THE ECOLOGICAL PLANET:

AN INTRODUCTION TO EARTHS MAJOR ECOSYSTEMS

COURSE GUIDE

Professor John Kricher

WHEATON COLLEGE

The Ecological Planet:

An Introduction to Earths Major Ecosystems Professor John Kricher
Wheaton College

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The Ecological Planet: An Introduction to Earths Major Ecosystems Professor John Kricher

Executive Producer John J. Alexander Executive Editor Donna F. Carnahan RECORDING Producer - David Markowitz Director - Matthew Cavnar COURSE GUIDE Editor - James Gallagher Design - Edward White

Lecture content 2008 by John Kricher Course guide 2008 by Recorded Books, LLC

72008 by Recorded Books, LLC

#UT119 ISBN: 978-1-4361-0596-5
All beliefs and opinions expressed in this audio/video program and accompanying course guide are those of the author and not of Recorded Books, LLC, or its employees.

Course Syllabus The Ecological Planet: An Introduction to Earths Major Ecosystems

About Your Professor/Introduction ...............................................................................4 Lecture 1 Lecture 2 Lecture 3 Lecture 4 Lecture 5 Lecture 6 Lecture 7 Lecture 8 Lecture 9 Lecture 10 Lecture 11 Lecture 12 Lecture 13 Lecture 14 Ecology and the Big Picture ..................................................................6 Earth and the Goldilocks Effect ...........................................................10 Distribution of Global Ecosystems.......................................................15 Climate and Ecology............................................................................19 Biogeography and Evolution................................................................23 Polar Ecosystems and Tundra ............................................................28 Boreal Forest .......................................................................................33 Temperate Deciduous Forest ..............................................................39 Grassland and Savanna ......................................................................44 Desert ..................................................................................................50 Tropical Rain Forest ............................................................................55 Marine Ecosystems .............................................................................61 Unique Coastal Ecosystems................................................................67 Current Issues in Global Ecology ........................................................75

Course Materials ........................................................................................................80

About Your Professor John Kricher

John Kricher is a professor of biology at Wheaton College, Norton, Massachusetts. His books include Galapagos: A Natural History, A Neotropical Companion, three ecology field guides (Eastern Forests; Rocky Mountain and Southwestern Forests; and California and Pacific Northwest Forests), and First Guide to Dinosaurs. John is a fellow in the American Ornithologists Union and past-president of both the Association of Field Ornithologists and Wilson Ornithological Society. He resides with his wife Martha Vaughan on Cape Cod.

Photo courtesy of John Kricher

Introduction
Earth is the only known ecological planet, a place with life. But life is diverse: plants, animals, microbes, millions of species, many interacting in complex ways and all of them influenced in myriad ways by the environments in which they are found. Ecology is the study of organisms as they relate to their environments, the scientific study of natural history. But what does that really mean? It means that because of the many physical, chemical, and atmospheric characteristics of Earth, diverse forms of life have evolved throughout the history of the planet, life that is shaped and reshaped by both physical and biotic forces. Life is forced to evolve because of Earths diverse conditions and the fact that, over time, conditions change. Ecology is the search for broad general patterns evident in the distribution and abundance of life. Further, ecology attempts to explain such patterns with empirical reasoning. Ecology, a word taken from the Greek oikos, meaning home, is the same root as the word economics. Like economics it studies complex systems, in this case ecosystems. Ecology is the broadest level of organization in biology, the discipline that deals with interactions among populations, ecological communities, and ecosystems. Earth is a planet with a complex climate, one that varies dramatically from equator to poles. As a consequence, life-forms adapted to polar regions are not well suited to equatorial areas and thus climate alone forces a high diversity to evolve among Earths numerous and variable organisms. Polar bears, for all their magnificent adaptations, are not adapted to survive in tropical conditions and the diverse array of trees comprising equatorial tropical rain forests would not long survive at higher latitudes. This course explores exactly why Earth supports life. It examines the various reasons why a small planet of about eight thousand miles diameter orbiting an average-sized star is uniquely suited to have a biosphere, a thin layer of living 4

matter surrounding its surface. Life abounds on both land and oceans but takes many different forms, mostly because climate is so variable. Beginning at the polar regions, the cold windswept Arctic and Antarctic, Professor Kricher will devote lectures to each of the Earths major ecosystems, called biomes. These include the various major forest types, the northern coniferous (or boreal) forests that surround the higher latitudes like a vast belt of spruce and fir trees, the immense temperate deciduous forests of North America, Europe, and Asia, and the tropical rain forest, the ecosystem with more species than any other. When moisture is insufficient to support forest, other kinds of biomes occur. These include grassland, called prairie in North America, as well as savanna, a combination of grassland and scattered trees that typifies much of East Africa. The driest of all biomes is desert, which is always water stressed but may be hot or cold depending on where it is located. The largest ecosystems on Earth are marine, the oceans and coastal areas where land meets sea. The ocean realm has ecosystems that differ by depth. The open ocean or pelagic zone is where the food webs of the sea begin, where billions of tiny plants, the phytoplankton, capture some of the Suns energy and thus support all of the other creatures of the seas. But most of the ocean below about six hundred feet is cold and dark, and the benthic zone, or sea bottom, is lit only by the bioluminescence of the life-forms that inhabit it. Coastal ecosystems such as salt marsh, mangrove swamp, and coral reefs are among the most ecologically valuable as they are highly productive. At the same time, they are among the most threatened. The final lecture in the course will focus on how ecology has matured into a pragmatic science that is as essential as economics for making sound judgments about how best to steward the ecological planet.

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Lecture 1: Ecology and the Big Picture

The Suggested Reading for this lecture is John Krichers A Field Guide to Eastern Forests (chapters 1 and 2).

Wildflowers at the edge of a wooded field.

cology is the scientific study of how life-forms interact and coexist. It is concerned not only with how living things adapt to each other, but also how they interact with and adapt to the nonliving components of the environment. Ecology is often defined simply as the study of organisms as they relate to their environments. Another definition is that ecology is the study of how various factors in the environment affect the distribution and abundance of organisms. Those factors are divided into two broad categories, abiotic and biotic. Abiotic factors are such things as temperature, precipitation, phosphorus, oxygen, salinity, and fire. Biotic factors are interactions such as predation, parasitism and pathogens, competition for resources, and mutually beneficial interactions. Ecology is taken from the Greek word oikos, meaning home, the same root from which the word economics is derived. Both disciplines, ecology and economics, study how things relate within complex systems. Economists study the flow of money through economic systems as well as how materials move in the form of goods and services. Ecologists focus on energy flow through ecosystems and study how atoms combine and recycle as life uses energy and material to maintain itself. The word ecology was coined in 1866 by Ernst Haeckel. He was attempting to form a word describing the economy of nature that Charles Darwin described in his monumental book On the Origin of Species (1859). 6

LECTURE ONE

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Nothing in ecology makes sense without environmental context. For instance, you can watch a kangaroo in a zoo cage but you will not be observing ecology. Kangaroos live in Australia, in open grassland and savanna habitats. To really learn about the ecology of a kangaroo, it is necessary to study the creature in the wild, in the Australian outback where groups of kangaroos eat, groom, defend their territories, mate, and reproduce. So it is with all other animals and plants as well. Ecology looks at living things in the context of the environment for which each is adapted. Ecology is often described simply as the scientific study of natural history, its roots reaching back to the early writings of the ancient Greek scholars who were curious about such questions as why fish look as they do, have scales, and swim in water, while birds look quite different, have feathers, and fly in air. But ecology is far more than descriptive natural history. It encompasses broader, more deeply penetrating questions. The living world does not yield its secrets easily. Ecology is the science that attempts to uncover patterns in nature and then to discover causal explanations for such patterns, explanations that account for the distribution and abundance of plants, animals, and microbes, and predict how various factors effect changes in such patterns. Ecologists study the big picture within the biological sciences: how an open field of grasses and wildflowers can, with time, develop into a closed, shadowy forest; why large predatory animals such as jaguars or great white sharks are so much rarer than their prey species; why the loss or gain of but a single species in a habitat can radically affect and alter that habitat while other species come or go with little, if any, discernible effects; why some species are abundant and broadly distributed and others are very local and rare; why some regions of the Earth are covered by rain forest while others are deserts; how essential minerals such as phosphorus and nitrogen move from the nonliving to the living components of what we call the environment. Ecologists today are concerned with such major global questions as the potential for climate alteration, the complex effects caused by pollutants, the increasing prevalence of invasive species, and the decline of global biodiversity. At the base of ecological study is the organism. Ecologists study organisms in the context of both their present environment and their evolutionary histories, meaning how they are adapted to survive within their environments. Organisms of the same species in nature are grouped into organizational units called populations; thus ecologists speak of the grizzly bear population at Denali National Park in Alaska or the right whale population in the Gulf of Maine in the Atlantic Ocean. When the focus is on questions pertaining to whole populations, the term population biology is often used. But no natural environment consists of but a single kind, a single species of organism. Thus various populations, called an ecological community, coexist within the same locality. Grizzly bears, along with caribou, Dalls sheep, and arctic ground squirrels, are each part of the arctic tundra animal community in Denali. Right whales, along with various oceanic birds such as gannets and greater shearwaters, plus other whale species such as humpback and minke whales, along with numerous fish species, oceanic invertebrates, and tiny plants called phytoplankton, are all part of the Atlantic pelagic (open sea) community. 7

Ecological communities, by necessity, interact with the nonliving, or abiotic environment, the air, water, and substrate. The combination of the living, the biotic, with the abiotic components of any habitat forms the most encompassing level of organization in the life sciences. The interactive association between a community of organisms and their physical and chemical environment is called an ecosystem. When ecology was in its infancy it was described as physiology (already an established laboratory science) applied to the environment, an attempt to explain how organisms function in an adaptive manner that permits some to survive in freezing cold and others to thrive in blazing heat. The ecosystem concept, first clearly articulated in 1935, was formulated to show that the interactions of multiple organisms with the physical environment in any given region forms a dynamic system worthy of its own study as a level of organization. Ecosystem boundaries are often fuzzy: a decaying log in a woodlot is an ecosystem, but on a larger scale, so is the woodlot in which the log is found, and on a larger scale still, so too is the regional forest of which the woodlot forms but a part. The largest ecosystem known is called the biosphere or ecosphere (the terms mean the same thing), the thin layer (perhaps twentyfive kilometers) of living matter that inhabits the crust and atmosphere of the Earth, thus far the only known area in the Universe to have an ecology, which brings us to the next lecture.

LECTURE ONE

Grizzly bear near a streambed in the Denali National Park, Alaska.

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FOR GREATER UNDERSTANDING

Questions
1. Is ecology the same thing as natural history study? How is ecology unique? 2. What are the differences among populations, communities, and ecosystems?

Suggested Reading
Kricher, John. A Field Guide to Eastern Forests. Boston: Houghton Mifflin Company, 1998.

Other Books of Interest

Attenborough, David. The Living Planet. Boston: Little, Brown, 1984. Bates, Marston. The Nature of Natural History. Princeton: Princeton University Press, 1990. Wilson, Edward O. The Diversity of Life. New York: W.W. Norton, 1992.

Lecture 2: Earth and the Goldilocks Effect

The Suggested Reading for this lecture is Peter D. Ward and Donald Brownlees Rare Earth.

nce upon a time a little girl named Goldilocks was very selective about eating porridge. It had to be just right, not too hot and not too cold. Astrobiologists like to compare Earth with the famous porridge of Goldilocks. Its characteristics, outlined in this lecture, make it just right to support life. Earth is 4.5 billion years old, nearly as old as the Sun. Earth formed along with the rest of the Solar System, the other planets, comets, and asteroids. The universe itself began about 13.8 billion years ago with an event physicists call a singularity, more popularly called the Big Bang. Earth is the third planet from the Sun, about 93 million miles away. The Suns characteristics, particularly its age and temperature, make Earth suitable for liquid water, the key to the origin of life. It is difficult to form scenarios for lifes evolution or for its persistence without liquid water because cells, which make up organisms, are mostly composed of water and all biochemical reactions occur in aqueous solution. While water may have once flowed on the planet Mars, it seems to be absent now and Mars appears to be a lifeless desert. Evidence from a variety of sources suggests that Jupiters moon Europa may contain a liquid ocean, perhaps salty, beneath a dense overtopping layer of ice evident on the moons surface. What might be crawling or swimming around in such an ocean? Earths only natural satellite, the Moon (the name is derived from the Greek for menstrual cycle), is unusual as planetary satellites go and its size and

LECTURE TWO

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presence may contribute to why life exists on Earth. It is proportionally large relative to its planet, indeed, proportionally the second largest moon in the Solar System. Earth and the Moon are unusual in that astronomers characterize them as virtually a biplanetary system, given the large proportional size of the Moon to the Earth. Thus, given Newtons insights regarding gravity, the Moon has a strong effect on the Earth. What if Earth had no moon? Yes, there would be no moonlit nights, many romantic songs would not have been written, eclipses would not occur, and wolves would have nothing to howl at. But the consequences of moonlessness could be more ecologically profound. The Moon, which today is on average 238,860 miles (384,400 km) from Earth, was considerably closer to Earth when it formed, though exactly how close is a matter of conjecture. Today the Moon is becoming more distant from Earth, receding at about three centimeters annually. But the proximity of the Moon to the Earth, and the proportionally large size of the Moon to the Earth, means that throughout its existence, the Moon has exerted a strong gravitational effect on its planet. Most of us realize that Earths tidal cycles are caused mostly by the influence of the Moon. Given that life may have originated in conditions prevalent in tide pools and other coastal environments, the Moon may have indirectly contributed to the first appearance of life on the planet. What is generally less well known, and what may be more important, is that the Moon likely stabilized the tilt of the Earth in space, what astronomers call Earths obliquity. If the Earths obliquity had undergone numerous substantial changes, making the planet basically wobble unpredictably, Earths climate would have undergone far more frequent, severe alterations, possibly too severe to permit complex multicellular life to evolve. Our Moons gravitational calming effect on Earth may have been of utmost importance to its future inhabitants. The Moon, a lifeless place, may have helped make life more possible on its larger neighbor. Earth has a strong magnetic field (the Van Allen belts) and that too is important for sustaining life. Earth is constantly being bombarded by potentially harmful radiation from space, most of it from the Sun, but also cosmic rays from space. The Sun emits what is called the solar wind, and radiation of this sort could certainly be harmful to life, particularly large multicellular life-forms. But we exist happily along with elephants and redwood trees, so how are we protected from cosmic rays and the solar wind? The answer is that Earth generates a strong magnetic field, called the magnetosphere. Like flak jackets around the planet, the belts intercept cosmic rays from space and solar wind particles from the Sun, affording a magnetic blanket of protection to Earth. The huge planet Jupiter also helps protect Earth and thus helps Earth support life. Jupiter is immense in comparison not only with Earth but with virtually all other planets in the Solar System. Its huge mass means that Jupiter exerts a very strong gravitational attraction on things that pass reasonably close to it, things like asteroids, comets, and meteors. There are many objects in space that have trajectories that occasionally cross the orbital path of Earth. The last of the dinosaurs, as well as many other life-forms, were victims of one of these objects at the close of the Mesozoic Era. There have 11

been other impacts on Earth, some major, both before and since the one that ended the Mesozoic Era. But it could have been worse and, to the degree that it wasnt, we probably have Jupiter to thank. The presence of a large planet such as Jupiter likely reduces the number of possible collisions of Earth with potentially catastrophic space debris. Jupiter acts as a kind of cosmic vacuum cleaner, sweeping the inner Solar System of potentially dangerous objects. While it is true that previous devastating events on Earth have been followed by the evolution of new life-forms, if massive collisions were to be far more frequent, such calamities could prevent the formation of complex ecological communities. Earths ecology is strongly seasonal. We take the seasons pretty much for granted. Summer, fall, winter, spring: each is a part of our year. Even someone with the most casual understanding of the natural world knows that the rhythms of nature are closely attuned to seasonal cycles. Most ecological patterns on Earth are affected in some way by the existence of seasons. It is thus fair to conclude that seasonality has been an important determinant of the ecosystem and biodiversity patterns of the planet. So why does Earth have seasons? Earth is tilted 23 degrees and 27 minutes, or 23.45 in its orbit. Planets vary in their axial tilts. Axial tilt causes seasons. As Earth progresses around its orbit its tilt places the Northern Hemisphere toward the Sun for part of the orbit while, at the same time, the Southern Hemisphere points away from the Sun. When this situation occurs, more direct solar radiation falls on northern latitudes than southern latitudes. North of the equator, days are long, The tilt of the Earth. while south of the equator, days are short. It is the northern summer and the southern winter. During the northern winter the situation is, of course, exactly reversed. More light falls on Buenos Aires than on Minneapolis. Snow falls on Minneapolis. Seasonal changes of Earth cause latitudinal differences in day length, affect patterns of temperature and precipitation, and are, indeed, the major determinant of climate on the planet. These patterns are at the very essence of how ecosystems function, a force profoundly influencing how species adapt. Think about the annual migration of thousands of bird species around the globe as well as overland migrations of caribou, wildebeest, and so many others. Think about the amazing physiological changes that result in such events as fall 12

LECTURE TWO

James Kaler

color changes in leaves and hibernation in animals. Each of these elegant ecological patterns, and many more, occurs because of a 23.5 axial tilt. It is indeed a significant tilt. The position of Earths continents and the fact that their positions have changed throughout history also has a strong effect on Earths ecology and the evolution of organisms. Look at a map showing the positions of the oceans and the various landmasses. Look at South America and notice how the East Coast of South America looks almost as though it could fit rather well against the West Coast of Africa. If the globe shows relief, notice how the Himalayan Mountains form a rugged boundary at the border of India and Asia. Australia sits by itself, alone, an immense island continent in the southern Pacific Ocean. The occurrence of cataclysmic events such as earthquakes and volcanic eruptions informs us that Earths geology is active. Unlike Earths Moon, or planets such as Venus and Mars, Earth is dynamic, continuously rearranging its surface because of processes occurring in its interior. One of the great scientific discoveries of the twentieth century was that the crust of Earth itself is changeable, and that as it changes the continents that rest atop it actually move in relation to one another. The term for this process of change is plate tectonics. Continental movement caused by plate tectonics has mighty consequences for Earths ecology. Without such constant movement, Earths climatic history would have been very different and less variable. As the continents move about the surface of the planet they effect changes in ocean currents, air currents, and climate in general. As more coastline is exposed, coastal shallowwater species such as corals tend to proliferate. But when coastline is minimized, as when continents fuse, extinctions of such organisms seem common. Separation of the continents acts to geographically separate organisms, stimulating evolutionary change, allowing evolution to proceed along varied and different pathways from continent to continent, island to island. The isolation of Australia, for example, led to the diversification of marsupial mammals, making Australia unique as the land of marsupials. Likewise, eucalyptus trees of over six hundred species occur in Australia and nowhere else (except when transplanted). Part of the great biodiversity of mammals throughout the Cenozoic may be attributable to continental separation stimulating high levels of speciation among groups isolated from one another. The Earth has changed continuously and dramatically since its origin, an evolving planet in an evolving universe. The scale of change has varied both with time and in area, but any careful consideration of the history of life on Earth shows that change is the rule, not the exception. Life exists only because it has the potential to change as circumstances around it change. The tapestry through time that has been woven by life on Earth has generated many millions of species, a temporal montage of fascinating ecosystems, most of which have forever been relegated to history.

