Tropical Ecology

thanks.

Introduction

What Is Tropical Ecology?

Asking the question, What is tropical ecology? may seem akin to asking questions such as, Who is buried in Grant’s tomb? Tropical ecology is the study of the ecology of tropical regions. But so what? Consider these questions: First, what is ecology? What are its paradigms, what are its principles, and what is its place in the biological sciences? What unique insights has ecology added to biology? Second and perhaps more important, is tropical ecology unique within the study of ecology? In other words, if you studied the ecology of the eastern deciduous forest, the arctic tundra, the Mojave Desert, or the seas that surround Antarctica, what paradigms and principles might you miss because none of these ecosystems are tropical? Take a few minutes to think about these questions, and then read on.

Readers of this book should be familiar with ecology and thus need little coaching as to what ecology is. It is, in a nutshell, the study of how organisms exist, adapt, and interact within their environments within the context of both abiotic and biotic factors. It is studied at widely different spatial and temporal scales. Ecologists reveal and attempt to explain patterns of species richness and explore how top-down and bottom-up forces influence the structure and stability of food webs, structuring whole ecological communities. Ecologists use experimental techniques and mathematical models to study competition, predation, mutualism, and the assembly, persistence, and stability of ecological communities. Ecology has matured in that disciplines such as landscape ecology, restoration ecology, and conservation ecology all use theory from basic ecology. Each of these areas of applied ecology has made an impact on issues in tropical ecology.

The various topics cited in the previous paragraph are the grist for ecological study and make up the usual suspects in ecology texts. Once one understands these topics, one is considered proficient in ecology. Do these topics form the paradigms of ecology? Not really. They are best considered principles, areas of study around which the discipline is organized. Ecology arguably has no paradigms. (I argue this point throughout my book The Balance of Nature: Ecology’s Enduring Myth.) On close inspection, ecology is a prominent branch of evolutionary biology. Not much of what ecologists study makes sense without placing the data firmly within evolutionary context. This reality will become apparent in the study of tropical rain forest ecology, especially regarding species’ origins, interactions, and interdependencies. The paradigm within which ecology is included is organic evolution, and it, along with cell theory, are the only real paradigms of biology. Cell theory and evolution are what biology is. All the rest is detail.

Where does this leave tropical ecology? What will you learn by going to Panama, or Ecuador, or Indonesia, or Queensland, Australia, that could not be learned by going to central Spain, or Norway, or Japan, or New Jersey? Yes, the species are different, but what of it? The pattern of seasonality is different; precipitation patterns are different; lots of things are different. What of it?

This what of it? is the subject of this book. By the conclusion of your study, you should be competent to voice an educated opinion regarding the question of just how unique tropical ecology is. Learn and enjoy the journey.

Overview

When Charles Darwin was fresh out of college, he visited a tropical rain forest in eastern Brazil as part of his famous voyage aboard the HMS Beagle. Like others who came before and since, his first impressions were vivid. He wrote of this experience in the tropical rain forest:

When quietly walking along the shady pathways, and admiring each successive view, I wished to find language to express my ideas. Epithet after epithet was found too weak to convey to those who have not visited the intertropical regions the sensation of delight which the mind experiences. (Darwin 1845)

In other words, Darwin was pretty overwhelmed by what he saw, felt, heard, even what he smelled. He commented on the hazy air within the forest, evidence of the high humidity. A tropical rain forest does leave you with a vivid first impression. It gets your attention both rationally and emotionally. But years later, a much older Charles Darwin had a different perspective when he was composing his autobiography:

In my journal I wrote that whilst standing in the midst of the grandeur of a Brazilian forest, it is not possible to give an adequate idea of the higher feelings of wonder, admiration, and devotion which fill and elevate the mind. I well remember my conviction that there is more in man than the mere breath of his body. But now the grandest scenes would not cause any such convictions and feelings to rise in my mind. (Darwin 1887)

What changed Darwin was, of course, evolution. When Darwin returned from the Beagle voyage, he quickly became convinced of the truth of organic evolution. All present species have arisen from previous species and were not specially created. This was soon followed by his discovery of the mechanism for evolutionary change, natural selection, and so he began to view the world differently. Of course, not everyone agreed with Darwin. Some rejected his theory outright; others agreed that evolution might be true but did not agree that a blind mechanism called natural selection caused species to evolve into different forms. From 1859, when On the Origin of Species was published, until Darwin’s death at the age of 73 in 1882, he was strongly attacked in speeches and in print over his assertions concerning evolution.

What does all this Darwinian history have to do with tropical ecology? The great twentieth-century evolutionary biologist Theodosius Dobzhansky, who spent a fair amount of time collecting fruit flies in the tropics, said Nothing in biology makes sense except in the light of evolution (Dobzhansky 1973, p. 449). Darwin’s big idea has become the most important paradigm in biology, and without it, the immense complexities that characterize a tropical rain forest or any other ecosystem would seem hopelessly beyond rational explanation.

Therefore, much of this book will be about evolution as it is evident in the tropics, and in rain forests in particular. There is nothing unique about this approach, as I fail to see how it could be otherwise for any kind of ecological study. If this book were about the ecology of Antarctica, evolution would still be the organizing paradigm. The noted ecologist G. Evelyn Hutchinson once titled a book about general ecology The Ecological Theater and the Ev olutionary Play. I doubt that many who visit a rain forest or any other tropical ecosystem today need fear losing their sense of awe (as Darwin apparently did) simply because of acquiring an understanding of how the process of evolution is manifest in the multitudes of morphologies and interactions that characterize the ecological community. Darwin, when elderly, apparently lost his awe of nature, perhaps embittered by all the criticism directed toward him. You need not worry, however. Learning the science behind the landscape really adds to awe—it does not detract from it. This book aims to teach the science behind the landscape.

As author of the book you hold, I am in a unique position. It’s my book, and I wrote it my way. Other books about the tropics take different approaches from what you will read within this volume. This book reflects my views from study and field experience (particularly in ornithology) over my four decades as an ecologist. Let me explain something of how the book came to be.

