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Mammals of Bolivia, taxonomy and distribution.

Anderson, Sydney

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MAMMALS OF BOLIVIA, TAXONOMY AND DISTRIBUTION SYDNEY ANDERSON BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY NUMBER 231 NEW YORK: 1997 Recent issues of the Bulletin may be purchased from the Museum. Lists of back issues of the Bulletin, Novitates, and Anthropological Papers published during the last five years are available free of charge. Address orders to: American Museum of Natural History Library, Department D, Central Park West at 79th St., New York, New York 10024. TEL: (212) 769-5545. FAX: (212) 769-5009. E-MAIL: [email protected] . MAMMALS OF BOLIVIA, TAXONOMY AND DISTRIBUTION SYDNEY ANDERSON Curator Emeritus, Department of Mammalogy American Museum of Natural History BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 231, 652 pages, 785 figures, 21 tables Issued April 29, 1997 Price: $52.00 a copy Copyright C) American Museum of Natural History 1997 ISSN 0003-0090 CONTENTS Abstract ....... .............................................................. 3 Introduction .............................. . . ..................... 3 Acknowledgments ....................... 5 Objectives and Organization ............. ...................................... 6 Taxonomic Concepts. 6 List of Taxa ... ... ................................................ 8 Keys to the Mammals of Bolivia ............................................... 21 Methods and Materials .............. 61 Faunal Analysis and Biogeography ............ 64 Geographical Position and Topography .............. 64 Climate .............. 66 Vegetation ........... 66 Faunal Analysis ........... 72 Management and Conservation ............ 79 Abbreviations and Acronyms ............ 80 Mammalogical Gazetteer of Bolivia .............. 82 History of Collecting ............ 120 Chronology ........... 120 Persons and Institutions ..................................................... 126 Systematic Accounts .............. 138 Hypothetical List ........... 516 References ........... 521 Tables ........... 569 Index to Scientific Names of Mammals ........... 623 2 ANDERSON: MAMMALS OF BOLIVIA ABSTRACT Studies of published records and of about 36,900 specimens of native mammals from Boliv- ia reveal that at least 327 species occur there. Probably more than 20 other species, either new to science or new to Bolivia, remain to be dis- covered. In addition to these species names, some 44 additional subspecies names are used. Most of the subspecies names reflect taxonomic history more than detailed knowledge of geographic van- ation. In this report, the taxonomic status of each taxon within Bolivia is noted. Scientific names that have been used for Bolivian specimens are given for each species and subspecies, and all known publications that have specifically referred to Bolivian specimens are cited, along with a few other works selected for special reasons. Speci- mens are listed, and localities are mapped, includ- ing the prediction of the probable distribution of each species within Bolivia. Illustrated keys are based primarily on external, cranial, and dental characters and include 10 domestic and intro- duced species in addition to native species. The number of species present in most local areas ranges from about 50 to 180. Analysis indicates four major faunal areas: lowland tropics, lowland temperate zone (including the chacoan area), for- ested yungas, and highlands (altiplano). The high- land and lowland faunas are almost mutually ex- clusive; the break between temperate and tropical is indistinct. INTRODUCTION This work is dedicated to the hypothesis- testers of this world. Everything concluded here is subject to further testing. I expect much of this information to be substantiated, but not everything. Bolivia is a country of amazing biological diversity, with faunal and floral richness aris- ing from its geographic location in the center of South America and its great topographic range. Its position at the juncture of the trop- ical and temperate zones has allowed the Bo- livian area to draw from and contribute to the faunal complements of those zones over mil- lions of years of evolution as well as over the shorter time span of ecological changes in the last 10,000 years. The task of disen- tangling the effects of these long- and short- term processes makes biogeography both dif- ficult and fascinating. The uplifting of the high Andes, resulting from the collision of two. tectonic plates in the earth's crust in the last 10 to 15 million years, is a relatively recent event in the long sweep of geological time (Roeder, 1988). Some of the changes and their possible ef- fects on the eventual development of present- day savannahs in Beni were presented by Hanagarth (1993). Most of what is known of fossil mammals in Bolivia relates to pre-Pleistocene time and an earlier fauna. There are later deposits near Tarija (Ameghino, 1902; Boule and Thev- enin, 1920; Hoffstetter, 1963). For other in- formation see MacFadden et al. (1985, 1994), Hoffstetter (1986), Marshall (1987), Marshall and Muizon (1988), and Janis (1993). Although the details of changes in local climates and vegetation occurring over lon- ger periods are poorly known, we can safely conclude that the relative extents of different habitats have fluctuated. At a given time, tropical forests might be continuous and widespread and enclose smaller and more dispersed areas of grassland, savanna, or oth- er habitats. At other times, tropical forests might be reduced to smaller dispersed patch- es (see Patton et al., 1990, for a discussion of vicariant versus gradient models of faunal evolution in the Andean region). The organization of this faunal report is similar to that of an earlier report on the mammals of Chihuahua, Mexico (Anderson, 1972). The theoretical and practical advan- tages and disadvantages of faunal studies that I discussed there apply with equal or greater force to the present situation. Bolivia and Chihuahua are similar in sev- eral respects. The southwestern part of each is of greater elevation than other parts. A coastline lies to the west of both, but is sep- arated by a relatively narrow belt in another political division with a largely different hab- itat. However, the size of Chihuahua is only about 23% the size of Bolivia. In 1972, 6600 Chihuahuan specimens were available for 3 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY study. This quantity is equal only to about 18% the 36,900 available for study in Bolivia in 1994. And yet, the stage of development of taxonomic knowledge for Bolivia was not as advanced in 1994 as it was for Chihuahua in 1972. In some ways the taxonomy for Bo- livian mammals is comparable in develop- ment to that of western North America at about the beginning of the 20th century, be- fore the publication of revisions for such widespread, common, and diverse genera as Microtus (Bailey, 1900), Perognathus (Os- good, 1900), and Peromyscus (Osgood, 1909). See also discussions of taxonomic comparability of the continents by Mares (in Mares and Genoways, 1982: 11) and by Pine (op. cit.: 27). Most of the abundant and di- verse South American genera, such as Ako- don, Proechimys, Oligoryzomys, and Oryzo- mys, as well as rarer genera, such as Tho- masomys and Rhipidomys, need critical and comprehensive revisions at the level of spe- cies and subspecies. These revisions can pro- vide basic data needed for analyses of rela- tionships at intermediate and higher taxo- nomic levels. I have found time to deal with only a few of these problems and have done what I could to encourage and expedite the work of others on these taxa, although it is frustrating, in a way, to leave so many of them unresolved. This is very much a work in progress, not the final word on taxonomy and distribution of Bolivian mammals. During the years since 1980, when I be- gan intensive work on this manuscript, tax- onomic advances changed our understanding of Bolivian taxonomy sufficiently to cast doubt on some identifications made in earlier years. I have not been able to restudy all of these specimens, or I was uncertain of their identities the first time I studied them, and so have listed them as unknown species un- der the appropriate genus. Some unidentified specimens are known to exist but I have not examined them; others are incomplete or oth- erwise difficult or impossible to identify as to species. I hope that listing these 1293 specimens (about 3.5%) will bring unsolved problems to the attention of future students and expedite their finding relevant material. I should note, also, that my research notes (on some 10,000 5 by 8-in. cards, with mu- seum catalog numbers and other data) are available in the archives of the Department of Mammalogy and may be consulted there. The mammalian fauna of South America, in general, has been studied less intensively than that of North America, as noted above. The major faunal summary for South Amer- ican mammals is still the annotated checklist by Cabrera (1958, 1961a). Regional or na- tional accounts are few. Books on Suriname (Husson, 1978) and Chile (Osgood, 1943) are the only detailed recent accounts of an entire national mammalian fauna. A sum- mary of the literature for Peru published by Soukup (1960-1961) and a booklet for Ar- gentina by Olrog and Lucero (1981) in "field guide" format, with general range maps and illustrations but without supporting docu- mentation, are examples of what has been available. Recently published popular ac- counts on the living mammals of the nearby provinces of Salta and Tucumain in Argentina (Mares et al., 1989; Barquez et al., 1991) and on the mammals of tropical and southern regions (Emmons and Feer, 1990; Redford and Eisenberg, 1992) have contributed to knowledge of South American mammals. At the request of Bolivian colleagues, a list of scientific names of Bolivian mammals was published in Bolivia to provide a work- ing preliminary summary (Anderson, 1985b), as well as lists of selected large mammals in parks and reserves (Cardozo et al., 1988). An annotated list, with informa- tion on general geographic distribution with- in Bolivia for each species and with illus- trated keys to Bolivian mammals was pre- pared in 1987 for publication in Spanish and was published later (Anderson, 1993). These lists provided a comparative basis for a series of short reports of additional distributional records (Yensen and Tarifa, 1993; Ibaniiez et al.,1994; Salazar et al., 1994; Yensen et al., 1994) and a more comprehensive discussion of biogeography (Salazar et al., in press). A number of shorter technical reports on taxonomy and distribution have been pro- duced in the course of the larger faunal study. These are cited in the appropriate spe- cies accounts. Since collecting by field parties from the American Museum of Natural History re- sumed in 1963 (after earlier work in 1915, 1926, and 1929), we have made specimens 4 NO. 231 ANDERSON: MAMMALS OF BOLIVIA and information from our work available to qualified scholars. These materials have been used in various studies (e.g., Hershkovitz, 1966; Davis, 1973, 1976; Eger, 1977). During the years of preparation of this re- port, draft copies of various sections or spe- cially extracted summaries of data (e.g., on the mammals of Chuquisaca, Tarija, Pando, Beni, the Cochabamba Valley, the yungas of La Paz, the altiplano, Rodentia, Chiroptera, certain genera, etc.) have been provided to Bolivian institutions and to investigators there and elsewhere. These institutions in- clude Coleccion Boliviana de Fauna (formed by combining the collections of the Instituto de Ecologia and the Museo Nacional de His- toria Natural), Centro de Datos para la Con- servacion, Instituto Boliviano de Biologia de Altura (all in La Paz), and Centro Nacional de Enfermidades Tropicales and Parque Zoologico de Fauna Sud Americana (in Santa Cruz). Information on Bolivian bats was published (Anderson et al., 1982) and later brought up to date by Anderson and Webster (1983) and by Anderson (1991). In some cases, catalog numbers are also given in lists or in text where changes of identification have occurred or specimens from one presumed series have been subse- quently placed in more than one taxon. Fu- ture workers may need to know exactly which specimens are involved. ACKNOWLEDGMENTS Before his death in 1953, G. H. H. Tate began to write an account of "the living land mammals of Latin America and the West In- dies." He did not complete the work. A copy in the archives of the Department of Mam- malogy provided some ideas on what might be useful in the introductory remarks for var- ious taxa. Each of the many authors whose works are cited below has contributed to this report, as have the many field collectors listed in the sections on history, each to a greater or lesser degree. Many small contributions, in the ag- gregate, amount to a significant part of the total body of knowledge compiled and syn- thesized here. The curators and other persons responsible for all the collections cited have been most helpful. I am grateful to Dr. Leslie Marcus for extracting data from my comput- er files and then preparing the final distri- bution maps with a computer. Walter de Gruyter and Co. granted permission to quote from Koopman (1994) on bats. Dr. Kurt Bauer of Vienna, Austria, sent information on the journey of J. Natterer in 1829. No- buko E. Pine translated pertinent parts of a number of Japanese articles. My wife, friend, field assistant, and pho- tographer, R. Justine Anderson, helped throughout this protracted project. Ted Dan- forth volunteered to prepare the initial draft of the gazetteer. The late Marie Lawrence was of great help in bibliographic matters. Nancy Olds collaborated in the lab and the field, and as an Exxon Fellow obtained and compiled many of the measurements for ta- bles. Muriel V. Williams provided highly competent secretarial services, including nu- merous reshufflings and revisions of com- puter files. A host of students and other vol- unteers have assisted in the lab in ways too numerous to mention. In addition to essential and important support for curatorial, techni- cal, secretarial, administrative, and library services, the American Museum of Natural History provided direct financial support for fieldwork and travel to study other collec- tions through the Koopman Taxonomic Mammalogy Fund and the Bolivian Expedi- tions Fund. Fieldwork in 1964 and 1965 was support- ed in part by the U.S. Army Medical Re- search and Development Command through grant number DA-MD-49-193-63-G82. The principal purpose was to conduct a survey of mammals and their ectoparasites, but many other vertebrates and arthropods were col- lected as well. Collaborative fieldwork in most of the years from 1985 to 1993 was supported in part by grants from the National Science Foundation to the American Museum of Nat- ural History, the University of New Mexico, and the University of California at Davis, in- cluding the following: BSR-83-16740, BSR- 84-08923, BSR-86-12329, BSR-89-20617, BSR-90-24816, and INT-92-12839. Another grant was received from the National Insti- tutes of Health (DRR-RRO8139). Graduate students working with us obtained grants, largely on their own initiatives, from the 5 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY American Society of Mammalogists, Sigma Xi, the Hayward Foundation, and the Tinker Foundation and the Mellon Foundation (via the Latin American Institute of the Univer- sity of New Mexico). Semifinal drafts of this paper were reviewed by David M. Arm- strong, Janet K. Braun, Karl E Koopman, Michael A. Mares, Bruce D. Patterson, James L. Patton, Ronald H. Pine, Jorge Salazar B., and Terry L. Yates. OBJECTIVES AND ORGANIZATION The primary objective of this volume is to summarize the present state of knowledge (and, by implication, its important reciprocal, ignorance) of the taxonomy and distributions of native mammals in present-day Bolivia. The reader needs to look elsewhere for illus- trations in color, illustrations of whole ani- mals, or general summaries of biology of the species treated here. Information contained in those sources is not summarized, although a few are cited for the convenience of read- ers. The focus is on taxonomy and distribu- tion, on new information, and on specimens or studies within Bolivia. Some notes on top- ics other than taxonomy and distribution are included when they are based on work within Bolivia. In most such cases, although the data are of limited scope, the paucity of such information warrants its inclusion. For ex- ample, under the category of "reproduc- tion," a few scattered records of litter size, months of pregnancies, and presence of young may be given. Fieldwork and collecting by Bolivians, in collaboration with foreign workers, have in- creased in recent years. Because taxonomic studies are continuing, we can look forward to continued progress in understanding the distribution and taxonomy of Bolivian mam- mals. But considering what needs to be learned, work has just begun. Account after account will mention or imply more unre- solved questions than resolved ones. The keys, tables of measurements, synonymies, maps, lists of specimens, and other infor- mation here offered should provide a basis for further work. In piecing together the details of geo- graphic distributions of animals or plants, it is not sufficient to have only a list of local- ities from specimen labels or other sources. One needs to know the detailed itineraries of the individual collectors. Sometimes routes of travel are described in original fieldnotes. In other cases, when notes were not avail- able, the itinerary needed to be reconstructed by comparing dates and locality designations for an entire collection. If the collector was available, he or she might be consulted di- rectly, and I have done this in several cases. Published maps or descriptions of expedi- tions are helpful in other cases. In every case, geographic and taxonomic detective work is required. For example, there are dozens of places named San Ram6n in Bolivia. In fact there may be as many as three different lo- cations named San Ramon in one province within a single department. A section titled Mammalogical Gazetteer of Bolivia provides further detail on all known collecting local- ities. Unresolved problems are also noted. Subsequent sections-Collecting in Bolivia and Persons and Institutions-also help es- tablish precise localities. Acronyms for col- lections housing Bolivian specimens are in- cluded. All cited material is listed in the Refer- ences except some papers in which names above the level of species were proposed. These may be found in other sources such as Wilson and Reeder (1993). TAXONOMIC CONCEPTS The use of the genus as an essential part of a hierarchical classification dates from the arbitrarily established beginning of scientific nomenclature in zoology (Linnaeus, 1758). A genus is a taxon at a slightly higher level in a classification than a species. In theory, if a genus is regarded as a group, then the idea of a genus containing only one species seems redundant. However, about one-half of living mammalian species are placed in gen- 6 NO. 231 ANDERSON: MAMMALS OF BOLIVIA era with only one species. These are the monotypic, or at least monospecific, genera. Monotypic genera are reasonable in terms of the traditional use of a phenetic criterion by which the degree of morphological differ- ence is considered. Other criteria may be considered, for example phylogeny or strict monophyly are more important in a cladistic approach to classification. For most mam- malian genera, no careful cladistic analysis has been done and thus no well established hypothesis of detailed phylogenetic relation- ships exists. The future completion of such analyses and hypotheses is desirable. How- ever, for practical taxonomy now, I am in- clined to continue using the existing classi- fication until additional and reasonably strong evidence, as opposed to a mere dif- ference of opinion, is presented in favor of a change. This view is eclectic, not strictly phenetic or cladistic, and, in the interests of nomenclatural stability, my taxonomy is cau- tiously conservative. Another consideration is the convenience for readers when an au- thor uses some readily available and familiar classification, and then documents only the departures therefrom. For this reason, the classification used here is generally that of Wilson and Reeder (1993). Their classifica- tion did not deal with subspecies. Some of the implications and ramifications of these different taxonomic approaches were dis- cussed earlier (Anderson, 1974, 1985a) and need not be repeated here. Another recent re- view of taxonomic trends in mammalogy is that of Engstrom et al. (1994). The concept of species in systematic biol- ogy has an interesting and varied history and no one definition can be applied to all groups. Furthermore, different biologists may have different concepts for the same group of or- ganisms. For Recent mammals, I think that the so-called "biological species concept" of actually or potentially interbreeding popula- tions of individuals has more advantages than any alternative concept. I hold this view in full awareness that the actual entities we deal with are individual specimens and that inter- breeding is rarely observed, but is an infer- ence from morphological data. Most species have relatively small geo- graphic ranges and may be expected to exhibit little geographic variation. However the ma- jority of species that have larger geographic ranges may be expected to exhibit some sig- nificant degree of geographic variation. Sub- species, with trinomial names, have been used generally for formal recognition of selected contiguous geographic populations that have both some degree of morphological homoge- neity and some degree of difference from oth- er subspecies. The literature displays a wide range of taxonomic opinions about what de- grees of distributional contiguity of local pop- ulations within a subspecies-or what degree of morphological (karyological, biochemical, etc.) homogeneity-should be recognized by formal subspecies. Likewise, opinions vary on the degree of difference that should be re- quired before separate subspecies are recog- nized, or whether the concept should simply be abandoned. The concept has not been abandoned, and I do not propose to do so (again, I refer to an earlier discussion in An- derson and Nelson, 1965). A recently pub- lished set of papers (Kimbel and Martin, 1993) nicely summarizes the ongoing saga of species concepts in taxonomy. Now, having stated my intention to use genera, species, and subspecies, let me em- phasize several qualifications and limitations, in the context of current knowledge of the taxonomy of Bolivian mammals. Every tax- on recognized here, from the subspecies on up, should be regarded as a taxonomic hy- pothesis subject to, or in most cases in crit- ical need of, further testing. Every student who sits down and attempts to identify a specimen using this publication should un- derstand that he or she is testing the current taxonomic and biogeographic hypotheses. The specimen may be of a species or sub- species not previously known from Bolivia. It may be of a species unknown to science. It may be from a local population with some- what different characters than any examined before. It may demonstrate that the morpho- logical limits of the species, as presently un- derstood, need to be redefined. It may dem- onstrate that one or more of the characters used in the keys are wrong or need modifi- cation. The essential point is that taxonomy is not a static body of knowledge, but a dy- namic process leading to better understand- ing, not to absolute truth. 1997 7 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY LIST OF TAXA All native species (321, including Homo sapiens) of mammals represented by speci- mens in collections are listed here and pre- sented in the following accounts. Also in- cluded (and noted as such in this list) are ten domestic species, two that are native to the Americas and eight that have been intro- duced from the Old World. At least three oth- er introduced species (of the genera Mus, Rattus, and Lepus) exist as feral populations. Imbedded among the accounts of known spe- cies, and in a separate summary following these accounts, are references to species of postulated occurrence, which are not includ- ed in this list. The sequence of orders, families, subfam- ilies, and tribes is mostly that of Wilson and Reeder (1993). Orders are arranged phylo- genetically, i.e., with more "primitive" groups placed before more "derived" or "specialized" groups. The published litera- ture reflects uncertainty about this somewhat traditional sequence and has resulted in var- Order Marsupialia Family Didelphidae Subfamily Caluromyinae Genus Caluromys C. lanatus lanatus .................. Genus Glironia G. venusta ........................ Subfamily Didelphinae Genus Chironectes C. minimus minimus ................ Genus Didelphis D. albiventris albiventris ............ D. albiventris pernigra ............. D. marsupialis marsupialis .......... Genus Gracilinanus G. aceramarcae.................... G. agilis buenavistae ............... G. agilis chacoensis ................ G. agilis unduaviensis .............. Genus Lutreolina L. crassicaudata crassicaudata ...... Genus Marmosa M. lepida ......................... M. murina ........................ ious placements of orders such as the Xe- narthra, Cetacea, and others. In Wilson and Reeder (1993), families and subfamilies are arranged alphabetically in some orders and phylogenetically in others. These "phyloge- netic" arrangements must be somewhat ar- bitrary because the phylogeny produced by evolution is a branching tree and not a linear sequence. The arrangement of taxa at the ge- neric, subgeneric, specific, and subspecific levels is alphabetical. This arrangement is for the convenience of readers, most of whom know the alphabet better than they know the currently postulated phylogenetic relation- ships at this level. The List of Taxa has page references and thus can serve both as a checklist and as a table of contents. Numbers of localities and numbers of specimens are given also, to pro- vide an overview of commonness and rarity. For species with more than one subspecies, the totals are for the entire species rather than the separate subspecies. Locality Total Pg. total Spec. 140 140 141 143 144 145 146 147 148 148 149 150 150 7 20 2 2 2 8 25 62 77 249 2 25 3 68 4 10 2 7 7 13 8 NO. 231 1997 ANDERSON: MAMMALS OF BOLIVIA 9 Locality Total Pg. total Spec. Genus Marmosops M. dorothea ......... ................. 151 23 46 M. impavidus ......... ................ 152 5 6 M. noctivagus keaysi ...... ............ 154 17 45 M. parvidens ......... ................ 154 1 1 Genus Metachirus M. nudicaudatus bolivianus ............ 155 22 45 Genus Micoureus M. constantiae budini ...... ........... 156 55 142 M. constantiae constantiae ..... ........ 156 M. regina .......... .................. 157 11 13 Genus Monodelphis M. adusta .......... .................. 158 1 2 M. brevicaudata ....... ............... 158 6 12 M. domestica ......... ................ 159 25 85 M. emiliae .......... ................. 159 1 1 M. kunsi ........... .................. 160 3 3 M. osgoodi .......... ................. 160 3 4 Genus Philander P. opossum canus ....... .............. 162 71 186 Genus Thylamys T. macrurus ......... ................. 163 1 1 T. pallidior .......... ................. 163 19 48 T. pusillus .......... .................. 164 14 48 T. venustus .......... ................. 165 52 181 Family Caenolestidae Genus Lestoros L. inca ............................... 166 1 1 Order Xenarthra Family Bradypodidae Genus Bradypus B. variegatus ......... ................ 167 18 51 Family Choloepidae Genus Choloepus C. hoffnanni ......... ................ 168 2 2 Family Dasypodidae Subfamily Chlamyphorinae Genus Chlamyphorus C. retusus retusus ....... .............. 169 3 6 Subfamily Dasypodinae Tribe Dasypodini Genus Dasypus D. kappleri beniensis ...... ............ 170 3 3 D. novemcinctus novemcinctus .......... 171 42 429 D. septemcinctus ....... ............... 172 5 9 Tribe Euphractini Genus Chaetophractus C. nationi .......... .................. 173 8 24 C. vellerosus ......... ................ 174 9 12 C. villosus .......... ................. 174 3 8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Locality Total Pg. total Spec. Genus Euphractus E. sexcinctus boliviae ...... ............ 175 23 44 Tribe Priodontini Genus Cabassous C. unicinctus squamicaudis ..... ........ 177 4 4 Genus Priodontes P. maximus ........................... 177 10 16 Tribe Tolypeutini Genus Tolypeutes T. matacus .......... ................. 178 14 29 Family Myrmecophagidae Genus Cyclopes C. didactylus catellus ...... ............ 179 8 36 Genus Myrmecophaga M. tridactyla tridactyla ...... .......... 180 15 22 Genus Tamandua T. tetradactyla ........ ................ 181 46 129 Order Chiroptera Family Emballonuridae Genus Peropteryx P. kappleri .......... ................. 184 2 5 P. macrotis macrotis ...... ............ 185 7 16 Genus Rhynchonycteris R. naso ............. ................. 185 17 55 Genus Saccopteryx S. bilineata .......... ................. 186 15 75 S. Ieptura ........... ................. 187 3 12 Family Noctilionidae Genus Noctilio N. albiventris affinis ....... ............ 189 41 522 N. leporinus rufescens ...... ........... 190 26 50 Family Mormoopidae Genus Pteronotus P. (Chilonycteris) personatus personatus 191 1 1 P. (Phyllodia) parnellii rubiginosus ..... 192 3 3 P. (Pteronotus) gymnonotus ............ 192 1 1 Family Phyllostomidae Subfamily Phyllostominae Genus Chrotopterus C. auritus ............ ................ 193 7 7 Genus Lonchorhina L. aurita .............. ............... 