The New Neotropical Companion

The New Neotropical Companion

The New Neotropical Companion

John Kricher

PRINCETON UNIVERSITY PRESS

PRINCETON AND OXFORD

Copyright © 2017 by Princeton University Press

Published by Princeton University Press, 41 William Street, Princeton, New Jersey 08540

In the United Kingdom: Princeton University Press, 6 Oxford Street, Woodstock, Oxfordshire OX20 1TR

press.princeton.edu

All Rights Reserved

ISBN (pbk.) 978-0-691-11525-2

Library of Congress Cataloging-in-Publication Data

Names: Kricher, John C., author.

Title: The new neotropical companion / John Kricher.

Other titles: Neotropical companion

Description: Princeton : Princeton University Press, 2017. | Revised edition of: A neotropical companion. 2nd ed., rev. and expanded. c1997. | Includes bibliographical references and index.

Identifiers: LCCN 2016027794 | ISBN 9780691115252 (pbk.)

Subjects: LCSH: Ecology—Latin America. | Natural history—Latin America. | Natural history—Latin America—Pictorial works.

Classification: LCC QH106.5 .K75 2017 | DDC 577—dc23 LC record available at https://lccn.loc.gov/2016027794

British Library Cataloging-in-Publication Data is available

This book has been composed in Minion Pro and Myriad Pro

Printed on acid-free paper. ∞

Printed in China

10 9 8 7 6 5 4 3 2 1

To my parents, who introduced me to the world we live in and to the wonders of life on Earth.

Contents

Preface

Why a New Neotropical Companion?

It was first published in 1989 and it soon became known to travelers, both literal and armchair, as The Little Green Book (plate A). The it was the first edition of a relatively humble book that bore the title A Neotropical Companion. The subtitle was more explanatory: An Introduction to the Animals, Plants, and Ecosystems of the New World Tropics. It was a small book that fit easily into the pocket of a field jacket or a backpack. Illustrations were limited in number, comprising only a few black-and-white line drawings. The text of the book attempted to capture the ecological allure of the tropics, and in particular what ecologists were learning about the complexity and splendor of the uniquely diverse rain forest biome. At that time most of my field experience was confined to Middle America, Belize in particular, plus Venezuela, Peru, and Ecuador, to which I had taken a few research trips, and thus the book omitted much information about the central Amazon Basin. Nonetheless the book resonated with travelers to the American tropics as well as students taking college-level courses in tropical ecology (many of which included some brief field time at a tropical location). I think the real reason the book found success was the remarkable subject matter it discussed. As I wrote the book I was continually amazed that so much hard-earned scientific insight (real boots on the ground ecology) about the tropics was still essentially confined within scientific circles rather than readily available to interested lay readers and introductory students. I don’t mean how to identify this plant or that bird. I mean how the myriad species of tropical organisms interact to form a complex web of interdependency. I knew about this uniquely complex ecology (I described it as a Gordian knot) from firsthand experience as well as from my extensive reading of the scientific literature. How could this stuff be any more interesting? It was so cool! People ought to know about such things.

Plate A. The Little Green Book. Photo by John Kricher.

I wrote at a time when field research in tropical ecology had really begun to burgeon. More and more researchers were in the tropics making that region their lifetime focus, and new field stations were appearing throughout the tropics, in particular in the New World, or Neotropics. The research was ongoing, the data were pouring in, and data analyses were revealing all manner of fascinating ecological and evolutionary insights. I just had to tell some of that story, and so I did.

In the mid-1990s my editor at Princeton University Press asked me to consider expanding the Little Green Book, adding more breadth and detail. In addition to black-and-white line drawings there would be several groupings of photographs, all reproduced in color. By then I had the benefit of many more trips to the tropics, including several to the Brazilian Amazon. So the Little Green Book morphed into a bigger book, both in dimensions and page count, expanding to 451 pages. It contained far more coverage of the region and lots more basic information and examples. The second edition of A Neotropical Companion was published in 1997 and has remained useful to students, travelers, and general readers.

Why a New Edition Now, and Why Call It New?

In the 1997 edition coverage expanded and, as in the 1989 edition, I really did strive to be a real companion to my readers and write in layperson’s terms without becoming overly reliant on scientific jargon. I tried to tell in a basic and uncluttered way just how the ecology of the tropics works and why it is so remarkable. But I also added a considerable amount of academic material, and in accordance with that, citations from the scientific literature, some 800 of them, were interspersed throughout the book. Nonetheless, some academics thought it to be too general. A few professors told me it was just not rigorous enough for use in their courses (forgetting, I suppose, that it was never meant as a textbook). Some academically focused folks also commented that they thought I was too colloquial, inserting myself too often into the book. Other, more general readers had the opposite reaction and thought the writing to be a bit too much like that found in a textbook, at least in a few places, and said it was a bit arcane for a field guide, even regarding a complex subject like tropical ecology. Fortunately, many (hopefully most) readers accepted my attempt to balance academics and basic natural history, and thus the book has enjoyed a strong continued readership to this day. It has been translated into Spanish, through the efforts of the Birders Exchange Program of the American Birding Association; several thousand copies of the Spanish edition have been handed out free of charge to Latin American scientists, conservationists, and students, and it is available as a free download from the ABA website. It pleases me to know the book has been useful to so many.

My next project was major in scope: an outgrowth of A Neotropical Companion but far more extensive. Scientific knowledge and new insights about the tropics continued to burgeon and there was a clear need for a rigorous and comprehensive book that would provide a solid academically based introduction to not only the Neotropics but also the global tropics in general. Thus I elected to write such a book and did so. That effort resulted in Tropical Ecology, a comprehensive upper-level college textbook, published by Princeton University Press in 2011. This was meant, wholeheartedly, to be a rigorous and thorough text. The book seems to have found its intended audience among the offerings of colleges that include Tropical Ecology courses within their curricula.

