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The giraffe biology ecology evolution and behaviour 1st Edition Bryan Shorrocks
The Ecology and Evolution of Tropical Biodiversity
Jan Sapp
Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide. Oxford is a registered trade mark of Oxford University Press in the UK and certain other countries.
Published in the United States of America by Oxford University Press 198 Madison Avenue, New York, NY 10016, United States of America.
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, by license, or under terms agreed with the appropriate reproduction rights organization. Inquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above.
You must not circulate this work in any other form and you must impose this same condition on any acquirer.
CIP data is on file at the Library of Congress
ISBN 978–0–19–063244–1
1 3 5 7 9 8 6 4 2
Printed by Sheridan Books, Inc., United States of America
For Camille Limoges
CONTENTS
Acknowledgments ix
Interviews xi
1. The Other World 1
2. Legends 11
3. Romancing the Rainforest 25
4. Regeneration 40
5. Is Evolution Different in the Tropics? 52
6. Niche Construction 62
7. Rhythms of the Forest 78
8. On the Waterfronts 93
9. The New Deal 104
10. Ecology in Disequilibrium 114
11. The Central Enigma 126
12. Liberated from Fashionable Science 136
13. Territories, Taxonomy, and Time 151
14. Nineteen Eighty-Nine 165
15. Biodiversity in Heat 174
16. A Continent in the Canopy 185
17. At the Root of Diversity 197
18. The Other World Today 208
Notes 215 Index 267
ACKNOWLEDGMENTS
My interest in this history has its genesis in my book about coral reef environmental science: What is Natural? Coral Reef Crisis. In 2007, I contacted Ira Rubinoff, then Director of the Smithsonian Tropical Research Institute, whom I had first met in 1996 at the Eighth International Coral Reef Symposium, which the institute hosted in Panama. I told him I was interested in writing another book on the history of tropical biology as soon as I completed the project I was working on— the history of microbial evolutionary biology. He asked me if I’d consider writing the history of STRI. In the following years, his successor, Biff Bermingham, gave me access to the facilities, invited me to become a research associate, and facilitated my research and interactions with Smithsonian biologists.
Many people have collaborated in this project by generously providing their time for interviews and/or reading draft chapters, correcting errors, and offering suggestions. George Angehr, Allen Herre, Stephen Hubbell, Jeremy Jackson, Nancy Knowlton, Eugene Morton, and Ross Robertson read specific draft chapters. Ira Rubinoff also offered helpful comments. I thank Carole McKinnon for her readings of the manuscript and support throughout the whole project. I am also thankful to Egbert Leigh for his support throughout the writing of this book and for his close and careful reading of the entire manuscript, which was simply invaluable.
I am grateful for the help of Nancy Korber at the Fairchild archives, the help of Mary LeCroy in the Department of Ornithology of the American Museum of Natural History, Anthony Walker for the adventure of floating across the rainforest canopy and among the trees on the Sherman Forest Crane in Panama, Jorge Aleman, and the Smithsonian Tropical Research Institute for the use of photos. I am grateful to the Smithsonian Tropical Research Institute and the Social Sciences and Humanities Research Council of Canada for their support of this work.
Thomas Barbour Papers, Pusey Library Archives, Harvard University, Boston.
Frank Chapman Papers, Department of Ornithology Archives, American Museum of Natural History, New York.
Eugene Eisenmann Papers, Department of Ornithology Archives, American Museum of Natural History, New York.
David Fairchild Papers, Center for Tropical Plant Conservation, Fairchild Tropical Botanic Garden, Miami, Florida.
Smithsonian Institution Archives, Washington, DC.
Andrew W. Mellon Foundation Archives, New York.
George Angehr
Peter Ashton
Lisa Barnett
Eldredge Bermingham
James Bever
Charles Birkeland
Anthony Coates
Rachel Collin
Laurel Collins
Timothy Collins
James Collins
Richard Condit
Richard Cook
Barbara Davis
Robert Dressler
Kerry Dressler
Robin Foster
Allen Herre
Stephen Hubbell
Jeremy Jackson
Gabriel Jacome
INTERVIEWS
Nancy Knowlton
Egbert Leigh
Harris Lessios
Olga Linares
Lena Lombardo
Scott Mangan
Eugene Morton
Leonor Motta
Jorge Motta
Aaron O’Dea
Dolores Piperno
Patricia Rand
Ira Rubinoff
Roberta Rubinoff
Ross Robertson
William Robertson
Henry Stockwell
William Wcislo
Donald Windsor
Joseph Wright
Coexistence
CHAPTER 1
The Other World
The traveller and the naturalist have combined to praise, and not infrequently to exaggerate the charms of tropical life—its heat and light, its superb vegetable forms, its brilliant tints of flower and bird and insect . Each strange and beautiful object has been described in detail … But so far as I am aware, no one has yet attempted to give a general view of the phenomena which are essentially tropical, or to determine the causes and conditions of those phenomena. The local has not been separated from the general, the accidental from the essential; and, as natural result, many erroneous ideas have become current as to what are really the characteristics of the tropical as distinguished from the temperate zones.