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FOR GREATER UNDERSTANDING

Questions
1. Why might life be rare in the solar system, confined essentially to Earth? 2. What major characteristics of Earth are essential for supporting life?

Suggested Reading
Ward, Peter D., and Donald Brownlee. Rare Earth. New York: Copernicus, 2000.

Other Books of Interest

Upgreen, Arthur. Many Skies: Alternative Histories of the Sun, Moon, Planets, and Stars. New Brunswick, NJ: Rutgers University Press, 2005. Ward, Peter D., and Donald Brownlee. The Life and Death of Planet Earth: How the New Science of Astrobiology Charts the Ultimate Fate of Our World. New York: Owl Books, 2004.

LECTURE TWO

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Lecture 3: Distribution of Global Ecosystems

The Suggested Reading for this lecture is Robert G. Baileys Ecoregions: The Ecosystem Geography of the Oceans and Continents.

uppose you want to see all ten thousand or so of the worlds extant bird species. What would you have to do? In a word, travel. You would look for parrots in the tropics and penguins in the Antarctic oceans. And those are simply two extremes. Earths distinct seasonality combined with ever-drifting continents over millions of years has resulted in widely separated landmasses where unique organisms have evolved to survive in varying climates. The result of this reality is that ecosystems vary dramatically from one point in the world to another. The goal of this lecture will be to provide an overview of ecosystem distribution and discuss how ecological processes vary from one region to another. We begin with the most obvious characteristic of ecosystem distribution, which is that Earth has two major kinds of ecosystems, aquatic and terrestrial. The seas are vast but they do not encompass the entire planet. This is no small point. Because continents and islands exist, the evolution of life on Earth evolved to be far more diverse than it would have been had life been confined entirely to the seas. Consider insects, for example. Though just under a million insect species have been described, it is estimated that between two and thirty million actually exist. Regarding beetles alone (order Coleoptera) there is thought to be over a million species. Insects have a strong influence on numerous terrestrial ecosystems, ranging from plague proportions in the case of outbreaks of locusts, to disease vectors, to essential plant pollinators. But there are no insects in the oceans. Insects are terrestrial. They evolved from early terrestrial arthropods and today, as in the past, represent the most abundant kind of organism on our planet. Without land, there would be no insects, and thus the most diverse kind of animal ever evolved would not exist.

Pictured at the right are five exotic members of the more than 30,000 species belonging to the family scarabaeidae (scarab beetles). These specimens, found in Latin America, are from the subfamily rutelinae (shining leaf chafers).

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Meet the Beetles

On the other hand, the most species-rich group of vertebrates is the group known as Osteichthyes, the bony fish. While some of these inhabit fresh water, and thus are in a way terrestrial, most are marine and there are some twenty thousand known species. The vastness of the oceans has served to permit much speciation, and fishes are an example. The next most diverse group of vertebrates, as mentioned above, are the ten thousand bird species. There are about forty-five hundred mammal species. In marine waters, the equivalent of insects is the Crustacea, the lobsters, crabs, and shrimp. But they are collectively far less diverse than insects, with about thirty thousand species, including some that inhabit fresh water. The 23.5 tilt of Earths axis, responsible for its seasonality, combined with the scattered positions of the continents, makes Earths climate quite variable from poles to equator. For example, Earth would be more temperate in general was it not for the fact that the continent of Antarctica is directly over the South Pole. When Antarctica drifted to that position about 36 million years ago, during the Tertiary period, it resulted in the world becoming more temperate, with areas of forest being gradually replaced by grassland and savanna. We can generalize by identifying three broad climatic regions: polar (Arctic and Antarctic), temperate, and tropical. Of course these regions are not sharply separated but tend to meld together where they overlap. An example is southern Florida, where it is best to describe the climate as subtropical. Physical conditions of climate, particularly temperature and patterns of precipitation, largely determine characteristics of terrestrial ecosystems. Ecologists have long understood that if you know the average temperature and average amount of annual precipitation, you get an accurate predictor of the kind of ecosystems that will dominate in that region. For example, a region that experiences less than ten inches of rainfall annually will be desert. If that region is warm, it will be hot desert, usually dominated by various succulent shrubs and cactuses. But if the region experiences a cold winter, including regular snowfall, then it will be cold desert, dominated by coldadapted shrubs such as sagebrush. The coldest of the desert-type ecosystems is so cold that we recognize it as unique, the tundra. True tundra is found only in polar regions and atop high mountains. Major terrestrial ecosystems are called biomes. Inside the Arctic Circle is the Arctic Tundra biome, a vast ecosystem of sedges and lichens and numerous tiny flowering plants. This is the land of caribou and lemmings. Tundra is not common in Antarctica, a land more cold and desolate but with a thriving marine ecosystem offshore.

LECTURE THREE

Caribou graze on the Alaskan tundra.

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The temperate zone begins beyond the Arctic and Antarctic circles. In the north it begins with the Boreal Forest biome that extends like a ring of spruce and fir trees around the globe, encompassing much of Canada, northern Europe, and Siberia. It is also found along mountain ranges, particularly in the American West. This biome is dominated by cone-bearing coniferous trees, most of which are evergreen. It is often called the sprucemoose biome. The next major biome is Deciduous Forest, comprising broad-leaved trees that typically drop their leaves before the cold of winter sets in. The autumnal changing colors of the leaves makes this biome one of the most aesthetic of natures ecosystems. Much of eastern North America as well as Europe and parts of China are dominated by deciduous forests of oaks, maples, sycamores, and elms, as well as numerous other trees and shrubs. The third major forest biome is tropical and it consists of a range of forest types from dry woodlands such as typify much of the Australian outback, to rich Tropical Rain Forest such as is found throughout the Amazon Basin, as well as parts of central Africa, Asia, Indonesia, and northeastern Australia. The Tropical Rain Forest is characterized by having more species of plants and animals per square hectare than any other of the worlds terrestrial biomes. Tropical Rain Forest is equatorial, and occurs between the Tropic of Cancer and Tropic of Capricorn, encompassing about a 47 latitudinal band. There are also biomes of grassland, savanna, and desert. These occur in areas with less precipitation or much more seasonal precipitation than is typical of forest biomes. Grassland once typified the American prairie, but much of the original prairie is now converted to agriculture. The steppes of Russia is another region dominated by mixed species of grasses. Savanna, which is well developed in much of Africa and Australia, consists of grassland with various amounts of trees scattered within. Deserts are found where moisture is least available and are usually dominated by shrubs. Hot deserts have succulents, such as cactuses in the Western Hemisphere and euphorbias (which closely resemble cactus) in places such as Africa. Fire-adapted shrubs and highly seasonal precipitation characterize a biome called Chaparral, which is found in much of central and southern California, Chile, and throughout the Mediterranean region. The biome concept is normally not applied to the worlds oceans. Rather, their life zones differ by depth and proximity to land. For example, the littoral zone is found near coasts, where water is shallow. The pelagic zone is the area of deep open ocean and the benthic zone is the life zone at the depths of the seas, on the bottom of the ocean floor. Why study biomes? Because they demonstrate how plants and animals are adapted for various kinds of climates. And because the amount of the Suns energy captured by vegetation differs remarkably from one biome to another. Since virtually all life on Earth depends on photosynthesis, such differences, and the reasons for them, are important. Climate and its effects on ecosystems will form the subject of the next lecture.

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FOR GREATER UNDERSTANDING

Questions
1. Why is the fact that Earth has continents important in the biodiversity of life on the planet? 2. What is a biome? Name the major terrestrial biomes.

Suggested Reading
Bailey, Robert G. Ecoregions: The Ecosystem Geography of the Oceans and Continents. New York: Springer, 1998.

Other Books of Interest

Aber, John D., and Jerry M. Melillo. Terrestrial Ecosystems. San Diego: Harcourt Academic Press, 2001. Barbour, Michael G., and William Dwight Billings. North American Terrestrial Vegetation. Cambridge: Cambridge University Press, 1988. Walter, Heinrich. Vegetation of the Earth: In Relation to Climate and the Eco-physiological Conditions. New York: Springer-Verlag, 1973.

LECTURE THREE

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Lecture 4: Climate and Ecology

The Suggested Reading for this lecture is Heinrich Walters Vegetation of the Earth: In Relation to Climate and the Eco-physiological Conditions.

hen I was a student in high school I was assigned The Grapes of Wrath, the classic novel authored by John Steinbeck. It is a poignant tale of the Joad family and their daunting trek across 1930s America, following Route 66 from dust-strickened Oklahoma to what they hoped would be the promised land of southern California. The Dust Bowl, a term describing the effects of a protracted and severe Midwestern drought, had a profound effect on the ecology of the region. But why did it happen? In 2004, a team of researchers using a complex atmospheric-land general circulation model determined that before the onset of the drought, sea-surface temperatures both in the Pacific and Atlantic oceans had deviated significantly from normal: the Atlantic warmer than it should have been, and the Pacific cooler. In particular, the colder-than-normal Pacific sea-surface temperature altered the air circulation in the upper troposphere of the atmosphere such that rainfall was suppressed throughout the Great Plains. The warmer-than-normal Atlantic sea-surface temperature created different conditions, but those resulted in blocking moisture from the Gulf of Mexico. The combination resulted in the severe and protracted drought. Records from tree-ring data suggest that the Great Plains has experienced similar droughts once or twice a century over the past four centuries. Such a frequency is sufficient to impose significant ecological stress on the ecosystems of the region. And that is why climate is the number one factor in determining ecosystem characteristics.

Dust storm approaching Stratford, Texas, April 18, 1935.

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NOAA, George E. Marsh Album

The linkage between climate and ecology has long been understood. In On the Origin of Species Charles Darwin wrote, Climate plays an important part in determining the average numbers of a species, and periodical seasons of extreme cold or drought, I believe to be the most effective of all checks. The checks to which Darwin referred were deaths of organisms that were not adapted to climatic shifts, the very essence of natural selection. There are dust storms on Mars and immensely high winds on Jupiter. It can thus be said that both of those planets have a climate. Earths moon, in contrast, has nothing happening above its rocky, dusty surface. It lacks a climate because it lacks an atmosphere. Climate results from the interaction of the atmospheric gases, whatever they might be, with solar radiation, landforms, and oceans. Because Earth has all of these things, climate is complex on Earth and strongly influences ecosystem composition. Because Earth is a sphere, solar radiation strikes the planet most directly at the equator and much more obliquely at the poles. Because of this fact, sunlight is more concentrated per unit area at the equator and thus that region of the world heats up more than regions to the north or south. Polar regions receive the least concentrated sunlight and thus are the coolest areas on the planet. The result of differential heating is the formation of large convection cells that characterize various latitudes. These cells converge in various places such as the Tropics of Cancer and Capricorn. Deserts are typical at latitudes 23.5 north and 23.5 south, because where the convection cells converge, they are essentially devoid of moisture, having lost it before convergence. The movement of convection cells as well as ocean currents is further complicated by the Earths rotation from west to east, a phenomenon known as Coriolis effect. A body on Earth moves most quickly at the equator, and as it moves away from the equator it is thus deflected east or west as a result of Earths rotation. If the object is moving north it will be deflected to the east and, if it is moving south, to the west. This fact, for example, creates the generally clockwise circulation of ocean currents in the Northern Hemisphere and the counter-clockwise circulation in the Southern Hemisphere. A look at the Earth from an ecological perspective will quickly reveal that there are several huge terrestrial areas on the planet that can be described in the broadest sense: forests, deserts, grasslands, for example. A somewhat closer look produces a finer resolution. Forests can be evergreen or deciduous, or a mixture of both kinds of trees, and trees may be predominantly broad-leaved species or needle-leaved species. Grasslands can be interspersed with trees, in which case they are called savannas, or be composed entirely of herbaceous plants, grasses, and various wildflowers. In the United States, such grassland ecosystems are called prairies. Deserts are typically arid, often extremely so, with organisms such as succulent cacti and longeared jackrabbits, whose very anatomies reflect obvious adaptations to environments where water is scarce and heat may be oppressive. Such largescale ecosystems are called biomes and each is characterized by distinctive species of plants and animals. But what causes biomes? The answer is climate. Biomes differ from one another primarily because of climatic variation from one place to another. Climate quite blindly selects for 20

LECTURE FOUR

differing arrays of adaptations from one climatic region to another. Two variables, temperature and precipitation, largely determine the type of biome. Amazingly, if one knows the average annual temperature and the average annual amount of precipitation, one can predict with great accuracy just what type of biome will be present. For example, if the mean annual precipitation is somewhere between 300 and 400 cm (118 to 157 inches) and the mean annual temperature is from about 20 to 30 C (68 to 86 F), the biome will be lush tropical rain forest. But if the mean annual precipitation is only, say, 100 cm, even if the mean annual temperature remains between 20 and 30 C, the biome will be savanna. Deserts are uniformly arid, receiving less than 50 cm precipitation annually, often much less. But some deserts are hot, as warm or warmer than rain forests while other deserts are cold, as cold as any place in the temperate zone. Arctic tundra, the realm of the caribou and snowy owl, is both dry and cold, so cold that permafrost (frozen soil) endures throughout the year. Tundra typically receives about 50 cm of precipitation per year and has a mean annual temperature of only 10 C. The importance of mean annual temperature and mean annual precipitation in determining which terrestrial biome will prevail in any one location was first revealed by Leslie R. Holdridge in 1947, who developed a system of ecological classification that he called life zones. The Holdridge system has passed the test of time and is still used today to classify ecosystems. Topological features such as mountains exert a strong influence on climate, creating what ecologists call rainshadows. Consider what happens in the Cascade Mountains of Oregon, for example, if you were to travel from west to east. You would begin in the cool, moist temperate rain forest of tall Douglasfirs and western redcedar trees, with a lush understory of bigleaf maple and various shrubs. In all likelihood, it would be raining. Ascending the mountain up its western slope, the trees would become shorter with increasing exposure to wind and cold. Spruces and firs would replace the temperate rain forest, some stunted and twisted by exposure. Once on the eastern slope of the mountain the predominant forest would be composed of ponderosa pines, and would be decidedly drier than the temperate rain forest. What is important is that the ponderosa pine forest occurs at exactly the elevation where, on the western slope of the mountain, there had been temperate rain forest. Finally, you descend into desert, the sagebrush desert of the Great Basin. The marked difference between the west and east slopes of the mountain is due to the fact that prevailing winds come from the Pacific Ocean, carrying evaporated moisture. As these winds encounter the tall Cascades, they are forced upward. As the air rises, it cools, condensing the moisture held within and producing rainfall in generous amounts. This precipitation supports the lush temperate rain forest. By the time the wind has crossed over the peaks of the Cascades, it is largely depleted of its moisture. It is therefore not possible to support temperate rain forest on the east slope, though there is sufficient moisture to support ponderosa pine forest. Finally, even this moisture is used and the air becomes so dry that only desert can occur. In our next lecture well see how separation of the continents, biogeography, also plays an essential role in global ecology. 21

FOR GREATER UNDERSTANDING

Questions
1. What characteristics of Earth affect climate? 2. Why does climate vary with latitude?

Suggested Reading
Walter, Heinrich. Vegetation of the Earth: In Relation to Climate and the Eco-physiological Conditions. New York: Springer-Verlag, 1973.

Other Books of Interest

Chapin, F. Stuart, Pamela A. Matson, and Harold A. Mooney. Principles of Terrestrial Ecology. New York: Springer, 2002. Odum, Eugene P., and Gary W. Barrett. Fundamentals of Ecology. 5th ed. Belmont, CA: Thomson, 2005.

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Lecture 5: Biogeography and Evolution

The Suggested Reading for this lecture is C. Barry Cox and Peter D. Moores Biogeography: An Ecological and Evolutionary Approach.

he famous ecologist G. Evelyn Hutchinson once authored a book with the intriguing title The Ecological Theatre and the Evolutionary Play. What he meant to convey was that an understanding of the principles of organic evolution is prerequisite to really comprehending the patterns evident in the worlds ecosystems. And it is essential to understanding that evolution results in different organisms inhabiting different continents. It is obvious that todays world consists of several immense continents separated to various degrees by oceans. In addition to continents, there are numerous islands, ranging from large ones such as Madagascar and Borneo to extremely tiny islands whose names few would recognize. Some islands such as South Georgia (near the Antarctic Circle) are rather isolated, while others, termed archipelagos, are in close proximity, such as the Hawaiian and Galapagos islands. Each continent and each island contains unique species, some of which migrated and colonized, and some of which evolved there. The study of biogeography compares various regions of Earth that are geographically distinct and examines the processes responsible for the differences. In some cases, such as the famous Darwins finches of the Galapagos Islands, organisms evolve and are unique to a single region. These sorts of species are termed endemic. In other cases, such as the cattle egret, which colonized the Americas from Africa, species are recognized as invasive. In either case, they become part of the ecology of the region. Consider a walk in a rain forest along the Napo River in eastern Ecuador compared with a similar walk in a rain forest in Queensland, Australia, half a world away. Initially, both rain forests would appear very similar. The climate would be hot and
Geospiza magnirostris (Large Ground Finch) An illustration of two large ground finches from the Charles and Chatham Islands, Galapagos Archipelago, as printed in John Goulds (18041881) The Zoology of the Voyage of H.M.S. Beagle, Part III: Birds (London: Smith, Elder & Co., 1841).

