Oliver P. Pearson: scientist, statesman, gentleman

Douglas A. Kelt, Enrique P. Lessa, Jorge Salazar-Bravo, and James L. Patton

Oliver Payne Pearson was born on 21 October 1915, in Philadelphia, Pennsylvania, to Olive Payne Corning Pearson and Forrest Garfield Pearson. He was the third surviving son (a first son had died in infancy), and his mother, despairing of not having a daughter, gave him her name, slightly modified (5 years later she finally did have a daugher). Oliver P. Pearson passed away 4 March 2003, leaving the world poorer in his absence. In his 87 years he became a decisive force promoting our understanding of mammalian biology, with notable advances in physiological and reproductive , predator- prey interactions, , zoogeography, and perhaps most notably to some, the general biology of small in temperate and Andean South America. Oliver Pearson, “Paynie” to many that knew him, graduated from Swarthmore College in 1937. He attended Harvard University for both his M.S. (1939) and Ph.D. (1947), and was then hired by UC Berkeley as an Instructor in (1947-48), entering the ranks of the tenured series the following year. He was an Assistant Professor of Zoology (Dept. of Zoology) and Assistant Curator of Mammals (Museum of Vertebrate Zoology, MVZ) from 1949 to 1955, gaining tenure and advancing to Associate Professor of Zoology and Associate Curator of Mammals in the MVZ in 1955. In 1957 he resigned his tenured position, maintaining status at UC Berkeley as a Lecturer in Zoology and as a Research Associate at the Museum. This allowed him to focus his attentions on his studies of voles and predation (see below), and to spend a year as a Visiting Professor of Ecology at the University of Buenos Aires (1964- 65). When Alden Miller, Director of MVZ, died in 1965, Paynie returned to the tenure series, this time as full Professor of Zoology (which he retained until his real and very active retirement in 1971), and as Acting Director of MVZ (1966-67) and then Director (1967-71). Additionally, Pearson served as Vice Chairman of the Zoology Dept. in 1953-54, and then as Acting Chairman in 1968. He was never highly fond of these administrative roles, as they detracted from the time he had to dedicate to fieldwork and writing. In 1944 he married Anita Kelley, who was to be his companion in all his subsequent travels. When awarded an honorary doctoral degree (“Doctor Honoris Causa”) from

Pp. xx-xx in Kelt, D. A., E. P. Lessa, J. Salazar-Bravo, and J. L. Patton (eds.). xxxx. The quintessential naturalist: honoring the life and legacy of Oliver P. Pearson. University of California Publications in Zoology xx:xx-xx.

  University of California Publications in Zoology the University of La Plata, Pearson famously stated “Yo soy un simple atrapador de ratones, y nada hubiese sido posible sin Anita” (“I am a simple mouse-trapper, and nothing would have been possible without Anita”) (UC Berkeley News, 7 March 2003). They had four children, Carol Ann (1947), Peter Kelley (1949), Sandia Coming (1956), and Alison Payne (1958), and toted them along on subsequent field trips. Oliver Pearson was an early convert to science; he set his first trapline at age 8 or 10, capturing a short-tailed shrew (Blarina brevicauda; see Pearson, 1982). As a Research Assistant in the Philadelphia Academy of Natural Sciences, Paynie participated in 2 trips to Panama with his undergraduate mentor, Robert K. Enders. They published on the taxonomy of small mammals from these trips (Enders and Pearson, 1939, 1940; Pearson 1939) while pursuing numerous studies on the reproductive of North American mustelids, canids, and other groups (e.g., Pearson and Enders, 1943a, 1943b, 1944; Enders and Pearson, 1943a, 1943b; Pearson 1944 ; Pearson and Bassett 1944; Enders et al. 1946; Pearson and Bassett, 1946). At Harvard, collaboration with his fellow student, Peter Morrison, led to a paper in Science, on “The metabolism of a very small ” (Morrison and Pearson, 1946). A subsequent paper on this topic (Pearson, 1947) was later recognized as a “Citation classic” in Current Contents (Current Contents (Agriculture, Biology, & Environmental Science) 35:16 (27 August 1984)). The field trips to Panama may have whetted Pearson’s appetite forthe Neotropics, as he pursued subsequent fieldwork in Peru (1939-40, 1946, 1952, 1955, 1967, 1972), Colombia (1950), and Argentina (1955, 1964-65, then annually 1978-1999). Work in the Peruvian Andes was productive and fertile ground for an energetic young field biologist. Essentially uncharted terrain, the mammals of this region were known only moderately well taxonomically, but poorly at best in ecologic terms. Pearson’s long-standing interest in tuco-tuco’s (Ctenomys) began with a description of the species known from Peru (Sanborn and Pearson, 1947), followed 13 years later by a short monograph on Peruvian tuco’s (Pearson 1959[1960]); his long-standing interest in this group was memorialized with the naming of Ctenomys pearsoni (Lessa and Langguth, 1993; see Tomasco and Lessa, this volume). Additionally, 1 species of Andalgalomys (A. pearsoni (Myers, 1977)) and 1 genus (Pearsonomys Patterson, 1992) have been named in recognition of Oliver Pearson. Meanwhile, Pearson pursued groundbreaking work on the ecology and reproductive biology of viscachas (Lagidium viscacia; Pearson, 1947, 1948); at the time this was only the second extensive study on any hystricomorph , and for this study the Pearson’s camped at 16,000 ft in the Caccachara Valley (Fig. 1, 2, 3). Pearson later (1951) synthesized his observations on highland mammals. Upon returning from his first Andean trip in 1940, he “found it impossible to identify many specimens of Phyllotis”; he “soon formed the opinion that the real number of species was much less than indicated by the existing nomenclature” and that “a revision of the genus therefore seemed desirable.” He embarked on this daunting task, and in 1958 he published the definitive treatment of the genus (Pearson, 1958:quotations from p. 394). Pearson also wrote a paper in 1975 on “an outbreak of mice” that set the bar for rigor in natural history observation; all subsequent studies of rodent outbreaks (“ratadas”) have followed Pearson’s lead in this regard. Thus, in the span of just a couple of decades, Pearson reported on natural history, population ecology, reproductive biology, biogeography, and . Few are the contemporary biologists with such breadth! Although this volume emphasizes mammals, it is worth noting that Oliver Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 