194 1 1 Genus Macrophyllum M. macrophyllum ........ ............. 194 7 9 Genus Micronycteris M. (Barticonycteris) daviesi ............ 195 1 1 M. (Micronycteris) megalotis megalotis .. 196 2 3 M. (M.) minuta ....... ................. 197 7 9 M. (Trinycteris) nicefori ...... ......... 197 1 2 10 NO. 231 ANDERSON: MAMMALS OF BOLIVIA Locality Total Pg. total Spec. Genus Mimon M. crenulatum longifolium ..... ........ 198 5 6 Genus Phyllostomus P. discolor discolor ....... ............ 199 10 31 P. elongatus ........... ............... 200 15 24 P. hastatus hastatus ....... ............ 201 46 132 P. stenops boliviensis ....... ........... 202 6 8 Genus Tonatia T. brasiliense .......... ............... 203 3 3 T. carrikeri ............ ............... 203 1 1 T. saurophila maresi ....... ........... 204 5 5 T. sylvicola sylvicola ....... ........... 205 16 30 Genus Trachops T. cirrhosus cirrhosus ....... .......... 207 15 21 Genus Vampyrum V. spectrum ........... ............... 207 1 1 Subfamily Lonchophyllinae Genus Lonchophylla L. thomasi ............ ............... 208 9 12 Subfamily Glossophaginae Genus Anoura A. caudifer ............. .............. 209 20 42 A. cultrata ............ ............... 210 1 2 A. geoffroyi geoffroyi ....... ........... 211 25 83 A. geoffroyi peruana ....... ........... 211 Genus Choeroniscus C. intermedius ......... ............... 212 1 1 C. minor ........... .................. 212 3 3 Genus Glossophaga G. soricina soricina ....... ............ 213 124 728 Genus Lichonycteris L. obscura ............ ............... 215 1 1 Subfamily Carolliinae Genus Carollia C. brevicauda .......... .............. 216 100 413 C. castanea ............ .............. 218 22 123 C. perspicillata ......... .............. 219 180 1597 Genus Rhinophylla R. pumilio ............. ............... 222 10 45 Subfamily Stenodermatinae Genus Artibeus A. anderseni ............ .............. 223 68 183 A. glaucus glaucus ........ ............ 225 33 100 A. glaucus gnomus ....... ............. 226 A. hartii .............. ............... 226 6 6 A. jamaicensis fallax ....... ........... 228 123 720 A. lituratus lituratus ....... ............ 230 135 932 A. obscurus ............ .............. 232 133 381 Genus Chiroderma C. salvini salvini ...................... 235 10 16 11 1997 12 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 231 Locality Total Pg. total Spec. C. trinitatum trinitatum ...... .......... 236 20 61 C. villosum villosum ....... ............ 237 32 105 Genus Mesophylla M. macconnelli macconnelli ............ 238 17 34 Genus Platyrrhinus P. brachycephalus ..................... 239 5 7 P. dorsalis ............. .............. 239 12 78 P. helleri incarum ........ ............. 241 87 426 P. infuscus .............. ............. 242 17 39 P. lineatus lineatus ........ ............ 243 34 175 P. lineatus nigellus ............. 244 P. vittatus ............ ................ 244 2 9 Genus Pygoderma P. bilabiatum magna ....... ........... 245 18 53 Genus Sphaeronycteris S. toxophyllum ......... ............... 246 9 11 Genus Sturnira S. erythromos ......................... 246 44 237 S. lilium lilium ......... ............... 248 146 669 S. magna ............ ................. 250 7 20 S. oporaphilum ........ ............... 251 38 310 S. tildae .............................. 252 16 42 Genus Uroderma U. bilobatum bilobatum ...... .......... 254 97 374 U. bilobatum thomasi ....... ........... 254 U. magnirostrum ........ .............. 255 69 192 Genus Vampyressa V. bidens ............................. 257 5 11 V. pusilla thyone ........ .............. 258 4 8 Genus Vampyrodes V. caraccioli major ....... ............. 259 4 21 Subfamily Desmodontinae Genus Desmodus D. rotundus .......... ................ 259 95 392 D. youngii ........... ................ 261 13 20 Genus Diphylla D. ecaudata .......... ................ 262 2 2 Family Thyropteridae Genus Thyroptera T. discifera discifera ....... ........... 264 1 14 T. tricolor tricolor ....... ............. 264 6 18 Family Vespertilionidae Genus Eptesicus E. andinus ........ ................... 265 2 3 E. furinalis chapmani ....... ........... 266 36 128 E. furinalis furinalis ....... ............ 267 E. furinalis montosus ....... ........... 267 Genus Histiotus H. montanus laephotis ...... ........... 268 5 7 H. velatus ........... ................. 268 4 4 ANDERSON: MAMMALS OF BOLIVIA Locality Total Pg. total Spec. Genus Lasiurus L. blossevillii frantzii ....... ........... 269 12 15 L. cinereus villosissimus ...... ......... 270 10 11 L. ega ega ........... ................ 271 19 38 Genus Myotis M. (Leuconoe) albescens ...... ......... 272 36 282 M. (L.) levis dinellii ....... ............ 274 5 8 M. (L.) oxyotus oxyotus ...... .......... 275 6 12 M. (L.) riparius ......... .............. 275 3 4 M. (L.) simus .......... ............... 276 4 48 M. (Selysius) keaysi keaysi ..... ........ 277 10 12 M. (S.) nigricans nigricans ..... ........ 278 99 1544 Genus Rhogeessa R. tumida ............ ................ 280 1 1 Family Molossidae Genus Eumops E. auripendulus auripendulus ........... 282 11 49 E. bonariensis beckeri ...... ........... 282 16 396 E. glaucinus glaucinus ...... ........... 283 8 25 E. hansae ............ ................ 284 2 2 E. perotis perotis ....... .............. 285 6 52 E. perotis trumbulli ....... ............ 285 Genus Molossops M. abrasus ........... ................ 285 1 1 M. planirostris planirostris ..... ........ 287 8 25 M. temminckii temminckii ...... ........ 287 17 59 Genus Molossus M. molossus crassicaudatus ............ 289 90 1839 M. rufus ............. ................ 290 17 67 Genus Nyctinomops N. aurispinosus ........ ............... 291 3 10 N. laticaudatus europs ...... ........... 292 13 130 N. laticaudatus laticaudatus ............ 292 N. macrotis ........................... 293 3 3 Genus Promops P. centralis occultus ....... ............ 293 3 5 P. nasutus ancilla ........ ............. 294 3 4 Genus Tadarida T. brasiliensis brasiliensis ...... ........ 295 11 67 Order Primates Family Callitrichidae Subfamily Callimiconinae Genus Callimico C. goeldii .... ......................... 296 2 3 Subfamily Callitrichinae Genus Callithrix C. melanura .......... ................ 298 13 31 Genus Cebuella 1997 13 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Locality Total Pg. total Spec. C. pygmaea .......................... 299 1 1 Genus Saguinus S. fuscicollis weddellii ...... ........... 301 9 22 S. imperator imperator ...... .......... 302 1 1 S. labiatus labiatus ....... ............. 302 8 16 Family Cebidae Subfamily Alouattinae Genus Alouatta A. caraya .......... .................. 304 17 75 A. guariba beniensis .......... ......... 305 2 6 A. seniculus sara ........... ........... 307 35 128 Subfamily Aotinae Genus Aotus A. azarai azarai .................... 309 49 145 A. azarai boliviensis ....... ............ 309 A. nigriceps .......................... 310 3 4 Subfamily Atelinae Genus Ateles A. chamek .............. 311 33 93 Subfamily Callicebinae Genus Callicebus C. brunneus .......................... 313 6 9 C. donacophilus donacophilus .......... 314 18 86 C. donacophilus pallescens ............. 315 C. modestus .......................... 315 1 2 C. olallae ............................ 316 1 1 Subfamily Cebinae Genus Cebus C. albifrons cuscinus .................. 317 24 59 C. albifrons unicolor .................. 317 C. apella pallidus ..................... 319 57 174 C. apella paraguayanus ................ 320 Genus Saimiri S. sciureus boliviensis ...... ........... 322 54 232 Subfamily Pitheciinae Genus Pithecia P. irrorata irrorata ....... ............. 324 3 3 Family Hominidae Genus Homo H. sapiens ........................... 325 Order Carnivora Family Canidae Genus Atelocynus A. microtis .......... ................. 325 4 4 Genus Canis (domestic, introduced) C. familiaris ......... ................. 326 12 15 Genus Cerdocyon C. thous entrerianus ....... ............ 327 40 116 Genus Chrysocyon C. brachyurus ........ ................ 328 5 4 14 NO. 231 ANDERSON: MAMMALS OF BOLIVIA Locality Total Pg. total Spec. Genus Pseudalopex P. culpaeus andina ....... ............. 330 16 23 P. gymnocercus gymnocercus ........... 331 12 43 Genus Speothos S. venaticus venaticus ...... ........... 332 3 8 Family Felidae Genus Felis F. (Felis) catus (domestic, introduced) ... 332 2 2 F. (Herpailurus) yaguarondi eyra ....... 333 6 24 F. (Leopardus) pardalis steinbachi ...... 334 26 68 F. (L.) wiedii boliviae ...... ........... 335 12 24 F. (Lynchailurus) pajeros garleppi ...... 336 4 6 F. (Oncifelis) geoffroyi euxantha ........ 337 10 22 F. (Oreailurus) jacobita ...... ......... 337 9 14 F. (Puma) concolor acrocodia .......... 339 23 38 F. (P.) concolor osgoodi ...... ......... 339 Genus Panthera P. onca palustris ........ .............. 341 22 28 P. onca peruviana ....... ............. 341 Family Mustelidae Subfamily Lutrinae Genus Lutra L. longicaudis enudris ...... ........... 343 17 30 Genus Pteronura P. brasiliensis paranensis ...... ........ 344 2 6 Subfamily Mephitinae Genus Conepatus C. chinga rex .......... ............... 345 18 24 Subfamily Mustelinae Genus Eira E. barbara barbara ....... ............ 346 26 71 E. barbara peruana ....... ............ 346 Genus Galictis G. cuja luteola ......... .............. 347 10 10 G. vittata andina ........ .............. 348 6 9 Genus Mustela M. frenata boliviensis .................. 349 5 6 Family Procyonidae Subfamily Potosinae Genus Bassaricyon B. alleni .............. ............... 349 4 4 Genus Potos P. flavus chapadensis ....... ........... 351 15 39 Subfamily Procyoninae Genus Nasua N. nasua boliviensis ....... ............ 352 58 278 N. nasua dorsalis ........ ............. 352 N. nasua spadicea ....... ............. 353 Genus Procyon P. cancrivorus cancrivorus ............. 354 13 30 1997 15 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Locality Total Pg. total Spec. Family Ursidae Genus Tremarctos T. ornatus .......... .................. 354 9 19 Order Odontoceti Family Platanistidae Genus Inia L. geoffrensis boliviensis ............... 359 12 18 Order Perissodactyla Family Equidae (domestic, introduced) Genus Equus E. asinus ........... .................. 360 1 2 E. caballus .......... ................. 360 1 1 Family Tapiridae Genus Tapirus T. terrestris spegazzinii ...... .......... 360 24 35 Order Artiodactyla Family Suidae (domestic, introduced) Genus Sus S. scrofa ........... .................. 361 2 3 Family Tayassuidae Genus Catagonus C. wagneri .......... ................. 362 4 5 Genus Tayassu T. pecari albirostris ....... ............ 363 36 213 T. tajacu tajacu ........ ............... 364 57 316 Family Camelidae Genus Lama L. (Lama) glama (domestic, native) ..... 366 2 3 L. (L.) guanicoe guanicoe ..... ......... 366 9 11 L. (Vicugna) pacos (domestic, native) ... 367 7 17 L. (V.) vicugna ........ ............... 368 25 50 Family Cervidae Subfamily Odocoileinae Genus Hippocamelus H. antisensis ......... ................ 369 6 12 Genus Mazama M. americana ........... 370 24 53 M. bricenii chunyi ....... .............. 372 2 2 M. gouazoupira gouazoupira ........... 373 34 83 Genus Odocoileus 0. (Blastocerus) dichotomus ..... ....... 374 16 62 0. (Odocoileus) virginianus peruvianus 375 1 0 0. (Ozotoceros) bezoarticus leucogaster 376 2 5 Family Bovidae (domestic, introduced) Subfamily Bovinae Genus Bos B. taurus .............. ............... 376 1 1 Subfamily Caprinae Genus Capra C. hircus .............. ............... 377 1 1 16 NO. 231 1997 ANDERSON: MAMMALS OF BOLIVIA 17 Locality Total Pg. total Spec. Genus Ovis 0. aries ............. ................. 377 1 1 Order Rodentia Suborder Sciurognathi Family Sciuridae Genus Sciurus S. ignitus argentinius ...... ............ 378 59 149 S. ignitus boliviensis ....... ............ 379 S. ignitus ignitus ........ .............. 379 S. spadiceus .......... ................ 380 48 183 Family Muridae Subfamily Murinae (introduced, commensal) Genus Mus M. musculus .......... ................ 382 66 731 Genus Rattus R. rattus ............ ................. 383 57 328 Subfamily Sigmodontinae Tribe Oryzomyini Genus Microryzomys M. minutus .......... ................. 385 10 28 Genus Neacomys N. spinosus spinosus ...... ............ 386 43 124 Genus Nectomys N. squamipes garleppii ...... .......... 388 13 24 Genus Oecomys 0. bicolor bicolor ....... .............. 389 32 193 0. concolor ......... ................. 389 3 3 0. mamorae ......... ................. 390 44 132 0. roberti ........... ................. 391 2 3 Genus Oligoryzomys 0. andinus .......... ................. 392 6 14 0. chacoensis ........ ................ 392 20 144 0. destructor ......... ................ 393 53 181 0. flavescens group, sp. B ............. 395 33 239 0. microtis .......... ................. 396 126 618 Genus Oryzomys 0. capito ............................. 399 78 313 0. legatus ........... ................. 400 13 45 0. levipes ........... ................. 401 36 248 0. nitidus ........... ................. 402 103 508 0. subflavus ......... ................. 404 56 320 0. yunganus ......... ................. 405 8 11 Genus Rhipidomys R. couesi austrinus ....... ............. 407 20 72 R. leucodactylus rex ....... ............ 408 4 4 R. nitela ............ ................. 408 3 3 Genus Thomasomys T. aureus aureus ........ .............. 409 11 20 T. daphne australis ....... ............. 410 12 44 18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 231 Locality Total Pg. total Spec. T. ladewi ............................. 410 4 12 T.oreas ............... 411 1 2 T. taczanowskii .......... ............. 411 1 2 Tribe Akodontini Genus Akodon A. aerosus baliolus .................... 413 37 206 A. albiventer berlepschii ............... 415 45 191 A. boliviensis ......................... 416 54 342 A. dayi .............................. 418 41 257 A. fumeus ............................ 419 50 253 A. lutescens caenosus .................. 421 53 371 A. lutescens lutescens .................. 422 A. lutescens puer ...................... 422 A. mimus .............. 423 28 533 A. pervalens .............. ............ 424 6 10 A. siberiae ........................... 425 6 35 A. simulator tartareus ................. 426 10 122 A. subfuscus subfuscus ................. 427 16 75 A. toba .............................. 427 10 91 A. varius ............................. 428 42 345 Genus Bolomys B. amoenus ............. 430 4 13 B. lactens lactens ..................... 431 6 15 B. lenguarum tapirapoanus ............. 431 116 913 Genus Chroeomys C. andinus dolichonyx ................. 434 9 25 C. jelskii bacchante ................... 435 17 113 C. jelskii ochrotis ..................... 435 C. jelskii pulcherrimus ................. 436 C. jelskii sodalis ...................... 436 Genus Lenoxus L. apicalis boliviae ........ ............ 436 5 35 Genus Oxymycterus 0. hucucha ........................... 437 4 5 0. inca doris ......................... 437 27 91 0. inca iris ........................... 438 0. paramensis jacentior ............... 439 55 352 0. paramensis nigrifrons ............... 439 0. paramensis paramensis ............. 440 Tribe Scapteromyini Genus Kunsia K. tomentosus tomentosus .............. 440 3 4 Tribe uncertain Genus Pseudoryzomys P. simplex ............... 441 5 6 Tribe Phyllotini Genus Andinomys A. edax edax ............ ............. 442 23 69 Genus Auliscomys A. (Maresomys) boliviensis boliviensis ... 444 9 48 ANDERSON: MAMMALS OF BOLIVIA Locality Total Pg. total Spec. A. (Auliscomys) pictus ................. 444 11 30 A. (A.) sublimis leucurus ............... 445 24 175 A. (A.) sublimus sublimis ............... 445 Genus Calomys C. callosus callosus ................... 447 135 2829 C. laucha ............................ 449 4 21 C. lepidus carillus .......... .......... 450 44 248 C. lepidus ducillus .......... .......... 451 C. musculinus ........................ 451 26 99 C. tener ............... 452 3 3 C. venustus ............. 453 30 85 Genus Chinchillula C. sahamae .......................... 454 6 15 Genus Eligmodontia E. puerulus ........................... 455 24 77 Genus Galenomys G. garleppii .......................... 456 4 12 Genus Graomys G. domorum domorum ......... ........ 458 41 255 G. domorum taterona ......... ......... 459 G. griseoflavus griseoflavus ............ 460 15 210 G. pearsoni dorbignyi ................. 461 1 2 Genus Neotomys N. ebriosus ebriosus ................... 462 9 26 N. ebriosus vulturnus .................. 462 Genus Phyllotis P. caprinus ........................... 462 7 58 P. chilensis ........................... 463 9 66 P. osilae osilae ....................... 465 39 198 P. osilae phaeus ........... ........... 465 P. wolffsohni ......................... 465 36 201 P. xanthopygus rupestris ............... 467 73 493 Genus undescribed ....................... 469 1 2 Tribe uncertain Genus Holochilus H. sciureus amazonicus ................ 470 49 263 Tribe Ichthyomyini Genus Chibchanomys C. unidentified ........................ 471 1 1 Suborder Hystricognathi Family Erethizontidae Genus Coendou C. bicolor simonsi .......... .......... 472 13 16 C. prehensilis boliviensis ............... 473 14 50 Family Chinchillidae Genus Chinchilla (probably extinct in Bolivia) C. chinchilla ......................... 473 3 12 Genus Lagidium L. viscacia cuscus ....... .............. 476 47 138 L. viscacia cuvieri ....... .............. 476 L. viscacia perlutea ....... ............ 477 19 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Locality Total Pg. total Spec. Genus Lagostomus L. maximus inmollis ....... ............ 478 7 16 Family Dinomyidae Genus Dinomys D. branickii ........... ............... 478 2 3 Family Caviidae Subfamily Caviinae Genus Cavia (domestic and wild) C. tschudii nana ......... ............. 480 45 149 Genus Galea G. musteloides auceps ...... ........... 481 68 266 G. musteloides demissa ...... .......... 481 G. musteloides musteloides ...... ....... 482 G. spixii campicola ........ ............ 483 4 13 Genus Microcavia M. niata niata .......... .............. 484 12 46 M. niata pallidior ........ ............. 484 Subfamily Dolichotinae Genus Dolichotis D. salinicola salinicola ...... .......... 485 8 12 Family Hydrochaeridae Genus Hydrochaeris H. hydrochaeris hydrochaeris .......... 486 28 65 Family Dasyproctidae Genus Dasyprocta D. punctata boliviae ....... ............ 488 55 248 D. punctata urucuma ....... ........... 489 D. punctata yungarum ...... ........... 489 Family Agoutidae Genus Agouti A. paca paca .......... ............... 490 43 146 Family Octodontidae Subfamily Ctenomyinae Genus Ctenomys C. boliviensis boliviensis ...... ......... 492 27 486 C. conoveri ........................... 493 10 15 C. frater frater .......... .............. 494 8 36 C. frater mordosus ........ ............ 494 C. goodfellowi .......... .............. 495 2 4 C. leucodon ........... ............... 495 8 29 C. lewisi ............. ................ 496 2 57 C. minutus ............ ............... 496 1 3 C. opimus opimus ........ ............. 497 30 167 C. steinbachi .......... ............... 498 14 86 undescribed taxa ........ .............. 499 Subfamily Octodontinae Genus Octodontomys 0. gliroides ........... ............... 499 10 44 20 NO. 231 ANDERSON: MAMMALS OF BOLIVIA Locality Total Pg. total Spec. Family Abrocomidae Genus Abrocoma A. boliviensis ......................... A. cinerea cinerea ..................... Family Echimyidae Subfamily Dactylomyinae Genus Dactylomys D. boliviensis ......................... Subfamily Echimyinae Genus Echimys E. didelphoides ....................... Genus Isothrix L bistriata bistriata ................... Subfamily Eumysopinae Genus Mesomys M. hispidus........................... Genus Proechimys P. brevicauda ........................ P. hilda .............................. P. longicaudatus ...................... P. simonsi............................ P. steerei ............................ Genus Thrichomys T. apereoides fosteri ................... Family Myocastoridae Genus Myocastor M. coypus popelairi ................... Order Lagomorpha Family Leporidae Genus Lepus (introduced, feral) L. europaeus ......................... Genus Oryctolagus (domestic, introduced) 0. cuniculus .......................... Genus Sylvilagus S. brasiliensis gibsoni ................. S. brasiliensis inca .................... S. brasiliensis paraguensis ............. 500 501 501 502 503 504 505 507 508 509 510 512 513 513 514 515 515 516 2 7 6 2 41 13 2 2 3 3 73 8 30 8 15 1 4 3 4 485 10 153 13 71 1 S 1 1 0 0 35 86 KEYS TO THE MAMMALS OF BOLIVIA After a preliminary identification of a specimen is obtained by using the keys, fur- ther testing should be done by checking the distributional map, the tables of measure- ments, and the individual account. In difficult cases, comparison with specimens in collec- tions and with newer literature will be need- ed. The following 28 illustrated keys are in- tended to aid in the identification of native and introduced species of mammals known or hypothesized to occur in Bolivia now or formerly (i.e., within historic times). Hypoth- esized species not actually represented by a specimen in a museum collection are indi- cated by an asterisk (*). A number of species that I might include if I were rewriting the key now (1995) are not here. The first key is 21 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY to orders and families and it refers the user to the appropriate family key if more than one species of the family is involved. The illustrations are intended to help the user who is not already familiar with the technical terminology or with the distinction emphasized in the key. Both external and in- ternal (cranial and dental) features are used, so it is necessary in some cases to have a specimen consisting of both skin and cleaned skull. Calipers and lenses for magnification are also needed in some cases. If there is doubt about a given choice, proceed to both of the alternatives and see if subsequent choices seem relevant. If they do not seem relevant, then the alternative original choice is indicated. Ideas and illustrations for the keys have been derived from many sources. It is not feasible to acknowledge every specific source, but major sources were the follow- ing: Tate (1933, Marmosa revision) Goodwin and Greenhall (1961, bats of Trinidad, drawings used here as figs. 65, 69-70, 103-104, 110, 114, 132, 135-138, and 185-186) Anderson (1972, Chihuahuan keys, drawings used here as figs. 172, 225, 261-262, 290-291) Rouk and Carter (1972, drawings from the de- scription of V. brachycephalus used here as fig. 157) Pine (1973a, Brazilian key) Eger (1977, Eumops revision, source of figs. 190- 192) Husson (1978, Surname, drawings used here as figs. 3, 34, and 35) Langguth and Anderson (1980, Uruguayan key, drawings by Fran Stiles appear here as figs. 249-250, 253-254, 259, 272-274) Hall (1981, drawings by Victor Hogg used here as figs. 43-46, 116-119, 133-134, 189, and 193-194) Albuja (1982, Ecuadorian bat key) Anderson and Jones (1984, drawings by Marie Dauenheimer, used here as figs. 9 and 218) Linares (1986, Venezuelan bat key) Olds (1988, the genus Calomys) List of Numbered Keys 1. General key to orders and families 2. Didelphidae 3. Emballonuridae 4. Noctilionidae 5. Mormoopidae 1 1 1 1 1 1 1 6. Phyllostomidae 7. Thyropteridae 8. Vespertilionidae 9. Molossidae L0. Callitrichidae and Cebidae L1. Myrmecophagidae L2. Dasypodidae L3. Canidae L4. Procyonidae L5. Mustelidae L6. Felidae 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. Tayassuidae Camelidae Cervidae Bovidae Leporidae Sciuridae Muridae Echimyidae Octodontidae and Abrocomidae Chinchillidae Caviidae Erethizontidae KEYl1 ORDERS AND FAMILIES OF LIVING MAMMALS OF BOLIVIA I Incisor teeth reduced to a single large and persistently growing pair in both upper and lower jaws (fig. 1); no small sec- ondary pair behind the upper pair; wide gap between incisors and grinding teeth .............. .Order Rodentia, 23 Incisor teeth range in number from none to five upper and none to four lower pairs, but are never a single pair as de- scribed above ...... ..... 2 2(1) Front limbs modified as wings; bones of hand and fingers greatly elongated; wing membranes connect fingers with each other and with body (fig. 2); bones of skull relatively thin and fused so that few sutures are visible; skull shorter that 54 mm . . Order Chiroptera, 34 Front limbs not wings; forelimbs not as described; bones of skull relatively heavy and many sutures visible; skull in many species longer than 54 ...... 3 3(2) Ten upper incisors and eight lower inci- sors (fig. 3); first hind toe projecting lat- erally (in one species connected by web) and without claw or nail (figs. 4, 97, and 98) .... Order Marsupialia, Family Didelphidae, Key 2 22 NO. 231 ANDERSON: MAMMALS OF BOLIVIA Fewer incisors above and below; first hind toe present or not; if present, somewhat opposable or not (usually not), and with claw or nail ... ...... 4 4(3) Front limbs are paddle like; no hind limbs; distally, tail horizontally flattened; ex- clusively aquatic ......5..... Not as above ... ........ 6 5(4) Tail with rounded margin (fig. 5); anterior grinding teeth, usually noticeably more worn than posterior ones; teeth with cusps or lophs and not simple and con- ical; skull heavy, bones dense, zygo- matic arch massive (fig. 7); possibly reach extreme northern Bolivia in the Abuna River ..................... *Order Sirenia, Family Trichechidae, Trichechus inunguis Tail with two lateral flukes (fig. 6); teeth numerous, mostly simple and terminally more or less conical; skull with thin bones, zygomatic arch incomplete (fig. 8), snout long and slender .... Order Odontoceti, Family Platanistidae, Inia geoffrensis 6(4) Teeth absent or simple, subcylindrical, and without enamel, no incisors present .............. Order Xenarthra, 7 Teeth present and not subcylindrical, enamel present, incisors present or ab- sent .......... 10 [Order Xenarthra begins here] 7(6) No teeth whatsoever; mouth relatively small, tongue very long and vermiform; eats ants and termites .............. . . Family Myrmecophagidae, Key 11 Teeth present; mouth of moderate size, tongue not extremely long and vermi- form; food not exclusively ants and ter- mites .... ......... 8 8(7) Dorsal surface well haired and without ar- mor of plates; limbs relatively long; claws long, laterally compressed and curved for hanging from branches; movements slow and deliberate; climbs trees; eats leaves ........... 9 Dorsal surface in most species without much hair and with a bony carapace or armor of plates; limbs relatively short and stout; claws heavily built for digging (fig. 9); movement sometimes rapid; ground dweller, burrower, eats insects ... . . . . F a m il y D a s y p o d id a e , K e y 1 2 9(8) Anterior upper tooth smaller than next tooth and space between them less than length of crown of second tooth; alve- olar length of upper tooth row less than 30 mm; three claws on forefoot; tail not rudimentary (but it is short, less than one-tenth of length of head and body); hair relatively short and crisp; face whitish with brown circumorbital areas ............ .Famil y Bradypodidae, Bradypus variegatus Anterior upper tooth larger than next tooth and space between them greater than length of second tooth; alveolar length of upper tooth row more than 30; two claws on forefoot; tail rudimentary; hair long; face brownish ........ Family Choloepidae, Choloepus hoffmanni 10(6) One or more toes of each foot have a con- spicuous hoof or (in camelids) foot with two large padlike toes, each with a nail- like hoof at the front; large gap (fig. 10A) between grinding teeth and inci- sors at front of lower jaw and length of jaw from articular process (fig. 10) more than 85 mm .............. 11 No toe has a conspicuous hoof (but toes usually have claws or nails), never with only two conspicuous toes; no large di- astema in lower tooth row, or, if so, then jaw shorter than 85 ............ 17 11(10) Main axis of foot lies in middle of central 1997 23 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 7~~~~~~~ 18 19t7 22 4S 24 25 30 v 31 33 I4G, toe that is larger than other toes; upper incisor teeth present; skull as shown in figs. 16 or 17 .......... Order Perissodactyla, 12 Main axis of foot lies between two toes of about equal size; some species have no upper incisors; skull as shown in figs. 20, 21, or 26-29 ............ ........... Order Artiodactyla, 13 [Order Perissodactyla begins here] 12(11) A single large rounded hoof on each foot (fig. 11); nose not a flexible proboscis although lip is somewhat flexible (fig. 13); nasal bones relatively long anteri- orly and nasal opening on skull not un- usually displaced posteriorly (fig. 16); domesticated ...... Family Equidae More than one conspicuous hoof on each foot (four front, three rear, fig. 12); nose elongated (fig. 14); nasal opening of skull displaced posteriorly (fig. 17) ... Family Tapiridae, Tapirus terrestris [Order Artiodactyla begins here] 13(11) Snout piglike, flattened, with rim, and somewhat rounded (fig. 15); canine teeth large, upper incisors present and not reduced to a single lateral pair in adults ..... 14 Snout not as above; canine teeth absent or relatively small, upper incisors absent or reduced in adults to a single lateral pair ..... 