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Elton, Charles Sutherland

ELTON, CHARLES SUTHERLAND

(b. Manchester, England, 29 March 1900; d. Oxford, England, 1 May 1991)

animal ecology, conservation.

Often considered the father of animal ecology, Elton published the field’s first general synthesis, formulating principles that would shape subsequent concepts and methods. He founded a small research center, the Bureau of Animal Population, at Oxford University in 1932, and directed it until its dissolution upon his retirement in 1967. The bureau became an internationally significant center for research and scholarly exchange, influencing two generations of ecologists. Elton was the founding editor of the Journal of Animal Ecology, a position he held for two decades. He also contributed to Great Britain’s nature conservation policies and agencies, and to defining the relevance of ecological science to economic and environmental concerns.

Early Life and Education . Charles Elton was the youngest of three sons of middle class, scholarly, and socially well-connected parents. Their father Oliver was an Oxford-educated professor of English at Liverpool University: a literary scholar, translator, and critic. Their mother Letitia Maynard (née MacColl) was a writer.

Geoffrey, the eldest son, attended Cambridge, served in the army during World War I, worked in fisheries research and finally as an English teacher. As children, Geoffrey and Charles rambled the countryside, studying nature near Liverpool and during holidays in the Malvern Hills. Years later Charles recalled that without Geoffrey’s inspiration, he might never have become an ecologist. When Geoffrey died unexpectedly in 1927, Elton dedicated his first book to him, and also vowed to create a research institution in his memory.

Elton attended Liverpool College, and trained with the Army Cadet School (Royal Engineers), but escaped battlefield service. He entered New College, Oxford in 1919, graduating in 1922 with first-class honors in zoology. These Oxford years proved to be formative: He was mentored by the anatomist Edwin S. Goodrich and tutored by the zoologist Julian Huxley. Alexander Carr-Saunders was also influential, with Elton eventually applying his sociological ideas to animal populations. Studies near Liverpool, in the Lake District, and at Oxford—of ponds, an estuarine stream, and sand dunes—confirmed his interest in ecology. In Victor Shelford’s Animal Communities in Temperate America (1913), he found models for the study of ecological communities.

Elton’s relationship with Huxley was fruitful: Through him he gained his first experience with Arctic ecology, and he wrote his first book, Animal Ecology(1927), at Huxley’s behest. Huxley’s efforts to expand Oxford zoology beyond anatomy and embryology also created opportunities for Elton (and others) to pursue their interests in ecology. But the relationship was sometimes tense: Peter Crowcroft recorded Elton’s terse comment, “I don't really mind when Julian steals my ideas. But I strongly object when he takes out a notebook and writes them down in front of me” (1991, p. 2).

After graduating in 1922, Elton remained at Oxford; he was at first employed part-time as a departmental demonstrator, then was promoted in 1929 to permanent status as university demonstrator. He retained this position until 1936, when he became university reader in animal ecology—a title he retained until retirement in September 1967.

In the early 1920s polar exploration had captured the British imagination, inspiring Oxford scientific expeditions to Spitsbergen (now Svalbard). These expeditions, part of an Oxford tradition of scientific study in remote locations, were seen as providing not just training in field sciences, but opportunities to build character and a spirit of teamwork. They would also prove essential to Elton’s formation as an ecologist. He accompanied expeditions in 1921, 1923, and 1924, undertaking ecological surveys of arctic islands. In the latter two expeditions Elton served as ecologist and chief scientist. The relatively small number of species in northern ecological communities—a simplicity that presented the possibility of their full description— was of special interest to him. This complemented another environmental perspective that Elton gained through his northern experience. George Binney, the expedition leader, introduced aircraft and aerial photography as research tools, and used these to develop overviews of the northern landscape. This “view from above” would long influence Elton’s perspective on nature: When describing ecological communities his perspective would be that of an elevated observer, able to perceive all the relations between species and their environment, and, in principle at least, to construct a comprehensive account of an ecological community. Although his Arctic research ended with a 1930 expedition to Lapland, these experiences would shape his perspective on ecology throughout his career.

Elton went on to play a large role in organizing expeditions, serving from 1927 to 1932 as the first chairman of the Oxford Exploration Club, and two more years as treasurer. The club organized thirteen expeditions before World War II, providing formative experiences for many later prominent ecologists and conservationists, including Max Nicholson, Nicholas Polunin, and Colin Trapnell.

Populations and Communities . While still in his early twenties, Elton drew on these experiences to construct a series of novel ecological perspectives. In 1923 he and botanist Victor Summerhayes published a paper, based on the 1921 expedition, in which they described the habitats and plant and animal communities found on Spitsbergen and Bear Island. Concluding that animal communities could be best described in terms of the food relations between species, they seized the opportunity presented by the relatively small number of species on Bear Island to present in diagram form a complete overview of their ecological interactions. In effect, they synthesized their knowledge of the natural history of the island by representing it in terms of the feeding relationships linking all animal species.

