STUDIES IN NEOTROPICAL MAMMALOGY Essays in Honor of Philip Hershkovitz

Edited by Bruce D. Patterson and Robert M. Timm

December 31, 1987 Publication 1382


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STUDIES IN NEOTROPICAL MAMMALOGY Essays in Honor of Philip Hershkovitz

Phiup Hershkovttz




STUDIES IN NEOTROPICAL MAMMALOGY Essays in Honor of Philip Hershkovitz

Edited by

Bruce D. Patterson

Division of Mammals

Field Museum of Natural History

Chicago, Illinois 60605-2496

Robert M. Timm

Division of Mammals

Field Museum of Natural History

Present address:

Museum of Natural History

Department of Systematics and Ecology

University of Kansas

Lawrence, Kansas 66045

Accepted for publication July 30, 1985 December 31, 1987 Publication 1382



New Series, No. 39

Studies in Neotropical Mammalogy:

Essays in Honor of Philip Hershkovitz

Bruce D. Patterson and Robert M. Timm, Editors

© 1987 Field Museum of Natural History Library of Congress Catalog Card Number: 87-82549


Table of Contents

Preface vii

A Biographical Sketch of Philip Hershkovitz, with a Complete Scientific Bibliography 1

Bruce D. Patterson

A History of the Recent Mammalogy of the Neotropical Region from 1492 to 1850 11

Philip Hershkovitz

A New Superfamily in the Extensive Radiation of South American Paleogene Marsupials 99

Rosendo Pascual and Alfredo A. Carlini

An Additional 14-Chromosome Karyotype and Sex-Chromosome Mosaicism in South American

Marsupials 1 1 1

Milton H. Gallardo and Bruce D. Patterson

Notes on the Black-Shouldered Opossum, Caluromysiops irrupta 117

Robert J. Izor and Ronald H. Pine

Feeding Habits of the Opossum {Didelphis marsupialis) in Northern Venezuela 125

Gerardo A. Cordero R. and Ruben A. Nicolas B.

Notes on Distribution of Some Bats from Southwestern Colombia 133

Michael S. Alberico

Distributional Records of Bats from the Caribbean Lowlands of Belize and Adjacent Guatemala

and Mexico 137

Timothy J. McCarthy

New Species of Mammals from Northern South America: Fruit-Eating Bats, Genus Artibeus

Leach 163

Charles O. Handley. Jr.

Seasonality of Reproduction in Peruvian Bats 173

Gary L. Graham

Tent Construction by Bats of the Genera Artibeus and Uroderma 187

Robert M. Timm

Comparative Ultrastructure and Evolutionary Patterns of Acinar Secretory Product of Parotid

Salivary Glands in Neotropical Bats 213

Carleton J. Phillips, Toshikazu Nagato, and Bernard Tandler

Distribution of the Species and Subspecies of Cebids in Venezuela 231

Roberta Bodini and Roger Perez- Hernandez

Host Associations and Coevolutionary Relationships of Astigmatid Mite Parasites of New World

Primates. I. Families Psoroptidae and Audycoptidae 245

Barry M. OConnor Notes on Bolivian Mammals. 2. Taxonomy and Distribution of Rice Rats of the Subgenus Oli-

goryzomys 261

Nancy Olds and Sydney Anderson

New Records and Current Status of Euneomys (Cricetidae) in Southern South America 283

Jose L. Ydhez, Juan C Torres-Mura. Jaime R. Rau, and Luis C. Contreras

Morphological Variation, Karyology, and Systematic Relationships of Heteromys gaumeri (Ro-

dentia: Heteromyidae) 289

Mark D. Engstrom. Hugh H. Genoways, and Priscilla K. Tucker

Species Groups of Spiny Rats, Genus Proechimys (Rodentia: Echimyidae) 305

James L. Patton

An Assessment of the Systematics and Evolution of the Akodontini, with the Description of New

Fossil Species of Akodon (Cricetidae: Sigmodontinae) 347

Osvaldo A. Reig


Biogeography of Octodontid Rodents: An Eco-Evolutionary Hypothesis 40 1

Luis C. Contreras, Juan C. Torres-Mura, and Jose L. Ydnez

Population Dynamics and Ecology of Small Mammals in the Northern Chilean Semiarid

Region 413

Peter L. Meserve and Eric Le Boulenge

Demography and Reproduction of the Silky Desert Mouse (Eligmodontia) in Argentina 433

Oliver Pearson. Susana Martin, and Javier Bellati

Baculum of the Lesser Andean Coati, Nasuella olivacea (Gray), and of the Larger Grison, Galictis

vittata (Schreber) 447

Edgardo Mondolfi

Origin, Diversification, and Zoogeography of the South American Canidae 455

Annalisa Bert a

Comparative Cytogenetics of South American Deer 473

Angel E. Spot or no. Nadir Brum, and Mariela Di Tomaso

Faunal Representation in Museum Collections of Mammals: Osgood's Mammals of Chile 485

Bruce D. Patterson and Clare E. Feigl

Taxonomic Index 497

Subject Index 505



In the early 1 980s, we discussed the possibility of a testimonial volume for Philip Hershkoviiz with Larry Marshall, then with the Department of Geology, Field Museum. As the senior mammal- ogist at Field Museum and a student of South American mammalogy for almost half a century, Hershkovitz had generously provided invaluable advice and assistance to each of us in the early stages of our careers. We felt a Festschrift in his honor might repay a portion of our debts to him and, at the same time, serve as an independent, lasting tribute to his life-work.

