AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 97:207-211 (1995)

Obituary: Warren Glenford Kinzey (1935-1 994)

A.L. ROSENBERGER Department of Zoological Research, National Zoological Park, Smithsonian Institution, Washington, DC 20008-2598

Warren Kinzey died peacefully during the of academe. That magic was not reserved for early hours of October 1, 1994, at home in his students alone. Warren was always Tarrytown, New York. Many of us knew this scouting talent, advising, cajoling, and en- was coming. Warren was suffering from couraging anyone with like interests to amyotrophiclateral sclerosis (ALS),LouGeh- carry out a project. rig‘s disease. His brother-in-law, Steven, at Warren’s magnetism as a teacher derived Warren’s side until the final moments along from an effervescent mix of scholarship and with Julie Kelly, Warren’s wife, said Warren charm. For decades, his youthful vigor, went to the last as he did everything, with warmth, and generosity drew students of the dignity of the great teacher that he was. many kinds and calibers into his office, Warren’s life was as full as he could possi- where the real teaching would begin. Under- bly fill it. Hours before the onset of the bout graduates and beginning graduate students of pneumonia that hospitalized him during were especially welcome. He invited all into the final week of his life, Warren was on the his home for late-term parties around phone to Eric Delson scheduling a trip in to Christmas, where he proudly showed off Manhattan. He planned to hear a talk by family, played a mean game of table tennis, one his own Ph.D. students, upon her return entertained with slide shows, and intro- from thesis research in Madagascar. A duced us to the lore and personalities-he month before, he completed a new manu- seemed to know everyone-of physical an- script on saki monkeys co-written with thropology. There we saw his personal di- Marilyn Norconk. Last June he shipped me mension, his love for singing, his passion for 16 long chapters on the natural history of ice dancing. Alas, it was from his wheelchair New World monkeys, a -by-genus ac- that Warren watched the medal rounds of count, for a book he was editing. Warren his last Winter Olympics together with a would soon call me back to relate proudly small group of friends. Always teaching, he that “New World : Ecology, Behav- marveled as he explained the triple axle ior and Evolution” had gone off to the pub- jump, dissecting it with the knowledge of a lisher. All that activity was made possible by functional morphologist and the exuberant a devoted entourage of family, friends, and spirit of a fan. He noted how this maneuver, graduate students, caring for him and work- unthinkable when he skated competitively, ing with him as his muscles wasted from is now possible because progress in the ana- disease. Most of all it was Julie Kelly who tomical sciences enabled an extension of enabled Warren to continue the enjoyment species-specific, human locomotor perfor- of travelling, writing, meeting with stu- mance. The irony was felt by all, for science dents, spending time with his family, and could do little to preserve Warren’s own bio- leaving all, including himself, prepared for mechanics as the disease progressed. death% untimely truth. Warren would have Warren was determined to be an anato- been 59 years old on the last day of October, mist when he arrived at the University of 1994. Chicago’s anthropology program in 1957, Warren is remembered for his pioneering but the discipline was then still undeveloped research on New World monkeys as well as there. Clark Howell, then at Chicago, re- his genius as a mentor. He inspired and nur- called Warren in his early twenties, “coming tured many young people and guided them down from Minnesota” where he had done into productive careers, both in and outside his bachelor’s and master’s degrees. Intel-