In 1989, Princeton University Press published my book titled A Neotropical Companion. The book was modest in size and had a bright green cover, which led to its nickname, the Little Green Book. It amazed me while writing that book how much wonderful information ecologists had garnered in their collective research efforts and how little of this really cool stuff was finding its way to more popular-based audiences, including into the hands and minds of students. My goal was to be selective and make some of that amazing information available.

The success of the Little Green Book led to a major revision and expansion of A Neotropical Companion that appeared in 1997, again published by Princeton University Press. This volume was bigger, had new illustrations and some color photographs, and, most importantly, covered more topics in greater detail, with much additional treatment of South America.

Eager to learn from my readers, I included my e-mail address in the revised volume, and many responded. I have had a few errors of both omission and commission pointed out to me, as well as many comments as to how the book could be improved. Some want more maps; some want more illustrations; some want greater emphasis on particular topics. But the most-repeated comment (the title of the book notwithstanding) is that the book is too narrow, as it covers only the Neotropics and omits the Old World tropics of Africa, Southeast Asia, and Australia/New Guinea. How does the Neotropics compare with other tropical regions of Earth? How are these regions biogeographically distinct, one from another? Are the ecological processes that characterize Neotropical ecosystems the same as those in other regions?

Thus, it seemed both to me and to my editors at Princeton University Press that it was time for yet another revision and expansion, this time going the full Monty toward a more comprehensive book covering the ecology of the global tropics. And that is how Tropical Ecology came to be.

Ah, but as always, the devil is in the details. My previous book was specifically about the Neotropics, and the vast majority of my field experience has been in that region. True, I have visited Africa (Tanzania), Southeast Asia (Indonesia), as well as the Australian tropics in Queensland and Darwin, but most of my reading and field study is from Central and South America and the Caribbean. And, I dare say, much if not most of the published literature continues to be from studies in the Neotropics. So I cannot pretend to take an even-handed approach to the global tropics. This book will expose my bias, as it reflects my confidence level in my expertise—and that bias remains clearly toward the Neotropics and, in particular, birds. But such a bias has a pragmatic component in that many students will likely use this text in conjunction with an actual field experience, a college course that includes a trip to experience a tropical region. And most North American college courses in tropical ecology visit some area within the Neotropics. My continued emphasis on that region will benefit such courses.

The organization of the work is mine, my way to tell the story of tropical ecology. But the information contained herein is largely the work of others, the thousands of researchers in the field and lab whose work, often repetitive and always exacting, deals with some aspect of tropical ecology. For the most part, I rely on primary source material, papers from journals—some of which, like Biotropica, publish only papers about tropical ecology, and some of which, like Ecology and American Naturalist, publish many papers that deal with the tropics. I also include many citations from Science and Nature, both of which publish numerous major papers on various aspects of tropical biology. Each topic covered is illustrated by selected examples, so obviously many worthy papers are not cited. My apologies to these authors, but this book is not meant as a comprehensive review of all topics tropical. It is nice to have an embarrassment of riches when it comes to information about the tropics, and I have had to carefully pick and choose. Examples were by and large chosen for their overall interest, potential significance, clarity, and robustness. And, since researching the second edition of A Neotropical Companion, I am amazed by the remarkable increase in the number of tropical studies represented in professional publications of all kinds. Another point worth emphasizing is that my area of expertise is inornithology, the study of birds, so whoever reads my text will encounter quite a few examples that involve birds. But consider that birds are fairly easy to observe in the field, and thus much data about species interactions in the tropics does, indeed, focus on birds.

The first question any author faces is, Who is the audience for this book? This work is aimed at students and others who take a serious interest in learning about tropical ecology. It assumes a basic knowledge of introductory biology, including ecology and the tenets of evolutionary biology. I present an overview of each topic and then note where information gaps exist and where controversies have arisen. The book is meant to be provocative and poses many questions, the answers to which are still to be revealed or are presently being hotly debated. Some of the studies I describe come to different conclusions about the same topic. Science is always a work in progress, and this book strongly takes that approach.

All science books should be subject to peer review, and this book certainly has enjoyed that advantage (see the Acknowledgments). Others, expert in various areas of tropical biology, have read chapter drafts and made numerous critical comments. I have taken many of these suggestions, but I have also chosen not to take some. No two tropical biologists would ever write the same book. Our views are influenced by our biases, varied interests, and experiences, to say nothing of our intellectual outlooks. We make judgments, and we often have to agree to disagree on various points. Any problems identified with this text should be directed to me (jkricher@wheatonma.edu), and I take sole responsibility for any errors that may be detected.

Some material that has remained robust has been taken and often updated from A Neotropical Companion, but most of the material is new to this book. That said, any book such as this is old before it is published. Right now, any number of good studies are occurring or have occurred that alter some of what is written on these pages. That is the nature of science, the strength of science, the very essence of science. Science is cumulative, and anyone entering into the study of a field of science profits from acquiring a firm foundation from which to begin critical study. I hope this book will serve that purpose for its readers.

Organization of This Book

This book consists of 15 chapters, a challenge for a one-semester course, but hopefully the writing is sufficiently engaging that the task of moving through the chapters will prove to be relatively pleasant. There are numerous figures, maps, tables, and color photographs throughout to add to the clarity of the text.

I begin (Chapter 1) with a broad overview of what and where the tropics are. This provides some information about how tropical ecology emerged from global exploration into the realm of scientific study. It also describes something of the climatic variables that are the overarching determinants of tropical ecosystem characteristics, and it sets the stage for what is to come throughout the book.

Next (Chapter 2) I discuss biogeography and evolution in the tropics. For those already well versed in these topics, this chapter may serve as a helpful review. But for many, and I expect most students, it will be essential in explaining how and why tropical regions differ, as well as in clarifying basic evolutionary principles, particularly speciation.

Then it is time to enter a rain forest (Chapter 3) and examine the physical structure of this unique ecosystem. This chapter discusses the various characteristics of rain forest vegetation, from tree shapes to buttressed roots. It should be the primer that will allow any student to see the many details and general characteristics common to all rain forests.

Chapters 4 and 5 both explore ideas as to why tropical regions and rain forests in particular are so rich in their numbers of species. Biodiversity is one of the most significant areas of tropical research, and thus two chapters are devoted to research focused on this complex topic. Chapter 4 discusses how diversity is partitioned within and between habitats and considers whether the high diversity of tropical regions is an example of high rates of speciation (or low rates of extinction). What factors allow the continued existence of such complex communities? Chapter 5 looks at the range of hypotheses attempting to explain the uniquely high tree species richness in tropical forests, one of the most difficult theoretical problems in tropical ecology.