In the entire history of Field Museum, only three testimonial volumes had been produced in honor of museum scientists. Each recognized the contributions of men who were both preeminent scientists and museum administrators: Wilfred H. Osgood, Chief Curator of Zoology, 1921-1941; Karl P. Schmidt, Chief Curator of Zoology, 1941- 1956; and Rainer Zangerl, Chief Curator and Chairman of Geology, 1962-1974. Although Hershkovitz has never served in an upper-level administrative capacity, his contributions to the museum through distinguished and continuing re- search clearly qualified him for this honor.

However, plans for a testimonial volume in Fieldiana: Zoology did not materialize until No- vember 1 983. By that time, Marshall had assumed a new position at the University of Arizona and Hershkovitz had just celebrated his 74th birthday. Given realistic editing and publication schedules, we were faced with the prospect of producing the volume nearly midway between traditionally cel- ebrated anniversary dates. Nevertheless, such tim- ing is somehow fitting: Hershkovitz the man is both extemporaneous and unconventional.

Another notable departure from the Festschrift tradition is evident from the table of contents: Hershkovitz himself is a contributor! On many occasions Hershkovitz had lamented the lack of a historical review of South American mammalogy. During the present information explosion, scien- tists are hard-pressed to keep up with current de- velopments of direct relevance to their research; much less are they afforded the occasion to amble through historical records in Latin, German, French, Spanish, and Portuguese, even though these records are full of interesting and relevant infor- mation. As a result of his 50 years in the discipline, Hershkovitz may be unique in his broad knowl- edge of both historical literature and current re- search on Neotropical mammals. The editors

therefore prevailed upon him to write such a his- torical survey to complement and enhance this volume. We convinced him that, by assembling a historical analysis of the subject, he would provide a tremendous service to younger workers.

Other contributions to the volume came from friends and colleagues of Hershkovitz. All share an interest in the distribution, taxonomy, and nat- ural history of Neotropical mammals, and each one was inspired to honor Hershkovitz with their contribution. Each of the contributions focuses on those fields of Neotropical mammalogy to which Hershkovitz has contributed most significantly.

We owe thanks to numerous persons connected with this volume. First and foremost, Tanisse Be- zin, Managing Editor of Field Museum Press, de- serves recognition. Her keen eye for grammar and style eliminated numerous editorial inconsisten- cies forwarded by the volume editors. Graham Harles, Field Museum Press copy editor, provided skillful editing and proofreading. The Scientific Editor for Fieldiana, Timothy Plowman, endured countless interruptions during production of this volume and served as corresponding editor for our own papers. Translations of abstracts into Spanish and Portuguese were kindly provided by Myriam Ibarra (an Ecuadorean ichthyologist) and Debra Moskovits (a Brazilian ecologist), who offered these as their own tributes to Hershkovitz. Assistance in assembling the indices was provided by Mary Anne Rogers.

Finally, we are enormously indebted to a ded- icated body of reviewers, who critically evaluated papers in this volume. Their constructive advice and recommendations made editorial tasks far lighter. The editors gratefully thank: W. T. Atyeo, P. V. August, K. Benirschke, W. A. Clemens, J.

A. Davis, W. B. Davis, M. R. Dawson, M. D. Engstrom, J. Fooden, G. L. Forman, M. H. Ga- llardo, A. L. Gardner, H. H. Genoways, W. E. Glanz, M. S. Hafner, D. Hunsaker II, R. J. Izor, J. A. W. Kirsch, K. F. Koopman, M. A. Mares, R. E. Martin, T. J. McCarthy, G. G. Musser, P. Myers, J. L. Patton, O. P. Pearson, R. H. Pine, W.

B. Quay, L. Radinsky, O. J. Reichman, D. S. Rog- ers, R. W. Thorington, Jr., W. D. Tumbull, J. H. Wahlert, S. D. Webb, C. Wemmer, J. O. Whitaker, Jr., D. E. Wilson, R. G. Wolff, and A. E. Wood, in addition to anonymous reviewers of our own papers.