Since the publication of Tropical Ecology I have come full circle, back to wondering what to do about what was once the Little Green Book. The information in A Neotropical Companion needs to be updated. Science is always a work in progress, and that is certainly true of tropical ecology. There remains, in my view, a need for a general book for nonacademically focused readers and travelers that with a broad brush describes New World tropical ecology and that serves to interpret and explain the most biologically diverse terrestrial ecosystems on Earth.

And thus it is that I chose to revise A Neotropical Companion and call it The New Neotropical Companion. It is new. It has been written to be much less academic in tone. I have adapted some of the former edition to this new edition and, as well, borrowed liberally from my 2011 text, Tropical Ecology, converting the academic writing to a more user-friendly and generalized treatment. There is an abundance of color illustrations that adds immeasurably to the utility of the book. You not only read about the Neotropics in these pages, you see the Neotropics. The writing is up to date, with full discussions of some of the newest and coolest scientific insights. Some of what was in the previous edition is essentially unchanged, some is somewhat changed, much is very changed. And, of course, there are numerous insights in the NNC not in the previous edition because they were not known then. That is the nature of science.

And that is why there is a New Neotropical Companion.

Acknowledgments

The list grows. In the previous two editions of A Neotropical Companion I have acknowledged many folks who have in various ways contributed to my continuously growing knowledge base about the Neotropics. These are still the folks who in so many ways made this book possible. The list includes former students, many friends with whom I have traveled, and the talented guides from whom I have learned. I have received travel support from Wheaton College (Norton, MA), where I teach, as well as from the American Birding Association, for whom I have led tours and run workshops at various Neotropical venues. I express my gratitude to all of these people for their assistance and companionship over the years. In particular and thinking back to how it all began, I thank, yet again, Fred Dodd of International Zoological Expeditions for introducing me to Belize in 1978 and for providing me with a delightful addiction that is satisfied only by being in the realm of palm trees and toucans.

The New Neotropical Companion stands out from its predecessor volumes in one obvious way. The book is stunningly illustrated with some of the finest nature photography there is. I say this unabashedly, because many of the images to which I refer were not taken by me. I approached various friends asking if they would permit me to use their work in my book. No one turned me down. Photos began arriving, lots of them. I only wish I could have included more. It was almost physically painful to leave some of them out. I know you will want to just thumb through the book and revel in the photos—and never mind about the text. But I do hope you get around to reading the book after admiring the multiple images that make it special. If one picture really is worth a thousand words, this volume is indeed a major expansion. It has come of age.

Therefore, thank you so much to James Adams, Steve Bird, Beatrix Boscardin, Edison Buenaño, Diana Churchill, David Clapp, Peter Crosson, Fred Dodd, Carl Goodrich, Bruce Hallett, Ed Harper, Jill Lapato, Bruce and Carolyn Miller, Gina Nichol, Nancy Norman, Dennis Paulson, Scott Shumway, Clay Taylor, Andy Whittaker, Alex Wild, Sean Williams, and Kevin Zimmer. Your long lenses and photographic talents have brought the art of tropical nature to these pages.

Since the publication of the second edition of A Neotropical Companion, and while collecting information for this volume, I have been fortunate to travel with some exceptional guides including Domiciano Alveo, Carlos Bethancourt, Edison Buenaño, Damien and Camilio Montanez, Olger Licuy, Marcelo Padua, Benjamin Schwartz, and Jose Rafael Soto. My good friend Raul Arias de Para has made my continuing tropical education a particular joy as I have made multiple and memorable visits to his Canopy Tower, Canopy Lodge, and Canopy Camp. Indeed, in my experience, the entire Canopy Family is as good as it gets. Muchas gracias, mis amigos.

I thank my friends Tony White and Elwood Bracy for introducing me to the Bahamas and its remarkable habitats and wildlife, and Tony again for arranging a day in the field with Paul Dean, and Woody for showing me Abaco.

Dick Payne, Mike Ord, and Terry Moore were terrific traveling companions, inviting my wife Martha and me to join them for what was a wondrous tour of Ecuador organized by Jane Lyons of Mindo Bird Tours (lots of hummingbirds).

In appreciation for help with Lepidoptera ID, I thank Marj Rines and Sheri Williamson.

Over the years I have received e-mails and letters from individuals who have made suggestions for what might be included in a new edition and who have pointed out various errors or omissions, and even a few grammatical hiccups. I thank each of you who took the time and interest in this book to offer that help to me.

Since the publication of the first edition I have enjoyed a highly cordial relationship with Princeton University Press. Again I thank my editor, Robert Kirk, for his continued patience, guidance, and support. I am proud to be a Princeton author. And thanks also to Robert for putting together such a fine team to make this book happen. I am grateful to Karen L. Carter for stewarding the project through the complex conversion of manuscript to book; to Ryan Mulligan for handling the details pertaining to the multiple photographs; and to the ever-so-amazing Amy K. Hughes, whose copyediting efforts and unflagging attention to detail have made this book so much better. Only I know that. You, the reader, are the beneficiary. So, thanks big time, Amy. It was a pleasure.

Just a quick acknowledgment to my grandchildren, Liam and Rory O’Toole, for the fun we have together. They are really going to enjoy the pictures in this book. Though Liam is only six years old and Rory only four, both know what a toucan is.

My wife, Martha Vaughan, has supported and encouraged me in numerous ways as we journey through life. During the course of various book projects she has been a meticulous copy editor and the best of field companions, and she does remarkably well in patiently putting up with ongoing authorial angst. Ah, but when next she asks me whether I have finally completed that book, I will happily answer, Well, as a matter of fact, yes, I have. And that will make her happy, and that is what I most like to do.

How to Use This Book

Readers already familiar with previous editions of A Neotropical Companion will immediately understand the tone and objectives of this book. This edition is not written in an academic style, even more deliberately so than its predecessor. Most important, it is not about how to identify various plants and animals of the tropics. When I visited Antarctica a few years ago I brought with me a book that illustrated literally every species of mammal and bird I was likely to find in Antarctica (mostly, of course, in the waters around the continental landmass). I had but to match the animal with its picture and bingo, a name! I saw most of the species listed. I did not have to worry about reptiles and amphibians, as there are none in Antarctica. The insects are few to none as well. But that is because Antarctica is very cold and windy, a basically inhospitable place to terrestrial life in general. Not so with the tropics.

