HUTCHINSON, G. EVELYN

(b. Cambridge, England, 30 January 1903; d . London, England, 17 May 1991)

population and theoretical ecology, limnology, biogeochemistry.

Hutchinson pioneered modern ecology, limnology, and biogeochemistry in the United States. He was the leading developer of theoretical ecology in the middle years of the twentieth century. In addition, he provided superb graduate training for many of the next generation of leading ecologists. Although Hutchinson was born and educated in England, and did research in South Africa, India, and Italy as well as in the United States, he worked and taught at Yale University for most of his long scientific career.

Origins and Education . Cambridge, England, influenced many of Hutchinson’s views of science and the teaching of science. Hutchinson himself wrote in his partial autobiography, The Kindly Fruits of the Earth (1979), that throughout his early years Cambridge was inhabited by genius, much of which he was exposed to by his family. His father, Arthur Hutchinson, was a mineralogist who taught at Cambridge University, and was later the master of a Cambridge college. His mother, Evaline Shipley Hutchinson, was a feminist and writer.

He attended Gresham's, a public school (in the English sense) in Norfolk, which emphasized science and mathematics. While there, Hutchinson belonged to every science club and conducted his own studies of the taxonomy and behavior of aquatic insects. He published his first paper, about a swimming grasshopper, at age fifteen, launching his seventy-year publishing career.

Hutchinson entered Cambridge University in 1921 to study zoology, but was already well trained in chemistry, physics, and mathematics. This background in physical sciences influenced his future ecological research. He took advantage of the university’s open-ended graduate program, particularly the opportunity to choose among its many famous professors—including F. Gowland Hopkins, Joseph Needham, and J. B. S. Haldane, all at that time in biochemistry. In zoology, G. P. Bidder influenced him the most.

While at Cambridge, Hutchinson became an expert on several groups of aquatic insects, particularly the Corixidae or water boatmen and their sound-producing organs. His observations were published in Edward A. Butler’s A Biology of the British Hemiptera-Heteroptera(London: Witherby, 1923). Henceforth his depth of knowledge on corixids and other groups of water bugs served him well, both in his limnological research and in his more theoretical ecological studies.

Early Postgraduate Research . Although Hutchinson received a “first,” the highest grade, in Parts I and II of the Zoology Tripos (final exams), he chose not to stay on for further graduate work at Cambridge University, but to accept a Rockefeller Higher Education Fellowship to do research at the Stazione Zoologica in Naples. His studies of the endocrine system of the octopus were largely unsuccessful, but Italian culture became an important part of his life; he returned later to do important research on Italian lakes—and even to give limnological lectures in Italian.

Hutchinson began his first professional teaching and research position in 1926, at the University of the Witwatersrand in Johannesburg, South Africa. While there he married his fellow Cambridge zoology student, Grace E. Pickford, who became a well-known fish endocrinologist and Yale professor. Together they carried out, and later published, extensive limnological research on the pans or dry lakes of South Africa. This was not only seminal work, including both physical and biological parameters, but also involved ecological questions. As a result of this work, Hutchinson discovered his new research field: limnology, the study of lakes and other freshwater systems. Lancelot Hogben, then professor of zoology at the University of Cape Town, who sponsored their limnology research, informed Hutchinson of a fellowship open at Yale University. He applied late, by cable. The fellowships were all taken, but he received a newly vacated instructorship instead. This unlikely event proved to be the beginning of his long and extremely productive career at Yale.

Beginnings at Yale . Evelyn Hutchinson and Pickford arrived at Yale in 1928. He taught many undergraduate courses, including embryology. This was the field of Yale’s most eminent scientist, Ross Granville Harrison, who became Hutchinson’s supporter in these early years and to whose encouragement he credited his eventually highly successful research career. Ordinarily a young professor in the United States would try to make his research name in one particular field. Hutchinson never did this. His publications in the first five years after arriving at Yale were in a bewildering variety of fields: the taxonomy of water bugs (Neonectidae and Corixidae); the branchial gland of the octopus; the limnology of South African pans (with Pickford); evidence from cores of climatic change in high altitude lakes in (Indian) Tibet (with Hellmut de Terra); South African Onychophora (a phylum-linking invertebrate group that fascinated him); and the relation between oxygen deficit and productivity of lakes. He also did a major project on arid lakes in Nevada while waiting for his divorce from Pickford in 1933. Hutchinson always credited department head Harrison for encouraging him in his various research endeavors instead of insisting on one area of research. Importantly, Harrison supported him when the question of his tenure at Yale arose. Not all of his colleagues were equally impressed by his early research in so many areas.

