Emerson, Alfred Edwards

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(b. Ithaca, New York, 31 December 1896;

d. Huletts Landing, New York, 3 October 1976), biogeography, ecology, entomology, evolutionary biology, systematic biology.

Emerson was a leading authority on the classification, anatomy, and biogeography of termites, noted for his writing on the concept of the “superorganism.” He used termite nest structure and termite colony behavior as important traits in his classifications. He also made significant contributions to the “Chicago school” of ecology and debates over evolutionary speciation, drawing generalizations from data and observations on adaptations, behavior, geographical distribution, and physiology.

Early Life and Education . Emerson was born in Ithaca, New York, on 31 December 1896, the youngest of four children of Alfred Emerson, a classical archaeologist then teaching at Cornell University, and Alice Louisa (Edwards) Emerson, a concert pianist and instructor in the history of music. He grew up in Chicago after his father joined the staff of the University of Chicago’s Art Institute as curator of antiquities. Family life revolved around the arts, and Emerson developed an early interest in music. Thoughts of a career in music, however, gave way to a fascination with biology, after he was placed in charge of building and maintaining a poultry farm at the Interlaken School, Rolling Prairie, Indiana, which he attended from 1910 to 1914. Emerson later described himself as the Emerson family’s “scientific mutant.” In 1914 he returned to Ithaca and enrolled at Cornell University for its agricultural program, but he soon found that the program lacked the intellectual challenge he desired.

After taking courses in all of Cornell’s scientific departments, he decided to major in entomology, studying at the premier entomology department in the United States under Professors John Henry Comstock and James G. Needham. The Comstock school of entomology focused on an evolutionary approach to insect systematics that included analysis of embryological, ecological, and behavioral characters. In the Cornell “Insectary,” students observed living organisms in their natural environments. Needham was also a pioneer in the emerging fields of ecology and limnology, introducing students to insect life in its dynamic aquatic environments. Emerson built on this strong base in his own career. He also formed a close friendship with the Comstocks and Needhams. Professor Comstock’s wife, Anna Botsford Comstock, was a leading figure in the Nature Study movement in the United States. Emerson attended Cornell with Karl Patterson Schmidt, a lifelong friend and collaborator, who was later curator of herpetology at the Chicago Natural History Museum, now the Field Museum.

Career in Termite Systematics . After receiving his BS from Cornell in 1918, Emerson served in the U.S. Army during World War I, but he did not see combat. Following his discharge in December 1918, he made the first of several field trips to the tropics, visiting the New York Zoological Society’s Station at Kartabo in British Guiana. The trip proved to be a pivotal moment in his life, when the New York Zoological Society naturalist and pioneering marine biologist William Bebee suggested that Emerson focus his research on termites, setting the course of his scientific career. Emerson completed the MA at Cornell in 1920 and returned to Kartabo for additional fieldwork that year and again in 1924, serving as assistant director of the station and amassing a large taxonomic collection and behavioral observations that would be the basis for years of research. In 1925 he was awarded the PhD from Cornell, with a dissertation on “The Termites of Kartabo, Bartica District, British Guiana,” which was subsequently published in Zoologica. In 1921 he had been appointed an instructor at the University of Pittsburgh; after receiving his doctorate, he advanced to associate professor there from 1925 to 1929. He was also a Guggenheim Fellow in 1925 and 1926, conducting research at laboratories in Italy and Sweden on the ontogenetic and phylogenetic origin of the castes of termites.