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muggy, possibly raining part or most of the day. The forest would be complex and dense, tall trees with wide buttresses at their bases. The crowns of the trees would widen into a canopy and the tree branches would be covered by numerous epiphytes, plants such as orchids that attach and grow on a tree without parasitizing it. Ones impression of both of these forests would be that they are much alike. But upon closer inspection it would soon be learned that the trees, mammals, insects, birds, indeed even the fungi are quite distinct in each region. The forests are both within the tropical rain forest biome, and their major structural characteristics evolved in response to the fact that both are situated in a climate that is hot and rainy throughout the year. The science of biogeography recognizes that the ecology and general appearance of widely separated ecosystems with similar climates will be similar even though the species have very different evolutionary histories. Tropical rain forest has been present since the time of the dinosaurs, the Jurassic period of the Mesozoic Era, some 150 million years ago, but those forests would bear little in common with those of today other than the heat, humidity, and rainfall. The geologic history of Earth reaches back some 4.5 billion years, but for our purposes, we may begin 248 million years ago at the beginning of the Mesozoic Era. The Earth at that time consisted of one supercontinent, Pangaea. But soon after the Mesozoic began, Pangaea began to break apart, initially into two supercontinents, Laurasia to the north and Gondwana to the south. Evolutionary patterns began to take different trajectories between Laurasia and Gondwana and have been doing so ever since. Australia has been long isolated from the rest of what was once part of the huge southern continent of Gondwana. Near the end of the Mesozoic Era, some 66 million years ago, Australia split from Antarctica (Australia and Antarctica had split from the other southern continents earlier), and was alone as a continent. Marsupial mammals such as kangaroos happened to be thriving on Australia, but placental mammals had not colonized the now-isolated continent. Thus marsupials were the sole mammalian occupants of the entire continent with millions of years to adapt to a wide range of habitats and lifestyles. Many species of marsupials proliferated on Australia. Placental mammals (other than a few bat species that colonized by flying to Australia) remained largely absent until humans arrived with their dogs only about fortyfive thousand years ago. The absence of placental mammals resulted in the evolution of marsupials such as sugar gliders that look compellingly like placental flying squirrels, a mole-like marsupial that looks and acts like a placental mole, marsupial mice, and various marsupial bandicoots

LECTURE FIVE

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A Sugar Glider (Petaurus breviceps) in flight at the Lone Pine Koala Sanctuary in Brisbane, Queensland, Australia.

that resemble placental rabbits and hares. This evolutionary phenomenon of two distantly related kinds of organisms evolving similar phenotypes is termed convergence or convergent evolution. Convergences are common throughout the worlds ecosystems. Indeed, convergence is so common that it often presents evolutionary biologists with difficulty. Is apparent similarity among certain organisms a result of sharing a recent common ancestor, or is it a case of ecological convergence, a result of natural selection molding very different genotypes into similar phenotypes? Genetic analyses such as DNA sequencing help answer such questions. Ratites are a group of large flightless birds: the ostrich, the rheas, the cassowaries, the emu, and the kiwis. The term ratite refers to the absence of a bony carina or keel on the sternum (breastbone) for the attachment of flight muscles. For many years it was debated as to whether or not the ratites are all genetically closely related or are an example of convergence. They do not fly, yet are widely separated, occurring on different southern continents. The argument for convergence seemed strong. The argument weakens, however, when one realizes that cassowaries are forest dwellers, unlike ostriches, emus, and rheas that all live in open savannas, and kiwis are unique in being nocturnal. In other words, the various ratites do not occupy such similar habitats that one would expect selection pressures to result in similar body forms. Recent work on DNA hybridization as well as DNA sequencing has shown convincingly that the ratite birds are not convergent but all stem from a common ancestor in ancient Gondwana. The ratite distributions offer convincing support for the reality of plate tectonics and resultant continental drift. A key issue in ecology is the question of what, exactly, is a species. If you travel throughout East and Central Africa you will surely see elephants. As large and unmistakable as these ponderous beasts are, it may surprise you to learn that, until quite recently, taxonomists were mistaken about exactly how many species of elephants currently inhabit Africa. It was assumed that there is but a single species, the African elephant, Loxodonta africana, when in fact there are two. Recent studies suggest strongly that there is a second species of African elephant, the African forest elephant, Loxodonta cyclotis. Compared with L. africana, now renamed the African savannah elephant, the forest elephant is smaller in body size, has rounded, not pointed ears, and straighter tusks. The results of the molecular analysis suggest that forest and savannah elephants are each as distinct from one another as lions are from tigers. Ecologists thus determine species by a combination of factors: anatomy, ecology, and genetics. The most widely used definition of a species is that it is a population that is reproductively isolated from other populations. Cichlids are a group of colorful freshwater fish commonly kept by fish enthusiasts in freshwater aquariums. These diverse bass-like fish are found in tropical waters around the world. There are about three hundred species in the Americas, including one that occurs as far north as Texas. But, by far, most species of cichlids are found in East Africa, clustered in three lakes that formed in the Great Rift Valley: Lake Victoria (less than four hundred species), Lake Tanganyika (two hundred species), and Lake Malawi (three hundred to five hundred species). Why are there so many species of African cichlids?

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The oldest of the three African lakes is Lake Tanganyika, which formed 9 to 12 million years ago, and the youngest is Lake Victoria, whose origin is between two hundred fifty thousand and seven hundred fifty thousand years ago. Considering only Lake Tanganyika, if one assumes it to be 12 million years old, the highest estimate of its age, and it has two hundred cichlid species, that is a speciation rate of about one species every sixty thousand years. However, researchers are in agreement that the amazing diversity of cichlids in the African rift lakes arose recently, within the past few million years, demonstrating that speciation can occur with impressive rapidity.

Two of the many varieties of cichlids found in Africa. At the left is a male Malawi cichlid found in Lake Malawi. At the right is a male Zaire Blue found in the waters of Lake Tanganyika in the Democratic Republic of the Congo (formerly Zaire), about 100 miles north.

Studies of DNA of the cichlids from Lake Victoria demonstrated that this cluster of species is genetically very closely related, showing that they all recently evolved from a common ancestor. What is stunning is that the total genetic variation among these four hundred species is less than that found throughout Homo sapiens, humans. In other words, the genetic distance between you and a stranger you might meet is greater than that between two separate species of cichlids from Lake Victoria. Given the estimated rates at which mutations occur, researchers believe the entire assemblage of cichlid species in Lake Victoria to have arisen within two hundred thousand years. Thats equivalent to about one species every five hundred years. The evolutionary process whereby one kind of organism, in this case an ancestral cichlid, evolves into numerous species, each adapted in such a way as to be uniquely specialized, is called adaptive radiation. There are numerous examples of adaptive radiation both in the fossil record and among extant animals and plants. Evolution is the process by which biodiversity is generated. Now it is time to see how it varies from one biome to another.

LECTURE FIVE

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FOR GREATER UNDERSTANDING

Questions
1. What is convergent evolution, and how does the Earths current continental distribution affect convergent evolution? 2. What criteria are used to distinguish a species?

Suggested Reading
Cox, C. Barry, and Peter D. Moore. Biogeography: An Ecological and Evolutionary Approach. Oxford: Blackwell, 1993.

Other Books of Interest

Adams, Douglas, and Mark Carwardine. Last Chance to See. New York: Harmony Books, 1990. Few, Roger. The Atlas of Wild Places. Washington, DC: Smithsonian Institution and Press, 1994. Kingdon, Jonathan. Island Africa. Princeton: Princeton University Press, 1989. Weiner, Jonathan. The Beak of the Finch. New York: Alfred A. Knopf, 1994.

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Lecture 6: Polar Ecosystems and Tundra

The Suggested Reading for this lecture is E.C. Pielous A Naturalists Guide to the Arctic.

uppose you are standing on the North Pole. And suppose at another time you are standing on the South Pole? What, besides the fact that the first location is 90 north and the second 90 south, is the difference between them? The answer is that at the North Pole you are standing on ice and at the South Pole you are on a continent, Antarctica. The North Pole is located on the Arctic Ocean, a sea of ice that because of global climate change is currently melting at its margins. Antarctica, once part of Gondwana, is the coldest continent on Earth, located as it is at the southern pole. Ice that surrounds this polar continent is also retreating. Though bitterly cold and in the dark or twilight for much of the year, both polar regions have in common that they are ecologically productive. In other words, they each capture a respectable amount of the Suns energy by photosynthesis. But neither does this terrestrially. The key to polar productivity is the oceanic food web during the summer growing season. Ocean currents bring up essential chemical nutrients to the surface when the sun shines for a few months on the surface waters. In both polar regions great whales feast on the bounty of invertebrates and fish. Many seabirds, including various penguin species at southern latitudes and murres and puffins at northern latitudes also partake of the seasonal bounty.

LECTURE SIX

Earths Polar Regions Two satellite composite images of the Earths polar regions are shown above. At left is the Antarctic continent; at right, a false-colored view of the Arctic Ocean. Both images were generated in 2006.

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On land it is a different story. The growing season is short and in extreme situations, as are found in interior Antarctica, virtually no plants exist. But there is one kind of ecosystem found not only in the high Arctic but also at high elevations on mountains that is permanent and diverse, adapted to the climatic extremes of wind and cold. This ecosystem is called tundra. Suppose you are standing in New Hampshire at an elevation of about fiftythree hundred feet on Mt. Washington. Or suppose you are visiting Rocky Mountain National Park in Colorado, on Trail Ridge Road at about the eleven thousand foot elevation. Or perhaps you are wandering along the gravel road in Deadhorse, Alaska, well inside the Arctic Circle. Or maybe you are just lost in northern Siberia. Regardless, in each case you would be on tundra. Tundra is the biome that is found in the coldest, driest places on the planet, where conditions are sufficiently severe so as to prevent the existence of trees, except in extreme dwarf form. Vegetation consists of lichens and mosses, grasses and wildflowers, scattered shrubs, and little else. Tundra is open and wind swept, in a climate so dry that even the protective blanket of winter snow, so common in the boreal forest, appears sparingly, if at all, on tundra. Organisms of this biome endure the worst weather conditions and enjoy the briefest growing season of anywhere on Earth. Latitudinally, the tundra biome, named Arctic tundra, occurs generally north of the Arctic Circle (at about 66 north latitude), though low tundra, where tundra mixes with boreal forest, begins at about 72 to 73 north latitude. Tundra exists as a band around the northernmost of the worlds continents and is generally similar in North America and Eurasia. Tundra can also be found above treeline at the summits of tall mountains. Montane tundra is usually called Alpine tundra, to distinguish it from latitudinally based tundra, though the two are much alike in appearance. Tundra is highly limited in Antarctica. Obviously tundra regions are cold much of the year, with a markedly short growing season. In the case of Arctic tundra, temperature is affected by the low angle of solar radiation. Even during the warmest time of the year, midsummer, temperatures reach only about 15 C. In the case of alpine tundra, exposure from elevation tends to keep the temperature low even though the angle of solar radiation may be much more direct than in the Arctic. Winds are a constant component of tundra climate. Wind speed is frequently high, especially in the Arctic, where it can routinely reach 65 km/hr. Snow is the usual form of precipitation for much of the year, but, perhaps surprisingly, in the Arctic snowfall amounts tend to be small. Precipitation is sufficiently sparse as to be within the range of that of a desert. Alpine tundra areas experience considerably more precipitation than Arctic areas. The most notable characteristic of Arctic soils is permafrost. This means that part of the soil is permanently frozen, preventing its use by plants. Roots cannot penetrate into permafrost. In summer, the upper levels of the soil thaw, providing sufficient moisture for the rapidly growing plants. Because of the thawing and refreezing of the upper layers of soil, there is an intrinsic instability in Arctic soils that results in the formation of hummocks, which are elevated areas, and polygons, geometric patterning that often spans wide areas. Polygons form from constant thawing and freezing of water in the upper layers 29

of soil, a process that eventually sorts the larger fragments of rocks from the smaller rubble, depositing the larger fragments on the borders of the polygon, which itself may be several meters in diameter. Alpine tundra soils do not usually experience permafrost. They are generally young, having been created with the rising of the mountain range. These soils typically are rocky with lots of gravel. Large rock fields, called fell-fields, are common, and form areas of habitation for such creatures as marmots and pikas. In general, the biodiversity of both alpine and Arctic tundra is markedly less than that found in other biomes, with the exception of certain hot deserts. Reduced biodiversity may result from the severity of physical conditions that prevail in tundra regions. In the high Arctic, it is profoundly cold, windy, and dark for most of the winter, conditions that are generally unsuitable for plants and animals. Nonetheless, the region is inhabited. Life-forms have evolved that succeed in the harsh climate. In Arctic tundra, the presence of high winds and a contracted growing season (only sixty to one hundred days) selects for prostrate plants, literally lying low to minimize evaporative water loss as well as to grow rapidly when conditions permit. Many perennial wildflowers, some called cushion plants for their shape, grow among the reindeer lichen (Cladonia spp.), dwarf willows, and dwarf birches. Sedges often dominate wet areas. These vegetation patterns are also generally true of alpine tundra. It is notable that numerous species of perennial wildflowers found in the Arctic also occur in alpine tundra. Animals abound during the summer growing season in the Arctic. Small rodents called lemmings experience repeated cycles of abundance and decline and, along with Arctic hares, serve as important food sources for wolves, arctic fox, grizzly bear, snowy owl, and rough-legged hawk. Many species of shorebirds, sandpipers, and plovers nest during the short Arctic growing season, along with loons, grebes, and various species of ducks and geese. Herds of caribou (called reindeer in Europe) graze on the tundra with

LECTURE SIX

Geometrically shaped hummocks are clearly visible in this image of the arctic tundra area in the Northwestern Territories, Canada.

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shaggy musk oxen. Many insects occur on tundra and any visitor to the Arctic tundra in summer soon becomes familiar with just how abundant mosquitoes can become. Tundra is fragile. This is because the growing season is so short that it requires many years for vegetation to recolonize and grow in areas subjected to disturbance. Unlike in the temperate zone or in the tropics, where vegetation development following disturbance is generally quite rapid, the opposite is true on tundra. The fragility of the tundra as an ecosystem was cause for concern when a pipeline was built to transport oil from the North Slope of Alaska, Prudhoe Bay, several hundred miles south to the port city of Valdez. The pipeline was carefully engineered to prevent damage to the landscape and, in various areas, it was either raised or buried to permit caribou herds to pass unimpeded during their annual migration. In general, the pipeline has been successful in not damaging delicate tundra ecology. Proposals to drill for oil within the Arctic Wildlife Refuge, an area entirely on tundra supported by permafrost, have met with skepticism because of the potential damage that could be inflicted on the ecosystem. In Russia and the Ukraine, where environmental protection laws are far less than those of North America, tundra areas have been subjected to damage far in excess than any experienced in North America. The Alaska oil pipeline is shown snaking through Atigun Pass (elevation 4,643 feet) in the Brooks mountain range. The Dawson Road is at the right Direct damage of the photo. from human use has been augmented by air and water pollution from numerous factories that deal with mining and oil production. Alpine tundra areas have been negatively affected by human use for activities such as skiing and hiking. One of the worst threats to alpine tundra is the increased use of off-road vehicles, which do significant damage to the fragile plants. Polar ecosystems illustrate how organisms evolve diverse adaptations to cope with extreme climate. Deserts, as will be clear in another lecture, are similar in that regard.

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FOR GREATER UNDERSTANDING

Questions
1. Why does tundra exist primarily in the Arctic and not in Antarctica? What is tundra? 2. Why are polar areas ecologically productive even with the short growing season?

Suggested Reading
Pielou, E.C. A Naturalists Guide to the Arctic. Chicago: University of Chicago Press, 1994.

Other Books of Interest

Miles, Hugh, and Mike Salisbury. Kingdom of the Ice Bear. Austin, TX: University of Texas Press, 1985. Sage, Bryan. The Arctic and Its Wildlife. New York: Facts on File, 1986. Shirihai, Hadoram. The Complete Guide to Antarctic Wildlife. Princeton: Princeton University Press, 2002.

LECTURE SIX

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Lecture 7: Boreal Forest

The Suggested Reading for this lecture is E.C. Pielous The World of Northern Evergreens.

The northern third of the boreal forest, where trees are stunted and shrub-like One of the many moose residents browsing in the woods at the Algonquin Provincial Park in central Ontario, Canada. due to the severity of climate, is traditionally called taiga (from a Russian word), but the word taiga is now often used to name the entire biome, thus boreal forest and taiga are commonly considered interchangeable. Much of the region of the northern coniferous forest has been subject to recent glaciation, resulting in poor soils and an abundance of lakes and bogs, created by the scraping action of the immense, heavy glaciers. A bog is different from a lake in that it has no inflow and outflow of water but is, instead, an isolated basin that eventually fills with peat, becoming increasingly acidic. In North America, the Boreal Forest begins roughly at the Canadian border, though it is also found in northern New England and northern Michigan, 33

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he Boreal, or Northern Coniferous Forest, is a vast biome that encircles the far northern latitudes, covering just over 10 percent of Earths terrestrial area. It is strongly identified with the wild, yodeling call of the common loon, which nests on the seemingly innumerable lakes that dot the boreal forest (thanks in large part to recent glaciation). It is a land of relatively low tree species richness, with millions upon millions of spruces, firs, and pines, intermingled with larches (which are deciduous conifers) and a few broad-leaved tree species such as aspens, birches, and willows. Often termed the spruce-moose biome, the largest of the worlds deer does, indeed, thrive throughout the biome, feeding on aquatic vegetation within the numerous bogs that interrupt the unbroken vastness of the forest of Christmas trees. Species such as moose, wolverine, lynx, goshawk, boreal owl, hawk owl, and red crossbill are among those found not only in the boreal forests of North America but also in Old World boreal forests as well. Plant species are not the same but are closely similar, with, for example, Norway spruce replacing white spruce in Europe.