Alfred Russel Wallace, Tropical Nature 18781
This book is about how biologists have grappled with the evolution and ecology of the great species diversity in tropical rainforests and coral reefs. Tropical rainforests are home to 50% of all the plant and animal species on earth, though they cover only about 2% of the planet. Coral reefs hold 25% of the world’s marine diversity, though they represent less than 1 percent of the world’s marine environment. The increase in species diversity from the poles to the tropics has remained one of nature’s greatest enigmas for more than two hundred years. Why are there so many species in the tropics? How can so many species coexist there?
At a time when rainforests and coral reefs are rapidly shrinking, when the earth is facing what has been called the sixth mass extinction, understanding the evolutionary ecology of the tropics is everyone’s business.2 This book is written for anyone who is interested in the ecology and evolution of life on earth. Readers require little prior knowledge of the science as we follow the development of some of the major theories and controversies that have shaped research on the evolution and ecology of tropical diversity to the present day.
The tropics, which had inspired the evolutionary theories of Charles Darwin and Alfred Russel Wallace in the nineteenth century, remained a largely unknown and unruly world for evolutionary biology and ecology during much of the twentieth century. It was not certain if principles that worked for temperate zones would apply to
the tropics. Indeed, leading biologists have long debated whether uniquely “tropical” principles were required to understand evolutionary and ecological processes. Every conceivable evolutionary scenario has been posited. Some evolutionary biologists have suggested that evolution in the tropics is faster, more “progressive,” and more creative than elsewhere. Yet others have suggested to the contrary that evolution in the tropical rainforest is in fact probably slower, and that tropical rainforests are more like museums that merely hold ancient species, many of which evolved elsewhere but are now extinct. Some have maintained that Darwinian processes of evolution based on the struggle for existence and survival of the fittest may not apply to the evolution of the great diversity of tree species in tropical rainforests; others have suspected that they would. As we shall see, most of these issues have only begun to be resolved in recent years.
However, the “mystery of mysteries” for many tropical biologists was not the origin of species, but the coexistence of species. It was not why there are so many species in the tropics, but how so many species can coexist. Are there more habitats in the tropics? Does every tropical species in rainforests and coral reefs have an exclusive niche that enables it to avoid competitive exclusion? Are species more specialized in the tropics? These were the questions at the interface of ecology and evolution—questions of evolutionary ecology. Discerning the unfamiliar patterns and processes in the chaos of the jungle and the glittering bustle of life on coral reefs has proved extraordinarily challenging. A variety of hypotheses has been proposed for species coexistence in the wet tropics and various creative methods devised to test them.
Two main competing visions have persisted to the present day. One maintains that tropical rainforests and coral reefs are extraordinarily complex communities of tightly integrated species that have coevolved and built up niche within niche and therefore possess a predictable species composition in a stable environment. The other vision could not be more different. It holds that tropical communities are not tightly integrated communities maintained by complex species interactions, but rather are more like chaotic species assemblages, and their composition is largely the result of chance. Debates over these two paradigms, as we shall see, have been fierce.
Despite the fundamental importance of the tropics to all of life on earth, tropical biology has evolved relatively slowly and often with difficulties—economic, political, and environmental. The world’s distribution of biologists largely mirrors economic wealth, not ecological wealth. The richest biotic communities are in the poorest and often the least stable countries. Tropical rainforests present the perils of heat and humidity, of malaria, yellow fever, hepatitis, Leishmaniasis, dengue fever, parasitic worms, ticks, and poisonous snakes for those who explore their rich nature. Little wonder the wet tropics have been given the contradictory epithets of “El Dorado” and “Green Hell.”3
The history of tropical biology in this book is situated in the sociopolitical and natural world of Panama. Panama’s location nine degrees from the equator, its species-rich forests, and unique geological history, with coastlines on the Pacific and Caribbean, have made it a natural laboratory for ecologists and evolutionary
biologists. It has been ground zero for testing hypotheses and studying problems of species coexistence in tropical rainforests.
The development of research on tropical biology in Panama is a story of another remarkable coexistence that existed between biologists aiming to study nature’s diversity and the Panamanian and American governments. The construction of the Panama Canal has long been hailed as one of the greatest triumphs in engineering of the past century. The development of biological research in Panama is a lesserknown aspect of American history, yet it is an extraordinary saga of establishing and maintaining biological research in the contexts of a politically turbulent and often dangerous century there.
For American naturalists early in the last century, establishing a research station in the tropics was, in the first instance, a matter of access. Barro Colorado Island, located in the middle of the Panama Canal, offered that accessibility. When a research station on the island first opened in 1924, it consisted only of a modest building where naturalists could live for a while and work in the surrounding rainforest. There was nothing like it anywhere. Funded largely out of the pockets of its founding naturalists and their friends, it quickly became one of the most important places in the world devoted purely to the study of tropical nature.