Figure 1. Campsite in Caccachara Valley, Dept. Puna, Peru. 1946. Photo courtesy of Anita Pearson.

Figure 2. Oliver Pearson holding a Figure 2. Anita Pearson in the Peruvian home-made livetrap. Caccachara Valley, Andes, 1946. Peru, 1946.  University of California Publications in Zoology

Pearson also published seminal papers on birds and herpetiles during this period. This included papers on hummingbird ecology and metabolism (5 papers), flight speeds in Allen’s hummingbird (Pearson, 1960a), torpidity in small birds (Pearson, 1960b), tinamou (Nothoprocta) natural history and reproduction (Pearson and Pearson, 1955), and spacing behavior in sparrows (Zonotrichia; Pearson 1979). He also reported on lizard ecology (Pearson, 1954) and thermoregulation (Pearson, 1977, Pearson and Bradford, 1976), and he included reptiles and amphibians in his well-known Andean altitudinal gradient (Pearson and Ralph, 1978). When Paynie and Anita turned their attention to southern Argentina, they channeled their energies to what was then a very poorly known fauna. Although Osgood (1943) and Mann (1978) had provided thorough treatments of Chilean mammals, the ecology and natural history of mammals of Patagonia and the Valdivian temperate rainforest region were known more from anecdotes than science; this was soon to change, in large part through the efforts of the Pearson’s, although their attention here was rapidly followed by others, both in Chile and Argentina. The Pearson’s obtained an apartment in Bariloche, which they used as a base for biennial trips to study the small mammals both in Valdivian temperate rainforest and Patagonian steppe. Following the Peruvian “ratada” they observed in the 1970s, they were receptive to the importance of major food pulses on wild small mammal populations, and they quickly turned their attention to the local bamboo, Chusquea culeo, documenting both the growth and population biology of this species (Pearson et al., 1994) while waiting for the next flowering. The waiting proved to be a standing topic of humor for Paynie (see comments in Mastozoología Neotropical 10:195-205), but the mass flowering finally occurred in 2000 (Pearson, 2002), and the remarkable impacts on small mammals are documented in this volume (Sage et al., 2006). During the lengthy wait for the bamboo, Paynie and Anita made tremendous strides towards understanding all facets of this fauna, from basic field identification (Pearson, 1995) and natural history (Pearson, 1983, 1984) to ecology (Pearson and Pearson, 1982), reproductive and foraging ecology (Pearson, 1988, 1992; Pearson and Pearson, 1989), demography and population biology (Pearson et al., 1987), community and biogeographic ecology (Pearson and Pearson, 1982; Pearson, 1994), systematics (Pearson, 1984; Pearson and Smith, 1999), and historical ecology (Pearson, 1987; Pearson and Christie 1993; Pearson and Pearson, 1993). In short, they did what they do so well – a fair amount of everything! During intervening periods in California, Pearson pursued his well-known studies on the effects of predators on small mammal communities, especially California voles (Microtus californicus; Pearson, 1964, 1966, 1971), reinvigorating this field with his empirical and conclusive evidence for the role of predators in population cycles. Two of these (Pearson 1966, 1971) have been extensively cited in the literature (Fig. 4). He also spearheaded one of the most comprehensive studies on bat reproductive ecology (Pearson et al., 1952); this paper has been cited at least 125 times (according to ISI; Table 1). Oliver Pearson was a gifted communicator. Robert Baker (this volume) comments on the clarity of Pearson’s oral presentations, noting that he was “fascinated with his class and style in presentations of papers” and that “he had such control over the English language and word order that many of his sentences were poetic.” The same was true of his written prose, which combined piercing knowledge of the he studied with clarity and focus, liberally infused with his well known sense of humor. Pearson’s passion for empirical studies on Neotropical small mammals, and his Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 

Oliver P. Pearson Publication Record 8 100 200 179 180 157 80 160 6 140 125 111 60 120 99 4 100

40 80 Annual Productivity Annual 60 (stars) Citations ISI 2 Productivity Cumulative 20 40