15 14(13) Four toes on hind feet (fig. 18); upper ca- nines curving outward, three pairs of upper incisors (fig. 20), lower third mo- lar longer than 35 mm, first lower pre- molar sometimes present; tail small but obvious and somewhat curled; domes- ticated ... Family Suidae, Sus scrofa Two or three (fig. 19) toes on hind feet; upper canines directed downward (fig. 21), two pairs of upper incisors, m3 considerably shorter than 35, pl never present; tail scarcely evident externally ....... .Family Tayassuidae, Key 17 15(13) No horns or antlers at any time; hind part of body contracted, knee-joint low, thigh appears distinct from body (fig. 22); metapodials of cannon bone in each leg somewhat separated distally (fig. 24), their articular surfaces with limited keel, but not strongly grooved or pulley-shaped; cutaneous pad on each of two toes; nails on upper sur- faces (rather than hoof encasing the last bone of each toe); two pairs of canini- form teeth in upper jaw (one a pair of incisors, the other a pair of canines), usually a pair of canines below, canines larger and more hooked in males, three pairs of lower incisiform teeth; deep medial posterior notch of palate extends considerably farther forward than lat- eral notches do (fig. 26) ........... ........ .Family Camelidae, Key 18 Horns or antlers may be present in one or both sexes, continuously or at certain times of the year; thigh not so distinctly separate from body (fig. 23); cannon bone with little distal separation of two metapodials (fig. 25); articular surfaces strongly grooved or pulley-shaped; ca- niniform teeth absent, four pairs of low- er incisiform teeth (lateral pair actually canines), anterior end of medial poste- rior notch of palate even with or pos- terior to lateral notches (fig. 27) . . 16 16(15) Vacuity conspicuous between lacrimal and nasal bones (fig. 28); molariform teeth brachydont (relatively low- crowned, fig. 30, young animal, unworn tooth); antlers of bony material and 24 NO. 231 ANDERSON: MAMMALS OF BOLIVIA yot3 4 36X536~~ 7 ,35 3 9.- -(~;4IB~ 44 "S / , 1 4 1<-- ; 447 X42 X r- ~4 4 9 with no permanent covering, present in males at least part of year, sometimes branched . . Family Cervidae, Key 19 Vacuity inconspicuous or not present (fig. 29); molariform teeth more hypsodont (high-crowned, fig. 31, young animal, unworn tooth); horns (with bony core and covering of horn), if present, not shed at times, never branched, in some species present in both sexes; domestic animals ... Family Bovidae, Key 20 17(10) Canine teeth absent, major gap (diastema) between incisors and grinding teeth at back of jaws (fig. 32); soles of feet cov- ered with hair (fig. 33) ....... Order Lagomorpha, Family Leporidae, 21 Canine teeth present and often noticeably larger than adjacent incisors and pre- molars, no large diastemal gap; soles of feet not covered with hair ......... 18 18(17) Total number of upper and lower incisors: 12 (3 pairs above, 3 pairs below) and outer upper incisors (13) larger than oth- er pairs (I1 or 12, fig. 34); neither pair of limbs especially elongated (except in Chrysocyon) . . Order Carnivora, 19 Total number of upper and lower incisors: 8 (2 pairs above, 2 pairs below) and 12 smaller than I1 (fig. 35); one pair or both pairs of limbs noticeably elongated .............. .Orde r Primates, 40 [Order Carnivora begins here] 19(18) Molariform teeth relatively low and rounded (fig. 36), none of them with sharp bladelike edge ........... 20 Some molariform teeth bladelike, specifi- cally in adults, the last upper premolar (P4, fig. 37) and first lower molar (ml); in young the carnassial teeth are P3 and p4 .... .......... 21 20(19) Tail very short, scarcely noticeable; large (head and body about 1.2 to 1.4 m; fig. 38) ........................... Family Ursidae, Tremarctos ornatus Tail from about 45% of head and body length to 1.3 times that length; small (head and body length about 400 mm) to medium (780 mm) ............. ....... .Famil y Procyonidae, Key 14 21(19) Color a pattern of dark spots on paler background (in five species) or relative- ly uniformly brownish or reddish dor- sally (in two species); claws sharp and retractable (most noticeably on front feet, fig. 39, A retracted, B extended) ... . . . . . . . . F a m i l y F e l i d a e , K e y 1 6 Color never a pattern of dark spots on pal- er background; claws duller and not re- tractable ... ..... 22 22(21) Relatively long legs, height at shoulders usually more than 40% of length of head and body (fig. 40), or if not, then tail less than 40% of length of head and body (fig. 41); ears often relatively large and erect; facial part of head rel- atively longer, eye usually about mid- way between end of nose and ear open- ing; on skull, distance between orbit and anterior nares (fig. 43A) more than interorbital breadth (fig. 43B); braincase not extending noticeably behind ears (fig. 44) .. Family Canidae, Key 13 Relatively shorter legs (percentage noted above usually less than 40, or if not, then tail is more than 40% of length of head and body, fig. 42); ears relatively shorter and more rounded; facial part of head relatively shorter, eye usually nearer to end of nose than to ear open- ing; on skull, distance between orbit and anterior nares less than interorbital breadth (fig. 45); braincase extending noticeably behind ears (fig. 46) ...... ........ .Famil y M ustelidae, Key 15 25 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY loo 57 [Order Rodentia begins here] 23(1) Infraorbital canal not conspicuously en- larged (figs. 48, 49); tail present, long or short but clearly visible; ears tend to be rounded; head and body length never more than 320 mm ............. 24 Infraorbital canal conspicuously enlarged (fig. 47); tail absent or present in vari- ous lengths; ears tend to have a slight posterior concavity of the margin; head and body in some species longer than 320 . .................. 25 24(23) Tail always long and well haired, not scaly; arboreal and diurnal; infraorbital canal small (fig. 49); more than three teeth in upper molariform row (fig. 50) ......... Family Sciuridae, Key 22 Tail of various lengths but never covered with long hair (usually with some short hair), scales clearly visible; most spe- cies not especially arboreal, most noc- turnal; infraorbital canal not so small (fig. 48); only three teeth in upper mo- lariform row (fig. 51) ............. .......... Family Muridae, Key 23 25(23) Tail not visible or scarcely visible, less than 10% of length of head and body; three toes on hind feet large, first and fifth digits much reduced ........ 26 Tail clearly visible (more than 10% of length of head and body; more than three large toes on hind foot, first and fifth digits not disproportionally small ............................. .2 9 26(25) Head and body longer than 510 mm; hind foot longer than 100; skull longer than 100; greatest distance across upper mo- lariform teeth more than 16 ...... 27 Head and body shorter than 510; hind foot shorter than 100; skull shorter than 100; greatest distance across first upper mo- lariform teeth less than 16 mm ...... .......... .Fam ily Cavildae, Key 27 27(26) Head and body longer than 900 mm, skull longer than 200; last upper tooth larger than other three molariform teeth together (fig. 52) .... Family Hydrochaeridae, Hydrochaeris hydrochaeris Head and body shorter than 900; skull shorter than 200; last upper tooth small- er than other three molariform teeth to- gether (figs. 53, 54) ............ 28 28(27) Jugal and part of maxillary expanded, forming a thick plate with rough sur- face; skull relatively broad (fig. 53); pelage brown and with rows of whitish or yellowish spots or lines; five toes on hind foot ...................... ..... .Family Agoutidae, Agouti paca Zygomatic arch normally developed, no large plate, skull relatively slender (fig. 54); pelage blackish, without rows of spots and lines; three toes on hind foot ... . . . . ... . . F a m ily D a s y p r o c tid a e , Dasyprocta and Myoprocta 29(25) Tail relatively well haired throughout its length and with dorsal crest of coarser, longer hairs (many more than 30 mm) toward end of tail; feet and ventral side of tail never white; upper cheek teeth persistently growing and with tightly pressed transverse laminae (fig. 55) . . ...... .Fam ily Chinchillidae, Key 26 Tail in most species scaled and with few hairs (or, in Octodontomys, longer haired distally but with few hairs longer than 30; feet, ventral side of tail white; head and body length about 170, thus smaller than any chinchillid ...... 30 30(29) Large, head and body longer than 350 mm; skull longer than 64 ........ 31 Smaller, head and body shorter than 350; skull shorter than 64 mm ........ 33 31(30) Pelage includes heavy sharp quills; skull bulges upward between eyes (fig. 56); tail prehensile, with special hairless area on top near end (fig. 57) .......... ... . . F a m ily E r e th i z o n tid a e , K e y 2 8 26 NO. 231 ANDERSON: MAMMALS OF BOLIVIA 62 63 ~A 66 6 64 Pelage without quills; skull relatively flat on top; tail not prehensile; if haired, not with special hairless area at end . . 32 32(31) Brown, not spotted; tail conspicuously scaled, more than half as long as head plus body; five hind toes, webbed, ex- cept the fifth toe (fig. 58) ... Family Myocastoridae, Myocastor coypus Brown or black, with whitish spots and stripes (fig. 59); scales of tail concealed by hair, tail less than half the length of head and body although longer than hind foot; four toes on each foot, un- webbed . .... Family Dinomyidae, Dinomys branickii 33(30) Upper cheek teeth relatively simple, shaped like figure eight, or a rounded "L," enamel around edges of flattened occlusal surface only (fig. 60) ..... ........ Families Abrocomidae and Octodontidae, Key 25 Upper cheek teeth more complex, not as above, with enamel folds and isolated enamel islands in middle of occlusal surface (fig. 61B; fig. 61A shows skull of Proechimys) and rooted (not persis- tently growing) .................. ....... Family Echimyidae, Key 24 [Key to Chiroptera begins here] 34(2) With distinct nose leaf (fig. 62; reduced in Sphaeronycteris [fig. 160] and in vam- pire bats [Desmodontinae], which have large, sharp-edged, pointed, cutting up- per incisors [fig. 64]) ............. ... . . F a m ily P h y llo s t o m id a e , K e y 6 Without nose leaf (fig. 63) and with upper incisors not as described for vampires ............................. .3 5 35(34) Tail relatively heavy and extending no- ticeably beyond edge of interfemoral membrane (fig. 65); membranes thick; legs relatively short; feet heavy, hairs extend beyond claws of toes, no suction discs on feet .................... ... . . . . . . F a m i l y M o l o s s i d a e , K e y 9 Tail absent or present but not as above (although it may project above surface of interfemoral membrane or slightly beyond its edge, the latter in Thyrop- tera,which have suction discs on feet) ........ . .................... 36 36(35) Gap between premaxillary bones at front of skull (fig. 66), incisors of right and left well separated; no discs on base of thumb or sole of hind foot ...... ....... 37 No distinct gap between premaxillary bones (fig. 67), incisors relatively close together at midline; may or may not have discs on thumb and hind foot . . . ............................. .3 8 37(36) Pocket may be present in leading wing membrane (fig. 69); sometimes skull has distinct postorbital process (fig. 68 arrow); hind leg slender; end of tail pro- jecting above upper surface of middle of interfemoral membrane or tail mostly ventral to this membrane .......... ... . . F a m ily E m b a llo n u r id a e , K e y 3 No pocket as described; never has distinct postorbital process; hind leg moderate in build; tail reaching to edge of wide interfemoral membrane ........... ... . . F a m ily V e s p e r t ilio n id a e , K e y 8 38(36) Distinct suction discs on base of thumb (fig. 70) and sole of hind foot; (in the two species now known from Bolivia) size small, head and body shorter than 75 mm ........................ ...... .Famil y Thyropteridae, Key 7 No suction discs; size medium, head and body longer than 75 ............ 39 39(38) Hind feet enlarged (longer than 19 mm); lips full (fig. 71); skull relatively broad (width more than half of length) ..... ........ .Famil y Noctilionidae, Key 4 27 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Hind feet shorter than 19; lips not full (fig. 72); skull relatively narrow (width less than half of length) .............. ...... Family Mormoopidae, Key S [Order Primates begin here] 40(18) Large; bipedal; tailless; only two upper and two lower premolars on each side (fig. 73, shows upper teeth) ....... ... Family Hominidae, Homo sapiens Small to medium (up to 675 mm in length of head and body); tail conspicuous; three upper and three lower premolars on each side (fig. 74, shows upper teeth) ........ Families Callitrichidae and Cebidae, Key 9 KEY 2 GENERA AND SPECIES OF DIDELPHIDAE IN BOLIVIA I Tail length ranges from nearly as long as to much longer than head and body; size varies, large or small ........ 6 Tail about half of length of head and body; skull always shorter than 45 mm; hind foot shorter than 30 mm; head and body always shorter than 200 mm ... .................. Monodelphis, 2 [Monodelphis is poorly known; for ad- ditional notes see text] 2(1) Head and body longer than 100 mm; skull longer than 30 mm; hind foot longer than 17 ......... ....... 3 Head and body shorter than 100; skull shorter than 30; hind foot shorter than 17 .... ........... 5 3(2) Tail furred ...... Monodelphis emiliae Tail not furred throughout its length . . 4 4(3) Lateral pelage rusty and contrasting sharply with grayish dorsal pelage; hind foot about 21 mm long ........... ......... Monodelphis brevicaudata Lateral pelage not rusty but gray or yel- lowish and blending into mid dorsal pelage color; hind foot about 19 ..... ............ Monodelphis domestica 5(2) Rostrum slender (fig. 75); larger, skull longer than 23.5 in adults, head and body longer than 75 .............. ... Monodelphis adusta or M. osgoodi Rostrum less slender (fig. 76); smaller, skull shorter than 23.5; head and body shorter than 75 . . Monodelphis kunsi 6(1) Fur intermixed with long (more than 20 mm) coarse guard hairs projecting be- 85 / t >l) 8 7 86 yond the woollier, softer underhairs; larger as adults, skull longer than 83 ... . . . . . . . . . . . . . . . ... D i d e lp h i s , 7 Fur not as above (few if any hairs longer than 20); smaller as adults, skull shorter than 83 ................... 8 7(6) Ears usually white, at least at edges; face with contrasting blackish (crown, band through each eye) and whitish areas (fig. 77); skull usually more slender for a given age and sex (fig. 79, adult male) ... . . . . . . . . . . . D id e lp h is a lb iv e n tr is Ears usually black; face without much contrast between darker and lighter ar- eas (fig. 78); skull less slender (fig. 80, adult male) ... Didelphis marsupialis 8(6) Webs between toes of hind feet; fur of back with striking pattern of dark mid- line stripe and saddle like blotches; skull relatively broad (fig. 81) ..... ... ... . . . .... C h ir o n e c te s m in im u s No webs between toes of hind foot; fur of back not as above, more or less uniform in color; skull relatively narrow (figs. 82, 83) ... ........ 9 9(8) Distinct pale spot in blackish surround- ings above each eye ............ 10 Not as above ................ 11 10(9) Pelage dark gray or blackish and extend- ing for about 50 to 75 mm on the tail; 28 NO. 231 ANDERSON: MAMMALS OF BOLIVIA bottom of tail near base fully haired; distinct supraorbital processes (fig. 82) ............... Philander opossum Pelage usually brownish and extending less than 30 on the tail; tail essentially naked below from anus to tip (tail rel- atively longer and more slender and feet more slender, but direct comparison needed to visualize these); no distinct supraorbital processes (fig. 83) ...... .......... Metachirus nudicaudatus 11(9) Middorsal streak present on head from muzzle to between ears, dark with paler areas on either side, or pale with dark areas on either side ............ 12 No distinct facial streak ........... 14 12(11) Middorsal facial streak dark; tail not haired for entire length; head and body of adults longer than 210 mm; hind foot about 40 in adults; supraorbital process- es large and projecting (fig. 84) ..... ................... Caluromys 13 Middorsal facial streak grayish; tail densely haired for its entire length; head and body shorter than 210; hind foot about 30; supraorbital processes in form of conspicuous shelf over each orbit (fig. 85) ......... Glironia venusta 13(12) More than half of length of dorsal surface of tail well haired ................ ............... Caluromys lanatus Less than half (about one-third) of length of dorsal surface of tail well haired . . ............. *Caluromys philander 14(11) Ears short, rounded, barely projecting above fur; ears laid forward, reaching barely half the distance to the eyes (fig. 86); eyes relatively far forward on head; snout short . Lutreolina crassicaudata Ears longer, clearly projecting beyond fur; ears laid forward, reaching beyond half the distance to the eyes (fig. 87) (the following key to "Marmosa," now di- vided into five genera, modified from that of Tate, 1933) ............. 15 15(14) Scales on tail annular and posterior mar- gin of each scale; scale nearly straight (fig. 88); mammae both pectoral and ab- dominal; bullae relatively large (fig. 90A); postpalatal bridge narrow (fig. 90B), palate highly fenestrated, lateral foramina large (fig. 90C); pattern of pelage usually tricolored, sides demar- cated relatively sharply from both back and belly; tail short, thickened with fat in some seasons; ears relatively larger (17 to 26 mm long); feet relatively smaller (14 to 17), ear is usually more 91 B 94 a G m 95 than 1.35 times the length of hind foot .................... .Thylamy s, 16 Scales arranged spirally and are generally rhomboidal (fig. 89); mammae abdom- inal; bullae smaller (fig. 91A); postpal- atal bridge broader (fig. 91B); palate usually less fenestrated (fig. 91C); tri- color pattern not developed; tail vari- able in length, never incrassated; ear length usually less than 1.35 times length of hind foot ............. 18 16(15) Dorsally grayish; bullae especially large (fig. 90A) ........... .... 17 Dorsally brownish; bullae not so large (fig. 92) ....... Thylamys venustus 17(16) Braincase relatively larger; interorbital re- gion not so narrow (fig. 93) ...... ............. .. Thylamys pallidior Braincase relatively small; interorbital re- gion narrow (fig. 94) ............. ... . . . . . . . . . . . . . . T h y l a m y s p u s i lla 18(15) Large to moderate in size (head and body 150 to 180 mm in length, skull longer than 38); fur grayish, often rather wool- ly; feet (hind foot length 24 to 29) and claws large and strong; external anterior and posterior pads of hind foot united (fig. 97); tail long (190 to 220); scales spiral, 9 to 14 rows per cm of tail 1997 29 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY length; skull strongly ossified; bullae small (2.9 to 3.3 mm across); postpal- atal bridge broad (4.2 to 4.6); greatest width across bullae (fig. 99B) more than 2.13 times inclusive distance across bulla and petrosal bone (fig. 99A); teeth large (maxillary tooth row, MI through M3, 7.0 or more) .... Micoureus, 19 Small to moderate in size; fur rarely gray, never woolly; feet and claws smaller; pads separate (fig. 98); tail seldom very long; scales more than 16 per cm; skull strongly to slightly ossified; bullae larg- er; bridge narrower; ratio lower; teeth smaller ..... ........ 20 19(18) Pelage long (longest dorsal hairs about 15 mm), somewhat waved, brownish gray, most ventral hair basally gray ....... ................ Micoureus regina Pelage shorter (longest dorsal hairs about 12 or 13 mm), straighter, less dark brownish, ventrally ochraceous, with few gray-based hairs ............. ............ Micoureus constantiae 20(18) Feet relatively small, never very broad; pad at base of digits 3 and 4 of hind foot considerably smaller than that of digits 2 and 3; the three minute hairs accompanying each caudal scale flat- tened, appressed, provided with a me- dian dorsal keel, somewhat petiolate at the insertion, often black; skull nar- rowed; nasals exceeding premaxillae anteriorly, base of nasala at maxillo- frontal suture very little or not at all ex- panded; no pointed supraorbital pro- cesses; teeth often small ......... .................. Marmosops, 21 Feet proportionally larger, or if short, then rather broad; pads at base of digits 2 and 3 and of 3 and 4 subequal; caudal scale hairs not as described above, rare- ly black; skull usually broader; nasals never exceeding premaxillae anteriorly; base of nasals at maxillofrontal suture abruptly expanded; pointed supraorbital processes present although sometimes small; teeth proportionally larger . . 23 21(20) Size small (total length less than 215 mm, generally 180 to 190) ............. Marmosops impavida or M. parvidens Size moderate to large; teeth larger (Ml- M3 more than 6.1) ............. 22 22(21) Large; fuscous, reddish, or brown, never grayish; tail long, brownish; ventral white area sometimes narrowed by gray-based hairs; ears small to medium; skull with supraorbital beading (fig. 100) ........ Marmosops noctivaga Moderate in size; brownish gray; hairs long, coarse, lusterless; white area of underparts not narrowed by lateral or gray-based hairs, white area about 30 mm wide; ears large; no supraorbital beading (fig. 101) ................ .............. .Marmosop s dorothea 23(20) Relatively unfenestrated palate (fig. 95); mammae abdominal ... Marmosa, 24 Fenestrae large (fig. 96); both pectoral and abdominal mammae present ........ ................ .Gracilinanus , 25 24(23) Larger, head and body longer than 120 mm; M1-M3 longer than 5.3 ..... ................ .Marmos a murina Smaller, head and body shorter than 120; M1-M3 shorter than 5.3 .......... .................. .Marmos a lepida 25(23) Brownish gray above, buffy white below; dorsal surface of shank same color as hind foot ...... Gracilinanus agilis Deep cinnamon color above, cinnamon washed below; dorsal surface of shank deep fuscous, hind foot buffy brown; guard hairs numerous ............. ......... .Gracilinanu s aceramarcae KEY 3 GENERA AND SPECIES OF EMBALLONURIDAE IN BOLIVIA The sac-winged bats or Emballonuridae are relatively small bats (forearm up to 66 mm long) without special outgrowths on muzzle. The second digit of wing has no phalanges and the third has two phalanges, of which the proximal is flexed upon dorsal surface of metacarpal when at rest. The tail projects above the uropatagium or interfe- moral membrane, not from its free edge. The premaxillae are incomplete and the first up- per incisors widely separated. An opening is often present on the propatagium near the el- bow, from which an odoriferous substance is secreted. 1 Postorbital processes obscured by wide su- praorbital ridges (fig. 102); wing sac, if present, opening on ventral surface of wing; color white; fur long and silky; thumb very short and mostly contained in membrane, first phalanx shorter than met- acarpal, claw rudimentary; distinct pouch- es on lower side of uropatagium (fig. 103); 30 NO. 231 ANDERSON: MAMMALS OF BOLIVIA 102 10 3 ( < s S-~~Y,s, S,W 104 tail mostly ventral to this membrane .... ........... *Subfamily Diclidurinae, 2 Postorbital processes long and curved (often broken off in cleaning skulls, fig. 104); sac, if present, opening on dorsal surface of wing; color not white; fur not long and silky; thumb not mostly contained in membrane, its first phalanx about as long as metacarpal, its claw not rudimentary; no pouches on lower side of uropatagium; tail lies within uropatagium for most of the tail's length ................... ........ Subfamily Emballonurinae, 3 2(1) Forearm 60 to 69 mm long; maxillary tooth row length 7.5 to 8.4 .............. ........ ......... *Diclidurus albus Forearm 70 to 73; maxillary tooth row length 9.4 to 9.6 ... *Diclidurus ingens 3(1) Calcar much longer than tibia (fig. 105), about half the length of forearm; wing sac absent; small tufts of whitish fur along the forearm (fig. 106), dorsal pelage grizzled (silvery tips of hairs on darker background) ..... ................ Rhynchonycters naso Calcar equal to tibia or shorter; wing sac usually present; no tufts of whitish hair along forearm; pelage not grizzled as not- ed ............. 4 4(3) Wing membrane attaches to foot near base of outer toe (fig. 107); mesopterygoid fossa broad in front, a median projection at back of palate .................. ... . . . . . ... . . . . * C o r m u r a b r e v ir o s tr is Wing membrane attaches to foot near ankle (fig. 108); mesopterygoid fossa narrowed in front, no medianposterior palatal pro- jection ............. 5 5(4) Dorsal surface with two more or less distinct wavy longitudinal pale lines on darker background (fig. 109); wing sac close to forearm near elbow; upper surface of ros- trum of skull with a slight median groove ... . . . . . . . . . . . . . ... . . S a cc o p t e r y x , 6 Dorsal lines absent; wing sac small and near anterior border of antebrachial membrane; upper part of rostrum inflated, convex, no median groove ......... Peropteryx, 7 6(5) Dorsally black (or in worn pelage brownish); wing membranes black; wing sac, at least in males, remarkably large; forearm lon- ger than 43 mm; length of upper tooth row (C-M3) more than 6.7 .............. ... . . . . . . .... . . S a c c o p te ry x b ilin e a ta Dorsally brown; wing sac not large; forearm shorter than 43; tooth row shorter than 6.7 ... . ... . . . . . . . . . . S a c c o p te r y x le p tu r a 7(5) Forearm usually longer than 45 mm; skull longer than 16; upper tooth row (C-M3) longer than 6.5; width across molars great- er than 6.9 ....... Peropteryx kappleri Forearm usually shorter than 45; skull short- er than 16; tooth row shorter than 6.5; width across molars less than 6.9 .... ... . .. . . . . . . . . . . P e r o p te r y x m a c r o tis KEY 4 SPECIES OF NOCTILIONIDAE IN BOLIVIA Enlarged hind limbs enable Noctilionidae or bulldog bats to catch and eat insects and small fish. The hind feet are more than half the length of the tibia. This family has only one genus and two species. Upper lips are large and form a hood over the mouth. Ears are long and pointed. Fur is short and often brightly colored. Wing membranes are at- tached to the back somewhat above the usual lateral position. Skull is broad and has a sag- ittal crest. 1 Forearm longer than 70 mm; combined length of tibia and hind foot relatively longer, over 70% of the length of forearm; hind foot lon- ger than 30 ....... Noctilio leporinus Forearm shorter than 70; length of tibia and 31 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY hind foot less than 70% of length of forearm; hind foot shorter than 30 ............. ................... Noctilio albiventris KEY 5 SPECIES OF MORMOOPIDAE IN BOLIVIA This family was separated from the Phyl- lostomidae by Smith (1972). Lower lip with platelike outgrowths. 1 Wing membranes originating on middorsal line (fig. 110), giving a naked-backed ap- pearance; rostral breadth (fig. 111) always greater than length of maxillary tooth row (fig. 111) .............. Subgenus Pteronotus, Pteronotus gymnonotus Wing membranes not originating on mid- dorsal line but from more lateral position; rostral breadth equal to or less than length of maxillary tooth row ............. 2 2(1) Basioccipital narrowly constricted between au- ditory bullae (fig. 112); condylobasal length more than 16 mm; forearm longer than 50; tympanic ring covering about one-third of auditory bulla (fig. 112) .... Subgenus Phyllodia, Pteronotus parnellii Basioccipital not narrowly constricted between auditory bullae (fig. 113); condylobasal length less than 16; forearm shorter than 50; tympanic ring covering more than half of auditory bulla (fig. 113) ............. ............ Subgenus Chilonycteris, Pteronotus personatus KEY 6 GENERA AND SPECIES OF PHYLLOSTOMIDAE IN BOLIVIA The American leaf-nosed bats of the fam- ily Phyllostomidae usually have a vertically projecting flaplike nose leaf (reduced in some genera). They lack the postorbital pro- cesses seen in most emballonurid skulls and the chin flaps of the mormoopids. The pre- maxillae are complete and fused to each oth- er and to the maxillae,and their palatal branches isolate two lateral foramina. The second finger has a well-developed metacar- pal and a small phalanx e third finger has three bony phalanges. 1 Upper incisors are large, pointed, sharp- edged bladelike teeth (fig. 64); postcanine teeth reduced in number to two or three 127 pairs above and below and all reduced in size, smaller than the upper incisor or ca- nine; nose leaf reduced to dermal ridges above nostrils ..................... ........ .Subfamily Desmodontinae, 36 Upper incisors not as above; postcanine teeth not both fewer than four in a tooth row and all smaller than any upper canine or incisor ..... 2 2(1) Tongue very long (fig. 114); upper surface of lower lip in the center divided by deep groove; head long and narrow (width less than 46% of length); cusps and commis- sures of upper molars often so reduced that the W-pattern is absent ........ 29 Tongue not unusually long; upper surface of lower lip not divided by a deep groove; head width greater than 46% of head length; cusps and commissures of upper molars variable, often with W-pattern ev- ident ......................... 3 3(2) Interfemoral membrane reduced to a very narrow band along legs and posterior part of body and covered with fine prominent hair; calcar indistinct or absent; no tail ev- ident (fig. 115); crown of molars with dis- tinct longitudinal groove . . Sturnira, 61 Interfemoral membrane not as above, but moderately or well developed at least lat- 32 NO. 231 ANDERSON: MAMMALS OF BOLIVIA erally; calcar distinct; tail present or ab- sent; crown of molars not as described above ... 4 4(3) Muzzle short and broad; no external tail; tooth rows arcuate (fig. 116) ........ ...... Subfamily Stenodermatinae, 38 Muzzle not so short and broad; tail distinct; tooth rows not distinctly arcuate (fig. 117) ............. 5.................. S 5(4) Zygomatic arches incomplete (fig. 118); up- per molars narrow and without W-pattern; two lower premolars; chin with large rounded protuberance in the center sur- rounded by two or more bumps (several smaller ones in Carollia or a larger one on either side in Rhinophylla).......... ............ Subfamily Carolliinae, 6 Zygomatic arches complete (fig. 119); upper molars with W-pattern; chin without large rounded protuberance in center ...... ......... Subfamily Phyllostominae, 9 6(5) First upper premolar much smaller than the second (fig. 120); third upper molar (M3) oval and mostly horizontal, with reduced cusps (fig. 120); teeth generally simplified; tail absent; tips of dorsal hairs slightly darkened; size small, forearm shorter than 37.5 mm ..... Rhinophylla pumilio First upper premolar approximately the same size as the second (fig. 121); M3 not oval and with more distinct cusps (fig. 121); tail present; tips of dorsal hairs distinctly darker than more proximal parts of hairs, either reddish or dark gray; forearm longer than 33.5 ........................ .... ..................... Carollia, 7 7(6) Lower jaw viewed from above has outer in- cisors obscured by cingula of canines (fig. 122) and/or upper tooth row straight (fig. 124); forearm longer than 41.3; condylo- basal length more than 20 .......... .... ............. Carollia perspicillata Lower jaw viewed from above has outer in- cisors visible (fig. 123); upper tooth row bowed in or with distinct notch or "step" in labial outline (figs. 125, 126; forearm length less than 41.8; condylobasal shorter than 20 .... ......... 8 8(7) Labial outline of upper tooth row has distinct notch or step, owing to labial side of sec- ond premolar being more lingual than la- bial edge of first molar (fig. 125); second lower premolar (p2) about twice as high as first molar, occlusal surface of first low- er molar (ml) with straight profile (fig. 127); forearm shorter than 37.5 mm; con- dylobasal length less than 18 ........ .................. Carollia castanea Labial outline of upper tooth row evenly curved (fig. 126); p2 not about twice as high as ml, surface of ml with discrete cusp or cusps (fig. 128); forearm longer than 37.5; condylobasal length more than 18 .............. Carollia brevicauda [Phyllostominae begin here] 9(5) External tail absent or rudimentary; fore- arm longer than 75 mm ......... 10 Tail present; forearm usually shorter than 75 mm . ................. 11 10(9) Two lower incisors; tail rudimentary; forearm 77 to 83 mm long ........ .. ........... Chrotopterus auritus Four lower incisors; no tail; forearm about 105 .......... Vampyrum spectrum 11(9) Tail in membrane and extending to its bor- der (fig. 129) ......... ... 12 Tail short and not reaching beyond the middle of the membrane (fig. 130; in some species end of tail projects above the membrane) ............ 13 12(11) Forearm shorter than 43 mm (usually 34 to 39); distal part of uropatagium with peculiar dermal papillae in longitudinal rows, most evident ventrally (fig. 131); wing membrane from distal half of tib- ia; calcar about same length as tibia; nose leaf less than 15 high ....... ... . . . M a c ro p h y llu m m a c ro p h y llu m Forearm longer than 43 (usually 47 to 52); uropatagium without peculiar papillae; wing membrane from ankle opposite calcar; calcar about two-thirds as long as tibia; nose leaf more than 15 high (fig. 132) ...... Lonchorhina aurita 13(11) Two lower incisors (i.e., one pair) . . 14 Four lower incisors .............. 18 14(13) Nose leaf more than half as long as ear ... . . . . . . . . . . . . M im o n c r e n u la t u m Nose leaf less than half as long as ear . . ... ..... .... . . . . T o n a t i a , 1 5 15(14) Forearm shorter than 40 mm; condylobas- al length of skull less than 19 ..... ... . . ... ... .. . . T o n a t i a b r a s i l i e n s e Forearm longer than 40; condylobasal length of skull more than 19 ..... 16 16(15) Forearm shorter than 49 mm; condylobas- al length of skull less than 22 ..... ... . . . . . . . . ... . . . T o n a t i a c a r r i k e r i Forearm longer than 49; condylobasal length of skull more than 22 ..... 17 17(16) Ears connected by a low, but often indis- tinct, band; ear length 33 mm or more (measured from basal notch to tip); postorbital area of skull very constrict- 33 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 136 141 '4 142 145 1 4 6iI~f\ 46 WBr 149 ed, much narrower than width of ros- trum just above base of canines (fig. 133); forearm nearly naked ....... ................. Tonatia sylvicola Ears completely separate; ear length 32 or less; postorbital constriction poorly de- veloped, about the same as the width of the rostrum just above the base of the canines (fig. 134); forearm furry ..... ................... Tonatia bidens 18(13) Forearm shorter than 60 mm .......... ~~~~Micronycteris, 19 . . . . . . . . . . . . . . . . . ........ irntes,1 Forearm longer than 60 ........... 25 19(18) Ears connected by high, notched, often in- distinct band, P3 about same size as P4 ............................. 20 Ears connected by low, unnotched band or band absent, P3 and P4 not equal in size ............................. 23 20(19) Forearm longer than 40 mm ....... 21 Forearm shorter than 40 ............ ........ Subgenus Micronycteris, 22 21(20) Two pairs of upper incisors ......... *Micronycteris (Xenoctenes) hirsuta One pair of upper incisors .......... ... Micronycteis (Barficonycters) daviesi 22(20) Interauricular band slightly notched me- dially (fig. 135); venter brown; calcar distinctly longer than hind foot; length of interfemoral membrane more than twice length of tail; upper premolars (P3, P4) about equal in height ..... . ... . . . . . M ic r o n y c te ris m e g a lo tis Interauricular band deeply notched medi- ally (fig. 136); venter gray-white; calcar shorter than hind foot; length of uro- patagium less than twice length of tail; P3 distinctly lower than P4 ......... ... . . . . . . . . . . M ic r o n y c te ris m in u ta 23(19) Third metacarpal longest ............ ........ . .....Micronycteris nicefori Fifth metacarpal longest .......... 24 24(23) Forearm longer than 42.5 mm ....... ............. .*M icronycteris behni Forearm shorter than 42.5 .......... ........... .*M ronycteris sylvestris 25(18) Nose leaf with margin of lancet finely toothed (fig. 137) . . Trachops cirrhosus Nose leaf with margin entire (fig. 138) ............................. .26 26(25) Calcar distinctly shorter than hind foot and forearm longer than 67 mm ..... ... . . . . . . . . . . P h y llo s to m u s s te n o p s Not as above ...... ...... 27 27(26) Forearm longer than 75 mm; condylobasal length more than 30; zygomatic breadth more than 18 Phyllostomus hastatus Forearm shorter than 75; condylobasal length less than 30; zygomatic breadth less than 18 ..... ..... 28 28(27) Sagittal crest absent or weakly developed; calcar about one-half of length of tibia and shorter than hind foot ........ ... ... . . . . ... P h y llo s to m u s d is c o lo r Sagittal crest well developed; calcar over three-quarters of length of tibia and lon- ger than hind foot ............... ... . . . . . . . . P h y llo s to m u s e lo n g a tu s [Glossophaginae-Lonchophyllinae begin here] 29(2) Interfemoral membrane extending to about ankle (fig. 139); p2/3 ...... 30 Interfemoral membrane very narrow, ex- tending to just below knee (fig. 140) ex- cept in A. caudifer; dental formula i2/0, cl/l, p3/3, m3/3 ........ Anoura, 34 30(29) Lower incisors present; dental formula i2/2, cl/i, p2/3, m3/3 ........... 31 Lower incisors, at least in adults, ab- sent .33 sen ................. 33 31(30) Upper incisors of almost equal size, form- ing continuous row almost from canine to canine (fig. 141); zygomatic arch complete .... Glossophaga soricina Outer upper incisors conspicuously small- 34 NO. 231 .. 11 1 3 8 ANDERSON: MAMMALS OF BOLIVIA er than inner ones and separated by spaces from these and from canines (fig. 142); zygomatic arch incomplete . . 32 32(3 1) Base of hairs darker than the tips; upper premolars with rounded bases, project- ing, and of equal size ............ ............. *Lionycteri& spurrelli Basal two-thirds of hairs pale buffy white, distinctly paler than the brown tips; up- per premolars with bases elongate, nar- row, and unequal in size .......... .... ......... Lonchophylla thomasi 33(30) Upper incisors evenly and widely spaced between canines (fig. 143); wing mem- brane from base of outer toe; dental for- mula i2/0, cl/l, p2/3, m2/2 ........ .............. Lichonycteris obscura I1 and 12 separated by a distinct space from each other and from canines (fig. 144); wing membrane from foot proxi- mal to base of outer toe; dental formula i2/0, cl/l, p2/3, m3/3 ............ ........... (see text), Choeroniscus 34(29) Distinctive pl, larger than others, blade- like (fig. 145); upper canine large and with longitudinal sulcus on anterior face ... .............. Anoura cultrata Premolars much alike, pl not enlarged or bladelike (fig. 146); upper canine not en- larged and its anterior face flat .... 35 35(34) Interfemoral membrane about 1.5 to 4 mm wide, 1.5 at midline, furred dorsally, with dense fringe of hairs on free mar- gin; tail absent; forearm length 40 to 48; skull length 24 to 27; P4 with medial internal cusp; M2 with anterointernal cusp and crest; zygomatic arch some- times incomplete; calcar not evident ................. Anoura geoffroyi Interfemoral membrane width 3.5 to 5 at midline; tail present; forearm 35 to 40; skull 21 to 24; P4 without median inter- nal cusp; Ml without anterointernal cusp and crest; arch usually complete; calcar about 3 to 4 ...... Anoura caudifer [Desmodontinae begin here] 36(1) Interfemoral membrane rudimentary, con- fined to mere fringe along legs, with rather long hairs; two pair of upper in- cisors, outer upper incisor minute, near middle of inner side of canine; one low- er premolar and two lower molars on each side ........ Diphylla ecaudata Interfemoral membrane moderately devel- oped, when stretched its middle part reaches distinctly below knees; one pair of upper incisors, two lower premolars and one lower molar on each side ... ... . ....... . . . ... . . D e s m o d u s , 3 7 37(36) Wing tips white between digits 3 and 5 and from second phalanx to margin; length of thumb distinctly less than that of the hind foot; three upper cheekteeth; forearm about 53 mm long; lower inner incisors entire, outer incisors bilobed (fig. 147) ....... Desmodus youngii Wmgs without white tips; thumb about as long as hind foot; two upper cheekteeth; forearm usually longer than 55; all lower incisors bilobed (fig. 148) ......... ... . . . . . . . . . . . . D e s m o d u s r o t u n d u s [Stenodermatinae begin here] 38(4) First lower premolar relatively small (less than one-third the occlusal area of next premolar, fig. 149A, separated by a dis- tinct gap from the next premolar (fig. 149B) and with greatest breadth across lower canines more than 2.8 mm; nar- row nasal emargination extending back to, or nearly to, level of orbit (fig. 150); eyes relatively large (fig. 151A) ..... .................. .Chiroderm a, 39 First lower premolar more than one-third the occlusal area of next premolar, no gap between these two teeth (fig. 149C; or if a gap present, then breadth across lower canines less than 2.8, otherwise breadth more than 2.8); no narrow nasal emargination; eyes not so enlarged (fig. l51B) ....... ........ 41 39(38) Larger, skull longer than 24 mm; forearm longer than 44; zygomatic breadth more than 14.5 ........... .... 40 Smaller, skull shorter than 24; forearm shorter than 44; zygomatic breadth less than 14.5 ... Chiroderma trinitatum 40(39) With middorsal whitish stripe; skull longer than 26 mm; forearm longer than 50; cal- car longer than 8 . . Chiroderma salvini Without middorsal stripe; skull shorter than 26; forearm shorter than 50; calcar shorter than 8 . . Chiroderma villosum 41(38) Without middorsal stripe ..... ..... 51 With middorsal stripe ...... ...... 42 42(41) Interfemoral membrane at midline extends beyond body about 10 mm; inner upper incisors bilobed and lobes about equal in size (lateral may be somewhat small- er) ..... .......... 43 Interfemoral membrane does not extend appreciably beyond body at midline; in- ner upper incisors not bilobed, or if bi- 1997 35 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY < u 153 0 0 156 t ~~15 5 160< i 162 163 - 161 A B 166 164 167 7 165 lobed, the lateral lobe much smaller than medial lobe; nasal opening heart- shaped; dorsal rim with medial emar- gination (fig. 152) ............ 45 43(42) Small, forearm about 39 mm long; I1 more than twice the height of 12 and converging distally; two lower incisors ............... Vampyressa bidens Larger, forearm 41-46; I1 less than twice the height of 12; four lower incisors . . ................... Uroderma, 44 44(43) Rostrum narrower and depressed in dorsal profile (fig. 153); mesethmoid narrow (fig. 155A); facial stripes contrasting with dark color of head; ear edged with yellowish white (bright yellow in life); pelage grayish brown ............. .............. Uroderma bilobatum Rostrum heavy and deep, dorsal profile of skull from crown to tip of snout nearly straight (fig. 154); mesethmoid mark- edly expanded laterally (fig. 156); facial stripes poorly developed; conch of ear without white or yellowish edging; pel- age yellowish brown ............. .......... Uroderma magnirostrum 45(42) Dorsal pelage a rich reddish brown and with a very conspicuous white middor- sal stripe; no small third upper molar (M3), last lower molar (m3) relatively small, its length about one-third or less the length of the large adjacent molar ............ .Vampyrode s caraccioli Dorsal pelage brown but less reddish and middorsal stripe less conspicuous be- cause narrower; small m3 present; m3 length more than one-third the length of m2 ....... .. Platyrrhinus, 46 46(45) Larger, forearm longer than 53 mm, skull longer than 30, head and body longer than 81 ... ...... 47 Smaller, forearm shorter than 54, skull shorter than 30, head and body shorter than 81 ... ...... 48 47(46) Pelage paler, lacking frosted appearance of underparts, dorsal and facial stripes inconspicuous, reduced fringe of hairs along trailing edge of uropatagium, tips of wings pale; occlusal outline of m3 longer than broad; skull shorter than 32 mm ......... Plyrrhinus infuscus Pelage dark blackish brown, dorsal stripes prominent, fringe of hairs conspicuous, ventral pelage with hairs tipped in gray- ish-white; skull longer than 32; condy- lobasal length greater than 29; occlusal outline of m3 broader than long ..... .............. .Platyrrhin us vittatus 48(46) Size small, head and body shorter than 61 mm, forearm length 42 or less; condy- lobasal length of skull less than 20.1, skull shorter than 23.5; more or less dense fringe of hairs on free edge of uropatagium ...... ..... 49 Size larger, head and body longer than 61, forearm length 40 or more; condylobasal length of skull more than 20, skull longer than 23.5; without dense fringe of hairs on free edge of uropatagium ...... 50 49(48) Two accessory cusps on anterior margin of p2 (fig. 157C); darker; rostrum shorter and broader (fig. 157B); cranium more inflated; zygoma wider; rostral depres- sion in profile view greater (fig. 157A) . . ... . P la ty rrh in u s b ra c h y c ep h a lu s Usually only one accessory cusp on p2 (fig. 157F); skull seems relatively more slender (fig. 157E); rostral depression less pronounced (fig. 157D) ....... ... . ... . . . . . . . . P la ty r r h in u s h e lle ri 50(48) Pelage dark, two conspicuous buffy facial stripes, less dense fringe on uropata- gium; skull longer than 26.5 mm; m2 with conspicuous cuspule between pro- toconid and metaconid, m3 with well- developed labial cingulum ........ .............. .Platyrrhin us dorsalis 36 NO. 231 ANDERSON: MAMMALS OF BOLIVIA Pelage not usually so dark (except in ni- gellus), four conspicuous white facial stripes (except in nigellus where they are present but medial pair buffy and poorly defined and lateral pair obso- lete), denser fringe of hairs on uropa- tagium; skull shorter than 26.8; m2 without cuspule, m3 without cingulum .... Platyrrhinus lineatus (including Platyrrhinus nigellus) 51(41) Distinct white spot (several millimeters across) at base of leading edge of wing; face peculiar in being either unusually short, concave (fig. 158), and with cir- cular nose leaf, or in being very heavy, cuboid, and broad in rostrum (fig. 159) ............................. 52 No distinct white spot at base of wing; face not as above ....... ....... 53 52(51) Face short, rostrum depressed into facial concavity (fig. 158); nose leaf rounded (fig. 160) ......... Sphaeronycteris toxophyllum Face with large cuboid rostrum (fig. 159); nose leaf pointed at top (fig. 161) .... ............ Pygoderma bilabiatum 53(51) Skull relatively more delicate, mastoid breadth less than 9 mm; breadth across upper canines at cingulum less than 4.7; first and second upper incisors differ- entiated-first more than twice the height of second, medially convergent, and bilobed or not; second bilobed or not ......... 54 Skull relatively more massive, mastoid breadth more than 9; breadth across ca- nines at cingulum more than 4.7; I1 bi- lobed until well worn ... Artibeus, 55 54(53) Face with dark eye stripe and paler stripes above and below it; teeth larger, gap be- hind first lower premolar inconspicuous (fig. 162), less than half the length of that tooth; first molar not noticeably narrower than long (fig. 162)...... ....... ....... Vampyressa pusilla Face without distinct stripes; margins of ear yellowish; teeth relatively small and delicate, gap behind first lower premo- lar at least half its length (fig. 163); first molar above and below much longer than wide (fig. 163) .............. ........... Mesophylla macconnelli 55(53) Larger, forearm longer than 55 mm; skull longer than 24; canine breadth (between outer edges of cingula of right and left upper canines) more than 7 ...... 56 Smaller, forearm shorter than 55; skull shorter than 24, canine breadth less than 7 ... .......... 58 56(55) Blackish, relatively uniform in color, no conspicuous facial stripes; forearm shorter than 63 mm; skull shorter than 28.5; canine breadth less than 8.4 .... ................ .Artibeu s obscurus Brownish, with pale supraorbital stripes; forearm longer than 63, skull longer than 28.5, canine breadth more than 8.4 ... . ... ... . . . . . . . . . . . . . ... . . . 5 7 57(56) Facial stripes nearly white, very conspic- uous; forearm longer than 68 mm; third upper molar absent; supraorbital rim el- evated and postorbital process conspic- uous (fig. 164) ... Artibeus lituratus Facial stripes grayish, not so strikingly in contrast with adjacent areas of face; forearm shorter than 68; third upper molar usually present; supraorbital rim not noticeably elevated and postorbital process less conspicuous (fig. 165) ... ... . ... . . A rtib e u s j a m a ic e n s is f a lla x 58(55) Smaller, forearm 41 mm or shorter; skull shorter than 20; canine breadth less than 5.5; no third upper molar ........ 59 Larger, forearm 40 or longer; skull longer than 20; canine breadth at cingula more than 5.5; third upper molar present ... ... . . . . . . . ..... . . . . . . .. . . . ... 6 0 59(58) Braincase relatively shorter, more highly vaulted, more elevated above facial area as seen in lateral profile (fig. 166); top of rostrum flattened, bordered laterally by distinct angular ridge (A) below which is flattened wall of bone (B) that turns sharply outward at anterior edges; calcar shorter than 4.2 mm ....... ... . . . . . . . . . . . . A r t ib e u s a n d e r s e n i Braincase relatively elongate, less vaulted, less elevated above facial area (fig. 167); top of rostrum less distinctly flat- tened, more gradually rounded laterally and with a pit rather than flattened area as described beneath this rounded edge in anterior part of orbit .......... ... . . . . . . . . . . . . . . A r t i b e u s g l a u c u s 60(58) Blackish, with four whitish facial stripes; forearm about 40 mm long; third upper and lower molars present; mastoid breadth less than 11.3 ............ ... A r t i b e u s h a r t i i Brownish, without facial stripes; with pal- er leading edge and tips of wing; fore- arm about 50; usually no third lower molar; mastoid breadth more than 11.3 .... *Artibeus (Koopmania) concolor 37 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY [1688+ 3169 170 / 177 178 179 80 [Sturnirinae begin here] 61(3) Forearm longer than 47 mm; metacarpal of third finger (measured from outer curve of wrist) longer than 44; condy- lobasal length greater than 21.5 ... 64 Forearm shorter than 47; metacarpal of third finger shorter than 45; condylo- basal length less than 21.8 ....... 62 62(61) Condylobasal length less than 19.4 mm .............. Sturnira erythromos Condylobasal length more than 19.4 . . ............................. 63 63(62) Occlusal view of first upper molar roughly square (fig. 168); posterior lingual cor- ner angular; cusps on lingual sides of lower molars pointed and relatively tall (fig. 170); M2 much smaller than M1 ................... Sturnira lilium Ml less squared (fig. 169); posterior lin- gual corner often rounded and with a gently sloping side; cusps more rounded and shorter, forming a shallow and broad occlusal surface (fig. 171); M2 only slightly smaller than Ml ..... ............. Sturnira oporaphilum 64(61) Forearm longer than 55 mm; condylobasal length greater than 24 ............ .................. Sturnira magna Forearm shorter than 55; condylobasal length of skull less than 24 ......... ... . . . . . . ... . . . . . . . S t u r n i r a t i l d a e KEY 7 SPECIES OF THYROPTERIDAE IN BOLIVIA 1 Larger, length of forearm more than 38.7 mm; length of skull more than 15.6 .... ................ *Thyroptera lavali Smaller, forearm less than 38.7; skull less than 15.6 ....... ........ 2 2(1) Venter white or partly white; dorsum darker; tail projecting 5 to 8 mm beyond uropa- tagium; usually two cartilaginous lobules on posterior edge of calcar ......... ................. .Thyropte ra tricolor Venter not white, only slightly paler than dorsum; dorsum usually not so dark; only last two vertebrae project about 2 mm be- yond membrane; only one cartilaginous lobule on calcar . . Thyroptera discifera KEY 8 GENERA AND SPECIES OF VESPERTILIONIDAE IN BOLIVIA 1 Tail membrane at least partly covered with hair on dorsal surface (fig. 172) ..... ...................... .Lasiuru s, 2 Tail membrane nearly hairless ....... 4 2(1) Hair on tail membrane confined to part near body; pelage yellowish; one upper premolar ............ Lasiurus ega Hair extending on membrane to near edge; pelage reddish or frosted gray; two upper premolars ............ 3 3(2) Forearm longer than 45 mm; grayish ... ... . . . . . . . . . . . . . L a s i u r u s c i n e r e u s Forearm shorter than 45; reddish ...... ... . . ... . . . . . . . L a s i u r u s b l o s s e v il lii 4(1) Ears much longer than head......... ... . .... . . . . . . . . . . . . . H i s t i o t u s , 5 Ears about equal to or shorter than head ............................. ..6 5(3) Inturning lobe at base of anterior margin of ear less developed (fig. 173); length of skull usually less than 18.4 mm ... ... . . . . . . . . . . . . H is t io t u s m o n ta n u s Inturning lobe large and triangular, reach- ing well beyond tip of snout when ears are folded forward (fig. 174) ...... ... . . .... . . . . . . . . H i s t i o t u s v e la t u s 6(4) Canine and last premolar of upper jaw separated by two small teeth (may need 38 NO. 231 171 ANDERSON: MAMMALS OF BOLIVIA 181 182 189 190 187 188 192 193 194 magnification to see, fig. 175, shows M. oxyota) ... Myotis, 7 Canine and last premolar of upper jaw in contact, not separated by intermediate teeth (fig. 176, shows Eptesicus) ..... ............................. 13 7(6) Sagittal crest usually present (fig. 177), often well developed; canine breadth usually more than breadth at postorbital constriction or if about equal, then post- orbital breadth usually more than 3.7 mmn ........................... 8 Sagittal crest usually absent (fig. 178), if present poorly developed; canine breadth usually less than breadth at postorbital constriction, or if about equal, then postorbital breadth usually less than 3.7 ...... ............ 10 8(7) Fur on upper surface of uropatagium ex- tends at least halfway from knee to foot along tibia, sometimes reaching foot; P3 may be crowded but is in tooth row (fig. 181) ........... Myotis keaysi Fur on uropatagium does not reach knee; P3 usually crowded to lingual side of tooth row (figs. 182 and 183) ..... 9 9(8) Fur extremely short (2 mm on back), yel- lowish orange, and contrasting with blackish of membranes; breadth at post- 185 186 orbital constriction wide, about 3.8 mm; P3 scarcely visible in lateral view (fig. 182) ............... Myotis simus Fur longer (3 mm on back); darker brown- ish or blackish, less contrasting with color of membranes; breadth at postor- bital constriction narrow, about 3.5; P3 crowded lingually but clearly visible in lateral view (fig. 183) ............ .. ............... Myotis riparius 10(7) Uropatagium fringed (may require mag- nification to observe) and often with pale border; forearm usually 37 to 40 mm long; greatest length of skull usu- ally 14.5 to 16 ........ Myotis levis Uropatagial fringe rarely present, no pale border; size in some cases smaller 1 11(10) May have slight fringe on uropatagium, visible with magnification; dorsal fur dark (often black) with tips of many hairs white or yellow, giving a frosted appearance ....... Myotis albescens No fringe (except M. oxyotus may have sparse fringe); in dorsal fur, color of distal part not confined to the tip but extending one-third to one-half of hair length and either contrasting with more proximal part or contrasting very little with base, or both types of coloration in the same pelage ............... 12 12(11) Fur long (5 to 6 mm), bicolored; forehead comparatively steeply sloping (fig. 179); size large, forearm about 40, skull more than 14.5 long . . Myotis oxyotus Fur variable, very weakly bicolored, often blackish; forehead variable (fig. 180); forearm usually shorter than 40; skull shorter than 14.5 . . Myotis nigricans 13(7) Two pairs of upper incisors (fig. 176); larger, skull longer than 14.5 mm; fore- arm longer than 35 .... Eptesicus, 14 One pair of upper incisors (fig. 184); smaller, skull shorter than 14.5; forearm shorter than 35 ... Rhogeessa tumida 14(13) Smaller, greatest length of skull less than 16.8 mm; forearm shorter than 42; color blackish .... Eptesicus furinalis Larger, greatest length of skull more than 16.8; forearm longer than 42; color brownish .... Eptesicus andinus KEY 9 GENERA AND SPECIES OF MOLOSSIDAE IN BOLIVIA Bats of the family Molossidae are recog- nizable by the thick tail, about half of which 39 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY extends beyond the edge of the well-devel- oped uropatagium (except in flight when the membrane extends nearer the tip of the tail). They have narrow wings and fly rapidly. The foot is broad with sensory hairs.extending beyond the claws. The ears are thick, their anterior margins close together--in some species actually joined across the forehead. Flight membranes are all thick. 1 Antitragus distinctly constricted at its base (fig. 185) ................... 2 Antitragus not distinctly constricted at its base (fig. 186) ................. 5 2(1) Upper incisors caniniform, tips well sep- arated (fig. 187); palate conspicuously domed; four lower incisors (lens need- ed); base of fifth metacarpal furry .... ...................... Promops, 4 Upper incisors triangular, tips closely ap- pressed and forming a beaklike struc- ture (fig. 188); palate only slightly domed; two lower incisors; base of fifth metacarpal less hairy ... Molossus, 3 3(2) Larger, forearm longer than 45 mm; skull longer than 19 ..... Molossus rufus Smaller, forearm shorter than 45; skull shorter than 19 . . Molossus molossus 4(2) Larger, forearm longer than 51 mm; skull longer than 19 ... Promops centralis Smaller, forearm shorter than 51; skull shorter than 19 .... Promops nasutus 5(1) Deep vertical grooves in upper lips (fig. 186) .......... 6 No conspicuous vertical grooves in upper lips ........... 9 6(5) Larger, forearm 48 mm or longer; skull longer than 18 ..... ..... 8 Smaller, forearm 48 or shorter; skull shorter than 18 ..... ..... 7 7(6) Ears joined at midline of head, reaching beyond nose; antitragus higher than width at base; pelage reddish brown . . .......... Nyctinomops laticaudatus Ears not joined at midline, not reaching beyond nose; antitragus wider at base than high; pelage darker and less red- dish brown .... Tadarida brasiliensis 8(6) Larger, forearm longer than 50 mm; skull longer than 20.5; ears longer than 18 mm; color grayish brown .......... ............. Nyctinomops macrotis Smaller, forearm shorter than 50; skull usually shorter than 20.5; ears shorter than 18; color brown to blackish ..... ......... Nyctinomops aurispinosus 9(5) Ears joined in midline .... Eumops, 10 Ears clearly separate ... Molossops, 14 10(9) Smaller, forearm less than 55 mm (usually 37 to 49); skull shorter than 22 ...... ... Eumops bonariensis or E. hansae Larger, forearm longer than 55; skull lon- ger than 22 ........1..... 1 11(10) Ears long (35 to 44 mm); tragus large, broad and square; basisphenoid pits deep and elongate (fig. 189); mastoid breadth less than 52% of condyloinci- sive length ........ Eumops perotis Ears short (17 to 34); tragus small, point- ed or square; basisphenoid pits not so deep (fig. 193); mastoid breadth more than 52% of condyloincisive length 12 12(1 1) Larger, skull longer than 28 mm (males) or 27 (females); ear heavily keeled (fig. 190, arrow); dorsal pelage cinnamon with buff basal band ............. ... . . . . . . . . .... * E u m o p s d a bb e n e i Smaller, skull shorter than 28 (males) or 27 (females); ear not heavily keeled (fig. 191); pelage snuff brown to sepia (with white basal band) or blackish brown ..... ........ 