B. D. Patterson R. M. TiMM Chicago, Illinois

A Biographical Sketch of Philip Hershkovitz, with a Complete Scientific Bibliography

Bruce D. Patterson

Philip Hershkovitz was bom October 12, 1909, in Pittsburgh, Pennsylvania, to Aba Hershkovitz and Bertha Halpem. He was the second of four children and their only son. He attended primary and secondary schools in Pittsburgh, graduating from Schenley High School in February 1927. In 1929 he enrolled at the University of Pittsburgh where he majored in zoology, serving as an Un- dergraduate Assistant in that department during 1 930-1 931. Having exhausted Pittsburgh's course offerings in zoology and seeking to pursue a career in mammalogy, he was advised to transfer to another school with an expanded curriculum (Har- vard University, University of Michigan, or Uni- versity of California, Berkeley). In his junior year (1931), he transferred to the University of Mich- igan at Ann Arbor because of its proximity to his home. There he became an Undergraduate Assis- tant in the Museum of Zoology, working under the supervision of Professor and Curator Lee R. Dice during 1931-1932. He supplemented the meager earnings of this position with taxidermy jobs, which supported him during the early years of the Great Depression.

His first fieldwork was undertaken during the summer of 1932. He went to the San Marcos re- gion of Texas to collect blind cave salamanders {Typhlomolge rathbuni) for Professor Uhlenhuth of the University of Maryland Medical School. Having a principal interest in mammals, he want- ed to collect small mammals in areas surrounding the caves, but Dice could spare no traps for him and told him to purchase some in Texas.

While hitchhiking from Ann Arbor to Texas, Hershkovitz stopped to visit friends in Chicago. There, a chance visit to Field Museum of Nat- ural History secured him the traps and supplies he needed and seemingly set the course of his later

career. Colin Sanborn, then Curator of Mammals during Wilfred Osgood's tenure as Chief Curator of Zoology (1921-1941), befriended Hershkovitz and loaned him the necessary supplies. As a con- sequence, the mammals that Hershkovitz collect- ed in Texas that first of many field seasons were deposited in the Field Museum collections. He now maintains that his chance visit to Field Museum in 1932 indelibly fixed that institution as the place at which to pursue his career goals.

Hershkovitz's formal education was delayed by the worsening economic situation during 1 933. No longer able to afford tuition, he sought advice on subsistence during the Depression, and was told that Ecuador and Paraguay were undoubtedly the least expensive countries in this hemisphere in which to live. Transportation costs decided the issue, and in 1933 he set sail via the Grace Line from New York to Guayaquil, Ecuador for the whopping sum of $600, one-way.

He stayed in Ecuador until 1937. During this period, he mastered Spanish and learned how to live off the land in the Neotropics. His boots dis- integrated after six months' time and thereafter he went barefoot. He assembled a fine collection of Ecuadorean mammals for the Museum of Zool- ogy, University of Michigan, supporting his activ- ities in part by selling horses bought on the Pe- ruvian frontier.

He then returned to the University of Michigan where he again enrolled as an undergraduate, grad- uating in 1938 with a Bachelor of Science degree. By this time. Dice had moved from the Museum of Zoology to the Laboratory of Vertebrate Ge- netics, and William H. Burt had assumed the cu- ratorship in the Museum. Hershkovitz spent the years 1938-1941 as a graduate student enrolled at the University of Michigan, working on his Ecua-


dorean collection under Burt's direction. From 1939-1941, he was supported in this work by a Graduate Assistantship. In 1940 he received his Master of Science degree and immediately entered the doctoral program.

Two years before the expected completion of his doctoral program, the Curator of Reptiles and Amphibians at the Museum of Zoology, Helen Gage, told Hershkovitz about the Walter Rath- bone Bacon Travelling Scholarship of the United States National Museum. This program was cus- tomarily reserved for postdoctoral support, but Mrs. Gage strongly urged him to apply immedi- ately. Thus encouraged, Hershkovitz submitted a brief proposal for work in the Santa Marta region of northern Colombia; his compliance with Mrs. Gage's wishes in this matter was so perfunctory that he failed to include a map of the proposed itinerary. But Remington Kellogg at the National Museum had long wished to obtain a Bacon Schol- ar for the Mammal Division and asked Hersh- kovitz to send the omitted material. Much to his surprise, Hershkovitz was awarded the scholar- ship and left Ann Arbor immediately for Wash- ington. He spent two months there studying the then very poor collection of Neotropical mam- mals. Afterward he spent two years in Colombia (1941-1943) collecting mammals, other verte- brates, and ectoparasites. The resulting collection was the National Museum's first large and repre- sentative Neotropical mammal accession.

In 1943 Hershkovitz's work was interrupted by World War II, and he returned to Ann Arbor to enlist in the Armed Services. He was assigned to the Office of Strategic Services (OSS) and served from 1943-1946 in the European Theater. While serving in France, he met Anne Marie Pierrette, whom he married in 1946. The two returned to the United States, where in 1946 and 1947 he continued his Bacon Scholarship studies of Co- lombian mammals in Washington. The first of three children (Francine, Michael, and Mark) was bom in 1947.