Later Hutchinson did the same with his own graduate students’ choice of research projects; there were many paths to follow in Hutchinsonian ecology. His graduate students worked on a great number of organisms, as distinct from Harrison’s Research School, which did so much definitive work on salamander development. In the fifties alone, Willard Hartman’s dissertation was on sponges, Larry Slobodkin’s on Daphnia, Thomas Goreau’s on coral, Peter Klopfer’s on Anatidae (ducks and geese), Alan Kohn worked on Hawaiian cone shells, and Robert MacArthur on birds, the wood warblers of Maine. Some graduate students did research on biogeochemistry or on more theoretical aspects of ecology.

Hutchinson’s graduate students, right from the start, worked on many different problems. They used a variety of techniques, some invented by the graduate students themselves. Although Hutchinson certainly discussed and influenced their research, he rarely put his names on their papers, not even the famous seminal paper in systems ecology by his postdoctoral student Ray Lindeman. He was never the “Herr professor”; most of his graduate students called him Hutch or Evelyn. The graduate students who arrived in the 1930s, Gordon A. Riley, Edward S. Deevey, W. T. Edmondson, and Maxwell J. Dunbar, worked on diverse research problems; all had outstanding scientific careers. Riley and Deevey were both Yale professors before moving to other universities and starting their own research groups. The next generation, those of the 1940s, also included two future Yale professors, John L. Brooks and Willard Hartman, as well as other very well-known ecologists including Fred Smith, Slobodkin, and Klopfer.

Thus, in spite of the great number of Hutchinson graduate students over four decades, Hutchinson’s was a research group rather than a classic Research School like Harrison’s in embryology at Yale. Hutchinson and his research students were connected by a research philosophy, not by particular tools, organisms, one set of problems, or only one aspect of ecology.

The Yale North India Expedition . In 1932 an opportunity arose for Hutchinson to go on the Yale North India Expedition, which traveled to Kashmir and to present-day Ladakh in the Himalayas. German geologist Hellmut de Terra led the expedition; Hutchinson was the head biologist. He collected all kinds of insects, aquatic life, animals, plants, and fossils and recruited specialists from all over the world to identify those specimens not belonging to his own special groups. A Yale undergraduate student, W. T. (Tommy) Edmondson, who already knew rotifers, worked and published with Hutchinson on the North Indian collections of that group. He later became a Hutchinson graduate student and an influential limnologist. Hutchinson also arranged for the publication of all this taxonomic data.

Hutchinson’s own interests on this expedition, however, were largely ecological. For example, how could all the lake biota live at the very high elevations of Ladakh? He studied lakes at more than 5,000 meters. He took physical and chemical measurements, conducted experiments, and made much of the needed equipment himself en route, all of which he described in many letters to Pickford. He wrote to her that he was “pushing on with experimental ecology … because that is where the results will come from that are really exciting now.” His research provided new insights into biogeography and paleolimnology, as well as much new data on high elevation ecology. A crustacean, for example, was often the top predator; many of these lakes did not support fish. Long before he published his famous ecological niche model, Hutchinson was searching for niche dimensions important in the lives of these lake organisms. For example, did ultraviolet light levels correlate with the lake levels at which different planktonic crustaceans were found? He was continually testing such hypotheses.

The Yale North India Expedition proved to be a milestone in Hutchinson’s life in a different respect as well: it was the beginning of his literary career. He wrote a book about his experiences, The Clear Mirror(1936). It got excellent reviews. The back cover of a later edition reads, “In 1932 a young biologist … recorded his observations with the eye of a trained scientist and the style and sensitivity of a novelist.” The book contains strikingly beautiful descriptions of the colors of the landscape, the lakes and their inhabitants.

Limnology and Hutchinson’s First Graduate Students . Not yet thirty, Hutchinson had already conducted research on four continents: Europe, Africa, Asia, and North America. Back in the United States he worked on a lake close to Yale, Linsley Pond, as did several of his first graduate students, Deevey, Edmondson, and Riley. All of them eventually started long-lasting research groups of their own. Edmondson arrived first, but Riley was Hutchinson’s first official graduate student. Riley came to Yale in 1935 to work with Ross Harrison, but found Harrison’s group of graduate students too large and new problems too scarce. By that time Hutchinson was teaching a graduate course in limnology. Riley wrote, “One lecture was enough to make me sit up straight and bright-eyed and to struggle to assimilate every word of his thick British accent. He was dynamic and obviously very bright. … Within a week I knew where I wanted to be. Unsolved problems stuck out all over the place” (1984).

Hutchinson was heavily involved in the biogeochemistry of lakes at that time, studying nitrogen, phosphorus, carbon, and other cycles. Riley’s research problem involved copper cycling in Linsley Pond and other Connecticut lakes. In those early years Hutchinson was in the field with his students, working on the lake and in the laboratory. As Riley described it, the two of them took to the lake as soon as it was ice-free in the same inflatable rubber boat Hutchinson had used in Ladakh. It leaked and they both got cold and wet. The best part, Riley remembered, was the talk: “a rare opportunity to further my education … with a man whom I have always regarded as having the keenest and best informed mind of any scientist I have known” (1984).