13 13(12) Tragus small and pointed (fig. 192); dorsal pelage blackish brown; basisphenoid pits shallow (fig. 193, arrow); mastoid breadth less than 49% of skull length ............. .Eumop s auripendulus Tragus broad and square; dorsal pelage snuff brown to bister with white basal band; basisphenoid pits well defined (fig. 194 arrow); mastoid breadth more than 52% of skull length .......... ................ .Eumop s glaucinus 14(9) Forearm length about 45 mm; total length 90-100; dorsum cinnamon red ...... ... . . . . . . . . ... M o lo s s o p s a b r a s u s Forearm shorter than 40 .......... 15 15(14) Forearm longer than 31 mm; skull longer than 16; extensive areas of white on venter; four lower incisors; large square antitragus ... Molossops planirostris Forearm shorter than 33; skull shorter than 16; throat pale; two lower incisors; small narrow antitragus ........... ............. .Molossop s temminckii KEY 10 GENERA AND SPECIES OF CALLITRICHIDAE AND CEBIDAE IN BOLIVIA 1 Two molars above and below on each side (fig. 195 shows Callithrix); generally small, head and body shorter than 250 mm) . Family Callitrichidae, 15 40 NO. 231 ANDERSON: MAMMALS OF BOLIVIA 201 Three molars above and below on each side (fig. 196 shows Callimico) .... 2 2(1) Flattened unguis or nail on most or all digits; size medium to large (head and body length rarely less than 300 mm); various colors, most species not black ............... Family Cebidae, 3 Nail only on hallux, other digits bearing claws (fig. 197); size small (head and body length less than 250); pelage mostly blackish ....... Subfamily Callimiconinae, Callimico goeldii 3(2) Tail not fully prehensile, hairy throughout its length; head plus body length gen- erally between 300 and 500 mm; total length usually less than 1 m ...... 4 Tail fully prehensile, with naked skin on ventral surface near tip (fig. 198); head plus body length generally more than 500 mm; total length of adults rarely less than 1 m ................. 11 4(3) Orbits large (fig. 199) .............. ....... Subfamily Aotinae, Aotus, 5 Orbits of normal size (fig. 200) ..... 6 5(4) Occurring north of the rio Madre de Dios; with 51 or 52 chromosomes in the dip- loid set; not easily distinguished by ex- ternal features or skull ............ .................. Aotus nigriceps Occurring south of the rio Madre de Dios; 2n = 49 or 50 chromosomes ...... ... . . . . ... . . ........ ..Ao t u s a z a r a i 6(4) Incisors not long and protrusive, not wide- ly separated from canines ........ 7 Incisors long and protrusive (fig. 201), close together and widely separated from canines .................... ......... .Subfamil y Pitheciinae, 10 7(6) Lower canines long, projecting distinctly above tooth row (figs. 202, 203); with considerable sexual dimorphism . .. 8 Lower canines short, barely projecting above other teeth (fig. 204); no obvious sexual dimorphism ............... Subfamily Callicebinae, Callicebus, 4 species, see text 8(7) Skull elongate and with large occipital re- gion (fig. 205); oval interorbital open- ing; tail length about 1.3 times head plus body length; not at all prehensile . ... . . . . . . . . . . . S a i m i r i s c i u r e u s Skull not especially elongate, braincase normally rounded (fig. 206); interorbital septum complete; tail length about the same as head plus body, semiprehensile ........................ .Cebu s, 9 9(8) Head tufted with dark brown or black erect hairs on crown forming lateral ridges or a complete cap; dark bands down sides of face, meeting beneath chin; limbs generally darker than body .................... .Cebu s apella Head not tufted, smooth brown hairs form broad patch set well back on crown; face, forehead, throat, shoulders, upper arms, and chest are whitish; hands and feet usually paler than body ...... .................. .Cebu s albifrons 10(6) Pelage relatively short and soft; hairs ra- diate from whorl on the top of head, directed sideways to form bushy lateral tufts; beard long and thick ........ ... . . . . . . . . . . * C h ir o p o t e s a lb in a s u s Pelage long and shaggy, forms hood and cape .......... Pithecia irrorata 11(3) Braincase large and rounded (fig. 202); foramen magnum directed downward; hyoid not especially enlarged; lower jaw not unusually deep (fig. 202); black with no distinctive markings of another color ....... Subfamily Atelinae, 14 Braincase small and flattened (fig. 203); foramen magnum directed backward; hyoid enlarged, lower jaw unusually deep (fig. 203) ........ Alouatta, 12 12(11) Pelage maroon or copper red ....... ................ .Alouat ta seniculus Not as above ............... 13 1997 41 195 196 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 13(12) Dark brown; hairs tipped with light brown ................. Alouatta guariba Black (adult males) or olive-buff or yel- lowish (females and young) ....... .. . ............. Alouatta caraya 14(11) Long shaggy hair, directed forward above the eyes to form peak . . Ateles chamek Dense, short, plushy hair; head rounded ............. *Lagothrix lagotricha 15(1) Lower jaw U-shaped; lower canines are much longer than incisors (fig. 207, hence "long-tusked") .. Saguinus, 16 Lower jaw narrow and V-shaped; lower incisors and canines form a prominent semicircular row of six teeth with the canines scarcely higher than incisors (fig. 195, "short-tusked") ....... 19 16(15) Facial skin white beneath long white mus- tache, otherwise black ............ ................ mystax group, 17 Facial skin and hair black except for short white hairs of mustache; mottled black and buff saddle separates fore and hind parts (trizonal pattern) ............ .............. Saguinus fuscicollis 17(16) Grayish buff, speckled; mustache long (60 mm); tail generally reddish with dark gray tip ........ Saguinus imperator Blackish in large part; mustache outlines lips or is larger but not exceptionally long; tail mostly black .......... 18 18(17) Tail entirely black . . *Saguinus mystax Tail reddish or orange on ventral side near base; underparts of body also reddish or orange; white patch on crown ..... ................ Saguinus labiatus 19(15) Head plus body length less than 175 mm; condylar process of jaw extends poste- riorly (fig. 208); neutral gray-brown owing to black and buff banding of hairs; tail banded dorsally, less than 250 long ........... Cebuella pygmaea Head plus body length more than 175 ; con- dylar process of jaw less extended (fig. 209); pelage brown with white hip-patch- es; tail black and not ringed, tail longer than 250 ....... CaUithrix melanura KEY 11 GENERA AND SPECIES OF MYRMECOPHAGIDAE IN BOLIVIA The anteaters or Myrmecophagidae are toothless, have relatively pointed snouts, small mouths, and long tongues. The forelimbs are strong and there is no zygomatic arch. 1 Hair coarse and long; tail bushy, not prehen- 7-- 2_ _-7K91 Ur--- ~ ~~ ~~~ = -: -3 sile; size large, head plus body longer than 700 mm; skull longer than 200 and ros- trum proportionally long (fig. 210) ... ............ .Myrmecophaga tridactyla Hair not coarse or long; tail not bushy, partly naked at tip, prehensile; head plus body length less than 700; skull shorter than 200, rostrum proportionally not so long (figs. 211, 212) .... ...... 2 2(1) Four claws on forefoot; fur, relative to the size of the animal, short and smooth; length of head plus body more than 300 mm; rostrum and nasals not extremely short (fig. 211); length of skull more than 70 ......... Tamandua tetradactyla Two claws on forefoot; fur dense, soft, and silky; length of head and body less than 300; rostrum and nasals extremely short when compared to braincase (fig. 212); length of skull less than 70 ........... ............... .Cyclopes didactylus KEY 12 GENERA AND SPECIES OF DASYPODIDAE IN BOLIVIA The armadillos or Dasypodidae are readily distinguished by their external "armor," their 42 NO. 231 ANDERSON: MAMMALS OF BOLIVIA simplified, peglike, persistently growing teeth without enamel coverings, and their short, stout limbs with strong, curved claws. l Snout long and slender; ears close together on top of head (fig. 213); longest claw on forefoot about half of length of ear; palate very long (fig. 215A), length behind tooth rows more than half of the length of the tooth rows ..................... . . Subfamily Dasypodinae, Dasypus, 2 Snout heavier; ears set wide apart on sides of head (fig. 214); longest claw of fore- foot noticeably longer than half the length of ear (in some cases longer than ear); palate shorter (fig. 216A), length behind tooth rows less than half length of tooth rows ............... 2(1) Conspicuous long, thick, brown hair; scutes of carapace not visible without parting hair ....... *Dasypus pilosus Very few hairs; scutes of carapace clearly visible ............... 3 3(2) Nasal bones longer than 35 mm; strikingly enlarged scutes on patellar region (fig. 217); usually five toes on forefeet .... ................. Dasypus kappleri Nasal bones shorter than 35; scales of pa- tellar region not much larger than other scales on hind leg; usually with four claws on forefeet ............... 4 4(3) Larger, total length more than 700 mm; length of skull more than 88; length of head shield more than 80; usually eight or more movable bands on the back; larger scutes on anterior and dorsal shields of carapace about 3-4 mm from center to center, smaller, and more closely spaced (fig. 219) .......... ............ Dasypus novemcinctus Smaller, total length less than 700; length of skull less than 88; length of head shield less than 80; usually fewer than eight movable bands on back; largest scutes about 5-6 from center to center, larger, and more widely separated (fig. 220) ....... Dasypus septemcinctus 5(1) Hairy, a noticeable quantity of long hairs on banded area of back, many extend- ing beyond adjacent band; most anterior upper tooth is in premaxillary bone (fig. 216B); head wide, width of skull more than half its length (fig. 216) ...... ........ Subfamily Euphractinae, 6 Few dorsal hairs on banded area of back and these not extending beyond adja- cent band; most anterior upper tooth is in maxillary bone; width of skull less than half its length .............. 9 6(5) Many scutes of head shield exceed 10 mm in greatest diameter; scales of nape band anteroposteriorly longer than wide (fig. 214) ... Euphractus sexcinctus Few scutes of head shield exceed 10; scales on nape band somewhat variable but often transversely wider than long ................ .Chaetophract us, 7 7(6) Smaller, head and body length less than 250 mm, condylonasal length less than 80; ear proportionally long, extending to first immobile row of scutes in scap- ular shield; width of cranial shield less than 90% of its length; dorsum varie- gated brown and pale tan ......... ......... .Chaetophract us vellerosus Larger, head plus body length more than 250 in large adults, condylonasal length more than 80; ear long or shorter; head shield ratio more than 90% of its length ...... 8 8(7) Dorsum tan, covered with long, brown and pale tan hairs; ear long, approxi- mately 30 mm; restricted to high An- dean grasslands ................. ... .. ... . . . . C h a e to p h r a c tu s n a tio n i Dorsum dark brown with sparse, long, black and brown hairs; ear shorter ... ... . . . . . . . . C h a e to p h r a c tu s v illo s u s 9(5) Small, head plus body length less than 200 mm; distinct rump plate, nearly vertical, not blending imperceptably with the dorsal scaled area ....... ... . . . . S u b fa m ily C h la m y p h o r in a e , Chiamyphorus retusus Head and body longer than 200; no dis- tinctly separate vertical rump plate . .10 10(9) Nasals shorter than 20 mm; tail shorter than head; two to four, usually three, movable bands in carapace, bands clear- ly distinct from continuous shields of front and rear parts of carapace (fig. 9); able to roll into compact ball (fig. 218); longest claw on forefoot shorter than 30 mm ... ....... Tribe Tolypeutini, Tolypeutes matacus Nasals longer than 20; tail longer than head; many more than three movable bands and these seem to grade into the immovable areas at front and rear; lon- gest claw longer than 30 ......... .. . . . . . . . . . . T r i b e P r i o d o n t i n i , 11 11(10) Tail with scattered scales separated by ar- eas of skin often as wide as the scales themselves; skull shorter than 110 mm; head plus body length less than 400; 1997 43 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY teeth not so flattened laterally as to be sometimes twice as long as wide .... ................... Cabassous, 12 Tail not with most scales separated by ar- eas of skin; gigantic, skull longer than 110 (up to at least 180); head plus body length longer than 400; some teeth so flattened as to be twice as long antero- posteriorly as wide ............... ....... Priodontes maximus giganteus 12(11) Smaller, interorbital breadth less than 23 mm; ear shorter than 20 and with fleshy anterior edge; mandible distinctly curved (fig. 221); transverse diameters of all but first and last teeth greater than anteroposterior diameters; skull rela- tively more slender (fig. 223) ..... ............ *Cabassous chacoensis Larger, interorbital breadth more than 23.5 mm; ear longer than 25 and without thickened anterior edge; mandible not distinctly curved (fig. 222); skull rela- tively broader (fig. 224); transverse di- ameters of teeth usually less than anter- oposterior diameters .............. ............. Cabassous unicinctus KEY 13 GENERA AND SPECIES OF CANIDAE IN BOLIVIA 1(2) Skull usually with bulging forehead as seen in profile (fig. 225; size, bodily propor- tions, color, length of hair, and other fea- tures highly variable, depending on breed); domestic ...... Canis familiaris Skull with straighter profile, less bulging forehead (fig. 226; size, proportions, color, length of hair, and other features vary with species, but not much within species); wild ..... .......... 2 2(1) Either the size of a large dog and unusually long-legged (fig. 227) or the size of a small dog and unusually short-legged (fig. 41); length of skull greater than 195 mm or less than 130 ........ .... 3 Both size and proportions moderate (as in a dog of medium size); breadth across upper canines at alveolus (fig. 228A) between 25 and 35; length of skull between 195 and 130 .......... 4 3(2) Large, long-legged (fig. 227), height at shoulder about 850 mm; ears longer than 60 from notch; white patch on throat and black stripe below white (general color reddish brown); tail longer than head; lon- gest hairs longer than 70; noticeable mane of longer hair on neck; breadth across up- 225 226 ---\ ICBJ 227 229 D *'231 232 per canines more than 35 ........... ............. .Chrysocyon brachyurus Small, short-legged (fig. 41), height at shoul- der less than 350; ears small, shorter than 60 from notch; no white and black patches on throat, head and neck paler than more posterior and ventral areas (general color reddish brown); tail short, about the length of head; hair short and coarse, no fine un- derfur, longest hairs about 30; no mane; breadth across canines less than 35 ..... ................. .Speothos venaticus 4(2) Distinct rusty patches on sides of front and hind limbs and on and behind ears; larger, upper canine breadth more than 27 mm; length of skull more than 165; fourth upper premolar longer than 16.8; lives at high el- evations ........ Pseudalopex culpaeus If pelage has a generally reddish hue, the rusty patches noted are absent or indistinct and reddish color extends along belly as well as on legs; smaller, upper canine breadth less than 27; skull shorter than 165; P4 shorter than 16.8; usually found at lower elevations .............5 5(4) Angular process narrow (fig. 229); breadth at postorbital constriction about the same as (fig. 23 1D), or noticeably less than (fig. 44 NO. 231 ANDERSON: MAMMALS OF BOLIVIA 228), breadth at interorbital constriction (fig. 231C) .......... ...... 6 Angular process broad (fig. 230); breadth at postorbital constriction greater than at in- terorbital constriction (fig. 232) ...... ................... Cerdocyon thous 6(5) Pelage paler, almost white below; legs bicolor (whitish on one side), not black; ears lon- ger than 50 mm from notch; smaller, con- dylobasal length about 135; interorbital (fig. 231C) and postorbital constrictions (fig. 231D) about equal; nasals extending about same distance posteriorly as maxil- laries ...... Pseudalopex gymnocercus Pelage dark, not whitish below; legs almost pure black; ears shorter than 50 (fig. 233); larger, condylobasal length (fig. 226) about 160; breadth at postorbital constric- tion noticeably less than at interorbital constriction (fig. 228); nasals usually do not extend so far posteriorly as maxillaries (fig. 228B) ...... Atelocynus microtis KEY 14 GENERA AND SPECIES OF PROCYONIDAE IN BOLIVIA 1 No dark rings on tail; three premolars above and below; palate parallel-sided, upper ca- nines nearly as far apart as molars (fig. 234); mandible unusually deep (fig. 236), entire lower tooth row shorter than great- est depth of jaw ......... Potos flavus Tail with or without rings; four premolars above and below; palate narrowing anteri- orly (fig. 235); jaw slender, tooth row much longer than depth of jaw (fig. 237) ..... 2 2(1) Rostrum elongated and compressed laterally, distance from front of eye to front of teeth more than 50 mm (fig. 238); upper fourth premolar (P4) and first upper molar (Ml) usually longer anteroposteriorly than broad (fig. 240) ........ Nasua nasua Rostrum not elongated and not laterally compressed; distance from front of eye to front of teeth less than 50 (fig. 239); P4 and Ml as broad mediolaterally as antero- posteriorly or broader (fig. 241) ..... 3 3(2) Eyes surrounded by black pelage that forms a "mask" across the face; tail with rings of black, shorter than length of head plus body; large, breadth across upper canines (fig. 44C) more than 25 mm; mastoid large (fig. 242); claws relatively blunt, not much curved (fig. 244); all digits extend more than 15 beyond their basal connec- tions . Procyon cancrivorus No distinct black markings, pelage nearly 257 258 uniform dorsally; tail not conspicuously ringed but may show faint alternating dark and paler areas; tail longer than length of head plus body; smaller, breadth across upper canines less than 25; mastoid small (fig. 243); claws relatively sharp and curved (fig. 245); web of skin between short toes reaching nearly to base of distal pad (fig. 246) ...... Bassaricyon alleni KEY 15 GENERA AND SPECIES OF MUSTELIDAE IN BOLIVIA Dorsal pelage brown, relatively short and sleek and with no distinctly contrasting marks on back or head (although throat may have whitish patch); head same color as back; five upper teeth behind canines .............. .Subfami ly Lutrinae, 2 Dorsal pelage not as above; three or four up- per teeth behind canines ........... 3 2(1) Tail without a prominent keel on each side; basal part of toes webbed, usually to base of distal phalanx (fig. 247 shows right front foot); condylobasal length less than 130 mm; frontal area short; skull relatively 45 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY broad posteriorly (fig. 249) ......... .................. Lutra longicaudis Tail somewhat flattened, with prominent keel on each side; toes webbed more exten- sively, always to the nails (fig. 248); con- dylobasal length of skull more than 120; frontal area long; skull relatively narrow posteriorly (fig. 250) ............... ............... Pteronura brasiliensis 3(1) Pelage of contrasting white markings on black background (fig. 251); claws of forefoot long and strong (fig. 252), the longest more than 14 mm; only three up- per teeth behind canine (fig. 253A), the back tooth (= Ml) large, its length about the same as its width (fig. 253B); no post- orbital processes (fig. 255); little extension of bony palate behind level of teeth, ex- tension less than distance between teeth (fig. 253) ..... Subfamily Mephitinae, Conepatus chinga Pelage not of contrasting white on black; front claws shorter than 14; four upper teeth behind canines, the back tooth about twice as wide mediolaterally as from front to back and with a medial constriction (fig. 254B); postorbital processes present (fig. 256); bony palate extending well be- hind teeth, usually more than the distance between teeth (fig. 254C) ........... ............ Subfamily Mustelinae, 4 4(3) Tail longer than 300 mm and longer than half the length of the head plus body (fig. 42); breadth across upper canines greater than 25; condylobasal length of skull (fig. 46A) more than 100; head usually paler than blackish back but some individuals pale golden over most of body, head with- out striking markings of blackish and whitish or cream color ... Eira barbara Tail shorter than 300 and in some cases shorter than half the length of head plus body; condylobasal length less than 100; head not noticeably paler than back, head with striking markings of blackish and whitish or cream color ............. 5 5(4) Condylobasal length less than 50 mm; total length less than 400; blackish part of head is top part, venter is whitish or yellowish (fig. 257) ........... Mustela frenata Condylobasal length more than 50; total length more than 400; blackish part of head is bottom part, extending up over nose and backward onto venter, top of head is grizzled plus between these areas is a paler cream-colored stripe (fig. 258) ........................ Galictis, 6 6(5) Larger, head plus body length more than 450 mm; condylobasal length of skull more than 75, breadth across upper canines more than 17; weight usually greater than 1.2 kg ... ... Galictis vittata Smaller, head plus body length less than 450; condylobasal length less than 75; up- per canine breadth less than 17; weight about 1 kg .... .. Galictis cuja KEY 16 GENERA AND SPECIES OF FELIDAE IN BOLIVIA 1 Braincase smaller, its greatest width (fig. 259B) usually less than 43.5 mm; color pattern varies greatly among individuals but never consists of black spots on yel- lowish or orange background; domestic species ................. Felis catus Braincase larger, its width usually more than 43.5 (if less than 43.5, then has black spots on orange background); color varies among species; wild species ........ 2 2(1) Size large, head plus body longer than 1100 mm; hind foot longer than 165; condylo- basal length (measurement shown in fig. 46A) more than 115; length of mandible (fig. 260) more than 80 ............ 3 Size medium, head plus body shorter than 1100; hind foot shorter than 165; condy- lobasal length less than 115; length of mandible less than 80 ............. 4 3(2) Whiskers growing from solid black patch, solid white patch between black patch and nose; dorsal pelage of adults a relatively uniform brownish above (young have spots); breadth across upper canines usu- ally less than 55 mm; anterior flange on parietal often overlaps frontal (fig. 261), but flange sometimes absent or bones fused and flange not evident ........ ... . . . . . . . . . . ... . . . . . F e lis c o n c o l o r Not so marked; pelage a striking pattern of black spots on orange background (except in occasional melanistic individuals in which the spots are almost obscured); breadth across upper canines usually more than 55; suture between parietal and fron- tal without flange noted (fig. 262) ...... ... . . . .... . ..... .... . P a n th e r a o n c a 4(2) Dorsal profile of skull only slightly arched (fig. 263), nasals extending relatively far forward (fig. 265); dorsal pelage a rela- tively uniform reddish, brownish, or gray- ish, with no conspicuously contrasting markings; at least part of the dorsal sur- face with "salt and pepper" effect ..... ... . . . . . . . . . . . . . . . . F e l is y a g u a r o n d i 46 NO. 231 ANDERSON: MAMMALS OF BOLIVIA Dorsal profile highly arched (fig. 264); nasals terminating well back of premaxillary ex- posing anterior edges of incisive alveoli in dorsal view (fig. 266); dorsal pelage. less uniform in color, usually with contrasting dark markings (spots, stripes, blotches) on paler background (in occasional melanistic individuals, markings are faintly visible un- der close examination even if general ap- pearance is of solid black, without "salt and pepper" effect) ..... .......... 5 5(4) Cranium long and narrow; notch or groove at junction of anterior and posterior cham- bers of bulla (fig. 267); anterior premolar toes in at front (fig. 268); pelage thick; color ashen, with widely spaced coffee- colored or darker spots . . Felis jacobita Cranium relatively broader; less distinct notch or groove (fig. 269); anterior pre- molar more in line with posterior premolar; pelage not so thick, color variable, usually not resembling above description ..... 6 6(5) Eyes relatively large (fig. 270), with maxil- lary rim of orbit more delicate and flared outward, or, if eyes do not seem relatively large, then size small, condylobasal length of skull less than 85 mm; length of head plus body less than 550; the largest upper molariform tooth (P4) shorter than 11 mm7 Eyes relatively smaller (fig. 271), maxillary rim seems more robust and less flared out- ward; P4 longer than 11 ........... 8 7(6) Hair on dorsum of neck pointing backward; smaller, length of P4 less than 11 mm, breadth across upper canines less than 21; braincase oval ......... *Felis tigrina Hair on dorsum of neck pointing forward; larger, length of P4 more than 11, breadth across upper canines more than 21; brain- case more rounded (fig. 270) ........ ... . . . . ... . . . . . . . . . . . . . . F e l i s w i e d i i 8(6) Larger, breadth across upper canines more than 29 mm; skull longer than 105; P4 longer than 12.7; some markings have a dark border and paler center (on even pal- er background color) .... Felis pardalis Smaller, breadth across upper canines less than 29; skull shorter than 105; P4 shorter than 12.7; few if any two-toned patches with dark borders ................. 9 9(8) Small anterior maxillary tooth (P2) usually (80% of cases) absent (fig. 272A); tym- panic bulla rounded anteriorly (fig. 272B); profile of nasal bones turns downward near its middle (fig. 273); markings on lower legs heavier than on more dorsal parts of body; fewer spots; middorsal crest of long hairs from behind shoulders to base of tail ............ Felis pajeros Small tooth (P2, P1 not present) always present (fig. 269A); bullae more extended and constricted anteriorly (fig. 269B); na- sal bones straighter in profile (fig. 274); markings on lower legs not heavier than on more dorsal areas; more spots; no mid- dorsal crest of long hairs ........... ..................... .Feli s geoffroyi KEY 17 GENERA AND SPECIES OF TAYASSUIDAE OF BOLIVIA 1 The lower second molar (m2, the second tooth on the mandible counting from the back; fig. 275A) longer than 17 mm; total length of mandible (from condyle to an- terior tip; fig. 275B) greater than 200; ratio of depth of bone below m2 to the total length of mandible less than 1 to 6 ..... ... . ... . . . . . . . . . . C a t a g o n u s w a g n e ri The lower second molar (m2) shorter than 17; total length of mandible less than 200; ratio of depth of bone below m2 to length of mandible more than 1 to 6 ....... ... . ... . . . . . . . . . . . . . . . . T a y a s s u , 2 2(1) Alveolar length of lower cheekteeth more than 75 mm; mandible longer than 165; 1997 47 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 284 285 28 0 28728 rIJ I r W.- . } 288 Y 28 diastema (fig. 275D) in lower tooth row greater than 29; alveolar length of lower three premolars (fig. 275P) greater than 29; D greater than P; D times P greater than 860 .. .. Tayassu pecari Alveolar length of lower cheekteeth less than 75; mandible shorter than 165; D less than 29; P less than 29; D less than P; D times P less than 860 ... Tayassu tajacu KEY 18 GENERA AND SPECIES OF CAMELIDAE IN BOLIVIA This key is adapted from that of Cabrera (1931). The variation in the four species of the genus Lama is sufficient to make identi- fication uncertain in some cases even when the entire skull and jaw are present. There is, in fact, a difference of opinion as to whether they should be treated as distinct species. With broken or fragmented bones such as are found in archaeological sites, the problem is greater. 1 Lower incisors tapered to the ends of their roots, greatest width about one-fifth to one-fourth the length of tooth (fig. 276) except in some alpacas with unusually long incisors; size larger, greatest width of skull of adult more than 125 mm; color varies from black to white, usually not colored as is vicuna .............. 2 Lower incisors of adults elongate and nearly parallel-sided (fig. 277), width about one- tenth length, and persistently growing (ex- cept in extremely old individuals or young ones in which milk teeth taper slightly, have widths about one-sixth of lengths, and have open roots); size small, greatest width of skull of adult less than 125; build delicate; dorsal pelage pale reddish brown, conspicuously whitish area on the body near base of neck ...... Lama vicugna 2(1) Rostrum and lower jaw short; height of man- dible at the front of fourth premolar (fig. 278A) almost equal to the distance from this point to the large mental opening (fig. 278B); submental opening (fig. 278C), usually beneath p4; domestic ........ ................ .Lama pacos, Alpaca Rostrum and jaw more elongate; height of mandible in front of p4 notably less than the distance from this point to the large mental opening (fig. 279); submental open- ing usually beneath front of ml ...... 