Consider birds, for example, a group with which I have considerable familiarity. On my Antarctica trip I saw almost all of the bird species present, and the list was fewer than 50 species. If I told you there are nearly 350 species to be found in the Neotropics, and you compared that with the number of Antarctic bird species, you would quickly see why it would take a much bigger book to cover just the bird species of the Neotropics, to say nothing of mammals and other animal groups. But in reality, there are nearly 350 species of hummingbirds (family Trochilidae) alone in the Neotropics. Several other bird families (Furnariidae, the ovenbirds and woodcreepers), Tyrannidae (the tyrant flycatchers), and Thraupidae (tanagers and related species) have similarly high species richness. Add to those all the other bird species, and you have about one-third of the world’s approximately 10,000 bird species, present and accounted for in Central and South America, most of them in either the Andes or the Amazon regions. So it becomes immediately obvious that it would take many volumes to catalog each and every bird, mammal, reptile, and amphibian species of the region. Add insects and other invertebrates, whose diversities are in the high thousands (likely well over a million), and it soon becomes hopeless to generate any sort of comprehensive field guide to the identification of Neotropical animals. Just to thoroughly catalog the butterflies of the relatively small country of Costa Rica, for example, requires two thick volumes. The distribution and identification of birds found in Ecuador, another small country, also requires two volumes.

And consider plant species. If you visit the lush cove forests of the central Appalachians, such as those found in Great Smoky Mountains National Park, you can, with skill and patience, find about 25 to 35 tree species in a single tract of 2 hectares, or about 5 acres of forest. (Note: hereafter in this book, we’ll use the abbreviations ha for hectares and ac for acres.) Take that numerical range, multiply it by 10, and you get the picture for parts of Amazonia. That’s right, more than 300 tree species may be present in just 2 ha. And many of them look very similar, requiring expert training in tree identification techniques. It is thus obvious that no book purporting to discuss the general ecology and natural history of the Neotropics could possibly focus on identification of each and every species.

The good news is that you do not by any stretch of the imagination have to be competent in identifying all of the trees, birds, or bugs in order to comprehend, appreciate, understand, and enjoy what you are seeing and experiencing as you walk a rain forest trail. Of course some degree of identification knowledge is very useful: Is that a howler monkey or a capuchin? Is that a toucan or a parrot? Is that tree a legume? Is that flower a heliconia? Thus I have selected to illustrate and discuss examples of widespread organisms that tend to be consistently encountered in many places throughout the Neotropics.

There are two words to keep in mind as I accompany you through the pages of this book: observation and interpretation. I have learned to see the world through the eyes of an ecologist, to read the landscape, to see and comprehend interactions among species. Ecologists typically identify patterns in nature that serve as initial jumping-off points that lead to investigative science. One global pattern, for example, is the distribution of species groups, such as trees, mammals, birds, and beetles. By far the majority of species from these various groups are found in equatorial regions, and species numbers (measured in units of area, such as number of breeding species per hectare) decline sharply as you move away from equatorial regions. Polar regions, the most inhospitable of terrestrial environments, have the fewest species (recall the Antarctica example above). This is a broad and consistent pattern, one that interested Charles Darwin (he talks about it in his most famous book, On the Origin of Species). Ecologists try to learn what the factors are that force such patterns, the causal factors. This example, called a latitudinal diversity gradient, will be discussed in much greater detail later in this book.

Plate B. This is the Saffron Playboy (Xanthiris flaveolata). At first glance it appears to be a butterfly. But, no, it is a day-flying moth, common to southern Amazonian rain forests. Wow, what coloring! It almost glows. That brightness suggests that the insect is noxious to its potential predators, birds. It is likely exhibiting warning coloration (described in chapter 11). But no one has yet shown it to be unpalatable to birds. The pattern of coloration suggests such a conclusion, based on numerous other examples. It awaits further study. Ecologists identify wide-ranging patterns and, using the methods of science, explain why they might be adaptive to the organisms. Oh, and note the tiny fly on its left wing. Who knows what that’s doing there? Photo by John Kricher.

My goal will be to teach you how to spot patterns, to observe, to see, and to understand a tropical ecosystem as an ecologist does. I will describe in some cases how ecologists have cleverly wrested information from nature that has resulted in much greater understanding of infrastructure of tropical ecosystems. As you move through the chapters you will begin thinking like an ecologist. Your focus will be less on putting a name on a particular organism that you encounter and more on understanding what it might be doing and what it might be interacting with (plate B).

The book is written in a reader-friendly, colloquial style that I have attempted to keep relatively free of technical jargon and embedded annotations and citations. A further-reading list is included for each chapter; these are listed before the index. The entries highlighted in boldface are those used directly in the chapter, while the others enhance the lessons of the chapter and take you deeper into the subject matter.

Unlike its two predecessors, the New Neotropical Companion is prolifically illustrated with color photographs throughout. That is huge! Ecology is a visual science. Describing something in words or with a few drawings goes only so far. Seeing it in a color photograph goes so much further. With the kindness of some of the best photographers I know, I have been able to put together a visual tour of the Neotropics to accompany my narrative. The photos tell the story too. Obviously I have not been able to do more than offer a representative sample of what you might see, but it’s a darned good sample. Test it in the field.

My textbook, Tropical Ecology (Princeton University Press, 2011), I will tell you unabashedly, may be of interest if you seek a broader, more rigorous, and technical college-level treatment of the subject matter.

But if you’re heading off to the New World tropics or just want to sit down and read about this extraordinary region of the world: Welcome to the New Neotropical Companion.