The following year Elton examined another phenomenon commonly associated with northern animal species. On the voyage home from the 1923 expedition, he had bought a book by the Norwegian biologist Robert Collett, Norges Pattedyr. He translated it, and studied Collett’s discussion of northern animal populations, including his credulous accounts of the population explosions, migrations, and mass drownings of lemmings (stories, as eventually became evident, that were more folklore than fact). These convinced Elton that dramatic fluctuations were a characteristic feature of northern animal populations. Their existence, he thought, must overturn the widespread but erroneous assumption that populations maintain a state of balance. Understanding why these fluctuations occur could also illuminate the mechanisms by which populations are regulated. In a 1924 paper, “Periodic Fluctuations in the Numbers of Animals: Their Causes and Effects,” Elton surveyed an extensive literature, including Collett, as well as C. Gordon Hewitt’s The Conservation of the Wildlife of Canada (1921), which discussed fluctuations in Canadian populations of fur-bearing animals. Believing that the cause of these fluctuations lay in external factors, and specifically variations in climate, he explored the possibility of correlating them with an eleven-year cycle of sunspot activity. These fluctuations, he also argued, both had economic implications and were relevant to understanding the action of natural section.

In subsequent years Elton pursued a series of research projects, applying diverse methods but with a common focus on the regulation of animal populations. The first project began in 1925, when the Hudson’s Bay Company hired him as a consultant. The position owed itself to George Binney’s intervention; more generally, it reflected the company’s interest in predicting variations in fur-bearing animal populations. For five years Elton immersed himself in the company’s London archives. Its fur trade records extending back to 1736 would, he hoped, throw light on fluctuations in fur-bearing animals in Canada. Supplementary data were provided by the fur records of the Moravian Missions in Canada, extending from 1834 to 1925. In a second project, also beginning in 1925, he conducted field studies of mice and vole populations in Bagley Wood near Oxford, tracking their reproduction and mortality. In this project Elton collaborated with other biologists, including John R. Baker and ecological geneticist Edmund B. Ford; they styled themselves the “Mouse Gesellschaft.” This project ended in 1928, the same year Elton began a study of vole populations that drew not just on his own fieldwork, but on reports from a wide range of observers, collected through a self-styled “intelligence system.” This project would continue until 1939.

Animal Ecology . While pursuing these studies on animal populations, Elton also wrote his first book, Animal Ecology, published in 1927. He had already demonstrated by his papers on animal communities and populations his interest in synthetic perspectives, but the immediate impetus for this book came from Huxley, who invited him to contribute to his series on biology. Their shared ambition, shaped in the context of a research community still focused on laboratory study of animals, was to demonstrate that ecology could also be studied scientifically. Animal ecologists, though, needed their own distinctive concepts, so as to be able to cease borrowing them from plant ecology (as they had with the concept of succession, for example).

Elton drew together decades of accumulated observations, and explained how these could be understood in terms of a small number of principles governing the structure and function of animal populations and communities. These included: the arrangement of species in food chains and cycles (or webs); the niche, as a description of an animal’s place in its community, especially its relation to food and predators; and the principles of food size and pyramid of numbers, meaning that large numbers of small prey tend to be at one end of a food chain, and small numbers of large predators at the other. Such ideas were not entirely new—food chains of course were familiar, and Charles Darwin and other naturalists had discussed the niche concept. Elton, however, placed these ideas in new contexts; he explained, for example, the relation of an animal’s size to its position on the food chain, or the value of viewing its niche in terms of functional (feeding) relationships.

Animal Ecology was noteworthy for its clear style and simplicity of language. Its significance also lay in its demonstration that ecological communities, in all their diversity and complexity, could be understood in terms of basic principles, that were based, in turn, on a classification of species as producers or consumers—an economic view of the natural order that was widely persuasive. This view of nature as an economy epitomized the ties of analogy and inspiration linking Elton’s views of animal communities and human society. Humans, indeed, were at the heart of Elton’s ecology: as he explained, his book was “chiefly concerned with … the sociology and economics of animals” (p. vii). He often drew the parallels implied by this comment, perhaps most notably when he explained the niche concept in terms of an analogy between the “occupation” of a badger and that of a vicar (p. 64). In drawing these ties Elton exhibited the influence of Carr-Saunders, whose work Elton had studied closely. In The Population Problem, published in 1922, Carr-Saunders had applied a neo-Malthusian perspective to human populations, examining how societies differentiated in terms of traditions or level of development proved able or unable to restrict their increase in population, and so to live within the limits of their resources. To some extent, Elton read Carr-Saunders’s sociology into nature; the sociologist’s explanations of social and economic cycles, classes and conflicts influenced Elton’s discussion of animal populations and relations between species.