About this time, he was contacted regarding the opening of a curatorial position at Field Museum in Chicago, an opportunity he eagerly hailed for several reasons: ( 1 ) The comprehensive collections of Neotropical mammals at Field Museum would be a tremendous resource for what he had already decided would be his life's work; (2) he had the highest regard for W. H. Osgood, who as a prin- cipal authority on South American mammals would be a great personal resource on which to draw; (3) the press of family responsibibties made

continuation of his graduate studies untenable; and (4) aspirations to a curatorial position had been the raison d'etre of his graduate program; a cur- atorial position made the graduate degree sujjer- fluous. Thus he jumped at the offer of employment at Field Museum, knowing full well that it marked the end of his graduate program at Michigan. Like many similar institutions, Michigan had a final year-in-residence degree requirement. Unfortu- nately, Osgood died in June of 1947, and what might have been a remarkably productive ap- prenticeship under Osgood never came to pass.

Upon his arrival at Field Museum, Hershkovitz found an uncurated backlog of some four or five years of accessions. Nevertheless, he wasted little time in returning to the field, prompted in part by postwar housing shortages in Chicago. (One can almost hear him now, telling the Museum's Di- rector Clifford Gregg that the nearest affordable housing was in Bogota!) In 1 948 he and his family moved to Colombia where he resumed his inven- tory of the mammals of that country. He remained in Colombia until the press of curatorial duties and a gently delivered ultimatum from Sanborn finally recalled him to Chicago in 1952.

The collections he made in Colombia, first for the National Museum, then for Field Museum, were to be the heart of all his subsequent research. But unlike others studying the mammal faunas of specific geographical regions, Hershkovitz found it unsatisfying to assess the systematics of Colom- bian mammals without following them across na- tional boundaries. Studies of a species or species group in Colombia led him to evaluate its context within genera, families, and even orders; and the remarkable diversity of Colombia's mammal fau- na led him into most major groups and most Neo- tropical subregions. In the course of his career, he has published dozens of generic, tribal, and fa- milial revisions, covering all 1 2 orders of Neo- tropical mammals. Few spatial and temporal boundaries have withstood the onslaught of his studies of Neotropical mammals. As examples one can point to the cosmopolitan Catalog of Living Whales {\9()6)—2iiXtr all, most cetaceans do occur in South American waters— and studies of Oli- gocene and later fossils (1974, 1982).

One senses that the Department of Zoology dur- ing Hershkovitz's early years was a stimulating, harmonious one. Chief Curator Karl P. Schmidt took an almost paternal interest in junior staff and served as a confidant on the most personal of mat- ters. In addition to Colin Sanborn, who was most considerate of his junior curator's interests and


talents, Hershkovitz shared mammalogical prob- lems and topics with Dwight Davis, Curator of Anatomy, and Bryan Patterson, Curator of Ver- tebrate Paleontology. During the early and mid- 1950s, Hershkovitz established a vigorous and productive research program and participated in all aspects of departmental affairs.

However, upon Schmidt's retirement in 1957, Austin S. Rand became Chief Curator of Zoology, and neither Rand nor Hershkovitz did much to disguise their antipathy for one another. Over the ensuing years, Hershkovitz increasingly detached himself from museum operations, culminating with Joseph Moore's appointment as Curator of Mam- mals in 1961, and Hershkovitz's appointment that year to Research Curator. No one before or since has held this title at Field Museum. Hershkovitz formally retired in 1971, although his work has continued unabated as Curator Emeritus. During his career, he assisted countless students in mam- mal projects, but has served on only a single grad- uate committee, that of Jack Fooden, now himself a renowned biologist and primate specialist at Field Museum.

Few scientists can claim the independence in research that is indicated in Hershkovitz's bibli- ography. Of his approximately 300 scientific, pop- ular, and encyclopedia articles, only three repre- sent collaborative efforts. The first, with William P. Harris, an important benefactor of the Museum of Zoology of the University of Michigan, was suggested by Burt in recognition of Harris's inter- ests in squirrels and in token repayment for his patronage of the museum. The second, with Paul Rode, came about one afternoon in the Museum National d'Histoire Naturelle in Paris when Hershkovitz offhandedly suggested that designat- ing a lectotype might solve a nomenclatural prob- lem that Rode had encountered in his research. Rode insisted that Hershkovitz share authorship on the resulting paper. Later, after further study in the United States, Hershkovitz arrived at a con- trary opinion and wrote a paper, with Rode as coauthor, correcting their earlier one.

Independent thought is also exemplified by the sometimes heated debates in which Hershkovitz has participated over the years. His published re- views and the discussion sections of many of his papers record his clearly enunciated views on such topics as the role of penial morphology in rodent taxonomy, the age and derivation of the South American fauna, panbiogeography, evolution of pelage coloration, and the systematic position of certain species (e.g., Dolichocebus). While such

firmly held views brand him as something other than conciliatory or diplomatic, they accurately reflect his abiding passion and zest in science. Un- fortunately, some acerbic exchanges had the effect of stifling the scientific dialogue to which they were offered (e.g., penial morphology).