0 1995 WILEY-LISS, INC. 208 A.L. ROSENBERGER lectually critical and stubbornly determined locomotion. The papers appeared in a spe- to pursue his own path, Warren avoided the cial volume of the AJPA in 1967, which he impression of arrogance with his “boyish edited. look, great big grin and infectious smile,” the While writing up his thesis work during expression that many of us in the profession the late 1960s and into the 1970s, Warren came to know. He soon moved on to the Uni- was establishing a family and a teaching ca- versity of California at Berkeley, to follow reer, and beginning to seed primatology and Sherry Washburn and to complete his Ph.D. paleoanthropology with a diverse new gen- in anatomy, which he received in 1964. War- eration of scholars. As he sought to build an ren’s thesis research on the endocrinology of anthropological research identity, the tug of reproduction in rats was a topic of conve- hominid evolution and fossil primates led nience, but it also became a foundation of his back to dentition after a ten-year hiatus. He lifelong interest in dietary adaptation. Like- published two interesting, related papers in wise, in the first phase of his research career the journal Nature early in the 1970s. One he worked several fronts in disparate disci- quantified the evolutionary rates of canine plines-human variation, func- crown size reduction in hominids. The other tional morphology, endocrinology, hominids, examined jaw shape in primates as a back- biometrics, allometry-exploiting his broad ground to the diagnosis of Hominidae, a background in anatomy, anthropology, zool- project stimulated by the issues surround- ogy, and biostatistics while gradually focus- ing “Ramapithecus”and its supposedly par- ing on the dentition as a main research spe- abolic dental arcade. In doing this work, cialty. Warren recognized that another primate Although he was widely admired for his presented that rare combination of small ca- influence on the growth of New World mon- nines and parabolic jaws, the genus Callice- key studies, 12 years of publishing were to bus. The wheels began to turn and in several elapse before Warren’s first platyrrhine pa- years Warren would be off to Amazonian per appeared, a modest little study quantify- , seeking to learn more about a species ing the lack of sexual dimorphism in in which small canines, a non-aggressive de- canine teeth. Nonetheless, one pa- meanor, and a monogamous social organisa- per of that earlier period stands out as a tion were coupled adaptively. Researching quiet, highly original prelude to what would this pattern eventually brought forth a radi- become a pillar of Warren’s scientific suc- cal alternative to the Washburn-Devore cess. It was under the tutelage of the inspir- open-country baboon model of human ori- ing dentist and odontologist, Albert Dahl- gins, radical in terms of its ecological and berg, working in a basement laboratory at sociobiological implications, as well as its the University of Chicago, that Warren’s in- taxonomic derivation. I imagine Warren terest in teeth began to form. Decades later, also found delight in delivering an alterna- Warren would be doubly amused that his tive caricature of the archetypical early long-forgotten first “tooth paper,” published hominid male as a cooperative, nurturing, in 1962 with Dahlberg, was not only an ex- paternalistic figure within an intimate fam- perimental study on microscopic tooth ily-group setting, one not given to the vehi- wear-actually, the effects on Warren of cles of ferocious bluff and gender-based chewing sand-but also that it was written chauvinism. in French. Warren’s career crystallized after his ar- Warren’s debut as a force in primatology rival at the City College of New York in occurred in 1965, when he was an assistant 1970. He was thrilled to be among a critical professor of anthropology, zoology, and mass of physical anthropologists united by a anatomy at the University of California at convenient subway system. His circle of Davis. In a brilliant reading of the contem- friends and fellow scholars in New York City porary research scene, he convened prima- included Cliff Jolly, Fred Szalay, Eric Del- tology‘s first major thematic symposium in son, Ian Tattersall, Bob Sussman, Ralph the United States. The topic was primate Holloway, and Pat Luckett; up at Yale, El- WARREN GLENFORD KINZEY (19351994) 209 wyn Simons, David Pilbeam, and Alison eaters hypothesis. In this 1974 paper, Richards; and on Long Island, Jack Stern, “Ceboid models for the evolution of the hom- Randy Susman, and John Fleagle, who were inoid dentition,” Warren showed that proto- assembling an anatomy faculty with intel- hominids may have been hard-object fruit lectual roots tracing back to Chicago and eaters living in the trees. Warren’s own deep interest in locomotion. Warren was drawn deeply into fieldwork Warren was quick to help found the New to solidify