Elton was somewhat unusual among ecologists in considering evolution—most ecologists at this time did not, and even he decided that the relevant chapter in Animal Ecology required some explanation. His interest undoubtedly reflected Huxley’s influence, as well as his youthful reading of Darwin and Alfred Russell Wallace. Yet, like many British and American naturalists, he found unpersuasive strict Darwinian accounts that relied solely on natural selection. Some features, such as color, or differences between closely related species, could not, he believed, be explained in terms of selective advantage. Natural selection, he concluded, varies in its intensity, is only one of a number of agents of evolution, and sometimes fades into irrelevance. Population ecology, in fact, could illuminate this varying role of selection: the abundance of a species—whether it was crowded or scarce— would shape the selective pressures imposed by its environment, or, indeed, determine whether an animal might migrate elsewhere, thereby effectively upending Darwinian assumptions by selecting its environment, rather than the reverse.

Animal Ecology epitomized the approach Elton would follow in his research over the next few decades. The book opened with the disarming claim that “Ecology is a new name for a very old subject. It simply means scientific natural history” (p. 1). This captured Elton’s intent: Throughout the book he discussed not just the concepts, but the practice of ecology, emphasizing that it was preeminently a field science, requiring careful observation of animals in their habitats. Animal Ecology also established a research agenda, outlining the topics requiring attention, including the significance of physical and biological factors to ecological interactions, and the implications of space and time. It also demonstrated the links between topics usually considered separately. For example, the pyramid of numbers and niche concepts showed how to relate research on feeding by individual organisms to the study of population size and animal community structure. More generally, his view of how ecological communities are organized provided a basis for organizing work in the discipline—the order of ecological knowledge and practice paralleling the order of nature.

The view presented in Animal Ecology was eventually superseded by other approaches to theory, methods, and modeling. Nevertheless, two generations of ecologists testified to the influence of the framework presented by Elton, and his principles became staples of ecological training. The book was reprinted nine times and translated into several languages.

In two subsequent books, Animal Ecology and Evolution, published in 1930 (based on three 1929 lectures at the University of London), and The Ecology of Animals, appearing in 1933 as part of a series of popular accounts of biology, Elton presented his ideas to wider audiences. During these years his own views regarding theory were evolving, as well. Reflecting his wariness of efforts to represent the complexity of nature in mathematical terms, Animal Ecology had contained little discussion of work by theorists such as Alfred Lotka and Vito Volterra. Nevertheless, he would eventually find their work helpful, particularly the argument that not just environmental factors, but also interactions between or within species (including competition, the significance of which he had once viewed skeptically), could cause populations to fluctuate. He added a brief discussion of recent theoretical work to the second edition of Animal Ecology in 1935.

In 1928 Elton married Rose Montague (1906–1997). The match was dissolved amicably in 1937. Late that year, he married poet Edith Joy Scovell. They had two children, Catherine (1940) and Robert Andrew (1943). The marriage of Charles and Joy was happy—it was once described as a “true conversation.” Elton’s decision to make family life a priority may explain his reluctance to travel over much of the rest of his career. (He also disliked crowds, and his refusal to attend large meetings was perhaps also a factor.)

The Bureau of Animal Population . By the early 1930s Elton had developed a sophisticated perspective on animal populations and communities. The perspective would prove highly productive, yet it also embodied a tension. In Animal Ecology he had presented ecological communities as integrated, self-regulating entities tending towards equilibrium. At the same time, these communities could be highly unstable: the “balance of nature,” he insisted, was a myth, and populations constantly varied as a result of disease, migration, or environmental change. It was a tension not unlike that facing Elton himself: living in a society that valued stability, yet that gave many only uncertainty and insecurity—a reality that a young and ambitious scientist of uncertain status in his own university could not but be aware.

British ecologists had to struggle for decades to obtain support for their research. Elton was no exception. Temporary positions at Oxford and elsewhere had supported him during the 1920s, but in 1930 the most significant one, his consultancy with the Hudson’s Bay Company, fell victim to the economic crisis. Yet in the same year a new opportunity emerged. At Binney’s instigation, American industrialist Copley Amory enlisted Elton to help organize his 1931 Conference on Biological Cycles, held at Matamek, Quebec. The conference was well attended, testifying to the interest in fluctuating populations, particularly of species with economic significance—several of which were in a state of ruinous scarcity in that region. As conference secretary Elton met many of the most active scientists in the field, including Canadian wildlife officials and biologists, and American game biologist Aldo Leopold, with whom he would develop a lengthy and friendly professional relationship. But he had another interest in participating, as well: the opportunity to discuss his research plans with potential funders. This effort was successful, and Elton returned home with a grant of 564 pounds per year for two and three-quarter years from the New York Zoological Society. These funds enabled Elton to persuade Oxford to give provisional approval for his most ambitious enterprise: the Bureau of Animal Population.