Minnesota, and Wisconsin. A unique extension of boreal forest, the Appalachian Extension occurs at high elevations southward from Maine to the Great Smoky Mountains of North Carolina and south to Georgia. In the American West, the boreal forest intermingles with the coniferous forests found within the Rockies, Cascades, and Sierra Nevada, as well as with the temperate rain forest of the Pacific Northwest. For example, white spruce, abundant throughout the boreal forest, is replaced by closely related Engelmann spruce throughout the high elevations of the Rocky Mountains. These two species are so genetically similar that they occasionally hybridize. The growing season becomes increasingly shortened as one moves north in latitude through the boreal forest, ranging from about 120 days in southern regions, to as short as 90 days or less at treeline, where the forest gives way to arctic tundra. In general, mean annual temperature ranges from about 4 C to 5 C. In a few places, temperature extremes from the heat of summer to the cold of winter range as much as 100 C. In other words, summer could bring a high temperature of, say, 30 C, and winter a low of, say, 70 C. Winters are almost always long and cold, the average daily low temperature being about 15 C, with most precipitation falling as snow. In general, precipitation averages between about 40 and 50 cm annually and is fairly constant from month to month, though falling as rain in summer and snow in winter. In winter, the frozen ground prevents plants from taking up moisture from the soil, thus the trees must enter into winter dormancy. Northern tree species undergo a physiological process (that varies somewhat among species) called winter hardening, enabling them to withstand the subzero temperatures without Boreal landscape in northern Finland. experiencing damage to delicate tissues. The conical shape typical of spruces and firs acts to aid the trees in shedding a burden of snow, which could potentially accumulate until branches break from the weight of the snow. Soils are typically acidic, thin, and nutrient poor. The former word used to describe boreal soils was podzol, but that word has been replaced by spodosol in recent years. Spodosols are acidic due, in part, to the breakdown of needle leaves, which tends to add hydrogen ions to the upper soil layers. Boreal soils are well leached, the minerals deposited in a lower B horizon well below the surface. The upper soil horizon is usually a grayish color. Because the growing season is short, there is a significant buildup of leaf litter, making the ground soft underfoot. Bogs are particularly acidic due both to 34

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the influx of decomposing litter as well as the prevalence of sphagnum moss, which enhances the acidic nature of bogs. Some boreal soils have permafrost, where the soil remains frozen throughout the year. Throughout much of North America, two species of trees, white spruce and balsam fir, are the dominant upland species of the boreal forest. In wet areas, black spruce abounds along with various willows and tamarack (American larch). Areas subjected to recent fire support stands of aspens, birches, and/or jack pine. In the Eurasian boreal forest these species are replaced by others, but the appearance of the forest is essentially the same. Bogs, most created by glacial retreat, are generally abundant throughout much of the boreal forest. Bogs provide habitat for species of insectivorous plants such as sundews and pitcher plants, as well as many other specialized plant species, including sphagnum moss, the dominant bog species throughout most of the biome. In the most northern regions of boreal forest the trees are much reduced in stature due to the severity of the winter weather. It is here that trees such as spruces and firs take on a shrub-like appearance called krummholz, a word that means twisted wood. The spreading, shrubby growth form is caused by the fact that the tree branches beneath the snow survive, but those that protrude above it are killed by wind chill. Thus the tree grows mostly horizontally, producing a shrubby, often prostrate form. Beyond the zone of krummholz, trees cannot exist due to the severity of winter weather, and it is there that the boreal forest yields to arctic tundra. Characteristic mammal species include red squirrel, ermine, varying hare, lynx, wolverine, grizzly and black bears, woodland caribou, and moose. Some of these, such as the ermine and the hare, change pellage seasonally, being white in winter and mottled tan in summer. In winter, bears locate dens where they enter into a deep sleep, though not a true hibernation. Mammals such as hare and lynx remain active throughout winter months, the former trying to avoid predation by the latter. Beavers are particularly abundant in many boreal forest regions. Porcupines are common rodents in boreal forests throughout North America, but are absent from Eurasia. Many bird species, such as common loon, wood warblers, thrushes, and flycatchers, are strongly migratory, but others, such as the pine grosbeak, boreal chickadee, three-toed woodpecker, spruce grouse, and great gray owl are resident, even during the cold, snowy winter months. Reptiles and amphibians experience a reduced biodiversity in the boreal forest compared with more southern latitudes. This is most likely because these animals are ectothermic and cannot survive easily in an ecosystem where cold temperatures predominate for most of the year. There is a major insect flush in the spring, as many insect species emerge from over-wintering eggs and pupae. Mosquitoes and black flies, a significant nuisance to humans as well as other animals, are often dramatically abundant. Timber harvesting is widespread in boreal forests in North America and much of Eurasia. The wood is used for construction as well as for paper pulp. Clearcutting, a lumbering practice where a large section of forest is completely cut, is a common practice in many regions. Clearcutting is controversial 35

among conservation biologists because it destroys large tracts of forest, though in some regions it seems to succeed in allowing for sustainable harvest. Clearcut areas are typically reseeded to promote rapid regrowth of trees, which can then be harvested in as short a time as forty to sixty years. Clearcuts thus go through a process of ecological succession that makes them suitable habitats for species that require brushy areas. Looking at the landscape on a large area scale, areas of clearcuts within otherwise mature boreal forest, each clearcut in a different state of regrowth, can actually enhance biodiversity by providing a patchwork quilt of different habitats. Because these habitats change with time, one area maturing, one newly clearcut, ecologists call such a forest a shifting mosaic of habitats. The shifting mosaic concept is developing into a model for how timber harvesting can be accomplished in such a way as to enhance preservation of biological diversity. Tree harvesting, as noted above, should ideally ensure sustainable yield and, in North America, that is generally the case. The most serious loss of boreal forest is currently in Russia, where far more forest is cut annually than is permitted to regrow, resulting in an overall loss of forest that is resulting in an expansion of barren ground tundra into areas traditionally occupied by forest. Boreal forests in North America are affected by increased human usage for recreational purposes. Activities such as power boating may exert significant stresses on species such as common loon, which has experienced significant population declines in some areas. Boreal animals such as beaver and lynx have been historically exploited for the fur trade. One boreal mammal, the caribou or reindeer, has been domesticated in Europe. Temperate Rain Forest: A Unique Coniferous Forest The Pacific Northwest of North America, in the states of California, Oregon, and Washington, and the Canadian province of British Columbia, is the site of a coastal forest of tall sitka spruce, western hemlock, western red cedar, grand fir, and common Douglas-fir, arguably the most majestic of the Earths forests, the temperate rain forest. In California, this forest also includes the imposing redwood, a tree that often exceeds 300 feet in height. This coniferous forest, composed of the most statuesque of all North American tree species, is a land of tall trees and high rainfall, up to 350 cm (138 inches) annually. The combination of coastal fog along the California and Oregon coasts as well as the high precipitation, particularly in Washington and British Columbia, provides the necessary moisture to support an unusually tall assemblage of tree species, collectively termed the temperate rain forest. In many ways the temperate rain forest is ecologically similar to the boreal forest, just larger and far more lush. Trees routinely exceed 200 feet in height and may have trunks fully eight feet wide at their bases. It is normal for the trees to live from between 400 to 700 years, though some may attain a life span of one thousand years. This forest is also unique in many areas because it is an old-growth forest, one in which the trees have not recently, if ever, been felled by ax or saw. The complex physical structure of an oldgrowth temperate rain forest makes it different from the small stature, more 36

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rapidly growing spruce-fir forests that typify most of the boreal region. Because of the oldgrowth nature of much of the temperate rain forest, there has been significant controversy surrounding logging practices. Tracts of forest have been clearcut in The giant redwood tree named General Sherman dwarfs tourists at many areas and the the Sequoia National Forest near Visalia, California. The tree has a forest that is regrown maximum diameter of 36.5 feet, is 274.9 feet tall, and is believed to be about 2,200 years old. either from seed or by planting seedling trees will itself be harvested in as short a time as sixty years, thus preventing the eventual reestablishment of old-growth forest. Clearcutting also has the potential to increase erosion of soil, polluting streams that provide essential habitat for salmon and other species. The intrinsic beauty as well as ecological significance of oldgrowth forests have made many urge that they be conserved. Because of the logging issue, old-growth temperate rain forests have been subjects of much ecological study. The northern populations of spotted owl have been at the center of intense debate (the owl made the cover of Time magazines June 25, 1990, issue) because the species, which is considered threatened, apparently is restricted to old-growth forests for nesting sites. In addition, a small, robin-sized seabird called the marbled murrelet, common on the offshore waters from British Columbia through northern California, flies from the sea to nest atop the tall trees of the temperate rain forest. This birds nesting site was utterly unknown until a nest was discovered atop a Douglas-fir in northern California in 1974. The red tree vole, sometimes called the red phenacomys, seems to live its life in a single Douglas-fir, feeding on the needle leaves and nesting within the boughs of the huge tree. Whole populations and numerous generations of red Moss-covered trees in the Olympic National Park near Port phenacomys reside withAngeles, Washington. The park boasts ocean shoreline and tall mountain peaks, including Mount Olympus at 7,965 feet. The in a single tree.
annual rainfall in the park is 56.5 inches.

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FOR GREATER UNDERSTANDING

Questions
1. Why is much of the boreal forest dominated by needle-leaved evergreen trees? 2. What differences separate the boreal forest from the temperate rain forest, also a forest of needle-leaved trees?

Suggested Reading
Pielou, E.C. The World of Northern Evergreens. Ithaca, NY: Comstock, 1988.

Other Books of Interest

Henry, J. David. Canadas Boreal Forest. Washington, DC: Smithsonian Institution and Press, 2002. Kricher, John. A Field Guide to California and Pacific Coast Northwest Forests. Boston: Houghton Mifflin, 1998. Mathews, Daniel. Cascade Olympic Natural History. Portland, OR: Raven Editions, 1988. Storer, Tracy I., and Robert L. Usinger. Sierra Nevada Natural History. Berkeley: University of California Press, 1963.

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Lecture 8: Temperate Deciduous Forest

The Suggested Reading for this lecture is John Krichers A Field Guide to Eastern Forests.

astern North America, as well as much of Europe and temperate Asia, has forest characterized by broad-leaved deciduous trees. The similarity among these forests can be striking. Sycamores in China are scarcely distinct from those in North America. This is not a coincidence but rather the historic result of continental drift, the trees having evolved before the division of ancient Laurasia, the northern supercontinent. Oaks, maples, sycamores, beeches, and hickories all typically lose their leaves in synchrony as summer turns to autumn. Leaf drop is an adaptation to the impending winter, when the air temperature will become sufficiently cold to freeze the soil, making uptake of water impossible for the trees. Leaf drop is a precursor to physiological dormancy, permitting the trees to endure the prolonged cold of winter without catastrophic water loss. A temperate deciduous forest typically contains a mixed assemblage of deciduous trees along with some evergreens such as hemlocks and pines. Such a forest is annually cyclic, leaves gradually opening and unfolding in spring, with summer, the growing season, the time of maximum photosynthesis. As autumn arrives, chlorophyll, the pigment that reflects green (thus making the leaf look green to us) and is of crucial importance in photosynthesis, deteriorates, and other pigments, masked until then, make leaf colors turn various shades of red, yellow, and brown. The array of fall colors, from the blazing oranges of sugar maple to the subtle browns of oaks, makes this forest one of the most splendid to behold. In winter, aside from the scattered conifers that mostly retain their needle-like leaves, the forest has a stark look, the branches barren of their leaves. Deciduous forests are typically stratified into fairly clear layers determined by the heights of the resident plants. There is a canopy layer defined by the crowns of the tallest trees, typically sixty to eighty feet above
Colorful leaves enhance a beautiful autumn day in New England.

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the ground. Usually there will be a subcanopy or understory of trees such as dogwoods or sassafras. Beneath them will be a shrub layer of various viburnums, blueberries, huckleberries, rhododendrons, or other shrubs. There is also a fern/herb layer where various wildflowers grown among the fern species and other plants collectively called ground-cover. This biome is represented in the northern hemisphere throughout most of eastern North America, Europe, much of China, and Japan. A similar forest is also found, with very different species, in southern Argentina, southern Australia, Tasmania, and New Zealand. The key to understanding the Temperate Deciduous Forest is growing season. This is the number of days when conditions of temperature and moisture are favorable for sustained growth. As you might expect, the growing season becomes shorter in the northern hemisphere as you move in latitude from south to north. For example, the growing season is about 250 days in much of the Deep South (Mississippi, Georgia, Louisiana), about 200 days in North Carolina, 175 days in New York, and only 140 days in Ontario, Canada. The most important variable of climate is temperature, which varies from high in the summer months to often below freezing for much of the winter months. Precipitation is less variable, but, because of the cold air temperatures of winter, it often falls as snow, which is unavailable to plant roots until it melts and the ground thaws. Temperate forest is found through a fairly wide range of mean annual temperatures, from about 3 C to 18 C. Mean annual precipitation, depending upon locality, ranges from about 150 cm to 250 cm. Deciduous forests drop leaves in fall and thus there is a substantial buildup of ground litter that is then worked on by decomposer organisms. This litter layer community is active in mulching the leaves, releasing minerals that will enter the soil and subsequently be taken back into the vegetation, the essence of recycling. Soils are usually fairly well drained, typically mixtures of sand, silt, and clay. As water moves through the soils it leaches minerals from the top layers of soil and deposits them in deeper layers. Thus the soil has its own pattern of stratification, called horizons, due to the interaction of climate, precipitation, and vegetation. Tree roots easily penetrate into the deeper horizons where minerals are most concentrated. Because of leaching, as well as the chemistry of the vegetation itself, the soils are typically acidic. 40

LECTURE EIGHT

USDA Forest Service

The biodiversity of temperate deciduous forests is impressive, though it never rivals that of equatorial rain forests. Permanent residents include many nut consumers, such as squirrels, chipmunks, raccoons, wild turkeys, and blue jays. Deer, skunks, and foxes are abundant and wide ranging, and bears are common in certain areas. In most regions, however, large predators such as bears, bobcats, and wolves have been eliminated. Vegetation diversity is complex in deciduous forests. In Eastern North America, for example, different regions are dominated by different tree species. In the Southeast, for example, is the Southern Hardwood Forest, a rich forest of species such as various magnolias, Virginia live oak, common persimmon, pecan, redbay, and pawpaw. This species assemblage is in strong contrast to that of the Northern Hardwood Forest comprising mostly yellow birch, sugar maple, American beech, eastern hemlock, and eastern white pine. The most species-rich area of deciduous forest is in the Appalachians, the Cove Forests of the Great Smoky Mountains of eastern Tennessee. Over thirty tree species may occur in close proximity, including such species as white basswood, Carolina silverbell, tulip tree, yellow buckeye, and sugar maple. Ecologists have struggled to understand what factors determine species composition in various regions within the overall deciduous forest biome. One prevalent view was that such forest associations were interdependent and thus strong interactions among, say, American beech and sugar maple forced both species to associate. A school of study called phytosociology, largely started by European ecologists, struggled to find order among plant associations and reveal predictive rules of assembly. But other ecologists objected, suggesting that the co-occurrence of such species was largely coincidental, a result of each species having similar physiological requirements and similar distributions. If anything, many species likely competed among themselves and with other species to reach adult size and successfully reproduce. This view of the plant community was termed individualistic, meaning that no two plant communities are more than superficially similar. Studies of pollen profiles in bogs and lakes, as well as carefully done statistical studies on plant distributions, have confirmed the individualistic model. Plant pollen accumulates (without decomposing) through thousands of years in the layers of mud found in bogs and lakes. Such accumulations allow a look back through time, reaching back to the time of glaciation, some 20,000 years ago. What is seen is that todays plant associations did not move as a unit when glaciation forced northern species to the south and, when glaciers eventually melted, plants did not migrate north in the same associations as now exist. Forests throughout much of eastern North America were extensively cut during the years of European colonization, but these areas, used for agriculture and pasture, were largely abandoned when the West opened to colonization in the mid-1800s. After abandonment, vegetation strongly tended to follow a generally predictable series of changes as weedy invader species were progressively replaced by various grasses and shrubs, then colonizing trees, and then so-called climax tree species typical of the mature forest. This process is called ecological succession and has been the subject of 41

much study. Ecologists have learned that succession results from numerous species having evolved life cycles that are optimal at various levels of light intensity and disturbance. This view has resulted in viewing deciduous forests as patchworks of varying disturbance histories rather than as massive, homogeneous tracts. Biodiversity is strongly seasonal in deciduous forests, with wildflowers and ferns evident only in summer months, though they survive as root systems below ground during the winter. The most pronounced seasonality is found in the animal community. Some species, such as many insects and mammals, overwinter in a dormant or semi-dormant stage. Insects survive in egg cases and as dormant pupae and some mammals may hibernate or enter deep torpor. Pond animals such as frogs and toads lie dormant in the mud throughout the winter chill. Bird species offer the most dramatic example of seasonal changes in biodiversity. With the opening of leaves in spring comes a flush of insects, especially caterpillars, flies, and beetles. It is then that many species of migrant birds, the thrushes, warblers, orioles, and tanagers, return from their tropical wintering grounds to nest in the broad-leaved forests. Many of the most industrialized areas of the planet are in the latitudinal range of temperate deciduous forest. As such, the forest is subject to cutting to make room for commercial ventures or housing as well as subjected to air and water pollution. Acid rain, for example, is a byproduct of industrialization, coming from emissions of industrial plants as well as automobiles and other vehicles with internal combustion engines. As mentioned earlier, the Eastern Deciduous Forest of North America was dramatically cut and cleared during the European settlement of the continent, as agriculture pushed westward. Much of what was pasture and farmland a century ago has reverted back into forest cover today. However, much of that area is being subdivided for shopping malls, industrial parks, golf courses, and housing developments, all of which represent relatively permanent alterations to the landscape. Ecologists use the term fragmentation to describe the process of chopping a sizeable tract of forest into ever-smaller parcels, thus isolating populations on small forest islands. This realization has led to the emergence of a branch of ecology called landscape ecology, and has contributed to yet another growing field called restoration ecology. Landscape ecology focuses on how landscape patches interact to affect such things as biodiversity. For example, if two woodlots are isolated by fragmentation, they become forest islands, and each may be too small to support animal species that typically require large areas. But if a woodland corridor is left intact connecting the two fragmented forest lots, in effect they function as though the actual area is much larger. As another example, ecologists have observed the nest failure of birds such as wood thrush from small, fragmented woodlots. But in spite of repeated nest failure, thrushes remain in such woodlots. This is because they move as surplus individuals from source populations in large forests and attempt to nest in sink habitats such as fragmented woodlots. Restoration ecology attempts to mimic natures processes and thus restore damaged or fragmented habitats.

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42

FOR GREATER UNDERSTANDING

Questions
1. In what ways does length of growing season influence the adaptations of temperate forest trees? 2. What was phytosociology and what was it attempting to do? Why is it no longer used in ecology?

Suggested Reading
Kricher, John. A Field Guide to Eastern Forests. Boston: Houghton Mifflin Company, 1998.

Other Books of Interest

Braun, E. Lucy. Deciduous Forests of Eastern North America. New York: Hafner, 1972. Brooks, Maurice. The Appalachians. Boston: Houghton Mifflin, 1965. Robichaud, Beryl, and Murray F. Buell. Vegetation of New Jersey. New Brunswick, NJ: Rutgers University Press, 1973. Sutton, Ann, and Myron Sutton. Eastern Forests. New York: Alfred A. Knopf, 1985.

