean-type vegetation Regions with Mediterranean-type climates are characterized by hot, dry summers and mild winters, with strongly seasonal rainfall that occurs primarily in the cooler part of the year. The vegetation is dominated by dense, low, mostly evergreen or summer-deciduous shrubs. There is a striking

supercial similarity in the vegetation, whether in California, Chile, Australia, South Africa, or the Mediterranean region, as seen here for (a) chapparal in southern California and for (b) similar vegetation, called maquis, on the Greek island of Corfu. Each region, however, has its unique set of species. 321

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CHECKPOINTS
By the time you nish reading this chapter, you should be able to answer the following questions: 1. What is a biome, and what factors affect the distribution of biomes on Earth? Given that tropical rainforests contain such a diversity of species, why are tropical soils unsuitable for agriculture? How is an African savanna different from a North American grassland? What role does re play in grasslands? Which best characterizes a deserthigh temperature or low precipitation? How have plants adapted to desert living? How does the appearance of the temperate deciduous forest biome change from one season to the next? What accounts for these changes?

2.

3.

4.

divide up the biosphere into biomes, and many scientists argue that it is better not to assume that discrete units exist, but rather to consider how ecosystems change as one moves across the gradients in climate, soils, geology, and topography. We will adopt a middle approach, using the generally recognized biomes as reference points along gradients. But in reality, change is often so gradual that it is difcult to know on the ground when one leaves one biome and enters another. In our discussion, we will emphasize vegetation, but it must not be forgotten that heterotrophs also play essential roles in the biomes.

5.

Life on the Land

The scheme we present for classifying the natural vegetation of the world is based on the work of the late A. W. Kchler, of the University of Kansas (Figure 322). Study of this map

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322 Biomes of the world The information in these maps and the accompanying key was originally supplied by A.W. Kchler of the University of Kansas, one of the leading authorities on the distribution of biomes. Because of the global coverage of the maps, the scale is relatively small and the content is generalized. Any given biome is not always uniform, and all of the biomes include considerable variations in vegetation.The boundaries between the biomes may be sharp, but more often they are blurred, consisting of broad zones of transition from one type of vegetation to another.

Life on the Land

1 2 3 4 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Temperate deciduous forests Temperate mixed forests Subtropical mixed forests Taiga Northwestern coniferous forest Alpine tundra and mountain forests Mixed west-coast forests Arctic tundra Ice desert Grasslands Savannas Mediterranean scrub Deserts and semideserts Juniper savanna Southern woodland and scrub Tropical mixed forests Monsoon forests Rainforests

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reveals some broad patterns, but also complexities. In the Northern Hemisphere, the sequence from north to south, from ice desert to treeless Arctic tundra, to northern coniferous forest (taiga), and to temperate forest or grassland is evident, with each type occupying a vast area. This sequence is not apparent in the Southern Hemisphere because of the small land mass in the subarctic and temperate zones. We see also that large areas of the Earth are occupied by arid or semiarid vegetationdesert and savannain both the tropical and temperate zones. Within the tropics (the zone between the latitudes of 23.5 north and 23.5 south), one nds the quintessential tropical ecosystemtropical rainforestas well as areas of seasonally dry forest types, such as monsoon forest and alpine mountain forest. These global patterns are largely explained by the latitudinal gradients in temperature and the distribution of precipitation inuenced by the latitudinal movement of air masses (Figure 323). Moisture and temperature are not the only important factors, but they strongly restrict what can grow in a particular area. We do not nd rainforests without rain, and desert plants that tolerate drought and intense sunlight cannot survive along the fog-bound coasts of Ireland. The tilt of the Earths axis relative to the plane in which it moves around the sun determines the amount and seasonal variation of solar energy that reaches the land surface. The higher latitudes, both north and south, have strongly seasonal climates with short days in winter and long days in summer. In the regions closer to the North or South Poles, the input of solar energy dips so low that temperatures fall below freezing for extended periods. Most plants can tolerate temperatures just above freezing, and some can adjust to tolerate

freezing temperatures, but only plants with special structural and physiological adaptations, such as the timing of growth and owering, can survive prolonged subzero temperatures. Evaporation by solar energy is the means by which water enters the atmosphere to be returned to the surface as precipitation. The worldwide distribution of precipitation, however, is extremely variable. Regions differ markedly both in total amounts of precipitation received and in the seasonal distribution of precipitation, with profound effects on ecosystems. Because the oceans are the primary source of evaporated water, the central regions of the continents tend to be dry, and in some cases extremely dry. Precipitation is also very low in the latitudinal band between 15 and 30 in both hemispheres (Figure 323). High-pressure zones dominate this zone and the air is generally stable, with many cloudless days and relative calm, explaining a traditional name for this zonethe doldrums. Within the tropics, some areas have high precipitation with little seasonal variation. Intense solar radiation causes high evaporation, a buildup of cloud masses, and nearly daily rainfall. The regional movement of air masses ensures a constant supply of moisture to maintain the cycle. In other tropical regions, however, rainfall is strongly seasonal with alternating periods of prolonged drought and intense rainfallbest typied by the monsoon climate.

Both Elevation and Latitude Determine Ecosystem Properties

In general, air temperature decreases with increasing height above the ground, a consequence of the fact that it is pri-

Green arrows: Moist air rises and cools high precipitation 60 N Westerlies 30 N

Northeast trade winds 0

Equatorial doldrums

Direction of Earths rotation Brown arrows: Dry air descends and warms low precipitation

Southeast trade winds 30 S Westerlies 60 S

323 Global air currents The Earths surface is covered by belts of air currents, which determine the major patterns of wind and rainfall. In this diagram, the blue arrows indicate the direction of movement of the air within the belts. The green arrows indicate regions of rising air, which are characterized by high precipitation, and the brown arrows indicate regions of descending air, which are characterized by low precipitation. The dry air descending at latitudes of 30 north and south is responsible for the great deserts of the world. The prevailing winds on the Earths surface, indicated by the black arrows, are produced by the twisting effect of the Earths rotation on the air currents within the individual belts.

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marily the absorption of short-wave energy at the Earths surface that captures the heat energy of the sun. The sunwarmed surface radiates long-wave radiation back into the atmosphere so that the air closest to the ground is warmest. The warming is enhanced because water vapor, carbon dioxide, and some other gases, as well as particulate matter, absorb the re-radiated heat, warming the air and radiating heat back to the surface. As a result of these processes, the usual pattern is for the temperature of the air to decrease with height above the ground. On average, the air temperature decreases by about 6.4C for each 1000 meters of elevation (3.5F for each 1000 feet). This is just an average, howeverlocal anomalies and departures from the average are common and are important in understanding weather phenomena. A result of the decrease in temperature with altitude is that locations on the Earth that are topographically higher will have signicantly cooler temperatures, on average, than locations at the same latitude at lower elevations. This effect is slightly counterbalanced by the fact that the air is thinner (because pressure also decreases with elevation) and the path of the sunlight through the absorbing atmosphere is shorter, so that surface heating on a clear day on a high mountain will be greater than on a spot at sea level at the same latitude. The cooling with increasing altitude has profound climatic consequences when air masses move from regions of low elevation across mountains. As the air mass is forced upward, it is cooled. Because cool air cannot contain as much moisture as warm air, moisture-rich air masses will produce precipitation as they are carried upward. The highest rainfalls on Earth are in places where this effect is particularly strong, the best examples being places where high mountains occur on islands or continental coasts. For example, Mount Waialeale on Kauai, Hawaii, is such a mountain. The average annual precipitation at the highest elevations on the windward slope is over 1200 centimeters (40 feet). This is extreme, but mountains typically have both cooler and wetter climates than the adjacent lowlands, a fact well known to anyone with an interest in downhill snow sports.

Descending, warming air Rising, cooling air Prevailing wind

Rain shadow

324 Rain shadow The effect of coastal mountains on patterns of precipitation in the Northern Hemisphere. As winds come off the water, the air is forced upward by the contour of the land, cools, and releases its moisture in the form of rain or snow. As the air descends on the far side of the mountains and becomes warmer, its capacity to hold water increases, producing a rain shadow.

The tendency of mountains to extract moisture from air masses affects the downwind climate. As the air descends, it will warm, and its capacity to contain moisture will correspondingly increase, so that even if downwind areas provide some moisture to the atmosphere by evapotranspiration, there will be limited precipitation. For example, the leeward side of Mount Waialeale receives only about 50 centimeters of annual precipitation. This downwind dryness is the rainshadow effect (Figure 324). It is particularly strong along the Pacic coast of North America, with zones of aridity or semiaridity on the east side of mountain chains from the northern United States down into Mexico. The climatic changes that one observes when climbing a mountain mimic in several important ways the changes that occur as one travels north from the equatormainly with respect to temperature and the occurrence of freezing conditions (Figure 325). In general, a change in mean atmospheric temperature corresponding to an increase in latitude of 1 occurs with each increase in elevation of

Snowcapped peak Snow Tundra Tree Altitude

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Taiga Temperate deciduous forest

Tropical rainforest Equator Latitude

Polar ice

Pole

325 Relationship between altitude and latitude In the Northern Hemisphere, we can experience a similar sequence of dominant plant life and its associated animal life by either traveling north for hundreds of kilometers or ascending a mountain. This relationship between altitude and latitude was rst pointed out by Alexander von Humboldt. To experience a similar sequence of vegetation in the Southern Hemisphere, we could ascend a mountain. However, by simply traveling south, we would never encounter vegetation corresponding to the taiga and the tundra of the Northern Hemisphere. Can you explain why?