20

0 0 0 1940 1950 1960 1970 1980 1990 2000 Year Figure 4. Publication record for Oliver P. Pearson. Following pulses in the 1940s and in 1960, he maintained a consistent output throughout his life, as indicated by the monotonically increasing cumulative productivity curve. Citations, as taken from the ISI database (Thompson Scientific; http://scientific.thomson.com), are indicated with stars. willingness to help students and colleagues whenever possible, left a powerful legacy. He was not interested in trivial science, and he always strived for productive and useful research. When one of us (DAK) sent him a draft manuscript documenting chromosomal variation in Eligmodontia and confirming the separation of E. morgani and E. puerulus (Kelt et al., 1991), his primary comment was a question – although the underlying science was useful and interesting, how could a field biologist distinguish these genetic entities with a live in hand? Finally, Oliver Pearson was not only a scientist; he also was an accomplished artist. After one of his early trips to Panama he prepared a woodcut showing a field biologist conducting nocturnal censuses with a headlamp (Fig. 5). Many years later he and Anita were surprised at the dearth of scenic postcards from the Patagonian region, and Oliver proceeded to convert some of his scenic photographs to cards that he sold to merchants in Bariloche. Oliver Pearson was an honorary member of the American Society of Mammalogists (1979), the Cooper Ornithological Society (1979), the Sociedad Argentina para el estudio de los Mamíferos (1983), and the Comite Argentino de Conservación de la Naturaleza (date?). He was elected as a Fellow of the California Academy of Sciences in 1964. He served as a trustee for ASM from 1969 through 1984, helping to solidify the financial underpinnings of that society. He held a similar role on the Investment Committee for the Cooper Ornithological Society from 1972 to 1987. He also was an elected Director  University of California Publications in Zoology

Figure 5. Block print of an intrepid field biologist hunting kinkajous (Potos flavus). This print was hand-carved by Oliver Pearson in ca. 1942.

for the ASM over 17 years between 1952 and 1990, for the COS in 1969-71, and for the San Francisco Zoological Society in 1959-60. He also served as a Councillor for the Save the Redwoods League. He served as Vice President for ASM from 1969 to 1972. Reflecting his years of dedication to the ASM, he received the Hartley H.T. Jackson Award “for long and outstanding service to ASM” in 1984. And, as noted above, Pearson was awarded an honorary doctoral degree from the University of La Plata in 2000, as much for his contributions to the knowledge of the fauna of Argentina as for his seminal role in the development of ecological and mammalogical science in that country. The following year, he was honored at the First Biennial Meeting of the Ecological Societies of Argentina and Chile (Bariloche, 2001); as Adrian Monjeau put it (pers. comm. 2006), this was “a public recognition as one of the ‘fathers’ of Ecology in Argentina.”

Concluding comments. Oliver Pearson was the field biologist’s field biologist. He combined a keen intellect with a probing curiosity. He also knew how to “think like a Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 

Table1. Publications by Oliver Pearson with > 50 citations in the ISI database (Thompson Scientific; http://scientific.thomson.com), arranged in chronological order.

No. Article Citations (ISI) Pearson, O. P. 1944 Reproduction in the shrew (Blarina brevicauda Say). American 111 Journal of Anatomy 75:39-93. Pearson, O. P. 1947 The rate of metabolism of some small mammals. Ecology 179 28:127-145. Pearson, O. P. 1948 Metabolism of small mammals, with remarks on the lower 73 limit of mammalian size. Science 108:44. Pearson, O. P., M. R. Koford, and A. K. Pearson 1952 Reproduction of the lump-nosed bat (Corynorhinus rafinesquei) 125 in California. Journal of 33:273-320. Pearson, O. P. 1954 Habits of the lizard Liolaemus multiformis multiformis at high 68 altitudes in southern Peru. Copeia 2:111-116. Pearson, O. P. 1963 History of two local outbreaks of feral house mice. Ecology 75 44:540-548. Pearson, O. P. 1966 The prey of carnivores during one cycle of mouse abundance. 157 Journal of Animal Ecology 35:217-233. Pearson, O. P. 1971 Additional measurements of the impact of carnivores 99 on California voles (Microtus californicus). Journal of Mammalogy 52:41-49. Pearson, O. P., and D. F. Bradford 1976 Thermoregulation of lizards and toads at high altitudes in 62 Peru. Copeia 1976:155-170 Pearson, O. P., and J. L. Patton 1976 Relationships among South American phyllotine 51 based on chromosome analysis. Journal of Mammalogy 57:339-350  University of California Publications in Zoology

Oliver Pearson reviewing papers at his home office, 1979.

mouse” as the following vignette illustrates. Pearson once co-taught a brief course on small mammal ecology at the Universidad Austral (Valdivia, Chile). One participant of this course (Ricardo Martínez) provided the following recollection:

“We [students] set 400 live traps . . . to get some mice for preparing study skins. Back at camp, I was lucky to find that Dr. Pearson was just leaving to have a walk, thus I asked him if I was welcomed and the gentleman of course told me that was OK. We talked about everything and he asked me if we had set the traps. He explained that during summer it was rather difficult to get some mice, but just in case he was thinking of setting some. Suddenly, he stopped at some quila [bamboo] near the trail and he took from his parka pocket a small Museum Special trap and got some oats from the other pocket, and rather embarrassed he spit on the oats and showed me how to prepare the bait, and he set the trap between the canes. We continued walking and then he stopped and began to search with his hands on the grass along the trail, he showed me my first mouse “runway”; he set the trap, and we continued walk- ing. He set eight Museum Special traps. Early the next day we [students] checked our 400 Sherman traps to retrieve just one chucao [a bird]. Feeling pretty battered we went back to camp just to found that Dr. Pearson got FIVE mice: 2 A. olivaceus, 1 A. longipilis and 2 O. longicaudatus...(63% catch success).”

