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 0 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 0 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 0 0 0 0 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 0 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 0 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 0 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' 0 I 3.0-3.5 dP' M^ 0 II 4.5-5.0 dP' M' 0 III 6.2-6.7 P' M' 0 IV 7.9-8.7 P' M* 0 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 0 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 0 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. panamensis. Meanwhile, L. e. xanthinus was rec- ognized in the Yucatan Peninsula (Jones et al., 1973; Bimey et al., 1 974). The yellow bat is poorly represented from the remainder of Central Amer- ica, which includes Honduras (Goodwin, 1942b; LaVal, 1969; Greenbaum &. Jones, 1978), Costa Rica (Goodwin, 1946; Starrett & Casebeer, 1968; Gardner et al., 1970; LaVal & Fitch, 1977), and Panama (Handley, 1966). Where designated, the subspecies panamensis has been applied to these preceding Central American localities, although Hall (1981) did not acknowledge panamensis north of Costa Rica. Elevational data are similar to those for L. borealis. Ingles (1958) reported two L. ega from Quintana Roo. Jones et al. (1973, p. 23) translated Ingles's locality from Spanish as "Puer- to Morelos" when it was actually a collection site only 16 km east of the state border with Yucatan, along the highway from Valladolid (Yucatan) to Puerto Morelos (Quintana Roo). Alvarez and Ra- mirez-P. (1972) cited an additional Caribbean lowland record from southeastern Campeche. This account provides the first L. ega records from Be- lize. Eight yellowish bats were captured (April, May) over streams, a river, and a swimming pool. Another was netted ( 1 8 August) at about 5 m above MCCARTHY: DISTRIBUTION OF BATS 151 the ground while circhng a building located in grass- sedge savanna. Lasiurus intermedius intermedius (H. Allen, 1862) Specimen Examined— BELIZE. Toledo: Crique Jute, 1 9 (cm). The range of this subspecies of the large yellow bat extends southeastward from Mexico to Hon- duras (Handley, 1960; Carter et al., 1966), El Sal- vador (Hellebuyck et al., 1985), and Guatemala (Carter et al., 1 966). Lasiurus intermedius has been recorded in Mexico from the northern Yucatan Peninsula and Chiapas northwestward to Texas along the eastern coast and to Sinaloa on the Pa- cific side. Recorded elevations range from lowland to highland (1620 m) habitats. A single specimen of L. intermedius from northern Quintana Roo (Bimey et al., 1974) provided the only record for that Mexican state. This Belizean specimen rep- resents the first record for the country. The above specimen was obtained on 30 March over the stream Crique Jute surrounded by sec- ondary vegetation. Bauerus dubiaquercus (Van Gelder, 1959) Specimens Examined— BELIZE. Cayo: 1.6 km NW Augustine, Rio Frio, 1 2 (rom). Toledo: 2. 1 km NNE Salamanca Camp, Columbia Forest, 1 6 (CM). The published localities of the rarely encoun- tered Bauerus dubiaquercus are scattered from the Islas Tres Marias (Nayarit), Jalisco, and southern Veracruz in Mexico to eastern Honduras and Cos- ta Rica (Engstrom & Wilson, 1981; Dinerstein, 1985). Mainland elevations range from approxi- mately 460 to 1450 m and appear to represent mid-elevation and montane forest habitats (Pine, 1 966; Pine et al., 1 97 1 ; Engstrom & Wilson, 1981; Dinerstein, 1985). These first occurrences of Bau- erus in Belize extend northward a scattered dis- tribution along the northern Caribbean lowlands in Central America. J. Kamslra and J. Fragoso collected (8 July) one specimen inside the main Rio Frio cave, located in a deciduous seasonal forest at approximately 410m. The second Bauerus was netted (26 March) along an open forestry track in an evergreen forest at about 180 m. Engstrom and Wilson (1981) and Martin and Schmidly (1982) evaluated the taxonomic status of Antrozous (Bauerus) dubiaquercus and con- cluded the chromosomal, cranial, postcranial, and phallic differences between this bat and Antrozous (Antrozous) pallidus were sufficient to recognize Bauerus as a distinct genus. I follow their conclu- sions and agree that the species is monotypic since the mainland sample size that previously was as- signed to A. d. meyeri Pine, 1971, was limited to a total of five specimens representing both sexes. Family MOLOSSIDAE Eumops auripendulus auripendulus (Shaw, 1 800) Specimens Examined— BELIZE. Orange Walk: Orange Walk Town, 1 ? (cm); Tower Hill, B.S.I, compound, 1 9 (fmnh). The recorded distribution of Eumops auripen- dulus auripendulus includes both moist uplands and drier lowland coastal and plateau areas, rang- ing from eastern Oaxaca, Tabasco, Quintana Roo, and Belize, through Guatemala, western Hondu- ras, El Salvador, western Nicaragua, Costa Rica, and Panama, into South America (Eger, 1974; Greenbaum & Jones, 1978). Villa-R. (1956) and Eger (1974), respectively, reported this free-tailed bat from Quintana Roo and Belize (Belize District: Rockstone Pond). This account provides the sec- ond and third records for Belize. The Orange Walk specimen consists of a man- dible and partial skull, which were recovered from an owl {Tyto alba) roost in a church tower. The second specimen was discovered (July) alive by L. G. Hoevers, after it apparently was attacked by a bird. Eumops bonariensis nanus (Miller, 1900) Specimens Examined— BELIZE. Orange Walk: Orange Walk Town, 2 ?? (cm). Eger (1977) summarized the few available Mid- dle American localities for this small mastiff bat, which are limited to southeastern Mexico (south- em Veracruz, Tabasco, Yucatan), eastern Hon- duras, and Panama. These and additional locali- ties in Panama (Dolan & Carter, 1979) and Nicaragua (Hall, 1981) are restricted to coastal lowland environments. This is the first recording of Eumops bonariensis for Belize. 152 HELDIANA: ZOOLOGY Entire specimens oi Eumops bonariensis as yet are unavailable from Belize. Documentation is based on two fragmented sets of maxillary tooth- rows, which were sifted from regurgitated rubble beneath an owl {Tyto alba) roost in a church tower. Moiossus ater nigricans (Miller, 1 902) Specimens Examined— GUATEMALA. Izabal: 25 km SSW Puerto Barrios, 2 33, 4 99 (tcwc). The black mastiff bat is a common inhabitant of roof spaces throughout its lowland Middle American range, from western (Sinaloa) and east- em (Tamaulipas) Mexico southeastward into South America. This species has been reported from the Caribbean lowlands of Quintana Roo (Jones et al., 1973) and Belize (Murie, 1935; Pendergast, 1979). These Moiossus ater from Izabal are the first rec- ord for that department. Apparently, all of these specimens were collected on 1 5 February over a stream by D. C. Carter and his field party. Addendum While this paper was in press, other papers and additional information concerning bats in Belize came to my attention. Two recent papers provide new country records of the glossophagines Li- chonycteris obscura (Hill, 1985) and Hylonycteris underwoodi (McCarthy & Blake, 1987), which increase the known bat fauna to 68 species. Both records are from Toledo District. McCarthy and Blake (1987) also reported the occurrence of the following bats: Rhynchonycteris naso, Saccopteryx bilineata, Balantiopteryx io, Noctilio leporinus, Pteronotus parnellii, Micronyctehs megalotis, M. nicefori, M. schmidtorum, Tonatia evotis, Mimon cozumelae, Phyllostomus discolor, Trachops cir- rhosus, Chrotopterus auritus, Vampyrum spec- trum, Glossophaga soricina, Carollia brevicauda, C. perspicillata, Sturnira lilium, Uroderma bilo- batum, Vampyressa pusilla, Artibeus jamaicensis, A. lituratus, A. phaeotis, A. watsoni, Centurio se- nex, Natalus stramineus, Rhogeessa tumida, Bau- erus dubiaquercus, and Eumops underwoodi. Hill, J. E. 1 985. The status of Lichonycteris degener Miller, 1931 (Chiroptera: Phyllostomidae). Mam- malia, 49(4): 579-582. McCarthy, T. J., and M. Blake. 1 987. Noteworthy bat records from the Maya Mountains Forest Re- serve, Belize. Mammalia, 51(1): 161-164. A bat specimen from Stann Creek was reported (Miller & Allen, 1928, p. 180) as Myotis nigricans nigricans. I examined this specimen at USNM and found it to be an immature Eptesicus furinalis. It is included in this paper under the species account for the latter species. Miller, G. S., Jr., and G. M. Allen. 1928. The American bats of the genera Myotis and Pizonyx. Bulletin of the United States National Museum, 144: 1-218. Silva-Taboada and Koopman (1964, p. 3) re- ported specimens of Tadarida laticaudata (= Nyc- tinomops laticaudatus) from Corozal District. Most of the bat species discussed in an unpublished dis- sertation by A. M. Cartwright were also reported by Kirkpatrick et al. (1975) and Cartwright and Kirkpatrick (1977). The remaining identifications (Cartwright, 1977, pp. 240, 242-246, 250, 251), which were from Belize District, included Rhyn- chonycteris naso, Saccopteryx bilineata, Carollia brevicauda, C. perspicillata, Sturnira lilium, Arti- beus lituratus, A. phaeotis, Des modus rotundus, Eptesicus furinalis, Rhogeessa tumida, and Mo- iossus moiossus. Baker et al. (1985, p. 236) re- ported cytogenetic data from specimens of Rho- geessa tumida that I collected in Belize District. Baker, R. J., J. W. Bickham, and M. L. Arnold. 1985. Chromosomal evolution in Rhogeessa (Chi- roptera: Vespertilionidae): Possible speciation by centric fusions. Evolution, 39(2): 233-243. Cartwright, A. M. F. 1977. Patterns of Neotrop- ical chiropteran reproduction including histological and ecological aspects of bats collected in Belize. Ed.D. diss.. Ball State University, Muncie, Ind., 278 pp. Silva-Taboada, G., and K. F. Koopman. 1964. Notes on the occurrence and ecology of Tadarida laticaudata yucatanica in eastern Cuba. American Museum Novitates, 2175: 1-6. A mammal checklist was included in a resource paper on Belize (Hartshorn et al., 1984). The list of bats supposedly was a compilation of known and expected species. The result is inaccurate and undocumented. The reader is referred to the checklist in the Appendix as correct. Hartshorn, G., et al. 1984. Belize. Country En- vironmental Profile. A Field Study. United States MCCARTHY: DISTRIBUTION OF BATS 153 Agency for International I>evelopment, San Jose, Costa Rica. 151 pp. Additional specimens of Micronycteris mega- lotis (2: Belize District, Cayo District), Artibeiis toltecus (11: Cayo District), Centurio senex (2: Cayo District), and Diphylla ecaudata (3: Cayo District) were found in the mammal collection of Royal Ontario Museum. D. J. Tallman collected speci- mens of M. megalotis ( 1 ) and Mimon cozumelae (2) from Orange Walk District, which were de- posited in Bell Museum of Natural History, Uni- versity of Minnesota. I secured further voucher specimens (AMNH) of Mimon cozumelae (1), Vampyrodes caraccioli ( 1 ), Vampyressa pusilla ( 1 ), and Bauerus dubiaquercus (1) from Toledo Dis- trict. Certain specimens (Sturnira lilium and Rho- geessa tumidd) that were catalogued by Dobson (1878, pp. 540, 246) were listed as collected in "Honduras." These were obtained by D. Dyson and H. Cuming between November 1844 and late 1845. During that time, "Honduras" correspond- ed to the present region that extends from southern Quintana Roo, Mexico, southeastward to northern Honduras. Many early collectors did not differ- entiate between the area of Belize ("British settle- ment in Honduras") and that of the Republic of Honduras ("Spanish Honduras"), but recorded only "Honduras" or "Bay of Honduras" without further locality data. The above specimens did not originate from present day Honduras, but were collected in Belize. Additional specimens of Mi- cronycteris megalotis (Dobson, 1 878, p. 479) from the "Bay of Honduras" and Rhynchonycteris naso (Dobson, 1878, p. 368) from "Honduras" remain orphaned records of the historical literature. Uroderma bilobatum was reported (Sanchez-H. et al., 1986) from southern Quintana Roo while this volume was delayed. Ten specimens were col- lected at Ruinas de Kohunlich (18°23'N; 88''42'W), about 16 km W Estevez on the Belizean border. Four other species {Pteronotus davyi, Mormoops megalophylla, Tonatia evotis, T. minuta) were documented for the first time from Quintana Roo, from localities within 35 km of the northern bor- der of Belize. The known bat fauna of Quintana Roo is now represented by 36 species. Sanchez-H., O., G. Tell£z-G., R. A. Medelun, and G. Urbano-V. 1986. New records of mammals from Quintana Roo. Mexico. Mammalia, 50(2): 275- 278. Acknowledgments Grateful acknowledgments are due H. Flowers, O. Rosado, R. Belisle, and E. O. Bradley, E>e- partment of Forestry, Belize, and H. Topsey, W. Branche, E. A. Graham, and J. Palacio, Depart- ment of Archaeology, Belize, for the necessary per- mission to conduct fieldwork. The Department of Agriculture, Belize, provided opportunities and support while I headed the Vampire Bat Education and Control Program (1975-1978). Division of Mammals, Field Museum of Natural History, pro- vided partial equipment needs (1974) and partial funding ( 1979). Mellon North American Mammal Research Institute, O'Neil Fund, and Section of Mammals, Carnegie Museum of Natural History, supported further fieldwork (1982, 1984). Nu- merous colleagues and friends have been of direct assistance to my needs in Belize, and the following are but a few: E. Awe, P. Boreham, H. Bums, W. D. Burton, J. Cab, G. Cal, G. C. Canto, J. Chun, M. Craig, L. de la Torre, L. Dieckman, R. Foster, P. W. Freeman, P. Gamble, E. Garcia, J. P. and M. Garcia, H. H. Genoways, E. Gomez, L. Hen- derson, R. W. Henderson, L. G. and E. Hoevers, A. M. Hutson, R. J. Izor, T. Juring, E. King, K. Leacock, D. Owen-Lewis, N. MacKenzie, G. T. McCarthy, H. Pastor, R. H. Pine, A. Rabinowitz, M. L. Reed, C. J. Rushin, K. and T. Salisbury, R. Schmidt, B. M. Silva, M. F. Valentine, L. Waight, L. Wilkins, and A. C. S. Wright. Various regiments and the Royal Air Force of Her Majesty's British Forces provided certain logistical support over the years. My fieldwork in Parque Nacional Tikal, El Pe- ten, Guatemala, was made possible while assisting D. J. Howell and J. G. Cant during their respective visits to the park. M. Dary-R., Universidad de San Carlos, and F. Polo-Sifontes, Instituto de An- tropologia y Historia, graciously assisted and per- mitted my work in 1979. J. R. Martinez R. kindly provided a copy of his thesis. The courtesies rendered to me by the personnel of the following institutions while providing either information, the loan of specimens, or assistance during visits are greatly appreciated: American Museum of Natural History; British Museum (Natural History); Carnegie Museum of Natural History; Field Museum of Natural History; Flor- ida State Museum, University of Florida; Museum of Natural History, University of Kansas; Mu- seum of Zoology, Louisiana State University; The Museum, Michigan State University; Royal On- tario Museum; Texas Cooperative Wildlife Col- 154 HELDIANA: ZOOLOGY lection, Texas A «& M University; The Museum, Texas Tech University; United States National Museum of Natural History. I am grateful to N. A. Bitar, F. J. Bonaccorso, D. C. Carter, M. D. Engstrom, A. M. Hutson, J. Kamstra, R. L. Pe- terson, and E. L. Tyson for sharing information concerning their field efforts in either Belize, El Peten, Izabal, or Quintana Roo. Besides J. R. Choate, J. K. Jones, Jr., K. F. Koopman, and R. M. Timm, two anonymous re- viewers enhanced earlier manuscripts with their comments and editorial skills. E. Mendez kindly reviewed my Spanish abstract. Verification of the state boundaries in the Yucatan Peninsula was provided by T. Dachtera, National Geographic Society, and G. de la Torre M., Secretaria de Pro- gramacion y Presupuesto, Mexico. M. A. Schmidt, R. A. Rollin, V. Risoli, and S. Cramer typed var- ious versions of the manuscript. S. M. Velez guid- ed the manuscript during my absence. This report is a contribution of the Mammals of Belize Pro- gram. Literature Cited Allen, J. A. 1910. Additional mammals from Nica- ragua. Bulletin of the American Museum of Natural History, 28:87-115. Alston, E. R. 1879-1882. Biologia Centrali-Ameri- cana. Mammalia. Taylor and Francis, London, xx + 220 pp., 22 pis. Alvarez, T., and J. Ramirez-P. 1972. Notas acerca de murcielagos mexicanos. Anales de la Escuela de Ciencias Biologicas, Mexico, 19: 167-178. Andersen, K. 1908. 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Handbook of Middle American Indians, vol. 1. University of Texas Press, Austin, 570 pp. Wright, A. C. S., D. H. Romney, R. H. Arbuckle, and V. E. Vial. 1959. Land in British Honduras. Report of the British Honduras Land Use Survey Team. Co- lonial Research Publication, London, 24: 1-327. Gazetteer The numbers for localities are plotted in Figure 1 . MEXICO QUINTANA ROO GUATEMALA El PETfeN 1. Bacalar 18°43'N; 88°22'W 2. Parque Nacional Tikal 1 7°20'N; 89°39'W 3. Poptun 16°2rN; 89''26'W 158 FIELDIANA: ZOOLOGY Fig. 1. Distribution of localities in the districts of Belize; Quintana Roo, Mexico; and El Peten and Izabal, Guatemala. The numbers refer to those listed in the Gazetteer. This map does not display the Caribbean lowlands of Guatemala and Mexico in their entirety. MCCARTHY: DISTRIBUTION OF BATS 159 IZABAL 4. Puerto Barrios 15°43'N; 88°36'W BELIZE CoROZAL District 5. Chan Chen 18°26'N; 88»27'W 6. Corozal 18°24'N; 88°24'W 7. Patchakan 18°24'N; 88''29'W 8. Santa Clara 18°18'N; 88°30'W Orange Walk District 9. Honey Camp Lagoon 18°03'N; 88°27'W 10. Orange Walk Town 18°05'N; 88°34'W 11. San Antonio, Rio Hondo 18°1 1'N; 88°39'W 12. Tower Hill, Belize Sugar Industries (B.S.I.) 18°02'N; 88''34'W Belize District 13. Belize City 17°30'N; 88°12'W 14. Churchyard 17°18'N; 88°33'W 15. Mussel Creek 17°39'N; 88°24'W 16. Rockstone Pond (Altun Ha) 17°46'N; 88°22'W 17. San Pedro, Ambergris Caye 17°55'N; 87°58'W 18. Tropical Park 17°28'N; 88''23'W 16°47'N: 30. Teakettle 17°13'N; 88°5rw 31. Xunantunich 17«t)5'N; 89°08'W Stann Creek District 32. Kendal 16°49'N; 88°22'W 33. Melinda 17°00'N; 88°18'W 34. Quam Bank, Cockscomb Basin 88°28'W Toledo District 35. Aguacate 16°10'N; 89°06'W 36. Big Fall 16''15'N; 88°53'W 37. Blue Creek 16°12'N; 89°03'W 38. Crique Jute and Salamanca Camp (Forestry Camp) 16°16'N; 89°0rw 39. Crique Negro, Columbia Forest 16°17'N; 89°02'W 40. Cuevas Creek and Jacinto Creek Bridges, at Punta Gorda Road 16°09'N; 88°53'W 41. Forest Home 16'^8'N; 88°50'W 42. Nimli Punit 16°20'N; 88''48'W 43. Orange Point 16°04'N; 88°49'W 44. Pueblo Viejo 16°13'N; 89°09'W 45. Punta Gorda 16°07'N; 88°48'W 46. Rice Station (Agricultural Station) 16°08'N; 88°51'W 47. San Antonio 16°15'N; 88°02'W 48. San Lucas (deserted) 16°05'N; 89°06'W 49. San Pedro Columbia 16°17'N; 88°58'W 50. Santa Elena 16°14'N; 89'X)6'W 51. Union Camp 16°24'N; 89°08'W Cayo District 19. Augustine 16°58'N; 88°59'W 20. Baldy Beacon, Bald Hills 17°0rN; 88°47'W 21. Banana Bank 17°15'N; 88''48'W 22. Barton Creek at Western Hwy. 17°13'N; 89''57'W 23. Central Farm and Listowel 17°irN; 89°00'W 24. C.I.T.A.,Sibun River at Indian Creek 17''16'N; 88''34'W 25. Macaw Bank 17°05'N; 89'^4'W 26. Roaring Creek 17°15'N; 88''47'W 27. San Antonio 17°05'N; 89°0rW 28. San Luis 16°54'N; 89°00'W 29. SibunCamp 17°05'N; 88°39'W Appendix This district checklist of the bat fauna of Belize is based on published accounts. The citations refer to the initial taxonomic treatments of specimens. Districts are arranged from north (left) to south (right). Abbreviations are as follows: Cz = Coro- zal; OW = Orange Walk; Bz = Belize; Cy = Cayo; SC = Stann Creek; Td = Toledo. In order to give an accurate list of bats, it is necessary to present certain discrepancies that have appeared in the literature. Dobson (1878) referred to certain early sp>ecimens that may have origi- nated from Belize. One reference (Dobson, 1878, 160 FIELDIANA: ZOOLOGY p. 520) to "Half-Moon Key, Honduras" for a spec- imen of Artibeus perspicillatus (= jamaicensis) be- longs to Belize, since this specimen was collected by O. Salvin (see Salvin, 1864). Sanderson (1941, p. 228) recalled "Anoura sp." in his descriptive narrative of a visit to then British Honduras. This species was not identified (Hershkovitz, 1951) in the Sanderson bat collection. Diaemus youngi was cited from Belize (Villa-R., 1966, p. 340), but R. L. Peterson (pers. comm.) stated that the specimen in question was actually from Guyana. Specimens of 17 species of bats were listed in Disney (1968) without locality data. These specimens, which are housed in British Museum (Natural History) and Royal Ontario Museum, are all from Cayo Dis- trict. J. L. Eger (pers. comm.) identified the ques- tionable specimen of Molossus bondae in Disney ( 1 968, p. 7) as M. molossus. Quinones et al. ( 1 978, p. 559) reported six species, which I collected and identified, without the exact locality information other than "the Maya Mountains region." This locality is 1 km NW Augustine, Cayo District. Sixty-six bat species are recognized in Belize. Species Cz OW Bz Cy sc Td References Rhynchonycteris naso X Murie, 1935, pp. 17-18; Disney, 1968, p. 7 Saccopteryx bilineata X X X X Sanborn, 1937, p. 331; Hershkovitz, 1951, p. 553; Disney, 1968, p. 7; Pendergast, 1979, p. 10 Saccopteryx leptura X This paper Peropteryx kappleri X Cartwright & Kirkpatrick, 1977, p. 466 Peropteryx macrotis X Hershkovitz, 1951, p. 553 Centronycteris maximilliani X Sanborn, 1941, p. 372 Balantiopteryx io X X Kirkpatrick et al., 1975, p. 330; Cartwright & Kirkpatrick, 1977, p. 466 Diclidurus virgo X This paper Noctilio leporinus X X X X This paper Pteronotus davyi X X X Disney, 1968, p. 7; this paper Pteronotus parnellii X X X Hershkovitz, 1951, pp. 553-554; Smith, 1972, p. 74; Cartwright & Kirkpatrick, 1977, p. 466; Quinones et al., 1978, p. 559; Pendergast, 1979, p. 10 Pteronotus personatus X This paper Mormoops megalophylla X X X X This paper Micronycteris brachyotis X X This paper Micronycteris megalotis X X X This paper Micronycteris nicefori X This paper Micronycteris schmidtorum X X X This paper Lonchorhina aurita X X This paper Macrophyllum macrophyllum X X This paper Tonatia bidens ... X X This paper Tonatia evotis X X Sanborn, 1941, pp. 372-373; Disney, 1968, p. 7 Disney, 1968, p. 7; this paper Tonatia minuta X X Mimon cozumelae X X X This paper Mimon crenulatum X X Ruiz, 1983, p. 374; this paper Phyllostomus discolor X This paper Phylloderma stenops X This paper Trachops cirrhosus X X X Sanborn, 1941, p. 374; Pendergast, 1979, p. 10; this paper Chrotopterus auritns X This paper Vampyrum spectrum X This paper Glossophaga commissarisi X X Webster & Jones, 1982, p. 5; this paper Glossophaga soricina X X Villa-R., 1966, p. 231; Disney, 1968, p. 7; Cartwright & Kirkpatrick, 1977, p. 466; Howell, 1977, p. 510; Quinones el al., 1978, p. 559; Pendergast, 1979, p. 10 Continued on next page MCCARTHY: DISTRIBUTION OF BATS 161 Species Cz OW Bz Cy SC Td References CaroUia brevicauda Carollia perspicillata Sturnira lilium Uroderma bilobatum Vampyrops helleri Vampyrodes caraccioli Vampyressa pusilla Chiroderma villosum Artibeus intermedius* Artibeus jamaicensis* Artibeus lituratus* Artibeus phaeotis X X X X Artibeus toltecus Artibeus watsoni X X X X Centurio senex Desmodus rotundus X X X X X X X X Diphylla ecaudata Natalus stramineus Thyroptera tricolor Myotis elegans Myotis keaysi X X X X X X X X X X Eptesicus furinalis Lasiurus horealis Lasiurus ega Lasiurus intermedius Rhogeessa tumida X X X X X X X X X X X X X X X Bauerus dubiaquercus Nyctinomops laticaudatus Eumops auripendulus Eumops bonariensis Eumops glaucinus Eumops underwoodi Molossus ater X X X X X X X X X X Molossus molossus Molossus sinaloae X X X X Hershkovitz, 1951, pp. 555-556; Villa-R., 1966, p. 270; Disney, 1968, p. 7; Pine, 1972, p. 42; Quinones et al., 1978, p. 559; Pendergast, 1979, p. 10 Hershkovitz, 1951, p. 555; Disney, 1968, p. 7; Pine, 1972, p. 42; Quinones et al., 1978, p. 559; Pendergast, 1979, p. 10 Disney, 1968, p. 7; Pendergast, 1979, p. 10 Disney, 1968, p. 7; Davis, 1968, p. 697; Pendergast, 1979, p. 10 This paper This paper Peterson, 1966, p. 676; this paper This paper Davis, 1984, p. 14 Dobson, 1878, p. 520; Andersen, 1908, p. 263; Gaumer. 1917, p. 298; Disney, 1968, p. 7; Davis, 1970b, p. 118; Pen- dergast, 1971, p. 102 Murie, 1935, p. 19; Hershkovitz, 1951, pp. 556-557; Disney, 1968, p. 7; Pendergast, 1979, p. 10 Disney 1968, p. 7; Davis, 1970a, p. 398; Pendergast, 1979, p. 10 This paper Davis, 1970a, p. 393; Pendergast, 1979, p. 10 This paper Hershkovitz, 1951, p. 557; Disney, 1968, p. 7; Cartwright & Kirkpatrick, 1977, p. 466; Quinones et al., 1978, p. 559; Pen- dergast, 1979, p. 10 This paper This paj)er Sanborn, 1941, p. 382 This paper Hershkovitz, 1951, pp. 557-558; LaVal, 1973a, p. 25; Quinones et al., 1978, p. 559 Disney, 1968, p. 7; this paper This paper This paper This paper LaVal, 1973b, p. 29; Kirkpatrick et al., 1975, p. 331 This paper Murie, 1935, p. 19 Eger, 1974, p. 5; this paper This paper Eger, 1977, p. 42 Eger, 1977, p. 55 Murie, 1935, p. 19; Pendergast, 1979, p. 10 Murie, 1935, p. 19 Hershkovitz, 1951, p. 559; Disney. 1968, p. 7 * Davis (1984) examined the Artibeus "lituratus" complex in Middle America and restored Artibeus intermedius J. A. Allen to specific status. Specimens cited in the publications listed for A. jamaicensis and A. lituratus should be reevaluated. 162 HELDIANA: ZOOLOGY New Species of Mammals from Northern South America: Fruit-Eating Bats, Genus Artibeus Leach Charles O. Handley, Jr. ABSTRACTS The larger species of Artibeus of the Amazon Basin are defined, and a new giant species is named and described from Venezuela and Colombia. Artibeus fallax, A. Hercules, and A. pla- nirostris are regarded as subspecies of Artibeus jamaicensis, by far the most variable of the larger Artibeus of the region. The smaller Artibeus are keyed and arranged in six species groups. A new dwarf species is described from Brazil, Ecuador, Guyana, Peru, and Venezuela. Distribution and diversity of the smaller species are discussed. Artibeus cinereus, once thought to range throughout Central America and much of South America and to include all of the smaller taxa except A. concolor and A. hartii, is restricted to include only the nominate form and A. quadrivittatus of the lower Amazon Basin and adjacent coastal areas. With these additions and changes in status, at least nine species of Artibeus now are known to occur in northeastern South America. Las especies de gran tamaiio de Artibeus de la Cuenca del Rio Amazonas son definidas y una nueva especie gigante de Venezuela y Colombia es nombrada y descrita. Artibeus fallax, A. hercules, y A. planirostris son consideradas como subespecies de Artibeus jamaicensis, que es el mas variable de los grandes Artibeus de la region. Una clave es preparada para las especies de Artibeus menores, y las especies son arregladas en seis grupos. Una nueva especie enana de Brasil, Ecuador, Guyana, Peru, y Venezuela es descrita. La distribucion y la diversidad de las especies menores son discutidas. Artibeus cinereus, que antes se penso estaba distribuida en Centro America y una gran parte de Sudamerica, y que incluyera todas las taxa mas pequeiias (a excepcion de A. concolor y A. hartii), es ahora restringuida para incluir solamente la especie nominal y A. quadrivittatus a la Cuenca baja del Rio Amazonas y a las areas costeras adyacentes. Con estas adiciones y cambios de "status," por lo menos nueve especies de Artibeus ya son conocidas y se encuentran en el nordeste de Sudamerica. Sao definidas as especies maiores de Artibeus que ocorrem na Bacia Amazonica, e uma especie nova, gigante, e descrita. Artibeus fallax, A. hercules, e A. planirostris sao consideradas subes- pecies de Artibeus jamaicensis, certamente a especie mais variavel dos Artibeus maiores da regiao. Uma chave para os Artibeus menores, os quais foram designados a seis grupos de especies, e fomecida. Uma especie nova aiia e descrita do Brasil, Equador, Guiana, Peru, e Venezuela. A diversidade, e as distribui^oes geograficas destas especies, sao discutidas. Artibeus cinereus, o qual acreditava-se abranger toda America Central e grande parte da America do Sul, alem de incluir todos taxa menores com excessao de A. concolor e A. hartii, e reduzido a um unico taxon, restrito ao sul da Bacia Amazonica e as suas areas adjacentes. Incluindo as adi9oes e mudan^as de status propostas neste trabalho, sao reconhecidas, atual- mente, ao menos nove especies de Artibeus na regiao nordeste da America do Sul. From the National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. HANDLEY: NEW SPECIES OF ARTIBEUS 163 Introduction Mammals and their ectoparasites were collected in Venezuela between 1965 and 1968 by the Smithsonian Venezuelan Project (SVP), supported in part by a contract (DA-49-MD-2788) of the Medical Research and Development Command, Office of the Surgeon General, U.S. Army. Nu- merous papers have described the ectoparasites and mammals of the Project. Throughout these papers undescribed species of mammals have been referred to by alphabetical designations. Some of these have been named subsequently by Handley and Ferris (1972), Handley and Gordon (1980), and Handley (1984). This paper deals with fruit- eating bats of the genus Artibeus Leach. The cranial measurements reported here were taken as outlined by Handley (1959, p. 98). Hind foot, tibia, calcar, and forearm were measured on dry museum si)ecimens or on specimens preserved in alcohol; all other external dimensions were measured on fresh specimens in the field. All mea- surements are in millimeters. Coloration was de- termined under Examolites (Macbeth Corp., New- burg, NY 12533) with natural light excluded. A New Giant Artibeus It is now generally agreed that in and around the Amazon Basin there are three large species of Artibeus. Handley (1976) recognized them as: (1) /1./m//^;>jo51« Gray— smaller, molars 3/3, rostrum arched, postorbital process poorly developed, fur long, coloration blackish, facial stripes faint or ab- sent, interfemoral membrane (IM) naked; (2) A. jamaicensis Leach — larger, molars 3/3, rostrum arched, postorbital process poorly developed, fur short, coloration gray-brown, facial stripes present but not sharply defined, IM naked; and (3) A. li- turatus Olfers— larger, molars 2/3, rostrum flat- tish, postorbital process well developed, fur short, coloration chocolate brown, facial stripes promi- nent and well defined, IM hairy. However, as shown by Koopman (1978) and Honacki et al. (1982), there is no consensus on the delimitation of these species. The difficulty in defining the species arises pri- marily from the fact that Artibeus jamaicensis is unusually variable geographically in morphology. The other species show very little variation in this region. Artibeus jamaicensis is large in the Ama- zon Basin, so large in fact that the subspecies there, A. j. fallax Peters and A. j. Hercules Rehn, until recently have been aligned by most authors with the universally large A. lituratus, although they differ from it in many characteristics other than size. To the southeast of the Amazon Basin {A. j. planirostris Spix) and to the north of it (A. j. trin- itatis Andersen), A. jamaicensis is dramatically smaller, in fact similar in size to A. fuliginosus. Everywhere east of the Andes A. jamaicensis has 3/3 molars; west of the Andes and in Central America it has 2/3 molars. Specimens in the SVP collection show that the large Artibeus jamaicensis fallax and small A. j. trinitatis apparently intergrade in the Llanos of Venezuela where the habitat is marginal for A. jamaicensis and where it is an uncommon bat. Furthermore, intergradation between the small, 1 2-molar A. j. trinitatis and the slightly larger, 10- molar A. j. aequatorialis Andersen of the north- west coast of South America can be seen in spec- imens from northern Colombia. These two zones of intergradation are of crucial importance in the nomenclature oi Artibeus, for they serve to link "/I. jamaicensis^'' of the West Indies and Central America and "A. planirostris" of eastern South America. They are especially im- portant in the present context because of the dis- covery of a fourth large Artibeus, superficially sim- ilar to but larger than A. j. fallax, occurring together with it in southern Venezuela and with the small A. j. trinitatis in western Venezuela and northern Colombia. It can be recognized as follows: Artibeus amplus new SF>ecies HoLOTYPE— USNM 440932, adult female with suckling young, skin and skull, collected 1 5 April 1968 by Norman E. Peterson, F. P. Brown, Jr., and J. O. Matson at Kasmera, 2 1 km SW Machi- ques, Estado Zulia, Venezuela, 270 m, in a damp cave in a cliff across the Rio Yasa from the Kas- mera Biological Station, eastern foothills of the Sierra de Perija. Original number, svp 22086. Etymology— Latin amplus, large, referring to the large size of this bat, one of the largest Artibeus. Distribution— Northern foothills of the Cen- tral Andes in Colombia; lower eastern slopes of the Sierra de Perija and the Venezuelan Andes in western Venezuela; and the vicinity of Cerro Dui- da and the low mountains of southeastern Bolivar in southern Venezuela. It probably occurs in ad- 164 HELDIANA: ZOOLOGY jacent parts of Guyana and Brazil as well. The SVP collectors found A. arnplus near streams and in other moist areas (98%); in evergreen forest (90%) and in forest openings such as yards and orchards (10%). Most specimens were mist netted (86%), but some ( 1 4%) were found roosting in the twilight zone of caves. Elevational range, 24-1200 m. Holdridge life zones (Ewel & Madriz, 1 968): Trop- ical humid forest (10%), Tropical very humid for- est (22%), Premontane humid forest (12%), Pre- montane very humid forest (2%), Premontane rain forest (4%), Lower montane very humid forest (10%), and Lower montane rain forest (40%). Ridge slopes and valley floor in the area where the ho- lotype was collected were clothed with second growth evergreen forest, while lawns, shrubbery, banana and papaya plants, and scattered grapefruit trees characterized the grounds of the biological station. Description— Size large (forearm 70.0, greatest length of skull 31.3, maxillary toothrow 11.2 — averages of Venezuelan specimens). Coloration of fur as in sympatric Artibeus jamaicensis (dorsum blackish brown to brown; facial stripes present but obscure; underparts blackish brown, usually frost- ed with white; underarms with abundant long, usually whitish hairs); ears dark fuscous to black, paler basally; lips and noseleaf blackish; mem- branes blackish; wing tips undifferentiated or gray- ish, never white. Horseshoe of noseleaf bound down mediobasally; legs and interfemoral mem- brane slightly hairy, the latter particularly medio- ventrally, where hairs extend as a short fringe be- yond edge of membrane; forearm long. Skull superficially like that of Artibeus jamai- censis, but relatively longer and narrower; rostrum long and flattish; supraorbital ledges subparallel and together with postorbital processes often poor- ly develojDed or even ill-defined; zygomata not very flared from skull, usually subparallel to one another, and in side view, thin and fragile; posterolateral angle of skull not particularly flared; palate rela- tively narrow and toothrows ovoid in outline; postpalatal extension usually long, narrow, and parallel sided; dentition as in A. jamaicensis, ex- cept I' only weakly bilobed; dental formula 2/2- 1/1-2/2-3/3 X 2 = 32. This bat is the only known host of Strebla paramirabilis Wenzel and Tri- chobius assimilis Wenzel (Diptera: Streblidae), so it can be distinguished from other Artibeus by its parasites as well as its morphology. Measurements of the holotype, an adult female: total length 101, tail vertebrae 0, hind foot (dry) 17, ear from notch 25, forearm 69.2, tibia 24.1, calcar 6.2, weight 70.4 g. Greatest length of skull 31.9, zygomatic breadth 18.3, postorbital breadth 7.7, breadth of braincase 13.3, depth of braincase 1 1.6, length of maxillary toothrow 1 1.2, postpal- atal length 9.8, palatal breadth outside of M^ 12.9, rostral breadth at base of canines 8.3. See Table 1 for additional measurements. Comparisons— Four large species of Artibeus occur in Venezuela, all of them together in the southern part of the country. Among these, Arti- beus amplus and A. jamaicensis are most alike; but despite the superficial resemblance, the two can be distinguished by many characters, both ex- ternal and cranial. All A. amplus examined have the lower edge of the noseleaf horseshoe bound down, while about 95% of A. jamaicensis from the same localities have it free; all A. amplus have the interfemoral membrane slightly hairy and fringed medially, hwX A. jamaicensis never does; and while A. jamaicensis often has the wings white-tipped, A. amplus never does. Cranially, A. amplus differs from A. jamaicensis in having a longer, narrower skull; longer, somewhat more ffattened rostrum (most easily seen in dimensions of rostral shield); less arched nasals; margins of supraorbital nearly parallel, rather than converging posteriorly, and usually not as well developed; zygomata thinner and more fragile and usually subparallel rather than diverging markedly posteriorly; posterolater- al angle of skull not so flaring; palate narrower and toothrows usually less nearly circular in outline; and postpalatal extension usually longer and nar- rower, parallel sided (not flaring posteriorward). The two species are hosts of different species of parasitic streblid ffies. Specimens Examined— Total 55. COLOMBIA. Antioquia: La Tirana, 33 km SW Zaragoza, 520 m (2 usnm). VENEZUELA. Apure: Nulita, Selvas de San Camilo, 29 km SSW Santo Domingo, 24 m (2 usnm). Bolivar: 21 to 33 km NE Icabani, 775-851 m (6 usnm); Km 125, 85 km SSE El Dorado, 826-1165 m (5 usnm). T.F. Amazonas: Belen, Rio Cunucunuma, 56 km NNW Esmeralda, 1 50 m (9 usnm); Cabecera del Caiio Culebra, Cerro Duida, 40 km NNW Esmeralda, 1 1 40-1 200 m (2 1 usnm); Caiio Culebra, Cerro Duida, 50 km NNW Esmeralda, 800 m (2 usnm); Tamatama, Rio Ori- noco, 2 km above Boca del Casiquiare, 135 m (2 usnm). Zulia: Kasmera, 21 km SW Machiques, 270 m (3 USNM, 1 ucv); 15 km W Machiques (1 amnh); Novito, 19 km WSW Machiques, 1 135 m (1 usnm). Remarks— In previous publications of SVP, Ar- tibeus amplus has been known as ""Artibeus sp. D". HANDLEY: NEW SPECIES OF ARTIBEUS 165 Table 1 . Measurements of adult Artibeus ampins and A. jamaicensis. For each measurement, line 1 includes the mean plus or minus two standard errors, line 2 the extremes, and line 3, in parentheses, the number of specimens measured. All specimens are from Venezuela unless otherwise stated. Total length 100.4 ± 3.88 93-104 (5) 89.9 ± 2.22 80-100 (21) 86.4 ± 2.90 77-95 (19) 83.8 ± 4.14 73-91 (10) 88.1 ± 1.62 80-93 (17) 86.3 ± 1.60 82-90 (8) Hind foot (dr>) Ear Forearm Greatest length Zygomatic breadth 15.9 ± 0.40 14-18 (19) 15.4 ± 0.32 15-16 (10) 17.8 ± 0.32 16-19 (17) 18.3 ± 0.32 18-19 (8) 16.9 ± 0.30 16.0-17.7 (12) 17.0 ± 0.28 16.6-17.6 (6) 19.1 ± 0.14 18.7-19.6 (13) 19.3 ± 0.32 18.6-20.2 (8) Postorbital breadth Artibeus amplus, males and females, Zulia and Colombia 18.4 ± 0.60 23.7 ± 1.28 70.8 ±1.78 31.4 ± 0.26 18.6 ± 0.20 17-19 22-26 68.6-75.3 31.0-31.9 18.1-18.8 (7) (7) (7) (7) (7) A. amplus, males and females, T.F. Amazonas and Bolivar 18.3 ± 0.30 23.0 ± 0.98 69.1 ± 0.90 31.2 ± 0.24 18.4 ± 0.14 17-20 18-26 65.0-73.2 30.3-32.8 17.4-19.1 (22) (21) (22) (29) (30) A. jamaicensis, females, Zulia 22.6 ± 0.96 61.1 ± 0.64 27.7 ± 0.24 17-25 58.9-64.2 26.7-28.5 (19) (19) (19) A. jamaicensis, males, Zulia 22.2 ± 0.98 59.3 ± 0.92 27.4 ± 0.14 20-25 56.2-61.4 27.1-27.7 (10) (10) (10) A. jamaicensis, females, T.F. Amazonas 24.6 ± 0.46 66.8 ±1.12 30.7 ± 0.28 23-26 62.1-70.1 29.4-31.3 (17) (17) (14) A. jamaicensis. males, T.F. Amazonas 24.6 ± 0.52 65.4 ± 1.42 30.7 ± 0.26 24-26 62.4-68.6 30.2-31.4 (8) (8) (8) 7.9 ± 0.20 7.6-8.3 (7) 7.8 ± O.IO 7.3-8.4 (31) 6.8 ± 0.08 6.5-7.1 (19) 6.8 ± 0.14 6.3-7.0 (10) 7.5 ± 0.10 7.1-7.7 (14) 7.6 ± 0.16 7.2-7.9 (8) A New Dwarf Artibeus The taxonomy of the smaller Artibeus is in a state of flux. As recently as 35 years ago all of the smaller species except A. concolor Peters and A. hart a Thomas were believed to be variants of .