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Alexander von Humboldt

Alexander von Humboldt (17691859) was perhaps the greatest scientic traveler who ever lived and was certainly one of the greatest writers and scientists of his era. A native of Germany, Humboldt ranged widely across the interior of Latin America from 1799 to 1804 and climbed some of its highest mountains. Exploring the region between Ecuador and central Mexico, Humboldt was the rst to recognize the incredible diversity of tropical life and, consequently, the rst to realize just how vast a number of species of plants and animals there must be in the world. In his travels, Humboldt was impressed with the fact that plants tend to occur in repeatable groups, or communities, and that wherever there are similar conditionsrelating to climate, soil, or biological interactionssimilar groupings of plants appear. He also discovered a second major principle the relationship between altitude and latitude. He found that climbing a mountain in the tropics is analogous to traveling farther north (or south) from the equator. Humboldt illustrated this point with his well-known diagram of the zones of vegetation on Mount Chimborazo in Ecuador, which he climbed. On this mountain, he reached the highest elevation on record attained by any human being up to that date. On leaving Latin America in 1804, Humboldt visited the United States for eight weeks. He spent three of these weeks as Thomas Jeffersons guest at Monticello, talking over many matters of mutual interest. It is thought that Humboldts enthusiasm for learning about new lands encouraged Jeffersons own great scheme for the exploration of the western United States. Thus, it is tting that Humboldts name is commemorated in the names of several counties, mountain ranges, and rivers in the American West.

approximately 100 meters. As a result, climbing a tall mountain can take you through a sequence of ecosystem types that will resemble what might occur if you went north toward the Arctic. For example, in the southwestern United States, the highest mountains are topped by conifer-dominated forests, then, at higher elevations, by tundra-like meadows and scree elds, and eventually by a zone of perpetual snow cover. In the tropics, the climatic changes are just as dramatic, but the vegetation of the cold upper elevations does not necessarily look like anything you will ever see in Alaska or Siberia. This does not refute the generality, but shows only that seasonality is an important aspect of climate.

Soils and Fire Inuence Regional Patterns of Distribution

The sketch we have just given of the major factors shaping biome types omits many other important inuences on plant growth. Of these, two deserve mention because of their importance at regional levelssoils and re. Soil, as the medium of plant growth, determines how much water and what amounts of necessary nutrient elements are available to plants. The kind of soil present within a region is deter-

mined, in part, by climate but also by the parent material in which the soil forms and by the length of time that a region has been stable and therefore has had surfaces exposed to the processes of soil weathering. Australia, for example, has been geologically stable over millions of years, and largely because of this, it has many areas that are nutrient poor, with deciencies of essential elements and especially phosphorus. It is thought that these deciencies partly explain unique features of Australian vegetation, such as the abundance of sclerophyllous (leathery-leaved) species, most notably the many species of Eucalyptus. In contrast, the continental glaciers of the Pleistocene that covered much of northern North America and Eurasia renewed the topsoil with fresh material, as bedrock was ground up by the glacier and redistributed in the form of glacial till and as windborne dust (loess) was carried far beyond the edges of the glaciers. Geologically speaking, such glacier-derived or glacier-affected soils are very young, so they have high levels of essential elements. Fire is particularly important for biomes that have alternating wet and dry seasons, such as grasslands, open savannas, or the chaparral of California, and much less so in evergreen, moist areas, such as tropical forests. In some

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regions, res determine the line between two vegetation types, as between Eucalyptus woods or savannas and moist tropical forests in eastern Australia. Here the area of moist forest has progressively been reduced as indigenous people have set res to improve grazing. For at least 1.4 million years, human beings have had the ability to start and keep articial res, which they have done extensively in some areas, especially during the last 20,000 years or so. In regions that have burned, most species of organisms survive, and in fact they often depend on res for the renewal of their populations. Wildres, which are set by lightning, generally consume masses of vegetation that have built up in the moist season, and these res may burn each dry season. In regions where lightning is particularly frequent, such as the state of Florida, there may be many wildres. Along with res set accidentally or sometimes even deliberately by humans, wildres in the western United States may burn hundreds of thousands of square miles of re-prone forests, woodlands, brushlands, and grasslands each year. Sometimes there are disastrous consequences for cities, towns, or individual houses or structures isolated in areas that burn periodically, and even the loss of lives. Hundreds of res may break out in a single day over the region, greatly taxing the ability of reghters to battle them effectively. As people have built homes in areas where res are a normal part of regeneration and renewal, the tendency is to suppress res. This suppression can lead eventually to the accumulation of large amounts of dry vegetation that act as tinder when they are ignited. When these res occur along the edges of cities, their effects are greatly magnied, and they may cause great loss to human life and to property. Better methods of managing vegetation, either by controlled burns or other methods of removing the accumulated biomass, will be necessary if these res are to be limited in number and scope in the future. Controlled burning by setting prescribed res works best in areas characterized by low-intensity res. In areas subject to high-intensity res, such burns can rage out of control, as happened around Los Alamos, New Mexico, in 2000. All methods of suppressing res affect both the appearance of the forests or other vegetation types and the well-being of the biodiversity they contain. The application of re-suppression methods has become the subject of arguments, often erce, among proponents of different approaches to the problem. Understanding the frequency of res in a particular area provides facts useful in predicting and managing future res. This information can be obtained by examining re scars on trees; analyzing sediments in lakes, ponds, or other wet areas; or studying the written history of particular areas. Thus, we have learned that lodgepole pines (Pinus contorta), which occur frequently at higher elevations throughout the western United States, usually burn at intervals of 250 to 400 years. Following a re, these pines can then germinate and grow well in the bare soil, outcompeting other trees locally. Lodgepole pines also have cones that remain closed and attached to the branches until they are heated. The heat melts the thick, waxy substance coating the cones and

releases the seeds at a time most suitable for their germination and establishment. The huge re in Yellowstone National Park in the summer of 1988 was basically an example of a natural re cycle in the park, burning mainly through lodgepole pines (Figure 326). Subsequent examinations showed that a similar cycle of res had occurred around the year 1700. Fires have many signicant ecological effects. They release stored nutrients, such as phosphorus, calcium, and potassium, from the woody or perennial vegetation back into the soil. This stimulates the regrowth of plants, which temporarily have access to large amounts of nutrients. Some plants exist as seeds stored in the soil, and they germinate and ower only after a re. Certain perennials grow luxuriantly and ower only after re releases necessary amounts of nutrients, and the plants have sufcient light. Particularly on relatively infertile soils, res may be essential for the renewal and regrowth of the vegetation. At the same time, soil erosion may be accelerated, which is why management agencies are often anxious to sow cover crops on burned slopes in the wake of a re. Fires not only remove vegetation, but the intense heat associated with res can cause soil particles to become water repellent so that water runs off rapidly, accelerating erosion still further.

326 Lodgepole pines Following the disastrous 1988 re in Yellowstone National Park, lodgepole pine (Pinus contorta) seedlings became established at the burn site.

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Fire and its consequences in particular areas will be discussed further in the biome descriptions that follow.