Chapter 1  Welcome to the Torrid Zone

Plate 1-1. Sunrise in the Torrid Zone. Photo by John Kricher.

Plate 1-2. The face of a Jaguar (Panthera onca). Photo by John Kricher.

Into the Torrid Zone

The lure of the tropics, the desire to visit a land of relentless heat and humidity, of scorching sun and torrential rain, may at first seem a bit hard to explain (plate 1-1). But it’s not. It can be explained in one image: plate 1-2.

Seeing a Jaguar (Panthera onca) for the first time, in its element, its home, its piece of rain forest, is worth any long bumpy and dusty ride, a few annoying mosquitos, muddy boots, an airport delay, or any other minor inconvenience typical of modern travel. Observing a real live Jaguar in the wild provides a remarkable connectivity with Earth’s natural world that is simply unrivaled. You have really seen something special. And there is so much more. The majesty of myriad imposing tall rain forest trees and the chance of encountering some of the multitudes of creatures that dwell within those forests is an experience that is nothing short of precious (plate 1-3). The tropics, a land of heat and humidity historically termed the Torrid Zone, contains most of the world’s species of, well, pretty much everything: plants, birds, mammals, reptiles, insects, you name it. And that is what this book is about. We are going to visit the Neotropics. Let’s begin with some geographical perspective.

Beginning about 248 million years ago, just after the end of the Permian period and the Paleozoic era, the world’s continents began drifting apart, a process that continues today. This separation is the result of a dynamic geological process known as plate tectonics. Continents made primarily of granite ride passively atop large and slowly moving plates of basalt (which compose the earth’s crust) kept in motion by the convective heat of the planet itself. The result, in a nutshell, has been that widely separated continents now contain markedly different groups of organisms. That is because 248 million years allow for a lot of evolutionary change, for new species to diverge and evolve, for whole new groups of organisms to develop. For example, the primates of the New World tropics (the Neotropics), the monkeys, marmosets, and tamarins of Middle and South America, are distinctly different from the Old World monkeys and apes, though they all (along with us), of course, ultimately share common ancestry in the order Primates (plate 1-4).

Plate 1-3. Tropical rain forest is the most structurally complex and biodiverse terrestrial ecosystem in the world. Photo by John Kricher.

Plate 1-4. These Humboldt’s White-fronted Capuchins (Cebus albifrons) are representatives of the New World monkeys, distinct from the Old World monkeys. Photo by Andrew Whittaker.

Plate 1-5. This colorful Saffron Toucanet (Pteroglossus bailloni), from southeastern Brazil, is in the family Ramphastidae, a group of birds found only in the Neotropics. Photo by Andrew Whittaker.

Plate 1-6. Tropical forests are generally warm and wet throughout the year, mostly due to the constancy of direct solar radiation. Photo by John Kricher.

Hornbills, large birds with colorful, elongate, bananalike bills, are found only in Africa and Asia. However, an anatomically similar but only distantly related group of birds, the toucans, toucanets, and aracaris (plate 1-5; discussed in chapter 8), is found only in the American tropics. Through these and numerous other examples, biogeographers have identified well-separated geographic realms comprising largely distinct floras and faunas.

North America is in the Nearctic biogeographic realm, which is primarily temperate in climate. Europe and northern Asia are in the Palearctic realm and are likewise mostly temperate. The African realm and Australasian realms (including islands such as Borneo and New Guinea) are largely tropical in climate, though with large areas of hot desert. The realm known as the Neotropics begins in central Mexico, extends through the Caribbean region, and reaches to the tip of South America. Although temperate at both its northern and southern extremes, the realm of the Neotropics is largely tropical. Here’s why.

The bulk of the Neotropic land area lies between the Tropic of Cancer to the north and the Tropic of Capricorn to the south, with the equator in the middle. The names Cancer and Capricorn refer, of course, to constellations of the zodiac through which the sun appears to trace its annual course. Because Earth is tilted on its axis, by 23.44°, it is a seasonal planet, its north side facing the sun part of the year (the northern summer months) and its south side facing the sun part of the year (the northern winter months). Thus on the dates of the summer and winter solstices the sun is either 23°26’22 (about 23.44°) north (directly over the Tropic of Cancer) or 23°26’22 south (directly over the Tropic of Capricorn). For those of us at northern latitudes, the sun appears low in the sky in winter and high, virtually overhead, in summer. On the dates of the equinoxes heralding the official beginning of spring and, six months later, autumn, the sun sits directly over the equator.

It is thus obvious that the part of Earth receiving the most solar radiation (i.e., direct sunlight) within the course of a year is the region lying between the Tropics of Cancer and Capricorn, a 46.88° belt that essentially defines what we call the tropics. The fact that the sun is never more than 23.44° north or south of the equator is the major reason the tropics exist. Earth receives different amounts of solar radiation depending upon latitude. But if you are in the Torrid Zone, no matter where on Earth, you get a lot of direct sunlight throughout the year and thus, unless you are at a high elevation, you experience a lot of heat. It is therefore consistently warm in the lowland tropics (plate 1-6).

North of the Tropic of Cancer and south of the Tropic of Capricorn you enter the temperate zone, an area of more extreme annual climate variability. North of the Arctic Circle (66°33’39" N) you enter the northern polar zone, historically termed the Frigid Zone, a region of extreme climatic stress that supports an ecosystem called tundra, realm of musk oxen and polar bears. The same, of course, is true climatically when you cross the Antarctic Circle (66°33’39" S) and enter the southern polar zone, the land of vast ice and diverse penguins.

Immediately beyond the Torrid Zone latitudes you move into the subtropics (plate 1-7). It is not unusual for parts of Florida to experience winter frosts, but nonetheless the southern areas of Florida, including Everglades National Park, are ecologically subtropical. Species of typically tropical plants such as mangroves and the Gumbo Limbo tree (Bursera simaruba) are found here. Many physical characteristics of typical tropical forests (such as the presence of high levels of epiphytic plants and buttressed roots) are evident to various degrees in coastal forests as far north as the Georgia–South Carolina border (plates 1-8–9). This is because the warm oceanic currents of the Gulf Stream (discussed below) allow the coastal Southeast to remain relatively balmy throughout the year, extending the subtropics northward (plate 1-10).