Riley’s research later turned from limnology to biological oceanography. He did research in that field as a faculty member at Yale and later headed a large department at Dalhousie University in Halifax, Nova Scotia. Riley made a trenchant comment in his memoir about Hutchinson’s philosophy of doing ecology that permeated all of his own plankton work: “He maintained that populations needed to be studied in terms of dynamic processes—rates of production and consumption and ways these are affected by ecological factors” (1984).

Deevey also participated in this lake research and scientific discussions with Hutchinson. He, however, was initiating a new field of ecology in the United States, paleoecology, including the history of human impacts on lake environments. He was taking cores from Linsley Pond to study the fossil pollen and macrofossils that had been deposited there over a long period. He left the Botany Department and went to zoology to work with Hutchinson, who encouraged Deevey’s attempt to integrate botany, zoology, paleoanthropology, and paleolimnology. Such a comprehensive study had never been done before. When Carbon 14 dating became available just after World War II, this provided an independent method for determining the chronology of such lake cores. Paleoecology became Deevey’s major research field. He and Hutchinson worked together on a core Hutchinson had collected on the North India expedition. Hutchinson and his later research associates, particularly Ursula M. Cowgill, continued to work in this field in other parts of the world as well.

Edmondson was the third of the early graduate students. He had a desk in Hutchinson’s laboratory as an undergraduate and became an expert on rotifers, microscopic but complex aquatic invertebrates with interesting behavior and life cycles. As a graduate student Edmondson studied growth rates and population biology of lake rotifers, making several important discoveries. He was the first Hutchinson student to bring limnology to a West Coast university. Edmondson and a large group of graduate students at the University of Washington were responsible for the ecological studies of Lake Washington that made possible the reversal of pollution in that large Seattle lake.

Dunbar also came to work with Hutchinson in the 1930s, on a fellowship from Oxford University. Later he completed a PhD at McGill University in Montreal, working in the Canadian Arctic but keeping in touch with Hutchinson about his plankton studies. He had a distinguished career in arctic research. Dunbar related, in an interview with this author, that once when Hutchinson was visiting Oxford, he and Charles Elton were seen “plotting together on the Broad… two people responsible for most of ecology.” Elton’s 1927 book on animal ecology had greatly influenced Hutchinson, but this was the first time they had actually met.

Hutchinson wrote many more papers concerning lake biota. He is known as the source of modern ecological ideas in American limnology and as the synthesizer of his own work and that of many others in this field. His four-volume Treatise on Limnology is the bible of this field in English. The first volume, published in 1957, covered the geography, physics, and chemistry of lakes; the following volumes included plankton and other lake biota, including plants (Volume III). The fourth volume was published posthumously, but Hutchinson had finished the text. The Treatise is also a superb guide to the worldwide literature in limnology, which has become part of the broader field of ecology. Hutchinson never really left limnology. Many of Hutchinson’s later theoretical papers concern the population and community ecology of lake biota. The American Society of Limnology and Oceanography gives out the G. Evelyn Hutchinson Award every year.

Research in Biogeochemistry . The field of biogeochemistry is one that Hutchinson was instrumental in bringing to the United States. He was very much impressed by the scientific work of the Russian scientist V. I. Vernadsky. Hutchinson had his work translated to make his ideas about the biosphere and about the chemistry of organisms available in English. Hutchinson and his graduate students in the 1940s and 1950s did a great deal of work in this field. Probably the best known of them is Howard T. Odum, who studied the biogeochemistry of strontium. He went on to become an important innovator in the new field of systems ecology. Hutchinson himself wrote papers about the biogeochemistry of phosphorus, nitrogen, carbon, aluminum, and thiamine. He was probably the first to discuss carbon dioxide in the atmosphere; he wrote a major paper on the biochemistry of the terrestrial atmosphere. His study of aluminum in many species of Lycopodium(club mosses) was his only major research on higher plants.

Two well-known Hutchinson papers were published in 1948: “On Living in the Biosphere,” a general approach to the significance of nutrient cycles on this planet, and “Circular Causal Systems in Ecology.” The latter was closely related to cybernetics, a major subject of the Macy Conferences in New York in the 1940s. Hutchinson was a participant in these conferences, together with Margaret Mead, Gregory Bateson, Norbert Wiener and other notable social and biological scientists from a variety of fields.