To catch mice, one needs to think like mice, and in this regard, Pearson had few peers. He was driven by a passion to decipher how they lived, how they died, and Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 

Cartoon by Alison Pearson, depicting her father’s tireless pursuit of small mammals, bamboo, and the interactions among them.

what they did in the intervening times. He eagerly shared his knowledge with all takers, and his influence on colleagues and students alike was both enduring and endearing. Shortly after returning from Argentina in November 2000, Paynie suffered an embolism in his lower leg. A vein graft was not successful, and when given the option of another attempt vs. amputation, Paynie opted for the latter. This likely was influenced by the fact that the bamboo in southern Argentina was flowering (see Sage et al., this volume), and he desperately wanted to see this. Not long thereafter, Dick Sage offered to collect animals in the Austral fall of 2001; Paynie thanked Dick, but said “WE are going!” As Dick reported to us, “So we went, he, Anita, his daugher Ali and her friend, and myself. It was a great expedition!” (it was during this trip that Pearson was formally recognized at the joint meeting of the Argentine and Chilean Ecological Societies). Although Paynie could not navigate the woods well, the abundance of small mammals in response to the immense pulse of food (bamboo seeds) was phenomenal (see Sage et al., this volume). Dick Sage reported (pers. comm. to DAK) that “we were camped amidst the clumps of (now-dead) bamboo, and the mice were plainly visible at night in the clumps of canes over our tents. I had mice dropping down onto my tent during three of the four nights we were camped there.” This irruption comprised one snapshot in a cycle that likely will not be repeated for many decades. Paynie’s drive and dedication to observing and documenting this phenomenon was well captured in his daughter’s depiction of Paynie -- our friend, colleague, and mentor -- maintaining his diligent watch from somewhere in the heavens. Oliver Pearson passed away in March of 2003. In respect for his life’s work and interest, his ashes were spread in 3 areas that were special to him. Some of these were in the region of the Berkeley Hills where he had studied small mammals and predator- prey interactions, and some around his home in Orinda. Finally, Dick Sage carried some to Bariloche, where he and Miguel Cristie laid them to rest amongst regions of 10 University of California Publications in Zoology

Patagonia where he and Anita had labored so fruitfully. As recorded by Dick Sage and Miguel Christie (2003),

“. . . durante un remanso en medio de una fuerte tormenta propia del solsticio de invierno, tuvimos el honor es esparcir sus cenizas entre los lugares que él mas amaba. Inmendiatamente de concluida la sencilla ceremonia se levantó un fuerte viento y una procession de unos 30 cóndores sobrevoló el lugar. ¡Fue emocionante! Tras nuestra despedida, los grandes Señores de los Cielos Patagónicos honraban a otro grande con un merecido saludo de bienvenida, como diciendo: ‘Nuestra casa es tu casa’ . . . y lo será por siempre.”

Literature Cited

Enders, R. K., and O. P. Pearson 1939 Three new rodent subspecies of Scotinomys from Chirique, Panama. Notulae Naturae of the Academy of Natural Sciences of Philadelphia 34:1- 4.

Enders, R. K., and O. P. Pearson 1940 A new subspecies of Reithrodontomys mexicanus from Panama with remarks on R. mexicanus cherrii. Notulae Naturae of the Academy of Natural Sciences of Philadelphia 60:1-2.

Enders, R. K., and O. P. Pearson 1943a The blastocyst of the fisher. Anatomical Record 85:285-287.

1943b Shortening gestation by inducing early implantation with increased light in the marten. American Fur Breeder, January:18-19.

Enders, R. K., O. P. Pearson, and A. K. Pearson 1946 Certain aspects of reproduction in the fur seal. Anatomical Record 94:213- 227.

Kelt, D. A., R. E. Palma, M. H. Gallardo, and J. Cook 1991 Chromosomal multiformity in Eligmodontia (Muridae, ), and verification of the status ofE. morgani. Zeitschrift für Säugetierkunde 56:352-358.

Lessa, E. P., and A. Langguth 1993 Ctenomys pearsoni, n. sp. (Rodentia, Octodontidae), del Uruguay. Resultados Comunicados Jornadas Ciencias Naturales, Montevideo, Uruguay 3:86- 88.

Mann, G. F. 1978 Los pequeños mamíferos de Chile: marsupiales, quirópterors, edentados y roedores. Guyana (Zoología) 40:1-342. Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 11

Morrison, P. R., and O. P. Pearson 1946 The metabolism of a very small mammal. Science 104:287-289.

Myers, P. 1977 A new phyllotine rodent (genus Graomys) from Paraguay. Occasional Papers of the Museum of Zoology, University of Michigan 676:1-7.

Osgood, W. H. 1943 The mammals of Chile. Field Museum of Natural History, Zoological Series 30:1-268.

Patterson, B. D. 1992 A new genus and species of long-clawed mouse (Rodentia: Muridae) from temperate rainforests of Chile. Zoological Journal of the Linnean Society 106:127-145.

Pearson, A. K., and O. P. Pearson 1955 Natural history and breeding behavior of the tinamou, Nothoprocta ornata. Auk 72:113-127.

Pearson, A. K., O. P. Pearson, and I. A. Gomez 1994 Biology of the bamboo Chusquea culeou (Poacea: Bambusoideae) in southern Argentina. Vegetatio 111:93-126.

Pearson, O. P. 1939 Three new small mammals from eastern Panama. Notulae Naturae of the Academy of Natural Sciences of Philadelphia 6:1-5.

1944 Reproduction in the shrew (Blarina brevicauda Say). American Journal of Anatomy 75:39-93.

1947 The rate of metabolism of some small mammals. Ecology 28:127-145.

1948 Life history of mountain viscachas in Peru. Journal of Mammalogy 29:345- 374.

1951 Mammals in the highlands of southern Peru. Bulletin of the Museum of Comparative Zoology 106(3):117-174.

1954 Habits of the lizard Liolaemus multiformis multiformis at high altitudes in southern Peru. Copeia 2:111-116.

1958 A taxonomic revision of the rodent genus Phyllotis. University of California Publications in Zoology 56:391-477.

1960 Biology of the subterranean rodents, Ctenomys, in Peru. Memorias del Museo de Historia Natural ‘Javier Prado’ 9:1-56. 12 University of California Publications in Zoology

1960a Speed of the Allen hummingbird while diving. Condor 62:403.