Hershkovitz has focused his research on Neo- tropical mammals, their origin, evolution, dis- persal, classification, nomenclature, and system- atics. Specialists in these fields are well aware of his impact. However, he is perhaps most widely known for his work on three general topics of Neotropical mammalogy: faunal origins, meta- chromism, and New World monkeys. It would be folly to attempt to review all of his research, and more definitive appraisals on selected topics can be found scattered throughout this volume. How- ever, some comments on these general issues seem in order.

As late as his revision of phyllotine rodents ( 1 962), Hershkovitz adhered to traditional notions of the derivation of certain South American taxa, notably "cricetid" rodents, from North and Mid- dle American stocks. This hypothesis of origins has been advocated most articulately by George G. Simpson, Bryan Patterson, and Rosendo Pas- cual, and more recently by Larry G. Marshall and S. David Webb. However, in the early 1960s, Hershkovitz was approached by Rupert Wenzel, Curator of Insects at Field Museum, who ques- tioned him on the evidence for Plio-Pleistocene origins of the Neotropical cricetids. Wenzel's stud- ies of the ectoparasites of Panamanian mammals suggested much earlier. South American origins. His interest piqued, Hershkovitz reviewed avail- able evidence, synthesizing continental drift (which was then becoming established in geological cir- cles) and neontological studies of mammals (es- pecially those of Hooper and Musser, which showed a relatively sharp dichotomy between sim- ple and complex penis-types of cricetids). He con- cluded that continental drift permitted a much greater role for paleotropical stocks in South American faunal origins than was allowed by the Simpsonian school, which in turn pointed to a much greater time period for independent evo- lution. Interestingly, and perhaps even character- istically, Hershkovitz concluded that South Amer- ican rodents were not only not derived from North American stocks, but instead gave rise to them. These views were published in 1966, 1969, and 1972.

Hershkovitz's theory of metachromism, or de- terministic evolution of pelage coloration through


the loss of one or the other or both classes of hair pigments (eumelanins and phaeomelanins), was first pubHshed in 1968. Since then he has used it repeatedly in describing geographic variation in platyrrhine monkeys (e.g., 1977). However, the origins of this concept stem from his earlier work on the Sciunds granatensis group in northern Co- lombia where populations of squirrels thoroughly isolated from one another show similar progres- sions of pelage patterns. Few workers other than Neotropical primatologists (and not all of these) have accepted his interpretations, although the theory is potentially applicable to a variety of oth- er, mostly diurnal taxa showing pelage pattern variations. While Timothy Lawlor detailed some theoretical misgivings with the theory in a 1969 paper in Evolution (rebutted by Hershkovitz in 1970), to my knowledge it has not been substan- tially refuted. The theory is eminently testable: refutation would simply entail showing that pelage pattern variation of taxa arranged by metachro- mism is not congruent with well-established phy- letic patterns.

Finally, some explanation seems warranted for Hershkovitz's current devotion to primates. In- deed, many recent workers unschooled in mam- malian systematics think of him as a primatolo- gist. Nothing could be further from the truth, as he hastens to point out. He had published several articles on primates in the course of working up his Colombian collections, but gave these taxa no special attention until the 1960s. Then govern- ment funding for primate studies soared, largely because of interest in biomedical applications, es- pecially for the complex and taxonomically con- fused Callitrichidae. For almost 20 years, Hersh- kovitz has focused first on the Callitrichidae and Callimiconidae, now on the Cebidae. His slower progress through these groups is attributable to the vast body of current knowledge about them; his 1977 and subsequent works serve as model syntheses of skin and skull morphology with bio- chemistry, karyology, ethology, serology, and ep- idemiology. By his own estimation, monkeys do not culminate his studies of Neotropical mam- mals, but rather represent a large and complex group to be covered in his attempt to treat all South American mammals. After seven years of work on Volume II of his primate monograph, he has near- ly completed generic revisions of cebids lacking prehensile tails and is beginning comparative stud- ies of their organ systems. In 1 984 he submitted another grant proposal for this work, totaling one- half million dollars in direct costs. His is not a

modest work; it has been described by Pine ( 1 982; Vol. 6, Spec. Publ. Ser., Pymatuning Lab. Ecol.) as "the most heroically monumental revisionary monograph ever devoted to a Neotropical group." In 1984, Hershkovitz turned 75 years old. The 14 years he spent in the field in South America have served him well, for he seems younger than many men 15 years his junior. His tireless energy is best indicated by his habitual use of stairs rather than elevators (even his two divisional offices are three floors apart), a continuing program of field- work (most recently in Brazil during 1986 and 1987), and a museum workday that extends from 9 a.m. to 6 p.m., uninterrupted by coffee breaks or even lunch. Visitors to his home, now within walking distance of the Museum, know of his office there which relieves the chronic insomnia of ad- vancing years. He is an outstanding cook, a genial host, a trusted and valued friend, and an awe- somely productive scientist.

Publications of Philip Hershkovitz 1938

1. A new caecilian from Ecuador. Occasional Papers, Museum of Zoology, University of Michigan, 370:1-3.

2. Two new squirrels fi-om Ecuador. Occasion- al Papers, Museum of Zoology, University of Michigan, 391:1-6 (with W. P. Harris).