Hutchinson was in charge of the Survey of Contemporary Knowledge of Biogeochemistry. As part of that survey he published a 550-page article titled “The Biogeochemistry of Vertebrate Excretion” (1950), largely about guano and guano islands and also bat caves in many parts of the world. It is quite a Darwinian book in terms of data and letters collected from sea captains and guano workers and from obscure published sources. He reworked the data in many cases and added his own ideas. It is a fascinating book, quite different from his other works.

By 1948 Hutchinson was well known in both limnology and ecology, but probably best known for his research in biogeochemistry. At this time he was offered a high position by the U.S. Geological Survey to do research on biogeochemistry or whatever he wished. He was tempted by this offer, but it did not include graduate students. He turned it down because he felt that his graduate teaching had succeeded in turning out outstanding young ecologists who were rapidly becoming leaders in their fields. He wanted to stay at Yale to continue his teaching; seven of his graduate students had already completed PhDs by that time. Thirteen more, in addition to postdoctoral students, arrived in the 1950s, and an additional eighteen completed their PhDs in the 1960s and early 1970s. The later years included women graduate students, several of whom, including Karen Glaus Porter, Maxine Watson, Donna Haraway, and Alison Jolly, went on to important academic and research careers. Hutchinson turned down at least six other academic positions as well, and remained at Yale until he was eighty-eight.

The Beginnings of Systems and Radiation Ecology . In the 1940s, work by Hutchinson and his students was responsible for the initiation of two new fields in ecology, systems ecology and radiation ecology. In 1942 his postdoctoral student Raymond Lindeman had written “The Trophic Dynamic Aspect of Ecology,” the acknowledged seminal paper in the field that became ecosystem or systems ecology. Hutchinson contributed ideas and quantitative methods to Lindeman’s paper. He was also instrumental in getting it published in Ecology after it was originally turned down as too theoretical by two reviewers, both established leaders in the field of limnology at that time. The paper was published, with an addendum by Hutchinson, after Lindeman’s death in early 1942. Systems ecology became a major area of research, particularly in the 1950s and 1960s. It included the International Biological Program, which studied productivity and related ecological topics in many different ecosystems.

The first experiments on the use of radioactive isotopes in ecological research were published in 1947 by Hutchinson and his graduate student, Vaughan T. Bowen: “A Direct Demonstration of the Phosphorous Cycle in a Small Lake.”

Radiation ecology also became a major research area involving diverse ecosystems, many workers, and Atomic Energy Commission funding. His former graduate student Odum did important large research projects in this field, but Hutchinson himself, who preferred smaller projects and other types of funding, moved on.

Experimental and Mathematical Ecology; Population Growth . What about experimental and theoretical ecology, the field in which Hutchinson was most influential? Recall his comment to Pickford about the importance of ecological experiments during the 1932 Yale North India Expedition. Hutchinson was already working in this field, however, even before that expedition. He published a 1932 paper titled “Experimental Studies in Ecology: I. The Magnesium Tolerance of Daphnidae and its Ecological Significance.” Most of Hutchinson’s early papers were published in European hydrobiology journals. There was little work of this kind being carried out in the United States at the time.

Hutchinson had been reasonably well trained in mathematics and was long an advocate for its use in ecology. He made this clear in a 1940 review he had written of Bio-ecology, a book in which the plant ecologist Frederic Clements and the animal ecologist Victor Shelford had tried to synthesize the field. Hutchinson was critical of this attempt, particularly because mathematical methods of analysis of ecological processes were completely omitted. Clements and Shelford expressed grave doubts that

mathematical and statistical methods could be applied to the complex processes studied in ecology. Mathematics became important in Hutchinson’s own research group, in terms of population growth. Lawrence B. (Larry) Slobodkin (PhD 1951) did innovative experiments and studied population dynamics of the water flea, the microscopic cladoceran crustacean Daphnia. F. E. (Fred) Smith (PhD 1950), who studied benthos populations, also did experiments on Daphnia and later worked out equations for its population growth in relation to population density.

Hutchinson and his students had predecessors, particularly George F. Gause and Raymond Pearl, on the study of population growth (and decline). Gause had produced a population (logistic) growth curve for Paramecium and Pearl had studied population growth in Drosophila, both in the 1930s. Pearl also worked on human population growth curves. Hutchinson, always interested in the history of his sciences, thoroughly reviewed the earlier work in An Introduction to PopulationEcology (1978.) He wrote here about the logistic curve of population growth:

The important thing about any simple and easily understood formulation such as the logistic is that it can provide a stepping stone to a number of other, less simple, less general, but more accurate theories. This is in fact one of the significant uses of general theories in science. (p. 31)

By the 1950s mathematics had become an essential part of experimental ecology in Hutchinson’s group. Hutchinson himself introduced the concept of the time lag, a complication to the logistic equation. For example, the process of reproduction may not be instantaneous, as in Daphnia when the parthenogenic female liberates newly hatched babies from her brood pouch. Hutchinson credited the original equation including a time lag in population growth to economists, but he brought it into ecology in his 1948 Macy Conference paper, “Circular Causal Systems in Ecology.”

