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Home , Ernst Mayr, George Gaylord Simpson, Paleozoology, Sewall Wright

Memorial to 1902-1984

EVERETT C. OLSON Department o f , University o f California, Los Angeles. California 90024 paleontologist, geologist, biogeographer, svs- tematist. evolutionist, linguist, historian, and philoso­ pher: George Gaylord Simpson was each of these, and more. He was a brilliant and complex man. with per­ sonal and scientific abilities that are yet to be fully comprehended. His mastery of the written word, his eclectic interests and capacity for synthesis have made him unique and the most influential natural historian of the twentieth century. His death, in Tucson. Ari­ zona. October 6. 1984. is a great loss to and the whole broad area of . George is an impossible person to capture simply and in a few paragraphs, not only because of the multiple, parallel, and often crossing themes which run through his scientific career, but also because he shied away from oral commitment of his thoughts, preferring to trust in the written word, of which he was a master. In his own words, from his Unconventional Autobiography (1978):

I do not value the spoken word as a means of serious communication. I do not like to lecture . . . I do not even like oral discussion of anything like a technical subject. Yet he did master considerable fluency in several foreign languages and with a few friends, over coffee, or more usually dry martinis, he would open up. once in a while discussing matters of some importance. Much of what we know of his thoughts must, however, be gained from his writing from approximately 750 technical reports, mono­ graphs. books, and commentaries on the work of others. A common pattern repeats itself throughout—first a glimmer of an idea of interest in a short paper, often followed by unrecorded intense study; enlargement and development of the substantive basis and interpretation in a longer paper: finally, a full-fledged treatment, sometimes followed some years later by a further revision. With this modus operandi as a guide. I will in later paragraphs cite and attempt to evaluate some of his most important contributions, but first some comments on his career which provided the milieu for his studies. George rose rapidly in his chosen profession of vertebrate . He was born in Chicago. Illinois, in 1902 to Joseph Alexander and Helen Julia (Kinney), and soon thereafter the moved to Denver. Colorado. George, after his early years, began his college career at the University of Colorado in 1922. After a short stay there, he continued his education at Yale, where he received his Ph.D. under Richard Swan l ull in 1926 at the age of 24. By this time he had already published several papers, unusual for a graduate student at that time. His first love was Mesozoic , and in this area he attained his first prominence. In 1923 he married Lydia Pedroja. Four daughters issued from this union. Helen. Francis. Patricia (deceased), and Elizabeth, each of whom remained close to George. In I938. after dissolution of the early marriage. George and Anne Roe. a prominent psychologist, were married and Anne became a colleague, companion, and strong influence throughout the remainder of his career. 2 THF GEOLOGICAL SOC1FTY OF AMERICA