the morphology-behavior inter- York Regional Primate Group to capitalize face from 1974 onward, with encouragement on this diverse, expanding brain trust and, from his old friend Bob Sussman. He made most of all, to offer students a stage to wit- Peru and then his summer and mid- ness and join at the exchange of ideas among winter destinations in a quest to study dif- professionals. City University of New York ferent species of the monogamous Callice- (CUNY) has since hosted a monthly Thurs- bus in as many habitats as possible. day evening seminar on primatology and pa- Social organization, feeding behavior, and leoanthropology, possibly the longest-run- dental morphology nourished Warren’s first ning gathering of its kind in the country. questions, but his appetite for projects con- The recent formation of the NYCEP consor- tinued to grow, for he read broadly, collabo- tium owes much to the seeds of discussion rated easily, and was always interested in Warren helped sow 20 years ago. new ideas and people. As the study of pri- Warren’s deepening commitment to mate locomotion gained momentum in the platyrrhines began in the early 1970s, when mid-l970s, Warren was ready with data on New World monkeys were an entirely open the positional behavior of and Ce- field of study. They offered an essentially buella. When the Pleistocene refugium hy- unknown radiation as a context for forging pothesis gained attention, Warren, a stick- his personal synthesis of S.L. Washburn’s ler for details, an adroit mapmaker, and now teachings, which inverted primatology’s pre- travelling widely in South America, was disposition for the science of bodies over be- keen to follow up its implications for under- haviors. Washburn emphasized the living standing the geographic distribution of as the primary information base of platyrrhines. As vocal playbacks developed functional anatomy, rather than the bones into a field tool, Warren investigated the so- and muscles by themselves. Warren blended cial and ecological meaning of the booming this view with the evolutionary perspective long calls of titi monkeys. of the anatomist Le Gros Clark, whose work During the last 10 years of his life, in the he admired, especially Clark‘s ideas of evo- buoyant years of what should have been lutionary trends and total morphological midlife, Warren returned to the nagging pattern. questions he left unanswered decades be- Warren’s two decades of research and fore. With Marilyn Norconk, he began an writing on platyrrhines was an artfully intense program of fieldwork exploring the scripted programme. The New World mon- adaptive significance of food hardness and keys demanded study because they are in- biochemistry in the little-known saki and trinsically interesting; because of their di- uakari monkeys. The methods they devel- dactic significance for hominid evolution; oped to test the physical properties of the because fieldwork on them could be accom- foods primates eat are becoming standard plished at relatively low cost; and because tools of the fieldworker’s kit. That Warren the study of non-human primates needed to would press this issue some 15 years after be legitimized as part of anthropology if pri- he first mulled it over in the literature testi- matology, as a discipline, was to be sustain- fies to the determination Clark Howell saw able in the social sciences and provide teach- in the same man 30 years younger. ing jobs for the students that would be While managing this benchmark series of coming out. His research agenda paid off field studies from afar, Warren did a two- quickly and well. One of warren’s early in- year stint in Washington, DC, as program sights led to an extension of Jolly’s seed- director of physical anthropology at the Na- 210 A.L. ROSENBERGER monkeys. Reluctant at first to be singled out this way, Warren eventually relinquished his modesty. Although he was then very weak and almost immobile, Warren enjoyed the meeting thoroughly. He mused about its significance for the field and the potential impact of this growing cadre-a mix of gen- erations and disciplines-talking to one an- other about platyrrhines. Always looking forward, Warren asked if we could possibly do this annually, without a sliver of regret that he would not make the next one. He knew that knowledge evolved and that peo- ple, its agents of change, needed to be en- couraged always. And that he always did.