The Smithsonian Institution administered the station beginning in 1946, but the station languished somewhat until the early 1960s, when, stimulated by renewed interest in evolution, ecology, and conservation in the tropics, it was transformed into a major research institute with its own scientific staff and with hundreds of visiting researchers annually. It grew rapidly beyond the island and extended to comprise marine biological stations on both coasts of the isthmus.
My aim is not to provide an overview of all the research conducted on tropical biology in Panama. It is rather to understand the way that tropical biology evolved in context, and the great transitions that punctuate its history. We follow its evolution from naturalists’ descriptions of the diverse species and their habits and habitats, to the formulation of general theories of biodiversity. We discuss the competing hypotheses in regard to species coexistence in the tropics, the controversies surrounding them, and diverse ways of testing them. In so doing, we observe the transition from individual research efforts to the formation of interdisciplinary and international teams and networks that study some of the most important problems in evolutionary ecology and most pressing conservation issues of our times. The keystone problem is understanding the basis of tropical diversity.
THE WORLD’S GREAT PATTERN
Naturalists for centuries perceived life in the tropics as belonging to an alien “other” world. In the fifteenth and sixteenth centuries, European representations of the tropics were founded on the imaginings of artists, based on the recollections of sailors and soldiers. The tropics of Brazil were depicted as a strange, unfamiliar world inhabited by chimeric forms with humanlike faces, and gargoyle-like heads set on peculiar bodies, and in which monstrous serpents rose from the depths (see Figures 1.1–1.4).4
, 1557–1558. Courtesy of the Library of Congress, USA.
Figure 1.1 Sloth. From André de Thevet, Les Singularitez de la France Antartique
Figure 1.2 Toucan. From André de Thevet, Les Singularitez de la France Antartique, 1557–1558. Courtesy of the Library of Congress, USA.
Figure 1.3 “Su,” probably based on a garbled description of an anteater. From André de Thevet, Les Singularitez de la France Antartique, 1557–1558. Courtesy of the Library of Congress, USA.
The French Royal Academy of Science sent a scientific expedition to the tropics in 1735 to measure the circumference of the Earth and to test a conjecture of Isaac Newton’s that the Earth was not a perfect sphere, but rather bulged around the equator and flattened at the poles.5 They arrived in Quito, Ecuador, the next year, and after completing their measurements in 1743, Charles Marie de la Condamine and his colleagues returned to France by the longer and more dangerous route up the Amazon River, conducting the first scientific exploration there.6
Specimens of wonderful forms of often economically important plants were brought back to European imperial powers from expeditions in the eighteen and nineteenth centuries.7 Joseph Banks’s voyages to the South Pacific and Brazil with Captain James Cook, and those he sent around the globe, made the Royal Botanic Gardens, Kew, perhaps the preeminent gardens of their kind in the world.8 But the tropics signified much more than a collection of more or less economically valuable plants when naturalists turned to study nature’s processes.
The greatest pattern in natural history on earth, the species diversity gradient— the increase in species richness from the poles to the tropics— was apparent to eighteenth- and nineteenth- century explorers. Alexander von Humboldt, who travelled extensively in Latin America—Mexico, the Andes, and the Orinoco Basin of Venezuela—between 1799 and 1805, attributed the rich species diversity of the
Figure 1.4 Sir Walter Raleigh witnesses a crocodile, depicted as a sea serpent, devour a crew member on the Orinoco River, 1595. From Theodor de Bry, Américas, 1625. Courtesy of the Library of Congress, USA.
tropics to the lack of a freezing winter. “Thus, the nearer we approach the tropics the greater the increase in the variety of structure, grace of form, and mixture of colors, as also in perpetual youth and vigour of organic life. This increase may readily be doubted by those who have never quitted our own hemisphere, or who have neglected the study of physical geography.”9
Humboldt’s writings brought the complexity, beauty, and wonder of the tropics to the attention of European naturalists. His accounts of his travels to the New World inspired Charles Darwin, then a student in Edinburgh, to plan his own expedition to the tropics. His voyage on the HMS Beagle—from 1832 to 1836 circumnavigating the globe is legendary. In his Journal of Researches on that voyage, he compared the primeval tropical rainforest of Bahia Brazil to another planet:
Epithet after epithet is found too weak to convey to those, who have not visited the intertropical regions, the sensations of delight which the mind experiences … The land is one great wild, untidy, luxuriant hothouse How great would be the desire in every admirer of nature to behold, if such were possible, another planet; yet to every one in Europe, it may be truly said, that a distance of a few degrees from his native soil, the glories of another world are open to him.10
Two years after his voyage on the Beagle, Darwin developed his theory of evolution by natural selection, that is, the “survival of the fittest.” But he did not publish it until twenty years later, when, in 1858, he received a letter from Alfred Russel Wallace from the Malay Archipelago containing a manuscript describing essentially the same theory.11 Inspired by the chronicles of Humboldt and Darwin, Wallace had spent twelve years living in the tropics; eight of them (1854−1862) were in the Malay Archipelago (Malaysia, Singapore, the islands of Indonesia, and New Guinea), where he developed his theory of evolution by natural selection independently of Darwin. Wallace also posed some of the central questions that came to define tropical biology.