The effect of disturbance is explored next (Chapter 6). Periodic natural disturbance may result in a shifting mosaic of different aged patches of forest such that high species richness is maintained over large areas. This chapter deals in part with gap phase dynamics. Gaps in a forest are created by forces ranging from single fallen trees to major blowdowns caused by storms or fire. Gaps are exposed to high levels of sunlight and thus have a different ecology from closed rain forest, where canopy trees strongly attenuate the light striking the forest floor. Last, this chapter considers the topic of secondary succession, which is of major importance in assessing how rain forest regrows following disturbance.

The richness of species in tropical forests results in numerous complex forms of interactions among species—some rather casual, some sufficiently complex that the species are evolutionarily interdependent. Chapter 7 presents examples of some of the most striking biotic interactions involving such topics as fruit consumption as it relates to seed dispersal and the propensity of many tropical species to evolve mutualistic interactions. This chapter includes a focus on coevolution, when two or more species become mutually interdependent.

Food webs and the adaptations associated with them form the core of Chapter 8. This chapter looks at various top-down and bottom-up interactions that contribute to structuring food webs and trophic dynamics in rain forests. Topics include the evolution of characteristics such as cryptic and warning coloration.

The high rate of photosynthesis evident in tropical rain forests results in high rates of primary productivity, and this is the subject of Chapter 9. How much carbon do lush tropical forests take in and how much do they emit (as carbon dioxide)? The carbon flux of tropical forests is an important area of research. As current climate change continues due to ongoing addition of atmospheric carbon dioxide, what is the potential of tropical forests to act as carbon sinks, absorbing and storing excess carbon? This chapter explores some of the sometimes contradictory research about how tropical forests might act to mitigate the effects of increasing atmospheric carbon dioxide.

Rich tropical forests often occur on poor soils. Chapter 10 examines nutrient cycling and tropical soils. It describes how vital nutrients such as phosphorus, nitrogen, and calcium cycle in tropical ecosystems and the essential function of decomposer organisms. This chapter includes discussion of various kinds of tropical soils and contrasts forests on floodplains, where annual flooding renews soil fertility, with those on terre firme, off floodplains.

Chapters 3 through 10 all focus on the ecology of the tropical rain forest. The remaining five chapters survey other tropical ecosystems and also examine topics having to do with conservation science in the tropics.

Areas of grassland with scattered trees are called savanna. Chapter 11 discusses tropical savannas and dry forest ecosystems. This discussion examines two views of how savannas and dry forests coexist in the tropics. One is that grassland, savanna, and dry forest represent a moisture gradient effect (where grassland is found in the most arid areas, and dry forest where there is more annual precipitation), and the other views them as alternative stable states dependent on local conditions. The importance of fire and grazing are also discussed.

Chapter 12 surveys other major tropical ecosystems, beginning with montane ecosystems and continuing to riverine forests and coastal mangrove forest This chapter also considers the connectivity between montane and lowland ecosystems from the standpoint of biodiversity maintenance.

Humans evolved in tropical Africa, and Chapter 13 discusses human ecology in the tropics. Topics include hunter-gatherer communities, simple agriculture in the tropics, and some of the traditional ways that humans have affected the overall ecology of the tropics.

The last two chapters deal with conservation issues in the tropics. Chapter 14 focuses on forest fragmentation and biodiversity. Chapter 15 discusses deforestation and forest degradation, as well as the effects of fire on tropical forests. It also examines emergent pathogens and invasive species and concludes with an overview of a current debate among ecologists about the likely future of tropical forests. While not meant to be comprehensive, these two chapters taken together cover the most fundamental issues and questions facing tropical conservation science.

1   What and Where Are the Tropics?

Beginnings

The world was once a larger place, or so it seemed. Vast areas lay unexplored, and following the Renaissance, European explorers began to fan out across the Earth in a nationalistic search for conquest, treasure, and lands to claim for their respective countries. With the onset of global colonialism and imperialism came an age of exploration. By the eighteenth and nineteenth centuries, many new regions were becoming known, and many of these were in the tropics.

Science, in the philosophical sense, traces its roots back to the emergence of science in ancient Greece, the Middle East, and Asia. But in a more contemporary sense, science really began during the Renaissance, when curiosity led to inductive and deductive reasoning, observational and experimental analysis, and most importantly, empirical investigation and materialistic explanations for natural phenomena. The study of the tropics was formalized only after the ages of global exploration and discovery and when the scientific method was used to investigate phenomena.

(a)

(b)

PLATE 1-1

(a) CHARLES DARWIN

(b) ALFRED RUSSEL WALLACE

The exploration of the world’s tropics by European nations brought with it much specimen collection and cataloging. Thus, the roots of tropical ecology lie most deeply in museum cabinets housing vast numbers of carefully labeled plants and animals brought back by many devoted and indefatigable naturalists including such figures as Charles Darwin and Alfred Russel Wallace (Plate 1-1). When Captain James Cook chose to have two naturalists, Joseph Banks and Douglas Solander included on the voyage of the Endeavor (1768), he set an immensely valuable precedent. It is that action that ultimately led to Darwin’s being invited on the Beagle voyage (1831-1836) and thus indirectly was responsible for the Origin of Species (1859).

One of the most noteworthy contributors to early knowledge of the tropics was Alexander von Humboldt (Plate 1-2), sometimes called the founder of the science of biogeography (Jackson 2009). Biogeography is the study of how organisms vary among regions, even when climate is similar For example, there are no apes in the New World tropics, but there are no monkeys with prehensile tails in the Old World tropics. This sort of example reflects differing evolutionary histories among regions. Biogeography will be a focus of the next chapter. Humboldt led successful and bold expeditions to difficult places ranging from Russia to northern South America. He published a massive five-volume tome titled Kosmos in which he attempted to summarize the entire scope of scientific knowledge of the world.