In 1929 Emerson was appointed associate professor of zoology at the University of Chicago, where he remained until his retirement in 1962. In 1934 he advanced to full professor. This highly regarded department had been founded in 1892 as a research center by the noted embryologist Charles Otis Whitman, who also directed the Marine Biological Laboratory at Woods Hole, Massachusetts. At the time of Emerson’s appointment, the Department of Zoology was chaired by embryologist Frank R. Lillie. Colleagues in the department included Sewall Wright, a pioneer in the application of statistical analysis to genetics and evolutionary biology, and Warder Clyde Allee, an ecologist interested in the evolutionary significance of animal societies. Emerson was an active participant in the development of a revitalized undergraduate curriculum in biology at Chicago, known as the “New Plan.” In earlier years, undergraduate biology education had languished at the university. Under Lillie’s leadership, a series of four basic courses in biological sciences were developed and taught by all department faculty. While research remained the highest priority of the department, undergraduate and graduate teaching were now required of all professors. Faculty and student interaction was also fostered by meetings of the “Zoology Club,” a weekly social gathering followed by a lecture and discussion. During his career, Emerson also held visiting appointments as an instructor in the biology laboratory at Cold Spring Harbor in 1922, at the University of California at Berkeley in the summer of 1949, and as Distinguished Professor of Natural Science at Michigan State University in the spring of 1960.

Emerson published more than one hundred articles on the natural history, classification, evolution, and bio-geography of termites. He was probably the most prolific of the termite systematists, publishing species descriptions as well as revisions of higher taxa. He was a research associate of the Chicago Museum of Natural History from 1942 and the American Museum of Natural History from 1940 until his death. In 1935 he conducted field work at the Canal Zone Biological Area on Barro Colorado Island in Panama, now the Smithsonian Tropical Research Institute. During his career, he conducted fieldwork in North, Central, and South America; the Caribbean; Asia; the Indian subcontinent; Pacific islands; Australia; Africa; and Europe. His collection contained specimens of 1,745 species, some 91 percent of the known taxa; it was acquired by the American Museum of Natural History. Emerson was deeply interested in termite physiology, morphology, and behavior. His observations focused on the social behavior and complex division of labor into separate castes in the termite colony, a stable environment that termites create, monitor, regulate, and defend. He was especially interested in using such behavioral traits as geographically specific nest-building techniques to differentiate taxonomic groups. Emerson called the structure of termite nests “frozen behavior”—a character that could be observed, illustrated, and weighed as accurately as any anatomical character. He demonstrated that species of Apicotermes termites that looked similar could be differentiated by their nest-building habits. He also studied the other organisms that had evolved within the environment created by the termite colony. Emerson also distinguished between stable, nonfunctional characters and rapidly evolving functional characters in his classifications. Emerson’s work was highly influential in integrating behavioral characters into systematic and evolutionary biology.

Emerson was also an expert on the geographical distribution of termite taxa, publishing such works as “The Biogeography of Termites” in 1952. His significant publications on the geographical range and evolution of termites ensured that termite—and insect—distribution was an important component of the emerging field of biogeography. Emerson was among the group of thinkers who developed the “evolutionary synthesis” in the 1940s, integrating modern genetic theory with Charles Darwin’s theory of evolution of species. He was a founding member of the Society for the Study of Evolution, and he participated in symposia surrounding the centennial of the publication of Darwin’s On the Origin of Species in 1859.

The Chicago School . At the University of Chicago, Emerson, Warder Clyde Allee, and Thomas Park became the leading figures in the “Chicago school” of ecology, which shared a conceptual framework that emphasized the importance of cooperation in the natural world and focused on the population, not the individual, as the unit of ecological study and evolutionary selection. The Chicago school became primarily known through their 1949 seminal work, Principles of Animal Ecology, which synthesized contemporary ecological thought and highlighted the emerging field of population biology. The volume was known popularly as “The Great AEPPS” for its authors: Warder Clyde Allee, Alfred Edwards Emerson, Orlando Park, Thomas Park, and Emerson’s old friend, Karl Patterson Schmidt. Allee, Emerson, and Thomas Park were on the University of Chicago faculty. The Park brothers were both students of Allee; Orlando Park then joined the faculty at nearby Northwestern University. Schmidt, who attended Cornell with Emerson, was then curator of herpetology at the Chicago Natural History or Field Museum. The five biologists formed the core of the “Ecology Group,” an informal seminar that met on alternate Monday evenings to discuss contemporary ideas. Thomas Park focused his research on the experimental analysis of population phenomenon, and his 1939 paper to the group, “Concerning Ecological Principles,” stimulated work on the volume. For the next decade, the five met on Sunday evenings to carefully review and discuss each chapter.