-i. cinerens Gervais. Since then, first one and then another of the supposed subspecies of .4. cinereus has been shown to be independent species. Today only A. bogotensis Andersen, A. glaucus Thomas, A. pumilio Thomas. A. quadrivittatus Peters. A. rosenbergi Thomas, and A. watsoni Thomas re- main associated with A. cinereus (Honacki et al., 1982). However, except for^. quadrivittatus, these do not properly belong with A. cinereus either. Artibeus glaucus and A. bogotensis intergrade in Ecuador and form an Andean-northern South American species sympatric with A. cinereus in southern Venezuela. Artibeus glaucus thus has two subspecies, the nominate form and A. g. bogoten- sis. Artibeus watsoni Thomas of northwestern South Ameinca and Central America is closely related, but intergradation with A. g. glaucus or A. g. bo- gotensis has not been obsei^ed. Artibeus pumilio is an enigmatic taxon. Many museum specimens bear the name A. pumilio, but perhaps the only specimen properly associated with the name is the holotype. This specimen may be only an odd variant of one of the other species, but not of the species described here. For the pres- ent, A. pumilio must be regarded as unplaceable. The same can be said for A. rosenbergi, charac- teiized by a curiously long, narrow skull such as can be found occasionally in large samples of most species o{ Artibeus. Because of their equivocal sta- tus, neither A. pumilio nor A. rosenbergi is includ- ed in the appended list of species and key. The characteii sties and status of these taxa will be the subject of another paper. Thus, .A. cinereus now has been shorn of all of its supposed subspecies except A. c. quadrivittatus. Its supposed range has been reduced from encom- passing most of Central Ameiica and tropical South America to occupying only the Amazon Basin (possibly only the lower basin) and adjacent coast- al areas. Sympatric with A. cinereus in much of its range is a distinctive dwarf species which can be known as: 166 HELDIANA: ZOOLOGY Table 1. Continued. Braincase breadth Braincase depth Maxillary Postpalatal toothrow length Width at molars Width at canines Tibia 13.5 ± 0.20 13.2-14.0 (7) Artibeus amplus. males and females, 1 1.0 ± 0.26 1 1.2 ± 0.16 9.7 ± 0.20 10.6-11.6 11.1-11.5 9.3-10.0 (7) (7) (7) Zulia and Colombia 13.2 ± 0.26 8.4 ±0.16 12.7-13.5 8.2-8.8 (7) (7) 25.9 ± 1.28 24.1-28.2 (6) 13.3 ± 0.12 12.9-14.0 (30) A. amplus, males and females, T.F. Amazonas and Bolivar 11.2 ±0.06 11.2 ±0.10 9.8 ±0.14 13.3 ± 0.10 8.6 ± 0.08 10.7-11.5 10.7-11.8 9.1-10.6 12.8-13.9 8.3-8.9 (30) (31) (28) (30) (30) 24.8 ± 0.38 23.1-26.2 (22) 12.3 ± 0.10 12.0-12.8 (19) 10.2 ± 0.16 9.5-10.9 (19) A. jamaicensis. females 10.0 ±0.12 8.6 ±0.14 9.4-10.5 8.2-9.3 (17) (18) , Zulia 12.1 ± 0.16 11.2-12.7 (19) 7.6 ± 0.10 7.2-8.0 (19) 22.4 ± 0.50 20.1-23.8 (19) 12.1 ± 0.14 11.8-12.4 (10) 10.3 ± 0.18 10.0-10.9 (10) A. jamaicensis. males, 10.1 ± 0.16 8.4 ± 0.12 9.7-10.4 8.2-8.8 (8) (10) Zulia 12.2 ± 0.18 11.8-12.6 (10) 7.7 ± 0.12 7.4-7.9 (9) 21.7 ± 0.52 20.1-23.0 (10) 13.2 ± 0.12 12.9-13.5 (14) 10.8 ± 0.18 10.2-11.5 (14) A. jamaicensis. females, T.F 11.4 ±0.20 9.4 ±0.18 11.0-12.0 8.9-10.0 (13) (14) . Amazonas 13.7 ± 0.22 13.2-14.4 (13) 8.6 ± 0.12 8.3-8.9 (13) 24.1 ± 0.46 22.3-25.4 (17) 13.4 ± 0.22 12.8-13.7 (8) 11.0 ± 0.18 10.7-11.4 (8) A. jamaicensis, males, T.F. 11.4 ±0.18 9.3 ± 0.24 11.1-11.8 8.7-9.8 (8) (8) Amazonas 13.8 ± 0.30 13.0-14.3 (8) 8.8 ± 0.12 8.6-9.1 (8) 23.3 ± 0.52 22.0-24.4 (8) Artibeus gnomus new species HoLOTYPE— USNM 387534, adult female, skin and skull, collected 14 June 1966 by A. L. and M. D. Tuttle at El Manaco (= Km 74), 59 km SE El Dorado, Bolivar, Venezuela, 150 m, in a mist net in an orchard. Original number, svp 9298. Etymology— Latin gnomus, diminutive fabled being, dwarf, alluding to the small size of this species, one of the smallest Artibeus. Distribution— The Amazon Basin and bor- dering regions; from northern Amazonas Territory (14 km SSE Pto. Ayacucho) and northern Bolivar State (28 km SE El Manteco) in Venezuela and northern Guyana, to Para (Belem) and Mato Gros- so (Serra do Roncador), Brazil, and Loreto (Santa Rosa), Peru. SVP collectors netted A. gnomus mostly in moist sites (92%) in evergreen forest (52%) or openings such as savannas (25%) and yards and orchards (23%). Elevations range 1 19- 161 m in Venezuela, sea level to 530 m in Brazil. Holdridge life zones: Tropical dry forest (22%), Tropical humid forest (67%), Tropical very humid forest (2%), and Premontane humid forest (9%). Description— Body size small (forearm aver- ages 36-38, greatest length of skull 18.5-18.7, and maxillary toothrow 5.7-6.0). Dorsal coloration gray-brown to brown; underparts paler; facial stripes very white and sharply defined. Soft parts coloration in life (usnm 36 1 742, male, Belem, Bra- zil): ear narrowly edged with yellow, brightest to- ward base; antitragus entirely yellow; tragus yel- low, brightest distally and on posterior basal lobe; noseleaf and horseshoe gray-brown medially, cream color laterally; lips and chin gray-brown; iris brown; forearm and fingers brownish flesh col- or; wings blackish, except membrane between fin- gers II and III transparent, grayish; interfemoral membrane sooty brown; legs and feet dark brown; claws horn color. Face short; shape and propor- tions of ears, noseleaf, horseshoe, lips, chin, and interfemoral membrane as in Artibeus cinereus; noseleaf minutely hirsute; lower edge of horse- shoe free; basal part of forearm hairy; hind ex- tremities (except for short hairs on feet) appear naked. Skull small, short, and broad; zygomata sub- parallel; rostrum narrow, very short, moderately HANDLEY: NEW SPECIES OF ARTIBEUS 167 Table 2. Measurements of adult male and female (combined) Artibeus gnomus and A. glaucus bogotensis. For each measurement, line 1 includes the mean plus or minus two standard errors, line 2 the extremes, and line 3, in parentheses, the number of specimens measured. All specimens are from Venezuela. Total length 47.5 ± 1.40 44-54 (13) 52.2 ± 0.86 49-56 (19) Hind foot (dry) Ear Forearm Greatest length Zygomatic breadth A. glaucus bogotensis. Km 125, 85 km SSE El Dorado 10.6 ± 0.22 17.4 ±0.38 39.6 ± 0.60 20.3 ± 0.10 11.6 ±0.10 10-11 16-19 36.8-41.9 19.4-21.2 10.8-12.1 (19) (19) (19) (50) (44) Postorbital breadth Artibeus gnomus, Rio Supamo, Los Patos, and EI Manaco 9.5 ± 0.28 16.9 ± 0.62 36.7 ± 0.54 18.5 ± 0.18 11.0 ±0.18 9-10 14-19 34.0-38.3 17.9-19.1 10.4-11.2 (13) (13) (13) (14) (8) 4.9 ± 0.10 4.5-5.2 (14) 5.0 ± 0.04 4.6-5.3 (51) deep and arched, and much swollen posterolater- ally (part on rostral shield, part within orbit, above eye); excavation for orbital nerve large and deep; braincase short and deep, with swelling at pos- terodorsal apex interrupting junction of sagittal and lambdoidal crests; postpalatal extension rel- atively short; internal edge of pterygoid fossa strongly ridged, narrowing mesopterygoid fossa and cupping pterygoid fossa which opens straight back; vomerine ridge visible in mesopterygoid fossa; va- cuities in roof of posterior nares much anterior to mesopterygoid fossa and not easily seen; outline of maxillarv toothrows nearly circular; upper ca- nine small (especially in basal diameter); M' with accessory internal ridge on lateral cusps, and with relatively wide talon; m, present (75 of 79 speci- mens examined). Measurements of the holotype, an adult female: total length 47, tail vertebrae 0, hind foot (dry) 9, ear from notch 18, forearm 36.5, tibia 12.6, calcar 4.9, weight 10.5 g. Greatest length of skull 18.2, zygomatic breadth 10.8, postorbital breadth 4.8, breadth of braincase 8.5, depth of braincase 7.2, length of maxillary toolhrow 5.5, postpalatal length 6.5, palatal breadth outside of M' 7.1, rostral breadth at base of canines 4.6. See Table 2 for additional measurements. Comparisons— ^r//Z)e«5 gnomus differs from A. concolor and A. hartii in many ways, but most significantly in lack of M\ From all other small Anibeus {A. anderseni, A. cinereus, and A. glaucus bogotensis) that occur within its range, A. gnomus can be distinguished by its possession of mj. Among the specimens examined, m, is consistently absent in these other taxa while it is consistently present in A. gnomus (except in southern Venezuela, where it is absent from both mandibles in four of 53 specimens and from one mandible only in two others). In addition, A. gnomus differs from all of the sympatric taxa in its more prominent white facial stripes; more colorful ears, noseleaf, and lips; average browner, less grayish coloration of pelage; shorter face and rostrum (except when compared with A. concolor); more swollen supraorbital re- gion; average larger and deeper orbital nerve ex- cavation (sometimes equally large and deep in A. g. bogotensis); and more cupped pterygoid fossa, with internal ridge so enlarged as to significantly narrow the mesopterygoid fossa. Artibeus gnomus differs from the sympatric taxa individually in several other ways. It is much smaller than A. concolor (forearm averages 36-38 vs. 46-48). In contrast to A. hartii it has notched inner upper incisors, brownish rather than dark chocolate coloration, and a wide, unfringed inter- femoral membrane. Compared with A. aruierseni (including the holotype, fmnh 21331), ^. gnomus is similar in size (slightly larger than Rio Madeira A. anderseni); has rostrum much deeper, more arched, narrower, and shorter; face not dished; orbit larger; zygomata more nearly parallel; and vacuities in roof of p)OSterior nares far forward of mesopterygoid fossa, rather than opening in it or close to it. At Belem, Brazil, both Artibeus gnomus and A. cinereus were numerous and were often taken in the same nets. There, fresh specimens of the two species were compared. Artibeus gnomus is smaller in size, and has a smaller head and shorter face; facial stripes much brighter, more sharply defined, and more prominent; ears, noseleaf, and lips more brownish, less grayish; ear edgings, antitragus, and tragus bright yellow, rather than cream; and nose- leaf edged with cream, rather than plain gray- brown. Furthermore, it has zygomata more nearly parallel; rostrum deeper and shorter; supraorbital area much swollen and its edges nearly parallel; and smaller teeth. 168 HELDIANA: ZOOLCX}Y Table 2. Continued. Braincase breadth Braincase depth Maxillary toothrow Postpalatal length Width at molars Width at canines Tibia 8.5 ± 0.14 8.1-9.0 (14) 9.0 ± 0.06 8.5-9.5 (48) Artibeus gnomus, Rio Supamo, Los Patos, and El Manaco 7.4 ± 0.14 5.7 ± 0.06 6.3 ± 0.10 7.5 ± 0.12 4.9 ± 0.08 13.2 ± 0.46 7.1-8.0 5.5-5.9 6.0-6.7 7.1-7.9 (14) (14) (14) (14) A. glaucus bogotensis, Km 125, 85 km SSE El Dorado 7.9 ± 0.06 6.5 ± 0.04 7.0 ± 0.08 8.0 ± 0.06 7.2-8.3 6.0-6.8 6.5-7.5 7.5-8.7 (48) (51) (47) (50) 4.6-5.1 (14) 5.1 ± 0.06 4.9-5.6 (50) 11.2-14.4 (13) 13.6 ± 0.36 12.3-15.8 (19) In southern Venezuela Artibeus gnomus is sym- patric with A. glaucus bogotensis. Compared with Venezuelan specimens and with the holotype (bm 99.11.4.35) of this taxon, A. gnomus is much smaller and shorter faced; has a deeper, shorter rostrum; disproportionately wider zygomatic spread; and smaller teeth. In addition to comparisons of /I. gnomus with sympatric species, two other small Artibeus need to be considered: 1. Artibeus g. glaucus —This species occurs nearby in the Andes. It (including the holotype, BM 94.8.6.13) possesses m„ and its skull has the basic shape of /i. gnomus. However, it is much larger and darker in color, has the hind extremities much hairier, the supraorbital region usually less swollen, and the pterygoid fossa much less cupped and opening to the mesopterygoid fossa. 2. Artibeus fvatsoni— West of the Andes and ex- tending into Central America is another small species, A. watsoni Thomas, which like A. gnomus possesses m,. It (including its holotype, bm 0.7. 1 1 . 1 9) is larger than A. gnomus; has larger teeth; longer rostrum, with reduced supraorbital swell- ing; shallower and less well-defined orbital nerve excavation; and like A. glaucus has the pterygoid fossa not cupped and opening into the mesopter- ygoid fossa (which consequently is not narrowed by the inner pterygoid ridge). Discussion— The ten small species of Artibeus recognized here can be associated in six species groups: 1. Artibeus concolor Group— Amazon and up- per Orinoco basins and Guianas. Includes only Artibeus concolor. 2. Artibeus hartii Group— Mexico and Central America, across northern South America to Trin- idad, and south to Peru east of the Andes and to Ecuador west of the Andes. Includes only Artibeus hartii. 3. Artibeus glaucus Group— Mexico, Central America, and South America to Mato Grosso and Peru. Includes Artibeus glaucus (with two subspe- cies, A. g. bogotensis and A. g. glaucus), A. gnomus, and A. watsoni. 4. Artibeus toltecus Group— Mexico and Cen- tral America. Includes Artibeus aztecus Andersen and Artibeus toltecus Saussure, each with several subspecies. 5. Artibeus cinereus Group— Guiana region, coastal Brazil, and lower Amazon Basin (dubious- ly also upper Amazon Basin). Includes only Ar- tibeus cinereus, with A. c. quadrivittatus as a sub- species. 6. Artibeus phaeotis Group — Mexico, Central America, and South America to upper Amazon Basin and western Ecuador. Includes Artibeus an- derseni Osgood and Artibeus phaeotis Miller, ' with several subspecies. Diversity in the small Artibeus is greatest in east- ern South America, where representatives of five of the six groups occur and where three of the groups are endemic. Altogether six species occur in and around the Amazon Basin, while only one is known with certainty in the central portion of the Basin; there are three in the lower Amazon ' Until recently (Koopman, p. 152, in Honacki et al., 1982) it has not been generally recognized that Artibeus phaeotis and A. ravus are conspecific. They inlergrade in eastern Panama and western Colombia. Both names date from Miller (1902). Although A. ravus was named first, on an earlier page, A. phaeotis became embedded in the literature as the name of this species. HANDLEY: NEW SPECIES OF ARTIBEUS 169 and on the southern fringes in Brazil and Bohvia, four or five in southern Venezuela, and five in east- em Peru, Ecuador, and Colombia. In contrast, only three of the species groups occur in Central Amer- ica, and only one of them is endemic there. Several distributional patterns are represented in the complex of Amazonian species. Artibeus concolor is found throughout the Basin but scarce- ly beyond it; A. cinereus occurs in the lower Am- azon and along the coast for some distance north and south of the river; A. anderseni is known from the upper Amazon and an isolated area in northern Colombia; A. glaucus is higher up, in the Andes, and eastward around the northern edge of the Ba- sin in Venezuela; the range of A. hartii resembles that of A. glaucus, but extends on into Central America; and the dwarf /I. gnomus has a p)eculiar circular range, completely ringing the Amazon Ba- sin but apparently not extending into its interior. Key to the Smaller Species of Artibeus 1. Molars 3/3 (mj large) 2 r. Molars 2/3 (mj minute) or 2/2 3 2. V notched; facial stripes absent; coloration pale brown; interfemoral membrane broad and naked; forearm 43-52 mm Artibeus concolor 2'. I' not notched; facial strip>es present; color- ation dark chocolate brown; interfemoral membrane narrow and fringed; forearm 36- 42 mm Artibeus hartii 3. Supraorbital region much swollen; molars 2/3 (2/2 m A. g. bogotensis and occasionally in the others) . . . Artibeus glaucus Group, 4 3'. Supraorbital region little, or not at all, swol- len; molars 2/2 7 4. Rostrum short and moderately arched; pter- ygoid fossa cupijed and opening back, causing mesopterygoid fossa to be narrowed; forearm 34-38 mm Artibeus gnomus 4'. Rostrum long and much or only moderately arched; pterygoid fossa not cupped, opening into and not narrowing mesopterygoid fossa 5 5. Rostrum much arched; orbital nerve exca- vation shallow and often ill-defined; dorsum pale brownish; ears pale; forearm 35-41 mm Artibeus watsoni 5'. Rostrum moderately arched; orbital nerve excavation deep and well defined 6 6. Molars usually 2/3; dorsum dark grayish or blackish; ears dark; forearm 38-42 mm . . . Artibeus glaucus glaucus 6'. Molars 2/2; dorsum pale brownish or grayish; ears pale; forearm 37-41 mm Artibeus glaucus bogotensis 7. Interfemoral membrane narrow and fringed; coloration blackish Artibeus toltecus Group, 8 7'. Interfemoral membrane broad, naked; col- oration brownish 9 8. Larger, forearm 42-48 mm Artibeus aztecus 8'. Smaller, forearm 37-41 mm Artibeus toltecus 9. Rostrum deep and arched; palate long and moderately wide Artibeus cinereus 9'. Rostrum shallow and flattened; palate short and very wide Artibeus phaeotis Group, 10 10. Maxillary toothrow 5.2-6.2 mm; rostrum often tilted up anteriorly Artibeus anderseni 10'. Maxillary toothrow 6.7-7.1 mm; rostrum usually not tilted up anteriorly^ Artibeus phaeotis Specimens Examined— /irf/Aeiii anderseni — BRAZIL. Amazonas: Borba, Rio Madeira (1 amnh). Rondonia: Porto Velho (2 amnh, 2 fmnh, including holotype of ^. anderseni); Sto. Antonio do Hauayara (4 amnh). COLOMBIA. Bolivar: Ca- tival. Upper Rio San Jorge, 120 m (16 fmnh). Antioquia: Aljibos, 26 km S and 22 km W Zara- goza, 630 m (2 usnm); nr. La Tirana, 24 km S and 22 km W Zaragoza, 520 m (2 usnm). ECUAIX)R. Napo: Rio Suno (Abajo) (4 amnh). Pastaza: Mon- talvo, Rio Bobonaza ( 1 fmnh); Rio Pindo Yacu (1 fmnh); Rio Yana Rumi (1 fmnh). PERU. Hua- nuco: Monte Alegre (1 amnh). Loreto: Boca Rio Curaray (1 amnh); Boca Rio Peruate, Rio Ama- zonas, 90 m (1 FMNH); Lagarto, Alto Ucayali (1 amnh); Mazan (1 amnh); 59 km W Pucallpa (1 usnm); Puerto Indiana, Rio Amazonas (2 amnh); Rio Morona (Quebr. Pushaga), Alto Amazonas, 220 m (2 FMNH); Rio Yavari Mirim (Quebr. Es- peranza), 200 m (2 fmnh); Santa Cecilia, Rio Man- iti, Iquitos, 110 m (3 fmnh); Santa Luisa, Rio Nanay, Iquitos, 160 m (1 fmnh); Sarayacu, Rio Ucayali (1 amnh). Pasco: San Juan, Oxapampa, ^ Couplet 10 will separate Artibeus anderseni and A. phaeotis in South America and in southern Central America, but it will not distinguish A. anderseni from Mexican A. phaeotis nanus. In such a comparison, A. anderseni can be recognized by its relatively broader skull. 170 FIELDIANA: ZOOLOGY j 274 m (3 usnm). Departamento (?): Yuhucumayo, 1200 ft [= Puno: Yahuaramayo, 366 m?] (1 mcz). Artibeus cinereus c/iiere«5— BRAZIL. Amazo- nas: Sta. Clara, Vila Bela Imperatriz [nr. Parintins] (1 amnh). Para: Fordlandia, Rio Tapajos (2 amnh); Maracano, Rio Jamunda [= Nhamunda?], Faro (5 amnh); Rio Yumunda, Faro ( 1 bm). Artibeus cinereus quadrivittatus — BRAZIL,. Maranhao: Juryassu [= Turia9u?] (1 bm). Para: Belem (10 usnm); Benevides (1 bm); Para [= Be- lem] (1 bm); Ilha do Taiuna, Rio Tocantins (3 amnh). Pemambuco: Pemambuco [= Recife] (2 bm). Rio Grande do Norte: Natal (1 usnm). SUR- IN AME. Surinam ( 1 bm). VENEZUELA. Bolivar: Hato San Felipe, Serrania de Nuria (1 ucv); Hato San Jose, 20 km W La Paragua, 300-324 m (2 usnm). Artibeus glaucus bogotensis — COLOMBIA. Cundinamarca: Bogota (2 bm): nr. Bogota ( 1 bm); Curiche, nr. Bogota (2 bm, including holotype of A. bogotensis); Fomeque (1 amnh); Fusagasuga (2 mcz); Rio Negro, nr. Bogota (2 bm). GUYANA. Kanuku Mts. (3 bm). VENEZUELA. Bolivar: El Manaco, 59 km SE El Dorado, 150 m (3 usnm); Hato San Jose, 20 km W La Paragua, 300-324 m (3 USNM); 23 to 45 km NE Icabaru, 824-851 m (3 USNM); Km 125, 85 km SSE El Dorado, 826-1 165 m ( 1 20 usnm); Rio Supamo, 50 km SE El Manteco, 1 50 m (2 usnm). T.F. Amazonas: Belen, Rio Cu- nucunuma, 56 km NNW Esmeralda, 150 m (1 usnm); Caiio Culebra, Cerro Duida, 50 km NNW Esmeralda, 800 m (3 usnm). Artibeus glaucus glaucus— BOIAW A. Santa Cruz: Buenavista, 400 m (1 fmnh). ECUADOR. Napo: Baeza (1 bm). PERU. Cuzco: Collpa de San Lorenzo, Quincemil, 700 m (1 1 fmnh); Hda. Ca- dena, Quincemil, 1000 m (9 fmnh). Junin: Chan- chamayo, 1000 m (2 bm, including holotype oi A. glaucus); Huacapistana ( 1 f>4Nh). Puno: Rio Inam- bari, 670 m (3 amnh); Santo Domingo (1 amnh); Yahuaramayo, 366 m (1 bm, 1 usnm). Artibeus gnomus-ToXdA 104. BRAZIL. Mate Grosso: Serra do Roncador, 264 km N (by road) Xavantina, 533 m (17 usnm). Para: Belem, Sta. A, IPEAN (7 usnm); Belem, Utinga (5 usnm); Be- lem, Benevides (2 usnm). ECUADOR. Pastaza: Canelos, upper Rio Bobonaza (1 amnh). GUY- ANA. E. Berbice District: Wikki River (3 usnm). Mazaruni-Potaro District: Issano Road, 1 2 mi W of Bartica-Potaro Road (1 usnm). PERU. Loreto: 59 km SW Pucallpa (1 usnm); Santa Rosa, Alto Ucayali (10°42'S, 73''50'W) (2 amnh). VENE- ZUELA. Bolivar: El Manaco, 59 km SE El Do- rado, 150 m (12 usnm); Km 38, SE El Dorado, 100 m (1 ucv); Los Patos, 28 km SE El Manteco, 1 50 m (4 usnm); Rio Supamo, 50 km SE El Man- teco, 150 m (1 usnm); Salto Chalimaha, Rio Pa- ramichi, Rio Paragua (1 ucv); Salto Ichun, Rio Paragua (2 ucv). T.F. Amazonas: Belen, Rio Cu- nucunuma, 56 km NNW Esmeralda, 150 m (2 usnm); Boca Mavaca, 84 km SSE Esmeralda, 1 38 m (1 usnm); Caiio Leon, Cerro Duida, 325 m (1 amnh); Capibara, Brazo Casiquiare, 106 km SW Esmeralda ( 1 usnm); Esmeralda, Cerro Duida, 325 m (3 amnh); 14 to 65 km S, SSE, and SSW Pto. Ayacucho, 119-161 m (16 usnm); Rio Mavaca, 1 08 km SSE Esmeralda, 1 40 m (7 usnm); San Juan, Rio Manapiare, 163 km ESE Pto. Ayacucho, 155 m (6 usnm); Tamatama, Rio Orinoco, 135 m (7 usnm). Artibeus phaeot is— HoXoXyTpcs o{A. phaeotis and A. ravus, plus hundreds of other specimens from Mexico, Central America, and NW South Amer- ica. Artibeus pumilio— PERU. Loreto: Masisea, Tushemo, Rio Ucayali, 328 m (1 bm, holotype of A. pumilio). Artibeus watsoni— PANAMA. Chiriqui: Boga- va, 250 m (5 bm, including holotype of A. watsoni); Progreso (34 usnm); Puerto Armuelles (2 usnm). Remarks— In previous publications of SVP, Artibeus gnomus has been known as '''' Artibeus sp. A". Acknowledgments Among the persons who helped me put together this paper I am especially grateful to Sally DeMott, who measured the SVP skulls; Linda Gordon, who compiled the tables and worked with me on the comparisons; and Jane Ailes Small, who read and criticized the manuscript and did the word pro- cessing. Curators of several collections kindly per- mitted me to study specimens under their care in the preparation of these descriptions: American Museum of Natural History (AMNH), British Museum (Natural History) (BM), Field Museum of Natural History (FMNH), Museum of Com- parative Zoology, Harvard University (MCZ), and Universidad Central de Venezuela (UCV). The SVP collection is in United States National Mu- seum of Natural History (USNM); a portion of its specimens have been returned to Venezuela. HANDLEY: NEW SPECIES OF ARTIBEUS 171 Literature Cited EwEL, J. J., AND A. Madriz. 1968. Zonas de Vida de Venezuela. Ministerio de Agricultura y Cria, Caracas, 265 pp., map. Handley, C. O., Jr. 1959. A revision of American bats of the genera Euderma and Plecotus. Proceedings of the United States National Museum, 110: 95-246. . 1976. Mammals of the Smithsonian Venezue- lan Project. Brigham Young University Science Bul- letin, Biological Series, 20(5): 1-91. . 1984. Newspeciesof mammals from northern South America: A long-tongued bat, genus Anoura Gray. Proceedings of the Biological Society of Wash- ington. 97: 513-521. Handley, C. O., Jr., and K. C. Ferris. 1972. De- scriptions of new bats of the genus Vampyrops. Pro- ceedings of the Biological Society of Washington, 84: 519-523. Handley, C. O., Jr., and L. K. Gordon. 1980. New species of mammals from northern South America. Mouse possums, genus Marmosa Gray, pp. 65-72. In Eisenberg, J. F., ed.. Vertebrate Ecology in the North- em Neotropics. Smithsonian Institution Press, Wash- ington, D.C. HoNACKJ, J. H., K. E. Kjnman, and J. W. Koeppl, eds. 1982. Mammal Species of the World. Allen Press, Inc., and the Association of Systematic Collections, Lawrence, Kansas, 694 pp. Koopman, K. F. 1978. Zoogeography of Peruvian bats with special emphasis on the role of the Andes. Amer- ican Museum Novitates, 2651: 1-33. Miller, G. S., Jr. 1902. Twenty new American bats. Proceedings of the Academy of Natural Sciences of Philadelphia, 54: 389-412. 172 HELDIANA: ZOOLOGY Seasonality of Reproduction in Peruvian Bats Gary L. Graham ABSTRACTS The reproductive conditions of 3,489 specimens were used to determine seasonal patterns of pregnancy and parturition in Peruvian bats. More species that are trophic generalists yield birth records for the dry season than do trophic specialists. Relatively more highland than lowland species have births recorded for both seasons (dry and wet). Presumably, trophic generalists and highland species experience less seasonal variation in food supplies compared to the other groups. A larger percentage of nectarivorous species than frugivores have birth records for the dry season; the reverse is true for the wet season. These patterns are associated with greater floral resource abundance during the dry season and a greater abundance of fruit resources during the wet season. Las condiciones reproductivas de 3489 especimenes fueron usadas para describir patrones estacionales de embarazo y alumbramiento en murcielagos peruanos. Mas especies de gener- alistas alimenticios tienen registros de nacimiento durante los dos estaciones (seca y mojada) que tienen especialistas. Relativamente mas especies desde tierras altas que desde tierras bajas tienen registros de nacimiento durante la estacion seca. Possiblamente, generalistas alimenticios y especies de tierras altas sufrir menos variacion estacional de provisiones alimentos que los otros grupos. Un mayor porcentaje de especies nectivoras que frugivoras tienen registros de nacimiento durante la estacion seca pero, el opuesto exista para la estacion mojada. Estos patrones son asociados con un mayor abundancia de recursos de flores durante la estacion seca y con un mayor abundancia de recursos de frutas durante la estacion mojada. As condi96es reprodutivas de 3489 especimes de morcegos p)eruanos foram usadas para descrever seus padroes de parturi^ao. Os morcegos de habitos alimentares generalizados pos- suem uma propor9ao maior de especies que possuem registros de partos durante a epoca da seca, do que os morcegos com habitos alimentares especializados. Proporcionalmente mais especis de morcegos de areas elevadas, do que das planicies, possuem registros de partos durante as duas epocas (seca e chuvosa). Morcegos generalizados e os de areas elevadas provavelmente experimentam menos varia^ao epocal em a quantidade da comida, do que os morcegos das planicies ou morcegos com habitos alimentares especializados. Uma porcentagem maior das especies nectarivoras, do que das especies frugivoras, possuem registros de partos durante a epoca da seca, sao a regra ao inves da exce9ao entre os morcegos que consumen fruta. Estes padroes podem ser interpretados como adpata96es de individuos a recursos alimentares que sao regularmente, ou ocasionalmente, obteniveis durante a epoca da seca. From the Department of Biology, University of New Mexico, Albuquerque, New Mexico 87131. GRAHAM: PERUVIAN BATS 173 Introduction Although the timing of reproductive events is an important Hfe history adaptation, there are few studies of the reproductive phenology of South American bats (Racey, 1982). Most of these stud- ies are of individual species and cover only part of the year. Gestation, parturition, lactation, and weaning should be timed to correspond with vari- ations in the abundance and diversity of food sup- plies (Fleming et al., 1972; Bradbury & Vehren- camp, 1977; Bonaccorso, 1979; Wilson, 1979; August & Baker, 1 982). These variations are known to be seasonal in most of the Neotropics (Janzen, 1967, 1973; Foster, 1982; Wolda, 1982; Smythe, 1982; Terborgh, 1983). Which reproductive event is actually synchronized with peak food supplies is likely to be determined by the relative cost of each event. Lactation is the most costly period for most females, but weaning and dispersal pose the greatest survival problem for young bats (Wilson, 1979; Racey, 1 982). These authors, and Tuttle and Stevenson (1982), agree that the weaning of young is the most critical period for individuals of most bat populations. Most species of Neotropical bats that have been studied are polyestrous (Fleming et al., 1972 Thomas, 1972; Bradbury & Vehrencamp, 1977 Myers, 1977; Wilson, 1979; Bonaccorso, 1979 Humphrey & Bonaccorso, 1979). Each year, fe- male bats confront the problem of timing two (rarely three in a few vespertilionids) periods of lactation and weaning with variations in food availability. Individuals of most species handle this problem by producing their young so that the first is weaned at the beginning of the wet season as fruit supplies are reaching their peak, and the second later in the wet season when fruit avail- ability reaches a second peak or remains relatively high (Wilson, 1979; Tuttle «& Stevenson, 1982). Exceptions to this general pattern of seasonal poly- estry have been observed in most studies (Fleming et al., 1972; Thomas, 1972; Myers, 1977; Myers & Wetzel, 1983; Bonaccorso, 1979; see Mares & Wilson, 1 97 1 ; Bradbury & Vehrencamp, 1 977; and August & Baker, 1982 for good discussions of dry season birth periods). The purpose of this study is to examine some of the details of dry season reproduction in Pe- ruvian bats. I demonstrate that the different feed- ing assemblages and the faunas of different zoo- geographical regions differ in the proportion of species with birth records during the dry season. I also offer suggestions as to how dry season births may be adaptive. Methods The bat specimens from Peru housed in the col- lections of the American Museum of Natural His- tory, Field Museum of Natural History, Louisiana State University Museum of Zoology, National Museum of Natural History, and the Texas Co- operative Wildlife Collection were examined in late 1977 and early 1978 for information on re- production. Data were taken directly from Tuttle (1970) and Bowles et al. (1979) for specimens not in the above collections but included in their re- ports. Information on reproductive condition of bats was also obtained from the notes of A. L. Gardner and J. L. Patton. Fieldwork at several localities in Peru, conducted from June through August 1977 and from June through November 1978, enabled me to collect and record the repro- ductive data for many of the bats now in the Lou- isiana State University collection. Diet informa- tion was taken from the literature (Heithaus et al., 1975; Paradiso, 1975; Gardner, 1977) and is given in Graham ( 1 983). Nomenclature follows Graham (1983) except for Artibeus glaucus, which I now consider a distinct species. Recorded data consisted of species identifica- tion, locality, elevation, sex, age, and macroscopic reproductive condition. Individuals with incom- pletely ossified phalangeal epiphyses were classi- fied as juveniles. Females were designated as preg- nant or lactating if this information was included on specimen tags or indicated by dissection of fluid preserved females. The crown-rump lengths of embryos (including extra-embryonic membranes) were either measured (in mm) or taken from spec- imen tags. Length of testes was also noted when it was recorded on sp>ecimen tags, but was not used to establish reproductive patterns because of un- certainties in the relationship between testes size and sexual activity (Taddei, 1976; Thomas, 1972). Monthly pregnancy frequencies are based only on those female SF>ecimens for which I am confident that the presence (or absence) of embryos had been properly recorded. This limitation excludes most of the specimens collected prior to 1960. I classified each species with adequate data (those with evidence of at least one birth period) into those that can give birth in the dry (May-Septem- ber) and/or wet (October-April) seasons. I as- 174 FIELDIANA: ZOOLOGY sumed that a birth between 1 May and 3 1 August also indicates weaning of the young during the dry season. Species with births recorded only for the month of September (usually a dry month) were not included in either breeding season, since these young could be weaned either as the dry season ended or as the wet season began. The above clas- sifications were accomplished by determining the distribution of births across all months, as indi- cated by the distribution of juveniles, lactating fe- males, and embryo sizes. If a single birth was in- dicated for a given season, then that species was identified as having the potential to produce young during that season. The presence of juveniles or lactating females was assumed to indicate partu- rition during the month prior to capture. If a fe- male was carrying a large, near-term embryo, par- turition was assumed to take place in that month. On the other hand, if the embryo was very small relative to the adult body length, birth was as- sumed to take place four months later for most species, three months later for small insectivorous species (Findley & Wilson, 1 974), and five months later for emballonurids (Bradbury & Vehrencamp, 1977) and Desmodus rotundus (Wilson, 1979). Classification of a species as one that can give birth during the dry season does not necessarily mean that it does so each year. I was unable with this method to determine the actual frequency of births during either season. A dry season classi- fication simply means that individuals can at least occasionally produce young during the dry season. The lack of birth records for a given species for either season can be the result of an actual absence of birthing during that season, or it may be due to an inadequate sample from that season. If I as- sume that the lack of a breeding record for a given season is not due to inadequate sampling, then I can use the G test (Sokal & Rohlf, 1981) to assess the significance of the differences between the groups of species compared. This is a safe as- sumption, since most of the specimens were col- lected during the Peruvian dry season (the season with which this study is principally concerned) and because the groups of bats predicted to have the greatest proportion of dry season births are the groups most poorly sampled (see Appendix and below). Regions and Climate I follow Koopman's ( 1978) division of Peru into three zoogeographical regions. He lists SF)ecies as lowland if they were collected below 1000 m east of the Andes and highland if collected above this elevation. Pacific coastal species are those col- lected either along the arid coast or adjacent An- dean foothills (including the mesic areas of north- western Peru). The Pacific coastal plain is characterized by low (< 50 mm) annual rainfall, most of which falls between December and April. The mesic areas on the northwestern Andean slopes experience great- er rainfall over an extended period. Rainfall east of the Andes (fig. 1 ) is seasonal and abundant, with the greatest amounts deposited between the ele- vations of 1 000 and 3000 m (Bowman, 1916). The wet season begins in October and continues through April and the dry season begins in May and ex- tends through September. Most investigators agree that food resources are primarily modulated by rainfall seasonality (Jan- zen, 1967, 1973; Frankie et al., 1974; Ricklefs, 1975; Buskirk & Buskirk, 1976; Wolda, 1978a,b, 1982; Foster, 1982; Smythe, 1982; Terborgh, 1983). Many of the following generalizations on seasonal changes in food supplies for bats of mid- dle and higher elevations are based on the as- sumption that Peruvian plants and insects respond to environmental conditions in a manner similar to closely related organisms in Costa Rica, Pan- ama, or Puerto Rico (as described by the above authors). Terborgh's (1983) study of the changes in fruit, flower, and insect supplies in Manu Na- tional Park indicates that in the lowlands of south- eastern Peru, fruit abundance and diversity usually increase in October with the onset of the wet sea- son, peak in November and December and again in March, and decrease to the lowest level in May and June. Not only is the first peak greater but it also includes a greater proportion of plants in the family Moraceae (Terborgh, pers. comm.), which have fruit favored by bats (Gardner, 1 977). Flower abundance peaks in the dry season, but some flow- ers are present throughout the year (Terborgh, 1983). At middle to higher elevations, the phe- nology of fruit and flower species is probably sim- ilar to lowland species, but less seasonal (Ter- borgh, 1977; Nevling, 1971). Changes in insect resources are more difficult to generalize because the species in different size classes (Smythe, 1982; Wolda, 1982) and in different habitats (Bradbury & Vehrencamp, 1976; Terborgh, 1983; Janzen, 1983) have different periods of peak abundance and diversity. If the supply across size classes and habitats is considered, then it is probably rather high throughout the year. I will not discuss the GRAHAM: PERUVIAN BATS 175 600. 500. 400. 5 300. I 200_ 100. Month Fig. 1. Average rainfall amounts at Iquitos (lowland) and Yurac (middle el- evation), Department of Loreto, Peru. Data are taken from a map published in 1971 by the Servicio Nacional de Me- teorologia e Hidrologia in Peru and are averaged from 1 0 years of records. seasonal changes in food levels west of the Andes in Peru because less is known of this region. Results More than 3,400 female specimens divided among 109 species were included in this study. Twins were recorded for only one female, a Car- ollia perspicillata, collected in November bearing two well-developed embryos (24 and 29 mm). Specimens were collected almost exclusively be- tween May and December, with June through Au- gust having the largest samples (see sample sizes in Appendix). The reproductive records are also unevenly distributed among the species (Appen- dix). These sampling problems made it difficult to identify the reproductive patterns for most species. If, however, the percentages of all the females that were pregnant are determined for each species for each month, many species have data that fit the pattern of seasonal polyestry. This pattern is well illustrated by the pregnancy curve ofCarollia per- spicillata (fig. 2). Pregnancy levels peak in Septem- ber and December and are followed by periods of parturition, as suggested by the October drop in the frequency of pregnant females and by the ju- veniles collected in January (Appendix). The be- ginning of the second reproductive cycle of the season is also indicated by the females of Carollia castanea (l), Vampyrops dorsalis (7), and Artibeus planirostris (1) that were simultaneously lactating and pregnant in November and December (Ap- pendix). On the other hand, the pregnancy curve of Myotis (fig. 2) and the records of juveniles and lactating females of Carollia (fig. 2) and Artibeus (Appendix) for May-September indicate that births can occur outside of the wet season. Individuals of some species (i.e., Desmodus rotundus, Myotis nigricans, and perhaps Glossophaga soricina and Artibeus planirostris; see Appendix) may be able to produce young during any month of the year. Table 1 lists the number of species in each fam- ily or subfamily with births recorded during the seasons. I was able to identify birth periods for 79 (72.5%) of the 109 species listed in the Appendix. Most species (63 of the 79. 79.7%) have birth rec- ords for the wet season but a surprisingly large proportion of all of the species (46 out of 79, 58.2%) have records of dry season parturition. The ab- sence of birth records during the wet season 176 HELDIANA: ZOOLOGY Carollia perspiclllata 60-1 0 0 0 3 3 24 38 100 18 15 23 15 Fig. 2. Changes in pregnancy levels of Carollia perspiclllata and Myotis nigricans. The numbers across the top refer to the sample sizes for each month. The ordinate represents the percentage of all females for each month that were recorded as pregnant. (Mormoopidae) and the dry season (Furipteridae and Thyropteridae) may represent real periods of no births but may also be the result of inadequate sampling. Are there groups of bats that have a greater ten- dency than other groups toward parturition during the dry season? If seasonal fluctuations in food supplies are less in the highlands than in the low- lands, then proportionately more highland than lowland species could be expected to produce and wean young during the dry season and, if most species are polyestrous, during both seasons. Thir- ty-one of the 58 lowland species (53.4%) and 17 of the 27 highland species (63.0%) have records of dry season births (table 2). Although the 9.6% difference is in the predicted direction, it is not significant {P > 0.25). The highland region has a greater proportion of species with birth records from both seasons (51.9% vs. 34.5%, P < 0.05). The highland Sturnira bogotensis that were lac- tating and pregnant in June and August (Appen- dix) provide evidence that some species with dry season birth records are polyestrous, becoming pregnant again during the dry season. Bats of the Pacific coastal and slope region also show a strong tendency toward dry season births. If trophic generalists (those that consume more than one type of food, such as fruit and insects) can switch to another resource when one type be- comes scarce, they should be less vulnerable to seasonal fluctuations in their food supply than are trophic specialists (those species that use only one major type of food). This reduced seasonality of food resources should be reflected by a tendency for births to occur during the dry season and, if polyestry is common, for both seasons. Nine of the 1 1 generalist species (81.8%) and 23 of the 68 specialists (33.8%, table 2) have records of indi- viduals that have given birth during the dry sea- son, and relatively more generalists (63.6%) than GRAHAM: PERUVIAN BATS 177 Table 1. Number of species of Peruvian bats with births recorded for each season. Family Dry Season* Wet Both Total spe- ciest Emballonuridae 1 3 4 Noctilionidae 2 1 1 2 Mormoopidae Phyllostomidae Phyllostominae Glossophaginae Carolliinae 2 6 6 2 6 4 5 3 3 2 2 9 7 5 Stumirinae 4 6 4 6 Stenodermatinae 10 22 10 22 Desmodontinae 1 2 1 2 Furipteridae Thyropteridae Vespertilionidae Molossidae 7 5 1 1 7 5 4 2 1 1 10 8 Totals 46 63 30 79 * The number of species with records of birth in either the dry or the wet season or in both. A species is listed in all three columns if it has birth records for both sea- sons. t The total number of species that have data indicating birth periods. The sum of the dry and wet season records does not equal the total, because some species have rec- ords for both seasons. specialists (29.7%) have records for births during both seasons (table 2). These large proportional differences are significant (P < 0.01 and P < 0.05, respectively). Since floral resources are greatest in the dry sea- son when fruit resources are lowest, nectarivorous species should show a greater tendency for indi- viduals to produce and wean young during the dry season compared to frugivores. Six of the seven (85.8%) bats that are at least partially nectarivo- rous have birth periods in the dry season com- pared with less than half (1 7 of 40, 42.5%) of those that are partially frugivorous (table 2). This large relative difference is significant (P < 0.05). Dry season production and weaning of young is prob- ably the common pattern for most bats that eat nectar. This is indicated by the large number of juveniles (13) and lactating females (35) recorded for this group from May through August. Since the resource abundances are reversed during the wet season, the pattern of frugivorous sp>ecies hav- ing relatively more birth records (39 of the 40 species that are partially frugivorous, 97.5%) than nectarivores (3 of the 7 species that are partially nectarivorous, 42.9%) was expected {P < 0.001). Percentages of the species in these trophic groups that have birth records for both seasons are similar (40.0% and 42.9% for frugivores and nectarivores, resp)ectively, P > 0.50). The number of insectivorous species that have birth records for each season is almost equal (17 for the dry, 1 8 for the wet). This is the expected pattern if insect abundance and diversity remain high throughout the year. If a species is seasonally polyestrous, the prob- ability of recording a birth for the wet season for that species is greater, since both young are pro- duced during that season. If most species are sea- sonally polyestrous, then the majority of species should have wet season birth records. All of the ecological groups mentioned above (except the nectarivores) show considerable evidence of par- turition during the wet season (table 2). Discussion Seasonal polyestry is the dominant pattern in other studies of Neotropical bat reproduction (Hemingetal., 1972; Thomas, 1972;Taddei, 1976; LaVal & Fitch, 1977; Myers, 1977; Bonaccorso, 1979; Wilson, 1979; August & Baker, 1982). The majority of bats in Peru, especially frugivores, give birth during the wet season. Although this pattern is expected if most species are seasonally polyestrous most of the time, this conclusion is compromised by the possibility that some species may be monestrous, producing and weaning their young during the wet season. I was unable, for most species, to distinguish between these two re- productive patterns. However, the females col- lected from November and December that were simultaneously lactating and pregnant provide evidence for seasonal polyestry. Females that fol- low this pattern wean their first young concur- rently with the onset of the heavy rains, as fruit supplies are reaching their peak, and wean their second young near the middle of the wet season when food resources remain plentiful. In Peru, as in many of the other Neotropical areas discussed in the above studies, young bats are at least occasionally bom and weaned in the dry season. These bats are members of zoogeo- graphical and ecological groups whose tendencies toward dry season births can be predicted. If food availability for a particular group of bats does not fluctuate greatly throughout the year relative to a second group, then the first group should include a larger percentage of species with birth records for the dry season and, if the species are polyes- 178 HELDIANA: ZOOLOGY Table 2. Number of species of Peruvian bats with births recorded for each season, by geographical region and ecological group. Season* Region Dry Wet Both Total and group % (N) % (N) % (N) species* Geographical Regions Lowlands 53.4(31) 79.3 (46) 34.5(19) 58 Highlands 63.0(17) 88.8 (24) 51.9(14) 27 Pacific coastal 80.0 (8) 60.0 (6) 30.0 (3) 10 Trophic Groups Insectivores 58.6(17) 62.1 (18) 22.2 (6) 29 Frugivores 38.7(12) 100 (33) 38.7(12) 33 Nectarivores 66.7 (2) 33.3(1) ... 