Rainforests
It makes sense to begin our exploration of biomes by asking what kind of system is found where the fewest factors are limitingwhere temperatures are relatively constant and never below freezing, where rainfall is consistent and therefore soil moisture ample throughout the year, and where geology and topography allow deep soil formation and retention of at least an adequate supply of nutrients. These conditions favor high and consistent photosynthesis and therefore maximum plant growth. Plants can expand their canopies rapidly and, without the constraints of freezing and drought, can have large or thin leaves for maximum absorption of light. The result is strong competition for light. Plants that fall behind in capturing sunlight are quickly overtopped. Over evolutionary time, this has favored a proliferation of woody species capable of obtaining their place in the sun by one means or another. The wet lowland tropics therefore are dominated by tropical rainforestsdense forests consisting of species that are active all year round. The plants are either evergreen (that is, their leaves are always actively photosynthesizing) or leaess for only very brief periods. The dominance of trees inuences all other aspects of the rainforest. Much of the sunlight is absorbed by the dense canopies, leaving relatively little light to penetrate to the forest oor. As a result, undisturbed tropical rainforest usually has only a sparse, patchy herbaceous understory consisting of species with special adaptations for dealing with low light intensity. Seedlings and saplings of shade-tolerant trees are often the most abundant plants in the ground layer. Because of the dominance of trees in the rainforest, the biomass in the system that is of value to herbivores is mostly in the tree canopies. This, and the favorable climate,

explains the abundance of arboreal (tree-dwelling) animals, such as gibbons and howler monkeys, which exhibit a way of life highly specialized to exploit the fruits, insects, and nutritious new growth of the treetop environment. Trees in all regions (unless subject to air pollution) have algae, mosses, and lichens on their branches and bark, but in the wet tropics, conditions favor an abundance of plants specially adapted to grow on trees. These plants, known as epiphytes (meaning on plants), grow on the branches of other plants that grow in the illuminated zone far above the forest oor (Figure 327). Epiphytes can survive without soil by collecting rain as well as nutrients from animal droppings, plant litter, dust, and dead insects. There are epiphytes from many plant families, but two well-known groups are the orchids and the bromeliads; there are also many epiphytic ferns. Lianas, or woody vines also known as climbers, represent another strategy for exploiting trees for support. Lianas are rooted in the soil, but grow up along the trunks of the trees into the canopy. Some lianas start as epiphytes and drop stems to the soil where they take root. Strangler gs (Ficus spp.) begin growth as epiphytes, dropping roots to the ground and growing to envelop their host trees. The g eventually becomes an independent, hollow tree, while its original host often dies. Tropical rainforests have extraordinary species richness. A typical hectare (2.47 acres, or about 2.5 football elds) of boreal coniferous forest will have 1 to 3 tree species, a rich temperate deciduous forest 10 to 20 species, but tropical rainforests commonly have 100 species, and some have 200 or more. Putting this in more concrete terms: in a typical temperate forest, there is a very good chance that if you select a tree, the tree closest to it will be the same species; in tropical rainforest, the tree closest to your selected tree is more likely to be some other species. Ecologists are still searching for a full explanation of why tropical rainforests are so diverse. It seems reasonably certain that uninterrupted

327 Epiphytes By collecting and storing water and nutrients from the surrounding air, rain, and dust, epiphytes create little patches of soil from accumulated debris. Bromeliads, seen here as leafy tufts growing along the tree branches, are among the most common of the epiphytes. The leaves of bromeliads merge at their bases to form watertight tanks that, in the larger species, can hold as much as 45 liters of rainwater. These pools of water are microcosms of bacteria, protozoa, larvae, insects, and insect-eaters. Many rainforest mosquitoes breed exclusively in bromeliad tanks. The bromeliads absorb water from their built-in reservoirs and are also supplied with nutrients from the debris.

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evolution is part of the reason, so that over time more species accumulate. This could be more or less chance, or it could represent the action of evolutionary pressures for the subtle differentiation of species to ll different ecological roles. Other theories stress that seed predators and pathogens might act disproportionately on the most common species, or environmental stability may make the extinction of smaller populations less likely. Like all natural systems, rainforests are dynamic and constantly changing. Because of the abundant rainfall, natural res are rare. But tree fall, usually due to wind, is common and is the main natural mechanism by which rainforest is disturbed. Those tropical rainforests that lie within the

reach of hurricanes can suffer widespread canopy loss, but such catastrophes are not essential for tree fall. Although rainforest trees are well protected against wind damage by a variety of mechanisms, including rot-resistant wood and buttressing (Figure 328), sooner or later they succumb. Even if it is only initially a single tree that falls, it often brings down other trees around it by direct impact, uprooting, or because the canopy of the falling tree is connected to others by lianas. The result is a large gap in the canopy. Light can then penetrate to the forest oor, and a strong pulse of growth by herbs, shrubs, seedling trees, and resprouted trees follows. Any sapling trees that survived the falling of their large neighbors can experience a growth

(a)

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328 Tropical rainforest (a) The interior of a rainforest in Costa Rica. The broadleaved plants with red owers are Heliconia irrasa. (b) The diversity of the trees in the forest, which may reach several hundred species per hectare, is revealed when individual trees burst into bloom, such as these trees in the coastal rainforest of Brazil. (c) A buttressed tree in the rainforest of Ecuador. Note the woody vines known as lianas on the trunk.
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spurt that allows them to extend up into the canopy. Some species of tropical rainforest trees require such sunny patches to become established. Temperate areas with cooler climates also can have climates with high rainfall. These rainforests of higher latitudes, like those along the eastern side of Australia, in Tasmania, and in parts of New Zealand, have much lower species diversity but share the quality of dense tree growth.

Rainforests Are Rapidly Being Destroyed

Tropical rainforests, which already have been cut back to about half of their original extent, are being lost rapidly as a result of human activity. Since they are estimated to contain half of the worlds species of plants and other organisms, this loss is extremely serious. A traditional pattern of human use, known as shifting cultivation, mimics the natural dynamics of the rainforest by opening small gaps in the canopy, cultivating crops for a few years, and then abandoning them. These areas then undergo succession and return to natural forest cover. This kind of land use is thought to be sustainable, but only so long as population levels are low relative to the land being exploited. With increasing populations, rotation time is shortened. But the greatest threat to the tropical rainforest is the conversion of forest to cropland and open pastures. In areas less suitable for agriculture, forestry practices often involve removing more trees than can be replaced by natural succession. Creating large gaps in the tropical rainforest makes it susceptible to re to a

degree unprecedented in history, as was tragically demonstrated by the destructive res of 19971998 in the Kalimantan region of Indonesia, where over 5 million hectares of forest were destroyed. Destruction of rainforest of this kind will lead to the loss of many species of plants, animals, and microorganisms. The destruction of tropical ecosystems would be more defensible if the results were always highly productive agricultural lands. But experience shows otherwise. Tropical soils often prove to be fragile. Tropical vegetation is lush, but in large areas of the tropics, upland soils tend to lose fertility rapidly when cleared for agriculture. Much of the fertility in tropical rainforest ecosystems is tied up in the vegetation. Soils are generally acidic and decient in calcium, phosphorus, and other nutrients. Plant roots tend to spread out in a thin layer no more than a few centimeters below the soil surface, and they rapidly transfer the nutrients released by the decomposition of fallen leaves and branches back into living plants. Below this thin layer of topsoil, which is easily destroyed during the process of clearing away the trees, there is virtually no organic matter. Fertilization may not help much because tropical soils often x the phosphorus into insoluble forms. For all of these reasons, the cultivation of tropical soils presents formidable problems. Nevertheless, the tropical forests of the world are being cut and burned at an ever-increasing rate, often to produce elds that become completely useless to agriculture within a few years (Figure 329). It is estimated that by the middle of this century, only about 5 percent of the area originally covered

329 Erosion Of the native vegetation that once covered Madagascar, an island in the Indian Ocean 420 kilometers east of the African mainland, only 10 percent remains, and that is fast disappearing. On Madagascar, as in other tropical areas, the soils exposed by clearing vegetation are often not suitable for sustainable agriculture. The abundant rains rapidly wash the topsoil off increasingly barren landscapes, choking the rivers with silt, as seen here, and sometimes causing oods and landslides. The sea around Madagascar is often rust-red from the loss of the islands weathered red soil, and it is difcult to produce enough food to sustain the people of the world's fourth-poorest country.

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by tropical rainforests will survive, and those relatively small patches will be extremely vulnerable to further harm.

Savannas and Deciduous Tropical Forests

Not all areas in the tropics have high or continuous rainfall. In regions in which there is a prolonged dry season alternating with a rainy period, tropical rainforest gives way to other forest types tolerant of drought. These forests can be grouped under the general term of tropical seasonal forests. For example, in central and southern Africa, the Indian subcontinent, and Southeast Asia, there are extensive areas of monsoon forest, and similar climates and forests are found in northern Yucatn Peninsula in Mexico, in northern Colombia and Venezuela, in Ecuador, and in eastern Brazil (Figure 322). These climates produce enough rain to permit vigorous tree growth and dense and productive forests, but the dominant trees are deciduous, losing their leaves during the dry season. Elsewhere in areas of the tropics with seasonal rainfall, there are other types of dry forests, such as the short-tree forests of tropical Mexico and Central America, or the biologically distinct dry forests of Madagascar. Tropical mixed forests, in which both evergreen trees and shrubs are present, occur locally in eastern and southern Brazil and northern Australia (Figure 322). The disappearing tropical rainforest has caused alarm among conservationists, but dry tropical forests also have extraordinarily high levels of biodiversity and are equally deserving of protection. Although pine trees are often associated with northern forests, pines also occur in the tropics and subtropics. Most of Florida and extensive areas throughout the southeastern United States are covered by subtropical mixed forests. In these forests, pines (Figure 3210) and some evergreen broadleaf trees are mixed with deciduous trees; the precipitation occurs mainly in the summer. With further decreases in rainfall, a point is reached at which moisture is not sufcient to sustain a continuous tree

3210 Subtropical mixed forest Slash pine (Pinus elliottii) is one of the widespread evergreen conifers that occurs in the subtropical mixed forests of the southeastern United States. Conifers tend to dominate on soils that are either nutrient-poor or seasonally ooded, or both.

canopy. Trees and shrubs are scattered individually or grow in groups in a grassland matrix, and, in places, woody plants nearly disappear. Such areas are called savannas, and they show many variations worldwide. Savannas usually have much lower annual rainfall than tropical rainforestsfrequently in the range of 90 to 150 centimeters a year. There is also a wider range in average monthly temperatures, due to the seasonal drought and the sparse covering of vegetation. In the tropics, savanna trees are broad-leaved deciduous and evergreen trees, which may occur singly or in groves, and some savannas are dominated by shrubs (Figure 3211).

