Allee, Warder Clyde

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(b. near Bloom-ingdale, Indiana, 5 June 1885; d. Gainesville, Florida, 18 March 1955)

ecology, animalbehavior. For the original article on Allee see DSB, vol. 17, Supplement II.

Allee was a central figure in the development of a distinctive school of ecology that flourished at the University of Chicago during the first half of the twentieth century. According to historian Gregg Mitman, ecology at Chicago developed quite independently of Darwinian evolution and Mendelian genetics, being much more heavily influenced by ideas drawn from developmental biology, physiology, and animal behavior. Victor Shelford, Allee’s mentor at the University of Chicago, argued that the study of ecological communities should be conducted without regard to evolutionary problems, focusing instead on the physiological responses of animals to their environments. Allee’s dissertation on the behavior of aquatic isopods (Asellus communis) investigated the effects of dissolved oxygen, carbon dioxide, and other environmental stimuli on the orientation behavior of these small invertebrates in ponds and flowing streams. Populations living in ponds oriented differently from those living in streams with strong currents. Allee discovered that by manipulating environmental conditions he could make isopods

collected from ponds behave like those taken from streams, and vise versa. This seemed to rule out hereditary differences between populations and suggested that orientation behavior was a direct response to environmental factors.

Later Research. When he returned to the University of Chicago as an assistant professor after World War I, Allee broadened this experimental, physiological approach to study the causes of animal aggregations. Allee sometimes referred to this approach to the study of populations as “mass physiology,” and he argued that the unconscious cooperation or “proto-cooperation” found in loose aggregations of isopods and other simple animals was the starting point for the complex, cooperative behaviors found in truly social animals, including humans. Understanding and improving human society became a powerful motivation for Allee‘s ecological and behavioral research.

Allee‘s physiological approach to ecology and behavior was partially transformed by his later interactions with his colleague Alfred Emerson during the 1930s. Emerson, who studied termites and other social insects, brought an evolutionary perspective that had been absent in Allee‘s earlier research. According to Mitman, Allee did not abandon his earlier commitments to physiology and development, but assimilated aspects of Emerson‘s evolutionary thinking. What unified Allee, Emerson, and other University of Chicago ecologists was a deep commitment to the population as a fundamental evolutionary unit. During the late 1930s, Allee increasingly emphasized natural selection as a causal factor in the evolution of behavior, but he believed that it acted primarily on groups of individuals rather than on the individuals themselves. Allee was able to argue that cooperation was a group adaptation that evolved because more cooperative groups had greater success than less cooperative groups. This commitment to group selection allowed Allee to view even phenomena such as dominance and social hierarchies in cooperative terms. In both cases he argued that these social interactions reduced conflict, which had survival value for the population as a whole. This perspective on social behavior was later widely abandoned by evolutionary biologists, particularly during the late 1960s and 1970s.

Social Implications and Legacy. Throughout his career Allee emphasized the implications of biology for understanding human social behavior. His studies of animal aggregations and the evolution of cooperation meshed neatly with his social activism and his political and religious opposition to war. Raised as a Quaker, Allee remained a strict pacifist throughout his life. His renunciation of war and vocal support for conscientious objectors during World War I made him a target of criticism and some public persecution. During the 1940s Allee argued that his behavioral research provided evidence against a biological basis for war and strongly supported a theory of sociality based upon cooperation among individuals. According to Mitman, Allee viewed ecologists as “social healers” whose research could provide a naturalistic basis for ethics. Mitman argues that after World War II, this idea of ecologists as social healers was swept aside by Eugene Odum and other ecosystem ecologists who embraced an engineering perspective and were more interested in environmental issues than social ones. During the 1950s Odum‘s textbook, Fundamentals of Ecology, rapidly eclipsed Principles of Animal Ecology (1949), which Allee had written with his colleagues at the University of Chicago. Despite their striking differences in perspective, Odum agreed with Allee about the importance of group selection and the evolutionary prevalence of cooperation even among very simple organisms.