1960b Torpidity in birds. Bulletin of the Museum of Comparative Zoology 124:93-103.

1964 Carnivore-mouse predation: An example of its intensity and bioenergetics. Journal of Mammalogy 45:177-188.

1966 The prey of carnivores during one cycle of mouse abundance. Journal of Animal Ecology 35:217-233.

1971 Additional measurements of the impact of carnivores on California voles (Microtus californicus). Journal of Mammalogy 52:41-49.

1975 An outbreak of mice in the coastal desert of Peru. Mammalia 39:375-386

1977 The effect of substrate and of skin color on thermoregulation of a lizard. Comparative Biochemistry and Physiology 58:363-358

1979 Spacing and orientation among feeding golden-crowned sparrows. Condor 81:278-285

1982 Round Table – Research Priorities in South America. P. 537 in Mammalian Biology in South America (Mares, M. A., and H. H. Genoways, eds.). The Pymatuning Symposia in Ecology, Special Publication Series, 6. Pymatuning Laboratory of Ecology, University of Pittsburgh, Pittsburgh, Pennsylvania, USA.

1983 Characteristics of a mammalian fauna from forests in Patagonia, Southern Argentina. Journal of Mammalogy 64:476-492.

1984 Taxonomy and natural history of some fossorial rodents of Patagonia, southern Argentina. Journal of Zoology 202:225-237.

1987 Mice and the postglacial history of the Traful Valley of Argentina. Journal of Mammalogy 68:469-478.

1988 Biology and feeding dynamics of a South American herbivorous rodent, Reithrodon. Studies on Neotropical Fauna and Environment 23:25-39.

1992 Reproduction in a South American mouse, longipilis. Anatomical Record 234:73-88.

1994 The impact of an eruption of Volcan Hadson on small mammals in Argentine Patagonia. Mastozoologia Neotropical 1:103-112. Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 13

1995 Annotated keys for identifying small mammals living in or near Nahuel Huapi National Park or Lanin National Park, southern Argentina. Mastozoologia Neotropical 2:99-148.

2002 A perplexing outbreak of mice in Patagonia, Argentina. Studies on Neotropical Fauna and Environment 37:187-200.

Pearson, O. P., and C. F. Bassett 1944 Size of the vulva in the fox and its relation to fertility. Anatomical Record 89:455-459.

1946 Certain aspects of reproduction in a herd of silver foxes. American Naturalist 53:45-67.

Pearson, O. P., and D. F. Bradford 1976 Thermoregulation of lizards and toads at high altitudes in Peru. Copeia 1976:155-170

Pearson, O. P., and M. I. Christie 1993 Rodent guano (amberat) from caves in Argentina. Studies on Neotropical Fauna and Environment 28:105-111.

Pearson, O. P., and R. K. Enders 1943a Duration of pregnancy in marten, mink, and related species. American Fur Breeder

1943b Your mating of foxes should be timed with great care. Black Fox Magazine, December:xxxxxxx

1943 Ovulation, maturation and fertilization in the fox. Anatomical Record 85:69-83.

1944 Duration of pregnancy in certain mustelids. Journal of Experimental Zoology 95:21-35.

Pearson, O. P., M. R. Koford, and A. K. Pearson 1952 Reproduction of the lump-nosed bat (Corynorhinus rafinesquei) in California. Journal of Mammalogy 33:273-320.

Pearson, O., S. Martin, and J. Bellati 1987 Demography and reproduction of the silky desert mouse (Eligmodontia) in Argentina. Fieldiana: Zoology, new series 39:433-446. 14 University of California Publications in Zoology

Pearson, O. P. and A. K. Pearson 1982 Ecology and biogeography of the southern rainforests of Argentina. Pp. 129-142 Mammalian Biology in South America (Mares, M. A., and H. H. Genoways, eds.). The Pymatuning Symposia in Ecology, Special Publication Series, 6. Pymatuning Laboratory of Ecology, University of Pittsburgh, Pittsburgh, Pennsylvania, USA.

1989 Reproduction of bats in Southern Argentina. Pp. 549-566 in Advances in Neotropical Mammalogy (Redford, K. H, and J. F. Eisenberg, eds.). Sandhill Crane Press, Gainesville, Florida, USA.

1993 La fauna de mamiferos pequeños de Cueva Traful I, Argentina: pasado y presente. Praehistoria 1:211-214 (PREP-CONICET, Buenos Aires, Argentina).

Pearson, O. P., and C. P. Ralph 1978 The diversity and abundance of vertebrates along an altitudinal gradient in Peru. Memoria del Museo de Historia Natural “Javier Prado” 18:1-97.

Pearson, O. P., and M. F. Smith 1999 Genetic similarity between Akodon olivaceus and Akodon xanthorhinus (Rodentia: Muridae) in Argenitina. Journal of Zoology 247:43-52.

Sage, R. D., and M. I. Christie 2003 “Mi casa es tu casa”: un homenaje a Oliver P. Pearsoni. Mastozoología Neotropical 10:196-199.

Sage, R. D., O. P. Pearson, J. Sanguinetti, and A. K. Pearson 2006 Ratada 2001: A rodent outbreak following the flowering of bamboo (Chusquea culeou) in southern Argentina / Ratada 2001: Una irrupción de roedores siguiendo al florecimiento de bambú (Chusquea culeou) en el sur de la Argentina. University of California Publications in Zoology xxx:xx- xx.