Humboldt’s travels in South America took him from rain forests along the Orinoco River to high-elevation Andean slopes as he traveled from Venezuela through the northern Andes. He visited central Mexico, Cuba, and the United States. Humboldt climbed some of the higher Andean peaks, reaching elevations of 5,800 meters (19,000 feet). It was Humboldt who first discovered and described the unique nocturnal oilbird (Steatornis caripensis) (see Chapter 7) in the caves around Caripe, Venezuela.

PLATE 1-2

ALEXANDER VON HUMBOLDT

Humboldt and his botanist colleague Aimé Bonpland described elevational changes evident onAndean slopes. They realized that as climate changed by elevation, so did plant and animal communities, and thus they were the first to elucidate what is termed the life zone concept. Much later, this concept was formalized by Leslie R. Holdridge (1947). A life zone is an ecological area defined by climatic variables such as mean annual temperature, total annual precipitation, and the ratio of mean annual evapotranspiration (a measure of heat) to mean total annual precipitation. (Transpiration is a term referring to the evaporative water loss from plants. Plants are adapted to pull water from soil via roots, and eventually that moisture is lost by evaporation from the surface parts of the plant.) In the Andes (as with all mountains), these variables change with elevation and result in differing ecosystems such as páramo (an ecosystem of wet grassland and shrubs) at high elevations and lush rain forest at low elevation (Plates 1-3 and 1-4). Holdridge’s life zone concept will be further discussed later in this chapter.

Naturalists were often moved to comment on the rich and diverse nature of flora and fauna within tropical rain forests. Their observations, by today’s standards, were largely (and understandably) anecdotal, though not lacking in important details. The modern student of tropical ecology will profit from reading some of the narratives of figures such as Thomas Belt, Henry Walter Bates, the aforementioned Darwin and Wallace, as well as others. For example, Henry Walter Bates discovered the form of animal mimicry (described in Chapter 8) that bears his name. These early explorers displayed much tenacity in capturing detail, transcribing vast amounts of information, and collecting and transporting specimens often under difficult circumstances.

HISTORICAL REFERENCES

These books, all still available (at least in libraries), were authored by some of the pioneers of tropical ecology and are strongly recommended as insightful, entertaining, and inspiring resources.

Bates, H. W. 1892. The Naturalist on the River Amazons. London: John Murray. Classic account of Amazonian natural history, and quite wonderful.

Beebe, W. 1918. Jungle Peace. London: Witherby. Beebe’s journey from the West Indies to the rain forest (jungle) of Guyana makes for engrossing reading, including a wonderful account of the odd Hoatzin bird (Chapter 12).

—. 1921. Edge of the Jungle. New York: Henry Holt. Like the previous volume,

contains short, delightful essays on tropical ecology. Classic.

Belt, T. [1874] 1985 reissue. The Naturalist in Nicaragua. Chicago: University of Chicago Press. One of the best of the classic exploratory accounts, focused entirely on Central America.

Chapman, F. M. 1938. Life in an Air Castle. New York: Appleton-Century. Highly readable with much information, particularly on tropical birds.

Darwin, C. R. [1906] 1959. The Voyage of the Beagle. Reprint. London: J. M. Dent and Sons. One of the best classic accounts of travel throughout South America. Many reprinted editions are available. Must reading.

Wallace, A. R. 1869. The Malay Archipelago. London: Macmillan. This is Wallace’s most famous book, an engrossing account of his eight years of travel throughout what is now Indonesia. The book abounds with information about wildlife as well as the various peoples Wallace encountered.

—. 1895. Natural Selection and Tropical Nature. London: Macmillan. This delightful book contains vivid descriptions, in glorious Victorian prose, of Wallace’s experiences in Amazonia.

Waterton, C. [1825] 1983. Wanderings in South America. Reprint. London: Century Publishing. A very entertaining narrative by a rather eccentric but perceptive explorer.

PLATE 1-3

SNOW ON PÁRAMO

Snow is common at high elevations in equatorial regions. Here, bunch grasses of the high páramo life zone are partially snow-covered. From Ecuador.

PLATE 1-4

FOREST CANOPY

The dense canopy of a low-elevation, humid tropical moist forest, such as this one in the Arima Valley in Trinidad, is typically irregular, with some emergent tree species rising above others. Forest gaps created by fallen trees also add to canopy heterogeneity.

Henry Walter Bates versus Curl-Crested Aracaris

The following text from Bates’s account in The Naturalist on the River Amazons illustrates the curl-crested aracari (Pteroglossus beauharnaesii), a species of toucan, and the memorable encounter that Bates had with a flock of these birds (Figure 1-1).

The Curl-crested Toucan (Pteroglossus Beauharnaisii)—Of the four smaller toucans, or arassaris, found near Ega, the Pteroglossus flavirostris is perhaps the most beautiful in its colours, its breast being adorned with broad belts of rich crimson and black; but the most curious species by far is the Curl-crested, or Beauharnais Toucan. The feathers on the head of this singular bird are transformed into horny plates, of a lustrous black colour, curled up at the ends, and resembling shavings of steel or ebony wood, the curly crest being arranged on the crown in the form of a wig. Mr. Wallace and I first met with this species on ascending the Amazons, at the mouth of the Solimoens; from that point it continues as a rather common bird on the terra firma, at least on the south side of the river, as far as the Fonte Boa, but I did not hear of its being found further to the west. It appears in large flocks in the forests near Ega in May and June, when it has completed its moult. I did not find these bands congregated at fruit trees, but always wandering through the forest, hopping from branch to branch amongst the lower trees, and partly concealed amongst the foliage. None of the arassaris to my knowledge make a yelping noise like that uttered by the larger toucans (Ramphastos); the notes of the curl-crested species are very singular, resembling the croaking of frogs. I had an amusing adventure one day with these birds. I had shot one from a rather high tree in a dark glen in the forest, and entered the thicket where the bird had fallen to secure my booty. It was only wounded, and on my attempting to seize it, set up a loud scream. In an instant, as if by magic, the shady nook seemed alive with these birds, although there was certainly none visible when I entered the jungle. They descended towards me, hopping from bough to bough, some of them swinging on the loops and cables of woody lianas, and all croaking and fluttering their wings. (Bates 1892, pp. 335-336)

(a)

(b)

FIGURE 1-1

(a) Curl-crested toucan. (b) Mobbed by curl-crested toucans.