Plate 1-7. The American Alligator (Alligator mississippiensis), here shown in a tableau along with several Yellow-bellied Slider turtles (Trachemysscripta), is a subtropical reptile that is the northernmost representative of its family. This photo is from coastal Georgia, near Savannah. In the Neotropics, there are species similar to alligators called caimans, along with several crocodile species, one of which, the American Crocodile (Crocodylus acutus), just reaches southern Florida. Photo by John Kricher.

But Where, Exactly, Are the Neotropics?

Because of the mild climate provided by the warm and complex current known as the Gulf Stream, the Caribbean islands all are part of the Neotropical realm, even though some (such as the Bahamas and Bermuda) are outside of the Torrid Zone. Therefore, the islands of the Bahamas, the Greater and Lesser Antilles, the West Indies, and so forth, are all in the Neotropics. So if you have taken a trip to Abaco, Jamaica, Puerto Rico, Martinique, or Cuba, you have visited a Neotropical place.

Much of northern and western Mexico is either subtropical or temperate (because of the influence of mountain ranges) in climate, but the lowland eastern slope of Mexico adjacent to the warm Caribbean Sea (Veracruz, for example) is warm and humid. Indeed, it is in this region that you find the most northerly extension of Neotropical evergreen moist forests. The remainder of Middle America, Belize, Guatemala, Costa Rica, Nicaragua, Honduras, El Salvador, and Panama, is Neotropical, and each of these countries contains typical tropical ecosystems.

Plate 1-8. A tree branch covered by epiphytic plants is typical of tropical forests throughout the world. However, this branch belongs to a Southern Live Oak (Quercus virginiana), found not within the Torrid Zone but near Savannah, Georgia, in the subtropics. Characteristics of tropical forests are evident to varying degrees in subtropical forests. Photo by John Kricher.

Plate 1-9. Root buttresses are characteristic of many tropical trees but are not confined to tropical trees. This strongly buttressed tree is a Bald Cypress (Taxodium distichum), photographed in the temperate subtropics near Savannah, Georgia. Photo by John Kricher.

Plate 1-10. The Zebra Longwing (Heliconius charithonia) is widespread in the Neotropics and subtropics, ranging from Texas and Florida through South America. It is one of the only heliconius butterflies to reach North America. Photo by John Kricher.

Finally, and perhaps most obviously, we come to South America. This huge continent contains the largest tract of remaining tropical rain forest in the world, the immense Amazon Basin. While the country of Brazil holds claim to most of Amazonia, all neighboring countries share part of it. Thus you can visit diverse tropical lowland evergreen forest not only Brazil but also in Bolivia, Peru, Ecuador, Colombia, Venezuela, Guyana, Suriname, French Guiana, Paraguay, and parts of northern Argentina.

Note that the southern regions of South America in Chile and Argentina are south of the Torrid Zone and have a temperate, seasonal climate.

It’s Not All Tropical Even in the Torrid Zone

Imagine that you are exiting your hotel in Guayaquil, Ecuador, a sea-level port city, for a brief ride to the airport. You walk out from the air-conditioned lobby into hot and decidedly humid air. It’s muggy and feels very tropical. After a short and scenic flight, your aircraft touches down in Quito, about 267 km (166 mi) away, as the Andean Condor flies. You have left the sultry sea-level climate of Guayaquil for the clear Andean air of Quito, which is located at about 2,800 m (9,200 ft) above sea level. The climate feels decidedly temperate, cooler and drier. That’s because it is.

Plate 1-11. Alexander von Humboldt was the first explorer to carefully document how life zones change with elevation and thus elucidate the importance of mountains, such as the Andes, shown here in Ecuador, to the diversity of ecosystems in the tropics. Photo by John Kricher.

Plate 1-12. Snow in the tropics? Tussock grass and low-growing plants poking out from newly fallen snow might suggest an ecosystem such as the moors of England and Scotland. But this is just outside of Quito, Ecuador, at an elevation of about 3,355 m (11,000 ft), in the Andes Mountains. Photo by John Kricher.

Humboldt on the Rain Forest

The German-born naturalist and explorer Alexander von Humboldt led an expedition to Central and South America from 1799 to 1804. He wrote of his first impressions of rain forest:

An enormous wood spread out at our feet that reached down to the ocean; the tree-tops, hung about with lianas, and crowned with great bushes of flowers, spread out like a great carpet, the dark green of which seemed to gleam in contrast to the light. We were all the more impressed by this sight because it was the first time that we had come across a mass of tropical vegetation But more beautiful still than all the wonders individually is the impression conveyed by the whole of this vigorous, luxuriant and yet light, cheering and mild nature in its entirety. I can tell that I shall be very happy here and that such impressions will often cheer me in the future. (Quoted in Meyer-Abich 1969.)

Latitude alone does not determine the tropics. Elevation is also a critical variable. The youthful and dynamic Andes Mountains run the western length of South America, extending from Tierra del Fuego at the southern tip of the continent all the way north and east through Venezuela, ending in the gentle northern and central ranges of the island of Trinidad. The great European explorer Alexander von Humboldt (1769–1859) was the first to describe in detail how habitats change with elevation (plate 1-11). He is credited with having elucidated the concept that was later formalized and called the ecological life zone (detailed in chapter 2). Humboldt realized that climate characteristics change with elevation and that climate, as you might have surmised by now, is the most important variable in determining what sort of habitat or ecosystem will be present. Humboldt documented how a zone of lowland tropical forest (i.e., rain forest) gradually transitions into cloud forest with increasing elevation and how, above cloud forest, trees become increasingly stunted until a zone is reached called páramo, a cold and windswept ecosystem of tussock grass and dwarfed trees. Snow is common at this elevation. When it comes to ecological montane (i.e., mountainous) life zones, the latitude may be well within the Torrid Zone, but elevation makes all the difference. Indeed, if you have a strong enough arm you could stand on the equator high in the Andes and toss a snowball east into the hot and steaming jungles of Amazonia far below (plate 1-12).