George’s first professional position was at the American Museum of in New York in 1926. He progressed rapidly from the position of assistant curator to head of the Department of Geology and Paleontology. I.ike several others at the Museum he was a concurrent professor at in charge of graduate students and closely associated with members of the renowned school of of that institution. Complicated relationships after an incapacitating accident in the Amazon Basin of Brazil, which immobilized him for an extended period of time, led to his resignation from the American Museum and acceptance of a position as Agassiz Professor in the Museum of Comparative , , which he held from 1959 to 1970. The Harvard years were productive, but because of misunderstandings, never fully satisfactory. While still at the American Museum. George and Anne had purchased a home and land in New Mexico, near the San .luan Basin. This served as a base for summer field work on early mammals and stratigraphy and a refuge for work and contemplation. It was fondly called Los Pinavetes and relinquished only in 1967 because of serious health problems that plagued both Anne and George. In 1967 George became professor of zoology at the in Tucson, and no! long thereafter severed his connections with Harvard. He resigned from the University of Arizona after reaching his 80th year of life. As he told me. “It seems the right thing to do.” During his long career. George was honored as a recipient of many honorary degrees from prestigious institutions around the world, by election to an array of scientific and philosophical societies, and by the reception of awards and medals for special aspects of his work. Particularly valued were his membership in the National Academy of Sciences of the and reception of the coveted Penrose Medal of the Geological Society of America in 1952. George’s studies of Mesozoic mammals, which held his interest for many years, were partially succeeded by intensive collecting and field and laboratory analyses of early Tertiary mammals, especially those of the San Juan Basin. The American Museum Scarritt expeditions to Patagonia, of which George was a prominent member, resulted in his lifelong fascination and study of the stratigraphy, morphology , and svstem- atics of these mammals of the “lost world.” One of George’s latest books. Discoverers of the Lost World, treated all of the many other paleontologists who opened up this world, but left his own contribution to be known from his other publications. Behavior and evolution was an embryonic subject when Anne and George enlisted the efforts of colleagues in psychology and evolutionary biology to present a symposium on the subject. Behavior and Evolution. edited by Anne and George, issued from this effort and was a seminal volume in this now greatly expanded field. Work on mammals of the San Juan Basin led George to recognize the need for biometrical analyses. In characteristic fashion he mastered the field and with Anne Roe. in 1939. published Quantitative Zoology, a book that changed the direction of thinking of many young paleontologists. Much later, in I960, a revision with as a third author, became a touchstone of biometry in paleontology, laying a base from which more sophisticated analyses could be developed. During the 1930s. George’s readings of work by . , and J.B.S. Haldane, his associations with the Columbia , and his intimate knowl­ edge of historical evolution led him to an intensive foray into substantive and theoretical genetics. was an ever-present inspiration. The threads of genet­ ics. of organismic evolutionary change, increasing comprehension of the geologic time scale, and George’s capacity to synthesize came together in what to me. at least, was his most stimulating and pervasive theoretical work: Tempo and Mode in Evolution. The MEMORIAL TO GEORGE GAYLORD SIMPSON 3 book was published in 1944 as George was closing his career in military intelligence, but had been in part formulated during the fervor of the 1930s and early 1940s which saw the emergence of the synthetic theory of evolution, under the guiding hands of Theodosius Dobzhansky, , and at Columbia. The paleontological com­ ponent was in Tempo and Mode, and the book brought many contemporary paleon­ tologists into the mainstream of evolutionary thinking. Seeing the importance of genetics in evolution. George attempted, in Tempo and M o d e , a synthesis in which he demonstrated clearly that the record and genetical theory were in no way incompatible, a departure for at least many paleontologists who were still groping for something more than Neo-Darwinistic explanations. Coming as it did at the right time. Tempo and Mode sent waves of stimulation among paleontologists, as documented in the Princeton Symposium volume Genetics, Paleontology and Evolu­ tion (1949). edited by G. Jepsen, E. Mayr, and G. Simpson. Tempo and mode were the vogue by then, but receded as many drew back from “too much speculation,” only later to undergo a strong resurgence in the 1960s as massive new evidence, both organismic and molecular, accumulated. A follow up to Tempo and Mode was the authoritative volume The Major Features of Evolution (1953). much more comprehensive than its predecessor but in some ways more conservative. Of course. George’s early work on genetics, rates of evolution, . and has become a focal point for both admiration and criticism, some of the latter vitriolic. But the current excitement engendered by his ideas, whether pro or con. is a tribute to George’s brilliance and persistent influence. Although his writing extends to many groups of . George’s studies of evolution and , with the notable exceptions of those on penguins, were primarily based on mammals. His systematics. exemplified in The Principles o f Classi­ fication and the Classification of Mammals (1945) and Principles of (1961), have this flavor. His concepts of biogeography, with an interest extending back to his graduate days at Yale, are primarily applicable to Cenozoic mammals and continents. A fascination with the sine que non of mammalian evolution, the Equidae, led his to a volume entitled (1951), which not only gives an evolutionary perspective on the family Equidae, but also incorporates information on the history of modern breeds as well. This, in brief, is George Gaylord Simpson, the scientist supreme, the eclectic individual with bulldog tenacity, and the man who did so much to change thoughts on life and evolution during the twentieth century. Major influences upon the literate public came from his many books, based, it is true, on his researches but broadened into areas of history, philosophy, sociology, and theology as they viewed the role of man in an evolutionary world. The Meaning of Evolution, translated into many languages. This View of Life, The World of an Evolutionist. and Why and How, Some Problems and Methods in Historical Biology tell some of the story. Most of all. however, his life and ways of thinking, his approach to science, his estimates of himself and other scientists, and his personality are portrayed vividly in Concession to the Improbable: An Uncon­ ventional Autobiography (1978). For all that has made him a legend. George was a person with the same doubts, the joys of being correct and winning, a benevolence for an incisive criticism of others, and impatience with critics of his own work that plague us all. He made many friends, many of whom also shied away from close contact at one time or another, if for no other reason than the shear strength of his intellect. He was not always comfortable to be with. Paul O. McGrew was one of his closest friends, along with Bryan Patterson, and I too spent many delightful hours with George and Anne at their New York apartment and as 4 THF. GEOLOGICAL SOCIFTY OF AMERICA they stopped enroutc to New Mexico and called to say come down to chat over martinis and coffee at the Palmer House. 1 think that Paul McGrew was probably the most at ease with George, for they communicated in an outdoor world of nature that they both understood so well. For all the ill health that dogged George for a major part of his adult life, he maintained an avid interest in the pursuit of knowledge until his very last days. Out of a sick bed in 1983. he plunged into completing work on books and papers and on seeing colleagues long after all but the strongest would have relaxed. Perhaps the greatest tribute that can be paid to his work is that it has been the point of departure for so much that is actually or purported to be new in evolution and systematics. Sometimes the criticism of his concepts have been severe, often unjustified, and often capped with misunderstanding. But these too are a tribute. Frequently criticism has missed the mark by being cast shallowly in comparison to the breadth of under­ standing and historical perspective that is the hallmark of George’s work. Somehow one cannot but be reminded of the century and more of flaying at and the continuing insights that have emerged both from the genius and errors of his conclusions.

