FIEL m .± NA Zoology NEW SERIES, NO. 39 STUDIES IN NEOTROPICAL MAMMALOGY Essays in Honor of Philip Hershkovitz Edited by Bruce D. Patterson and Robert M. Timm December 31, 1987 Publication 1382 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY Information for Contributors to Fieldiana . and researc n.anala'il'is tVoin : ^pace permits. The Journal Ccirries a page charge ot $63 iiaction ihcivot. L oiiinnuiions iioiu suuT, research associales. and invited authors will be con- uion regardless of ahiliix to pav paee charges, but the full charge is mandatory for nonaffiliated authors of unsohcued manuscripts. Pav ; 'f page charges qualifies a paper for expedited p. mg. which reduces the publication time. Manuscripts should be submitted to Dr. Timothy Plowman. Scientific Editor, Fieldiana, Field Museum of Natural History, Chicago, Illinois 60605-2496, USA. 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Chanecs in page STUDIES IN NEOTROPICAL MAMMALOGY Essays in Honor of Philip Hershkovitz Phiup Hershkovttz FIELDIANA Zoology NEW SERIES, NO. 39 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 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY HELDIANA: ZOOLOGY 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 ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA 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 V 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 VI Preface 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- PATTERSON: BIOGRAPHY OF PHILIP HERSHKOVITZ 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 HELDIANA: ZOOLOGY 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 PATTERSON: BIOGRAPHY OF PHILIP HERSHKOVITZ 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. 1940 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. 1941 7. The South American harvest mice of the ge- nus Reithrodontomys. Occasional Papers, Museum of Zoology, University of Michi- gan, 441:1-7. FIELDIANA: ZOOLOGY 1944 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. 1945 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). 1947 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. 1948 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. 1949 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. 1950 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. 1951 24. Mammals from British Honduras, Mexico, Jamaica and Haiti. Fieldiana: Zoology, 31(47):547-569. 1953 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. 1954 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. PATTERSON: BIOGRAPHY OF PHILIP HERSHKOVITZ 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. 1955 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. 1956 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. 1957 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. 1958 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. FIELDIANA: ZOOLOGY 59. The metatarsal glands in white-tailed deer and related forms of the Neotropical region. Mammalia, 22(4): 5 3 7-546. 1959 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. 1960 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. 1961 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. 1962 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. 1963 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. 1965 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. 1966 85. Catalog of living whales. Bulletin of the United States National Museum, 246: 1-259. PATTERSON: BIOGRAPHY OF PHILIP HERSHKOVITZ 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. 1967 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. 1968 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- stracts, May 1968. 1969 103. Comments on Cynocephalus Boddaert ver- sus Galeopithecus Pallas. Z.N.(S.) 1 792. Bul- letin of Zoological Nomenclature, 25(6):202- 203. 1 04. The evolution of mammals on southern con- tinents. VI. The Recent mammals of the Neotropical Region: A zoogeographic and ecological review. Quarterly Review of Bi- ology, 44(1): 1-70. 1970 105. The decorative chin. Field Museum of Nat- ural History Bulletin, 41(5):7-10. 106. Dental and periodontal diseases and abnor- malities in wild-caught marmosets (Pri- mates— Callithricidae). American Journal of Physical Anthropology, 32(3):377-392. 107. [Review] Taxonomy and Evolution of the Monkeys of Celebes (Primates: Cercopithe- cidae); Bibliotheca Primatologica, No. 10; Karger. Folia Primatologica, 13(l):75-76. 108. Metachromism like it is. Evolution, 24(3): 644-648. 1 09. Notes on Tertiary platyrrhine monkeys and description of a new genus for the Late Mio- cene of Colombia. Folia Primatologica, 12: 1-37. 110. Errata: Notes on Tertiary platyrrhine mon- keys and description of a new genus for the Late Miocene of Colombia. Foha Primato- logica, 12:1-37(1970). 111. Cerebral fissural patterns in platyrrhine monkeys. Folia Primatologica, 13:213-240. 112. [Review] The Squirrel Monkey; Academic Press. Journal of Mammalogy, 51(4):836- 839. 113. Supplementary notes on Neotropical Ory- zomys dimidiatus and Oryzomys hammondi (Cricetinae). Journal of Mammalogy, 51(4): 789-794. FIELDIANA: ZOOLOGY 1971 1977 1 14. Stapedial processes in tympanic cavities of capuchin monkeys (Cebus). Journal of Mammalogy, 52(3):607-609. 115. Basic crown patterns and cusp homologies of mammalian teeth, pp. 95-150. In Dahl- berg, A. A., ed., Dental Morphology and Evolution. University of Chicago Press, Chi- cago. 116. A new rice rat of the Oryzomys palustris group (Cricetinae, Muridae) from north- western Colombia, with remarks on distri- bution. Journal of Mammalogy, 52(4):700- 709. 126. Comment: Pan and Panthera or Oken's Lehrbuch? Z.N.(S.) 482. Bulletin of Zoolog- ical Nomenclature, 33(3/4): 135-1 36. 127. [Review] Catalogue of Primates in the Brit- ish Museum (Natural History). I. Families Callitrichidae and Cebidae; British Museum (Natural History). Folia Primatologica, 28: 315. 128. Living New World Monkeys (Platyrrhini). With an Introduction to Primates. Volume I. University of Chicago Press, Chicago, xiv +1117 pp. 1972 1 1 7. Notes on New World monkeys. Internation- al Zoo Yearbook, 12:3-12. 118. The Recent mammals of the Neotropical Region: A zoogeographic and ecological re- view, pp. 31 1-431. In Keast, A., F. C. Erk, and B. Glass, eds.. Evolution, Mammals and Southern Continents. State University of New York Press, Albany. 1974 119. The ectotym panic bone and origin of higher primates. Folia Primatologica, 22:237-242. 120. A new genus of Late Oligocene monkey (Cebidae, Platyrrhini) with notes on post- orbital closure and platyrrhine evolution. Folia Primatologica, 21:1-35. 1975 121. [Review] Taxonomic Atlas of Living Pri- mates; Academic Press. American Journal of Physical Anthropology, 41:155-156. 122. The scientific name of the \tsstv Noctilio (Chiroptera), with notes on the chauve-sou- ris de la Vallee d'Ylo (Peru). Journal of Mammalogy, 56(l):242-247. 123. Comments on the taxonomy of Brazilian marmosets (Callithrix, Callitrichidae). Folia Primatologica, 24:137-172. 1976 124. The taxonomic status of """Noctilio ruber Rengger." Mammalia, 40(1): 164-166. 125. Comments on generic names of four-eyed opossums (family Didelphidae). Proceed- ings of the Biological Society of Washington, 89(23):295-304. 1979 1 29. Races of the emperor tamarin, Saguinus im- perator Goeldi (Callitrichidae, Primates). Primates, 20(2):277-287. 1 30. The species of sakis, genus Pithecia (Cebi- dae, Primates), with notes on sexual dichro- matism. Folia Primatologica, 31:1-22. 1981 131. Comparative anatomy of platyrrhine man- dibular cheek teeth dpm4, pm4, ml with particular reference to those oT Homunculus (Cebidae), and comments on platyrrhine origins. Folia Primatologica, 35:179-217. 132. Philander and four-eyed opossums once again. Proceedings of the Biological Society of Washington, 93(4):943-946. 1982 133. Supposed squirrel monkey affinities of the late Oligocene Dolichocebus gaimanensis. Nature, 298(5870):20 1-202. 134. Subspecies and geographic distribution of black-mantle tamarins Saguinus nigricollis Spix (Primates: Callitrichidae). Proceedings of the Biological Society of Washington, 95(4):647-656. 135. Neotropical deer (Cervidae). Part I. Pudus, genus Pudu Gray. Fieldiana: Zoology, n.s., 11:1-86. 136. The staggered marsupial lower third incisor (I3), pp. 191-200. In Buffetaut, E., P. Jan- vier, J. C. Rage, and P. Tassy, eds., Phylo- genie et Paleobiogeographie. Livre jubiliare en I'honneur de Robert Hoffstetter. Geobios, memoire special 6, Lyon. PATTERSON: BIOGRAPHY OF PHILIP HERSHKOVITZ 1983 137. Two new species of night monkeys, genus Aotus (Cebidae, Platyrrhini): A preliminary report on Aott4s taxonomy. American Jour- nal of Primatology, 4:209-243. 138. On the validity of the family-group name Callitrichidae (Platyrrhini, Primates). Mam- malia, 48:153. 1 39. Taxonomy of squirrel monkeys, genus Sai- miri (Cebidae. Platyrrhini): A preliminary report with description of a hitherto un- named form. American Journal of Prima- tology, 6:257-312. 140. [Review] Mammalian Biology in South America. M. A. Mares and H. H. Genoways, eds. Ecology, 65(6): 1944-1 945. 1985 141. 1986 142. A preliminary taxonomic review of the South American bearded saki monkeys, genus Chi- roptes (Cebidae, Platyrrhini), with the de- scription of a new subspecies. Fieldiana: Zo- ology, n.s., 27:1-46. [Review] Handbook of Squirrel Monkey Re- search. L. A. Rosenblum and C. L. Coe, eds. Quarterly Review of Biology, 61:286-287. 143. The piebald saki. Field Museum of Natural History Bulletin, 57(2):coverplate + 24-25. 1987 144. Uacaries, New World monkeys of the genus Cacajao (Cebidae, Platyrrhini): A prelimi- nary taxonomic review with the description of a new subspecies. American Journal of Primatology, 12:1-53. 1 45. First South American record of Coue's marsh rice rat, Oryzomys couesi. Journal of Mam- malogy, 68(1): 152-1 54. 146. The titi. Field Museum of Natural History Bulletin, 58(6): 11-15. 147. The taxonomy of South American sakis, ge- nus Pithecia (Cebidae, Platyrrhini): A pre- liminary report and critical review with the description of a new species and a new sub- species. American Journal of Primatology, 12:387-468. In Press 148. More on the Homunculus Dpm4 and ml and comparisons with Alouatta and Stirto- nia (Primates, Platyrrhini, Cebidae). Amer- ican Journal of Primatology. 10 HELDIANA: ZOOLOGY A History of the Recent Mammalogy of the Neotropical Region from 1492 to 1850 Philip Hershkovitz ABSTRACTS The history of Neotropical mammalogy began with the first voyage of Christopher Colum- bus in 1492. The earliest notices were purely anecdotal, recorded by Spanish chroniclers from the mouths of the sailors on their return from voyages of discovery during the 1 5th and 1 6th centuries. Colonization of the Guianan and Brazilian coasts during the 1 7th century provided opportunities for inventories and descriptions of the mammals by trained European naturalists and physicians. The systematization and scientific naming of the known Brazilian species by Carolus Linnaeus in 1758 were based primarily on the mammals described in the 17th century monograph of Brazilian biota by Georg Marcgraf The actual collection and preservation of mammals for study, however, began in the 18th century with the Brazilian-bom Alexandre Rodrigues Ferreira. The 18th and first half of the 19th century was an explosive and romantic period of independently or govemmentally sponsored scientific expeditions for field observa- tions, collections, preservations, and taxonomic studies of the specimens shipped to European museums and private collectors. Outstanding among the naturalists who made significant con- tributions to mammalogy during this period are Alexander von Humboldt, Johann Baptist Ritter von Spix (Brazil), Maximilian Prinz Wied-Neuwied (Brazil), Johann Natterer (Brazil), Sir Robert Herman Schomburgk and Richard Schomburgk (Guyana), Claudio Gay (Chile), Johann Jakob von Tschudi (Peru), Felix de Azara (Paraguay), Rudolph Rengger (Paraguay), Alcide Charles Victor d'Orbigny (Argentina, Bolivia), and Charles Robert Darwin (Patagonia and Galapagos). Their itineraries, collections of mammals, taxonomies, and some field notes are included in the accounts of these and other noteworthy naturalists. By the middle of the 1 9th century, the mammalian fauna of South America became the best known of any continent with exception of the western European part of Eurasia. The problems of origins and distribution of Neotropical mammals intrigued scholars from among the earliest chroniclers down to pre- evolutionary Darwin. Their concepts on these subjects are briefly discussed. La historia de mastozoolo^a neotropical empieza con el primer viaje transatlantico de Cris- tobal Colon en 1492. Poco despues de desembarcarse de sus viajes de regreso los descrubridores y conquistadores del Mundo Nuevo en los siglos quince y diez y seis contaron a los cronistas espaiioles de las cosas curiosas que encontraron. Colonizacion de las costas guyanas y brasileras durante el siglo diez y siete ofrecio oportunidades a los naturalistas y medicos europeos residentes para le van tar inventarios de los mamiferos y anotar y hacer informes sobre sus observaciones. La sistematizacion y el nombramiento cientifico de las especies brasileras conocidas por Carolus Linnaeus en 1758 fueron basadas primariamente sobre los mamiferos descritos y figurados por Jorge Marcgraf en su monografia del siglo diez y siete. La coleccion y preservacion efectiva de mamiferos para el estudio empezo en comienzos del siglo diez y ocho con el "Viajem Filosofica" de Alejandro Rodriguez Ferreira, brasilero de nacimiento. From the Division of Mammals, Department of Zo- ology, Field Museum of Natural History, Chicago, Illi- nois 60605-2496. HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 1 1 El siglo diez y ocho y las primeras decadas del siglo diez y nueve senalaron un periodo explosive y romantico de expediciones cientificas fomentadas por gobiemos europeos, o por naturalistas particulares con los objectos de hacer observaciones sobre la fauna, tomar notas de campo, y recoger y preservar ejemplares para estudios taxonomicos en los museos extranjeros. Entre los naturalistas europeos que hicieron contribuciones de consequencia a la mastozoologia neo- tropical en este epoca, se cuentan Alejandro von Humboldt, Juan Baptista Ritter von Spix (Brasil), Maximiliano Principe de Wied-Neuwied (Brasil), Juan Natterer (Brasil), Lord Roberto Herman Schomburgk y Ricardo Schomburgk (Guyana), Claudio Gay (Chile), Juan Jacobo von Tschudi (Peru), Felix de Azara (Paraguay), Rodolpho Rengger (Paraguay), Alcidio Carlos Victor d'Orbigny (Argentina, Bolivia), y Carlos Roberto Darwin (Patagonia y Galapagos). Compren- dido en este informe son los itinerarios, listas de mamiferos coleccionados y observados, taxonomias, y algunas experiencias de campo de los naturalistas mentados, y de otros digno de atencion. A mediados del siglo diez y nueve, la fauna mamifera de Sud America llego a ser la mejor conocida de todos los continentes del mundo menos Europa. Problemas de origen y reparticion geografica de los mamiferos del Mundo Nuevo estimularon la imaginacion de sabios desde los primeros cronistas del Descubrimiento hasta el joven Darwin pre-evolutionario. Los conceptos sobre estos temas son brevemente discutidos. A historia da mastozoologia neotropical, come90u com a primeira viagem transatlantica de Cristovao Colombo, em 1492. Os primeiros relatorios, puramente anedotais, foram registrados pelos cronistas espanhois, logo apos o regresso das viagens de descobrimento durante os seculos XV e XVI. As colonizacoes da costa Guianense e Brasileira durante o seculo XVII, ofereceram amplas oportunidades a naturalistas e medicos, de treinamento Europeu, para inventoriar e descrever os mamiferos encontrados. A sistematica e a nomenclatura cientifica das especies Brasileiras conhecidas por Carolus Linnaeus em 1758 basearam-se primariamente nos ma- miferos descritos por Georg Marcgraf, em sua monografia do seculo XVII. No entanto, as colecoes e preservacoes de mamiferos para estudos come^aram, efetivamente, no seculo XVIII, com a "Viajem filosofica" do Brasileiro, Alexandre Rodrigues Ferreira. Marcaram o seculo XVIII, e as primeiros decadas do seculo XIX, um periodo explosivo e romantico nas expedicoes cientificas. Estas foram patrocinadas tanto por naturalistas indepen- dentes, como por govemos Europeus, a fim de realizarem observacoes sobre a fauna e colecoes para estudos taxonomicos nos museus Europeus. Entres os naturalistas Europeus que distin- guiram-ce em suas contribuicoes aos estudos de mamiferos neotropicais durante esta epoca, sobressaem: Alexandre von Humboldt, Juan Baptista Ritter von Spix (Brasil), Maximilian Principe de Wied-Neuwied (Brasil), Johan Natterer (Brasil), Sir Robert Herman Schomburgk e Richard Schomburgk (Guiana), Claudio Gay (Chile), Johan Jakob von Tschudi (Peru), Felix de Azara (Paraguai), Rudolph Rengger (Paraguai), Alcides Charles Victor d'Orbigny (Argentina, Bolivia) e Charles Robert Darwin (Patagonia e Galapagos). Os itinerarios, as listas de mamiferos observados e colecionados, as taxonomias, e algumas notas de campo encontram-se incluidos nos relatorios aqui apresentados sobre estes e outros naturalistas importantes. Nas meadas do seculo XIX, a fauna mamifera sul-americana tomouse a melhor conhecida de todos OS continentes, exceto a da Europa. Os problemas de origem e da distribuicao geografica dos mamiferos neotropicais estimularam a imaginacao de varios estudiosos, desde os primeiros cronistas ate o pre-evolucionario Darwin. Seus conceitos sobre estes temas sao brevemente discutidos. Organization II. Voyages of Discovery: 1 5th and 16th Centuries 14 I. Introduction 13 III. Spanish Chroniclers of New The Neotropical Region Defined . . 14 World Discoveries 14 12 HELDIANA: ZOOLOGY IV. First Mammals: Anecdotal Period 16 Island Mammals of the Discoverers 16 Mainland Mammals of the Discoverers 18 V. Brazil: Mammalogy Through 1 8th Century 21 Andre Thevet (1503-1592) 21 Georg Marcgraf (1610-1644) 21 Alexandre Rodrigues Ferreira (1756-1815) 21 VI. Brazil: Mammalogy to Middle of 1 9th Century 27 Introduction 27 Johann Baptist Ritter von Spix (1781-1826) and Carl Friedrich von Martius (1794-1866) 27 Maximilian Prinz von Wied-Neu- wied (1782-1867) 31 Johann Natterer (1787-1843) .... 34 VII. GuiANAs: Mammalogy to End of 1 8th Century 38 Pierre Barrere (1690-1755) 38 Jose Gumilla (d. 1750) 38 Jacques Nicolas Bellin (1703-1772) 38 Edward Bancroft (1744-1821) .... 38 Philippe Fermin (1720-1790) .... 39 Monsieur Bajon (1763?) 40 John Gabriel Stedman (1744-1797) 40 VIII. GuiANAs: Mammalogy of First Half of 19th Century 43 Sir Robert Herman Schomburgk (1804-1865) and Richard Schomburgk (181 1-1891) 43 IX. Alexander von Humboldt ( 1769- 1859) AND AlME BONPLAND (1773-1858) 51 X. Paraguay 57 Felix de Azara (1746-181 1) 59 Johann Rudolph Rengger (1795-1832) 64 XI. Chile 64 Giovanni Ignazio Molina (1737-1829) 64 Eduard Friedrich Poeppig (1798-1868) 65 Claudio Gay (1800-1873) 65 XII. Peru 65 Johann Jacob von Tschudi (1818-1889) 65 XIII. Patagonia 71 Alcide Charles Victor d'Orbigny (1802-1857) 71 Charles Robert Darwin (1809-1882) 77 XIV. Georges Louis Leclerc de Buffon (1707-1788) 87 XV. Faunal Origins and Distribution 87 Jose de Acosta (1539-1600) 87 Antonio Vazquez de Espinosa (1560/1575-1630) 90 Carolus Linnaeus (1707-1778) ... 90 Georges Louis Leclerc de Buffon (1707-1788) 90 Johann Andreas Wagner (1797-1861) 91 Maximilian Prinz von Wied- Neuwied (1782-1867) 91 Johann Jacob von Tschudi (1818-1889) 91 Charles Robert Darwin (1809-1882) 91 XVI. Inventories to Middle of 1 9th Century 91 System Naturce of Linnaeus, 1758, 1766 91 Histoire Naturelle of Buffon, 1750-1789 92 Synopsis Mammalium of Schinz, 1844 92 XVII. Summary 92 XVIII. Acknowledgments 94 XIX. Literature Cited 94 I. Introduction The gradual accumulation of knowledge of Neotropical mammals is recorded here from the time of the first voyage of discovery by Christo- pher Columbus in 1492 to the middle of the 19th century, or just before the Darwinian revolution in biological thought. The knowledge was mainly of species or kinds, the numbers of kinds, their behavior, habitat, geographic distribution, and re- lationship to man. Early voyagers to the New World followed by naturalist-travelers gathered the data used later by philosophers and scientists for the development of biological principles. Only the most important and better-known contributors are discussed here. At least as many more personages could be included in a more extended account. HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 13 The Neotropical Region Defined The Neotropical Region, as defined by its mam- malian fauna, includes all South America, Middle America except the dry and temperate zones of Mexico, continental islands of coastal Middle and South America, and the oceanic Bahamas, West Indies, Galapagos, and Falklands (Hershkovitz, 1972, p. 326). With few exceptions, modem names for Neo- tropical countries and geographic features are used throughout the text. The map (fig. 1) shows the South America of the colonial period with colonial or precolonial names for political subdivisions and geographic features. II. Voyages of Discovery: 15th and 16th Centuries The inhabited islands found by Columbus on his first voyage across the Atlantic Ocean in 1492 were thought to be near the mainland of China or India. The islanders welcomed the ships' crews with food and drink, but the great stores of pre- cious metals, stones, and artifacts the travelers ex- pected to find were not seen. Nevertheless, the voyagers claimed the islands for the Spanish crown and returned with accounts told to awaiting re- porters of their discoveries, including their de- scriptions of plants and animals of economic value or imputed medicinal virtues. Zoological results of the four transatlantic voy- ages commanded by Christopher Columbus— the first (1492-1493) and second (1493-1496) to the Antilles, the third (1498) to the Antilles and Ven- ezuela, and the last (1 502-1 504) to Middle Amer- ica—included reports of a variety of mammals. The kinds seen were identified with such familiar Old World forms as lion, tiger, bear, fox, dog, ferret, rabbit, deer, boar, goat, sheep, rodent, mon- key, and ape. Characterizations given were less descriptions of external morphology than of gen- eral mien, gross habitat, behavior in response to human confrontations or predation on human property, gastronomic qualities, and use, if any, in medical treatment, ceremonial rites or magic, or as household pets. Those who followed Columbus in the discovery and exploration of the mainland returned with additional bits of information on mammals noted by the attendant Spanish chroniclers. Among the more important of these voyagers of discovery were Pinzon, who followed Columbus to the Ven- ezuelan coast in 1 500, and Amerigo Vespucci, who sailed first with Ojeda to Brazil in 1499 and in- dependently again in 1 502 and 1 503 in the service of Portugal. Pedro Cabral, however, had already claimed Brazil for Portugal in 1 500 on his way to India. In 1516 Juan Diaz de Solis discovered the estuary of the Rio de la Plata, and Sebastian Cabot, in the service of Spain, sailed in 1526 up the Rio Parana. Vasco Nunez de Balboa accompanied En- ciso to Panama in 1510, and in 1513, with Fran- cisco Pizarro, crossed the isthmus to behold the vast Pacific Ocean. Pizarro visited Panama again in 1531, recrossed the isthmus, and sailed south along the west coast of South America to the dis- covery and conquest of Peru. Cabeza Alvarez Nu- nez de Vaca arrived in Buenos Aires in 1541 and continued overland into Paraguay. Pedro de Val- divia visited Venezuela in 1 530, Peru in 1 532, and Chile in 1540, 1541, and 1552. The explorations of Colombia by Gonzalez Jimenez de Quesada from 1536 to 1539 and again in 1569 to 1571 signaled the end of the period of discovery and conquest. III. Spanish Chroniclers of New World Discoveries The recorders or chroniclers of New World dis- coveries, conquests, happenings, and natural phe- nomena were the clerics and scribes who accom- panied the explorers or awaited their return to Spain for recording the news. Most of the accounts or records remained unpublished, but some of the manuscripts are reportedly preserved in the ar- chives of Spain or the Vatican. The chroniclers whose published narratives contain interesting in- formation on mammals include the following. Peter Martyr of Anghiera (1455-1526), Italian by birth, and the first and most prestigious chron- icler of the Discovery, was a member of the Royal Spanish Council of the Indies, Prothonotary of the Catholic Church, correspondent of Popes, confi- dant of Christopher Columbus, and friend of sea captains, clergymen, and other contemporary voy- agers to the New World. News he received from his informants constitutes the first records of New World discoveries. His chronicles, known as the Decades and addressed to the Pope, began to ar- rive at the Vatican in 1 494. The first Decade de Orbe Novo, with first notices of American mam- mals, was published in 1516, but pirated Italian 14 HELDIANA: ZOOLOGY Fig. 1. Map, South America of the Colonial period from the Stevens (1726) translation of Herrera y Tordesillas. editions appeared in 1504 and 1507. The fourth Decade was published in 1521, and the complete set of eight of the projected 10 appeared posthu- mously in 1587. Gonzalo Fernandez de Oviedo y Valdes (1478- 1557) was appointed royal chronicler of news sent directly to him by provincial governors and other New World officials. Included were Oviedo's own observations and results of investigations during his residence as representative of the Spanish Crown in the Provinces of Darien, Panama, Gua- temala, Cuba, and Santo Domingo. He published HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY IS the first part of his Historia de las Indias in 1 526, other parts in 1535 and 1 547. The entire work was printed between 1851 and 1855 in Madrid. The Spanish Jesuit Jose de Acosta ( 1 539-1 600) wrote his Historia Natural y Moral de las Indias during a residence in Peru from 1571 to 1 587 and saw it published in 1 590. Acosta's philosophical inquiries extended to all asjjects of nature in the New World and greatly influenced the thinking of his contemporaries. Antonio Vazquez de Espinosa (b. between 1560 and 1575, d. 1630), a Carmelite missionary, lived many years in Spanish America, most of them in Peru and Mexico. His natural history notes are compiled from many sources, including his per- sonal observations and testimony of people he met in travels connected with his clerical duties. The forgotten manuscript of his Compendium was dis- covered in the Vatican library by Charles Upson Clark in the early part of the 20th century. Clark's English translation of the work was published in 1 942 by the Smithsonian Institution, and his tran- scription of the original Spanish in 1 948 by the same institution. Antonio de Herrera y Tordesillas (1559-1625), historiographer to the King of Spain, compiled the General History of the Vast Continent and Is- lands of America from archived reports by the New World discoverers and conquistadores, gov- ernors, clergy, colonists, and travelers. He also borrowed heavily from published accounts, in- cluding those of other chroniclers. There is no in- dication that his notices on mammals were based on personal observations. The first edition of Herrera's History was published in 1601, another in 1 60 1-1 6 1 5. These and a 1 728 Spanish edition in the Library of Congress are cited in the bibliography. I have not seen these works. The Stevens translation, published 1725-1726, was used here. Whatever the quality of the translation, I find no fault with the descriptions of mammals, and the stories about them are in line with similar accounts in other sources. IV. First Mammals: Anecdotal Period Island Mammals of the Discoverers The first Columbian voyage, in 1492, resulted in the discovery of the Antillean islands of Cuba, Hispaniola, and part of the Bahaman Archipelago. According to Peter Martyr, who reported results of the voyage in his first Decade (1504, 1516), quadrupeds were not seen, but three kinds of "rabbits" were said to occur in Hispaniola (Haiti and Dominican Republic). The same animals, ac- tually caviomorph rodents, were described later by Oviedo during his residence in Santo Domingo. The following accounts are freely translated or paraphrased from the Spanish of the Paraguayan (1944-1945) edition of Oviedo's work. Hutia, the first "rabbit" (1944, libro XII, cap. I), is smaller than the ordinary rabbit, its ears smaller and tail ratlike. The hutia is said to be dark grayish in color and very good eating. It was hunted and killed by the barkless dogs of the na- tives, but is no longer found, except rarely. Gerrit S. Miller (1929, p. 12), studied the re- mains of mammals in kitchen middens of the Sa- mana Bay region, Dominican Republic, and con- cluded that the original description of the hutia "would apply as well to the species of Plagiodon- tia, and presumably also to the Isolobodons [sic] that there seems to be no reason to doubt that these were the animals Oviedo had in mind." The quemi, second of the "rabbits" (1944, libro XII, cap. II), is said to be blackish like the hutia and similar in form, but larger like an ordinary hound. Natives of the island who saw and ate the animal found it savory. Oviedo believed them ex- tinct. All attributions to the quemi, according to Mil- ler ( 1 929, p. 13), agree with those of a "large rodent whose remains I found in the caves near St. Mi- chel, Haiti, in 1925. Consequently, I proposed for it the generic name Quemisia. The presence of the same creature in the Boca del Infiemo kitchen midden appears to confirm my guess." The mohuy "rabbit" (1945, libro XII, cap. Ill), is smaller than the hutia, a paler brown or grayish in color, its flesh highly esteemed by the island's caciques and noblemen. The pelage, unlike that of the hutia, is stiff", sharply pointed, and erect. Ovie- do saw no mohuy, but knew persons on the island who did and reportedly regarded its flesh as better than that of the other "rabbits." "There be little if any doubt," says Miller ( 1 929, p. 13), "that the animal Oviedo thus described was Brotomys voratus ... its remains have been found in every kitchen midden that has been ex- amined in the Dominican Republic. . . . The ac- count of stiff", pointed, erect-standing hairs of the back seems especially applicable to a relative of the South American spiny-rats." The cori, a fourth "rabbit," described by Oviedo 16 HELDIANA: ZOOLOGY (1945, libro XII, cap. IV), is almost certainly the domestic guinea pig. Miller (1929, p. 14) ques- tioned whether the guinea pig was pre-Columbian or a Spanish introduction. He inclined to the sec- ond alternative "chiefly because remains of the animal have been found in only one midden." It appears, however, that one Simone Verde, who accompanied Columbus on his first voyage, men- tioned in a letter dated 20 March 1494 (cf. Martyr in Gaffarel, 1907 trans, p. 12, footnote 2; p. 14, footnotes 1 , 2) the existence on the island of a black and white dormouse-like animal without tail. The guinea pig or cui, domesticated in Peru, was carried by pre-Columbian Indians for food and barter and introduced into islands and many parts of mainland South America where cavies do not naturally occur. Many of them, such as completely isolated colonies I saw in Colombia near Bogota, had become feral, their coloration having reverted to the wild or agouti pattern. Other Hispaniolan mammals mentioned by Oviedo are the barkless domestic dogs and house rats, the latter certainly brought by the Spaniards. Apart from the extinct insectivore Nesophontes, Miller found no remains of mammals the size of mice or rats in kitchen middens or owl pellets. Two additional native West Indian mammals observed by Oviedo in 1 523 or 1 524 in Cuba differ from those of Hispaniola. My paraphrased trans- lation of Oviedo's Spanish descriptions follows. The guabiniquinax is somewhat larger than a rabbit, its feet similar, the tail long and ratlike, the pelage smoother than that of a badger, the skin white, the flesh savory. It lives and breeds in the mangroves along the coast. To capture it, the In- dians position their canoes beneath the mangroves where it nests, then shake the tree to cause the animal to fall into the water where it is seized. The animal as described above is certainly a form of Capromys, but Oviedo continues as fol- lows: "The animal is the size of a hare, looks like a fox, its color is dark brown mixed with reddish, the tail hairy and the head shaped like that of a ferret. It abounds along the Cuban coast." The characterization and habitat are obviously out of place and probably were meant to be included with the description of the ayre, the second of the Cu- ban mammals reported by Oviedo. Herewith my paraphrased translation of his description of that animal. The ayre is reddish brown, the size of a rabbit with pointed muzzle, its flesh exceedingly tough. Notwithstanding, the natives cook or roast as many of the animals as they can capture, for they are abundant. But no matter how long the meat may be cooked or roasted, it is no less tough to chew. This characterization seems to fit the insectivore Solenodon. On the other hand, the flesh of Cap- romys, as of most if not all caviomorphs, is tender and, as a rule, delectable. From his correspondents Oviedo received no- tice of still another mammal, the guacabitinax, an inhabitant of the islands near those of Las Perlas in the Golfo de San Miguel and the Isla de las Culebras or Gorgona, off" the southwest coast of Colombia. The name, not to be confused with the preceding, and the description and details of the animal's habits, are unmistakably those of the paca {Agouti paca Linnaeus). Manatees sighted at sea at various times by Co- lumbus and his men were believed to be mer- maids, albeit ugly ones. Martyr's narrative of a captive manatee as given in the available French translation of his third Decade (Gaffarel, 1907) is composite. The account by Herrera of the same manatee (in Stevens's translation, 1725, vol. 1, p. 278) appears to hew closer to the original source of information: The Spaniards at this Time found a new Sort of Fish, which was a considerable ad- vantage to them; tho' in those Parts there is much Variety. It is call'd Manati, in shape like a Skin they use to carry Wine in, having only two Feet at the Shoulders, with which it swims, and it is found both in the Sea and in Rivers. From the Middle it sharpens off" to the Tail, the Head of it is like that of an Ox, but shorter, and more fleshy at the Snout; the Eyes small, the Colour of it grey, the Skin very hard, and some scattering Hairs on it. Some of them are twenty Foot long, and ten in Thickness. The Feet are round, and have four Claws on each of them. The Females bring forth like the Cows, and have two Dugs to give suck. The Taste of it is beyond Fish; when fresh it is like Veal, and salted like Tunny-Fish, but better, and will keep longer; the Fat of it is sweet, and does not grow rusty. Leather for Shoes is dress'd with it. The Stones it has in the Head are good against the Pleurisy and the Stone. Sometimes they are taken ashore, grazing near the Sea, or Rivers, and when young they are taken with Nets. Thus the Cazique Caramestex took one, and fed it twenty-six Years in a Pond, and it grew sensible and tame, and would come when call'd by the Name of Mato, HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 17 which signifies Noble. It would eat what- soever was given it by Hand, and went out of the Water to feed in the House, would play with the Boys, let them get upon him, was pleas'd with Musick, carry'd Men over the Pool, and took up ten at a Time, without any Difficulty. Martyr's third Decade mentions many "rab- bits" and deer encountered in 1 5 1 6 by Andre Mo- rales on the forested Isla Rica (now San Jose) of the Archipielago de las Perlas in the Golfo de Pa- nama. The deer, most likely Mazama gouazoubira permira Kellogg, 1 946, and rabbits (Sylvilagus sp.) were said to be so abundant that Spaniards could shoot them with arrows from horseback. The re- tiring tapeti, Sylvilagus brasiliensis, the only rabbit known from mainland Panama and the Pearl Is- lands, would have been an unlikely target for the equestrian Spaniards. Mainland Mammals of the Discoverers First knowledge of mainland American mam- mals was contained in reports of the Paria Pen- insula, Venezuela, by Columbus on his third trans- atlantic voyage in 1498 and Vicente Yaiiez Pinzon, who followed in the tracks of Columbus. Martyr's first Decade carried the news of their encounters with the common opossum (Didelphis marsupi- alis), sloths, armadillos, anteaters, deer (Odocoi- leus, Mazama), peccaries, tapir, kinkajou (Potos flavus), barkless dog {Canis familiaris), jaguar, puma and their color varieties, vampire(?) bats, and red howler monkey (Alouatta seniculus). On his fourth and last voyage (1 502-1 504), Co- lumbus explored the Atlantic coast of Middle America from the Golfo de Uruba to Guatemala. Spanish emissaries charged with establishing set- tlements followed quickly. Mammals reported by them and noted in Martyr's second and third Decades, and by Oviedo, include the common opossum, bats, monkeys, three-toed sloths, ant- eaters, armadillos {Dasypus novemcinctus), white- tail deer, red brocket, collared and white-lipped peccaries, squirrels, a composite of carnivorous species identified as raposas (including Didel- phis?), zorros (foxlike Camivora), lobos (Dusicyon or Lutra), rabbits (Sylvilagus brasiliensis), "hares" (Dasyprocta sp.. Agouti paca, and perhaps the newly introduced European hare or rabbit). The domes- tic dog, like that first seen in the Antilles, was barkless. An encounter with vampire bats by Pinzon's men is reported by Martyr in the first Decade. Vampires are also mentioned in the second De- cade in connection with Enciso's disastrous ex- periences in the Darien and in the third Decade in accounts of the animals of the Golfo de Uruba. The following characterization of a vampire bat by Herrera (in Stevens's translation, 1726, vol. 2, p. 7 1 ) is a translation from the original sources in Spanish. "This venomous Creature has one quality that tho' it bites one man among an hundred one Night, the next Time it only bites in the very same Place, tho' the Person bit be among two hundred; which it does either on the Toes, the Fingers, or the Head, and much Blood runs from it." That the same vampire bat should visit the same person sleeping in the same place on successive nights may not be unusual. An experience of mine in 1935 on the Rio Napo in Ecuador is of interest in this regard. Two Indian families and I, alto- gether 10 persons including a five-year-old girl, traveled three days upstream in a large dugout canoe. The river was low and we could bivouac on sandbars at the end of each day's travel. On each of the three nights, a vampire bat visited the little girl, scraped the skin of her nose, and fed on the trickling blood. No other member of the party was attacked. It seems improbable that the same bat should have found the same victim at each of the three different bivouacs. Perhaps the child slept more soundly than the others of the party, or her blood was more attractive to the vampires which abounded in the region. The last of Martyr's eight Decades includes de- scriptions of Spanish settlements in the Golfo de Paria, Venezuela. In addition to those mammals previously mentioned by Martyr (above) are the lesser anleater (Tamandua tetradactyla), capuchin monkey (Cebus apella or C. nigrivittatus), peccary, deer (Odocoileus virginianus), jaguar {Felis onca), spotted cats {Felis pardalis and/or F. wiedii), wea- sels (Mustela frenata), skunk (Conepatus chinga), porcupine (Coendou prehensilis), and manatee {Trichechus manatus). Oviedo described the same animals of the region in greater detail, but with no additions of sp>ecies. Vazquez de Espinosa, who in 1628 presumably visited the northern Venezuelan coast and the town of Santo Tomas above the mouth of the Rio Orinoco, reported the same mammals as well as squirrels (Sciurus aestuans) 18 FIELDIANA: ZOOLOGY and many kinds of monkeys. He claimed that Isla Margarita, off the Venezuelan coast, was overrun with rabbits {Sylvilagus floridanus). Many of the larger mammals of Colombia in the territories of the Muso and Colima Indians north of Bogota were already known by 1544. With bats and other small mammals omitted, more kinds were reported by Herrera than could be re- corded today from the same region on the basis of extant specimens preserved in museums. Her- rera, in the English version by Stevens (1726, vol. 6, p. 191), states: There are a great number of grey Swine [Tayassu pecan] that have the Navel on the Back, and a smaller sort of several Colours [Tayassu tajacu] much like wild boars. Ti- gers (Felis onca) not numerous but very fierce; Lions (Felis concolor) that do no harm, except only among the Cattle and two other sorts of Tigers that were inoffensive besides another sort that are always in the water, like Greyhounds, and all their four feet are like those of a Goose [Lutra annectens]. The black wild cats [Felis yagouaroundi] seize the Hens, carry them away under one of their front legs and run away on the other three. The black Bears [Tremarctos ornatus] like those in Spain, do no hurt but only to the small Cattle. The Ant-Bears [Myrme- cophaga tridactyla] when they go, lay their Tail, which is long, on their Heads, winding them about their Necks, and so walk from Ant-hill to Ant-hill, stretch out their Tongues near half a Yard which are soon cover'd with Pismires, then they draw them back and eat them. There are Dantas [Tapirus pinchaque or T. terrestris]. Deer [Odocoileus virgini- anus] like ours in Europe, and others red like wild Goats [Mazama rufina or Mazama americana], and the Bezoar stones found in them are best. The Guadatinajas [Agouti paca] are like Hares; and the Zorillas [Di- delphis marsupialis] or little foxes, that have a purse under their Belly, in which they carry their Cubs, the ever so many, are very mis- chievious to the Henroosts. The little Crea- tures call'd Umazia [Marmosa] have a dug growing out for every one of their young, and they stick to it till bred up. The Ar- madillo [Dasypus novemcinctus] which has been spoken of having five claws in each Forefoot, with which it throws up the Earth, is tame and eaten. The Perico Ligero [Bra- dypus variegatus] is three hours climbing a Tree, goes about in the Night, gives a cry every time it lifts a Foot, and is half an Hour, between every Step, is as big as a Barbary monkey, and fierce, yet does no harm. There are cats (?) that sleep all the Day, and all the Night catch Birds and Mice. The Pizma [Na- sua nasua] about as big as a large Lap Dog, has a bad countenance, a long Snout, its voice like a Bird, defends itself against Dogs, and the Spaniards call them Badgers. The Hedgehogs [spiny echimyids] are like those in Spain, the largest like Porcupines [Coen- dou sp.] darting out their Prickles. There are many sort of Apes, squirrels. Elsewhere, in the Province of Bogota, Herrera (in Stevens's translation, 1726, vol. 6, p. 77) notes "innumerable apes, monkeys, ferrets [marsupi- als?], squirrels, weasels [Mustela frenata], deer [Odocoileus virginianus], roebuck [Mazama rufi- na] and Rabbits [Sylvilagus brasiliensis] . . . but not Hares." Manatees were reported from the Rio Magdalena. From coastal Colombia, at Zaragoza, 30 leagues from Caceres in the lower Rio Cauca Valley, Vaz- quez de Espinosa records jaguar, puma, danta (Tapirus terrestris), oso (Myrmecophaga or Ta- mandua), cuchumbi (Nasua), armadillo (Dasy- pus), raposa (Dusicyon thous), chucha (Didelphis marsupialis), "three" species of sahinos or pec- cary, perico ligero (Bradypus variegatus), nutria (Lutra or Chironectes), and guadatinaja (Agouti paca). Acosta's long residence in Peru made him fa- miliar with some of the mammals in the vicinities of Cuzco and Lima and others about which he may have learned from travelers or records. He described sahinos (peccaries), dantas (tapir), ar- madillos, perico ligero (three-toed sloth), osos (anteaters), otoronco (bear), chinchilla, vizcacha, cui (guinea pig). The "liebres verdaderas" or true hares are certainly the introduced European hare. He affirmed that conejos or rabbits (Sylvilagus brasiliensis) occur in the Reino de Quito (Ecua- dor). Acosta declared there were monkeys of all kinds throughout America, but those he described were Middle American. At Capira near Nombre de Dios, Panama, he saw monkeys (presumably spider monkeys) swing by their tails from a tree on one side of a stream to another tree on the opposite HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 19 side. Where the river was too wide for this ma- neuver, the monkeys of the troop, he related, crossed by forming a hanging chain holding on to each other's tail, then swinging so that the endmost could grab the branch of a tree on the other side of the river and let all the others clamber up. The anecdote is less a fabrication than an ex- aggeration. Individual howlers and spider mon- keys, usually the alpha male or an old suckling female, may bridge a narrow gap in the canopy pathway by holding with its prehensile tail the branch on one side of the gap and with it swinging the body to catch, with outstretched arms, the nearest branch of the far side. Monkeys too small or weak to hurdle the gap run or scramble over the bridging back of their elder. I have seen strong young adults take advantage of the same conve- nience. Acosta also narrated the tale of a monkey that resided in the palace of a provincial governor. As related to him, the simian was trained to fetch wine from the town tavern. The animal would set off with the empty wine pitcher in one hand, the wine money gripped in the other. Not before the pitcher was filled to the brim did the sage monkey release his coins to the tavern keeper. There were times on these errands when taunting street ur- chins chased and hurled stones at the monkey. Annoyed by this sport, the simian halted, set down the pitcher, and returned the stones with sufficient force and accuracy to rout his tormentors. Retriev- ing his pitcher, he moved on serenely to deliver the wine at the palace. Peruvian "sheep" or camelids were greatly ad- mired by the Spaniards when first seen. Acosta's interesting account of them was suitably appre- ciated by Herrera, and the English translation by Stevens (1726, vol. 4, p. 36) is quoted herewith. There are no such Vicunas and Sheep in New Spain [Mexico] As those of Peru, and those Sheep are Tame, and very serviceable; but the Vicunas are wild, and have no Horns, the like of them not to be seen in the whole World, but only in Peru and Chile, bigger than Goats, but smaller than Calves, their Colour almost Murrey, breeding on the highest Mountains, in cold and desert Places, which they call Punas. They go in flocks, run swiftly, and when they see any Men, fly and drive their Young before them. Of their Wooll are made very valuable Mantles, which never lose their Colour, because it is natural; they are said to be good for Inflam- mations in the Kidneys, as