Sanborn, C. C., and O. P. Pearson 1947 The tuco-tucos of Peru (Genus Ctenomys). Proceedings of the Biological Society of Washington 60:135-138. Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 15

Appendix 1. Publications by Oliver P. Pearson.

This listing, compiled by the editors with assistance from Anita K. Pearson and the staff of MVZ, includes all publications htat were authored or co-authored by Paynie. These begin with results from his first Neotropical field trips. Collaboration with his mentor, Robert Enders, is apparent, as is an applied focus on breeding ecology and physiology of furberaing mammals. This rapidly yields to his better-known studies on vertebrate physiology, South American mammals, and a rare integration of systematics, ecology, biogeography, and natural history.

1939

Pearson, O. P. Three new small mammals from eastern Panama. Notulae Naturae of the Academy of Natural Sciences of Philadelphia 6:1-5.

Enders, R. K., and O. P. Pearson. Three new rodent subspecies of Scotinomys from Chirique, Panama. Notulae Naturae of the Academy of Natural Sciences of Philadelphia 34:1-4.

1940

Enders, R. K., and O. P. Pearson. A new subspecies of Reithrodontomys mexicanus from Panama with remarks on R. mexicanus cherrii. Notulae Naturae of the Academy of Natural Sciences of Philadelphia 60:1-2.

1942

Pearson, O. P. On the cause and nature of a poisonous action produced by the bite of a shrew (Blarina brevicauda). Journal of Mammalogy 23:159-166.

1943

Pearson, O. P., and R. K. Enders. Ovulation, maturation and fertilization in the fox. Anatomical Record 85:69-83.

Pearson, O. P. The status of the vicuna in southern Peru, 1940. Journal of Mammalogy 24:97.

Enders, R. K., and O. P. Pearson. The blastocyst of the fisher. Anatomical Record 85:285-287.

Pearson, O. P., and R. K. Enders. Duration of pregnancy in marten, mink, and related species. American Fur Breeder 16 University of California Publications in Zoology

Enders, R. K., and O. P. Pearson. Shortening gestation by inducing early implantation with increased light in the marten. American Fur Breeder, January:18-19.

Pearson, O. P., and R. K. Enders. Your mating of foxes should be timed with great care. Black Fox Magazine, December:xxxxxxx

1944

Bassett, C. F., F. Wilke, and O. P. Pearson. The effect of mating time on productivity in foxes. Black Fox Magazine, January: 12-13, 27, 29.

Pearson, O. P., and R. K. Enders. Duration of pregnancy in certain mustelids. Journal of Experimental Zoology 95:21-35.

Pearson, O. P. Reproduction in the shrew (Blarina brevicauda Say). American Journal of Anatomy 75:39-93.

Pearson, O. P., and C. F. Bassett. Size of the vulva in the fox and its relation to fertility. Anatomical Record 89:455-459.

Bassett, C. F., F. Wilke, and O. P. Pearson. Relative productivity under different systems of mink mating. The Fur Journal 10:10-12.

Bassett, C. F., O. P. Pearson, and F. Wilke. The effect of artificially increased length of day on molt, growth, and priming of silver fox pelts. Journal of Experimental Zoology 96:77-83.

1945

Pearson, O. P. Longevity of the short-tailed shrew. American Midland Naturalist 34:531-546.

1946

Pearson, O. P., and C. F. Bassett. Certain aspects of reproduction in a herd of silver foxes. American Naturalist 53:45-67.

Enders, R. K., O. P. Pearson, and A. K. Pearson. Certain aspects of reproduction in the fur seal. Anatomical Record 94:213-227.

Pearson, O. P. Scent glands of the short-tailed shrew. Anatomical Record 94:615- 629.

Morrison, P. R., and O. P. Pearson. The metabolism of a very small mammal. Science 104:287-289. Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 17

1947

Pearson, O. P. The rate of metabolism of some small mammals. Ecology 28:127- 145.

Pearson, O. P., and A. K. Pearson. Owl predation in Pennsylvania, with notes on the small mammals of Delaware County. Journal of Mammalogy 28:137-147.

Sanborn, C. C., and O. P. Pearson. The tuco-tucos of Peru (Genus Ctenomys). Proceedings of the Biological Society of Washington 60:135-138.

1948

Pearson, O. P. Metabolism and bioenergetics. Scientific Monthly 66:131-134.

Pearson, O. P. Metabolism of small mammals, with remarks on the lower limit of mammalian size. Science 108:44.

Pearson, O. P. Life history of mountain viscachas in Peru. Journal of Mammalogy 29:345-374.

1949

Pearson, O. P. Reproduction of a South American rodent, the mountain viscacha. The American Journal of Anatomy 84:143-173.

1950

Pearson, O. P. The submaxillary glands of shrews. Anatomical Record 107:161- 169.

Pearson, O. P. The metabolism of hummingbirds. Condor 52:145-152.

Pearson, O. P. Keeping shrews in captivity. Journal of Mammalogy 31:351-352.

1951

Pearson, O. P. Mammals in the highland of southern Peru. Bulletin of the Museum of Comparative Zoology 106(3):117-174.

Pearson, O. P. Additions to the fauna of Santa Cruz Island, California, with description of a new subspecies of Reithrodontomys megalotis. Journal of Mammalogy 32:366-368. 18 University of California Publications in Zoology

1952

Pearson, O. P., M. R. Koford, and A. K. Pearson. Reproduction of the lump-nosed bat (Corynorhinus rafinesquei) in California. Journal of Mammalogy 33:273- 320.