A SELECTED BIBLIOGRAPHY OF WARREN G. KINZEY Brozek J and Kinzey WG (1960) Age changes in skinfold compressibility. J. Gerontol. 15(1):45-51. Dahlberg AA and Kinzey WG (1962) Etude micro- scopique de l’abrasion et de l’attrition sur la surfaces des dentes. Bull. Groupe. Int. Recherche Sci. Stoma- tol. 5(2):242-251. Garber PA and Kinzey WG (1992) Feeding adaptations in New World Primates: an evolutionary perspective. Am. J. Phys. Anthropol. 88:411-413. Kinzey WG and Srebnik HH (1962) Maintenance of Fig. 1. Warren Glenford Kinzey (1935-1994). pregnancy in protein-deficient rats with short-term injections of ovarian hormones. Proc. SOC.Exp. Med. 114t158-160. Kinzey WG (1967) Preface. Symposium on primate loco- tional Science Foundation. In this capacity, motion. Am. J. Phys. Anthropol. 26(2):115-118. he continued to promote primatology with Kinzey WG (1968) Hormonal activity of the rat placenta in the absence of dietary protein. Endocrinology8f2)2): wisdom and zeal and managed to signifi- 26C-270. cantly enlarge support for student research. Kinzey WG (1969) Tubal transport of fertilized ova in Those of us who knew Warren well under- protein deficient rats. Endocrinology 84(5):126& stood the joy he took in this and why he 1267. considered it one of his finest personal ac- Kinzey WG (1969) The effect of protein deficiency on complishments. uterine sensitivity in the pseudopregnant rat. Acta Endocrinologica61(2);232-238. In 1990, shortly after Warren left DC to Kinzey WG (1970) Maintenance of pregnancy in protein- return to CUNY and City College, the mys- deficient rats with dietary supplements of methio- terious early symptoms of ALS began to ap- nine. Roc. SOC.Exp. Biol. Med. 133(2):449451. pear. True to his character-inquisitive, de- Kinzey WG (1970) Endocrine function of the placenta in termined, and consistant-Warren made thiamine-deficient rats. Proc. SOC.Exp. Biol. Med. ALS another research project, and the sub- 134U):l-l. Kinzey WG, Sands L, and Bonds P (1970) Maintenance ject of death was added to his teaching rep- of pregnancy in absence of dietary protein with pro- ertoire. All along he spoke candidly about gesterone. Proc. SOC.Exp. Biol. Med. 13#1):72-73. the physical and emotional aspects of his Kinzey WG (1970) Rates of evolutionary change in the experience, particularly with his medical hominid canine teeth. Nature 225:296. students. Kinzey WG (1970) The basic rectangle of the mandible. In February, 1994, some sixty of us gath- Nature 228:28%290. Kinzey WG (1971) The evolution of the human canine ered at the Smithsonian’s National Zoologi- tooth. Am. Anthropol. 3(3):680-694. cal Park in Washington, DC, to honor War- Kinzey WG (1971) Male reproductive system and sper- ren with a two-day conference on New World matogenesis. In ESE Hafez (ed.): Comparative Repro- WARREN GLENFORD KINZEY (1935-1994) 211 dudion of Nonhuman Primates. Springfield: Charles Kinzey WG (1984)The ecological role of the Callitri- C. Thomas, pp. 85-114. chidae. Am. J. Phys. Anthropol. 64:419449. Kinzey WG (1972)Canine teeth of the monkey Callice- Kinzey WG (1984)The dentition of the pygmy chimpan- bus: lack of sexual dimorphism. Primates 13(4):365- zee. In RL Susman (ed.): The Pygmy Chimpanzee. 369. Evolutionary Biology and Behavior. New York: Aca- Kinzey WG (1973)Reduction of the cingulum Ceboidea. demic Press, pp. 65-88. In MR Zingeser (ed.): Symp. IVth Intern. Congr. Pri- Kinzey WG (1984)Titi monkey family life. In DW Mac- matol., vol. 3. Basel: S. Karger, pp. 101-127. donald (ed.): The Encyclopedia of . Oxford: Kinzey WG (1974) Ceboid models for the evolution of Equinox, pp. 36C367. hominoid dentition. J. Hum. Evol. 3:193-203. Kinzey WG (1986)New World primate studies: What’s Kinzey WG, Rosenberger AL, and Ramirez M (1975) in it for anthropology? Ann. Rev. Anthropol. 