Most eminent naturalists of the nineteenth century honed their skills through participation in voyages of discovery. For example, Thomas Henry Huxley and Joseph Hooker, both close friends of Darwin, each made an extensive voyage for the express purpose of expanding his horizons. These voyages were insightful, but they normally did not permit careful, systematic study within a given region.

It was not until the twentieth century that systematic study of regions within the tropics was really possible. What was needed were permanent field stations where scientists could go and conduct their work.

In 1923, following the completion of the mammoth engineering project known as the Panama Canal, a hilltop area of about 1,500 hectares (3,706 acres) on the Panamanian isthmus remained above water, a newly formed island surrounded by the newly created Gatun Lake. This was named Barro Colorado Island. Since 1946, it has been a field station administered by the Smithsonian Institution, and it has served numerous researchers, a function that continues to the present day. As you continue through this book, you will notice numerous citations regarding Barro Colorado Island, usually referred to simply as BCI.

William Beebe

William Beebe (1877-1962) (Plate 1-5) did, far more than change the direction of research in tropical ecology. He had a long and highly distinguished career as an explorer, scientist, and writer. He spent most of his career associated with the New York Zoological Society. His productivity was amazing, having authored 24 books and about 800 scientific articles over the course of his career. Beebe began as an ornithologist, and one of his first major books bore the title The Bird, Its Form and Function (1906). Arguably his strongest contribution to ornithology (he had many) was his classic book A Monograph of the Pheasants, published in four volumes from 1918 to 1922.

PLATE 1-5

WILLIAM BEEBE

Beebe’s tropical research was prolific and began in British Guiana (now Guyana) in 1916. He traveled widely, including to the Galapagos Islands, a trip that resulted in another memorable book, Galapagos: World’s End (1924). He also authored two books about tropical ecology, one of which, Jungle Peace (1918), greatly impressed Theodore Roosevelt, himself an explorer of the Neotropics.

In 1928, Beebe established a field station in Bermuda, an event that led to another major chapter in his career, that of oceanic explorer. Beebe was frustrated that animals hauled up from the depths of the seas were not only dead on arrival but also mutilated by the pressure change experienced while being hauled to the surface. After consultation with Theodore Roosevelt, Beebe eventually focused on a submersible that he called a bathysphere, a thick metal chamber that could withstand the great pressures of the ocean depths and that could be lowered and raised from a ship. He subsequently made about 30 descents, accompanied in the tiny chamber by the bathysphere’ s inventor, Otis Barton. In 1934, Barton and Beebe reached a depth of 922.93 meters (3,028 feet), an amazing accomplishment that was not equaled for many years.

Beebe was a popularizer of science but, at the same time, a very productive scientific researcher. It has been said of him that he really founded the science of tropical ecology.

Another unique field station devoted to tropical biology is Simla, in the Arima Valley on the island of Trinidad. While Simla is not one of the most prominent sites today, it is noteworthy for its founder, the naturalist-explorer William Beebe, and for its initial purpose. Though Trinidad appears to be part of the West Indies, it is biogeographically distinct, as it is actually on the continental shelf, part of Venezuela, just as the island of Martha’s Vineyard is really part of Massachusetts. Thus, the flora and fauna are less West Indian than they are South American. Recognizing this, Beebe established Simla in 1950. Beebe purchased the land at an advanced age (he was 73) and began the research station. His strong belief was that researchers must work on live animals and not merely collect specimens. At that time, specimen collection for museums essentially dominated tropical research; thus, Beebe’s approach helped change the direction of ecological work from specimen collection to detailed ecological studies of organisms in the field.

Many tropical field stations exist today and more are being established. In the Western Hemisphere tropics, in addition to Barro Colorado Island, some of the most prolific research sites are La Selva (Costa Rica), Cocha Cashu (Peru), Manu Biological Preserve (Peru), Minimal Critical Size of Ecosystems (Brazil), Los Tuxtlas (Mexico), and Luquillo (Puerto Rico). The Tropical Ecology Assessment and Monitoring Network (TEAM) includes 122 sites throughout the tropical world, from Colombia to China (see http://www.teamnetwork.org/en/about). The Organization for Tropical Studies (usually referred to as OTS) maintains not only the La Selva Biological Station in Costa Rica (Plate 1-6) but also biological stations at Las Cruces and Palo Verde. Many other research stations also are found throughout the tropics, some of which will be noted for studies described within this book.

In many areas within the tropics, huge challenges ranging from marginal living conditions, lack of facilities, and political instability still face the field worker. Nonetheless, it is fair to say that many if not most field sites now have sufficient infrastructure and facilities to permit cutting edge research to be conducted. What was once the age of exploration and description has become the age of experimental design, data collection, and hypothesis testing.

PLATE 1-6

The OTS field station at La Selva.

Modern Tropical Ecology

Many college-level tropical ecology courses now include field opportunities at one or more research stations somewhere within the world’ s tropical regions. The very fact that undergraduate and graduate students now routinely visit and work at tropical research stations has vastly increased tropical ecological research.

PLATE 1-7

The harpy eagle (Harpia Harpyja) released on Barro Colorado Island, Panama, is monitored by radio tracking.

Professional researchers work many months at a time at tropical research stations with well-equipped laboratories, computers with Internet access, global positioning systems, and so on (Plate 1-7). Expeditions into remote areas where services are few to none still occur, however, as much of the tropics still demands field work away from major research stations. For a wonderful and insightful look into how tropical research is conducted and the dedication and drive of the researchers, read The Tapir’s Morning Bath by Elizabeth Royte (Boston: Houghton Mifflin, 2001). It chronicles the life and dedication of researchers at Barro Colorado Island.

In the Western Hemisphere, most tropical researchers focus on the Neotropics, their work taking them anywhere from Mexico to southern Amazonia as well as to the Caribbean islands. In the Eastern Hemisphere, much research is carried on inAfrica and Australasia. The intensity of tropical research varies considerably from one country to another, and thus there are some areas that are far less studied than others (Stocks et al. 2008).

Professional papers detailing tropical research are routinely published in leading science journals such as Science and Nature. In addition, journals such as American Naturalist, Ecology, Oecologia, Conservation Biology, and many others routinely include papers about research from the tropics.