3 Piscivores 100 (1) ... 1 Vampires 50.0(1) 100 (2) 50.0(1) 2 Frugivore/insectivores 60.0 (3) 80.0 (4) 40.0 (2) 5 Nectarivore/insectivores 100 (3) 66.7 (2) 66.7 (2) 3 Piscivores/insectivores 100 (1) 100 (1) 100 (1) I Omnivorest 100 (2) 100 (2) 100 (2) 2 Trophic Specialists:}: 33.8 (23) 79.4 (54) 29.7(19) 68 Trophic Generalists§ 81.8(9) 90.9(10) 63.6 (7) 11 * See notes to Table 1 . N = Number of species; % = % of total species. t Includes species that eat more than two major types of items, t Includes insectivores through vampires (see text, p. 1 77). § Includes frugivores/insectivores through omnivores. trous, for both seasons. Two groups of Peruvian bats, highland species and trophic generalists, pos- sibly experience less seasonal fluctuation in food supplies relative to lowland species and trophic specialists. However, the reasons for the damp- ened fluctuations differ with each group. At high elevations in Peru, even during the dry season, clouds cover the forests most of the time (Terborgh, 1977; pers. obs.), providing moisture that is largely unavailable to lowland plants. This yearlong availability of moisture presumably keeps plant productivity and ultimately food availability from fluctuating as greatly as in the lowlands. This reduced fluctuation in food supplies is reflected by a weak trend toward relatively more highland birth records for the dry season and by a stronger high- land trend for birth records for both seasons. Species with generalized feeding habits are thought to be able to switch resources when one becomes reduced. This dampens the effect of sea- sonal changes in food availability and explains their larger percentages of birth records for the dry season and for both seasons. Fleming et al. ( 1 972), Heithaus et al. ( 1 975), and Bonaccorso ( 1 979) pro- vide data from Costa Rica and Panama that il- lustrate seasonal switches made by several of the same species included in my study as feeding gen- eralists (i.e., Phyllostomus discolor and Glossoph- aga soricina). Relatively more nectarivorous than frugivorous species have individuals with dry season birth and weaning records, but the records for both seasons are not different. These were the expected patterns, since both groups presumably experience seasonal variations in their food supplies; floral resources are greatest and fruit resources lowest during the dry season (Terborgh, 1983). Hence, dry season births may actually be the rule for bats that con- sume nectar but the exception for frugivores. The reverse is true for the wet season. These seasonality differences explain the almost equal proportions of species with birth records for both seasons. Thus, for trophic generalists, highland, and nec- tarivorous bat species, many females produce their first young in the dry season and become pregnant again, producing their second young during the wet season. Obviously, many more data are need- ed to clarify these patterns and to address the pos- sibility of differential survival of young between the two seasons. In Peru, the onset and termination of the rainy season can vary annually (Terborgh, 1983). Food supplies track these variations (Terborgh, 1983; Foster, 1982), and so may bat reproduction. I do not believe that climatic variability is the principal factor responsible for the births recorded in Peru between May and August because the ecological groups that I compared (frugivores and nectari- GRAHAM: PERUVIAN BATS 179 vores, trophic specialists and generalists) showed different seasonal birth patterns. If unusual dry season rains were the principal factor, then equal proportions of the different ecological groups should have responded by producing young during the normally dry season. My data support the hypothesis that dry season births are adaptations for producing and perhaps weaning young when food resources are available predictably or occasionally during this time. These patterns need to be confirmed by long term studies of actual bat communities, and of the resources used by individuals at both highland and lowland localities and during both seasons. Acknowledgments I am grateful to the many people who assisted me in Peru and in the United States. Antonio Brack E., Eric Cardich Briceno, Richard Bustamante M., and Susana Moller-H. of the Direccion General Forestal y de Fauna, Ministerio de Agricultura, Lima, provided the necessary collecting permits. Hernando de Macedo R. and Ramon Ferreyra of the Museo de Historia Natural "Javier Prado" were also helpful. I thank Arturo Koenig R., Man- uel A. Plenge, and Gustavo del Solar for their hospitality and help in making this study possible. I appreciate the assistance provided by Faucet and Aero Peru. Without the reliable assistance of my Peruvian associates, Manuel Sanchez, Klaus Wehr, and Reyes Rivera, the fieldwork would have been much more difficult. Karl F. Koopman (American Museum of Nat- ural History), Don E. Wilson (National Museum of Natural History), David J. Schmidly and Wil- liam B. Davis (Texas Cooperative Wildlife Col- lection), and Patricia W. Freeman (formerly of Field Museum of Natural History) were very help- ful during trips to their museums. Koopman and James L. Patton (Museum of Vertebrate Zoology) loaned fluid-preserved bats. James B. Cope (Jo- seph Moore Museum), Alfred L. Gardner (Na- tional Museum of Natural History), and Patton were kind enough to send me unpublished manu- scripts and field notes. For valuable assistance in the field I thank Linda J. Barkley, J. William Eley, Gary R. Graves, John P. O'Neill, Theodore A. Parker, III, Thomas S. Schulenberg, and Morris D. Williams. The excellent suggestions on the manuscript by Philip Myers were appreciated. I thank Susan T. Graham for her support during the several revisions of this work. 1 gratefully acknowledge the financial support of the LSUMZ Peruvian fieldwork by John S. Mcllhenny, Irving and Laura R. Schweppe, E. W. Mudge, and the late Babette M. Odom. Travel to other museums was made possible in part by an LSU Foundation-Graduate Student Travel Award. Literature Cited August, P. V., and R. J. Baker. 1982. Observations on the reproductive ecology of some Neotropical bats. Mammalia, 46: 177-181. BoNACcoRso, F. J. 1979. Foraging and reproductive ecology in a Panamanian bat community. Bulletin of the Florida State Museum, Biological Sciences, 24: 359^08. Bowles, J. B., J. B. Cope, AND E. A. Cope. 1979. Bi- ological studies of selected Peruvian bats of Tingo Maria, Departamento de Huanuco. Transactions of the Kansas Academy of Sciences, 82: 1-10. Bowman, I. 1916. The Andes of Southern Peru: Geo- graphical Reconnaissance Along the Seventy-Third Meridian. H. Holt and Co., New York, 336 pp. Bradbury, J. W.. and S. L. Vehrencamp. 1 976. Social organization and foraging in emballonurid bats. I. Field studies. Behavioral Ecology and Sociobiology, 1:337- 381. . 1977. Social organization and foraging in em- ballonurid bats. IV. Parental investment patterns. Be- havioral Ecology and Sociobiology, 2: 1 9-29. BusKiRK, R. E., and W. H. Buskjrk. 1976. Changes in arthropod abundance in a highland Costa Rican forest. American Midland Naturalist, 95: 288-298. FiNDLEY, J. S., AND D. E. WiLSON. 1974. Observations on the Neotropical disk-winged bat, Thyroptera tri- color Spi\. Journal of Mammalogy. 55: 562-571. Fleming, T. H., E. T. Hooper, and D. E. Wilson. 1972. Three Central American bat communities: Structure, reproductive cycles, and movement patterns. Ecology, 53: 555-569. Foster, R. B. 1982. The seasonal rhythm of fruitfall of Barro Colorado Island, pp. 151-172. In Leigh. E. G., Jr., A. S. Rand, and D. M. Windsor, eds.. The Ecology of a Tropical Forest. Smithsonian Institution Press, Washington, D.C.. 468 pp. Frankie, G. W., H. G. Baker, and P. A. Opler. 1974. Comparative phenological studies of trees in tropical wet and dry forests in the lowlands of Costa Rica. Journal of Ecology, 62: 881-919. Gardner, A. L. 1977. Feeding habits, pp. 293-350. In Baker, R. J., J. K. Jones, Jr., and D. C Carter, eds.. Biology of bats of the New World family Phyllosto- matidae. Part II. Special Publications, The Museum, Texas Tech University, 13: 1-364. 180 HELDIANA: ZOOLOGY Graham, G. L. 1983. Changes in bat species diversity along an elevational gradient up the Peruvian Andes. Journal of Mammalogy, 64: 559-571. Heithaus, E. R., T. H. Fleming, and P. A. Opler. 1975. Foraging patterns and resource utilization in seven species of bats in a seasonal tropical forest. Ecology, 56:841-854. Humphrey, S. R., and F. J. Bonaccorso. 1979. Pop- ulation and community ecology, pp. 409-441. In Ba- ker, R. J., J. K. Jones, Jr., and D. C. Carter, eds., Biology of bats of the New World family Phyllosto- matidae. Part III. Special Publications, The Museum, Texas Tech University, 16: 1-441. Janzen, D. H. 1967. Synchronization of sexual repro- duction of trees within the dry season in Central Amer- ica. Evolution, 21: 620-637. . 1973. Sweep samples oftropical foliage insects: Effects of seasons, vegetation types, elevation, time of day, and insularity. Ecology, 54: 687-708. . 1983. Insects: Introduction, pp. 619-645. In Janzen, D. H., ed., Costa Rican Natural History. The University of Chicago Press, Chicago, 816 pp. Koopman, K. F. 1978. Zoogeography of Peruvian bats with special emphasis on the role of the Andes. Amer- ican Museum Novitates, 2651: 1-33. LaVal, R.K., AND H.S. Fitch. 1977. Structure, move- ments and reproduction in three Costa Rican bat com- munities. Occasional Papers, Museum Natural His- tory, University of Kansas, 69: 1-27. Mares, M. A., and D. E. Wilson. 1971. Bat repro- duction during the Costa Rican dry season. Bio- Science, 21: 471^77. Myers, P. 1977. Patterns ofreproduction of four species of vespertilionid bats in Paraguay. University of Cal- ifornia Publications in Zoology, 107: 1-41. Myers, P., and R. M. Wetzel. 1983. Systematics and zoogeography of the bats of the Chaco Boreal. Mis- cellaneous Publications, Museum of Zoology, Uni- versity of Michigan, 165: 1-59. Nevling, L. I., Jr. 1971. The ecology of an elfin forest in Puerto Rico, 16. The flowering cycle and an inter- pretation of its seasonality. Journal of the Arnold Ar- boretum, 52: 586-613. Paradiso, J. L., ED. 1975. Walker's Mammals of the World, 3rd ed., vols. 1 and 2. Johns Hopkins Uni- versity Press, Baltimore, 1,500 pp. Racey, p. a. 1 982. Ecology of bat reproduction, pp. 57-104. In Kunz, T. H., ed.. Ecology of Bats. Plenum Press, New York, 425 pp. RiCKLEFS, R. E. 1975. Seasonal occurrence of night- flying insects on Barro Colorado Island, Panama Canal Zone. Journal of the New York Entomological Society, 83: 19-32. Smythe, N. 1982. The seasonal abundance of night- flying insects in a Neotropical forest, pp. 309-318. In Leigh, E. G., Jr., A. S. Rand, and D. M. Windsor, eds.. The Ecology of a Tropical Forest. Smithsonian Insti- tution Press, Washington, D.C., 468 pp. SoKAL, R. R., AND F. J. Rohlf. 1981. Biometry, 2nd ed. W. H. Freeman, San Francisco, Calif, 859 pp. Taddei, V. A. 1976. The reproduction of some Phyl- lostomidae (Chiroptera) from the northwestern region of the State of Sao Paulo. Boletim de Zoologia, Uni- versidade de Sao Paulo, 1: 313-330. Terborgh, J. 1977. Bird species diversity on an An- dean elevational gradient. Ecology, 58: 1007-1019. . 1983. Five New World Primates. Princeton University Press, Princeton, N.J., 260 pp. Thomas, M. E. 1972. Preliminary study of the annual breeding patterns and population fluctuations of bats in three ecologically distinct habitats in southwestern Colombia. Ph.D. diss., Tulane University, New Or- leans, La., 161 pp. TuTTLE, M. D. 1970. Distribution and zoogeography of Peruvian bats, with comments on natural history. University of Kansas Science Bulletin, 49: 45-86. TuTTLE, M. D., AND D. Stevenson. 1 982. Growth and survival of bats, pp. 105-150. In Kunz, T. H., ed., Ecology of Bats. Plenum Press, New York, 425 pp. Wilson, D. E. 1979. Reproductive patterns, pp. 3 1 7- 378. In Baker, R. J., J. K. Jones, Jr., and D. C. Carter, eds., Biology of bats of the New World family Phyl- lostomatidae. Part III. Special Publications, The Mu- seum, Texas Tech University, 16: 1-441. WoLDA, H. 1978a. Fluctuations in rainfall, food and abundance oftropical insects. Journal of Animal Ecol- ogy, 47: 369-381. . 1978b. Fluctuations in abundance of tropical insects. American Naturalist, 112: 1017-1045. . 1982. Seasonality of Homoptera on Barro Col- orado Island, pp. 319-330. In Leigh, E. G., Jr., A. S. Rand, and D. M. Windsor, eds., The Ecology of a Tropical Forest. Smithsonian Institution Press, Wash- ington, D.C., 468 pp. GRAHAM: PERUVIAN BATS 181 s I a o - & g i I i I I 1 Ci — — § "" ^^ ^ ^ _ -^ ^^ ,«i^ ^^ .^-^ ir> ■* o VO 7^ ^m^ M f^ ir> (N ^^ o f^ ■* o OS O 00 f<^ vo iri r> Tf 00 .^ik^ ^-^ ^-^ ,^i^ ■* 00 (N 00 . . o ^^ f«^ O "5 ^ c ■«: B '^ c: ft, vj H K ^ ^ ^ C .s ft. ft.^ S^ S^ ft. ^ U 5 3 .J U Qu U o: :©:: :o: • • .; • ■ • -ooo- • o u X hJ hJ U u ^ ^ ^ U u cu u X u .J -j u u u u .'^ .'^ c -2 -2 111 Z ft. ft^ 3 3 S3 I o o o o - c s: c c "OOP o 1 1? ? ? 3 If t?.s 5 S 5 3'^§ a ."> .'^ .'o .'^ .'^ .'^ 2 C C C C C C - i» ii» w ^ ^ %j ^ o 5; a: a: (S; ^ 6^ O ft. s: k: p o c c c 2 P P P ■? 5 C a a 5 :* .Vj .U 3 .5 o to ^ 3 s a bo ft, ts c 2 •^-2 2 S -2 S c c ^ — -2 -2 -2 -2 -2 -2 ;: 13 := o ti^^^^^ 3 3a pot • 0 e-. Q^QQ^c^. QQQ ^ : o : : o — O Tf c^ v y—s^-s ^-s On — . 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3 ^ (^ GRAHAM: PERUVIAN BATS 185 I' ^ e-. ^ e-. C-. e-- Q ^ Q ^ Q Q c^. Q . m — • g2 O O s a M 9 ^J^^ f^:^:^^ s s s I ^ ^ 1^ ? S S -S .'^ c ^ ^ 'J "2 ^ <3 53 CiO Ci. g to to » c^ i 5 5 s a a; uj t*j uj ttj a •c 3 •o Ka a, -c CL, m c II « o S ^ T3 00 c s < £ x^ o E w « 00 S o .2 u ^ u tu Q. a Q.-= JQ 2 •o U (A - Si ifi c a* ti C = E C/5 186 FIELDIANA: ZOOLOGY Tent Construction by Bats of the Genera Artibeus and Uroderma Robert M . Timm ABSTRACTS Herein, I describe new styles of tents cut and utilized by Artibeus anderseni, A. glaucus, A. gnomus, A. phaeotis, A. toltecus, A. watsoni, Uroderma bilobatum, and U. magnirostrum; review and summarize the literature on tent use by Artibeus and Uroderma; and discuss the effectiveness of tents as diurnal roosts. Artibeus anderseni alters the shape of Heliconia leaves by cutting the lateral nerves and interconnected tissue extending out from the midrib. Artibeus glaucus cuts the basal lateral nerves in Xanthosoma, causing the two sides of the leaf to collapse downward around the midrib. Artibeus phaeotis cuts the lateral nerves and interconnected tissues in both banana and Heliconia imbricata; the basal cuts veer out from the midrib such that a distinctive V-shaped enclosure is formed by the hanging leaf Artibeus toltecus cuts the basal nerves on Anthurium, causing the sides of the leaf to fold down around the midrib to form a pyramid- shaped tent. Artibeus watsoni was found to make four distinctive styles of tents, including simple V-shaped cuts on bifurcated palms, cuts of a few side veins on aroids to produce a rounded pyramid, elongate J-shaped cuts on banana and Heliconia, and polygonal cuts on Carludovica pal mat a. Artibeus watsoni has the greatest repertoire in tent styles, and uses the most diverse array of plant species and leaf shapes. Two styles of tents constructed by Uroderma bilobatum are reported for the first time, one on the large pinnately leafed palm Scheelea rostrata and the second on banana. The common denominator between the Uroderma bilobatum tents reported herein and those previously described is that all are on large, broad leaves and all have a distinctive V-shaped pattern cut by the bats. Uroderma magnirostrum also creates an inverted elongate V-shaped tent on pinnately leafed palms. All New World tent-makers described to date are tropical members of the phyllostomid subfamily Stenoderminae. Each species of tent-making bat has one or more distinctive style of tent. Bats select leaves of specific shapes, sizes, and angles for tent construction. Most species appear to be obligate tent-roosters. Tents provide bats with a cryptic diurnal roost site, in addition to providing shelter from both the sun and rain and an early warning to the approach of predators. Aqui yo describo nuevos estilos de carpas cortadas y utilizadas por Artibeus anderseni, A. glaucus, A. gnomus, A. phaeotis, A. toltecus, A. watsoni, Uroderma bilobatum, y U. magnirostrum reviso y hago un sumario de la literatura acerca del uso de carpas por filostomidos; y discuto la efectividad de las carpas como perchas diumas. Artibeus anderseni altera la forma de las hojas de Heliconia cortando las nervaduras centrales y tejido interconectado que se extiende From the Division of Mammals, Field Museum of Natural History, Chicago, Illinois 60605-2496. The au- thor's present address is Museum of Natural History and Department of Systematics and Ecology, University of Kansas, Lawrence, Kansas 66045. TIMM: ARTIBEUS AND URODERMA 187 exteriormente desde la nervadura central. Artibeus glaucus corta la base de las nervaduras laterales en Xanthosoma, causando asi que los dos lados de la hoja colapsen hacia abajo al rededor de la nervadura central. Artibeus phaeotis corta las nervaduras laterales y tejidos in- terconectados en banana y Heliconia imbricata; los cortes basales viran hacia afuera desde la nervadura central de tal modo que una distintiva cavidad en forma de V es formada por la hoja colgante. Artibeus toltecus corta las nervaduras basales de Anthurium, adi que los lados de la hoja doblen en redor de la nervadura central de tal modo que una cavidad es formada en forma de una piramide. Artibeus watsoni fue encontrada haciendo cuatro distintos estilos de carpas, incluyendo simples cortes en forma de V en palmas bifurcadas, cortes en unas pocas nervaduras laterales en araceas para producir una piramide redondeada, cortes alargados en forma de J en banana y Heliconia. y cortes poligonales en Carludovica pal mat a. Artibeus watsoni tiene el mas grande repertorio en estilos de carpas y usa la mas diversa serie de plantas y formas de hojas. Dos estilos de carpas construidas por Uroderma bilobatum son reportadas por primera vez; una en la larga y pinnada hoja de palma Scheelea rostrata y la segunda en banana. El comun denominador entre carpas de Uroderma bilobatum repnirtadas aqui y aquellas previa- mente descritas es que todas usan hojas grandes y anchas y todas tienen un distintivo patron en forma de V cortado por los murcielagos. Uroderma magnirostrum tambien corta una carpa en forma de V-invertido en hojas pinnadas de palmas. Todos los filostomidos cortadores y utilizadores de carpas descritos del Nuevo Mundo son membros tropicales de la subfamilia Stenoderminae. Cada especie tiene uno o mas estilos distintos de carpas. Los murcielagos escogen hojas de formas especificas, y constroen sus carpas en angulos peculiares. La mayor parte de las especies parecen utilizar las carpas obligatoriamente. Las carpas oferecen una percha oculta durante el dia, asi como un abrigo del sol, de la lluvia, y de predadores. Neste trabalho, (1) descrevo novos estilos de tendas cortadas e utilizadas por Artibeus an- derseni, A. glaucus, A. gnomus, A. phaeotis, A. toltecus, A. watsoni, Uroderma bilobatum, e U. magnirostrum; (2) reviso e resumo a literatura sobre o uso de tendas pelos morcegos da familia Phyllostomidae, e (3) discuto a eficiencia de tendas como alojamentos diumos. Artibeus an- derseni altera a forma das folhas de Heliconia, cortando as veias laterais e os tecidos interligados que estendem da veia central. Artibeus glaucus corta as veias basilares laterais em Xanthosoma, causando com que os dois lados da folha caiam contra o centro. Artibeus phaeotis corta as veias laterais e os tecidos interligados nas folhas de bananas e de Heliconia imbricata. Os cortes basilares partem da veia central, formando um abrigo distinto em forma de "V". Artibeus toltecus corta as veias basilares de Anthurium, causando com que os lados da folha dobrem em volta da veia central, criando um abrigo em forma de piramide. Artibeus watsoni constroi quatro tipos diferentes de tendas, incluindo simples cortes em forma de "V" em folhas de palmeiras bifurcadas; cortes em algumas das veias laterais em folhas de trepadeiras (resultando em piramides redondas); cortes alongados, em forma de "J", em folhas de bananas e de Heliconia; e cortes poligonos em fdlhas de Carludovica palmata. Artibeus watsoni possue o maior repertorio de estilos de tendas, e usa o conjunto mais di verso de especies de plantas e de configura96es de folhas. Dois estilos de tendas construidas por Uroderma bilobatum sao descritos pela primeira vez; um nas fdlhas grandes da palmeira Scheelea rostrata, e outro nas folhas de bananas. Fatores comuns entre as tendas construidas por Uroderma bilobatum aqui descritas, e as descritas previamente, sao a forma distinta em "V" cortada pelos morcegos, e o uso de folhas grandes e largas para a construfao das tendas. Uroderma magnirostrum tambem constroe tendas em forma de "V" invertido nas folhas de palmeiras. Todos morcegos construidores de tendas no Novo Mundo p)ertencem a subfamilia Steno- derminae (familia Phyllostomidae), e cada especie exibe um ou mais estilos caracteristicos de constru9ao. A maioria destas especies de morcegos parecem alojar-se obrigatoriamente em tendas, as quais oferecem nao so um abrigo diumo camuflado, mas tambem prote9ao contra sol, chuva, e predadores. 188 HELDIANA: ZOOLOGY Introduction The use of cut leaves for diurnal roosting sites by bats was first described by Thomas Barbour (1932), who reported on diurnal roosts of Uro- derma bilobatum near the Panama Canal. He found these bats roosting under the leaves of two culti- vated palms identified as Livistona chinensis and Prichardia pacifica. Not only were these bats found roosting under palm fronds, but they had also al- tered the leaf to produce a diurnal roosting struc- ture. Barbour (1932, p. 307) stated that "by nip- ping the ridges of the plications on the under side the leaf is weakened and as the bitten spots are skillfully and serially distributed the leaf finally is sufficiently weakened so that the distal portion droops sharply downward." Chapman (1932, p. 555) discovered Artibeus watsoni roosting under the cut veins of a bifurcated palm, Geonoma cu- neata (reported as G. decurrens), on Barro Colo- rado Island, Panama, and first called these mod- ified leaves "tents." Ingles (1953) also reported A. watsoni constructing tents in the palm Geonoma oxycarpa (reported as G. binervia) on Barro Col- orado Island. Goodwin and Greenhall (1961, p. 262) found Artibeus cinereus roosting "under cut leaves of palm trees and on the under side of ba- nana leaves" and Uroderma bilobatum roosting under cut leaves of the carat palm, Sabal glau- cescens, and coconut palm, Cocos nucifera, on Trinidad. Ectophylla alba was reported by Timm and Mortimer (1976) to alter the leaves of five species of Heliconia in Costa Rica; the bats se- lected specific leaves for both size and angle to the ground. Artibeus jamaicensis was found by Foster and Timm (1976) roosting under the cut leaflets on a pinnately leafed palm, Scheelea rostrata, in a tropical dry forest in Costa Rica. Recently, Timm ( 1 984) reported tent construction by another phyl- lostomid, Vampyressa pusilla, in Costa Rica, and Koepcke (1984) found Mesophylla macconnelli utilizing similar tents in Peru. Only one Old World bat, Cynopterus sphinx (Pteropodidae), has been reported to alter the shape of palm leaves to pro- duce a diurnal roosting structure (Goodwin, 1 979). Reviews of roosting site selection by bats were provided by Tuttle (1976) and Kunz (1982). To date, seven species of phyllostomids {Arti- beus cinereus, A. jamaicensis, A. watsoni, Ecto- phylla alba, Mesophylla macconnelli, Uroderma bilobatum, and Vampyressa pusilla) have been re- ported to modify leaves of plants to produce diur- nal roosting sites herein referred to as tents. All are tropical members of the phyllostomid subfam- ily Stenodermatinae. The phyllostomid genus Artibeus, which in- cludes some 1 5 species, is widespread in the Noo- tropics from northern Mexico southward to Ar- gentina and Chile. These bats range in size from 10 g (A. anderseni and A. watsoni) to 70 g (A. lituratus). Uroderma, a closely related genus of medium-sized stenodermatines that includes only two species, is found from southern Mexico through the Amazon Basin of South America. The better known of the two species, U bilobatum, weighs from 13 to 21 g. Herein I describe tent construction and utili- zation by Artibeus anderseni, A. glaucus, A. gno- mus, A. phaeotis, A. toltecus, A. watsoni, Uroderma bilobatum, and U. magnirostrum; describe several new styles of tents; review and summarize the lit- erature on tent use by Artibeus and Uroderma; discuss the effectiveness of tents as diurnal roosts; and suggest directions for future research. Methods Descriptions of Study Areas Costa Rica— Bosque Brrancia, also known lo- cally as Bosque Blanco, is located 0.8 km west of Cuarto Cruces on the south side of the Pan Amer- ican Highway (Route 1) in Guanacaste Province, in the Pacific lowlands of western Costa Rica. Bos- que Brrancia is a restricted area of undisturbed lowland forest classified as Tropical Moist Forest; the dominant vegetation includes A nacardium ex- celsum and Scheelea rostrata. This stand of forest probably has not been logged previously, and rep- resents a close approximation to the original (pre- Columbian) forests of this part of Guanacaste. Further descriptions of this forest can be found in Janzen (1971) and Wilson and Janzen (1972). Parque Nacional de Corcovado is located on the Osa Peninsula of southwestern Costa Rica, Puntarenas Province (between 08°27'N and 08°39'N, and 83°25'W and 83°45'W); the eleva- tion ranges from sea level to 400 m. Corcovado lies within the Tropical Wet Forest Life 21one (Holdridge, 1967), with lowland evergreen forest being the dominant forest type. Mean annual rain- fall is 3,800 + mm and the wettest months are from August through November; mean monthly tem- peratures range from 25.0°C to 26.5°C. Vegetation and habitat types at Corcovado have been de- TIMM: ARTIBEUS AND URODERMA 189 scribed by Herwitz (1981) and Hartshorn (1983). Areas surveyed included both primary forest and secondary scrub along the coast. The La Selva Biological Station is the field sta- tion of the Organization for Tropical Studies lo- cated 1 km SW of Puerto Viejo de Sarapiqui, Heredia Province in the Caribbean lowlands of northeastern Costa Rica (10°27'N,84°00'W); ele- vation ranges from 29 to 100 m. Mean annual rainfall is 3,990 mm, with the wettest months being November, December, and February; mean monthly temperatures range from 24.5°C (Decem- ber) to 26.1°C (April). La Selva lies within the Tropical Wet Forest Life Zone, with lowland ev- ergreen forest being the dominant forest type. Vegetation and habitat types of La Selva have been described by Slud (1960), Holdridge et al. (1971), Sawyer and Lindsey (1971), and Hartshorn ( 1 983). One unusual feature of the subcanopy of the La Selva forest is the diversity and abundance of dwarf palms (Hartshorn, 1983), especially the broad- leaved species, Geonoma congesta and Asterogyne martiana. These species are regularly utilized by Artibeus watsoni for tent construction. The tran- sect surveys, which included all Artibeus tents ob- served, were restricted to primary forest. The Uro- derma tents described from this site were restricted to an open banana patch. Palo Verde (Refugio Nacional de Fauna Sil- vestre Dr. Rafael Lucas Rodriguez Caballero) is a wildlife refuge located 2 km S and 12 km E of Bolson, in the Pacific lowlands of Guanacaste Province, northwestern Costa Rica (10''30'N, 85°20'W); the elevational range is from 3 to 183 m. Palo Verde lies within the Tropical Dry Forest Life Zone, with lowland deciduous forest and riv- erine swamp forest being the dominant forest types. The vegetation and habitat types of Palo Verde have been described by Slud ( 1 980) and Hartshorn (1983). Mean annual rainfall is 1,700+ mm, with the wettest months being April, May, September, and October; mean monthly temperatures range from 26.0°C (November) to 29.7°C (April). The immediate vicinity of the survey at this site was in a mosaic of primary forest and secondary scrub that included considerable bananas scattered throughout. The newly expanded Parque Nacional Braulio Carrillo is located in northeastern Costa Rica (be- tween 10°05'N and 10°25'N, and 83°54'W and 84'^5'W); the elevation of the park ranges from 100 to 2600 m. Braulio Carrillo is located on the eastern, Caribbean slope of Volcan Barba in He- redia Province. The elevational range at which Ar- tibeus toltecus and associated tents were observed was from 700 to 1400 m, within the Premontane Rain Forest Zone with midelevational evergreen forest and tall palms being the dominant forest types. Mean annual rainfall at this elevational range is perhaps as much as 5,000 mm, although no exact measurements are yet available. The vege- tation and habitat types at the midelevational ranges also have yet to be described. On 14-15 April 1986, 3 km of trail ranging from 700 to 1 100 m were surveyed for bat tents. Additionally, an intensive netting effort with Richard K. LaVal and Don E. Wilson was conducted in this area over a 7-day period to determine what species of bats were present and their relative abundance. At Finca Las Cruces (2 km S of San Vito, Pun- tarenas Province, 08°45'N, 82°58'W, 1200 m) in the Premontane Wet Forest-Rain Forest transition area, approximately 2 km of trail leading down to the Rio Jaba was surveyed for tents on 1 3 August 1982. Ecuador— Cascada San Rafael lies 17 km (by road) west of the village of Reventador, Napo Province, in northeastern Ecuador (00°5.8'S, 77°34.4'W). Rainfall averages 1,500 to 2,000 mm; average temperatures range from 18°C to 22°C. The elevation is 1200 m. Reventador lies within the Humid Subtropical Life Zone. Lagarto Cocha and Zancudo Cocha are military encampments along the Rio Aguarico of Ama- zonian Ecuador named for prominent lagoons. Both areas are undisturbed primary lowland rain forests classified as Moist Forest, with an annual rainfall averaging from 2,000 to 3,000 mm; the elevation is approximately 200 m. Peru— Hacienda Amazonia lies just north of the confluence of the Alto Rio Madre de Dios and the Rio Pinipini in the department of Madre de Dios, southeastern Peru (12°58'S, 71'>09'W), just north of Atalaya. The Hacienda is located just east of Parque Nacional del Manu in the Upper Trop- ical Zone on the eastern foothills of the Andes. On 25 July 1985 Barbara L. Clauson searched the ridge above the Hacienda for bat tents at an ele- vation of 825 m in primary rain forest that had received some selective timber harvest; she re- turned to this site again on 3 November 1985. Cerro de PantiacoUa lies above the Rio Palotoa, 10-15 km NNW of Shintuya, in the department of Madre de Dios, southeastern Peru (12°35'N, 71''18'W). On 15 November 1985 Clauson searched a steep sloping ridge at 600 m. The sur- 190 HELDIANA: ZOOLOGY Table 1 . Individual measurements (in cm or degrees) collected from four tents constructed by Artibeus anderseni on Heliconia in Ecuador. Blade Petiole Basal height Length Width Angle Length Angle Remarks 176 158 154 140 34 38 36 27 40 15 50 80 75 88 60 80 55 80 70 200 220 260 140 1 adult male A. anderseni 3 male A. anderseni rounding forest was primary rain forest in the Up- per Tropical Zone on the eastern Andean foothills. Reference specimens of the bats have been de- posited at Field Museum of Natural History, Chi- cago; Escuela Politecnica Nacional, Quito, Ecua- dor; and Universidad Nacional de Costa Rica and Servicio de Parques Nacionales, San Jose, Costa Rica. Voucher specimens of plants have been de- posited in the herbaria at Field Museum, Duke University, Missouri Botanical Garden, and/or University of Wisconsin. Accounts of Species Artibeus Artibeus anderseni Osgood, 1916 Artibeus anderseni occupies an extensive range in western Amazonia; however, little is known of its biology. This species has long been considered a junior synonym of A. cinereus. In resurrecting A. anderseni as a distinct species, Koopman (1978, p. 14) stated, "Besides its shorter face and more abrupt forehead, A. anderseni apparently always lacks the last lower molars, which A. cinereus in western Amazonia almost always has." I concur with Koopman in recognizing A. anderseni as a distinct species. In late October and early November of 1983, the trails and forest surrounding the military en- campments were searched at Lagarto Cocha and Zancudo Cocha in eastern Ecuador. Artibeus an- derseni was found to alter the shape of leaves of several small, forest Heliconia species to produce diurnal roosting structures. To create a tent from a Heliconia leaf, the bat severs the lateral nerves and interconnecting veins that extend along both sides of the midrib. The cuts ran along the central 90% of the leaf from near (0 to 14 cm) the base to near (10 to 20 cm) the tip. Nerves and inter- connected tissues were severed, but not complete- ly, so that they did afford some support for the sides of the leaf. Cuts ran parallel to the midrib for most of its length, but did flare outward slightly toward the base. The lateral nerves were cut from 3 to 8 mm from the midrib; the midrib was not cut. Claw marks where the bats roosted started 50 cm from the base in one tent and ran for 1 6 cm distally; in another they started at 70 cm and ran for 10 cm distally. Measurements of the blade length, blade width, blade angle, petiole length, petiole angle, and basal height of four tents are provided in Table 1 . At Zancudo Cocha one Heliconia tent was un- occupied for two days in succession, then on the third day was occupied by an adult male with enlarged testes, an adult lactating female, and a juvenile male Artibeus anderseni (fig. 1). Another Heliconia tent was unoccupied. At Lagarto Cocha 13 tents were found in Hel- iconia. One was occupied by three subadult males not in breeding condition; a second tei t contained a single adult male with enlarged testes. Artibeus cinereus (Gervais, 1856) Artibeus cinereus, Gervais's fruit-eating bat, is found on the islands of Grenada, Trinidad, and Tobago, and throughout the Amazon Basin and adjacent coastal areas. Surprisingly little has been published on roosting behavior or ecology of this widely distributed species. On Trinidad, Goodwin and Greenhall (1961, p. 262) stated, "It roosts in small colonies of a few individuals under the cut leaves of palm trees and on the under side of ba- nana leaves." On Tobago, Husson (1954, p. 64) reported a single male Artibeus cinereus "hanging in a banana tree in cultivated country near the shore." TIMM: ARTIBEUS AND URODERMA 191 Fig. 1. Photograph of an adult male, adult female, and juvenile Artibeus anderseni roosting in a Heliconia leaf tent. Details of the cut side nerves can be seen along the midrib of the leaf Artibeus glaucus Thomas, 1 893 Artibeus glaucus is found at midelevalions along the eastern slopes of the Andes from Venezuela to Peru. The status of the name glaucus has long been in a state of flux; it often has been considered a subspecies of A. cinereus. I concur with Handley (1987) in regarding .4. glaucus as a distinct species. A single old adult male Artibeus glaucus (fmnh 124844) was observed roosting under a cut Xan- thosoma leaf at Cascada San Rafael, Ecuador, on 21 September 1983. Four cut leaves on separate plants were observed in close proximity to each other; a fifth cut leaf was observed approximately 10 to 1 5 m to the south. Only the one cut leaf was occupied by the single bat. An adult female A. glaucus was netted in the vicinity that evening. All tent leaves were cut down at the time and four were measured. On 26 November, two more cut leaves were found in this Xanthosoma population; one contained three A. glaucus. The Xanthosoma in which the bats were roosting were part of a population of Xanthosoma that occupied approx- imately 1 ha on a steep east-facing hillside. The basal lateral nerves from 2 to 5 (usually 3) of the Xanthosoma leaves were severed near the midrib, causing the two sides of the leaf to collapse downward around the midrib. The midrib was not cut. Four of the five cut leaves were measured in September; one found in November was measured (table 2). The leaves that were selected by Artibeus glaucus for tents all had the midrib running ap- proximately parallel to the ground, whereas the majority of unaltered leaves in the p>opulation stood at more vertical angles. Artibeus gnomus (Handley, 1987) Artibeus gnomus, the dwarf fruit-eating bat, is found in a peculiar circular range ringing the west- em edge of the Amazon Basin. Although this species has a wide distribution from Venezuela and Guyana to Peru, it was only recently recog- 192 HELDIANA: ZOOLOGY Table 2. Individual measurements (in cm or degrees) collected from five tents constructed by Artibeus glaucus on Xanthosoma in Ecuador. Blade Length Width Angle Petiole length Basal height Remarks 57 68 60 140 230 3 basal cut veins on each side, 2-3 cm from midrib 53 • • • 40 • • • 230 3 basal cut veins on each side, 1.5-3.0 cm from midrib 55 62 40 • •■ 230 2 veins cut left, 4 cut right side, 2-3 cm from midrib 64 36 45 137 300 3 basal veins cut on each side, 1.5-3.0 cm from midrib; adult male A. glaucus hanging 16 cm from base 61 66 20 145 275 5 basal veins cut on each side, 3.5-4.0 cm from midrib; 3 ^. glaucus nized as a distinct species, and little biological in- formation is available (Handley, 1987). On 15 November 1985 Barbara L. Clauson found a single adult male Artibeus gnomus roost- ing under a cut Monstera lechleriana leaf. The Monstera was growing as an epiphyte approxi- mately 10 m off the ground on a tree on a sloping hillside at 600 m elevation at Cerro de Pantiacolla, southeastern Peru. The single cut leaf was green and healthy and hung horizontally. No other cut leaves were observed in the immediate vicinity. The altered leaf was 70 cm long and 38.5 cm at its widest point. All lateral nerves along the basal nearly two-thirds of the leaf were severed imme- diately adjacent to the thick midrib; this included the basal 12 nerves on one side and 14 on the other. The midrib was severed at 44.5 cm from the base which caused the apical third of the leaf to droop downward perpendicular to the midrib. All nerves proximal to the midrib cut were sev- ered. The lateral nerves along the apical, drooping 25.5 cm were unaltered. The tent resulting from these cuts was quite en- closed, being formed by the sides of the leaf col- lapsing downward around the midrib and the dis- tal third of the leaf folding down, perpendicularly to the midrib. The lone Artibeus gnomus was roosting 9 cm toward the base from the severed midrib of the leaf. Artibeus jamaicensis Leach, 1821 Artibeus jamaicensis, the Jamaican fruit-eating bat, is found throughout much of tropical Central America, the northern half of South America, and the Greater and Lesser Antilles. At many localities this is one of the most common species of bats encountered and consequently has received more study than any other phyllostomid. Artibeus jamaicensis was reported roosting un- der the cut leaflets of Scheelea rostrata in Costa Rica by Foster and Timm (1976). Scheelea ros- trata is a large, pinnately leafed palm with the leaflets extending out at right angles from the hor- izontal rachis. Leaflets within the middle 1.3 m region of the frond were cut at varying distances that increased going up to the center of the cut area, then decreased. "As a result . . . the distal parts of the leaflets folded perpendicularly, hung vertically below the frond, and formed a broadly lanceolate tent" (Foster & Timm, 1976, p. 266). Although several Artibeus jamaicensis occupied the roost, only two males were captured, one an adult with enlarged testes, and the second a smaller male not in breeding condition. Artibeus jamai- censis apparently has a harem mating system, in which a single breeding male defends a roost used by several females and their offspring; nonbreed- ing males may be found either singly or in small groups (Morrison, 1979; Kunz et al., 1983). Ar- tibeus jamaicensis has been found roosting in a wide variety of situations, including caves, hollow trees, buildings, and under unaltered leaves (Tut- tle, 1976), and thus is certainly not an obligate tent-roosting species, as apparently are the smaller species oi Artibeus. Artibeus phaeotis (Miller, 1 902) Artibeus phaeotis, the pygmy fruit-eating bat, is found from central Mexico to northern South America (Timm, 1985). Most accounts of habitat for pygmy fruit-eating bats mention their being netted in close proximity to stands of bananas, Musa X para^/5/aca (Ramirez-Pulidoetal., 1977; Watkins et al., 1972). Davis (1970) suggested that they might roost under the leaves of bananas. During the summer of 1982, Artibeus phaeotis TIMM: ARTIBEUS AND URODERMA 193 Fig. 2. Top, Dorsal view of a banana leaf {Midsa x paradisiacd) showing the cut nerves running aloQg the midrib and flaring out toward the base; bottom, tent of Art ibeus phaeotis made from a banana leaf was observed roosting under the leaves of banana and Heliconia imbricata at Palo Verde and La Selva, respectively, in Costa Rica. In all cases the leaves had been altered to produce tents. Art ibeus phaeotis constructs roosts in both ba- nanas and Heliconia imbricata by biting the lateral nerves and interconnecting tissue that extend at right angles from the midrib, causing the blade to fold over in a V-shaped enclosure. The two sides of the leaf collapse downward, hanging beneath the midrib (fig. 2). Nerves and interconnected tis- sues are not completely severed, thus the sides of the leaf provide some support for the entire length of the cut. The cuts ran from the base of the leaf to near the tip (table 3). Near the base, the cuts flared out from the midrib to the sides to form an elongate J-shaped pattern. The uncut tip and basal portion of the leaf provide additional strength. The undersides of roost leaves are obscured from view from almost all angles except from directly be- neath the tent. To characterize Musa clumps, each of which presumably represented an individual plant, the number of stalks (ramets) per clump was counted; the height of each ramet was estimated to the near- est half meter. This was done for all clumps of Musa in the patches that could be located, both those with tents and without bat tents. When a bat tent was located, the following measurements were taken in centimeters: the angle of the petiole (mid- way from stalk to blade), angle of blade (midway on blade), petiole length, blade length, blade width, height of base of blade, height of tip of blade, height of roost, isolation distance (distance from nearest solid object on the same vertical level), length of uncut basal portion of blade, and length of uncut distal portion. I also noted whether the roosts were in direct sunlight or in shade (table 3). Also for all tents the number of stalks on that column was counted, and the age of the leaf rel- ative to other expanded leaves was recorded. The oldest (lowest leaf) on a plant was assigned the number 1 , then the rest counted up from there. I scored 232 individual stalks of Musa, which 194 HELDIANA: ZOOLOGY Table 3. Individual measurements (in cm or degrees) collected from 19 tents constructed by Artibeus phaeotis on leaves of banana, Musa x paradisiaca, in Costa Rica. Petiole Length Blade Angle Width Baiwl . height Height Tip Roost Isolation Uncut Length Angle Base Tip 43 40 110 -27 20 196 156 182 81 0 15 S3 38 136 -14 19 230 146 225 120 10 45 43 42 146 -40 22 213 170 228 125 1.5 48 49 12 120 -55 23 250 180 240 53 5 8 43 S2 100 -16 18 205 200 70 0 30 SO 55 120 -3 21 200 165 210 25 3 8 S4 50 142 -36 20 235 160 230 115 1 44 40 75 80 + 15 19 190 195 206 80 0,25 20 36 68 175 -12 26 320 260 320 >300 0 1.5 60 70 195 +25 30 360 400 400 > 300 0,35 0 44 66 130 +23 25 250 260 290 120 0 40 41 50 94 +5 20 140 155 170 40 0,40 25 34 45 97 -22 20 205 175 200 100 0 0 SO 68 130 0 23 180 220 230 60 0 2 S6 60 147 -37 27 260 180 240 35 0 28 33 55 85 -12 21 205 215 220 18 0 5 16 52 60 +28 13 130 140 35 0 12 46 50 129 + 15 21 250 220 260 120 0 65 40 54 88 + 15 14 170 180 220 0 3 were in 4 1 clumps scattered in 1 00 m of second growth forest. The mean number of ramets per clump was 5.4; the range from 1 to 23. I located 26 leaves that were cut by bats, a ratio of roughly one tent per 8. 