Charles Otis Whitman

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(b. North Woodstock, Maine, 14 December 1842; d. Chicago, Illinois, 6 December 1910)


Whitman grew up on a farm, where at an early age he became interested in natural history, particularly pigeons. He was exceptionally skilled as a self-taught taxidermist and built up quite a museum in his father’s house. By tutoring and teaching in private schools he earned enough to enter Bowdoin College in 1865, receiving the B.A. in 1868. From 1868 to 1872 he was principal of Westford Academy in Massachusetts, and in 1872–1874 he taught at English High School in Boston. The latter post was of crucial importance for his career because it was in Boston that Whitman came under the influence of Louis Agassiz. As a result he was a participant in the first course in marine biology on Penikese Island, conducted by Agassiz in June and July 1873.

In 1875 Whitman went to Europe, first to Dohrn in Naples and then to Leuckart in Leipzig, where he learned the modern methods of microscopy and embryology. His Ph.D. dissertation (he received the degree in 1878) was on the embryology of Clepsine (Glossiphonia). In 1879 he went for two years to the Imperial University of Tokyo, as professor of zoology. Since eight of his students later became well-known zoologists, four of them holding major chairs, he has rightly been called the father of zoology in Japan. From November 1881 to May 1882, Whitman was again at the Zoological Station in Naples, working on the embryology, life history, and classification of the dicyemids, publishing a standard reference work on these mesozoans in 1883. One of the most productive periods in his life was the period 1882–1886, when he was assistant in zoology at the Museum of Comparative Zoology at Harvard, under Alexander Agassiz. From 1886 to 1889 Whitman served as director of the Allis Lake Laboratory at Milwaukee, Wisconsin, where he founded the Journal of Morphology, the first periodical in America devoted to zoology and anatomy. It served as a model for other publications founded in later years.

In 1889 Whitman accepted the chair of zoology at the newly founded Clark University in Worcester, Massachusetts, where he stayed until 1892. In that year he and most of his colleagues in the science departments moved to the newly established University of Chicago.

Whitman played a leading role in the founding of the Marine Biological Laboratory at Woods Hole in 1888 and served as its first director (1893–1908), developing the policies that have made this institution such a signal success. He resigned because his research on the heredity and behavior of pigeons was seriously impeded by his spending every summer at Woods Hole.

Being extremely unselfish, a person of complete integrity, and dedicated to science, Whitman played an important role as an organizer and first director of new institutions, and as the founder and first editor of new journals. Many of them are still flourishing, such as the Woods Hole Marine Biological Laboratory, the American Society of Zoologists (founded in 1890 as the American Morphological Society), Journal of Morphology, and Biological Bulletin.

Whitman’s research was of unusual breadth. For instance, he became interested in leeches as material for embryological research: but he soon turned to the study of their anatomy and taxonomy, and finally of behavior, all of these interests resulting in publications. He found abundant evidence that the development of the leech egg was completely predetermined but–of course–not preformed in a homunculus-type way. He fought the extreme celllineage (“mosaic”) interpretation, stressing the contribution to development made by the interaction of cells. Whitman’s exceptionally careful work and perceptive interpretation had a profound impact on the embryology of his period. His discovery of the sensilla, segmental sense organs in the leeches, greatly facilitated the study of leech morphology and taxonomy.

When, after 1900. the great split occurred between evolutionists who ascribed evolutionary change to the pressure of a few major mutations and those who ascribed it to selection, Whitman was emphatically on the side of selection. He found numerous (now known to be polygenic)characters in his pigeon crosses that did not mendelize in the simple manner claimed by de Vries and Bateson. For more than ten years he bred pigeons, hybridizing 200 domestic varieties and 40 wild species. The results of his crosses are recorded in two posthumously published volumes (1919). Seeing the similarity of variation in related species and the unmistakable trends of evolutionary change from the most primitive to the most advanced species, Whitman developed a theory of orthogenetic evolution. At first held to be totally erroneous, it is now considered far less of a failure since it has been realized that the potential for variation in a phyletic line is very narrowly prescribed by the existing genotype. Whitman’s emphasis was ahead of its time. Some of his studies dealt with the analysis of sexual dimorphism and led to Oscar Riddle’s endocrinological research.

Whitman was one of the pioneers of ethology. His paper “Animal Behavior” (1898) contains many well-chosen examples of innate (nonlearned)behavior. In his later work he analyzed particularly the relation between innate and learned behavior and the ability of animals to adjust their behavior to new experiences. His posthumous “The Behavior of Pigeons” (1919) is an extraordinarily detailed analysis. In particular, courtship and breeding behaviors of some forty species are compared. With Oskar Heinroth’s pioneering work on ducks (1911), it was the first extensive study in comparative ethology.


I. Original Works. A full bibliography of Whitman’s contributions (67 titles) is given by Lillie (see below). The more important ones are “The Embryology of Clepsine,” in Quarterly Journal of Microscopical Science, 18 (1878), 215–315: “A Contribution to the Embryology, Life History and Classification of the Dicyemids,” in Mitteilungen aus der Zoologischen Station zu Neapel, 4 (1883), 1–89: “A Contribution to the History of the Germ Layers in Clepsine,” in Journa of Morphology, 1 (1887), 105–182: “A Series of Lectures on Bonnet and the History of Epigenesis and Preformation,” in Biological Lectures. Marine Biological Laboratory, Woods Hole, Mass (1894), 205–272: “Animal Behavior,” ibid. (1898), 285–338: and the posthumous works edited by Oscar Riddle (vols. and II) and Harvey A. Carr (vol. III): “Orthogenetic Evolution in Pigeons,” Publications. Carnegie Institution of Washington, no 257 (1919), vol. I; “Inheritance, Fertility, and the Dominance of Sex and Color in Hybrids of Wild Species of Pigeons,” ibid., vol.II: and “The Behavior of Pigeons,” ibid., vol.III.

II. Secondary Literature. Informaion on Whitman’s life and work is in C. B. Davenport, “The Personality, Heredity and Work of Charles Otis Whitman,” in American Naturalist, 51 (1917), 5–30: F. R. Lillie, “Charles Otis Whitman,” in C. O. Whitman memorial volume, Journal of Morphology, 22 (1911), xv–lxxvii; and E. S. Morse, in Biographical Memoris. National Academy of Sciences, 7 (1912), 269–288.

Ernst Mayr

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Charles Otis Whitman, 1842–1910, American zoologist, b. Woodstock, Maine, grad. Bowdoin, 1868, Ph.D. Univ. of Leipzig, 1878. From 1892 he was professor of zoology at the Univ. of Chicago. He founded (1887) and edited the Journal of Morphology and was a founder and director (1888–1908) of the Marine Biological Laboratory at Woods Hole, Mass. He wrote on animal behavior, embryology, evolution, microscopical anatomy, and other subjects. Posthumous Works (3 vol.) appeared in 1919.