The first professional society devoted to tropical research was the Bombay Natural History Society, founded in 1883. The International Society for Tropical Biology was founded in 1956 (Chazdon 2002). The Association for Tropical Biology and Conservation, founded in 1963, publishes the journal Biotropica. The papers published in Biotropica include studies about all tropical regions, but the majority of the work reported is about the Neotropics. Another important journal is the Journal of Tropical Ecology, first published in 1985. This journal, like Biotropica, publishes research papers about all tropical regions, but most papers tend to be about Africa and Asia. Taken together, a recent survey showed that the highest percentage of papers in those two journals were from studies conducted in Brazil, Costa Rica, Mexico, Panama, Malaysia, Puerto Rico, Australia, French Guiana, Venezuela, Ecuador, United States, Peru, Indonesia, Colombia, and India, in that order (Stocks et al. 2008) (Figure 1-2). The dearth of studies from African nations is obvious, as is the dominance of studies from the Neotropics.

The Tropical Rain Forest—A Pioneering Book

The study of tropical ecology became formalized in 1952 when P. W. Richards, a botany professor from Great Britain, published The Tropical Rain Forest: An Ecological Study (Cambridge, UK: Cambridge University Press). As the title suggests, this book focused on rain forests but also included discussions of coastal mangrove forests and savannas. Reflecting the field of ecology as it was at that time, the book is largely descriptive, but it does an admirable job of comparing global equatorial forests and identifying common patterns—particularly of physiognomy, climate, and soils—that they share to varying degrees (Figures 1-4 and 1-5). What is obviously missing from the discussion is treatment of the complex roles of animals in rain forest ecology. Pollination and seed dispersal—both of which, in the tropics, are usually dependent on animals—are not listed in the index, for example, and are barely touched upon. Richards’s book was nonetheless the first serious attempt at summarizing knowledge about rain forest ecology as well as synthesizing concepts. It paved the way for much research, and it is still valuable and insightful. This classic book was revised and a second edition was published in 1996 by Cambridge University Press.

FIGURE 1-2

This graphic depicts the number of studies published in two professional journals, Biotropica and Journal of Tropical Biology, from 1995 to 2004. It shows how many studies were conducted in each of various countries. Note that most studies are focused in the Neotropics. (Numbers on bars are actual numbers of published studies.)

The View from the Canopy

FIGURE 1-3

This sketch illustrates the use of a large industrial crane to examine tropical forest canopies: a is the tower, b is the counterjib, c is the counterweight, d is the operator’s cab, and e is the gondola, where a researcher would be located.

Tropical research has been facilitated in recent years by towers, walkways, and cranes that allow direct access to the forest canopy. It has been frustrating to tropical ecologists to realize that much of the biological diversity found in tropical forests is confined to the forest canopy, the dense and irregular layer of foliage representing numerous tree species, usually located 30 to 40 meters (about 98 to 130 feet) from the ground. Scores of arthropod and other invertebrate species as well as birds, mammals, reptiles, and amphibians spend most or all of their lives well above ground level. The same is true for epiphytes, the myriad species of bromeliads, orchids, cacti, and other plant species that live on the bark of tree trunks and branches. (Epiphytes live on bark and branches of other plants but are not directly parasitic.) But ecologists now do have more access to the forest canopy. The construction of high towers, some associated with tall, emergent trees, allows one to be at canopy level and make observations. Even more access is provided by canopy walkways that connect various trees or that are supported by metal towers. In some areas, tall cranes, such as are used in building construction, allow mobile access to the canopy (Parker et al. 1992) (Figure 1-3; Plates 1-8 to 1-11).

PLATE 1-8

CANOPY WALKWAY AND FOREST

This tall and lengthy canopy walkway at Sacha Lodge in Ecuador provides safe and easy access that permits study of species largely confined to the forest canopy.

Canopy research has helped not only to document patterns of biodiversity but also to demonstrate how tropical tree species vary in their response to physiological variables, such as elevated carbon dioxide levels and the production of biogenic aerosol compounds (Ozanne et al. 2003).

PLATE 1-9

HIGH CANOPY WALKWAY

Viewed from the ground, the canopy walkway is seen crossing a major forest gap.

PLATE 1-10

VIEW FROM CANOPY WALKWAY

The expanse and structural complexity of Ecuador’s lowland forest is readily evident, as observed from a canopy walkway.

PLATE 1-11

WOODEN CANOPY TOWER

Canopy towers allow more restricted access than walkways but, if strategically placed, afford excellent opportunities for canopy study. Note the broad branching pattern of the tree. The tower is essentially built around the tree. From Sacha Lodge, Ecuador.

Geographic Definition of the Tropics

In today’s world, the tropics are the region located between the Tropic of Cancer (23°27’N) and the Tropic of Capricorn (23°27’S), a latitudinal band of approximately 47°. This belt of latitude encircles the Earth at its widest circumference. At either extreme, ecosystems become subtropical. Tropical regions represent about half of Earth’s surface and thus has an immense effect on Earth’s climate (Huber 2009). For much of Earth’s history, even before the initial evolution of flowering plants (angiosperms) in the mid-to late Cretaceous period (144 to 65.5 million years ago), Earth has been even more tropical than it is today (see Appendix). Fossil palm trees and crocodiles have been unearthed in what is now Alaska and Siberia (Huber 2009). This is because the climate of Earth, for a variety of reasons discussed in the next chapter, is not constant. In times when climate was warmer, the distribution of the tropics was globally broader than is the case today.

FIGURE 1-4

This is a sketch of a forest profile in what was then British Guiana and is now the independent nation of Guyana. This is adapted from one of several classic forest profiles included in Richards’s famous book The Tropical Rain Forest.

FIGURE 1-5

Also adapted from Richards’s classic volume, this is a profile of a mixed Dipterocarp forest in Borneo. Compare this with Figure 1-4. Both forests are structurally similar but contain entirely different tree species.