“The luxuriance and beauty of Tropical Nature is a well- worn theme,” Wallace wrote in his book Tropical Nature in 1878. Therein he not only described the insects, reptiles, mammals, and birds, and the lush plant life, he also aimed to understand what was unique to tropics, what general phenomena were essentially tropical, and to determine the cause and conditions of these phenomena. Animal life and plant life were generally more abundant and varied within the tropics than in any other part of the globe, Wallace observed. “Endless eccentricities of form, and extreme richness of colour are its most prominent features.”12 The cause of these “essentially tropical features,” he said, were not to be found in the simple influence of solar light and heat, but rather in their constancy, not only throughout the year, but over eons, not having been much affected by successive glacial periods that destroyed many life forms in temperate zones. The “equatorial lands,” he surmised, “must always have remained thronged with life.” They were “a more ancient world.”
Wallace suspected that evolution in tropical zones was different from that in temperate and frigid zones. In the latter, the kinds of characteristics that could evolve were constrained by a constant struggle for existence against the vicissitudes and severities of climate. This was not so in the tropics: “The struggle for existence as against the forces of nature was there always less severe,—food was there more abundant and more regularly supplied,— shelter and concealment were at all times more easily obtained.”13 Evolution in the tropics was largely a matter of “those complex influences of organism upon organism” and the uninterrupted nature of evolution over eons. These were “the main agents in developing the greatest variety of forms and filling up every vacant place in nature.” Wallace put his tropical conceptions in a nutshell:
The equatorial zone, in short, exhibits to us the result of a comparatively continuous and unchecked development of organic forms; while in the temperate regions, there have been a series of periodical checks and extinctions of a more or less disastrous nature, necessitating the commencement of the work of development in certain lines over and over again. The equatorial regions are then, as regards their past and present life history, a more ancient world than that represented by the temperate zones, a world in which the laws which have governed the progressive development of life have operated with comparatively little check for countless ages, and have resulted in those infinitely varied and beautiful forms— those wonderful eccentricities of structure, function, and of instinct— that rich variety of colour, and that nicely balanced harmony of relations— which delight and astonish us.14
All of this was wholly guesswork, to be sure, but tropical biologists over the next century discussed and addressed all of these ideas in an effort to understand the greater richness in tropical biodiversity compared to that of temperate and cold lands. It was far from certain how tropical environments affected the evolutionary potentiality of life there. Belief that the species richness of the tropics was the result of continuous uninterrupted evolution, a world that was little affected by the climatic disturbances of the temperate zone, was widespread. So too was the idea that other different processes of evolution might operate there.
PANAMA
American naturalists, like their European counterparts, also set out on expeditions in the nineteenth and twentieth centuries, making collections of specimens, which they brought back to be classified and curated at the US National Herbarium in Washington, DC, the Museum of Comparative Zoology at Harvard, the Peabody Museum at Yale, or at the American Museum of Natural History in New York.15 By the turn of the twentieth century, the United States had acquired a dispersed tropical empire as Spain relinquished control of Cuba, Puerto Rico, the Philippines, and Guam to the United States following the Spanish-American War.
But it was Panama that became the center of interest for American science in the tropics. The building of the canal brought engineers and geologists to Panama as well as biologists, especially entomologists from the US Department of Agriculture (USDA) who studied mosquito vectors of diseases.16 Control over infectious disease was critical to the success in the building of the canal. The earlier attempt by the French, who had acquired the rights to build the canal across the isthmus of what was then the Republic of Colombia in 1878, was disastrous. Digging began in 1881 and ended in tragedy eight years later, by which time more than twenty- two thousand workers had died, mostly from yellow fever and malaria.17
The US Congress authorized President Theodore Roosevelt to buy out the French interests. It aimed for a treaty that would pay Colombia a percentage of the tolls and give itself a six-mile- wide zone along the canal. When the Colombian Senate voted down the offer in 1903, Roosevelt supported a Panamanian separatist movement, which would allow the United States to build the canal. Following Panama’s secession from Colombia that year, a new treaty was quickly signed that gave the United States ownership of the canal and sovereignty over a zone extending 8 km (five miles) on each side of the centerline of the canal (with the exception of Panama City and Colón). The Canal Zone, inhabited by canal employees, the American military, and their families, would be wholly government run, with schools, churches, hospitals, libraries, and a commissary.