3211 Savanna A savanna in East Africa with giraffes surrounded by a herd of impala. The transitional nature of this biome, relative to the characteristic vegetation of the tropical rainforest biome and the desert biome, is evident in the grasses, shrubs, and short trees (acacias) seen here.

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Savannas cover large areas of East Africa and are also found on all continents on the margins of rainforests, where the rainfall is seasonal and limiting (Figure 322). The thorn forest (cerrado) that covers wide areas in Brazil belongs to the savanna biome. In savannas, because of the scattered distribution of taller woody plants, the ground is generally well illuminated, and perennial herbs (mostly grasses) are common. Bulbous plants, which are able to withstand periodic burning, can be abundant. Because of the dense cover of perennial herbs made possible by the high seasonal rainfall, there are few annual herbs. Epiphytes are also rare because of the stress of the prolonged dry season. Trees that occur in savannas often have thick bark as protection against re. They are well branched, but they are seldom more than 15 meters tall. Many trees, such as the acacias, are protected by stout spines to discourage grazing (Figure 317a). Their leaves are generally smaller than those of the evergreen trees of the rainforest, and so they lose less water by transpiration. Fire, which is common in all tropical savannas, is a strong inuence on the relative abundance of trees, shrubs, and grass. Where soils and climate allow dense grass cover to develop, re can be frequent, sometimes occurring annually. With such high re frequencies, trees are limited to protected situations, such as along rivers or in rocky areas. Thus a climate already challenging for trees can favor re, which will further restrict and perhaps even completely eliminate trees over large areas. Savannas and similar seasonally deciduous tropical and subtropical plant communities grade into tropical rainforests as the rainfall becomes higher and its seasonal distribution more even, and corridors of tropical rainforest (called gallery forest) extend out into the savannas along rivers. At their poleward margins, savannas and related plant communities grade into deserts. Savannas extend into temperate areas, occurring in the region between the prairies and temperate deciduous forests, as well as in the region between prairies and taiga in North America and over much of eastern Mexico, Cuba, and southernmost Florida. Savannas in these cooler, moister climates are thought to be dependent on re for their maintenance. Many believe that the savanna vegetation was greatly expanded in pre-Colombian times because of burning by the indigenous people. Deliberate burning was also practiced in other parts of the world, often to improve grazing for domestic stock, a practice that continues today in many places.

Deserts
With further decreases in rainfall, a point is reached at which extended drought prevents the plant cover from being continuous in time or space or both, resulting in deserts. Bare patches of soil appear, and dense vegetation is conspicuously restricted to drainages, or areas where local runoff accumulates, such as at the base of rocky cliffs, or around the rare springs or seeps, as in the desert palm oases. The most important factor creating the dry conditions is the relative stability of the atmosphere in desert regions. The great

deserts of the world are all located in the zones of atmospheric high pressure that ank the tropics at about 30 north latitude and 30 south latitude, and they extend poleward in the interior of the large continents (Figure 322). Many deserts receive less than 20 centimeters of rainfall per year. In the Atacama Desert of coastal Peru and northern Chile, the average rainfall is less than 2 centimeters per year. Averages, however, do not tell the whole story, because as average rainfall decreases, the variability from year to year increases. In a humid region, a severe drought would be 50 percent less rain than average, but in a desert a drought can be no precipitation at allsometimes for several years. The plants and animals of extremely arid regions must be able to tolerate these periods of extreme dryness and to exploit the good years when they occur. Extensive deserts are located in North Africa and in the southern part of the African continent, where the Namib Desert is inhabited by some of the worlds most extraordinary organisms, including Welwitschia (Figure 1838). Other deserts occur in the Near East, in southeast Mongolia and northern China, in western North America and western and southern South America, and in Australia. The Sahara, which extends all the way from the Atlantic coast of Africa to the Arabian peninsula, is the largest desert in the world. Seventy percent of the entire continent of Australia is covered in semiarid or arid areas. Less than 5 percent of North America is desert, and much of that is not extreme (Figure 3212). The temperatures in many desertsthe hot desertsare very high because the skies are usually completely clear and little heat is taken up in evaporation and transpiration. Summer temperatures of more than 36C are common in some deserts. The same conditions that make deserts hot in the day cause them to cool rapidly at night. With little or no cloud cover, generally clear air because of limited moisture, and no tree canopies to trap outgoing radiation, the heat stored during the day is lost through re-radiation, and temperatures drop precipitously as soon as the sun sets. Human beings can die from exposure in deserts, not only from extreme heat but also from cold. Although all deserts are dry, not all of them are characterized by continuously high daytime temperatures. There are also cold deserts. An example is the Great Basin Desert of western North America, which lies between the Sierra NevadaCascade Mountain system and the Rocky Mountains (Figure 3212d). Here there are only a few weeks of high temperatures each year, and winters can be bitterly cold, with some of the scant precipitation falling as snow. The annual distribution of rainfall in deserts generally reects that of the adjacent areas. On the equatorial side, it rains in the summer; on the poleward side, it rains in the winter. Between the two, as in the lowlands of Arizona, there may be two annual peaks of precipitation. As a result, such an area generally has two periods of active plant growthone in the winter and one in the summerand different plants are active in each period. In general, the patterns of activity of desert plants reect the origins of the plants. Characteristically, plant species that have migrated

Deserts

3213

(a)

(b)

(c)

(d)

3212 Desert Shown here are some representative plants of the principal deserts of North America. (a) The Sonoran Desert stretches from southern California to western Arizona and south into Mexico. A dominant plant, the giant saguaro cactus (Carnegiea gigantea), is often as much as 15 meters high, with a widespreading network of shallow roots.Water is stored in a thickened stem, which expands, accordionlike, after a rainfall. (b) To the east of the Sonoran is the Chihuahuan Desert; one of its principal plants is the agave, or century plant (Agave deserti), a monocot. (c) North of the Sonoran is the Mojave Desert, with its characteristic plant, the Joshua tree (Yucca brevifolia). This plant was

named by early Mormon colonists who thought that its form resembled that of a bearded patriarch gesticulating in prayer.The Mojave contains Death Valley, the lowest point on the continent (90 meters below sea level), only 130 kilometers from Mount Whitney, with an elevation of more than 4000 meters. (d) The Mojave blends into the Great Basin Desert, a cold desert bounded by the Sierra Nevada to the west and the Rockies to the east. It is the largest and bleakest of the American deserts. The dominant plant is sagebrush (Artemisia tridentata), shown here with the snow-covered Sierra Nevada in the background.

from areas where they grew actively in the winter continue doing so in the desert. Similarly, plant species that have migrated into the deserts, from areas where they grew during periods of summer rainfall, continue to grow actively in the summer in the desert, wherever a sufcient amount of summer rainfall makes this possible.

Desert Plants Are Adapted to Low Precipitation and Extremes of Temperature

High variability in moisture conditions means that desert plants must be able either to survive the dry season or dry

periods in a dormant condition or to have some mechanism of averaging out variable moisture conditions. The life history of annual plants is ideally suited to survival through droughts of unpredictable length. It therefore makes sense that annuals are better represented, both in number and in kind, in the deserts and semiarid regions of the world than in any other biome. Annuals can germinate and complete their life cycles in the open areas of ground during the limited periods when water is available. The seeds of these plants are able to survive extended droughts in the soil. Then, when rains thoroughly wet the soil, the seeds can germinate rapidly, ower, and set seeds before the soil moisture

3214

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Global Ecology

3213 Creosote bushes One of the most characteristic plants of the Mojave, Sonoran, and Chihuahuan deserts of North America is the creosote bush (Larrea divaricata), which has small, leathery water-conserving leaves. Creosote bushes in the Mojave Desert may form circular or elliptical clones because of new branch production at the periphery of stem crowns and the segmentation and death of older stem segments, resulting in a ring of satellite bushes around a central bare area.The latter usually accumulates a mound of sand, which may reach a depth of about half a meter. Some clones may attain extreme ages: the King Clone, shown here, is estimated to be nearly 12,000 years old. Such ancient clones started from seeds that germinated near the end of the last glacial expansion.