Tropical Worlds Long Ago: Rain Forest in Denver and a Very Large Snake from South America

Denver, Colorado, is located along the front range of the Rocky Mountains, an area well within the temperate zone. Forests of spruce, fir, pine, and aspen characterize the region. But had Denver existed 64.1 million years ago (a time called the Paleogene), not long after the mass extinction that ended the Cretaceous period, a very different forest would be evident—a tropical rain forest (see Appendix). At a site called Castle Rock, near Denver, fossil plants (particularly leaves) have been unearthed, plants that are clearly tropical (Johnson and Ellis 2002) (Figure 1-6). The diversity of tropical plants at this fossil site shows that tropical conditions then prevailed at a region far from today’s tropics. Plant diversity had recovered quickly (within 1.4 million years) after the mass extinction that ended the Mesozoic era.

FIGURE 1-6

This drawing illustrates the morphotypes of tropical leaves found near Castle Rock, Denver, dating back 64.1 million years. Only the leaves in the upper right are nondicot leaves. By this time, modern plants, dominated by dicots, dominated terrestrial plant communities.

PLATE 1-12

ANACONDA

The largest snakes found in the tropics are the pythons (Pythonidae) of Africa and Australasia and the boas and anaconda (Boidae) of the Neotropics. The yellow anaconda (Eunectes murinus) of the Neotropics is arguably the world’s largest extant snake, reaching a length of 9 meters (about 29.5 feet) (Plate 1-12). The great lengths reached by pythons and boas are facilitated by the warm tropical climate in which they live (Head et al. 2009). Snakes are poikilothermic, ectothermic vertebrates, meaning that their body temperature and thus their metabolism is dependent on ambient air temperature. (Poikilothermic and ectothermic are different in meaning. Poikilothermic means specifically that body temperature changes as ambient temperature changes, whereas ectothermic means that the animal does not have physiological ability to regulate its body temperature to stay warm.) Mathematical models have been developed that correlate the difference in maximum body size of snake species occurring in different regions with mean annual temperature of each region. The warmer the mean annual temperature is, the larger the snakes are able to grow. Thus, it was of great interest when a fossil boid snake dated at 58 to 60 million years old was unearthed from a site in northeastern Colombia and the fossils (vertebrae) indicated that the snake reached an estimated length of 13 meters (about 42.6 feet). This serpent, named Titanoboa, was estimated to have weighed about 1.27 tons. That’s a lot of snake. When included within the model that correlates today’s boid snakes and air temperature, the results suggest that Titanoboa required a minimum annual mean temperature of between 30°C to 34°C (86°F to 93.2°F) to survive. This would make the climate at that time about 6°C to 8°C warmer than it is currently and suggests that a much higher level of carbon dioxide (a gas that contributes strongly to retention of atmospheric heat) was present in the atmosphere when Titanoboa lived.

These two examples show that the distribution and intensity of tropical climate have changed throughout Earth’s history, as it appears to be doing today.

Well before the evolution of flowering plants, the world was largely tropical. In the Jurassic period (206 to 144 million years ago), the giant long-necked sauropod dinosaurs lived in a largely tropical world, where there was no polar ice and where the average global temperature as well as oxygen and carbon dioxide levels exceeded those found today (Plate 1-13).

PLATE 1-13

CAMPTOSAURUS, APATOSAURUS, STEGOSAURUS, DRYOSAURUS, CAMARASAURUS AND ALLOSAURUS (LEFT TO RIGHT)

Each of these dinosaurs cohabited tropical regions in western North America during the late Jurassic period. The tropics then would bear scant resemblance to the tropics of today.

During the Cenozoic era (65 million years ago until the present), a time of proliferation of insects, flowering plants, and mammalian diversity, global climate cooled, beginning strongly during the Miocene epoch (23.8 to 5.3 million years ago) and accelerating during the Pleistocene (1.8 million to 8,000 years ago), commonly referred to as ice ages (see Appendix). During times of cooling, tropical forests presumably shrank in area, replaced by other more seasonal tropical ecosystems such as grassland, savanna (an area where grasses predominate but with varying amounts of scattered trees), dry forest (forests that experience significant dry season), or deciduous forest (forests where most trees synchronously shed leaves for part of the year). Earth is currently considered to be within an interglacial period, and there is strong evidence for climate change due to human activities, a topic to be considered later in this book (Figures 1-7 and 1-8).

FIGURE 1-7

Satellite image of Earth, showing the band of tropics around the equator.

FIGURE 1-8

Political map of Earth, showing nations that occur within the tropics.

Because of the geological process of plate tectonics (described in the following chapter), the Earth’s continents, united 248 million years ago at the boundary of the Paleozoic and Mesozoic eras, have gradually fragmented and separated, with large areas of ocean isolating them to varying degrees. This separation into massive continents and varying-sized islands has resulted in evolutionary consequences, particularly the evolution of endemic species. An endemic species is a species whose range is confined within a limited geographic area. For example, the imposing Komodo dragon lizard (Varanus konodoensis) is endemic to Indonesia, specifically to the Lesser Sunda Islands, most particularly to Komodo (Plate 1-14). Biogeographers recognize various biogeographic realms throughout the planet. Plate tectonics and biogeography will be discussed in greater detail in the next chapter.

Middle America and South America, along with the various islands of the Caribbean and tropical Atlantic Ocean, constitute the Neotropic realm, the so-called New World tropics. The Paleotropic realm, or Old World tropics, comprises Africa and Madagascar (Malagasy Republic), southern India, and Australasia. The latter is further divided into the Southeast Asian tropics (extreme southern China, Myanmar [Burma], Vietnam, Laos, Thailand, Malaysia, Sumatra, Borneo, Philippines) and the Australian tropics (including Sulawesi and other islands of eastern Indonesia as well as New Guinea). The point of separation between the biogeographic realms of tropical Asia and Australia is called Wallace’s Line, after Alfred Russel Wallace, who first described it (see Chapter 2). Biogeographers differ on exactly where to place Wallace’s Line because plant species are not as sharply separated by it as animal species (Whitmore 2002). The southern half of India is within the tropics, and it is unique because India separated from Gondwanaland (a massive southern continent once consisting of South America, Africa, Madagascar, India, Antarctica, Australia, and New Zealand) in the early Cretaceous period (about 140 million years ago) and drifted as an island until it united with Asia about 40 million years ago, in the early Cenozoic. Likewise, Madagascar has been isolated from the rest of Africa to the point where it has numerous endemic species such as the primate group known as lemurs (Lemuridae).