So What Actually Are the Tropics?

The climate of the tropics will form the main topic of the next chapter, but for now, know this: a tropical climate is consistently warm but variably wet. Much of the area within the tropic zone is not rain forest. There are, for example, deserts in the tropics, including the desolate Atacama Desert of coastal Peru and Chile. The African Serengeti, which is a vast savanna of grassland and scattered trees, is famous for the forced annual migrations of herds of large animals such as wildebeest, which must move seasonally to find water. Much of central and southern Brazil is cerrado, an ecosystem ranging from dry forest to open grassland depending upon seasonal moisture input and occurrence of natural fire (chapter 14).

Therefore, a large part of the tropics experiences a seasonal climate in which the seasons vary between rainy and dry rather than hot and cold. Tropical areas have relatively little annual variability in air temperature (and the temperature is generally quite warm), but rainfall amounts may vary dramatically throughout the year, something to bear in mind when you plan a trip. Tropical ecologists speak of wet and dry seasons rather than winter (implying cold) and summer (implying heat). However, in many lowland rain forests rainfall is sufficiently abundant on a daily basis throughout the year that the dry season is scarcely expressed, and thus the forest remains constantly wet and lush year-round. This is true rain forest.

Most people who visit the Neotropics tend to seek a rain forest experience and thus should be prepared for heat, humidity, and a bit of rain, sometimes quite a bit of rain, often daily.

Plate 1-13. Dense, complex tropical rain forest is home to more species than any other terrestrial ecosystem on Earth. Photo by Beatrix Boscardin.

Welcome to the Jungle: A Quick Overview

Because of the relatively constant presence of heat, rain, and humidity, much of the Neotropics is biologically lush, profligate with species of plants and animals (plate 1-13). It is this profusion of diverse life that attracts so many ecologists as well as ecotourists to tropical destinations and has long fascinated explorers and naturalists, including Charles Darwin, Alfred Russel Wallace, and Theodore Roosevelt. Life-forms are not randomly distributed on Earth. Tropical terrestrial ecosystems occupy only about 7% of Earth’s surface but are believed to hold more than 50% of the world’s terrestrial plant and animal species. There is a very basic observation nested in this reality: tropical climate is conducive to supporting diverse life, more so than any other climate to be found on the planet. As climate becomes less warm, less moist, less equable, fewer species are to be found. The term biodiversity, referring generally to the sum total of species present in an area or region, has come into common usage over the past decades. Indeed, one of the principal concerns of ecologists is that the present century will see a severe reduction in global biodiversity, including many species of tropical organisms (chapter 18, and see Conservation Issues, below).

The profusion of life in the tropics takes physical form in the concept of the rain forest biome once commonly referred to as jungle. Tarzan, as most of us know, lived happily in the African jungle, moving easily from one place to another by swinging on vines while hollering loudly. As a child in the early 1950s I used to watch a television show titled Ramar of the Jungle, about a medical doctor facing heroic weekly situations as he plied his way through the jungles of Africa and India. It wasn’t a great show but it did get me interested in the tropics. And one of my first really good introductions to tropical plants and animals was Ivan Sanderson’s classic Book of Great Jungles (1965).

So what is a jungle? The term (derived from the Sanskrit word jangala) has always been associated with dense tropical forests and usually invokes visions of plant growth so prolific as to be virtually impenetrable. The mind’s eye sees massive trees draped with thick vines, a mysterious and somewhat foreboding blanket of vegetation hosting a strange cacophony of birds and insects. But that vision is not entirely accurate, and rain forest and jungle do not mean quite the same thing.

It is true that there are many places in the tropics where the vegetation is so profuse as to require the skilled use of a tool such as a machete to move through it. But these are typically areas of relatively recent disturbance (chapter 7), such as cleared areas that then receive abundant sunlight that promotes rapid and prodigious plant growth. Where there is mature, old-growth rain forest the sense of impenetrability is much reduced. It is actually no more difficult to maneuver through rain forest than through most temperate-zone forest. Large trees are relatively widely spaced, and the lack of light at the ground surface (because of the dense leafy canopy above) prevents much in the way of plant growth that would impede movement along the forest floor.

Tropical rain forest is tall, lush, and most of all diverse with species. (It will be described in detail in chapter 3). It is without doubt the ecosystem that most visitors to the Neotropics seek to experience. Fortunately, there are still many places where that is easily possible.

Visiting the Neotropics

When A Neotropical Companion was first published (in 1989) there were relatively few well-known and reliable tourist facilities within the Neotropics. The famous Asa Wright Nature Centre on Trinidad was one of them. This one-of-a-kind guesthouse, now much expanded, offers easy access to lush forest abounding in tropical wildlife (about 2,200 plant species, 617 butterfly species, and 400 bird species) and has been visited by thousands of naturalists since the 1950s. It remains today one of the premier destinations in which to experience the Neotropics.

Since 1989 Neotropical ecotourism has burgeoned and, unsurprisingly, so has the availability of fine accommodations throughout the region. It is now possible to book a stay at any number of highly comfortable and commodious lodgings (far too many to list in this book) in virtually any Neotropical country, each of which generally offers a tasty (and safe) cuisine, hot showers, clean and comfortable rooms, and, most important, highly competent local guides. One example is the Canopy Tower, located in Soberania National Park, Panama, very near the Panama Canal and just a short drive from Panama City (plate 1-17). This unique facility, located high atop Semaphore Hill Road, is an old radar installation that has been completely renovated, upgraded, and converted to an ecotourism facility of the highest quality. It offers an outstanding rain forest experience that includes daily guided tours to such places as the famous Pipeline Road, one of the most important field sites in historic and ongoing studies of Neotropical ecology (and located very near Barro Colorado Island, one of the most important research stations in the Neotropics). As with most other lodges, it is easy to book reservations on the Internet for the Canopy Tower (and its sister facilities, the Canopy Lodge, Canopy Camp, and Canopy B&B, take your choice).