Pearson, O. P. Notes on a pregnant sea otter. Journal of Mammalogy 33:387.

1953

Pearson, O. P. Metabolism of hummingbirds. Scientific American 188:69-42.

Pearson, O. P. Use of caves by hummingbirds and other species at high altitudes in Peru. Condor 55:17-20.

Pearson, O. P., and P. Baldwin. Reproduction and age structure of a mongoose population in Hawaii. Journal of Mammalogy 34:436-447.

1954

Pearson, O. P. Habits of the lizard Liolaemus multiformis multiformis at high altitudes in southern Peru. Copeia 2:111-116.

Pearson, O. P. Shrews. Scientific American 191(2):66-70.

Pearson, O. P. The daily energy requirements of a wild Anna hummingbird. Condor 56:317-322.

Pearson, A. K., and O. P. Pearson. Natural history and breeding behavior of the tinamou, Nothoprocta ornata. Auk 72:113-127.

1956

Pearson, O. P. A toxic substance from the salivary glands of a mammal (short- tailed shrew). Venoms 44:55-58.

1957

Pearson, O. P. Additions to the mammalian fauna of Peru and notes on some other Peruvian mammals. Breviora 73:1-7. Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 19

1958

Pearson, O. P. A taxonomic revision of the rodent genus Phyllotis. University of California Publications in Zoology 56:391-477.

Pearson, O. P. Certain aspects of the biology of mouse runways. Science 128:1148- 1148.

1959

Pearson, O. P. A traffic survey of Microtus-Reithrodontomys runways. Journal of Mammalogy 40:169-180.

1960

Pearson, O. P. Habits of harvest mice revealed by automatic photographic recorders. Journal of Mammalogy 41:58-74.

Pearson, O. P. The oxygen consumption and bioenergetics of harvest mice. Physiological Zoology 33:152-160.

Pearson, O. P. Biology of the subterranean rodents, Ctenomys, in Peru. Memorias del Museo de Historia Natural ‘Javier Prado’ 9:1-56.

Pearson, O. P. Speed of the Allen hummingbird while diving. Condor 62:403.

Pearson, O. P. A mechanical model for the study of population dynamics. Ecology 41:494-508.

Pearson, O. P. Habits of Microtus californicus revealed by automatic photographic recorders. Ecological Monographs 30:231-249.

Pearson, O. P. Torpidity in birds. Bulletin of the Museum of Comparative Zoology 124:93-103.

1961

Pearson, O. P. Flight speeds of some small birds. Condor 63:506-507.

1962

Pearson, O. P. Survival value of vibrissae. Journal of Mammalogy 43:105-106. 20 University of California Publications in Zoology

1963

Pearson, O. P. History of two local outbreaks of feral house mice. Ecology 44:540- 548.

1964

Pearson, O. P. Carnivore-mouse predation: An example of its intensity and bioenergetics. Journal of Mammalogy 45:177-188.

Pearson, O. P. Metabolism and heat loss during flight in pidgeons. Condor 66:182- 185.

1966

Pearson, O. P. The prey of carnivores during one cycle of mouse abundance. Journal of Animal Ecology 35:217-233.

1967

Pearson, O. P. La estructura por edades y la dinamica reproductive en una poblacion de ratones de campo, Akodon azarae. Physis 27:53-58.

Pearson, O. P. Review: Comparative biology of reproduction in mammals. Journal of Mammalogy 48:335.

1968

Pearson, O., N. Binszstein, L. Boiry, C. Busch, M. Di Pace, G. Gallopin, P. Penchaszadeh, and M. Piantanida. Estructura social, distribucion espacial y composicion por edades, de una poblacion de tuco-tucos (Ctenomys talarum). Investigaciones Zoologicas Chilenas 13:47-80.

1971

Pearson, O. P. Additional measurements of the impact of carnivores on California voles (Microtus californicus). Journal of Mammalogy 52:41-49.

1972

Pearson, O. P. New information on ranges and relationships within the rodent genus Phyllotis in Peru and Ecuador. Journal of Mammalogy 53:677-686 Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 21

1974

Pearson, O. P. Ecology and behavior. Chairman’s introduction: Symposium of the Zoological Society of London 34:109-111

1975

Pearson, O. P. An outbreak of mice in the coastal desert of Peru. Mammalia 39:375- 386

1976

Pearson, O. P. and D. F. Bradford. Thermoregulation of lizards and toads at high altitudes in Peru. Copeia 1976:155-170

Pearson, O. P. and J. L. Patton. Relationships among South American phyllotine rodents based on chromosome analysis. Journal of Mammalogy 57:339-350

Pearson, O. P. and A. K. Pearson. A stereological analysis of the ultrastructure of the lungs of wild mice living at low and high altitude. Journal of Morphology 250:359-368

1977

Pearson, O. P. The effect of substrate and of skin color on thermoregulation of a lizard. Comparative Biochemistry and Physiology 58:363-358

1978

Pearson, O. P., and C. P. Ralph. The diversity and abundance of vertebrates along an altitudinal gradient in Peru. Memoria del Museo de Historia Natural “Javier Prado” 18:1-97.

1979

Pearson, A. K. and O. P. Pearson. Granular pneumocytes and altitude: a stereological evaluation. Cell and Tissue Research 201:137-144

Pearson, O. P. Spacing and orientation among feeding golden-crowned sparrows. Condor 81:278-285 22 University of California Publications in Zoology

1982

Pearson, O. P. Distribución de pequeños mamíferos en el Altiplano y los desiertos del Perú. Pp. 263-284 in Salinas, P. (ed.), Zoología Neotropical: Actas de VIII Congreso Latinoamericano de Zoología, Mérida, Venezuela.