At first, the United States was on track to fail, just as France had, as yellow fever returned and killed many workers. Then, in 1905, Colonel William Gorgas, Chief Sanitation Officer of the Canal Zone, unleashed one of the most extensive sanitary campaigns in history.18 More than four thousand people worked on his mosquito
brigade in Panama. They drained ponds and swamps or covered all sources of standing water with kerosene to prevent mosquitos from laying eggs, fumigated areas infested with mosquitos, and isolated disease- stricken patients with screening and netting. They managed to eradicate yellow fever in the Canal Zone by 1906, and to contain malaria during the ten- year period of canal construction.
When the construction of the Panama Canal began in 1904, American naturalists appealed for a biological survey to document native flora and fauna and their distribution in the Canal Zone prior to the completion of the Panama Canal, just as such provisions had been made prior to the building of the Suez Canal. Those appeals were finally heeded in 1910, after various American scientific organizations requested that the Smithsonian Institution undertake the work, and the proposal was taken directly to President William Howard Taft.19 Collections of specimens from the survey were taken to the new Natural History Building of the Smithsonian in Washington, DC.
Opened in 1914, the Panama Canal was considered “the greatest engineering feat of the modern age,” cutting the sailing distance from New York to San Francisco by about 12,000 km (seven thousand miles).20 American success in constructing the canal took advantage of the Chagres River, which John Stevens, the head engineer of the canal project, proposed be dammed. It was the largest dam of its kind and resulted in the creation of Gatun Lake, which took ships through 33 km (twenty-one miles) of their 77 km (forty-eight-miles) transit across the isthmus.
BARRO COLORADO ISLAND
The flooding of lowlands formed several islands, formerly hills in a valley. Barro Colorado was the largest of them. Before the flood, the name Barro Colorado had referred to a few settlements where the flow of the river revealed the red mud of its shores. All the old settlements had fallen beneath the waters of Gatun Lake by 1914. Barro Colorado Island has an area of 3,840 acres (over five square miles) and a diameter of approximately 3.5 miles; it rises to a height of about 450 ft above the lake and is forested throughout.
“Red Mud Island” was a gross misnomer; it belied its rich, green, lush forest. The main jungle roof was about 30 m (100 ft) high. The waters did not rise suddenly; there was no deluge. Gatun Lake was four years in filling. Still, the island, it was said, resembled Noah’s Ark because of the great diversity of animals that took refuge there from the rising flood: puma, ocelots, tapirs, sloths, monkeys, deer, agoutis, peccary, and various species of small mammals.21 The local nimrods declared the island to be a “hunter’s paradise.”22
To American naturalists searching to develop research in tropical biology in the 1920s, Barro Colorado Island seemed to be the perfect spot for a jungle laboratory. It was in the Panama Canal Zone over which the United States had sovereignty. The Zone had hospitals, a commissary, and other facilities that would be great assets for building and maintaining an island laboratory. The island was two miles across the
lake from a railway station at Frijoles, a fifty-minute train ride from Panama City. One could leave the island and be in the city two hours later.
The Barro Colorado Island Laboratory was certainly not the first station in the tropics. There was a long tradition of colonial governments establishing gardens in their tropical holdings. The Dutch government had established palatial gardens and biological laboratories in Buitenzorg in Java (now Bogor) in 1817.23 Botanists there introduced to Java such plants as the Australian Eucalyptus , tobacco, maize, and Liberian coffee. Those gardens also played an important role in the introduction of quinine produced from the bark of the Cinchona tree, originally from Peru, used for treating malaria. Research there was also conducted on diseases that threatened economically important plants such as coffee and sugar cane. The British government also had maintained a fine garden and laboratory buildings in Calcutta since 1786 and in Ceylon since 1821. The French founded the Saigon Zoo and Botanical Gardens in 1865, but all of these colonial gardens were far removed from the jungle.
The Barro Colorado Island Laboratory was different. It was situated in the midst of an isolated rainforest, and it was not concerned with problems of economic importance to the United States or Panama. It was the most prominent tropical research station in the world by the late 1920s, and it grew into a major tropical research institute focussed on studying the ecology, evolution, and natural history of tropical life.24 How that island laboratory came to be, how it was managed and emerged into prominence, is a tale of complex relations among biological explorers and entrepreneurs, philanthropists, scientific organizations, and government institutions.
“I
got Barro Colorado Island.”
CHAPTER 2
Legends
James Zetek, April 19, 19231
When the research station on Barro Colorado Island opened its doors in March 1924, it was a modest, all-in-one building, a laboratory and dormitory in the jungle where small parties could live and work. Some naturalists referred to the island itself as a “natural laboratory.” It was seen as a piece of “pristine primeval forest,” “the Naturalist’s Paradise,” a veritable “Garden of Eden,” luxuriant in its primitive growth, and a source of biological inspiration.2 It quickly emerged as the most scientifically productive tropical biology station in the world.