The AEPPS text is credited with stimulating much population biology and community ecology in the 1950s and 1960s. It also created the model for later ecology textbooks, such as Eugene P. Odum’s 1953 Fundamentals of Ecology. Emerson’s most distinctive contribution was to highlight the importance of studying behavior in the social insects. The Chicago school members believed that only group selection could account for many of the ecological and evolutionary processes they observed in populations. They were influenced by their colleague Sewall Wright’s interdemic model of selection in which evolution is the result of natural selection operating on a set of demes or small semi-isolated groups. They also focused on the concept of cooperation, arguing that competition, at the group level, could be either beneficial and therefore a form of cooperation, or it could be dysfunctional. Their work on cooperation had been stimulated by the 1939 book Bio-Ecology, by Frederick Clements and Victor Shelford, which coined the term disoperation to describe interactions between organisms that had immediate harmful effects. In his work on termites, Emerson argued that evolution could only be understood by studying the action of selection on populations, rather than individuals, and used the concept of cooperation to explain the origins of sociality and social organizations in insect communities. Emerson believed that cooperation was superior because it created a division of labor, allowed selection to act at a group level, and tended to create homeostasis. The cooperative and productive work of the Chicago school of ecology was itself a demonstration of Emerson’s belief than organisms functioned best in groups.

The Concept of the Superorganism . Emerson is perhaps best known for his use of the concept of the “superorganism” in evolutionary biology, a concept first developed by the noted ant specialist William Morton Wheeler in the early twentieth century. Wheeler had taught at the University of Chicago prior to Emerson’s arrival. Both men were influenced by their observations and studies of complex insect societies that differentiated members through a division of labor. Emerson integrated thinking on homeostasis and equilibrium theory with Darwin’s principle of natural selection to develop his theory of biological and social evolution. In publications such as his 1952 essay, “The Superorganismic Aspects of the Society,” Emerson used the behavioral and structural characters of the colony, as well as the anatomical and physiological characters of individual organisms. For Emerson, the social group—in his research, the entire termite colony—was the unit of selection in evolution, not the individual organism. Competition within and between species, he argued, occurred at multiple levels within an ecological system and created either cooperation or conflict. He concluded that reproductive behavior and the division of labor were the fundamental group adaptations that controlled the social evolution of the group. In 1950s essays such as “Dynamic Homeostasis: A Unifying Principle in Organic, Social, and Ethical Evolution” and “Homeostasis and Comparison of Systems,” he argued that the social insects were the exemplification of “dynamic homeostasis,” that is, their group level adaptations led to a greater overall functional efficiency and ecological equilibrium, a concept that he believed would prove to be a unifying principle of evolutionary biology. Emerson’s concept of the “superorganism” had little long-term impact on a field that was to move away from holistic approaches to research, focusing on more specialized studies. However, Emerson’s work was an important influence in the “units of selection” debate that stimulated much work in evolutionary biology throughout the second half of the twentieth century.

Emerson also extended his analysis of the evolution of social dynamics to human populations in essays such as “Biological Sociology” in 1941, “Biological Principles of Human Social Integration” in 1948, “Human Cultural Evolution and Its Relation to Organic Evolution of Termites” in 1962, and in 1973 “Some Biological Antecedents of Human Purpose.” Emerson acknowledged a difference between animal and human social evolution, based on human ability to communicate through learned symbols, but argued that such communication was subject to the same forces as genes. Emerson and Allee were regarded as highly principled men, and their scientific views were integrally related to their social views. Indeed, during World War II, they publicly argued against the misuse of Darwin’s principle of the survival of the fittest to justify nationalist aggression and genocide. They maintained that cooperation was more often the key to survival and well-being among social organisms, especially humans. Allee’s views, influenced by his Quaker religion, held that cooperation was an absolute good. Emerson’s concept of cooperation was based on a relativistic natural ethics that held that the good and right were those things and actions that functioned to ensure efficient homeostasis for all members of the society. Human ethics served as an integrating mechanism at many levels in human societies that led toward greater homeostasis. Both Allee and Emerson advocated social democracy, based on peaceful intergroup competition, similar to Wright’s interdemic selection. Emerson rejected the communism and socialism of his day, arguing that competitive capitalism, under strong social control, produced better results.