are Quilts made of the Wooll, because they moderate the Heat, and the same in the Gout; and in them the Bezoar Stones are found. The abundance and ubiquity of llamas may have inspired some Spaniards to attempt to raise Old World camels in Peru. According to Acosta, some brought from the Canary Islands were bred for a while. Sebastian Cabot's journal of conquest and ex- ploration of the Province of Rio de La Plata, then consisting of modem northern Argentina, cisan- dean Bolivia, and southeastern Brazil, included data on natural history. As recorded by Herrera, the mammals seen were the hairy armadillo {Chaetophractus sp.) and several other kinds, ca- vies {Cavia), swamp deer (Blastocerus dichotomus), pampa deer {Blastoceros bezoarticus), brockets (Mazama sp.), tapirs {Tapirus terrestris), peccaries (any or all of the known species), howler monkeys (Alouatta), canids (Dusicyon), lesser anteater (7a- mandua tetradactyla), jaguar (Felis onca), and puma (Felis concolor). Southern Brazilian mam- mals in particular included deer, peccary, tapir, "rabbits" with small, round ears {Dolichotisl), paca {Agouti paca), armadillo, sloth (Bradypus torqua- tus), opossum {Didelphis albiventris), monkeys, and coastal seals, most likely Arctocephalus australis. Vazquez de Espinosa adds capybaras, armadil- los (tatu and quirquincho specified), and guanacos. In the vicinity of Chuquisaca (La Plata), Bolivia, the missionary notes brockets {Mazama), vicufia, guanaco, dark gray wildcats known as oscollos, jaguar called "otorongo," puma locally called poma, a large beast called lilisto with a horselike head that lures cattle and humans, a ferret called siqui {Mustela frenatal), skunks or anatiria {Co- nepatus), bear {Tremarctos ornatus), antbears (probably Tamandua), vizcacha {Lagidium), and cuis {Cavia porcellus). The occurrence of sea lions {Otaria flavescens) and fur seals {Arctocephalus) on both southern continental coasts was mentioned by Vazquez de Espinosa. The sea lions along the coast of Are- quipa, Peru, he reported come out of the water onto the rocks and make low sounds at night. The animals were hunted by the Indians for their hides. In northern Chile, the natives of Arua and Ata- cama converted the hides into balloon-like floats for support of their seagoing fishing rafts. The conquest of Chile by Pedro de Valdi via in 1 54 1 provided the chroniclers with additional in- formation on mammals. Vazquez de Espinosa re- 20 HELDIANA: ZOOLOGY ported huemul {Hippocamelus bisulciis), "fallow deer" (spotted fawns of huemul), guanaco, and vicuna in the vicinity of Osomo. According to the same authority, the Rio Guasco valley (29°S) har- bored "squirrels" (chinchillas) with very fine fur. V. Brazil: Mammalogy Through 18th Century Andre Thevet (1503-1592) The French missionary Andre Thevet arrived in 1555 in Rio de Janeiro, the principal port of a French colony in the ephemeral France Antarc- tique. Thevet returned to France via the Antilles a year later, and the accounts of his travels were published in 1557 or 1558. Father Thevet's cu- riosity about all he saw in the New World knew no bounds, and he became an avid collector of Indian artifacts, local birds, and insects. Not all objects and events described in his book con- formed to popular European prejudices or gen- erally accepted misconceptions. The work stirred up considerable debate and was rejected by many not prepared to accept the realities that opossums had pouches or that the bodies of American In- dians were not densely furred. The Brazilian mammals described or men- tioned by Thevet include the locally common opossum (Didelphis albiventris), tapeti (Sylvilagus brasiliensis), agouti {Dasyprocta leporina, declared good eating), peccaries, deer (probably Mazama), coati {Nasua nasua), tapir (Tapirus terrestris), ca- puchin monkey {Cebus apella), golden tamarin {Leontopithecus rosalia), armadillos, jaguar (Felis onca), and deer-hunting canids (Speothos"?), but no lions or wolves. The three-toed sloth was abun- dant, but never observed eating or drinking. The- vet adds, however, that there are those who believe the beast sustains itself solely by the small, slender leaves of a very high tree called amahut. Georg Marcgraf (or Marggrav or Marggraf] (1610-1644) Most illustrious of the pre-Linnaean naturalist- explorers of Brazil was Georg Marcgraf Bom in Liebstad, Saxony, educated in Holland with em- phasis on astronomy and botany, he sailed for Brazil in 1638 on a scientific expedition led by Johann Moritz, Count of Nassau-Siezen. The par- ty, which included the young physician Piso (1611- 1678), landed in Pemambuco. Explorations were restricted to northeastern Brazil in the present states of Pemambuco, Paraiba, and Rio Grande do Norte. Among MarcgraPs accomplishments were the construction of an astronomical observatory, the first of its class in the New World, and a mono- graphic study of the plants and animals of the region. After turning over his notes and illustra- tions to Moritz, for preparation and publication, the naturalist sailed for Africa, where he died shortly after arrival. MarcgraPs monumental His- toriae Rerum Naturalia Brasiliae, a part of Willen Piso's Historia Naturalis Brasilia, was published in 1648 in Amsterdam. Of the mammals of the northeastern region of Brazil described by Marcgraf, 32 were native species, the others introduced. Their detailed de- scriptions and life history notes, together with crude but useful woodcuts (fig. 2), were among the pri- mary references on which Linnaeus based bino- mials in the 10th (1758) and 12th (1766) editions of his Systema Naturce. The mammals are listed in Table 1 by the in- digenous names used by Marcgraf and their cur- rent scientific names. Provenance of the forms which served as types for binomialists, mainly Linnaeus, was restricted for taxonomic purposes to Pemambuco by Thomas (1911). Alexandre Rodrigues Ferreira (1756-1815) The first Brazilian naturalist of European ex- traction, Alexandre Rodrigues Ferreira, was bom in Salvador, Bahia. He pursued higher studies in Portugal, received his doctorate in 1779 from the University of Coimbra, and was then appointed Naturalist of the Museu Real d'Ajuda in Lisbon. He retumed to Brazil in 1783 commissioned by the museum to collect samples of plants, animals, and minerals and to record all matters of scientific and political interest within his scope. The expe- dition, or "Viagem Filosofica," explored the prov- inces of Grao Para, Rio Negro, Mato Grosso, and Cuiaba from 1 783 to 1 792 (fig. 3). Rodrigues Fer- reira retumed to Lisbon the following year. The scientific materials collected in Brazil, with notes and illustrations, were deposited in the Mu- seu d'Ajuda. Included were 4 1 7 species of animals represented by 592 specimens. Of these, 76 spec- imens represented 65 species of mammals. The whole collection was confiscated by the invading armies of Napoleon and taken to Paris for study HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 21 •c 1 2-* 03 so 'a$5M /a/acM Linnaeus, 1758 2 Hydrochaeris hydrochaeris Linnaeus, 1 766 2 Sciurus aestuans Linnaeus, 1 766 Dasypus septemcinctus Linnaeus, 1758 Dasypus novemcinctus Linnaeus, \1 5% 2 Tolypeutes trici net us Linnaeus, 1758 2 Felis tigrina Schreber, 1775 Coendou prehensilis prehensilis. Linnaeus, 1758 2 Pteronura brasiliensis Gmelin, 1 788 2 Blastoceros bezoarticus Linnaeus, 1 758 Blastoceros bezoarticus Linnaeus, 1758 F(e//5 onca Linnaeus, 1758 2 Felis onca (melanistic) 2 Felis concolor Linnaeus, 1771 * Editors' Note: Here and elsewhere in this paper. Article 51(c) of the International Code of 2kx)logical Nomen- clature, governing the use of parentheses in scientific names, is not followed. by Etienne Geoffroy St.-Hilaire of the Museum National de Histoire Naturelle in Paris. Monkeys constituted a sizeable part of the loot, and the following were described as new by Etienne Geoffroy St.-Hilaire in 1812 and by others as not- ed in brackets; the current form of each name is used: Callithrix jacchus penicillatus, Callithrix jacchus geoffroyi [Humboldt], Callithrix jacchus aurita, Callithrix humeralifer, Callithrix argentata melanura, Saguinus labiatus, Saimiri ustus [I. Geoffroy], Callicebus amictus, Callicebus person- al us, Pithecia monachus, Alouatta fusca, Cebus apella cirrifer. Cebus flavus, and Lagothrix la- gothricha canus. Mounted specimens of previ- ously named forms also brought to Paris from the Lisbon museum included Callithrix jacchus Lin- naeus, Leontopithecus rosalia Linnaeus, Chiro- potes satanas Hoffmannsegg, Brachyteles arach- noides E. Geoffroy, Inia geoffrensis Blainville, and probably others lost or discarded. Except for the descriptions by the French zo- ologist, the specimens and manuscripts of Rod- rigues Ferreira were largely neglected during the naturalist's lifetime. The several portions of the memoirs published posthumously were heavily edited. In 1972, however, the entire Viagem Fi- losofica, in two text volumes and two of colored plates, was published by the Conselho Federal de Cultura of the Brazilian Ministry of Education and Culture. Treatment of mammals in the zoological mem- oir was a model of its kind for the times. Each species was described, with bibliographic refer- ences for the ones better known, external char- acters and what was learned of habitat, habits, reproduction, utilization by man, and gastronomic rating. With respect to the last, Rodriguez Ferreira grouped the Brazilian mammals according to those used most widely for food (peccary, deer, tapir, paca, agouti), those eaten only by Indians and some HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 23 ' W' KOIFIRO |)A AIAC.KM l'llA POR AlfXANDRK RODRU.I Is l-F.RRHRA. \l MA hISI ANCIA APROXIMADA l>E Vf J7'.' kM (17M \- Fio. 3. Map of Brazil showing routes (bold lines) of Alexandre Rodrigues Ferreira, during the " Viagem Filosofica, 1783-1792; from Rodrigues Ferreira (1972). 24 FIELDIANA: ZOOLOGY Table 2. Mammals illustrated in the Viagem Filosoftca by Rodrigues Ferreira (1971), Plate no. Brazilian name Current scientific name F gure 118 Gamba Didelphis marsupialis Linnaeus 119 Macaco-da noite Aotus sp. 120 Zogue-zogue; uapuca Callicebus moloch Hoffmannsegg 121 Parauacu Pithecia monachus E. Geoffroy 122 Cuxiu Chiropotes satanas chiropotes Humboldt 123 Cuxiu-preto Chiropotes satanas satanas Hoffmannsegg 124 Guariba-vermelho Ahuatta seniculns Linnaeus 125 Guariba-da-mao-ruiva Alouatta belzebul Linnaeus 126 Mico-de-cheiro Saimiri ustus \. Geoffroy 127 Quata-de-cara-vermelha Ateles paniscus Linnaeus 128 Barrigudo-cinzento Lagothrix lagothricha Humboldt 129 Sauitinga Callithrix argentata argentata Linnaeus 130 Saui dourado Callithrix humeralifer chrysoleuca Wagner 131 Saui Callithrix jacchus penicillata E. Geoffroy 132 Saui-de-mao-ruiva Saguinus midas midas Linnaeus 133 Tamarin Saguinus midas tamarin Link 134 Saui-de-bigode-branco Saguinus labiatus labiatus E. Geoffroy 135 Tamandua-mirim Tamandua tetradactyla Linnaeus 136 Tamanduai Cyclopes didactylus Linnaeus 137 Tamanduai Cyclopes didactylus Linnaeus 138 Preguifa-de-tres-dedos Bradypus variegatus Schinz 139 Tatu-galinha Dasypus novemcinctus Linnaeus 140 Tatu peba Euphractus sexcinctus Linnaeus 141 Guaraxaim Procyon cancrivorus F. Cuvier 142 Janauira Speothos venaticus Lund 143 Guara Chrysocyon brachyurus Illiger 144 Quati Nasua nasua Linnaeus 145 Jupara Potosflavus Schreber 146 Furao Galictis vittata Schreber 147 Irara Eira barbara Linnaeus 148 Ariranha Pteronura brasiliensis Gmelin 149 Maracaja Felis geoffroyi d'Orbigny and Gervais 150 Jaguartirica Felis pardalis Linnaeus 151 Su9uarana Felis concolor Linnaeus 152 Jaguar Felis onca Linnaeus 153 On9a preta Felis onca Linnaeus 154 Peixe-boi, male & female Trichechus inunguis Natterer 155 Caitetu Tayassu tajacu Linnaeus 156 Veado vermelho Mazama americana Erxleben 157 Cariacu Odocoileus virginianus cariacou Boddaert 158 Quatipuru- vermelho Sciurus igniventris Wagner 159 Quatipuru-preto Sciurus spadiceus Olfers 160 Quatipuru-louro Sciurus igniventris Wagner 161 Rato-d'agua Nectomys squamipes Brants 162 Prea Cavia aperea Erxleben 163 Cutia-vermelha Dasyprocta leporina Linnaeus 164 Cutia-preta Dasyprocta fuliginosa Wagler 165 Acutiuaia Myoprocta exilis Wagler 166 Paca Agouti paca Linnaeus 167 Cuandu Coendou prehensilis Linnaeus 168 Uiara Inia geoffrensis Blainville 169 Tucuxi Sotalia fluviatilis Gervais and Deville white residents (anteaters, armadillos, sloths, por- cupines, monkeys, jaguar), and animals not eaten by humans (marsupials, melanistic felids, squir- rels, capybara). Bezoar stones and certain parts of the animal, usually tegumentary, were also cited for their medicinal merits, particularly as anti- venins for headaches and female sterility, or as aphrodisiacs. A memoir on the peixe boi or river manatee (Tricheciis inunguis Natterer) provides detailed in- formation on such topics as hunting, harpooning, reproduction, size, weight, blubber, butchery, HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 25 Fio. 4. Four monkeys of the "Viagem Filosofica" collections: upper left, parauaco (Pithecia monachus E. Geoffroy), possibly the holotype; upper right, saui-de-bigode-branco {Saguinus labiatus labiatus E. GeofTroy), possibly the holotype; lower left, mico-de-cheiro (Saimih ustus I. Geoffroy), possibly the holotype; lower right, saui (Callithrix jacchus penicillata E. Geoffroy), possibly the holotype; from Rodrigues Ferreira (1972). 26 HELDIANA: ZOOLOGY preservation, and market value of the flesh. The author decried the slaughter of the young and not- ed the disappearance of manatees in certain lakes. Of all Brazilian mammals described or merely listed in the Viagem Filosofica, those depicted in color in the 50 plates (each 1 9 x 29 cm) are rep- resentative. They are listed in Table 2 by plate number with their Brazilian and current scientific names. The animals were postured as prepared by taxidermists (fig. 4). Many of the monkeys are those later described by E. Geoffroy. VI. Brazil: Mammalogy to Middle of 19th Century Introduction Growth of science in South America during the first third of the 19th century shifted from the Spanish colonies, with their wars for independence and internal political turmoil, to the relatively sta- ble Portuguese colony of Brazil. Following the in- vasion of Portugal by the Napoleonic armies, the royal family fled to Brazil and made Rio de Janeiro its capital and center of cultural activities. During previous years Brazil had been closed to foreigners to prevent the mines of precious metals and min- erals from passing out of control of the ruling Por- tuguese. Dom Joao VI, however, opened the ports and changed the environment to one befitting an enlightened monarch in residence. Cultural insti- tutions, including museums, libraries, and uni- versities, were built, and scientific investigations were promoted. Betrothal of the Archduchess Leo- poldina, daughter of the Emperor of Austria, with Dom Pedro, Crown Prince of Portugal and Brazil, became the most important single factor in the advancement of science in the New World during the first half of the 1 9th century. The entourage of the bride on her voyage to Brazil included some of the best and most adventurous of the younger scientists of Austria and Bavaria. The Viennese naturalists of the party included the field collector Johann Natterer, and from the court of Munich, the zoologist Spix and the bot- anist Martins. Two years earlier, in 1815, the most accomplished of the naturalist-travelers, Maxi- milian Prinz Wied zu Neuwied of Prussia, arrived on the scene. Modem Brazilian mammalogy begins with the scientific accounts of the collections and travels of these naturalists. Johann Baptist Ritter von Spix (1781-1826) and Carl Friedrich von Martins (1794-1866) The German naturalist Johann Baptist Ritter von Spix first studied for the priesthood, but after two years his attention turned to medicine and natural history. His doctorate was earned in 1 806. That same year he was appointed assistant in the Museum of the Munich Academy of Science, with responsibility for the organization of the zoolog- ical collections. In 1816 he was ordered by the King of Bavaria to undertake a two-year scientific expedition to Brazil, together with the museum's assistant in botany, Carl Friedrich von Martins. The two departed on 10 April 1817 through the port of Trieste, and after considerable delay, they arrived in Rio de Janeiro on 15 July 1817. The exuberance and variety of the native plant life in eastern Brazil at first awed and bewildered the two young naturalists. Everything they saw was new to them, and all they could possibly collect and preserve was easily reached along the trails they traveled from Rio de Janeiro to Minas Gerais and beyond. Real or fantasized dangers lurking in what they imagined as dark, brooding, impene- trable forests restrained their urges for stepping ofl" the beaten path. The strange and wonderful wild- life encountered on the roads was enough to gratify their utmost expectations and inspired them to record their impressions in ecstatic prose. On the trip from Ipanema, Sao Paulo, to Vila Rica, Minas Gerais, they described, as translated into English by Lloyd in equally romanticized and tortured prose, the numerous flocks of little monkeys [that] run whistling and hissing to the recesses of the forest; the cavies, running about on the tops of the mountains, hastily secrete themselves under loose stones; the American ostriches (Emas), which herd in families, gallop at the slightest noise, like horses through the bush- es, and over hills and valleys, accompanied by their young; the dicholopus {Seriemas), which pursues serpents, flies, sometimes sinking into the grass, sometimes rising into the trees, or rapidly climbing the summits of the hills, where it sends forth its loud deceitful cry, resembling that of the bustard; the terrified armadillo {Tatu Canastra, Peba, Bola) runs fearfully about to look for a hid- ing place, or, when the danger presses, sinks into its armour; the ant-eater {Tamandud, Bandeira mirim) runs heavily through the HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 27 ^^ ^'Salvador (Bahia) Januaria liPbrto de Salgado) aOiamantina \>^ ^-■^buro Preto (Villa ricQ) Rio de Janeiro Sao Poulo Kane von Brasilien mit dem eingezeichneten Reiseweg von Johann Baptist von Spix und Carl Friedrich Philipp von Martius anlaBlich ihrer Expedition in den Jahren 1817-1820. Fig. 5. Map of Brazil showing routes of the Spix and Martius expedition (1817-1820); only principal stations plotted; from Tiefenbacher(1983). plain, and, in case of need, lying on its back, threatens its pursuers with its sharp claws. Far from all noise, the slender deer, the black tapir or a pecari, feed on the skirts of the forest. Elevated above all this, the red-head- ed vulture (urubii) soars in the higher re- gions; the dangerous rattle-snake {Casca- vel), hidden in the grasses, excites terror by its rattle; the gigantic snake sports suspended from the tree with its head upon the ground; and the crocodile resembling the trunk of a tree, basks in the sun on the banks of the pools. After all this has passed during the day before the eyes of the traveler, the ap- proach of night, with the chirping of the grasshoppers, the monotonous cry of the goat-sucker {Jodo corta pdo), the barking of the prowling wolf, and of the shy fox, or the roaring of the ounces, complete the singular picture of the animal kingdom in these peaceful plains. For the next three years, the zoologist and bot- anist explored the eastern states of Brazil from Sao Paulo and Minas Gerais north to Para. Most of July and August of 1 8 1 9 was spent in Belem (Para). On 2 1 August they shipped up the Rio Amazonas, making stopovers at the mouth of the Rio Tocan- tins, the Rio Xingu ( 1 September), Santarem on the Rio Tapajos ( 1 8 September), Obidos (23 Sep- tember), Parintins, and Vila Nova da Rainha (1 October). The mouth of the Rio Madeira was passed 1 5 October, and on 22 October they landed at Barra do Rio Negro (Manaus). Travel upstream 28 HELDIANA: ZOOLOGY continued in November with a stop at Tefe (for- merly Ega) on 26 November. Spix then traveled alone up the Solimoes to Tabatinga at the Peru- vian border, arriving 9 January 1820. Martius, for his part, ascended the Rio Japura to Araracuara in eastern Colombia. Spix returned to Manaus on 3 February 1820. On 1 1 February he ascended the Rio Negro to Barcelos and was back again in Manaus 28 Feb- ruary to continue his travels downstream to Be- lem, where he arrived on 16 April. He embarked on 14 June 1820 for Europe from Rio de Janeiro (fig. 5). In the Reise. Spix and Martius (1828, p. 541) made up an impressive list of the mammals of the sertao (scrub country) of Campos Gerais de Sao Felipe in the angle between the Rio Sao Francisco and its eastern tributary, the Rio Verde Grande, northern Minas Gerais. The data were evidently compiled uncritically from a number of sources, including local informers, personal observations, and publications based on the Wied-Neuwied (1826) collections. Their use and misuse of names are too involved to unravel here. Except for the missing bats (given elsewhere by Spix, 1823) and some small rodents, it is unlikely that a similar or larger number of mammalian species of the area, based on actual specimens, could be made today. The sertao mammals of the Spix and Martius ex- pedition are listed in Table 3 by current scientific names of the species only, with the Spix and Mar- tius equivalents omitted. In his journey up the Amazon, Spix noted habits of the inia {Inia geoffrensis) (Spix &, Martius, 1831, p. 1 1 1 9) and of the manatees (Trichechus inunguis) (Spix & Martius, 1831, p. 1122). The results of the expedition are recorded in several publications, including the Simiarum et Vespertilionum Brasiliensium by Spix (1823). The account of the nearly three-year journey or Reise in Brazil by Spix and Martius (1823-1831) is re- plete with observations on the biology, geography, geology, paleontology, mineralogy, meteorology, and the various human cultures and industries of the parts of the country they traveled. Many kinds of mammals are mentioned, but except for bats and monkeys, few of them were collected. The zoological material actually collected con- sisted of thousands of invertebrates and 498 species of vertebrates, of which 34 were monkeys and 15 bats. Altogether, according to Avila Pires (1974, p. 139), 85 species of mammals were collected. Spix (1823) reported only on the monkeys and bats and illustrated in color the types of all species. Table 3. Mammals of the sertao of Campos Gerais de Sao Felipe, Minas Gerais, recorded by Spix and Mar- tius (1828, p. 541, footnote 3). Current scientific names to species only are used. The Spix and Martius usage of local, German, and scientific names is too confused for tabulation. The arrangement is phylogenetic. Marsupialia Caluromys philander Linnaeus Didelphis marsupialis Linnaeus Primates Callithrix jacchus Linnaeus Cebus apella Linnaeus Alouatta fusca E. GeofTroy Alouatta caraya Humboldt Edentata Tamandua tetradactyla Linnaeus Myrmecophaga tridactyla Linnaeus Bradypus torquatus Desmarest Bradypus variegatus Schinz Dasypus novemcinctus Linnaeus Tolypeutes tricinctus Linnaeus Priodontes maximus Kerr Euphractus sexcinctus Linnaeus Carnivora Dusicyon thous Linnaeus Chrysocyon brachyurus Illiger Nasua nasua Linnaeus Procyon cancrivorus G. Cuvier Conepatus chinga Molina Eira barbara Linnaeus Pteronura brasiliensis Gmelin Felis wiedii Schinz Felis tigrina Schreber Felis pardalis Linnaeus Felis concolor Linnaeus Felis onca Linnaeus Felis yagouaroundi E. Geoffroy Perissodactvla Tapirus terrestris Linnaeus Artiodactyla Mazama gouazoubira Fischer Mazama americana Erxleben Blastoceros bezoarticus Linnaeus Lagomorpha Sylvilagus brasiliensis Linnaeus RODENTIA Sciurus aestuans Linnaeus Wiedomys pyrrhorhinos Wied-Neuwied Echimys and/or Proechimys species? Myocastor coypus Molina Kerodon rupestris Wied-Neuwied Cavia aperea Linnaeus Dasyprocta leporina Linnaeus Agouti paca Linnaeus Coendou insidiosus Kuhl Chaetomys subspinosus Olfers HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 29 d. c 5? 11 c 5 1^ 2 i" •S 30 HELDIANA: ZOOLOGY Table 4. Monkeys (Primates) of the Spix and Martius Expedition described by Spix (1823); the arrangement is phyiogenetic. Current name Spix and Martius synonym Figure Cebuella pygmaea Spix, 1 823 Callithrix jacchus jacchus Linnaeus, 1758 Callithrix jacchus penicillatus E. Geoffroy, 1812 Saguinus bicolor bicolor Spix, 1823 Saguinus fuscicollis fuscicollis Spix, 1823 Saguinus mystax mystax Spix, 1823 Saguinus nigricollis nigricollis Spix, 1 823 Saguinus oedipus geoffroyi Pucheran, 1845 Callicebus cupreus Spix, 1823 Callicebus personatus personatus E. Geoffroy, 1812 Callicebus personatus nigrifrons Spix, 1 823 Callicebus personatus melanochir Kuhl, 1820 Callicebus torquatus torquatus Hoffmannsegg, 1 807 Callicebus cinerascens Spix, 1823 Aotus vociferans Spix, 1 823 Actus azarae infulatus Kuhl, 1820 Pithecia monachus monachus E. Geoffroy, 1812 Pithecia pithecia pithecia Linnaeus, 1 766 Chiropotes satanas chiropotes Humboldt, 1812 Cacajao melanocephalus ouakary Spix, 1823 Alouatta caraya Humboldt, 1812 Alouatta belzebul discolor Spix, 1823 Alouatta fusca Spix, 1823 Alouatta seniculus stramineus Humboldt, 1812 Cebus albifrons unicolor Spix, 1 823 Cebus apella libidinosus Spix, 1823 Cebus apella macrocephalus Spix, 1823 Cebus apella xanthosternos Wied-Neuwied, 1 820 Lagothrix lagothricha lagothricha Humboldt, 1812 Lagothrix lagothricha carta E. Geoffroy, 1812 Brachyteles arachnoides E. Geoffroy, 1 806 Jacchus albicollis Spix, 1 823 Midas oedipus (varietas), Spix, 1823 Callithrix gigot Spix, 1823 Callithrix amicta E. Geoffroy, 1812 Nyctipithecus felinus Spix, 1823 Pithecia hirsuta Spix, 1823; Pithecia inusta Spix, 1823 Pithecia capillamentosa Spix, 1823 Brachyurus israelita Spix, 1 823 Mycetes barbatus Spix, 1823 Cebus gracilis Spix, 1823 Cebus cucullatus Spix, 1823; Cebus xanthocephalus Spix, 1823 Gastrimargus infumatus Spix, 1823 Gastrimargus olivaceus Spix, 1 823 Brachyteles macrotarsus Spix, 1 823 most life-size. Separate reports on all groups of animals collected by Spix have been brought to- gether in a Festschrift in his honor edited by Tie- fenbacher (1983). The mammals are treated by Kraft (1983). The 31 presently recognized species and sub- species of monkeys ( 1 5 new) and the 1 4 recognized species of bats (six new) are listed in Tables 4 and 5 by current names with synonyms in parentheses. Maximilian Prinz von Wied-Neuwied (1782-1867) Maximilian Prinz von Wied-Neuwied was bom in Prussia and studied biological sciences at the University of Gottingen under the famous natu- ralist-anthropologist Blumenbach. His ambition to travel and study nature in South America was realized when he sailed for Rio de Janeiro from England the first week of May 1815, and arrived on 17 July. After a few excursions in the surroundings of Rio de Janeiro, Wied-Neuwied left for Cabo Frio on 15 August 1815, stopping at many fazendas and villages along the way. He left Cabo Frio on 8 September for Sao Salvador dos Campos dos Goitacazes (now simply Campos) on the Rio Pa- raiba, and arrived on 25 September. After more excursions and more collections in the state of Rio de Janeiro, he crossed the Rio Itabapoana on 26 November into the state of Espirito Santo. A con- siderable amount of time was devoted there to explorations of the Rio Doce region. February 1816 saw Wied-Neuwied in Bahia, where he occupied himself until May 1817. The coastal town of Bel- monte, where he arrived in August 1816, was the base for explorations of Botocudo Indian territory. In December 1816 Wied-Neuwied established II- heus as center for travel westward to Sao Pedro de Alcantara, now Itabuna, and the border of Mi- nas Gerais. On 1 May Wied-Neuwied embarked at Salvador for Lisbon, then transshipped to Ger- many through an English port. Wied-Neuwied's itinerary is difficult to track be- HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 31 Table 5. Bats (Chiroptera) of the Spix and Martius Expedition described by Spix (1823); the arrangement is phylogenetic. Current name Spix and Martins synonym Rhynchonycteris naso Wied-Neuwied, 1820 Noctilio albiventris albiventris Desmarest, 1818 Noctilio leporinus leporinus Linnaeus, 1 758 Tonatia bidens Spix, 1823 Trachops cirrhosus Spix. 1823 Glossophaga sohcina Pallas, 1 766 Carollia perspiciUata Linnaeus, 1758 Artibeus planirostris Spix, 1 823 DiphyUa Spix, 1823 Diphylla ecaudata Spix, 1 823 Thyroptera Spix. 1 823 Thyroptera tricolor Spix, 1823 Eptesicm brasiliensis Desmarest, 1823 Promops nasutus Spix Molossus ater E. Geoffroy, 1 805 Proboscidea rivalis Spix, 1 823; Proboscidea saxatilis Spix, 1 823 Noctilio albiventer Spix, 1823 Noctilio rufiis Spix, 1823 Glossophaga amplexicaudata Spix, 1823 Vampynts soricinus Spix, 1823 Molossus fumarius Spix, 1823 Molossus ursinids Spix, 1823 cause of his many roundabout journeys and short excursions with too few dates for fixing comings and goings. To add to the difficulty, the names of many localities he visited no longer exist or were never plotted on any official map; a few names have changed. Bokermann's (1957) gazetteer of nearly all localities of the Reise, with page refer- ences to their mention in Wied-Neuwied's works, is indispensable for study of the naturalist's op- erations in Brazil. Wied-Neuwied was interested in all aspects of nature, but the fauna and Indians engaged most of his attention. His species accounts are models of precision, his descriptions detailed, and com- parisons where needed are made with published descriptions by Humboldt, Azara, Buffon, and others. The bibliographic references to the species are complete. Observations of habitats and repro- duction are carefully recorded, and geographic range is usually given with circumspection. Wied- Neuwied's account of Geoffi"oy's tufted-ear mar- moset (his Hapale leucocephalus) is an example (my translation): I found it in the state of Espirito Santo. I am unable to determine if it extends north of the Rio Doce or beyond as I could not hunt often in the dark forests of this river because of the Botocudo Indians. I can therefore state that the habitat of this species lies between 20° and 21" south latitude. The animal is common in the forests of the Rio Espirito Santo, especially in the outlying bush and the mangue bush {Conocarpus and Av- icennis) bordering the river, as well as in the low palm {Allagoptera pumila and others)- covered sandy coastal districts not far from the mouth of the Espirito Santo. . . . The following excerpt of Wied-Neuwied's (1826, p. 161) observations on the golden lion tamarin (Leontopithecus rosalia rosalia Linnaeus) brings together his observations on distribution, habits, habitat, food, and reproduction: The sahuim vermelho is nowhere abundant; we saw only single individuals or family groups, particularly in the Serra da Inua, the forests of Sao Joao, and in the hilly forest surrounding Ponta Negra and Gurupina. The animal lives just as well on bushy sandy plains as in the high mountain forests. It feeds on fruits and insects and hides from strangers by disappearing into the leafy tree- tops. One or two young are produced at a birth. The female carries the offspring on her back or at her breasts [when suckling] until they are strong enough to follow her on their own. . . . Any excitement causes them to erect the long hair surrounding their faces. In gen- eral, however, their habits are similar to those of other sahuis. Wied-Neuwied also accurately delimited the distribution of the subspecies Leontopithecus ro- salia chrysomelas and added information on hab- its and reproduction. Wied-Neuwied notes (1826, p. 1 59) that "sahuis bom in Europe are carried by the father but I have never seen this here." Although generally careful in interpreting his 32 HELDIANA: ZOOLOGY ". ^.... Fig. 7. Some animals of the Wied-Neuwied Brazilian expedition: upper left, Hapale chrysomelas Wied-Neuwied (= Leontopithecus rosalia chrysomelas), possibly the holotype; upper right, Mus pyrrhorhinos Wied-Neuwied (= fViedomys pyrrhorhinos), possibly the holotype; lower left, Desmodus rufus Wied-Neuwied (= Desmodus rotundus E. Geoffroy); lower right, Felis macroura Wied-Neuwied (= Felis wiedii Schinz), possibly the holotype; from Wied- Neuwied (1822-1831). data, Wied-Neuwied could arrive at unwarranted conclusions. Among the bats collected, the leaf- nosed Phyllostomus hastatus was largest and for this reason was regarded as a blood-sucking vam- pire, although Wied-Neuwied found only insects and no blood in the stomach of this or any other bat he had examined. After confessing he had nev- er seen a bat feed on blood, he correctly blamed the large bats seen fluttering around the pack mules at night for causing them to appear next morning covered with blood. Convinced in his judgment, he described the wartlike excrescences around the mouth of innocent phyllostomine bats as adap- tations for blood-sucking. Ironically, Wied-Neu- wied (1824, 1826) later described and figured the external and dental characters of a bat he named Desmodus rufus, unaware it was a real blood-suck- ing vampire. Wied-Neuwied noted, however, that he had no opportunity to observe the live animal, because it had been captured and prepared as a specimen by assistants during his absence. The food and habits of this bat, he believed, were no different from those of other bats. The mammals of Wied-Neuwied's Brazilian ex- HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 33 pedition are described or recorded in several pub- lications. Those under Wied-Neuwied's own name are found in Isis ( 1 820, 1821), the Reise nach Bra- silien in two volumes ( 1 820, 1821), the Abbildun- gen zur Naturgeschichte Brasi liens ( 1 822-1 83 1 , see fig. 7 for some samples), and the four-volume Bei- trdge zur Naturgeschichte von Brasilien. The first volume of the last title is on reptiles, the second on mammals ( 1 826), the third and fourth on birds. Some diagnoses and binomials that Wied-Neu- wied proposed for new forms received duly ac- knowledged advance publication by Kuhl (1820) and Schinz (1821). Authorship of such newly named forms continues to be attributed to Wied- Neuwied, according to Articles 1 1 and 50 of the International Code of Zoological Nomenclature. In the few cases where Kuhl or Schinz proposed names other than those used by Wied-Neuwied, authorship is determined by priority. The actual number of mammals collected by Wied-Neuwied is unknown. According to him, they represented 82 species, but the number recognized today as valid is 7 1 . The specimens were preserved in his private museum, but duplicates were dis- tributed to the natural history museums of Berlin, Frankfurt, Leiden, and Paris. After Wied-Neu- wied's death, the remainder of the collection was sold, and the American Museum of Natural His- tory in New York acquired a part in 1869. Avila Pires ( 1 965, p. 3) affirms that fewer than 600 spec- imens of the original collection are registered in the catalogue of mammals of the New York in- stitution. Of these, only 38 skins and 16 skulls are of South American origin. Included are holotypes (or syntypes) of Didelphis aurita Wied-Neuwied, Didelphis cinerea Temminck, Molossus plecotus Wied-Neuwied, Phyllostoma brevicaudum Wied- Neuwied, Vespertilio leucogaster Wied-Neuwied, Vespertilio naso Wied-Neuwied, Hypudeus dasy- trichos Wied-Neuwied, and Mus pyrrhorhinos Wied-Neuwied. Table 6 lists all mammalian species recorded by Wied-Neuwied. Current names are used; syn- onyms used by Wied-Neuwied are included. Johann Natterer (1787-1843) Johann Natterer, bom near Vienna, was well schooled in the sciences, especially biology, and in modem languages and illustration. Natterer's father, the imperial falconer and collector of birds and insects, taught him to hunt and preserve an- imals as museum specimens. In 1 8 1 6 he was em- ployed as assistant in the Imperial Natural History Museum of Vienna and in 1817 was appointed member of an expedition to investigate the Bra- zilian biota. He arrived in Rio de Janeiro on 5 November accompanied by Mikan and Pohl, both naturalists, and Schott, a botanist. Within a year Mikan, Sochor, a hunter, and two artists who were to accompany Natterer, retumed to Europe. Pohl and Schott retumed in 1821. Natterer was primarily a bird collector, but his interest in collecting extended to mammals, other vertebrates, insects, and parasitic helminths. He traveled light and, as a rule, worked alone or with few native helpers (Ihering, 1 902). He collected in most of the eastem coastal states and in Mato Grosso and the Amazonian region between the Rios Tapajos and Madeira and in the Rio Negro basin north of the Rio Amazonas (fig. 8). His main base for the first five years was Ipanema, Sao Pau- lo. His itinerary— with goings and comings, side trips, short stopovers in some sites, long delays in others— was arranged chronologically by Pelzeln (1871,1883) into "Reisen" (or journeys), with dates given for all points visited, and is summarized below. Only general areas or terminal points and inclusive dates are given. Johann Natterer's Brazilian Reisen, 1817-1835. I. Rio de Janeiro, 5 November 1817 to 1 No- vember 1818. II. Eastem Sao Paulo, 2 November 1818 to March 1820. III. Southern Sao Paulo to boundary between Rio Grande do Sul and Rio de Janeiro, July 1820 to 1 February 1821. IV. Rio de Janeiro, Sao Paulo, 1 February to September 1822. V. Northern Sao Paulo, Goias, eastem Mato Grosso, Minas Gerais, October 1822 to 31 December 1824. VI. Mato Grosso, January 1825 to July 1829. VII. Mato Grosso, Rio Madeira, and upper trib- utaries to Borba in Amazonas (Capitania Rio Negro), 15 July 1829 to June 1830. VIII. Borba to Rio Negro, Rio Casiquiare, Ven- ezuelan border, retum to Barcelos and Bor- ba, June 1830 to 31 August 1830. IX. Rio Negro from Barcelos to Rio Branco, 5 September 1831 to 2 July 1832; Barra do Rio Negro, 29 August 1832 to 7 July 1834; Rio Tapajos, August 1834. X. Para, Maranhao, Rio Grande, Paraiba, Per- 34 FIELDIANA: ZOOLOGY ■ ■ ■ /yj/__/"/»«-»r ornt .\tm^ni6er A*!// <»«.»<' ' 6fx Ar^rintr fS'2/ /«y.//r/«- /«;'/< 17'f/l'ri. ./mm /^ifA/jt . 1ntft,*l /xV 1/:^/- /S'y? .-r -IZ Fig. 8. Map of Brazil showing routes of Johann Natterer (bold line); from [brother of Johann] Natterer (1833, Oken's Isis, heft VI, pi. 14). nambuco, Bahia, Rio de Janeiro, September Natterer's enormous collections were sent to the 1834 to September 1835 (no mammal col- Vienna museum and, except for the birds and lections). mammals, were never fully reported. His friend Sailed for Europe 15 September 1835. Andreas Wagner (1797-1861) described most of HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 35 Table 6. Brazilian mammals recorded by Wied-Neuwied (1826) with some figured in the Abbildungen (1822- 1831); the arrangement is phylogenetic. Current name Wied-Nenwied synonym Figure Marsupiaua Afarmosa murina Linnaeus, 17S8 Marmosa cinerea Temminck, 1 824 Philander opossum frenata Olfers, 1818 Didelphis marsupialis aurita Wied-Neuwied, 1826 Chiroptera Rhynchonyaeris naso Wied-Neuwied. 1820 (Reise) Centronyaeris maximiliani Fischer, 1 829 Peropteryx macrotis Wagner. 1843 Diclidurus albus Wied-Neuwied, 1819 Noctilio leporinus Linnaeus, 1 758 Xfacrophyllum macrophyllum Wied-Neuwied, in Schinz, 1821 Phyllostomus hastatus Pallas, 1867 PhvUostomus obscunis Wied-Neuwied, in Schinz, i821 Glossophaga soricina Pallas, 1 766 Anoura caudifera E. Geoffrey, 1818 CaroUia brevicauda Wied-Neuwied, 1821 Carollia perspicillata Linnaeus, 1 758 Artibeus liturcaus 0\fcT%, 1818 Desmodus rotundus E. Geoffiroy, 1810 Myotis albescens E. Geoflfroy, 1806 Myotis nigricans Wied-Neuwied, in Schinz, 1821 Eumops perotis Wied-Neuwied, in Schinz, 1 82 1 Primates Callithrix jacchus penicillatus E. Geoffix)y, 1812 Callithrix jacchus geoffroyi Humboldt, 1812 Leontopithecus rosalia chrysomelas Kuhl, 1 820 Leontopithecus rosalia rosalia Linnaeus, 1758 Callicebm personatus persoruUus E. GeoflSroy, 1812 Callicebus personatus melanochir Wied-Neuwied, 1820 (Reise) Alouatta caraya Humboldt, 1812 Alouatta fusca E. Geoffroy, 1812 Cebus apella nigritus Goldfiiss, 1 809 Cebus apella robustus Kuhl, 1 820 Cebus apella xanthostemos Wied-Neuwied, 1820 (Reise) Brachyteles arachnoides E. Geoflfroy, 1806 Edentata Tamarulua tetradactyla Linnaeus, 1758 Myrmecophaga tridactyla Linnaeus, 1758 Edentata Bradypus torquatus Desmarest, 1816 Didelphys myosuros Temminck. 