1 ramets. Fourteen clumps had cut leaves; if a clump had cut leaves it had a mean of 1.9 cut leaves (range 1 to 2). Of the 26 altered leaves, complete data were taken on 19. The re- maining tents were decomposing; in a few cases the bats had not completed the tents. On the par- tially completed tents, only one side of the midrib was cut along one-quarter to one-third the distance of the blade, but did not cause the side to collapse. Only 2 of the 1 9 complete tents were located di- rectly in the sun. Bat tents were found in clump sizes ranging from 1 to 23 (mean = 6.2). The av- erage height of all plants (N = 232) was 2.3 m. The average height of plants with tents was 2.0 m (N = 16). One adult male with enlarged testes and six pregnant females were watched over a 3 -day p)e- riod in one of the roosts. Additionally, two solitary nonpregnant female Artibeus phaeotis roosted un- der separate leaves (fig. 3). Artibeus phaeotis appears to select banana leaves with specific characteristics. Usually these are the oldest fully expanded leaves, just over 2 m above the ground, with the center of the midrib nearly horizontal to the ground and positioned far enough from nearby stems and branches to limit access by predators. Roost sites generally are located in the shade of surrounding forest trees where ap- parently they are protected by the forest overstory from wind, blowing rain, and sunlight. Although bananas are not native to the New World, they are now common throughout the range of Artibeus phaeotis and probably provide roost sites in other localities. Heliconia, Calathea, and broad-leafed palms are uncommon at Palo Verde, hence are not readily available to A. phaeotis for tent sites there. Artibeus phaeotis used only banana leaves for tent making at Palo Verde, but con- structed similar tents in Heliconia imbricata at La Selva. Heliconia imbricata is the largest species of Heliconia in Costa Rica, and its leaves are similar in size and shape to banana. The tents in Heliconia were similar in all respects to those in banana leaves. An adult male and adult female were found roosting together under a single Heliconia leaf in late June. Villa-R. ( 1 967) found a single specimen roosting near the mouth of a small cave in Mexico. Artibeus toltecus (Saussure, 1 860) Artibeus toltecus, the lowland fruit-eating bat, is found along the coasts of eastern and western Mex- ico from Nuevo Leon and Sinaloa south through Central America and perhaps to extreme north- western Colombia. This species appears to be re- stricted to midelevational slopes, and in Costa Rica TIMM: ARTIBEUS AND URODERMA 195 Fig. 3. Photograph of an adult female Artibeus phaeotis roosting in a banana leaf tent. Details of the cut side nerves can be seen along the midrib of the leaf. I have found it only from 650 to 1 500 m in ele- vation. In April 1986 I observed a single Artibeus tol- tecus roosting under a cut leaf of Anthurium ca- peratum in Braulio Carrillo National Park, north- eastern Costa Rica. The Anthurium was growing as an epiphyte on a small tree at 800 m (IVi km S, 11 km E of San Miguel, 10°17'N, 84'>05'W). One leaf on the plant was altered; it was IVi m off the ground and the midrib hung parallel to the ground. Four or five lateral nerves were cut basally on each side, causing the sides of the leaf to fold down around the midrib. A break of the midrib at its midpoint caused the distal half of the leaf to droop down (fig. 4). Seven additional tents of this style were observed on Anthurium in this area, ranging in elevation from 700 to 1400 m. It is assumed that they were made by A. toltecus, the only small species of Artibeus we netted there, al- though these tents were not occupied. Six tents were observed in a 3-km transect ranging from 700 to 1 100 m in elevation. Davis (1944) reported that Artibeus toltecus roosts imder banana leaves, although he did not indicate that the bats were modifying the leaves. Davis (1944, p. 378) stated: . . . they had regularly established roosts tin- der the large, drooping leaves of the banana trees, each one easily recognized by the man- ner in which the vane of the leaf hung limply suspended from the midrib. The closely ap- pressed vanes of the leaf, plus the natural darkness within the depths of the grove, af- forded good concealment. These bats, too, were wary and that feature coupled with the nature of their retreat caused considerable difficulty in procuring specimens. In light of Davis's description of the roost sites of A. toltecus in banana leaves and my own obser- vations on A. toltecus, I suspect that this species was creating tents similar to those I observed for A. phaeotis in Costa Rica. The tents formed by A. phaeotis in Musa (see fig. 2) are similar in ap- 196 HELDIANA: ZOOLOGY Fig. 4. Top, Ventral view of an Anthurium caperatum leaf showing the cut nerves along the base of the leaf and the broken midrib; bottom, tent of Art ibeus toltecus. TIMM: ARTIBEUS AND URO DERMA 197 pearance to those described by Davis (1944) for A. toll ecus. Artibeus toltecus has also been reported in caves (Davis et al., 1964; Jones, 1966; Jones & Alvarez, 1 964), and Goodwin ( 1 934, p. 1 2) reported a single specimen collected "in one of the buildings (church?) at San Lucas . . . [the] rest of the con- gregation seemed to be Glossophaga." Artibeus watsoni Thomas, 1 90 1 Artibeus watsoni, Thomas's fruit-eating bat, is one of the smaller members of the genus Artibeus and found from southern Veracruz south through Central America to northern South America. It appears to be restricted to lowland and midele- vation humid forests. During the summers of 1974, 1982, 1984, and 1 986, numerous individuals of Artibeus watsoni were seen roosting under 19 different sj)ecies of broadleafed plants at several localities in Costa Rica. At Parque Nacional de Corcovado, Costa Rica, trails were surveyed on three separate oc- casions for the presence of tents made by Artibeus watsoni, in June and August 1982 and again in August 1984. In mid-June 1982, the following groupings of Artibeus watsoni were observed: two (both adults, a male and pregnant female), two (pregnant female and one not captured), two (not captured), and six hanging singly (of which two were captured and found to be adult males). Ad- ditionally, several tents on banana and Heliconia were marked for relocation later in the summer. A tent marked on Heliconia imbricata was relo- cated 60 days later. The tent was still intact, al- though it was beginning to break down; a single A. watsoni v/zs using it. All other marked tents had decomposed. On 10 August 1982 I found 90 tents constructed by Artibeus watsoni along the trail through Cor- covado's "Monkey Woods." These tents were made from the following species of plants: Musa x par- adisiaca (49, 54%), Anthurium ravenii (13, 14%), unidentified aroid (11, 12%), Heliconia imbricata (9, 10%), Heliconia latispatha (1, 1%), Heliconia sp. (3, 3%), and Calathea insignis (4, 4%). Tents located on Anthurium ravenii were most often found clumped, with an average of 2.6 tents per plant, whereas in the other species of plants it was uncommon to find more than one tent per indi- vidual plant. Bats were found singly (five) or in three groupings of four, three, and two individuals. Four of the single bats were all adult males. A trail running up to a ridge top was surveyed from 9 through 1 1 August 1982, with the following results: 25 tents found of which 16 were on Hel- iconia imbricata (64%), 8 on Calathea insignis (32%), and 1 on Carludovica palmata (4%). Three tents were occupied by two (sexes unknown), one male, and one female. Near the mouth of the Rio Llorona on 8 and 9 August I counted the following groups of bats: eight (three adult females, three young, and two not caught), two (adult female with volant young), two (sexes unknown), and three singles (one a nonreproductive adult female). Ad- ditional tents were observed in banana, coconut palm (Cocos nucifera), Calathea insignis, and Car- ludovica cf. drudei. In August 1 984, 1 found 63 tents constructed by Artibeus watsoni. These were distributed on the following plants: Anthurium ravenii (36, 57%), Heliconia sp. (14, 22%), Musa x paradisiaca (7, 1 1%), Calathea insignis (3, 5%), Carludovica pal- mata (\, 1.6%), IVelfia georgii (1 , 1.6%),and(/eo/i- oma sp. (1, 1.6%). Only 3 of the 63 tents were occupied; one had two bats and two each had sin- gle bats. As I noted in 1982, tents on Anthurium ravenii were often clumped on the same plant with an average of 2.5 tents per plant. At La Selva in July 1982, 43 Artibeus watsoni tents were located over a 5 -day period in the fol- lowing species of plants: Asterogyne martiana (33, 77%), Geonoma congest a (6, 14%), Geonoma cu- neata (2, 5%), and an unidentified species of Cy- clanthaceae (2, 5%). One adult male A. watsoni was found under an Asterogyne martiana tent on the first day. On the fifth day an adult female with young was found under another A. martiana tent that had been unoccupied for the previous fovu" days, as was a third adult (not captured) under another A. martiana tent. All other tents were un- occupied. In 1974 1 surveyed approximately 10 km of trails at La Selva and found 29 tents on the following species of palms: Asterogyne martiana (19, 66%), Bactris wendlandiana ( 1 , 3%), Geonoma congesta (2, 7%) and Geonoma cuneata (7, 24%); all were unoccupied. Foster and Timm (1976) reported tents in these palms, although they were not able to associate bats with the tents. My recent studies at La Selva have confirmed that these tents were made by A. watsoni. At Finca Las Cruces in mid-August 1982, 13 tents constructed by Artibeus watsoni were located; 198 HELDIANA: ZOOLOGY 2 '^ e o •a 00 Si eo| C * (/J c* . k» a T •2 o I a c g . u C f 3 ■-> P « « to x: CO ♦^ o S li -3 e o V TIMM: ARTIBEUS AND URODERMA 199 Fig. 6. Top, Dorsal view of the cyclanth, Carludovica palmata, showing the polygonal cuts; bottom, tent of Artibeus waisoni on C. palmata. 200 HELDIANA: ZOOLOGY Fig. 7. Photograph of adult female Artibeus watsoni and her subadult offspring roosting in a Carludovica leaf tent. Details of the polygonal cuts and folds can be seen in the background. one was occupied by an adult male. The tents were distributed on the following species of cyclanths: Asplundia euryspatha (6, 46%), Carludovica drudei (4, 31%), and Cyclanthus bipartitus (3, 23%). Artibeus watsoni uses a variety of species of plants and a wide array of leaf shapes for diurnal roosts. I have found four distinct styles of tents at a single locality (Corcovado). These styles include the sim- ple V-shaped cuts on bifurcated palms (fig. 5), cut- ting a few side veins on aroids to produce a round- ed pyramid, the elongated J-shaped cuts on banana and Heliconia leaves, and the polygonal cuts on Carludovica (figs. 6-7). For each distinct leaf shape, the cuts create a well-concealed diurnal roost. Ar- tibeus watsoni probably is an obligate tent-rooster, as it has only been found roosting under cut leaves. On several instances a bat occupied the same tent, or tents in close proximity, for two to three days in succession. Those tents might then remain unoccupied for several days in succession. Dis- turbed bats generally flew directly to another tent from 20 to 50 m away, or attempted to return to the tent where originally found. Tents generally are found clumped, both on a single plant if leaf morphology and age are appro- priate, and in restricted areas. Up to five tents have been found on a single Anthurium ravenii, and when present the mean number of tents was 2.5. At Parque Nacional de Corcovado, Choe and Timm (1985) found that Artibeus watsoni showed strong preference for Anthurium ravenii leaves that were medium sized, low within the plant, and grew closer to the ground than average A. ravenii leaves. Also at this site, Boinski and Timm (1985) doc- umented that squirrel monkeys {Saimiri oerstedi) were major predators on A. watsoni, with the adult male monkeys being the most successful at cap- turing bats. Additionally, double-toothed kites {Harpagus bidentatus) followed troops of foraging squirrel monkeys, using them as "beaters." When tent-making bats were flushed by the monkeys and escap)ed, they were routinely captured and con- sumed by the attending double-toothed kites. Artibeus watsoni has long been known to cut palm tents for diurnal roosts, although prior to this study little had been published on roosting TIMM: ARTIBEUS AND URODERMA 201 Fig. 8. Dorsal view of Scheelea rostrata showing the leaflets cut by Uroderma bilobatum to form a tent 202 HELDIANA: ZOOLOGY behavior of this species. The elongate J-shaped cuts made on bifurcated palms were first described and illustrated by Chapman (1932, p. 555). He stated that "both vanes of the leaf whence the bat flew were cut diagonally to the midrib of the leaf, so that their terminal portions drooped downward to form a tentlike shelter." Chapman appropri- ately termed these three-sided diurnal roosts "tents," and I have expanded the use of the word tents to include all modified leaves by bats. Barbour's (1932) original description of Uro- derma bilobatum cutting palm leaves for roosts also provides a secondhand report (p. 308) by H. C. Clark stating that "Clark has just found for the first time a youngish coconut palm, a single leaf of which was being cut by bats of the genus Uro- derma in a very similar way." The common use of young coconut palms {Cocos nucifera) by Ar- tibeus watsoni in Costa Rica, coupled with the total lack of evidence that Uroderma bilobatum uses juveniles of this palm, leaves that small, or roosts that close to the ground, suggests that the tents seen by Clark were in fact made by A. watsoni. Artibeus watsoni also is abundant on Barro Col- orado Island, Panama. Apparently no voucher specimens of the bats were preserved at the time. Allen (1939, p. 69) reported "a specimen of A. watsoni that was hanging by day from the under- side of a banana leaf" Perhaps the natural-looking folds caused by the cuts running parallel to the midrib were not noticed at the time. Ingles (1953) reported on tents of A. watsoni in two species of Geonoma on Barro Colorado Island; one tent was occupied by three individuals. Thomas's fruit-eat- ing bat has been found roosting in an artificial tent, an inverted hanging box. Wilson (1970) reported that several females raised young in the comer of a suspended box on Barro Colorado Island. Uroderma Uroderma bilobatum Peters, 1865 Uroderma bilobatum has been given the dis- tinctive "common" name of Peters's tent-making bat. Tents constructed by Uroderma bilobatum were seen at three separate localities in Costa Rica during the summers of 1982, 1984, and again in 1986, the first at Bosque Brrancia near Cuarto Cruces in the Pacific lowlands of northwestern Costa Rica, the second at Corcovado on the Osa Peninsula, and the third at La Selva in the Carib- bean lowlands. On 25 August 1982 a colony of Uroderma bi- lobatum was roosting under a modified frond of the palm Scheelea rostrata at Bosque Brrancia. The colony included an adult male with enlarged testes and four adult females. The Scheelea rostrata frond in which the colony of Uroderma bilobatum roosted was a mature leaf, approximately 6.5 m in length. The bats were hanging approximately 4.5 m off"the ground; most were clustered together, although a few were spread out over 50 cm of the frond. The cut leaflets started at about 3.5 m off" the ground and proceeded up the frond for the next 2.5 m (fig. 8). The general pattern of the cut leaflets was a tapering effect, with the cuts on the lowermost leaflets being farthest from the midrib. Leaflets along the proximal 2 m and the distal 50 cm were unmodified. Only the midrib of the leaflets was cut. Each leaflet had a distinctive V-shaped fold at its base where it was attached to the midrib. The bats were hanging from the leaflets rather than the midrib. From the dorsal aspect of the leaf, the proximal portion of the tent (cut leaflets) extended 50 cm further down on the right side than on the left to include 10 basal leaflets whose opposites on the left were unaltered. The basalmost cut leaflet was cut 34 cm from the midrib. Proceeding distally, the length of the unmodified basal portion of each leaflet decreases. The basalmost cut leaflet on the left was cut 19 cm from the midrib. The overall appearance of the tent was a sharp, convergent taper for the next meter. Following this section, there was a 7 5 -cm section in which the cuts were close to the midrib (within 3 cm). On the distal- most 30 cm of the tent, the leaflets were cut closer to the midrib on the left side than on the right. Similar tents, each housing a colony of Uroderma bilobatum, were found in a large stand of Scheelea rostrata at Corcovado in 1984, and William A. Haber (pers. comm.) informed me that he has seen similarly cut leaves in the same species of palm at Cahuita (09°44'N, 82°49'W) in the Caribbean low- lands of southeastern Costa Rica in 1 984. In June of 1982 and again in March of 1986, I found numerous banana leaves cut by Uroderma bilobatum just to the north of the field station at La Selva. The midrib on vertical leaves was cut to the extent that the distal portion of the leaf collapsed downward to form a two-sided tent (fig. 9). Severing the midrib on vertical leaves had the effect of folding the leaf back upon itself creating a tight, dark crevice at the fold where the bats roosted (fig. 1 0). In addition to severing the mid- rib, the bat cut a large V-shaped pattern running TIMM: ARTIBEUS AND URODERMA 203 Fig. 9. Left, Dorsal view of a banana leaf showing the cut midrib and the large V-shaped cuts through the side nerves; right, tent of Uroderma bilobatum made from a banana leaf. from the midrib to the base of the leaf. The side veins and interconnected tissues were partly sev- ered. However, because the leaf stood nearly ver- tical, these V-shaped cuts did not cause further folding of the leaf The only cut creating the tent was that of the midrib. In 1982 five tents in widely separated banana leaves were located (table 4). One was occupied by eight Uroderma bilobatum, which included one adult male with enlarged testes and seven females. In 1986 eight tents were observed in the same banana patch. On this occasion eight U. bilobatum 204 FIELDIANA: ZOOLOGY TIMM: ARTIBEUS AND URODERMA 205 tents were clustered in three clumps of bananas. Only one tent was occupied; it contained 1 3 bats. This folded broad-leaved style of tent is un- doubtedly the tent style illustrated by Walker ( 1 960, p. 30) in his photograph of roosting Uroderma bilobatum, although he did not describe it nor mention where it was observed. Interestingly, I have searched several dozen banana and larger Heliconia groves throughout Costa Rica and Ec- uador specifically looking for this style of tent, and none were observed. Uroderma bilobatum is an abundant and widespread species in the lowlands, but employment of this particular style of tent appears spotty as I have not observed it elsewhere. In Panama, Barbour (1932) found that colony size under a single cut leaf of Prichardia pacifica varied from a few bats to 56. Prior to Barbour's discovery that Uroderma altered leaves, Goldman (1920, p. 199) stated of these bats in Panama. In the forest near Gatun Uroderma biloba- tum was located several times, a few in a place, clinging during the day in clusters to the midribs on the under sides of large palm leaves. They usually choose darkened spots where the leaf was folded over, or over- hanging pinnae shut out much of the light. Burt and Stirton (1961) reported U. bilobatum in El Salvador roosting beneath the leaves of bananas and coconut palms. I strongly suspect that the bats had altered these leaves and that these authors had failed to notice it or failed to associate the hanging bats with the damage to the leaves. Bloedel (1955, p. 234) stated of Uroderma bilobatum in Panama that: I observed these bats only in their palm-leaf tents. ... In the latter part of March most females have nursing young, and are roost- ing in clusters of 20 to 40, while the males are separated from them, usually solitary or in small groups of from 2-5. In Trinidad, Goodwin and Greenhall (1961, p. 254) found them on the under side of the fan-shaped leaves of certain palm trees, especially the carat palm (Sabal glaucescens). . . . The bat makes a series of cuts across the pleated surface of a leaf, causing half of the leaf to bend at an angle to form a protected retreat. Table 4. Individual measurements (in cm or de- grees) collected from four tents constructed by Uroderma bilobatum on banana, Musa x paradisiaca, in Costa Rica. Blade Petiole Cut midrib Length Width angle from base Remarks 210 95 40 117 Not occupied 220 90 70 90 8 U. bilobatum 80 1 U. bilobatum 80 Not occupied 210 95 Not occupied They were found roosting in colonies of 1 0 or more individuals. In Suriname, Husson (1962, p. 161; 1978, p. 143) collected three pregnant female U. bilobatum "in a plantation where they were found hanging on the imder side of a leaf of [a] so-called 'paloeloe,' Ravenala guyariensis." In Nicaragua, Jones ( 1 964, p. 507) collected four female U. bi- lobatum that "hung together about 10 feet above the ground in the 'tent' formed by a cut palm frond. Each was pregnant with a single embryo." In Gua- temala, Dickerman et al. (1981, p. 409) reported, "Palm leaf tents were frequently found occupied by one to seven individuals, but nursing females were usually found alone or with juveniles." Leaves selected by Uroderma bilobatum for tents are all large, and of a variety of shapes. The large V-shaped pattern cut into the leaves is a charac- teristic of Uroderma tents. Artibeus watsoni also uses a variety of leaf shapes for tents; however, the nature of the cuts and style of tent created vary with leaf shape. Uroderma bilobatum, on the other hand, makes patterned cuts that appear to be an innate response to large leaves, regardless of the shape. As noted in the banana tents, a single cut across the midrib creates the tent, and the V-shaped pattern had no effect upon tent shape; this perhaps represents wasted effort by the bats. There are a few records of Uroderma bilobatum being found roosting in a hollow tree and one "un- der the eave of house" (Davis, 1968, p. 695). In all cases these have been of single individuals, and I suspect represent either recently dispersed young that have yet to join a breeding colony or bachelor males. Uroderma bilobatum roosting under a cut Prichardia leaf was illustrated by Kunz (1982). A colony of U. bilobatum roosting under a banana leaf tent was illustrated by Keller ( 1 986). Mac- donald's ( 1 984, p. 806) photograph of two tent- making bats under a Heliconia tent is erroneously labeled Uroderma bilobatum. These bats are ac- 206 HELDIANA: ZOOLOGY Fig. 11. Tent of Vroderma magnirostrum on the pinnately leafed palm Astrocaryum muru- muru. tually a small species of Artibeus, probably A. phaeotis; the tent style and size and coloration of the bats are typical oi A. phaeotis. Uroderma magnirostrum Davis, 1968 Although Vroderma magnirostrum is a widely distributed bat found from Mexico to Bolivia, it was not recognized as a species distinct from U. bilobatum until 1968 (Davis, 1968), and few spec- imens are represented in collections. When W. B. Davis described this new species he commented that although it was widely distributed only seven specimens had been collected prior to the wide- spread use of mist nets in the 1960s, and that all specimens available to him had come from local- ities less than 1000 feet in elevation. He stated that "These facts strongly suggest basic differences in the habits of the two species and that those bats with a deep rostrum are not 'tent-makers' as are members of the species Uroderma bilobatum" (Davis, 1968, p. 678). There have been no reports on the behavior or ecology of U. magnirostrum. On 25 July 1985 Barbara L. Clauson discovered a colony of two male and three female Uroderma TIMM: ARTIBEUS AND URODERMA 207 magnirostrum roosting under the cut leaflets of the pinnately leafed palm, Astrocaryum murumuru. The single occupied tent found was on the ridge above Hacienda Amazonia at 825 m in south- eastern Peru. The entire colony was collected by John W. Fitzpatrick. On 3 November Clauson re- turned to the site to measure ,»ie leaf and noted an additional cut leaf in the same plant. When first observed, four bats were hanging to- gether and one was hanging several centimeters away. When observed an hour later, all five were hanging together in a tight cluster approximately 7.5 m off the ground. The bats were hanging from the leaflets rather than the midrib, approximately 200 cm from the tip of the leaf The colony in- cluded one adult male with enlarged testes, two adult females, and two subadults, one female and one male. The roosting structure of Uroderma magniros- trum was in a pinnately leafed palm (fig. 1 1 ). The bats severed the leaflets along the upper two-thirds of the leaf; those along the lower third were un- altered, as were the leaflets at the very tip. As the leaflets proceeded up the tent they were severed closer to the midrib forming an elongate, conver- gently tapering tent (fig. 1 1). The general appear- ance of the U. magnirostrum tent is similar to that described herein for U. bilobatum on the pinnately leafed palm Scheelea rostrata. The Astrocaryum frond in which the colony of Uroderma magnirostrum roosted was a mature leaf, approximately 6.1 m in length and 1.9 m in width at the widest point, with the petiole 1.1m long. The leaf left the trunk (d.b.h. .4 m) at 3 m from the ground and hung at an angle of approximately 50°. The bats were hanging approximately 7.5 m off" the ground. The cut leaflets started at 1.5 m from the lowest leaflet and proceeded up the frond for the next 2.9 m to nearly the tip (fig. 1 1). The cuts on the lowermost leaflets were furthest from the midrib. The lowest severed leaflets were cut up to 34 cm from the midrib, whereas the distal leaflets were severed only 2 cm from the midrib of the leaf The midribs of the leaflets were cut causing the distal portion of the leaflets to fold downward. Leaflets along the proximal 1.5 m and the distal .5 m were unmodified. The trunk, pet- iole, and midrib of this palm were covered with sharp, penetrating spines several centimeters in length. After 14'/2 weeks this tent was still alive and green, most of the leaflets appearing as fresh in November as they did in July. This Astrocaryum contained a second cut leaf that was unoccupied. This roost was also in a ma- ture leaf which was an older leaf than the occupied tent, with many broken, yellowed, and brown leaf- lets. The leaf was approximately 6 m in length and 2 m in width at the widest point, with a petiole .8 m long. The leaf left the trunk at 2.9 m from the ground and hung at an angle of approximately 60°. Cuts were distributed asymmetrically along the length of the leaf The cut leaflets on the left side of the leaf started 3.05 m from the lowest leaflet and proceeded up the frond for the next 1 .3 m, to .88 m from the tip. The cut leaflets on the right side of the leaf started 3.25 m from the lowest leaflet and proceeded up the frond for the next 1.49 m, to .49 m from the tip. The cuts on the lower leaflets were farthest from the midrib. The lowest severed leaflets were cut up to 35 cm from the midrib, whereas the distal leaflets were severed as close as 1.5 cm from the midrib. Leaflets along the proximal 3.05 m and 3.25 m and the distal .88 m and .49 m were unmodified. I propose the common name of Davis's tent- making bat for this species. Conclusions A review of the literature on tent-making bats contains some 32 primary references covering the 55-year period from 1932 through 1986. Surpris- ingly, we actually know very little about the bi- ology of these bats. As late as 1975 Eisentraut was yet doubting th^t bats were cutting leaves to make tents, stating: . . . observers maintain that the bats form these tentlike structures themselves, by making a series of holes running across the middle of a large palm leaf The bats then supposedly bend the outer half of the leaf around, so they can then rest inside this 'tent'. . . . On the basis of personal obser- vations in tropical regions in Africa, I tend to believe instead that these holes were made by insect larvae while the leaves were still rolled up. A storm can then easily break the leaf along the line of holes and form the tent roof which is so convenient for the bats (Ei- sentraut, 1975, p. 142). Eisentraut, by his own admission, had never seen a bat tent. I believe that if he had, he would have come to the same conclusion Thomas Barbour did 208 HELDIANA: ZOOLOGY nearly a half century earlier, that the bats and not insects were making the cuts. Although we have yet to actually observe bats cutting leaves to form the roosting structures de- scribed herein, I hope the volume of data pre- sented here and in my other works establishes for a fact that many species of small and medium sized stenodermines are indeed tent-makers. The ob- servations presented represent data collected from several hundred tents located over a 1 5 -year pe- riod. Several facts consistently emerge between my observations and those independently corrobo- rated by others. Bats of the genera Artibeus and Uroderma (as well as Ectophylla, Mesophylla, and Vampyressa) roost under cut leaves. These leaves may be on a wide variety of species of plants, but generally the shape of the leaves is similar. The shape of the cuts is very characteristic for each species of bat and the patterns and styles of tents created by the bat species are consistent. The behavioral repertoire associated with tent- making in bats certainly evolved more than once, as evidenced by the patchy distribution of tent- making species with the chiropteran suborders Megachiroptera and Microchiroptera. Within the Megachiroptera, a single species of tent-maker is known, Cynopterus sphinx. Within the Microchi- roptera, tent-makers are known only from one subfamily of the Phyllostomidae, the Stenoder- minae. The Stenoderminae constitute an ex- tremely speciose and diverse group of bats, with more than 30 species currently recognized. Tent construction within stenodermines may be a trait that evolved once, twice, or as many as three times. The Artibeus- Uroderma group are sister genera and form one clade of the tent- making repertoire. Sec- ondly, the Mesophylla- Vampyressa group are sis- ter genera (and perhaps should be considered con- generic) and would constitute the second clade. Finally, Ectophylla would constitute a third lin- eage. The relationship between these three lineages is uncertain and warrants further investigation. Knowing that bats modify the leaves of several species of plants to produce diurnal roosting struc- tures led to the following questions: (1) Are bats selecting specific species of plants for tents? (2) What styles of tents are cut by bats and do these differ between species? (3) Do bats select for a particular angle, size, or shape of leaf for diurnal tents? (4) Are leaves selected preferably in larger clumps or smaller clumps? (5) Are older or youn- ger leaves selected? (6) Are leaves of a particular height class selected? (7) Are leaves that are not adjacent to solid objects selected? (8) What do typical tents look like? (9) How and why did tent construction evolve? On occasion I have found "cheaters," species of bats roosting in a tent made by another species. Is cheating an evolved strategy of roost site selec- tion of some bats? Bats of the genera Artibeus and Uroderma ac- tively modify leaves to produce diurnal roosting structures, but by biting the tissue between veins along the midrib and leaving the midrib and most veins intact, do not kill the leaves. The resulting tent is available for use as a roost for an extended period of time; one was observed in use for more than 60 days. Bats select for specific sizes and shapes of leaves. Tents provide concealment from predators and protection from the rain, wind, and sun. This type of roost offers the additional ad- vantage that the bats are warned about the ap- proach of a potential predator, because even slight movements of the leaf stem or the leaf itself are transmitted as magnified vibrations to the roosting bat. Tents may provide bats with suitable roosting sites that would not otherwise be available in close proximity to prime food resources. One of the most productive areas for future re- search will be exploring aspects of the biology of these bats from an evolutionary perspective. Fu- ture subjects I will be addressing include the role of tent roosting in controlling ectoparasites and the correlation between complexity of tents and social systems in these bats. I believe that tent- making originated as an antipredation strategy and has since, secondarily, evolved to play a major role in controlling ectoparasites and in social be- havior. Many factors influence the choice of roost site selection by bats. Included among these are vul- nerability to predation, physical stability of the site, proximity to food sources, and general ap- propriateness of the nest microenvironment for the rearing of young. It seems likely that tent con- struction requires considerable time and energy expenditure by bats, attesting to intense selection pressures involved. Acknowledgments I thank Eduardo Lopez Pizarro and El Depar- tamento de Vida Silvestre and Fernando Cortes and Servicio de Parques Nacionales of Costa Rica for making this study possible. The Organization for Tropical Studies (OTS), Rebecca Butterfield, TIMM: ARTIBEUS AND URODERMA 209 William A. Haber, Gary Hartshorn, Charles E. Schnell, and Joe M. Wunderle are gratefully ac- knowledged for assistance with logistics. Robert J. Izor assisted with field logistics in Costa Rica and Peru, and John W. Fitzpatrick assisted in Peru. In Ecuador I thank the Comandancia General del Ejercito Ecuatoriano, the Corporacion Estatal Pe- trolera Ecuatoriana, and the Ministerio de Agri- cultura y Ganaderia for making our studies there possible. Luis Albuja, Ramiro Barriga, Angelitos Garrett, Myriam Ibarra, Gustavo Orces, and Don- ald J. Stewart provided logistic assistance in Ec- uador. Kerry A. Barringer, William C. Burger, Thomas B. Croat, Robin B. Foster, Barry Ham- mel, and Timothy Plowman provided identifica- tions or confirmed identifications of the plants. Barbara L. Clauson, Alfred L. Gardner, Lawrence R. Heaney, Karl F. Koopman, Thomas H. Kunz, Bruce D. Patterson, and Timothy Plowman pro- vided valuable suggestions on earlier drafts of the manuscript. My wife, Barbara, provided superb assistance with all aspects of this project, including providing several of the photographs used and all data on Uroderma magnirostrum. Rosanne Mie- zio prepared the illustrations. Nina Cummings, Ron Testa, and Diane White expeditiously and cheerfully executed my photography requests. This project was funded in part by grants from the Rice Foundation of Chicago, the National Science Foundation [INT-8303 194], National Geographic Society, and Field Museum of Natural History. I especially thank Mr. and Mrs. Arthur A. 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Bulletin of the American Museum of Natural History, 121: 49-148. . 1980. The birds of Hacienda Palo Verde, Guanacaste, Costa Rica. Smithsonian Contributions to Zoology, 292: 1-92. TiMM, R. M. 1984. Tent construction by Vampyressa in Costa Rica. Journal of Mammalogy, 65: 166-167. . 1985. .4m"/?^M.s pAaeo/w. Mammalian Species, 235: 1-6. TiMM, R. M., AND J. Mortimer. 1976. Selection of roost sites by Honduran white bats, Ectophylla alba (Chiroptera: Phyllostomatidae). Ecology, 57: 385-389. TuTTLE, M. D. 1976. Collecting techniques, pp. 71- 88. In Baker, R. J., J. K. Jones, Jr., and D. C. Carter, eds.. Biology of Bats of the New World Family Phyl- lostomatidae. Part I. Special Publications, The Mu- seum, Texas Tech University, 10: 1-218. ViLLA-R., B. 1 967. Los murcielagos de Mexico. Anales del Instituto de Biologia, Universidad Nacional Au- tonoma de Mexico, xvi + 49 1 pp. Walker, E. P. 1960. Studying our Fellow Mammals. The Animal Welfare Institute, New York, 1 74 pp. Watkins, L. C, J. K. Jones, Jr., and H. H. Genoways. 1972. Bats of Jalisco, Mexico. Sjjecial Publications, The Museum, Texas Tech University, 1: 1-44. Wilson, D. E. 1970. An unusual roost o{ Artibeus ci- nereus watsoni. Journal of Mammalogy, 51: 204-205. Wilson, D. E., AND D. H. Janzen. 