The story of the establishment of the jungle island laboratory is complex and clouded in confusion and controversy. Here we untangle that history through close study of the discourse, aims, and strategies of a small group of naturalist adventurers—a coalition of Harvard professors and biologists who worked for the USDA. In doing so, we shall see the unique and fragile ground on which such a research station stood when dedicated solely to the natural history of the tropics.
CITY TO FOREST
Our story begins far to the north in the midst of a cold, dark winter when a group of American naturalists, brought together by the National Research Council (NRC), held a series of meetings in Washington, DC, to promote scientific and economic development in tropical regions. The NRC had been established during the First World War to further scientific and technical services for the military, and it continued in a nonmilitary capacity during the 1920s and 1930s with an aim “to promote co-operative research among academic institutions and disseminate information in regard to research opportunities.”3
In January 1921, the biologists at those meetings formed the Institute for Research in Tropical America. Comprised of twenty- two member institutions, including universities, museums, botanic gardens, and other scientific organizations, it would aim to establish a tropical research station in Panama. A temporary executive committee was formed with Albert Hitchcock as chairman. Hitchcock, a botanist who worked for the USDA and the Smithsonian National Herbarium, was an experienced field naturalist who traveled the world to amass one of the world’s largest collections of grasses at the National Herbarium. Hitchcock had been to Panama in 1911 when the Smithsonian Institution organized a survey of the flora and fauna in the Canal Zone before the canal was completed (Chapter 1).4
Panama, with its American-governed Canal Zone, with hospitals, commissary, and other amenities, was indeed the obvious choice for a research station. It was connected by steamship to all parts of the world and could be reached by scientists from both sides of the continent, and it was relatively free of tropical diseases.5 However, the executive of the Institute for Research in Tropical America certainly had no intention of building a research station in the midst of a rainforest when they met to discuss plans at the Cosmos Club in Washington, DC, in October 1922. At that time, they were aiming for a research station in Panama City, one that would be comparable to what the British had in the Peradeniya Gardens of Ceylon, the Dutch in the Gardens of Buitenzorg, and the French in Saigon.6 Their idea was to erect a research station next to a site that had been designated for the construction of a laboratory for the study of tropical diseases— the Gorgas Memorial Laboratory in Panama City. They also aimed to have a marine station just outside of the city.
In the spring of 1922, Thomas Barbour travelled to Panama on behalf of the newly formed institute to investigate conditions.7 He was the right man for the job. Thirtyeight years old, he was already a seasoned scientific explorer with considerable experience working in the tropics, and he spoke Spanish. He had first visited Panama on a collecting expedition for Harvard’s Museum of Comparative Zoology in 1909. He had completed his doctorate at Harvard two years later, and then worked as curator of reptiles and amphibians at the Museum of Comparative Zoology. He would become its director in 1927.
Though his specialty was herpetology, Barbour was a general naturalist, a dying breed, interested in many aspects of natural history. He studied birds, insects, and especially butterflies; he was keenly interested in botany and plant introduction.8 He was also a man of means. His father, Colonel William Barbour, was founder and president of the Linen Thread Company, the largest company of that kind in the world, which employed some 2,000 people in Lisburn, Northern Ireland.9 Tom Barbour himself would attain considerable wealth on the stock market, and he would play a central part in establishing the Panama research station, funding it largely with his own money.
Barbour had also played a key role in the founding of the Harvard Botanical Gardens in Soledad, Cuba, a few years earlier. In 1919, he created an endowment fund for Harvard dedicated to “tropical research in economic botany,” and he funded the construction of Harvard House, which served as headquarters,
laboratory, and living quarters. 10 Until the Cuban Revolution, classes for Harvard students were held there in horticulture and collecting botanical specimens. Barbour was custodian of those botanical gardens for two decades beginning in 1927. 11
During his trip to Panama in the spring of 1922, Barbour met with Richard Strong, founding director of Harvard’s School of Tropical Medicine, to discuss the idea of building a research station near the proposed Gorgas Memorial Laboratory in Panama City.12 Strong was enthusiastic about the idea, but plans would shift from city to island forest after Barbour told a resident of Panama, James Zetek, an entomologist working for the USDA, of their plans. Several months later, Zetek would learn of the possibility of acquiring Barro Colorado Island as a preserve for scientific study.