43

Lecture 9: Grassland and Savanna

The Suggested Reading for this lecture is Stephen R. Jones and Ruth Carol Cushmans A Field Guide to the North American Prairie.

he sight of millions of bison grazing on endless prairie, the grasses waving in the gentle breeze, portrays, for the minds eye, a vision of the grassland biome in North America, a sea of grass. Grassland appears monotonous, populated with various grasses and wildflowers, with little in the way of trees and shrubs. Grassland Grasslands do not all look alike. Because of differences in annual moisture, some grasslands contain predominantly tall grasses, others mixed, and others short. This pattern is evident in North America, where the tallest grasses typify the most eastern regions. Approaching the Rocky Mountains, grassland becomes predominantly short, and eventually mixes with desert. Grassland, wherever it occurs, gives the impression of vast openness. There is a good reason why Montana, which is mostly grassland, is often called Big Sky Country. In grassland, the horizon is more sweeping than any other place except the open ocean. In North America, grassland dominates the central part of the continent, from southcentral Canada into eastern Mexico. States such as Montana, the Dakotas,
Top: The Flint Hills of Kansas contain the largest tract of remaining tallgrass prairie in the United States. Bottom: Tallgrass prairie is dominated by three main grasses: Big Bluestem, Little Bluestem, and Indiangrass. Subdominant grasses and forbs (non-woody, non-grass, flowering plants), though less abundant, contribute to the diversity of the prairie. Inset: A bee and a spider take advantage of the flowering grasses during the pollination cycle.

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All images: Teresa M. Woods/Konza Prairie Long Term Ecological Research (LTER) Program

Kansas, Nebraska, and Oklahoma are historically grassland dominated, and natural prairie occurs as far east as Ohio. Grassland is also the natural vegetation of the Central Valley of California. The biome occurs in many other places in the world: in South America, where, in Chile and Argentina, the grassland is called pampas. Throughout central Europe grassland is called steppe. In central and southern regions in Africa it is veld. In North America, it is called prairie. In general, mean annual precipitation is less in grassland than in forest, but there is a wide range. In the short grass prairie of Colorado, for example, the mean annual precipitation is a meager 10 inches (25 cm), but it ranges from 25 to 40 inches (65 to 100 cm) in the most eastern regions. In some prairie regions, mean annual precipitation is as low as 6 inches (15 cm). There is a gradient in precipitation from west to east, with increasing precipitation moving eastward. Thus tallgrass prairie is found in eastern regions, and much of the central prairie is mixed grassland. Because North American grasslands are confined to the interior of the continent, there are no mitigating climatic effects by the oceans, such as occur along the eastern and western seaboards. Extremes of temperature, indeed, extremes of weather, are common. Prairie experiences severe winter blizzards with drifting snow blown by incessant high winds. One of the characteristics of prairie most recorded in the journals of settlers and pioneer travelers during the nineteenth century was the constancy of wind. Tornadoes are commonplace throughout the summer months. In spring, one day may be mild while the next features a major snowfall. Natural fire, set by lightning (many thunderstorms occur throughout the summer months), is a major factor influencing prairie ecology. Without regular fires, in many areas prairie would eventually be replaced by forest. This pattern is evident in places such as Wind Cave National Park in South Dakota. The area where the park is located is called the Black Hills, named for the dark foliage of ponderosa pine forests that occur there. Without prescribed burning by the National Park Service, Wind Cave National Park, one of the last vestiges of natural grassland in the region, would be invaded by ponderosa pine and associated species. There is anthropological evidence that suggests that Native Americans regularly set fire to the grasslands, apparently understanding that doing so would maintain the area as grassland, which was desirable for hunting game and for moving from one place to another. Soils are typically rich in nutrients, with abundance of such elements as calcium, potassium, and phosphorus. The dense roots of the grasses effectively tap into this abundance of nutrients, allowing continual growth through the 120- to 200-day growing season. Because of the natural richness of the grassland soils, it is hardly surprising that agricultural activities dominate regions of grassland. Many species of grasses occur throughout the biome, but they differ from one region to another depending on how much precipitation the region receives. Tallgrass prairie in North America is dominated by species such as big bluestem (Andropogon gerardi ), which is a bunch grass. Bunch grasses, as the name implies, have stems that radiate from a central cluster. Big bluestem, under ideal growing conditions, can reach a height of just over three meters, 45

about ten feet. When General Custers troops were easily defeated at the Battle of the Little Bighorn in Montana, Sitting Bulls braves were totally hidden in the tall grass that Custers cavalry rode through, making it easy for the braves to attack the soldiers. Many grasses are sod forming, rather than bunch grasses. Species such as blue grama (Bouteloua gracilis) and buffalo grass (Buchloe dactyloides), which occur abundantly in shortgrass prairie, are similar to grasses that make up lawns, growing as a dense mat covering the soil. Grasses are adapted for rapid growth after fire or grazing. They often spread by underground stems called rhizomes, soon carpeting an area. The root system contains as much or more tissue as the above-ground shoots, making it easy for grasses to resprout after fire or grazing (consider how often one has to cut the lawn in summera lawnmower is essentially a form of grazing). Grasses are wind-pollinated, which is an effective means of pollen dispersal in an open land where wind normally prevails. Many wildflower species are mixed among grasses, including legumes such as various clovers, which take nitrogen from the atmosphere and incorporate it into usable form. The abundance of legumes further enhances the high fertility of the soil. Numerous species of composites, daisy-like plants, many of them with wonderfully colorful flowers, grow among the prairie grasses. One of the great pleasures of exploring real prairie is to enjoy the myriad of different kinds of wildflowers encountered there. In North America, several animals are closely associated with prairie, but the two that are most obvious are the American bison and prairie dogs. Bison, once among the most abundant of North American mammals, were hunted nearly to extinction in the nineteenth century. Bison were important grazers in the central regions of the prairie and many ecologists believe that regular grazing by bison helped to conserve the grassland, preventing invasion by non-grass species. Thus it is the combination of low precipitation, periodic fire, and grazing that combine in determining if grassland will persevere. There are two prairie dog species, the black-tailed and the white-tailed, and they are both keystone species in maintaining prairie ecology. Prairie dogs are really a kind of ground squirrel, a member of the rodent family, and other ground squirrels also are common throughout grassland, as are many species of mice. The black-footed ferret, a member of the weasel family and the rarest North American mammal, closely associates with prairie dog colonies. Other grassland mammals include coyotes, badgers, and pronghorns. Scattered throughout much of the central regions of North American prairie are lakes and marshes called prairie potholes. These are essential nesting grounds for many duck species such as canvasback and redhead, as well as numerous herons and shorebirds.
LECTURE NINE

Most natural prairie in North America has been replaced by agriculture. What was once natural grassland are now fields of wheat, rye, corn, sorghum, and other essential crops. Thus it is difficult to find natural prairie where once it may have seemed endless. There are scattered preserves that sustain natural prairie, but the total areas of these places are generally small. In addition, many alien species of grasses and wildflowers (weeds) have invaded natural prairie, diminishing the biodiversity that would otherwise prevail in these areas. 46

The spread of cattle and sheep ranching in North America has made a major impact on grasslands. In areas that are particularly sensitive, such as the short grass regions found in the southwest, natural grassland has become desert as a result of selective grazing by cattle. Cattle eat the grasses but ignore such noxious species as cacti and creosote bush, allowing these species to eventually replace the grasses. Savanna Savanna is an open habitat with scattered trees among various grasses. Trees, particularly acacias, may be scattered rather evenly among the grasses or be more restricted to certain areas within the expanse of grasses. Wind is often a major factor and periodic fire, set by lightning, is common throughout savannas. In Africa, large animal herds are an obvious feature of savanna, but an abundance of large animals is not a universal characteristic of savannas elsewhere. The vast herds of antelope, gazelles, wildebeest, zebra, and elephants that serve as food for lions, cheetahs, leopards, and hyenas are unique to Africa. Most of us associate savanna with east and southern Africa. But savanna occurs elsewhere. In North America, the Everglades of Florida is a type of wet savanna. There is much savanna throughout parts of South America, particularly the llanos of Venezuela, the cerrados of northern Brazil, and the pantanal of southern Brazil. Savanna can also be found in parts of Central America such as Nicaragua and Belize. Savannas are also found in parts of India and southeast Asia, and Australia. Savannas occur in regions with a strongly developed dry season. The severity of the dry season varies among savannas, but in East Africa, the dry season is sufficiently severe to require annual migrations of the large animals, as they move in search of water and fodder. Savannas generally receive between 50 and 150 cm of precipitation annually. During wet months, the grasses and trees are green and lush, but during dry season, the landscape turns brown. Tropical savannas experience a mean annual temperature from about 18 to 30 C, the same temperature range as is found with rain forests. The lack of rainfall during dry season and overall high temperature result in significant physiological stress on both plants and animals. It is unclear to ecologists exactly what factors determine the existence of savannas. Not all savannas occur in areas with a protracted dry season. As a result, some ecologists believe that savannas are also caused by nutrientpoor soils that, for various reasons, are insufficient to support lush forest. Prolonged human agricultural activities have been blamed for causing savanna formation. The argument is that human use has depleted nutrients in the soil, thus preventing regrowth of rain or seasonal forest. Savanna vegetation claims such depleted soils by default. Fire is another strong factor in savanna ecology. Periodic natural fires, just as they do with grasslands, aid in promoting regrowth of savanna grasses and sedges, at the expense of woody species that, in the absence of fire, might eventually come to dominate the region. Because most savannas are also areas of strong human influence, fire frequency may have increased due to human activity. 47

Historically, savannas have been increasing in area since the middle part of the Cenozoic Era. Global climate has become drier and more temperate, causing the spread of savanna where once there was unbroken rainforest. It is this climatic shift that was likely the major selection pressure in the emergence of hominids, ancestors of human beings, from East and Southern Africa. Without question the large herds of hoofed (ungulate) mammals that inhabit the African savanna are the animals most associated with savanna ecology. Savannas elsewhere in the world lack this amazing mammalian diversity. It is interesting to note that African rainforests do not support nearly the same mammal biomass as savannas. The large herbivorous mammals of African savannas feed either by browsing or grazing. Browsers, like giraffes and Thompsons gazelle, for example, feed mostly on leaves from shrubs and trees, while grazers, like zebra and wildebeest, rely mostly on grasses. There are numerous species of antelopes and gazelles throughout African savanna as well as zebra, wildebeest, giraffe, elephant, and wart hog. Elephants exert strong effects on savanna ecology. They damage trees such as baobobs as well as prevent their regeneration, thus eventually converting a savanna rich in trees to one of mostly grassland. Though vertebrates command most attention on African savanna, numerous insect species are ecologically important, ranging from locusts, which periodically experience population eruptions, to termites, whose tall mounds characterize savannas worldwide. Human beings, as mentioned above, likely owe their very existence to savannas. If one habitat deserves to be called natural as far as our species evolution is concerned, it is savanna. It has even been suggested that our fondness for lawns, for grassy areas with scattered trees, stems from a deeply ingrained familiarity with a savanna-like habitat. Savannas support a high biodiversity of unique species. In addition, these animals, as is the case in Africa as well as other areas, are dependent on being able to move around, to migrate, with the seasons. It is thus imperative that large areas of savanna be maintained as sanctuaries for the ungulates and associated species. These must be separated from areas where cattle and other non-native animals are grazed as the native animals can act as vectors for diseases such as sleeping sickness (vectored by the tsetse fly) and malaria (vectored by mosquitoes). Such diseases can devastate cattle, as they are not nearly as resistant to the pathogens as the native species that have shared a long evolution with the pathogen and thus developed natural immunity.
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LECTURE NINE

Elephants on the move along the savanna during the African spring.

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FOR GREATER UNDERSTANDING

Questions
1. How do variables such as precipitation, fire, and grazing combine to affect grassland? 2. What is the essential difference between grassland and savanna?

Suggested Reading
Jones, Stephen R., and Ruth Carol Cushman. A Field Guide to the North American Prairie. Boston: Houghton Mifflin, 2004.

Other Books of Interest

Brown, Lauren. Grasslands. New York: Alfred A. Knopf, 1985. Gleason, Henry A., and Authur Cronquist. The Natural Geography of Plants. New York: Columbia University Press, 1964. Reichman, O.J. Konza Prairie. Lawrence: University of Kansas Press, 1987.

Websites to Visit
1. The TIEE (Teaching Issues and Experiments in Ecology) journal website provides an excellent abstract of an article by professors Harmony J. Dalgleish (Kansas State University) and Teresa M. Woods (Utah State University) entitled The Effects of Bison Grazing on Plant Diversity in a Tallgrass Prairie http://tiee.ecoed.net/vol/v5/practice/dalgleish/abstract.html 2. Konza Prairie Biological Station (LETR) Long-Term Ecological Research at Kansas State University, Manhattan, KS, provides a comprehensive website about prairie ecology http://climate.konza.ksu.edu/konza

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Lecture 10: Desert

The Suggested Reading for this lecture is James A. MacMahons Deserts.

eserts look dry. The very word desert seems to convey a sense of aridity, of a land where water is at a premium. Indeed that is true. But there is much variety evident in the worlds deserts, which can range in appearance from undulating sand dunes with little apparent life, to colorful landscapes awash with the color of myriad blooming flowers. Some deserts, like the Atacama along the coast of Peru and Chile, resemble what we imagine the surface of a planet such as Mars to be, seemingly devoid of any life. Others, like the Great Basin Desert of western North America, are populated by a monotonous-looking assemblage of shrubs, such as Big Sagebrush. Still others, like the Sonoran Desert of Mexico, Arizona, and southern California abound with succulents, various cacti, and other species, some the size of trees. Deserts contain many unique species of plants and animals because inhabitants of deserts are subjected to strong selection pressures resulting in remarkable adaptations to the constant aridity. Some adaptations of desert organisms, like the water-holding barrel-shape of spiny cactuses, are readily evident while others, like the subtle behaviors of lizards to control their body temperatures or the ability of kangaroo rats to subsist without drinking liquid water, are much more subtle. Desert biomes occur in many places on Earth. Deserts found in the temperate zone are called cold deserts because they experience snow and cold temperatures in the winter months. Deserts closer to the equator are called hot deserts because they rarely experience snow. Shrubs tend to be the

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The vast Atacama Desert, near San Pedro de Atacama, Chile.

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dominant form of plants in cold deserts, but succulents join shrubs in hot deserts. In general, biodiversity is highest in hot deserts. Globally, many deserts occur around 30 north or 30 south latitude. This is because of major convection cells in the atmosphere that converged at those latitudes and that are previously depleted of moisture. Thus dry air is forced down, but precipitation does not occur, so deserts result. There are four deserts in North America: the Great Basin, the Chihuahuan, the Sonoran, and the Mojave. Of these, only the Great Basin is cold desert. It ranges from the Canadian border and Washington south to Nevada and parts of Arizona and eastern California. It is called the Great Basin as it is bordered to the east by the Rocky Mountains and to the west by the Cascade and Sierra Nevada mountain ranges. The Chihuahuan Desert is found throughout much of northern and central Mexico, crossing the U.S. border in Texas and eastern New Mexico. The Sonoran Desert, perhaps the most picturesque of any of the four North American deserts, contains many cactus species including the giant saguaro cactus and organpipe cactus. It is found from northwestern Mexico and the Baja Peninsula through southern Arizona and parts of southern California. The Mojave Desert is the smallest of the North American deserts, confined to southeastern California, including Death Valley and the Salton Sea. It is famous for its joshua trees, which belong to a group of desert-dwelling plants called yuccas. In South America, one finds the Atacama Desert along the coast of Peru and Chile, blocked from obtaining moisture from the huge Amazon Basin by the high Andes Mountains. There is desert in Africa both in the northern part of the continent, where the great Sahara Desert extends from Morocco through Egypt, as well as in the south, where the Karroo Desert is found. Much of interior Australia is desert, a huge area. The largest desert on Earth is the vast Gobi desert found throughout most of central Asia, throughout much of China and Mongolia. Deserts typically experience below 10 inches (25 cm) of annual precipitation. What moisture they do receive is seasonal, falling during a brief wet season. In the Sonoran Desert in southern Arizona, for example, rain falls briefly in the spring, often resulting in a flush of annuals, many species of colorful flowers that bloom in synchrony. Rain also occurs again sparingly in mid to late summer, when late afternoon thunderstorms are typical. The temperature range of deserts is wide, with annual mean temperatures from 23 F to 86 F (5 C to 30 C). The warmest temperatures of any place in the United States are typically recorded for Death Valley, California, in the Mojave Desert, where a temperature of 134.5 F (57 C) was once recorded, ranking as the second highest temperature recorded on Earth. Desert soils tend to be fertile, with an abundance of essential minerals. The sparse plant growth that typifies deserts is thus the result of a continual shortage of water. Some desert plants, such as honey mesquite, have long taproots that grow sufficiently deep to reach ground water. Others, like the many species of succulents, absorb large quantities of water during the brief times when it is available and then utilize it as needed during the long dry spells. In areas of high temperature, evaporative water loss from desert soils can result in the deposition of minerals on the soil surface. Minerals, dissolved in the 51

water, remain behind as the water evaporates. Salt lakes, where minerals are concentrated in bodies of standing water, are common in some desert regions. Because there is relatively little plant cover, erosion by wind and floodwaters is common in deserts. Such erosion sometimes exposes ancient sedimentary rocks that contain an abundance of fossils. In North America, such is the case in many locations, including Dinosaur National Monument in Utah and the Petrified Forest in Arizona. One of the richest fossil hunting grounds in the world is the Gobi Desert in China and Mongolia, where the remains of many dinosaurs and early mammals have been excavated. Given the natural fertility of many desert soils, it is little surprise that irrigation can convert a desert into productive agricultural land. Doing so, however, involves importing a great deal of water and thus tends to be costly. The high cost is increased even more in hot desert irrigation because of the high evaporative water loss caused by high temperature. Thus crops such as corn raised in such areas require even more water than they would in a cooler location. Deserts have a surprisingly high biodiversity. Many species of plants have adapted to thrive in deserts. Shrubs as well as large cactus species tend to be widely spaced, with shallow, extensive root systems that absorb moisture from a large area of soil. Leaves are typically small and waxy, traits that minimize water loss from heat and wind. Some plants have an abundance of thorns as well as toxic chemicals in their leaves to discourage herbivores. The pungent odors of shrubs such as creosote bush and big sagebrush are produced by such chemicals. Species such as cholla cactus actually use their dense thorns as a means of dispersal. These plants have long, chain-like branches divided in to sections called joints. A joint can easily become attached to a passing animal such as a deer or peccary. The joint, transported by the animal, eventually drops off or is picked off by the creature. Once on the ground, the joint can root and grow. Many desert plants, most particularly cactus and Old World euphobias, photosynthesize mostly through their stems. The palo verde tree of the Sonoran Desert is so named for its green bark. Though it is leafless for much of the year, it still can photosynthesize without experiencing the evaporative water loss that would come with leaves. Many desert plants, including creosote bush, mesquite, palo verde, and cacti, are pollinated by animals. Some, like creosote bush, produce small flowers, but others, like honey mesquite, become amazingly colorful when flowering, a real beacon for pollinators. In the case of the giant saguaro cactus, bats are the usual pollinators, visiting the large white flowers at night. Many insects are important desert pollinators, as are some hummingbird species. Though harsh physical conditions typify deserts, many intriguing biotic interactions are essential to ecosystem structure. For example, a seed of a saguaro cactus will not germinate unless it is on a slope and has some shade from a nearby rock or plant. The best place for a saguaro seed to germinate and grow is in the shade of a shrub or tree such as a palo verde. It is common to see giant cactuses emerging from the low crown of a shrub or palo verde, which has served as a nurse tree, furnishing essential shade during the early development of the cactus. Deserts harbor a high biodiversity of animals. Up to thirty bird species nest in cavities in saguaro cactuses, for example. Many desert animals are active 52