1 825 Didelphis marsupialis. Wied-Neuwied, 1826, not Linnaeus Vespertilio caJcaratus Wied-Neuwied, in Schinz, 1821, not Rafinesque, 1818 Vespertilio caninus Wied-Neuwied, in Schinz, 1821, not Blumenbach, 1797 Diclidurus freyreissii Wied-Neuwied 1822, Abbild. Noctilio dorsatus Desmarest, 1818; Noctilio unicolor Desmarest, 1818 Artibeus planirostris Spix, 1823 Glossophaga amplexicaudata E. Geoflftoy, 1818 Phyllostoma bernicaudum {sic) Wied-Neuwied, in Schinz, 1821 Phyllostoma brachyotos (sic) Wied-Neuwied, in Schinz, 1821 Phyllostoma superciliatum Wied-Neuwied, in Schinz, 1821 Rhinolophus ecaudatus Wied-Neuwied, in Schinz, 1821; ZJesmorfus rw^ Wied-Neuwied, 1824 Vespertilio leucogaster Wied-Neuwied, in Schinz, 1821 Hapale penicillatus kuhlii Wied-Neuwied, 1826 p. 142)* Hapale leucocephalus Kuhlii (sic), Wied-Neuwied, 1826t Mycetes niger Kuhl, 1820 Mycetes ursinus Humboldt, 1812, not Humboldt, 1805 Cebus cirrifer E. Geoffroy, 1812, not Cebus fatuel- lus Linnaeus ? Cebus flavus E. Geoffroy, 1812 Ateles hypothanthus Kuhl, 1820 Myrmecophaga jubata Linnaeus, 1766 Bradypus tridactylus Wied-Neuwied, 1826, not Linnaeus, 1758 36 HELDIANA: ZOOLOGY Table 6. Continued. Current name Wied-Neuwied synonym Figure Cabassous unicinctus Linnaeus, 1758 Euphractus sexcinctus Linnaeus, 1758 Dasypus novemcinctus Linnaeus, 1 758 Priodontes maximus Kerr, 1 792 Carnivora Dusicyon thous brasiliensis Wied-Neuwied, in Schinz, 1821 Chrysocyon brachyurus Illiger, 1815 Nasua nasua solitaria Wied-Neuwied, in Schinz, 1821 Procyon cancrivorus G. Cuvier, 1 798 Potosflavus nocturnus Wied-Neuwied, 1826 Eira barbara Linnaeus, 1758 Pteronura brasiliensis Gmelin, 1 788 Felis wiedii Schinz, 1821 Felis pardalis mitis F. Cuvier, 1820 Felis yagouaroundi eyra Fischer, 1814 Felis concolor Linnaeus, 1 77 1 Felis onca Linnaeus, 1758 SiRENIA Trichechus manatus lAnnditns, 1758 Perissodactyla Tapirus terrestris lArmaitxis, 1758 Artiodactyla Tayassu tajacu Linnaeus, 1758 Tayassu pecari Link, 1795 Mazama gouazoubira Fischer, 1814 Mazama americana Er\\ehcn, Mil Blastoceros bezoarticus Linnaeus, 1758 Blastocerus dichotomus Illiger, 1815 Lagomorpha Sylvilagus brasiliensis Linnaeus, 1758 RODENTIA Sciurus aestuans Linnaeus, 1 766 Wiedomys pyrrhorhinos Wied-Neuwied, 1821 (Reise) Oxymycterus rufus dasytrichos Wied-Neuwied, in Schinz, 1821 Proechimys myosuros Lichtenstein, 1818 Cavia aperea Erxleben, 1 777 Kerodon rupestris Wied-Neuwied, 1820 (Isis) Hydrochaeris hydrochaeris Linnaeus, 1 766 Dasyprocta leporina aguti Linnaeus, 1 766 Agouti paca Linnaeus, 1 766 Coendou insidiosus Olfers, 1818 Chaetomys subspinosus Olfers, 1818 Dasypus setosus Wied-Neuwied, 1 826; Dasypus gilvipes Illiger, 1815 Dasypus longicaudus V^icd-Neuwied, 1826 Dasypus gigas Cuvier, 1822 Canis azarae Wied-Neuwied, 1 823 Canis campestris Wied-Neuwied, 1826 A^asMfl 50c/a//5 Wied-Neuwied, 1826 Mustela gulina Wied-Ncuwied, 1821 Felix macroura Wied-Neuwied, 1 823 Felis pardalis. Wied-Neuwied, 1826 Felis yaguarundi, Wied-Neuwied, 1 826 Felis brasiliensis Wied-Neuwied, 1 82 1 Manatus americanus Link, 1795 Tapirus americanus Gmelin, 1788 Dicotyles torquatus Cuvier, 1817 Cervus simplicicornis Illiger, 1815 Cervus rufus Cuvier, 1817 Cervus campestris 'Wied-Nexxwied, 1826, not Cuvier, 1817 Cervus paludosus Desmarest, 1 822 Hypudeus dasytrichos Wied-Neuwied, 1826 * The name is a correctly formed trinomial but this form was not in use at the time, and Wied-Neuwied used no trinomials elsewhere in his publications on Brazilian mammals. t The name appears to be a trinomial although the patronymic, properly in the genitive, is not italicized. Most likely Wied-Neuwied meant to cite Kuhl for this and the preceding taxon as authority for his use of the names in question. It was common practice at the time to cite the author who replaced an earlier generic name with a different one. HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 37 the new mammalian species in a series of reports published in the Archivfur Naturgeschichte ( 1 842, 1843), in the Abhandlungen der Akademie .Vfiin- chen ( 1 847-1 849), and his supplementary volumes of Schreber's Sdugethiere (1840-1855). Finally, Pelzeln (1883) brought together most, if not all, available taxonomic, descriptive, and geographic data in a single report. Natterer intended to work up the entire collection himself, but died within a few years of his return to Europe. His journal, with notes on habits, reproduction, and anatomy of the Brazilian animals collected, was lost. Natterer collected 78 1 specimens of mammals, representing more than half (58%) of the currently known Brazilian genera and nearly as many (44%) of the species (table 3). Most poorly represented are bats, mice, and mouse opossums. Had Natterer been equipped with suitable traps and trammel nets known at the time but not used in fieldwork, he might have collected nearly all the mammalian genera and species now known to occur in Brazil. Still, his collection represented more species and included t\pes of more new species than had been collected in Brazil by anyone else in the century, or p)ossibIy at any time. The numbers of genera and species of mammals collected by Natterer, as identified by Pelzeln (1883), are listed in Table 7. The totals are com- pared with the numbers currently recognized, some genera having been increased and some species eliminated by synonymy. The revised numbers of genera and species are shown, in turn, as percent- ages of the estimated total numbers of currently known Brazilian genera and species of mammals. VII. Guianas: Mammalogy to End of 18th Century Pierre Barrere (1690-1755) The physician, botanist, and correspondent of the French Royal Academy of Sciences, Pierre Barrere, resided three years (1752-1755) in Cay- enne, with instructions to prepare a detailed report on the natural history of French Guiana. The work he finally published in 1 74 1 , however, is no more than an abbreviated glossary- of Guianan minerals, plants. moUusks, fishes, reptiles, birds, and mammals. The list of mammals was uncrit- ically compiled from Marcgraf and others. Species previously recorded by early chroniclers from the lower Rio Orinoco region which occur throughout the Guianas but were not mentioned by Barrere are the golden handed tamarin {Saguinus midas), red brocket (Mazama americana), red acuchi (A/y- oprocta exilis), tayra (Eira barbard), white-lipped peccary (Tayassu pecari), and silky anteater (Cy- clopes didactylus). Jose Gumilla (d. 1750) A natural history and geography of the Rio Ori- noco region in Spanish, published by Father Jose Gumilla, provides interesting, but largely erratic, descriptions of the countryside and human inhab- itants, but nothing of interest regarding native mammals. Gumilla's explorations of the interior led him to deny the reported existence of a con- nection between waters of the Orinoco and Negro rivers. Jacques Nicolas Bellin (1703-1772) The description of the Guianan possessions of France, Spain, Holland, and Portugal, from the Orinoco River to the Amazonas River, by Jacques Nicolas Bellin, published in 1763, contains infor- mation on natural history, but adds nothing note- worthy to the then-known mammalian fauna. Edward Bancroft (1744-1821) The English physician Edward Bancroft lived three years in Dutch Guiana, now Suriname, prac- ticing medicine and gathering notes for his Essay on the Natural History of Guiana. The work, pub- lished in 1 769, deals broadly with plants and an- imals, but the author's knowledge of mammals was mostly limited to hearsay, although he also made some observations on animals brought to him by natives or seen in captivity or during short walks into the countryside. Persistent reports of the existence of apes or ape-men in South America were recounted by Bancroft (p. 1 30) in these terms: The Orang-Outang of Guiana is much larger than either the African or the Oriental, if the accounts of the natives may be relied on; for I do not find that any of them have been seen by the White inhabitants of this coast, who never penetrate far into the woods. These animals, in all the different languages of the Natives, are called by names signi- 38 HELDIANA: ZOOLOGY Table 7. Numbers of mammalian genera and species collected by Johann Natterer in Brazil, 1817-1835, based on Pelzeln (1883), and compared with currently known totals. Table 7. Continued. Total Taxon Number reported by Pelzeln (1883) Current equiva- lent number cur- rently known for Brazil (esti- mated, 1984) Percent- age of current total col- lected by Natterer Marsupialia Genera Species 2 18 6 15 8 30 75% 50% Chiroptera Genera Species 10 48 28 40 60 125 47% 32% Primates Genera Species 12 45 14 28 16 50 87% 56% Edeimtata Genera Species 10 16 10 12 12 15 83% 75% Carnivora Genera Species 11 17 10 14 14 25 71% 56% PiNNIPEDIA Genera Species 2 2 0% 0% SlRENlA* Genera Species 1 1 1 1 1 2 100% 50% Perissodactyla Genera Species 1 1 1 1 1 1 100% 100% Artiodactyla Genera Species 4 7 4 6 5 7 80% 86% Lagomorpha Genera Species 1 1 1 1 1 1 100% 100% RODENTIA Sciuropmorpha Genera 1 Species 5 1 3 3 6 33% 50% Myomorpha (Murinae Genera 3 Species 1 7 excluded) 5 17 20 45 25% 24% Caviomorpha Genera Species 11 24 15 22 23 47 65% 47% Cetacea* Genera Species 2 2 2 2 2 2 100% 100% Taxon Total cur- rently known Percent- Number for age of reported Current Brazil current by equiva- (esti- total col- Pelzeln lent mated, lected by (1883) number 1984) Natterer Totals Genera 69 99 170 58% Species 202 156 358 44% * Fresh water only. fying a Wild Man. They are represented by the Indians as being near five feel in height, maintaining an erect position, and having a human form, thinly covered with short black hair; but I suspect that their height has been augmented by the fears of the Indians, who greatly dread them, and instantly flee as soon as one is discovered, so that none of them have ever been taken alive, much less at- tempts made for taming them. The Indians relate many fabulous stories of these ani- mals; and, like the inhabitants oi Africa and the East, assert, that they will attack the males, and ravish the females of the human species. Philippe Fermin (1720-1790) Philippe Fermin, the author of an account pub- lished in 1769 of the history, geography, and nat- ural objects of colonial Suriname, was one of those European men who "never penetrate far into the woods." Indeed, Fermin believed that all Euro- [)eans and Creoles were physically incapable of coping with the difficulties of surveying the natural fauna of the countryside, let alone the wilderness, or resisting the diseases generated by the "foul" air of forests and swamps. Notwithstanding this, Fermin compiled a fair list of the mammals. The didelphids included Didelphis marsupialis, Phi- lander opossum, and Marmosa spp. All three kinds of anteaters and the two- and three-toed sloths are mentioned. The two native squirrels, Sciurus aes- tuans and Sciurillus pusillus, are distinguished. Other rodents are the capybara, paca, a porcupine, cavy, spiny rats or echimyids (most likely of the genera Proechimys and Echimys), and a water rat HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 39 (probably Nectomys squamipes). The carnivores include tayra, otter, jaguar, puma, margay, ocelot, and two kinds of bush dogs (Dusicyon thous and possibly Speothos venaticus). Monkeys are Sa- guinus midas, Saimiri sciureus, Pithecia pithecia, Chiropotes sat anas, Cebus apella, Alouatta seni- culus, Ateles paniscus. and variants of some of them regarded as distinct species. African simians introduced with the slave trade and mentioned by Marcgraf are included. The ungulates are tapir, brocket {Mazama americana), and the collared and white-lipped peccaries. Regarding white-lipped peccaries, Fermin af- firms they form herds of as many as 300 individ- uals. Hunters, he states, tremble when they hear the sound of their clicking tusks. When attacked, only two avenues of escape are open: The first is a tree, if it can be climbed; the second and surest is standing ground and urinating, the odor of the urine, he affirms, being a powerful peccary repel- lant. Monsieur Bajon (1763?) The French physician, surgeon, and anatomist Bajon, with 1 2 years' residence in French Guiana, investigated climate, agriculture, natural history, and human diseases. The knowledge he gained was acquired firsthand, much of it new or supplemen- tary to what was already contained in the ency- clopedic volumes on natural history by Buffon and Daubenton. In the second of his two-volume work, Bajon (1778, p. 178) declared that, contrary to popular belief, the jaguar feared man and did not attack without provocation. His accounts of habits and detailed descriptions of intestinal morphology and female genitalia of peccaries supplement Dauben- ton's (in Buffon) gross anatomy of a male collared peccary. Bajon clarified the differences between the agouti (Dasyprocta leporina) and acouchi {My- oprocta exilis). He described the male agouti penis, with its peculiar complement of spines, erectile spears, and sharp blades. Descriptions with life history notes are given for the chien sauvage {Du- sicyon thous), eira {Eira barbara), and chien cra- bier (Procyon cancrivorus). Marsupials fascinated him, particularly the role of the pouch in females of the pean {Didelphis marsupialis), quatre-ouel (Philander opossum), and also the pouchless rat de bois {Marmosa sp.). The commonly held belief that each didelphid young is bom and develops at the end of a teat was rejected by Bajon, but despite numerous observations and dissections, he failed to solve the mystery of marsupial birth. Bajon's monographic account of the tapir {Tap- irus terrestris) includes detailed, but not always accurate, descriptions of anatomy, reproduction, development, behavior, food, vocalization, hunt, and human utilization. John Gabriel Stedman (1744-1797) A soldier of the Scots Brigade of the Nether- lands, John Gabriel Stedman arrived in Suriname in 1773 to help subdue the uprising of the African slaves. Most of the fighting was already over when he landed, so Stedman devoted much of his time to recording his observations of life in the country and wilderness. His Narrative, published in 1 796 in two volumes, contains much on the natural history of Suriname, with illustrations by his own hand (fig. 9). The mammals, some only listed, oth- ers described, often with anecdotes, are the fol- lowing. Stedman used local names, current bino- mials are in parentheses. Volume I, p. 14. Narwhal (Monodon monoceros). Sighted from shipboard at Devil's Island off Cayenne. ". . . appeared but six or eight feet in length, and its horn about four. . . . The narwhal ... is more frequently found in cold than warm climates. The female is said to be unprovided with that protuberance so re- markable in the male. It appears that some authors have confounded this animal with the sword-fish, to which however it does not prove to have the very smallest resemblance." The locality record for the circumpolar narwhal is unexpected, and no doubt erroneous. Nevertheless, Stedman's description is accu- rate albeit the dimensions given seem small. At the same time, Stedman provided a de- tailed description and good figure of a sword- fish or sawfish to prove it was not a sawfish he saw! Volume I, p. 153, pi. 16. Sicapo (Bradypus tri- dactylus). Volume I, p. 153, pi. 16. Dago luyaree (Choloepus didactylus). Volume I, p. 153. Ourang-outang. "I have never seen, nor heard described, while I was in this country. . . ." Volume I, p. 166, pi. 18. Micoo or mecoo {Cebus apella) (fig. 9). 40 HELDIANA: ZOOLOGY •^ jT^^-..^ _^ p* \^^^ )& 'J] J^ u* JC^ # 1 H^'^^!^ %sj ■ .^.:.A^ ■ '^^^ •c s u E t^ < c j= o s (> o it . ^L O J \ (MAY 1800) ^^ ( ■"^ s,y//f ^ ; Cerro Dulda \ / \ '^ --*^f\ ^^^^ — . Vv" jL Q La Esneralda^^ / jg ,• Ms) ^' H**"^" PORTAGE f~^ .i^-r O'^*""*'*"'^-?^ \ ( ^d— — 3" ^< ^^ y^ - 3° »^>^ ^ ^\ y"^ y^ \y< \. / —r' ^^/\J * "'^ / y^^"^ Sa "^vK ^}^^^ ^ Boca MaWcarJ v* /^ V— ^ / /<^ ^X- , fr^ / 1 /^\ *"# ~ — M L ( til 31 ■ *^ \ y^ / ^^ \ 1 f Al *N / ( • 1 _ Solano ^' ^^x^^ / ^^-^^ 2° / \ . „Jl ""^ "^o Negro/ ^ n^s^^ 2" ^\ ^\ *. Vi WJ r-^^ \ \ \ ^ ^^\} r\ A ^>^ V \. J \ 7 • ^ I ^Vr'^ \ ( / \" \ A V-P^ • tf 3V ""^^""^'^ \ \ ( \ ^ X * \0 ^ >. l^.'°\.Cucuy 3\ \ ( ^^y -^'j 1° ^^^"^^^ '^ > A^E^ Catarat. de^^ Cerro de la Nebllna_ _^« \ / ^ "^ 1° ^ :>a* '*"* X ^ ^ \i 1 r^ 1 ) 1 \ 67° 66° 6S- Fig. 1 2. Route of Humboldt and Bonpland in Amazonas, Venezuela, from the Rio Orinoco-Atabapo to the Rio Guainia-Negro via p>ortage between the Rio Temi and Rio Pimichin and the Rio Casiquiare connecting the Negro and Orinoco. expedition up the Rio Orinoco for verification of its reputed connection with the Amazonian Rio Negro. The exploration began on 27 March 1800 with a three-day inspection of a western tributary, the Rio Apure. The journey then continued up the mainstream to the Spanish mission of San Fer- nando de Atabapo near the confluence of the Rios Atabapo and Guaviare with the Orinoco. At this point, the travelers left the Orinoco and continued up the Atabapo to the tributary Temi, which they followed to the tiny mission of Yavita, arriving on 1 May. On 10 May, after portage to the Rio Pimichin, a tributary of the Guainia, they attained San Carlos de Rio Negro at the mouth of the Rio Casiquiare. The next day they headed up the Ca- siquiare and, after 10 days' travel by water, reen- tered the Orinoco on 21 May (figs. 1 1-12). Having confirmed the connection between the HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 53 waters of the Orinoco and Amazon rivers, the ex- plorers shipped 750 miles downstream to arrive at Angostura (Ciudad Bolivar) in mid-June 1800. After more work on the coast, Humboldt and Bon- pland departed Venezuela on 24 November 1800 for Havana, Cuba. They remained there until 1 7 March 1 80 1 , then sailed for Colombia with land- ings along the Rio Sinu on 25 March and Carta- gena on 30 March. The journey thereafter was devoted mainly to explorations of the Cordilleras of Colombia and Ecuador, then through moun- tains, deserts, and the upper Amazonia of Peru south to Lima. The few mammals observed or described during this part of the journey are men- tioned in Humboldt's (1805-181 1) Recueil. From Lima, Humboldt and Bonpland em- barked on 24 December 1 802 for Guayaquil and left 15 February 1803 for Mexico. Humboldt's lively Personal Narrative evokes vi- sions of Venezuelan life and landscapes from coastal plains to the headwaters of the Rio Ori- noco. The narrative is replete with descriptions of geography, ecology, astronomical orientations, widths, depths, and volumes of rivers, histories, languages and customs of Indians, Catholic mis- sions, missionaries, and the human interest trials and tribulations of the travelers. Information on mammals, however, is comparatively meager, but some interesting bits can be quoted or paraphrased from the Ross translation of the original French (Humboldt, 1884). Humboldt and Bonpland found manatees abun- dant in the Rio Orinoco and tributaries Meta and Apure, but absent above the cataracts of Mai- pures. Some of the animals they caught were 1 to 12 feet long and weighed 500 to 800 pounds. Humboldt's dissection of one (fig. 1 3) revealed "no vestige of nails on the external surfaces of the fins which were quite smooth, but little rudiments of nails appear at the third phalanx when the skin of the fins is taken off." The lungs, they observed, consisted of "large cells resembling immense swimming bladders; they [the lungs] are 3 feet long. Filled with air they have a bulk of more than a thousand cubic inches [Humboldt, Ross transla- tion, 1884, vol. II, p. 169]." Its distinction from T. manatus was not appreciated, however, until 1 883 when described by Natterer (in Pelzeln, 1 883). There is also considerable doubt that a clawless manatee does occur in the Rio Orinoco basin or anywhere outside the Amazonian watershed. Dolphins (Sotalia) were seen above and below the great cataracts of the Orinoco and often swam alongside the canoe. In the inundated forest of the divide between the waters of the Orinoco and Ne- gro, the travelers "were astonished by an extraor- dinary noise. On beating the bushes a shoal of toninas (fresh-water dolphins) four feet long sur- rounded our boat. They fled across the forest, throwing out those spouts of compressed air and water. . . ." Other Venezuelan mammals mentioned in the narrative include the expected jaguar, otter, deer, peccaries, capybara, and vampire bats. Monkeys, however, absorbed more of Hum- boldt's attention than other animals. He carried with him a number of live simians captured in the upper Rio Orinoco region for shipment to the Jar- din des Plantes in Paris via the Antillean island of Guadeloupe. The newly discovered bearded saki {Chiropotes satanas chiropotes Humboldt; fig. 14) died before transshipment, but its skin was saved and arrived in Paris. The type specimen of red howler, Simla urslna Humboldt (= Alouatta se- nlculus arctoides Cabrera) survived the journey, whereas the first-known douroucouli or night monkey {Aotus trivlrgatus Humboldt; fig. 14) suc- cumbed in Guadeloupe. Humboldt often mentioned the ubiquitous, highly visible howler or araguato {Alouatta seni- culus). At one time he saw from the road below troops of 30 to 40 individuals crossing through the trees. In a carefully deployed experiment in Ara- gua, he calculated the distance the howler's vo- calization could be heard as 800 toises (6 ft 4.73 inches x 800 = 5,1 15 ft) or nearly 1 mile (5,280 ft). Humboldt (Ross translation, 1884, vol. II, p. 453) recounts the Indian tale of bearded sakis (Chiropotes) and uacaries (Cacajao) of the Orinoco "placing themselves in a circle and, by striking the shell [of the Brazil nut pericarp] with a stone, suc- ceed in opening it so as to take out the triangular nuts." Although Humboldt dismissed the story as fabulous, he did believe that the monkeys cracked the shell of the Bertholletia nut with their teeth to obtain the meat which they devoured with gusto. Belief in the existence of a hairy man of the woods was practically universal. The missionary Father Gili gravely related to Humboldt the tale of a woman "in the town of San Carlos in the Llanos of Venezuela who much praised the gentle character and attentions of the man of the woods. She is stated to have lived several years with one in great domestic harmony, and only requested some hunters to take her back because she and the children (a little hairy also) were weary of living so far from the church and the sacraments." Hum- 54 FIELDIANA: ZOOLOGY IkbU. ^ % Fig. 1 3. The Orinoco clawless manatee, supposedly Trichechus inunguis Natterer left, lateral (1) and ventral (2) views; right, head from above (1), mouth, upper inner view (2), mouth, lower inner view (3), mouth, side view (4), and trunk, sagittal section (5); original illustrations by Humboldt; from Humboldt (1838). boldt resented that he and Bonpland "were every- where blamed, in the most cultivated class of so- ciety, for being the only persons to doubt the reality of the great anthropomorphic monkey of Ameri- ca." Humboldt's Recueil d 'Observations de Zoologie et d 'Anatomic Comparee, a collection of memoires published as a volume in 181 1-1812, deals with many species of invertebrates and vertebrates, but a large share of the text is about monkeys. One memoir with excellent illustrations by Humboldt is on the comparative anatomy of the hyoid bone and larynx of the cotton-top tamarin (Saguinus oedipus oedipus Linnaeus; fig. 1 4), and that of the red howler {Alouatta seniculus seniculus Lin- naeus), the Colombian squirrel {Sciurus granaten- sis granatensis Humboldt; fig. 14), birds, and croc- odiles, all from the Rio Magdalena region. Another memoir on the carnivores includes descriptions of Gulo quitensis (= Conepatus chinga quitensis Humboldt) from Quito, Ecuador, Mustela sinuen- sis (= Eira barbara sinuensis Humboldt), from the Rio Sinu, Colombia, and a discourse on other mustelids and the kinkajou {Potos Jlavus Schre- ber). The memoir on monkeys of the upper Rio Orinoco and connecting Rios Casiquiare and Ne- gro includes the original descriptions o^ Aotus tri- virgatus, Chiropotes satanas chiropotes, Cacajao melanocephalus, Callicebus torquatus lugens, La- gothrix lagothricha, and Cebus albifrons. A chap- ter on the monkeys of Colombia and the upper Amazonian region includes the description of a representative each of Cebus capucinus Linnaeus from the Rio Sinu, A teles belzebuth marginatus E. HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 55 V M. PSIT TAl- 1 S AR Al' RAN A. N" ^ N° Ml. SI n HIS (.RANATF.NSIS. _;?•«• V. vm. SIMIA OKDIHl S. -. •»« ^ Fig. 1 4. Monkeys and anatomical dissections from Humboldt (1811): upper left, Simia melanocephala Humboldt (= Cacajao melanocephalus), holotype; lower left, two views of Simia trivirgata Humboldt (= Aotus trivirgaius), holotype; upper right, Simia satanas Hoffmannsegg (= Chiropotes satanas satanas), lectotype; lower right, throat cartilages of Psiltacus araurana Linnaeus (= Ara araurana), Sciurus granatensis Humboldt, and Simia oedipus Linnaeus (= Saguinus oedipus oedipus). 56 HELDIANA: ZOOLOGY Table 8. New World monkeys (Platyrrhini) recorded by Humboldt (1812); the arrangement is phylogenetic. Current name Humboldt synonym Figure Callitrichidae Callithrix jacchus jacchus Linnaeus, 1758 Callithrix jacchus penicillata E. Geoffroy, 1812 Callithrix jacchus geoffroyi Humboldt, 1812 Callithrix jacchus aurita E. Geoffroy, 1812 Callithrix humeralifer humeralifer E. Geoffroy, 1812 Callithrix argentata melanura E. Geoffroy, 1812 Callithrix argentata argentata Linnaeus, 1771 Saguinus fuscicollis fuscus Lesson, 1840 Saguinus labiatus labiatus E. Geoffroy, 1812 Saguinus midas niger E. Geoffroy, 1 803 Saguinus midas midas Linnaeus, 1758 Saguinus oedipus oedipus Linnaeus, 1758 Leontopithecus rosalia rosalia Linnaeus, 1 766 Cebidae Saimiri sciureus cassiquiarensis Lesson, 1 840 Callicebus moloch moloch Hoffmannsegg, 1 808 Callicebus torquatus lugens Humboldt, 1811 Callicebus torquatus torquatus Hoffmannsegg, 1 808 Callicebus personatus personatus E. Geoffroy, 1812 Actus trivirgatus Humboldt, 1811 Actus azarae azarae Humboldt, 1811 Pithecia mcnachus monachus E. Geoffroy, 1812 Pithecia pithecia pithecia Linnaeus, 1 766 Chiropotes satanas satanas Hoffmannsegg, 1 808 Cacajac melanocephalus Humboldt, 1811 Alouatta caraya Humboldt, 1812 Alouatta seniculus arctoidea Cabrera, 1 940 Alouatta seniculus straminea Humboldt, 1812 Cebus capucinus capucinus Linnaeus, 1758 Cebus nigrivittatus nigrivittatus Wagner, 1 848 Cebus apella apella Linnaeus, 1 758 Cebus apella xanthosternos Wied-Neuwied, 1 820 Cebus apella nigritus Goldfuss, 1810 Lagcthrix lagcthricha lagothricha Humboldt, 1812 Lagothrix lagcthricha cana E. Geoffroy, 1812 Lagcthrix flavicauda Humboldt, 1811 Atetes paniscus chamek Humboldt, 1812 Ateles paniscus paniscus Linnaeus, 1 766 Ateles belzebuth belzebuth E. Geoffroy, 1 806 Ateles belzebuth marginatus E. Geoffroy, 1809 Brachyteles arachnoides E. Geoffroy, 1 806 Jacchus leucccephalus Geoffroy, 1812 Simla leonina Humboldt, 1805, not Shaw, 1800 Simla Ursula Hoffmannsegg, 1808 Not Simla sciurea Linnaeus Simla amicta Humboldt, 1811 Pithecia rufiventer E. Geoffroy, 1812; Simla leuco- cephala E. Geoffroy, 1812 Simla ursina Humboldt, 1805, not Bechstein, 1800 Simla hypoleuca Humboldt, 1811 Simla capucina Humboldt, 1812, not Linnaeus, 1758 Cebus barbatus Humboldt, 1812, attributed to E. Geoffroy Simla variegata Humboldt, 1812, not Kerr, 1 792 Simla cirrifera Humboldt, 1812; Cebus niger E. Geoffroy, 1812 13 14 13 13 Simla chuva Humboldt, 181 1, p. 340; 1812, p. 362, footnote 2 Geoffroy from lower Amazonia, Alouatta senicu- lus Linnaeus from the Rio Magdalena, and La- gothrix flavicauda Humboldt from northern Peru. In an addendum, Humboldt listed all platyrrhine monkeys known to 1812. They are arranged in Table 8 by current scientific names with Hum- boldt's synonyms. X. PARAGUAY The Paraguayan province, claimed by Spain, was first visited in 1526 by Sebastian Cabot and then explored by Cabeza Alvarez Nunez de Vaca in 1541. For the next two centuries, waves of mis- HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 57 Fig. 1 5. Map of Azara's Paraguay and adjacent parts of Brazil and Argentina; from Azara (1809). 58 FIELDIANA: ZOOLOGY sionaries and colonists penetrated to the remotest comers of the province in quest of climates or environments that resembled or could be trans- formed into the familiar ones of Spain. The monumental Histoire du Paraguay by the Jesuit missionary Pierre Francois Xavier de Char- levoix (1682-1761), published in 1757, describes the land that extended from the Atlantic to the eastern base of the Andes between latitudes 1 5° and 35° in the drainage basin of the Rio Parana- Paraguay. It relates the history of the province from the time of the conquest, describes native customs, conversions to Christianity, and estab- lishment of missions. The little of natural history in the text adds nothing about wild mammals not already recorded by others. Two decades later Fe- lix de Azara wrote the most complete natural his- tory account of the mammalian fauna of Paraguay for its time and ever since. Felix de Azara (1746-1811) The Spaniard Don Felix de Azara (1 746-1 811), an army engineer, was commissioned in 1781 to assist in defining the boundaries between Spanish and Portuguese territories. Unmapped territories between Brazil and Paraguay were assigned to Azara, but the Portuguese showed no interest in their share of the work. With time on his hands and a disposition toward the natural sciences, Azara devoted nearly the full 20 years, from 1781 to 1800, of his American residence to the study of geography, Guarani Indians, and the birds and mammals of Paraguay and northeastern Argen- tina between 24° and 36°S and about 54°3r to 56°W (or 60°W of Greenwich) (fig. 15). With no schooling in the natural sciences and no books for reference or guidance, Azara de- pended on his own resources. They proved ade- quate. Azara recorded his observations with care, precision, meticulous attention to detail, and rig- orous exclusion of speculation and fantasy. His anatomical descriptions, measurements, and ac- counts of behavior were based on animals ob- served in the wild or in captivity, usually in his own home or garden. Useful information received from others was credited to the informants. Pop- ular beliefs and hearsay were labeled as such. Without other sources of information, Azara used the Guarani names for most of the amimals he described and Spanish epithets for the remainder. The manuscript of the mammals or quadrupe- dos of Paraguay contained accounts of 66 species. Shortly after its completion, the author received a shipment of several volumes of a Spanish trans- lation of Buffon's Histoire Naturelle. Not surpris- ingly, Azara found in them much with which to disagree, but some of his adverse criticism was unfair. Azara knew Paraguayan mammals better than anyone else, but only a minority of the species were the same as the Neotropical species described in the Histoire Naturelle, and those that were the same did not always behave in the same way at different times or in different places. Azara sent a copy of the manuscript of the quad- rupedos to his brother, Jose Nicolas, then Spanish ambassador to Paris, who arranged for publication in that city after translation into French by M.-L.-E. Moreau de Saint-Mery. A year after his return to Spain in 1801, Azara secured publication in Ma- drid of the original Spanish manuscript with emendations and addition of 1 1 species, for a total of 77. Azara may not have been aware that as many as 62 of the 77 species he described were still un- known to science. His clear and precise charac- terization of each of the species or subspecies, however, provided contemporary cataloguers and systematists with the bases for the descriptions of 50 new species, many with their vernacular ap- pellations in the binomial. Actual specimens served as types for the remaining 1 2 species. The mammals described by Azara are listed be- low, with the scientific name of each given first followed by its local name(s). The page references are to Azara's works in French (Essais, 1801), Spanish (Apuntamientos, 1802), and the Voyage (1809). The last is a French translation in four volumes of Azara's travels in Paraguay with sep- arate atlas, but only the first volume and atlas contain information on mammals. Tapirus terrestris Linnaeus, 1758 Mborebi, Essais I, p. 1; Mborebi, Apunt., I, p. 1; Mborebi ou tapir. Voyage, p. 246. Tayassu G. Fischer, 1814 Coure ou Tayazou, Essais, I, p. 18; Cures o Tayaziis, Apunt., I, p. 14; Cure ou tayazii. Voyage, p. 248. Tayassu pecari albirostris Illiger, 1815 Tagnicati, Essais, I, pp. 2 1 , 25; Taiiicati, Apunt., p. 19; Tanicati, Voyage, p. 249. Bibliographic type of the subspecies. Tayassu tajacu Linnaeus, 1758 Taytetou, Essais, I, pp. 21,31; Taytetvi, Apunt., I, p. 23; Taytetu, Voyage, p. 249. HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 59 CERVIDAE Gazou, Essais. I, p. 43; Venados, Apunt., I, p. 29; Guazu, Voyage, p. 250. Blastocenis dichotomus Illiger, 1815 Gouazoupoucou, Essais, I, p. 70; Guazu-pucu, Apunt., I, p. 33; guazu-pucu. Voyage, p. 250. Bibliographic type of the species. Blastoceros bezoarticus leucogaster Goldfuss, 1817 Gouazouti, Essais, I, p. 77; Giiazu-ti, Apunt., I, p. 41; Guazu-ti, Voyage, p. 251. Bibliographic type of the subspecies. Mazama americana gouazoupita Fischer, 1814 Gouazoupita, Essais, I, p. 82; GUazu-pita, Apunt., I, p. 5 1 ; Guazu-pita, Voyage, p. 252. Bibliographic type of the subspecies. Mazama gouazoubira gouazoubira Fischer, 1814 Gouazoubira, Essais, I, p. 86; Giiazu-bira, Apunt., I, p. 57; Guazu-bira, Voyage, p. 252. Bibliographic type of the species. DIDELPHIDAE Micoures, Essais, I, p. 240; Fecundos, Apunt., I, p. 204; Feconds, Voyage, p. 281. Didelphis albiventris Lund, 1 840 Micoure premier, ou micoure propement dit, Essais, I, p. 244; Micure, Apunt., I, p. 209; Micure, Voyage, p. 283. Caluromys lanatus Olfers, 1818 Micoure second, ou Micoure laineux, Essais, I, p. 175; Lanoso, Apunt., I, p. 221; Lanoso, Voyage, p. 287. Holotype in alcohol, no. 528, Museo de Cien- cias Naturales, Madrid, captured 22 July 1789, by Felix d'Azara (Cabrera, 1916, Bol. Real Soc. espanola Hist. Nat., 16, p. 1). Lutreolina crassicaudata Desmarest, 1 804 Micoure troisieme, ou micoure a queue grosse, Essais, I, p. 284; Coligrueso, Apunt., I, p. 229; Coligrueso, Voyage, p. 290. Bibliographic type of the species. Marmosa pusilla Desmarest, 1 804 Micoure quatrieme, ou micoure a queue longue, Essais, I, p. 290; Colilargo, Apunt., I, p. 251; Colilargo, Voyage, p. 291. Bibliographic type of Marmosa macrura Ol- fers, 1818 (= M. pusilla Desmarest). Micoure sixieme, ou micoure nain, Essais, I, p. 304; Enano, Apunt., I, p. 262; Enano, Voy- age, p. 284. Bibliographic type of Marmosa pusilla Des- marest, 1804. Monodelphis brevicaudis Olfers, 1818 Micoure cinquieme, ou micoure a queue courte, Essais, I, p. 295; Colicorto, Apunt., I, p. 258; Colicorto, Voyage, p. 293. Bibliographic type of the species. MYRMECOPHAGIDAE Hormigueros, Apunt., I, p. 61. Myrmecophaga tridactyla Linnaeus, 1758 Gnouroumi, ou Yoquoui, Essais, \, p. 89; Nu- rumi o Yoqui, Apunt., I, p. 66; Nurumi ou tamandua. Voyage, pp. 253, 255. Tamandua tetradactyla Linnaeus, 1758 (fig. 16) Cagouare, Essais, \, p. 103; Cagiiare, Apunt., \, p. 74; Cagiiare, Voyage, pp. 253, 256; Atlas, pi. VII (tamandua noir), pi. VIII (Cag- uouare). FELIDAE Gatos, Apunt., I, p. 85. Felis onca Linnaeus, 1758 Yagouarete, £'55a/5, 1, p. 1 14; Yaguarete,^;?Mm., I, p. 91; Yaguarete, Voyage, p. 258; Atlas, pi. IX. Yagiiarete negro, Apunt., I, p. 114; Yaguarete noir. Voyage, p. 267. Felis concolor Linnaeus, 1771 Gouazouara, Essais, I, p. 133; Giiazuara, Apunt., I, p. 120; Guazuara, Voyage, p. 268. Felis geoffroyi D'Orbigny and Gervais, 1 844 Mbaracaya, ^/7Mn/., I, p. 147; Baracaya, Voyage, p. 271. Note: Said not to exist in Paraguay. Felis species? Negro, Apunt., I, p. 154; Chat noir. Voyage, p. 273. Felis pardalis Linnaeus, 1758 Chibigouazou, Essais, I, p. 152; Chibi-giiazii, Apunt., I, p. 132; Chibi-guazu, Voyage, p. 269. Herpailurus yagouaroundi eyra Fischer, 1814 (fig. 16) Yagouaroundi, Essais, I, p. 171; Yaguarundi, Apunt., I, p. 156; Yaguarundi, Voyage, p. 273, Atlas, pi. X (Yagouarondi, black phase); Eyra, Essais, I, p. 177; Eyra, Apunt., I, p. 159; Eyra, Voyage, p. 274 (red phase). Bibliographic type of the subspecies. Felis colocolo pajeros Desmarest, 1816 Chat pampa, Essais, I, p. 1 79; Pajero, Apunt., I, p. 160; Pajero, Voyage, p. 274. Bibliographic type of the species. Note: Said not to exist in Paraguay. 60 FIELDIANA: ZOOLOGY .(■ ').i.'..M.ire of the species. Myotis albescens E. Geoffroy, 1 806 Chauve-souris douzieme, ou chauve-souris brun-obscur, Essais, II, p. 294; Pardo ob- scuro, Apunt., II, p. 309. Bibliographic type of the species. Johann Rudolph Rengger (1795-1832) Azara was followed by Johann Rudolph Reng- ger, a Swiss pharmacist and naturalist, who arrived in Paraguay in 1819 and devoted himself to the study of its mammals. His six-year study culmi- nated in the Naturgeschichte der Saeugethiere von Paraguay, published 1830. A total of 59 species was described, including four as new of which only Calomys callosus and Proechimys longicaudatus survived revisions. Azara distinguished 77 species, or 1 8 more, but several are not strictly Paraguayan. Among the Paraguayan forms missed by Rengger but recognized by Azara are the murine opossum (Marmosa), hairy armadillo (Chaetophractus), three-banded armadillo (Tolypeutes), skunk (Co- nepatus), tucotuco (Ctenomys), four cricetine ro- dents, and two bats. Well over 100 species are presently known from Paraguay. No doubt Azara set standards for the high qual- ity and accuracy of Rengger's descriptions and be- havioral accounts. The wealth of information in the Naturgeschichte has hardly been tapped by modem mammalogists. XI. Chile Giovanni Ignazio Molina (1737-1829) Knowledge of Chilean land mammals as a re- gional fauna begins with publication of the Saggio in 1782 by the Jesuit priest Don Giovanni (Juan) Ignazio Molina, who lived in Chile the first 30 years of his life. Expulsion of the Jesuits from the country obliged Molina to emigrate in 1768 and settle in his ancestral Italy. What Molina knew about Chilean mammals he learned before 1768; much of what he wrote about them thereafter suf- fered from a decayed memory. Molina was a naturalist in the broadest sense 64 HELDIANA: ZOOLOGY and was familiar with the Systemce of Linnaeus. He was not, however, particularly dedicated to any one branch of science, and his descriptions of the Chilean mammals are, for the most part, vague, inaccurate, and sometimes composite. A few of his subjects were fanciful, and none of the re- mainder were closely examined. Nevertheless, by dint of elimination and stretches of the imagina- tion, modem mammalogists have come to agree- ment on the application of most of the Linnaean names proposed by Molina for the likeliest species he may have had in mind. Thirty kinds of mammals were described in the Saggio. According to Osgood (1943, p. 15), five of them are unidentifiable, four (armadillos) are extraterritorial, two are but one and the same, and one is duplicated. The 14 still valid, with names dating from Molina, 1782, are Lutra felina, My- ocastor coypus, Conepatus chinga, Galictis cuja, Dusicyon culpaeus, Felis guigna, Felis colocolo, Felis concolor puma, Spalacopus cyanus, Octodon degus, Vizcacia vizcacia, Pudu puda, Vicugna vi- cugna, and Hippocamelus bisulcus. Remaining species, notably the larger mammals, recorded by Molina were well known to early voyagers, chron- iclers, and naturalists and had already received Linnaean names. first volume (1847) of eight on zoology contains virtually all Chilean mammals known at the time. Fifty-four species are described, with accounts of habits, habitat, and geographic distribution of each. For the most part. Gay worked from actual spec- imens brought to him by natives or observed by him on his travels throughout the country. On his return to France, Gay included in his studies the Chilean material preserved in the Paris Natural History Museum. The species recorded by Gay include Marsupi- alia, 2 (4% of the total); Chiroptera, 7 (13%); Gar- ni vora, 12 (22%); Pinnipedia, 6(11%); Rodentia, 23 (43%; myomorphs, 24%, caviomorphs, 18%); Artiodactyla, 3 (5%). Among the 30 species re- corded by Molina, only 3 or 10% are rodents. Of the 20 Chilean species collected by Darwin, 12 or 60% are rodents. In this volume Patterson and Feigl recognize 93 living Chilean sF)ecies, of which 53 or 57% are rodents (33% myomorphs, 24% caviomorphs), and 1 or 1 1 % are bats. XII. Peru Johann Jacob von Tschudi (1818-1889) Eduard Friedrich Poeppig (1798-1868) The German naturalist Eduard Poeppig is known for his Reise in Chile, Peru, and on the Rio Ama- zonas during the years 1827-1832. The account of his travels, in two volumes, was published 1835-1836. The Chilean mammals recorded in- clude seals, sea lions, and elephant seals, the degu, Spalacopus cyanus Molina (Psammomys nocti- vagus Poeppig, a synonym), the coypu, and a small canid, probably Dusicyon griseus Gray. In Antuco, Province of Bio Bio, he encountered the pudu, huemul, and two species of bats, one described as Nyticyus varius (= Lasiurus borealis bonariensis Lesson & Gamot, 1827), the other as Nycticyus macrotus (currently Histiotus macrotis Poeppig, 1835). Claudio Gay (1800-1873) Between the years 1 844 and 1871, Claudio Gay, French naturalist and longtime resident of Chile, produced 25 volumes, including two of plates, on the history, geography, and biota of Chile. The The Swiss biologist Johann von Tschudi was bom in the town of Glarus and studied the sciences at Swiss, French, and German universities. In- spired by the accounts of the travels of Humboldt and Darwin in South America, Tschudi sailed on 27 February 1838 from Le Havre for Peru. The first landing on the continent was made 5 June 1 838 on the Chilean island of Chiloe. After a delay of about three weeks and many observations of the natural history of the island, von Tschudi reembarked for Callao, Pern, with short stopovers in Valdivia and Juan Femandez. From August 1838 through most of 1843, von Tschudi traveled over much of Peru. Of particular interest to him were the higher vertebrates and the physical factors controlling their geographic dis- tribution. He distinguished faunal zones based on mling ecological features. The major zones were Pacific coast, Andean altitudinal zones of westem and eastem slopes, and the tropical Amazonian selva. Apparently, no one had preceded von Tschudi in the recognition of definable biogeo- graphic areas in the New World. The narrative of von Tschudi's travels in Pern was published in 1846 in German, followed in 1847 by Thomasina Ross's English translation. HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 65 It li I ^1 si ll « 5 3 C U S Si, If J it ^6^ . 3 X) O w 66 HELDIANA: ZOOLOGY The scientific accounts of the mammals are found in a preliminary report (1844a) and first part of the Untersuchungen liber die Fauna Peruana, pub- lished later the same year ( 1 844b). Although von Tschudi attempted to provide the fullest account possible of Peruvian mammals, it appears he had little or no contact with the ma- jority of them. Most of his characterizations and life history accounts are taken from Humboldt, Spix, Wied-Neuwied, other European travelers and natives. Camelids, the dominant animals of the Peruvian landscape fascinated von Tschudi, and he wrote more about them than of other animals. His description of a vicuria hunt is quoted below from the Ross translation (Tschudi, 1 847, pp. 2 1 9- 220). The Indians seldom employ fire-arms in hunting the vicunas. They catch them by what they term the chacu. In this curious hunt, one man at least belonging to each family in the Puna villages takes a part, and women accompany the train, to officiate as cooks to the hunters. The whole company, frequently amounting to seventy or eighty individuals, proceeds to the Altos (the most secluded parts of the Puna), which are the haunts of the vicuiias. They take with them stakes, and a great quantity of rope and cord. A spacious open plain is selected, and the stakes are driven into the ground in a circle, at intervals of from twelve to fifteen feet apart, and are connected together by ropes fastened to them at the height of two or two and a half feet from the ground. The circular space within the stakes is about half a league in circumference, and an opening of about two hundred paces in width is left for en- trance. On the ropes by which the stakes are fastened together the women hang pieces of colored rags, which flutter about in the wind. The chacu being fully prepared, the men, some of whom are mounted on horseback, range about within a circuit of several miles, driving before them all the herds of vicurias they meet with, and forcing them into the chacu. When a sufficient number of vicunas is collected, the entrance is closed. The timid animals do not attempt to leap over the ropes, being frightened by the fluttering rags sus- pended from them, and, when thus secured, the Indians easily kill them by the tolas. These bolas consist of three balls, composed either of lead or stone; two of them heavy, and the third rather lighter. They are fas- tened to long, elastic strings, made of twisted sinews of the vicuria, and the opposite ends of the strings are all tied together. The Indian holds the lightest of the three balls in his hand, and swings the two others in a wide circle above his head; then taking his aim at the distance of about fifteen or twenty paces, he lets go the hand-ball, upon which all the three balls whirl in a circle, and twine round the object aimed at. The aim is usually taken at the hind legs of the animals, and the cords twisting round them they become firmly bound. It requires great skill and long prac- tice to throw the bolas dexterously, espe- cially when on horseback: a novice in the art incurs the risk of dangerously hurting either himself or his horse, by not giving the balls the proper swing, or by letting go the hand-ball too soon. The vicuiias, after being secured by the bolas, are killed, and the flesh is distributed in equal portions among the hunters. The skins belong to the Church. The price of a vicuna skin is four reals. When all the ani- mals are killed, the stakes, ropes, &c., are packed up carefully, and conveyed to another spot, some miles distant, where the chacu is again fixed up. The hunting is continued in this manner for the space of a week. The number of animals killed during that inter- val varies according to circumstances, being sometimes fifty or sixty, and at other times several hundred. During five days I took part in a chacu hunt in the Altos of Huayhuay, and in that space of time 122 vicurias were caught. With the money obtained by the sale of the skins a new altar was erected in the church of the district. The flesh of the vicuiia is more tender and better flavored than that of the llama. Fine cloth and hats are made of the wool. When taken young, the vicurias are easily tamed, and become very docile; but when old, they are intractable and ma- licious. At Tarma I possessed a large and very fine vicuria. It used to follow me like a dog whenever I went out, whether on foot or on horseback. The frequent hunting seems not to have the effect of diminishing the numbers of these animals. If in the vicinity of the villages where chacus are frequently established, they are less numerous than in other parts, it is because, to elude the pursuit of the hunters, HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 67 Table 9. Peruvian mammals according to Tschudi (1844a,b); current scientific names are used followed by Tschudi's synonym or misidentification, local names, and figure in this text; extralimital species are bracketed; arrangement of taxa follows Tschudi. Current wune Tschudi synonym or misidentification Local name Figure Aieles paniscus chamek Hum- boldt LagothrLx lagothricha poeppigi Schinz LagolhrLx flavicauda Humboldt Alotmtta seniculus Linnaeus [Mycetes rufimanus Kuhl] Cebus apella Linnaeus Cebus albifrons Humboldt [Cebus capucinus Linnaeus] Saimiri boliviensis peruviensis Hershkovitz Callicebus torquatus Hofimann- segg (subsp.?) [Callicebus personatus E. Geof- froyl Aotus nigriceps E)ollman [Chiropotes] Saguinus mystax mystax Spix Saguinus nigricollis Spix Saguinus fuscicollis Spix [Saguinus midas midas Lin- naeus] [Leontopithecus rosalia chryso- melas Kuhl] Chiroptera Phyllostomus elongatus E. Geof- fh)y Phyllostomus hastatus Pallas Phvlloslomus discolor Wagner, i843 Artibeus cinereus Gervais Stumira erythromos Tschudi Sturnira oporophilum Tschudi Glossophaga soricina Pallas Anoura geoffroyi peruana Tschudi Eptesicus innoxius Gervais Histiotus macrotus Poeppig Noctilio leporinus Linnaeus Noctilio albiventris Desmarest Tadarida brasiliensis I. Geoffroy Xfolossus molossus Pallas? Eumops auripendulus Shaw Molossus ater E. Geofiroy [Promops nasutus Spix] Carnivora Tremarctos omatus F. Cuvicr Nasua nasua montana Tschudi Potosflavus Schreber Eira barbara Linnaeus Ateles marginatus; Aieles ater; Ateles pentadactylus Lagothrix humboldti; Lagothrix canus Mycetes flavicaudatus (sic) Mycetes stramineus Alouatta belzebul Cebus robustus Chrysothrix sciureus Callithrix amictus Callithrix personatus Nyctipithecus trivirgatus IPithecia* [Midas labiatus] [Midas labiatus] [Midas labiatus] [Midas rufimanus] [Midas chrysomelas] Chuva; maquisapa; chamek; mahmonda; machucusillo; supaya Mono oki; choko Coro [= coto?] Macaquito Tocon Hatmnmasu Phyllostomus innominatum Tschudi Phyllostomus (Artibeus) pusil- lum Phyllostomus (Sturnira) oporo- philum Tschudi Glossophaga amplexicauda Glossophaga (Choeronycteris) peruana Tschudi Vespertilio innoxius Vespertilio ( Vesperugo) velatus Noctilio unicolor Noctilio affinis Molossus (Dysopes) naso Molossus (Dysopes) velox Molossus (Dysopes) ferox; Dy- sopes longimanus Molossus (Dysopes) myosuros Tschudi; Molossus anonymus Tschudi Dysopes fitmarius Ursus fivgilegus Tschudi Nasua socially, Nasua solitaria, Nasua leucorhynchos Tschudi Cercoleptes caudivolvidus Galictis barbara Hucamari Achuna, mishash Cushumbi Omeyro 17 68 HELDIANA: ZOOLCXJY Table 9. Continued. Current name Tschudi synonym or misidentification L4>cal name Figure Carnivora {continued) Mustela frenata agilis Tschudi Conepatus chinga Molina Lutra felina Molina Lutra montana Tschudif Dusicyon thous Linnaeus Felis concolor Linnaeus Felis onca Linnaeus Felis pardalis Linnaeus Feiis wiedii Schinz Felis yagouaroundi E. Geoffroy PlNNIPEDIA Otaria flavescens Shaw Marsupialia Didelphis marsupialis Linnaeus Metachirus nudicaudatus E. Geoffroy Philander opossum Linnaeus Marmosa noctivaga Tschudi Marmosa impavida Tschudi Marmosa murina Linnaeus Caluromys lanatus ornatus Tschudi RODENTIA Sciurus aestuans Linnaeus Sciurus pyrrhinus Thomas Sciurus stramineus Eydoux and Souleyet Sciurus spadiceus tricolor Tschudi Proechimys sp.? Chinchilla brevicaudata Water- house Lagidium peruanum Meyen Lagidium viscacia Molina^ [Octodon degus Molina] [Myocastor coypus Molina] Coendou bicolor Tschudi Dasyprocta leporina Linnaeus Dasyprocta variegata Tschudi Akodon boliviensis Meyen Phyllotis darwini Waterhouse Oryzomys longicaudatus de- structor Tschudi Oryzomys melanostoma Tschu- di Rhipidomys leucodactylus Tschudi Agouti paca Linnaeus Hydrochaeris hydrochaeris Lin- naeus Cavia porcellus Linnaeus Molina (Thiosmus) mapurita; Mephitis furcata; Mephitis amazonica Lutra chilensis Canis azarae Felis onza Felis macrura (sic = Felis ma- croura); Felis celidogaster Felis yaguaruruii Otaria jubata; Otaria ulloae Tschudi; Otaria aurita Hum- boldt (in Tschudi) Didelphys azarae Didelphys myosuros Poma, leon Choque china, yana cheque, tigre Uturunco Mucamuca, jarachupa 17 \Sciurus variabilis] [Echinomys leptosoma] Eriomys chinchilla Lagidium peruvianum {sic) Lagidium pallipes [Octodon cummingii] [Myopotomus coypus] Sphingurus {sic) bicolor Dasyprocta aguti Linnaeus Acodon boliviense Hesperomys darwini Hesperomys destructor Hesperomys melanostoma Hesperomys {Rhipidomys) leu- codactylus Coelogenys fulvus Hydrochoerus capybara Cavia cutleri Cutspi or cushpi 17 Cuy del monte HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 69 Table 9. Continued. Current name Tschudi synonym or misidentification Local name Figure Lagomorpha Sylvilagus brasiliensis Linnaeus Edentata Bradypus vahegatus Schinz [Bradypus torquatus lUiger] Dasypus novemcinctus Linnaeus Cabassous unicinctus Linnaeus Tamandua tetradactyla Lin- naeus Cyclopes didactylus Linnaeus Perissodactyla Tapirus terrestris Linnaeus Tapirus pinchaque Roulin Artiodactyla Tayassu tajacu Linnaeus Tayassu pecari Link Lama glama Linnaeus Lama pacos Linnaeus Lama guanicoe Miiller Vicugna vicugna Molina Mazama americana Erxleben Mazama gouazoubira peruana Tschudi Hippocamelus antisensis d'Or- bigny Lepus brasiliensis Bradypus infuscatus Dasypus 9-cinctus {sic) Dasypus tatuay {sic = tatouay) Myrmecophaga tamandua Myrmecophaga didactyla Quirquincho Tapirus americanus Tapirus villosus Dicotyles torquatus Dicotyles labiatus Auchenia lama Llama Auchenia paco Alpaca Auchenia huanaco Auchenia vicuna Vicuna Cervus rufus Cervus nemorivagus var. per- Liucho, venado uana Cervus antisiensis Tarush, taruga 17 * Sakis {Pithecia) evidently not seen by von Tschudi. His descriptions are of bearded sakis {Chiropotes) after Humboldt (1811), which do not occur in Peru, t May not be an otter, according to Thomas (1908, p. 393). % The species was known to occur in parts of formerly southwestern Peru now in Chile. they seek refuge in the Altos, where they are found in vast numbers. Several modem travelers have lamented the diminution of the vicunas, but without reason. In fonner times those animals were hunted more ac- tively than at present. Von Tschudi's journeys in the puna inspired him to poetic descriptions of the habits, particu- larly the visual propensities, of its denizens. Herds of vicuiias approached me with cu- rious gaze, and then on a sudden fled with the swiftness of the wind. In the distance I observed stately groups of huanacos turning cautiously to look at me, and then passing on. The Puna stag (tarush) slowly advanced from his lair in the mountain recesses, and fixed on me his large, black, wondering eyes, whilst the nimble rock rabbits (viscachas) playfully disported and nibbled the scanty herbage growing in the mountain crevices. (Tschudi, Ross translation, 1 847, p. 249) On descending the eastern slope of the Cordi- lleras to the subtropical zone inhabited by a greater variety of different kinds of mammals, von Tschu- di (Ross translation, 1847, p. 275) romanticized that: . . . the swift-footed roe [Mazama sp.] of the Cordillera roams here and dwells in the thickets, avoiding the warm forest. The dark brown coati {Nasua montana, Tsch.) howls and digs at the root of trees in search of food, the shy opossum crawls fearfully under the foliage; the lazy armadillo creeps into his hole, but the ounce [Felis onca] and the lion [Felis concolor] seldom stray hither to con- test with the black bear {Ursus frugilegus Tsch.) the possession of his territory. The 70 FIELDIANA: ZOOLOGY little hairy tapir (Tapirus villosus, Wagn.) ventures only at twilight out of his close am- bush to forage in the long grass. The systematic arrangement in the Untersu- chungen is said to include all mammals known at the time to occur in Peru. By von Tschudi's count, the fauna consists of 1 1 9 species in 48 genera. These totals include domestic animals, the intro- duced house mouse, some duplicated names of native species, and a number of others not known to occur in Peru. In terms of currently recognized species found in Peru, von Tschudi's combined lists (1844a, pp. 244-255; 1844b, pp. 6-20; 21- 264) consist of 87 species in 58 genera. The species are listed in Table 9 with von Tschudi's synonyms or misidentifications. Author attributions of the synonyms are omitted unless they are to von Tschudi himself Vernacular names, if given, are included. Extralimital species are shown in brack- ets. In the case of unrevised groups or where two or more subspecies occur in Peru without possi- bility of determining which were described by von Tschudi, only the specific names are given. XIII. Patagonia Alcide Charles Victor d'Orbigny (1802-1857) The French-bom Alcide d'Orbigny was educat- ed by his country's leading naturalists. His apti- tudes were recognized by authorities of the Mu- seum National d'Histoire Naturelle, and with that institution's financial and material assistance, he sailed for South America charged with making a scientific survey of the southern half of the con- tinent. Circumstances restricted his studies and collections of mammals almost entirely to Argen- tina and Bolivia. D'Orbigny left France 3 1 July 1 826 and arrived in Rio de Janeiro 24 September 1826 on his way to Montevideo where he landed on 29 September. The natural history of the region between Mal- donado east of Montevideo and Buenos Aires en- gaged his attention for several months. On 14 February 1827, d'Orbigny ascended the Rio Parana and arrived 1 5 March at the important fluvial port of Corrientes, capital of the province of the same name. With the town as base, d'Or- bigny explored the province throughout much of one year. On his return to Buenos Aires in April 1828, he made stops in Entre Rios and Santa Fe. Beginning June 1828 and continuing through 1829, his at- tentions were devoted to faunal studies in the provinces of Buenos Aires and Rio Negro. The chronology of the early part of 1829, as given by d'Orbigny (1835-1847) in the Voyage, confuses time spent in the two provinces with that spent in Corrientes. In any event, d'Orbigny was clearly in Buenos Aires and Rio Negro during the last half of 1 829. He returned to Montevideo in December 1 829 and on 29 December sailed on to Patagonia and Chile. Cape Horn was rounded on 19 January 1830 and Valparaiso, Chile, was reached 16 February. Because of the political unrest in the country, d'Or- bigny sailed to the then Bolivian port of Cobija, where he landed on 8 April; 20 April found him in Arica and Tacna, both ports then in Peru's pos- session. After some investigation of the coast, d'Orbigny left Tacna on 19 May for La Paz, the mountain capital of Bolivia, arriving there 28 or 29 May. For the next three years, d'Orbigny explored, mapped, and sampled the natural resources of the country. He crisscrossed Bolivia from La Paz east to the Paraguayan border and from Potosi in the south to the lower Rio Mamore in the north. D'Or- bigny's actual itinerary is almost impossible to track because of the inaccuracies of the then avail- able maps. Modem maps aided Pilleri and Arvy (1977) in their reconstmction of the itinerary in chronological sequence (fig. 1 8). A complete account of d'Orbigny's South Amer- ican joumey with observations on and descrip- tions of the geology, paleontology, living plants, animals, and Indians is contained in seven huge volumes published serially from 1 835 through 1847 in Paris under the title Voyage dans I'Amehque Meridionale. A full report on the mammals was reserved for the last, or perhaps a separate pub- lication, but a turn in d'Orbigny's fortunes inter- rupted the work. A number of colored plates of mammals believed new to science and a few short articles on others had already been published. So that all would not be lost, a synoptic systematic report on the mammals collected was published in 1 847 jointly with the distinguished mammal- ogist Paul Gervais, as number 2 of volume 4 of the Voyage. Brief notes on distribution and be- havior accompany the abbreviated descriptions of each species. The species are listed in Table 10 with abstracted locality data. Scientific names used are current with synonyms and misidentifications added. The specimens are deposited in the Mu- seum National d'Histoire Naturelle in Paris. HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 71 5 : ' '■ i =. I ? I •? :^ ^ < Is I § I 1 1 5 , 1 , ; = £ II ^ I I 1 1 s I §• z X i -i £ 3. ^ ^ > f- }. i. i^ EccSccccc'.cc :^ ;? j: c ?. ■?. ■?. f. ■?. a i ■>. I •e o CO 3 P I 4f 72 FIELDIANA: ZOOLOGY CAI.ITKIN M».pkn" '*• r M.I.ITIIKIN J..