Both historians and biologists agree that Allee‘s reputation was diminished by George C. Williams‘s Adaptation and Natural Selection (1966). In this highly influential work Williams championed individual selection and effectively discredited purported examples of group selection. Allee was one of the major targets of Williams’s critique. As a result, Allee left a somewhat contradictory legacy. In 1973 the Animal Behavior Society instituted an annual student research award in his name, but Edwin Banks later complained that many younger animal behaviorists were unaware of Allee‘s important contributions to the field. Some ecologists remain interested in the eponymous “Allee Effect” (that undercrowding can be as deleterious to organisms as overcrowding), and they acknowledge Allee‘s pioneering work on this phenomenon. But although Allee viewed aggregations as an example of unconscious cooperation leading to group adaptation, most ecologists in the early twenty-first century explain them in terms of individual fitness. Ironically, Elliot Sober, David Sloan Wilson, and other evolutionary theorists who have attempted to revive interest in group selection strongly disassociate their ideas from what they characterize as naive group selection arguments used by Allee.


Allee‘s papers are archived in the Special Collections Research Center of the University of Chicago Library. A complete bibliography of Allee‘s published work appears in Karl P. Schmidt‘s “Warder Clyde Allee, 1885-1955,” in Biographical Memoirs, published by the National Academy of Sciences.


Animal Aggregations: A Study in General Sociology. Chicago: University of Chicago Press, 1931.

The Social Life of Animals. New York: Norton, 1938.

With Alfred E. Emerson, Orlando Park, Thomas Park, et al. Principles of Animal Ecology. Philadelphia: Saunders, 1949.

Cooperation among Animals, with Human Implications. New York: Schuman, 1951.


Banks, Edwin M. “Warder Clyde Allee and the Chicago School of Animal Behavior.” Journal of the History of the Behavioral Sciences 21 (1985): 345–353.

Mitman, Gregg. “Dominance, Leadership, and Aggression: Animal Behavior Studies During the Second World War.” Journal of the History of the Behavioral Sciences 26 (1990): 3–6.

———.The State of Nature: Ecology, Community, and American Social Thought, 1900–1950. Chicago: University of Chicago Press, 1992.

Schmidt, Karl P. “Warder Clyde Allee, 1885-1955.” BiographicalMemoirs30 (1957): 2-40.

Sober, Elliot, and David Sloan Wilson. Unto Others: The Evolution and Psychology of Unselfish Behavior. Cambridge, MA: Harvard University Press, 1998.

Stephens, Philip A., W. J. Sutherland, and Robert P. Freckleton. “What Is the Allee Effect?” Oikos 87 (1999): 185–190.

Williams, George C. Adaptation and Natural Selection: ACritique of Some Current Evolutionary Thought. Princeton, NJ: Princeton University Press, 1966.

Joel B. Hagen

Allee, Warder Clyde

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(b. near Blooomingdale Indiana. 5 June 1885; d. Gainesville, Florida. 18 March 1955)


Clyde Allee, professor of zoology at the University of Chicago (1921–1950), was a leader of American ecology during the years in which it was effectively established as a subdiscipline of biology. His large number of students and his part in producing the textbook Principles of Animal Ecology (1949) were his most lasting influences on the field, although he was a pioneer of ecological studies of social behavior in animals.

Born on a farm to Mary Newlin and John Wesley Allee, Clyde grew up in the strong Quaker (Society of Friends) community of southern Indiana, He attended the Friends Academy in Bloomingdale and Earlham College in Richmond., Upon graduation in 1908, Allee studied with Victor Shelford at the University of Chicago, where he received the Ph.D. in 1912. That same year he married Marjorie Hill, also a member of the Society of Friends; they had three children. Profoundly religious (in an ethical, not mystical, manner), Allee became intellectually demoted to the theoretical problems of social behavior, cooperation in nature, and the evolution of human ethics. Science held a significant role in his philosophy, and he felt science and religion could mutually profit from an interchange of values. Also reflecting his values was the time he gave to scientific societies, especially the Ecological Society of America and the American Society of Zoologists, editorship of Physiological Zoology (1928–1954), collaboration with students and colleagues, and teaching. He was also a trustee of Eartham College and the Woods Hole Marine Biological Laboratory, In 1938, following a series of operations on a spinal tumor, he was paralyzed from the waist down and confined to a wheelchair for the rest of his life. The death in 1945 of Marjorie Allee, a successful author of children’s books and helpful collaborator on Allee’s books, also deeply affected him emotionally, yet he continued an extremely active academic life. He retired from Chicago in 1950, but took the chair-manship of the biology department at the University of Florida. He married Ann Silver in 1953, Allee died in 1955 shortly after entering the hospital with a kidney infection.