The bureau’s purpose was to study fluctuations in animal populations, with special reference to the role of disease. More specifically, Elton envisaged several functions. One was to enable research on the causes of these fluctuations, through projects such as his study of vole populations. A second was to collect and store population data, including the output from two new studies: the Canadian Arctic Wild Life Enquiry, and the Snowshoe Rabbit Enquiry. Each involved the coordination of hundreds of observers—game managers, police officers, and trappers— who, using questionnaires distributed with the assistance of the Canadian National Parks Bureau, reported on animal population changes in their area. Elton considered this approach superior to his archival studies: It could provide a picture of fluctuations over a large area, and the data were not inferred from fur returns, but based on direct surveillance by knowledgeable observers.

These questionnaire studies—dubbed “mail-order zoology”—were pursued until the late 1940s, generating a series of papers, most written by other bureau ecologists and staff, including Dennis Chitty, Helen Chitty, and Mary Nicholson. The projects accumulated evidence of cycles of abundance of lynx, rabbits, and other species; these data were then tabulated on maps, illustrating Elton’s view that population phenomena, including, among other factors, the consequences of migration and environmental change, may be best understood in spatial terms. Yet explanations for these cycles remained elusive, and Elton eventually came to doubt that this method could find them.

Beyond field research and analysis of population data, the small group that Elton assembled at the bureau pursued a variety of tasks, all focused on population phenomena. These included breeding of laboratory animals for experimental work, research on reproduction, mortality, and the pathology of diseases, development of census methods, and application of statistical and mathematical methods to ecological problems. Whereas Elton himself never made extensive use of mathematics, he recognized its importance, drawing on the work of Lotka and other mathematically inclined scientists, even as he occasionally complained that they gave too little assistance to ecologists hoping to apply their techniques to nature. The value he attached to mathematics was evident in the role he gave Patrick Leslie, who joined the bureau in 1935 as assistant in biomathematics. Leslie worked closely with ecologists, introducing them to statistical and modeling techniques.

In its early years, financial security for the bureau proved elusive. Besides the crucial support of the New York Zoological Society, the bureau received funds from other sources, including the Royal Society of London, the Agricultural Research Council (ARC), and the chemical firm ICI. Elton saw fundraising as a waste of time and a distraction from research, but he pursued the task energetically. During one year alone (1937–1938) the bureau listed support from thirteen external agencies—the product of more than 100 letters of request. These supporters, particularly the ARC, also pressed Oxford itself to provide stable funding, and in 1935 it began to do so, albeit with the proviso that the Bureau continue to pursue outside support. Elton was also named a senior research fellow of Corpus Christi College in 1936.

While at times unsteady financially, and with only a small staff, the bureau nevertheless gained an international reputation for population research, attracting visiting scientists and students from many countries. It was said that almost everyone of note in ecology made a “pilgrimage” to the bureau, such that it served, as Crowcroft described it, as “the venue of a thirty-five-year international conference” (1991, p. 12). Many students, British and foreign, also came to study, several as Rhodes Scholars. They found not just research opportunities, but an outstanding library, and opportunities for discussion and collaboration. They also enjoyed the close-knit, collegial atmosphere of the bureau, even if most were conscious of their status as colleagues or students, but not friends, of “the boss.” Elton himself did not travel extensively. In 1938, however, he and Joy toured Canada and the United States visiting colleagues, including Canadian biologists and conservationists, as well as Aldo Leopold in Wisconsin.

In 1932 the British Ecological Society, acknowledging the distinctive identity of Elton’s field, permitted him to establish the Journal of Animal Ecology. He would edit it until 1951. In addition to conventional research articles, the journal presented abstracts of reports published elsewhere, including the publications of local natural history societies. This exemplified Elton’s view that information collected by amateur but knowledgeable observers could also be an essential resource for ecologists—a view evident in his population research during the 1920s and 1930s, and that would be presented most expansively in his 1942 monograph, Voles, Mice, and Lemmings. He echoed this point in his communication with wider audiences. When in 1932, for example, the British Broadcasting Corporation invited him to present a series of radio lectures (these eventually appeared in book form as Exploring the Animal World, 1933), Elton encouraged his listeners to record systematically their observations of birds and other fauna.

Information provided by careful observers was, of course, only useful if it was organized. This was, as has been seen, one of the functions of the bureau, and this illustrates how Elton saw this task of organizing—of information, research collaborations, expeditions, publications—as a priority, and one which placed him at the center of diverse social networks combining amateurs and professionals. In addition, the bureau itself, as a storehouse of ecological information, and a crossroads in the professional terrain of ecology, reinforced his role in coordinating ecological research and knowledge. Together, these ambitious efforts constituted, in effect, partial fulfillment of plans for the organization of knowledge of the natural world along ecological lines that Elton had himself sketched on the last page of Animal Ecology.