— i^M.. I. . ■■! .'■■■!■ Fig. 1 9. Animals of the d'Orbigny Bolivian Expedition: upper left, Callithrix entomophagus d'Orbigny (= Saimiri boliviensis boliviensis I. Geoffroy and Blainville); upper right, Callithrix donacophilus d'Orbigny (= Callicebus do- nacophilus donacophilus); lower left, Felis geoffroyi d'Orbigny and Gervais (= Felis colocolo geoffroyi); lower right. Mephitis humboldtii (= Conepatus chinga suffocans Illiger); from d'Orbigny and Gervais (1847). HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 73 Table 10. Mammals of the southern half of South America, mostly Bolivia and Argentina, recorded by d'Orbigny and Gervais (1847); the arrangement is phylogenetic. Current name d'Orbigny and Gervais synonym Locality Figure Chiroptera Noctilio albiventris Desmarest, 1818 Noctilio leporinus rufipes d'Or- bigny, 1835 Tonatia sylvicola d'Orbigny, 1835 Artibeus planirostris Spix, 1 823 Desmodus rotundus E. Geoffroy, 1810 Myotis nigricans Wied-Neu- wied, 1821 Eptesiciis furinalis d'Orbigny and Gervais, 1 847 Myotis albescens E. Geoffroy, 1806 Myotis ruber E. Geoffroy, 1 806 Histiotus velatus I. Geoffroy, 1824 Tadarida brasiliensis I. Geof- froy, 1824 Molossus crassicaudatus E. Geof- froy, 1805 Primates Saimiri boliviensis boliviensis I. Geoffroy and Blainville, 1 834 Callicebus donacophilus donaco- philus d^Orhigny, 1835 Alouatta seniculus sara Elliot, 1910 Cebi4s apella paraguayanus Fischer, 1829 Carnivora Dusicyon gymnocercus Fischer, 1814 Chrysocyon brachyurus Illiger, 1815 Tremarctos ornatus F. Cuvier, 1825 Procyon cancrivorus nigripes Mivart, 1886 Nasua nasua solitaria Wied- Neuwied, 1821 PotosflavusSchxc\xT, 111 A Lyncodon patagonicus Blain- ville, 1842 Galictis cujafurax Thomas, 1907 Conepatus chinga suffocans Illi- ger, 1815 Lutra platensis Waterhouse, 1838 Noctilio affinis d'Orbigny, 1835 BOLIVIA: Moxos Province BOLIVIA: Chiquitos and Moxos provinces BOLIVIA: Yuracare territory, base of eastern Cordillera BOLIVIA: Chiquitos Province Not Vespertilio perspicillatus Linnaeus, 1758 Desmodus rufus Wied-Neuwied, 1824; Edostoma cinerea d'Or- bigny, 1835 Vespertilio hypothrix d'Orbigny and Gervais, 1847 Vespertilio isidori d'Orbigny and Gervais, 1847 Molossus rugosus d'Orbigny, 1835, not Molossus nasutus Spix, 1823 Molossus moxensis d'Orbigny, 1835; Molossus velox Tem- minck, 1827 Calithrix (sic) entomophagus d'Orbigny, 1835 Not Stentor stramineus E. Geof- froy Cebus fulvus var. Not Canis cancrivorus Desma- rest, 1820 Canis jubatids Desmarest, 1820 Not Procyon cancrivorus Cuvier, 1798 Nasua fusca Desmarest, part Cercoleptes caudivolvulus Schre- ber, 1774 Not Mustela brasiliensis Gme- lin, 1788 Mephitis castaneus d'Orbigny and Gervais, 1 847, not Me- phitis humboldtii Gray, 1837 BOLIVIA: Chiquitos BOLIVIA: Moxos ARGENTINA: Corrientes ARGENTINA: Corrientes ARGENTINA: Corrientes BOLIVIA: Chuquisaca ARGENTINA: Corrientes BOLIVIA: Moxos and Chiqui- tos provinces BOLIVIA: Chiquitos; Moxos; Santa Cruz BOLIVIA: Moxos Province BOLIVIA: Santa Cruz; Chiqui- tos; Moxos BOLIVIA: near Santa Cruz de la Sierra BOLIVIA: Chiquitos Tropical South America to 41°S BOLIVIA: Cochabamba; Chu- quisaca BOLIVIA: Chiquitos; ARGEN- TINA: Corrientes BOLIVIA: tropics to 30'^ BOLIVIA: foot of eastern Cordi- llera ARGENTINA: Rio Negro 19 19 19 ARGENTINA: Rio Parana in Provinces Buenos Aires and Corrientes 74 FIELDIANA: ZOOLOGY Table 10. Continued. Current name d'Orbigny and Gervais synonym Locality Figure Carnivora Felis colocolo pajeros Desma- rest, 1816 Felis geoffroyi d'Orbigny and Gervais, 1847 Felis concolor Linnaeus, 1771 Felis onca Linnaeus, 1758 PiNNIPEDIA OtariaflavescensShs^N, 1800 Arctocephalus australis Zimmer- mann, 1782 Mirounga leonina Linnaeus, 1758 Artiodactyla Mazama gouazoubira Fischer, 1814 Blastoceros bezoarticus Lin- naeus, 1758 Hippocamelus antisensis d'Or- bigny, 1834 Blastocerus dichotomus Illiger, 1815 RODENTIA Sciurus spadiceus Olfers, 1818 Eligmodontia typus F. Cuvier, 1837 Octodon degus Molina, 1782 Octodontomys gliroides, Gervais and d'Orbigny, 1 844 Ctenomys boliviensis Water- house, 1848 Ctenomys magellanicus Bennett, 1835 Microcavia australis Gervais and d'Orbigny, 1833 Galea flavidens Brandt, 1835 Dolichotis patagonum Zimmer- man, 1780 Dasyprocta azarae Lichtenstein, 1827 Cetacea Inia boliviensis d'Orbigny, 1834 [Pontoporia blainvillei Gervais and d'Orbigny, 1 844; not part of d'Orbigny collection] Lagenorhynchus cruciger Quoy and Gaimard, 1 824 Lissodelphis peroni Lacepede, 1804 Otaria jubata Schreher, 1776 Otaria porcina Molina, 1782 Phoca proboscidea Peron, 1817 Cervus simplicicornis Illiger, 1815 Not Cervus campestris F. Cu- vier, 1817 Cervus paludosus Desmarest, 1822 Not Sciurus igniventris Wagner, 1842 Not Ctenomys brasiliensis Blainville, 1826 Dasyprocta patachonica Desma- rest, 1820 Not Dasyprocta nigricans Wag- ner, 1842 ARGENTINA: from 35°-45'S ARGENTINA: Pampas to 44'S 19 BOLIVIA; ARGENTINA: to Straits of Magellan Tropical South America not be- yond 40^; ARGENTINA: Pampas; Serrania de Tandil ARGENTINA: S mouth Rio Negro ARGENTINA: coast; PERU: coast ARGENTINA: Rio Negro, near mouth Tropical South America to 28°S Lowland savannas to northern Patagonia BOLIVIA: La Paz; Cochabam- 20 ba; Chuquisaca; rarely below 3500 m ARGENTINA: Corrientes; BO- LIVIA: Chiquitos BOLIVIA: Chiquitos ARGENTINA: Corrientes CHILE: Santiago de Chile BOLIVIA: La Paz ARGENTINA: Corrientes; BO- LIVIA: Santa Cruz de la Sie- rra ARGENTINA: northern Pata- gonia ARGENTINA: Rio Negro BOLIVIA: Cochabamba; Chu- quisaca; La Paz ARGENTINA: northern Pata- gonia; Corrientes Tropical South America BOLIVIA: rivers of Moxos and 20 Chiquitos URUGUAY: Montevideo Atlantic Ocean (57»-76'«, E and S of Cape Horn) Atlantic Ocean (48°-64'«); At- lantic-Pacific Oceans around Cape Horn HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 75 ■' S, : i ^ 'b 1 1 \Vi «J M S V ' i ,^ \ I ^ ja ■•»■ ^ •c f- is ■* \ \ ■_ 1 tj J "3 11 ^ u \ i •1 ■■i I « 'm ■/. 1U "b t^ 1 1 ■Si <« ;§« ^ t "^ e SO 11 X S c^. •2 12 S 1 pa CO .!>•§ O > e s ected, soon found it. HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 77 lOV Coplapo'U a Coqulabo pLl- Coocepcirfn J9 \ l Buenos Aires ^-xJUjtontevldeo Y (SIERRA DF, ^fo Plata CHARLES DARUIN TOTAGE OF B.M.S. BEACLE SOUTH AMERICAN LOCALITIES (1832-1835) CBOHOS*- AXCRIPELACO^^ ,' -■w T. Good Success (■snia dc Good Uickf 80* Hp-y^^ ^/San Julian FALKLAND ISLANDS tllUA DEL ruioo Canai fagla J 1 L. 60* 50* 30* Fig. 2 ! . Map showing principal South American stations visited by Charles Darwin ( 1 832-1 835) on world cruise of H.M.S. Beagle (1832-1836). 78 HELDIANA: ZOOLOGY u HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 79 Table 1 1 . Mammals collected or observed by Oanvin in the Maldonado Region, Uruguay and parts of Argentina, and those recorded by Waterhouse (1838-1839); the arrangement is phylogenetic. Current name Waterhouse synonym or misidentification Locality Figure Marsupialia Didelphis albiventris Lund Lutreolina crassicaudata Des- marest Monodelphis dimidiata Wagner Chiroptera Tadarida brasiliensis I. Geoffroy Edentata Dasypus hybridus Desmarest Zaedyus pichiy Desmarest Chaetophractus villosus Desma- rest Tolypeutes matacus Desmarest Carnivora Dusicyon gymnocercus Fischer Felis colocolo pajeros Desmarest Galictus cujafurax Thomas Lutra platensis Waterhouse Conepatus chinga gibsoni Thomas Felis concolor acrocodia Gold- man Felis onca palustris Ameghino Artiodactyia Blastoceros bezoartiats Lin- naeus Lama guanicoe Muller RODENTIA Myomorpiia Oryzomys flavescens Waterhouse CaJomys laucha Olfers Eligmodontia typus Cuvier Holochilus brasiliensis darwini Thomas Reithrodon physodes typicus Waterhouse Akodon azarae Fischer Akodon colibreve Brants Scapteromys tumidus Water- house Oxymycierus rufus nasutus Waterhouse Caviomorpiia Cavia porcellus Linnaeus Hydrochaeris hydrochaeris Lin- naeus Didelphis azarae AucL Didelphis brachyura Auct. Not Dysopes nasutus Spix Dasypus minutus AucL Not Canis azarae Wied-Neu- wied Not Galictis vittata Schreber Not Cervus campestris Cuvier Mus bimaculatus Waterhouse; Mus gracilipes Waterhouse Mus elegans Waterhouse Mus arenicola Waterhouse Mus obscurus Waterhouse Cavia cobaia Auct. Hydrochoerus capybara Auct. URUGUAY: Maldonado URUGUAY: Maldonado URUGUAY: Maldonado URUGUAY: Maldonado URUGUAY: Maldonado ARGENTINA: Banda Oriental, Entre Rios ARGENTINA: Bahia Blanca (observed) ARGENTINA: Bahia Blanca (observed) ARGENTINA: La Plata (ob- served) ARGENTINA: Bahia Blanca URUGUAY: Maldonado URUGUAY: Maldonado ARGENTINA: Bahia Blanca (observed) ARGENTINA: the pampas (ob- served) ARGENTINA: in the Rio Pa- rana (observed) URUGUAY: Maldonado; AR- GENTINA: Bahia Blanca; Rio Negro ARGENTINA: Rio Negro (ob- served) URUGUAY: Maldonado ARGENTINA: Bahia Blanca ARGENTINA: Bahia Blanca ARGENTINA: Bahia Blanca URUGUAY: Maldonado URUGUAY: Maldonado URUGUAY: Maldonado URUGUAY: Maldonado URUGUAY: Maldonado URUGUAY: Maldonado URUGUAY: Maldonado 22 80 HELDIANA: ZOOLOGY Table 1 1 . Continued. Current name Waterhouse synonym or misidentification Locality Figure Caviomorpha (continued) Dolichotis patagonum Zimmer- man Vizcacia maximus Desmarest Ctenomys brasiliensis Blainville Lagostomus trichodactylus Brookes ARGENTINA: Rio Negro (ob- served) URUGUAY: Maldonado URUGUAY: Maldonado Darwin could not explain the viscacha's behavior. Pampas deer (Blastoceros bezoarticus) were abundant throughout the La Plata region. Darwin (1839, p. 55) saw very many small herds, containing from five to seven animals each, near the Sierra Ven- tana, and among the hills north of Maldo- nado. If a person crawling close along the ground, slowly advances toward a herd, the deer frequently, out of curiosity, approach to reconnoitre him. I have by this means killed, from one spot, three out of the same herd. Although so tame and inquisitive, yet when approached on horseback, they are ex- ceedingly wary. In this country nobody goes on foot, and the deer knows man as its en- emy only when he is mounted and armed with the bolas. The jaguar by some accounts is a man-killer, by others, fears man. Darwin (1839, p. 159) records several instances reported to him of man-killing jaguars of the Rio Parana region. The Beagle left the Rio Plata on December 1 833 for Puerto Deseado, or Port Desire, on the Pata- gonian coast. The mammals collected by Darwin and reported by Waterhouse (1838-1839), with descriptions and supplementary notes by Darwin, are listed in Table 1 1, with the Waterhouse syn- onyms (misidentifications included). Added are the few species Darwin mentioned in his Journal but did not collect. Unless otherwise indicated, all species are from the neighborhood of Maldonado, Uruguay. The geology and natural history of Patagonia investigated by Darwin included those of the Straits of Magellan and Tierra del Fuego (December 1 832- January 1833; May-June 1834), Puerto Deseado (Port Desire) (December 1833-January 1834), Santa Cruz (April-May 1834), and the Falkland Islands (March 1834). The Beagle itself (fig. 22) sailed up the Rio Santa Cruz to a point 1 40 miles from its mouth in the Atlantic Ocean to about 60 miles from the nearest arm of the Pacific Ocean on the opposite side of the cordillera. Darwin was greatly impressed by the number, variety, and great size of fossil mammals, mostly Pleistocene, exposed on the Patagonian plains. These, he (1839, p. 209) believed, were confir- mation of the "law" that existing animals in an area have a close relation in form with extinct species in the same area. The natural causes for extinction, however, eluded Darwin. After pro- posing and rejecting a number of explanations, the nonevolutionist Darwin (1839, p. 212) concluded that the whole series of animals, which have been created with f>eculiar kinds of organization, are confined to certain areas; and we can hardly suppose these structures are only ad- aptations to peculiarities of climate or coun- try; for otherwise, animals belonging to a distinct type, and introduced by man, would not succeed so admirably even to the exter- mination of the aborigines. On such grounds it does not seem a necessary conclusion that the extinction of species, more than their creation, should exclusively depend on the nature (altered by physical change) of their country. All that at present can be said with certainty, is that, as with the individual, so with the species, the hour of life has run its course, and is spent. The small number of extant large mammals and great number and variety of small mammals, also impressed Darwin (1839, p. 215). Patagonia, poor as she is in some resj)ects, can, however, boast of a greater stock of small rodents than p)erhaps, any other coun- try in the world. Several sjjecies of mice are externally characterized by large thin ears and a very fine fur. These little animals HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 81 swarm amongst the thickets in the valleys, where they cannot for months together taste a drop of water. They all seem to be can- nibals, for no sooner was a mouse caught in one of my traps than it was devoured by others. A small and delicately-shaped fox which is likewise very abundant, probably derives its entire support from these small animals. The guanaco was regarded as the characteristic quadruped of the Patagonian plains (Darwin, 1 839, p. 215). Herds of fifty or a hundred were common, and, as I have said, we saw one which must have contained at least five hundred. The puma with the condor in its train, follows and preys upon these animals. The footsteps of the former were to be seen almost every- where on the banks of the river [Santa Cruz]; and the remains of several guanaco, with their necks dislocated, and bones broken, showed how they had met their death. In March 1834, Darwin visited the Falkland Islands. On a tour he encountered large numbers of horses, cattle, swine, and rabbits {Oryctolagus cuniculus Linnaeus [= Lepus magellanicus Lesson and Gamot] in domestic and feral states. The an- imals had been brought by French colonists in 1 764. Darwin wrote (p. 249), The only quadruped native to the island, is a large wolf-like fox [Dusicyon (Iculpaeus) australis Kerr] which is common to both East and West Falkland. I have no doubt it is a peculiar species and confined to this archijjelago. . . . These wolves are well known . . . [for] their lameness and curiosity. ... To this day their manners remain the same. . . . As far as I am aware, there is no other in- stance in any part of the world, of so small a mass of broken land, distant from a con- tinent, possessing so large a quadruped pe- culiar to itself. Their numbers have rapidly decreased; they are already banished from that half of the island which lies to the east- ward of the neck of land between St. Sal- vador Bay and Berkeley Sound. Within a very few years after these islands shall have become regularly settled, in all probability this fox will be classed with the dodo, as an animal which has perished from the face of the earth. The mammals collected by Darwin in the Ar- gentine Patagonia (including Falkland Islands) and bordering parts of the Chilean Straits of Magellan are listed in Table 12. The Chilean leg of the cruise began in May 1 834 with the passage of the Beagle into the eastern mouth of the Straits of Magellan and ended July 1835 with departure from Copiapo in northern Chile. While the Beagle sailed up and down the Chilean coast, Darwin explored the coast, islands, archipelagos, and cordillera. He crossed the Andes on 21 March 1835 through the Portillo Pass south of Santiago, and proceeded to the town of Men- doza in Argentina. Differences observed between the biota of eastern and western versants of the cordillera impressed Darwin. The mountains, he (1839, p. 399) reasoned, have existed as a great barrier, since a period so remote that whole races of animals must subsequently have perished from the face of the earth. Therefore, unless we suppose the same species to have been created in two different countries, we ought not to expect any closer similarity between the organic beings on opposite sides of the Andes, than on shores separated by a broad strait of the sea. The correlation between geographic isolation and faunal peculiarity was noted in other circum- stances. Darwin (1839, p. 439) observed that next to lizards, mice appear to be able to support existence on the smallest and driest portions of the earth— even on islets in the midst of great oceans. I believe it will be found, that several islands, which possess no other warm-blooded quadruped, have small rodents peculiar to themselves. Ratadas or rat plagues in Chile also caught Dar- win's attention. One of the earliest recorded for Oryzomys longicaudatus longicaudatus, viewed through the eyes of Darwin (in Waterhouse, 1 838, p. 40), "overran the wooded country south of Con- cepcion, in swarms of infinite numbers." The mammals of Tierra del Fuego tallied by Darwin (1839, p. 300) included, besides cetaceans and phocids, one bat [not named but likely Histiotus montanus magellanicus Philippi], a mouse with grooved front teeth {Reithrodon of Waterhouse) and two other species, the tu- 82 HELDIANA: ZOOLOGY Table 12. Patagonian mammals collected by Darwin and recorded by Waterhouse (1838-1839); arrangement is phylogenetic. Current name Waterhouse synonym or misidentification Locality Figure Carnivora Dusicyon australis Kerr Dusicyon griseus Gray Felis colocolo pajeros Desmarest Artiodactyla Lama guanicoe Miiller RODENTIA Oryzomys longicaudatus magel- lanicus Bennett Akodon xanthorhinus Water- house Akodon canescens Waterhouse Auliscomys micropus Water- house Graomys griseojlavus Water- house Phyllotis xanthopygus Water- house Reithrodon physodes cunicu- loides Waterhouse Euneomys chinchilloides Water- house Myocastor coypus Molina Microcavia australis I. Geoffroy and d'Orbigny Dolichotis patagonum Zimmer- man Cetacea Lagenorhynchus cruciger Quoy and Gaimard Canis antarcticus Shaw Not Canis azarae Wied-Neu- wied Auchenia llama Desmarest Cavia patachonica Shaw Delphinus fitzroyi Waterhouse Falkland Islands ARGENTINA: PaUgonia ARGENTINA: Santa Cruz ARGENTINA: Patagonia ARGENTINA: Puerto de Hambre (Port Famine); CHILE: Straits of Magellan CHILE: Peninsula de Hardy ARGENTINA: Puerto Deseado (Port Desire); Santa Cruz ARGENTINA: Rio Santa Cruz, Santa Cruz ARGENTINA: Rio Negro ARGENTINA: Puerto Deseado (Port Desire); Santa Cruz ARGENTINA: Puerto Deseado (Port Desire); San Julian; Rio Santa Cruz, Santa Cruz ARGENTINA: Eastern entrance to Straits of Magellan ARGENTINA: Rio Chubut ARGENTINA: 41°S to Straits of Magellan ARGENTINA: Patagonia ARGENTINA: Golfo San Jose, 42°30'S, Chubut 22 cotuco (the greater number of these rodents are confined to the eastern and dry part), a fox, sea-otter, guanaco, and one deer [un- named but likely Hippocamelus bisulcus]. The latter animal is rare, and is not, I be- lieve, to be found south of the Straits of Magellan, as happens with the others. With respect to geographic distribution, Darwin (1839, p. 300), observing the general correspondence of the cliffs of soft sandstone, mud, and shingle, on the opposite side of the Strait, together with those on some intervening islands [was] strongly tempted to believe that the land was once joined and thus allowed animals so del- icate and helpless as the tucotuco, and Reithrodon to pass over. The tucotuco in question is Ctenomys magellan- icusfueginus Philippi (Osgood, 1943, p. 1 19). Dar- win (1839, p. 327) also mentioned the occurrence of the puma {Felis concolor) in Tierra del Fuego, and related something of its habits in other parts of Chile and Argentina. The type specimen of Darwin's zorro {Dusicyon fulvipes Martin), peculiar to the island of Chiloe, was discovered by Darwin (p. 34 1 ) on 6 December 1834 sitting on the rocks and so intently absorbed in watching the maneuvers of two ship's officers engaged in surveying, that I was able, by quietly walking up be- hind, to knock him on the head with my geological hammer. This fox, more curious or more scientific, but less wise, than the generality of his brethren, is now mounted in the museum of the Zoological Society. Sea otters {Lutrafelina Molina) were described by Darwin (in Waterhouse, 1838, p. 24) as ex- HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 83 ceedingly common amongst the innumerable channels and bays which form the Chonos Ar- chipelago. . . . they may generally be seen quietly swim- ming with their heads just out of water amidst the great entangled beds of kelp, which abounds on this coast. They burrow in the ground, within the forest, just above the rocky shore, and I was told, that they some- times roam about through the woods. This otter does not, by any means, live exclu- sively on fish. One was shot whilst running to its hole with a large volute-shell in its mouth; another (I believe the same species) was seen in Tierra del Fuego devouring a cuttle fish. But in the Chonos Archipelago perhaps the chief food of this animal, as well as that of the immense herds of great seals, and flocks of terns and cormorants, is a red- coloured crab (belonging to the family Mar- rouri) of the size of a prawn, which swims near the surface in such dense bodies, that the water appears of a red colour. This spec- imen weighed nine pounds and a half The vampire bat which Darwin (1839, p. 25) recognized as a species of d'Orbigny's genus Edos- toma (= Des modus) was singled out as often the cause of much trouble, by biting the horses on their withers. The injury is generally not so much owing to the loss of blood, as to the inflammation which the pressure of the saddle afterwards produces. The whole circumstance has lately been doubted in England; I was therefore fortun- ate in being present when one was actually caught on a horse's back. We were bivouack- ing late one evening near Coquimbo, in Chile, when my servant, noticing that one of the horses was very restive, went to see what was the matter, and fancying he could dis- tinguish something, suddenly put his hand on the beast's withers, and secured the vam- pire. In the morning, the spot, where the bite had been inflicted, was easily distinguished from being slightly swollen and bloody. The third day afterwards we rode the horse, with- out any ill effects. The Chilean mammals collected and others, only observed by Darwin, are listed in Table 13. Departing Chile on 12 July 1835, the Beagle sailed north along the Peruvian coast before turn- ing west to the Galapagos Islands. Darwin's ac- counts of the stopovers in Iquique, Callao. and Lima make no mention of indigenous mammals. The Galapagos Archipelago, it seemed to Dar- win ( 1 839. p. 454), was "a little world within itself; the greater number of its inhabitants both vege- table and animal being found nowhere else." Dar- win (p. 464) "endeavoured to make as nearly a p)erfect collection in every branch as time permit- ted" but the only land mammals he found were the Chatham Island rice rats described by Water- house as Oryzomys galapagoensis (fig. 23), and the introduced Rattus on James Island. Darwin's investigations of the Galapagos fauna, Ijarticularly the birds, lizards, tortoises, and cer- tain plants stirred old beliefs and generated new and conflicting thoughts. However, at the time he wrote his journal in October 1835, Darwin (1839, p. 474) made no attempt to come to any definite conclusions, as the sp)ecies have not [as yet] been accu- rately examined; but we may infer, that, with the exception of a few wanderers, the organic beings found on this archipelago are peculiar to it; and yet their general form strongly par- takes of an American character. . . . This similarity in type between distant islands and continents, while the sp)ecies are distinct, has scarcely been noticed. The circumstances would be explained according to the views of some authors, by saying that the creative power had acted according to the same law over a wide area. Writers on Darwin, quoting from his revised (1845) edition of the Journal, attribute to Darwin more foresight on the origin of species than is ap- parent in the first (1839) edition quoted here. At the time of its publication, two years delayed, Dar- win, still a creationist and believer in the immut- ability of species, had yet to know the identity or specific aflinities of the vast majority of the plants and animals he had collected. This knowledge served him later for definition and elaboration of thoughts expressed in the second and other revised editions of the Journal, but not in the first. The following impressions of the biota of the Galapagos Islands in the second edition (p. 372 of an 1 899 "authorized edition") and oft quoted in whole or in part by various authors, are absent in the first. The natural history of the islands is emi- nently curious and well deserved attention. Most of the organic productions are aborig- inal creations, found nowhere else; there is 84 FIELDIANA: ZOOLOGY Table 13. Chilean mammals collected or only observed by Darwin, and those identified by Waterhouse (1838- 1839); the arrangement is phylogenetic. Current name Waterhouse synonym or misidentification L4>cality Figure Marsupialia Marmosa elegans Waterhouse Chiroptera Histiotus montanus magellani- cus Philippi Myotis chiloensis Waterhouse Tadarida brasiliensis I. Geoffroy Desmodiis rotundus dorbignyi Waterhouse Carnivora Dusicyon culpaeus magellanicus Gray Dusicyon fulvipes Martin Dusicyon griseus Gray Lutra felina Molina Felis concolor Linnaeus Artiodactyla Hippocamelus bisulcus Molina RODENTIA Oryzomys longicaudatus longi- caudatus Bennett Oryzomys longicaudatus magel- lanicus Bennett Akodon olivaceus olivaceus Waterhouse Akodon olivaceus brachiotus Waterhouse Akodon xanthorhinus xantho- rhinus Waterhouse Abrothrix longipilis longipilis Waterhouse Phyllotis darwini darwini Water- house Reithrodon chinchilloides Waterhouse Abrocoma bennetti Waterhouse Spalacopus cyanus Molina Myocastor coypus Molina Octodon degus Molina Ctenomys magellanicus fueginus Philippi Not Dysopes nasutus Spix Not Canis azarae Wied- Neuwied Lutra chilensis Bennett Mus renggeri Waterhouse Abrocoma cuvieri Waterhouse Poephagomys ater Cuvier Octodon cummingii Bennett Valparaiso Tierra del Fuego (observed) Chiloe Valparaiso Coquimbo Copiapo; Straits of Magellan Chiloe 22 Copiapo; Straits of Magellan Chonos Archipelago Tierra del Fuego and central Chile to 10,000 ft elevation (observed) Tierra del Fuego (observed) Concepcion Puerto de Hambre, Straits of Magellan Valparaiso; Coquimbo Chonos; Chiloe Hardy Peninsula, Tierra del Fuego Coquimbo Coquimbo 22 Straits of Magellan Valparaiso; Aconcagua Valparaiso Chonos Archipelago Valparaiso Tierra del Fuego (observed) even a difference between the inhabitants of the different islands; yet all show a marked relationship with those of America, though separated from that continent by an open space of ocean, between 500 and 600 miles in width. The archijjelago is a little world within itself, or rather a satellite attached to America, whence it has derived a few stray colonists, and has received the general char- acter of its indigenous productions. Consid- ering the small size of these islands, we feel the more astonished at the number of their aboriginal beings, and at their confined range. Seeing every height crowned with its crater, and the boundaries of most of the lava- streams still distinct, we are led to believe that within a period, geologically recent, the unbroken ocean was here spread out. Hence, both in space and time, we seem to be brought somewhat near to that great fact— that mystery of mysteries— the first appear- ance of new beings on this earth. HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 85 "-(SI 86 HELDIANA: ZOOLOGY XIV. Georges Louis Leclerc de Buffon (1707-1788) Georges Louis Leclerc de Buffon was bom into wealth and devoted his life to scientific labors; he won recognition as the leading naturalist of his time. In 1739 he was appointed keeper of the Jar- din du Roi in Paris (now the Jardin des Plantes), which he turned into one of the most important centers of biological research during the 18th cen- tury. Buffon's lifetime work was a general natural history in 36 volumes. The first volume dealt with science in general, the second with man, the next 1 3 with nonhuman mammals ( 1 750-1 767). These were followed by nine volumes on birds, seven volumes (1789) supplementary to the preceding, and the last five on minerals, including fossils. Treatment of most species in the Histoire Na- turelle is usually monographic. Gross descriptions, including measurements and weights, are based on individuals received in the Jardin du Roi. Geo- graphic distribution of the species is included with the description. Habits observed in captivity and mentioned in the literature are recorded. Anatom- ical descriptions by Daubenton, Buffon's collab- orator, are of the skeleton, with soft parts and tegumentary structures of particular interest. Complete bibliographic references and synony- mies, including those to the 10th edition of the Linnaean Systema Natura, accompany each spe- cies account. Buffon drew together much if not everything known of a species, often an indiscriminate com- posite of species. Most of the information was compiled, some of it original. Many life history notes were received from correspondents, partic- ularly M. de la Borde, the royal physician resident in Cayenne, French Guiana. Another correspon- dent, M. Saint Lurrent of Trinidad, believed he had solved the mystery of marsupial birth (cf p. 40). At a certain stage of development, he in- formed Buffon, the embryonic op>ossum crawled from the uterus through a tube at the end of which it found a long teat to which it remained attached until fully developed. An easily verifiable discov- ery by Daubenton (in Buffon) was that tapirs have simple stomachs, not the complex ruminant type claimed by Bajon (above). Buffon reported that domestic cats kill but do not eat shrew- or short- tail opossums of the genus Monodelphis. House cats do indeed kill these animals and usually de- posit them whole in the middle of the path leading from the house to the garden. Most of the illustrations of mammals and all anatomical drawings of the Histoire Naturelle are original. A small sampling is reproduced here (figs. 24-25). Buffon was the first naturalist to recognize re- gional faunas as such and to discriminate between Old World species and different but similar ap- pearing or like-named species of the New World. He perceived the platyrrhine-catarrhine dichoto- my of primates, and the phylogenetic distance be- tween the groups. He further distinguished pre- hensile-tailed monkeys from non-prehensile-tailed species, and cebids from callitrichids by their un- gues and teeth. Buffon's sense of rivalry with the contemporary Linnaeus led him to find fault with and cast scorn on the binomial system used in the Systema Na- turae. Buffon argued for retention of vernacular names for species as well as a makeshift vernacular terminology for generic or supergeneric groups. Lack of a scientific system of nomenclature in Buffon's work, and the almost universal adoption of the Linnaean binomial system by contemporary and later authors caused the Histoire Naturelle to be regarded as no better than a layman's encyclo- pedia of science. It has been republished with many revisions in many editions and languages. It is unfortunate that Buffon's important contributions to life histories, morphology, and evolutionary bi- ology were largely ignored by Darwin and are little appreciated today. It seems that the greater luster credited to Darwin owes much to the dimming of Buffon's because of his lack of organization and consistency in his writings. XV. Faunal Origins and Distribution Early attempts to explain observed similarities and differences between Old and New World mammals all supposedly descended from occu- pants of Noah's ark, began with the 1 6th century philosopher and chronicler Acosla and in some quarters continues to this day. Jose de Acosta (1539-1600) Jose de Acosta argued that the animals of the New World had not been carried there by man. His evidence indicated that New World man brought nothing but himself over a land route. The possibility that animals migrating from the ark might have crossed the Atlantic Ocean by swim- HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 87 IIJI.IXI /W ,.♦ j^^ ,„ J"/ X-W /-.ff //y I.B 5.VOOI IV \ ri.(;.\|HK.\IKNl AvPBi.r. Msnt, i>K :«i»iT. /■//. 11// /..y i.A URAXDK riiAivr-soiRis mn- im- i.ajht h" h iu'vaxnb IK c- \i;; \i Fig. 24. Mammals figured in the Histoire Naturelle of Buffon: upper left, le saki (= Pithecia pithecia Linnaeus, male; from Buffon, 1767); upper right, le sagouin singe de nuit (= Pithecia pithecia Linnaeus, female; from Buffon, 1 789); lower left, la grande chauve-souris fer-de-lance de la Guyanne (= Phyllostomus hastatus Pallas; from Buffon, 1 789); lower right, le cabiai (= Hydrochaeris hydrochaeris Linnaeus; from Buffon, 1 764). 88 FIELDIANA: ZOOLOGY rtijii j-^ lit I'fFK l"|-K 111' fllll.l l>K. MIR IK llf, l.A <.l'^.^NNK /■/ Win ..^ f- Xm >U 11 'CXXl/.fja td> 1 n \ 1 1,/ ^ r 1 m 1i- B^^Bl 1 ^i»' Via '■ 1 , \*«.ll m )<': fwfl 1 ■^/ i '^fjS 1 '' ■'u t 1 r "*■■ ■ r 1 -,-:5V ^>^^%P vi.: ' 'utA m ^ ■^3^5^ »«yr'-^->— ^ :^; i^?5??S»«— , -ttt; IK I \1l\RIN NK«i«l L II* ossr.rsE i>K 1.A noxvi! iir. i.'Ar.ot'Arrr. Fig. 25. Mammals figured in the Histoire Naturelle of BufTon: upper left, la mouffette du Chili (= Conepatus chinga Molina); upper right, la grande marte de la Guyanne (= Eira barbara Linnaeus); lower left, le tamarin n^e (= Saguinus midas Linnaeus); lower right, hyoid apparatus and thyroid cartilage of the throat ofAlouatta seniculus Linnaeus); from Buffon (1789). HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 89 ming or island hopping was also dismissed be- cause, as Acosta pointed out, none of the animals was known to occur on oceanic islands. The leg- endary island of Atlantis, which might have been part of a former transatlantic archipelago, was treated as fable. Other conjectures discarded, Acosta resolved that New and Old World northern continents were or had been connected, or very nearly approximated, at their polar extremities. Differences between New and Old World species, he affirmed, could be explained by the disappear- ance of connecting Old World populations, mu- tations among the New World sijecies engendered by their isolation in different environments, or by degeneration. No accounting or explanations were needed for "imperfect" organisms such as rats, frogs, insects, or vermin in general. These, it was commonly held, arose spontaneously from decay- ing matter. Antonio Vazquez de Espinosa (1560/1575-1630) Antonio Vazquez de Espinosa agreed with the explanation of a northern migratory or connecting route but added (1948, chap. 36), in the awkward phrasing of Clark's translation, that near the Straits of Magellan in what is called Tierra del Fuego, which is still not well known or explored, and there are numerous other quarters where the mainland of the New World could have communicated with that of the Old, or at least have lain so close as to afford passage not merely for the peo- ples who settled the New World, but the various kinds of animals which live in them— many of species well known in Eu- rope and elsewhere, and others peculiar and unique in the New World, like the Peruvian sheep [llamas], the guanacos [regarded as the wild form of llamas], vicuiias and tarugas [Hippocamelus bisulcus]. Carolus Linnaeus (1707-1778) Linnaeus played no direct role in the develop- ment of Neotropical mammalogy apart from pro- viding scientific names for some species discov- ered or described by others. His impact on the scientific world, however, was enormous. Like the philosophers before him, he believed all species were created as they are now in one place from which they spread in search of the habitats for which they were specially created. This idea of a staging area as the center of origin and dispersal, still dominant today in the minds of some students of Neotropical mammalogy, was critically reex- amined by Buffon. Georges Louis Leclerc de Buffon (1707-1788) Buffon, on comparing faunas of New and Old Worlds, was impressed by similarities between some of the species and differences between others. His explanation for similar-apjjearing sp>ecies, like that of the chroniclers, was that a land connection permitted passage of animals from Old to New World. Differences between New and Old World species, he suggested, in agreement with the chron- icler Jose Acosta, could have resulted from de- generation, environmental pressures, or isolation of the New World derivatives. On the other hand, Buffon argued, species peculiar to the New World or without Old World analogs must have arisen in situ, an opinion already intimated by Vazquez de Espinosa. Buffon was the first naturalist to envision the mammals of the region as a community or fauna that might well have originated indeF>endently of other regional faunas. Noah's ark had no place in his concept of faunal origins, and he rejected as too short the scripturally based 6,000-year esti- mate of the earth's age. Buffon's ideas of organic evolution and multiple centers of origin were nov- el and prepared the minds of his and succeeding generations for the acceptance of Darwinian evo- lution. Linnaeus, the arch exponent of the fixity of species and their origin and dispersal from a single center, conceived the elements of his binomial sys- tem as symbols for nailing down his credo. The system was so good it proved to be the best yet devised for the expression of genetic relationships between species and the surest base for the con- struction of evolutionary sequences in nominate terms. On the other hand, Buffon, independent of the religious constraints of his time and evolu- tionist in thought if not always in words, never attained the stature of his contemporary for lack of a competing system, key, code, or standard that would bring cohesion to his rambling philoso- phies. 