is depleted. At the other extreme are species of plants, the succulents, that can endure the dry periods by storing water. Succulent plants are found in a wide range of plant families, but two groups are notable for having many succulent speciesthe cacti of the New World, essentially all of which are succulents, and the succulent euphorbs (Euphorbiaceae) of the Old World. Succulents are designed to absorb water while it is available and store it in special tissues. CAM photosynthesis provides another means of conserving the stored water. CAM plants absorb carbon dioxide only at night; their stomata remain closed during the day. By this means, they are able to take up carbon dioxide with much reduced water loss. Another feature of succulents is a morphology that allows them to change volume without damage (see the essay How Does a Cactus Function?). The success of the succulent life history is evidenced by the fact that succulents often are among the tallest and most robust plants of desertsthe tall saguaro cactus of the Sonoran Desert of the southwestern United States being a wellknown example (Figure 3212a). Other plantscreosote bushes are an exampledo not store large amounts of water, but they have the ability to tap into deep-water reserves by means of extensive root systems (Figure 3213). Because deep water is accumulated by slow downward percolation, it tends to vary much less than surface water and to be available during dry seasons and, in some situations, through many dry years. Tapping deep water would not work, however, if the plants were not able to use water efciently. Adaptations, such as small, leathery leaves with relatively few stomata and stems resistant to the loss of conductivity due to embolisms, restrict water loss and allow the plant to resist wilting when under extreme moisture stress. C4 photosynthesis is likewise more common among the plants of deserts and other seasonally dry, warm habitats than it is elsewhere. Such plants also have the

capacity to drop leaves and even entire branches in dry periods so that they can maintain their water balance. Then, when better conditions return, they can regenerate their canopies. Other plants have green stems rich in chlorophyll and thus are able to photosynthesize even when leaess. Perennial herbs face difcult challenges in the desert because of the extremely dry conditions, but grasses are found in many deserts, usually as widely scattered, dense bunches known as tussocks. Some plants have deeply buried bulbs that break their domancy when stimulated by rain, while others occur along arroyos or washes, where moisture is available near the surface. The amazing capacity of plants to adapt to desert conditions is exemplied by the graywhite wooly shrub known as Arizona honeysweet (Tidestromia oblongifolia), a C4 perennial herb common in the Death Valley region of California and Nevada. The maximum photosynthetic rate for this species is achieved at temperatures between 45 and 50C in full sunlight in midsummertemperatures that would be damaging or even fatal to plants of regions with higher moisture. As we have discussed, deserts grade into savanna-type communities. One such community is the widespread juniper (Juniperus spp.) woodlands of western North America (Figure 3214). Juniper woodlands occur in places that are usually drier than will support prairie but wetter and cooler than desert areas. In many places in the western United States, juniper, often found with pinyon pine, grows in areas adjacent to deserts but at higher elevations. Studies of pollen and macrofossils preserved in pack-rat middens show the degree to which vegetation types that seem like permanent features of the landscape actually are quite dynamic over time. During the pluvial periods of the Pleistocene epoch, at the times of maximum expansion of the continental glaciers, juniper woodland communities moved down into the lowland areas that presently support juniper-free deserts.

Deserts

3215

How Does a Cactus Function?

The barrel cactus Ferocactus acanthodes, which grows along the northwestern edge of the Sonoran Desert in southern California, gets its name because it is shaped like a barrel. It looks as if its stem has been folded like a fan, and it is covered with spines like a porcupine. Why should this desert plant present such an unusual and formidable appearance? When the barrel cactus is full of water, the folds swell and are barely visible, but when the plant dries, the folds are deep and the stem is able to contract without crushing the cells. This ridge-and-valley folding of the stem has other advantages, too. Deep inside the valleys between the folds are the stomata. The spines help to break up the wind currents, and thus the valleys serve as protected retreats where dry desert winds cannot easily reach to carry moist air away from the vicinity of the stomatal chambers.These formidable spines also help to protect the cactus from the rodents and birds that are in constant search of water, even going so far as to steal it from a succulent stem. In a study conducted by Park Nobel, of the University of California, Los Angeles, barrel cacti stored enough water in their succulent stems to permit them to open their stomata for about 40 days after the soil became too dry to furnish them with any additional water. Under such conditions, many of the ne roots are sloughed off, to prevent water loss to the soil. After seven months of drought, stomatal activity ceased, and the osmotic potential of the stem was more than double the value it had been during wet periods, despite the ability of the stem to fold and shrink. When rain nally returned, the shallow root systems (with a mean depth of only about 8 centimeters) took in water so rapidly that the stomata were fully functional again within 24 hours of rainfall. But these are not the only mechanisms the barrel cactus uses for conserving water. Like many other desert plants, the barrel cactus exhibits crassulacean acid metabolism (CAM). It opens its stomata only at night and so undergoes gas exchange with cooler air, which can hold less water than warmer air. Consequently, the plant loses less water to the atmosphere through transpiration. Its ratio of mass of water transpired to mass of CO2 xed is only about 70:1 for the entire year. This is considerably lower than for a typical C3 plant, which requires larger quantities of water to x an equivalent amount of carbon but has a higher maximum rate of photosynthesis. Because seedlings of the barrel cactus, unlike their parents, cannot tolerate extremely high temperatures and prolonged drought, they survive only in certain years and in protected microhabitats. By the age of 26 years, these plants have usually grown to only about 34 centimeters tall, adding about 10 percent to the mass of their stems each year.

Barrel cactus in a hydrated state (left), and in a dehydrated state (right).

(a)

(b)

3214 Juniper woodlands (a) Juniper (Juniperus osteosperma) savanna in the Great Basin near Wellington, Utah. (b) Cold winters

are characteristic of the pinyon-juniper savannas and woodlands of the Great Basin, as shown here south of Moab, Utah.

3216

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Global Ecology

Grasslands
Grasslands occur where the amount of rainfall is less than is needed to support vigorous tree growth but great enough to allow herbaceous plants, especially grasses, to dominate. There are many kinds of grasslands; the major variations relate to rainfall. Grasslands with the highest rainfall are dominated by tall grasses in closely spaced clumps or dense sods. Grasses in drier areas occur in widely spaced clumps, with sparse or virtually no vegetation between them. Grasslands occur from the tropics to the edges of the boreal regions and from sea level to high mountain meadows. They grade into savanna and desert and occur as patches in woodland and forest. They are most extensive, however, in the middle latitudes. These grasslands are characterized by cold winters and warm summers, with moderate to low rainfall and occasional periods of extreme dryness. Grasses, which regenerate at the beginning of each growing season from buds at or below ground level, have many ne roots. Over long periods of time, grasslands tend to build soils that are deep and rich in organic matter. Such soils are ideal for agriculture, and vast areas of grassland have been converted to cropland. In areas too dry for agriculture, grasslands have value as grazing lands (Figure 295). Grasslands generally occur over large areas in the interior portions of the continents, most notably in North America and across Eurasia (Figure 322). In North America, there is a transition from the more desertlike, western shortgrass prairie (the Great Plains), through the moister, richer, tallgrass prairie (the Corn Belt), to the eastern temperate deciduous forest (Figure 3215). Grasslands become progressively drier

with increasing distance from the Atlantic Ocean and the Gulf of Mexico, which are the major sources of moisture-bearing winds in the eastern half of the North American continent. Farther north, grasslands become moister again as evaporation decreases at relatively cool temperatures. Typical shortgrass prairie occurs under near-drought conditions. In years with greater moisture, taller grasses (but not the very tall ones of the tallgrass prairie) tend to predominate. Grasslands also occur as patches within forested biomes when soil conditions, such as shallow soil over bedrock, limit tree growth. In the central and eastern United States, such grassy patches are often called glades, and these are typically dominated by prairie plants (Figure 3216). Perennial bunchgrasses and sod-forming grasses dominate, but other perennial herbs are common. Although the growth of grassland plants is seasonal, there is little room for the development of annual herbs, and these are mostly limited to disturbed areas, such as prairie-dog towns or badger diggings. Annuals and introduced weeds are common in areas disturbed by humans, such as near buildings and along roadsides and railroads. Because most grasslands have more or less continuous cover and die back for part of the year, they carry re readily. Where woody plants and grasses coexist, res carried in grasses can top kill or otherwise injure trees and shrubs. As a result, re can push back woody plants and increase the dominance of grasses. Though such patterns are natural, humans can increase the frequency with which res occur. It is thought that burning of grasslands by Native Americans, for example, may have increased savanna vegetation.

(a)

(b)

3215 Grasslands The grasslands of North America include large regions of shortgrass and tallgrass prairie. (a) A female bison nursing her calf in the shortgrass prairie of Custer State Park, South Dakota.

(b) A June day on a tallgrass prairie in North Dakota. The cottonwood grove by the prairie creek is characteristic of this biome.

Temperate Deciduous Forests

3217

epoch, when they were widely hunted by our ancestors. Many of these grazing mammals, such as the American bison, have since been hunted almost to extinction. They survive today mainly in refuges, having given way to herds of domestic animals and cultivated elds.