3. A review of the rabbits of the andinus group and their distribution in Ecuador. Occasion- al Papers, Museum of Zoology, University of Michigan, 393:1-15.


4. Four new oryzomyine rodents from Ecua- dor. Journal of Mammalogy, 21:78-84.

5. Notes on the distribution of the akodont ro- dent, Akodon mollis, in Ecuador with a de- scription of a new race. Occasional Papers, Museum of Zoology, University of Michi- gan, 418:1-3.

6. A new spiny mouse of the genus Neacomys from eastern Ecuador. Occasional Papers, Museum of Zoology, University of Michi- gan, 419:1-4.


7. The South American harvest mice of the ge- nus Reithrodontomys. Occasional Papers, Museum of Zoology, University of Michi- gan, 441:1-7.



8. A systematic review of the Neotropical water rats of the genus Nectomys (Cricetinae). Mis- cellaneous Publications, Museum of Zool- ogy, University of Michigan, 58:1-88.


9. Designation d'un lectotype de Callithrix penicillatus (E. Geoffroy). Bulletin du Mu- seum National d'Histoire Naturelle, Paris 17(3):22 1-222 (with P. Rode).


10. A correction. Journal of Mammalogy, 28(1): 68 (with P. Rode).

1 1 . Mammals of northern Colombia. Prelimi- nary report no. 1 : Squirrels (Sciuridae). Pro- ceedings of the United States National Mu- seum, 97:1-46.


12. Mammals of northern Colombia. Prelimi- nary report no. 2: Spiny rats (Echimyidae), with supplemental notes on related forms. Proceedings of the United States National Museum, 97:125-140.

13. Mammals of northern Colombia. Prelimi- nary report no. 3: Water rats (genus Necto- mys), with supplemental notes on related forms. Proceedings of the United States Na- tional Museum, 98:49-56.

1 4. The technical name of the Virginia deer with a list of the South American forms. Pro- ceedings of the Biological Society of Wash- ington, 61:41-48.

1 5. Names of mammals dated from Frisch, 1 775, and Zimmermann, 1777. Journal of Mam- malogy, 29(3):272-277.


1 6. Technical names for the fallow deer and Vir- ginia deer. Journal of Mammalogy, 30(1): 94.

1 7. Generic names of the four-eyed pouch opos- sum and the woolly opossum (Didelphidae). Proceedings of the Biological Society of Washington, 62:11-12.

18. Technical names of the African muishond (genus Zorilla) and the Colombian hog-nosed skunk (genus Conepatus). Proceedings of the Biological Society of Washington, 62: 13-16.

1 9. Mammals of northern Colombia. Prelimi- nary report no. 4: Monkeys (Primates), with taxonomic revisions of some forms. Pro- ceedings of the United States National Mu- seum, 98:323-427.

20. Mammals of northern Colombia. Prelimi- nary report no. 5: Bats (Chiroptera). Pro- ceedings of the United States National Mu- seum, 99:429-454.

21. Status of names credited to Oken, 1816. Journal of Mammalogy, 30(3):289-301.

22. Tapirs: Strange mammals native to Asia and tropical America from Mexico south. Chi- cago Natural History Museum Bulletin, 20(9):6-7.


23. Mammals of northern Colombia. Prelimi- nary report no. 6: Rabbits (Leporidae), with notes on the classification and distribution of the South American forms. Proceedings of the United States National Museum, 100: 327-375.


24. Mammals from British Honduras, Mexico, Jamaica and Haiti. Fieldiana: Zoology, 31(47):547-569.


25. Zorilla I. Geoffroy and Spilogale Gray, ge- neric names for African and American pole- cats, respectively. Journal of Mammalogy, 34(3):378-382.

26. Four years on a zoological expedition in Co- lombia. Chicago Natural History Museum Bulletin, 24(l):3-4.

27. The reindeer— Important to man in fact and fancy. Chicago Natural History Museum Bulletin, 24(12):3-4.


28. Mammals of northern Colombia, Prelimi- nary report no. 7: Tapirs (genus Tapirus), with a systematic review of American species. Proceedings of the United States National Museum, 103:465-496.

29. What the groundhog undergoes to make a "holiday." Chicago Natural History Mu- seum Bulletin, 25(2):3-4.

30. Who's a cow? Chicago Natural History Mu- seum Bulletin, 25(7):5.


3 1 . Some ecological aspects of natural versus ar- tificial rehabilitation of a water basin area in Bogota, Colombia. Boletin del Instituto de U Salle, Bogota, 41(193/194):80-83.


32. South American marsh rats genus Holochi- lus with a summary of sigmodont rodents. Fieldiana: Zoology, 37:639-673.

33. [Review] Mammals, a guide to familiar American species. Chicago Natural History Museum Bulletin, 26(7):7.

34. Notes on American monkeys of the genus Cebus. Journal of Mammalogy, 36:449-452.

35. Status of the generic name Zorilla (Mam- malia): Nomenclature by rule or by caprice. Proceedings of the Biological Society of Washington, 68:185-192.