90 FIELDIANA: ZOOLOGY Johann Andreas Wagner (1797-1861) Johann Andreas Wagner, the foremost masto- zoologist of his generation and author of a mono- graph on the geographic distribution of mammals, summarized (1844, p. 13) the three current but disparate opinions on mammalian origin and dis- persal. First, all species were created in one and the same region and spread from there to all cor- ners of the earth. Second, the species could have been created in separate localities in the same or different regions. Finally, each species could have arisen spontaneously anywhere and developed ac- cording to its peculiar constraints. Zoogeographers of the early half of the 19th cen- tury divided the world into major faunal regions correlated primarily with climate. Wagner (1844) separated the earth into four provinces: the Nord- liche north of 30°N, the Mittlere between BCN and 30°S, excluding the Australische roughly be- tween 0°S-55°S and 1 30°W-200''W, and the South American Magellanische, south of 30°S. The South American portion of the pantropical Mittlere Province extended from Mexico southward to southern Brazil and central Chile. Wagner's de- scriptions of the provincial faunas included tab- ulations of their resfjective genera and included species. Maximilian Prinz von Wied-Neuwied (1782-1867) Scriptural constraints were not evident in the thinking of the field naturalists. Maximilian Prinz von Wied-Neuwied recognized the limitations of geographic range as a property of a species. Johann Jacob von Tschudi (1818-1889) Tschudi attempted to follow Wied-Neuwied in defining specific ranges, but nearly all were based on the presumption that the geographic range of the species coincided with ecological life zones. The ecological life zones of Peru described by von Tschudi on the basis of fauna, flora, and climate, are the first of their kind for any Neotropical re- gion. Charles Robert I>arwin (1809-1882) The young Darwin also recognized geographic limitations of distribution in the light of physical barriers such as mountains and large bodies of water. XVI. Inventories to Middle of 19th Century Systema Naturm of Linnaeus, 1758, 1766 The 1 0th edition of the Systema Naturce pub- lished in 1758 by the Swedish naturalist Carolus Linnaeus ( 1 707-1 778), marks the beginning of the consistent application of his binomial system of zoological nomenclature. According to the uni- versally accepted International Code of Zoological Nomenclature, names for animals published be- fore 1 758 are not available, no matter how clearly defined the species. Likewise, zoological names for species published after 1757 that are not binomial or do not satisfy all provisions of the Code are not available. The effect of the Code in practice is that species without Linnaean names are treated as un- known to science. The 1 0th edition of the Systema Naturce lists a total of 1 72 species of mammals, exclusive of ma- rine cetaceans, each with its binomen consisting of a defined generic and defined specific name. Subsequent revisions of the bases for the names revealed that some represented more than a single species, others were duplicates or synonyms, and a few were equivocal or belonged to unidentifiable animals. The revisions, however, made no signif- icant change in the total number of real mam- malian species known to Linnaeus in 1758. The 1 2th and last revised edition of the Systema Naturae by Linnaeus himself, published in 1766, lists a world total of 208 mammalian species. Ta- ble 14 compares the relative numbers of world. Neotropical, and Nearctic genera and species in the Linnaean 10th and 12th editions of the Sys- tema Naturce with the totals in Buffbn's Histoire Naturelle. Cetaceans are omitted because they are oceanic species known before the discovery of America. Primary sources for the definition and naming of the Linnaean New World species were speci- mens preserved in the Swedish museums, partic- ularly the Adolphi Friderici Regis Museum, and primary bibliographic references. Such references for the Neotropical mammals were the works of Marcgraf (1648), Anson (1748), Browne (1756), and Seba (1 734-1 765). For both Neotropical and HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 91 Table 14. Number of world and New World mammals known to Linnaeus (1758, 1766) and to Buffon (1750- 1 789) and their percentage of world total. Author Date World total Neotropica Nearctica Linnaeus 1758 35 genera 18(51%) 15(43%) 1 72 species 45 (26%) 23(13%) Linnaeus 1766 36 genera 20 (55%) 15(42%) 208 species 55 (26%) 28(13%) BulTon 1750-1789 251 species 78(31%) 47(19%) Nearctic species, Linnaeus cited Hernandez (1651) and Edwards ( 1 743- 1751), and for Nearctic species alone, Catesby (1731) and Kalm (1753). Histoire Naturelle of Buffon, 1750-1789 In volume 9 of his Histoire Naturelle, published in 1 76 1 , Buffon estimated a world total of ap- proximately 200 known mammalian species of which 1 30 were Old World, 70 New World. When the sp)ecies of all 1 3 volumes on mammals and the supplementary volumes are counted, the total is 251, of which 78 are Neotropical, 47 Nearctic. The greater number of species recognized by Buffon as compared with those of Linnaeus re- flects his better knowledge of mammals and wider use of the available literature. Neither authority searched the works of the chroniclers for descrip- tions or figures of New World mammals. Synopsis Mammalium of Schinz, 1844 The Schinz catalogue of Recent mammals of the world, published 1844, brings the inventory of mammalian species to near the cutoff date of this part of the history of Neotropical mammalogy. The totals indicate that about 50% of all New World mammalian species now known had been de- scribed. The vast majority of the remaining 50% described since the middle of the 19th century are as small as or smaller than common tree squirrels. With respect to Neotropical mammals, by mid- 1 9th century about 90% of known sp)ecies larger than common tree squirrels had been described. In contrast, no more than about 1 0% of the smaller forms, mainly marsupials, bats, and rodents, had been named. The Schinz catalogue is summarized in Table 1 5. A first glance at the figures of the first order, the Marsupialia, may suggest the list is skewed. Only one marsupial species, Didelphis virginiana, is known to occur in Nearctica. Schinz recorded three because the species had been ovemamed at the time. It is believed, however, that "under- named" or composite species, as well as over- named identical species, are more or less evenly distributed in all three columns. As a result, the bottom line totals, particularly the percentages, are fair estimates of the real number of species known to science at the time of Schinz's compilation. The percentages have not changed significantly since. XVII. Summary Knowledge of Neotropical mammals from 1492 to the mid- 17th century was mainly an ag- gregation of anecdotes often riddled with myth, folklore, and untested generalizations. Eurojjeans identified New World species with similar-ap- pearing or similar-behaving Old World species and used the same names for them. Descriptions of mammals were usually comparisons with familiar European animals; measurements, rarely given, were rough estimates. Habitat when mentioned was usually on the order of "forest," "plain," or "river." Descriptions of animals often included use as food or pets, medicinal merits, or value of rawhide or bones in the manufacture of artifacts. Habits were usually described in terms of reactions to man when hunted or in captivity, or as harmful or benign to his person or property. Mammals— the term had not yet been concocted— were the hairy beasts of the earth. Whales and manatees were fish and could be eaten on Fridays. Bats were something else, mostly vampires; mice and other small mammals were vermin, in a class with frogs and cockroaches. Mammal collecting during this period was gen- erally limited to capture of live individuals for domestication, as pets, or for exhibition in zoos, circuses, or fairs. Dead animals were sometimes skinned and stuffed or bottled in brandy. The crudely prepared or pickled specimens, if not live 92 HELDIANA: ZOOLOGY animals, often served as models for the woodcut drawings of early treatises on natural history. Some specimens were purchased for museums or cabi- nets of collectors, including those of Linnaeus, King Frederick Adolph of Sweden, Reaumur of Paris, the King of France (Jardin du Roi), or the shelves of the Dutch pharmacist, Albert Seba. Most of the Neotropical specimens probably originated in the South American possessions of Holland and France. The few crude attempts at classification of mam- mals during the 16th and 17th centuries were hard- ly more than random arrangements equivalent to shopping lists. Species, being individually created kinds, were unrelated to other created kinds, or simply arose spontaneously from putrefying mat- ter. The scientific study of mammals, or mammal- ogy, of the Neotropical Region began with the ex- plorations of northeastern Brazil by Georg Marc- graf and culminated with the publication in 1648 of his Historic Rerum Naturalium Brasilia. His accounts of the included 32 sp)ecies of mammals reveal the glimmer of an attempt at natural group- ings of kinds or the beginnings of a classification of Neotropical mammals. Insofar as is known, none of MarcgraPs animals were preserved. Lin- naean names for the species of the Historice were based on bibliographic references to their descrip- tions and figures (cf. fig. 2, table 1). The first expedition to the Neotropical Region actually committed to the collection and perma- nent preservation of mammals (and other objects) for scientific study was the Brazilian Viagem Fi- losofica, 1783 to 1792, conceived by the Portu- guese government and conducted by the Brazilian- bom naturalist Alexandre Rodrigues Ferreira. The large number of specimens gathered by the ex- pedition was deposited in Lisbon's Museu d'A- juda. The specimens of monkeys that had been carried away to the Paris Natural History Museum were studied by the French scientist Etienne Geof- froy St.-Hilaire. His descriptions were published without reference to source of material. Alexander von Humboldt followed on the heels of the Viagem Filosofica with his explorations of northwestern South America from 1799 through 1802. His expedition was highly successful and in scope has rarely been equaled by other "one-man" surveys of a large portion of the South American continent. The personal narrative of his travels inspired successive naturalist-travelers, most no- tably the explorers of Brazil, Spix and Martius, Maximilian Wied-Neuwied, and Johann Natterer. Table 1 5. Number of world, Neolropic, and Nearc- tic species (subspecies) of mammals known to Schinz ( 1 844); species common to both regions are included in both. Order World Neotropica Nearctica Marsupialia 138 31(22%) 4(3%) Insectivora 114 2(2%) 21 (18%) Chiroptera 326 110(34%) 21 (6%) Primates 281 73 (26%) 0(0%) Edentata 31 24 (77%) 0(0%) Camivora 303 58(19%) 41 (13%) Pinnipedia 39 1 1 (28%) 2(5%) Sirenia 5 2 (40%) 1 (20%) Perissodactyla 23 2(9%) 0(0%) Artiodactyla 186 1 1 (6%) 12(6%) Lagomorpha 52 4(8%) 14(27%) Rodentia 563 152(27%) 104(18%) Cetacea 4 2 (50%) 0(0%) Proboscidea 2 0(0%) 0(0%) Hyracoidea 5 0(0%) 0(0%) Totals 2,072* 482 (23%) 220(11%) * The estimated number of species in 1972 (Hersh- kovitz, p. 332, table 3) is Neotropica 810, Nearctica 442 or an approximate doubling since 1 844 in both regions, with a slightly greater increase (less than 2%) in Nearctica relative to Neotropica. Increase since then has been al- most exclusively Neotropical. Later there was von Tschudi, who traveled in Peru; d'Orbigny, who journeyed in Patagonia but did his finest and most lasting work in Bolivia on a scale almost equal to that of Humboldt's; and Darwin, who voyaged around the southern half of South America and the Galapagos Islands in H.M.S. Beagle. The brothers Schomburgk, motivated by Hum- boldt's trip up the Rio Orinoco to its connection via the Casiquiare Canal with the Rio Negro trib- utary of the Rio Amazonas, completed the trajec- tory by plotting the course of the upper Rio Negro to its connection with the Casiquiare. Their ex- plorations of the British Guianas and bordering parts of Brazil and Venezuela yielded the first large collection of Guianan mammals, all depos- ited in the Berlin Natural History Museum. Chilean mammals became fairly well known through the reports of Molina (1782), Poeppig (1836), and Gay (1847). The mammals of Para- guay, their distribution, habits, or biology in gen- eral, became better known through the efforts of Felix de Azara than those of other Neotropical countries. The 200-year period from MarcgraPs ( 1 648) re- port to the last of those of Schomburgk (1 848) was one of survey and inventory of South American mammals. The published reports and personal HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 93 narratives of travel provided much reliable data on geographic distribution, habitat, life histories, ecological backgrounds, itineraries and maps of the expeditionary routes, and stopping and col- lecting localities. Descriptions of the collected mammals, most of them by the naturalist-trav- elers themselves, were often based on skeletal, dental, and soft parts in addition to purely tegu- mentary characters. Their classifications were pu- tatively natural groupings on the ordinal, family, and, as a rule, the generic levels. The prevailing belief in the biblical version of creation and fixity of species, not confessed in writing, did not blind systematists of the period to evident relationships between species and their clusterings into supra- specific groups. Descriptions of species were, nevertheless, typological. Subspecies or geograph- ic races were, at best, vaguely conceived but de- scribed as species. The infrequent or rare use of trinomials was accidental or equivocal and not certainly intended for a clearly defined geographic race. The term usually used for deviates firom "types" was "variety." Controversies regarding origin of species or fau- nas centered on where, not how. Philosopher- chroniclers of the first era accepted Noah's ark literally as the one place of origin and disjiersal of the Recent fauna. Acosta may have been the first to suggest the former existence of intercontinental connections for passage of Old World animals into the New World. More and better knowledge of the world's fauna during the second era revealed the weaknesses or fallacy of the ark dogma. Staunch creationists such as Linnaeus pointed instead to a vaguely located region as the place from which all species dispersed to occupy predestined habitats for which they had been created. Other authorities like Buffon argued for multiple centers of origin, with sp)ecies origi- nating in the habitats for which they were adapted. Darwin also believed in multiple places of origin, or faunal regions separated by geographic barriers but with some trepidation. The belief in multiple creations was heretical. Inconsistencies between religious dogma and realities did not prevent Wied-Neuwied from rec- ognizing the geographic range of a species (or sub- species) as a property of that species. Another advance beyond scriptures was the concept of eco- logical life zones contributed by von Tschudi, who plotted them for Peru on the basis of plants, an- imals, and climate. The total of named Neotropical sjjecies of mam- mals counted ft-om 1758, the year of publication of the 10th edition of the Linnaean Systema Na- tura and starting date of zoological nomenclature, to mid- 1 9th century, exceeded by far that of the Nearctic region and any other equivalent area of the world. Neotropical mammals were also better known than those of other continents except west- em Europe. By mid- 1 9th century, about 90% of currently known Neotropical mammalian species larger than common tree squirrels had already been described, but no more than about 10% of the smaller forms. The great number and variety of Neotropical mammals (and animals generally) known to sci- ence by mid- 19th century and the accumulated knowledge gained from study of living and pre- served specimens in field and laboratory, much of it contributed by Charles Darwin, helped pscvt the way to the Darwinian revolution of the next half century. XYIII. Acknowledgments I am indebted to Benjamin W. Williams, As- sociate Librarian and Librarian of Rare Books, Field Museum of Natural History, for p)ermission to consult at pleasure in the Museum's Mary W. Runnells Rare Book Room the books needed for writing this article; and to Bruce D. Patterson, Robert M. Timm, Ronald H. Pine, Debra Mos- kovitz, and J. A. Gagliano for reviewing the manu- script. Map>s shown in Figures 11, 12, and 2 1 were prepared by the author with assistance of Mary Anne Rogers from accounts of the travelers cited and other sources. Photographic reproductions of the figures are by Field Museum of Natural His- tory Staff Photographer Ron Testa. Technical Assistants Barbara Brown and Mary Anne Rogers typed the manuscript and contributed in other ways toward its completion. XIX. Literature Cited AcosTA, JosE DE- 1590. Historia natural y moral de las Indias en que se tratan las cosas notables del cielo, elementos, metales, plantas y animales dellas y los ritos y ceremonias, leyes y gobiemo y guerras de los Indios. Juan de Leon, Seville, 535 + (36) pp. [not seen]. . 1894. Historia natural y moral de las Indias. Escrita por el P. Joseph de Acosta de la Compaiiia de Jesus. Publicada en Sevilla en 1 590 y ahora fielmente 94 nELDL\NA: ZOOLOGY reimpresa de la primera edicion. Ramon Angles, Ma- drid, 2 vols. Anghiera, see Martyr. Anson, George. 1 748. A Voyage Round the World in the Years MDCCXL, I, II, III, IV by George Anson . . . Esq. Compiled and published by Richard Walter. J. & P. Knapton, London. AviLA Pikes, Fernando Dias de. 1 965. The type spec- imens of Brazilian mammals collected by Prince Maximilian zu Wied. American Museum Novitates, no. 2209: 1-21. . 1 974. CaracterizafSo zoogeografica da Provin- cia Amazonica. I. Expedi^oes cientificas na Amazonia Brasiliera. Anales Academia Brasiliera de Ciencias, 46(1): 133-158. AzARA, Felix de. 1801. Essais sur I'histoire naturelle des quadrupedes de la Province du Paraguay. Trans- lated from the original Spanish by M. L. E. Moreau- Saint-Mery. 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Archiv fiir Naturgeschichte, 9(1): 365- 368. 98 HELDIANA: ZOOLOGY A New Superfamily in the Extensive Radiation of South American Paleogene Marsupials Rosendo Pascual and Alfredo A. Carlini ABSTRACTS Significant new mammals have been recovered from the Colhuehuapian mammal-bearing beds (latest Oligocene) exposed in the Gaiman region of Patagonia (Chubut Province, Argentina). Some fragmentary mandibles and isolated teeth belong to a new genus and species, Patagonia peregrina. The mandibular and dental specializations of this form are so distinctly convergent on those of some fossorial rodents that it is regarded as a distinct clade of South American marsupials. It represents the type of a new family, Patagoniidae, which is assigned to a new superfamily, Patagonioidea, which represents a natural evolutionary group in the same sense as other superfamilies of marsupials recognized by Simpson. Its systematic position within the superorder Marsupialia awaits comprehensive analysis of those enigmatic marsupials (Groe- berioidea and Argyrolagoidea) apparently most closely related to it. Varios nuevos y significativos mamiferos han sido recogidos de capas mamaliferas del Col- huehuapense (Oligoceno tardio) expuestas en la region de Gaiman, Patagonia (Chubut, Argen- tina). Algunos fragmentos mandibulares y dientes aislados pertenecen a un nuevo genero y especie, Patagonia peregrina. Esta forma presenta especializaciones mandibulares y dentarias tan distintamente convergentes hacia las de albunos roedores cavadores que es considerada como un distinto clado de marsupiales sudamericanos. Representa el tipo de una nueva Familia, Patagoniidae, que es asignada a la nueva Superfamilia Patagonioidea, porque representa un grupo evolutivo natural como los de otras Superfamilias de marsupiales reconocidas por Simp- son. Su posicion sistematica dentro del Superorden Marsupialia depende del analisis integrado de aquellos marsupiales enigmaticos (Groeberioidea y Argyrolagoidea) aparentemente mas estrechamente relacionados a el. Novos mamiferos foram recuperados dos leitos de Colhuehuapian (do alto Oligoceno), ex- postos na regiao de Gaiman, Patagonia (Provincia de Chubut, Argentina). Fragmentos man- dibulares e dentes isolados pertencem a um novo genero e especie, Patagonia peregrina. As especializa9oes mandibulares e dentais encontradas sao tao claramente convergentes as de alguns roedores fossorios, que esta forma e considerada uma classe distinta de marsupiais sulameri- canos. A especie representa o tipo de uma nova familia, Patagoniidae, a qual e designada a uma nova superfamilia, Patagonioidea, por formar um grupo evolutivo bem definido, como o formam as outras familias de marsupiais, reconhecidas por Simpson. A posi9ao sistematica dos Patagonioidea, dentro da superordem Marsupialia, aguarda uma analise mais compreensiva dos marsupiais ainda enigmaticos (como Groeberioidea e Argyrolagoidea) aparentemente e seus relativos mais proximos. From the Division Paleontologia Vertebrados, Museo de La Plata, Paseo del Bosque, 1900 La Plata, Argentina; and CONICET, Argentina. PASCUAL & CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS 99 Introduction The taxon described in this paper is yet another example of the great adaptive radiation and dis- persal of marsupials in South America. It repre- sents a second line of marsupials that is convergent on the rodent adaptive zone (cf Groeberioidea— Patterson, 1952; Simpson, 1970c; Clemens & Marshall, 1976). However, it is distinct from pre- viously named forms, not only phylogenetically but also ecologically. The new form does not suggest that marsupials attained the breadth and diversity of rodent ad- aptations, but it does show that marsupials oc- cupied the rodent adaptive zone in previously un- imagined ways. This new marsupial indicates that marsupial radiations in South America were al- most as broad and reached as great extremes as those in Australia. The find is consistent with the view that "A complete record of South American marsupials would certainly include a large number of taxa, probably some of high categorical rank, now unknown" (Simpson, 1970a, p. 59). This and other forms recently found in northwestern Ar- gentina (Pascual, 1980a, b, 1981, 1983) validate Simpson's prophetic suggestion that ". . . major parts of marsupial evolution were occurring in areas and facies inadequately sampled, if at all, by the known fossil deposits and the collections so far made" (Simpson, 1970a, p. 58). These deposits indicate the value of applying new sample-col- lecting techniques at mammal-bearing localities that are supposedly well known; it is only neces- sary to find new, appropriate facies. The new ecological type from the Paleogene pro- vides evidence to support Gould's (1983) view of "early experimentation, later standardization," with a consequent reduction in diversity. As in therians (Pascual et al., 1985) the diversification of South American marsupials took place princi- pally in the Paleogene. Measurements reported in Table 1 are depicted in Figure 3 and are given in millimeters. The ab- breviation MACN CH is used for the Museo Ar- gentino de Ciencias Naturales "Bernardino Ri- vadavia" (Buenos Aires), Coleccion Chubut. Classification Superfamily PATAGONIOIDEA nov. The only known family of this taxon is the Pat- agoniidae. The superfamily is sufficiently charac- terized by the diagnoses of that family and its only known species. Justification for its superfamilial rank is given in a later section on affinities. Family Patagoniidae nov. Type— Patagonia gen. nov. The only known ge- nus. Known Distribution— Late Oligocene. Col- huehuapian from Central Patagonia (Chubut Province, Argentina). Diagnosis.— Small marsupials with the same reduced number of lower teeth as the Groeberi- idae, but with a different dental formula: 1.1.0.3. Open-rooted and rodent-like lower incisor, oval in cross section, strongly curved, although not as much as in the Groeberiidae, and with the intra- alveolar portion differently arranged. The incisor extends lingually along the ventral border of the mandible to the root of the inflected crest beneath the last molar, where it forms a prominence sim- ilar to that of hystricognathous rodents, but ven- trally. Lower canine smaller, procumbent, appar- ently incisor-like and closed-rooted, separated from the cheekteeth by a short, crested diastema at al- veolar level. Lower cheekteeth rectangular in cross section, decreasing in size posteriorly, hypselo- dont, rootless, wholly surrounded by enamel, and slightly curved, with the concavity forward. Hor- izontal ramus of the mandible short and deep, with the highest part posterior, beneath the masseteric fossa, where the body of the mandible becomes strongly convex and inflected; deep pterygoid fos- sa, limited ventrally by a flange like that found in Argyrolagidae and in some Australian marsupials (e.g., Macropodidae); strong, salient coronoid pro- cess; masseteric fossa relatively deep but reduced, dorsally situated with a prominent masseteric crest; subvertical symphysis unfused, with nearly smooth symphyseal surfaces. PATAGONIA gen. nov. Etymology- From Patagonia, its geographical record. Type— Patagonia peregrina sp. nov. Known Range and Diagnosis— Same as that of the family. Patagonia peregrina sp. nov. Figures 1-3 Etymology— From Latin peregr inns, strange or rare. 100 FIELDIANA: ZOOLOGY «f *^ « Fig. 1 . Patagonia peregrina gen. et sp. nov. A-B, Stereopairs of MACN CH-865, a fragment of a right mandibular ramus with m,.,: A, occlusal view; B, posterior view; C-D, X-ray of fragments of two right mandibular rami with i,, alveolus of c,, and m,., complete (C, holotype; MACN CH-869) and with alveoli of i,, and c,, and m,., complete (D, MACN CH-865). Graphic scale = 2 mm. HoLOTYPE-MACN CH-869 (fig. 2A-B). Frag- ment of right mandibular ramus with three cheek- teeth, intra-alveolar portion of the incisor, and alveolus of the canine. Hypodigm — Holotype and the following: MACN CH-864, part of right mandibular ramus with first and second cheekteeth, part of alveolus of the third, and part of alveoli of incisor and canine; MACN CH-865, part of right ramus with three cheekteeth and alveoli of the incisor and PASCUAL & CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS 101 B 'C, alveolus alveolus m 1 m 2 m alveolus Fig. 2. Patagonia peregrina gen. et sp. nov. A-B, Stereopairs of MACN CH-869 (holotype), a fragment of a right mandibular ramus, with i,, alveolus of c,, and m,.,: A, labial view; B, lingual view. Graphic scale = 2 mm. C, Stereopairs of MACN CH-867, a fragment of a left mandibular ramus, with alveoh of i,, c, and mj, and m,_2; occlusal view. canine; MACN CH-866, part of left ramus with the second and third cheekteeth, and part of al- veoli of the first cheektooth and the incisor, MACN CH-867, part of the left ramus with the first and second cheekteeth, and part of alveoli of the third cheektooth, incisor, and canine; MACN CH-868, part of the right ramus with three cheekteeth and alveolus of the incisor; MACN CH-870, part of left ramus with first and second cheekteeth and part of the alveolus of the third; MACN CH-874, part of right ramus with the second cheektooth, alveoli of the first and third cheekteeth, and part of the alveolus of the incisor; MACN CH-875, part of right ramus with the second and third 102 HELDIANA: ZOOLOGY cheekteeth and part of the alveolus of the incisor; and MACN CH-876, three isolated upper(?) cheekteeth. Horizon and Locality— Both the holotype and the hypodigm come from the Trelew Member of the Sarmiento Formation (see Mendia & Bayar- sky, 1981) and are Colhuehuapian (Late Oligo- cene) in age. Apparently they were found in the upper unit, exposed on the south side of the Chu- but River valley, Chubut Province, Argentina (Central Patagonia; see Fleagle & Bown, 1983, pp. 242-244). Quite probably this corresponds to Simpson's "stratum F of Fig. 1," which is part of his "Trelew beds" (= "Trelewense"). The material was recovered by O. E. Donadio, M. Soria, J. G. Fleagle, and T. M. Bown (see Fleagle & Bown, 1 983) through dry-screening local deflation lag de- posits. Diagnosis— The only known species of the fam- ily. Description— Dentition— See Figures 1 A,C-D; 2-3. Each side of the lower jaw has one fully ro- dent-like gnawing incisor, only incompletely pre- served in the holotype; it is posteriorly bordered by a relatively shallow and conical alveolus (the tooth being absent in all specimens at hand) sep- arated from the medial one by bone and set at a relatively oblique angle (figs. IC-D; 2C). Homol- ogies of these teeth are uncertain, but the rodent- like medial tooth is surely an incisor, designated for description as i,. The shape and disposition of the second alveolus agrees with the procumbent canine of Polydolopidae (Epidolopinae; cf Paula Couto, 1952, 1961; Pascual & Bond, 1981) and Prepidolopidae(Pascual, 1980b, fig. 2D-E); it thus appears that this tooth is c,. This alveolus is fol- lowed by a short diastema at alveolar level, then three cheekteeth, all rectangular in cross section (with some differences among them) and in close approximation, forming a molariform series. They are surrounded by enamel on all sides and are not strictly lobate, nor are the trigonid, talonid, or original cusps clearly indicated, as occlusion with the uppers was mediated through practically flat areas. The dentine forms a shallow basin sur- rounded by the highest enamel layer, which is slightly higher on the lingual side. There is a slight- ly deeper anteroposterior wear groove, extending from the anterolabial comer to the posterolingual one (fig. lA). Grinding involved a longer propal- inal movement and a shorter ectental stroke. The homologies of these teeth with the more numerous ancestral series cannot be determined. Plausibly B /r- ^[ii^dEJE).--]^ Fig. 3. Patagonia peregrina gen. et sp. nov. Outline of a right mandibular ramus fragment, with alveoli of i, and c,, and m,., complete (MACN CH-865), showing the measurements of Table 1 . A, Labial view; B, occlusal view; C, cheekteeth series (m ,.3); D, lingual view. Graph- ic scale = 2 mm. they are homologous with those typically desig- nated m,_3 in marsupials and are so designated here, yielding the lower dental formula 1.1.0.3. which is provisionally homologized as i,, c,, m,.,. However, many specialized marsupials from the South American fossil record show tendencies (1) PASCUAL &. CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS 103 Table 1 . Dimensions of specimens of Patagonia peregrina gen. et sp. nov. (see fig. 3 for measurement references). Dimensions Specimen MACN CH-864 MACN CH-865 MACN CH-866 MACN CH-867 MACN CH-868 MACN CH-869 MACN CH-870 MACN CH-874 MACN CH-875 3.68 4.08 1.40 1.12 4.08 3.60 1.00 1.00 0.96 1.40 1.20 1.08 1.08 1.12 0.96 5.40 1.00 1.08 1.00 0.92 1.20 1.00 0.96 0.96 1.00 1.00 1.04 1.00 0.92 1.36 1.08 1.00 0.96 0.96 0.92 5.12 1.40 1.00 1.08 1.00 1.00 0.96 4.80 1.28 1.16 1.24 1.08 6.76 2.80 3.00 4.60 2.20 4.40 6.20 1.80 1.60 1.80 2.40 to elongate or modify either the pj (Polydolopidae and Parabderitini caenolestids; see Marshall, 1980) or the m, (Abderitinae and Palaeothentinae caen- olestids), and (2) to reduce (e.g., Caenolestidae) or lose (Polydolopidae) the m4. All teeth except the c, are completely hypselodont and rootless. The cheekteeth are slightly curved, with the concavity forward (fig. IC-D). Incisor ("/J— Incompletely preserved in the ho- lotype (fig. 2A-B). The anterior end is broken, but the posterior end is unaltered, showing an open pulp cavity with no sign of root formation. It is as elongate and curved as in the most specialized caviomorphs (e.g., Ctenomyidae), although not upcurved posteriorly. The extra-alveolar part would have been nearly vertical, although not so much as in the Groeberiidae (cf. fig. 4B,D). The tooth extends along the ventral border lingually, first beneath the m, and then lingually to far below the mj, terminating where the ventral border be- comes an inflected flange (MACN CH-865, fig. 1 B) reminiscent of kangaroos; the base of the alveolus shapes a superficial prominence similar to that of hystricognathous rodents, although enveloped by the inflected ventral border. It is approximately oval in cross section, with the long axis oriented dorsoventrally and with a flatter medial surface. Apparently enamel covers the entire tooth, but a noticeably heavier layer extends as a ventral band. Canine fie J— None of the specimens in the hy- podigm include this tooth, but its alveolus (figs. 1 C-D; 2C) is oblique, tapered, and relatively shal- low, indicating a closed-rooted tooth; its oblique orientation suggests that the occlusal apex was ap- pressed against the occlusal tip of i,. First Molar (m,)—The first molar is separated from the c, by a crested diastema (fig. 1 A) as long as m,. It is the largest cheektooth, almost rect- angular in cross section, with the longer lateral faces slightly concave; the lingual face sometimes bears a very shallow groove along the intra-alveo- lar portion. The anterior face is convex, occasion- ally somewhat pointed; the posterior face is almost flat, forming angles with the lateral faces slightly greater than 90°. Second Molar (^wj— The second molar is irreg- ularly quadrate, with the lateral faces slightly con- vex and the anterior and posterior ones flatter. The posterolingual angle is less than 90°, whereas the others are almost 90°. Its width is similar to that of m, (fig. lA). Third Molar (m^J—The third molar is the small- est cheektooth, being subtriangular in cross section rather than square. The anterior face is slightly convex, lingually flatter, and labially more strongly curved; the labial face converges posterolingually with the lingual face, forming a rounded pillar rather than a well-defined posterior face (fig. lA). Mandible— No nearly complete mandible is known, but parts of the horizontal ramus and base of the coronoid process are known. These parts indicate the mandible is extremely short and deep, like that of Groeberia minoprioi, although very different in other respects (cf. fig. 4B,D). The sym- physis is subvertical and unfused, with a nearly smooth symphyseal surface (i.e., normal in struc- ture instead of fused and forming the odd medial posterior projection peculiar to Groeberiidae). The depth of the mandible increases abruptly toward the mj. The deep masseteric fossa appears to be F)eculiarly confined to a dorsal position, as the masseteric crest is situated at a level between the alveolar rim and the lowest level of the rounded and inflected ventral border (figs. 1 B; 2 A). A sim- ilar condition is found in some Abderitinae caen- olestids (e.g., Parabderites bicrispatus; Marshall, 1976, fig. 8a), although in P. bicrispatus the man- dibular body is not as deep and the alveolar border 104 FIELDIANA: ZOOLOGY not as extensively inflected. The coronoid process has its root beneath the m,, forming a strong, sa- lient lamina (known only by its root), so that a conspicuous diagonal valley is formed between the coronoid and the alveolar border behind m, (fig. lA); a similar structure is present in Groeberia minoprioi (see Patterson, 1952, p. 41); the valley is open labially and lingually limited by a prom- inence similar to that present in Australasian Po- toroinae. In many respects this strong, salient, ascending ramus and correlated features are reminiscent of highly fossorial caviomorphs, such as burrow-in- habiting Ctenomyidae. Although the mandibular angle is not preserved in any of the specimens, it probably was inflected, as suggested by the inflec- tion of the ventral border, beginning at the level of mj, which defines a lingual flanged crest (figs. IB; 2B) similar to that producing the extremely inflected angle in the Macropodidae. This lingual ventral flanged crest seems to be the lingual border of an expanded and relatively deep pterygoid fos- sa, resembling that of argyrolagids (see Simpson, 1 970a). There is a relatively large alveolar foramen within the pterygoid fossa, level with the alveolar border and within a pit (fig. 3D), and a mental foramen beneath the anterior face of m, at the level of the alveolus of i, (fig. 2 A). Affinities As in the case of Groeberia (see Simpson, 1 970c), the conclusion that Patagonia is a marsupial rests on a combination of definite, negative, and indi- rect evidence. The most definite evidence for its being a marsupial is the inflected ventral border of the mandible and probably the related inflected angle. This evidence alone is inconclusive, as a few marsupials lack an inflected angle and a few placentals have one. However, no known placental has such an extended and upturned flange-shaped inflection, and even in marsupials it is rarely so well developed (e.g., Groeberiidae [Patterson, 1952]. Argyrolagidae [Simpson, 1970a,b], and the Australasian Macropodidae). Unlike Groeberia, Patagonia has other characters supporting its mar- supial affinities, namely the lower procumbent in- cisor-like canine. In the Epidolopinae (Pascual &. Bond, 1981) there are three procumbent lower teeth, the third being unquestionably the canine. Within the more advanced Polydolopidae (Poly- dolopinae), there are one or three procumbent lower teeth; in the latter case, evidence suggests they consist of two incisors and a canine, the me- dial incisor being quite reduced and the canine well developed, single, and closed-rooted. As in Groeberia the negative evidence is that Patagonia has no features precluding its reference to the Marsupialia. It does exhibit characters mak- ing reference to any Eutheria highly improbable. Its habitus is rodent-like, but its two differentially procumbent lower teeth rule out reference to the Rodentia. While the incisor is rodent-like in shape, it is oriented differently than that in rodents, ex- tending along the ventral border of the horizontal ramus, first below the m,, then lingually to other molars, without curving upward. It apparently shapes the ventral border of the mandible. In ad- dition, the short diastema extends at the level of alveoli. Among known rodents, only Paramyidae and Ischyromyidae developed diastemas at the al- veolar level, but even in these groups, the incisor extends as in other rodents, not as in Patagonia. A more-or-less rodent-like habitus was also char- acteristic of some notoungulates, especially among Typotheria and Hegetotheria, but insofar as known not so extreme in development as in Patagonia. Neither the enlargement of the incisor nor the re- duction of the cheekteeth is known in any prim- itive Paleocene notoungulates or in other South American "ungulates." Even later rodent-like no- toungulates were much less specialized than the Oligocene Patagonia. South American marsupials diverged very early into unique evolutionary lin- eages (see Simpson, 1970a-c, 1971, 1 980; Pascual, 1980a,b, 1981; Paula Couto, 1979; Reig, 1981). Patagonia peregrina is unquestionably a mar- supial because its unique and diagnostic combi- nation of characters are unknown in any eutherian. Nevertheless, it could be regarded as another of the extinct South American mammals considered by some as incertae sedis and by others as a tertium quid with regard to the eutherian-marsupial di- chotomy (McKenna, 1980; Reig, 1981). However, the marsupial affinities of other peculiar fossil mammals from South America remain unques- tioned, despite weaker support than that offered here for Patagonia. For example, the basis for con- sidering the Polydolopidae as marsupials is the combination of an inflected mandibular ramus, palatal vacuities, and a cheektooth formula of 1-3 1-4 P— -- and ^-r~z- These characters were formerly used to exclude the polydolopids from the Allothe- ria. But, as these characters are present in prim- itive therians outside South America, their diag- nosis of marsupials can be considered an "act of PASCUAL & CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS 105 Fig. 4. Labial (1) and occlusal (2) outlines of mandibles, showing the diflFerent development of incisor. A, Ar- gyrolagus parodii Rusconi; B, Groeberia minoprioi Patterson; C, Proargyrolagus bolivianus Wolff, D, Patagonia peregrina gen. et sp. nov. Graphic scale = 2 mm. faith based on . . . geography and stratigraphic po- sition rather than on . . . biology" (McKenna, 1 980, pp. 58-59). We beheve that assignment of poly- dolopids to marsupials represents the most par- simonious conclusion. Like the newly described Proargyrolagus boli- vianus (Wolff, 1984), Patagonia peregrina is another peculiar marsupial that appears in the fos- sil record without known ancestors (see Simpson, 1970c, p. 16) only to vanish again soon afterward: Groeberiidae (Divisaderan Age, Late Eocene); Pat- agoniidae (Colhuehuapian Age, Late Oligocene); Necrolesiidae (Santacrucian Age, Early Miocene); Argyrolagidae (Huayquerian to Uquian Ages, Late Miocene to Early Pleistocene). We believe there are cedent reasons to think of Proargyrolagus bo- 106 nELDL\NA: ZOOLOGY livianus Wolff, 1984, described as a Deseadan ar- gyrolagid, as possibly representing a distinct fam- ily of Argyrolagoidea. This raises the question of the position of Pat- agonia among the varied ranks of South American marsupials. The previous descriptions and illus- trations demonstrate that Patagonia peregrina has many peculiarities that are rare, differently devel- oped, or completely absent in other marsupials (cf fig. 4). The most striking of these are: 1 . Mandible extremely short and deep, with un- fused subvertical symphysis, dorsally posi- tioned masseteric fossa, ventral border in- flected at level of the mj, enveloping there the alveolus of the incisor. 2. Presence in each ramus of mandible of one rodent-like rootless incisor that extends lin- gually along ventral border of mandible to below the m,. 3. Presence in each ramus of one procumbent canine, single- and closed-rooted, scarcely separated from the incisor and with the oc- clusal apex probably appressed to the inci- sive apex. 4. Three rectangular and continuously growing cheekteeth arranged in close sequence. These and other less striking characters under- score the unique specializations of Patagonia per- egrina, leading to its assignment to a new family, Patagoniidae. But the distinctive combination of characters in the Patagoniidae identify it as a dis- tinct evolutionary group, that is, a different clado- genetic unit. Simpson (1945, 1970a, 1980) des- ignated natural evolutionary groups of marsupials as suF>erfamilies. Following this line of reasoning, Patagoniidae should be allocated to a new super- family, the Patagonioidea. What are the affinities of this new superfamily to other superfamilies within the superorder Mar- supialia? Any discussion of its affinities depends on the systematics of other taxa, many of which are problematic. The systematics of fossil and ex- tant South American marsupials, including the merits of recognizing Marsupialia as a superorder, are discussed by Simpson (1970a, 1971) and Pas- cual (1980b). The majority of South American marsupials represent the order Polyprotodonta; this is roughly equivalent to Ride's ( 1 964) Marsupicamivora, but also includes Ameghino's Paucituberculata (see Pascual, 1980b; contra Kirsch, 1977a,b; Reig, 1981). There is as yet no compelling argument to include any South American families within the Australasian order Diprotodonta (Reig, 1981), de- spite some suggestions to the contrary (e.g., Pas- cual & Herrera, 1 973, 1 975). While the allocations of these groups seems unambiguous, the positions of most remaining groups (e.g., Argyrolagidae, Necrolestidae, and Groeberiidae) remain uncer- tain. With some reservation, Kirsch (1977b) in- cluded the Necrolestidae in the polyprotodont Borhyaenoidea (as did Patterson, 1958), and the Groeberioidea and Argyrolagoidea within the Paucituberculata. Independently, Clemens and Marshall ( 1 976) also treated these animals as mar- supials, recognizing each as superfamilies: Argy- rolagoidea, Necrolestoidea, and Groeberioidea. Like Simpson, they made these assignments with disclaimers that the interrelationships of these groups were far from clear. Reig (1981, p. 60) not only questioned whether the Argyrolagidae (his Microtragulidae) were mar- supials, as none of its known characters are ty- pologically diagnostic, but conjectured probable affinities to the Anagalida. Further, without rig- orous analysis, he suggested that the Argyrolagidae could be treated as an independent order, pro- posing the name Argyrolagida. He concluded that only more intensive study or additional records could substantiate allocation of this order to the Metatheria or the Eutheria. Remains of Patagoniidae exhibit a unique mo- saic of characters, some of which are absent or differently developed in Groeberiidae and Argy- rolagidae. Despite their similarities, each of these taxa apF)ears prima facie to represent indep)endent evolutionary trends. To assess their interrelation- ships, common and distinctive characters of each must be carefully weighed. Remains of Argyro- lagoidea obtained in the same horizon and locality as the hypodigm of Patagonia peregrina should be particularly useful in this regard and are now under study. Ordinal and subordinal allocation of the Patagonioidea await this more comprehensive analysis. Known representatives of this taxon are so highly derived, as is the case with other peculiar marsupials, that their relationships to other mar- supial groups are obscure and can only be clarified by an expanded record of earlier forms. Ecology and Historical Biogeography Biological inferences of Patagonia are necessar- ily limited to the mandibular fragments thus far PASCUAL & CARLINL NEW SUPERFAMILY OF PALEOGENE MARSUPIALS 107 known. These demonstrate unique characters among marsupials, Hving or extinct, which are ob- viously related to a particular mode of life. No known eutherian possesses such mandibular fea- tures. Superficially it is similar to Groeberia, both being rodent-like marsupials: each has a short and deep mandible with a single enlarged, open-rooted incisor, deeply extended along the mandible, with the extra-alveolar part apparently nearly vertical. These represent functional not phylogenetic sim- ilarities, as similar states were attained by different routes: in Groeberia this tooth extended within an odd medial posterior projection of the symphysis, whereas in Patagonia the intra-alveolar portion is truly rodent-like, in being extended along the hor- izontal ramus (cf fig. 4B,D). No doubt both were powerfial gnawers as the lower incisor worked al- most vertically, much more so than in most ro- dents. The unknown face and snout of Patagonia was probably short and deep; whether it had two pairs of lagomorph-like upper incisors like Groe- beria remains unknown. Related to this gnawing specialization, both Groeberia and Patagonia show a short diastema near the alveolar level and a re- duced number of postincisive teeth, four in both; however, Patagonia has three cheekteeth, whereas Groeberia has four. The rodent-like habitus of Pat- agonia is especially advanced, because the three cheekteeth are truly hypselodont, rectangular- shaped in cross section, with at most only shallow lateral grooves representing the remnants of an- cestral bilobate cheekteeth. This combination of features suggests food was obtained by gnawing and prepared for swallowing by grinding. It represents extraordinary conver- gence on some desert-adapted and fossorial forms, such as the Octodontidae. The evolution of cheek- teeth toward a rectangular shape and numerically reduced sequence has been recognized as occurring within the Octodontoidea (from the Octodontidae to the Ctenomyidae; Pascual et al., 1965). The dental features of Patagonia are also convergent on those of the desert-dwelling African Bathyer- gidae, particularly to the sand rat Heterocephalus glaber, and to the North American Geomyidae. These convergent anatomical features suggest that Patagoniidae were probably fossorial marsupials. Anatomical convergence of Patagonia on des- ert-dwelling fossorial rodents is curious, because prevailing conditions in central Patagonia during the Colhuehuapian Age were not highly favorable to desert dwellers. The first record of platyrrhine monkeys in Patagonia occurs at the same locality and level (Fleagle & Sown, 1983) as Patagonia. Many other vertebrate remains recovered at this site (see Bordas, 1939; Donadio, 1983) suggest an environment of well-watered tropical woodlands. Conversely, however, both Argyrolagoidea and very advanced Cephalomyidae rodents from this site (currently under study) show dental features reminiscent of desert or at least drier environ- ments. Generally, the Colhuehuapian vertebrate fauna from central Patagonia (see Pascual, 1970; Pascual & Odreman Rivas, 1971; Marshall et al., 1983) is composed of both forest and open-coun- try types, presumably brought together in a sub- tropical savanna. Thus, the Patagoniidae, Ceph- alomyidae, and Argyrolagoidea occurred in apparently inappropriate environments, probably restricted to xeric patches in the subtropical sa- vanna mosaic. Because the Colhuehuapian Pata- gonioidea were already highly specialized for xeric habitats, they probably evolved earlier in the Pa- leogene. It therefore seems likely that ancestral forms existed in the Deseadan (Early Oligocene). Another highly specialized group of marsupials, the Argyrolagoidea, suggests this hypothesis. For- merly believed present in the record fi-om the Huayquerian (Late Miocene) to the Uquian (Early Pleistocene; see Marshall et al., 1983), argyrola- goids have now been reported from the Deseadan of Bolivia (Wolff, 1984), and here from the Col- huehuapian beds of central Patagonia. The pre-Deseadan record contains no potential ancestor for either Argyrolagoidea or Patagonioi- dea. Simpson (1970c, p. 17) proposed that "these groups (including Groeberioidea) evolved in what are now (and quite likely were then) the tropics and are picked up in our record only when they spread rather briefly to what was for them a mar- ginal area." It seems quite probable that the en- vironments responsible for their initial divergence were poorly or not represented in the known fossil record. Global diastrophic movements in the Late Eocene, and apparently related climatic and en- vironmental changes, are thought to be responsi- ble for the cosmopolitan turnover in Early Oh- gocene mammal communities (Kurten, 1971). This turnover also occurred in South America (Pascual, 1984). Mammal communities in the Deseadan (Early Oligocene) are substantially different from Eocene communities in composition (see Pascual et al., 1 985), apparently reflecting Stehlin's '^^grande coupure."" The apparently sudden occurrence of the Argyrolagoidea. and probably the Patagonioi- dea, in the Deseadan Age is probably another ex- ample of this global turnover. 