1972. Predation on Scheelea palm seeds by bruchid beetles: Seed density and distance from the parent palm. Ecology, 53: 954- 959. Appendix List of scientific names of plants mentioned in the text and used by Neotropical bats for tent con- struction. Anacardiaceae Anacardium excelsum (Bertero & Balbis) Skeels Araceae (cont'd.) Monstera lechleriana Schott Xanthosoma sp. Araceae Cyclanthaceae Anthurium caperatum Croat &. Baker Anthurium ravenii Croat & Baker Asplundia euryspatha Hari. Carludovica dmdei Masters TIMM: ARTIBEUS AND URODERMA 211 Cyclanthaceae (cont'd.) Carludovica palmata R. & P. Cyclanthus bipartitus Poit. Marantaceae Calathea insignis Petersen Musaceae Heliconia imbricata (Kuntze) Baker Heliconia latispatha Benth. Musa X paradisiaca L. Plenakospermum guyanense Endl. (syn. Ravenala guyanensis Petersen) Palmae Asterogyne martiana (H. Wendl.) H. Wendl. ex Hemsley Astrocaryum murumuru Mart. Bactris wendlandiana Burret Cocos nucifera L. Geonoma congesta H. Wendl. ex Spruce Geonoma cuneata H. Wendl. ex Spruce (syn. G. decurrens H. Wendl.) Geonoma oxycarpa Martius (syn. G. binervia Oerst.) Livistona chinensis (Jacq.) R. Br. ex. Mart. Prichardia pacifica Seem. & H. Wendl. Sabal mauritiiformis (Karsten) Griseb. &. H. Wendl. ex Griseb. (syn. S. glaucescens Lodd. ex H. E. Moore) Scheelea rostrata (Oersted) Burret Welfia georgii H. Wendl. ex Burret 212 FIELDIANA: ZOOLCX5Y Comparative Ultrastructure and Evolutionary Patterns of Acinar Secretory Product of Parotid Salivary Glands in Neotropical Bats Carleton J. Phillips, Toshikazu Nagato, and Bernard Tandler ABSTRACTS Secretory products produced by acinar cells of the parotid salivary glands of 1 5 species of Neotropical bats {Pteronotus parnellii, Phyllostomus elongatus, P. latifolius, Tonatia bidens, T. sylvicola, Trachops cirrhosus, Glossophaga soricina, Leptonycteris sanborni, Sturnira lilium, Artibeus jamaicensis, Ariteus flavescens, Eptesicus lynni, E. brasiliensis, Tadarida brasiliensis, and Molossus molossus) were compared by transmission electron microscopy. Extensive inter- and intrageneric differences were found in the ultrastructure of the mature acinar secretory granules. This variation in secretory cell product exceeded any previously reported intraordinal phenotypic variation at the cellular level, but was in keeping with previously reported bio- chemical data on salivary protein polymorphism in primates and rodents. Data from molecular biology and systematics lend support to the hypothesis that the microscopic variations are directly representative of genie differences among species. It also is postulated that intrageneric microscopic differences at least partly are due to neutral (nonfunctional) differences in molecular structure or charge (or both) rather than evolutionary selection. Among the phyllostomids, a general trend in parotid acinar cell product was found in Artibeus and Ariteus, in which a decrease in enzymatic content of the product could be correlated with ultrastructural differences. The secretory product in Artibeus and Ariteus also differed significantly from that of the genus Sturnira. and it is proposed that the phenotypic differences between Sturnira and the other two stenodermatines represent a major genetic difference of systematic importance. The ultrastruc- tural appearance and substructure of the parotid acinar secretory granules could not be con- sistently correlated with diet alone, although insectivorous-animalivorous SF)ecies have enzyme- rich, mostly electron-dense granules, whereas two fruit bats, Artibeus and Ariteus, have pale, enzyme-poor parotid granules. Productos de secrecion producidos por celulas acinares de las glandulas salivales de 1 5 esF>ecies de murcielagos neotTopicales (Pteronotus parnellii, Phyllostomus elongatus, P. latifolius, Tonatia bidens, T. sylvicola, Trachops cirrhosus, Glossophaga soricina, Leptonycteris sanborni, Sturnira lilium, Artibeus jamaicensis, Ariteus flavescens, Eptesicus lynni, E. brasiliensis, Tadarida bra- siliensis y Molossus molossus) fueron comparados mediante microscopio electronico de trans- mision. Extensivas diferencias inter- e intragenericas fueron encontradas en la estructura de granulos glandulares acinares maduros. Esta variacion en productos de celulas secretoras excedio From the Department of Biology, Hofstra University, Hempstead, NY 11550 (Phillips); and Department of Oral Biology, School of Dentistry, Case Western Reserve University, Cleveland, OH 44106 (Nagato and Tandler). Dr. Nagato's present address is Department of Oral and Maxillofacial Surgery, Ehime University School of Med- icine, Shizukawa, Shigenobu, Onsen-gun, Eshime 79 1 -02, Japan. PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 213 cualquier variacion fenotipica intraordinal previamente reportada a nivel celular, pero estuvo en armonia con datos bioquimicos previamente reporlados de polimorfismo en proteinas sa- livales de primates y roedores. Datos de biologia molecular y sistematica proveen apoyo a la hipotesis de que las variaciones microscopicas son direclamente representativas de diferencias geneticas entre especies. Es lambien postulado que diferencias microscopicas intragenericas, al menos parcialmente, son debidas a diferencias neulrales (no funcionales) en estructura molecular o carga (o ambas) mas bien que a seleccion evolutiva. Entre los filostomidos, una tendencia general en el producto parotido de las celulas acinares fue encontrado en Artibeus y Ariteus, en los cuales una disminucion en contenido enzimatico del producto p>odria estar relacionado con diferencias ultraestructurales. El producto secretado en Artibeus y Ariteus tambien se diferencio significativamente de aquel del genero Sturnira y es propuesto que las diferencias fenotipicas entre Sturnira y los otros dos stenodermatinos representan una mayor diferencia genetica de importancia sistematica. La apariencia ultraestructural y subestructura de los granulos secre- torios parotidos acinares podria no estar consistentemente correlacionada solo con la dieta, aunque especies insectivoras-animalivoras tienen granulos ricos en enzimas mayormente densos en electrones, mientras que los murcielagos frugivoros, Artibeus y Ariteus, tienen granulos parotidos palidos y pobres en enzimas. Produtos secretorios produzidos por acinos das glandulas parotidas de 15 especies de mor-' cegos neotropicos (Pteronotus parnellii, Phyllostomus elongatus, P. latifolius, Tonatia bidens, T. sylvicola, Trachops cirrhosus, Glossophaga soricina, Leptonycteris sanborni, Sturnira lilium. Artibeus jamaicensis, Ariteus flavescens, Eptesicus lynni, E. brasiliensis, Tadarida brasiliensis, e Molossus molossus) foram comparados atraves da microscopia de transmissao eletronica. Vastas diferengas intra- e intergenericas foram encontradas na ultra-estrutura dos granulos maduros dos acinos secretorios. Esta variafao no produto das celulas secretorias supera qualquer varia^ao fenotipica intraordinal previamente relatada para o nivel celular, porem concorda com as rela96es publicadas sobre dados bioquimicos do polimorfismo de proteinas salivares em primatas e em roedores. Dados sistematicos, e de biologia molecular, apoiam a hipotese que variances microscopicas sao diretamente representativas das diferen^as geneticas entre especies. Propoese tambem, que estas diferen^as microscopicas intragenericas sao ao menos parcialmente causadas por diferen9as neutras (i.e., naofuncionais) nas estruturas moleculares ou nas suas cargas eletricas (ou ambas), ao inves de serem consequencias da sele^ao evolutiva. Entre os morcegos da familia Phillostomidae, foi encontrado em Artibeus e em Ariteus um padrao geral nos produtos dos acinos parotideos, onde uma redu9ao do conteudo enzimatico' e correlacionado a diferencas nas ultraestruturas dos granulos produzidos. Os produtos secre- torios em Artibeus e em Ariteus sao significantemente diferentes dos produtos do genero Sturnira. e prop6e-se que as diferen9as fenotipicas entre Sturnira e os outros dois stenodermatinos representam uma grande diferen9a genetica, de importancia sistematica. A aparencia ultraes- trutural, e a subestrutura dos granulos secretorios dos acinos parotideos, nao se correlacionam consistentemente com a dieta por si, apesar de que as especies insetivoras-animalivoras possuem granulos ricos em enzimas e densos em eletrons, equanto que dois morcegos frugivoros, Artibeus e Ariteus, (wssuem granulos parotideos palidos e com poucas enzimas. Introduction Neotropical bats are extremely diversified; ex- isting species represent perhaps the most outstand- ing mammalian example of ecomorphological ad- aptation. Dentitions, jaw morphology, brains, kidneys, tongues, and digestive tracts are only a few examples among the anatomical features that have been investigated in recent years (e.g., Phil- lips, 1971; Forman, 1972; Phillips et al., 1977, 1984; Freeman, 1979, 1981; Eisenberg «t Wilson, 1978; Studier et al., 1983). The major salivary glands are yet another system that has attracted attention, primarily because histological, ultra- structural, and histochemical investigations have consistently revealed striking interspecific differ- ences and unusual histological and secretory fea- tures (Wimsatt, 1956; DiSanto, 1960; Junqueira & Fava de Moraes, 1965; Junqueira et al., 1967, 1973; Phillips et al., 1977; Mineda, 1977, 1978; 214 HELDIANA: ZOOLOGY Pinkstaff et al., 1982; Tandler & Cohan, 1984; Nagatoetal., 1 984; Tandler & Phillips, 1985; Phil- lips & Tandler, 1985, 1987; Tandler et al., in press). Mammalian salivary glands are highly complex organs that not only secrete digestive enzymes but also can secrete hormones, antibodies, and toxins, to name but a few known products (e.g., Tandler, 1972; Hand, 1980b). Data are available on the biochemistry of saliva in humans and common laboratory species; but very little is known about the specific biochemistry of the parotid acinar se- cretory granules except for laboratory rats in which some of the proteins have been characterized (Ro- binovitch & Sreebny, 1969; Ball, 1974; Wallach et al., 1975). The complex structure and function of salivary glands is underscored by data from studies of Neotropical bats, which recently have been analyzed by both transmission electron mi- croscopy and histochemistry. For example, the ac- cessory submandibular gland of Trachops cirrho- sus has been shown to differ histologically from any known mammalian salivary gland, with the exception of the same gland in Megaderma lyra and M. spasma. Megaderma lyra is an Old World ecological equivalent of Trachops (Phillips &. Tandler, 1985, 1987; Phillips et al., 1987). Both of these unrelated species feed on frogs (Lekagul & McNeely, 1977; Tuttle & Ryan, 1981), which possibly has been a factor in the convergent evo- lution of their submandibular glands. A previously unknown cellular organelle has been described in another Neotropical bat, Tonatia sylvicola (Na- gato et al., 1984). In this species, the presence of xmique crystalloid smooth endoplasmic reticulum in seromucous acinar cells is sex-linked (being found only in submandibular acinar cells in males); a steroid product produced by this organelle pos- sibly serves as a species-isolating mechanism or as part of a chemo-behavioral system, or both. Lastly, a comparative investigation of the secre- tory product in seromucous acinar cells in sub- mandibular glands of five species ofArtibeus has revealed that the ultrastructural characteristics of secretory products can have systematic implica- tions (Tandler et al., 1983, 1986). This study was of particular interest because the salivary gland data matched genie data independently derived by Koop and Baker (1983). For the present investigation we surveyed par- otid acinar cell secretory products in a selected group of 1 5 species of Neotropical bats. Our group comprised four families (Mormoopidae, Phyllo- stomidae, Vespertillionidae, and Molossidae) and included a group of species in which dietary habits ranged from insectivory and animalivory to om- nivory and frugivory. This investigation is the first comprehensive interspecific survey of secretory product ultrastructure, and addresses the follow- ing questions: ( 1 ) what is the range of variation in secretory products; (2) what are the evolutionary patterns in parotid secretory product; and (3) what systematic conclusions can be reached by com- parative ultrastructural analysis? Materials and Methods Numbers and sex of specimens used in the pres- ent study are given in the Appendix. All of these bats were collected during fieldwork in Mexico, Jamaica, and Suriname. Voucher specimens for all species and collecting localities are deposited in the mammal collections of either the Carnegie Museum of Natural History or The Museum, Tex- as Tech University. Bats typically were collected at night with mist nets, and were kept overnight without food until they were killed between 0900 and 1 200 the following morning. The animals were anesthetized with 0.25 ml of sodium pentobarbital (50 mg/ml, intraperitoneally) and the salivary glands removed, placed on dental wax, flooded with freshly mixed fixative, and diced into pieces measuring approximately 1 mm'. Two fixation protocols were used at different times during the project. Specimens collected in Mexico and Jamaica were fixed in 2% glutaral- dehyde in 0. 1 M phosphate buffer and then stored unrefrigerated in fresh fixative. The specimens col- lected in Suriname were fixed in a modified tri- aldehyde-dimethylsulfoxide (dmso) mixture, first described by Kalt and Tandler (1971) and later modified slightly for fieldwork (Phillips, 1 985). The trialdehyde fixative, consisting of 3% glutaralde- hyde, 1% paraformaldehyde, 0.5% acrolein, 2.5% DMSO, and 1 mM CaClj in a 0.05 M cacodylate buffer and sucrose at pH 7.2, proved superior to the simple glutaraldehyde fixative in that (1) a higher percentage of tissues proved to be ade- quately fixed for study, and (2) the mitochondria tended to remain intact instead of being disrupted. Although the composition of the fixatives was dif- ferent, we have not found any evidence that these differences introduced undesirable artifacts that would influence our analysis. Additional details about the techniques can be found in both Phillips (1985) and Nagato et al. (1984). To remove unbound aldehydes, the tissue blocks were subjected to prolonged washing in phos- PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 215 acinar lumen. acinar cell secretory graauLej striated Lntercalated acinus duct duct Fig. I . Schematic diagram of a secretory unit in a typical parotid gland. A cluster of secretory cells, arranged around a lumen to form an acinus, drains into an intercalated duct that in turn empties into a striated duct. In reality, more than one acinus may be associated with an intercalated duct, which itself may be branched. The acinar cells elaborate the primary saliva, which has plasma-like concentrations of electrolytes, and add their organic secrelory products to it. The precise role of the intercalated duct cells has not been established, but they probably play a role in secretion and transport. The principal function of the striated ducts is resorption of electrolytes, mainly sodium, from the primary saliva, rendering it hypotonic. phate-buffered sucrose. The blocks were postfixed for two hours in phosphate-buffered 2% OSO4, rinsed in distilled water, soaked overnight in cold aqueous 0.25 uranyl acetate, rinsed again in dis- tilled water, dehydrated in ascending concentra- tions of ethanol, passed through propylene oxide, and embedded in Epon-Maraglas (Tandler & Wal- ter, 1977). Thin sections were doubly stained with methanolic uranyl acetate (Stempak & Ward, 1 964) and lead citrate (Venable & Coggeshall, 1965). All sections were examined in a Siemens Elmiskop la transmission electron microscope (tem). Semithin sections ( 1 nm) used for orientation were stained with toluidine blue (Bjorkman, 1962) and exam- ined in a Zeiss Ultraphot. Tissue samples used for comparisons were se- lected carefully from among available tissue blocks. We generally avoided edges of tissues where me- chanical trauma often affects not only the cellular ultrastructure but also the microscopic appearance of the secretory granules. In selecting representa- tive "mature" granules we took into account the full array of inter- and intracellular variability as well as the often complex substructural geometry of the granules. Selection of the most representa- tive granules admittedly was subjective, but based on our experience, four principal criteria were applied: (1) cells selected for analysis showed no signs of shrinkage or swelling, and sensitive or- ganelles, such as mitochondria, were not distorted; (2) the appearance of the granule was not altered in any meaningful way by variations in fixation; (3) the appearance of the secretory granules had to be unrelated to their location within the spec- imen block; and (4) the development of the gran- ules could be traced (from Golgi complex to apical cytoplasm) without any major breaks in devel- opmental sequence. Results ' The basic histology of the parotid salivary gland was similar in all 15 sp)ecies examined; in each species the acinus was formed by a cluster of cells connected to the striated duct by intercalated ducts of varying lengths (fig. 1). At the tem level, acinar cell secretory products were found to differ in all species examined (figs. 2-5). These differences could not be related to granule ontogeny by com- paring "immature" Golgi-GERL (Golgi-endoplas- mic reticulum-lysosomes)-associated granules with "mature" granules in the apical cytoplasm. In our samples that consisted of both males and females, we found no evidence of sexual dimorphism in secretory granule substructure, and in our largest samples (5 to 1 0 specimens) we found no examples of individual or geographic variation that could not be attributed to typical inter- or intracellular variation or to granule geometry. Although vari- 216 FIELDIANA: ZOOLOGY PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 217 53 F •8 c u <3 55 tn "2 g § t3 4J 5s es ? c O s* ! J3 a S 2 i/i o o ■ 1 u d V u ^ U c •i 00 w c o u ■^ o CA CO r<^ _3 s r-' **^ u "3 c X to C 4J C CQ i/i (^ & O U ^ 1 k Q O ■S. o 1 S., 5 is £ T3 ^ 3 C o C8 CO o s s s X 3 ^ to c c §5 1 1 tw 8 a o c •->, V U 1 > ■•5 t 2 s ^ 3 c o Y-. '^ o \j ss ^ 5 ■"O vo' ■jj u eripheral light material had a layered structure (fig. 4, top). Eptesicus brasiliensis— As in E. lynni. the lim- iting membrane was subtended by layered mate- rial, usually in several plies, that extended into the dense matrix. These extensions were less tortuous than those in E. lynni and appeared to subdivide the granule interior (fig. 4, bottom). Tadarida brasiliensis— The secretory granules in this species exhibited a spectrum of patterns. The most common was relatively simple, with either short, flat, dense prisms or dense dots being suspended in a slightly less dense matrix. A few granules contained dense hollow spheres, whereas others had a mazelike configuration based on light and dark laminations (fig. 5, top). PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 219 Fig. 4. Higher magnification of mature secretory granules in parotid acinar cells in Eptesicus lynni (top) and E. hrasiliensis (bottom). Note the subtle but consistent differences in the granule substructure. Eptesicus lynni, x 63,000; E. brasiliensis, x 76,800. 220 HELDIANA: ZOOLOGY Fig. 5. Parotid acinar cell granules in Tadarida brasiliensis (top) are compared to those found in Molossus molossus (bottom). Note the range of variation in granule substructure in T. brasiliensis. Tadarida brasiliensis, x 26,000; Molossus molossus, x 24,000. PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 221 MoLOSSUs MOLOSSUS— The variable morphol- ogy of the granules in this species appeared to depend on their stage of maturation. Early gran- ules were small, with a moderately dense matrix in which were some prominent dense particles. As the granules matured (based on size, density, and spatial relationship to the Golgi complex), they enlarged and the particles decreased somewhat in density. The mature granules had a farinaceous matrix probably resulting from comminution of the dense particles (fig. 5, bottom). Discussion The idea of using comparative ultrastructural analysis of homologous, morphologically differ- entiated cells to study evolutionary pathways and to explore systematic relationships is new to the study of mammalian orders, but has been used in broader studies of vertebrates and invertebrates (Eakin, 1968; Rieger & Tyler, 1979; PhiUips & Tandler, 1987). Nevertheless, the potential value of ultrastructural comparisons has been demon- strated recently by studies of gastric mucosa, ret- ina, and submandibular salivary glands (Phillips et al., 1984; Feldman & Phillips, 1984; Tandler et al., 1983, 1986; Phillips & Tandler, 1987). The discovery of patterns of intersF>ecific differences in cellular architecture and cellular secretory prod- ucts is in keeping with previously successful use of histology in evolutionary and systematic mam- malogy (e.g.. Quay, 1954; Forman, 1972; Phillips & Oxberry, 1972; Sands et al., 1977; Naumova, 1981; Hood & Smith, 1982, 1983). Secretory cells, such as the parotid acinar cells used in the present study, seem to hold special promise for comparative investigation. Firstly, the entire secretory process— from nuclear dna to syn- thesis of proteins and complex carbohydrates and packaging of materials into secretory granules and their subsequent discharge— has been studied in- tensively over the past several decades (e.g., Jamieson & Palade, 1971; Castle et al., 1975; Pa- lade, 1975). Secondly, the secretory process seems to be relatively conservative— in the sense that basic pathways are the same in virtually all secre- tory cells— and, therefore, interspecific compari- sons are facilitated and interpretation is somewhat simplified (Phillips & Tandler, 1987). In the present study we limited our descriptive comparisons to "mature" secretory granules. These are the secretory granules that accumulate within the apical cytoplasm of the acinar cell; this product either is discharged from the cell into the acinar lumen (thus becoming part of the formative saliva) or, after an unknown storage interval, is broken down through autophagy and recycled within the cell. The glycoprotein components of the acinar secretory product are elaborated by the rough en- doplasmic reticulum acting in concert with the Golgi complex and are perhaps the most interest- ing feature from a comparative point of view. Syn- thesis of these glycoproteins begins with transcrip- tion of a very small segment of the genome into mRNA. The exportable proteins of the secretory granules could be regarded as providing a "win- dow" on the genome because, although all somatic cells have the same genome (Briggs & King, 1 952, 1957), only a small portion actually is operational in any given fully differentiated cell. A protein synthesized for export thus directly reflects a por- tion of the operational genome. Any polysaccha- ride components of the secretory granules prob- ably are at least one more step removed from the genome because synthesis of complex carbohy- drates is enzymatically determined and usually takes place within saccules in the Golgi complex (Tandler, 1978). Even so, the polysaccharides also have considerable potential for comparative anal- ysis. Production of secretory granules can be consid- ered in terms of ontogeny. The first "granules" typically are seen in direct proximity to the Golgi complex, where carbohydrates are linked to pro- teins and the membrane that will encase the gran- ule is synthesized in a process involving both the GERL and the Golgi complex itself (Tandler, 1978; Hand, 1 980a; Hand & Oliver, 1 984). Newly formed ("immature") granules differ greatly from "ma- ture" granules (see Castle et al., 1975) and were not used by us in describing the product for each of our species. Nevertheless, in view of species differences found by us, it is interesting to ask whether or not immature granules of one species might resemble mature granules in another species. Such similarities might be expected if hetero- chronic differences accounted for differences in mature product. However, because no such cross- species similarities were found by us between immature and mature secretory granules, hetero- chrony does not seem to account for sp>ecies dif- ferences, at least among the 1 5 bats examined here. No two Neotropical sfiecies examined here ex- hibited the same mature secretory granules in their parotid gland acinar cells, although in a few cases there was a degree of resemblance. This extreme variability is easily the greatest ever reported for 222 HELDIANA: ZOOLOGY homologous cells within an order or, as in the case of the Phyllostomidae, within a family of mam- mals (or any other vertebrates). To what can we attribute this striking finding? One consideration is fixation, which certainly affects the appearance of any cellular feature as viewed with transmission electron microscopy. In- deed, it can be said that the appearance of cells and their products essentially is the consequence of their intrinsic chemistry combined with the chemistry of the fixative at the moment in time when fixation occurred. Different fixatives and tis- sue processing can have profound effects on the microscopic appearance of secretory granules in salivary glands (Simson et al., 1978). However, we used consistent processing techniques and two similar fixatives. Our specimens of Ariteus and Leptonycteris both were fixed with 2% glutaral- dehyde, whereas all of our other specimens were fixed with a trialdehyde-DMSO fixative (Kalt & Tandler, 1971; Phillips, 1985). We were able to eliminate fixation as a source of variation because we also have examined specimens of Artibeus that had been fixed in both solutions, coincidentally with Leptonycteris, Ariteus, and all of the other species examined (see A. phaeotis parotid in Phil- lips et al., 1977). Generally speaking, given consistent prepara- tion techniques, microscopic differences in secre- tory granule substructure can be ascribed to bio- chemical differences among the granules. Microscopically detectable sequestration of indi- vidual types of macromolecules within secretory granules has been demonstrated only rarely (Ra- vazzola & Orci, 1980; Kousvelari et al., 1982) but clearly is the best available explanation of intra- granule substructure. Based on fundamental prin- ciples of biochemistry, it thus can be concluded that macromolecules packaged in the secretory granules most likely sort themselves out according to charge and steric effects, as well as chemical interactions, to yield a characteristic pattern for each species. However, it also should be noted that a homogeneous appearance of intragranular sub- stance does not in itself preclude sequestration of different enzymes within the granule. Separate lo- calization of different enzymes (a-amylase and chymotrypsinogen A or a-amylase and trypsino- gen) within pancreatic cell zymogen granules has been demonstrated with a combination of hrp (horseradish-p)eroxidase)-labeled and ferritin-la- beled antibodies (Ono et al., 1980). At present it is impossible to correlate exactly secretory granule substructure with particular chemical components such as certain enzymes or mucosubstances, so from microscopic images alone we cannot say precisely how the granules in our species differ chemically from one another. How- ever, some conclusions can be inferred from the literature. For example, an electron-dense image (see, for example, fig. 2), labeled classically as "se- rous," can be associated with granules rich in en- zymes. Such an image is typical in species such as laboratory rodents and primates, for which some data are available on the biochemistry of parotid saliva (e.g., Jacobsen «Sc Hensten-Pettersen, 1974). On the other hand, the presence of electron-dense "serous" granules does not preclude the presence of mucosubstances within the secretory granules. Pinkstaff'et al. (1982) reported that, although the parotid product in the little brown bat, Myotis lucifugus, was "serous" with standard histological techniques, both neutral and acidic mucosub- stances could be demonstrated histochemically. The parotid granules in Artibeus and Ariteus are interesting in this regard because electron-dense material is scarce (especially in Artibeus, fig. 3) and enzyme production is extremely low, at least in Artibeus (Junqueira et al., 1973). By way of con- trast, Sturnira lilium has largely electron-dense granules (fig. 3) and thus differs considerably from the other two stenodermatines; Sturnira produces saliva rich in enzymes (Junqueira et al., 1973). Although the parotid granules in Artibeus (and, by extension, Ariteus) are low in enzyme content, to what can we attribute their tem image? This is an intriguing question because Wimsatt (1956) re- ported that the gland was negative for mucosub- stances, whereas Radtke (1972) reported the pres- ence of sialomucins in parotid acinar cells. In part, this apparent disagreement is the result of differ- ences in techniques that cannot be resolved by transmission electron microscopy. What can be determined about the parotid sal- ivary glands in Neotropical bats that relates to their diets, evolutionary history, or systematic re- lationships? Clearly the ultrastructure of parotid secretory granules has systematic significance, be- cause no two genera (or species either) are exactly alike. This finding is in keeping with our studies of the submandibular gland in five species of Ar- tibeus (Phillips et al., 1977; Tandler et al., 1983, 1986). The seromucous cells in this gland were found to contain granules that allowed for three groupings of Artibeus species {A. cinereus-A. phaeotis, A. jamaicensis-A. lituratus, and A. con- color) that matched the independently derived genie data (from isozyme analysis) reported by PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 223 Koop and Baker ( 1 983). The differences in parotid acinar product in Eptesicus lynni and E. brasi- liensis (fig. 4) also are interesting from this per- spective because independent genie data from 1 9 presumptive loci show that E. lynni probably orig- inated from the E. fuscus species complex, sepa- rate from the origin of E. brasiliensis (Arnold et al., 1980). Eptesicus lynni was found to share only 62% of analyzed alleles with E. brasiliensis. It might seem surprising that the microscopic images of salivary gland secretory products are generically (and often specifically) distinctive, but variation of such a fine resolution actually is in keeping with numerous genetic studies of mam- malian saliva. Several genetic markers have been found in human saliva (Azen & Opp)enheim, 1 973; Ashton & Balakrishinan, 1974; Tan & Ashton, 1976), and salivary proteins in particular tend to be polymorphic (Azen, 1972, 1973). Additionally, both sex and strain differences in salivary proteins have been reported in laboratory mice (Ikemoto & Matsushima, 1984). Although none of these dif- ferences has been demonstrated microscopically (all are based on biochemical analysis alone), their occurrence nevertheless is significant to compar- isons among mammalian species. Although the sometimes subtle but consistent microscopic differences between species within the genera Phyllostomus, Tonatia, and Eptesicus (figs. 2, 4) are in keeping with our hypothesis of the sensitivity of secretory product to genie differ- ences, they cannot readily be related to any known ecological differences between species within each genus. It is altogether possible that such species differences do not represent direct evolutionary selection. Instead, the differing images might rep- resent species-specific protein polymorphisms of a type that would not significantly affect "perfor- mance" of the saliva even though differences in primary molecular structure or surface charges (or both) could be indirectly detected by transmission electron microscopy (Phillips & Tandler, 1987). The existence of such "nonfunctional" (and pre- sumably nonselected, i.e., "neutral") interspecific variation in a protein molecule has been demon- strated previously in the otherwise conservative hemoglobin molecule (Perutz, 1983). In this ex- ample the tertiary and quaternary structures ap- parently are conserved regardless of large numbers of functionally neutral amino acid substitutions in the primary structure. Although these substitu- tions in themselves most often have no effect on the functional capacity of the molecule, they nevertheless are known sometimes to change mo- lecular surface charges (Perutz, 1983). For the present we postulate that similar nonselected, non- functional variations could account for the micro- scopically detectable intrageneric differences in salivary proteins in Tonatia bidens and T. sylvi- cola, Phyllostomus latifolius and P. elongatus, and Eptesicus lynni and E. brasiliensis. This explana- tion seems more parsimonious than the alterna- tive, which would be to assume that the acinar cell component of the parotid saliva is "functionally" different in closely related, ecologically similar species. The extent or degree of microscopic dif- ferences in the mature acinar cell product ulti- mately might tell us more about relative times of divergence than about ecological differences among congeneric species of bats. Microscopic comparisons of salivary gland se- cretory products could be valuable to cladistic studies of chiropteran families, at least to judge from our data. The phyllostomines generally are regarded as the most primitive (least derived) of the Phyllostomidae (Smith, 1976), and their par- otid acinar products thus would qualify as plesio- morphic (following Henning, 1 966). The products in all five species examined (Phyllostomus elon- gatus, P. latifolius, Tonatia bidens, T. sylvicola, and Trachops cirrhosus) contained large amounts of electron-dense material (fig. 2), as did the se- cretory product in Pteronotus parnellii (a closely related mormoopid), Eptesicus lynni and E. bra- siliensis (Vespertilionidae), and Tadarida brasi- liensis and Molossus molossus (Molossidae), all of which serve as "outgroups." If the electron-dense, enzyme-rich secretory granules are regarded as plesiomorphic in micro- chiropteran bats, then it would be reasonable to regard a granule with less electron-dense material as apomorphic (derived). The glossophagine gen- era examined (Glossophaga and Leptonycteris) are representative of a phyllostomid evolutionary trend in which dentition, tongues, and associated mus- culature were modified for feeding on fruit, pollen, and nectar (Park & Hall, 1951; Phillips, 1971; Greenbaum & Phillips, 1974; Griffiths, 1982, 1983; Smith 8l Hood, 1 984). In these genera the electron- dense component has been reduced in comparison to the phyllostomines. Carollia perspicillata, which is omnivorous, fits into this category in that the parotid secretory granules contain relatively little electron-dense material. In this bat the secretory granules are very distinctive because the electron- dense inclusions are often in the form of cagelike geodesic structures (Phillips & Tandler, 1987; Tandler et al., in press). 224 FIELDIANA: ZOOLOGY Perhaps the most interesting systematic finding in our data lies within the nominal subfamily Stenodermatinae. These bats represent an evolu- tionary trend toward frugivory that includes ex- treme gastric adaptation at the gross, histological, histochemical, and cellular levels (Forman, 1972; Forman et al., 1979; Phillips & Studholme, 1982; Phillips et al., 1984). The pale parotid secretory granules in Artibeus and Ariteus are synapomor- phous, whereas the parotid granules in Sturnira are electron-dense, enzyme-rich, and more nearly like the plesiomorphic granules of the phyllosto- mines, in particular Trachops cirrhosus (figs. 2-3). This example is interesting because Sturnira al- ways has been something of an enigma. Although de la Torre (1961) allied this genus with Vampy- rops-\\\it stenodermatines and Smith (1976) in- cluded it with the "long-faced" stenodermatines, others (e.g., Walton & Walton, 1968) previously had placed the genus in a separate subfamily (Stur- nirinae) based on a variety of morphological fea- tures that seemed inconsistent with the other, more traditional, stenodermatines such as Artibeus. In- deed, Slaughter (1970) pointed out that Sturnira has some distinctive dental features that possibly link the genus to the glossophagines. The differ- ences in the parotid secretory granules thus are in keeping with a variety of other phenotypic differ- ences. While it is reasonable to suggest that secre- tory granule differences of this magnitude repre- sent a major interspecific difference in the operational segment of the genomes of homolo- gous parotid acinar cells, the eventual systematic value of such data will await availability of data about still other phyllostomid genera. Although our comparative data clearly docu- ment great microscopically detectable plasticity in the secretory product of parotid acinar cells, many questions about their evolution remain unan- swered. For example, what has been the role of diet? At first glance, our data suggest that insec- tivorous and animalivorous species have enzyme- rich, electron-dense granules, whereas frugivores have enzyme-poor, pale secretory granules. Yet, what about Sturnira lilium, which certainly in- cludes large amounts of fruit in its diet (Gardner, 1977)? At least three factors interfere with any effort to correlate parotid acinar cell product with diet. First, acinar cells are but one cell type among several that influence the biochemistry of parotid saliva. Second, and possibly more important, the par- otid is but one salivary gland among a host of glands (the submandibular and sublingual and mi- nor glands) that are located throughout the oral region and that contribute substances to the saliva. Some data suggest that different glands in different species might have been more responsive, in an evolutionary sense, to changes in diet. For ex- ample, it is the accessory submandibular glands of Trachops and Megaderma that are unique in histology and might correlate with feeding on frogs (Phillips &. Tandler, 1985; Phillips et al., 1987); the parotid in Trachops is similar to that of other phyllostomines. A third factor, for which we presently have no data, is the possibility that parotid acinar cells can respond in some way to diet at the individual level. Our sample sizes are large enough to convince us of the near uniformity of secretory granule mor- phology within a population. However, Schick et al. (1984) have recently published the first report of a secretory cell responding to dietary intake by a shift in enzyme production. This is the first such detailed molecular data known to us and, although pancreatic acinar cells in laboratory rats were the source of the data, the implications for compar- ative studies of salivary glands are worth noting. A final factor for consideration is the complex role that salivary glands play in the lives of mam- mals. Salivary glands in Artibeus, Chiroderma, and Ametrida might contribute to gastric cytoprotec- tion (Studier et al., 1983; Phillips et al., 1984). In Tonatia sylvicola an unusual organelle found in submandibular seromucous cells of males might be related to species isolation or sex recognition, or both (Nagato et al., 1984). If these examples are typical for bats, then digestion is but one of several major functions of salivary glands. In conclusion, this first systematic microscopic survey of a secretory cell product has demonstrat- ed a previously unknown, extraordinary degree of variation within a group of related sjjecies. Based on our findings, one might conclude that Neo- tropical microchiropteran bats will serve as a sig- nificant model for study of how secretory cells have evolved in mammals; meanwhile, data from comparative investigations will contribute to our knowledge of genie relationships among these an- imals. Acknowledgments Financial support for field and laboratory re- search that led to the data reported here came from a variety of sources, which we are pleased to ac- knowledge: Research Corporation Grant C-1251 PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 225 (to Phillips); NIH Grant DE 03455-02 (to Phil- lips); NSF Grant CDP-80 1-8653 (to Phillips); Hofstra University HCLAS Executive Committee grants (to Phillips); NIH Grant AM-08305 (to Tandler); NIH Grant RO DE 07648-01 Al (to Tandler & Phillips); the Alcoa Foundation (through Dr. Hugh H. Genoways, Carnegie Museum of Nat- ural History); and the Graham Netting Research Fund via a grant from the Cordelia S. May Char- itable Trust (through Genoways). In addition to recognizing our financial support, we also wish to thank several colleagues whose field companion- ship in such places as Suriname and Jamaica, hard work, and willingness to share time and ideas in their laboratories played a major part in our study. These are: Hugh H. Genoways, Carnegie Museum of Natural History; Gary W. Grimes and Dorothy E. Pumo, Hofstra University; Robert J. Baker, Texas Tech University; and Henry A. Reichart, formerly of STINASU, in Suriname. Field assis- tance was provided by K. M. Studholme, S. L. Williams, N. M. Sposito, J. Groen, R. L. Honey- cutt, B. Koop, M. Arnold, B. A. Oxberry, P. Bil- leter, J. Bickham, and J. Patton. We also acknowl- edge technical assistance of Carol Ayala, Case Western Reserve University, and typing by Linda Cossen, Special Secretarial Services, Hofstra Uni- versity. Artwork and lettering were done by Helen Tandler. 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Journal of Morphology, 99: 1 69- 209. 