Wytham Woods . Just as in the 1920s Elton formulated the perspectives that would guide his research over the following decade, the 1940s would be similarly formative. This time, however, he reconsidered, revised, even rejected, some of his earlier ideas. First, however, urgent events intervened. Shortly before war began in September 1939, Elton had proposed to the Ministry of Agriculture a study of the control of rodent pests endangering food stocks. This proposal—reflecting both Elton’s interest in the practical implications of ecology, and his now well-honed ability to pursue funding opportunities—received quick approval, and until 1947 the survey and elimination of rodents (mostly rats, but also the house mouse) preoccupied Elton and other bureau ecologists. This work, apparently such a sharp departure from his previous studies, in fact drew on the research he and his colleagues had done on rodent population dynamics, while branching into new directions, including animal behavior and the chemistry of poisons.

But in 1942 Elton also published a work distant in both conception and results from his work on rodent control. Voles, Mice, and Lemmings summarized years of research and collation of data relating to animal populations. Ranging widely, from Britain to Lapland to northern Canada, he demonstrated the existence of cycles in various species. Causes proved more elusive, with various possibilities considered—climate, interactions between populations, migration, environmental change—but with each able to account for only some observations. G. Evelyn Hutchinson of Yale University, who practiced an ecology more focused on theory and the testing of hypotheses, was both impressed by the book’s avalanche of empirical observations, and frustrated by its lack of a conclusion. He titled his review, “Nati Sunt Mures, et Facta est Confusio” (Mice are born, and the result is confusion).

Elton was himself moving toward at least partial agreement with Hutchinson. He eventually concluded that the approach followed in Voles, Mice, and Lemmings, of induction from data collected by observers scattered over a large area, was unlikely to generate satisfying explanations. Helen Chitty had maintained the Canadian wildlife questionnaire surveys while the rest of the bureau was preoccupied with rodent control, but in the late 1940s they were suspended.

Long before then—indeed, the same year that he published Voles, Mice, and Lemmings—Elton had begun exploring an alternative approach to ecological survey, involving studies of population dynamics, interactions between species, and habitats, with the ultimate goal of building a detailed understanding of the ecology of a specific site. This approach reflected his growing interest, shared with other British ecologists, including Arthur Tansley (who had coined in 1935 the term ecosystem), in breaking down the division between plant and animal ecology, with the aim instead of formulating a synthetic perspective on ecological communities and their habitats.

An opportunity soon arose to put these plans into effect. Elton had originally envisaged an ambitious ecological survey of Oxfordshire, Berkshire, and Buckinghamshire, but in 1943 a more limited and therefore more practical terrain became available when the thousand-acre Wytham Woods estate near Oxford was donated to the university. It possessed abundant possibilities for ecological studies: As Elton described the scene in 1966, “a rich series of habitats from open ground and limestone to woodland with many springs and marshes interspersed occupies a hill set in riverine surroundings” (p. 17). It also had the security of tenure necessary for long-term studies—a valuable feature, given British ecologists’ experience of losing research sites to development. A further advantage was the support now available from the Nature Conservancy, provided in recognition of the project’s status as a pilot program for the ecological survey of nature reserves. As it did for British ecology generally, the Conservancy provided Elton with the novelty of reliable research funding. (Incorporation within the university’s Department of Zoological Field Studies in 1947 had also given the bureau firmer institutional status.)

For two decades after the war, Wytham Woods was the site of intensive research, with Elton hosting field ecology courses, entertaining visiting scientists, supervising projects and graduate theses—providing, in short, the terrain on which he could apply his aptitude for organizing ecological research and information. These studies also contributed to the Wytham Biological Survey, an approach to data collection based on a system of habitat classification devised by Elton and Richard Miller, an American postgraduate.

In 1966 Elton summarized the findings of the first two decades of the survey in The Pattern of Animal Communities. As he did so, he distinguished his approach from that of ecologists who, overly “dazzled by the technical and mathematical triumphs of physics and chemistry,” in his view embraced quantitative approaches at the expense of understanding nature’s complexities (p. 374). Whereas, as has been observed, Elton valued mathematics, he never risked retreating into formulae. Always stressing the need for careful observation, often illustrating points through anecdotes drawn from the field, at heart he remained a field naturalist, with a strong aesthetic sense and a naturalist’s joy at observing life. And as he once noted, fieldwork was essential if one was to have time to think!

Nature Conservation . Elton’s affinity for nature was expressed not only through his research, but through his long-standing concern for the practical implications of ecology. These implications—fluctuations in the Canadian fur trade, or the impacts of rodents on food stocks, for example—commonly reflected, but went beyond, the priorities of his patrons. Elton was also a determined communicator, incorporating conservation concerns into his popular writings and radio talks.

During and after World War II Elton contributed to ecologists’ advocacy for nature conservation. He served on the British Ecological Society’s committee on nature conservation, and then worked with Tansley and other ecologists on the Wild Life Conservation Special Committee. Its 1947 report presented a plan for the Nature Conservancy, duly established two years later as the national agency responsible for ecological research, protection and management of nature reserves, and provision of advice on conservation. Like Tansley, John Baker (his Oxford colleague and Bagley Wood collaborator), and other British ecologists who were members of the Society for Freedom in Science, Elton combined these practical concerns with a belief in the need for science to remain independent of utilitarian compulsion. He continued until 1956 to serve on the Conservancy’s Scientific Policy Committee; in more subtle ways his impact was expressed through his influence on individuals such as Max Nicholson, the Conservancy’s dynamic Director-General.