108 nELDL\NA: ZOOLOGY It is remarkable that, to the numerous succes- sive parallel trends ("successive trends" or "iter- ation"; Simpson, 1953, pp. 248-259; 1961, p. 127) in the evolutionary history of South American mammals, especially from the Deseadan on, can be added the convergence of Oligocene patagoniid marsupials and Pliocene to Recent ctenomyid ro- dents on a common morphology. These conver- gences are products of similar responses to re- peated environmental conditions. The anatomical and functional similarities of Patagonia peregrina with the extant Ctenomys are so striking that we are tempted to call the former the "marsupial tuco- tuco." Acknowledgments All of the material studied here was discovered by 1983 and 1984 expeditions of the Museo Ar- gentino de Ciencias Naturales "Bernardino Ri- vadavia" (MACN), in which Lie. Oscar E. Don- adio and Lie. Miguel Soria (both of MACN) and the American paleontologists John G. Fleagle and Thomas M. Bown participated. Dr. Jose F. Bo- naparte, Chief of the Seccion Paleontologia Ver- tebrados, MACN, and responsible for these ex- peditions, generously put this and other marsupial material at our disposal. The x-ray plates were made by Dr. Roberto Guevara, Profesor de Odon- tologia, Universidad Nacional de La Plata, by the authority of the Dean, Dr. Oscar Barletta. We thank all of them very much. Literature Cited BoRDAS, A. F. 1939. Diagnosis sobre algunos mami- feros de las capas con Colpodon del valle del Rio Chu- but. Physis, 14:413-433. Clemens, W. A., and L. G. Marshall. 1976. Amer- ican and European Marsupialia. Pars 123. Marsupi- alia. In Westphal, F., ed., Fossilium Catalogus. I: An- imalia. W. Junk, The Hague, 1 14 pp. DoNADio, O. E. 1983. Los lacertilios del Colhuehua- pense de la provincia del Chubut, Argentina. Circular Informativa de la Asociacion Paleontologica Argen- tina, 11: 5-6. Fleagle, J. G., and T. M. Bown. 1983. New Primate fossil from Late Oligocene (Colhuehuapian) localities of Chubut Province, Argentina. Folia Primatologica, 41: 240-266. Gould, S. J. 1 983. Nature's Great Era of experiments. Natural History, 7: 12-21. KiRSCH, J. A. W. 1977a. The classification of marsu- pials, pp. 1-48. In Hunsaker II, D., The Biology of Marsupials. Academic Press, New York, San Francis- co, London, 537 pp. . 1977b. The comparative serology of Marsu- pialia, and a classification of marsupials. Australian Journal of Zoology, Supplementary Series, 52: 1-152. KuRTfeN, B. 1971. The Age of Mammals. Columbia University Press, New York, 250 pp. Marshall, L. G. 1976. Revision of the South Amer- ican fossil marsupial subfamily Abderitinae (Mam- malia, Caenolestidae). Publicaciones Museo Munici- pal de Ciencias Naturales de Mar del Plata "Lorenzo Scaglia," 2(3): 57-90. . 1980. Systematicsofthe South American mar- supial family Caenolestidae. Fieldiana: Geology, n.s., 5: i-vii, 1-145. Marshall, L. G., R. Hoffstetter, and R. Pascual. 1983. Mammals and stratigraphy: Geochronology of the continental mammal-bearing Tertiary of South America. Palaeovertebrata, Memoire Extraordinaire, Montpellier, 1983: 1-93. McKenna, M. C. 1980. Early history and biogeogra- phy of South America's extinct land mammals. In Ciochon, R. L., and A. B. Chiarelli, eds.. Evolutionary Biology of the New World Monkeys and Continental Drift. Plenum Publishing Corp., New York, xvi + 528 pp. Mendia, J. E., AND A. Bayarsky. 1981. Estratigrafia del Terciario en el valle inferior del rio Chubut. VII Congreso Geologico Argentino, San Luis (20-26 Sep- tiembre 1981), Actas 3: 593-606. Pascual, R. 1970. Evolucion de comunidades, cam- bios faunisticos e integraciones biocenoticas de los vertebrados cenozoicos de Argentina. Actas IV Con- gresso Latinoamericano de Zoologia (Caracas, 10-16 de Noviembre de 1968), 2 (Paleontologia Sudameri- cana): 991-1088. . 1980a. Nuevos singulares tipos ecologicos de marsupiales extinguidos de America del Sur (Paieo- ceno tardio o Eocene temprano) del Noroeste Argen- tino. Actas II Congreso Argentino de Paleontologia y Bioestratigrafia y I Congreso Latinoamericano de Pa- leontologia (Buenos Aires, 2-6 de Abril 1 978), 2: 151- 173. 1 980b. Prepidolopidae, nueva familia de Mar- supialia Didelphoidea del Eoceno sudamericano. Ameghiniana, 17(3): 216-242. . 1981. Adiciones al conocimiento de Bona- partherium hinakusijum (Marsupialia, Bonapartheri- idae) del Eoceno temprano del Noroeste Argentino. Anais II Congresso Latino-Americano de Paleonto- logia (26 a 30 de Abril, 1981, Porto Alegre-Brasil), 2: 507-520. . 1983. Novedosos marsupiales paleogenos de la Formacion Pozuelos (Grupo Pastes Grandes) de la Puna, Salta, Argentina. Ameghiniana, 20(3-4): 265- 280. . 1984. La sucesion de las Edades-mamifero, de los climas y del diastrofismo sudamericanos durante el Cenozoico: fenomenos concurrentes. Anales de la Academia Nacional de Ciencias Exactas, Fisicas y Naturales, 36: 1 5-37. PASCUAL 8c CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS 109 Pascual, R., and M. Bond. 1981. Epidolopinae subfam. nov. de los Polydolopidae (Marsupialia, Po- lydolopoidea). Anais II Congresso Latino- Americano de Paleontologia (26 a 30 de Abril, 1981, Porto Alegre- Brasil), 2: 479-488. Pascual, R., AND H. E. Herrera. 1973. Adiciones al conocimiento de Pliolestes tripotamicus Reig, 1955 (Mammalia, Marsupialia, Caenolestidae) del Plioceno superior de la Argentina. Ameghiniana, 10(1): 36-50. . 1975. Stilotherium Ameghino, 1887, el mas primitivo Caenolestidae conocido. Consideraciones sobre la transicion Didelphidae-Caenolestinae (Mar- supialia). Actas I Congreso Argentine de Paleontologia y Bioestratigrafia (12-16 Agosto, 1 974, Tucuman, Ar- gentina), 2: 417-430. Pascual, R., and O. E. Odreman Rivas. 1971. Ev- olucion de las comunidades de los vertebrados del Terciario argentine. Los aspectos paleozoogeograficos y paleoclimaticos relacionados. Ameghiniana, 8(3-4): 372-412. Pascual, R., J. PiSANO, and E. J. Ortega. 1965. Un nuevo Octodontidae (Rodentia, Caviomorpha) de la Formacion Epecuen (Plioceno medio) de Hidalgo (Provincia de La Pampa). Ameghiniana, 4(1): 19-30. Pascual, R., M. G. Vucetich, G. J. Scillato-Yane, AND M.Bond. 1985. Main pathways of mammalian diversification in South America, pp. 219-247. In Stehli, F., and S. D. Webb, eds.. The Great American Biotic Interchange. Plenum Publishing Corp., New York. Patterson, B. 1952. Un nuevo y extraordinario mar- supial deseadiano. Revista del Museo Municipal de Ciencias Naturales, Mar del Plata, 1: 39-44. . 1958. Affinities of the Patagonian fossil mam- mal Necrolestes. Breviora (Museum of Comparative Zoology), 49: 1-14. Paula Couto, C. de. 1952. Fossil mammals from the beginning of the Cenozoic in Brasil. Marsupialia: Poly- dolopidae and Borhyaenidae. American Museum Novitates, 1559: 1-27. . 1961. Marsupiais fosseis do Paleoceno do Bra- sil. Anais Academia Brasileira Ciencias, Rio de Ja- neiro, 33(3/4): 321-333. . 1979. Tratado de Paleomastozoologia. Aca- demia Brasileira de Ciencias, Rio de Janeiro, 590 pp. Reig, O. A. 1981. Teoria del origen y desarrollo de la fauna de mamiferos de America del Sur. Monogra- phiae Naturae (Publicadas por el Museo Municipal de Ciencias Naturales "Lorenzo Scaglia"), Mar del Plata, 1: 1-162. Ride, W. D. L. 1964. A review of Australian fossil marsupials. Journal of the Royal Society of Western Australia, 47(4): 91-131. Simpson, G. G. 1945. The principles of classification and a classification of mammals. Bulletin of the Amer- ican Museum of Natural History, 85: i-xxvi, 1-350. . 1953. The Major Features of Evolution. Co- lumbia University Press, New York, 248 pp. 1 96 1 . Life of the Past. Yale Paper Bound, Yale University Press, New Haven, Conn. 1970a. The Argyrolagidae, extinct South American marsupials. Bulletin of the Museum of Comparative Zoology, 139: 1-86. 1970b. Additions to knowledge of the Argy- rolagidae (Mammalia, Marsupialia) from the Late Ce- nozoic of Argentina. Breviora (Museum of Compar- ative Zoology), 361: 1-9. 1970c. Addition to knowledge of Groeberia (Mammalia, Marsupialia) from the Mid-Cenozoic of Argentina. Breviora (Museum of Comparative Zool- ogy), 362: 1-17. 1971. The evolution of marsupials in South America. Anais Academia Brasileira de Ciencias, Su- plemento, 43: 103-118. 1 980. Splendid Isolation. The Curious History of South American Mammals. Yale University Press, New Haven and London, IX + 266 pp. Wolff, R. G. 1984. A new Early Oligocene Argyro- lagid (Mammalia: Marsupialia) from Salla, Bolivia. Journal of Vertebrate Paleontology, 4(1): 108-1 13. 110 FIELDIANA: ZOOLOGY An Additional 14-Chromosome Karyotype and Sex-Chromosome Mosaicism in South American Marsupials Milton H. Gallardo and Bruce D. Patterson ABSTRACTS The karyotype of Rhyncholestes Osgood is described for the first time. The karyotype has 2n = 14 and is similar in most respects to karyotypes of similar number found in other American and Australasian genera in several families. The karyotype of somatic (bone marrow) tissues from male Dromiciops Thomas is presented for the first time; surprisingly, it differs from the 2n = 14 complement previously reported from female bone marrow and male gonads. The 2n = 1 3 karyotype found in bone marrow of male Dromiciops lacks a minute element thought to be the Y chromosome. This instance of somatic chromosome elimination represents the first case reported for American marsupials and presents an interesting parallel to sex-chromosome mosaicism among Australasian Peramelidae and Petauridae. El cariotipo de Rhyncholestes Osgood es descrito por primera vez. El cariotipo consta de 2n = 14 y es muy similar a cariotipos de igual mimero encontrados en otros generos americanos y australoasiaticos de varias familias. El cariotipo de tejidos somaticos (medula osea) de un Dromiciops Thomas macho es presentado por primera vez; sorprendentemente, difiere del complemento 2n = 14 reportado previamente de medula osea femenina y gonadas masculinas. El cariotipo 2n = 1 3 encontrado en medula osea del Dromiciops macho carece de un diminuto elemento que supuestamente corresponde al cromosoma Y. Este ejemplo de eliminacion so- matica de cromosomas representa el primer caso reportado en marsupiales americanos y pre- senta un interesante paralelo con el extenso mosaicismo de los cromosomas sexuales descrito entre las formas australoasiaticas. Descreve-se pela primeira vez, o cariotipo de Rhyncholestes Osgood. O cariotipo e de 2n = 1 4, e, na maioria de seus aspectos, assemelha-se aos cariotipos de numeros similares encontrados em outros generos americanos e austral^sios. O cariotipo de tecidos somdticos (da medula ossea) de Dromiciops Thomas machos e descrito pela primeira vez. Supreendentemente, este cari- otipo difere do complemento de 2n = 1 4, previamente descrito para a medula ossea das femeas e para as gonadas dos machos. No cariotipo de 2n = 13, encontrado na medula ossea de Dromiciops machos, falta um elemento miudo, possivelmente o cromossomo Y. Este e o primeiro exemplo documentado da elimina9ao somatica de um cromossomo em marsupiais americanos, e apresenta um paralelo interessante ao mosaico frequentemente encontrado nos cromossomos sexuais de outras formas austral^sias. From the Institute de Ecologia y Evolucion, Univer- sidad Austral de Chile, Casilla 567, Valdivia, Chile; and Division of Mammals, Field Museum of Natural His- tory, Chicago, IL 60605-2496. GALLARDO & PATTERSON: KARYOTYPES OF MARSUPIALS 1 1 1 Introduction Several unusual cytological features, including low diploid number (Hayman & Martin, 1969; Reig et al., 1977), paternally derived X inactiva- tion (Lyon, 1974a,b), multiple sex-chromosome systems (Hayman & Martin, 1969; Schneider, 1977), somatic elimination of sex-chromosomes (Schneider, 1977; Close, 1984), and sperm con- jugation (Biggers & Creed, 1962; Biggers & De- Lamater, 1965), have made marsupials interesting subjects of cytological research. These studies have clarified fundamental cytological mechanisms. Additionally, results of the research have shed Ught upon directions of chromosomal evolution and upon interrelationships of lineages (Hayman & Martin, 1969; Reig et al., 1977; Sharman, 1982). A 14-chromosome karyotype occurs in several distinct lineages in all living American families: Didelphidae (Reig et al., 1 977), Microbiotheriidae (Spotomo & Fernandez, 1971; Reig et al., 1972), and Caenolestidae (Hayman et al., 1971). This karyotype also occurs in several Australian mar- supial lineages (Hayman & Martin, 1969) and is therefore considered the primitive chromosome number for Metatheria (Reig et al., 1977). Direc- tion of chromosome evolution in Metatheria has proceeded via centromeric dissociations— with pericentric inversions superimposed on the basic Robertsonian mechanism— to give rise to the re- maining 2n = 1 8 and 2n = 22 karyotypes known for American forms (Hayman & Martin, 1969; Reig et al., 1977). Extremes of karyotypic varia- tion in Australasian marsupials are 2n = 10 to 32 (Schneider, 1977). Two autochthonous and endemic South Amer- ican genera, Rhyncholestes and Dromiciops, are especially interesting from an evolutionary view- point. Both are represented by a single species and occur only in the temperate Valdivian rainforests of southern Chile and Argentina. Rhyncholestes. one of three extant genera of Caenolestidae, is widely isolated from its relatives in the northern Andes and presents some striking morphological specializations. Dromiciops. thought by some to have special affinities with Australasian lineages (Sharman, 1982; Szalay, 1982), is the only hving genus of the otherwise extinct Microbiotheriidae (Marshall, 1982). Its affinities with other marsu- pial genera are currently uncertain. In this note we present the first somatic karyotypes of male Rhyn- cholestes raphanurus and Dromiciops australis. Additionally, we document the first instance of somatic sex-chromosome mosaicism in South American marsupials. Materials and Methods Seven specimens of D. australis (five males and two females) from Valdivia (39°32'S, 72°52'W), Osorno (4r06'S, 72°30'W), and Concepcion (37»26'S, 73°19'W) provinces, Chile, were ana- lyzed by the in vivo colchicine-hypotonic citrate technique using bone marrow as a source of mi- toses (Patton, 1967). Modifications of the same procedure were used for the one R. raphanurus collected at La Picada, Volcan Osorno (41*^6'S, 72''30'W); incubation with colchicine lasted 2.5 hours and a slightly more hypotonic solution of sodium citrate was used. A total of 4 1 9 mitotic plates was examined: 29 1 from male and 1 20 from female D. australis and 10 from R. raphanurus. Museimi specimens were deposited in the Collec- tion of Mammals, Instituto de Ecologia y Evolu- cion, Universidad Austral de Chile, and Field Mu- seum of Natural History. Results and Discussion Rhyncholestes raphanurus presents a 2n = 14 complement, consisting of three pairs of large metacentric, one pair of medium-sized metacen- tric, and two pairs of small metacentric autosomes. The sex-chromosomes are an acrocentric X and a minute Y (fig. 1). This karyotype differs morpho- logically from the didelphid 2n = 1 4 in not show- ing a clear break between chromosome groups A and B. It also differs in arm ratios (table 1) from the other living caenolestids, Lestoros and Caeno- lestes (see Hayman et al., 1971). Moreover, the interstitial region of the short arm of pair two shows an achromatic area, resembling a secondary con- striction, not described in other caenolestids (but see discussion in Sharman, 1982). Nevertheless, a 2n = 14 karyotype characterizes all three genera of Caenolestidae, which supports previous claims that this karyotype is primitive for Metatheria (Hayman & Martin, 1969; Hayman et al., 1971; Reig et al., 1 977) and reinforces the pattern of low karyotypic variation within marsupial families. Secondary constrictions can serve as chromo- some markers and are thus useful, in the absence 112 FIELDIANA: ZOOLOGY of banding data, for phylogenetic reconstruction. However, the secondary constriction evident in the karyotype of Rhyncholestes is unreported in other South American marsupials, although sec- ondary constrictions are widespread among Aus- tralasian marsupials (Hayman &. Martin, 1969). Considering commonality and in-group and out- group comparisons, we regard the secondary con- striction of Rhyncholestes as apomorphic. Thus, the similar structures of Australasian marsupials were apparently independently derived and can- not be traced back to some marsupicamivorous or other common ancestor. Chromosome counts from all four male D. aus- tralis consistently indicated 2n = 13 chromo- somes. The diploid number for females was 2n = 14 as was previously reported (Spotomo & Fer- nandez, 1971; Reig et al., 1972). No differences among our specimens from geographically isolated localities were detected, nor were secondary con- strictions evident. Electron microscope studies of sex-chromo- somes in spermatocytes of D. australis and the didelphid Marmosa elegans demonstrate striking similarities (Fernandez et al., 1979). These simi- larities suggest that a 2n = 14 karyotype should be present in D. australis, its Y chromosome should resemble that of M elegans, and both genera should exhibit an XX/XY sex-chromosome system. We have consistently found 2n = 13 chromo- somes in somatic tissues of male Dromiciops and 2n = 14 in female somatic tissue. The missing chromosome in males is dotlike and probably the Y chromosome (fig. 1). Translocation of the Y to an autosome is an unlikely mechanism for the dif- ferences between sexes because males have 2n = 14 in germinal cells and because the sex vesicle appears normal (Fernandez et al., 1979). While it is possible that such a small chromosome might be overlooked in one or a few chromosomal spreads, its universal absence in all counted plates makes this alternative highly unlikely. Available data favor a somatic elimination of the Y chro- mosome. Previous studies have shown that both consti- tutive and facultative heterochromatin can be de- leted from marsupial cells in vivo without appar- ent deleterious effects on cell replication and survival (Hayman & Martin, 1969). Most exam- ples of somatic elimination of sex-chromosomes in marsupials involve the X chromosome in dos- age compensation (e.g., perameUds and petaurids; Close, 1984). Mitotic figures from the testes of B (( a (» A-1 a B-1 II C-1 B-1 A-2 A-3 II •« C-2 XY D& Si! \l A-1 A-2 A-3 U lOjLL »'' A« ^^ c-1 c-2 XX iiii !3 W A-1 A-2 A-3 B-1 Ift 6A c-1 c-2 X Fig. 1 . Karyotypes from bone marrow cells of A, Rhyncholestes raphanurus, male; B, Dromiciops aus- tralis (2n = 1 4), female; C, Dromiciops australis (2n = 13), male. GALLARDO & PATTERSON: KARYOTYPES OF MARSUPIALS 113 Table 1 . Arm ratios Gong arm/short arm) of Rhyn- cholestes autosomes (ratios are based on 1 counted plates; sex chromosomes are acrocentric). Pair 1: 1.43 Pair 2: 1.53 Pair 3: 1.39 Pair 4: 1.44 Pair 5: 1.42 Pair 6: 1.46 Dromiciops do show the XY constitution. There- fore, male zygotes begin development as XY, and the Y is retained in the germinal cell line, but is lost in at least some somatic tissues. More studies will be needed to determine the extent of this mo- saicism in other tissues. We believe this instance of sex-chromosome mosaicism probably represents a parallel, inde- pendently derived case from that in Australasian forms. However, it could be used to support Sza- lay's (1982) assertion that Dromiciops is more closely related to Australasian lineages than any other American form, belonging in the Australa- sian cohort Australidelphia. In this regard it is noteworthy that Sharman's (1982) analysis of gross chromosomal morphology suggested that the 2n = 1 4 karyotype of Dromiciops (virtually identical to those of some burramyids, peramelids, and Vom- batus ursinns) might be highly similar to that of the common ancestor of Australian marsupials. This instance of sex-chromosome mosaicism also bears on Archer's (1976) contention that pera- melids, which also exhibit sex-chromosome mo- saicism, appear to be derivatives of didelphids in basicranial anatomy. Banding studies of chro- mosomal morphology in these groups are needed to help resolve these various suggestions. A "ratchet" model for the evolution of the Y chromosome and dosage compensation has been suggested (Charles worth, 1978). Initially an active chromosome, the Y is homologous to the X, but chiasmata formation (and thus recombination be- tween the two) is suppressed (e.g., Ohno, 1967). A gradual accumulation of deleterious mutations could account for its erosion over time, leading to minute size. In the didelphid Monodelphis dimid- iata. the synaptonemal complex is absent in the X-Y pairing region. Structural elements of the complex are present, but their assembly seems in- hibited by the shortness of the Y chromosome. It could be argued that, in Monodelphis and other metatherians with dotiike Y chromosomes, Y function is apparently reduced to sex determina- tion, unnecessary in at least some somatic tissues. We favor the evolutionary erosion of the Y chro- mosome and its lack of function in the bone mar- row tissue of Dromiciops australis as ultimate causes for this sex-chromosome mosaicism. Late replication of highly heterochromatinized DNA, a proximate mechanism for sex -chromosome mo- saicism suggested in dosage compensation (Hay- man &. Martin, 1974), may account for the acci- dental loss of the minute Y chromosome during mitotic divisions of somatic cells of Dromiciops. Acknowledgments We thank Brian K. Lang and Peter L. Meserve for assistance in obtaining specimens at La Picada. We received financial support from the Direccion de Investigacion, Universidad Austral de Chile (S- 83-03), Field Museum of Natural History, Amer- ican Philosophical Society (Johnson Fund # 1 646), and National Geographic Society (#2582-82). Dr. R. Fernandez kindly facilitated study of Dromi- ciops material. The constructive criticisms of J. A. W. Kirsch, P. Myers, and J. L. 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Pro- ceedings of the Royal Society of London, Series B, 187: 243-268. Marshall, L. G. 1982. Systematics of the South American marsupial family Microbiotheriidae. Field- iana: Geology, n.s., 10: 1-75. Ohno, S. 1967. Sex Chromosomes and Sex-Linked Cells. Springer- Verlag, Berlin, 192 pp. Patton, J. L. 1967. Chromosome studies of certain pocket mice, genus Perognathus (Rodentia: Hetero- myidae). Journal of Mammalogy, 48: ll-'il. Reig, O. a., R. Fernandez, and O. A. Spotorno. 1 972. Further occurrence of a karyotype of 2n = 14 chro- mosomes in two species of Chilean didelphoid mar- supials. Zeitschrift fur Saugetierkunde, 37: 37—42. Reig, O. A., A. L. Gardner, N. O. Bianchi, and J. L. Patton. 1977. The chromosomes of the Didelphi- dae (Marsupialia) and their evolutionary significance. Biological Journal of the Linnean Society, 9: 191-216. Schneider, L. K. 1977. Marsupial chromosomes, cell cycles, and cytogenetics, pp. 51-93. In Hunsaker II, D. D., ed.. The Biology of Marsupials. Academic Press, New York. Sharman, G. B. 1 982. Karyotypic similarities between Dromiciops australis (Microbiotheriidae, Marsupi- alia) and some Australian marsupials, pp. 711-714. In Archer, M., ed.. Carnivorous Marsupials, Vol. IL Royal 2toological Society of New South Wales, Sydney, 804 pp. Spotorno, O. A., and D. R. Fernandez. 1971. The chromosomes of the "monito del monte" Dromiciops australis Philippi. Mammalian Chromosomes News- letter 12(2): 40-41. SzALAY, F. S. 1982. A new appraisal of marsupial phy- logeny and classification, pp. 621-640. In Archer, M., ed.. Carnivorous Marsupials, Vol. II. Royal Zoological Society of New South Wales, Sydney, 804 pp. GALLARDO & PATTERSON: KARYOTYPES OF MARSUPIALS 115 Notes on the Black-Shouldered Opossum, Caluromysiops irrupta Robert J. Izor and Ronald H. Pine ABSTRACTS Caluromysiops is distinct from the three species of Caluromys in external, cranial, dental, skeletal, and phallic characters, although the two genera are certainly more closely related to each other than to any other extant genus. Much uncertainty remains regarding the ecology and distribution of this rare opossum. Caluromysiops es distinto de las tres especies de Caluromys en caracteres extemos, craneales, dentales, esqueletales y falicos, aunque los dos generos son por cierto mas cercanamente rela- cionados entre si que lo es ningun otro genero existente. Todavia hay mucha incertidumbre en relacion a la ecologia y distribucion de este rara raposa. Caluromysiops difere das tres especies de Caluromys en carateres extemos, craniais, dentais, esqueletais e falicos, embora sejam os dois generos claramente mais proximos entre si do que entre qualquer outro genero atualmente existente. A ecologia e a distribuifao desta rara especie continuam muito pouco conhecidas. Introduction The black-shouldered opossum, Caluromysiops irrupta Sanborn, is the rarest of the larger didel- phids. Its history as a subject of scientific study is peculiar, beginning with a very late discovery (195 1); also, many more specimens have been dis- played in zoos (15) than have been collected for museums directly from the wild (2). Despite the paucity of associated data and other shortcomings, zoo animals have been the source of some valuable information during this study. Materials and Methods All zoos known or suspected to have kept Cal- uromysiops were contacted for information on the From the Division of Mammals, Field Museum of Natural History, Chicago, Illinois 60605-2496. Dr. Pine's present address is Illinois Mathematics and Science Academy, Aurora, Illinois 60506-1039. acquisition, history in captivity, and eventual dis- position of animals. All known specimens pre- served in collections were examined by one or both of us, and all tag data were recorded. Results Small sample sizes have hampered previous work on this species, and have affected this study to some extent. Most published information is based on single specimens. Some of the characters described by Sanborn (1951) as diagnostic are in- dividually or ontogenetically variable and not re- liable for identification in all cases. For example, the extent of the hair on the dor- sum of the tail is distinctive in Caluromysiops, although not as extreme as originally described. On immature animals such as the holotype, the furred area reaches nearly to the end of the tail. Adults, however, lack fur on the distal 1 5-20 mm. IZOR & PINE: CALUROMYSIOPS IRRUPTA 117 A^ ■ -i; Fig. 1. Ventral and lateral views of the cranium and lateral view of the mandible of adult male Caluromysiops irrupta. fmnh 60698. Certain incisors have fallen out and have been lost. 118 FIELDIANA: ZOOLOGY Fig. 2. Adult female Caluromysiops irrupta with two young (one is specimen no. cvg M-17 BE 173). Photo courtesy of Edward T. Maruska and the Cincinnati Zoo. The portion of the tail covered dorsally with fur is still much more extensive than even in Calu- romys lanatus, in which only the proximal 50%- 70% is covered. Except perhaps for some Glironia, Caluromysiops is unique among didelphids in that the fur extends onto a distal unpigmented portion of the tail. The distal one-quarter to one-third of the tail fur is also white. In other genera of didel- phids, individuals with some distal portion of the tail skin unpigmented have fur of the tail confined to the proximal pigmented area of the tail skin. The most striking external feature of Caluro- mysiops is probably the pair of dark lateral and dorsal stripes. These typically arise on the back of the hand and run up the inner side of the forelimb onto the shoulder, where they reach their greatest width of 15-30 mm. They approach each other middorsally but usually do not merge, and run in narrowing parallel bands to the rump. In one old individual, cvg M-30 BE 95, which had been dis- played for six years and eight months at the Cin- cinnati Zoo, the pattern is obscured by a general grizzling. A common variant of the pattern has the back of the hand white, with the dark stripe beginning as a sharply delineated black band around the wrist. This feature may occur on one or both forefeet. As Sanborn and others have noted, some in- dividuals of the woolly opossums Caluromys der- bianus and C. lanatus have coloration suggesting the characteristic dorsal markings of Caluromy- siops. In the species of Caluromys, there is typi- cally a darker brown or reddish dorsal area which grades into the paler, grayer sides of the body. In some individuals, this darker region is bisected on the back of the head, neck, and shoulders by a middorsal gray streak. The supposed similarity to Caluromysiops, however, is not at all close. The darker dorsal areas in Caluromys are most sepa- rated in the place where in Caluromysiops they are closest to merging. Moreover, the individuals of Caluromys having the gray middorsal stripe IZOR & PINE: CALUROMYSIOPS IRRUPTA 119 Table 1 . Caluromysiops irrupta formerly exhibited in zoos. Sex Date arrived Date died Disposition of remains Acquisition data Bronx 2too (New York 2^ological Society) F 10 Sept. 1962 28 July 1969 Incinerated? F 20 Nov. 1963 26 Dec. 1964 amnh 208101 National Zoo (Smithsonian Institution, Washington, D.C.) 'Iquitos, Ecuador" (presumably Peru) M 4 Nov. 1969 M 31 Mar. 1971 12 Apr. 1971 (Sent to Lincoln Park Zoo, 11 Oct. 1972) usNM 396160 C. Chase, Miami From Oklahoma City Zoo Oklahoma City Zoo F 23 Nov. 1965 M 28 Jan. 1967 F 19 Dec. 1965 20 Oct. 1970 5 Aug. 1967 Discarded? (Sent to National Zoo, 31 Mar. 1971) Discarded? C. Chase, Miami Lincoln Park Zoo, Chicago F 18 Aug. 1972 M 18 Aug. 1972 (pouch young) F 18 Aug. 1972 (pouch young) M 11 Oct. 1972 7 4 Apr. 1973 25 Mar. 1973 16 Sept. 1974 Discarded? FMNH 121522 FMNH 60154 FMNH 60398 ... Brookfield Zoo (Chicago Zoological Society) M 21 Apr. 1970 19 Aug. 1971 FMNH 60698 R. Baudy, Center Hill, Fla. Tarpon Springs Zoo M (1 Aug. 1972 "rec'd in lab") USNM 397626 ... Cincinnati Zoo F 25 July 1965 F 25 July 1965 (young w/F) F 25 July 1965 (young w/F) M 1 July 1967 8 Mar. 1967 6 Nov. 1965 11 Dec. 1967 27 Feb. 1974 Discarded? CVGM-17BE 173 Discarded? cvG M-30 BE 95 Peru, via Animated Shippers, Miami Peru, via Animated Shippers, Miami Peru, via Animated Shippers, Miami Cuxio (= Cuzco?), Peru, via C. Chase, Miami Data from E. Maruska, M. Jones, J. Eisenberg, A. Dittmar, A. Hamer, C. Chase (all in litt.), and Collins (1973). AMNH = American Museum of Natural History, New York; usnm = National Museum of Natural History, Washing- ton, D.C; FMNH = Field Museum of Natural History, Chicago; cvg = personal collection of E. Maruska, Director, Cincinnati Txio. Table 2. Measurements of Caluromysiops irrupta. Sex No. Total Hind length Tail length foot Greatest Medial skull Basal palatal length Condylo- length length (incl. incisive (incl. (incl. Ear incisors) length incisors) incisors) 6 CVG M-30 BE 95 617 333.5 29 63.7 61.0 57.3 30.0 S USNM 397626 630+ 330 + 52 34 64.5 63.4 59.1 32.3 6 USNM 396 1 60 590 340 51 32 63.6 62.5 57.3 30.5 S FMNH 60698 63.4 60.6 56.5 30.5 9 AMNH 208101 570 310 47 37 62.6 60.7 56.4 29.7 120 FIELDIANA: ZOOLOGY also strongly tend to have the palest extremities, whereas Caluromysiops has extremities with broad blackish bands (on the inner side of the forelimbs and outer side of the hind limbs). Other differences in pelage include the Mar- mosa-like eye rings and the median facial stripe of all Caluromys, which are completely lacking in most Caluromysiops and only faintly suggested in a few. There is no feature of the color pattern indicating that Caluromys and Caluromysiops represent simple variants of a single evolutionary trend. Cranially, the extant didelphids present a rather restricted array of morphologies. All have the same dental formula. The skulls differ primarily in size, in the presence and arrangement of palatal vacu- ities, and in details of the masticatory apparatus such as sagittal crests, shape of the zygomata, and the postorbital processes. To our knowledge, a key to the skulls of the genera has never been con- structed. It is not surprising, therefore, that it is difficult to find trenchant cranial characters sup- porting the distinctiveness of Caluromysiops as a genus. In the context of the family's relative uni- formity, this does not necessarily argue against generic distinction. Pine, however, indicated in Honacki et al. (1982) that he prefers to regard Caluromysiops as a subgenus o^ Caluromys, most- ly because of similarity in skull shape. The dentition of Caluromysiops irrupta was de- scribed by Sanborn (1951) as having larger M'-- and m,., than Caluromys. He noted the absence in the holotype of M^, M^*, and m4 and attributed the lack of an M^ to its probable loss in the cleaning of the skull, but did not discuss the absence of the other molars. The holotype is a juvenile and the developing alveolus of the m4 is quite evident, so the tooth is probably unerupted. The larger size of the molars is generally a valid character distin- guishing Caluromysiops from Caluromys. Some individuals oi Caluromysiops may never have had the minute P', which is frequently lost in adults, but otherwise the dental formula conforms to that of the other didelphids. The single root of the usu- ally spicule-like P' differs from the condition in Caluromys, in which the tooth is double rooted, or at least very broad with an incipient division. There is a strong tendency in Caluromys for the small cusps on the labial stylar shelf to be subdi- vided into as many as nine small, low cusps. Cal- uromysiops typically has five such cusps, each being higher and more distinct than in Caluromys. Caluromys and Caluromysiops are united by the apparently derived character (Archer, 1 982) of clo- sure of the maxillary palatal fenestrae. This feature alone is sufficient to distinguish them from all oth- er living New World marsupials, with the possible exception of some Marmosa. Archer apparently erred in attributing such closure to Glironia. Cal- uromysiops is slightly farther along in the process than Caluromys, with only small, round, paired foramina remaining at the maxillopalatal suture. Species of Caluromys have more or less elongate foramina. Several cranial features of Caluromysiops sug- gest adaptations for strong biting forces. The sag- ittal crest in adults is very pronounced, and the zygomatic arches are robust and widely bowed outward. Rostral length is relatively shorter than in Caluromys, and the mandible is deeper, with the ascending ramus broader and more upright. This seems incongruous in view of the description by Janson et al. (1981) of nectarivorous behavior. Zoo animals, however, have readily accepted a varied diet including animal products (Collins, 1973), and the species probably only exploits nec- tar and pollen opportunistically. Cranial asymmetry is prevalent in our sample. About half of the skulls examined had some sort of deviation of the rostral axis relative to that of the braincase, or deflection of the sagittal crest from the midline. Table 2. Continued. Post- Breadth zygo- Depth Inter- post- Post- matic brain- Maxil- Man- orbital orbital orbital Zygo- brain- Length case Length Length lary dibular con- pro- con- matic case longer (incl. of mand. tooth- M'- tooth- striction cesses striction breadth width nasal bullae) mandible ramus row M' row 12.7 22.1 8.2 38.2 23.4 25.1 22.2 45.8 47.2 22.8 9.5 28.3 13.9 21.1 36.9 23.2 24.4 20.6 46.8 48.6 23.2 8.7 28.8 12.4 21.4 7.8 39.2 22.9 25.1 22.1 47.5 23.1 29.4 14.0 20.1 9.5 37.0 23.6 23.9 9.3 11.3 18.3 9.2 38.0 23.3 25.8 22.3 9.1 IZOR & PINE: CALUROMYSIOPS IRRUPTA 121 Postcranial anatomy of the black-shouldered opossum displays some interesting but as yet inexplicable differences from that of woolly opos- sums. The hind limbs of Caluromysiops are rel- atively much shorter than the forelimbs. The fore- arm is especially long. In addition, all of the skeletal elements are more heavily built than in Caluro- mys, with larger articular surfaces. Both genera exhibit a slightly offset articulation of the second metacarpal, which allows the animals to spread the second and third digits and grasp small branch- es between them. This schizodactylous grip, also found in phalangeroids, is useful for slow, delib- erate climbers which may back up along a branch rather than turn around to proceed headfirst. The tail has 30-31 vertebrae, compared to 36-38 in Caluromys, and has well-developed chevron bones throughout its length. Rosenthal ( 1 972, 1 975) noted that a female Cal- uromysiops was received at the Lincoln Park Zoo with pouch young, which 40 days later still lacked markings and body hair. Details of pouch anatomy were not provided. All of the didelphids examined to date have a more or less cleft glans penis. Biggers ( 1 966) noted that Caluromys derbianus differed from other species he examined in the greater extent of the cleft (half the length of the penis), in the contin- uation of medial urethral grooves to the apices, and in the rounded, slightly bulbous ends of the glans. The single available dissected-out specimen of a Caluromysiops penis (fmnh 60698) suffered some postmortem deterioration and may not be completely representative, but still shows clearly a very deeply split glans (ca. 4 cm) with distinctly enlarged, rounded tips. The urethral grooves also seem to extend nearly to the ends. These characters of the genitalia would seem to ally Caluromys and Caluromysiops. However, Caenolestes also has a deeply cleft glans penis (Os- good, 1921), and many Australian marsupials ex- hibit some version of the same phenomenon, so it may represent a shared primitive character. Moreover, a large majority of didelphid species have not been evaluated in this regard, and the significance cannot be properly assessed. Genitalia of mammals lacking bacula generally have been less studied, even though soft tissue structure can be equally informative (Woolley, 1982), and our cursory survey of preserved material indicates considerable undocumented variety. A remarkable feature, poorly preserved on fmnh 60698, but manifest on the protruding penial apex of FMNH 60398, is a dense covering of small (ca. 1 mm), comified, recurved spines. These are dis- tributed primarily on the rather rugose tip and medial sides of the glans, along the urethral groove. | Osgood (1921) described the glans of Caenolestes as rugose proximally and covered distally by small i circular papillae, but Biggers ( 1 966) noted no such structures on Caluromys or other didelphids ex- j amined. The taxonomic affinities of Caluromysiops ir- rupta have been controversial at both the generic and suprageneric levels. Cabrera (1958), Hersh- kovitz (in Marshall, 1982), and Pine (in Honacki et al., 1982) have suggested that its evident rela- tionship to Caluromys might be better expressed by including it in the latter genus. The present authors are divided on the question of whether this change would improve the current arrange- ment. Reig's (1955) assertion that this species belongs in the Microbiotheriidae has received adequate refutation (Segall, 1969; Szalay, 1982). Kirsch's (1977) attempt to subdivide the Didelphidae is undermined by the fact that his subfamily names Caluromyinae and Dactylopsilinae, as proposed, are nomina nuda. Given his uncertainty about the contents of the supposed subfamilies of didel- phids, this fact could spare future workers consid- erable confusion, although the names may have since become available inadvertantly in subse- quent publications. As most zoo animals have changed hands sev- eral times before reaching their final destinations, there is little likelihood of accurate field data ac- companying them. Among dubious origins re- ported for zoo-held Caluromysiops are Sao Paulo, Brazil, and Iquitos, Ecuador (sic). According to J. A. Davis, Jr. (in litt.), the latter animal "was said by the dealer to have been captured in a backyard on the outskirts of Iquitos, Peru"; see also Bridges (1968) and Davis (1965). Another purported lo- cality, Cuxio, Peru, has not been located and may represent a transcription error for Cuzco. There are only three unquestioned locality rec- ords, all from southern Amazonian Peru, as fol- lows: Peru: Depto. Cuzco; Prov. Quispicanchis, Quince Mil (13°16'S, 70°38'W), 680 m, fmnh 68336 (the holotype). Peru: Depto. Madre de Dios; Itahuania (12°47'S, 7 1°1 3'W), skull is fmnh 84426, skin is in the Museo Nacional de Historia Natural "Javier Prado", Lima. Peru: Depto. Madre de Dios; Manu National 122 FIELDIANA: ZOOLOGY Park, Cocha Cashu Biological Station ( 1 1°55'S, 7 1°1 8'W) (Janson et al., 1981; Terborgh et al., 1984; Emmons, 1984). These three localities are within 1 50 km of each other, along the western margin of the Amazon basin, between 400-700 m elevation. The only sympatric species of Caluromys recorded is C la- natus. Simonetta's (1979) report of a Caluromysiops near Leticia, Colombia, is a problem. Although we are unable to locate the original account, it is our opinion that this record is best discounted. The photograph appears to have been staged with a captive specimen, since the species is nocturnal (Collins, 1973; Janson et al., 1981; Terborgh et al., 1984). Leticia is at least 900 km from the three well-documented localities, and one of the mu- seum specimens we examined (usnm 397626) is known to have passed through Leticia from an unknown source en route to a zoo in Florida. Le- ticia is the location of a major animal dealership, and the point of exportation of many Amazonian species to the U.S. The dusky brown color on the crown of the head, which Simonetta suggests may differentiate his Colombian specimen subspecifi- cally, is variable in the material we examined, and is probably of no taxonomic importance. Conclusions Caluromysiops irrupta is a species which has often been erroneously or incompletely character- ized in the scientific literature. There are now enough specimens in collections to allow reason- ably complete treatments of its morphology, al- though it remains an almost complete ecological and behavioral enigma. Acknowledgments The authors thank those individuals and insti- tutions listed in Table 1 for their invaluable as- sistance in compiling these data, and for loans of specimens in their care. Anita McQuaig, Linda E. Pine, Nobuko Etoh Pine, Joyce Shaw, and Mary Reed helped with the manuscript. Joseph A. Davis and the editors and reviewers made many helpful suggestions. Literature Cited Archer, M. 1982. A review of Miocene thylacinids (Thylacinidae, Marsupialia), the phylogenetic position of the Thylacinidae and the problem of apriorisms in character analysis, pp. 445-476. In Archer, M., ed., Carnivorous Marsupials. Royal Society of New South Wales. BiGGERS, J. D. 1 966. Reproduction in male marsupials, pp. 251-280. In Rowlands, I. W.. ed., Symposia of the Zoological Society of London, 15: 1-559. Bridges, W. 1968. The Bronx Zoo Book of Wild An- imals. New York Zoological Society and Golden Press, New York, 8 unnumbered pp. + 304 pp. Cabrera, A. 1958. Catalogo de los mamiferos de America del Sur. I. (Metatheria-Unguiculata-Camiv- ora). Revista del Museo Argentino de Ciencias Na- turales "Bernardino Rivadavia": 2k>ologia (1957), 4: 1-307. Collins, L. R. 1973. Monotremes and Marsupials. Smithsonian Publication 4888, Smithsonian Institu- tion, Washington, D.C., 323 pp. Davis, J. A., Jr. 1965. Agreat year for rarities. Animal Kingdom, 68(5): 130-133. Emmons, L. H. 1 984. Geographic variation in densities and diversities of non-flying mammals in Amazonia. Biotropica, 16: 210-222. HoNACKJ, J. H., K. E. Kjnman, and J. W. Koeppl, eds. 1 982. Mammal Species of the World. Allen Press and Association of Systematics Collections, Lawrence, Kansas, 694 pp. Janson, C, J. Terborgh, and L. H. Emmons. 1981. Non-flying mammals as pollinating agents in the Am- azonian forest. Biotropica, 12(Suppl.): 1-6. KiRSCH, J. A. W. 1977. The comparative serology of Marsupialia, and a classification of marsupials. Aus- tralian Journal of Zoology, supp. sen, 52: 1-152. Marshall, L. G. 1982. Evolution of South American Marsupialia, pp. 251-272. In Mares, M. A., and H. H. Genoways, eds.. Mammalian Biology in South America. Special Publication Series, F*ymatuning Lab- oratory of Ecology, University of Pittsburgh, 6: 1-539. Osgood, W. H. 1921. A monographic study of the American marsupial, Caenolestes. Field Museum of Natural History, Zoological Series, 14: 1-156. Reig, O. A. 1955. Noticiapreliminarsobrelapresencia de microbiotherinos vivientes en la fauna sudameri- cana. Investigaciones Zool6gicas Chilenas, 2: 121-130. Rosenthal, M. A. 1972. Observations on the water opossum or yapok (Chironectes minimus). Proceed- ings 48th Annual Conference American Association of Zoological Parks and Aquariums held in Portland, Oregon, Oct. 1-5, 1972: 95-98. . 1975. Observations on the water opossum or yapok Chironectes minimus in captivity. International Zoo Yearbook, 15: 4-6. Sanborn, C. C 1951. Two new mammals from south- em Peru. Fieldiana: Zoology. 31: 473-477. Segall, W. 1 969. The middle ear region of Dromi- ciops. Acta Anatomica, 72: 489-50 1 . IZOR & PINE: CALUROMYSIOPS IRRUPTA 123 SiMONETTA, A. M. 1979. First record of Ca/Mrow>'5;op5 cies of Cocha Cashu Biological Station, Manu Na- from Colombia. Mammalia, 43: 247-248. tional Park, Peru. Fieldiana: Zoology, n.s., 21: 1-29. SzALAY, P. S. 1982. A new appraisal of marsupial phy- Woolley, P. A. 1982. Phallic morphology of the Aus- logeny and classification, pp. 621-640. //J Archer, M., tralian species of Anlechinus (Dasyuridae: Marsupi- ed., Carnivorous Marsupials. Royal Society of New alia): A new taxonomic tool?, pp. 767-781. /« Archer, South Wales. M., ed.. Carnivorous Marsupials. Royal Society of New Terborgh, J. W., J. W. FiTZPATRiCK, AND L. Emmons. South Walcs. 