Born in Chicago, Zetek had moved to Panama after graduating with an A.B. degree from the University of Illinois in 1911 to work as an entomologist for the Isthmian Canal Commission. In 1914, he married Maria Luisa Gutierrez, a member of a highly respected Panamanian family. Over the next six years, he held various positions, including professor of natural sciences at Panama’s Instituto Nacional, before beginning work for the USDA.13 Zetek was a member of the Ecological Society of America and head of a committee on conservation for the Panama Canal Zone, and he was knowledgeable about areas that might be studied. Hitchcock wrote to him from Washington on November 17, 1922, asking for information about sanitary and social conditions, regions to be studied, road conditions, railroads, and steamships.14
In early March 1923, Colonel William Erwin, Chief Land Inspector, and A. H. Becker, Land Agent of the Panama Canal Land-Lease Division, suggested to Zetek the idea of acquiring Barro Colorado Island as a scientific preserve and building a station there.15 Erwin and Zetek were very concerned about the depletion of the tropical forest in the Canal Zone because of the large number of leases (about 1,800) given to individuals for small- scale agriculture. Virgin forests were cut down and burned. Barro Colorado Island, an island of some four thousand acres, was considered to be the only spot along the entire canal where the tropical jungle was almost just as it had been before the canal was built. Four leases had been given to settlers on the island, from 1 to 5 ha each, but Erwin managed to put a halt to all others.16
Acquiring the island as a preserve and location for the proposed research station was an inspired suggestion, as Zetek saw it. He immediately relayed the idea to Hitchcock, chair of the Institute for Research in Tropical America, on March 4, 1923: “The land agent and his chief inspector, Messers Becker and Erwin, have very kindly suggested the possibility of getting a very large island in the canal, easily accessible and in virgin state… The only cost is the clearing for the laboratory site and the building. In the meantime no leases will be let until we know of your decision.”17
Zetek had not yet seen the island. He did so after he discussed the idea with Strong and two entomologists who were attending a ceremony to lay the first stone for the Gorgas Memorial Laboratory in Panama City— Cornell Professor Oskar Johannsen and Harvard University’s William Morton Wheeler.18 The four of them made a trip to the island on March 20, 1923. They stayed for less than an hour, but it was enough
time to convince them that it was a great spot for a station. Wheeler wrote the next day to his friend and colleague Tom Barbour about their visit to the island:
Yesterday Zetek, Johannsen, Dr. Strong and I spent an hour on the island which is reached by launch in 20 minutes from Frijoles on the railroad, not far from Colon. In a small clearing, less than an acre in area, I took 19 species of ants, Zetek took 10 species of termites (1 new species) and both of us took a dozen species of myrmecophiles and termitophiles (2 new genera, one beetle, very remarkable!). The vegetation is extraordinarily diverse. It is an ideal place for a lab. in every way The ground is dry and rises in hills, one of which (“Gigante”) is about 500 ft. above the level of Gatun Lake. A small bungalow with screened verandah for a lab., a good launch, a resident director (Zetek would be just the man), a cook and one or two competent negroes as assistants, the development of a few good trails across the island, and we should have an ideal zoological and botanical paradise. The king of all the tapirs lives on the island with many of his descendants, together with ocelots 9 ft. long and other beasts too numerous to mention.19
ZETEK’S LETTER TO GOVERNOR MORROW
On March 22, Zetek recommended to Hitchcock that the NRC and the Ecological Society of America write to the governor of the Canal Zone, Jay J. Morrow, requesting that the island be made a preserve for scientific research; he said that he himself would be willing to serve as island custodian.20 He wrote to Hitchcock again the next day, telling him that it was “absolutely foolish to build a concrete building next to the Gorgas Memorial. This would be the best way to kill the project. It would, in fact, become a Mausoleum ”21 Hitchcock thought it was a great idea and informed him that he would take the matter up with the NRC.
Zetek was under the impression that the NRC would have the funds to pay for the project. But the relationship of the NRC to the Institute for Research in Tropical America was solely administrative: It conducted elections for the institute’s executive committee, and it received and dispersed funds of the institute, as directed by the executive committee of the institute.22 That was it. There was no funding obligation. “It is not the policy of the Research Council to finance projects,” Hitchcock explained to Zetek on March 22, 1923. “They open a way for cooperation in various ways and give various sorts of support that are not financial.”23
Zetek quickly became disenchanted with Hitchcock when he informed him that funding was unlikely.24 Still, he wanted to get the island, and hopefully obtain funding from the NRC for it later. He wrote to Governor Morrow himself on March 27, 1923, as representative of the Ecological Society of America. He pointed to the NRC’s interest in establishing a research station in Panama and requested that the island be put aside as a protected area for scientific study.25 Morrow embraced the idea enthusiastically and on April 17, 1923, issued the following brief circular setting the island aside for use as a natural park for scientific study.
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Karl stood still, with eyes that swam. He began to speak, but ended with a shake of his head, as if something had choked him.
"To-morrow, dear friends," he muttered very low. "To-morrow. To-night I am tired, very tired and very happy. Long live George Trafford and his beautiful bride!" he said in stronger tones. "God bless them! God bless our poor country! God help me to rule"—but his voice had sunk again to a whisper and as he spoke he reeled against Saunders.
The latter held the massive but limp frame from falling, while someone produced brandy from a flask and poured a generous measure down the King's throat. Then the soldiers made a seat of their crossed weapons, and shoulder-high and supported by willing arms, Karl of Grimland was borne, half-fainting with exposure and fatigue, but serene of mind, to the winter palace of his beloved Weissheim.