36. On the cheek pouches of the tropical Amer- ican paca. Agouti paca (Linnaeus, 1766). Saiigetierkundliche Mitteilungen, 3(2):67-70.

37. Know your rabbits. Sports Afield, 134(6): 36-41,88.


38. Comments on Galerella Gray, Herpestes II- liger, Leucomitra Howell, Icticyon Lund, Lutreola Wagner, Oryctogale Merriam, Paracynictis Pocock. Opinion 384 Interna- tional Commission of Zoological Nomen- clature, 12(5):71-190(intext).

39. Critical remarks on the status of names in Oken's "Lehrbuch." Opinion 417, Interna- tional Commission on Zoological Nomen- clature, 14(l):33-35.


40. Comments on Canis dingo Meyer. Opinion 451, International Commission on Zoolog- ical Nomenclature, 15(17):335-336.

41. Comments on the validation of Muntiacus Rafinesque. Opinion 460, International Commission on Zoological Nomenclature, 15(26):467-468.

42. Comments on the generic name Mormoops Leach. Opinion 462, International Com- mission on Zoological Nomenclature, 16(1): 8-9.

43. Comments on Sciurus gambianus. Opinion 464, International Commission on Zoolog- ical Nomenclature, 16(3):36-39.

44. Comments on the validation of silvestris Schreber, 1777 [Felis {catus) silvestris].

Opinion 465, International Commission on Zoological Nomenclature, 16(4):49.

45. Comments on the validation of the name Phacochoerus Cuvier. Opinion 466, Inter- national Commission on Zoological No- menclature, 16(5):67-68.

46. Comments on the validation of the name Odobenus Brisson. Opinion 467, Interna- tional Commission on Zoological Nomen- clature, 16(6): 84-8 5.

47. The systematic position of the marmoset, Simia leonina Humboldt (Primates). Pro- ceedings of the Biological Society of Wash- ington, 70: 1 7-20.

48. The type locality of Bison bison Linnaeus. Proceedings of the Biological Society of Washington, 70:31-32.

49. A synopsis of the wild dogs of Colombia. Novedades Colombianas Museo de Historia Naturale Universidad del Cauca (Popayan), no. 3:157-161.

50. On the possible occurrence of the spectacled bear, Tremarctos ornatus(F. Cuvier, 1825), in Panama. Saugetierkundliche Mitteilun- gen, 5(3): 122-1 23.


5 1 . [Review] Biological investigations in the Sel- va Lacandona, Chiapas, Mexico. Quarterly Review of Biology, 33(1):67.

52. [Review] Mammals of the Anglo-Egyptian Sudan, by Henry Setzer. Quarterly Review of Biology, 33:82.

53. Technical names of the South American marsh deer and pampas deer. Proceedings of the Biological Society of Washington, 71: 13-16.

54. Type localities and nomenclature of some American Primates, with remarks on sec- ondary homonyms. Proceedings of the Bi- ological Society of Washington, 71:53-56.

55. Stabilization of zoological nomenclature by a "Law of prescription." Bulletin of Zoolog- ical Nomenclature, 15B(20/21):630-632.

56. A critique of Professor Chester Bradley's "Principle of conservation." Bulletin of Zoo- logical Nomenclature, 15B(25/28):9 11-913.

57. The status of secondary homonyms and the concept of permanent rejection. Bulletin of Zoological Nomenclature, 15B(37/38):1242- 1243.

58. A geographic classification of Neotropical mammals. Fieldiana: Zoology, 36(6):583- 620.


59. The metatarsal glands in white-tailed deer and related forms of the Neotropical region. Mammalia, 22(4): 5 3 7-546.


60. The scientific names of the species of ca- puchin monkeys (Cebus Erxleben). Proceed- ings of the Biological Society of Washington, 72:1-4.

6 1 . Two new genera of South American rodents (Cricetinae). Proceedings of the Biological Society of Washington, 72:5-10.

62. A new species of South American brocket, genus Mazama (Cervidae). Proceedings of the Biological Society of Washington, 72: 45-54.

63. A new race of red brocket deer {Mazama americana) from Colombia. Proceedings of the Biological Society of Washington, 72: 93-96.

64. The type locality of Felix concolor concolor Linnaeus. Proceedings of the Biological So- ciety of Washington, 72:97-100.

65. Nomenclature and taxonomy of the Neo- tropical mammals described by Olfers, 1818. Journal of Mammalogy, 40(3):337-353.


66. Supposed ape-man or "missing link" of South America. Chicago Natural History Museum Bulletin, 31(4):6-7.

67. [Review] The Mammals of North America. Chicago Natural History Museum Bulletin, 31(5):6-7.

68. Publication dates for names of the Anubis baboon. Journal of Mammalogy, 41 (3):402- 403.

69. Mammals of northern Colombia. Prelimi- nary report no. 8: Arboreal rice rats, a sys- tematic revision of the subgenus Oecomys, genus Oryzomys. Proceedings of the United States National Museum, 1 10:513-568.