The most prominent of Elton’s practical concerns related to the ecological consequences of the movement of species. To his geographical perspective and interest in migration, already evident in his research on wildlife population cycles, Elton added a fascination with (as an ecologist) and an alarm regarding (as a conservationist) what he called “the breakdown in Wallace’s realms”: the establishment of populations of plants and animals in new areas as a result (intentionally or otherwise) of human actions. In 1958 he published The Ecology of Invasions by Animals and Plants, where he explained why he considered this phenomenon deeply problematic: “we are living in a period of the world’s history when the mingling of thousands of kinds of organisms from different parts of the world is setting up terrific dislocations in nature” (p. 18). Presenting evidence that more diverse ecological communities are more stable and so better able to resist these dislocations, he concluded that conservation must encompass the preservation of diversity.

Elton retired in September 1967. To his chagrin, and in spite of his advocacy, the university took this occasion to close the bureau. This was partly the consequence of changing views regarding the organization of research at Oxford. It also reflected, however, how research in animal ecology had shifted away from Elton’s “scientific natural history.” The style of patient observation and data collection represented by the Wytham Biological Survey was less suited to newer trends in theoretical ecology, or to an increasingly competitive scientific world that demanded immediate results. Young ecologists such as Robert MacArthur were pursuing new approaches to theory, modeling and experimentation with populations; ecosystem ecology, with its focus on flows of energy and nutrients, was attracting much attention. To some, Elton’s continued focus on accumulation of data and induction of general principles had begun to appear as the relic of an earlier generation. Elton himself conducted relatively little research after retirement, although visits to South and Central America in 1965, 1968, and 1970 provided the basis for a study of the invertebrate ecology of the neotropical rain forest.

Yet Elton had an enormous influence on the field—a legacy testified to by countless students and colleagues. The concepts he elucidated: of food chains and cycles, the pyramid of numbers, and the niche, became staples of ecological training—so intrinsic to the grammar of the field that many no longer associated them with an actual individual. The broader approach they represented: of understanding ecological communities in terms of feeding relationships—generalized as flows of nutrients and energy—gained a similar status, and eventually underpinned the study of ecoenergetics and ecosystem ecology.

Elton’s influence was also expressed through his organizational vision, most immediately by the formation and nurturing of the Bureau of Animal Population into an institution that influenced the careers of dozens of ecologists. This vision was also evident in his contributions to constructing animal ecology as a distinct field of study, the formation of the Nature Conservancy as the chief postwar institution for British ecology, and the emergence of environmentalism through his writings and lectures. The more recent revival of his Ecology of Invasions, celebrated by some as a prescient commentary on the new field of “invasion biology,” testifies to his continuing significance.

Elton received many honors. In 1931 he was named a fellow of the New York Zoological Society, in 1953 a fellow of the Royal Society, and in 1983 an honorary fellow of the Institute of Biology in London. He also became in 1961 the first foreigner to be named an eminent ecologist by the Ecological Society of America, and in 1968 was named a foreign honorary member by the American Academy of Arts and Sciences. Additional awards included the Linnean Society Gold Medal (1967), the Darwin Medal of the Royal Society (1970), and the Tyler Award (1976). A succession of organizations named him an honorary life member: the Chicago Academy of Science (1946), The Wildlife Society (1949), the British Ecological Society (1960), the American Society of Mammalogists (1973), and the American Society of Zoologists (1985). He resided in Oxford with his wife Joy until his death in 1991. Elton’s unpublished materials are held by the Bodleian Library, Oxford University.

BIBLIOGRAPHY

WORKS BY ELTON

With Victor S. Summerhayes. “Contributions to the Ecology of Spitsbergen and Bear Island.” Journal of Ecology 11 (1923): 214–268.

“Periodic Fluctuations in the Numbers of Animals: Their Causes and Effects.” British Journal of Experimental Biology 2 (1924): 119–163.

Animal Ecology. London: Sidgwick & Jackson, 1927.

Animal Ecology and Evolution. Oxford, U.K.: Clarendon Press, 1930.

The Ecology of Animals. London: Methuen, 1933.

Voles, Mice, and Lemmings: Problems in Population Dynamics. Oxford, U.K.: Clarendon Press, 1942.

“Population Interspersion: An Essay on Animal Community Patterns.” Journal of Ecology 37 (1949): 1–23.

“Research on Rodent Control by the Bureau of Animal Population September 1939 to July 1947.” In Control of Rats and Mice, 3 vols., edited by Dennis Chitty. Oxford, U.K.: Clarendon Press, 1954.