Competition was becoming a central concern of ecologists. Garrett Hardin published his competitive exclusion principle in 1960, but it too had its predecessors. Many experiments had been done in the laboratory using two closely related species of flour beetle, Paramecium, and other organisms, showing that, depending on the set-up of the experiment, one species would outcompete the other. The type of food or some sort of refuge might be a deciding factor in such experiments. In Hardin’s dictum, complete competitors cannot coexist (in laboratory experiments or in nature).

The mathematical biologist to whom competition equations are usually attributed is Vito Volterra; his work goes back to 1926. Gause carried out laboratory experiments on competing organisms using Volterra’s work as a mathematical basis. Alfred J. Lotka was also involved in the early 1920s in the development of competition equations. They are often referred to as the Lotka-Volterra equations, although the two worked independently.

American ecologists paid little attention to this early work, perhaps because the more complete account of Gause’s experiments was published in French. It was only the 1940s and thereafter that competition in mathematical terms was again taken up, in large part due to Hutchinson’s amazing knowledge of the literature of ecology and limnology. He himself read French, German, and Italian, and found others to translate the important Russian work.

Much of this early work was done in the laboratory, using protozoans—particularly related Paramecium species—and species of flour beetles, Tribolium. All of these reproduce rapidly in the laboratory, and the environmental conditions of these small organisms were easy to regulate. The outcome of competition varied with these external conditions, and could be measured mathematically. Hutchinson was well aware of these studies; he, however, was most interested in competition and competitive exclusion—or coexistence—not in the laboratory but in the field.

Population Ecology in Nature . Slobodkin brought Robert MacArthur, whose previous training was in mathematics, into Hutchinson’s group of graduate students. When MacArthur and, later, another graduate student, Egbert Leigh, came to work with Hutchinson, he had first-rate mathematicians in his research group. Hutchinson and MacArthur published two joint papers about competition in birds and other animals in 1959.

MacArthur’s dissertation research (PhD 1958), known to every first-year ecology student, concerned the population ecology of the brightly colored warbler species that breed in the coniferous forests of northern New England. MacArthur took many hours of data on the feeding habits of five closely related warblers of the genus Dendroica in the spruce trees of Maine. Thousands of birdwatchers follow these species’ migrations every spring, but no one had conducted a quantitative study of their feeding or breeding habits.

The question MacArthur asked was a very Hutchinsonian one: how can so many closely related species coexist without competitive exclusion? MacArthur’s study answered this question. The warblers search for their food in different zones of the spruce trees; MacArthur was able to quantify the amount of time each species spent in each zone, as well as the overlap between different species. He also recorded the number of breeding pairs per 5 acres, and was able to write competition equations for four of these warbler species. The environment in which these warblers were feeding was sufficiently coarse-grained to allow for their continuing coexistence.

Many ecologists have since worked on similar problems in nature, working on a variety of organisms, aquatic and terrestrial, animals and plants. Among Hutchinson students, Alan J. Kohn worked on the many species of the mollusk genus Conus in Hawaii, and in the 1970s, Maxine Watson worked on competition and niche diversification among the members of the moss family Polytrichaceae, the haircap mosses.

The Multidimensional Niche . In 1957 Hutchinson published his most influential paper, in which he introduced his own niche concept. Its title, “Concluding Remarks,” came from his role in summing up a Cold Spring Harbor Symposium. Hutchinson later wrote pages of footnotes on the history of niche concepts in his population ecology book, but the quantitative aspects of niche study in ecology are not very old. Hutchinson presented a timely new niche concept that enabled many ecologists to evaluate niche relationships quantitatively.

Hutchinson called it the “n-dimensional niche” but pointed out that it would probably not be possible to quantify all the n dimensions of any particular species’ niche, in other words all the tolerances and needs of a particular species. Two or three niche factors, whether biotic or abiotic, may be enough to separate the niches of two interacting species. He gave a hypothetical example of temperature, food size, and branch density delimiting a three-dimensional niche of one species of squirrel; such a niche can be diagramed in terms of measured variables. Hutchinson made a very useful distinction between a species’ fundamental niche in the absence of competition, and its realized (postinteractive) niche in the presence of a competitor, in this case a second species of squirrel. Competitive exclusion could occur where the fundamental niches of two species overlapped, or the overlap area could be divided, producing separate realized niches for each species. These differences can be measured and expressed in terms of competition coefficients; species that have contiguous niches will show high values of these coefficients; for those whose n-dimensional niches hardly overlap, the value will approach 0. Niche breadth and niche overlap can also be mathematically determined.