Emerson also engaged in the discussions of science and religion in his 1960 article “The Impact of the Theory of Evolution on Religion.” Throughout his career, Emerson addressed popular, as well as scholarly, audiences, likely reflecting the influence of Anna Botsford Comstock and the Nature Study approach he had learned at Cornell. Indeed, Emerson’s first publication, “A Willy-Nilly Stepmother and Other Disasters,” appeared in 1917 in the Nature Study Review. He also wrote for such popular journals as Classroom Teacher, Natural History, and New Republic. In 1937 he coauthored a children’s book on termite biology and society, Termite City, with Eleanor Fish.

Emerson’s professional activities reflected his broad interests in contemporary biology. Emerson was an influential member of the Ecological Society of America, serving as editor of Ecology from 1932 to 1939, as well as secretary-treasurer in 1931 and president in 1941. He was an active member of the Society for Systematic Zoology, serving as its president in 1958, and of the Entomological Society of America, serving as vice president. He was a co-organizer of the Society for the Study of Evolution, and he served as its president in 1960. He was a fellow of the New York Zoological Society and the Animal Behavior Society. He was active in the American Association for the Advancement of Science, serving as Section F, Zoological Sciences, vice president in 1946, and in the Illinois State Academy of Science, serving as president from 1945 to 1946. Emerson also served on the Cornell University Committee on Systematics, Evolution, and Environmental Biology; the Atlantica Foundation, an organization that supported research, especially the Atlantica Ecological Research Station in Southern Rhodesia; and the Belgian American Educational Foundation, a group that supported technical schools, libraries, and scientific research. In recognition of his contributions to zoology, Emerson received an honorary DSc from Michigan State University in 1961. He was elected a Fellow of the National Academy of Sciences in 1962 and served as chair of the board of its Bache Fund. He received the Eminent Ecologist Award of the Ecological Society of America in 1967.

Family Life and Interests . In 1920 Emerson married Winifred Jelliffe, whom he had met at Cornell. She was the daughter of the prominent psychiatrist Smith Ely Jelliffe. They had two children, Helena Emerson (Wilkening) and William Jelliffe Emerson. After Winifred’s sudden death in 1949, while Emerson was a visiting instructor in biology at the University of California at Berkeley, Emerson married Eleanor Fish in 1950. An old friend, she had coauthored the children’s book Termite City with him; she died in 1971. Emerson’s younger sister, Gertrude (Mrs. Boshi Sen), lived in India and became editor of Asia Magazine. Because of her extensive connections there, Emerson visited the subcontinent several times and formed a friendship with Indian Prime Minister Indira Gandhi in his later years.

After retirement from the University of Chicago in 1962, Emerson was named professor emeritus and moved to Huletts Landing, New York. He made his second “around the world” tour, continuing his taxonomic collecting and ecological observations. He and his wife, Eleanor, returned to Chicago for several months each year to conduct research at the university’s libraries and laboratories, as well as to enjoy the music and arts offerings of the city. He continued to write and publish in his later years, especially on fossil termites. In 1976, the year of his death, he revised two papers for inclusion in External Constructions of Animals. He died in Huletts Landing, New York, on 3 October 1976, following a heart attack.


The Alfred E. Emerson Papers, 1917–1976, are held by the University of Chicago Library, Special Collections Research Center.


“A Willy-Nilly Stepmother and Other Disasters.” Nature Study Review 13 (1917): 198–199.