With Richard S. Miller. “The Ecological Survey of Animal Communities: With a Practical System of Classifying Habitats by Structural Characters.” Journal of Ecology 42 (2, 1954): 460–496.

The Ecology of Invasions by Animals and Plants. London: Methuen, 1958.

The Pattern of Animal Communities. London: Methuen, 1966.

OTHER SOURCES

Anker, Peder. Imperial Ecology: Environmental Order in the British Empire, 1895–1945. Cambridge, MA: Harvard University Press, 2001.

Chity, Dennis. Do Lemmings Commit Suicide? Beautiful Hypotheses and Ugly Facts. New York: Oxford University Press, 1996.

Cox, David L. “Charles Elton and the Emergence of Modern Ecology.” PhD diss. St. Louis, MO: Washington University, Department of Biology, 1979.

Crowcroft, Peter. Elton’s Ecologists: A History of the Bureau of Animal Population. Chicago: University of Chicago Press, 1991.

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

Hardy, Alister. “Charles Elton’s Influence in Ecology.” Journal of Animal Ecology37 (1968): 3–8.

Hutchinson, G. Evelyn. “Nati sunt mures, et facta est confusion.” Quarterly Review of Biology17, no. 4 (1942): 354–357. Review of Charles Elton’s Voles, Mice and Lemmings.

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

Lindström, Jan, Esa Ranta, Hanna Kokko, et al. “From Arctic Lemmings to Adaptive Dynamics: Charles Elton’s Legacy in Population Ecology.” Biological Reviews 76 (2001): 129–158.

Macfadyen, Amyan. “Obituary: Charles Sutherland Elton.” Journal of Animal Ecology61 (1992): 499–502.

Morrell, Jack. Science at Oxford 1914–1939: Transforming an Arts University. Oxford, U.K.: Clarendon Press, 1997.

Paviour-Smith, Kitty. “Elton, Charles Sutherland (1901–1991).” Oxford Dictionary of National Biography. Oxford: Oxford University Press, 2004. Available from http://www.oxforddnb.com.

Southwood, Richard, and J.R. Clarke. “Charles Sutherland Elton.” Biographical Memoirs of Fellows of the Royal Society 45 (1999): 129–146. Includes a complete Elton bibliography.

Stephen Bocking

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"Elton, Charles Sutherland." Complete Dictionary of Scientific Biography. 2008. Encyclopedia.com. 25 Sep. 2016 <http://www.encyclopedia.com>.

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"Elton, Charles Sutherland." Complete Dictionary of Scientific Biography. 2008. Retrieved September 25, 2016 from Encyclopedia.com: http://www.encyclopedia.com/doc/1G2-2830905648.html

Elton, Charles Sutherland

Elton, Charles Sutherland

British Biologist 1900-1991

Charles Sutherland Elton was born in Liverpool, England. Elton is considered the father of animal ecology. Ecology is the study of the relations between living creatures and their natural environment.

Elton was educated at Liverpool College and then at New College, Oxford. In 1922 he graduated from Oxford with a degree in zoology. At that time, most scientists studying animals emphasized their physical makeup and performed studies in laboratories. However, Elton was more interested in the scientific study of animals in their habitats. While still in college, Elton began this study, making numerous journeys to the Arctic.

In 1927 Elton published his first book, Animal Ecology. It was considered brilliant, and established many of the basic principles of modern animal ecology. He discussed food chains and the food cycle, niches, and the "pyramid of numbers." The pyramid of numbers is an observation about food relationships. A large number of plants feed a smaller number of animals, and these animals in turn provide food for an even smaller number of meat-eating animals. Elton addressed evolution in his 1930 book Animal Ecology and Evolution.

Elton concentrated his studies on how the number of animals in a population was affected by a changing environment. In 1932, he established the Bureau of Animal Population at Oxford, which for thirty-five years served as an important center for worldwide studies of ecology. In 1932 Elton became the editor for the newly established Journal of Animal Ecology.

In 1936 he was appointed to prestigious positions at Oxford and Corpus Christi College. Elton's research on mice populations enabled him to assist his country during World War II with the control of rodent pests. He published two books on rodents, Voles, Mice and Lemmings in 1942 and The Control of Rats and Mice in 1954. Other important books were The Ecology of Invasions of Animals and Plants (1958) and The Pattern of Animal Communities (1966).

In 1953 Elton was elected a member of the Royal Society of London and a foreign member of the American Academy of Arts and Sciences. He was awarded the Gold Medal of the Linnean Society in 1967 and the Royal Society's Darwin Medal in 1970. Elton retired from his studies in 1967.

Denise Prendergast

Bibliography

Elton, Charles S. Animal Ecology. London: Sidgwick & Jackson, Ltd., 1927. Reprint, London: Science Paperbacks and Methuen, 1966. Reprint: New York: October House, 1966. Reprint, Chicago: University of Chicago Press, 2001.

. Animal Ecology and Evolution. Oxford: Clarendon Press, 1930.