Rachel Carson brought the term ecology to the attention of the American public. But it was Hutchinson’s view of ecological niches that could be quantified, and his concept of niche diversification in a great variety of organisms and ecosystems, that spawned so much PhD research, so many ecological papers, new mathematical and statistical methods, and eventually experimental manipulations in organisms from birds to wood rats to protozoa to pond weeds and mosses. MacArthur, Richard Levins, and many other ecologists have written theoretical papers and books using Hutchinson’s and later related niche concepts.

As is the case with much theoretical ecology, Hutchinson’s niche concept and particularly the importance of competition as a major factor in structuring ecological communities, have been subject to criticism and rival theories, but ecological research using these concepts continues.

Biodiversity and Biogeography . For Hutchinson, a related major puzzle was the whole question of animal diversity. “Homage to Santa Rosalia” (1959), another of his very influential papers, explored this question and spurred the interest in and funding for biodiversity studies. Both the understanding and the preservation of biodiversity of organisms on this planet were a longtime concern of Hutchinson, and also, notably, of Edward O. Wilson. Both have paid attention to the remarkable, and still largely unknown, diversity of insects, particularly in the tropics. As Hutchinson concluded, the great diversification of animals, including insects, is highly dependent on the great variety of plants and their interactions with the animal world.

Hutchinson considered MacArthur his most brilliant student and was devastated at his early death at age forty-two. He was MacArthur’s mentor while he was a student as well as his codeveloper of ecological theory. MacArthur went beyond Hutchinson in the development of mathematical theory, in both population ecology and in biogeography, a field that Hutchinson had contributed to since his North India studies in 1932. Wilson and Hutchinson in their Biographical Memoir about MacArthur for the National Academy of Sciences wrote that as a “mathematician naturalist,” he had eventually formed many of the “parameters of ecology, biogeography and genetics into a common framework of fundamental theory.” The ideas and research of the 1950s and 1960s changed ecology from a descriptive science to a “structured predictive science that combined powerful quantitative theories with the recognition of widespread patterns in nature.”

Hutchinson and his earlier students, including Lindeman, Slobodkin, and Odum, initiated this change in large part. According to Martin Cody and Jared Diamond, the “revolution in ecology” was carried further by MacArthur’s work in the 1960s. Hutchinson’s 1975 paper in the book by these two authors is titled “Variations on a Theme by Robert MacArthur,” whom he quoted as writing

Scientists are perennially aware that it is best not to trust theory until it is confirmed by evidence. It is equally true, as Arthur S. Eddington [and also Hutchinson] pointed out, that it is best not to put much stock in facts until they have been confirmed by theory. (p. 492)

MacArthur had written that both the ecological theories and the facts had serious inadequacies, which provided stumbling blocks to progress in ecology. Hutchinson illustrated these problems in his paper. In the area of competitive exclusion, he concluded, facts and theories had worked out well, but in the study of population cycles, that had not yet happened. The theoretical ecolo-gist might consider “all possible models,” but it was the field or experimental ecologist’s responsibility to find out which of these models were realized in nature. In order for this to happen in the future, Hutchinson wrote that ecologists must be trained to have “a deep understanding of organisms” (1975, p. 515). Hutchinson’s classic remark about his mathematical theorist student follows: “MacArthur really knew his warblers.” At the beginning of the twenty-first century, experimental field ecologists are busy testing ecological theory with real data.

Writings and Prizes . In 1943 Hutchinson began a series of “Marginalia,” essays and reviews on a great many scientific subjects, constituting a special column in the American Scientist from that date until 1957. It was revived in the 1970s and thereafter. Many of these columns were reprinted elsewhere. Several books, The Itinerant Ivory Tower (1953), The Enchanted Voyage, and Other Studies(1962), and The Ecological Theater and the Evolutionary Play(1965), consisted of Marginalia as well as some new essays. Through the original articles in a journal intended for scientists in many fields, and through his books, Hutchinson became well known as an excellent science writer, and to some extent even as a literary figure. He was very much interested in literature, as was his second wife, Margaret Seal Hutchinson. The writer Rebecca West was a close friend of Hutchinson's. They first met as a result of her reading one of his Marginalia articles. They exchanged hundreds of letters (now at the Beinecke Library at Yale and at the University of Tulsa). When West died in 1983, Hutchinson became her literary executer. Margaret Mead was also a good friend; he helped to edit one of her later books.