SELECTED BIBLIOGRAPHY OF G. G. SIMPSON 1925-1928 Mesozoic mammals: I through XI: American Journal of Sciencc, v. X-XV. 1928 Catalogue of the Mesozoic Mammalia in the Geological Department of the British Museum (Natural History): British Museum (Natural History), London. 215 p. 1929 American Mesozoic mammals: Memoir of the Peabody Museum, Yale. no. 3, pt. 1. 171 p. 1931 A new classification of mammals: American Museum of Natural History Bulletin 59. p. 259-293. 1932 Miocene land mammals from Florida: Florida Geological Survey Bulletin 10. p. 7-41. 1933 The ear region and foramina of the cynodont skull: American Journal of Science, v. 26. p. 285-294. 1934 Provisional classification of extinct South American hoofed mammals: American Museum of Natural History Novitates, no. 750, p. 1-21. ____ Attending marvels of a Patagonian journal: MacMillan and Company. New York. 295 p. ____The Scarritt expedition of the American Museum of Natural History. 1930-1934: Science, (n.s.) v. 80. p. 207-208. 1935 The Tiffany fauna upper Paleocene: American Museum of Natural History Novitates. no. 795. p. 1 — 19: no. 816. p. 1-30: no. 817. p. 1-28. 1936 Horses and history: N atural History, v. X XXVIII. New York. p. 276-288. 1937 Patterns of phyletic evolution: Geological Society of America Bulletin, v. 48. p. 303-314. ____ Superspecific variation in the nature and in classification: from the viewpoint of paleontology: American Naturalist, v. I.XXL p. 236-267. ____ The Fort Union of the Crazy Mountain field. Montana and its mammalian faunas: United States National Museum Bulletin 169. 287 p. ____ Skull structure of the Multituberculata: American Museum of Natural History Bulletin I.XIII. p. 727-763. 1938 Osteography of the ear region in monotremes: American Museum of Natural History Novitates. no. 978. p. 1-15. MEMORIAL TO GEORGE GAYLORD SIMPSON 5

1939 (with Anne Roe) Quantitative zoology, numerical concepts and methods in the study of recent and fossil animals: McGraw-Hill Book Co.. New York and London. 414 p. 1940 Mammals and land bridges: Journal of the Washington Academy of Science. XXX. p. 137-163. ____ Types in modern taxonomy: American Journal of Science, v. CCXXXVII1. p. 413-431. ____ Studies on the earliest primates: American Museum of Natural History Bulletin LXXVI1. p. 185-212. ____ Antarctica as a faunal migration route: Proceedings of the Sixth Pacific Science Congress. 1939. p. 755-768. 1941 The role of the individual in evolution: Journal of the Washington Academy of Science. XXXI. p. 1-20. ____ The affinities of the Borhyaenidae: American Museum of Natural History Novitates. no. 1118, p. 1-16. ____ Large Pleistocene felines of North America: American Museum of Natural History Novitates. no. 1149. p. 1-16. ____ Range as a zoological character: American Journal of Science, v. CCXXXIX. p. 785-804. 1942 The beginnings of in North America: Proceedings of the American Philosophical Society, v. 86. p. 130-188. 1943 The discovery of fossil in North America: Journal of Paleontology, v. 17, p. 26-38. ------Mammals and the nature of continents: American Journal of Science, v. 241. p. 1-31. ------Criteria for genera, , and in zoology and paleozoology: Annals of the New York Academy of Science, v. 44. p. 145-178. 1944 Tempo and mode in evolution: Columbia University Press. New York. 237 p. 1945 Symposium on age of the distribution patterns of the arrangements in Drosophila pseudoobscura. (3) Evidence from fossils and from the application of evolutionary rate distribution: Lloydia, v. 8. p. 103-108. ____ The principles of classification and classification of mammals: American Museum of Natural History Bulletin 85, p. 1-350. 1946 Tertiary land bridges: Transactions of the New York Academy of Science, ser. 2. no. 8. p. 255-258. ____ Fossil penguins: American Museum of Natural History Bulletin 87. p. 100. 1947 The problem of plan and purpose in nature: The Science Monthly, v. 64. p. 481-495. ____ Holarctic mammalian faunas and continental relationships during the Cenozoic: American Museum of Natural History Bulletin 88. p. 613-688. ____ Evolution, interchange, and resemblance of the North American and Eurasian mammalian faunas: Evolution, v. 12. p. 218-220. 1948 The Eocene of the San Juan Basin. New Mexico: American Journal of Science, pt. 2. v. 246. p. 257-282. and p. 363-385. ____ The beginnings of the age of mammals in South America: American Museum of Natural History Bulletin 91, p. 1-232. 1949 in animals, in Genetics, paleontology and evolution. Jepsen. G. I... Mayr. E.. and Simpson. G. G.. eds.: Princeton University Press. Princeton. New Jersey, p. 205-228. ____ The meaning of evolution: Press. New Haven and London. 364 p. 1951 Horses: Oxford University Press. New York. 247 p. 6 T H F CiFOI OOICAI SOCIETY O F A M FR IC A