EPILOGUE
Down the great, white highway of the Rylvio Pass a bob-sleigh was speeding in the early hours of a perfect morning. The incense of dawn was in the air, and the magic of stupendous scenery uplifted the souls of the two travellers. Fantastic peaks of incomparable beauty rose up in majesty to meet the amazing turquoise of the heavens. Sparkling cascades of dazzling whiteness hung in streams of frozen foam from dun cliffs and larchcrowned boulders. The roadway down which the sleigh was coursing with unchecked speed wound like a silver ribbon at the edge of precipices, sometimes tunnelling through an arch of brown rock, only to give again, after a moment's gloom, a fresh expanse of argent domes and shimmering declivities. Perched high on perilous crags were ancient castles of grim battlements and enduring masonry, stubborn homes of a stubborn nobility that had levied toll in olden times on all such as passed their inhospitable walls. Below, in the still shadowed valley, were villages of tiny houses, the
toy campanili of Lilliputian churches, and a grey-green river rushing over a stony bed to merge itself in the ampler flood of the Danube.
"Oh, could anything be more perfect?" asked Gloria, who, as on the previous night, was doing duty at the wheel. There was a flush on her cheeks that was a tribute to the keen mountain air, and a sparkle in her dark eyes that told of welling happiness and a splendid conscious joy. Radiant as the morn, fragrant as the pine-laden air, she seemed the embodiment of a hundred vitalities crowded into one blithe being.
"We are on our honeymoon," returned Trafford, "and it would be a cold, dank day that could depress my soaring spirits. As it is, the impossible beauty of our surroundings is so intoxicating my bewildered brain that I am neglecting my duties as brakesman in a most alarming manner. We shall be over a precipice in a minute, if I don't master my exaltation of spirits."
"Perfect love casteth out fear," laughed Gloria.
"Is it perfect?" he asked.
"Absolutely—now, dear," she replied. "From the first you captured my fancy; that was why I did not lie to you in Herr Krantz's wine-shop. Then, when I thought you had killed Karl in the Iron Maiden, my heart grew sick and cold, for I believed you were, as the others, without ruth or mercy. The news that you had saved his life while pretending to take it, put new fire into my soul; but there was ever a war in my breast between true tenderness and the lust of power. I had inherited ambition from a long line of callous ancestors; my whole life had been a tale of scheming, deadening opportunism. And Bernhardt, as we know, with his great domineering personality, was as death to sentiment. And then, last night, well, you took the bit between your teeth and let yourself go. You over-rode my will, you set at nought my interests: you were master, and I handmaid, and my whole soul went out to you in admiration of your strength, and love of the way you used it."
Trafford drank in the words as he drank in the clear, sweet air of the mountain-side, and happiness—the heroic happiness that befel poets and warriors in the days of the world's youth, when men were demi-gods and
gods were demi-mortals—took him with golden wings and exalted him, so that the soaring mountain and the wheeling bird and the forest and the crag and the river were as his brothers and sisters, fellow members of the worshipful company of rejoicing creation.
Onward and downward they flew, while the beams of the rising sun climbed down the valley walls, ledge by ledge and rock by rock, turning brown cliffs to gold, and snowy slopes to diamond and silver. Already they were far below the supreme height of the Weissheim plateau, and the air, dry though it was in reality, seemed almost damp in comparison after the moistureless atmosphere of the lofty tableland they had quitted. The snow held everywhere, but it was the thin covering of an English hill-side in January, not the sumptuous and universal mantle of frost-bound Weissheim. The larch and fir of the uplands were giving gradual place to the stunted oak and the starved chestnut. A thin hedge maintained a scrubby, struggling line at the edge of the roadside, and on the southern slopes the fields were furrowed in endless terraces for the vine.
"We shall reach the frontier in an hour," said Trafford.
"And then?"
"Then we must quit our faithful 'bob.' The road ceases to run downhill at Morgenthal, and a bob-sleigh will not defy the laws of gravity even for the happiest couple in Christendom."
"Then what are our plans?" asked Gloria.
"Plans!" he echoed; "we have no plans. The poor, the unhappy, and the hungry have plans, for they must scheme to improve their condition. But you and I are rich in every gift, and life will be one delicious and unending bob-sleigh ride through gorgeous scenery and vitalising air."
The Princes sighed luxuriously. Then, after a pause—
"We must reach the valley some day," she said.
"Some day, yes," he acquiesced. "Some day the ride will be done, and the road end in the great shadows which no human eye can pierce. But that day will find us hand in hand, with no fear in our hearts, and ready for a longer, stranger, and even more beautiful journey."
As he spoke the valley widened out, and the hills on either side receded at a broad angle. The roofs of Morgenthal were plain to their gaze, and the tinkling of goats' bells broke the silence.
"Austria!" cried Gloria.
"Austria, Vienna, Paris, London," he said, "Southampton, and then the very first boat bound for little old New York. But in our hearts Grimland— always Grimland."
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