Hutchinson received all the honors and prizes available to him, throughout the many decades of his career; there is no Nobel Prize in Ecology. These included many honorary doctorates, including one from Cambridge University in his old age. He could almost certainly have sent his published papers to Cambridge and received an earned doctorate. Early in his career he was afraid of being turned down, as one of his fellow students had been; later, already famous in at least two fields of ecology, he was rather proud not to have one. He enjoyed the pomp of his 1981 Cambridge honorary degree, however, parading in the company of molecular biologist Max Perutz and other notable honorees.

Hutchinson received the Benjamin Franklin Medal in 1979 for “developing the scientific basis of ecology.” He was both proud and humbled to be in the company of previous winners who included Thomas Edison, Max Planck, and Albert Einstein. He was a member of the National Academy of Sciences (NAS) and a foreign member of the Royal Society. He put both memberships to good use when, as a NAS spokesman, he flew to London to talk to the Royal Society in the successful battle to preserve the remarkable biodiversity of the island of Aldabra.

Hutchinson had a serious though usually quiet role as an environmentalist, primarily through his research, throughout his career. He wrote in the American Scientist in 1943 as reprinted in his 1953 book:

The writer believes that the most practical lasting benefit science can now offer is to teach man how to avoid destruction of his own environment, and how … to find ways to avoid injuries that at present he inflicts on himself with such devastating energy. (p. 270)

His contribution to public awareness of these issues resulted in the NAS Cottrell Award for Environment. When he won the Daniel Giraud Elliot Medal in 1984, the citation read: “Hutchinson was admired as limnolo-gist, biochemist, ecologist, evolutionist, art historian and ranks among our zoological giants.”

Hutchinson is perhaps the only scientist to have been awarded the U.S. National Medal of Science twice. Hutchinson turned it down for political reasons when it was offered during the Nixon administration. It was offered again in 1991 after Hutchinson’s death. His nephew, Francis Hutchinson, came to Washington, D.C., from London to accept the award. Hutchinson’s final award during his lifetime was the 1986 Kyoto Prize in Basic Science, sometimes called the Japanese Nobel Prize. He was over eighty at that time. His third wife, Anne Twitty Goldsby Hutchinson, whom he married after Margaret’s death, accompanied him to Japan.

Renowned as the major inventor of modern ecology in the United States, and for his research in and synthesis of limnology, Hutchinson himself always cited his role in the education of graduate students as equally important. He officially retired from Yale in 1971 but still had graduate students thereafter, and he continued his work there until 1990. The 1971 intellectual family tree of his students and their students to three and even four generations created by Yvette Edmondson includes a great many of the best-known American ecologists. By the time of Hutchinson’s death there were six generations.

Hutchinson moved back to England after his wife Anne’s death in 1990 and died in London in 1991. Yale held a 100th birthday celebration of his life and work in 2003.

BIBLIOGRAPHY

WORKS BY HUTCHINSON

With Grace E. Pickford and J. F. M. Schuurman. “The Inland Waters of South Africa.” Nature 123 (1929): 832–834.

Letters from the Yale North India Expedition (1932). G. E. Hutchinson archives, Yale University Library. Written to Grace E. Pickford.

“Experimental Studies in Ecology: I. The Magnesium Tolerance of Daphnidae and its Ecological Significance.” Internationale Revue der gesamten Hydrobiologie und Hydrographie 28, no. 11–12 (1932): 90–108.

The Clear Mirror: A Pattern of Life in Goa and in Indian Tibe t. Cambridge, U.K.: Cambridge University Press, 1936.

Review of F. E. Clements and V. E. Shelford’s Bio-ecology. Ecology 21, no. 2 (1940): 267–268.

“Limnological Studies in Connecticut: IV. Mechanisms of Intermediary Metabolism in Stratified Lakes.” Ecological Monographs 11, no. 1 (1941): 21–60.

“The Biogeochemistry of Aluminum and Certain Related Elements.” Quarterly Review of Biology 18 (1943): 1–29.

With Vaughan T. Bowen. “A Direct Demonstration of the Phosphorous Cycle in a Small Lake.” Proceedings of the National Academy of Sciences of the United States of America33, no. 5 (1947): 148–153.

“Circular Causal Systems in Ecology.” Annals of the New York Academy of Science 50 (1948): 221–246.

“On Living in the Biosphere.” The Scientific Monthly 67, no. 6 (1948): 393–397.

“Survey of Existing Knowledge of Biogeochemistry: 3. The Biogeochemistry of Vertebrate Excretion.” Bulletin of the American Museum Natural History 96 (1950): 1–554.

“Copepodology for the Ornithologist.” Ecology 32 (1951): 571–577.

The Itinerant Ivory Tower: Scientific and Literary Essays. New Haven, CT: Yale University Press, 1953. Reprints from “Marginalia,” American Scientist, and one unpublished article.

“Concluding Remarks.” Cold Spring Harbor Symposium Quantitative Biolog y 22 (1957): 415–427.