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only at night when the air temperature is much cooler, minimizing evaporative water loss. Numerous rodent species inhabit deserts, some elegantly adapted anatomically and physiologically to the harsh desert conditions. The kangaroo rat subsists on water obtained from the seeds it digests. Jackrabbits have large ears that obviously serve to increase evaporation and cool the animal during the heat of the day. Wood rats, sometimes called pack rats, assemble large middens containing mostly seeds as well as other objects unrelated to nutrition. Reptiles adapt well to deserts, and numerous species of lizards and snakes can be found throughout the worlds deserts. One of the more specialized species is the sidewinder, a rattlesnake that moves in a sideways manner rather than forward, an adaptation to unstable, shifting desert sands. People have adapted throughout history to living in deserts and continue to do so. Many of the major oil-supplying nations of the world are located in the Middle East, now a region that is primarily desert. Thanks to sophisticated though costly irrigation techniques, countries such as Israel are able to grow many crops in some of the worlds most arid lands. Human use of deserts in North America is potentially stressful to regional water supplies. The only way to bring water to deserts is to tap what is deeply below ground or transport the water, via pipelines, from other areas. Thus as human populations grow in desert regions, as agricultural activities in such areas increase through irrigation, water is depleted from some other location. Because of increasing human populations in desert regions, encroachment by housing developments, malls, and industrial parks is threatening many deserts with fragmentation and species loss. Immigrants to desert regions frequently attempt to replace natural vegetation with lawns and other water-costly landscaping. Such luxuries as swimming pools, which tend to be commonplace among residents of desert regions, require significant amounts of water. Another threat to deserts is the increasing use of off-road vehicles. Like the Arctic tundra, desert plants are not able to recover easily from disturbance. It requires about eighty years for a saguaro cactus to attain a height of 6 to 8 feet, at which point it will produce its first blossoms. Full maturity of the plant is not reached until it is well over a century in age, and it can survive for up to 250 years. Nonetheless, the root system is very shallow, and a minor impact from an SUV or pickup truck can topple the tree in seconds. Unfortunately, many desert plant species are in demand as ornamentals. The illegal collection of various cactuses has become an increasing problem, requiring legal protection for species such as organpipe cactus and giant saguaros. Deserts, once thought of as wastelands, must be recognized for their unique ecological characteristics and afforded reasonable conservation protection.
A saguaro cactus emerged from the scant shade provided by a palo verde plant.
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FOR GREATER UNDERSTANDING

Questions
1. What are the primary adaptations of desert plants? 2. Why are deserts fairly different in different places?

Suggested Reading
MacMahon, James A. Deserts. New York: Alfred A. Knopf, 1985.

Other Books of Interest

Jaeger, Edmund C. Desert Wildlife. Stanford, CA: Stanford University Press, 1961. Kirk, Ruth. Desert: The American Southwest. Boston: Houghton Mifflin, 1973. Phillips, Steven J., and Patricia Wentworth Comus, eds. A Natural History of the Sonoran Desert. Tucson: Arizona Sonoran Desert Museum Press, 2000.

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Lecture 11: Tropical Rain Forest The Suggested Reading for this lecture is John Krichers A Neotropical Companion: An Introduction to the Animals, Plants, and Ecosystems of the New World Tropics. ropical rain forest is the most lush and diverse of the worlds terrestrial biomes. The word jungle is closely associated with tropical rain forest, but rain forest and jungle, though related, are really different habitats within the same biome. Jungles are disturbed areas where rain forest has been temporarily destroyed, either by cutting, windthrow, or some other factor, to be replaced by a thick tangle of vegetation that often grows so densely that a machete is necessary to cut a path through it. Jungles, if left alone, will eventually transform back into rain forest, a process called ecological succession.

Tropical rain forest in southern Venezuela.

Tropical rain forests are much more open inside and deeply shaded than jungles, composed of many species of trees, some of which grow to great stature, often in excess of 100 feet. Trees, which are broad-leaved and usually evergreen, are often rather slender, accentuating their apparent height. Branches radiate out high above ground level, like spokes on an umbrella, and roots emerge in large buttresses at the bases of the trees. At ground level rain forests are well shaded, as up to 99 percent of the light striking the canopy fails to reach the forest floor. Rain forest trees are often abundantly laden with an impressive assemblage of epiphytes, plants that live on other plants. These include orchids, various cacti, and bromeliads (with a pitcher-like array of spiky leaves resembling those of a pineapple). Vines also typify rain forests, including various figs (such as strangler fig), philodendrons, and looping, twisting lianas. The combination of numerous tree species plus diverse epiphytes and vines makes the physiognomy, or physical structure, of the rain forest perhaps the most structurally complex of any terrestrial ecosystem. Animals are generally hard to see well within rain forest, as the complex physical structure of the forest shelters its animal inhabitants very well. Mammals, ranging from canopy-dwelling monkeys to tapirs and tigers, can be 55

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very difficult to detect. Butterflies are conspicuous in places where sunlight strikes blooming flowers, attracting the colorful insects. Birdsong reveals the presence of many bird species and insect sounds include the almost constant whine of cicadas and other stridulating insects. Rain forest is found in equatorial regions throughout the world. Rain forests are best developed in proximity to the equator, where true rain forest prevails. Moving north or south from the equator, forests become more subject to strong seasonality and are called tropical moist forests. Rain forests and moist forests, taken together, can be found within about a 47 latitudinal belt, bounded by the Tropic of Cancer to the north and the Tropic of Capricorn to the south. The only actual rain forest in the United States is on the Hawaiian Islands, though the Everglades in south Florida rank as subtropical. The largest expanse of rain forest in the world is found in South America, in the Amazon Basin. Rain forest also occurs in Central Africa, Southern India, and Southeast Asia, Thailand, Indonesia, Malaysia, New Guinea, as well as Northeastern Australia. Constant heat and relatively constant high humidity characterize all rain forests. The growing season, the time in which plants can actively add tissue, is usually year-round. Typical daily high temperatures are around 88 F (31 C), sometimes higher, and nightly low temperature is usually around 72 F (22 C). This temperature range prevails during all months of the year. In other words, temperature is basically constant in rain forests, not seasonal, with cold winters and hot summers, as it is in the temperate zone. At ground level, humidity can reach as high as 95 percent, making the air feel very oppressive. All precipitation, and there is lots of it, falls as rain. In any area of rain forest or moist forest, it is usual for rain to fall anywhere from 130 to 250 days per year. In the Amazon Basin, precipitation ranges between 150 cm (nearly 60 inches) to 300 cm (118 inches) annually. Some areas experience up to 460 cm (180 inches) or more precipitation annually. Rain is typically seasonal, most of it occurring during the rainy season, though, in many places, it rains substantially during the dry season as well. North of the Equator, rainy season is usually from late summer through early January. The opposite prevails south of the Equator. Soils are often heavily eroded of their minerals in rain forest areas, due to the constant precipitation washing the minerals from the soil, a process called leaching. As water passes through soils it adds hydrogen atoms, making the soil increasingly acidic. Adding to the low soil fertility is the fact that many tropical soils are geologically ancient, and their great age has also resulted in accumulated leaching. It may seem ironic that lush areas of rain forest sit atop some of the Earths most infertile soils. However, recycling of minerals is rapid and efficient in rain forests and most of the minerals are rapidly pulled from the litter (leaves, twigs, roots, dead animal bodies) by the vegetation even before the chemicals have a chance to enter the soil and be washed away in erosion. Infertile tropical soils are often deep reddish in color due to the accumulation of iron and aluminum oxides, chemicals for which plants have little use. Most tropical soils also have a high content of clay, a microscopic kind of soil particle that makes the soil feel compacted, slippery, and gummy when 56

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wet. Not all tropical soils are infertile. In areas where there is recent geological activity, such as mountain raising or volcanism, tropical soils are young and mineral rich. More species of organisms comprise tropical rain forests than do any other ecosystem on Earth. One of the unique characteristics of rain forest is the high species richness that typifies it. Indeed, more than 50 percent of the worlds total species are thought to inhabit rain forests. Nearly 300 different species of trees have been identified within a single hectare (approximately 2.5 acres, not a large area) of some Amazonian rain forests. In contrast, in the most species-rich forests in the temperate zone, such as one finds in the Great Smoky Mountains in Tennessee, at most thirty species of trees will be encountered within a hectare. Typically, the number is lower than that. Added to the diversity of trees is the diversity of such groups as epiphytes and various vines, all adding to the profligate diversity of plant life in the forest. Animal life is also stunningly diverse. There are more bird species, more insect species, more frog species, more snake species in rain forest than in other kinds of ecosystems. In one area in southwestern Amazonia, 1,234 species of butterflies have been identified from within a two kilometer area. The total species richness of insects and other arthropods is unknown, but one estimate, based on carefully collected samples from single trees, suggests that as many as 20 to 30 million arthropod species may reside within the worlds rain forests. Vertebrate diversity is high as well. Within a single reserve in Amazonian Ecuador, eighty-one species of frogs have been documented, the same number of frog species found in the entire United States. Bats are highly diverse in rain forests around the world. In African, Asian, and Australian rain forests there are large bats often called flying foxes, for their dog-like faces. These impressive creatures dine mostly on fruit and are essential in dispersing the seeds of many fruit trees. In South and Central American rain forests, bats are found that consume insects, fruit, nectar, fish, frogs, birds, other bats, and even blood. The infamous vampire bat, which makes small, painless incisions in sleeping mammals and then laps the flowing blood, is found throughout the American tropics. In the tiny country of Belize, in Central America, a land with approximately the area of the state of Massachusetts, there are eighty-four bat species, compared with a total of forty found within the entire United States. Birds are among the most species rich of vertebrate groups in rain forests. Roughly twice as many species of birds, 1,695, are found in Colombia, in northern South America, than in all of North America. There are far too many species of rain forest birds to even attempt a summary but, like bats, they demonstrate many modes of feeding.

One of many species of Poison-Arrow Frog (Dendrobates ventrimaculatus) found in the Amazonian rain forests of Brazil, Colombia, Ecuador, French Guiana, and Peru. Adults reach a length of a little more than a half inch. It feeds mainly on ants. This photo was taken in the Parque Nacional Yasun in Ecuador.

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A chestnut-mandibled toucan (Ramphastos swainsonii) photographed in the Honduran rain forest.

The harpy eagle of South America, the largest of the worlds eagles, feeds on tree sloths and monkeys, which it plucks from the forest canopy during its swift flight. These are similar eagles to the harpy in Africa, tropical Asia, and New Guinea. The dinosaur-like flightless cassowarys of Australia and New Guinea feed mostly on large fruits taken from the rain forest floor, though they mix their vegetarian diet with occasional small animals. Colorful birds-ofparadise inhabit New Guinea and Australia, while equally brilliant manakins and cotingas, such as the Guianan cock-of-the-rock, are widely spread throughout the American tropics. Both feed largely on fruit and both groups of birds include species where males concentrate in arenas of their own construction and perform elaborate courtship dances to entice females. Many kinds of parrots live within rain forests, most highly colorful though often hard to see in the dense green foliage. Not all rain forest birds are gaudy: many species of antbirds are found in South America, most of them subtle shades of gray and brown. Some antbirds actually accompany army ant swarms and feed on the many arthropods attempting to flee the marauding ants. Many kinds of primates, virtually all of them arboreal, dwell within the worlds rain forests, ranging from lowland gorillas in Africa, orangutans in Sumatra and Borneo, to howler monkeys in the American tropics. Various hoofed mammals such as deer, pigs (or peccaries in the American tropics), and tapirs roam the forest interior, preyed on by jaguars (American tropics) and tigers (India and Asia).
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Snakes, including constrictors such as pythons and boas, are common within rain forest and surrounding ecosystems, and many species of poisonous snakes, pit vipers, true vipers, and cobras are found within rain forests. In central Africa, the lush rain forest is the haunt of the Gaboon viper, which reaches five feet in length but is very thick, sometimes over twelve inches in circumference, weighing as much as eighteen pounds. The snake has fangs two inches in length and very toxic venom. It is magnificently camouflaged among the leaf litter of the rain forest floor. 58

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Most people are aware that rain forests are being cut down around the world, often for wood to be used as fuel or for building, often for such activities as agriculture or cattle ranching. Loss of rain forest is of major concern to those who value biodiversity and believe it to be essential for ecosystem functioning. As rain forest is lost, so are species. Many rain forest species are rare, their distributions limited, and thus they are quite vulnerable to loss of habitat. Most remaining rain forest is in the vast Amazon Basin, an area where increasing human encroachment is ongoing, but one that is so immense that the majority of the overall forest remains, at least for now, intact. In contrast, rain forests of Central America, central Africa, and Asia are being lost very rapidly. Less than 5 percent of remaining rain forests on Earth are protected as national parks or reserves, so rain forests remain vulnerable to the chainsaw and bulldozer. When rain forest is cut, it is typically burned, releasing carbon dioxide into the atmosphere and contributing to global warming through Greenhouse Effect. Beyond that, rain forest is often replaced with ecosystems that take in far less carbon dioxide. Indeed, rain forest is often called a sink for carbon dioxide, as it takes in so much of it in the process of photosynthesis. Thus rain forest removal is a double-edged sword in the steady accumulation of carbon dioxide in the atmosphere. Rain forest loss is unlikely to be lessened, as the most rapidly growing human populations on Earth are in tropical regions. The conflict between humanitys perceived needs and the utility of preserving natural ecosystems is nowhere more challenging than in tropical regions. Rainfall is distributed quite unevenly over the course of a year in some tropical areas. In such places, the dry season is so severe that tropical rain forest cannot exist on the site. In its place is a type of ecosystem called either Tropical Dry Forest, Thorn Scrub Woodland, or Tropical Seasonal Forest. In marked contrast to lush rain forest, Tropical Seasonal Forest typically has a decidedly arid look to it. Trees are far smaller in stature than those that typify rain forest and the species richness, not only of trees, but of other life-forms as well, is less, often far less, than in rain forest. Woodlands consist of relatively few tree species, typically those with thorns, such as acacias, for example. Mean annual temperature ranges from about 20 to 30 C, the normal range for tropical ecosystems, but mean annual precipitation can range from as low as 50 to about 250 cm. What this means is that there is a moisture gradient from moderate (250 cm) to low (50 cm) and thus there is a range of ecosystem types within the overall biome itself. At the wettest end of the gradient, forests exist, ranging from broad-leaved evergreen to seasonally deciduous. The canopy is always low, typically no higher than 10 to 12 m (roughly 30 or 40 feet) and epiphytes and vines, so common within rain forests, are far less abundant, if present at all. The ground is often covered with grasses and small shrubs, many of which have leathery leaves. On the driest end of the moisture gradient, bordering that of a typical desert, the ecosystem consists of low stature trees often referred to as thorn scrub. Many animal inhabitants of Tropical Seasonal Forest, from insects to large mammals, are migratory, their perambulations determined by their need to find water.

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FOR GREATER UNDERSTANDING

Questions
1. What factors combine to make tropical rain forests the most species rich of all the worlds terrestrial ecosystems? 2. What are the major threats to rain forest biodiversity?

Suggested Reading
Kricher, John. A Neotropical Companion: An Introduction to the Animals, Plants, and Ecosystems of the New World Tropics. 2nd ed. Princeton: Princeton University Press, 1999.

Other Books of Interest

Forsyth, Adrian, and Ken Miyata. Tropical Nature: Life and Death in the Rain Forest of Central and South America. NY: Touchstone, 1987. Janzen, Daniel H. Costa Rican Natural History. Chicago: University of Chicago Press, 1983. Primack, Richard, and Richard Corlett. Tropical Rain Forests: An Ecological and Biogeographical Comparison. Oxford: Blackwell, 2005.