228 FIELDIANA: ZOOLOGY Appendix Specimens Examined Voucher specimens of each species and each locality are deposited in the collections of the Car- negie Museum of Natural History and The Mu- seum, Texas Tech University. All specimens ex- cept those of Leptonycteris sanborni (Jalisco, Mexico) and Ariteus flavescens and Eptesicus lynni (Jamaica) were collected in Brokopondo Province, Suriname. Pteronotus parnellii, 10 (5 S6, 5 29); Tonatia bi- dens, 3 {2 SS, 1 2); T. sylvicola, 2 {66); Trachops cirrhosus, 2 (1 3, 1 2); Phyllostomus elongatus, 3 (\6,2 22); P. latifolius, 1 (2); Glossophaga soricina, 3 (2 $6, 1 2); Leptonycteris sanborni, 2 (3<5); Sturnira lilium, 5 (2 35, 3 22); Artibens jamaicensis, 3 (22); Ariteus flavescens, 1 (2); Eptesicus lynni, 2 {$$); E. brasiliensis, 3 (22); Tadarida brasiliensis, 1 (S); Mo- lossus molossus, 8 (4 66, 4 22). PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 229 Distribution of the Species and Subspecies of Cebids in Venezuela Roberta Bodini and Roger Perez-Hernandez ABSTRACTS Thirteen species of Primates representing nine genera in the family Cebidae are found in Venezuela. The geographic distribution of these species exhibits four main patterns. Alouatta and Cebus are widely distributed in all parts of the country. Four genera, Saimiri, Callicebus, Cacajao, and Chiropotes are restricted to south-central Venezuela; their distribution is centered in the Amazonian lowlands. Aotus and Ateles are each represented by one species or subspecies in south-central Venezuela and another in northwestern Venezuela. Pithecia is mainly restricted to the Guianan highlands in eastern Venezuela, with a single known outlying locality in south- central Venezuela. Of the nine cebid genera in Venezuela, all occur in south-central Venezuela, four occur in northwestern and north-central Venezuela, and only three occur in eastern Ven- ezuela. Distribution maps for the 1 3 species in Venezuela are presented with exact localities of specimens. Nueve generos de cebidos representados por trece especies se hallan en Venezuela. La re- particion geografica de las especies se resuelve en cuatro patrones geograficos principales. De este modo, Alouatta y Cebus son extensamente repartido por todo el pais. La distribucion de los generos, Saimiri, Callicebus, Cacajao y Chiropotes esta restringida al sur-central de Vene- zuela con concentracion en las tierras bajas Amazonicas. Aotus y Ateles son representatados, cada cual, por una especie (o subespecie), en el sur-central, y otra especie in el noroeste del pais. Pithecia se reparte en las alturas guayanas venezolanas del este, y es conocido, a la vez, por un solo dato de captura del sur-central venezolano. En resumen, la totalidad de los nueve generos esta presente en el sur-central de Venezuela, cuatro de ellos en el noroeste y norte- central, y tres en la Venezuela oriental. La reparticion de las trece especies esta documentado por mapas. Treze especies de Primatas, representando nove generos da familia Cebidae, sao encontradas na Venezuela. A distribuifao geografica destas especies exibem quatro padroes principais. Al- ouatta e Cebus espalham-se por todas partes do pais. Quatro generos, Saimiri, Callicebus, Cacajao, e Chiropotes, limitam-se ao centro-sul de Venezuela; a Bacia Amazonica sendo seus focos de distribuifoes. Aotus e Ateles sao representados por uma especie ou subespecie cada no centro-sul da Venezuela, e por outra especie no noroeste do pais. Pithecia limita-se ao planalto Guianense, no leste da Venezuela, com apenas uma localidade conhecida no centro- sul do Pais. Em suma, todos nove generos de Cebidae ocorrem no centro-sul da Venezuela, quatro ocorrem no centro-norte e noroeste do Pais, e apenas tres ocorrem no regiao leste da Venezuela. Apresentam-se mapas de distribui96es das 13 especies na Venezuela, com locali- dades exatas dos especimes colecionados. From the Institute de Zoologia Tropical, Universidad Central de Venezuela, Apartado 47058, Los Chaguara- mos, Caracas 1041 -A, Venezuela. BODINI & PEREZ-HERNANDEZ: CEBIDS IN VENEZUELA 231 Introduction Primates of the family Cebidae are some of the most conspicuous mammals in the Neolropics, as they are diurnal, often forage in large troops, and may be quite vocal. However, we actually know very little about the exact distributions of most species of cebids. In Venezuela, several authors have discussed cebids as part of generic revisions (see Elliot, 1912; Hershkovitz, 1949, and later works; HiU, 1960, 1 962; Kellogg & Goldman, 1944) and in works on the status or collections of specific species (i.e., Bodini, 1983; Handley, 1976; Mon- dolfi & Eisenberg, 1979; Rudran & Eisenberg, 1982). Herein we summarize the distribution of cebids in Venezuela based on recent collections, speci- men records from several museums, and the lit- erature. Specimens reported on are housed in the following collections: Estacion Biologica Rancho Grande (EBRG); Museo de Biologia, Universidad Central de Venezuela (MBUCV); Museo de Cien- cias Naturales (MCN); and Museo de Historia Natural La Salle (MHNLS). Our objectives are to provide accurate locality records and distribution maps that may be utilized in future investigations. Generic Distributions The nine genera of cebids that inhabit Venezuela exhibit four main patterns of geographic distri- bution (figs. 1-7). (1) Two geneT^—Alouatta and Cebus— arc widely distributed in all paris of the coimtry. (2) Four genera— Sa/m/r/, Callicebus, Cacajao, and Chiropotes—UTe restricted to south- central Venezuela, centering on the Amazonian lowlands (see Eisenberg & Redford, 1 979). (3) Two genera— .4orw5 and Ateles—are each represented by one species or subspecies in south-central Ven- ezuela and another in northwestern Venezuela or northwestern and north-central Venezuela, which is the region of the northeastern spurs of the An- dean chain and the enclosed Maracaibo basin. (4) One genus— Pithecia— is mainly restricted to the Guianan highlands in eastern Venezuela, with a single known outlying locality in south-central Venezuela. In summary, of the nine cebid genera in Venezuela, all occur in south-central Venezuela (including the outlier record of Pithecia), four oc- cur in northwestern and north-central Venezuela, and only three occur in eastern Venezuela. Specific and Subspecific Distributions Saimiri In a recent revision of the squirrel monkeys, Hershkovitz (1984) recognized four species: Sai- miri boliviensis, S. oerstedi. S. sciureus, and S. ustus. All those squirrel monkeys found in north- em South America he referred to the single species S. sciureus (fig. 1 ). Squirrel monkeys from Ama- zonian Venezuela and adjacent Brazil and Colom- bia he regarded as 5. sciureus cassiquiarensis. Hershkovitz (1984) plotted several localities in Territorio Federal Amazonas, but listed only a single specific locality: "Casiquiare, Rio (mouth), 2°01'N, 67"X)7'W." Hill (1960) previously had mapped the distribution of 5. sciureus as occurring throughout all of Venezuela, although he listed only a single locality in the state of Bolivar (Camp Canaracuni, 4''36'N, 64''10'W). The name cassi- quiarensis Lesson is based on Humboldt's descrip- tion of a captive female from the banks of the Rio Casiquiare, Amazonas, Venezuela. Specimens Examined— Total 29. Bolivar: Ca- naracuni (4°36'N, 64«'10'W). Territorio Federal Amazonas: Campo Cacuri (4°49'N, 65°26'W); Cano Yureba, Rio Ventuari (3''35'N, 66°46'W); Rio Pu- runame, 40 km from union with Rio Orinoco (3°19'N, 65°15'W); Rio Ventuari (3°59'N, 67'^2'W); San Fernando de Atabapo (4°02'N, 67»37'W); San Juan de Manapiare (5°14'N, 66'^2'W). Aotus Night monkeys are found from Panama throughout much of Amazonian South America to Paraguay. Historically, it has generally been as- sumed that all night monkeys represented the sin- gle species Aotus trivirgatus. However, in a recent revision of the genus, Hershkovitz (1983) recog- nized nine allopatric species. A tenth species, Aotus hershkovitzi from Colombia, has recently been proposed by Ramirez-Cerquera (1983). Two species of Aotus, A. lemurinus and A. trivirgatus. have been reported from Venezuela (fig. 2). Aotus lemurinus griseimembra Elliot is known in Venezuela only fi-om the extreme northwestern region, the states of Cojedes and Merida (Hersh- kovitz, 1983), and the states of Tachira, Trujillo, and Zulia (Handley, 1976). Specimens Examined— Total 10. Zulia: Campo a Rosario, EHstrito Catatumbo (l''44'N, 67°03'W); 232 FIELDIANA: ZOOLOGY BODINI & PEREZ-HERNANDEZ: CEBIDS IN VENEZUELA 233 234 HELDIANA: ZOOLOGY BODINI & PEREZ-HERNANDEZ: CEBIDS IN VENEZUELA 235 Reserva San Manuel, Los Canaguatos (9*t)4'N, 71''56'W); Kunana, Rio Negro, Sierra de Perija (9*75'N, 72*25'W); Rio Tocuco (72''25'W). Aotus trivirgatus trivirgatus Humboldt is known from the region south of the Rio Orinoco in Ter- ritorio Federal Amazonas and the state of Bolivar. The Venezuelan national collections contain specimens from Cano Manapiare and Rio Anta- vari, and two literature records report specimens from the right bank of the Rio Caroni (INP- ARQUES, 1982;MARNR-DGS-POA-SFS, 1982). Our recent discovery of Aotus trivirgatus in Guyana to the east of the Rio Caura, well outside of the Rio Orinoco basin, led to an investigation of geographic variation in the species. Preliminary results suggest that these A. trivirgatus represent a new geographic race (Bodini & Ferreira, in prep.). Specimens Examined— Total 1 1. Bolivar: Cano Manapiare (7»1 1'N, 66°40'W); Rio Antavari (S'^O'N, 63°10'W). Territorio Federal Amazonas: Alto Manapiare (S'^S'N, 66°02'W); San Juan de Manapiare (S^M'N, 66°02'W); near Isla Cigarron, Rio Negro (r44'N, 67°03'W); recently collected in La Neblina, east of Rio Varia (0°59'N, 66°10'W). Callicebus The genus Callicebus is the sole representative of the subfamily Callicebinae and is found from northern South America south to northern Para- guay. Hershkovitz (1963) and Kinzey (1982) rec- ognized three species, Callicebus moloch, C per- sonatus, and C. torquatus, of which only C. torquatus is found in Venezuela. Callicebus torquatus lugens Humboldt is known in Venezuela south of the Rio Orinoco (fig. 3). Hill (1960) originally suggested that C. torquatus is found as far east as Guyana, although this was questioned by both Handley (1976) and Hersh- kovitz (1963), who reported Venezuelan speci- mens only from Territorio Federal Amazonas. Handley (1976) reported 31 specimens from southern Territorio Federal Amazonas. Bodini (1981) reported specimens from Maripa, Camp Canaracuni, and the Rio Antavari in Venezuela. Recent reports by Kinzey ( 1 982) and Bodini (1981) confirm the presence of C torquatus in Guiana Region. Specimens Examined— Total 26. Bolivar: Cana- racuni (4'36'N, 64°10'W); Maripa, 150 km from Ciudad Bolivar (7*22'N, 65'^9'W); Rio Antavari (5»20'N, 63M0'W); Alto Paragua (4°30'N, 63*WW). Territorio Federal Amazonas: Alto Cano Caname, Departamento Atabapo (3*'33'N, 67"06'W); Alto Ventuari (3°50'N, 67°04'W); Boca Cano Maica, Rio Ventuari (66°30'W); Cano Ya- gua, Cerro Cucurito, Departamento Atabapo (3''38'N, 66»25'W); Cacuri, Alto Ventuari (4°49'N, 65"26'W); La Esmeralda (3"X)8'N, 65°32'W); Rio Cunucunuma (3°10'N, 66°0rW); Rio Puruname (3°19'N, 65''15'W); La Neblina, east of Rio Varia (0°59'N, 66°10'W). Alonatta Alouatta is the only genus within the subfamily Alouattinae and is represented by some six species widely distributed in Central and South America. A single species, Alouatta seniculus, is abundant and widely distributed in Venezuela. Hill (1962) recognized three subspecies of A. seniculus as oc- curring in Venezuela. Alouatta seniculus seniculus Linnaeus is foimd in extreme northwestern Ven- ezuela, primarily in the states of Apure, Tachira, and Zulia, with a single record from Barinas (Ti- coporo Forest, 8'X)6'N, 70*'40'W). Alouatta seni- culus arctoidea Cabrera, called the "Caracas howler monkey" by Humboldt, inhabits all the coastal region from Falcon to the state of Miranda. Ca- brera (1 958) proposed Caracas as the type locality. A third subspecies, A. seniculus stramineus Hum- boldt, inhabits all of Venezuela south of the Rio Orinoco in Territorio Federal Amazonas and the state of Bolivar (fig. 4). In addition to these three subspecies, we believe a fourth, previously unrecognized, form exists and is widely distributed throughout the Venezuelan llanos. This undescribed subspecies is character- ized by coloration and size. The southern limit of its distribution is clearly defined by the Rio Ori- noco, but its northern and western limits in the Andean piedmont are as yet undetermined. Howler monkeys are extremely adaptable to a wide array of environments, and we believe they are found throughout Venezuela. Their apparent absence in certain regions probably reflects lack of collecting rather than true distributional gaps, as our records demonstrate for the state of Anzoa- tegui. The problem of current and historical dis- tribution and systematic relationships of the var- ious populations in Venezuela is in need of study. Specimens Examined— Total 80. Anzoategui: Los Cocos, Rio Caris (8''30'N, 64'^5'W). Apure: Caiio San Agustin, Selva de San Camilo (7"'19'N, 7r57'W); Hato El Frio (7°44'N, 68°54'W); Las Raicitas, El Saman (7°55'N, 68°40'W). Aragua: Asentamiento Los Castillos, Turagua (10°09'N, 67»3rW); Cumbre de Guacamaya (10°2rN, 67'40'W); Los Picachos, Rancho Grande ( 1 0*2 1'N, 236 FIELDL\NA: ZOOLOGY BODINI & PEREZ-HERNANDEZ: CEBIDS IN VENEZUELA 237 6T4l'W). Barinas: Reserva Forestal Ticoporo (8°09'N, 70°40'W). Bolivar: Guayoba, Rio Caura, 4 km from Maripa (7°20'N, 65°10'W); Hato Bella Vista, El Palmar (8°00'N, 62°00'W); Represa del Guri, Operacion Rescate (7°40'N, 63°00'W); San Martin de Turumban, Anacoco-Cuyuni (6°42'N, 61°02'W). Carabobo: Las Quiguas, San Esteban (10''25'N, eS^OrW). Cojedes: Hato Barbasco (9°04'N, 68°08'W); Hato Itabana, 38 km from Las Vegas (9°17'N, 68*'13'W); Las Queseras, El Baul (8'*25'N, 68°17'W); Montaria Las Lomas, San Car- los (9°38'N, 68°34'W). Distrito Federal: Hacienda El Limon (10°28'N, 67''1 7'W). Falcon: Sanare, De- partamento Silva (10°23'N, 68°25'W); Sierra San Luis (1 1°15'N, 69°10'W). Guarico: Corozo Pando (8°14'N, 67°17'W); Hato Mapurite, 40 km N of Calabozo (9°17'N, 67°24'W); Manapire, near San Antonio (9°1 7'N, 66''1 1 'W); Rio Tiznado (8°1 8'N, 67°48'W). Lara: Cumbre de Las Trojas, 45 km S of Cabudare (9°45'N, 69°07'W). Miranda: Rio Ne- gro (10°20'N, ee'lS'W); La Guzmanera, Guatopo (10'X)0'N, 66°15'W). Portuguesa: Agua Blanca (9°40'N, 69°07'W). Tachira: Cerro El Teteo, Bura- gua (7°30'N, 7 r57'W); La Fria (8°1 3'N, 72°14'W); Paramo Tama (7°27'N, 72°26'W). Territorio Fed- eral Amazonas: Cacuri, Alto Ventuari (4°49'N, 65°26'W); Caiio Yureba, Ventuari (3°35'N, 66°46'W); Rio Hacha, Alto Ventuari (3°47'N, 65''38'W); Rio Puriname, 40 km from confluence of Rio Orinoco (3°19'N, 65°15'W); San Juan de Manapiare (9°05'N, 66°02'W); Cerro La Neblina, E of Rio Varia (0°59'N, 66°10'W). Territorio Fed- eral Delta Amacuro: Caiio Caneima (9°05'N, 60°55'W); Guiniquina (9°10'N, 61°06'W). Zulia: Laguna de Manaties, Departamento Catatumbo (9°27'N, 72°02'W); La Victoria, Rio Negro (9°36'N, 72°1 5'W); Rio Guasare ( 1 1°02'N, 72°05'W); Sierra de Perija (9°00'N, 72°00'W). Chiropotes The bearded sakis comprise two species limited to northern South America (Cabrera, 1958; Hersh- kovitz, 1977;Mittermeier«Sc.Coimbra-Filho, 1981). One species is found in southern Venezuela rep- resented by a single subspecies, Chiropotes satanas chiropotes Humboldt (fig. 5). All of our records and those reported by Handley (1976) are from Territorio Federal Amazonas. Cruz Lima (1945) and Rudran and Eisenberg (1982) proposed the occurrence of this species in the state of Bolivar on the basis of Humboldt's description; Mondolfi (1976) reported observations of Chiropotes at Maripa and Caiio Maniapure (Bolivar), but we are aware of no specimens from this region. Specimens Examined— Total 29. Territorio Federal Amazonas: Cacuri, Alto Ventuari (4°49'N, 65''26'W); Caiio Yureba, Departamento Atabapo (3°3rN, 66''44'W); Caiio Yagua, Cerro Cucurito (3''3 1 'N, 66°44' W); Laguna de Chiripo, Caiio Blan- co (3°27'N, 66°40'W); Rio Ocamo, Alto Orinoco (2°44'N, 65°11'W); Puerto Ayacucho (5°36'N, 67''35'W); Rio Orinoco, S of San Fernando de Ata- bapo (4'^0'N, 67°38'W); San Fernando de Ata- bapo (4°02'N, 67°37'W); San Juan de Manapiare (5°14'N, 66'^2'W). Cacajao The genus Cacajao, or uakaris, contains two species which are found in northern South Amer- ica (Cabrera, 1958; Hershkovitz, 1972; Mit'ter- meier & Coimbra-Filho, 1981). One species, Ca- cajao melanocephalus, is found in Venezuela restricted to the upper Orinoco region of southern Territorio Federal Amazonas (fig. 5). Although few specimens exist in collections, they appear to be abundant. Specimens Examined— Total 2. Territorio Fed- eral Amazonas: Alto Cano Atacavi, Departamento Casiquiare (3°05'N, 67'>02'W); La Nebhna, E of Rio Varia (0°59'N, 66°10'W). Cebus The capuchin monkeys are found from Hon- duras south through Central America and the northern two-thirds of South America. Four species currently are recognized (Cabrera, 1958; Hersh- kovitz, 1972). Three species of Cebus are repre- sented in Venezuela: Cebus albifrons, C apella, and C. «/^rm//a/z^s (Hershkovitz, 1949, 1958). Three subspecies of Cebus albifrons are found in Venezuela (fig. 6). Cebus albifrons adustus Hershkovitz was described on the basis of three specimens from "near head of Rio Cogollo (Apon) at eastern base of Sierra de Perija, about 5 kilo- meters northwest of Machiques, Lake Maracaibo region, Zulia" (Hershkovitz, 1949, p. 369). We report additional specimens from Rio Guasare and Kasmera. This subspecies is restricted to the Sierra de Perija of extreme northwestern Venezuela and adjacent Colombia. Cebus albifrons leucocephalus Gray is found in extreme western Venezuela, in the region south of the Lago de Maracaibo basin. 238 HELDIANA: ZOOLOGY BODINI & PEREZ-HERNANDEZ: CEBIDS IN VENEZUELA 239 and the states of Apure, Merida, Tachira, and Zu- lia. Cebus albifrons unicolor Spix is found in ex- treme southern Venezuela in Territorio Federal Amazonas. Hershkovitz (1949) reported speci- mens from Marimonda, Rio Orinoco, and from Solano, Rio Casiquiare. Handley (1976) reported specimens from Rio Mavaca and Tamatama. All locality records for this subspecies in Venezuela are south of the Rio Ventuari. Specimens Examined— Total 15. Apure: Caiio San Augustin, Selva de San Camilo (7°19'N, 7r57'W). Merida: Palmichoso, S of Las Virtudes (9'^9'N, 70°57'W). Tachira: La Fria (8°13'N, 72°14'W). Territorio Federal Amazonas: Cano Yagua, Cerro Cucurito, Departamento Atabapo (B^SS'N, 66''25'W); near Boca Padamo, left side of Rio Orinoco (3°02'N, 65°13'W). Zulia: Kasmera, Perija (10°05'N, 72°45'W); Kunana (9°36'N, 72°15'W); Rio Bravo, Distrito Catatumbo (9°05'N, 72°22'W); Rio Guasare (1 1°02'N, 72°05'W). Cebus apella is represented in Venezuela by two subspecies (fig. 6). Cebus apella apella Linnaeus is restricted to Amazonian Venezuela, the Territorio Federal Amazonas, and is found along both banks of the upper Orinoco. Cebus apella margaritae HoUister is endemic to and restricted to Margarita Island. The 800-km gap between the ranges of these two subsp>ecies is striking and unexplained. Specimens Examined— Total 10. Nueva Espar- ta: Sierra de Copey (1 1°03'N, 63°56'W). Territorio Federal Amazonas: Alto Caiio Caname (3''22'N, 67°08'W); Caiio Yapacana (3°30'N, 66°45'W); San Fernando de Atabapo (4°02'N, 67°37'W). Cebus nigrivittatus is widely distributed in Ven- ezuela and represented by perhaps five subspecies (fig. 6) (Hershkovitz, 1949; Cabrera, 1958). Cebus nigrivittatus apiculatus was described by Elliot (1912) on the basis of specimens from La Union, Rio Caura, near its confluence with the Rio Ori- noco. It is distributed throughout central Vene- zuela south of the Orinoco between the Rio Caroni and the Rio Ventuari. Cebus nigrivittatus brunneus was described by J. A. Allen (1914) from specimens from Aroa, a station on the Bolivar Railway, Yaracuy, north- western Venezuela. Hershkovitz (1949) reported an additional specimen from the Paria Peninsula. These records plus our specimens suggest that C. n. brunneus is continuously distributed through- out the Cordillera de la Costa of extreme northern Venezuela. Cebus nigrivittatus nigrivittatus Wag- ner is restricted in Venezuela to the Amazonian region of Territorio Federal Amazonas. Cebus ni- grivittatus olivaceus Schomburg is found in south- eastern Venezuela. Hershkovitz (1949, p. 348) re- ports the type locality as "Vicinity of 'Our Village,' said to be situated at latitude 4°57'N., 61°rW., altitude 3,100 feet above sea level, southern foot of Mount Roraima." We believe that an undescribed subspecies, Ce- bus nigrivittatus subsp., is widely distributed throughout central and northern Venezuela north of the Orinoco. The status of this population is currently under study. The Orinoco Delta region, which Eisenberg and Redford ( 1 979) excluded from their consideration of biogeographic regions due to insufficient data, is now shown by the distributions of Alouatta, Cebus, and Pithecia to be clearly aligned with the Guyana highlands. The Llanos region now extends up to the western edge of the delta, but does not include it, as demonstrated by the distributions of Alouatta and Cebus. Specimens Examined— Total 75. Anzoategui: Los Cocos, Rio Caris (8°30'N, 64°05'W); 10 km W of Laguna de Unare (10°02'N, 65°12'W); Mor- ichal Largo [between Anzoategi and Monagas] (8°18'N, 63°15'W). Aragua: Rancho Grande (10°10'N, 67°19'W). Barinas: Reserva ForestalTi- coporo, on Barinas-Pedregal road (8°03'N, 70°18'W). Bolivar: Caiio La Urbina (7°15'N, 66°25'W); Carretera Caicara-S. Juan de Mana- piare, km 1 75 (6°02'N, 66°29'W); Carretera El Do- rado-Santa Elena, km 33 (6°12'N, 6ri4'W); Cur- aima. El Palmar (8°01'N, 61°26'W); El Dorado- Santa Elena, km 121 (5°18'N,6ril'W);Guayopo, Rio Caura, 14 km from Maripa (7°09'N, 65°10'W); Canaracuni (4°17'N, 64°05'W); Guri, Operacion Rescate (7°18'N, 63°00'W); Rio Antavari (5'i09'N, 63°05'W); Rio Marajano, Meseta de Jaua (4°08'N, 64°1 1'W); Rio Villacoa, 4 km N of mouth (6°16'N, 67'^5'W); San Martin de Turumban, Rio Cuyuni (6"'19'N, 61'^9'W). Carabobo: Bahia de Patanemo (10"'12'N, 67''26'W); Urama (10°12'N, 68°08'W). Cojedes: Cerro Azul, La Blanquera (8°26'N, 68°07'W); Montana Las Loma, San Carlos (9°1 7'N, 68°16'W); Pica, Las Vegas (9°1 5'N, 68°1 7'W). Dis- trito Federal: El Avila, Caracas (10° 14'N, 66°13'W); Hacienda El Limon (lO^B'N, 67°08'W). Falcon: Sanare, Distrito Silva (8''23'N, 68°12'W). Guarico: Hato Flores Moradas, Calabozo (8°23'N, 67"! 3'W); Hato Mapurite, 40 km N of Calabozo (9°08'N, 67°irW); Parmana (7°28'N, 65°18'W); San Jose de Tiznados (9°16'N, 67''16'W). Lara: La Pastora, 1 1 km SSW of Sanare (9°2rN, 70°07'W). Miran- da: Rio Negro (10''20'N, 66°17'W); La Guzma- nera, Guatopo (10°00'N, 66°15'W). Territorio Federal Amazonas: Alto Manapiare (5''13'N, 66°0rW); Alto Ventuari (4°45'N, 65°20'W); Caiio Yureba, Rio Ventuari (3°16'N, 66''21'W); Cano 240 FIELDIANA: ZOOLOGY ■ * m ** T r 2 5 ^ :: 2 • > Z Q. • =» « e e e e e u u u o o o ■ o o n ♦ < c o 3 •c BODINI & PEREZ-HERNANDEZ: CEBIDS IN VENEZUELA 241 < 242 HELDIANA: ZOOLOGY Mayaba, Rio Ventuari (4°07'N, 66°16'W); Cano Morrocoy, Alto Ventuari (5°08'N, 66°02'W); La Esmeralda (S-^S'N, 65°32'W); Ocamo, Rio Oca- mo (2°20'N, 65°15'W); Puruname, 40 km from Rio Orinoco (3°19'N, 65°15'W); La Neblina, E of Rio Varia (0°59'N, 66°10'W). Territorio Federal Delta Amacuro: Cano Araguabisi (9°12'N, 60*'27'W); Guiniquina (9''10'N, 61'X)3'W); Tobe- juba, Guayo (9°09'N, 6r25'W). Yaracuy: Agua Negra (10°14'N, 68''14'W); Carretera Boca de Aroa, 20 km from Palmasola (lO^Ol'N, 69°27'W). Ateles Spider monkeys are widely distributed from northeastern Mexico throughout tropical South America. Four species are recognized, of which only Ateles belzebuth occurs in Venezuela (fig. 7). Ateles belzebuth belzebuth GeofFroy is found in southern Venezuela, south of the Rio Orinoco; most records are from Territorio Federal Ama- zonas. Kellogg and Goldman ( 1 944) indicate a wide distribution in Guyana and report the Venezuelan localities of La Union, Rio Mato, and El Llagual (on both banks of the Rio Caura). Ateles belzebuth hybridus Geoffroy is known from northern and western Venezuela (Hershko- vitz, 1949; Cabrera, 1958; Kellogg & Goldman, 1944). Handley (1976) reported specimens from Apure and Trujillo in western Venezuela. Our specimens are from the states of Barinas, Tachira, and Zulia. Mondolfi and Eisenberg (1979) report- ed it from Cupira and Guatopo, state of Miranda, suggesting a discontinuous distribution on the coast. Specimens Examined— Total 16. Barinas: Re- serva Forestal de Ticoporo, Sabana de Anare (8°06'N, 70°40'W). Bolivar: Canaracuni (4°06'N, 64°10'W). Miranda: Cupira (10°10'N, 65°44'W). Tachira: La Fria (8°13'N, 72°14'W). Territorio Federal Amazonas: Cacuri, Rio Ventuari (4°49'N, 65°26'W); Rio Ocamo, Alto Orinoco (2°44'N, 65°11'W); Salto del Oso, Alto Ventuari (4''55'N, 65°25'W); San Juan de Manapiare (S^M'N, 66°02'W). Zulia: Rio Guasare ( 1 1°02'N, 72°05'W). Lagothrix Woolly monkeys have not been collected in Venezuela; however two subspecies of Lagothrix lagothricha are to be expected: L. lagothricha la- got hricha should be found in Territorio Federal Amazonas south of the Rio Ventuari, and L. I. lugens, in the Selva de San Camilo, state of Apure (Fooden, 1963; Hemandez-Camacho & Cooper, 1976). Pithecia Sakis are found only in northern South America; Hershkovitz (1979) recognized four monotypic species, of which only Pithecia pithecia Linnaeus is found in Venezuela. P. pithecia is found in Ven- ezuela south of the Rio Orinoco and throughout the Guianas and northeastern Brazil. In Venezuela specimens have been reported primarily from the extreme northeastern region, the state of Bolivar and Territorio Federal Delta Amacuro (fig. 5), with a single outlier locality record for Belen, Rio Cunu- cunuma, Territorio Federal Amazonas (3°39'N, 65°46'W) (Handley, 1976). If the distribution pro- posed by Hershkovitz (1979) and Mittermeier and Coimbra-Filho (198 1) is correct, P. pithecia should inhabit the entire region between the upper Rio Orinoco and the Rio Caroni, an enormous area for which no specimens have been recorded. Specimens Examined— Total 24. Bolivar: Rio Curumo (7°15'N, 61°20'W); Rio Grande (8°16'N, 6 ri 7'W); Gurisoco, El Palmar (8°62'N, 6 1°26'W); La Trinidad, El Palmar (7°12'N, 6r23'W); Guri, Operacion Rescate (7°18'N, 63°00'W); Rio Boto- namo, near Rio Cuyuni (6°59'N, 6rirW); San Martin de Turumban, Rio Cuyuni (6''59'N, 61°02'W). Territorio Federal Delta Amacuro: Al- tiplanicie de Nuria (7°50'N, 61°18'W); Yotacuay, SW of Cupiare (8°30'N, 61°00'W). Literature Cited Allen, J. A. 1914. New South American Monkeys. BviUetin of the American Museum of Natural History, 33: 653. BoDiNi, R. 1981. Musculatura locomotora de la viudita (Callicebus torquatus lugens). Sus implicaciones fun- cionales y iilogeneticas. Memoria de la Sociedad de Ciencias Naturales La Salle, 16(XL): 1-165. . 1983. Distribuci6n, status de la investigacion y conservaci6n de los cebidos en Venezuela, p. 1 39. In Symjwsio sobre Primatologia en Latinoamerica. IX Congreso Latinoamericano de Zoologia, Arequipa, Peru. Cabrera, A. 1957(1958]. Calalogo de los mamiferos de America del sur. Revista Museo Argentine de Cien- cias Naturales, "Bernardino Rivadavia," 4(1): xviii + 138 pp. Cruz Lima, E. 1945. Mammals of Amazonia. I. Gen- eral introduction and Primates. Contribution from the BODINI & PEREZ-HERNANDEZ: CEBIDS IN VENEZUELA 243 Museu Paraense Emilio Goeldi de Historia Natural e Etnografia, Belem do Para, Rio de Janeiro, Brasil, 274 pp., 42 pis. EiSENBERG, J. F., AND K. Redford. 1979. A biogeo- graphic analysis of the mammalian fauna of Vene- zuela, pp. 31-36. In Eisenberg, J. F., ed., Vertebrate Ecology in the Northern Neotropics. Smithsonian In- stitution Press, Washington, D.C., 271 pp. Elliot, D. G. 1912. A Review of the Primates. Vols. I and II. American Museum of Natural History, New York. FooDEN, J. 1963. A revision of the woolly monkey (genus Lagothrix). Journal of Mammalogy, 44: 213- 247. Handley, C. O., Jr. 1976. Mammals of the Smith- sonian Venezuelan project. Brigham Young Univer- sity Sciences Bulletin, Biological Series, 20(5): 1-91. Hernandez-Camacho, J., and R. W. Cooper. 1976. The nonhuman Primates of Colombia, pp. 35-69. In Thorington, R. W., Jr., and P. G. Heltne, eds.. Neo- tropical Primates: Field Studies and Conservation. National Academy of Sciences, Washington, D.C. Hershkovitz, P. 1949. Mammals of northern Colom- bia. Preliminary report No. 4: Monkeys (Primates), with taxonomic revisions of some forms. Proceedings of the United States National Museum, 98(3232): 323- 427. . 1958. Type localities and nomenclature of some American primates, with remarks on secondary hom- onyms. Proceedings of the Biological Society of Wash- ington, 71: 53-56. 1963. A systematic and zoogeographic account of the monkeys of the genus Callicebus (Cebidae) of the Amazonas and Orinoco river basins. Mammalia, 27: 1-79. . 1972. The Recent mammals of the Neotropical region: A zoogeographic and ecological review, 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, N.Y. 543 pp. 1977. Living New World Monkeys (Platyrrhi- ni) with an Introduction to Primates. University of Chicago Press, Chicago, 1117 pp. 1 979. The species of sakis, genus Pithecia (Ce- bidae, Primates), with notes on sexual dichromatism. Folia Primatologica, 31: 1-22. 1983. Two new species of night monkeys, ge- nus Aotus (Cebidae, Platyrrhini): A preliminary report on Aotus taxonomy. American Journal of Primatol- ogy, 4: 209-243. . 1984. Taxonomy of squirrel monkeys genus Saimiri (Cebidae, Platyrrhini): A preliminary report with description of a hitherto unnamed form. Amer- ican Journal of Primatology, 7: 155-210. Hill, W. C. O. 1 960. Primates. Comparative Anatomy and Taxonomy. IV. Cebidae, Part A. Edinburgh Uni- versity Press, London, 523 pp. . 1962. Primates. Comparative Anatomy and Taxonomy. V. Cebidae, Part B. Edinburgh University Press, London, 537 pp. INPARQUES. 1982. Guia de los Parques Nacionales y Monumentos Naturales de Venezuela. Ediciones Fundacion de Educacion Ambiental, Caracas. Kellogg, R., and E. A. Goldman. 1944. Review of the spider monkeys. Proceedings of the United States National Museum, 96(3186): 1-45. Kjnzey, W. G. 1982. Distribution of primates and for- est refuges, pp. 455-482. In Prance, G. T., ed.. Bio- logical Diversification in the Tropics. Columbia Uni- versity Press, New York, 7 1 4 pp. MARNR-DGS-POA-SFS. 1982. Informe Nacional de Fauna Silvestre. Ministerio del Ambiente y de los Re- cursos Naturales Renovable, Caracas. Mittermeier, R. a., and A. Coimbra-Filho. 1981. Systematic: Species and subspecies, pp. 29-109. In Coimbra-Filho, A. F., and R. A. Mittermeier, eds.. Ecology and Behavior of Neotropical Primates. Aca- demia Brasileira de Ciencias, Rio de Janeiro. MoNDOLFi, E. 1976. Fauna silvestre de los bosques humedos tropicales de Venezuela, pp. 113-181. In Hamilton, L. S., J. Steyermark, J. P. Veillon, and E. Mondolfi, eds., Conservacion de los bosques humedos de Venezuela. Sierra Club— Consejo de Bienestar Ru- ral, Caracas. MoNDOLR, E., AND J. F. EiSENBERG. 1 979. Ncw records for A teles belzebuth hybridus in northern Venezuela, pp. 93-96. In Eisenberg, J. F., Vertebrate Ecology in Northern Neotropics. Smithsonian Institution Press, Washington, D.C, 271 pp. Ramirez-Cerquera, J. 1983. Reporte de una nueva especit de Primates del genero Aotus de Colombia, p. 146. Symposio sobre Primatologia en Latinoamerica. IX Congreso Latinoamericano de Zoologia. Arequipa, Peru. RuDRAN, R., AND J. F. Eisenberg. 1982. Conservation and status of wild primates in Venezuela, pp. 52-59. In Olney, P. J. S., ed.. International Zoo Yearbook, Vol. 22. The Zoological Society of London, London, 488 pp. 244 FIELDIANA: ZOOLOGY Host Associations and Coevolutionary Relationships of Astigmatid Mite Parasites of New World Primates I. Families Psoroptidae and Audycoptidae Barry M. OConnor ABSTRACTS Coevolutionary patterns among mites of the families Psoroptidae and Audycoptidae and New World Primates are reviewed. Host records for primate parasites originally described from artificial situations are compared with field collections from Peruvian primates, with most host associations verified. A new sp)ecies of Audycoptidae, Saimirioptes hershkovitzi, is described from Cebus apella. The psoroptid subfamily Cebalginae is hypothesized to be a monophyletic group based upon 10 synapomorphies. Phylogenetic relationships within the Cebalginae are reviewed, with cospeciational histories supported for the genera Alouattalges and Schizopodalges and their hosts, and a more diffuse cospeciational pattern observed for the genera Cebalgoides, Cebalges, and Fonsecalges and their hosts. Historical relationships of the genus Procebalges remain problematical. Patrones coevolutivos entre acaros de las familias Psoroptidae y Audycoptidae y primates del Nuevo Mundo son revisadas. Registros de huespedes para parasitos de primates originar- iamente descritos de situaciones artificiales son comparados con colecciones de campo de primates peruanos, con la mayoria de asociaciones de huesped verificadas. Una nueva especie de Audycoptidae, Saimirioptes hershkovitzi, es descrita de Cebus apella. Los psoroptidos de la subfamilia Cebalginae son hipotetizados ser un grupo monofiletico basado en 10 sinapo- morfias. Relaciones filogeneticas entre los Cebalginae son revisadas, con historias coesp>ecia- cionales soportadas F>or el genero Alouattalges y Schizopodalges y sus huesp>edes, y un patron coespeciacional mas difuso observado en los generos Cebalgoides, Cebalges y Fonsecalges y sus huesjjedes. Relaciones historicas del genero Procebalges p>ermanecen problematicas. Padroes coevolucionarios entre os acarinos das familias Psoroptidae e Audycoptidae, e os primatas do Novo Mundo, sao revisados. Os registros de hospedes primatas de parasitas, que foram descritos em situa^oes artificials (cativeiros), sao comparados com cole96es de campo de primatas peruanos, e a maioria das associagoes hospedeiras atualmente registradas foram averiguadas. Uma nova esjjecie de Audycoptidae, Saimirioptes hershkovitzi, encontrada em Cebus apella e descrita. Baseando-se num estudo de 1 0 sinapomorfias, prop6e-se ser a subfamilia psoroptidea, Cebalginae, um grupo monofiletico. As rela96es filogeneticas entre os Cebalginae sao revisadas, e as historias de coesF>ecializa9ao entre os generos Alouattalges e Schizopodalges, e seus respectivos hospedes, sao confirmadas. Os padroes de coespecializa9ao entre os generos Cebalgoides, Cebalges e Fonsecalges, e seus hospedes, sao mais difusos. Os relacionamentos historicos do genero Procebalges permanecem incertos. From the Museum of Zoology and Department of Bi- ology, The University of Michigan, Ann Arbor, MI 48 109. OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 245 Introduction Coevolutionary patterns among hosts and par- asites have been the focus of much recent study and discussion (Brooks, 1979, 1981, 1985; Brooks & Glen, 1982; Futuyma & Slatkin, 1983; Nitecki, 1983). Phylogenetic analysis of evolutionary re- lationships among parasite groups or host groups can provide additional data sets (e.g., treating par- asite distributions as character states of their hosts), or such analyses can be used as tests for hypotheses regarding evolutionary relationships of the other lineages of associated organisms. These phyloge- netic analyses are especially useful when the par- asite groups are host specific and indicate little history of secondary colonization of new hosts. The associations between mites and primates are particularly amenable to such analysis because a number of acarine groups are specifically associ- ated with primates, and these lineages exhibit enough within-group diversity to allow the con- struction and comparison of phylogenetic hypoth- eses. I have previously detailed such hypotheses for several groups of astigmatid mites parasitic on primates (OConnor, 1984). Formulation of hypotheses regarding the his- tory of associations between primates and their associated mites requires three steps. First, the parasite taxa must be described and their natural host and geographic ranges discovered. Second, taxa above the species level in classically derived classifications must be tested for naturalness (i.e., monophyly). Finally, phylogenetic relationships among all taxa must be elucidated. In the present paper, each of these questions will be addressed for the associations among certain groups of as- tigmatid mites and New World Primates. For the reader interested in summary information regard- ing known host-parasite relationships for New World primates, an exhaustive list of literature records may be found in Hershkovitz (1977). One major difficulty in the application of this methodology to the study of the history of mite- primale associations is the scanty knowledge of the distribution and identity of the parasite sp)ecies and their natural host ranges. Although many aca- rine parasites of primates have been described, much of the material has originally come from zoos and primate research centers where the f>os- sibility of unnatural interspecific contact between host species makes the transfer of parasites a real problem. Many other sjjecies have been described from preserved host specimens in museum col- lections where contamination in the field or in the museum may also have been a problem. Finally, the actual field locality from which either parasites or hosts were collected is known for extremely few of the known primate parasites. The uncertainties involved in the host range and geographic distri- bution of so many of the known species of primate parasites render the phylogenetic analyses pro- posed earlier (OConnor, 1984) subject to some doubt. In 1 98 1 , 1 was invited by Philip Hershkovitz to collect parasites from specimens of a number of primate species which had been collected or ob- tained during field studies in Peru in 1 980. A very large number of mites was collected, providing a unique survey of the primate parasites in a small area in Peru and a test for previously reported host-parasite associations. These collections yield- ed specimens belonging to three families of astig- matid mites: Psoroptidae, Audycoptidae, and Atopomelidae. In the present paper, the collec- tions of Psoroptidae and Audycoptidae will be discussed. The collections of Atopomelidae, con- sisting of a number of described and undescribed species of the genus Listrocarpus, will be studied separately. r Materials and Methods Primate specimens were collected in the field by Hershkovitz or obtained from local individuals. These specimens were prepared by removing and simply drying the skins at the time of collection. Upon their arrival in the United States, one or more skins of each species collected were made available to me for parasite removal before the skins were sent for tanning. The following species, all identified by Hershkovitz, were examined (number examined in parentheses): Cebus apella (3); C. albifrons (1); Lagothrix lagothricha (1); L. flavicauda ( 1 ); Alouatta seniculus ( 1 ); Pithecia hir- suta (3); Callicebus moloch (1); Aotus nancymai (4); and Saimiri sciureus (5). Parasites were removed in two ways. First, all skins examined were vigorously brushed over white paper, with the dislodged parasites collected under a dissecting microscop>e. Finally, one skin of each species was soaked in water and mild soap until soft (the single specimen oi L. flavicauda was not soaked). These skins were then gently washed, the wash water filtered through a 200-mesh sieve (mesh 246 HELDIANA: ZOOLOGY openings 75 micrometers), and the residue ex- amined under a dissecting microscope. Mites col- lected were preserved in 70% ethanol for subse- quent study. In the laboratory, mites were cleared in lacto- phenol and mounted in Hoyer's medium, with some specimens retained in alcohol in the cases of large series. Voucher specimens of nominal species are deposited in Field Museum of Natural History, Chicago, and, when available, will be placed in the following institutions: Museum of Zoology, The University of Michigan, Ann Arbor; The United States National Museum of Natural History, Washington, D.C.; LTnstitut Royale des Sciences Naturelles, Brussels, Belgium; and the collection of F. S. Lukoschus, Katholieke Uni- versiteit, Nijmegen, Netherlands. Species Accounts tritonymphs. Among 1 1 protonymphs in the pres- ent collection, eight exhibit characteristics similar to the male tritonymphs (i.e., posterior opistho- somal lobes sclerotized; coxal fields III well scler- otized) while three exhibit characteristics of female tritonymphs (posterior lobes unsclerotized; scler- otization of coxal fields III much weaker). I inter- pret these differences as evidence for sexual di- morphism at the protonymphal instar in this species. Material Examined— Total 133. Twenty-nine females, 43 males (of which 3 1 were in tandem with female tritonymphs), 38 female tritonymphs, 8 male tritonymphs, 3 female protonymphs, 8 male protonymphs, 4 larvae from Cebus albifrons. PERU, Loreto: Nauta, Rio Tigre, 6 km above Rio Tigrillo; 18 December 1980; P. Hershkovitz (9264). Host now a tanned skin (fmnh 122795). Mites labeled bmcx: 81-081 1-3. No specimens were re- covered from Cebus apella. Family PSOROPTIDAE Seven species of mites in the family Psoropti- dae, subfamily Cebalginae, are known to parasitize New World primates. These species, with their known hosts and distributions, are listed below, with new records from the Peruvian collections of Hershkovitz indicated under "Material Exam- ined." Keys to most of these species may be found in Fain (1963c). Cebalgoides cebi Fain, 1963 Cebalgoides cebi Fain, 1 963, Bull. Ann. Soc. Roy. Ent. Belg., 99: 331. Fain, 1963, Bull. Inst. Roy. Sci. Nat. Belg., 39(32): 91. This species was briefly diagnosed by Fain (1963a) from specimens collected from Cebus al- bifrons which originated in "Amerique du Sud" and died in the Antwerp (Belgium) Zoo. It was more thoroughly described and illustrated by Fain (1963c), who listed specimens from Cebus albi- frons from Venezuela (type collection), C apella from "Amerique du Sud," and Leontocebus (Oed- ipomidas) oedipus (= Saguinus oedipus) from Co- lombia. All these hosts had died in the Antwerp Zoo. Fain (1963c) noted sexual dimorphism in tri- tonymphs of C cebi. The two protonymphs he examined exhibited the characteristics of the male Alouattalges corbeti Fain, 1963 Alouattalges corbeti Fain, 1963, Bull. Inst. Roy. Sci. Nat. Belg., 39(32): 122. Fain, 1966, Acarologia 8: 103. Rosalialges cruciformis Lavoipierre, 1 964, Acarologia, 6: 348. This species was briefly described from the ho- lotype female collected from a preserved specimen of Alouatta seniculus macconnelli which had been collected at Paramaribo, Surinam and preserved in the British Museum (Natural History) (Fain, 1 963c). Although only the holotype was described, several specimens were apparently recovered from this host. Almost simultaneously, Lavoipierre (1964a) described and figured the female based upon two specimens collected from an "Aotes'''' (sic) sp. which had died in San Francisco, Cali- fornia, after its importation from Peru. Fain (1966) provided illustrations of the female and illustrated but did not describe the male beyond length and width measurements. Material Examined— Total 15. Five females, 8 males, 2 female tritonymphs from Alouatta se- niculus. PERU, Loreto: Nauta, Rio Samiria; 18 November 1980; P. Hershkovitz (9050). Host now a tanned skin (fmnh 122789). Mites labeled bmoc 81-0809-5. No specimens were recovered from Aotus nancymai, suggesting the possibility that the two specimens collected by Lavoipierre (1964a) represent contamination. OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 247 Schizopodalges lagothricola Fain, 1 963 Schizopodalges lagothricola Fain, 1963, Bull. Ann. Soc. Roy. Ent. Belg., 99: 469. Fain, 1963, Bull. Inst. Roy. Sci. Nat. Belg., 39(32): 100. This species was described from numerous spec- imens from two juvenile Lagothrix lagothricha which died in the Antwerp Zoo. The origin of the hosts was stated as "Amerique du Sud" (Fain, 1 963b). Fain ( 1 963c) provided illustrations of male and female and indicated that the hosts originated in Colombia. Material Examined— Total 30. Eleven fe- males, 1 1 males, 3 tritonymphs, 2 protonymphs, 3 larvae from Lagothrix lagothricha. PERU, Lo- reto: Nauta, Rio Samiria; 15 November 1980; P. Hershkovitz (9032). Host now a tanned skin (fmnh 122790). Mites labeled bmcx: 81-0809-10. Cebalges gaudi Fain, 1 962 Cebalges gaudi Fain, 1962, Rev. Zool. Bot. Afr., 66: 160. Fain, 1963, Bull. Inst. Roy. Sci. Nat. Belg., 39(32): 81. This species was briefly described from numer- ous sp)ecimens collected from a preserved speci- men of Cebus capucinus with no locality infor- mation (Fain, 1962). Full descriptions and figures were provided later (Fain, 1963c). Material Examined— Four females from Ce- bus apella. PERU, Loreto: Nauta, Rio Samiria; 18 November 1980; P. Hershkovitz (9049). Host now a tanned skin (fmnh 122792). Mites labeled bmoc 81-0811-2. Fonsecalges johnjadini Fain, 1962 Fonsecalges johnjadini Fain, 1 962, Rev. Zool. Bot. Afr., 66: 161. Fain, 1963, Bull. Ann. Soc. Roy. Ent. Belg., 99: 468. Fain, 1963, Bull. Inst. Roy. Sci. Nat. Belg., 39(32): 86. This species was briefly described from several Caliithrixjacchus which had died in captivity (Fain, 1 962). Fain ( 1 963b) provided an illustration of the male, and Fain (1963c) gave a complete descrip- tion with figures of both male and female. The hosts, which were listed as eight Hapale jacchus (= Callithrix jacchus) imported from the "bassin de I'Amazone," died in Antwerp. Additional sjiec- imens were recorded from a specimen of Hapale jacchus leucocephalus (= Callithrix jacchus geof- froyi) collected in Bahia, Brazil, and preserved in Brussels. Material Examined— None. Fonsecalges saimirii Fain, 1963 Fonsecalges saimirii Fain, 1963, Bull. Ann. Soc. Roy. Ent. Belg., 99: 330. Fain, 1963, Bull. Inst. Roy. Sci. Nat. Belg., 39(32): 90. Fain, 1966, Acarologia, 8: 107. Dunnalges lambrechti Lavoipierre, 1 964, Acarologia, 6: 343. This species was briefly described from numer- ous specimens collected from two Saimiri sciureus from "Amerique du Sud" and which died in the Antwerp Zoo in 1959 and 1963 (Fain, 1963a). Fain (1963c) provided further descriptive infor- mation and indicated the original hosts had come from "Amazonie" and had died shortly after their arrival at the Antwerp Zoo. He also mentioned additional specimens from "Tamarins spp. ori- ginaires d' Amazonie." The inexact locality infor- mation leaves the actual identity of the type host in doubt following the revision of the genus Sai- miri by Hershkovitz (1984). Lavoipierre (1964a) described and illustrated this species as Dunnalges lambrechti, from "a long series of sjiecimens com- prising all stages" from several Tamarinus nigri- collis (= Saguinus nigricollis) from eastern Peru which died in San Francisco, California. Fain ( 1 966) illustrated parts of this species from the type specimens. Fain (1963c) did not observe sexual dimor- phism in the nymphs of this species, as only two tritonymphs and some shed tritonymphal cuticles were examined. Sexual dimorphism was noted in the tritonymphs during the present study. All lar- vae and protonymphs examined bear three pairs of sclerotized apophyses in the ventrolateral re- gion: a rounded apophysis immediately posterior to trochanter II, a pointed apophysis lateral to the base of leg III, and a pointed apophysis poster- iolaterally near the posterior lobes. Tritonymphs in which these apophyses are retained are here interpreted as males, while those in which all apophyses are lost are interpreted as females. Material Examined— Total 59. Three females from Saimiri sciureus macrodon. PERU, Loreto: Nauta, Rio Tigre, 5 km above Rio Tigrillo; 17 December 1980; P. Hershkovitz (9257). Host now 248 FIELDIANA: ZOOLOGY a tanned skin (fmnh 122810). Mites labeled bmoc 81-0809-15. Two females, 1 male from same host species and locality; P. Hershkovitz (9258, fmnh 122811, BMOC 8 1 -0809- 