A Treatise on Limnology. Vol. 1, Geography, Physics, and Chemistry. New York: Wiley, 1957.

“Homage to Santa Rosalia, or Why Are There So Many Kinds of Animals?” American Naturalist 93, no. 870 (1959): 145–159.

With Robert H. MacArthur. “A Theoretical Ecological Model of Size Distributions among Species of Animals.” American Naturalist 93, no. 869 (1959): 117–125.

“The Paradox of the Plankton.” American Naturalist 95 (1961): 137–147.

The Enchanted Voyage, and Other Studies. New Haven, CT: Yale University Press, 1962. Reprinted articles and three unpublished essays.

With Ursula M. Cowgill. “Chemical Examination of a Core from Lake Zeribar, Iran.” Scienc e 140, no. 5 (1963): 67–69. Hutchinson and Cowgill published on lake cores from all over the world. This is an early example.

“The Lacustrine Microcosm Reconsidered.” American Scientist 52 (1964): 334–341.

The Ecological Theater and the Evolutionary Play. New Haven, CT: Yale University Press, 1965.

“To Save [Lake] Baikal.” New York Times, 23 June 1966. One of many such letters to the editor about environmental issues.

A Treatise on Limnology. Vol. 2, Introduction to Lake Biology and the Limnoplankton. New York: Wiley, 1967.

“When Are Species Necessary?” In Population Biology and Evolution, edited by Richard Lewontin. Syracuse, NY: Syracuse University Press, 1968.

“The Biosphere.” Scientific American 223, no. 3 (1970): 45–53. A good introduction for students and general readers.

“G. Evelyn Hutchinson Celebratory Issue.” Edited by Yvette H. Edmondson. Limnology and Oceanography 16, no. 2 (1971). Includes a complete Hutchinson bibliography from 1918 to 1971.

A Treatise on Limnology. Vol. 3, Limnological Botany. New York: Wiley, 1973.

“Variations on a Theme by Robert MacArthur.” In Ecology and Evolution of Communities, edited by Martin T. Cody and Jared M. Diamond. Cambridge, MA: Belknap Press, Harvard University, 1975.

An Introduction to Population Ecology. New Haven, CT: Yale University Press, 1978. A very useful book for both biologists and historians.

The Kindly Fruits of the Earth: Recollections of an Embryo Ecologist. New Haven, CT: Yale University Press, 1979. A partial autobiography through the early days at Yale.

“Marginalia: What Is Science For?” American Scientist 71, no. 6 (1983): 639–644.

“Keep Walking.” Physiological Ecology Japan 24 (Special number, 1987): s81–s87. Lecture of the 1986 Kyoto Prize in Basic Science.

With Edward O. Wilson. “Robert Helmer MacArthur.” Biographical Memoirs 58 (1989): 318–327.

A Treatise on Limnology. Vol. 4, The Zoobenthos, edited by Yvette H. Edmondson. New York: Wiley, 1993.

OTHER SOURCES

Hagen, Joel B. An Entangled Bank: The Origins of Ecosystem Ecology. New Brunswick, NJ: Rutgers University Press, 1992.

Kingsland, Sharon. Modeling Nature: Episodes in the History of Population Ecology. Chicago: Chicago University Press, 1985.

Levin, Simon A. Fragile Dominion: Complexity and the Commons. Reading, MA: Perseus Books, 1999.

MacIntosh, Robert P. The Background of Ecology: Concept and Theory. Cambridge, U.K.: Cambridge University Press, 1985. Hutchinson and his work appear in nearly every chapter of this book.

Mills, Eric L. Biological Oceanography: An Early History, 1870–1960. Ithaca, NY: Cornell University Press, 1989.

Riley, Gordon A. Reminiscences of an Oceanographer. 1984, 166 pages. Unpublished; in possession of author.

Slack, Nancy G. “Botanical and Ecological Couples: A Continuum of Relationships.” In Creative Couples in the Sciences, edited by Helen M. Pycior, Nancy G. Slack, and Pnina G. Abir-Am. New Brunswick, NJ: Rutgers University Press, 1995.

———. G. Evelyn Hutchinson and the Invention of Modern Ecology. New Haven, CT: Yale University Press, forthcoming.

———. “G. Evelyn Hutchinson: From Cambridge, England School Boy to America’s Foremost Ecologist.” In The Beauty of the World: The Writings of G. Evelyn Hutchinson, edited by David Skelly. New Haven, CT: Yale University Press, forthcoming.

Slobodkin, Lawrence B., and Nancy G. Slack. “George Evelyn Hutchinson: 20th-Century Ecologist.” Endeavour 23, no. 1 (1999): 24–30.

Nancy G. Slack