Temperate Deciduous Forests

As we move poleward from the tropics, seasonality increases, and eventually, at about 35 degrees latitude, we encounter climates having long periods with temperatures below freezing. If rainfall is well distributed through the growing season, plant growth can ourish, and the tree form, which can compete successfully for sunlight, is favored for the same reasons that it is in the rainforests. The kinds of trees and the structure and function of the forests are, however, quite different. On fertile soils in these regions, the dominant trees lose their leaves at the end of the growing season and remain leaess through the winter. From an energetic and physical standpoint, this pattern can be explained by the fact that it requires less energy and involves fewer risks to produce a new leaf than to try to maintain an evergreen leaf through the rigors of the cold season. Even if it were possible to sustain photosynthetic function at a reasonable level, water is unavailable because the ground is frozen for much of the year, a situation some describe as physiological drought. Despite the fact that the climates in these regions vary between tropical heat and moisture in the summer and Arctic cold in the winter, it is traditional to describe them as temperate and the deciduous forests that are found there as temperate deciduous forests. Such forests were present over large areas in the Northern Hemisphere, but primarily because of the limited land area in the appropriate latitudinal bands they are almost absent in the Southern Hemisphere (Figure 322). Temperate deciduous forests are best developed in areas with warm summers and relatively cool

3216 Glade Coreopsis dominating a glade in temperate deciduous forest near St. Louis, Missouri. Such openings in the forest, which usually occur in areas of shallow soil, are often dominated by prairie plants that form continuous stands beyond the borders of the forest.

When disturbed, grasslands have often changed into either forests or deserts. Thus, the grasslands that once stretched from southern Arizona to western Texas, which were encountered by early European settlers, were largely overgrazed and converted to deserts or shrublands during the past century. The failure of farmers and ranchers to manage grasslands properly led to the dust bowl disasters of the central United States in the 1930s (Figure 3217). All of the great grasslands of the world were once inhabited by herds of grazing mammals associated with large predators. Such herds were widespread in the periods of maximum expansion of the glaciers during the Pleistocene

3217 Dust bowl The prairie soils were once so bound together with the roots of grasses that they could not be cultivated until adequate plows were developed. But once the plants were removed by overgrazing or careless cultivation, prairie soils rapidly deteriorated and were blown away by the wind. This photograph of a badly eroded farmyard, taken in Oklahoma in 1937, vividly recalls the dust bowl conditions that led many thousands of people to migrate away from the central United States. John Steinbecks novel The Grapes of Wrath was based on the experiences of these migrants.

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Global Ecology

3218 Temperate deciduous forest Representative plants of the temperate deciduous forests of North America. (a) A beech and maple forest in Michigan, photographed in the spring. The forest oor is carpeted with largeowered trillium (Trillium grandiorum). (b) In the fall, as seen here in a forest in the southern Appalachian mountains, the leaves of the maples and eastern sourwoods (Oxydendrum) turn a beautiful scarlet.

(a)

(b)

winters (Figure 3218). Annual precipitation generally ranges from about 75 to 250 centimeters and is either distributed evenly throughout the year or concentrated somewhat during the summer months. The temperature-driven annual cycle of growth in temperate forests is a salient feature of temperate deciduous forests (Figure 3219). In winter, the trees are leaess, and their metabolic activity is greatly reduced. In spring, a variety of herbaceous plants burst forth in profusion on the well-illuminated forest oor (Figure 3218a). Some species (the spring ephemerals) have leaves that emerge fully formed from bulbs or rhizomes and complete their period of activity in sunlight before the trees leaf out overhead. Others (for example, early and late summer species, and evergreen species) emerge more slowly and sustain photosynthetic activity longer and later during the shady conditions prevailing in summer. Summer-active species generally have leaves that are broader but thinner in cross section than those active only in spring, and they usually have smaller storage organs than species with shorter growing periods. Forest herbs vary in height from a few centimeters to more than one meter. Variation in leaf height appears to be related to the typical density of the competing foliage that different species encounter in their microenvironments. Most of the species that ripen their seeds in spring are dispersed by ants (page 469), which are active when few other dispersers are present. Most of the species that ripen seeds in the fall, however, are dispersed by birds, at a season that coincides with the height of fall migration. Very few annual plants occur in deciduous forests, probably because the dense plant cover in the moist understory puts seedlings at a strong competitive disadvantage. The soils in regions occupied by temperate deciduous forest are usually acidic, and they tend to contain moderate amounts of nutrients. For these reasons, such soils are easily depleted of nutrients after the forests are cleared, and they may become highly infertile following years of intensive cul-

tivation. Prairie soils are far more fertile and have characteristics much more favorable to sustained cultivation. One of the striking features of the temperate deciduous forest is the similarity of plants found in each of its three main regions in the Northern Hemisphere, often representing related species within the same genus. For example, the herbaceous plants of the deciduous forests of China and Japan resemble those of eastern North America more closely than either group resembles those of western North America. This was less true some millions of years ago, when a band of deciduous forest existed across North America.

Temperate Mixed and Coniferous Forests

Bordering the deciduous forests on the north (as noted above, the temperate forest is limited in the Southern Hemisphere) are mixed forests, in which conifers form an important element along with the deciduous trees. Such temperate mixed forests (Figure 3220) are characteristic of the Great LakesSaint Lawrence River region, much of the southeastern United States, eastern Europe, the northern and eastern border regions of Manchuria (in northeast China) and adjacent Siberia, eastern Korea, and northern Japan (Figure 322). They can be considered to be the intermediate condition between temperate deciduous forests to the south and the taiga to the north. Temperate mixed forests occur in areas with colder winters and more reliable snow cover than are found in areas of temperate deciduous forest. The coniferous component of deciduous or subtropical broadleaf forest also increases in areas with nutrient-poor soils or seasonally saturated soils to form a variant of mixed forest that occurs closer to the equator (Figure 3210). The absence of extensive deciduous and mixed temperate forests in western North America (excluding Mexico) is striking. Study of the fossil record shows, however, that the relative absence of deciduous and mixed forest genera and species is a recent phenomenonin the earlier Cenozoic era, these genera were abundant. Most of the deciduous

Temperate Mixed and Coniferous Forests

3219

3219 Annual growth cycle The four seasons in a temperate deciduous forest in Illinois.The trees leaf out early in spring and begin to manufacture food; they lose their leaves in autumn and enter an essentially dormant state, in which they pass the unfavorable growing

conditions of winter. Many herbs grow under the trees (see Figure 3218), and a number of them ower very early in spring, before the tree leaves reach full size and shade the forest oor. In spring, most of the trees produce abundant pollen, which is carried by the wind.

trees and associated herbs were eliminated in western North America during the latter half of the Cenozoic era as the amount of summer rainfall was greatly reduced. In their place now grow the mountain coniferous forests and mixed west-coast forests of western North America, which contain such trees as the coast redwood (Sequoia sempervirens; Figure 3221), the big tree or giant sequoia (Sequoiadendron giganteum), the Douglas r (Pseudotsuga menziesii), and the sugar pine (Pinus lambertiana; Figure 3113b), all of which

were much more widely distributed in the past. Similar kinds of vegetation are found in areas with similar climatic characteristics in Scandinavia, central Europe, the Pyrenees, the Caucasus, the Urals, southern Tibet, and the Himalayas northward to eastern Siberia. Vegetation types that resemble these are also found in areas in western South America, central New Guinea, southwestern New Zealand, the southern Arabian peninsula, Ethiopia, and the mountains of Central Africa (Figure 322). At higher elevations in these regions

3220

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Global Ecology

3220 Temperate mixed forest In this forest in southern Ontario, the evergreen conifers appear dark green and the deciduous trees are showing their fall colors.

3222 Alpine tundra As seen here on the Olympic Peninsula in Washington State, alpine tundra is comparable in many respects to the Arctic tundra found hundreds of miles to the north. In this area, however, forested slopes are found within a hundred meters or so of the alpine meadows.

are found the open grassland communities called alpine tundra (Figure 3222), which in North America are intermixed with mountain forests from the Brooks Range in southern Alaska southward into the Rocky Mountains, the Cascades, and the Sierra Nevada.

Mediterranean Scrub
Highly distinctive scrub communities have evolved from mixed deciduous-evergreen forests in areas with Mediterranean climates, which are characterized by cool, moist winters and hot, dry summers (Figure 3223). Such climates are found along the shores of the Mediterranean Sea, over a large part of California (and for a short distance to the north and south of that state), in central Chile, on the southern coast of Africa, and along portions of the coast of southern and southwestern Australia (Figure 322). The plants in these areasoften evergreen or summer-deciduous trees and shrubshave relatively short growing seasons that are restricted to the cool part of the year, when moisture is relatively abundant. They may lock up nutrients efciently in their evergreen leaves. In Mediterranean climates, the luxuriant growth of spring is followed by drought and dormancy during the summer. Shrublands cover vast areas in all the Mediterranean climate regions. These shrublands have a similar appearance when viewed at a distance, but each region has its own unique species, and therefore deserves its own name. In California and elsewhere in western North America, chaparral is the word for these mostly evergreen

3221 Mixed west-coast forest Redwoods (Sequoia sempervirens) are a prominent feature of the mixed west-coast forests of California. Watered by the frequent fogs of the region during the dry summers and protected from freezing temperatures by their proximity to the ocean, redwoods often form spectacular groves, many of which, like the one shown here in Muir Woods near San Francisco, are now protected in parks and reserves.