1 6). Twenty-four females, 8 males, 5 female tritonymphs, 8 male trito- nymphs, 5 protonymphs, 1 larva from same host species. PERU, Loreto: Nauta, Rio Tigre, 6 km above Rio Tigrillo; same date; P. Hershkovitz (9268, FMNH 1 228 1 6, bmoc 8 1 -0809- 1 8). Two fe- males from same host sjjecies and locality; P. Hershkovitz (9267, fmnh 1 228 1 5, bmoc 8 1 -0809- 19). Procebalges pitheciae Fain, 1963 Procebalges pitheciae Fain, 1963, Bull. Ann. Soc. Roy. Ent. Belg., 99: 332. Fain, 1963, Bull. Inst. Roy. Sci. Nat. Belg., 39(32): 96. This species was briefly described from speci- mens collected from a Pithecia monachus im- ported from "Amerique du Sud" and which died in the Antwerp Zoo (Fain, 1963a). The descrip- tions were completed and illustrations provided later (Fain, 1963c). Material Examined— Total 242. Nineteen fe- males, 28 males (7 in tandem with female trito- nymphs), 15 female tritonymphs, 1 protonymph from Pithecia hirsuta. PERU, Loreto: Nauta, Rio Samiria; 30 November 1980; P. Hershkovitz (9 1 1 5). Host now preserved as a tanned skin (fmnh 122797). Mites labeled bmoc 81-0809-7. Forty- seven females, 8 1 males ( 1 7 in tandem with female tritonymphs), 22 female tritonymphs, 8 male tri- tonymphs, 13 protonymphs, 8 larvae from same host and locality; P. Hershkovitz (9088, fmnh 122796, BMOC 81-0809-9). Family AUDYCOPTIDAE Three species of hair follicle inhabiting mites of the family Audycoptidae have been previously de- scribed from New World primates, all from squir- rel monkeys identified as Saimiri sciureus. With the recent recognition of several valid species in the genus Saimiri (Hershkovitz, 1984), the exact identification of the reported hosts is problemat- ical. Specimens representing a fourth sr>ecies were recovered from Cebus apella from the present col- lections. Audycoptes greeri Lavoipierre, 1 964 Attdycoptes greeri Lavoipierre, 1 964, Ann. Natal Mus., 16: 194. This species was described from females col- lected from the sinus-hair follicles of Saimiri sci- ureus collected in eastern Peru and kept in captiv- ity in California (Lavoipierre, 1964b). In the absence of more detailed collection information, the true host may have been either S. sciureus or S. boliviensis, as both occur in eastern Peru (Hersh- kovitz, 1984). Material Examined— Five females from "5fl/- muri sciurea" (sic), without further collection data, from the Lavoipierre collection. University of Cal- ifornia, Davis. Audycoptes lawrencei Lavoipierre, 1964 Audycoptes lawrencei Lavoipierre, 1964, Ann. Natal Mus., 16: 199. This species was described from the same hosts and habitat as Audycoptes greeri (Lavoipierre, 1964b), so the actual specific identity of the host remains uncertain, as indicated for greeri. Material Examined— Two females from "^a/- muri sciurea"" (sic), without further collection data, from the Lavoipierre collection, University of Cal- ifornia, Davis. Saimirioptes paradoxus Fain, 1968 Saimirioptes paradoxus Fain, 1968, Acarologia, 10: 286. This species was described from a single female, containing a larva, collected from a Saimiri sci- ureus which had died in the Antwerp Zoo (Fain, 1968). No information concerning the geographic origin of the host was given, making the specific identity of the host uncertain. Material Examined— None. Saimirioptes hershkovitzi, new species In the following description, all measurements are given in micrometers and are presented as ho- lotype (range of three measured specimens). Female (figs. 1-2)— Body elongate, cylindrical OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 249 Fig. 1 . Saimirioptes hershkovitzi, female. A, dorsum; b, venter. to somewhat flattened dorsoventrally; length in- cluding gnathosoma 4 1 5 (4 1 5—42 1 ), width at level of transverse coxal apodemes III 146 (135-146); entire body with transverse striations which are very thick anteriorly, thin posteriorly. Dorsum (fig. la)— Prodorsal sclerite narrow an- teriorly, much widened medially, narrowed pos- teriorly and fused internally with apodemes of cox- al fields II in most posterior region. Sclerite traversed by anterior transverse striations later- ally. Length of sclerite 70 (70-77). Paired dorsal protuberances present mesal to setae h and sh. with striations encircling protuberances along most of their length; lengths 29 (28-29). Idiosomal chae- totaxy as follows: scapular setae filiform, in a transverse line posterior to prodorsal sclerite, lengths sci 4 (4-5), see 39 (29-39); setae J, and /, very short and rounded, subdivided into two dis- tinct lobes; setae h and sh filiform, lengths 9 (6- 9); setae di subdivided into three parts, a ventral filiform part, length 26 (23-26), and two dorsal rounded lobes; setae d^ similar to ^2 but filiform part shorter, length 19 (14-19), and middle lobe more elongate; remaining setae filiform; /j 29 (28- 250 FIELDIANA: ZOOLOGY < > OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 251 29); /, ventrally positioned, 18 (18-19); ^^4 24 (19- 24); ^5 21 (no variation); l^ ventral, 5 (4-5); l^ ventral, 30 (27-32); a, ventral, 14 (no variation); a, 30 (27-30). Four pairs of idiosomal cupules present: ia dorsal between setae h and sh; im dorsal between and lateral to setae d2 and d^; ip ventral slightly anterior to seta /j; ih ventral, between and mesal to setae U and /j. Opisthosomal glands pres- ent, small, opening between and lateral to setae d^ and /,. Anus terminal. Opening to bursa copulatrix terminal, above anal opening. Venter (fig. lb)— Coxal apodemes well devel- oped. Anterior apodemes of coxal fields I straight, length 66 (57-66), fused medially to form a ster- num of length 23 (23-24). Anterior apodemes of coxal fields II curving mesally, length 75 (75-79); posterior apodemes of coxal fields II transverse, fused ventromedially with anterior apodemes of coxal fields III, extending dorsally, length of ven- tral portion 50 (46-50), length of dorsal portion 31 (28-31). Anterior apodemes of coxal fields III extending almost longitudinally from trochanters III to F>osterior apodemes of coxal fields II, then bending at right angles to fuse with the latter, length of longitudinal portion 49 (49-51), transverse portion 39 (39-44). Anterior apodemes of coxal fields IV very wide, anteriomedially directed, length 50 (50-53). Ovipore located between coxal fields III and IV; a crescentic epigynal apodeme posi- tioned anterior to ovipore; genital valve elongate, disappearing posteriorly under transverse flap; a pair of small thickenings along posterior, internal portion of valves; two pairs of vestigial genital papillae lateral to genital ojiening. Three pairs of filiform coxal setae present: cxI at posterior end of coxal fields I between coxal apodemes II, length 12 (9-12); cxIII on median apex of coxal apo- demes IV, length 9 (6-9); cx/P'mesad of the base of leg IV, length 7 (6-7). Genital setae absent. Gnathosoma (fig. 2g)— Gnathosoma elongate, somewhat widened posteriorly, then tapering to a point ventrally, length (excluding chelicerae) 62 (59-62), maximum width 26 (24-26). Chelicerae elongate, with small, toothed digits, length 46 (44- 46). Palps apparently two-segmented; basal seg- ment bearing two very short, filiform setae, 1 prox- imodorsal, 1 medioventral; distal segment with short solenidion and 5 sclerotized, pxjinted pro- jections. Subcapitulum with pair of short, filiform setae medioventrally; with pair of sclerotized, pointed projections positioned slightly anterior to subcapitular setae, and unpaired projection ante- riomedially; median rutellar lobes with sclerotized points medially. Legs (figs. 2a-f) — Legs I-II similar in structure, with all segments free, lengths 84 (80-90). Femora I-II bearing two ventral apophyses, one filiform seta vF. lengths: I, 9 (8-9); II, 33 (30-33). Genua I-II each with large ventral apophysis; setae cG expanded basally, then tapering, lengths 22 (20- 25), setae mG filiform, lengths 43 (40-45). Tibiae I-II each with large ventral apophysis; setae gT short, filiform, lengths 6 (5-6); solenidia 0 api- codorsal, lengths: I, 20 (18-20); II, 22 (21-23). Tarsi I-II each with pointed dorsal apophysis, two distal hooked apophyses; chaetotaxy of tarsi I-II similar; setae d as strongly hooked, apical claws, lengths 5 (no variation); e filiform, lengths 40 (43- 50); /filiform, lengths 7 (6-7); la filiform, lengths 6 (5-6); wa very short, not measurable; ra filiform, lengths 6 (5-6); solenidia «, blunt apically, lengths: I, 14 (no variation); II, 26 (24-26). Pretarsi I-II similar, total lengths 40 (37-40); pretarsi divided into long ambulacral stalk and rounded ambula- cral disc; ambulacral disc with condylophore guide and ventral "rays" (fig. 2b); condylophores atten- uate, appearing to divide at base of ambulacral disc. Legs III-IV similar, each with fused trochanter- femur bearing a large, ventral apwaphysis. Genua III-IV glabrous. Tibiae III-IV each with setae kT enlarged, bifurcate, and deeply rooted; tibia III with solenidion 0, length 6 (6-7), 0 IV absent. Tarsi III-IV each with apical, pointed apophysis, three setae: fi^ apical, lengths: III, 100(91-100); IV, 97 (86-97); r filiform, lengths 7 (6-7); w filiform, lengths 3 (no measurable variation). Types— Holotype and two paratype females from Cebus apella. PERU, Loreto: Nauta, Rio Samiria; 18 November 1980; P. Hershkovitz (9049). Host now a tanned skin (fmnh 122792). Mites labeled BMOC 81-0811-2. Holotype and 1 paratype de- posited in Field Museum of Natural History, Chi- cago; 1 paratype in Museum of Zoology, Univer- sity of Michigan, Ann Arbor. Systematic Position— Saimirioptes hershko- vitzi shares with 5". paradoxus, the tyF>e-sp)ecies and only other member of the genus, the presence of dorsal lobes between setae ^1 and t/j^ the similar form of setae t/, and /, (rounded, bifurcate, and without filiform part), and the expansion of the bases of setae cG of genua I-II. The new sF>ecies differs from the type-species in the relative lengths of the scapular setae (sce:sci 5-6:1 in S. hershko- vitzi, 1.5:1 in S. paradoxus); the shorter filiform fKJrtion of setae f/, (not reaching dy in S. hersh- kovitzi, extending past d^ in S. paradoxus); the shorter lengths of setae 4 (29 in S. hershkovitzi. 252 HELDIANA: ZOOLOGY 67 in S. paradoxus); the presence of setae U (absent in S. paradoxus); the longer pretarsi I-II (37-40 in S. hershkovitzi, 17 in S. paradoxus); and the greater lengths of setae d on tarsi III-IV (at least twice as long as the entire leg in S. hershkovitzi, less than half the leg length in S. paradoxus). Discussion The Peruvian collections reported upon here support the hypothesis that most of the records of primate-Cebalginae associations previously re- ported reflect natural host-parasite associations. The association between Alouattalges corbeti and Aotus species was not verified and remains ques- tionable. As there were no field collected repre- sentatives of the Callitrichidae examined during this study, the occurrence of cebalgine mites on these hosts remains generally untested in natural situations, with only Fonsecalges johnjadini re- corded from noncapti ve callitrichids (Fain, 1 963c). Having established that most records of pri- mate-cebalgine associations reflect natural asso- ciations, two additional hypotheses must be pro- posed and tested before any coevolutionary hypotheses may be tested. These preliminary hy- potheses concern the monophyly of the Cebalginae and the phylogenetic relationships of the taxa within the group. I have previously discussed these questions (OConnor, 1 984) but, due to space con- straints, was unable to detail the reasoning behind my conclusions. Monophyly of the Cebalginae Before any hypothesis of historical associations may be tested, at least one of the lineages must be hypothesized to be monophyletic (Brooks, 1981). In this study, the psoroptid subfamily Cebalginae, which comprises all psoroptid mites parasitizing New World Primates, must be tested for mono- phyly. In last defining this subfamily. Fain ( 1 963c) listed 21 character states for the group. Because the comprehensive morphological studies of Fain (1963c) were carried out before the methods of phylogenetic systematics became widely discussed and utilized, no distinction between ancestral and derived states was made in the diagnosis of the Cebalginae. In order to test whether the Cebalgi- nae represents a monophyletic group, I have ex- amined the characters listed by Fain (1963c) using outgroup comparison to polarize the states. I have previously hypothesized that the taxa comprising the four subfamilies of Psoroptidae which para- sitize the Primates form a monophyletic group (OConnor, 1984). These taxa include the Makial- ginae (sensu OConnor, 1984; i.e., including the Cheirogalalginae and Galagalgidae of Fain), par- asites of the Strepsirrhini; the Paracoroptinae, par- asites of African Cercopithecidae and Hominidae; the Nasalialginae, parasites of Asian Cercopithe- cidae; and the Cebalginae, from New World pri- mates. This grouping of taxa is regarded as the ingroup in the following analysis. Outgroups used in defining the character state polarities were the other subfamilies of Psoroptidae and earlier de- rivative groups in the Astigmata. Character Analysis 1 . Presence of retrograde apophyses on coxa! fields III. This state is unique to the cebalgine genera, with such apophyses not occurring in other taxa in the ingroup or outgroup. 2. Male body size much smaller than female. In the outgroups and other ingroups, males are similar in size to females or somewhat larger or smaller. The substantial reduction in body size of male Cebalginae is unique. 3. Female opisthosoma more or less squared posteriorly and laterally, reinforced with sclero- tized areas. This condition is exhibited by all Ce- balginae, but not in any outgroup. Among the in- groups, the squared body is unique to the Cebalginae, but sclerotized reinforcement is also found in the Nasalialginae and some Paracorop- tinae (Pangorillalges). Among outgroup taxa, sclerotized reinforcement is present in some Pso- ralginae (Edentalges), but again without the squared body form. 4. Male with legs III very modified, with 3-4 terminal segments fused and bearing medially di- rected projections. The modification of the third pair of legs in male Cebalginae is unique and pres- ent in all taxa. No similar modification occurs anywhere in ingroup or outgroup. 5. Reduction or loss of paranal suckers in males. Paranal suckers are present in males in the out- groups and ingroups. In the Cebalginae, the suck- ers are very reduced or absent. 6. Tarsus I with 2 apical solenidia. I regard the apical displacement of solenidion oj, in the Ce- balginae as derived. In the other ingroups and most outgroups, this solenidion is median or basal on OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 253 the tarsus. Apical displacement of this solenidion also occurs in the subfamily Psoroptinae and the monobasic Marsupialginae, conditions I regard as convei^ent. 7. Loss of dorsal seta ^,. Seta d^ is absent in all Cebalginae and retained in the other ingroups and most outgroups. This seta is also lost in the Lis- tropsoralginae and the psoroptid parasites of ro- dents (Echimyalges. Myoproctalges, and Coen- dalges), conditions I regard as convergent. 8. Reduction or loss of apophyses from the pos- terior tarsi of the female. Apophyses are present on the posterior tarsi of some of the outgroups such as the Listropsoralginae and are retained in some ingroups, the Makialginae and Paracorop- tinae. I consider the presence of these apophyses to be plesiomorphic for the primate-associated psoroptid lineage. In the Cebalginae, taxa in which females have well-developed posterior legs retain vestiges of these apophyses, while taxa in which the legs are reduced retain no traces. I consider this reduction to be a derived state for the Cebal- ginae and regard the loss of these structures in the Cebalginae and Nasalialginae as convergent. 9. Posterior edge of female opisthosoma with 2 pairs of long, strong setae. In most ingroups and psoroptid outgroups, seta /, is long and strong while seta di is shorter and thinner. Of the ingroups, setae l^ and d^ are equally well developed in the Cebalginae and Nasalialginae. However, in the Ce- balginae, setae /, and d^ are closely associated, usu- ally on a single projection, while in the Nasalial- ginae these setae are separate, on distinctly different projections. I regard these two conditions as con- vergent. I also regard as convergence the elonga- tion of seta di in some Psoralginae. 10. Dorsal seta l^ sometimes absent. Seta I4 is absent in all Cebalginae except Cebalgoides cebi. The seta is retained in that species, the other in- groups, and most outgroups. The loss of seta I4 could be regarded as a synapomorphy defining a group containing all Cebalginae except Cebal- goides. However, this hypothesis conflicts with groupings suggested by all other characters (see page 256). At this point, it is more parsimonious to regard the loss of seta U as a synapomorphy for the Cebalginae, with a reversal in Cebalgoides. 1 1 . Loss of retrograde apophyses on the ante- rior legs. Retrograde apophyses are present on the anterior legs in some of the outgroups, notably the Listropsoralginae and Marsupialginae in the Pso- roptidae and in the related families Audycoptidae, Rhyncoptidae, and Myocoptidae. Among the in- groups, these apophyses are retained in the Ma- kialginae but lost in the Cebalginae, Paracorop- tinae, and Nasalialginae. I regard this loss as characterizing these latter three groups as a natural unit and thus plesiomorphic for the Cebalginae. 12. Female bursa copulatrix subterminal or ventral. In the outgroups, the female bursa cop- ulatrix is terminal, as it is in the Cebalginae, Ma- kialginae, and Nasalialginae among ingroups. I re- gard the dorsal position of the bursa in the Paracoroptinae as derived. Thus, this character retains the plesiomorphic condition in the Cebal- ginae. 13. Base of the gnathosoma with retrograde apophyses. In some outgroups as well as in the Cebalginae, Makialginae and some Paracordpti- nae (i.e., Pangorillalges), the base of the gnatho- soma bears retrograde apophyses. These are ab- sent in certain outgroups (e.g., the Psoroptinae) and in Paracoroptes and Nasalialges among in- group taxa. I regard the retention of these apoph- yses in the Cebalginae as plesiomorphic. 14. Absence of retrograde apophyses on coxae I-II. Retrograde apophyses occur on coxae I-II only in the Makialginae. They do not occur on both coxal fields in any other ingroup or outgroup taxa, although the Listropsoralginae and Myocop- tidae have apophyses on coxae II. I regard, these structures as synapomorphies characterizing the Makialginae, and their absence in the Cebalginae as plesiomorphic. 15. Well-developed "claws" on tarsi I-II. Clawlike apophyses on the apices of tarsi I-II are present in many outgroup taxa, including both mammal and bird parasites. I regard the presence of this character state as ancestrally characterizing all Psoroptidia, and thus plesiomorphic for the Cebalginae. 16. Tarsus II with solenidion apically dis- placed. Solenidion oj is apically displaced in the Cebalginae, Paracoroptinae, and Nasalialginae. In most outgroups and in the Makialginae, the so- lenidion retains its ancestral, basal position. I re- gard this displacement as diagnosing a natural group consisting of the above three primate-as- sociated subfamilies and thus plesiomorphic for the Cebalginae. 17. Female tarsal chaetotaxy 7-7-6—4 or -5. Possession of 7 setae on tarsi I-II involves the loss of seta ba in the Paracoroptinae and Cebalginae and probably the Nasalialginae. Retention of 8 setae on these segments (including ba) is charac- teristic for most outgroup taxa and the Makial- ginae. I regard this state as diagnosing a natural group comprising the Cebalginae, Paracoroptinae, 254 HELDIANA: ZOOLOGY and Nasalialginae and thus plesiomorphic for the Cebalginae. 1 8. Angles of female opisthosoma generally with strong hooks. Such hooks are found in the cebal- gine genera Cebalges, Cebalgoides. and Fonse- calges, but not in other taxa in the outgroup or ingroup. I regard the presence of these structures as characterizing a monophyletic unit within the Cebalginae but not the group as a whole. 19. Posterior legs of female normal or atro- phied. Legs III-IV of the female are reduced in size and may exhibit fusion of segments in the genera Cebalges and Fonsecalges but not in other Cebalginae nor other members of the ingroup. Well-developed legs are characteristic of most out- group taxa, although similar reduction of the pos- terior legs occurs in some Psoroptinae and Pso- ralginae. I regard the atrophied legs of some Cebalginae as characterizing a smaller monophy- letic unit within the group and convergent with the outgroup taxa noted above. 20. Posterior legs of nymphal stages normal or short and atrophied and bearing a long seta. The reduction of the legs in the nymphs is found in all cebalgine genera except Procebalges. It is also found in a few outgroup taxa (e.g., the Psoroptinae and Psoralginae), but not in most outgroups or in other ingroup taxa. I regard this state as diagnosing a group within the Cebalginae and not characteriz- ing the group as a whole. 2 1 . Dorsal seta d^ sometimes absent. Seta d^ is absent in Schizopodalges lagothricola and Fon- secalges johnjadini (but not F. saimirii), and pres- ent in other Cebalginae, other ingroups, and most outgroups. The distribution of this derived state within the Cebalginae strongly suggests the inde- pendent loss of this seta in the two species sharing the state. This character at best diagnoses a group within the Cebalginae, but is more likely a case of convergence. In no way does this character diag- nose the group as a whole. Conclusions On the basis of the above analysis, I conclude that, of the 21 character-states listed by Fain (1963c) as diagnosing the Cebalginae, five repre- sent unique synapomorphies which diagnose the Cebalginae and occur in no other group (character- states 1-5), five are synapomorphies which diag- nose the Cebalginae but which also occur as con- vergent states in certain other taxa (character-states 6-10), seven are symplesiomorphies diagnosing larger groups which include the Cebalginae (char- acter-states 11-17), and four represent within- group apomorphies diagnosing smaller groups within the Cebalginae (character-states 1 8-2 1 ). The ten synapomorphies diagnosing the Cebalginae leave no doubt that the group is a natural one. Phylogenetic Relationships Within the Cebalginae I have previously presented a hypothesis of phy- logenetic relationships among genera in the Ce- balginae based upon 17 characters (OConnor, 1 984). This cladogram is reproduced here (fig. 3), with numbers on the cladogram referring to the derived states of the characters listed below. In the earlier study, space limitations prevented discus- sion of the different states of these characters and the reasons for interpreting their polarity. Addi- tionally, an error in character 1 2 appeared in the list of character-states. As I have hypothesized that the Cebalginae form a monophyletic group within a larger lineage comprising the Makialginae, Par- acoroptinae, and Nasalialginae, taxa in these three lineages were used as outgroups to polarize the character-states within the Cebalginae. 1. Female with coxal apodemes III-IV fused. In all taxa in the outgroups, coxal apodemes III- IV end freely. Within the Cebalginae, these apo- demes are fused together on either side in the gen- era Procebalges, Schizopodalges, and Fonsecalges. Although I regard the fused condition to be de- rived, conflicts with many other characters suggest that these apodemes have fused independently in the three genera. 2. Female epigynum fused with coxal apo- demes I. In most outgroup taxa, the female ovi- pore is located between coxal fields II and III. There has been a trend toward anterior displace- ment of the ovip)ore in a number of psoroptid lineages (e.g., Psoralginae, Listropsoralginae, Pso- roptinae), and the more anterior p>osition occurs in certain taxa among the primate associated lin- eage as well (e.g., Lemuralges in the Makialginae). Within the Cebalginae, this derived state occurs in the genera Procebalges and Schizopodalges. Be- cause this character-state distribution conflicts with many other characters, I regard the presence of this state in the two genera as convergence. 3. Male with paranal suckers lost. In all out- groups, the male retains paranal suckers. Within the Cebalginae, the suckers are retained only in OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 2SS V) O O O) ,^*— i...^. D >< o l« a -c o o N OD -C D o ~~j oo . ^ VI -2 < «/> 3 C 3 D5 D 00 O ^ O^ to 13 D -Q -Q Q) 0) U O 0) ^ — ■ D) • "^ C o o 00 u^ — ^ ■ 17 ■ 16 ■ 1 Fig. 3. Phylogenetic relationships among genera in the subfamily Cebalginae. Numbers refer to derived states of characters discussed in the text. Hosts of each mite taxon given in parentheses. Procebalges. I regard the loss of these structures as derived and characterizing the sister group of Procebalges. 4. Immature stages with pretarsi III-IV lost. Described immature stages of outgroup taxa all retain the pretarsus on legs III-IV. Within the Ce- balginae, only Procebalges retains this ancestral state. In all other Cebalginae, pretarsi III-IV are lost in the immature stages, a condition I regard as derived. 5. Gnathosomal apophyses lost in both sexes. As discussed in the previous section, I regard the presence of gnathosomal apophyses as ancestral in the Cebalginae. These structures are lost in both sexes in the genera Schizopodalges and Alouat- talges within the Cebalginae, a condition I regard as derived. I regard this state as a synapomorphy for these two genera and convergent with the sim- ilar state occurring in the genera Lemuralges (Makialginae), Paracoroptes (Paracoroptinae), and Nasalialges (Nasalialginae). I have treated loss of these apophyses in the female only as a separate character (16, below). 6. Female with seta 5 of tarsi I-II in the form of a strongly hooked claw. In the outgroup taxa, seta s is simple and filiform. Within the Cebalgi- nae, this state is retained in all taxa except Schi- zopodalges and Alouattalges where the seta is en- larged and hooked. I regard the latter state as derived and a synapomorphy for the two genera. 7. Male with a large spur on tarsus III. This condition is unique to the cebalgine genera Schi- 256 FIELDIANA: ZOOLOGY zopodalges and Alouattalges. In the outgroups and other ingroup taxa, such a spur does not exist. I regard the presence of this character-state as in- dicative of common ancestry of these two genera. 8. Male with ridges on leg III restricted to tibial element. Ridges on the distal portion of leg III of the male characterize the Cebalginae (see above). Such ridges do not occur in any outgroup. I regard the restriction of the ridges to a spur extending from the tibial portion of the leg as a derived state defining the group Schizopodalges + Alouattalges because this condition forms a functional complex with the spur on tarsus III (character 7) and a dorsal spur on femur IV. This latter character was not considered in the previous analysis (OConnor, 1984). The conjunction of the three spurs serves to lock legs III and IV together into a unit. 9. Female with recurved hooks on posterior border of opisthosoma. Strong hooks are present in this position in females of Cebalgoides, Ce- balges. and Fonsecalges. Weaker development in this area is characteristic of other cebalgine genera, while in the outgroups, no projections exist. I re- gard the possession of strong hooks as a derived condition defining a lineage comprising the three genera noted above. 10. Male with pretarsus III lost. Males retain a pretarsus on leg III in most outgroup taxa. This pretarsus is lost in the genera Galagalges and Chei- rogalalges (Makialginae), and also in the cebalgine genera Cebalgoides, Cebalges, and Fonsecalges. I regard the loss of pretarsus III as a synapomorphy for the latter three genera within the Cebalginae, with independent loss in the makialgine lineage comprising the former two genera. 1 1 . Female without opisthosomal sclerite. A median sclerite is present in females of most out- group taxa and is retained in all Cebalginae except Schizopodalges. I regard this loss as derived in the latter genus. Convergence in this character occurs with some outgroup taxa. This sclerotization is also lost in Gaudalges caparti, and Lemuralges (Makialginae), and Nasalialges (Nasalialginae). 12. Female with seta s of tarsi III-IV enlarged and clawlike. In the previous study (OConnor, 1984), a lapsus occurred in that the state "seta 5 of tarsi III-IV reduced" was listed as a derived condition for the genus Alouattalges. In fact, pos- session of a small, filiform seta s on these tarsi must be regarded as the ancestral condition, as it occurs in all outgroup taxa as well as in most Ce- balginae. In the genus Alouattalges, seta 5 is en- larged and recurved on tarsi III-IV, a condition I regard as the true derived state. 13. Male with opisthosomal lobes widely spaced. In most outgroup taxa, the opisthosomal lobes of the male are close together, a condition also found in most Cebalginae. These lobes are widely spaced in the genus Cebalgoides within the Cebalginae, a condition I regard as derived. Con- vergence occurs, with this state also occurring in Nasalialges (Nasalialginae). 1 4. Male with apodeme between genital and anal region strongly reduced. In the outgroup taxa, males do not possess a transverse apodeme between the genital and anal regions. A large apodeme is pres- ent in this position in most male Cebalginae. In the prior analysis (OConnor, 1 984), I stated that this apodeme was absent in Cebalgoides as indi- cated by Fain (1963c) and regarded this as a re- versal. Closer examination of a number of speci- mens indicates that a very small apodeme is present in some males of this taxon. I regard the presence of the apodeme as a derived condition for the Cebalginae, and its reduction or loss in Cebal- goides as a further derived state. 1 5. Female with tibiae-tarsi III-IV fused. In the outgroups, the tibia and tarsus of legs III-IV are freely articulated. This condition is present in most Cebalginae as well. In the genera Cebalges and Fonsecalges, these segments are fused in the fe- male, a condition I regard as derived within the Cebalginae. Similar fusions occur in some other psoroptid subfamilies but not within the primate- associated lineage. 16. Female with gnathosomal apophyses lost. Gnathosomal apophyses are retained in females of most outgroup taxa and are retained in some Cebalginae. These apophyses are lost in the female but retained in the male in the genus Fonsecalges. I regard this loss as independent of the loss of apophyses in the Schizopodalges-Alouattalges lin- eage, where the apophyses are lost in both sexes (character 5). 17. Female with pretarsi III-IV lost. Pretarsi III-IV are retained in the female in all outgroup taxa and in all Cebalginae except Fonsecalges. I regard this loss as derived and convergent with the similar loss in females in other psoroptid groups (e.g., some Psoroptinae, Psoralginae). History of Primate-Cebalgine Evolution The present knowledge of the diversity and host associations of the family Audycoptidae is not suf- ficient for formulating hypotheses concerning the OCO^fNOR: MITE PARASITES OF NEW WORLD PRIMATES 257 history of their host associations. However, given the phylogenetic relationships among cebalgine taxa presented in the cladogram (fig. 3), two meth- ods exist for using these relationships to make hy- potheses regarding the history of the associations between the New World primates and the Cebal- ginae. The first method would involve comparing the phylogenetic hypothesis for the parasite group with a similar hypothesis for the hosts. The hy- pothesis tested by this comparison is whether the current pattern of host-parasite associations di- rectly results from strict cospeciation between the hosts and their parasites. Given a phylogenetic hypothesis for each group, such a hypothesis is easily tested. The second type of conclusion which might be drawn from the parasite cladogram is a phylogenetic hypothesis concerning host relation- ships. This method is dependent upon the as- sumption of cospeciation, or at least noncoloni- zation, between the two lineages. The current state of knowledge concerning the phylogenetic relationships among New World pri- mates is somewhat confused. Many early hypoth- eses were based upon classical methodology in which ancestral and derived character-states were not differentiated in proposing hypotheses of re- lationships. Schwartz et al. (1978) summarized the state of knowledge only a few years ago by saying that "Platyrrhini apF>ears in and of itself to be a natural group, although both its wider relation- ships and the relationships among its members remain unclear" (p. 128). Much new information relating to this question was presented in symp>osia in 1978-1979 and published in a volume edited by Ciochon and Chiarelli (1980). In the following discussion, I refer to phylogenetic hypotheses for New World primates suggested by Rosenberger (1977) and contributors to the Ciochon and Chi- arelli volume as representing the most modem phylogenetic thinking regarding Platyrrhine rela- tionships. Returning to the question of cospeciation be- tween primates and cebalgine mites, the first dif- ficulty encountered is the lack of a consensus con- cerning phylogenetic relationships among all New World primates. Phylogenetic hypotheses based on the following types of data sets are in strong disagreement: dentition (Rosenberger, 1977; 3 separate hypotheses); integumentary characters (Perkins & Meyer, 1980); karyology (Chiarelli, 1980); immunological methods (Sarich & Cronin, 1980); and other immunological data and protein sequence data (Baba et al., 1 980). Despite the strong disagreement among these data sets, some patterns are common to several or all of these. These may be compared with the cladogram of mite relation- ships to test the cospeciational hypothesis. One host relationship which is supported by al- most all data sets is the relationship between Al- ouatta and the Atelinae (including Lagothrix). Of the hypotheses cited above, only the karyological evidence did not support this relationship, and then only because the karyotype of Lagothrix is so derived that Chiarelli ( 1 980) made no hypoth- esis as to its relationships. This relationship be- tween Alouatta and Lagothrix is mirrored by the sister group relationship between their parasites, Alouattalges and Schizopodalges, which is among the most strongly supported relationships among the Cebalginae. A second host relationship which is partially mirrored in the mile phylogeny is the relationship between the genera Cebus and Saimiri, which was supported by all data sets except the integumen- tary data (Perkins & Meyer, 1980). In this case, the mite phylogeny suggests a cospeciational pro- cess, but some additional hypotheses are required to explain the distributions of the genera Cebal- goides, Cebalges, and Fonsecalges. The relation- ships among these three parasite genera are com- patible with a cospeciational scenario if an early sjjeciation event between the Cebalgoides and the Cebalges-Fonsecalges lineages occurred in con- junction with a common ancestor of Cebus and Saimiri. Extinction of the Cebalgoides lineage on the Saimiri line, with cospeciation of the Cebalges- Fonsecalges lineage in both host lines, leads to the present distribution on the Cebidae. In all cases, colonization events are required to explain the presence of both of these lineages on the Callitrich- idae, given the probable monophyly of that taxon. Certain other relationships among the parasite taxa are not mirrored by host phylogenies. The sister group relationship between Procebalges and all other cebalgines is not reflected in any hyp)oth- esis of host relationships. The relationship of Pi- thecia to the Alouatta-Lagothrix lineage was sug- gested by both dental data (Rosenberger, 1977) and karyology (Chiarelli, 1 980). Interestingly, the integumentary data (Perkins &. Meyer, 1980) sug- gest that Pithecia retains the most plesiomorphic skin characteristics of any mite-bearing New World primate (Aotus was regarded as even more plesio- morphic). As cebalgine mites are skin inhabitants, it might be suggested that plesiomorphic skin re- tains plesiomorphic mites, a hypothesis which would require the ancestral possession of a Pro- cebalges lineage on other cebids with subsequent 258 HELDIANA: ZOOLOGY extinction on all but Pithecia in order to retain a basically cospeciational history. Discovery of ce- balgine mites on hosts related to Pithecia (Chi- ropotes, Cacajao) will provide a test of these hy- potheses. The utility of using the proposed mite phylogeny to infer a host phylogeny at higher levels than mentioned above depends upon the degree to which cospeciational patterns outweigh colonizations or extinctions in the hypothesized historical relation- ships. Among the entire psoroptid lineage para- sitizing all Primates, cospeciational patterns ap- pear to be supported in a large majority of cases (OConnor, 1984). However, noncosj)eciational patterns such as the distribution of the genus Le- muralges in the Makialginae, or the cebalgine par- asites of the Callitrichidae, make the use of these mites as consistent indicators of host phylogeny at least somewhat suspect. Given the large dis- parity in phylogenetic hypotheses generated from subsets of the overall character matrix for the New World Primates, this parasite data deserves to be at least considered by future workers in this area. Acknowledgments I would like to thank Philip Hershkovitz, with- out whose cooperation and interest this study could not have been attempted. I also thank Robert Timm and Bruce Patterson, Field Museum of Nat- ural History, for their hospitality and cooperation during the processing of host specimens and for their critical review of the manuscript. I thank J. H. S. Klompen, University of Michigan, for his comments on the manuscript. I thank Margaret van Bolt, Museum of Zoology, University of Michigan, for assistance with the illustrations. Literature Cited Baba, M., L. Darga, and M. Goodman. 1980. Bio- chemical evidence on the phylogeny of Anthropoidea, pp. 423-443. In Ciochon, R. L., and A. B. Chiarelli, eds.. Evolutionary Biology of the New World Monkeys and Continental Drift. Plenum Press, New York. Brooks, D. R. 1979. Testing the context and extent of host-parasite coevolution. Systematic Zoology, 28: 299- 307. studying the evolution of ecological associations. An- nals of the Missouri Botanical Garden, 72: 660-681. Brooks, D. R., and D. R. Glen. 1982. Pinworms and Primates: A case study in coevolution. Proceedings of the Helminthological Society of Washington, 49: 76- 85. Chiarelli, A. B. 1980. The karyology of South Amer- ican Primates and their relationship to African and Asian species, pp. 387-398. In Ciochon, R. L., and A. B. Chiarelli, eds.. Evolutionary Biology of the New World Monkeys and Continental Drift. Plenum Press, New York. Ciochon, R. L., and A. B. Chiarelli, eds. 1980. Evo- lutionary Biology of the New World Monkeys and Continental Drift. Plenum Press, New York, 528 pp. Fain, A. 1 962. Diagnoses d'acariens nouveaux. Revue de Zoologie et de Botanique Africaines, 66: 154-162. . 1 963a. Nouveaux acariens psoriques parasites de marsupiaux et de singes sud-americains (Psoralgi- dae: Sarcoptiformes). Bulletin et Annales de Societe Royale d'Entomologie de Belgique, 99: 322-332. . 1 963b. Un nouvel acarien producteur de gale chez un singe sud-americain. Bulletin et Annales de Societe Royale d'Entomologie de Belgique, 99: 467- 470. 1 963c. Les acariens producteurs de gale chez les lemuriens et les singes avec une etude des Psorop- tidae (Sarcoptiformes). Bulletin du Institut Royal des Sciences Naturelles de Belgique, 39(32): 1-125. 1 966. Les acariens producteurs de gale chez les lemuriens et les singes II. Nouvelles observations avec description d'une espece nouvelle. Acarologia, 8: 95- 114. . 1968. Notes sur trois acariens remarquables . 1981. Hennig's parasitological method: A pro- posed solution. Systematic Zoology, 30: 229-249. . 1985. Historical ecology: A new approach to (Sarcoptiformes). Acarologia, 10: 276-291. FuTUYMA, D. J., AND M. Slatkin, EDS. 1983. Cocvo- lution. Sinauer Associates Inc., Sunderland, Mass., 555 pp. Hershkovitz, P. 1977. Living New World Monkeys (Platyrrhini), vol. 1 . University of Chicago Press, Chi- cago, 1117 pp. . 1984. Taxonomy of squirrel monkeys genus Saimiri (Cebidae, Platyrrhini): A preliminary report with description of a hitherto unnamed form. Amer- ican Journal of Primatology, 7: 155-210. Lavoipierre, M. M. J. 1964a. A note on the family Psoralgidae (Acari: Sarcoptiformes) together with a description of two new genera and two new species parasitic on primates. Acarologia, 6: 342-352. . 1964b. A new family of acarines belonging to the Suborder Sarcoptiformes parasitic in the hair fol- licles of Primates. Annals of the Natal Museum, 16: 191-208. NiTECKi, M. H., ED. 1983. Coevolution. University of Chicago Press, Chicago, 392 pp. OConnor, B. M. 1984. Co-evolutionary patterns be- tween astigmatid mites and primates, pp. 186-195. In Griffiths, D. E., and C. E. Bowman, eds., Acarology VI, vol. 1. Ellis Horwood Ltd., Chichester, England. Perkins, E. M., and W. C. Meyer. 1980. The phy- logenetic significance of the skin of primates: Impli- cations for the origin of New World monkeys, pp. OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 259 331-346. In Ciochon, R. L., and A. B. Chiarelli, eds., R. L., and A. B. Chiarelli, eds., Evolutionary Biology Evolutionary Biology of the New World Monkeys and of the New World Monkeys and Continental Drift. Continental Drift. Plenum Press, New York. Plenum Press, New York. RosENBERGER, A. L. 1977. A'e«o//?r/jc and ceboid phy- Schwartz, J. H., I. Tattersall, and N. Eldredge. logeny. Journal of Human Evolution, 6: 461-481. 1978. Phylogeny and classification of the Primates Sarich, V. M., AND J. E. Cronin. 