. Ecology of Animals. London: Methuen, 1933. Reprint, London: Methuen; New York: John Wiley, 1950. Reprint, London: Science Paperbacks and Methuen, 1966.

. Exploring the Animal World. London: G. Allen & Unwin, 1933.

. Voles, Mice and Lemmings: Problems in Population Dynamics. Oxford: Clarendon Press, 1942. Reprint, Codicote, Herts.: Wheldon & Wesley; New York: Stechert-Hafner, 1965.

. Ecology of Invasion by Animals and Plants. London: Methuen, 1958. Reprint, London: Chapman & Hall, 1977. Reprint, Chicago: University of Chicago Press, 2000.

. Pattern of Animal Communities. London: Methuen; New York; John Wiley, 1966.

McMurray, Emily J., ed. Notable Twentieth-Century Scientists. Detroit: Gale Research Inc., 1995.

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Prendergast, Denise. "Elton, Charles Sutherland." Animal Sciences. 2002. Encyclopedia.com. 25 Sep. 2016 <http://www.encyclopedia.com>.

Prendergast, Denise. "Elton, Charles Sutherland." Animal Sciences. 2002. Encyclopedia.com. (September 25, 2016). http://www.encyclopedia.com/doc/1G2-3400500119.html

Prendergast, Denise. "Elton, Charles Sutherland." Animal Sciences. 2002. Retrieved September 25, 2016 from Encyclopedia.com: http://www.encyclopedia.com/doc/1G2-3400500119.html

Elton, Charles Sutherland

Elton, Charles Sutherland (1900–91), A British zoologist who studied animal communities and made major contributions to the development of ecological studies in general and of animal ecology in particular. One of his early books, Animal Ecology (first published in 1927) was very influential. He emphasized the importance of conserving species and habitats. He proposed the existence of niches, occupied by species at particular points in a food chain. From 1932 until 1967 he was director of the Bureau of Animal Population at the University of Oxford. Elton was elected a Fellow of the Royal Society in 1953 and was awarded many scientific prizes.

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MICHAEL ALLABY. "Elton, Charles Sutherland." A Dictionary of Zoology. 1999. Encyclopedia.com. 25 Sep. 2016 <http://www.encyclopedia.com>.

MICHAEL ALLABY. "Elton, Charles Sutherland." A Dictionary of Zoology. 1999. Encyclopedia.com. (September 25, 2016). http://www.encyclopedia.com/doc/1O8-EltonCharlesSutherland.html

MICHAEL ALLABY. "Elton, Charles Sutherland." A Dictionary of Zoology. 1999. Retrieved September 25, 2016 from Encyclopedia.com: http://www.encyclopedia.com/doc/1O8-EltonCharlesSutherland.html

Elton, Charles Sutherland

Elton, Charles Sutherland (1900–91) A British zoologist who studied animal communities and made major contributions to the development of ecological studies in general and of animal ecology in particular. One of his early books, Animal Ecology (first published in 1927) was very influential. He emphasized the importance of conserving species and habitats. He proposed the existence of niches, occupied by species at particular points in a food-chain. From 1932 until 1967 he was director of the Bureau of Animal Population at the University of Oxford. Elton was elected a Fellow of the Royal Society in 1953 and was awarded many scientific prizes.

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MICHAEL ALLABY. "Elton, Charles Sutherland." A Dictionary of Plant Sciences. 1998. Encyclopedia.com. 25 Sep. 2016 <http://www.encyclopedia.com>.

MICHAEL ALLABY. "Elton, Charles Sutherland." A Dictionary of Plant Sciences. 1998. Encyclopedia.com. (September 25, 2016). http://www.encyclopedia.com/doc/1O7-EltonCharlesSutherland.html

MICHAEL ALLABY. "Elton, Charles Sutherland." A Dictionary of Plant Sciences. 1998. Retrieved September 25, 2016 from Encyclopedia.com: http://www.encyclopedia.com/doc/1O7-EltonCharlesSutherland.html

Elton, Charles Sutherland

Elton, Charles Sutherland (1900–91) British zoologist and ecologist, who founded the Bureau of Animal Population at Oxford in 1932 and the same year became editor of the new Journal of Animal Ecology. The first zoologist to study animals in relation to their environment, he explored the nature of food chains and studied population fluctuations.

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"Elton, Charles Sutherland." A Dictionary of Biology. 2004. Encyclopedia.com. 25 Sep. 2016 <http://www.encyclopedia.com>.

"Elton, Charles Sutherland." A Dictionary of Biology. 2004. Encyclopedia.com. (September 25, 2016). http://www.encyclopedia.com/doc/1O6-EltonCharlesSutherland.html

"Elton, Charles Sutherland." A Dictionary of Biology. 2004. Retrieved September 25, 2016 from Encyclopedia.com: http://www.encyclopedia.com/doc/1O6-EltonCharlesSutherland.html

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