3 3(2) Ascending ramus of mandible broad, its an- teroposterior span at level of angular pro- cess (fig. 280A) much more than the dis- tance from this process to the condyle (fig. 280B); domestic . . Lama glama, Llama Ascending ramus of mandible narrow, its width at the level of the angular process about equal to or not much more than the distance from this process to the condyle (fig. 281); nearly extinct in Bolivia ..... ... . . . . . . . . L a m a g u a n ic o e , G u a n a c o KEY 19 GENERA AND SPECIES OF CERVIDAE IN BOLIVIA The Bolivian Cervidae are ruminant arti- odactyls with no upper incisors; upper ca- nines, if present at all, are small; without a flattened (piglike, fig. 15) muzzle or nearly horizontal slitlike nostrils (as in llamas, fig. 282), but with a distinct bare nosepad (fig. 283); and with branching antlers or spikes almost always present in males. 1 Preorbital pits in skull very deep (up to the length of one's fingernail in most cases, fig. 284), upper rim prominent ...... 2 48 NO. 231 ANDERSON: MAMMALS OF BOLIVIA Preorbital pits not especially deep (fig. 285), rounded upper border .............. 3 2(1) Antlers (when present) never with more than two tines; orbital rim relatively heavy, ex- panded dorsally and laterally so that greatest width of skull is at rim instead of at zygo- matic arch, orbit facing only slightly dorsal- ly (fig. 286); lower incisors fan out laterally so that width across incisors is almost as great as length of mandibular symphysis; pelage coarse and brittle, paler, flecked with black, legs paler; elevation 3000 to 4000 m .............. Hippocamelus antisensis Antlers (when present) usually with more than two tines; orbital rim not so heavy, less expanded, greatest width of skull at zy- gomatic arch, orbit facing noticeably more upward and forward (fig. 287); breadth across incisors only about 60% of the length of symphysis; pelage less coarse, darker, rufous brown (redder in summer), legs dark; inhabits elevations below 1000 m ........... Odocoileus dichotomus 3(1) Auditory bulla moderately inflated (fig. 288); antlers (when present) usually with more than two points ............... .............. Odocoileus virginianus Auditory bulla slightly inflated (fig. 289); antlers (when present) with one or more points ........... 4 4(3) Orbits relatively smaller, dorsoventral di- ameter of orbit (at widest point of orbit) is less than half the breadth between orbits (dorsal point of diameter dimension of each orbit as end point); antlers (when present) often with more than one point .............. Odocoileus bezoarticus Orbits relatively larger, dorsoventral diame- ter more than half distance between orbital rims; antlers never more than a simple spike ..... ...... Mazama, 5 5(4) Size small (head plus body shorter than 740 mm; skull shorter than 150; hind foot shorter than 175; length of dentary from articular process to anterior-most point less than 120 mm .... Mazama bricenii Size larger (head plus body longer than 740; skull longer than 150; hind foot longer than 175; length of dentary more than 120) ........... 6 6(5) Skull longer than 190 mm; greatest breadth of skull at orbit more than 85; length of dentary generally 150-160; pelage red- dish brown ....... Mazama americana Skull shorter than 190; greatest breadth of skull at orbit less than 85; length of den- tary about 140; pelage not reddish brown 30( 5 )30 ?,05 but grayish brown ................. ... . . . . . . . . . . . . M a z a m a g o u a z o u p ir a KEY 20 GENERA AND SPECIES OF BOVIDAE IN BOLIVIA The Bovidae of Bolivia are all introduced domestic animals. There are no native bovids in South America. Domestic herds and free- ranging individuals are common and impor- tant in the ecosystem. Skulls and other bones are often found. The inclusion of domestic as well as native species in these keys will help users to distinguish remains of native species from those of domestic ones. 1 Total length of skull (not including the horns if present) more than 350 mm ....... ... . . . . .. . . .. . . . . . . . . . . . B o s t a u r u s Total length of skull (without horns) less than 350 ....................... 2 2(1) Deep preorbital fossa; lacrimal-jugal suture passing across bottom of fossa (fig. 290) ... . . . . . . . . . . . . . . . . . . . . . . O v i s a r i e s No evident preorbital fossa; lacrimal-jugal suture passing along crest of preorbital ridge (fig. 291) ......... Capra hircus 49 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY KEY 21 GENERA AND SPECIES OF LEPORIDAE IN BOLIVIA 1 Small, head plus body length usually less than 380 mm; hind foot shorter than 85 mm; tail small and inconspicuous, shorter than 30 mm, dark, not in part white; su- praorbital process posteriorly not well sep- arated from frontal (fig. 292A) ...... ... native species, Sylvilagus brasiliensis Larger, head plus body usually longer than 380; hind foot longer than 85; tail con- spicuous although relatively short in com- parison to body, longer than 30, dark above and usually white below; supraor- bital process well separated from frontal posteriorly (figs. 293 and 294) ...... 2 2(1) Ear longer, length more than 100 mm from basal notch, with black patch on outer sur- face at tip; hind foot longer than 100 mm; head plus body longer than 550; supraorbital processes wide and more laterally projecting (fig. 293); mesopterygoid fossa wide (fig. 295); rostrum larger in proportion to rest of skull (fig. 293); introduced from Europe into Argentina, then spread into southern Bolivia ................... Lepus europaeus Ears shorter, length less than 100 from notch, without large black patches al- though edges of ears sometimes black; hind foot shorter than 100; head plus body shorter than 550 (except in some large do- mestic breeds); supraorbital processes nar- rower and less laterally projecting (fig. 294); mesopterygoid fossa not so wide (fig. 296); rostrum smaller in proportion to rest of skull (fig. 294); domestic, native to Europe, no feral population known in Bo- livia .. *Oryctolagus cuniculus KEY 22 GENERA AND SPECIES OF SCIURIDAE IN BOLIVIA Pelage reddish; large, length of head plus body usually more than 210 mm; occipitonasal length more than 50; hind foot (from back of heel to end of claws) longer than 50 ............ ...................... Sciurus spadiceus Brownish; smaller, length of head plus body usu- ally less than 210; occipitonasal length (fig. 297) less than 50; hind foot shorter than 50 . . ........................ Sciurus ignitus KEY 23 GENERA AND SPECIES OF MURIDAE IN BOLIVIA I Cusps of upper molar teeth arranged in three longitudinal rows (fig. 298); tail always conspicuously scaly and never well clothed in either long or short hairs; introduced species, usually com- mensal .... Subfamily Murinae, 2 Cusps of upper molar teeth not arranged in three rows, may be in two rows or flattened occlusally so that no tops of cusps remain (fig. 299); tail may or may not show scales conspicuously, in most species tail more or less clothed in hair; native species, some inhabiting houses and other buildings in the absence of Murinae ....................... ....... .Subfamil y Sigmodontinae, 3 2(1) Larger, hind foot longer than 28 mm; skull longer than 20; length of molar row more than 5 ......... Rattus rattus Smaller, hind foot shorter than 23; skull shorter than 20, length of molar row less than 5; upper incisors with notch in side view (fig. 300) .. Mus musculus 3(1) Zygomatic plate much narrowed, usually slanting backward from lower to upper border (fig. 301); infraorbital foramen usually large, well open (fig. 304); ros- trum lengthened (fig. 303; except in the recently discovered ichthyomyine, Chibchanomys, which has not been added to this key) .............. 4 Zygomatic plate not narrowed or less nar- rowed (fig. 302), and tilted more strong- ly upward (if narrowed, it is always tilt- ed upward, fig. 305); skull without above peculiarities .............. 8 4(3) Foreclaws prominent and long (fig. 306); rostrum especially elongate (fig. 307) .................. .Oxymycteru s, 5 Foreclaws not so prominent and long; ros- trum not so elongate ............ 7 5(4) Small, head plus body shorter than 120 mm; skull shorter than 32 (hucucha known only from Siberia Cloud Forest at elevation 2800 m) ... *Oxymycterus hiska or Oxymycterus hucucha Large, head plus body longer than 120; skull longer than 32 ............. 6 6(5) Larger, occipitonasal length of skull more than 36 mm in adults; generally below 2500 m elevation ... Oxymycterus inca Smaller, length of skull less than 36; gen- erally above 2500 m elevation .... .......... .Oxymycteru s paramensis so NO. 231 ANDERSON: MAMMALS OF BOLIVIA _~~~~~~320 ~~~~~~~~~3 _ e - 7(4) Tail and toes not whitish, pelage generally dark; whitish spot on chin; smaller, length of skull less than 30 mm; nasals not extending behind front of orbit (fig. 308A); interparietal small (fig. 308B), less than 2 mm; interorbital constriction more marked (fig. 308C) .......... .................. Akodon mimus End of tail whitish, hairs on toes whitish and contrasting with darker hairs on rest of foot and on dorsal part of head plus body; larger, length of skull more than 30; nasals extending posteriorly behind front of orbit (fig. 309A); interparietal large (fig. 309B), more than 2 mm front to back; interorbital constriction scarce- ly apparent (fig. 309C)............ ................. Lenoxus apicalis 8(3) Molars, especially Ml, "pentalopho- dont," with a distinctly connected me- soloph and mesostyle (fig. 310A), ten- dency toward enamel islands (fig. 311B) in adults; tail longer than head plus body length ..... Oryzomyini, 9 Molars "tetralophodont," without a con- nected mesoloph and mesostyle, teeth simple (fig. 312) or complex (fig. 313), tend not to have enamel islands; tail length variable ............... 32 9(8) Fur spiny or of bristles, head plus body length usually less than 95 mm, skull length less than 25; forests in lowlands up to at least 1500 m elevation ...... ... . . ... . . . . . . . N e a c o m y s s p in o s u s Fur not spiny or of bristles; small to large ............................. .1 0 10(9) Palate short, antenror margin of meso- pterygoid fossa lies anterior to line con- necting the posterior margin of molar alveoli (fig. 314); not reaching posterior part of tooth rows; without lateral pits in posterior part, zygomatic notch shal- low (fig. 316) ........ ......... 11 Palate reaching behind posterior part of tooth rows (fig. 315A); with conspicu- ous lateral pits (fig. 315B) except per- haps in Kunsia; zygomatic notch shal- low or deep (fig. 317) ..... ..... 16 11(10) Feet relatively wide, short, and with strong curved claws (fig. 322); dark patch on top of feet contrasting with paler toes and sides (fig. 319); tail pen- icillate (fig. 321) ................ ........... .Rhipidomy s (in part), 12 Feet relatively long and slender, and with weaker, less curved claws (fig. 318); tail not penicillate (fig. 320) ......... ... . . . . . . . . . . . . . . T h o m a s o m y s , 1 3 12(11) Smaller, head plus body length about 170 mm; length of skull about 29.5 .... ............... .Rhipidom ys couesi Larger, head plus body length about 200; skull about 42 .................. ... . . .... . R h ip id o m y s le u c o d a c ty lu s 13(11) Large, head plus body longer than 140 mm; skull length more than 34 .... ... ... . . . . . . . . T h o m a s o m y s a u r e u s Smaller, head plus body shorter than 140; skull shorter than 34 ........... 14 14(13) Larger, head plus body longer than 120 mm; skull longer than 31; hind foot lon- ger than 31; dorsum blackish, velvety .............. .Thomasom ys ladewi Smaller, head plus body shorter than 120; skull shorter than 31; hind foot shorter than 31; dorsum brown ......... 15 15(14) Bullae smaller (fig. 323A); parapterygoid fossa narrower (fig. 323B); venter darker; dorsum brown ............ .............. .Thomasomy s daphne Bullae larger (fig. 324); parapterygoid fossa wider (fig. 324); venter paler; dor- sum reddish brown ............... ............... .Thomasom ys oreas 16(10) Tail penicillate (fig. 321); outer surface of ears and dorsal surface of tail blackish, gray patches on top of feet (fig. 319), 1997 51 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 327 325\ 329 3 33328 330 332 333- 7~~~~~~~~~ \ 34 335 '336 338 contrasting with toes and sides of feet; head plus body length about 110 mm ................ Rhipidomys nitela Tail lightly haired, but not penicillate (fig. 320); ears and tail variable in color; feet without dark patches; size variable, small to large .............. 17 17(16) Larger, head plus body of adults longer than 160 mm; hind foot longer than 40; cheekteeth tending to become more or less flat-crowned early; the outer folds of upper molars isolated, or practically so, on crown surfaces as islands, early in life (figs. 325, 326) .......... 18 Smaller, head plus body shorter than 160; hind foot shorter than 40; cheekteeth not tending to become flat-crowned un- til late in life; cusps usually traceable; less tendency for isolation of outer folds as enamel islands .............. 19 18(17) Outer folds of upper molars isolate as broad islands (fig. 325); general dental pattern simple; M1 and M2 with not more than two isolated islands each . . ............... Kunsia tomentosus Outer folds of upper molars isolate as nar- row islands (fig. 326); general dental pattern more complex; MI and M2 eventually with three or four isolated is- lands each ..... Nectomys squamipes 19(17) Hind feet shorter and broader (fig. 322), D5 short; coronoid process of mandible longer (fig. 327); zygomatic notch shal- lower (fig. 329A) ..... Oecomys, 20 Hind feet longer and narrower (fig. 318), D5 longer; coronoid process shorter (fig. 328); zygomatic notch deeper (fig. 330) ... ........ 22 20(19) Venter whitish and sharply delineated from lateral and dorsal pelage; smaller, hind foot shorter than 24 mm; occipi- tonasal length less than 28.5; head plus body shorter than 115; alveolar length of upper molar tooth row less than 4.5 ... . . ..... . .... . . O e c o m y s b ic o lo r Venter not both whitish and sharply delin- eated; larger, hind foot longer than 24; occipitonasal length more than 28.5; head plus body longer than 115; alve- olar length of upper molar tooth row more than 4.5 .............. 21 21(20) Dorsal pelage grayish, grizzled or with mixed colors, general appearance not red- dish but sometimes tinged with yellowish, venter whitish and tinged with ochra- ceous or pinkish; skull larger at a com- parable age, interorbital region narrower (fig. 329); zygomatic plate wider (fig. 331A); anterior spine of zygomata longer (fig. 329B) ....... Oecomys mamorae Dorsal pelage usually somewhat reddish, mixed with black hairs, venter not as above, but buffy or with much basal gray showing; skull smaller at a com- parable age, interorbital region relative- ly broader (figs. 332, 333A, 334); zy- gomatic plate narrower (figs. 332, 333, 334), anterior spine shorter (fig. 334A) ................. .Oecomys roberti 22(19) Size small, head plus body shorter than 100 mm; hind foot generally shorter than 28 mm; skull shorter than 28 ... ... . ... . . . . . . . ... . . . . . . . . . . . 2 3 Size larger, head plus body longer than 100; hind foot generally longer than 28; skull longer than 28 ...... ...... 27 23(22) Size small, head plus body length gener- ally less than 80 mm; hind foot small, 20-21; dorsum very dark brown, ears dark brown; venter dark gray; skull (fig. 335) with shallow zygomatic notch; rel- atively narrow rostrum; expanded zy- gomatic arches; evenly rounded brain- case .Microryzomys minutus Head plus body length generally greater than 80; hind foot small to large (20- 52 NO. 231 ANDERSON: MAMMALS OF BOLIVIA 28); dorsum golden brown to brown, rarely very dark; venter white to gray, hairs with white tips; skull with well- developed zygomatic notch (figs. 336- 338); rostrum not especially narrow; zy- gomatic arches not greatly expanded; braincase not evenly rounded ..... ................. Oligoryzomys, 24 24(23) Tail relatively short, generally less than 120% of head and body length; tail 75- 116 mm long; hind foot generally short- er than 23; venter usually with a buffy wash; maxillary tooth row delicate, generally shorter than 3.31 ....... ............. Oligoryzomys microtis Tail relatively long, greater than 130% of head plus body length; tail generally longer than 120%; hind foot usually longer than 24; venter white or gray, rarely with a buffy wash; maxillary tooth row not obviously delicate, gen- erally greater than 3.3 .......... 25 25(24) Tail generally greater than 150% of head plus body length; dorsum medium brown with a grizzled appearance, hairs long and fluffy; interorbital region par- allel-sided (fig. 337); occurring at mod- erate to high elevations ......... 26 Tail generally less than 150% of head plus body length; dorsum tending to rufous, lined with black hairs; hairs moderately long, not fluffy; venter white; interor- bital region of skull with posteriorly di- vergent sides (fig. 338); occurring at low elevations in southeastern Bolivia .......... Oligoryzomys chacoensis 26(25) Darker, bottom of tail slightly paler than top; pelage shorter, width of basal gray band when hair parted on rump about 4.5 mm, total length of hair about 5.5 ........... Oligoryzomys destructor Paler, tail noticeably bicolored, bottom whitish to gray; pelage longer and softer, width of basal gray band when hair parted on rump about 5.5, total length of hair about 7.5 ........... ........... Oligoryzomys flavescens 27(22) Palate very long, extending well past the posterior edge of M3 (fig. 339A); zy- gomatic notch deep (fig. 340A); sphen- opalatine vacuities present (fig. 339B); deep lower jaw with a long coronoid process (fig. 341); teeth when moder- ately worn as shown in fig. 342 ..... ............ Pseudoryzomys simplex Palate not so long, extending just past M3 (figs. 343-346); zygomatic notch less deeply excised (as in fig. 347, or shal- 347 iI 349 350 352 lower); sphenopalatine vacuities small or absent (figs. 343, 344A, 345, 346); coronoid process not so long (fig. 327) ........ .(als o see text) Oryzomys, 28 28(27) Larger, head plus body longer than 145 mm, occipitonasal length greater than 35; hind foot longer than 33 ..... 29 Smaller, head plus body shorter than 145; occipitonasal length less than 35; hind foot shorter than 33 ............ 30 29(28) Reddish, darker, tail dark below, ears blacker; venter whitish in general ap- pearance but bases of hairs blacker; in- cisive foramina broader (fig. 343A); molar teeth larger (fig. 343B) ..... ... .. .. . . . . ... . . . O r y z o m y s l e v ip e s Yellowish, paler, tail paler below, ears less blackish; venter more buffy and bases of hairs gray; incisive foramina rela- tively narrow and longer (fig. 344C); teeth smaller (fig. 344B) .......... ... ........ . . . . O ry z o m y s s u b fl a v u s 30(28) Tail relatively longer, whitish below on basal part and contrasting strongly with darker dorsal part; incisive foramina longer (fig. 346); anterior zygomatic notch relatively deep and wide (fig. 347); squamoso-alisphenoid groove present (fig. 348) .. Oryzomys nitidus 1997 53 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tail relatively shorter, slightly paler be- low, but less contrasting with darkness of dorsal part; incisive foramina shorter (fig. 345); anterior zygomatic notch not so deep and wide; groove absent 31 31(30) Venter paler; enamel island in M2 absent (fig. 349); occurring in the lowlands be- low 500 m elevation ............. ................. Oryzomys capito Venter darker gray; enamel island in M2 present (fig. 350); occurring in the Yun- gas above 1000 m elevation ...... .............. Oryzomys yunganus 32(8) Dentition rather weak, molars narrow (figs. 312, 313), the folds usually not approaching each other and not well marked; teeth wear early, simplifying structure; parapterygoid fossa relatively narrow (fig. 351A), and mesopterygoid fossa relatively broad (fig. 351B), inter- parietal narrow (fig. 353) ......... ................... (akodonts), 33 Dentition not as above; parapterygoid fos- sa relatively broad (fig. 352), meso- pterygoid fossa relatively narrow; inter- parietal not narrow (fig. 354) ...... ....... (phyllotines, sigmodonts), 46 33(32) Nasals relatively short and upper incisors relatively procumbent (fig. 355A); ros- trum short and less sharply set off from the slope of the anterior part of zygo- matic arch; supraorbital borders poste- riorly divergent and angular to beaded (fig. 355B); anterior wall of braincase tending to angularity (fig. 355C); par- apterygoid fossa narrow (fig. 356C) . . ... . . . . . . . . . . . . . . . . . . B o l o m y s , 3 4 Nasals longer and incisors not procum- bent (fig. 353); rostrum not so short and usually more distinct; supraorbital edg- es less strongly divergent posteriorly, rounded or slightly angular; paraptery- goid fossa not so narrow (fig. 351) ... ... . . . . . . . . . . . . . . . . . . A k o d o n , 3 6 34(33) Smaller, head plus body shorter than 110 mm; hind foot shorter than 22; crown length of tooth row less than 4.5; pelage paler, with ochraceous hues and whitish venter; inhabiting highlands of the northern altiplano and valles areas ... ................ .Bolomys amoenus Larger, head plus body longer than 110; hind foot longer than 22; tooth row (fig. 356A) longer than 4.5; pelage darker gray or blackish ............. 36 35(34) Pelage dark, venter gray, feet blackish; upper incisors less procumbent and yel- lower; incisive foramen wider, shorter, not constricted posteriorly (fig. 356B); inhabiting lowlands below 800 m ele- vation .. Bolomys lenguarum Pelage not so dark, venter ochraceous, feet dark gray; incisors more procum- bent, whiter; incisive foramen narrow, longer, constricted posteriorly (fig. 357) ... . . . . . . ... . . . . . . B o lo m y s la c te n s 36(33) Smaller, crown length of maxillary tooth row usually less than 4.3 mm; greatest length of adult skull usually less than 28 ..... .......... 37 Larger, tooth row usually longer than 4.3; adult skull usually longer than 26 41 37(36) Bullae enlarged (fig. 358) ..... .... 38 Bullae not especially enlarged (fig. 359) ... . . . . . . .. .. .. . . . . . . . . . . . . . . 3 9 38(37) Pale and ochraceous, venter gray with buffy wash; smaller, maxillary tooth row shorter than 3.6 mm .......... ... . . . . . . . . . . . . C h r o e o m y s a n d in u s Darker, not ochraceous, dorsum with mix- ture of darker and paler hairs, venter whitish; maxillary tooth row (dimen- sion shown in fig. 356A) longer than 3.6 mm ......... Akodon albiventer 39(37) Small, hind foot shorter than 16 mm; greatest length of skull less than 24; maxillary tooth row shorter than 3.5; 54 NO. 231 ANDERSON: MAMMALS OF BOLIVIA living above 3000 m elevation .... ................ Akodon lutescens Larger, hind foot longer than 16; greatest length of skull 22 to 26; some living as low as 2000 m elevation ........ 40 40(39) Darker brown dorsum, dark gray venter without buffy wash; anterior margin of zygomatic plate sloping (fig. 360) .... ................ Akodon subfuscus Paler brown dorsum, buffy or ochraceous venter; anterior margin of zygomatic plate vertical (fig. 361) ....... ............... Akodon boliviensis 41(36) Colorful, white spots at base of ear, white venter contrasts with dark dorsum, ful- vous on nose, hind feet, and tail; inter- orbital region hour glass-shaped (fig. 362); bulla large (fig. 363); inhabiting higher elevations, above 3000 m ..... ................ Chroeomys jelskii Not colorful, relatively drab brownish or blackish; bulla small; generally at ele- vations below 3000 m ..... ..... 42 42(41) Blackish or dark brown, venter not greatly contrasting with dorsum; hairs relative- ly short, soft; tail not obviously or sharply bicolored ....... ....... 43 Not so dark, hairs generally longer, mix- ture of dark and pale hairs more notice- able on dorsum; tail more obviously bi- colored; venter pale gray ......... .... Akodon varius group (including A. varius, A. pervalens, A. toba, A. sylvanus, A. simulator) 43(42) Long rostrum (fig. 364); zygomatic arch and plate smaller anteriorly ...... 44 Normal rostrum (fig. 365, shows A. dayi) ............................. 45 44(43) Pelage darker; braincase less inflated, breadth of braincase less than 12 mm; smaller, condyloincisive length of skull less than 24.3; rostrum not so long ... .................. Akodon fumeus Pelage not so dark; braincase more inflat- ed, breadth of braincase more than 12; larger, condyloincisive length of skull generally more than 24.3; rostrum lon- ger and narrower ... Akodon siberiae 45(43) Hind foot longer than 22 mm (up to 30) .................... Akodon dayi Hind foot shorter than 27 .......... .................. Akodon aerosus 46(32) Molars with compressed folds (figs. 366, 367), occlusal surface flat; paraptery- goid fossa relatively deep (fig. 371) ... ............................. 47 Molars without compressed folds (figs. 368, 369), occlusal surface flat, cuspi- date, or some intermediate condition; parapterygoid fossa not so deep . . 48 47(46) Nose conspicuously rust-colored; hind foot about 25 mm long; upper incisors clearly grooved (fig. 370, showing a cross section of teeth); upper tooth row as in fig. 366; deep posterior palatine pits (fig. 371B); zygomatic notch rela- tively wide (fig. 372A); nasals widely flared anteriorly (fig. 372B); interorbital region unusually narrow (fig. 372C); jaw deep and coronoid process small (fig. 374); occurring at elevations above 2500 m ... Neotomys ebriosus Nose not conspicuously rust-colored; hind foot longer than 35; upper incisors not grooved; upper tooth row as in fig. 367; without deep palatal pits; zygomatic notch not so wide; nasals not widely flared; interorbital region not so narrow (fig. 373); jaw relatively slender (fig. 375), coronoid process curved and pointed; inhabiting elevations below 2500 m ... Holochilus sciureus 48(46) Tail variable from longer than to slightly shorter than head plus body length (the former more common) and with distinct- ly penicillate tip (less hairy than fig. 321, but more than in fig. 320) ......... 49 1997 55 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tail shorter than, or occasionally equal to, head plus body length; without a dis- tinctly penicillate tip (fig. 320) ... 54 49(48) Skull with constricted interorbital region and greatly flared zygomatic arches (fig. 376); teeth high-crowned; bullae less inflated (figs. 378, 379) Phyllotis, 50 Interorbital region posteriorly divergent (fig. 377), occasionally ridged; zygo- matic arches flared or not; teeth less high-crowned; bullae generally more inflated (fig. 380) ........... 51 50(49) Posteropalatal pits small and usually lat- eral to mesopterygoid fossa (fig. 378A); bullar tubes relatively long and tubular (fig. 378B); nasals usually pointed be- hind and extending behind the premax- illaries (fig. 381); ear smaller; tail less hairy ........... Phyllotis osilae [specimens from populations assigned to two other species, P. caprinus and P. wolffsohni will probably key out here to P. osilae; see text for other characters]. Posteropalatal pits anterior to fossa (fig. 379A); bullar tubes not unusually long and tubular (fig. 379); nasals usually less pointed and not extending behind premaxillae (fig. 382); ears large; hair- ier tail, with or without pectoral streak; rostrum not heavy ............... ... (formerly in P. darwini), Phyllotis chilensis or P. xanthopygus 51(49) Palms and soles with hairy pads (fig. 383); head plus body shorter than 100 mm; interorbital region weakly diver- gent posteriorly (fig. 384), not ridged; teeth low-crowned ............... .. ......... Eligmodontia puerulus Palms and soles without distinct hairy pads; size medium, head plus body lon- ger than 100; interorbital region strong- ly divergent posteriorly and with edges slightly elevated (fig. 377) ........ .................... Graomys, 52 52(51) Ventral pelage white, sharply demarcated from dorsal color; ear with white spot below its base; teeth low-crowned, tending to form lophs (fig. 385); anter- ocone may be divided (fig. 388A); no alisphenoid strut (fig. 390) ........ ................ Graomys pearsoni Ventral pelage gray; no conspicuous white spot at base of ear; teeth high-crowned (fig. 386), tending not to form lophs (fig. 387); anterocone entire (fig. 389); alis- phenoid strut present (fig. 391A)... .53 53(52) Olivaceous (as in juvenals of G. griseofla- vus) to rufous; hairs of ventral surface dark basally; bullae smaller; incisive fo- ramina longer; M3 relatively smaller (fig. 392); 2n = 28 .............. ... . . ... ....... G r a o m y s d o m o r u m Buffy to tawny; basal parts of hairs of un- derparts white to plumbeous; bullae larger; incisive foramina shorter; M3 relatively larger (fig. 393); 2n = 36-38 ... . . ...... . G ra o m y s g rise o fl a v u s 54(48) Teeth high-crowned, tendency to be pris- matic; anterocone entire (figs. 368, 369); greatly flared zygomatic arches; interorbital region greatly constricted (fig. 394) .... ...... 55 Teeth low-crowned (fig. 395), not prismat- ic; divided anterocone (fig. 388, shows C. callosus); zygomatic arches not great- ly flared; interorbital region generally not constricted ........ Calomys, 60 55(54) Upper molar teeth as in fig. 369; ml as in fig. 396; rostrum as in fig. 397; large (head plus body about 160 mm long in adults); distinct pattern of black, gray, and white markings .............. ... ... . . . . . . . C h in c h illu la s a h a m a e Teeth and rostrum not as above; size mod- erate; lacking distinct pattern of black, gray, and white markings ........ 56 56(55) Size moderate, head plus body usually NO. 231 56 ANDERSON: MAMMALS OF BOLIVIA longer than 140 mm in adults; pelage relatively uniform gray; rostrum as in fig. 302; first lower molar exceptionally complex (fig. 398); teeth prismatic (fig. 368) ........... Andinomys edax Size smaller, head plus body shorter than 140; pelage various; rostrum not as above; ml not as above; teeth not pris- matic ........... 57 57(56) Skull much bowed in dorsal profile (fig. 399); anterior roots of zygomatic arch widely spreading (fig. 394 shows this species) ........ Galenomys garleppi Skull less bowed; anterior roots of zygo- matic arch less widely spreading (fig. 400, shows A. sublimus) ....... 