1 980. South Amer- revisited. Yearbook of Physical Anthropology, 21: 95- ican mammal molecular systematics, evolutionary '33. clocks, and continental drift, pp. 399-422. In Ciochon, 260 HELDIANA: ZOOLOGY Notes on Bolivian Mammals 2. Taxonomy and Distribution of Rice Rats of the Subgenus Oligoryzomys Nancy Olds and Sydney Anderson ABSTRACTS There are at least three kinds of small, long-tailed rice rats of the subgenus Oligoryzomys (genus Oryzomys) in Bolivia. We use the names Oryzomys microtis (including O. fornesi), O. chacoensis, and O. longicaudatus for these taxa. The correctness of these names is less certain and resolution of the nomenclatorial questions awaits study of specimens from outside Bolivia. The ranges of O. chacoensis and O. microtis are at low elevations and probably overlap to some degree geographically. The range of O. longicaudatus is in the highlands. No sample from one locality includes specimens of more than one species; therefore, ecological and microgeographic differences at places where any two of the three species meet are unknown. Further study may reveal other and more cryptic species within Bolivia. No one measurement or other characteristic that we have studied will unequivocally distinguish all adult specimens of any one of the three kinds from Bolivia. Geographic variation probably occurs within as well as beyond Bolivia in at least two of the three species (the most uniform seems to be O. chacoensis), but more material is needed to describe such patterns. We refrain from using subspecies names in consideration of an ignorance of both geographic patterns of variation and the status of available names. The only name in the subgenus with a Bolivian type locality, O. chaparensis, is tentatively considered a synonym of O. microtis. Existen en Bolivia por lo menos tres clases de pequefias ratas arroceras de cola larga del subgenero Oligoryzomys (genero Oryzomys). Nosotros usamos los nombres de Oryzomys mi- crotis (incluyendo O. fornesi), O. chacoensis, y O. longicaudatus para estos taxa. La exactitud de estos nombres es menos cierta y la resolucion de problemas de nomenclature necesita estudio de esp>ecimenes de fuera de Bolivia. Los rangos de distribucion de O. chacoensis y O. microtis estan en bajas elevaciones y probablemente se superponen geograficamente en algun grado. Ninguna muestra de una localidad contiene especimenes de mas de una especie. Diferencias ecologicas y microgeograficas son desconocidas en lugares donde cualquiera de las dos o tres especies se encuentran. Ulterior estudio puede revelar otras y mas cripticas especies en el interior de Bolivia. Ninguna medida u otra caracteristica que nosotros hemos estudiado separara cla- ramente todos los especimenes adultos de Bolivia. Variacion geografica ocurre probablemente dentro asi como fuera de Bolivia en al menos dos de las tres especies (la mas uniforme parece ser O. chacoensis), pero mas material es necesario para describir tales patrones de variacion. Nos abstenemos de usar nombres subespecificos en consideracion de la ignorancia de los patrones de variacion y el estado de los nombres disponibles. El unico nombre en el subgenero con una localidad tipica en Bolivia, O. chaparensis, es tentativamente considerado como O. microtis. From the Department of Mammalogy, American Mu- seum of Natural History, New York, NY 10024. OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 261 Existem no minimo tres tipos de pequenos ratos-de-arroz, de cauda longa, pertencentes ao subgenero Oligoryzomys (genero Oryzomys) na Bolivia. A estes taxa damos os nomes de Ory- zomys microtis (incluindo O. fornesi), O. chacoensis, e O. longicaudatus. A precisao destes nomes e incerta, e resolu^oes da nomenclatura aguardam estudos de especimes nao bolivianos. O. chacoensis e O. microtis ocorrem em eleva?6es baixas, e as duas esjiecies provavelmenle coocorrem em algumas areas. Oryzomys longicaudatus ocorre em areas montanhosas. Amostras de um so local nao incluem mais do que uma esF)ecie, e nao se conhecem diferen9as ecologicas ou microgeograficas em areas onde duas ou mais especies possam coocorrer. Com futuros estudos, novas especies, mais ocultas, poderao ser encontradas na Bolivia. Nenhum unico carater que estudamos pode inequivocamente separar especimes adultos dos tres tipos de Oligoryzomys na Bolivia. Varia96es geograficas provavelmente ocorrem na Bolivia, bem como em outras areas, mas mais material e necessario para poder documentar quaisquer padroes geograficos (O. chacoensis parece ser a esp>ecie mais uniforme em aparencia). Dada a falta de conhecimento, ambos dos padroes geograficos, e da disponibilidade dos nomes, nao usamos nomes de subespecies. O unico subgenero com localidade de tipo na Bolivia, O. chaparensis, e aqui lentativamente considerado como sinonimo de O. microtis. Introduction Bolivian landscapes range from less than 300 m to more than 4000 m, and habitats range from the humid lowland Amazonian tropical forests and subtropical savannahs to the high barren plains and snow-capped peaks of the Andean altiplano (fig. 1 ). Habitats may change abruptly, often within only a few kilometers. The mammals of Bolivia are also diverse, and provide excellent opportu- nities for ecological and taxonomic studies of broad scope. However, the animals are poorly known (Mares & Genoways, 1982); before satisfactory general conclusions can be reached, the Bolivian species need to be clearly delimited, both mor- phologically and geographically. Mice of the genus Oryzomys occur throughout South America and are important members of small mammal communities (Myers & Carleton, 1981; Mares et al., 1981; Alho, 1982; O'Connell, 1982;Streilein, 1982a-c; Viega-Borgeaud, 1982). We examined critically one subgenus of Oryzo- mys, Oligoryzomys, in Bolivia to determine how many species are present and where they occur. For a general description of the subgenus, see Myers and Carleton (1981, pp. 9-12). The subgenus Oligoryzomys needs revision. There are few published studies of the more than 45 named forms (Tate, 1932; Ellerman, 1941; Ca- brera, 1961; Myers & Carleton, 1981; Honacki et al., 1982). Myers and Carleton (1981) studied Oligoryzomys from Paraguay, where they recog- nized three species: Oryzomys nigripes. O. cha- coensis, and O. fornesi. They also clarified no- menclatorial questions relating to the 'name Oryzomys nigripes. Since Paraguay borders Boliv- ia, this recent study was used as our starting point. We used the same measurements and comparable analyses. We assumed that O. chacoensis and O. fornesi occurred also in southeastern Bolivia, near the Paraguayan border. If O. nigripes occurs in Bolivia, the most probable place for it is in eastern Santa Cruz, from which no specimens are now available. At least six names have been used in the liter- ature or in museum collections for Bolivian sjdcc- imens of the subgenus Oligoryzomys: Oryzomys longicaudatus and O. stolzmanni (of the high- lands); O. chaparensis (from the lowlands of Co- chabamba); O. nigripes (used for all forms); O. delicatus (used for a few specimens at middle el- evations); and 0./7ave5ce/ts (lowland). These names have been used with considerable uncertainty in the past (see summaries in Tate, 1932, and espe- cially Myers & Carleton, 1981). One cause of this problem is that the original descriptions are vague or apply equally well to more than one species of Oligoryzomys. The taxonomic confusion can be resolved by detailed study of adequate numbers of museum specimens, and the nomenclatorial confusion re- solved by comparisons with type specimens. Re- newed interest in South American mammals has resulted in more specimens, which will help in these tasks. 262 FIELDIANA: ZCX)LOGY Brasi Chile Argentina V _ - II Fig. 1 . Map of Bolivia showing the departments and the 500- and 3000-m contour lines. Be, Beni; Ch, Chuquisaca; Co, Cochabamba; LP, La Paz; Or, Onaro; Pa, Pando; Po, Potosi; SC, Santa Cniz; Ta, Tarija. OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 263 Methods We examined specimens in the collections of the American Museum of Natural History (AMNH); Academy of Natural Sciences in Phila- delphia (ANSP); California Academy of Sciences (CAL): Field Museum of Natural History (FMNH); Museum of Vertebrate Zoology, University of Cal- ifornia (MVZ); Museum of Zoology, University of Michigan (UMMZ); and United States National Museum of Natural History (USNM). Measurements were obtained as follows: exter- nal measurements are from the original labels or field notes, when available, or remeasured (fluid- preserved specimens only). The following cranial measurements were taken to the nearest 0.0 1 mm on a craniometer (see Anderson, 1 968) following Myers and Carleton (1981) and Musser (1979): ZN = depth of zygomatic notch LR = length of rostrum from tip of nasals to posterior edge of zygomatic notch GLS = greatest length of skull ZB = zygomatic breadth BB = breadth of braincase BIC = breadth of interorbital constriction LIF = length of incisive foramen LPB = length of palatal bridge LMl, LM2, LM3 = crown lengths of upF>er mo- lars WMl, WM2, WM3 = crown widths of upjjer molars MM = greatest breadth across molars (labial edges) LMX = crown length of upp>er toothrow LNP = length of nasal projection LD = length of diastema LB = length of bulla HB = height of bulla TL = total length, including tail T = tail length HP = length of hind foot, including claw E = length of ear, from notch Specimens were sorted by age, using dental cri- teria as outlined here (following Myers & Carleton, 1981): Age Class I: M' not erupted or newly erupted, M- unworn. Age Class II: M^ slightly to moderately worn, but not flat; M- slightly worn; enamel island formed by the isolation of the internal part of the mesoflexus of M-. Age Class III: M ' flat or slightly concave; enam- el island of above well isolated; M' and M- substantially worn. Age Class IV: M^ concave; enamel island oblit- erated; teeth well worn, but main cusps still discernible. Age Class V: M ' and M- flat or concave; folding pattern obliterated. Individuals in age class I were examined, mea- sured, and included in mapping geographic ranges, but were excluded from statistical treatment of character variation. Sp)ecimens examined are listed by locality and museum catalogue number in the Appendix. Lo- calities are plotted in Figure 2. Statistical analyses were done using the computer facilities of the City University of New York (CUNY) and programs from SAS (Statistical Analysis System) Institute, Inc. (1982). Taxonomy Our knowledge of geographic variation and the status of some of the names is sketchy at best, so we do not use subspecific names. The name Ory- zomys (Oligoryzomys) longicaudatus stolzmanni was first used by Hershkovitz (1940, p. 81), by inference for Bolivian populations, but we are not certain that O. stolzmanni (Thomas, 1894; type locality Huambo, 3700 ft, department of Ama- zonas, Peru) and O. longicaudatus (type locality restricted to Valparaiso, Chile) are conspecific or that the Bolivian specimens are conspecific with either. The resolution of these problems awaits further study beyond Bolivia. We noticed no diflerence between Bolivian spec- imens from Beni and Brazilian SF)ecimens from the vicinity of the typ>e locality of O. microtis (Low- er Rio Solimoes, 50 mi above mouth), and thus the subspecific name Oryzomys microtis microtis might be applied to Bolivian specimens. However, we need to know more about geographic variation in the species (see Remarks under O. microtis). Some other names that may refer to consjsecific populations and thus be relevant as possible sub- specific epithets are as follows: 1. Oryzomys destructor from lowland eastern Peru has been assigned to longicaudatus but may prove to be consp)ecific with microtis; if so, de- structor is the senior synonym. If destructor and microtis are conspecific, the northern Bohvian mice 264 HELDIANA: ZOOLOGY Fig. 2. Distribution of Bolivian Oligoryzomys: • = Oryzomys microtis; O = O. longicaudatus; X = O. chacoensis. might be known as Oryzomys destructor destruc- Bolivia on geographic and ecological grounds, al- tor, or perhaps O. destructor microtis. though no specimens are presently available. Myers 2. Oryzomys fornesi from northern Argentina and Carleton (1981) referred specimens from San might also be expected to occur in southeastern Joaquin, Beni, to O. fornesi. OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 265 3. Oryzomys mattogrossae from Brazil just to the east of Bolivia is conspecific with O. microtis and has its type locality nearest the department of Beni, where most of the Bolivian specimens have been taken; thus, for geographic reasons, it must be considered in any future subspecific determi- nations. The uncertain specific status of utiariten- sis from the same type locality as mattogrossae is noted below. If it is conspecific with mattogrossae, utiaritensis should be regarded as a synonym thereof at the subspecies level also. We concur with Myers and Carleton (1981) in the recognition of two species groups of Oligory- zomys. These are the smaller-bodied, small-toothed Oryzomys microtis. O.Jlavescens, and O. delicatus, and the larger-bodied, larger-toothed O. chacoen- sis, O. longicaudatus, and O. nigripes. In the Species Accounts section, synonymies cover only Bolivian records. Other relevant names are discussed elsewhere in the text. Results of Statistical Analyses Sex and Age Variation We examined sex and age variation in Oryzomys microtis. O. longicaudatus. O. chacoensis. and O. flavescens (from Uruguay). Males are slightly larg- er in general; the average size differences, consid- ering all characters for each species, are 2.7%, 1 .6%, 2.5%, and 2. 1%, respectively. We examined sexual dimorphism in all measurements for each species by one-way and two-way analyses of variance of sex and of sex and age for each species. Results of two-way analyses of variance of sex and age (using all specimens of tooth wear class II or greater) on specimens of O. microtis from the department of Beni, Bolivia, are presented in Table 1 . Our results are roughly comparable to those of Myers and Carleton (1981). We did not separate sexes in fur- ther statistical analyses, although we watched for unbalanced sex ratios in samples when interpret- ing results. Age variation is more difficult to assess, as no detailed study has been published. Mice of this subgenus probably continue to grow for most of their lives (see Myers &. Carleton, 1981), although the rate slows with age. We performed least-squares regression analyses (General Linear Models pro- cedures of SAS) on five variables: greatest length of skull (GLS), total length (TL), zygomatic breadth (ZB), length of the first upper molar (LMl), and length of the hind foot (HF), by species, to plot i graphically the relationship between relative age I and size. Based on study of these graphs, we de- cided to include age classes II-V in further anal- yses, as did Myers and Carleton (1981). Most spec- I imens were juveniles (age class I) or young adults (age classes II-III). Few specimens of age classes IV-V were present among the species we exam- ined. See Tables 2 and 3 for the mean adult ages of the specimens studied. Principal Components We analyzed principal components (with the Princomp procedure of SAS on a correlation ma- trix) using measurements taken on individuals of all sp)ecies (the approximate numbers of specimens are in tables 2 and 3, some specimens were ex- cluded because of missing measurements). When plotted, the first principal component tends to sep- arate the smaller species {O. microtis and O. fla- vescens) from the larger species {O. chacoensis and O. longicaudatus). The percentage of the totJil vari- ance accounted for by the first three components is 68.7%. Discriminant Analysis We performed several different discriminant analyses (using the SAS programs Discrim, Step- disc, and Candisc). A stepwise discriminant anal- ysis chose the following characters (in order of selection): tail length, breadth of interorbital con- striction, diastema length, length of incisive fo- ramina, rostral length, bullar length, length of hind foot, molar breadth, length of nasal projec- tion, length of zygomatic notch, total length, zy- gomatic breadth, length of palatal bridge, and length of maxillary toothrow. Using the SAS program Discrim, we were able to test the posterior probability of group mem- bership. In all analyses, we used as "known" groups samples of the species Oryzomys microtis from Beni, Bolivia; O. flavescens from Uruguay; O. lon- gicaudatus from the department of La Paz. Boliv- ia; and O. chacoensis from Paraguay. Plots of the individual mice on the first two canonical corre- lates showed little overlap among the four species. We assumed that the following "unknowns" be- longed to one of the four "known" groups. We submitted as "unknowns" the holotypes of O. mi- 266 HELDIANA: ZOOLOGY crotis, O. mattogrossae, and O. chaparensis; the paratype of O. chaparensis; and two specimens from Beni assigned by Myers and Carleton (1981) to O. chacoensis (see Remarks under O. microtis). The holotypes of O. microtis and O. mattogrossae were assigned to the O. microtis species sample, with a posterior probability of more than 95%. The holotype of O. chaparensis was also assigned to O. microtis (P = 0.987). The paratype of O. chaparensis was assigned to O. microtis (P = 0.985). Several other specimens that we wanted to test, including the type of O. delicatus, had missing values and could not be used in the analysis. We recognize that the validity of the taxonomic con- clusions based on these analyses depends on the correctness of the initial assumption, and that it needs further testing. Not all measurements were available for all skulls, but in each comparison the largest possible subset was used. Species Accounts Subgenus Oligoryzomys Bangs, 1 900 Diagnosis— Within the genus Oryzomys, the subgenus Oligoryzomys is distinguished by small size and delicate structure throughout, tail rela- tively long, hind foot long and slender; skull small, delicate, interorbital region narrow, outer edges of frontals squarish but unbeaded, braincase smooth and unridged, zygomatic plate narrow and with slight forward projection, molar teeth and incisors small and delicate but with cusp pattern like other Oryzomys (description adapted from Bangs, 1 900). Oryzomys of the subgenus Microryzomys are also small and delicate, but they differ from Oligory- zomys in having a more slender rostrum, shallow- er zygomatic notch, shorter and more rounded braincase, sphenofrontal foramen and squamo- soalisphenoid groove present, and karyotype with a low FN/2n ratio (Myers & Carleton, 1981, p. 12). Oryzomys chacoensis Myers and Carleton, 1981 Oryzomys chacoensis Myers and Carleton, 1 98 1 , p. 19 (typ)e locality "419 km by road NW Villa Hayes [alongside the Trans Chaco Highway], Dept. Bo- queron, Paraguay"). Diagnosis— C>ryzc>my5 chacoensis Myers and Carleton (1981, p. 20) was diagnosed as "A me- Table 1 . Nongeographic variation in Oryzomys mi- crotis. Results of analysis of variance of Bolivian O. mi- crotis. Char- Error F (inter- acter d.f. F(sex) F (age) action) ZN 98 3.75 0.44 0.52 LR 89 26.31*** 1.87 1.35 GLS 81 22.69*** 1.77 0.75 ZB 86 13.65*** 1.19 1.33 BB 92 6.57* 0.41 2.55* BIC 96 0.74 1.79 1.22 LIF 95 4.64* 1.71 0.51 LPB 93 4.32* 0.81 1.01 LMl 98 0.08 2.38* 0.76 LM2 99 1.38 5.37*** 5.57*** LM3 95 4.23* 1.93* 1.82 WMl 98 0.51 1.38 1.68 WM2 99 0.82 1.43 1.06 WM3 95 0.17 1.90* 1.24 MM 95 3.07 3.08** 1.81 LMX 94 1.53 2.38* 1.36 LNP 87 15.05*** 1.26 1.73 LD 96 10.87** 1.23 1.58 LB 92 5.45* 0.48 0.71 HB 91 1.77 0.41 0.83 TL 91 5.36* 1.38 1.21 T 91 0.02 1.65 2.34* HF 92 6.41* 0.95 0.67 E 48 0.02 1.53 0.84 * P < 0.05; ** P < 0.01; *** P < 0.001. dium-sized species of the subgenus Oligoryzomys unique in its whitish underside with hair white to the base on the chin and throat, relatively long ears having hairs on inner surface with unusually short or absent dark basal bands, small but dis- tinctive tufts of orangish hairs anterior to the ears, and karyotype with 2n = 58, FN = 74." Distribution in Bolivia— We examined spec- imens (see Appendix) from the departments of Cochabamba, Santa Cruz, and Tarija (fig. 2). Myers and Carleton (1981) reported two specimens from Beni. These specimens have been restudied and are here reassigned to Oryzomys microtis (see Re- marks under that species). Habitat of O. chacoen- sis is grassland and thomscrub. The highest known elevation is 640 m (Rio Lipeo). General Description and Comparisons— Cranial and external measurements are listed in Table 2. The dorsal pelage is rufous and heavily lined with black hairs. The venter is white and sharply distinguished from sides, which are clearer than dorsum. Frequently a thin orange line sep- arates the sides from the belly. The cheeks are slightly paler than the rest of the face. The tail is dark gray, weakly bicolored, and long relative to the body. The hind feet are whitish above. Juve- OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 161 Table 2. Mean, standard deviation, and range (in mm) for Oryzomys chacoensis and O. longicaudatus. O. chacoensis O. longicaudatus Bolivia Paraguay Argentina Bolivia Character (N = 15) (N = 16) (N = 5) (N = 34) ZN 1.15 ± 0.28 1.30 ± 0.20 1.15 ± 0.12 1.02 ± 0.20 (0.88-1.59) (0.90-1.61) (0.94-1.24) (0.57-1.36) LR 7.27 ± 0.45 7.39 ± 0.57 7.67 ± 0.34 7.14 ± 0.72 (6.62-7.94) (6.57-8.86) (7.25-8.07) (4.49-8.15) GLS 24.90 ± 0.82 24.52 ± 1.09 26.12 ± 0.32 25.01 ± 1.39 (23.80-26.69) (22.52-27.12) (25.79-26.43) (21.06-27.29) ZB 13.10 ± 0.59 12.76 ± 0.62 13.64 ± 0.09 13.17 ± 0.65 (11.94-13.98) (11.91-14.27) (13.54-13.71) (11.55-13.98) BB 11.33 ± 0.41 11.16-0.34 11.39 ± 0.06 11.25 ± 0.37 . (10.42-11.87) (10.59-11.59) (11.35-11.45) (10.58-11.96) BIC 3.89 ± 0.20 3.76 ± 0.10 3.86 ±0.13 3.63 ± 0.22 (3.52-4.27) (3.64-3.95) (3.76-4.04) (3.35-4.17) LIF 4.62 ± 0.26 4.71 ± 0.39 5.07 ± 0.29 4.68 ± 0.27 (4.25-5.20) (4.17-5.56) (4.64-5.31) (4.23-5.26) LPS 4.14 ± 0.22 3.98 ± 0.26 4.22 ± 0.31 3.97 ± 0.21 (3.65-4.54) (3.50-4.65) (3.75-4.52) (3.49-4.41) LMl 1.68 ± 0.10 1.63 ± 0.07 1.74 ± 0.5 1.59 ± 0.09 (1.47-1.84) (1.54-1.79) (1.70-1.83) (1.42-1.81) LM2 1.09 ± 0.06 1.10 ± 0.07 1.14 ± 0.07 1.05 ± 0.04 (0.96-1.18) (1.00-1.28) (1.08-1.26) (0.97-1.15) LM3 0.84 ± 0.05 0.80 ± 0.04 0.84 ± 0.04 0.81 ± 0.05 (0.74-0.92) (0.73-0.88) (0.78-0.88) (0.67-0.92) WMl 1.07 ± 0.07 1.03 ± 0.03 1.11 ± 0.05 1.04 ± 0.06 (0.93-1.24) (0.96-1.08) (1.05-1.17) (0.91-1.15) WM2 1.02 ± 0.07 1.00 ± 0.04 1.04 ± 0.06 0.99 ± 0.06 (0.86-1.14) (0.93-1.06) (0.97-1.11) (0.87-1.09) WM3 0.87 ± 0.05 0.85 ± 0.03 0.90 ± 0.05 0.83 ± 0.06 (0.78-0.97) (0.78-0.92) (0.84-0.94) (0.67-0.95) MM 4.66 ± 0.20 4.57 ± 0.17 4.70 ± 0.13 4.61 ± 0.27 (4.32-4.99) (4.34-5.09) (4.58-4.87) (4.03-5.08) LMX 3.60 ±0.12 3.50 ± 0.13 3.71 ± 0.15 3.46 ± 0.14 (3.29-3.78) (3.23-3.68) (3.61-3.98) (3.20-3.77) LNP 1.43 ± 0.24 1.59 ± 0.17 1.49 ± 0.17 1.40 ± 0.25 (1.04-1.77) (1.35-2.07) ~ (1.29-1.64) (0.99-2.04) LD 5.66 ± 0.34 5.63 ± 0.44 5.89 ± 0.24 6.04 ± 0.47 (5.17-6.33) (5.02-6.44) (5.63-6.18) (4.96-7.07) LB 3.54 ± 0.14 3.60 ± 0.14 3.75 ± 0.03 3.47 ± 0.20 (3.28-3.81) (3.28-3.82) (3.72-3.78) (3.07-3.84) HB 2.75 ± 0.18 2.73 ± 0.15 2.72 ± 0.06 2.76 ± 0.32 (2.43-3.04) (2.40-2.96) (2.66-2.76) (2.23-3.32) TL 227.93 ± 12.45 219.50 ± 12.29 229.00 ± 8.34 217.68 ± 16.88 (204-252) (194-241) (215-237) (172-252) T 134.27 ± 10.26 126.06 ± 8.46 138.00 ± 5.70 124.15 ± 12.18 (112-150) (112-139) (130-145) (87-145) HF 25.33 ± 1.33 24.38 ± 1.54 23.40 ± 2.30 25.13 ± 1.20 (23-28) (23-29) (21-26) (22-28) E 16.70 ± 1.33 15.07 ± 1.69 18.40 ± 1.14 15.03 ± 1.72 (14-18) (12-17) (17-20) (12-18) AGE 2.65 ± 0.82 2.31 ± 0.68 2.55 ± 0.87 3.04 ± 0.88 (2-4) (2-4) (2-4) (2-5) 268 HELDIANA: ZOOLOGY niles are grayer, as are all young Oligoryzomys in Bolivia, but this is especially noticeable on the venter, which has mixed gray and white hairs. In- cisive foramina extend posteriorly to the anterior edge of M' or slightly beyond. Alisphenoid strut (a strut of the alisphenoid bone that covers the lateral part of the alisphenoid canal, see Musser, 1 982, p. 29) is generally absent (table 4). The sides of the interorbital constriction are divergent pos- teriorly (less parallel-sided than in O. longicau- datus) (fig. 3). The following quotes are from Myers and Carleton (1981, first and second quotes, p. 2 1 , third quote, p. 24): Oryzomys chacoensis differs from O. cha- parensis Osgood (1916) primarily in color pattern: the type of chaparensis is much darker and less hispid dorsally, yellowish ventrally, lacks the orange tufts anterior to the ears, and has a grayish throat. The distal portions of the nasals of the holotype flare laterally to an extent not seen in chacoensis. amined the two darker ones. When skulls are com- pared, the two darker mice are among the older and larger individuals in the series. They seem to have relatively broader braincases than most, but there is no character shared by these two that is not also seen in one or more of the others. In a discriminant analysis (SAS program Discrim) the posterior probability of membership (in the four reference species) allies these two mice with O. chacoensis (ansp 18187, P = 0.997, and ansp 18188, P = 0.954 with chacoensis). We have as- signed these mice to O. chacoensis. Since Rio Li- pto is in the area where these two species meet, further study there should reveal whether sym- patry exists and if species differences remain dis- tinct. Specimens from Argentina (listed in the Ap- pendix) are similar in coloration to both Para- guayan and Bolivian specimens, but are woolier. This sample extends the known range of O. cha- coensis into the department of Jujuy, northwestern Argentina. Oryzomys chacoensis can be distinguished from fornesi, with which it occurs sympat- rically, by its larger size (maxillary toothrow usually > 3.3mm, ears usually > 15mm), characteristic karyotype, lack of preputial glands, and in most specimens by its lack of buff" on the belly. . . . the hind feet of chacoensis are relatively short compared to those of the more terres- trial fornesi. In reference to the last point, however, our cal- culations of length of hind foot relative to length of head and body (using data from tables 2 and 3) are about 26% for both O. chacoensis and O. mi- crotis (including fornesi). Comparison of speci- mens also reveals no noticeable difference. Remarks— This species is clearly distinct from O. microtis (here including O. fornesi). In most morphometric characters, O. chacoensis grossly resembles O. longicaudatus from Bolivia and O. nigripes from Brazil. Our specimens from Santa Cruz show some variation in coloration: two specimens (amnh 247772-247773) are pale. The specimens from Tarija resemble the Paraguayan samples more closely than do the specimens from Santa Cruz. A series of 13 sp)ecimens from Rio Lipeo in- cludes two with darker pelage that resemble O. longicaudatus. We measured these mice and ex- Oryzomys longicaudatus (Bennett, 1832) Mus longicaudatus Bennett, 1832, p. 2 (type locality "In trees in Chile," restricted to Valparaiso by Ca- brera, 1961, p. 391). Oryzomys longicaudatus: Thomas, 1898, p. 3 (Aguai- renda Mission, San Francisco, perhaps not O. lon- gicaudatus); Thomas, 1926, p. 194 (Tupiza). Oryzomys Stolzmanni: Thomas, 1902, p. 130 (Cha- ruplaya, Choro); Neveu-Lemaire and Grandidier, 1 9 1 1 , p. 9 (Charuplaya, Choro). Oryzomys ^X>;flavescens groMTp: Thomas, 1925, p. 578 (Carapari, perhaps not O. longicaudatus). Oryzomys stolzmanni stolzmanni: Sanborn, 1950, p. 2 (Rio Aceramarca, Cocapunco, Nequejahuira, Okara, Pitiguaya, Pongo). Diagnosis— Bolivian specimens referred to this species differ from other Bolivian Oligoryzomys in that they are larger (especially in the size of the teeth) than O. microtis and lack the buffy wash on the venter; are grayer-bellied, darker, and less griz- zled dorsally than O. chacoensis; and occur at gen- erally higher elevations (at least in northern Bo- livia). Distribution in Bolivia— O. longicaudatus is found in the valleys and mountains of the Andes from at least middle elevations (1200 m, Entre Rios) up to 3720 m (Poopo). Specimens from near Camiri and Cuyambuyo in southern Bolivia are from elevations of 780 to 1000 m. Specimens (see Appendix) have been reported or examined from OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 269 Fig. 3. Dorsal and ventral views of skull of (left), Oryzomys longicaudatus stolzmanni; (middle), O. chacoensis, and (right), O. microtis microtis. Specimens in the American Museum of Natural History. Scale at lower right represents 10 mm. the departments of Chuquisaca, Cochabamba, La Paz, Oruro, Polosi, Santa Cruz, and Tarija (see map, fig. 2). General Description and Comparisons— Cranial and external measurements are presented in Table 2. The belly is gray, rarely washed with buff; upper parts are a dark brown lined with black hairs, and often the sides are more rufous. A thin orange lateral line may be present. Nose and face are dark with paler cheeks. Often there are pale spots just behind the ears. Hind feet are pale above. The tail is brown above, weakly to strongly bi- colored, and relatively long compared to that of any other Bolivian Oligoryzomys. Incisive foram- ina generally extend to the anterior edge of M' or slightly behind. Alisphenoid strut is generally ab- sent (table 4). The interorbital area is slightly more constricted on the average than that of O. cha- 270 HELDIANA: ZOOLOGY coensis, and has relatively parallel sides (see fig. 3). Remarks— Oryzomys longicaudatus may be a composite of more than one species. Oryzomys destructor and O. stolzmanni were considered sub- species of 6>. longicaudatus by Cabrera ( 1 96 1 ) and full species by Soukup (1961). Gardner and Patton (1976) found four karyotypic variants in speci- mens they assigned to O. longicaudatus, and com- mented that these may represent four separate species. One of their karyotypic variants, no. 2, is represented by 1 7 specimens from Balta, depart- ment of Loreto (now Acayali), Peru, at an eleva- tion of about 300 m. Myers and Carleton (1981, p. 26) mentioned that the karyotype of these spec- imens as reported by Gardner and Patton (1976) agreed better with results they obtained for O.for- nesi (= O. microtis) than for either O. chacoensis or O. longicaudatus. We have examined these specimens and judge that they are clearly different from O. longicaudatus and belong with either O. microtis or O. destructor (Balta is near the type locality of O. destructor). The inclusion of karyo- type no. 3 under O. longicaudatus, represented by one female from Venezuela, also needs verifica- tion. We have five specimens from Caracato, de- partment of La Paz (amnh 248977-248981), that are quite different from other Bolivian Oryzomys longicaudatus. The pelage closely resembles that of O. nigripes from Paraguay in color and texture; it is paler and more obviously lined with black hairs dorsally than the pelage of typical O. lon- gicaudatus. They are also slightly larger bodied, but four of the five are old individuals (age classes IV-V) and this may account for the larger size. Because of the small sample size and the single locality, we chose not to distinguish this popula- tion taxonomically. It is possible that Andean populations may oc- cupy areas that are rather widely scattered and physically isolated in different valleys or side can- yons. If so, there may be considerable geographic variation. More specimens are needed to detect and describe such patterns. The sjjecimens from near Cuyambuyo are at comparatively low elevations (980 to 1000 m) for O. longicaudatus, although the species occurs at progressively lower elevations farther south (Os- good, 1916). The two localities near Cuyambuyo are only about 60 km from Rio Lipeo, where O. chacoensis occurs. The SAS Discrim program analysis allied the largest adult specimen from near Cuyambuyo (ummz 155891) with O. longicauda- tus (F = 0.956). There are problems in distinguish- ing O. longicaudatus and O. chacoensis solely by cranial morphology in all areas. Oryzomys microtis Allen, 1916 Oryzomys (Oligoryzomys) microtis Allen, 1 9 1 6, p. 525 (type locality Lower Rio Solimoes, 50 mi above mouth [80 km from its confluence with Rio Negro, Amazonas, Brazil]). Oryzomys chaparensis Osgood, 1 9 1 6, p. 205 (holotype and paratype, type locality Todos Santos, on Rio Chapare, Department of Cochabamba, Bolivia); Gyldenstolpe, 1932, p. 25 (Todos Santos); Myers and Carleton, 1981, p. 38 (Todos Santos); all in reference to the same specimens. Oryzomys fornesi Massoia, 1973, p. 22 (type locality Naineck, Dept. Rio Pilcomayo, province of For- mosa, Argentina); as used by Myers and Carleton, 1981, p. 25 for five specimens fVom San Joaquin, Beni, Bolivia. Diagnosis— Allen described Oryzomys microtis as being readily distinguished from other Olig- oryzomys by its pale coloration, relatively small ears, and tail less than half total length. However, the tail of the holotype is more than half of the total length, a discrepancy noted by Goodwin (1953). Massoia (1973) diagnosed O. fornesi, here considered a junior synonym of O. microtis, as the smallest species of the subgenus in Argentina, hav- ing the ears covered with short ochraceous hairs, the pterygoids short (shorter than the molar se- ries), the interorbital constriction narrow, and the incisive foramina not extending to the molar se- ries. In Bolivia, it can be distinguished by its small body size, toothrow generally less than 3.3, gray- buffy color of the belly, and relatively short tail in proportion to body. Distribution in Bolivia— Specimens (see Ap- pendix) from the lowlands of Bolivia, in the de- partments of Beni, Cochabamba, Pando, La Paz, and Santa Cruz have been examined. Habitat in- cludes marshes and wet forests up to an elevation of 1 800 m (Guanay, see fig. 2). General Description and Comparisons — Smallest of the Bolivian Oligoryzomys, with small teeth, toothrow generally less than 3.3 (averaging 3.10). Massoia ( 1 973) gave general measurements useful in distinguishing this species from other Ar- gentine species of Oryzomys (Oligoryzomys) as fol- lows: length of hind foot generally less than 24, length of ear generally less than 13, breadth of braincase less than 10.8, and length of incisive foramina generally less than 4.5. All of his mea- OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 271 Table 3. Mean, standard deviation, and range (in mm) for Oryzomys microtis and O. flavescens. O. microtis O. flavescens Bolivia Brazil Peru Uruguay Character (N = 67) (N = 20) (N = 26) (N = 20) ZN 1.01 ± 0.17 0.98 ±0.18 0.84 ±0.13 1.07 ± 0.15 (0.53-1.46) (0.78-1.48) (0.61-1.18) (0.79-1.32) LR 6.82 ± 0.51 6.82 ± 0.42 6.60 ± 0.41 6.75 ± 0.43 (5.17-8.16) (6.16-7.46) (5.96-7.36) (5.89-7.39) GLS 23.54 ± 1.06 23.79 ± 0.88 23.27 ± 0.82 23.41 ± 0.94 (21.24-25.78) (22.47-25.04) (21.75-24.53) (21.66-24.81) ZB 12.28 ± 0.51 12.40 ± 0.44 11.86 ± 0.53 12.05 ± 0.49 (11.42-13.73) (11.59-13.32) (11.01-12.93) (11.25-12.76) BB 10.78 ± 0.36 10.71 ± 0.23 10.47 ± 0.32 10.70 ± 0.26 • (9.38-11.78) (10.35-11.06) (9.49-10.87) (10.29-11.22) BIC 3.76 ± 0.17 3.72 ± 0.14 3.63 ±0.11 3.35 ±0.17 (3.38-4.19) (3.44-3.93) (3.45-3.94) (2.95-3.77) LIF 3.96 ± 0.27 4.01 ± 0.22 4.00 ±0.18 4.59 ± 0.35 (3.38-4.53) (3.65-4.37) (3.73-4.33) (3.54-5.20) LPB 4.04 ± 0.32 4.06 ± 0.22 3.98 ± 0.26 3.77 ± 0.19 (3.14-4.76) (3.64-4.38) (3.42-4.51) (3.51-4.06) LMl 1.46 ± 0.10 1.43 ± 0.07 1.51 ± 0.06 1.47 ± 0.06 (1.22-1.64) (1.32-1.56) (1.39-1.66) (1.37-1.60) LM2 0.95 ± 0.06 0.95 ± 0.05 0.93 ± 0.08 1.01 ± 0.06 (0.81-1.09) (0.84-1.06) (0.67-1.04) (0.83-1.09) LM3 0.70 ± 0.07 0.68 ± 0.05 0.66 ± 0.05 0.73 ± 0.04 (0.54-0.85) (0.59-0.78) (0.55-0.77) (0.66-0.80) WMl 0.94 ± 0.06 0.93 ± 0.06 0.99 ± 0.04 0.97 ± 0.04 (0.78-1.08) (0.80-1.04) (0.91-1.10) (0.93-1.07) WM2 0.88 ± 0.06 0.89 ± 0.06 0.93 ± 0.04 0.93 ± 0.05 (0.70-1.01) (0.78-0.99) (0.82-1.00) (0.86-1.03) WM3 0.73 ± 0.06 0.74 ± 0.06 0.80 ± 0.05 0.78 ± 0.04 (0.55-0.85) (0.58-0.85) (0.71-0.93) (0.70-0.85) MM 4.33 ± 0.21 4.39 ± 0.23 4.33 ± 0.18 4.26 ±0.15 (3.83-4.93) (3.99-4.81) (4.07-4.70) (3.98-4.50) LMX 3.10 ± 0.17 3.07 ± 0.16 3.14 ± 0.13 3.22 ±0.12 (2.64-3.43) (2.75-3.34) (2.93-3.39) (2.96-3.47) LNP 1.32 ± 0.22 1.25 ± 0.32 1.20 ± 0.20 1.33 ± 0.14 (0.91-1.74) (0.75-1.70) ■ (0.81-1.62) (1.16-1.68) LD 5.75 ± 0.41 5.78 ± 0.36 5.32 ± 0.40 5.35 ± 0.35 (4.82-6.62) (5.13-6.51) (4.49-6.01) (4.67-5.90) LB 3.29 ± 0.14 3.18 ± 0.19 3.01 ± 0.12 3.30 ±0.15 (2.94-3.57) (2.90-3.63) (2.75-3.27) (2.91-3.53) HB 2.46 ± 0.20 2.10 ± 0.23 2.22 ±0.12 2.64 ± 0.23 (1.96-2.87) (1.72-2.45) (1.96-2.38) (2.02-2.94) TL 185.88 ± 11.12 185.00 ± 10.60 179.77 ± 9.16 198.56 ± 17.66 (165-214) (164-202) (160-196) (160-223) T 101.00 ± 6.67 95.44 ± 7.88 101.69 ± 8.13 110.37 ± 11.05 (87-116) (75-105) (77-115) (85-127) HF 22.81 ± 1.21 21.68 ± 1.34 21.37 ± 1.60 25.21 ± 1.85 (19-25) (20-25) (18-25) (20-28) E 14.49 ± 1.04 (12-16) 13.97 ± 1.05 (12-16) AGE 2.57 ± 0.73 2.35 ± 0.59 2.97 ± 0.88 2.18 ± 0.35 (2-5) (2-4) (2-5) (2-3) 272 FIELDIANA: ZOOLOGY surements fall within the extremes of our values for O. microtis, except for measurements of the toothrow (LMX), in which his are on the upper end of our range. Values of cranial and external measurements are listed in Table 3. The venter is white mixed with gray to gray-bufFy and clear huf- fy. The dorsum is rufous brown, lined with black hairs, and has paler sides. Often there is no clear demarcation between sides and belly. There are no white spots behind ears. Juveniles are similar to adults, but show a grayer belly. The tail is dark above and weakly bicolored. Occasionally orange- tipped hairs lie anterior to the ear. Hind feet are white above. Incisive foramina generally extend posteriorly to the anterior edge of M' and not beyond. In comparison with adults of O. cha- coensis and O. longicaudatus, O. microtis has a shorter skull, longer braincase, shorter incisive fo- ramina, a relatively broader interorbital constric- tion with divergent sides, and smaller, narrower teeth; also, an alisphenoid strut is more often pres- ent (table 4). Remarks— This species is similar to Oryzomys flavescens of Uruguay and Argentina (see Myers & Carleton, 1981). Specimens of both species were reported from the same locality, Capitan Solari, in the province of Chaco, northern Argentina, by Contreras and Berry (1983). Measurements of 6>. flavescens from Uruguay are listed in Table 3 (see also Langguth, 1 963). In comparison with O. mi- crotis, specimens of O. flavescens have slightly larger teeth, larger bullae, longer incisive forami- na, and are generally larger. Oryzomys flavescens needs to be more clearly defined. Massoia and Fomes ( 1 967) once synonymized O. flavescens with O. nigripes, but according to the later analyses of Massoia (1973) and Myers and Carleton (1981) these taxa are not consjiecific. Uruguayan speci- mens oi O. flavescens are clearly different from O. nigripes from eastern Paraguay. Future studies may reveal additional relationships, including the pos- sibility that O. microtis and O.fiilvescens of Cen- tral America and northern South America are con- specific (Handley, 1976, referred Venezuelan specimens to O. fiilvescens). The name Oryzomys microtis flyrnesi might be used to reflect the conspecific status oiflyrnesi and microtis. This subspecific name would apply to at least the five specimens identified by Massoia (1973), pending more detailed studies that would test whether geographic differences warrant con- tinued subspecific recognition. Until the relationships of the Andean ""longi- caudatus-group" are clarified, it is also possible Table 4. Presence or absence of alisphenoid strut in five species of Oryzomys (Oligoryzomys) expressed as a percentage; number of specimens in parentheses. Alisphenoid strut Species Present Absent O. chacoensis 2.4(1) 97.6(41) O. longicaudatus 5.6 (2) 94.4 (34) O. microtis 35.8 (54) 64.2 (97) O. flavescens 3.2(1) 96.8 (30) • O. nigripes 8.3 (3) 91.7(33) that one or more of the earlier names currently assigned to this group actually belong with Ory- zomys microtis, and have priority. For example: O. stolzmanni (named in 1894, type locality listed under synonymy) and O. destructor (named in 1 844, type locality: "the house mouse of the 'Plan- tagen at the border of the forest," eastern Peru, above 6000 ft, according to Tate, 1932, p. 9; re- stricted by Cabrera, 1961, to the haciendas along the Rio Chinchao in the department of Huanuco, between 900 and 1000 m) have been associated with O. longicaudatus. These type localities are at low elevations, which suggests the possible affinity of these forms with microtis rather than with lon- gicaudatus. We have examined a series from low elevation in the department of Pasco, Peru, that is clearly O. microtis. This locality is near the type locality of destructor (Cole, 1984). If future study of the holotype of destructor proves it to be conspecific with O. microtis, the name of the species should be O. destructor instead of O. microtis. We have examined the type specimens of Ory- zomys microtis (amnh 37091, type locality Lower Rio Solimoes, 50 mi above mouth, Brazil), O. mattogrossae (amnh 37542, type locality Utiarity, Mato Grosso, Brazil), O. utiaritensis (amnh 37541, type locality Utiarity, Rio Papagaia, Mato Grosso, Brazil), and O. delicatus (amnh 7317/5925, type locality Trinidad, West Indies). We judge that the small lowland Bolivian specimens are O. microtis and also represent the same species as specimens referred by Myers and Carleton to O. fornesi (O. microtis has priority). The Bolivian specimens dif- fer slightly from Massoia's description of O. for- nesi in having a shorter average toothrow, other- wise in being slightly larger. If O. fornesi and O. microtis are conspecific, it is possible that this dif- ference in measurements reflects his small sample size (N = 5), or that there may be significant geo- graphic variation between Argentine and Bolivian populations. We further treat O. mattogrossae as a synonym of O. microtis (O. mattogrossae is not OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 111 included in the synonymy above because the name has never been used in print for a Bolivian spec- imen). As mentioned by Myers and Carleton ( 1 98 1 ), O. utiaritensis may belong with O. nigripes. More specimens from Mato Grosso are needed to establish the characteristics of the species there. Oryzomys microtis was considered a synonym of O. delicatus by Alho ( 1 982) and Pine ( 1 973); how- ever, because of the slightly damaged condition of the skin and skull of the holotype of O. delicatus and the remoteness of its type locality (on the is- land of Trinidad, Allen & Chapman, 1 897), we defer any decision on this question. We have reexamined the two specimens from Beni that Myers and Carleton (1981) referred to Oryzomys chacoensis (usnm 39 1 297, 46074 1 ) and have assigned them to O. microtis. These speci- mens are young adults (age class II), have external measurements that fit well with O. microtis and that are rather small for O. chacoensis (TL 200, 200; T 111, 110; HF 19, 24, respectively), and have toothrows measuring 3.34 and 3.40, respec- tively. The toothrows fall on the upper end of the range for O. microtis and the lower end of the range for O. chacoensis (tables 2-3). We then compared their measurements through a discriminant anal- ysis with the samples ofO.flavescens, O. microtis, O. chacoensis. and O. longicaudatus. The posterior probability of membership for both was greatest for O. microtis (usnm 391297, P = 0.937; usnm 460741,/' = 0.953). We refer the holotype (fmnh 21330) and the one paratype (amnh 40787) o^ Oryzomys chapa- rensis to Oryzomys microtis. The holotype is the largest and one of the oldest individuals we have identified as O. microtis. Its greatest skull