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Lecture 12: Marine Ecosystems

The Suggested Reading for this lecture is Sean Connells Marine Ecology.

he worlds oceans are the largest and most voluminous of Earths ecosystems. The major variables of importance in determining ocean ecology are the amount of solar radiation received and the abundance of available elemental nutrients such as phosphorus and nitrogen. The topic of oceanic productivity will be discussed in the next chapter. What is important here is to understand the basic organization of oceanic ecosystems. Oceans do not exhibit life zones that are comparable to those found terrestrially. It is helpful in understanding the great diversity of oceanic life to recognize patterns in zonation as determined by depth. Consider that the oceans average depth is about 2.33 miles (3.7 km), and that the deepest trenches in the oceans exceed 7 miles (11.2 km), substantially deeper than Mt. Everest is tall. Physical conditions change radically from surface waters to deep waters. In addition, conditions near shore and over the continental shelf are different from those far from shore, in open ocean. Pelagic Zone When you are on a ship on the open ocean, far from shore, and you look at the rolling waves of the sea, you are looking at the Pelagic Zone. Suppose you had access to a submersible, such as a diving bell, a device that could

The Major Zones in the Marine Ecosystem

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take you many fathoms (a fathom is a nautical term that means six feet) below the surface. What would you find? You begin your journey at the ocean surface, where sunlight strikes the sea. This is where photosynthesis is occurring, so it is called the euphotic zone, the zone of light. If you were to drag a plankton net you would collect many kinds of phytoplankton, much of it single-celled diatoms, dinoflagellates, and other minute plants. You would need a microscope to see these tiny plants well, though they are ecologically equivalent in what they do to the trees of forests and the grasses of prairies. They are the primary producers of the oceanic realm. Some phytoplankton, collectively called nannoplankton, are so small that they elude capture in plankton nets and must be centrifuged from water to concentrate them for microscopic observation. Feeding on the phytoplankton hordes are various kinds of zooplankton, tiny animals, some single-celled, but most multicellular. Some, like copepods, which are crustaceans related to lobsters and crabs, are permanent members of the zooplankton community, but others are larval stages of animals such as barnacles, crabs, and jellyfish. Zooplankton form the food base for numerous kinds of small fish, which in turn support larger fish and other organisms including tuna, mackerel, sharks, sea turtles, whales, and porpoises. The term plankton is taken from the same Greek root as the word planet, meaning wanderer. Plankton organisms are so small that they have limited mobility, if any at all, against the currents, placing them, for all intents and purposes, at the mercy of the currents. All of the fish, cetaceans, turtles, and larger invertebrates such as squid and shrimp are collectively referred to as nekton, a term that refers to their ability to move effectively against the currents. Much of the life in the ocean is confined to about the first 600 feet or so (about 200 meters), where light penetrates. If phytoplankton is abundant, light will attenuate at a more shallow depth and, if phytoplankton is in low density, such as in some tropical seas, light will penetrate more deeply. Animals, some of which migrate to the surface from deeper waters on a nightly basis, concentrate in the surface waters because thats where most available energy is found. Phytoplankton (or any other plants) cannot photosynthesize beneath the euphotic zone. Once your diving bell passes through the last twilight of the euphotic zone you have entered the permanent darkness of the mesopelagic zone. You would notice far fewer kinds of fish, but what you would see would astound you. Even in the darkness you would see some of the fish, because they light up. Numerous species are bioluminescent, illuminated by tiny lights similar to those found on fireflies (which are not really flies at all, but beetles). Bioluminescence is a form of cold light created by the oxidation of certain protein molecules. Most of the bioluminescent fish are tiny, no more than the size of a quarter. These include schools of lantern fish and hatchetfish. Joining them in the water column would be various squid and shrimp. With luck, a huge sperm whale might dive past you as it plumbs the depths in search of its principal prey, the giant squid. Deeper still, your diving bell is withstanding many hundreds of pounds of pressure per square inch as you enter the bathypelagic zone, the deepest part of the ocean, profoundly dark and very cold. As is the case with the 62

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mesopelagic zone, the permanent inhabitants of this zone must rely on organic matter from the euphotic zone to eventually drift downward. A fish that dies in the surface waters is consumed well before it strikes the bottom several miles below. You might see the skeleton of a tuna or swordfish drifting past your diving bell but little more of the deceased creature. Energy is at a premium at such a great depth. Some amazing fish inhabit the deepest depths of the oceans. Many are small, their color predominantly black, though with bioluminescent patterning, particularly in the head region. Some, popularly called swallowers and gulpers and viperfish, have immense jaws with needle-like teeth. A swallower has an abdomen that can expand to hold a fish longer than it is. There are also species of deepwater sharks and rays as well as deepwater squid and octopus. Deep-sea anglerfish are chunky, with huge mouths that snap up prey attracted to the bioluminescent lure that they wiggle to entice prey to within capture distance. Some species of deep-sea anglerfish have a curious life cycle such that males transform into parasitic worm-like animals that attach to the body of a female anglerfish. This odd characteristic is adaptive and points to an interesting ecological characteristic of the bathypelagic zone. It is adaptive because these animals, like most of the bathypelagic creatures, are relatively rare, existing in small populations within an immense volume of habitat. In an environment with so little organic matter, most populations will be limited. When a male angler Illustration of a humpback anglerfish attaches to a female and (Melanocetus johnsonii) becomes an ectoparasite, at least Source: Brauer, A. Die Tiefsee-Fische: Tiefsee-Expedition the female has a source of sperm Valdivia, 189899. Jena: Berlin, 1906. accompanying her. When she sheds eggs, she need not search the vastness of the ocean depths for a mate. Hes already attached to her. Benthic Zone The R.M.S. Titanic, subject of several popular motion pictures, many books, and an ambitious and successful underwater search, rests on the benthic zone of the North Atlantic Ocean. The term benthic refers to bottom. It can be applied widely, to lakes, shallow bays, or the ocean depths. Once it was believed that no life could exist on the ocean bottom. It seemed incredulous that an environment so deep, so permanently cold, with no light, and under thousands of pounds per square inch of water pressure, could support living things. Such a view was, to say the least, nave. Another fanciful idea was that the ocean bottom was uniformly covered by a curious organic jelly called Bathybius, a living organic slime of sorts. Though such a suggestion may sound like the plot of an old episode of Star Trek, it required 63
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a serious oceanic expedition, the Challenger Expedition, from 1872 to 1876, to disprove it. When your diving bell has reached bottom, you observe the soft sediments that cover the ocean floor, sediments that are far younger than the ocean itself, many of them the countless tiny shells of pelagic forms of amoebas comprising part of the permanent zooplankton. There are organisms moving about or protruding from the sediments. You might see such things as brachiopods, clam-like creatures that were far more diverse and abundant in the Paleozoic Era, before the dinosaurs evolved. Or waving serenely in the slow bottom currents, you might see crinoids, plant-like, stalked echinoderms related to sea stars and sand dollars, whose ancestry also reaches back to the very appearance of multicellular animals in the fossil record. The ocean floor may be virtually covered with active brittle stars or bizarre-looking sea cucumbers, all roaming the ocean floor. Perhaps you might see the odd-looking tripod fish, which supports itself on two spiny extensions of its pectoral fins and one from its caudal (tail) fin, making it look like a tripod. Should you find a dead whale or other carcass on the sea bottom, it would be swarming with opportunistic scavengers. Among the oddest of the deep sea fish, hagfish are jawless fishes that may be among the only remaining lineages of some of the first of the vertebrates. Their closest relatives are lampreys, also jawless, some of which inhabit fresh water. Hagfish are also called slime eels because they exude a dense mucous through skin pores. They are extraordinarily flexible, capable of tying themselves in knots (and untying themselves). Hagfish mass at carcasses of dead organisms and the thick mucous that sometimes clouds the water probably is adaptive in discouraging other competitors such as sharks from consuming the carcass. In addition, perhaps you would see elongate fish called grenadiers or ratfish that compete with hagfish for access to whatever organic matter they can find. Even at the greatest depths of the oceans, you can study ecology. Littoral Zone Coastal areas, where the presence of continents makes the sea shallow, support complex ecosystems. The littoral zone, a term that can apply to lakes and estuaries as well as to oceans, is the shallow area at the edge of a body of water, where light penetrates all the way to the bottom, making photosynthesis possible throughout the water column. For this reason, macroscopic plants such as kelp can grow in abundance, joining phytoplankton in capturing solar radiation. The littoral zones of the worlds oceans support many kinds of ecosystems. These include kelp forests, prime fishing grounds such as the Grand Banks of the North Atlantic, and even tropical coral reefs.
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Littoral zones are ecologically diverse and productive because physical conditions are conducive to biodiversity. Not only is there light throughout most, if not all, of the zone, but proximity to shore means that nutrients are regularly introduced, transported from rivers to the sea, fertilizing the shallow waters, making them more productive. Currents and tides mix the water, ensuring availability of nutrients and an abundance of dissolved oxygen. Many kinds of fish and invertebrates inhabit littoral waters, as well as cetaceans and seabirds. The littoral zone is a prime area of reproduction for 64

many forms of life in the sea, and shallow waters can serve as ideal nurseries for juvenile fish. Offshore currents help distribute larval forms of fish and invertebrates, including many that inhabit the intertidal zone. Intertidal Zone The place where sea meets land is the intertidal zone. The definition refers to the area exposed between low and high tide. Intertidal zones are composed primarily of marine organisms, including some of the hardiest examples. Like the littoral zone, it is an area of diverse habitats. It may be rocky, sandy, or composed mostly of mud, exposed at high tides. Intertidal zones may be associated with such coastal ecosystems as salt marshes and tropical mangrove swamps. Rocky intertidal zones have many kinds of algae, often called seaweeds, among which are various barnacles, mussels, crabs, snails, and other lifeforms. There is usually a fairly clear zonation from low to high water, with only the hardiest creatures able to tolerate prolonged exposure to air. Just look at a dock piling exposed at low tide, for example, and you will likely see brown and red algae still partly submerged and partly mixed with mussels, above which, totally exposed, will be a zone primarily of barnacles. Sandy and muddy intertidal zones are A rock on a beach near Kalaloch, Washington. frequently too unstable to support The rock, seen at low tide, exhibits typical intermany organisms on their surfaces, but tidal zonation. they both have a diverse infauna of burrowing worms and mollusks as well as others. In some areas, however, there are impressive forests of kelps among which there are many kinds of fish and invertebrates. Shallower waters support plants such as eelgrass, one of the only vascular plants (thus not an alga) that thrives in salt water. Eelgrass flats are often associated with salt marshes and support many kinds of marine organisms, including such odd fish as seahorses. Intertidal zones have many of the same ecological advantages as littoral zone habitats: an abundance of oxygen, light, and elemental nutrients well mixed by the tides. But intertidal zones are also subject to intense natural disturbances, as tides fluctuate and as storms batter coastal areas. Thus there are always areas of disturbance and recolonization within intertidal zones. The life cycles of organisms have evolved to adapt to such realities.

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FOR GREATER UNDERSTANDING

Questions
1. What are the major zones of the sea and how does zonation by depth compare with factors that determine terrestrial biomes? 2. What areas of the oceans are most productive and support the highest biomass, and why?

Suggested Reading
Connell, Sean. Marine Ecology. Oxford: University of Oxford Press, 2007.

Other Books of Interest

Bertness, Mark D., Steven D. Gaines, and Mark E. Hay, eds. Marine Community Ecology. Sunderland, MA: Sinaur, 2000. Hardy, Sir Alister. The Open Sea: Its Natural History. Boston: Houghton Mifflin, 1970. Russell, F.S., and C.M. Yonge. The Seas. London: Frederick Warne, 1975.

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Lecture 13: Unique Coastal Ecosystems

The Suggested Reading for this lecture is Mark D. Bertnesss Atlantic Shorelines: Natural History and Ecology.

here are a number of marine ecological systems that appear only in specialized places on the Earth. Their uniqueness is due to such forces as the latitude, landform, wind, and ocean currents. Other factors may be the nearness of freshwater estuaries, extreme tidal variations, and human intervention. Salt Marsh A marsh is defined as a wetland where grasses and sedges predominate. Freshwater marshes are often characterized by such species as cattails (Typha spp.). A salt marsh, as the name implies, is a coastal marsh where tidal influence is strong. Consequently the marsh is regularly inundated by salt water. However, a salt marsh is also affected by fresh water carried by rivers as they flow to the sea. Thus a salt marsh has brackish water, its waters carrying variable concentrations of salts, depending upon the degree to which fresh and salt water have mixed in the marsh and depending upon the tidal cycle. Salt marshes are usually in close proximity to estuaries. The combination of marsh and estuary is the ideal habitat for many kinds of marine organisms, particularly juvenile life-cycle stages of fish and invertebrates that will later join the fauna of the open sea. Thus salt marshes have a valuable role in the conservation of marine fisheries. The ecological worth of salt marshes has not been appreciated until relatively recently. Both on the East and West coasts, salt marshes were historically regarded merely as havens for mosquitoes. Consequently, they were extensively drained and converted to various human uses. The so-called Back Bay region of Boston, Massachusetts, now home to many thousands of people, is well named. It was once a bay, extensively bordered by salt marsh. Those

Bride Brook and Coastal Salt Marsh, near East Lyme, Connecticut

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ecosystems were long ago either filled or drained or some combination of both. A visitor to Boston today would never suspect that much of the city was once bay and salt marsh. The same story is true of much of San Francisco, California. Salt marshes are characterized by grasses and sedges. These species are collectively called halophytes, as they are tolerant of high concentrations of salt. Salt marshes in eastern North America consist mostly of Spartina grasses. One species, Spartina alterniflora, called salt marsh cordgrass, is the most physiologically hardy of any of the marsh grasses and thus tends to dominate areas of highly salty water, where tidal influence is greatest. Salt marsh cordgrass grows well under virtually anoxic conditions and with high salt exposure. No other salt marsh plant can survive under such oxygen limitation, thus salt marsh cordgrass is able to persist by virtue of its physiological ability to adapt to extremely low oxygen levels. When in competition with other salt marsh grass species under controlled conditions, where the substrate is not anoxic, cordgrass is easily outcompeted. A closely related species, S. patens, called salt meadow hay, is usually found growing at higher elevation, beyond alterniflora. Mixed among the S. patens is spikegrass (Distichlis spicata) and various glasswort species (Salicornia spp.). Other species grow in zones higher in elevation, beyond patens, thus, like is the case with the intertidal zone, salt marshes show a pattern of zonation as tolerance levels to saline water varies among the resident species. Beyond the salt hay zone there is typically a zone of black rush (Juncus gerardi) followed by a zone of marsh elder (Iva frutescens). Salt marshes of the Pacific coast also are characterized by Spartina at the waters edge, but the species is S. foliosa. It is followed by a similar zonation pattern to that seen in the East, though the species are somewhat different. Salt marshes, though low in plant biodiversity, are highly productive ecosystems, accomplishing much photosynthesis throughout the growing season. In addition to the salt marsh grasses and related plants, many minute diatoms coat the mud with a golden sheen. Their combined photosynthesis further enhances the overall marsh productivity. The high productivity of the salt marsh supports many kinds of invertebrates (crabs, shrimp, snails, mussels, worms) as well as animals such as minks, raccoons, and numerous bird species such as herons, egrets, rails, and shorebirds. Many insect species, particularly such nuisances as mosquitoes and biting flies, also abound in salt marshes, where they are fed upon by various dragonflies and birds such as tree swallows (Tachycineta bicolor) and purple martins (Progne subis). Fiddler crabs are common in salt marshes. Males have one claw that is much larger than the other, giving the group its general name fiddler crab. Studies have demonstrated that fiddler crabs have a mutualistic relationship with salt marsh cordgrass. The extensive burrow network of the crab colony aids in bringing aeration to the roots as well as slowing down the accumulation of dense sediment (called peat), which limits the growth of the cordgrass. The presence of the stand of cordgrass acts to stabilize the mud, a huge advantage for the crabs. Without the cordgrass, the tidal currents would continually agitate the mud, making it difficult for the crabs to have a stable colony. 68

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Salt marshes export much of their productivity to neighboring estuaries. Grasses that are products of the growing season die back in winter, surviving the cold and ice of winter as roots in the mud. The above-ground shoots, often sheared off by ice or tidal forces, are moved by tidal action, eventually reaching estuaries where they are colonized by a host of microbial organisms that form a food resource for larval invertebrates and fish. Thus the marsh grasses are also an important energy resource for the estuary. Salt marshes act as natural sponges, absorbing nutrients, which are recycled to the growing marsh plants. Some organisms such as the ribbed mussel (Geukensia demissa) are keystone species in concentrating nutrients. These filter-feeding mollusks, which grow abundantly among the salt marsh cord grasses, feed by filtering suspended particles such as plankton from water during periods of immersion. In doing so, they remove and concentrate nutrients such as nitrates and phosphates. These are deposited on the marsh mud in the form of compact pseudofeces that serve as small pellets of fertilizer. Many marsh species, including various clams and oysters, also filter water to remove suspended food particles. Pollutants, ranging from pesticides such as DDT to heavy metals such as mercury, are also removed by organisms and concentrated in the marsh. This is the mode by which these esoteric and often highly dangerous compounds begin to bioconcentrate as they enter natural food webs. Natural pollution products such as the toxins produced by minute algae (dinoflagellates) responsible for red tide also can bioconcentrate in marshes due to activities of filter feeders. Mangrove Forest Mangroves are a diverse assemblage of coastal tropical tree species, all tolerant of high salt concentration. There are thirty-four species worldwide, mostly distributed throughout the vast tropical Pacific. A few species occur abundantly in tropical America. Mangroves are not a taxonomic group any more than all salt marsh plants are, instead representing numerous plant families. Mangroves are all defined by their physiological tolerance to immersion in salt water and thus are the tropical equivalent of salt marsh grasses, sedges, and associated plant species. Like salt marsh plant species, mangroves are halophytes. Mangrove forests typify tropical coastlines and may line rivers where there is tidal influence. Like salt marsh grasses, they thrive in high saline environments, but are outcompeted by plants in low salinity areas, where otherwise mangroves could survive. In general, mangrove species are excellent colonizers and quickly invade and grow in areas subjected to the effects of tropical storms such as hurricanes. Most mangroves are small trees, rarely exceeding 10 to 20 meters (33 to 66 feet) in height. Some grow as dense shrubs. Mangrove root systems grow well in soft coral sands as well as thick, anoxic mud. The forest itself provides a complex coastal habitat for many species of animals. Red mangrove (Rhizophora mangle) is an abundant species in tropical regions throughout the world. It typically has extensive prop roots anchoring it firmly in the shifting sands. The roots are abundantly equipped with lenticels, openings that admit air needed for the roots to survive when immersed. A superb colonizing species, red mangrove seeds actually germinate while still 69

attached on the parent plant. The seedlings look like long, pencil-like pods hanging vertically from the tree. Once dropped in water, they can survive for a long period, transported by the currents. As a seedling absorbs seawater it soon floats vertically rather than horizontally, bobbing with the current until it strikes land, where it immediately grows a root to anchor it. Red mangroves form dense stands on exposed sand and mudflats, taking advantage of sediment deposition by coral reefs. As other mangroves colonize, a new mangrove island, or caye, is formed. Black mangrove (Avicennia germinans) often grows densely in thick, anoxic mud. It manages to secure sufficient oxygen because its shallow, horizontal roots send up vertical shoots, called pneumatophores, which allow adequate amounts of air to enter the root system. The prop roots of mangroves provide excellent habitat for many marine animals. Various sponges, anemones, tunicates, mollusks, and worms find purchase among the roots. Numerous fish and crabs find shelter and food as well. Many species of coral reef fish spend part of their juvenile life cycle among the mangroves. In this manner, mangrove swamps function like salt marshes and estuaries, providing nurseries for juvenile marine animals. Similarly, mangrove leaves, thick and waxy, eventually drop on to the sediment or into the sea, where they provide an essential energy source. Red mangrove is highly productive, its rate of photosynthesis so high that it can add up to eight grams dry organic matter per square meter per day. How is this energy utilized? Once leaves drop, they act as sites for microbial colonizers (bacteria, fungi, various protozoans) that, in turn, serve as a food base for larval sea animals. A fish can ingest a particle of mangrove leaf, digest the various organisms on it, and spit out the leaf particle or allow it to pass through its alimentary canal. In either case, the leaf fragment now back in the water is soon recolonized and serves yet again as a food base for microbes. Thus the energy of the mangroves is moved, in small and continuous amounts, into sustaining the marine food web of the coral reef and surrounding seagrass. It is essential to recognize how the food webs of what appear to be various different kinds of ecosystems are, in reality, closely linked.