Pearson, O. P. and A. K. Pearson. Ecology and biogeography of the southern rainforests of Argentina. Pp. 129-142 in: The Pymatuning Symposia in Ecology, Vol. 6: Mammalian Biology in South America (Mares, M. A., and H. H. Genoways, eds.). Special Publication Series, Pymatuning Laboratory of Ecology, University of Pittsburgh, Linesville, Pennsylvania.

1983

Pearson, O. P. Characteristics of a mammalian fauna from forests in Patagonia, Southern Argentina. J. Mamm. 64:476-492.

1984

Pearson, O. P. Taxonomy and natural history of some fossorial rodents of Patagonia, southern Argentina. Journal of Zoology 202:225-237.

1985

Pearson, O. P. Predation. Pp. 535-566 in Biology of New World Microtus (Tamarin, R. H., ed.). Special Publication, American Society of Mammalogists 8.

Pearson, O. P. and M. I. Christie. Los tuco-tucos (genero Ctenomys) de los parques nacionales Lanin y Nahuel Huapi, Argentina. Historia Natural 5:337-344

1987

Pearson, O. P. Mice and the postglacial history of the Traful Valley of Argentina. Journal of Mammalogy 68:469-478.

Pearson, O., S. Martin, and J. Bellati. Demography and reproduction of the silky desert mouse (Eligmodontia) in Argentina. Fieldiana:zoology, new series 39:433-446.

1988

Pearson, O. P. Biology and feeding dynamics of a South American herbivorous rodent, Reithrodon. Studies on Neotropical Fauna and Environment 23:25-39. Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 23

1989

Pearson, O. P. and A. K. Pearson. Reproduction of bats in Southern Argentina. Advances in Neotropical Mammalogy, pp. 549-566.

1991

Pearson, O. P. and M. I. Christie. Sympatric species of Euneomys (Rodentia, ). Studies on Neotropical Fauna and Environment 26:121-127.

1992

Pearson, O. P. Reproduction in a South American mouse, Abrothrix longipilis. Anatomical Record 234:73-88.

Pearson, O. and H. A. Lagiglia. “Fuerte de San Rafael”: una localidad tipo ilusoria. Revista del Museo de Historia Natural de San Rafael (Mendoza) 12:35-43.

1993

Pearson, O. P. and M. I. Christie. Rodent guano (amberat) from caves in Argentina. Studies on Neotropical Fauna and Environment 28:105-111.

Pearson, O. P., and A. K. Pearson. La fauna de mamiferos pequeños de Cueva Traful I, Argentina: pasado y presente. Praehistoria 1:211-214 (PREP-CONICET, Buenos Aires, Argentina).

1994

Pearson, A. K., O. P. Pearson, and I. A. Gomez. Biology of the bamboo Chusquea culeou (Poacea: Bambusoideae) in southern Argentina. Vegetatio 111:93-126.

Pearson, O. P. and G. J. Kenagy. Reproduction. In Seventy-five years of Mammalogy (1919-1994). Special Publication No. 11, the American Society of Mammalogists (Birney, E.C., and J.R. Choate, eds.), pp. 271-287.

Pearson, O. P. The impact of an eruption of Volcan Hadson on small mammals in Argentine Patagonia. Mastozoologia Neotropical 1(2):103-112. 24 University of California Publications in Zoology

1995

Pearson, O. P. Annotated keys for identifying small mammals living in or near Nahuel Huapi National Park or Lanin National Park, southern Argentina. Mastozoologia Neotropical 2:99-148.

Pearson, O. P. In memoriam: Wilbur Brooks Quay, 1927-1994. Auk 112:489-491.

1997

Pearson, O. P. Monticulos “Mima”. Revista Patagonia Silvestre/Snap No. 4, 2 p.

1999

Pearson, O. P., and M. F. Smith. Genetic similarity between Akodon olivaceus and Akodon xanthorhinus (Rodentia: Muridae) in Argenitina. Journal of Zoology 247:43-52.

2000

Kenagy, G. J. and O. P. Pearson. Life with fur and without: experimental field energetics and survival of naked meadow voles. Oecologia 122:220-224.

2002

Pearson, O. P. A perplexing outbreak of mice in Patagonia, Argentina. Studies on Neotropical Fauna and Environment 37:187-200.

2005

Chan, Y. L., E. A. Lacey, O. P. Pearson, and E. A. Hadly. Ancient DNA reveals Holocene loss of genetic diversity in a South American rodent. Biology Letters 1:423-426.

2006

Sage, R. D., O. P. Pearson, J. Sanguinetti, and A. K. Pearson. Ratada 2001: A Rodent Outbreak Following the Flowering of Bamboo (Chusquea culeou) in Southern Argentina / Ratada 2001: Una Irrupción de Roedores Siguiendo al Florecimiento de Bambú (Chusquea culeou) en el Sur de la Argentina. Pp. Xx-xx in The quintessential naturalist: honoring the life and legacy of Oliver P. Pearson (Kelt, D. A., E. P. Lessa, J. Salazar-Bravo, and J. L. Patton, eds.). University of California Publications in Zoology xxx:xxx-xxx. Kelt et al.: Oliver P. Pearson, Scientist, Statesman, Gentleman 25

In press

Pearson, A. K., O. P. Pearson, and P. L. Ralph. Growth and activity patterns in a backyard population of the banana slug, Aridimax columbianus. Veliger. 26 University of California Publications in Zoology