1984. Annotated checklist of bird and mammal spe- 124 FIELDIANA: ZOOLOGY Feeding Habits of the Opossum {Didelphis marsupialis) in Northern Venezuela Gerardo A. Cordero R. and Ruben A. Nicolas B. ABSTRACTS The food items in the annual diet of the opossum {Didelphis marsupialis) in northern Ven- ezuela are reported by season, sex, and dental age. One hundred eight opossums were sampled in 21 different sites on a monthly basis from March 1983 to March 1984. The number of food items recorded varies seasonally. By volume, animal foods (63.5%) are more important than plant foods (22.9%) throughout the year. Birds (2 1 .5%), mammals (1 5.3%), insects (14.8%), and fruits (12.8%) are the most prominent foods, by volume. Feeding habits of males and females do not differ significantly. However, diets of young and old animals are different. Se seiialan los componentes de la dieta anual del rabopelado {Didelphis marsupialis) en el norte de Venezuela por epoca del ano, sexo y edad. El muestreo se hizo mensualmente colec- tandose 108 animales desde Marzo 1983 a Marzo 1984 de 21 localidades diferentes. El numero de componentes de la dieta varia estacionalmente. En terminos de volumen, los alimentos de origen animal (63.5%) son mas importantes que los de origen vegetal (22.9%) a traves del aiio. Las aves (21.5%), los mamiferos (15.3%), los insectos (14.8%) y las frutas (12.8%) son las alimentos mas sobresalientes, en terminos de volumen. Los habitos alimentarios de los machos y las hembras no difieren significativamente. Sin embargo, las dietas de los animales jovenes y viejos son diferentes. Relata-se os componentes da dieta anual do gamba {Didelphis marsupialis) no norte da Venezuela, por epoca, sexo e idade. Amostras foram coletadas mensalmente de marfo de 1983 a marfo de 1 984, e de 2 1 locais diferentes, para um total de 108 animais examinados. O nvimero dos componentes da dieta varia sasonalmente. Em termos de volume, os alimentos de origem animal (63,5%) sao mais importantes do que os de origem vegetal (22,9%) atraves do ano. Aves (21,5%), mamiferos (15,3%), insetos (14,8%), e frutos (12,8%) foram os alimentos mais abun- dantes por volume. Apesar dos habitos alimentares nao diferirem entre machos e femeas, a dieta dos animais jovens difere da dieta dos adultos. Introduction of at least seven of 70 species are known (Fleming, 1972; Hunsaker, 1977; Atramentowicz, 1982; Feeding habits of neotropical didelphid mar- Streilein, 1982; Charles-Dominique, 1983; cf. supials are poorly known, in spite of their high Kirsch & Calaby, 1977). However, the informa- diversity and broad geographical distribution. Diets tion reported for most species is based on quali- tative data. This paper reports the food items in- gested by opossums {Didelphis marsupialis) in From the Facultad Ciencias Institute de Zoologia northern Venezuela throughout the year by sea- Tropical, Apartado 47058, Caracas 1041 -A, Venezuela. son, sex, and dental age. CORDERO & NICOLAS: FEEDING HABITS OF OPOSSUMS 125 Study Area Fieldwork was conducted mainly in the Barlo- vento region of the State of Miranda and within the city of Caracas and its surroundings in north- em Venezuela (10°00'-l(y30'N, 66°00'-67°00'E). The climate is highly seasonal, with a humid pe- riod of nine months (May-January) and a dry pe- riod of three months (February-April) in Barlov- ento and seven months of rainfall (May- November) and five months of drought (Decem- ber-April) in Caracas. Annual mean temperature of Barlovento is 26° C versus 20.6° C in Caracas and its surroxmdings. Rainfall is 2,053 mm at Bar- lovento and 1,011 mm at Caracas. Elevations sampled range from 40 m to more than 1 ,000 m above sea level. According to the Holdridge Life Zones (Ewel et al., 1976), the vegetation of Bar- lovento is primarily a humid tropical forest, whereas that of Caracas is mostly in premontane humid forest. Materials and Methods The sample of 108 opossums was assembled fi"om March 1 983 to March 1 984, either from road- kills or hunting. Fifty-two animals were taken from nine localities at Barlovento, whereas 56 speci- mens were taken from 1 2 localities in or near Ca- racas. Body measurements, sex, and dental age of each animal were recorded. Age determination was based on tooth eruption and wear (Petrides, 1 949; Tyndale-Biscoe & MacKenzie, 1976), permitting their grouping into seven age classes (Cordero, un- publ. data, see Appendix 1). Stomach contents were analyzed according to Korschgen's (1980) rec- ommendations. Each stomach and its contents were placed in a fine sieve ( 1 -mm diameter mesh screen) and thoroughly washed under running water in order to separate fine from coarse material. After measuring the entire volume of the contents, each item was separated under a dissecting microscope and its volume recorded. A reference collection was used for the identification of insects. Results Opossum Foods and Seasonal Variation Six (5.6%) of the 108 stomachs we examined were empty. Numbers of stomachs with items were: dry season, 1 6(15.7%) and wet season, 86 (84.3%). Data for these 102 stomachs appear in Table 1 and Figure 1 . Percentage of volume and frequency of occurrence are shown for each class of items in Table 1. Considerable seasonal variation exists in the number of food items recorded. During the dry season, the most important food items are mam- mals, birds, and insects. In the wet season, birds are more important by volume than mammals or insects, and fruits seem to be of greater impor- tance. Gastropods are ingested in a higher pro- portion during dry season than wet season. Snakes, toads, and earthworms are consumed only in the latter period. Food of animal origin is more important (63.5% by volume) than plant food (22.9%) in the diet of opossums throughout the year. By volume, birds (21.5%), mammals (15.3%), insects (14.8%), and fi-uits (12.8%) are the principal foods ingested by opossums. In terms of frequency, insects (49.1%), fiiiits( 18.6%), birds (12.7%), and mammals (8.8%) contribute to the annual diet. Domestic cats {Felis catiis) and rats (Rattus rat- tus) were considered as prey items of opossums because no dipteran carrion larvae were observed in stomach contents. However, unidentified mam- malian remains are more important than those of cat and rat by both volume and frequency. Birds ingested by opossums were either chickens (Gallus sp.) or young birds which were more numerous in the wet season; during the dry season, chickens were recorded as carrion. Avian material account- ed for 12.7% of the stomach contents and 21.4% by volunie. Snakes and toads are consumed at low levels in relation to their abundance in study sites, suggesting that these food items are of little im- portance for opossums in northern Venezuela. Insects of at least nine families occurred in 49.0% of the stomachs, with an annual volume of 14.8%. Beetles and grasshoppers accounted for the ma- jority of insects consumed. Slugs (Veronicellidae) were recorded in the rainy season (1.7% by volume), whereas Vulimulidae are important in the dry season (6.3% by volume). Centipedes and earthworms were poorly repre- sented in the stomachs. Fruits such as Psidium guajava and Giiazuma ulmifolia are very important in the diet of opos- sums. By both volume and frequency of occur- rence, ftiiits are more important in the rainy sea- son. Miscellaneous foods such as garbage (paper, plastic bags, felt, thread filaments), particulate ma- 126 FIELDIANA: ZOOLOGY Table 1. Percentages of volume (V) and frequency (F) of food items of opossums in northern Venezuela in 1983 and 1984, by season and for the year. Wet season Dry season Annual Food items % V %F %V %F %V %F Animals 65.95 90.65 51.22 114.40 63.48 97.03 Mammalia 15.42 5.81 14.75 20.19 15.31 8.82 Felis catus 4.14 2.32 3.44 1.96 Rattus rattus 0.41 7.69 0.07 0.98 Mammal remains 11.28 3.49 14.34 12.50 11.80 5.88 Aves 23.52 13.95 11.27 7.69 21.46 12.74 Gallus sp. 6.42 2.32 5.34 1.96 Young birds 10.04 3.49 8.3S 4.90 Bird remains 7.06 8.14 11.27 7.69 7.77 5.88 Reptilia 0.41 1.16 0.34 0.98 Snake remains 0.41 1.16 0.34 0.98 Amphibia 1.97 1.16 1.64 0.98 Bufo sp. 1.97 1.16 1.64 0.98 Insecta 14.82 47.65 14.70 63.45 14.81 49.01 Coleoptera 7.36 23.24 3.69 23.07 6.76 21.56 Passalidae 0.21 1.16 1.20 7.69 0.28 1.96 Scarabaeidae 5.07 11.63 1.54 7.69 4.48 9.80 Coccinelidae 1.54 7.69 0.26 0.98 Curculionidae 0.31 2.32 0.26 1.96 Meloidae 0.31 1.16 0.26 0.98 Carabidae 0.12 1.16 0.10 0.98 Remains 1.34 5.81 1.12 4.90 Orthoptera 5.12 16.28 0.87 7.69 4.41 14.71 Acrididae 5.12 16.28 0.87 7.69 4.41 14.71 Cursores 0.64 2.32 8.09 25.00 1.89 5.88 Blattaria 0.43 1.16 8.09 25.00 1.72 4.90 Phasmida 0.21 1.16 0.17 0.98 Lepidoptera 1.66 4.65 2.05 7.69 1.72 5.88 Larvae 1.66 4.65 2.05 7.69 1.72 5.88 Homoptera 0.04 1.16 0.03 0.98 Cicadidae 0.04 1.16 0.03 0.98 Mollusca 1.71 8.14 6.25 7.69 2.47 5.88 Veronicellidae 1.71 8.14 1.42 4.90 Vulimulidae 6.25 7.69 1.05 0.98 Chilopoda 1.73 5.81 0.15 7.69 1.46 4.90 ■ Annelida 1.20 5.81 1.00 4.90 Lumbricidae 1.20 5.81 1.00 4.90 Carrion 5.17 1.16 4.10 7.69 4.99 3.92 Dendrophidion parcarinatum 5.17 1.16 4.30 0.98 Gallus sp. 4.10 7.69 0.69 2.94 Plants 22.05 43.01 27.26 32.69 22.92 39.22 Fruits 14.19 20.92 6.15 7.69 12.84 18.63 Psidium gtmjava 5.70 13.95 6.15 7.69 5.77 13.73 Guazuma ulmifolia 6.21 2.32 5.17 1.96 Passiflora sp. 1.91 3.49 1.59 1.96 Mangifera sp. 0.37 1.16 0.31 0.98 Grass remains 0.83 2.32 0.69 1.96 Plant remains 7.03 19.77 21.11 25.00 9.39 18.63 Miscellaneous 2.38 19.76 8.71 12.40 1.98 18.62 Paper trace 11.63 trace 9.80 Plastic bags trace 4.65 trace 6.20 trace 4.90 Felt trace 1.16 trace 0.98 Thread filaments trace 2.32 trace 6.20 trace 2.94 Particulate Material 9.62 12.79 21.52 18.75 11.62 11.76 CORDERO &. NICOLAS: FEEDING HABITS OF OPOSSUMS 127 GRASS 0.7 % SNAKES 0.3% CENTIPEDES 1.5% EARTHWORMS 1.0% Fig. 1. Proportionate annual volumes of major groups of items from 102 stomach contents of opposums from northern Venezuela between March 1983 and March 1984. terial, and plant remains comprised 2.0%, 1 1.6%, and 9.4% by volume, respectively. Garbage items were only recorded for those animals collected in or near Caracas. Variation of Food Items by Sex Feeding habits of male and female opossums are compared in Table 2. By volume, males con- sume mainly fruits (22.8%), birds (17.1%), plant remains (15.4%), and insects (14.5%), whereas fe- males consume mammals (31.4%), birds (14.5%), insects (11.6%), and fruits (8.8%). However, by frequency of occurrence, males consume primarily insects (30%), fruits (19.2%), and plant remains (15.6%); females consume insects (28.6%), plant remains (12.2%), mammals (10.2%), and fruits (10.2%). Both comparisons by means of a Mann- Whitney U test indicate no significant differences between the sexes. Table 2. Food items, by sex, in terms of volume (V) and frequency (F). Males Females (N = = 53) (N = 31) Food items % V %F % V %F Mammalia 9.6 3.6 31.4 10.2 Aves 17.1 6.0 14.5 8.1 Reptilia 0.7 1.2 Amphibia 3.3 1.2 Insecta 14.5 30.0 11.6 28.6 Mollusca 2.3 6.0 4.0 6.1 Chilopoda 1.2 3.6 0.08 2.0 Annelida 0.6 2.4 1.8 4.1 Carrion 2.0 2.0 Fruits 22.8 19.2 8.8 10.2 Plant remains 15.4 15.6 7.6 12.2 Miscellaneous 0.7 3.6 5.2 6.1 Particulate material 12.4 7.2 13.0 10.2 N = Sample size. 128 FIELDIANA: ZOOLOGY A Mann-Whitney U test was also used to com- pare volumes of principal food groups (mammals, birds, amphibians, fruits, insects, and plant re- mains) in the diets of male and female opossums; no significant differences were detected. Despite this, the composition of the diet suggests that males are more arboreal than females. However, a / test comparing the capture frequencies of both sexes on the ground in a 26-hectare grid indicates no significant differences {P > 0.05; Cordero, unpubl. data). The grid contained 18x18 National live- traps, with a distance of 30 m between stations and rows, and was run from December 1981 to May 1984. Variation of Food Items by Age Food of opossums by age classes appears in Ta- ble 3. Note that the number of food items increases as animals become older. By volume and fre- quency of occurrence, animals of younger ages (I, II, III, and IV) consume mainly invertebrates, fruits, and plant remains, while older animals (ages V, VI, and VII) take those items plus mammals and birds, which become more important as the animal ages. Diets of opossums were compared by successive ages, that is, II with III, III with IV, and so on, by Mann-Whitney U tests. No significant differences were detected. However, when diets of young and old animals were compared, significant differences were demonstrated. Nothing has been published on age-related diet variation for D. virginiana, D. albiventris, or any other marsupial. Discussion These results provide a preliminary view of the annual diet of Didelphis marsupialis in north- western Venezuela. This study shows that opos- sums, while omnivorous, are more carnivorous and insectivorous than herbivorous or frugivo- rous. However, we accept these patterns guardedly because they may represent methodological arti- facts: (1) most of our specimens (84.3%) were tak- en in the wet season, so that trophic habits during the dry season are imprecisely known; and (2) the rinsing step in processing stomach contents may have inadvertently washed away traces of fruit pulp that might have been studied using other methods. Our results indicate that insects, fruits, birds, and mammals figure prominently in the annual diet. These figures contrast with those reported by oo ■ • ■* • • 1/^ CTn O 00 00 ■ • O • • 00 r^ 00 r~ (^ . • r~ • . so — »o 00 00 Ov 00 00 vO PO rn »0 (N rn • ■^ ^_ ^ r-' — — • vd r~-' o O 00 d — ■ . . . . vo VO ■O ! : : : i~^' 00 iri 1^ ■<)...«-) — Tf f^ ^ r) o\ O m . . . . O ; ; ; ; O : : : : 8 c .2 ••£ „ •c 2 S « Ji = 3 ^ t» o ? o • - cu S cu CORDERO & NICOLAS: FEEDING HABITS OF OPOSSUMS 129 Molins de la Sema and Lorenzo (1 982) in a study of stomach contents of 47 Didelphis marsupialis sampled from February 1981 to May 1982 in the lowlands of Sierra de Perija in the State of Zulia, northwestern Venezuela. In their study, the order of importance of food items, by frequency, is as follows: plant leaves (68.3%), fruits (56.2%), rep- tiles (42.6%), insects (29.2%), amphibians (28.8%), birds (14.3%), mammals (15.1%), mollusks ( 1 2.2%), and seeds ( 1 1 .4%). The effects of seasonal and habitat differences in the two studies may ex- plain these differences, since the main vegetation types of the lowlands of Perija are dry and humid tropical forests, with eight months of rainfall (April-November) and four months of drought (December-March). Other studies have also shown that opossums feed on vertebrates. The volume we recorded for mammalian prey (15.3%) is low in comparison with diets determined for the Virginia opossum {Didelphis virginiana), except for Lay's ( 1 942) 7% value. Hopkins and Forbes (1980) also recorded cats and rats in low frequencies and volumes in the diets of opossums in Oregon. Similarly, do- mestic chickens figured prominently in the diet of our specimens and have been reported as prey or carrion of D. virginiana in New York (Hamilton, 1951, 1958), Missouri (Reynolds, 1945), Iowa (Wiseman & Hendrickson, 1950), Michigan (Taube, 1947), and Kansas (Sandidge, 1953). In contrast, snakes and toads were taken infrequent- ly, paralleling the results of Blumenthal and Kirk- land (1976), who reported traces of amphibians in the diets of Pennsylvania Didelphis, and of Wise- man and Hendrickson (1950), who showed rep- tiles have a frequency of 1% in the diet of Iowa opossums. The importance of insects in the diet of our animals is somewhat lower than that pre- viously reported for opossums in Michigan (30.4%; Gardner, 1982, citing Dearborn, 1932), Missouri (34.2%; Reynolds, 1 945), and Kansas (42.2%; San- didge, 1953). However, the volumes we report are higher than those in literature records for New York (Hamilton, 1951, 1958), Oregon (Hopkins & Forbes, 1980), and Pennsylvania (Blumenthal & Kirkland, 1976). Records for other inverte- brates are also similar to those in existing literature reports (e.g., Taube, 1947; Hamilton, 1951, 1958; Reynolds, 1945; Sandidge, 1953). Our data and literature records indicate that Di- delphis species have similar diets, embracing a wide range of food items. More detailed studies, espe- cially of food-use in relation to availability, will be needed to establish the degree of euryphagy. Acknowledgments This study was partly granted by CONICET Project SI- 1158. We thank J. Ojasti for sugges- tions and review of the manuscript. We greatly appreciate the editorial assistance of B. Patterson. The staff members of the Estacion Experimental Rio Negro, Universidad Simon Rodriguez pro- vided logistical support during fieldwork. L. Du- que and R. Martinez helped us in the identification of snakes and slugs, and E. Pannier provided some stomach contents. To all of them, our thanks. Literature Cited Atramentowicz, M. 1982. Influence du milieu sur I'activite locomotrice et la reproduction de Caluromys philander (L). Revue d'Ecologie Appliquee (Terre Vie), 36: 373-395. Blumenthal, E. M., and G. L. Kirkland. 1976. The biology of the opossum, Didelphis virginiana in south- central Pennsylvania. Proceedings of the Pennsylvania Academy of Science, 50: 81-85. Charles-Dominique, P. 1983. Ecology and social ad- aptations in didelphid marsupials: Comparison with eutherians of similar ecology, pp. 395-422. In Eisen- berg, J. F., and D. G. Kleiman, eds., Advances in the Study of Mammalian Behavior. Special Publication of the American Society of Mammalogy, no. 7. EwEL, J. J., A. Madriz, and J. A. Tosi. 1976. Zonas de vida de Venezuela. Fondo Nacional de Investiga- ciones Agropecuarias, Caracas, 265 pp. Fleming, T. H. 1972. Aspects of the population dy- namics of three species of opossums in the Panama Canal Zone. Journal of Mammalogy, 53: 619-623. Gardner, A. L. 1982. Virginia opossum {Didelphis virginiana), pp. 3-36. In Chapman, J. A., and G. A. Feldhamer, eds., Wild Mammals of North America. Johns Hopkins University Press, Baltimore. Hamilton, W. J. 1951. The food of the op>ossum in New York State. Journal of Wildlife Management, 15: 258-264. . 1958. Life history and economic relations of the opossum {Didelphis marsupialis virginiana) in New York Slate. Cornell University Agricultural Station Memoir, 354: 1-48. Hopkins, D. D., AND R. B. Forbes. 1980. Dietary pat- terns of the Virginia oix)ssum in an urban environ- ment. The Murrelet, 61: 20-30. HuNSAKER, D. 1977. Ecology of New World marsu- pials, pp. 95-156. In Hunsaker II, D., ed.. Academic Press, New York. KiRSCH, J. A. W., andJ. H. Calaby. 1977. The species of living marsupials: An annotated list, pp. 9-26. In Stonehouse, B., and D. Gilmore, eds.. The Biology of Marsupials. The Macmillan Press Ltd., London and Basingstoke. 130 FIELDIANA: ZOOLOGY KoRSCHGEN, L. J. 1980. Procedures for food-habits analyses, pp. 1 13-127. /n Schemnitz, S. D., ed.. Wild- life Management Techniques. The Wildlife Society, Washington, D.C. Lay, D. W. 1 942. Ecology of the opossum in eastern Texas. Journal of Mammalogy, 23: 147-159. MOLINS DE LA SeRNA, M., AND J. LORENZO PrIETO. 1982. Alimentacion del rabipelado {Didelphis marsupialis) de la Sierra de Perija. Acta Cientifica Venezolana, 33: 410. Petrides, G. a. 1949. Sex and age determination in the opossum. Journal of Mammalogy, 30: 364-378. Reynolds, H. C. 1 945. Some aspects of the life history and ecology of the opossum in central Missouri. Jour- nal of Mammalogy, 26: 361-379. Sandidge, L. L. 1953. Food and dens of the opossum {Didelphis virginiana) in northeastern Kansas. Kansas Academy of Science, 56: 97-106. Streilein, K. E. 1982. Ecology of small mammals in the semiarid Brazilian Caatinga. I. Climate and faunal composition. Annals of Carnegie Museum, 51: 79- 107. Taube, C. M. 1947. Food habits of Michigan opos- sums. Journal of Wildlife Management, 11: 97-103. Tyndale-Biscoe, C. H., and R. B. Mackenzie. 1976. Reproduction in Didelphis marsupialis and D. albi- ventris in Colombia. Journal of Mammalogy, 57: 249- 265. Wiseman, G. L., and G. D. Hendrickson. 1 950. Notes on the life history and ecology of the oi>ossum in south- east Iowa. Journal of Mammalogy, 31: 331-337. Appendix 1. supialis. E>ental age classes for Didelphis mar- Tooth Age Age eruption Wear class (months) dP' M' I 3.0-3.5 dP' M^ II 4.5-5.0 dP' M' III 6.2-6.7 P' M' IV 7.9-8.7 P' M* V 10.9-11.7 P' M- P^ M'-2 VI 12.8-14.1 P' M* P' M^ VII > 16.1 Source: G. A. Cordero (unpublished data). CORDERO & NICOLAS: FEEDING HABITS OF OPOSSUMS 131 Notes on Distribution of Some Bats from Southwestern Colombia Michael S. Alberico ABSTRACTS Noteworthy range extensions are presented for Noctilio albiventris, Rhinophylla alethina, Sturnira aratathomasi, and Lonchophylla handleyi, including the second Colombian report for the last. A previous report of Molossops brachymeles is clarified as representing M. abrasus. Se presentan algunas notables extensiones del rango de distribucion para las especies Noctilio albiventris, Rhinophylla alethina, Sturnira aratathomasi y Lonchophylla handleyi, este ultimo siendo el segundo reporte para Colombia. Un reporte anterior de Molossops brachymeles se clarifica como representativo de M. abrasus. Apresentam-se notaveis exten^oes mas distribui96es das especies Noctilio albiventris, Rhin- ophylla alethina, Sturnira aratathomasi, e Lonchophylla handleyi, esta ultima sendo apenas o segundo registro para a Colombia. Clarifica-se o registro anterior de Molossops brachymeles como representativo de M. abrasus. Introduction Despite considerable interest in Neotropical mammals, southwestern Colombia remains poor- ly understood in this respect. This is mainly a result of a lack of adequate collections caused by the inaccessible nature of much of the zone. Early collecting expeditions to which we owe much of our knowledge were undertaken around the turn of the century by personnel of the American Mu- seum of Natural History and summarized by Allen (1916). Bats were typically underrepresented in these early collections because of inadequate col- lecting techniques in use at the time. Now, with the aid of Japanese mist nets, we are able to obtain more complete samples of bat communities. In this report I present results of a continuing col- lecting effort during the past five years in this poor- ly known region, extending the known distribution From the Departamento de Biologia, Universidad del Valle, Call, Colombia. of Noctilio albiventris, Lonchophylla handleyi, Rhinophylla alethina, Sturnira aratathomasi, and Molossops abrasus. All specimens mentioned were collected in mist nets, prepared as standard study skins with skulls, and deposited in the mammal collection of the Departamento de Biologia, Universidad del Valle, Cali, Colombia (UV). Distribution Noctilio albiventris The lesser bulldog bat was recently reviewed by Davis (1976) and by Hood and Pitocchelli (1983). Both mapped the distribution as including eastern Colombia across the Llanos and Amazonas and the northern Caribbean coast. Davis (1976) re- ported the altitudinal range of the species as ex- tending up to 1 , 1 00 m. We have found this species to be common in the upper Cauca valley, between ALBERICO: DISTRIBUTION OF COLOMBIAN BATS 133 the Cordillera Central and the Cordillera Occi- dental of the Andes, where the elevation reaches this approximate limit. Fifteen specimens from the Departamento (= state) del Valle del Cauca and adjacent Departamento del Cauca were com- pared with the descriptions and measurements of all subspecies recognized by Davis (1976). This population is indistinguishable from N. a. minor in all characters examined and undoubtedly fol- lows the Rio Cauca south from the Caribbean low- lands. A similar southern extension is most prob- able in the valley of the Rio Magdalena to the Departamento de Huila, but has yet to be con- firmed by collections. Specimens Examined— Cauca: Rio Palo, 18 km S, 5 km E Puerto Tejada, 3°04'N, 76°22'W, 1,050 m (3 92, UV3 1 3, 324, 325); Valle del Cauca: 2 km S, 4 km W Candelaria, 3°23'N, 76°23'W, 1,000 m (1 (5, UV676); Universidad del Valle (Melendez Campus), 8 km S Cali, 3°22'N, 76°32'W, 1,000 m (5 S6, UV2602, 2603, 2604, 2608, 2609; 2 99, UV2605, 2607); 13 km S, 1 km E Cali, 3°22'N, 76°32'W, 1,000 m (2 66, UV2620, 2611; 1 9, UV2612). Lonchophylla handleyi This species was described on the basis of spec- imens from Peru and southern Ecuador by Hill (1980), who suggested that some individuals in existing collections might be misidentified as L. robusta. Lonchophylla handleyi was first reported for Colombia by Alberico and Orejuela (1982), who collected a single individual from near the Ecuadorian border at 850 m. A specimen recently collected from the Departamento del Valle del Cauca at 480 m provides the second record for Colombia. Both specimens are larger (greatest length of skull, 28.4 and 28.6 mm, respectively) than the largest L. robusta reported by Hill (1980) for Peru and Ecuador and are larger than any L. robusta in our collections from western Colombia. Both Colombian specimens of L. handleyi are from the lower slope Andean forests, probably one of the last habitats to be intensively sampled for mammals in this country. The presence of this species in a relatively narrow elevational band within this habitat type attests to the importance of continued collecting in the Pacific slope of the Andes in southwestern Colombia. Specimens Examined— Nariilo: 5 km E Junin, l''20'N, 78°08'W, 850 m (1 6, UV3007); Valle del Cauca: Rio Cajambre, approx. 60 km S Buena- ventura, 3°20'N, 77°00'W, 480 m (1 9, UV3694). Rhinophylla alethina This species was described based on specimens from western Colombia in the Departamento del Valle del Cauca (Handley, 1 966) and until recently was known only from the type locality. Albenco and Orejuela (1982) reported it from Narifio near the Ecuadorian border and suggested that it might have a broader geographic range than previously thought, which was confirmed by Baud (1982) who reported the species for Ecuador. Our collections show R. alethina to be relatively common in the Pacific lowlands and the adjacent lower slopes of the western Andes up to 850 m. That this species was only recently described and remains poorly known is undoubtedly due to insufficient collect- ing in the forests of this zone. Specimens Examined— Nariflo: 5 km E Junin, 1°20'N, 78°08'W, 850 m (3 66, UV3029, 3033, 3036; 5 99, UV3030, 3031, 3032, 3034, 3035). Valle del Cauca: Alto Anchicaya, 35 km S, 20 km E Buenaventura, 3°34'N, 76°54'W, 400 m (2 66, UV3166, 3167); Rio Azul, 5 km N, 25 km W Darien, 3°59'N, 76°44'W, 560 m (1 9, UV3391); Rio Cajambre, approx. 60 km S Buenaventura, 3°20'N, 77°00'W, 480-520 m (l 6, UV3702; 1 9, UV3703); Rio Cahma, 13 km N, 14 km E Bue- naventura, 4°00'N, 76°59'W, 40 m (1 9, UV2809). Stumira aratathomasi In their description of this species, Peterson and Tamsitt (1968) reported three specimens, the ho- lotype from the Departamento del Valle del Cauca in western Colombia and two from an unknown locality in Ecuador. They stated that it might be restricted to the Pacific side of the Andes. Thomas and McMurray ( 1 974) provided measurements for the holotype and six individuals collected near the type locality and suggested that this species may be common at high elevations in the western An- des of Colombia. Our recent collections extend the known range some 150 km to the north in the Cordillera Occidental and, more importantly, re- cord the presence of S. aratathomasi in the Cor- dillera Central, where it was previously unknown. This species appears to inhabit medium to high elevation forests which are relatively continuous 134 FIELDIANA: ZOOLOGY in Colombia, and its occurrence both farther to the north and in the Cordillera Oriental is likely. Specimens Examined— Valle del Cauca: Cor- dillera Central: Hacienda "Los Alpes," 6 km S, 1 1 km E Florida, 3°16'N, 76°09'W, 2,400 m (1 9, UV3482); Cordillera Occidental: Betania, 10 km N, 15 km W Bolivar, 4°26'N, 76''19'W, 1,800 m (1 9, UV3876); Parque Nacional "Los Farallones de Cali," 1 km S, 1 6 km W Cali, 3°22'N, 76°4 1 'W, 2,600 m (1 9, UV3373); Paso de Galapagos, 8 km N, 4 km E El Cairo, 4°50'N, 76°12'W, 1,800 m {2 66, UV4131,4133). Molossops abrasus This species was reported for Colombia by Al- berico and Naranjo-H. (1982) as M. brachymeles, based on specimens from the Cauca valley in northern Valle del Cauca. Although often referred to by this latter specific epithet (see Cabrera, 1958; Freeman, 1981), the holotype of Dysopes abrasus from Brazil has been shown to represent this species (Husson, 1962; Carter & Dolan, 1978). The Co- lombian record extends the known distribution of M. abrasus in western South America from An- dean Peru some 1 ,600 km to the north. Specimens Examined— Valle del Cauca: 1 1 km S, 2 km W Cartago, 4''39'N, 75°56'W, 930 m (2 66, UV2451, 2452; 1 9, 2453). Acknowledgments This report is the result of the combined efforts of many friends and students, too numerous to mention by name, who have collaborated either by accompanying the author in the field, by sharing specimens collected during other activities, or both. However, a few individuals have contributed more than could be expected in the normal turn of events, and their support in the field and out has been especially important in the present study: Eduardo Velasco, Gloria Giral, Alonso Gonzalez, Guiller- mo Cantillo, and Luz Marina Alberico. To these, the author is most appreciative. Literature Cited Alberico, M., and L. G. Naranjo-H. 1982. Primer registro de Molossops brachymeles (Chiroptera: Mo- lossidae) para Colombia. Cespedesia, II: 141-143. Alberico, M., AND J. E. Orejuela. 1982. Diversidad especifica de dos comunidades de murcielagos en Na- rino, Colombia. Cespedesia, Suplemento no. 3(4 1-42): 31-40. Allen, J. A. 1916. List of mammals collected in Co- lombia by the American Museum of Natural History expeditions, 1910-1915. Bulletin of the American Museum of Natural History, 35: 191-238. Baud, F. J. 1982. Presence de Rhinophylla alethina (Mammalia, Chiroptera) en Equateur et repartition actuelle du genre en Amerique du Sud. Revue Suisse de Zoologie, 89: 8 1 5-82 1 . Cabrera, A. 1958. Catalogo de los mamiferos de America del Sur. Revista del Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Ciencias Zoologicas, 4: 1-307. Carter, D. C, and P. G. Dolan. 1978. Catalogue of type specimens of neotropical bats in selected Euro- pean museums. Special Publications, The Museum, Texas Tech University, 15: 1-136. Davis, W. B. 1976. Geographic variation in the lesser noctilio, Noctilio albiventris (Chiroptera). Journal of Mammalogy, 57: 681-107. Freeman, P. W. 1981. A multivariate study of the family Molossidae (Mammalia, Chiroptera): Mor- phology, ecology, evolution. Fieldiana: Zoology, n.s., 7: 1-173. Handley, C. O., Jr. 1966. Descriptions of new bats {Choeroniscus and Rhinophylla) from Colombia. Pro- ceedings of the Biological Society of Washington, 79: 83-88. Hill, J. E. 1 980. A note on Lonchophylla (Chiroptera: Phyllostomatidae) from Ecuador and Peru, with the description of a new species. Bulletin of the British Museum (Natural History), Zoology Series, 38: 233- 236. Hood, C. S., and J. Pitocchelll 1983. Noctilio al- biventris. Mammalian Species, 197: 1-5. HussoN, A. M. 1962. The bats of Suriname. Rijks- museum van Natuurlijke Historic, Leiden, 58: 1-282. Peterson, R. L., and J. R. Tamsitt. 1968. A new species of bat of the genus Sturnira (family Phyllosto- matidae) from northwestern South America. Life Sci- ences Occasional Papers, Royal Ontario Museum, 12: 1-8. Thomas, M. E., and D. N. McMurray. 1974. Ob- servations on Sturnira aratathomasi from Colombia. Journal of Mammalogy, 55: 834-836. ALBERICO: DISTRIBUTION OF COLOMBIAN BATS 135 Distributional Records of Bats from the Caribbean Lowlands of Belize and Adjacent Guatemala and Mexico Timothy J. McCarthy ABSTRACTS Thirty new species records are presented for the bat fauna of Belize, along with secondary records for eight bats that had been recorded previously from that country. Contiguous lowland localities in Guatemala provided new department records: nine for El Peten, five for Izabal, and two for Alta Verapaz. The El Peten records include the first confirmation of Vampyrum spectrum in Guatemala. One state record for Quintana Roo, Mexico, is reported. These species represent the genera Saccopteryx, Balantiopteryx, Diclidurus, Noctilio, Pteronotus, Mormoops, Micronycteris, Lonchorhina, Macrophyllum, Tonatia, Mimon, Phyllostomus, Phylloderma, Trachops, Chrotopterus, Vampyrum, Glossophaga, Uroderma, Vampyrops, Vampyrodes, Vam- pyressa, Chiroderma, Artibeus, Centurio, Diphylla, Natalus, Myotis, Eptesicus, Lasiurus, Bau- erus, Eumops, and Molossus. Range extensions are acknowledged for Saccopteryx leptura, Diclidurus virgo, Noctilio leporinus, Micronycteris nicefori, Macrophyllum macrophyllum, Phyl- lostomus discolor, Vampyrum spectrum, Glossophaga commissarisi, Uroderma bilobatum, Vam- pyrodes caraccioli, Artibeus toltecus, and Bauerus dubiaquercus. A checklist of the bat fauna of Belize, which stands at 66 species, is presented. Se registran 30 especies que no habian sido citadas antes para la fauna de murcielago de Belice, con registros secundarios para ocho murcielagos ya conocidos de ese pais. En ciertas localidades contiguas de las tierras bajas de Guatemala, se obtuvieron nuevos registros depar- tamentales: nueve de El Peten, cinco de Izabal, y dos de Alta Verapaz. Los registros de El Peten incluyen la primera confirmacion de Vampyrum spectrum en Guatemala. Ademas, se presenta un nuevo registro estatal para Quintana Roo, Mexico. Las especies obtenidas estan segregadas en los generos Saccopteryx, Balantiopteryx, Diclidurus, Noctilio, Pteronotus, Mormoops, Mi- cronycteris, Lonchorhina, Macrophyllum, Tonatia, Mimon, Phyllostomus, Phylloderma, Trach- ops, Chrotopterus, Vampyrum, Glossophaga, Uroderma, Vampyrops, Vampyrodes, Vampyressa, Chiroderma, Artibeus, Centurio, Diphylla, Natalus, Myotis, Eptesicus, Lasiurus, Bauerus, Eu- mops, y Molossus. Para cada una de las siguientes especies de murcielagos se anota el alcance geografico de su distribucion conocida: Saccopteryx leptura, Diclidurus virgo, Noctilio leporinus, Micronycteris nicefori, Macrophyllum macrophyllum, Phyllostomus discolor, Vampyrum spec- trum, Glossophaga commissarisi, Uroderma bilobatum, Vampyrodes caraccioli, Artibeus tolte- cus, y Bauerus dubiaquercus. Se incluye una lista de 66 especies que representan la fauna de murcielagos de Belice. Apresenta-se records de 30 novas especies de morcegos para Belice, e de oito especies pouco conhecidas no pais. Areas adjacentes, na Guatemala, providenciaram novos records para: El From the Department of Mammalogy, American Mu- seum of Natural History, Central Park West at 79th Street, New York, NY 10024. MCCARTHY: DISTRIBUTION OF BATS 137 Peten (nove especies), Izabal (cinco especies), e Alta Verapaz (dois especies). Os records de El Peten incluem as primeiras confirma^oes de Vampyrum spectrum na Guatemala. Um novo record para Quintana Roo, Mexico, e incluido. Estas especies representam os generos Saccop- teryx, Balantiopteryx, Diclidurus, Noctilio, Pteronotus, Mormoops, Micronycteris, Lonchorhina, Macrophyllum, Tonatia, Mimon, Phyllostomus, Phylloderma, Trachops, Chrotopterus. Vam- pyrum, Glossophaga, Uroderma, Vampyrops, Vampyrodes, Vampyressa, Chiroderma, Artibeus, Centurio. Diphylla, Natalus, Myotis, Eptesici4s, Lasiurus. Bauerus, Eumops, e Molossus. Re- conhece-se extensoes nas areas onde sao encontrados Saccopteryx leptura, Diclidurus virgo, Noctilio leporinus, Micronycteris nicefori, Macrophyllum macrophyllum, Phyllostomus discolor, Vampyrum spectrum, Glossophaga commissarisi, Uroderma bilobatum, Vampyrodes caraccioli, Artibeus toltecus. e Bauerus dubiaquercus. Apresenta-se uma lista da fauna de morcegos em Belice, que agora conta com 66 especies. Introduction Inventories of bat communities in Mexico and Central America have increased significantly dur- ing the last twenty-five years (Jones et al., 1977). Although the resulting data have enhanced our knowledge of the distributions and the zoogeo- graphical relationships of species, incomplete sur- veys exist for certain regions. The northern low- lands along the Caribbean coast of Honduras, Guatemala, Belize, and Quintana Roo, in Mexico, is one such region. Travel within this coastal ver- sant has improved with agricultural and settle- ment expansion. The isolation of Belize from its neighbors has been reduced with the construction of roads in Guatemala's frontier of El Peten and Mexico's former territory of Quintana Roo. A paved road from Izabal now connects El Peten and Belize with the Pan-American Highway in western Guatemala. Road development continues within Belize for all-weather travel. Belize is situated within the Caribbean lowland drainage of northern Central America. Contiguous with Belize on this eastern slope is the eastern portion of the department of El Peten to the west and, to the south, the department of Izabal, both of Guatemala. Southern Quintana Roo of penin- sular Mexico borders to the north (see fig. 1 and Gazetteer). The topography of these Caribbean lowlands extends from the lower ranges (600 m and below) of the eastern Sierra de Chama, the Sierras de las Minas, the Sierra de Santa Cruz, the Sierra del Meredon, and the Montarias del Mico in Alta Verapaz and Izabal, and the Maya Moun- tains of southern Belize and southeastern El Peten to the low undulating relief of southern Quintana Roo. The Maya Mountains represent a heavily eroded Paleozoic formation that now ranges at the top from 671 to 853 m in elevation, with the high- est peak at 1113m (Wright et al., 1959). Annual rainfall in portions of Izabal averages from 3,000 to nearly 5,000 mm (Portig, 1976). Over 4,500 mm of rainfall was reported from the most south- em coastal area of Belize. North and northwest- ward of the Maya Mountains, rainfall decreases appreciably to less than 1 ,500 mm in north-central El Peten and northern Belize, where less than 1 ,400 mm was recorded near the Quintana Roo border (Walker, 1973). The severity of this northward reduction of rainfall is intensified by the increased lack of surface drainage into the Yucatan Penin- sula of Mexico. Because the limestone shelf of northern Belize has geological affinities with the Yucatan Peninsula (Wright et al., 1 959), the south- em limit of this peninsula can be considered the fault line extending from north of the Maya Moun- tains westward through the northem shore of Lake Peten-Itza, El Peten (Wadell, 1938; West, 1964). Effectively, the northem plain of Belize and north- em El Peten are portions of the Yucatan Peninsula. The northward shift from alluvial soils to shallow calcareous soils, along with the mentioned cli- matic changes, create edaphic conditions that af- fect the composition and the structure of the vege- tation that can be supported (Lundell, 1 934, 1 937; Standley & Record, 1936; Wright el al., 1959; Pen- nington & Samkhan, 1968). The potential effect of this transitional physiography on the distribu- tion and relative abundance of bats in this Carib- bean lowland region will require further inventory studies. This paper documents 30 new records for Belize. A checklist of the known bat fauna for this country is annotated in the Appendix. Sixty-six species are cited. Included here are also records from nearby localities for El Peten, Izabal, and Alta Verapaz, Guatemala, and Quintana Roo, Mexico. Nine species records from EI Peten, five records from 138 HELDIANA: ZOOLOGY Izabal, and two records from Alta Verapaz in- crease the number of reported species for these departments to 35, 31, and 40, respectively (see Jones, 1966; Carter etal., 1966; Rick, 1968; Smith, 1972; LaVal, 1973a; Martinez R., 1980; Mc- Carthy, 1982). Jones et al. (1973) and Bimey et al. (1974) summarized the records for 31 species from Quintana Roo, and this paper provides one additional record. Materials and Methods The bats that I collected during the years 1974- 1984 in Belize and El Peten (Parque Nacional Ti- kal), Guatemala, were obtained principally with mist nets set at ground level; aerial netting and the use of a bat trap were limited. Unless otherwise stated, mist netting was carried out during the first half of the night. A limited number of specimens were obtained with hand nets or plastic funnel traps at roost sites. Specimens were prepared as standard museum skins with skulls and/or skele- tons, or as fluid-preserved specimens. These vouchers are housed in Field Museum of Natural History, Chicago (FMNH); The Museum, Mich- igan State University, East Lansing (MSU); Car- negie Museum of Natural History, Pittsburgh (CM); and American Museum of Natural History, New York (AMNH). A survey of 45 museum collections in the United States, Canada, Mexico, Guatemala, and England resulted in additional specimens from Belize, El Peten, Izabal, Alta Verapaz, and Quintana Roo. Pertinent specimens (147) have been included in this report from the following institutions [collec- tors in brackets]: American Museum of Natural History, New York [N. Sullivan]; British Museum (Natural History), London, England (BM) [R. H. L. Disney; P. Williams; A. M. Hutson; R. E. Steb- bings]; Carnegie Museum of Natural History [N. A. Bitarj; Field Museum of Natural History [L. de la Torre]; Florida State Museum, University of Florida, Gainesville (FSM) [F. J. Bonaccorso]; Museum of Zoology, Louisiana State University, Baton Rouge (LSUMZ) [D. M. Uy]; Royal On- tario Museum, Toronto, Canada (ROM) [R. L. Peterson; J. Kamstra; J. Fragoso]; Texas Coop- erative Wildlife Collections, Texas A«&,M Uni- versity, College Station (TCWC) [D. C. Carter; M. D. Engstrom]; Texas Tech University, Lubbock (TTU) [P. Diamond]; and United States National Museum of Natural History, Washington, D.C. (USNM) [E. L. Tyson]. Systematic arrangement of species accounts and nomenclature, unless otherwise indicated, follow Jones et al. (1977) and Handley (1980). Disney (1968) did not provide data for the first records of Pteronotus davyi, Tonatia minuta, and Eptesi- cusfurinalis from Belize. Those data are presented in the respective accounts of this report, with ad- ditional records. Further secondary records from Belize of Mimon crenulatum, Trachops cirrhosus, Glossophaga commissarisi, Vampyressa pusilla, and Eumops auripendulus are also included. All of the species accounts are discussed in the context of their range and elevational distributions in Mexico and Central America. Hall (1981) was the primary reference for this unless cited otherwise. Forest types in Belize follow Wright et al. ( 1 959), whose classification was partially based on the sea- sonal formation series (Beard, 1 944), which refers to structural appearance. The correct political alignments between the states of the Yucatan Pen- insula are inconsistent among a number of pub- lished maps. The state boundary between Quin- tana Roo and Campeche on the map in Figure 1 (see also Gazetteer) is based on a number of Gov- ernment of Mexico (Secretaria de Programacion y Presupuesto) maps, including "Carta Topografica, Merida" (1:1 ,000,000; 1 979 and 1 983) and "Mapa Geografica" (1:5,000,000; 1980). Species Accounts Family EMBALLONURIDAE Subfamily EMBALLONURINAE Saccopteryx leptura (Schreber, 1 774) Specimen Examined— BELIZE. Toledo: 2. 