Allee came under two strong influences in his doctoral education, and the manner in which he combined them remained the mark of his career and contribution to ecology. As Shelford’s student he was in the circle of biologists making Chicago a center of research on ecological succession; their descriptive studies were tied in part to Shelford’s attempt to create a physiology of environmental factors. Another influence was F. R. Lillie, departmental chairman at the University of Chicago and also director of the laboratory at Woods Hole, Massachusetts, where Alice spent his summers. There he learned the techniques of the experimental zoologists such as Lillie, C. O. Whitman, and J. Loeb. The experimentalists’ dependence on tropisms as the new paradigm for animal behavior was attractive because it was a program for laboratory progress, and Allee began a long-running series of experimental investigations into what physical factors affect the movements and behavior of a small aquatic crustacean. He taught at a succession of universities for the next decade, always returning to Woods Hole for summer research and teaching. The synthesis he produced was to use the lab-derived results about physical factors to understand the animal’s behavior and place in nature. He also argued that tropisms were insufficient to account fov adaptive peculiarities in nature, thus moving away from strictly physiological and behaviorist explanations. Because he designed experiments in the laboratory to shed light on the larger ecological setting, he became an influential figure in a young science.

By 1921 Allee was back at Chicago and deeply involved in showing how distribution of animals is a response to physical factors. His work was distinguished within ecology by his emphasis on the behavior that produces the distribution. Thus he began his most famous work, on animal aggregation, by opening a specifically ecological research program, described in Animal Aggregations; A Study in General Sociology (1931), with the search for the general principles that lead to aggregation and. ultimately. the structure of biotic communities. He saw all animals existing in a social network, reproducing and interacting ecologically, manifesting a level of sociality more loosely organized than the highly specialized insect societies that dominated earlier theories. Extending the classifications of societies by A. V. Espinas, P. Deegener, and especially W. M. Wheeler, Allee organized the subject around the type or degree of integration of the social group, and considered a far greater range of aggregations to have social significance.

In his career he directed the research of about forty graduate students, who themselves took important posts in American ecology. Together, their physiological experiments demonstrated survival and growth benefits for members of groups, and through the 1930’s he expanded into social effects on learning. Following the impetus of T. Schjelderup-Ebbe’s far-reaching theory of “peck-order,” Allee’s group by the 1940’s centered more on the conditions and ecology of social hierarchies and less on the simpler physiological questions.

Allee himself became more and more concerned with principles and the practice of ecology, producing several well-known books. The Social Life of Animals (1938) presented his theory that cooperation, by which he meant automatic mutual interdependence, is a fundamental trait of life, a principle as important as struggle for existence. Claiming that cooperation is found throughout the biological world, he extrapolated the development of innate tendencies to human altruism in an attempt to use the biology of sociality to improve human society. He denied the supremacy of competition as the law of nature. Within behavioral ecology, however, his grand principle of universal tendencies did not lead to a productive research program. Instead, after 1950, Niko Tinbergen became the major figure in the field with his emphasis on the individual advantage found within social behaviors, including cooperation and apparent altruism.

Allee’s great institutional contribution was leading the group that became an identifiable Chicago school of ecology. Together with A. K. Emerson, O. Park, T. Park, and K. P. Schmidt, he produced a standard textbook, Principles of Animal Ecology, that presented the heart of their view that the ecological community is a metaphorical organism, with homeostasis and an evolutionary history. The distinctive Allee stamp was the weight given physical factors, organized around the idea that shared principles structure all communities. It was an intentional extension of the earlier descriptive approaches and the classifying of types and biogeographical regions; and it was distinct from the Eltonian school’s concentration on food webs, competition, and predation.


A detailed biography appears in Karl P. Schmidt. “Warder Clyde Allee, 1885–1955,” in Biographical Memoirs. National Academy of Sciences, 30 (1957), 2–40, which includes a complete bibliography of Allee’s works. The Joseph Regenstein Library, University of Chicago, has a small collection of Allee’s papers, including research notes, notebooks, correspondence of the Committee on the Ecology of Animal Populations of the National Research Council, and manuscripts. A larger collection of his correspondence is in the Lilly Library, Eartham College, Richmond, Indiana, The Northwestern University Archives, Evanston, Illinois, holds correspondence concerning his lectures for the Norman Wait Harris Lecture Series, which became The Social Life of Animals.

William C. Kimler

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