58 58(57) Tail shorter than 70 mm; upper incisors sometimes faintly grooved, somewhat proodont, white or pale yellow .... .............. Auliscomys sublimis Tail longer than 70; upper incisors slightly yellower ... ........ 59 59(58) Upper incisors with longitudinal grooves, yellow or orange, not proodont (fig. 401); head plus body shorter than 125 mm; ear length from notch shorter than 25; soles of feet pale ............. ................ Auliscomys pictus Upper incisors without longitudinal grooves, pale yellow, distinctly proo- dont (fig. 402); head plus body of adults longer than 125; ears longer than 25; well clothed with hair and with tuft of long hair at base; soles of hind feet blackish .... Auliscomys boliviensis 60(54) Edges of supraorbital region divergent posteriorly (fig. 377) and with distinct ledges in adults ..... .......... 61 Edges of interorbital region parallel or di- vergent posteriorly, occasionally with slight beading, but never heavy ridges ............................. 63 61(60) Greatest length of skull in adults more than 24 mm; hind foot longer than 20; max- illary tooth row generally longer than 4.1; head plus body length generally greater than 90; total length greater than 200; occurring at moderate elevations 1000 to 2700 m.... Calomys venustus Greatest length of skull less than 26; hind foot shorter than 21; maxillary tooth row shorter than 3.8; head plus body length less than 90 ............. 62 62(61) Maxillary tooth row generally 3.7 to 4.0 mm; greatest length of skull of adults generally greater than 24 mm; total length generally greater than 170 mm and less than 210 mm; venter light to dark gray, no reddish tint above or be- low; alisphenoid strut not generally present, 2n = 50 . . Calomys callosus Maxillary tooth row generally 3.2 to 3.6; greatest length of skull of adults gen- erally less than 24 mm; total length gen- erally less than above; venter often dark and body frequently with a reddish or chestnut color; alisphenoid strut usually present, 2n = 66 .... Calomys tener 63(60) Tail shorter than 60 mm .......... 64 Tail longer than 60, may be equal to head plus body length ............... 65 64(63) Dorsal pelage marbled in appearance, fluffy; hairs of venter gray-based; tail very short, less than one-half of head plus body length; rostrum narrow; max- illary tooth row longer than 3.2 mm; occurring at elevations above 3000 m ................. .Calomy s lepidus Dorsal pelage not marbled, short hairs; venter white; tail more than one-half of head plus body length; rostrum not nar- rowed; maxillary tooth row shorter than 3.3; occurring in the Chaco of south- eastern Bolivia, at elevations of less than 1000 m ....... Calomys laucha 65(63) Tail approximately equal to head plus body length; dorsum golden brown lined with black hairs; sides of interor- bital region divergent posteriorly; oc- curring at moderate to high elevations (2000 to 3500 m) in the eastern slopes .............. .Calomy s musculinus Tail shorter than head plus body; dorsum brown; sides of interorbital region par- allel; occurring at high elevations in the vicinity of Lake Titicaca ......... ................ *Calomys sorellus KEY 24 GENERA AND SPECIES OF ECHIMYIDAE IN BOLIVIA 1 Cheekteeth broader than 3 mm; reentrant folds deep and persistent (fig. 403A); pal- ate constricted anteriorly to less than 2 mm breadth (fig. 403B); digits relatively long and slender; claws of forefeet resem- ble keeled nails over a pad about as long as the claw .... ... Subfamily Dactylomyinae, Dactylomys boliviensis Cheekteeth narrower than 3 mm; reentrant folds may isolate as narrow islands in adults (fig. 61B); palate narrower anteri- orly but not greatly constricted, breadth between alveoli at least 2 mm; digits and 57 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY claws of forefeet not as described above .......... (also see Mesomys in text) 2 2(1) Cheekteeth smaller, with narrow folds that typically become isolated as islands in adult; length of first upper cheektooth less than 2.5 mm; occlusal length of upper mo- lariform tooth row less than 10; feet not specially adapted for climbing, narrower, width of hind foot less than 10; claws less curved, blunter ....... Proechimys sp. [at least five species occur in Bolivia, see text for further details] Cheekteeth heavier, with more persistent folds, the general effect complex, folds isolating late or not at all; length of occlu- sal surface of first upper cheektooth more than 2.5; occlusal length of upper molar- iform tooth row more than 10; feet adapt- ed for arboreal life, broader, width of hind foot more than 10 in adults; claws more curved, sharper ....... ..... 3 3(2) Fur soft, tail abundantly haired, almost bushy; base of tail reddish brown, distal two-thirds black; top of head has paler brownish patch contrasting with blackish surrounding areas .... Isothrix bistriata Fur bristly or spiny, scalation of tail evident, not well haired; color not as above on tail and head .... ........ 4 4(3) Area around nose chestnut-colored; body rather uniform in color; long hairs grow- ing from tips of ears; tail with scattered hairs, scales mostly visible; spines small- er; tail about as long as head plus body, without distal tuft .................. ............... Echimys didelphoides Area around nose not chestnut-colored; without long hairs on tips of ears; spines heavier; tail longer than head plus body, with tuft at end . . *Echimys macrurus KEY 25 GENERA AND SPECIES OF ABROCOMIDAE AND OCTODONTIDAE IN BOLIVIA 1 Ears large, longer than 15 mm; forefeet not especially enlarged, longest claw shorter than 5; skull not heavily ridged ..... 2 Ears small, shorter than 10; forefeet en- larged for digging (fig. 404), longest claw more than 5; skull heavily ridged, especially the zygomatic arches ..... Subfamily Ctenomyinae, Ctenomys, 4 [also see text and map; additional taxa re- main to be described, and these are not included in this key] 2(1) Lower teeth simplified (fig. 406); tail bushy; rostrum less slender (fig. 408) . . ............. .Octodontomy s gliroides Lower teeth complex (fig. 405); tail haired but not bushy; rostrum slender (fig. 407) . . Family Abrocomidae, Abrocoma, 3 3(2) Tail short, less than 50% of length of head plus body; pelage pale gray; bullae enor- mous, projecting posteriorly beyond oc- cipital (fig. 409), gap separating bullae at midline less than 1 mm; known from Tarija .......... Abrocoma cinerea Tail long, more than 50% of length of head plus body; pelage brownish gray; bullae large, not projecting beyond occipital (fig. 407), gap more than 1 mm; known from western Santa Cruz ........... ... . . . . . . .. . . ..Ab r o c o m a b o l i v i e n s i s 4(1) Size large, hind foot longer than 55 mm; greatest length of skull more than 65; pelage coarse, shaggy, reddish brown; in- habiting the Gran Chaco in southeastern Bolivia ......... Ctenomys conoveri Size smaller, hind foot shorter than 55; greatest length of skull less than 65; pel- age not as above ........ 5... 5(4) Pelage short, few hairs of dorsum longer than 15 mm; nasal bones relatively short, broad, and truncated anteriorly (fig. 410) ........... 6 Pelage long, most hairs of dorsum longer than 15; nasal bones longer, narrower, and not truncated anteriorly (fig. 411) .....8 6(5) Size small, hind foot shorter than 35 mm; head plus body length less than 175; greatest length of skull less than 45; ros- trum short (fig. 410); inhabiting eastern Santa Cruz ...... Ctenomys minutus Size larger, hind foot longer than 35; head plus body length more than 175; greatest length of skull more than 45 ....... 7 7(6) Pelage blackish; postorbital processes far- ther forward (fig. 412); inhabiting western Santa Cruz ...... Ctenomys steinbachi Pelage brown; postorbital processes farther back (fig. 413); widely ranging in Santa Cruz ...... Ctenomys boliviensis 8(5) Pelage pale reddish tan (in unworn pelage); bullae large (fig. 414) ............. ................. .Ctenomys opimus Pelage varying from tan to blackish; bullae not expanded (fig. 415) ........... 9 9(8) Dorsal pelage pale reddish tan; upper in- cisors unusually procumbent (fig. 416) and pale; occurring to the south of Lake Titicaca .. . Ctenomys leucodon Dorsal pelage brown to blackish; upper in- cisors not unusually procumbent (figs. 58 NO. 231 ANDERSON: MAMMALS OF BOLIVIA 417, 418) and pale; occurring in the Val- les Area from the city of Potosi south- eastward ..... 10 10(9) Pelage usually with distinct cinnamon hue; width of an upper incisor greater than 3.3 mm; incisors more procumbent (fig. 417); postorbital process of jugal more anterior (fig. 419); braincase less vaulted (fig. 421) ... .. Ctenomys lewisi Pelage dark brown; width of an upper in- cisor less than 3.3; incisors less procum- bent (fig. 418); postorbital process more posterior (fig. 420); braincase more vaulted (fig. 422) .... Ctenomys frater KEY 26 GENERA AND SPECIES OF CHINCHILLIDAE IN BOLIVIA 1 Pale gray, with neither yellowish belly nor black and white markings on head; length of skull less than 65 mm; interorbital breadth less than 15; bullae inflated, visi- ble as conspicuous areas (width more than 10) on top of skull (fig. 423) and nearly meeting at midline below; probably ex- tinct in Bolivia . . Chinchilla chinchilla Color not as above; larger, length of head plus body more than 350; skull longer than 65; interorbital breadth more than 15; bullae less inflated, not visible as large ar- eas on top of skull, and not nearly meeting (separation more than 4) below ...... 2 2(1) Black and white areas on head (fig. 424), head bulky, belly whitish; tail relatively shorter, length usually less than 200 mm; larger, head plus body more than 435; oc- cipitonasal length of skull more than 90; bullae less inflated and paroccipital pro- cesses projecting prominently below (fig. 425); inhabitant of Gran Chaco region at elevations of less than 500 m ....... ... . . . . . . . ... . . L a g o s to m u s m a x im u s No striking markings on head, belly yellow- ish; more delicate in build; tail longer, length usually more than 200; smaller, head plus body shorter than 435; occipi- tonasal length less than 90; bullae more inflated and paroccipital processes barely projecting below bullae (fig. 426); inhab- itant of rocky areas in high mountains . . ... . . . . . . . ... . . . . L a g i d iu m v is c a c c ia KEY 27 GENERA AND SPECIES OF CAVIIDAE IN BOLIVIA I Larger, head plus body longer than 350 mm; skull longer than 80 mm; ears longer than 90; relatively long-limbed, radius longer than humerus, hind foot longer than 70; nasals attenuated (fig. 427A); interorbital region broad, breadth at constriction more than 18 (fig. 427); posterior margin of bony palate a long "V" with anteriormost point at about the level of front of third molariform tooth ........ Subfamily Dolichotinae, Dolichotis salinicola Smaller, head plus body shorter than 350; skull shorter than 80; ears shorter than 90; relatively short limbed, radius shorter than humerus, hind foot shorter than 70; nasals not so attenuated (fig. 428A); interorbital region narrower (fig. 428B), at constric- tion less than 18; posterior margin of pal- ate a broader "U" with anteriormost point at level of middle of fourth (last) molari- form tooth (figs. 441, 442) ......... .............. .Subfami ly Caviinae, 2 2(1) Teeth less simplified, breadth of occlusal surface more than 2.5 mm, posterior lobe of upper cheekteeth with clear and deep outer reentrant fold (fig. 429); averages larger, head plus body longer than 215; skull length 50 or more, zygomatic breadth greater than 31; large rounded fo- 59 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY ramen in lacrimal (fig. 433); rostrum deep, incisors relatively recumbent (fig. 436); wild or domesticated ... Cavia tschudii Teeth more simplified, breadth of occlusal surface less than 2.5, posterior lobe of up- per cheekteeth without clear and deep out- er reentrant fold (fig. 430); averages smaller, head plus body shorter than 270; skull length less than 60, zygomatic breadth less than 34; lacrimal not with large rounded foramen (figs. 431 and 432); rostrum not so deep, incisors may be more procumbent (figs. 434 and 435); generally paler in pelage .... ....... 3 3(2) Incisors pigmented orange at front, each up- per incisor broader than 1.4 mm; incisors relatively recumbent (fig. 435); orbital branch of maxilla completely interrupted by lacrimal (fig. 437); lacrimal foramen relatively small and less lateral in position at front of orbit (fig. 432); skull less bowed in profile (fig. 439); length of pa- rietal usually more than 15; not so pale and grayish, more agouti appearance in pelage; gular skin with bare area in middle .......................... Galea, 4 Incisors not pigmented but whitish and nar- row, each upper incisor less than 1.4 wide; incisors procumbent (fig. 434); orbital branch of maxilla not completely inter- rupted (fig. 438); lacrimal foramen rela- tively large and more lateral (fig. 431); skull bowed in dorsal profile (fig. 440); bullae relatively larger (fig. 440); parietal length 15 or less; feet heavier, breadth across pad of hind foot more than 8, across front foot more than 6; pale and grayish; gular skin completely covered with hair .............. Microcavia, 5 4(3) Whitish eye ring larger; mesopterygoid fossa deep and narrow, with more or less di- verging sides (fig. 441); first prism of Ml smaller than second; zygomatic arches spread widely (fig. 443); interorbital breadth averages about 16 mm; averages smaller, occipitonasal length of skull about 52; hind foot usually about 39 ...... .................. .Galea musteloides Eye ring smaller; mesopterygoid fossa broad and with parallel sides (fig. 442); zygo- matic arches spread less widely (fig. 444A); interorbital constriction narrower (fig. 444B); averages larger, occipitonasal length of skull about 57; hind foot of adults usually about 42 ... Galea spixii 5(3) Skull smaller, greatest length (average for adults) about 44.3 mm; dorsal profile of skull more rounded (fig. 440A); meso- pterygoid fossa pointed (fig. 446); incisors proodont (fig. 434) ... Microcavia niata Skull larger, greatest length (average for adults) about 46.8; profile straighter (fig. 440B); mesopterygoid fossa rounded (fig. 445); incisors orthodont (similar to those in fig. 435); occurs in Argentina, not yet reported from Bolivia .............. ............... .*M icrocavia australis KEY 28 SPECIES OF ERETHIZONTIDAE IN BOLIVIA Quills substantial, with white tips that contrast with dark areas (fig. 447A); larger, occipiton- asal length of adult skull more than 85 mm; alveolar length of upper tooth row more than 20.5; forehead highly inflated (fig. 448A); pos- terior margin of palate broadly U-shaped (fig. 449A) ............. Coendou prehensilis Quills more variable in size and in ratios of length to diameter, quills on shoulders and surrounding areas relatively long and slender (fig. 447B); quills have dark tips; smaller, occipitonasal length less than 90; alveolar length of upper molar tooth row less than 20.5; forehead less inflated (fig. 448B); posterior margin of palate somewhat V-shaped (fig. 449B); width of upper incisor less than 3.2 ...... Coendou bicolor NO. 231 60 ANDERSON: MAMMALS OF BOLIVIA METHODS AND MATERIALS I have seen or have records of more than 36,900 specimens of Bolivian mammals. A single identifiable specimen with reliable data is sufficient evidence to plot a dot on a distribution map. However, the adequacy of samples depends on the question(s) being asked. Material for study may seem abun- dant, but although Bolivia is a large and di- verse country with more than 320 species of native mammals, more species remain to be discovered. The mean number of specimens per species is about 115; however, numbers vary from 1 to 2914 and the median number is only 26. Clearly, species are not equally represented in the total sample, and most are poorly represented. Actual rarity in some cases, small ranges within Bolivia, little col- lecting in the appropriate range or habitat, 1ooo0L 0 0 0 r Pk 10 1 1 10 and inadequate methods of collecting all have contributed to this poor representation. Further indication of the situation is the num- ber of localities per species, which ranges from 1 to 169 with a median of 11. A "series" is defined for present purposes as a sample of one or more specimens of one species from one place. The number of such samples is about 7300. Thus, the average number of specimens per series is only five. Specimens in 2836 series of rodents were tal- lied for figure 450. Over half of all series consist of only one or two specimens. For a given species, the series also tend to be distributed in the same highly skewed way; the most common number of specimens in a series is one, then two. Because of the disproportionate influence of a few large val- 100 1000 NUMBER OF SPECIMENS PER SERIES Fig. 450. Graph to indicate the small size of most samples of mammals available for study in collections. A series is defined as the specimen(s) of one species from one locality. Data for 2836 series of Bolivian rodents were tabulated for this graph. The most common series size is one specimen; and more than half of all series (1583 of 2836) consist of only one or two specimens. lI lI 11-I 0 0 A * - I eIelL-!lll-- tI1 . 61 1997 I "1 717 1 F BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 0 Al co U, 0 100 1 10 100 1000 NUMBERS OF LOCALITIES 0 AND SPECIMENS 0 Fig. 451. Cumulative semilogarithmic graph showing the percentage of species of mammals rep- resented by different numbers of specimens and localities for Bolivia. For example, 80% of species are represented by more than four localities and 80% are represented by more than six specimens; the reciprocals are 20% represented by fewer than five localities and 20% by fewer than seven specimens. ues on arithmetic means, these means are greater than the medians. For example, the average number of specimens per series is five and the median is two. For bats, the av- erage number of specimens per series is 5.4; for rodents it is 5.9; and for other mammals it is 2.8. The largest series consists of 1706 specimens of Calomys callosus from San Joaquin. This species of rodent was found to carry the Machupo virus that caused a hem- orrhagic fever that killed about 10% of the people in that village during an epidemic in the early 1960s (Johnson et al., 1965). This large series of mice was collected during studies at San Joaqufn. No other series num- bering more than 1000 exists. The entire rep- resentation for only two other species ex- ceeds 1000. These are the common bats My- otis nigricans and Molossus molossus. The size of the samples of these two species also is largely the result of single series of 465 and 413, respectively, obtained in the early stages of the study of hemorrhagic fever at San Joaquin when researchers investigated the possibility that bats might be transmitting the disease. To study geographic variation, a number of series from different parts of the range of a species and consisting of at least 10 adult specimens per series are needed. The number 10 is only a rough minimal approximation. More are needed when measurements or counts are highly variable or sexual dimor- phism is pronounced. Only 14% (389 of 2836) of the series of rodents consists of 10 or more specimens. Few species are repre- sented by series adequate for this type of study and even fewer species have been crit- ically studied from this standpoint, by me or anyone else. Neither the numbers of specimens nor lo- calities per species are normally distributed; both are highly skewed, with lower values predominating. The percentages of species represented by different numbers of localities and specimens are shown in figure 451. Values at the 50th percentile are the medians of 11 localities 62 NO. 231 ANDERSON: MAMMALS OF BOLIVIA 30 20 10 (A q 1:1 XA 5 1 0 10 20 30 NUMIBER OF LOCALITIES Fig. 452. Semilogarithmic graph showing the number of localities per species of Bolivian mammals. In general, the most common number (for 24 species) is one locality, then two (23 species), then three (22 species), etc. Data for 240 species known from fewer than 30 localities are plotted; 82 additional species are known from 30 to 169 localities. For values from 30 to 39, the average number of species is 2.3; for the 40s the average is 1.7; for 50s, it is 1.5; for 60s, it is 0.3; etc. The downward trend seen in the graph continues for the scattered values above 30 localities. and 26 specimens per species. The values at any other percentile can be read approxi- mately from the graph. The area of Bolivia is 1,098,581 km2 (In- stituto Nacional de Estadistica, Atlas censal de Bolivia, 1982). The total of 36,900 spec- imens thus represents one specimen per 30 km2. In comparison, an unusually detailed North American faunal study of mammals in the state of Arizona (Hoffmeister, 1986) was based on 42,278 specimens representing 138 native species. The area of Arizona is 295,144 km2, so there was one specimen per 7 km2. Arizona is, therefore, between four and five times better represented in collec- tions of mammals and correspondingly better known than Bolivia. The number of speci- mens per species on the average in Arizona is 306, compared to 115 in Bolivia. The numbers of species that are repre- sented by different numbers of localities are summarized in figure 452. Note the predom- inance of small samples. We now have a much better idea of what species are present and their geographic dis- tributions within Bolivia than we had a few years ago, but much more remains to be learned about taxonomy and distribution. Other biological aspects, such as ecological relationships with the physical environment and with other species of both plants and an- imals, behavior, population levels and fluc- tuations, reproduction, genetics, parasitology, epidemiology, public health, conservation bi- ology, wildlife management, and economic impacts are virtually unexplored and offer exciting fields for study. I hope that the pres- ent progress report on taxonomy and distri- bution will provide some ideas and incen- tives for this future work. rI I I I I I 0e.0 0 * * 0 0 0 0- p 0 0 0 - 0 I I *- 1 0 0 I I I Le.... I I 63 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY FAUNAL ANALYSIS AND BIOGEOGRAPHY Geological, topographic, climatic, and vegetational factors are among the major en- vironmental influences on mammalian distri- butions and populations. These will be out- lined only in a general way, to provide a background for the discussion of mammalian biogeography. Some diverse mammalian habitats are illustrated in figures 453 to 469. These photos, arranged by elevation, were taken from 5000 m in the Cordillera Oriental of the Andes down to 200 m in the Ama- zonian lowlands. GEOGRAPHICAL POSITION AND TOPOGRAPHY Bolivia extends in latitude from approxi- mately 10 to 230 south of the equator. These latitudes, and great topographic relief, are major factors in maintaining diverse climates and mammalian habitats. These habitats range from humid, high (i.e., tall) tropical forest in the north to areas of perpetual snow and glaciers on the cordilleras that are rela- tively barren biologically. Bolivia can be roughly divided into lowlands and highlands. The northern and eastern two-thirds of Bo- livia lie at elevations less than 500 m above sea level, some less than 200 m. When one considers that the distance from the lowest parts of Bolivia to the Atlantic Ocean is about 2000 km via the Amazon or 1500 km via the Rio de la Plata, the limited amount of topographic relief in much of eastern South America, from the Andean foothills to the ocean, is impressive. In Bolivia, this flat- ness, in combination with considerable rain- fall, results in seasonal flooding of rivers and inundation of large tracts of forests. The other third, the highlands in the south- west, rises abruptly from the lowlands and reaches to more than 6000 m in the cordil- leras of the Andes. The eastern edge of the Andes is a relatively steep escarpment rising from 500 to more than 3000 m, at some plac- es over distances of less than 100 km. At other places, it has been dissected by erosion to yield a series of ridges and valleys. As a result of this complex topography and pre- vailing and seasonal winds and precipitation, small areas as varied as perpetually humid Fig. 453. Lake at divide on the road between Antequilla and Pelechuco (1448/6910), above elevation 5000 m, September 1980. Rocky habitat of Chroeomys jelskii and Lagidium viscacia. NO. 231 64 ANDERSON: MAMMALS OF BOLIVIA Fig. 454. Houses east of Antequilla (1452/6920), at elevation of about 4800 m, September 1982. Heavily grazed by alpacas, little vegetation evident. Habitat of Auliscomys pictus, Auliscomys sublimis, and Calomys lepidus. cloud forests and dry valleys are intermin- gled in complex ways. In addition to the peaks of the cordilleras, the high country in- cludes the relatively level altiplano. This area lies mostly in Bolivia but extends into Peru, Argentina, and Chile. In North American terms, the altiplano is roughly the size of the state of California and lies at about 13,000 ft elevation. The Altiplano is relatively dry, and aridity increases from north to south. Fig. 455. Station at La Cumbre (1622/6803), elevation 4675 m. Photographed by Tate in 1926. Habitat of Neotomys ebriosus, three species of Auliscomys, and Calomys lepidus. 65 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 456. Grassy vegetation in ungrazed area within the fence around the ruins at Tiwanacu (1633/ 6839), elevation 3840 m. Few ungrazed areas are to be seen on the Altiplano. We did not trap here but would expect to find Akodon boliviensis and Calomys lepidus. CLIMATE In the long sweep of geological history, major climatic changes have occurred. Cur- rent mammalian distributions largely reflect recent conditions and changes. A period of drought began rather abruptly about the year 1000 AD, at least in the highlands. This drought, documented from the study of ice cores from the Quelccaya ice cap in Peru and from sediment cores from Lake Titicaca, probably caused the collapse of the "politi- cally integrated, expansive state society" of Tiwanaku (Ortloff and Kolata, 1993). We can surmise that fluctuations in aridity through- out the Pleistocene and earlier allowed the ranges of various species to expand or com- pelled them to contract, although few data are available to document former ranges. Even today our knowledge of the environ- mental conditions that may restrict the ranges of species now living in Bolivia is quite lim- ited. Average annual rainfall ranges from more than 3000 mm (3056 recorded as average over a 5-year period at Chimore, elevation 300 m, Cochabamba) to less than 100 mm (51 recorded as average over a 14-year pe- riod at Chiguana, elevation 3684 m, Potosif) in the southern part of the Altiplano (Unzue- ta, 1975: 310, 312). Mean annual temperatures in degrees Cen- tigrade range from greater than 240 (24.2° in a period of 22 years at Cobija, Pando) to less than 7° (6.20 over 10 years at Penas, eleva- tion 3986 m, La Paz). VEGETATION In figure 470, I have mapped six major vegetation zones (as did Salazar et al., in press; see further discussion of their work under Faunal Analysis below). Some other useful sources of vegetational and general ecological data are Holdridge et al. (1971), Unzueta (1975), UNESCO (1981), and Gey- ger and Arze (1982). The designation of six rather than a smaller or larger number of zones is arbitrary. No zone is uniform. Local conditions, such as the small wet areas or bogs on the Altiplano known as bofedales, are important to various mammals. Large parts of the lowlands, especially in the Beni, are subject to seasonal flooding (bosques de 66 NO. 231 ANDERSON: MAMMALS OF BOLIVIA Fig. 457. Vegetation on bank of the rio Desaguadero (1740/6732), about elevation of 3700 m, August 1984. Habitat of Akodon albiventer, Phyllotis xanthopygus, and Oligoryzomys andinus. inundacion). The limited number of areas with rocks and caves in the lowlands, may be the reason that bats of the family Mor- moopidae are known only from the region near the Serrania Huanchaca. Fossorial mam- mals such as tuco-tucos favor areas with rel- atively deep and friable soils. These are just a few examples. Mostly, limiting factors for species have not been postulated except in the most general terms. Fig. 458. Vegetation, including cactus, at Pongo (1620/6756) in the Unduavi Valley at elevation 3690 m. Photographed by Tate in 1926. Habitat of Oligoryzomys sp. B, Akodon mimus, Chroeomys jelskii, and Auliscomys pictus. 67 1997 6BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY ~~~~~~~~~~~~~~~~~~~~~, ..- %Z. ..- Fig. 459. Rocky hillsides and short scattered shrubs near Tarabuco (1910/6455), elevation ca. 3300 m, July 1983. Habitat of Phyllotis xanthopygus. Fig. 460. View from the valley floor of the rio Caracato (1659/6749), elevation 2900 m, April 1982. Habitat of Octodontomys gliroides on the arid hillsides and Thylamys pallidior on arid hillside and in marsh on valley floor. Desmodus rotundus was the only bat captured here. 68 NO. 231 ANDERSON: MAMMALS OF BOLIVIA Fig. 461. Forested slopes at Nequejahuira (1619/6752), elevation 2450 m. Photographed by Tate in 1926. Habitat of Mustelafrenata, Lenoxus apicalis, Oryzomys levipes, Thomasomys aureus, and Oxymycterus paramensis. 