The Asa Wright Nature Centre and Simla

The Asa Wright Nature Centre bears the name of a courageous and formidable woman, Asa Wright, who, along with her husband Newcombe Wright, developed a lush plantation in the Arima Valley within the northern mountain range in Trinidad, where they grew coffee, cacao, and various citrus species (plate 1-14). Asa and Newcombe Wright hosted many illustrious visitors at their Spring Hill Estate, as the house and property that became the center was then known. Most famous among them was the eminent naturalist, author, and explorer William Beebe (1877–1962). When Beebe was 73 he purchased land in the Arima Valley near the Spring Hill Estate and moved there, starting a field station in 1950, which he named Simla. Beebe welcomed researchers to the tropics and encouraged them to work with live animals, not just collect specimens for museums. Ornithologists such as David and Barbara Snow came frequently and contributed immensely to the understanding of tropical bird ecology and evolution (as will be discussed often in this book).

Large numbers of visitors enjoy the spacious veranda at Spring Hill Estate as well as the many trails that provide access to the Arima Valley and its magnificent forest (plate 1-15). When Newcombe Wright died in 1967 an effort was made to secure the property for education and conservation. With the help of numerous individuals and organizations, including the renowned bird artist Donald Eckelberry and the philanthropist Erma J. Fisk, the Asa Wright Nature Centre was established.

Asa Wright remained a resident of the house until her death in 1971 (plate 1-16). Today the property includes in excess of 400 ha (approx. 1,000 ac) of protected land, including the Dunston Cave, perhaps the easiest place in the Neotropics to see the remarkable Oilbird (Steatornis caripensis; chapter 10). The spacious veranda of the original house overlooking the Arima Valley is one of the best places in the tropics to see and enjoy numerous birds and other species. And yes, complimentary rum punch is served every afternoon, in the fine tradition established by Asa Wright.

Plate 1-14. The magnificent Arima Valley, as seen from the veranda of the Asa Wright Nature Centre. Photo by John Kricher.

Plate 1-15. The famous veranda at the Asa Wright Nature Centre. Photo by John Kricher.

Plate 1-16. Asa Wright’s original living room at Spring Hill Estate, now the Asa Wright Nature Centre, has been and continues to be enjoyed by thousands of visitors over the years. Photo by John Kricher.

Canopy Walkways and Towers

It was long ago realized that many species of tropical animals spend most of their energy foraging in the forest canopy. But that means that everything from colorful butterflies to numerous birds may be 30 m (98 ft) or more away from the ground-based observer. What if you could actually get up into the canopy of a tropical forest, among the foliage, what would you see? The answer is becoming more and more evident, as various research stations and ecotourism lodges erect canopy towers and walkways that allow visitors some limited but significant access to this remarkable zone of life (plates 1-18–19). Carefully engineered and constructed canopy towers and walkways allow you to look directly into fruiting trees and watch monkeys, toucans, parrots, tanagers, and many other species as they forage. There is nothing quite like it.

The View from Above

As astronaut John Glenn passionately stated while he was rocketing high above Earth at the commencement of his orbital mission in Friendship 7 in 1962, Oh, the view is tremendous. That holds true for canopy walkways and towers throughout the tropics.

Plate 1-17. The wraparound observation deck on the Canopy Tower near the Panama Canal in Panama allows for a full, 360° panorama of the landscape at treetop level. Monkeys, toucans, and other tropical animals are easy to observe. Photo courtesy of Canopy Family.

Plate 1-18. This wooden canopy tower at Sacha Lodge in Ecuador literally circles a huge emergent tree, affording views at various levels as you climb the tower. Photo by John Kricher.

Plate 1-19. The canopy walkway at Sacha Lodge allows one to safely peruse a large swath of tropical forest from treetop level. Photo by John Kricher.

My first visit to a canopy walkway was at the ACEER Foundation, the Amazon Center for Environmental Education and Research (plate 1-20). It is located in one of the most species-rich areas in upper Amazonia, along the Amazon River, about 160 km (100 mi) east of Iquitos, Peru. The site includes a superbly engineered canopy walkway about 0.4 km (0.25 mi) in length, an elaborate arboreal pathway interconnected with 14 emergent trees, permitting one to literally walk through the rain forest canopy. Each of the trees used in the walkway is fitted with strong wooden platforms allowing several people to stand and look out at the canopy. The narrow spans between the tree platforms are built rather like suspension bridges, supported by strong metal cable and mesh-lined at the sides to provide total security and safety. The spans vibrate a bit, especially when more than one person is walking across. One of the spans, when I visited, was nearly the length of a football field, affording a breathtaking, if shaky, look at the rain forest below. The first of the platforms is about 17 m (55 ft) above the forest floor, but the spans eventually take you to a platform that is fully 36 m (118 ft) above the ground. From that privileged position, you gaze upon a panorama of unbroken rain forest for many, many miles. The view is tremendous.

From within the canopy you get an immediate, almost overwhelming impression of the richness of the rain forest. Trees are anything but uniform in height—and of so many species that you wonder whether any two along the walkway’s length are the same, or if every tree is different from every other. You notice the many different leaf sizes and shapes and see that some leaves are damaged by leaf-cutter ants, the insects having traveled 30 m (98 ft) up the tree bole to collect food for their subterranean fungus gardens. Now you can really look at the fine details of epiphytic plants such as orchids and bromeliads. You can see down into the cistern-like bromeliads and learn what kinds of tiny animals inhabit these microhabitats high above the forest floor. You note the uneven terrain below and realize that the canopy is by no means continuous but is punctuated by frequent openings of various sizes, called gaps. A male Collared Trogon (Trogon collaris) is perched 6 m (20 ft) below the walkway. How odd it is to actually look down on such a creature. A male Spangled Cotinga (Cotinga cayana), a stunning turquoise bird whose plumage seems to shimmer with iridescence in the full sunlight, sits in display at eye level (plate 1-21).