The Northernmost ForestsTaiga and Boreal Forest

3221

3223 Mediterranean scrub Chaparral in the mountains near Los Angeles, California. This sort of plant community, consisting of broad-leaved, drought-resistant, and often spiny evergreen shrubs, occurs only in limited areas of the world, but it has evolved independently on ve continents in areas having Mediterranean (summer-dry) climates. (See also Figure 321.)

and typically very dense shrublands. The equivalent vegetation around the Mediterranean Sea is called maquis (Figure 321), in Chile it is known as matorral, in South Africa, fynbos, and in Australia, mallee (kwongan is used in western Australia). The prolonged summer drought and the dense growth make the chaparral-type shrublands very susceptible to re, and re is a prominent ecological factor in Mediterraneantype vegetation, both at present and in the past before the appearance of humans. But what was once a natural disturbance is today often seen as a catastrophe. The main problem is the intrusion of cities and towns into areas of highly ammable natural vegetation. This problem is particularly severe in southern California, where homes extend far up into the chaparral and the interval between res is long. Turnover in homeownership is high, and while a disastrous re will stimulate much discussion at the time, the talk slowly ebbs, the matter remains unresolved, and a new generation of homeowners experiences the next major conagration. As is true of the deserts of the world, each area of Mediterranean climate is geographically isolated, and each has its own distinctive assemblage of plants and animals. The degree of ecological convergence is high, however. Seasonal drought enhances the importance of edaphic (soilrelated) and biotic variation, and small differences in precipitation often have profound effects on the vegetation and animal life present in the area. Hence, these areas often have high proportions of extremely local species of plants and animals, many of them now in great danger of extinction. In their modern form, these areas have already been profoundly changed by people. Much of their vegetation is now in a very different condition from that occurring before people occupied the areas with their grazing animalsfor

example, todays areas might be inhabited by more shrubs and fewer trees than formerly, or by more spiny and poisonous plants.

The Northernmost Forests Taiga and Boreal Forest

Moving poleward from the mixed coniferous forest, the seasonal contrast becomes more pronounced, with very short and extremely cold days in winter and a short growing season of long days. In these conditions, trees still dominate, because during the relatively brief summer, growth conditions are favorable, and evergreen conifers are abundant. But the shortness of the growing season makes the energetics of the deciduous habit less favorable. It is evidently more efcient, during the short season, to retain an evergreen needle that can begin photosynthesis as soon as conditions allow rather than lose valuable time growing leaves from buds. Deciduous trees do occur in the most northern forests, but they often are restricted to particular situations, such as the margins of streams where frost does not penetrate so deeply into the ground. And a deciduous conifer, larch (Larix spp.), is abundant in some taiga areas. These northern coniferous forests are often referred to by the Russian name taiga; in North America they are also called the boreal forest. In both cases, these forests are characterized by a persistent cover of snow in the winter. In the southern reaches of the taiga, the trees are taller and more luxuriant, often reaching 75 meters or more in height (Figure 3224). In its main, northern area, however, the trees are shorter, and thousands of square kilometers are covered by this uniform forest, with relatively few species of plants and animals (Figure 3225). Taiga extends over much of Russia, Scandinavia, and northern North America (Figure 322).

3222

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Global Ecology

3224 Pacic coast rainforest Douglas rs (Pseudotsuga menziesii) growing on the Olympic Peninsula in Washington State. Epiphytic mosses, liverworts, Selaginella, and lichens often grow luxuriantly on the trees.

Taiga occurs in the interior of large continental masses at the appropriate latitudes. In such regions, extreme temperatures range from 50 to 35C. Taiga is anked on the south by montane forests (as in western North America), deciduous forests, savannas, or grassland, depending on the

amount of precipitation in the region. Because continental masses do not occur at the appropriate latitudes in the Southern Hemisphere, taiga is absent there. Due to the inuence of the prevailing westerlies blowing over relatively warm ocean currents between 40 and 50 north latitude, the western portions of North America and Eurasia are characterized by milder climates than their eastern portions. Consequently, taiga is found somewhat farther north toward the Pacic coast than it is along the Atlantic coast in North America, and the same is true of the individual distributions of many kinds of plants and animals that inhabit the biome. The northern limits of taiga are ultimately determined by the severity of the Arctic climate, reaching a limit where the maximum monthly temperature is approximately 10C. In the extensive northern reaches of the taiga, most of the precipitation falls in the summer; the cold winter air in these regions has a very low moisture content. The annual total precipitation usually amounts to less than 30 centimeters. The rate of evaporation is low, however, and lakes, bogs, and marshes are common (Figure 3225). More than three-quarters of the northern reaches of the taiga is underlain by permanent ice, or permafrost, usually within less than a meter of the surface; the permafrost tends to trap surface water and form lakes. Fires are common in the taiga, and they result in generally warmer, more productive sites for at least 10 to 20 years afterward, due to the local melting of the permafrost. In general, taiga soils are highly acidic, very low in nutrients, and poorly suited for agriculture. Species of a few genera of trees are common in the northern taiga, including spruce (Picea), larch (Larix), r (Abies), and poplar (Populus). Among the more common shrubs are dewberries (Rubus), Labrador tea (Rhododendron), willows (Salix), birches (Betula), and alders (Alnus). Pines (Pinus) can be common, usually in drier areas. The members of all these genera of trees and shrubs are ectomycorrhizal (Figure 291), and they occur in dense stands con-

3225 Northern taiga The northern taiga, which covers hundreds of thousands of square kilometers in the cooler part of the north temperate zone, is dominated by white spruce (Picea glauca) and larch (Larix). This photograph was taken in northern Manitoba, Canada. In the more northern part of their range, the trees are smaller than seen here.

Arctic Tundra

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sisting of only one or a very few species. Perennial herbs are common, and mosses and lichens are especially prevalent, often forming luxuriant masses. Annual plants, except in areas where there is human disturbance, are essentially absent. At its northern limits, taiga grades unevenly into tundra. In both these biomes, the days are long during the relatively short growing season; north of the Arctic Circle, the sun does not set during at least part of the summer. Because of the seasonally abundant light and favorable temperatures, cool-season cultivated plants, such as cabbages (Brassica oleracea var. capitata), may grow rapidly in cleared areas in the taiga, attaining large sizes in a remarkably short period of time. Yet the infertile, highly leached soils of the taiga do not allow most forms of agriculture. Coniferous forests extend far to the south of taiga, along the Pacic coast of North America. These magnicent evergreen coniferous forests (Figure 322) occur where there is a pronounced summer drought, but high and persistent rainfall during the cooler seasons. Because photosynthesis is limited by lack of moisture during the warm season, deciduous trees are at a disadvantage and are usually found only along stream banks. The evergreen conifers, however, can synthesize carbohydrates all year round and, because of their massive size, can store water and nutrients for use during the dry season. Their thick barks and high crowns protect them from all but the most intense res characteristic of the region.

Arctic Tundra
The Arctic tundra is a treeless biome that extends to the farthest northern limits of plant growth (Figure 3226). It occupies an enormous area: fully one-fth of the Earths

land surface (Figure 322). The majority of tundra can be found in the Arctic, mostly above the Arctic Circle, although it extends farther south along the eastern sides of the continents than along the western sides. The Arctic tundra essentially constitutes one huge band across Eurasia and North America, with alpine tundra, more closely related to the adjacent mountain forests, extending southward in the mountains (Figure 3222; see also Figure 325). Some species of plants that occur in the Arctic tundra have wide circumpolar ranges. Toward the poles, the winter season lengthens relative to that of summer. Although daylength in summer increases north of the Arctic Circle the sun does not set for days this does not compensate for the long period of little or no sun. This portion of the Earth operates at a strong solar energy decit, radiating more energy back to space than is received in direct sunlight. Frosts can occur throughout the year, and winters are bitterly cold, with high winds that will desiccate any plant that rises above the snow level. At low temperatures, blowing snow acts as an abrasive that will ruthlessly attack any exposed tissues. The limited supply of solar energy is insufcient to thaw more than a shallow layer of soil. The result is permafrost permanently frozen soil. The annual freezing and thawing of the surface has many effects unknown in milder climates, such as landscapes broken into polygonal blocks ranging from 3 to 30 meters across, and small hills, or pingos, up to 50 meters high, formed with a core of ice. The soils are acidic to neutral, low in nutrients, and generally poor for agriculture. Even though precipitation is usually less than 25 centimeters per year, much of it is held near the surface by underlying permafrost, and the ground is usually wet.