15121- Vertical clinging and leaping in a neotropical anthro- 148. poid. Nature 255:327-328. Kinzey WG and Easley SP (1986)Territorial shift in the Kinzey WG, Rosenberger AL, Heisler PS, Prowse DL, yellow-handed titi monkey. Am. J. Primatol. 11:307- and Trilling JS (1977)A preliminary field investiga- 318. tion of the yellow handed titi monkey, Callicebus tor- Kinzey WG (ed.) (1987)Primate Models for the Evolu- quatus torquatus in northern Peru. Primates 18:159- tion of Human Behavior. Albany: State University of 181. New York University. Kinzey WG (1977)Diet and foraging behavior of Callice- Kinzey WG (1987) Introduction. In WG Kinzey (ed.): bus torquatus. In TH Clutton-Brock (ed.): Primate Primate Models for the Evolution of Human Behavior. Ecology: Studies of Feeding and Ranging Behavior in Albany: State University of New York University, pp. Lemurs, Monkeys and Apes. London: Academic Press, vii-Wi. pp. 127-151. Kinzey WG (1987)A primate model for human mating Kinzey WG (1977)Positional behavior and ecology in systems. In WG Kinzey (ed.): Primate Models for the Callicebus torquatus. Yrbk. Phys. Anthropol. 20:46& Evolution of Human Behavior. Albany: State Univer- 480. sity of New York University, pp. 105-114. Kinzey WG (1978)Feeding behavior and molar features Kinzey WG, Norconk MA, and Alvarez-Corder0E (1988) in two sympatric species of titi monkey. In DJ Chivers Primate survey of eastern Bolivar. Primate Conserva- and J Herbert (eds.): Recent Advances in Primatology. tion 9:66-70. vol. 1, Behaviour. London: Academic Press, pp. 373- Kinzey WG and Norconk MA (1990)Hardness as a basis 385. of fruit choice in two sympatric frugivorous primates. Kinzey WG and Gentry AH (1978)Habitat utilization in Am. J. Phys. Anthropol. 81.5-15. two species of Callicebus. In RW Sussman (ed.): Pri- Kinzey WG (1992)Dietary and dental adaptations in the mate Ecology: Problem-Oriented Field Studies. New . Am. J. Phys. Anthropol. 88149-14. York: John Wiley, pp. 89-100. Kinzey WG and Norconk MA (1993)Physical and chem- Stark N, Kinzey WG, and Pawlowski P (1980)Soil fertil- ical properties of fruit and seeds eaten by Pithecia and ity and distribution. Proc. Symp. Tropical Chiropotes in Surinam and . Am. J. Prima- Ecology, Kala Lumpur: Tropical Development. pp. tol. 14:207-227. 101-111. Mittermeier RA, Kinzey WG, and Mast RB (1989)Neo- Kinzey WG (1981)The titi monkey, genus Callicebus. In tropical primate conservation. J. Hum. Evol. 18.597- A-F Coimbra Filho and RA Mittermeier (eds.): Ecol- 610. ogy and Behavior of Neotropical Primates, vol. 1. Rio Norconk MA and Kinzey WG (1994)Challenge of neotro- de Janeiro; Brasilian Academy of Sciences, pp. 241- pica1 frugivory: travel patterns of spider monkeys and 276. bearded sakis. Am. J. Primatol. 34:171-18. Kinzey WG (1982) Distribution of primates and forest Kinzey WG and Cunningham EP (1994)Variability in refuges, In GT Prance (ed.): Biological Diversification platyrrhine social organization. Am. J. Primatol. 34: in the Tropics. Proceedings of the Fifth International 185-198. Symposium of the Association for Tropical Biology. Peetz A, Norconk MA, and Kinzey WG (1992)Predation New York University Press, pp. 455482. by a jaguar on howler monkeys (Alouatta senzculus)in Kinzey WG and Wright PC (1982)Grooming behavior in Venezuela. Am. J. Primatol. 28:223-228. the titi monkey, Callicebus torquatus. Am. J. Prima- Rosenberger AL and Kinzey WG (1976)Functional pat- tol. 3:267-275. terns of molar occlusion in platyrrhine primates. Am. Kinzey WG and Robinson JG (1983) Intertroop loud J. Phys. Anthropol. 45t281-298. calls, home range size and spacing in Callicebus tor- Robinson JG, Wright PC, and Kinzey WG (1987)Monog- quatus. Am. J. Phys. Anthropol. 60:539-544. amous cebids and their relatives. In BB Smutts, DL Kinzey WG and Becker M (1983)Activity pattern of the Cheney, RM Feyfarth, RW Wrangham, TT Struh- masked titi, Callicebus personutus. Primates 24:337- saker (eds.): Primate Societies. Chicago: University of 343. Chicago Press, pp. 44-53.