“The Termites of Kartabo, Bartica District, British Guiana.” Zoologica 6 (1925): 291–459; 7 (1926): 69–100.

Second Year College Sequence in Botany, Zoology, and Physiology. Syllabus for Zoology. Chicago: University of Chicago Press, 1931.

With Eleanor Fish. Termite City. Chicago: Rand McNally, 1937.

“Biological Sociology.” Journal of the Scientific Laboratories, Denison University 36 (1941): 146–155.

“Biological Principles of Human Social Integration.” Intercultural Education News 9 (1948): 8–9.

With Warder Clyde Allee, Orlando Park, Thomas Park, and Karl P. Schmidt. Principles of Animal Ecology. Philadelphia: W.B. Saunders Co., 1949.

“The Biogeography of Termites.” Bulletin of the American Museum of Natural History 99 (1952): 217–225.

“Phylogeny of Social Behavior as Illustrated by the Termite Genus Apicotermes.” Bulletin of the Ecological Society of America 33 (1952): 66.

“The Supraorganismic Aspects of the Society.” In Structure et Physiologie des Societes animals. Colloquium International de Centre National de la Recherche Scientifique, no. 34, 1952.

“The Biological Foundations of Ethics and Social Progress.” In Goals of Economic Life, edited by A. D. Ward. New York: Harper and Brothers, 1953.

“Dynamic Homeostasis: A Unifying Principle in Organic, Social, and Ethical Evolution.” Scientific Monthly 78 (1954): 67–85.

“Homeostasis and Comparison of Systems.” In Toward a Unified Theory of Human Behavior, edited by Roy R. Grinker Sr. New York: Basic Books, 1956.

“Ethospecies, Ethotypes, Taxonomy, and Evolution of Apicotermes and Allognathotermes (Isoptera: Termitidae).” American Museum Novitates 1771 (1956): 1–31.

“The Impact of the Theory of Evolution on Religion.” In Science Ponders Religion, edited by Harlow Shapley. New York: Appleton-Century-Crofts, 1960.

“The Impact of Darwin on Biology.” Acta Biotheoretica 15 (1962): 175–216.

“Human Cultural Evolution and Its Relation to Organic Evolution of Termites.” In Termites of the Humid Tropics, Proceedings of the New Delhi Symposium, 4–12 October 1960. Paris: UNESCO, 1962.

“A Revision of the Tertiary Fossil Species of the Kalotermitidae (Isoptera).” American Museum Novitates 2359 (1969): 1–57.

“Some Biological Antecedents of Human Purpose.” Zygon 8, no. 3–4 (1973): 294–309.

“Termite Nests: A Study of the Phylogeny of Behavior” and “Changes in the Biological Basis of Termite Ethology with More Recent Bibliography.” In External Constructions by Animals, Benchmark Papers in Behavior, edited by Nicholas Collias and E. Collias. Stroudsburg, PA: Dowden, Hutchinson and Ross, 1976.


Guide to the Alfred E. Emerson Papers, 1917–1976. Chicago: University of Chicago Library, Special Collections Research Center, 1999.

Mitman, Gregg. “From Population to Society: The Cooperative Metaphors of W. C. Allee and A. E. Emerson.” Journal of the History of Biology 21, no. 2 (1988): 173–194.

Newman, Horatio H. “History of the Department of Zoology in the University of Chicago.” Bios 19 (1948): 215–239.

Park, Thomas. “Alfred Edwards Emerson, Eminent Ecologist—1967.” Bulletin of the Ecological Society of America 48 (1967): 104–107.

“Prof. Alfred Emerson, Pioneer in Ecology, Dies.” Chicago Tribune, 5 October 1976, B7.

Wilson, Edward O., and Charles D. Michener. “Alfred Edwards Emerson, December 31, 1896–October 3, 1976.” National Academy of Sciences Biographical Memoirs 53: 159–177. Washington, DC: National Academy Press, 1982.

Pamela M. Henson