1 km NNE Salamanca Camp, Columbia Forest, 1 9 (cm). The known distribution of this small sac-winged bat north of Panama extends through Costa Rica and Nicaragua to Chiapas along the Pacific ver- sant. The presence of predominantly lowland Sac- copteryx leptura in southern Belize represents a country record and an extension of its distribution along the Caribbean side from southeastern Nic- aragua. Small bats were observed foraging up to heights of 13-13.5 m during the twilight period of the evening. Flight appeared to be concentrated within MCCARTHY: DISTRIBUTION OF BATS 139 a small, open area below the lower canopy of ev- ergreen seasonal forest. A short mist net was hand- hoisted to capture (24 March) this adult specimen. Saccopteryx bilineata was collected shortly after the capture of S. leptura. Balantiopteryx io Thomas, 1 904 Specimens Examined— GUATEMALA. EI Pe- ten: Poptun, Finca Ixobel, \S 66, 1 8 99 (cm). The restricted distribution of Balantiopteryx io ranges from the Gulf lowlands of Veracruz, Oa- xaca, and Tabasco to the lowlands of Belize and eastern Guatemala. Kirkpatrick et al. ( 1 975), Cart- wright and Kirkpatrick (1977), and Sanborn (1936) represent the previous records for Belize and Iza- bal. The Poptun locality represents the first record for El Peten. The specimens reported here were collected ( 1 2 June) by N. A. Bitar as they exited from a cave surrounded by secondary forest. The distribution of this colonial species may be restricted in part by the availability of adequate cave habitats as roosting sites. Subfamily DICLIDURINAE Diclidurus virgo Thomas, 1903 Specimen Examined— BELIZE. Cayo: 1.5 km SSW Roaring Creek, 1 6 (fmnh). The white bat is represented by relatively few localities in Middle America, which extend from western (Nayarit) and eastern (southern Veracruz) Mexico through Central America. Specimens from southwestern El Peten were reported by Jones (1966). The single specimen from Belize repre- sents a northward range extension in the Carib- bean lowlands from northwest Honduras (Carter & Dolan, 1978) and a record for the country. The single bat apparently was roosting on the trunk of a fig tree (Ficus insipida) overhanging a pool along the Roaring Creek River. It was cap- tured (May) by C. Tzul after being observed on a number of occasions roosting near, but not among, a group of Rhynchonycteris naso. Jones ( 1 966), Starrett and Casebeer ( 1 968), and Handley ( 1 976) commented on the high foraging habits of Dicli- durus. Similar to the molossid bats, Diclidurus probably concentrates its foraging efforts at levels well above the tree canopy and beyond the reach of conventional collecting techniques, except fire- arms. This may explain why there are few speci- mens available in collections. Goodwin ( 1 969) considered Diclidurus virgo at best not more than subspecifically different from D. albus. Both species were recognized by Ojasti and Linares (1971), who questioned Goodwin ( 1 969) because they believed that his South Amer- ican comparative material represented D. virgo and not D. albus. Family NOCTILIONIDAE Noctilio leporinus mastivus (Vahl, 1797) Specimens Examined— BELIZE. Cayo: Banana Bank, 1 <5, 1 9 (fmnh); Barton Creek, at Western Hwy., 1 9 (fmnh). Stann Creek: Melinda, Stann Creek River, 1 6 (fmnh). Toledo: 1.2 km E Agua- cate, Aguacate River, 2 66 (cm), 1 9 (bm); Big Fall, vicinity Rio Grande Bridge, 1 9 (fmnh); Salamanca Camp, 1 9 (bm). The fishing bat occurs along the riparian habi- tats of river systems, inland lakes, and coastlines in primarily lowland regions from northwestern (northern Sinaloa), eastern (southern Veracruz), and peninsular (Yucatan) Mexico throughout Cen- tral America (Davis, 1973;Hellebuycketal., 1985). Dickerman et al. (1981) reported a locality for Noctilio leporinus from Alta Verapaz as in the Ca- ribbean drainage when it was clearly in the Rio Usamacinta drainage of the Gulf lowlands. The Belizean localities extend northward the recorded occurrence of A^. leporinus from Izabal and north- western Honduras (Carter et al., 1966). All of the specimens were obtained (March, April, May, July, August) over rivers and a pond except for one individual, which was mist netted (28 August) low over a pasture adjacent to a flood- ed river. This bat was foraging primarily for insects since its feces contained the chitinous remains of these prey. Additional fishing bats from the lo- calities in Cayo and Stann Creek districts were captured, banded, and released. This bat was com- mon along the South Stann Creek drainage. Cocks- comb Basin. A specimen belonging to M. Craig, Belize Audubon Society, was collected at Indian Church (Lamanai), New River Lagoon, Orange Walk District. An old specimen of M leporinus in the collections of British Museum (Natural His- tory) was registered in 1 909 without pertinent field data. The two peninsular records from Campeche (Jones et al., 1973) and Yucatan (Bimey et al., 1974) were obtained in coastline habitat along the 140 FIELDIANA: ZOOLOGY Gulf of Mexico. Although subsurface drainage predominates north of Belize into Quintana Roo, shallow inland "lagunas" are fairly common and probably support Noctilio populations. Family MORMOOPIDAE Pteronotus davyi fulvus (Thomas, 1 892) Specimens Examined— BELIZE. Cayo: Central Farm, 1 <5 (cm), 1 $ (fmnh); Ontario, 5.5 km W Teakettle, 1 S (fmnh); Unitedville, 9 km WSW Teakettle, 1 <5 (fmnh). Orange Walk: Tower Hill, B.S.I, compound, 3 99 (fmnh). Toledo: Aguacate, 1 (3 (cm); 1.2 km E Aguacate, 1 $ (bm), 1 $ (cm); Rice Station, 2 6$ (fmnh); 0.4 km W Rice Station, 1 $ (fmnh); San Antonio, 1 $ (fmnh). GUATE- MALA. El Peten: Parque Nacional Tikal, 1 $ (msu). Smith (1972) summarized the majority of the capture localities for this subspecies of naked- backed bat, which ranges from northwestern (So- nora), northeastern (Tamaulipas), and peninsular (Yucatan) Mexico southeastward to Honduras and El Salvador, but omitted the only record for Belize (Disney, 1968). Parque Nacional Tikal is the first record for El Peten, and the Belizean specimens provide additional records for Belize. Disney ( 1 968) did not present data for his single Pteronotus davyi specimen. This male was ob- tained ( 1 November) in Cayo District, at Listowel along the Belize River, and is housed in British Museum (Natural History). The subsequent spec- imens reported here were collected (October-De- cember, May, July, August) in open areas, bor- dering on vegetation and buildings, and over water. The specimen from El Peten was captured (25 March) along a trail in upland deciduous forest. An additional P. davyi from Tikal was captured, banded, and released. Pteronotus personatus psilotis (Dobson, 1878) Specimens Examined— BELIZE. Toledo: 1 .2 km E Aguacate, Aguacate River, 1 9 (bm), 4 33, 1 9 (CM); Big Fall, 1.5 km WSW Rio Grande Bridge, 1 $ (fmnh); 0.8 km NW Blue Creek, 1 9 (fsm); Crique Jute, 1 $ (fmnh); Crique Lagarto, 1 km NW San Antonio, 1 9 (fmnh); Jacinto Creek, at Punta Gorda Road, 1 3 (msu); 0.4 km W Rice Station, 1 9 (fmnh); Salamanca Camp, 1 9 (usnm); San Antonio, 1 9 (fmnh); 0.9 km WNW San Pedro Columbia, 1 9 (fmnh). The distribution of Pteronotus personatus psi- lotis extends from western (southern Sinaloa) and eastern (Tamaulipas) Mexico southeastward to Honduras and El Salvador (Smith, 1972), with Caribbean lowland localities in Campeche (Jones et al., 1973), El Peten (Jones, 1966), and Alta Ver- apaz (Jones, 1966). Elevations range from 123 to 984 m. These localities from southern Belize are the first records for the country. Fifty-three percent of the small moustache bats were collected (March, May, July) over open water; the remainder were foraging (January, April, Au- gust, December) in open areas adjacent to build- ings or corralled cattle. Mormoops megalophylla megalophylla Peters, 1864 Specimens Examined— BELIZE. Belize: 6.6 km N Churchyard, 1 9 (cm). Cayo: 1.6 km NW Au- gustine, Rio Frio, 1 3 (ttu). Stann Creek: Melinda, 1 3 (fmnh). Toledo: Forest Home, 1 3 (msu); Pueb- lo Viejo, 1 3 (fmnh). GUATEMALA. Izabal: 25 km SSW Puerto Barrios, 1 3 (tcwc). The leaf-chinned bat has been reported through- out Mexico, Guatemala, El Salvador, and Hon- duras (Smith, 1972). Davis and Carter (1962), Jones (1966), and Taibel (1977) provided lowland records for El Peten and Alta Verapaz. Elevations range from near sea level to 2270 m. These lo- calities are the first records for Belize and Izabal. Except for one Belizean specimen, which was captured (9 June) in a cave, these leaf-chinned bats were associated (March, April, December) with open areas bordering on forest or orchard edges, including pine savanna. One Mormoops specimen, which was registered into the British Museum (Natural History) collections in 1892, may have been obtained in the vicinity of Belize City. The Guatemalan specimen was collected by D. C. Car- ter. Family PHYLLOSTOMIDAE Subfamily PHYLLOSTOMINAE Micronycteris brachyotis (Dobson, 1878) Specimens Examined— BELIZE. Cayo: 1 km NW Augustine, 2 $S (fmnh). Toledo: Crique Ne- gro, Columbia Forest, 1 3 (bm). The first Middle American specimen of Micro- McCARTHY: DISTRIBUTION OF BATS 141 nycteris brachyotis was initially reported from Nic- aragua as M. syhestris by Goodwin (1946), but was correctly identified by Sanborn (1949). Sub- sequent records are from the Gulf-Caribbean low- lands of southern Veracruz (Medellin L. et al., 1983), Oaxaca (Schaldach, 1964), Chiapas (Davis et al., 1964), and El Peten (Jones, 1966; Rick, 1 968; McCarthy, 1 982); and the Pacific-Caribbean versants of Costa Rica (Howell & Burch, 1974; Starrett, 1976; LaVal & Fitch, 1977) and Panama (Handley, 1966; Fleming etal., 1972; Bonaccorso, 1979). Reported elevations range from 40 to 594 m. The present specimens are the first records of the yellow-throated bat for Belize. Two specimens were captured (29 July) as they exited from a cave into a low deciduous seasonal forest, and a third bat was taken (28 May) along a path in an evergreen seasonal forest. The two specimens from Cayo, which I obtained while as- sisting a histoplasmosis survey, were listed as "M. bardyoiis" in a preliminary report (Quinones et al., 1978, p. 559) and no specific locality data were provided. Micronycteris megalotis mexicana Miller, 1898 Specimens Examined— BELIZE. Corozal: San Antonio, 2 km NW Corozal, 1 6 (fmnh). Orange Walk: San Antonio, Rio Hondo, 2 55, 1 2 (fmnh). Toledo: Aguacate, 1 6 (cm); Big Fall, 2 km E Rio Grande Bridge, 1 2 (bm); Cuevas Creek Bridge, 10 km NW Punta Gorda, 1 5, 1 2 (bm), 1 S (amnh), 1 2 (msu); Nimli Punit, 1 2 (cm); Rocky Run Ranch, 4.8 km NW Punta Gorda, 1 3, 1 2 (bm); Union Camp, 2 22 (bm); Vista Hermosa Ranch, 3.7 km WNW Punta Gorda, 1 2 (cm). GUATEMALA. El Peten: Parque Nacional Tikal, 1 <5 (fmnh). The distribution of this subspecies of big-eared bat extends from western (Jalisco), eastern (south- em Tamaulipas), and peninsular (Yucatan) Mex- ico, along the Pacific coastal and highland regions, to Costa Rica. Gardner et al. ( 1 970) suggested that the southern extent of Micronycteris megalotis mexicana is in the Cordillera Talamanca of Costa Rica. This species has been recorded most often at lowland-moderate elevations, up to 2870 m. Specimens from Isla Cozumel, Quintana Roo, rep- resent the only record for Quintana Roo (Jones et al., 1973). The records of A/, m. mexicana which are reported here are the first for Belize and El Peten. Belizean specimens were obtained (May, July, August, November) in diurnal roost sites (shallow caves and limestone chambers, bridge approach- ments, abandoned rum factory boiler) and col- lected in forest habitats (riparian marsh, evergreen and semi -evergreen, deciduous semi -evergreen, and deciduous seasonal). The Tikal specimen was captured (6 June) roosting in a passageway of an excavation tunnel within a ruin complex. A second juvenile male was captured, banded, and released (29 July) in escobal palm (Cryosophila argentea) forest, 1.9 km SE Tikal Reservoir. Micronycteris nicefori Sanborn, 1 949 Specimen Examined— BELIZE. Toledo: 0.4 km NE Aguacate, 1 2 (fmnh). Handley ( 1 966) documented the first specimens of Micronycteris nicefori north of South America, from Panama. Subsequently, it has been reported from southeastern Nicaragua (Baker & Jones, 1975) and both the dry Pacific (Starrett, 1976) and wet Caribbean (LaVal, 1977) lowlands of Costa Rica. These Central American localities range from near sea level to over 100 m. This first record from Belize also represents a significant Central Amer- ican range extension along the Caribbean versant. The M. nicefori specimen reported here was mist netted on 1 5 December along a track in hilltop, evergreen seasonal forest. Micronycteris schmidtorum Sanborn, 1935 Specimens Examined— BELIZE. Corozal: Pat- chakan, 2 22 (fmnh). Orange Walk: 1 .3 km W San Antonio, Rio Hondo, 1 6 (fmnh). Toledo: Big Fall, 1 km E Rio Grande Bridge, 1 6 (cm). Micronycteris schmidtorum was described (San- bom, 1935) from specimens collected in the Ca- ribbean lowlands of Izabal. An additional Gua- temalan specimen was recorded in the Pacific piedmont (Dickerman et al., 1981). The remaining Central American records represent both the Pa- cific and Caribbean lowland slopes from Honduras (Sanbom, 1941), Nicaragua (Davis et al., 1964; Baker & Jones, 1 975), Costa Rica (Starrett & Case- beer, 1968; Fleming et al., 1972; Howell & Burch, 1974; LaVal & Fitch, 1977), and Panama (Han- dley, 1 966). Specimens from Yucatan assigned to M. schmidtorum by Villa-R. (1966) were reiden- tified as M. megalotis by Jones et al. (1973). An identification of M. schmidtorum (Jones et al., 1 973) for a specimen from Isla Cozumel, Quintana Roo, was questioned by Hall (1981) because this 142 FIELDIANA: ZOOLOGY specimen previously was identified as M. mega- lot is (Jones & Lawlor, 1 965). I examined this spec- imen (University of Kansas 9 1 539) and agree that it is M. schmidtorum. The northern distribution of this big-eared bat extends to the Caribbean coast of the Yucatan Peninsula. The specimens reported here are the first records for Belize. At Parque Nacional Tikal, one juvenile and two adult females, which were captured (30 July) in a hollow tree (Bursera semirouba) of an upland de- ciduous seasonal forest, were photographed, band- ed, and released. This site was revisited during the following March, but no Micronycteris were found. These individuals of M. schmidtorum were the first seen in El Peten. Similarly, Sanborn (1935) and Starrett and Casebeer (1968) reported indi- viduals from tree hollows. The Belizean specimens were captured (February, September, November) in the orchard vegetation of a village, along a sec- ondary forest edge, and in riparian secondary vegetation. Lonchorhina aurita aurita Tomes, 1863 Macrophyllum macrophyllum (Shinz, 1821) Specimens Examined— BELIZE. Cayo: Sibun River at Indian Creek, 1 $ (fmnh). Toledo: Big Fall, 1.7 km NE Rio Grande Bridge, 1 $ (cm). Tabasco, Mexico, represents the northernmost occurrence for the long-legged bat, which is known from both the Caribbean and Pacific regions of Central America. Primarily a lowland inhabitant, Macrophyllum macrophyllum ranges from 40 to almost 600 m. These specimens represent a Ca- ribbean lowlands range extension from north- western Honduras (Valdez «& LaVal, 1971) and the first records for Belize. Harrison and Pendleton (1974), Gardner (1977), and Dickerman et al. (1981) indicated that long- legged bats may be closely associated with aquatic habitats. Similarly, the Belizean specimens were obtained ( 1 7 March, 1 April) from along the Sibun River, although not directly above water, and over the surface of the Rio Grande. The first bat was taken at approximately 0340 in a stand of shade trees, dominated by cohune palms (Orbignya co- hune), at the edge of an open pasture. Specimens Examined— BELIZE. Stann Creek: 5.3 km WNW Quam Bank, Cockscomb Basin, 1 9 (CM). Toledo: 0.8 km NW Blue Creek, 1 3, 1 2 (amnh); Crique Jute Village, 1 9 (cm); Crique Ne- gro, Columbia Forest, 1 S (bm), 1 $ (usnm); 2. 1 km NNE Salamanca Camp, Columbia Forest, 3 66 (CM). GUATEMALA. El Peten: Poptun, Finca Ixobel, 2 66 (cm). Lonchorhina aurita was first recorded in Middle America from Panama (Miller, 1912). Subsequent collecting has found this cave-dwelling bat north- ward through Central America to southeastern (southern Veracruz, Oaxaca, Tabasco) and pen- insular (Quintana Roo) Mexico. Predominately lowland, this distinctive leaf-nosed bat extends up to more than 1500 m in representative habitats. Jones et al. (1973) reported the only record from Quintana Roo, while specimens from Izabal (San- bom, 1936) are apparently the next Caribbean ver- sant record north of eastern Costa Rica (Nelson, 1965); records from Nicaragua and Honduras are lacking. The specimens examined for this account are the first records from Belize and El Peten. All specimens from Belize were captured (March, April, May, August) in deciduous seasonal and evergreen seasonal forests. The Guatemalan bats were captured by N. A. Bitar as they exited from the cave discussed in the Balantiopteryx io ac- count. Tonatia bidens bidens (Spix, 1823) Specimens Examined— BELIZE. Cayo: Rio Frio, 1 .6 km W Augustine, 1 9 (cm). Toledo: Nimli Punit, 1 6 (cm); Orange Creek, 1.5 km SW Punta Gorda, 1 6 (msu); 2. 1 km NNE Salamanca Camp, Columbia Forest, 1 6 (cm); 2.2 km NNE Sala- manca Camp, Columbia Forest, 1 9 (cm). Goodwin (1946) first recorded Tonatia bidens in Central America from the Pacific lowlands of Costa Rica. Other humid lowland records include both the Caribbean and Pacific versants of Pan- ama, continuing along the Caribbean corridor of Nicaragua, Honduras, and Guatemala. The north- ernmost record is from eastern Chiapas (Medellin L., 1983). The Guatemalan records are from the Caribbean lowlands of El Peten (McCarthy, 1982) and Izabal (Carter et al., 1966). Elevations range from near sea level to around 660 m. The present specimens constitute the first records from Belize. Four adult males were taken (March, April) over a creek in a low transitional forest, in a high ev- ergreen seasonal forest, and in a deciduous sea- sonal forest. A subadult male was captured (24 September) in the courtyard of a Mayan archae- ological site located in a high deciduous seasonal forest. MCCARTHY: DISTRIBUTION OF BATS 143 Tonatia evotis Davis and Carter, 1978 Specimen Examined- GUATEMALA. El Pe- ten: Parque Nacional Tikal, 1 6 (fmnh). Davis and Carter ( 1 978) described Tonatia evo- tis on the basis of its smaller size in comparison to T. sylvicola; a female from Izabal was designated as the holotype. El Peten is part of a Gulf-Carib- bean distribution which extends from southern Veracruz, Tabasco, Chiapas, and Campeche to Be- lize, and continues along northern Honduras (Da- vis & Carter, 1978). Martinez R. (1980) recorded an additional eastern Guatemalan locality in Aha Verapaz. All recorded elevations are less than 100 m. The T. evotis from Tikal represents the first record for El Peten. Two adult males and one pregnant female were mist netted (20 February, 29 and 25 March) in Tikal along the Uaxactun Road, at a permanent water pool in escobal palm forest, and in an upland deciduous seasonal forest. One male and the fe- male were banded and released. Tonatia minuta Goodwin, 1 942 Specimens Examined— BELIZE. Cayo: 1.1 km W Augustine, 1 2 (fmnh); Central Farm, at Belize River, 1 9 (fmnh); 1.2 km E Macaw Bank, 1 2 (fmnh). Toledo: Big Fall, 1.7 km NE Rio Grande Bridge, 1 2 (msu); San Lucas, 1 2 (msu). This small Tonatia was originally described from the Caribbean coast of Nicaragua as T. nicaraguae (Goodwin, 1942a). Its Middle American distri- bution is lowland ( 1 5 to 6 1 m) along Caribbean and Pacific versants, from southern Veracruz (Lackey, 1 970) to El Peten, Guatemala (McCarthy, 1982) and Belize (Disney, 1968), continuing through Honduras (LaVal, 1969; Valdez & LaVal, 1971; Greenbaum & Jones, 1978), Nicaragua (Jones et al., 1971; Greenbaum & Jones, 1978), and Costa Rica (Gardner et al., 1 970; LaVal, 1 977), to Panama (Davis et al., 1964; Handley, 1966). This account represents additional records for the small round-eared bat in Belize. Disney (1968) reported no data for the first To- natia minuta specimen from Belize, which was a female collected (25 November) in Cayo District, at Listowel, along the Belize River. This specimen was deposited in British Museum (Natural His- tory). The additional specimens reported here were captured (November, January, February, April, May) over rivers or in a deciduous seasonal forest. The name minuta is applied in accordance with the discussion by McCarthy ( 1 982). Gardner ( 1 976) referred to a personal communication with C. O. Handley, Jr., who suggested that all small Tonatia (including minuta) represent a single species, T. brasiliense. Because the taxonomy is poorly under- stood, a systematic review of this group would be useful. Mimon cozumelae Goldman, 1914 Specimens Examined— BELIZE. Belize: Churchyard, Sibun River, 1 2 (fmnh). Cayo: "Mountain Pine Ridge", 2 33, 1 2 (bm); 0.8 km W Augustine, 1 6 (cm); 1 km NW Augustine, 2 $6 (fmnh); Barton Creek, at Western Hwy., 2 $S, 3 22 (fmnh). Toledo: vicinity Aguacate, 2 $S, 2 22 (cm), 1 (5 (fmnh); Crique Negro, Columbia Forest, 1 2 (bm); Pueblo Viejo, 1 3, 1 2 (fmnh); 2. 1 km NNE Salamanca Camp, Columbia Forest, 2 6S (cm); 2.2 km NNE Salamanca Camp, Columbia Forest, 1 <5 (cm); vicinity Union Camp, 2 5(5, 1 2 (bm), 2 22 (cm). This spear-nosed bat ranges from southeastern (northern Oaxaca, southern Veracruz) and pen- insular (Yucatan, Quintana Roo) Mexico south- eastward along the humid Caribbean side of Cen- tral America. Specimens from Isla Cozumel, Quintana Roo, provided the original description for this species (Goldman, 1914). Recorded ele- vations extend to 495 m. The Belizean localities reported here are the first records for the country. Mimon cozumelae were collected (January, March, May, July, August, September, December) along the edge of deciduous and semi-evergreen seasonal forests bordered with pasture, on riparian flood plains, over rivers, along paths in high de- ciduous, semi-evergreen seasonal forests, and in caves. Schaldach (1964), Villa-R. (1966), and Hall (1981) considered cozumelae a subspecies of ben- nettii. I tentatively accept cozumelae at the specific level. Minion crenulatum keenani Handley, 1 960 Specimens Examined— BELIZE. Cayo: Listow- el, Baking Pot, 1 S (fmnh). Toledo: Crique Negro, Columbia Forest, 1 6 (usnm). There are few records for Mimon crenulatum keenani from Middle America. The distribution of this distinctive spear-nosed bat extends along the Caribbean versant, from Panama (Handley, 144 FIELDIANA: ZOOLOGY 1966; Bonaccorso, 1979), Costa Rica (Gardner et al., 1970; LaVal, 1977), Nicaragua (Greenbaum & Jones, 1978), Belize (Ruiz, 1983), El Peten (McCarthy, 1982), and Campeche (Jones, 1964) to the Gulf lowlands of eastern Chiapas (Medellin L., 1 983). All recorded elevations range below 265 m. These specimens are the second and third rec- ords from Belize. The first record (Ruiz, 1 983) was obtained near Blue Hole, 14 km SE Belmopan, Cayo District. One Mimon crenulatum was captured (8 Oc- tober) in a house after it flew through an open window. The house was situated along the Belize River in an agricultural area. The second specimen was netted (29 March) along a path in evergreen seasonal forest. E. L. Tyson collected the specimen from Toledo District. Phyllostomus discolor verrucosus Elliot, 1905 Specimens Examined— BELIZE. Toledo: Cri- que Lagarto, 1 km NW San Antonio, 1 S (fmnh); 1 km NNE Salamanca Camp, Columbia Forest, 1 (3 (cm). GUATEMALA. Alta Verapaz: Lanquin, Lanquin Cave, approx. 1 49 km WSW Puerto Bar- rios, 1 (5, 1 5 (fmnh). Records of Phyllostomus discolor extend from southern (Oaxaca, Veracruz) Mexico along both the Pacific and Caribbean corridors of Central America. Records are more common at lower el- evations, less than 600 m. The new records from southern Belize provide a limited range extension northward from eastern Izabal (Sanborn, 1936). An adult from Crique Lagarto was captured ( 1 January) along the edge of low secondary forest bordering this settlement. The head of the bat was covered with yellow pollen. The second specimen was netted (21 March) in secondary vegetation, which resulted from slash-bum agriculture. Whit- ish pollen dusted the face, chest, and ventral wing surfaces. A male subadult Phyllostomus discolor that was taken ( 1 3 July) along a fenceline of sec- ondary vegetation between two pastures, 1 .9 km ENE Rio Grande Bridge, Big Fall, Toledo District, was photographed, banded, and released. L. de la Torre apparently captured (3 June) the two Phyl- lostomus from Alta Verapaz inside the entrance of Lanquin Cave. I tentatively follow Jones et al. (1977) in as- signing the specimens of Phyllostomus discolor from the Caribbean lowlands to the subspecies verrucosus. Sanborn (1936, p. 98) recognized ver- rucosus subspecifically, stating the "available mea- surements of rf/5co/or would place them much clos- er to verrucosus."'' He suggested the Panamanian P. d. discolor are assignable to verrucosus based on larger size. Felten (1956) and ^urt and Stirton (1961) concurred with his statement by referring a large series from El Salvador to verrucosus; with the availability of greater series of specimens, Da- vis and Carter (1962) indicated they could not recognize two subspecies of P. discolor in Central America and northern South America, acknowl- edging only P. d. verrucosus. Handley ( 1 966) ap- parently disagreed as he recognized the subspecies discolor in Panama. Multivariate analysis of mor- phological data (Power & Tamsitt, 1 973) suggested this species might be monotypic. Phylloderma stenops septentrionalis Goodwin, 1940 Specimens Examined— BELIZE. Toledo: Cri- que Negro, Columbia Forest, 1 2 (usnm); 2. 1 km NNE Salamanca Camp, Columbia Forest, 2 $S (CM). This rarely encountered species has been re- corded north of Panama from the Caribbean coast of Costa Rica (LaVal, 1977), the highlands of Hon- duras (Goodwin, 1940), the Caribbean lowlands of Guatemala (McCarthy, 1982), and the Gulf lowlands of Chiapas (Carter et al., 1966). Limited elevational data are from lowland to approxi- mately 1320 m. The specimens oi Phylloderma stenops from Belize represent the eighth, ninth, and tenth specimens north of Panama and the first records from Belize. All specimens were mist netted (March, Decem- ber) in similar evergreen seasonal forest habitats. E. L. Tyson collected the specimen from Crique Negro. Handley ( 1 966) regarded the Panamanian spec- imens to be Phylloderma stenops stenops, and those from northward into Middle America were thought to be subspecifically different from the nominal species. LaVal (1977) did not designate a subspe- cies for his Costa Rican specimen. Trachops cirrhosus coflini Goldman, 1925 Specimens Examined— BELIZE. Orange Walk: Richmond Hill (Goat Hill), 8.9 km SSW Orange Walk Town, 1 3, 1 $ (cm). Toledo: 2.2 km NNE Salamanca Camp, Columbia Forest, 1 $ (cm). MCCARTHY: DISTRIBUTION OF BATS 145 GUATEMALA. Izabal: 25 km SSW Puerto Bar- rios, 1 $ (tcwc). This lowland subspecies of the fringe-lipped bat is recognized from eastern (southern Veracruz) and southeastern (eastern Oaxaca) Mexico southeast- ward to Nicaragua. Recorded elevations are from near sea level to approximately 330 m. Jones (1966), Rick (1968), and McCarthy (1982) pro- vided records for El Peten. The description of this subspecies was based on specimens from eastern El Peten (Goldman, 1925). The first Belizean rec- ords were reported from Belize District in the vi- cinity of Belize City (Sanborn, 1941) and Rock- stone Pond (Pendergast, 1979). The specimen from Izabal is the first record for that Guatemalan de- partment. D. C. Carter obtained the single specimen from Izabal on 19 February. The additional Belizean specimens were mist netted (March, April) in de- ciduous marsh and evergreen forests. Chrotopterus auritus (Peters, 1856) Specimens Examined— BELIZE. Toledo: vicin- ity Crique Negro, Columbia Forest, 1 9 (fmnh); 1.6 km NNE Salamanca Camp, Columbia Forest, 1 9 (fmnh). Chrotopterus was first reported in Central Amer- ica from El Salvador (Burt & Stirton, 1961). Sub- sequently, this carnivorous bat has been recorded from southern (southern Veracruz, northern Oa- xaca, Chiapas) and peninsular (Yucatan, Quintana Roo) Mexico southeastward throughout Central America at lowland and upland elevations (40 to over 1 880 m). Chrotopterus auritus has been re- ported from Quintana Roo (Jones et al., 1 973) and El Peten (Rick, 1968; McCarthy, 1982). These specimens from southern Belize provide the first records for the country. The Belizean specimens were netted (10 April, 28 July) in an evergreen seasonal forest at ground level along a path and at a height of about 13.7m over an intermittent stream bed. Both were active during the morning hours, 0418 and 0330, re- spectively. The subspecific name Chrotopterus auritus au- ritus has been applied to Middle American pop- ulations (Jones et al., 1971). Carter and Dolan (1978) stated the type specimen for Vampyrus au- ritus Peters, 1856, actually was collected in Santa Catarina, Brazil, not in Mexico. The discussion by Carter and Dolan (1978, p. 37) suggested that Pe- ters based his description on one or more speci- mens from Brazil and compared these with a spec- imen from an unrecorded locality in Mexico as the "verwandten Art aus Mexico." Handley ( 1 966) doubted that subspecies were recognizable. Vampyrum spectrum (Linnaeus, 1758) Specimen Examined— BELIZE. Toledo: Santa Elena, 1 S (fmnh). Two localities in southern Veracruz, Mexico (Goldman, 1917; Navarro L., 1 979) are the north- westernmost records of the false vampire bat's Middle American distribution, which continues in Nicaragua (Dobson, 1 878; Allen, 1910), Costa Rica (Casebeer et al., 1963; Armstrong, 1969; Gardner et al., 1970; Howell &. Burch, 1974; Vehrencamp et al., 1977; LaVal & Fitch, 1977), and Panama (Handley, 1966; Peterson & Kirmse, 1969; Bo- naccorso, 1979). Although primarily lowland in distribution, its highest recorded elevation was about 1815m. The occurrence of Vampyrum spec- trum in the Caribbean lowlands of Belize is doc- umented by this specimen. There appears to be no definite record of this carnivorous bat from Guatemala (Jones, 1966). Dobson (1878, p. 471) recorded "Guatemala" as part of the Central American range for Vampyrum, but did not list any examined specimens. Alston (1879-1882, p. 39) stated Dobson (pers. comm.) saw specimens from Guatemala, although Alston realized the collector, O. Salvin, had not obtained specimens of Vampyrum; hence, the identification of this species is doubtful. Five false vampire bats were mist netted on three separate dates in Parque Nacional Tikal, El Peten. Two females were cap- tured during the dry season (22 and 24 March) in an upland deciduous seasonal forest, in the vicin- ity of Central Plaza of the archaeological site, and at a permanent water pool in escobal palm forest, 2.6 km SE Central Plaza. Two females and one male were netted during the wet season (22 July) at a location along an archaeological transect in escobal palm forest, 1 km SE Tikal Reservoir. All of these bats were released after being observed, measured, and/or photographed. These individ- uals provide the first record for Guatemala and, along with the specimen from Belize, bridge an intermittent distribution that now extends north- ward toward peninsular Mexico. The Vampyrum spectrum from Belize was cap- tured (8 April) during the early morning (0300) in 146 FIELDIANA: ZOOLOGY an open field. We were "trapping" Desmodus ro- tundus during a vampire bat control effort in the village. This large bat was captured after it made a number of low passes over horses and mules, which were encircled by mist nets. The bat died while enroute to captivity via an assistant. The Central American population of Vampy- rum was described as a distinct subspecies, V. s. nelsoni (Goldman, 1914), but Handley (1966) ar- gued that the species was monotypic. Subfamily GLOSSOPHAGINAE Glossophaga commissarisi commissarisi Gardner, 1962 Specimens Examined— BELIZE. Belize: Rock- stone Pond, 2 SS, 3 99 (rom). Toledo: Aguacate, 1 9 (fmnh), 1 9 (cm); Big Fall, 1 km SE Rio Grande Bridge, 2 $S (cm); Forest Home, 1 9 (fmnh); 2.8 km NNW Punta Gorda, 1 9 (fmnh). GUATE- MALA. Izabal: 25 km SSW Puerto Barrios, 7 SS, 6 99 (tcwc). Webster and Jones (1982) summarized the dis- tribution for this subspecies of nectivorous bat, which was documented from eastern (Veracruz) and southern (Oaxaca, Chiapas) Mexico and southern Belize southeastward throughout Central America. Hellebuyck et al. (1985) recently re- ported records from El Salvador. The specimens from Izabal are the first records from this Gua- temalan department. The specimens from Belize District extend northward the distribution of Glos- sophaga commissarisi along the Caribbean low- lands. According to D. C. Carter's field notes, the ma- jority of the Guatemalan Glossophaga commis- sarisi were mist netted (February, March) over a stream and in the adjacent undisturbed forest. Many of these nectivorous bats were captured in association with night-blooming "bat flowers" bordering on a stream. The Belizean specimens reported (Webster & Jones, 1982) from Lubaan- tun, Toledo District, were collected ( 1 8 April) in a disturbed semi-evergreen seasonal forest. Ad- ditional specimens were secured (January, July, September, December) in secondary and orchard vegetation of villages, in riparian secondary vege- tation, and from the hollow of a mamey tree (Pou- teria mammosa). Subfamily STENODERMATINAE Uroderma bilobatum molaris Davis, 1968 Specimen Examined— MEXICO. Quintana Roo: 2 km N, 8 km W Bacalar, 1 $ (tcwc). Davis (1968) recognized this subspecies of the tent-making bat from the Gulf-Caribbean versant of southern Veracruz, Tabasco, northeast Oaxaca, northern Chiapas, Belize, Honduras, Nicaragua, Costa Rica, and northwest Panama. Disney (1968) and Pendergast (1979) also reported the occur- rence of Uroderma bilobatum from Belize. The specimen reported here represents the first record for Quintana Roo and a marginal range extension into the Mexican peninsula of Yucatan. The above specimen was taken in a net on 6 August by M. D. Engstrom along a path leading to an inland lagoon. Vampyrops helleri helleri Peters, 1866 Specimens Examined— BELIZE. Cayo: Banana Bank, 5 99 (fmnh); 0.8 km W Macaw Bank, 1 6 (fmnh). Toledo: Big Fall, 1.9 km ENE Rio Grande Bridge, 1 9 (amnh), 1 9 (cm), 1 $ (msu); Crique Negro, Columbia Forest, 1 $ (bm); Forest Home, 1 (5 (fmnh), 1 (5 (msu); Salamanca Camp, 1 S (bm), 1 (5 (fmnh), 1 9 (usnm); 1.8 km NNE Salamanca Camp, Columbia Forest, 1 9 (fmnh); vicinity Union Camp, 1 9 (bm), 2 99 (cm). The Middle American records of this fruit bat indicate a distribution from sea level to elevations of over 1 300 m and a range from southeastern Mexico (southern Veracruz, Oaxaca, Tabasco) throughout Central America. Lowland records have been reported from El Peten (Rick, 1968) and Izabal (Carter et al., 1966). This account con- stitutes the first records from Belize. Eighty-seven percent of the Vampyrops helleri specimens were captured along or in proximity to waterways. Eleven additional individuals were re- leased at Banana Bank, where a concentration of stenodermatines (Sturnira, Uroderma, Vampyres- sa, Chiroderma, Artibeus, and Vampyrops) was observed. The remaining localities were in upland evergreen seasonal forest and in disturbed village vegetation. A specimen in the collection of St. John's College, Belize City, was collected by E. L. Tyson in Columbia Forest. I follow Dickerman et al. (1981) for the taxo- nomic assignment of the subspecific epithet. MCCARTHY: DISTRIBUTION OF BATS 147 Vampyrodes caraccioli major G. M. Allen, 1908 Specimens Examined— BELIZE. Toledo: Agua- cate, 1 (5 (CM); Big Fall, 1 .9 km ENE Rio Grande Bridge, 1 S (cm), 1 6 (fmnh); Big Fall, 2.1 km E Rio Grande Bridge, 1 S (bm); Crique Negro, Co- lumbia Forest, 1 S (bm), 1 5, 1 9 (msu); Salamanca Camp, 1 S (usnm); 1.6 km N Salamanca Camp, Columbia Forest, 1 S (fmnh); 2. 1 km NNE Sala- manca Camp, Columbia Forest, 4 66, I 9 (cm); San Antonio, 1 9 (fmnh). The published distribution of Vampyrodes car- accioli major northwestward of Costa Rica and Panama is confined to the Gulf-Caribbean low- lands as far as southern Mexico (Oaxaca, southern Veracruz, Chiapas); elevational data are less than 300 m. The records from Belize extend the range of this stenodermatine north of Izabal (Sanborn, 1936). The Belizean localities represent habitats of ri- parian lowland and upland evergreen seasonal for- ests and village secondary vegetation. The capture dates cover both the dry and wet seasons (March, April, May, July-September, December). I follow Carter and Dolan (1978) for the correct spelling of Vampyrodes caraccioli. Vampyressa pusilla thyone Thomas, 1 909 Specimens Examined— BELIZE. Cayo: 1.6 km NW Augustine, 3 66, I 9 (cm); Banana Bank, 1 9 (fmnh); Blancaneaux, 8.3 km NNE Augustine, 1 9 (fsm). Toledo: vicinity Aguacate, 1 9 (bm), 3 99 (cm); 1.2 km E Aguacate, 1 3, 1 9 (cm); Big Fall, 1 km E Rio Grande Bridge, 1 9 (cm); Big Fall, 2. 1 km E Rio Grande Bridge, 1 6 (cm); Big Fall, 1 .9 km ENE Rio Grande Bridge, 1 5, 1 9 (cm), 1 9 (fmnh); Crique Negro, Columbia Forest, 1 6 (msu), 1 6 (usnm); Forest Home, 1 6 (msu); Pueblo Viejo, 1 9 (fmnh); 1 .6 km NNE Salamanca Camp, Co- lumbia Forest, 1 3, 2 99 (fmnh). The general distribution of the little yellow-eared bat extends from southern (Oaxaca, southern Ve- racruz, Chiapas) and peninsular (Campeche) Mex- ico and continues southeastward along the Carib- bean slope to both the Pacific and Caribbean corridors of southern Nicaragua, Costa Rica, and Panama, into South America. Elevational data are primarily lowland, from sea level up to a recorded 2200 m. Peterson (1966) reported the only record of Vampyressa pusilla in Belize, from Rockstone Pond, Belize District. There are also previous rec- ords from El Peten (Rick, 1 968) and southeastern Campeche (Jones et al., 1973). This account pro- vides additional records of this species. These specimens of Vampyressa pusilla were collected (February-May, July-September, De- cember) in moist habitats, the majority of which were associated directly with riparian vegetation or in village and pasture-edge vegetation situated near rivers. Evergreen seasonal forest provided an upland habitat. Chiroderma villosum jesupi J. A. Allen, 1900 Specimens Examined— BELIZE. Cayo: Banana Bank, 1 3, 5 99 (fmnh). Corozal: Chan Chen, 1 6 (fmnh). Toledo: Big Fall, vicinity Rio Grande Bridge, 1 6 (fmnh); Big Fall, 1 .7 km NE Rio Grande Bridge, 1 9 (msu); Big Fall, 1 .9 km ENE Rio Grande Bridge, 1 3, 1 9 (cm); San Antonio, 1 6 (fmnh); 1 km WNW San Pedro Columbia, 1 9 (fmnh). GUA- TEMALA. EI Peten: Parque Nacional Tikal, 1 6 (fmnh). The Middle American occurrence of Chiroder- ma villosum has been documented in southern (Oaxaca, southern Veracruz, Chiapas) and pen- insular (Campeche, Quintana Roo) Mexico, Gua- temala, Nicaragua, Costa Rica, and Panama. Hel- lebuyck et al. (1985) recently reported this fruit bat from El Salvador. Locality records reach from the coastal lowlands to upland habitats at 1 300 m. Southeastern Campeche (Jones et al., 1973) and northern Quintana Roo (Bimey et al., 1974) are previous Caribbean lowland localities, in addition to these first records from Belize and El Peten. All but one of the Belizean Chiroderma were associated either directly with or in the vicinity of riparian evergreen or semi-evergreen seasonal for- ests (April, May, August, September, December). One individual was captured (15 November) in village orchard vegetation. Five additional indi- viduals were released at Banana Bank. The Tikal specimen was captured (24 March) along the per- manent water pool mentioned in the Tonatia ev- otis account. Artibeus toltecus toltecus (Saussure, 1 860) Specimens Examined— BELIZE. Cayo: vicinity Augustine, 2 66, 4 99 (fsm); 1 .6 km NW Augustine, Rio Frio, 1 3, 1 9 (fmnh), 5 66 (ttu), 4 66 (cm); "Rio On," ? km N Augustine, 1 9 (ttu); 1.1 km S Baldy Beacon, Bald Hills, 3 99 (cm); vicinity San Luis, 7.1 km SSW Augustine, 1 9 (ttu). Toledo: 148 FIELDIANA: ZOOLOGY Orange Point, 1 2 (fmnh); Pueblo Viejo, 3 9$ (fmnh); Union Camp, 5 S6, 4 92 (cm). In his revision of the small Artibeus of Middle America, Davis (1969) recognized the range of Artibeus toltecus toltecus from southern Tamau- lipas, Mexico, southeastward along the mountain- ous region of the Gulf versant, upland of southern Mexico, Guatemala, Honduras, Nicaragua, and Costa Rica. He did not examine Panamanian spec- imens. Handley (1966) summarized the Pana- manian localities for /I. toltecus. This bat primarily occurs at elevations between 328 and 1640 m, although elevations near sea level were recorded (Davis, 1969). Consequently, the occurrence of ^. toltecus in the Maya Mountain range of southern Belize and southeastern El Peten was not unex- pected. These Belizean localities represent the first northern Caribbean lowland records. The Belizean localities range in elevation from near sea level to approximately 720 m. Artibeus toltecus is more common at the higher elevations. These dark-colored Artibeus were captured (De- cember-February, April, June, September) in hab- itats of deciduous seasonal forest, semi-evergreen seasonal forest, transitional forest, and pine forest- savanna. The subspecies toltecus is applied, based on the proximity of Belize to its distribution as defined by Davis (1969). Centurio senex senex Gray, 1 842 Specimens Examined— BELIZE. Belize: 1 .4 km S San Pedro, Ambergris Caye, 1 3, 1 9 (fmnh). Cayo: 1.6 km NW Augustine, Rio Frio, 1 <5 (ttu); vicinity Augustine, Rio On, 1 9 (ttu); Blanca- neaux, 8.3 km NNE Augustine, 1 9 (fsm); Central Farm, 1 3, 1 9 (fmnh); Teakettle, Young Gal Road at Belize River, 1 3, 1 9 (fmnh); Xunantunich, 1 $ (fmnh). Corozal: 1.2 km E, 1.6 km N Corozal, 1 (5 (LSUMZ). Orange Walk: 1.6 km NW San An- tonio, Rio Hondo, 1 9 (fmnh). Toledo: Big Fall, 1 .9 km ENE Rio Grande Bridge, 1 S (cm); Crique Negro, Columbia Forest, 1 5, 1 9 (usnm); Forest Home, 1 9 (amnh); vicinity Union Camp, 2 99 (BM), 1 9 (CM). GUATEMALA. Alta Verapaz: Lan- quin, vicinity Lanquin Cave, approx. 149 km WSW Puerto Barrios, 1 $ (amnh). Izabal: 25 km SSW Puerto Barrios, 1 3, 5 99 (tcwc). The recorded distribution of the wrinkle-faced bat extends from western (southern Sinaloa), northeastern (southern Tamaulipas), and penin- sular (Campeche and Quintana Roo) Mexico and continues southeastward through Central America at principally lower to upland elevations (sea level to 1882 m). The records given here are the first for Belize, Alta Verapaz, and Izabal. The distribution of this unusual bat in Belize reflects apparent ecological flexibility. Centurio se- nex has been captured in low littoral forest and mangrove swamp edge on the coastal sand strip of Ambergris Caye, to about 720 m in evergreen and semi-evergreen seasonal forest on the south- ern slope of the Maya Mountains. Evergreen sea- sonal and transitional forests, secondary forest, and agriculturally disturbed areas provide additional habitats. This bat was captured throughout the year. Two males and one female were mist netted and released at Orange Point, Toledo District. Brother N. Sullivan collected (15-17 January) the specimen from Alta Verapaz, but I assume the bat was captured outside of Lanquin Cave. The spec- imens from Izabal were obtained (February, March) by D. C. Carter and field party. Field data are limited, but four Centurio were captured over a stream. Diphylla ecaudata Spix, 1823 Specimens Examined— BELIZE. Cayo: vicinity Augustine, 1 S (rom); San Antonio, 1 6 (fmnh). Toledo: Crique Jute, 1 $ (amnh); San Antonio, 1 9 (fmnh); Santa Elena, 1 9 (fmnh). The distribution of Diphylla ecaudata appears primarily restricted along the Gulf side and in the Yucatan Peninsula of Mexico southeastward throughout Central America, where this bat occurs from the coastal lowlands up into the mountainous highlands (1880 m). The hairy-legged vampire bat has been recorded from El Peten (McCarthy, 1 982) and Quintana Roo (Jones et al., 1973). The spec- imens reported here are the first records from Be- lize. Four of the localities represent village environ- ments where Diphylla was captured (April, July, August, December) along with Desmodus rotundus during vampire bat control activities. Mist netting was carried out in direct immediacy to domestic livestock and homes. The feeding activities of £>/- phylla in these villages were not documented, al- though one blood meal was obtained for analysis. P. Boreham, Imperial College Field Station, En- gland, reported (in litt.) a weak precipitin reaction for a mammal host from the blood meal sample without a response for bird or reptile. It is not known if this blood meal was obtained in the vil- McCARTHY: DISTRIBUTION OF BATS 149 lage (Santa Elena). Gardner (1977) summarized the sanguivorous preference ofDiphylla as for pri- marily avian hosts. The hairy-legged vampire from Augustine was apparently taken (22 February) in a deciduous seasonal forest. Family NATALIDAE Natalus stramineus saturatus Dalquestand Hall, 1949 Specimens Examined— BELIZE. Cayo: 1.6 km NW Augustine, Rio Frio, 2S6,2 9i (fsm); 0.8 km W Augustine, 2 66, 3 22 (cm); 1.5 km N Augustine, 5 22 (cm); Sibun Camp, Hummingbird Hwy. at Silver Creek, 1 2 (fmnh). Orange Walk: Richmond Hill (Goat Hill), 8.9 km SSW Orange Walk Town, 1 2 (cm). Stann Creek: Kendal, 1 6 (fmnh). Toledo: vicinity Aguacate, 1 3, 3 22 (cm); 1.2 km E Agua- cate, 1 2 (cm); Vista Hermosa, 3.7 km WNW Punta Gorda, 8 66, 6 22 (fmnh). The northern range of Natalus stramineus sa- turatus extends from both northwestern (Sinaloa) and northeastern (Nuevo Leon) Mexico, including the Yucatan Peninsula, southeastward through Central America where the number of records for this species is noticeably reduced beyond Guate- mala to Panama. Although predominately a low- land species, elevations were recorded as high as 2400 m. The presence of the funnel-eared bat in Belize was anticipated, as it appears to be well reported throughout the Gulf-Caribbean versant. Those specimens obtained (April, August, Sep- tember) at roost sites in Belize were from caves. Other capture localities include low riparian forest and open areas bordering on forest, in orchard habitats, and alongside a building. Family VESPERTILIONIDAE Subfamily VESPERTILIONINAE Myotis elegans Hall, 1962 Specimens Examined— BELIZE. Belize: Belize City, Landivar, 1 2 (amnh), 1 3, 1 2 (fmnh), 1 6 (msu); Mussel Creek, 7.5 km W Burrell Boom, 1 6, 1 2 (fmnh). LaVal (1973a) summarized the lowland distri- bution of Myotis elegans. ranging from the Gulf (eastern San Luis Potosi, Veracruz), Pacific coastal (Chiapas), and peninsular (southeastern Cam- peche) regions of Mexico to Honduras, Nicaragua, and northeastern Costa Rica. Subsequent records were reported from the Pacific side of Costa Rica and the Caribbean lowlands of El Peten (LaVal, 1977; McCarthy, 1982). The majority of eleva- tions are less than 1 20 m, ranging to 750 m. These additional Caribbean lowland localities are the first records from Belize. Two elegant Myotis were netted (1 July) along a tractor track, in low riparian vegetation domi- nated by bamboo and thistle palms. Four indi- viduals were obtained (January, February, May, December) at a coastal locality in low vegetation bordering on disturbed mangrove (Rhizophora mangle, Avicennia germinans) habitat. Eptesicus furinalis gaumeri (J. A. Allen, 1897) Specimens Examined— BELIZE. Belize: Belize City, Landivar, 1 2 (cm). Cayo: Central Farm, 2 22 (cm), 5 66, 16 22 (fmnh), 2 66,19 (TTu); Little Vaquero Creek, 9.3 km NNW Augustine, 1 5, 1 2 (fsm); Ontario, 5.5 km W Teakettle, 1 2 (fmnh); Teakettle, 1 6 (fmnh). Corozal: Estero Lagoon, 4 km W Patchakan, 1 3, 1 2 (fmnh); Santa Clara, 1 2 (fmnh). Orange Walk: Honey Camp Lagoon, 1 6, 2 22 (fmnh); Tower Hill, B.S.I, compound, 3 66 (CM), 1 5, 4 22 (fmnh); 2 km SSW Tower Hill Bridge, 1 2 (cm). Stann Creek: Melinda, 3 22 (fmnh); Dangriga (Stann Creek), 1 <5 (usnm). Toledo: Or- ange Creek, 1.5 km S Punta Gorda, 1 2 (msu); Punta Gorda, 1 2 (msu). The Mexican distribution of Eptesicus furinalis gaumeri ranges from the western (Jalisco) and the eastern (San Luis Potosi) versants southeastward to South America. Davis (1965), Disney (1968), and Starrett and Casebeer (1968) reported records from all of the Central American countries except El Salvador. Lowland elevations range from near sea level to 1800 m, the majority being below 500 m. This tropical brown bat has been reported from El Peten (Rick, 1968; McCarthy, 1982) and Quin- tana Roo (Jones et al., 1973). The localities here are additional records for Belize. Disney ( 1 968) did not present locality data for his two specimens of Eptesicus furinalis. Both were males, captured (16 November, 29 December) in Cayo District, near Central Farm and Esperanza (4.5 km W Central Farm). These are located in British Museum (Natural History). An additional 1 96 individuals were captured from three of the localities reported here; the majority of these were ISO FIELDIANA: ZOOLOGY banded and released during a behavioral study. The majority was found in direct association with buildings, utilizing the infrastructure of the walls or floors and the space behind window shutters as roost sites. Individuals have been taken over water (creeks and a swimming pool) at three localities and in riparian vegetation along two lagoons. Lasiurus borealis (Muller, 1776) Speomens ExAMnsfED— BELIZE. Orange Walk: Tower Hill, B.S.I, compound, 1 9 (fmnh). Stann Creek: 5.3 km NNW Quam Bank, Cockscomb Basin, 1 9 (cm). GUATEMALA. El Peten: Parque Nacional Tikal, 1 9 (fmnh). The subspecies teliotis ranges southward from both the western and eastern regions of Mexico to Oaxaca and the northern Yucatan Peninsula. Specimens of Lasiurus borealis from the Guate- malan central highlands were assigned by Jones (1966) to the Central American subspeciesyra«/z/7, based on Handley (1960). Carter et al. (1966) as- signed specimens from both lowland and highland localities in Chiapas Xofrantzii, suggesting that the region of the Isthmus of Tehuantepec represents the break heXwecn front zii and teliotis. Hall (1981) concurred with this arrangement. Similarly, Jones et al. (1973) suggested that southern Mexico, in- cluding the Yucatan Peninsula, may represent a zone of intergradation between frantzii and teli- otis. Few specimens of L. borealis are available from El Salvador (Burt & Stirton, 1961), Honduras (Goodwin, 1942b), Nicaragua (Davis & Carter, 1962— as L. b. teliotis), Costa Rica (Goodwin, 1 946; Gardner et al., 1 970) and Panama (Handley, 1 966). Recorded elevations (near sea level to about 2540 m) are primarily low or moderate (< 1 155 m). Koopman (1959) reported the only record from Quintana Roo. This account represents the first records for Belize and eastern Guatemala from El Peten. The red bats captured in Belize (April, May) were netted over a stream and a swimming pool. The Tikal specimen was taken (30 July) while it was flying in an open area near a large man-made reservoir. I hesitate to assign a subspecific designation be- cause I see no practical purpose in doing so until adequate series of specimens from throughout the range of Lasiurus borealis become available. Han- dley (1960) had fewer specimens of L. borealis at hand for a proper evaluation of subspecific vari- ation. Consequently, the limits of the distributions for the recognized subspecies remain unresolved. Lasiurus ega (Gervais, 1855) Specimens Examined— BELIZE. Belize: Trop- ical Park, Mi. 14.5 Western Hwy., 1 S (fmnh). Orange Walk: Tower Hill, B.S.I, compound, 2 6i (FMNH), 1 (3 (cm). Stann Creek: 5.3 km WNW Quam Bank, Cockscomb Basin, 2 66, I 9 (cm). Toledo: Big Fall, 1.7 km NE Rio Grande Bridge, 1 9 (cm); Orange Creek, 1 .5 km SW Punta Gorda, 1 6 (msu). Similar to Lasiurus borealis, the distribution for the two recognized subspecies of the yellowish bat is not well understood. While L. e. panamensis was recognized along the Pacific versant of Chia- pas (Baker &. Patton, 1 967) and Guatemala (Dolan & Carter, 1979;Dickermanetal., 1981), Goodwin ( 1 969) identified panamensis from the moderate elevations of the Gulf drainage in northern Oaxaca and suspected L. e. xanthinus may occur in the drier Pacific portion of that state. Baker et al. ( 1 97 1 ) determined the variation in karyotypes and pelage color of L. ega from near Brownsville, Texas, re- sembled those from eastern coastal and southern Mexico and referred the Texas specimens to L. e. panamens