1951 The : Evolution, v. 5. p. 285-298. 1952 Probabilities of dispersal in geological time, in The problem of land connections across the south Atlantic with special reference to the Mesozoic, A symposium: American Museum of Natural History Bulletin 99. p. 163-176. 1953 Life of the past: Yale University Press. New Haven and London, 198 p. ____ The major features of evolution: Columbia University Press, New York. 434 p. ____ Evolution and geography: Condon Lectures. Oregon State of Higher Education. Eugene. Oregon. 63 p. 1956 Zoogeography of West Indian land mammals: American Museum of Natural History Novitates. no. 1749. p. 1-28. 1957 Australian fossil penguins, with remarks on penguin evolution and distribution: Records of the South Australian Museum, v. XIII. p. 51-70. 1958 Behavior and evolution, in Behavior and evolution. Roc. A., and Simpson. G. G.. eds.: Yale University Press. New Haven, p. 507-535. 1959 Anatomy and morphology: Classification and evolution 1859 and 1959: Proceedings of the American Philosophical Society, v. 103. no. 2. p. 286-306. Mesozoic mammals and the polyphyletic origin of mammals: Evolution, v. 13. p. 405-414. _ The nature and origin of supraspecific taxa: Cold Spring Harbor Symposia on Quantitative Biology, v. 24. p. 225-271. 1960 (with Anne Roe and Richard Lewontin) Quantitative zoology (revised edition): Harcourt Brace and Company. New York. 440 p. ____Diagnosis of the classes Reptilia and Mammalia: Evolution, v. 14. p. 388-392. ____ Man's evolutionary future: Zoologische Jahrbücher. Bd. 88. p. 125-134. 1961 Principles of animal taxonomy: Columbia University Press, 247 p. ____ Lamarck. Darwin and Butler: Three approaches to evolution: The American Scholar, v. 30, p. 238-249. 1964 This view of life. The world of an evolutionist: Harcourt, Brace and World. Inc.. New York. 308 p. ____ Organisms and molecules in evolution: Science, v. 146. p. 1535-1538. 1965 The geography of evolution. Collected essays: Chilton Books. Philadelphia and New York. 249 p. 1966 The biological nature of man: Science, v. 152. p. 462-478. 1969 The first three billion years of community evolution, in Diversity and stability in ecological systems: Brookhaven Symposia in Biology, no. 22. p. 162-177. 197! A review of the pre-Pliocene penguins of New Zealand: American Museum of Natural History Bulletin 144. 223 p. 1972 Conspectus of Patagonian fossil penguins: American Museum of Natural History Novitates. no. 2488. p. 1-37. 1975 Recent advances in methods of phylogenetic inference, in Phylogeny of the primates, a multidisciplinary approach. I.uckett. U. P.. and S/aley. F. S.. eds.: Plenum. New York. 349 p. 1978 Concession to the improbable, an unconventional autobiography: Yale University Press. New Haven and London. 291 p. 1980 Splendid isolation. The curious history of the South American mammals: Yale University Press, New Haven and London. 226 p. 1983 Fossils and the history of life: Books. Inc.. New York. 239 p. 1984 Discoverers of the lost world: Yale University Press. New Haven and London. 222 p.

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