PLATE 1-14

The Komodo dragon, which sometimes reaches a length of nearly 3 meters (9.8 feet), is endemic to the Lesser Sunda Islands of Indonesia.

Each of these realms contains unique assemblages of plant and animal species (Primack and Corlett 2005)

Tropical Climate

In two words, the tropics (especially where there is lush forest) are warm and wet (Figure 1-9). Tropical regions experience high average temperature, and most (dry grassland and deserts being exceptions) experience substantial precipitation in the form of rainfall. Tropical ecosystems experience seasonality in the form of varying and often predictable amounts of rainfall throughout the year. In contrast to the seasonal pattern that typifies the temperate zone, the tropics vary primarily in precipitation amounts, not temperature fluctuation. There are no periods of protracted cold, no frost, no snow in tropical ecosystems except at high elevations. But there are distinct and pronounced dry and wet seasons. If the dry season is moderate, forests will remain evergreen throughout the year. If the dry season is protracted and severe, forests will have less biomass and more deciduous species, or the ecosystem will not be forest but will be savanna, shrubland, or grassland. The ecosystem type of a given region is also strongly influenced by soil characteristics, a point to be considered later in the text.

Leslie Holdridge and the Life Zone Concept

One of the most significant papers in the science of climatology and ecology was published in Science by Leslie R. Holdridge in 1947. The title of the paper, Determination of world plant formations from simple climatic data, addressed a profound truth: It is possible by knowing three broad climatic variables to predict what major kind of terrestrial ecosystem will prevail in any given geographical region. Those variables are (1) mean annual biotemperature (defined as the average air temperature after values below 0°C or above 30°C are removed); (2) total annual precipitation; and (3) ratio of mean potential evapotranspiration (a function of moisture and temperature) to mean annual precipitation. Holdridge illustrated the concept with a triangle of hexagons, each of which is positioned according to where it fits among the variables, each of which represents one side of the triangle (Figure 1-10).

FIGURE 1-9

These graphs contrast the climate in tropical Brazil with that of temperate Canada. Note the strong seasonality of the tropics with regard to precipitation but the steady temperature that prevails in the tropics. The opposite pattern characterizes the temperate zone.

FIGURE 1-10

This is the famous Holdridge diagram that illustrates the relationship between ecosystem types as determined by latitude, elevation, and combination of precipitation and temperature.

FIGURE 1-11

Diagram showing the relationship between ecosystem type, mean annual temperature (°C), and mean annual precipitation (centimeters). Note that tropical rain forest occurs where both climatic variables are highest.

Holdridge’s 38 life zones were broadly divided into polar, subpolar, boreal, cool temperate, warm temperate, subtropical, and tropical. Eight life zones are represented in subtropical and eight life zones in tropical latitudinal regions. The tropical life zones are tropical desert, tropical desert scrub, tropical thorn woodland, tropical very dry forest, tropical dry forest, tropical moist forest, tropical wet forest, and tropical rain forest (Figure 1-11).

Consider the difference between tropical rain for est and tropical dry forest, both of which may occur at the same latitude. Dry forest occurs where there is a significant dry season for part of the year. Tropical dry forest (Plates 1-15 and 1-16), under Holdridge’s classification, is defined as an ecosystem that experiences a mean annual biotemperature of greater than 24°C (75.2°F), has a potential evaporation ratio of 1:2 (defined as the ratio of annual potential evapotranspiration [PET] to mean total annual precipitation—this means that dry ecosystems in tropical regions have higher ratios and humid ecosystems lower ratios), is subhumid, and has an average total annual precipitation of between 100 and 200 centimeters (39.4 to 78.7 inches). In contrast, rain forest has a mean annual biotemperature also greater than 24°C but has a potential evaporation ratio of only 0.125 to 0.25, is superhumid, and experiences greater than 800 centimeters (315 inches) of average total annual precipitation. Notice that Holdridge’s classification makes true rain forest an extremely wet ecosystem. In fact, most rich lowland tropical forests routinely called rain forests tend to fall into what, in Hold-ridge’s classification, would be termed tropical moist or tropical wet forests (Figure 1-11; Plate 1-17).

PLATE 1-15

Where rainfall is highly seasonal, tropical ecosystems are often dry forest, with many species of deciduous trees. Forest stature is generally small, and trees are usually widely spaced. From Venezuela.

PLATE 1-16

Northern Australia is an area of vast dry forest with tall, scattered termite mounds. This is the ecosystem typically called outback.

Some tropical ecosystems may be much drier than dry forest, with less annual precipitation, most of it highly seasonal. Tropical thorn woodland (under Holdridge’s classification) includes ecosystems that are dominated by grasses with varying densities of trees scattered throughout. These kinds of ecosystems are called savannas and are found in most tropical regions. The best known savannas occur in Africa, where the last of the world’s megafauna (diversity of large animals) is still found (Plates 1-18 and 1-19). Savannas and dry forests (Plate 1-20) will be discussed in detail in Chapter 11.

It is important to realize that Holdridge’s life zones are not merely latitudinal but also altitudinal. A walk from low Andean rain forest up to an elevation of about 3,500 meters (about 11,483 feet) will take you through several life zones, from hot, humid, dense forest to cold, windswept paramo, a vastly different ecosystem. Likewise, life zones would markedly change if you were in Tanzania and scaled Mount Kilimanjaro (whose summit is 5,895 meters, or 19,341 feet).

PLATE 1-17

Rapidly growing tree species such as Cecropia trees, shown here in the foreground, typify the forest edge, where vegetation structure is dense. This would be considered tropical moist forest in the Holdridge classification. From Trinidad.

Differences in elevation result in differences in average temperature and precipitation, and that, in turn, results in different ecosystems. Within the South American country of Venezuela, for example, several major elevational ecosystem types are apparent. At highest elevations (what would be described as alpine), a wet and windswept shrubland called páramo (Figure 1-12) is found. Lower in elevation is forest of low-stature gnarled trees, and lower still, is cloud forest, a forest shrouded in cool mist much of the time. Below cloud forest may be semievergreen forest, deciduous forest, or thorn forest, depending on soil and water availability and local seasonal patterns (Figure 1-12). Tropical rain forest occurs in lowland areas where there is ample precipitation