70. On the South American small-eared zorro Atelocynus microtis Sclater (Canidae). Field- iana: Zoology, 39(44):505-523.

71. On the nomenclature of certain whales. Fieldiana: Zoology, 39(49):547-565.

72. "This is a mammal." Chicago Natural His- tory Museum Bulletin, 3 2(6): 3.


73. Suriname zoological expedition. Chicago Natural History Museum Bulletin, 33(4):3, 7-8.

74. Bats and their menus. Chicago Natural His- tory Museum Bulletin, 33(8):2-3, 5-8.

75. Evolution of Neotropical cricetine rodents (Muridae) with special reference to the phyl- lotine group. Fieldiana: Zoology, 46:1-524.


76. A systematic and zoogeographic account of the monkeys of the genus Callicebus (Cebi- dae) of the Amazonas and Orinoco River basins. Mammalia, 27(l):3-79.

77. [Review] Primates. Comparative Anatomy and Taxonomy. Vol. V, Cebidae, part B., A Monograph; Edinburgh University Press. American Journal of Physical Anthropolo- gy, 21(l):92-98.

78. [Review] Primates. Comparative Anatomy and Taxonomy. Vol. V, Cebidae, part B., A Monograph; Edinburgh University Press. American Journal of Physical Anthropolo- gy, 2 1(3):39 1-398.

79. Notes on South American dolphins of the genera Inia, Sotalia and Tursiops. Journal of Mammalogy, 44(1):98-103.

80. The nomenclature of South American pec- caries. Proceedings of the Biological Society of Washington, 76:85-88.

81. The Recent mammals of South America. Proceedings of the XVI International Con- gress of Zoology, Washington, D.C., Aug. 20-27, 1963.

82. Comments on the proposed suppression of Zorilla I. Geoffroy, 1826. Z.N.(S.) 758. Bul- letin of Zoological Nomenclature, 20(4):242- 244.


83. Primate research and systematics. Science, 147(3662):1 156-1 157.

84. The importance of taxonomy in primate re- search and care. Illinois Society for Medical Research— Chicago Branch— Animal Care Panel Bulletin, 39:2 pp.


85. Catalog of living whales. Bulletin of the United States National Museum, 246: 1-259.


86. Taxonomic notes on tamarins, genus Sa- guinus (Callithricidae, Primates), with de- scriptions of four new forms. Folia Prima- tologica, 4:381-395.

87. On the identification of some marmosets, family Callithricidae (Primates). Mamma- lia, 30(2):327-332.

88. What ever happened to hairy man? Science, 153:362.

89. Comments on the proposal for conservation oi Pan Oken, 1816, and Panthera Oken, 1816. Bulletin of Zoological Nomenclature, 23(2/3):67-69.

90. [Review] Evolutionary and Genetic Biology of Primates, vol. II; Academic Press. Amer- ican Biology Teacher, 28(7):564-565.

91. Comments on the proposed suppression of Meles montanus Richardson, 1829, and M. jeffersonii Harlan, 1825. Z.N.(S.) 1639. Bul- letin of Zoological Nomenclature, 22(5/6): 336-337.

92. On the status of Procyon brachyurus Wieg- mann and P. obscurus Wiegmann. Z.N.(S.) 1640. Bulletin of Zoological Nomenclature, 22(5/6):338.

93. South American swamp and fossorial rats of the Scapteromyine group (Cricetinae, Mu- ridae) with comments on the glans penis in murid taxonomy. Zeitschrift fiir Saugetier- kunde, 31(2):81-149.

94. Status of the black-footed ferret in Wyo- ming. Journal of Mammalogy, 47(2):346- 347.

95. Comments on the proposal on Zorilla by Dr. Van Gelder and the counter proposal by Dr. China. Z.N.(S.) 758. Bulletin of Zoological Nomenclature, 2 3(2/3): 74-7 5.

96. Museum taxonomy serves medical research. Bulletin of the Field Museum of Natural History, 37(9):4-7.

97. Mice, land bridges and Latin American fau- nal interchange, pp. 725-751. In Wenzel, R. L., and V. J. Tipton, eds.. Ectoparasites of Panama. Field Museum of Natural History, Chicago.


98. (Review] Evolutionary and Genetic Biology of Primates, vol. I; Academic Press. Amer- ican Biology Teacher, Nov. 1967:665.

99. Reply to Mayr's comment on the proposed preservation oi Pan from Oken, 1816. Z.N.(S.) 482. Bulletin of Zoological Nomen- clature 24(5): 1 p.

1 00. Dynamics of rodent molar evolution: A study based on New World Cricetinae, family Mu- ridae. Journal of Dental Research, Suppl. to 46(5):829-842.


101. Metachromism or the principle of evolu- tionary change in mammalian tegumentary colors. Evolution, 22(3):556-575.

102. [Review] Dynamics of rodent molar evolu- tion: A study based on New World Cricet- inae, family Muridae. Oral Research Ab-