LECTURE THIRTEEN

Above- and below-water view of mangroves in Bangladesh.

Mangrove knees (pneumatophores) at low tide on the Yap Islands of Micronesia in the Pacific.

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Left: US National Oceanic and Atmospheric Administration Right: Photograph courtesy of Eric Guinther

Mangroves, like salt marshes, have high ecological value. It is thus regrettable that throughout the world mangrove forests are being cleared by such activities as dredging, channeling, or cutting as a source of wood. Huge tracts of Southeast Asian mangrove forest were destroyed by herbicide spraying during the Vietnam War, over three decades ago. In addition to serving as essential nurseries for many marine species, and in buffering the land against the forces of tropical storms, mangrove forests form important nesting areas for tropical waterbirds such as pelicans, boobies, herons and egrets, spoonbills, and frigatebirds. In addition, long distance migrant landbirds often overwinter in the food-rich mangrove forests. Mangrove forests persist in North America in southern Florida and along the Florida Keys. The endangered American crocodile (Crocodylus acutus) is one of many species dependent on the mangrove ecosystem. Coral Reef Coral reefs are found throughout the clear marine waters of the worlds tropics. Among the oldest ecosystems on Earth, coral reefs, though not with the same assemblages of species found today, have existed virtually since the Cambrian Period, over 500 million years ago. Reef organisms are among the most abundant in the marine fossil record. Today, coral reefs support the highest species richness of any marine ecosystem. Like their terrestrial counterparts, tropical rainforests, corals reefs support myriad species and accomplish high rates of photosynthesis. Reefs are both diverse and productive. All coral reefs are confined to clear, warm tropical seas. Cool water or sediment deposition will kill coral reefs. Coral reefs require a temperature of at least 18 C (64.4 F). Most coral species are in Indo-Pacific waters, where over 700 species can be found. In comparison, Atlantic reefs are much less species rich, with only about sixty species. This difference in species richness between the older and larger Indo-Pacific tropical oceans and the Atlantic/Caribbean is true of fish as well. There are about 500 fish species in the Bahamas, compared with 2,000 in the Philippines and 1,500 in Australias Great Barrier Reef. The difference in richness between the two

A pillar coral at the Florida Keys National Marine Sanctuary.

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oceans likely is scale related with regard to both time and area. The IndoPacific is a much older ocean, with considerably greater area than the Atlantic. Both of these factors would suggest more speciation in the Indo-Pacific. However, while the total species richness of fish is lower in the Caribbean than in the Pacific, the number of fish species that can be found, per unit area, within a given reef in either region is about the same. In other words, you would find approximately equal numbers of fish species per 100 meters of coral reef in the Caribbean and Pacific, but in the Pacific, as you moved from one reef to another, you would find new species. In the Caribbean, you would find many of the same species you had already encountered. Coral is a kind of animal related to sea anemones and, somewhat more distantly, to creatures such as jellyfish and hydras. They are part of the phylum Cnidaria (sometimes called Coelenterata), known for their tentacles that possess cnidoblasts or stinging cells. All reef corals are colonial, growing together in matrixes that are composed mostly of calcium carbonate secreted by the animals themselves. The reef structure is also augmented by algae that secrete calcium carbonate and, in the Indo-Pacific, by giant clams. Corals are cup-like, tiny animals called polyps that capture minute organisms suspended in the water column, using their tentacles with stinging cells.

Anatomy of a coral polyp.

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Reef corals have an extraordinarily high primary productivity, fixing about as much carbon as rain forests that have huge amounts of plant biomass. But where are the plants on the reef? Corals are, after all, animals. The answer to why it is that coral reefs are so productive can be sensed by looking at the shapes of the corals themselves. In shallow, well-lit waters, they resemble the branching patterns so typical of plants. A close look within the coral matrix reveals the presence of numerous tiny, one-celled plants called zooxanthellae. These plants and their coral hosts are mutualistic in their interaction toward one another. Zooxanthellae augment the growth of the coral colony and supply much carbohydrate because they are active photosynthesizers. Measurements of the intake and output of carbon dioxide on a reef indicate that corals with zooxanthellae, which are known as hermatypic corals, are, for all intents and purposes, functioning physiologically as plants. They take in more carbon dioxide than they respire, all due to the presence of the zooxanthellae. Coral reefs are structurally complex ecosystems, characterized by both horizontal and vertical zonation patterns. Horizontal zonation is determined by the intensity of wave action from the windward to the leeward side of the reef. Vertical zonation is a result of depth and light penetration. Reefs may form as 72

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fringing reefs, barrier reefs, or atolls. Fringing reefs typically surround islands, built by the corals in the shallow sediments where the island rises. Barrier reefs run parallel to coastlines, and atolls are circular reefs that remain where an island, now subsided, once existed. In one of his first major scientific works, Charles Darwin correctly surmised that the various kinds of coral reefs all form from the same basic process of coral growth. In other words, an atoll was once a fringing reef but, as the island subsided, the activity of the millions of coral animals maintained the reef at the waters surface, even as the island disappeared beneath the sea. Ecologists have learned that corals are highly competitive with one another. Like plants, some corals overtop and shade out other species. Still others poison their competitors when in direct contact with them. Given the clear competition among coral species, it may seem surprising that reefs can maintain a high species richness of corals, but they do. The reason is that disturbance is a relatively constant feature of coral reefs. Disturbance by wave action or storms (such as hurricanes) can quickly change the dynamics of competitive interactions on reefs. In this way, coral reefs are similar to ecosystems such as rainforests, where periodic disturbances of varying magnitudes are a key component to the persistence of high species richness. Some disturbance agents can reek havoc on coral reefs. One in particular is the sea star (starfish) Acanthaster planci, the socalled crown-of-thorns. Population outbreaks of this species can result in the destruction of whole reefs. It is unclear what causes such outbreaks. Like rainforests, coral reefs are being lost due to various human activities. In many areas, over-fishing is a major problem, but other activities such as dredging and mining can cloud the water to the degree that corals can no longer endure. Chemical pollution is also of major concern. In recent years it has been learned that meteorological events such as El Nio negatively affect coral reefs. El Nio is a periodic and generally unpredictable change in global climate caused by the migration of a high-pressure system in the central Pacific. Patterns of current flow change, precipitation patterns change, and the result is that some ecosystems suffer serious negative impact. Unfortunately, coral reefs do badly in El Nio years. Globally, many scientists believe coral reefs are generally in decline.

Colorful reef fish Pennantfish, Pyramid butterflyfish, and Milletseed butterflyfishschool in great numbers at Rapture Reef, off the northwestern Hawaiian islands.

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FOR GREATER UNDERSTANDING

Questions
1. What are the most essential ecological functions of salt marsh and how are these functions similar to mangrove forest processes? 2. Why is the coral reef so species rich and so highly productive? What are the current threats to coral reef ecosystems?

Suggested Reading
Bertness, Mark D. Atlantic Shorelines: Natural History and Ecology. Princeton, NJ: Princeton University Press, 2006.

Other Books of Interest

Kaplan, Eugene H., and Susan L. Kaplan. A Field Guide to Coral Reefs: Caribbean and Florida. Boston: Houghton Mifflin, 1999. . A Field Guide to Southeastern & Caribbean Seashores. Boston: Houghton Mifflin, 1999. Shumway, Scott. The Naturalists Guide to the Atlantic Seashore: Beach Ecology from the Gulf of Maine to Cape Hatteras. Helena, MT: Falcon, 2008. Teal, John, and Mildred Teal. Life and Death of the Salt Marsh. New York: Ballantine, 1991.

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Lecture 14: Current Issues in Global Ecology

The Suggested Reading for this lecture is Edward O. Wilsons The Future of Life.

cology is sometimes called a We must nd new ways subversive science because it to provide for a human seems to commonly espouse society that presently has views antithetical to wellestablished disciplines such outstripped the limits of as economics. Conservation global sustainability. can historically be viewed as a sociopoliti~Peter Raven, President cal movement that places intrinsic value on American Association for the nature. Ecological science was not foundAdvancement of Science, 2002 ed to promote conservation, though ecologists typically have sympathies toward the natural environment. As ecology has matured as a predictive discipline, approaches to conservation have become increasingly based on science and thus ecology has been at the forefront of such approaches. The field of conservation biology is now recognized as one of the sub-disciplines of ecology. Ecologists have also developed applied and restoration ecology as sub-disciplines of ecology, so ecological principles are being widely applied in ways that were not imagined a few decades ago. Conflicts arise about how best to utilize, value, and preserve what are perceived as natural environments. Humans, at least in Western culture, often consider themselves as somehow apart from nature, rather than part of the

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evolutionary process from which nature results. The philosophical roots of the Western view of nature are historically deep and may be at least partly explained by the traditional Judeo-Christian theological view of man and nature. Technology driven by science and economics coupled with human population growth have exerted strong anthropogenic effects on global ecosystems. This has resulted in a global loss of topsoil and agricultural land, depletion of the ozone layer, addition to the atmosphere of greenhouse gases, and ongoing major losses of biodiversity. Humans demand a uniquely large percentage of Earths net primary productivity and renewable fresh water. Earth can be viewed as a global ecological commons, where resources such as soil, air, water, and wildlife are the common resources of humanity. Concerns are focused on how best to manage global resources for sustainability. The worlds fisheries are in dramatic decline because, as fundamentally a commons-type resource, there has been inadequate study, regulation, and agreement about how best to manage fisheries. Different nations take different approaches, but stocks continue to decline. It is necessary to abandon the concept of a free commons and replace it with one in which all interested parties are subject to strong regulation regarding degree of exploitation. The major environmental problem facing citizens in this century is how best to ensure that Earths ecosystems remain sustainable such that we continue to enjoy the diverse ecosystem services essential to the functioning of the biosphere. Economic analysis of natural ecosystem function compared with gains derived from human development demonstrates that it is often in the long-term economic best interest of society to preserve natural ecosystems. Environmental ethics has emerged from the pragmatic need to recognize and value the natural environment and its component species. Still in its infancy as a discipline, environmental ethics attempts to rigorously provide a rationale for how applied conservation biology ought to be practiced to the collective good of humanity. Biodiversity loss is arguably the single greatest problem in conservation biology. Biodiversity can be expressed as numbers of species, genetic diversity among populations, or ecosystem diversity. It is not known how many species actually inhabit Earth today, but it is clear that biodiversity is in decline. The current rate of extinction far exceeds the current rate of speciation, so Earth is losing species. There are many causes for biodiversity loss, including habitat loss, pollution and contamination, and over-exploitation, such as is seen in modern industrialized fisheries operations or the decline of great ape populations in Africa. Another major cause for biodiversity loss is the increasing number of invasive species that, by various means, are entering ecosystems in which they did not evolve. These exotic species often out-compete or predate native species to the point of driving them to extinction. Global climate change is also affecting diversity patterns. Natures services, the natural ecosystem functions upon which all life, including humanity, depends, are diverse and numerous, but their importance was not really recognized until recently. Ranging from purification of air and 76

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water, cycling and movement of nutrients, generation and preservation of soils, and renewal of soil fertility, to seed dispersal, pollination of crops and other vegetation, to maintenance of biodiversity, it is clear that the functioning of natural ecosystems is essential to human welfare. But how essential? If put in the light of economic analysis, just how much, in dollar value, are natures services worth? A team of researchers led by Robert Costanza (Costanza, et al., 1997) used numerous databases to estimate that natures services, presuming that humans had to somehow pay for them, would be worth about 33 trillion dollars annually. It is noteworthy that this estimate is about double the gross world product, the total of all gross national products, which, at that time, was about $18 trillion. In 2002, updated estimates suggested a rough average of $38 trillion, with a range of between $18 and $61 trillion. The wide range reflects the extraordinary difficulty of attempting to measure the macroeconomic worth of all ecosystem goods and services. Some economists have criticized the effort by Costanza et al. for a number of reasons, including the assertion that the extrapolations made were inconsistent with principles of microeconomic theory (Balmford, et al., 2002). In response, a team of nineteen researchers, including economists and ecologists, examined over three hundred specific case studies in an attempt to compare what they termed marginal values of goods and services delivered by a biome when relatively intact, and when converted to typical forms of human use. Robert Costanza, who authored the previous study, was part of the research team. Their results were consistent in showing that natural ecosystems have the potential for greater societal economic gain than do ecosystems converted for narrow economic objectives. In one example it was clear that reduced impact logging in Malaysian tropical forests did not provide the immediate economic benefits to individuals that would be obtained by high-intensity unsustainable logging, which is what is normally done there. However, unsustainable logging reduced social and global benefits through loss of forest products (other than timber), flood protection, carbon stocks, and endangered species. The total economic value of the forest was about 14 percent greater when managed to be sustainable, using reduced impact logging techniques. In a second example, a mangrove ecosystem in Thailand was converted to aquaculture (shrimp farming), something that is occurring in many places in the worlds tropics. There was no question that short-term economic interests were well served by such a conversion. However, when social benefits of leaving the mangrove ecosystem intact were included in the economic analysis, the result changed. Benefits such as the sequestration of carbon, storm protection, and protection for fish added much more value to the intact mangrove forest than to the aquaculture ponds. The estimate showed that the intact mangrove forest was worth about 70 percent more than the aquaculture. Most ecologists agree that climate change is the most significant factor now influencing Earths ecology. Climate change has always occurred throughout Earths history for many reasons that have nothing whatsoever to do with humans. But todays climate change is unique in that human activities do, in

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fact, seem to be forcing much of the climate change, much of which is due to greenhouse effect. It is well known that greenhouses admit light and trap heat. Some atmospheric gases have an analogous effect to that of the glass in a greenhouse. Gases such as water vapor, methane, nitrous oxide, and carbon dioxide act to block heat energy from passing easily through the atmosphere. Rather than a quick, easy transit from Earth to space, the heat is trapped, retained within the atmosphere for a relatively long period. The more greenhouse gases there are, the more this effect of trapping heat is manifest. This phenomenon, the retention of heat energy by certain atmospheric gases, has come to be termed the greenhouse effect. It is tremendously important in mitigating rapid temperature fluctuations on Earth and it has contributed in an essential way to making the Earth a habitable planet. Recall that Earth is an example of the Goldilocks effect, situated at precisely the right distance from the Sun for water to exist in liquid form. One profound benefit of the presence of oceans of liquid water is that greenhouse gases, and in particular, carbon dioxide, can be absorbed into the oceans, and, in the case of carbon dioxide, by a series of purely physical reactions converted to insoluble carbonate, taken out of circulation. The importance of this reality cannot be overemphasized. Without the oceans, any buildup in carbon dioxide, such as from volcanic emissions, for example, would not be correctable and the atmosphere would continually increase in carbon dioxide concentration that, in turn, would trap more and more heat. Eventually, this process would run away and the amount of heat trapped would be sufficient to raise the temperature of the planet beyond that which life could endure. Such is apparently the case with the planet Venus. Although the oceans can and do absorb carbon dioxide, it is clear that since the onset of the Industrial Revolution, atmospheric carbon dioxide concentration has increased steadily and today it is about 385 ppm (parts per million) by volume (compared with 315 ppm in 1960). This increase has correlated with the growing use of fossil fuel and, particularly in the latter part of the twentieth century, with increased global deforestation. These two factors have combined to release a significant amount of carbon dioxide, a process that is ongoing and which is altering the atmosphere to the degree that Earth is warming and climate changing. The result of climate change will be to alter ecosystem properties and species interactions globally. Some species, such as the polar bear of the Arctic, may be seriously endangered because of its dependency of shelf ice on which to hunt seals and give birth. Penguins in the Antarctic are becoming isolated between inland breeding colonies and coastal feeding areas, reducing their reproductive potential. Everywhere on the planet where ecologists measure populations or ecological processes, changes seem to be occurring, many of them negative, though not all. In a sense, this is an exciting time for ecology as it is challenged to respond to the realities of human culture and its collective and powerful influence on this, the ecological planet.

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FOR GREATER UNDERSTANDING

Questions
1. Why was ecology once described as the subversive science? What should be the relationship between ecology and economics? 2. What are the major factors that are today altering ecosystems on Earth? Which of these is likely to have the greatest influence?

Suggested Reading
Wilson, Edward O. The Future of Life. New York: Alfred A. Knopf, 2002.

Other Books of Interest

Botkin, Daniel B. Discordant Harmonies: A New Ecology for the Twenty-First Century. Oxford: Oxford University Press, 1992. Levin, Simon A. Fragile Dominion: Complexity and the Commons. NY: Basic Books, 2000.

Articles of Interest
Balmford, A., et al. Economic Reasons for Conserving Wild Nature. Science. Vol. 297, pp. 950953, 2002. Costanza, Robert, et al. The Economic Value of the Worlds Ecosystems. Nature. Vol. 387, pp. 253260, 1997.

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COURSE MATERIALS

Suggested Readings: Bailey, Robert G. Ecoregions: The Ecosystem Geography of the Oceans and Continents. New York: Springer, 1998. Bertness, Mark D. Atlantic Shorelines: Natural History and Ecology. Princeton, NJ: Princeton University Press, 2006. Connell, Sean. Marine Ecology. Oxford: University of Oxford Press, 2007. Cox, C. Barry, and Peter D. Moore. Biogeography: An Ecological and Evolutionary Approach. Oxford: Blackwell, 1993. Jones, Stephen R., and Ruth Carol Cushman. A Field Guide to the North American Prairie. Boston: Houghton Mifflin, 2004. Kricher, John. A Field Guide to Eastern Forests. Boston: Houghton Mifflin Company, 1998. . A Neotropical Companion: An Introduction to the Animals, Plants, and Ecosystems of the New World Tropics. 2nd ed. Princeton: Princeton University Press, 1999. MacMahon, James A. Deserts. New York: Alfred A. Knopf, 1985. Pielou, E.C. A Naturalists Guide to the Arctic. Chicago: University of Chicago Press, 1994. . The World of Northern Evergreens. Ithaca, NY: Comstock, 1988. Walter, Heinrich. Vegetation of the Earth: In Relation to Climate and the Eco-physiological Conditions. New York: Springer-Verlag, 1973. Ward, Peter D., and Donald Brownlee. Rare Earth. New York: Copernicus, 2000. Wilson, Edward O. The Future of Life. New York: Alfred A. Knopf, 2002. These books are available online through www.modernscholar.com or by calling Recorded Books at 1-800-636-3399.

COURSE MATERIALS

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