(a)

(b)

3226 Arctic tundra (a) Arctic wet coastal tundra near Prudhoe Bay, Alaska, in late summer. The reddish brown plants are the grass Arctophila fulva, the green ones are the sedge Carex aquatilis. Note that the water table is above the surface because of the presence of permafrost; such conditions are characteristic of the Arctic tundra.

(b) Arctic tundra at Barrow, Alaska. A clone of the sedge Eriophrum angustifolium is growing out into a solid patch of another species of the same genus, Eriophrum scheuchzeri. Vegetative reproduction, like that in E. angustifolium illustrated here, is characteristic of many tundra plants.

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CHAPTER 32

Global Ecology

Fixed nitrogen is generally in very short supply, and there are only a few species of legumes and other plants with symbiotic bacteria that x atmospheric nitrogen. The evaporation rate is low because of the relatively high moisture content of the air and low temperatures. In contrast, some tundra areas are so dry as to constitute true polar deserts. Most of the land north of 75 north latitude is a desert or semidesert, with few plants taller than 5 centimeters. This is a result of the very low precipitation in both summer and winter and the very cold winter temperatures. For plant growth to occur at all, the mean temperature must be above freezing for at least one month of the year. The growing season (the time from one killing frost to the next) in many areas of Arctic tundra is less than two months. Several genera of low shrubs, including birch (Betula), willow (Salix), blueberry (Vaccinium), and Labrador tea (Rhododendron), are common in the tundra, and there are a number of genera of perennial herbs but only one native annual species (Koenigia islandica). Many of the plants that grow in the Arctic tundra, including a number of the grasses and sedges, are evergreen, thus able to initiate photosynthesis soon after adequate light, moisture, and temperature become available. Plant height is controlled primarily by snow depth in winter; large woody plants are absent because of their relatively high energy allocation to unproductive stem tissue, which cannot be supported given the short, cool growing season. A number of tundra plants have relatively large, showy owers, the production of which costs the plants considerable quantities of energy. Such owers hold energy-rich rewards for their pollinatorsnecessary rewards, given the low temperatures that prevail at high latitudes. Vegetative propagation is characteristic of many of the perennials, and this may be correlated with the uncertainties of setting seed during the brief Arctic summer. Much of the biomass of tundra plantsfrom half to as much as 98 percentis underground, consisting not only of roots but also of rhizomes and other kinds of underground stems. North of the Arctic tundra is ice desert, where physical conditions are even more extreme and vegetation is absent. Ice desert is characteristic of the interior of Greenland; Svalbard, a small group of islands off Norway; and Novaya Zemlya, two islands off the north coast of Siberia. Much of Antarctica, not mapped in Figure 322, is also covered by ice.

Tropical Rainforests Have a Great Diversity of Species, with Few Individuals per Species
Tropical rainforest, in which neither water nor low temperature is a limiting factor, is by far the richest biome in terms of number of species. The trees are evergreen and characterized by medium-sized leathery leaves. A poorly developed layer of herbs grows on the forest oor, but there are many vines and epiphytes at higher levels. Tropical soils are often acidic and very poor in nutrients; such soils lose their fertility rapidly when the forest is cleared.

Savannas and Deciduous Tropical Forests Occur Where Rainfall Is Seasonal

Mostly tropical and subtropical communities that are characterized by a seasonal drought are termed savannas, subtropical mixed forests, monsoon forests, tropical mixed forests, and southern woodland and scrub. The trees and shrubs of these communities are wholly or partly deciduous, shedding their leaves during times of drought. Herbaceous perennials are common. Savannas also occur between the prairies and temperate deciduous forests and between the prairies and the taiga in North America. Subtropical mixed forests cover most of Florida and the Coastal Plain of the southeastern United States. In these forests, pines and other evergreen trees grow intermixed with deciduous trees.

Desert Plants Are Adapted to Low Precipitation and Extremes of Temperature

Away from the equator, tropical and subtropical communities grade into deserts and semideserts, which are characterized by low precipitation and often by high daytime temperatures during at least part of the year. Succulent plants and annual herbs are common in deserts.

Grasslands Occur Where Precipitation and Fire Interact to Support Grasses but Not Trees
Grasslands, which intergrade with savannas, deserts, and temperate forests, are characterized by a general lack of trees, except along streams. The most productive soils for temperate agriculture are grassland soils.

Temperate Deciduous Forests Are Made up of LeafShedding Trees and Many Types of Perennial Herbs
In the temperate deciduous forests, most of the trees lose their leaves during the cold (usually snowy) winters, when moisture may be unavailable for growth. Many genera are common to the temperate deciduous forests of eastern North America and eastern Asia. Temperate deciduous forests are bordered by temperate mixed and coniferous forests to the north, in which conifers play an important role.

SUMMARY
Biomes Are Terrestrial Ecosystems Characterized by Distinctive Vegetation
The distribution of biomes is a result of complex interactions among the distribution of heat from the sun, air-circulation patterns, and geologic features. These factors cause wide differences in temperature and precipitation from place to place and season to season. In addition to climate, differences in the surfaces of the continents, such as soil composition and altitude, affect the kinds of plant and animal life found in the various biomes on Earth.

Mediterranean Scrub Is Characterized by Evergreen, Drought-Resistant Shrubs or Trees That Form Thickets
Distinctive scrub communities, called chaparral in North America and maquis in the Mediterranean region, have evolved in the ve widely separated areas of the world with a

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SUMMARY TABLE Some Characteristics of the Earths Principal Biomes

BIOME Rainforests T E M P E R AT U R E A N D P R E C I P I TAT I O N High temperature and high rainfall year round. CHARACTERISTIC PLANTS Broad-leaved evergreen trees, epiphytes, and lianas. Grasslands with scattered broad-leaved deciduous or evergreen trees or shrubs. M I S C E L L A N E O U S F E AT U R E S The biome with the greatest diversity of species. Infertile soils. Periodic burning is an important aspect.

Savannas and deciduous tropical forests Deserts

High temperature and seasonal drought.

Precipitation generally very low except for occasional peaks; maximum temperature varies with the type of desert. Moderately low precipitation; cold winters and warm summers. Moderate precipitation evenly distributed; cool winters and warm summers. Moderately low precipitation and moderately cold winters.

Succulents such as cacti; annual herbs.

Adaptations include small leaves, thick cuticles, and photosynthetic rates with high maximum temperatures. Heavily exploited for agriculture.

Grasslands

Perennial bunchgrasses and sod-forming grasses. Deciduous trees and many perennial herbs.

Temperate deciduous forests Temperate mixed and coniferous forests Mediterranean scrub Taiga

The dominant herbaceous plants vary with the seasons. Occur as a transition zone north of the deciduous forest. Also found in areas with nutrient-poor soils or with less seasonal environments. Called chaparral in California and maquis around the Mediterranean Sea. Soils are highly acidic and very low in nutrients. Permafrost may be present.

Mixtures of deciduous trees and conifers.

Cool, moist winters and hot, dry summers.

Evergreen or summer-deciduous, droughtresistant trees and shrubs in dense thickets. Forest of evergreen trees.

Moderately low precipitation and cold winters, although in the Pacic Northwest the winters are very wet. Very low precipitation in both summer and winter; very cold winters.

Arctic tundra

Low shrubs, grasses, sedges, and lichens.

Permafrost present throughout. Much of the biomass is underground.

Mediterranean climatea dry summer and a cool, rainy winter growing season. Such communities occur in western North and South America, around the Mediterranean, in the Cape Region of South Africa, and in southwestern Australia.

ration, tundra and taiga soils are relatively moist and highly leached of nutrients.

The Taiga Is Characterized by Forests of Evergreen Trees

The taiga is a vast northern coniferous forest that extends in unbroken bands across Eurasia and North America and down the Pacic coast to northern California. In its southern reaches, the taiga is dominated by tall trees with a lush growth of bryophytes and lichens; northward, it consists of vast monotonous stretches of forest with very few tree species.

QUESTIONS
1. Describe the inuence of latitude and altitude on the distribution of organisms on Earth. 2. Describe the effect of mountains on local precipitation. 3. Compare tropical and temperate forests in terms of the numbers of species found in each, and in the size and appearance of the trees. 4. Explain why annual plants are better represented, both in number and in kind, in the deserts and semiarid regions of the world than anywhere else. 5. Compare the relative amounts of nutrients found in forest and grassland soils. 6. How are the evergreen conifers of the Pacic Northwest of the United States and Canada adapted to the winter-wet/ summer-dry environment of that region? 7. What are the principal differences between taiga and tundra? What role does permafrost play in these biomes?

The Arctic Tundra Has Low-Lying Shrubs and Grasses but No Trees
North of the taiga is the tundra, a treeless region that also extends around the Northern Hemisphere, mostly above the Arctic Circle, in a band that is broken only by bodies of water. Both the northern reaches of the taiga and all of the tundra are underlain by permafrost. Because of this permafrost, and especially because of the low rates of evapotranspi-