Plate 1-20. Early morning in the canopy at the Amazon Center for Environmental Education and Research (ACEER), looking down. Photo by John Kricher.

A tree near one of the platforms is in heavy fruit, hundreds of small orange berrylike fruits peppering the branches. Fruit trees normally attract a crowd, and this one is no exception. Colorful tanagers of six different species fly in to feast on the fruits, at most just 3 m (10 ft) away from us. Equally gaudy aracaris and toucanets join the tanagers. Two sedate, long-haired saki monkeys (Pithecia spp.; plate 1-22), apparently a female and an adolescent, stop at the fruiting tree. The monkeys’ long, bushy tails hang limply below the branch on which they sit, as these simians do not have prehensile tails, as do their forest cohabitants, the howler, spider, and woolly monkeys (see chapter 16 for more on monkeys).

Plate 1-21. A male Spangled Cotinga at eye level, a view that is impossible unless you have access to the rain forest canopy. Photo by Gina Nichol.

The simians soon realize they are not alone. The female sees us and rubs her chin on the branch. She stands fully erect and emits a short, demonstrative hoot to warn us to come no closer. She needn’t worry. We are not about to leave the security of the walkway. We marvel at how monkeys have adapted the requisite skills to move effortlessly through such a tenuous three-dimensional world as the rain forest canopy. A frenetic Amazon Dwarf Squirrel (Microsciurus flaviventer), a chipmunk-size evolutionary relative of the northern acorn collectors, scurries with nonchalance on the underside of a branch over 30 m (98 ft) from the ground below. A thought occurs, and recurs many times: from the ground, we’d never know this little animal was up here.

The canopy walkway affords a unique and broad window into the life above the forest understory. It is exciting to visit it, to be on it at dawn, when the forest below is still clothed in mist, or to watch the sun set over what seems like an endless vista of rain forest. But it also affords an opportunity for the kind of research that needs to be done to accurately ascertain an understanding of the rhythms of life in this essential habitat.

Ecotours and Lodges

The Canopy Tower and its associated sites the Canopy Camp and Canopy Lodge (in Panama), noted above, are all wonderful destinations in the Neotropics. And there are numerous other fine lodges and facilities. A quick but thorough Internet search will turn up many possibilities for places to stay, depending upon which countries you wish to visit and how long you wish to stay. For example, if you want to sail the Amazon River, you can book passage on a comfortable, well-appointed boat and take a river tour from Iquitos, Peru, or Manaus, Brazil.

Another approach to visiting the Neotropics, also easily accomplished on the Internet, is to book an ecotour with an established tour company. Many such companies exist, and they offer customized itineraries tailored to show you the most wildlife in the shortest time period, all the while attending to your comfort and safety. Many of these tour companies routinely base their tours at outstanding facilities such as the Asa Wright Nature Centre (Trinidad), the Lodge at Pico Bonito (Honduras), the Canopy facilities (Panama), Monteverde and Selva Verde (Costa Rica), Chan Chich Lodge (Belize), Explorama (Peru), and Sacha Lodge (Ecuador), a list far from exhaustive. Most will categorize their tours as to whether or not they are strenuous, relaxed, and so forth. Many companies specialize in birding tours, but even highly focused birding tours rarely neglect the many other amazing animals as well as plants that are part of the tropical landscape. If you elect to take an ecotour, once you meet your guide everything else is pretty much done for you. You are told when to be at breakfast, how long you will be in the field, what to wear, what to expect, and there is usually a nightly session (either before or after dinner) devoted to summarizing the day’s observations and briefing you about tomorrow’s activities. It is a great way to learn a lot in a short time span.

Plate 1-22. Face to face with a saki monkey. Photo by Sean Williams.

Finally, if sufficiently independent, you might want to go it alone. The Pan American Highway extends through Mexico and Central America, stopping within the Darién in Panama. It picks up again in Colombia and extends south from there through Ecuador, Peru, and Chile. I don’t recommend that you drive the length of the Pan Am Highway unless you are highly adventurous. It is much easier to pick a destination, fly there, and rent a vehicle if you choose to explore on your own. Be aware that if you are on your own you must take care to assure all aspects of your comfort and safety. In most places, knowing a reasonable amount of Spanish is essential (or Portuguese if you are in Brazil).

What about Bugs, Spiders, and Snakes?

It is the question I am asked the most often. If I go anywhere in the Neotropics, am I going to be consumed by hordes of biting insects (that are sometimes vectors for malaria and other serious maladies), carried off by army ants, or bitten by a venomous spider or snake?

The patterns of arthropod abundance as well as the ecology of army ants will be treated in detail (chapter 10), as will snake distribution and abundance (chapter 16). But for this overview, suffice it to say that indeed, mosquitoes, ticks, and biting flies of various species are present, on occasion in abundance. Mosquitoes tend to be most numerous in rainy season, while forest ticks sometimes abound in dry season. It is wise to bring ample repellent (yes, with Deet, though some less noxious organically based repellents suffice in many situations) and to keep yourself reasonably well covered when you can plainly see that biting insects are present. If you are going into a region where forms of malaria are known to occur, consider consulting your physician about taking a malaria-prevention drug beforehand. Caution also suggests getting inoculated for yellow fever prevention, as that mosquito-vectored virus occurs in many Neotropical areas. Some countries require that you be inoculated against yellow fever before they permit you to enter. See the Appendix for more on this subject.

Spiders and scorpions are generally little cause for any alarm, but a few species can inflict sufficient toxin that one needs to observe prudent caution when around them. Most people seem to have a more or less natural aversion to picking up spiders and scorpions—and rest assured, the spiders and scorpions prefer it that way. But also be aware that spiders and their