1. Altiplano (and associated mountains and valleys). The predominant vegetation in this region is known as the puna. The puna is sunnier than the pdramos [at high ele- vations in the northern Andes]; it also has a greater daily temperature range .... [A]bove 4,000 m there is a night frost on more than 300 nights a year. From 4,700 m, there are alternating nightly frost and daily thaw throughout the year. The ecoclimatic conditions are such that plant-growth is interrupted. Moreover, the length of the dry season, added to the cold, helps to limit the growth and development of plants whose adaptation to the very harsh ecoclimatic conditions can be seen in the biological types. This adaptation is of two morphological and anatomical types. They display a disproportionate development of under- ground organs, with a reduction or absence of stems, [the presence of] cushion forms, and leaf reduction accompanied by abundant pilosity, spinescence and succulence. Lastly, many woody plants of the puna are resinous, and are thus much sought after for fuel, which explains their destruction and rarity (UNESCO, 1981: 95). The region as mapped in figure 470 includes the Andean plateau proper, the surrounding mountains (the Cordillera Occidental along the Chilean border and the Cordillera Ori- ental on the eastern side), which are devoid of vegetation on the snow-clad peaks, and various valleys. Most of the region is above 3000 m elevation. The few trees are mostly of introduced species of eucalyptus. A de- tailed study of vegetation in the vicinity of Ulla Ulla was published by Seibert (1993). Figures 453 to 459 are of localities above 3000 m elevation. 2. Yungas. These are the forested eastern slopes of the Andes mountains, which range in elevation roughly from 3000 to 500 m. "In the ... Bolivian Andes, the forests of the east facies of the mountains ... are known as the ceja de la montana .... [Sluch a range of altitude must contain several types of forest" (UNESCO, 1981: 92). The forest is mostly not deciduous. In the most humid areas there is a distinctive cloud-forest with tall ferns, deep humus on the forest floor, much moss, and trees festooned with epiphytes. These slopes are at places deeply dissected, and be- cause of prevailing winds and resulting rain- shadow effects, local areas within short dis- tances may differ widely in aridity and in vegetation. Figures 460 to 465 are of local- ities in the yungas and valles. Figure 464 shows one of the more arid valleys. 3. Tropical and Subtropical Forest (Bosque Amazonico). The forests of the low- er Yungas grade into the lowland (less than 500 m elevation) subtropical forests. The strictly tropical areas (i.e., where it never freezes) are in the northern part of the de- partment of Pando. Here are the largest trees, the greatest number of species of trees, and the trees yielding rubber and Brazil nuts. The photograph in figure 467 was taken at 350 m elevation. 4. Savannah (Savana). These are open ar- eas, largely grassland, with scattered trees or patches of forest (islas de bosque). In the southern part of the region, the forest exists in patches, as the dispersed vegetation phase. From south to north the amount of forest in- 69 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 462. Coca plantations on slopes at La Florida (1621/6746), elevation 1780 m. Photographed by Tate in 1926. Habitat of Gracilinanus agilis, Marmosops dorothea, Akodon aerosus, and Rhipidomys couesi. creases until the grassland becomes the dis- persed phase, remaining only in isolated patches known as pampas or chacos (e.g., Pampa de Heath near the Peruvian border and Pampa de Meio on the rio It6nez, Chaco Lejo). Incidentally, the term "chaco" is used also for smaller clearings in forests. When most of the trees are palms, the term "Palm Savannah" is used. The ecology of the sa- vannahs of Beni in the longer-term geologi- Fig. 463. View from Coroico (1610/6744), elevation 1715 m, May 1982. Forested or partly cleared slopes in the yungas of La Paz department. NO. 231 70 Ri unm IM I 0; W11.111- ..... ANDERSON: MAMMALS OF BOLIVIA Fig. 464. Arid vegetation east of Comarapa (1758/6429), elevation 1700 m, September 1984. Habitat of Graomys domorum, Promops nasutus, Anoura geoffroyi (feed at flowers of cacti). Fig. 465. Illustration of "cut and bum" agricultural practice commonly used in many parts of Bolivia, at about elevation 1000 m in Santa Cruz department, 1984, exact location not recorded. 1997 71 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 466. Relatively arid area at the northern edge of the Gran Chaco on the road west of Robore (1816/6007), elevation 475 m, October 1984. Habitat of Ctenomys minutus, Thrichomys apereoides, and Chaetophractus vellerosus. cal context was analyzed by Hanagarth (1993). 5. Subhumid Forest (Bosque Chiqui- tano). This region is intermediate between the wet Amazonian forests to the north and the dry Chacoan forests to the south. 6. Chaco. The trees that are present in the Chaco are smaller and scrubbier than trees farther north. Figure 466 shows a Chacoan area west of Robore. Faunal Analysis SPECIES DENSITIES: Six one-degree sample areas (A to F in fig. 470) were selected along a transect; tallies were made of numbers of species with ranges (as mapped for individ- ual species in subsequent accounts) occurring in at least part of each quadrat. These num- bers, plotted in figure 471, compare the spe- cies densities estimated among sample areas and among bats, rodents, and other mam- mals. The combined totals for all mammals and for each sample area are also shown. Clearly, for all groups, the Altiplano (area A) has a lower species density than other areas; this paucity is most extreme for bats. Ro- dents comprise 36% of all Bolivian species, bats about 31%, and species of all other or- ders combined 33%. Thus, the ratio of bats to rodents is about 1 to 1.2. However, the ratio of numbers of species of bats to num- bers of rodents in one-degree sample areas D, E, and F is about 2.5 to 1. The reason for this reversal in relative abundance is that ro- dents, on the average, have smaller geo- graphic ranges than do bats, a fact with im- plications discussed elsewhere (Anderson, 1991). The relative abundances of individuals of different species in different places and at different times are additional biological di- mensions scarcely touched upon here. Fur- ther study is needed. A graph showing the cumulative number (decade by decade) of presently recognized living native species of mammals known by reported specimens from present-day Bolivia is shown in figure 472. By 1990, the number had reached about 300. Three projections (A, B, and C) beyond that date are shown. Ad- ditional species will be from three sources: those new to science, those already known from outside of Bolivia but newly discovered within Bolivia, and those already known and NO. 231 72 ANDERSON: MAMMALS OF BOLIVIA Fig. 467. Village of Rurrenabaque (1428/6734), elevation 350 m, September 1980. This is where the rio Beni leaves the final foothills of the yungas. Habitat of Akodon dayi. named but currently regarded as subspecies or synonyms. Taxonomic revisions that lump formerly recognized species and extinctions of Bolivian populations tend to reduce the number of recognized species living in Bo- livia. The chinchilla (Chinchilla chinchilla) may now be extinct in Bolivia and the rate of extinction is likely to accelerate in the coming years. Within 20 or 30 years the rate of extinction may increase and the rate of discovery decrease enough to cause the known living Bolivian fauna to peak and be- gin to decline. Projections B and C are min- imum and maximum estimates, while A shows an increase that was certain in 1991 because of specimens already in collections and taxonomic revisions that were underway. Experience through 1994 suggests that pro- jection C is closer to reality than A or B. About 78 species are presently on the hy- pothetical list. Some of these, although listed because of their proximity to Bolivia, prob- ably will never be found in Bolivia. My own rough estimate is that about one-half of them will be found. A published faunal list (Pacheco et al., 1993) for the Manu Biosphere Reserve-lo- cated some 150 to 200 km west of the Bo- livian border in the drainage basin of the rio Madre de Dios and its left-hand tributary, the rio Manu, at elevations from 365 to 3450 i-provides interesting comparative data. Of the 190 species listed, 160 (or 84%) occur also in Bolivia. Only 30 do not, or at least they have not been found there yet. Salazar et al. (in press) discussed the bio- geography of Bolivian native mammals, us- ing published data (Anderson, 1993) and un- published data from my files. Their conclu- sions are of considerable interest. The relative density of collecting efforts in different parts of the country was mapped by Salazar et al. (in press; see also fig. 473 in which all collecting localities are shown). Areas near cities and major roads and rivers are represented best. Although the data ana- lyzed by Salazar et al. included records of more than 32,000 specimens in collections from hundreds of localities, 47% of the one- half-degree quadrats in the country were not represented by even one specimen. In a symposium on the montane forests of Peru (Young and Valencia, 1992), great bio- diversity (many species) and the occurrence of relatively small areas having high per- centages of endemic species were noted for various taxonomic groups. According to Al- wyn H. Gentry (p. 11), a single forested 73 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 468. At the edge of the Bafiados del Izo- zog (1828/6207) in the dry season, elevation 290 m, October 1984. In the wet season, large areas are inundated. Seasonal inundation occurs in many parts of the lowlands. Akodon toba and Cal- omys callosus occur in grassy habitat nearby on slightly higher ground. Panthera onca occurs in the general area. ridge (data from Ecuador but probably also true in Peru and Bolivia) can have as many as 100 locally endemic plants. Diana Silva (p. 34) noted that "very few" of the 750 spe- cies of spiders found at eight study sites in montane forests occurred at more than one site, and thus she postulated both small rang- es and high local endemism. John P. O'Neill (p. 47) commented that more than 1700 spe- cies of birds are known from Peru, that over 55% of these occur above elevations of 1000 m, that 112 species are endemic to Peru, and that 106 of these have Andean distributions. Species density and endemism in Bolivian mammals are discussed below. Biogeographic units are commonly ana- r1g. 40o. rorest on DanJ oI no itenez De- tween Ilha Floras and Larangeiras, Brazil, about elevation 200 m, 7 August 1964. The latter is at 1313/6209. Habitat of Holochilus sciureus, Ory- zomys capito, and Didelphis marsupialis. lyzed and discussed by ecologists and other biologists. Many systems or schemes for de- fining, recognizing, and naming such units have been proposed. The history, technical details, and arguments for and against the systems need not concern us here. Vegetation is often the major basis for selecting such units. The selection process is somewhat sub- jective because conditions are not uniform throughout any given unit and the boundaries are not so sharp as neat lines on a map may suggest. Even if an unambiguous basis for defining units is formulated, data needed to apply the definition are limited in quantity and accuracy. Nonetheless, the process is in- formative in a general way. Six major biogeographic units in Bolivia were delineated by Salazar et al. after con- sidering both the areas used by earlier au- thors and the limitations of the methodology. 74 NO. 231 ANDERSON: MAMMALS OF BOLIVIA 60°W lo) S Fig. 470. Map of Bolivia showing six one- degree-square sample areas (A to F) along a tran- sect from the high Altiplano to the Amazonian lowlands. Six major vegetation zones are: 1) the Altiplano and Interandean Valleys (Altiplano y Valles Interandinos); 2) Yungas (the forested slopes on the east side of the mountains); 3) Trop- ical Rain Forest (Bosque Amazonico); 4) Savan- nah (Savana); 5) Subhumid Forest (Bosques Chi- quitanos); and 6) the Chaco (Ergueta and Salazar Bravo, 1991, and other sources). The species den- sities of groups of mammals in these sample areas are shown in figure 471. The units are listed and described above. The species occurring in each unit were tallied, and the numbers of species, numbers of en- demic species, and degrees of similarity be- tween the mammalian faunas of the units were compared. The relative sizes of average adult individuals of the species and their tro- phic roles in the six units were also dis- cussed. Some differences in patterns between taxonomic groups, for example bats as com- pared to rodents, were also analyzed. Mam- malian biogeography and Bolivian conser- vation were given special consideration. Ex- cluding from the count humans and intro- duced species, 316 species of native mammals were known then from Bolivia. The numbers of species present in the six units were, respectively, 53, 114, 152, 114, 143, and 114. Thus, the Amazonian forest exhibits the greatest species diversity. Sixteen recognized species were consid- A B C D E F Fig. 471. Graph showing the numbers of spe- cies of bats, rodents, and other groups (stippled area within graph for all species at the top), and all mammals in the six sample areas shown in figure 470. ered to be Bolivian endemics (i.e. confined to Bolivia). Certainly, some of these endem- ics will be found outside of Bolivia by fur- ther collecting or will be found by further taxonomic study to intergrade with some oth- er species now recognized that occurs out- 300o U1) ILL 0 ILI m z 200O 100o 1850 1900 1950 2000 Fig. 472. Cumulative number of living spe- cies of native mammals known to inhabit Bolivia. A record of increasing knowledge. Values for fu- ture decades, predicted as discussed in text, are shown as A, B, and C. I I I _ .... * -B....~~e. : B -l/ , I ~~ ~~~I II 75 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 473. Map showing all (1259) collecting localities for mammals. Separate maps were gen- erated for bats, rodents, and other mammals. The only conspicuous difference in the pattern was that there are few localities for bats on the Alti- plano. Otherwise, the same clusters of localities near major cities and along avenues of transpor- tation, along rivers in the lowlands of northern Bolivia and major roads elsewhere, were evident on each map. This pattern is clear on the present map. side of Bolivia. Although both of these pro- cesses tend to reduce the number and per- centage of species that are thought to be Bo- livian endemics, other processes tend to increase the number of endemics. Among these processes is the discovery of species new to science. Certainly more species await discovery, especially in the Yungas. Probably the percentage of endemic species within a given unit is highest for the Altiplano, fol- lowed by the Yungas. For a detailed discus- sion of the concept of endemism see Ander- son (1994). I speculate that the net change in percentage of Bolivian species that are judged to be endemic to Bolivia will ap- proach in the future a level of equilibrium slightly higher than that at present. Before continuing the discussion of pat- terns in mammalian distributions, the nature of the basic data needs to be reviewed. Lo- calities represented by specimens were plot- ted on a map for each species. Then I looked at the map, thought about what I knew about the animal (principally within Bolivia but also in surrounding regions) and about con- ditions within Bolivia, and drew a line or lines showing my estimate of the limit of the distribution within Bolivia for each species. In spite of the inherent subjectivity involved, I think that these estimates are about the best that can be made, until we learn more. The confidence and the precision with which that line can be drawn depends on how much is known about (1) the actual oc- currence of the species (i.e. the plotted lo- calities), (2) the habitat requirements of each species, and (3) the habitats specifically available in each local area. Habitat require- ments and local habitats are known, but only in a very general way. The fact that most species are known from few localities, as dis- cussed elsewhere, means that even the "hard data" represented by dots on the maps are quite limited. The reader who wishes to dis- count my subjective boundaries has the dots to contemplate. In addition to the inherent subjectivity in estimating boundaries, there is potential circularity in reasoning about pat- terns. If a species is known only from local- ities in the Gran Chaco, and I know roughly where the limits of the chacoan habitat are, I may draw the estimated boundary for that species near the habitat limits. To contend that there is a suite of chacoan species that have common limits then tends to be redun- dant or the reasoning a bit circular. Alternatively, a more objective procedure might be considered. Suppose that for each species map a convex polygon is drawn around the existing dots and a procedure for extralimital data is incorporated. Then, an- other polygon is drawn outside the first, with the distance between the two polygons equal to the average distance between the dots. The outer polygon becomes the "objective" es- timate of the boundary for each species. I postulate that the general patterns of distri- butions that are evident in the more subjec- tive analyses would remain. To obtain a visual summary of distribu- tional limits within Bolivia for each of three groups of mammals (bats, rodents, and oth- ers), all limits estimated for the individual species were superimposed on one map for each group (figs. 474 to 476). Areas of rel- atively rapid faunal change are indicated by 76 NO. 231 ANDERSON: MAMMALS OF BOLIVIA 60°W lo' S 10 S 7~ + s + X20 S% SCALE 0 200 km. Fig. 474. Map of Bolivia with the boundaries of the geographic ranges of all species of bats superimposed. Any clustering of boundaries is ev- idence of relatively rapid faunal change. the clustering of boundary lines. Most boundaries for all three groups cluster in a broad band around the eastern slopes of the Andes, an area known as the Yungas and Valles. As noted elsewhere, this area is not simply a location where lowland and high- land species reach their respective south- western and northeastern limits. Inspection of the maps for individual species reveals that many species are confined to the Yungas and Valles area. Conversely, for example, the maps reveal few species of bats and few boundaries on the Altiplano. This approach to analysis of faunal areas uses animal distributions to reveal patterns. In a way, this is the reciprocal of another common approach (as used by Salazar et al., in press; and many other authors) in which areas are first defined and delineated by veg- etation, topography, and precipitation, or on the basis of other environmental parameters. Then the distribution of mammalian fauna (or whatever other part of the total fauna is of interest) is compared to these previously defined areas to determine which are more and which are less similar. An additional approach to the study of dis- tributional areas (areography) was used by Armstrong et al. (1986) in an analysis of the Fig. 475. Map of Bolivia with the boundaries of the geographic ranges of all species of rodents superimposed. mammalian fauna of the Plains states in North America. Four suites of species with more or less common patterns of distribution were plotted, each suite on a separate map. For Bolivia, I adopted this approach by cre- ating four separate maps. In figure 477, I plotted all of the species that occur in the 68°W -- 60°W 10 S Fig. 476. Map of Bolivia with the boundaries of the geographic ranges of all species of mam- mals other than bats and rodents superimposed. 77 1997 + 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 10° S 68°W 60°W + + M\ X, 10°S Fig. 477. Map of Bolivia with the boundaries of the Chaco suite of species superimposed. Gran Chaco southeast of Bolivia and that reach their northern limits within Bolivia. All of the species that occur on and that reach their limits on the Altiplano or near its east- ern edge are shown in figure 478. Species that occur in the Amazonian region north of Bolivia and that reach southern or south- western limits in Bolivia are shown in figure 479. Finally, species which, within Bolivia, 68° W ~ 60°W + ++ lO° S W + 4J+ t+ X ++ + + ++ + + t +S~~+ + + + 4 + t~~~~ + *+X + SCALEF 0 200 km. Fig. 478. Map of Bolivia with the boundaries of the Altiplano suite of species superimposed. Fig. 479. Map of Bolivia with the boundaries of the Amazonian suite of species superimposed. are more or less confined to the Yungas and Valles area are shown in figure 480. No suite of species was discernible for either of the other two types of areas delineated in figure 471 and used by Salazar et al. (in press), namely the Savannah and Chiquito Forest. The species occurring in these areas almost invariably occur also in other areas. The lines for approximately 50 of the 136 species in 68°W l60°W + * + +9 10°S ~~~~~~~+ <+ 20' S SCALE 0 200 km. Fig. 480. Map of Bolivia with the boundaries of the Yungas suite of species superimposed. 78 NO. 231 ANDERSON: MAMMALS OF BOLIVIA the Amazonian suite cross the border into Argentina or Paraguay to the south and thus indicate that the ranges of these species ex- tend south of Bolivia. About 20 Amazonian species reach their southern limit near the edge of the Gran Chaco. The general pattern for the Amazonian suite is a gradual drop- ping out of species as one leaves the tropical Amazonian high forest in the north, passes through progressively cooler and drier Sa- vannah and Chiquito Forest, and finally en- ters the thorn scrub of the Chaco. The numbers of each mammalian order that either do or do not fit one of these four selected patterns are given in table 1. Most species (226 of 296 species analyzed, or 76%) can be reasonably assigned to one of the patterns. A few poorly known species were excluded from this analysis because it was not reasonable to say whether they con- formed to any of the patterns. Bats and pri- mates are principally Amazonian. Rodents have a much higher proportion of species in the Chaco, the Altiplano, and the Yungas and Valles than in other areas. Almost half of the species of Carnivora do not easily fit any one of the four patterns. MANAGEMENT AND CONSERVATION Some interactions between native mam- mals and people have been noted briefly in various accounts of species. Among interac- tions that may be regarded negatively from the human point of view are the transmit- tance of organisms that cause disease in hu- mans or in domestic mammals; damage to crops; damage to buildings, ditches, or other human facilities; and the killing of domestic animals or even humans on rare occasions. In relation to disease, the role of Calomys callosus as a host of the Machupo virus has been mentioned. Certainly other mammals act as hosts to other organisms or viruses that can produce disease in humans, such as yel- low fever, leishmaniasis, and plague, but lit- tle specific information has been obtained. The vampire bat, Desmodus rotundus, has been known to transmit to cattle the virus that causes rabies, and rabies can occur in other species of mammals, including hu- mans. Various small rodents, especially the intro- duced house mouse and rat, Mus musculus and Rattus rattus, are known to damage crops both in the field and in storage after harvest. Reports of predators killing domestic ani- mals are only anecdotal. Certainly large predators such as the jaguar (Panthera onca), the puma (Felis concolor), and the spectacled bear or jucumari (Tremarctos ornatus) do sometimes kill domestic animals. The small- er cats and the foxes may take chickens or other small domestic animals. Although we know such predation occurs, we have no quantitative data to use in its evaluation. Other interactions are regarded as benefi- cial. Among these are the use of wild mam- mals as food, use of skins for their fur or for leather, and the recreational or tourist values of sport hunting, nature photography, or wildlife watching. The larger rodents, including the capybara (Hydrochaeris hydrochaeris), the hochi pin- tado (Agouti paca), and the hochi colorado (Dasyprocta punctata), are commonly hunt- ed in rural lowland areas. The larger pri- mates, Alouatta, Ateles, and Cebus, are like- wise hunted for food and, where hunted in- tensively, populations have been reduced or eliminated. The peccaries (Tayassu pecari and T. tajacu) are widely hunted for both food and their skins. Two species of small deer of the genus Mazama (M. americana and M. gouazoupira) are relatively abundant and are favorite sources of food in the low- lands. The tendency, illustrated above, to classify animals or their actions as either good or bad on some particular human scale is generally naive from a broader biological or philo- sophical perspective. Firstly, a given action may be good, bad, or neutral, depending on circumstances and the perspective from which it is viewed. Secondly, we do not know enough about the current status of most species or their complex ecological re- lationships to evaluate their actions effective- ly. Thirdly, human societies, not only in "de- veloping" countries but everywhere, have not yet learned how to balance short-term and long-term values, how to equitably and productively allocate costs and benefits, or how to achieve and maintain a sustainable life-style, civilization, or ecosystem. In fact, 1997 79 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY it is not generally agreed that it is possible to do these things or even desirable to try. The human population in Bolivia has about doubled since I first worked there in 1963, and current trends, if continued, will double the population again in the next 30 years. Clearly this will affect wildlife, al- though in what ways exactly is not so clear. No one is closely monitoring changes in pop- ulations of most of the native species of mammals. Only for the vicunia, Lama vicug- na, are some general data available. This spe- cies has been protected in special areas, has survived, and in fact has increased from the low level reached several decades ago. The chinchilla, Chinchilla chinchilla, probably was exterminated in Bolivia by 1963. The guanacoe, Lama guanicoe, is known to have survived only in one local area at the edge of the Gran Chaco and may soon become extinct there. The white-tailed deer, Odocoi- leus virginianus, was recorded in 1974 in the vicinity of Pelechuco, but no current infor- mation on its status is known to me. The pampa deer (Odocoileus bezoarticus) and the other large deer (the marsh deer, 0. dicho- tomus) have probably been reduced in range and in numbers by hunting, but no specific or current information is available. The tar- uca (Hippocamelus antisensis) of the high Andes may be subject to overhunting; again, no data are available. The Chacoan peccary (Catagonus wagneri) is at risk; its population is probably declining throughout its range and overhunting could eliminate it from the Bolivian fauna. Conservationists' attention has turned to the importance of general biodiversity, rather than focusing only on the risks that a few charismatic megavertebrates face (Scott et al., 1987). Increasing knowledge is raising Bolivia's rank among countries exhibiting megadiversity (Yensen et al., 1994); howev- er, rank is a tentative thing considering our ignorance about the faunas of many nations. The future for native mammals in Bolivia will be decided by Bolivians. Mammals are just one small part of the larger natural sys- tem. For a general review of the state of the environment and related problems see Free- man et al. (1980) and E. Geyger and C. Arze (eds., 1982). More Bolivians are aware of conservation problems now than a few years ago, and I wish them well. They can use whatever international help they are offered. I hope that they will do better than what was done historically in North America. Here the bison was nearly exterminated and the pas- senger pigeon and Carolina parakeet were exterminated before effective conservation efforts were begun (see Shaw and Schmidly, 1994). I have merely outlined the broadest view of the subject of conservation and have not suggested specifics, which would be be- yond the scope of the present work. The present summary of the mammalian fauna and the distribution of its species provides a basic, but general, picture of an incredibly diverse fauna and may help in future plan- ning for its conservation. ABBREVIATIONS AND ACRONYMS Some of the standard abbreviations for collections (Yates et al., 1987) are shortened further here to save space and costs because they are cited hundreds or thousands of times in lists of specimens, e.g., AMNH for the American Museum of Natural History is shortened to AM in lists. ACB Atlas Censal de Bolivia, 1982 AMNH or AM American Museum of Natural History, New York Academy of Natural Sciences, Philadelphia British Museum (Natural His- tory), London BZM CAS CBF CDC CENETROP CITES CMNH or CM Berlin, Zoologisches Muse- um, Humbolt Universitat California Academy of Sci- ences, San Francisco Colecci6n Boliviana de Fau- na, La Paz, formerly in IE and MNLP Centro de Datos para la Con- servacion, La Paz Centro Nacional por Enfer- medades Tropicales, Santa Cruz Convention on International Trade in Endangered Species Carnegie Museum of Natural ANSP BM 80 NO. 231 ANDERSON: MAMMALS OF BOLIVIA CNHM EBB EBD ENDE ENTEL FMNH or FM GEOBOL GPS IBBA ICZN IE IGM IML IUCN LACM LSUMZ or LSU MACN MAPA MARU MCN MCZ MHA MLP MNK History, Pittsburgh, Pennsyl- vania Chicago Natural History Mu- seum Estacion Biologica del Beni Estacion Biologica Dofnana, Sevilla, Spain Empresa Nacional de Energia Empresa Nacional de Telef6n- ico Field Museum of Natural His- tory, Chicago Servicio Geologico de Bolivia Global Positioning System (satellite electronics) Instituto Boliviano de Biolo- gia Altura, La Paz International Code of Zoolog- ical Nomenclature Instituto de Ecologia, La Paz Instituto Geografico Militar, Mapa de la Reputblica de Bo- livia (1:3,000,000); IGM5, 1: 500,000 maps Instituto Miguel Lillo, Tucu- man International Union for Con- servation of Nature Los Angeles County Museum Louisiana State University Museum of Zoology, Baton Rouge Museo Argentino de Ciencias Naturales, Buenos Aires Mapa de la Reputblica de Bo- livia (1:1,500,000), 1947, by R. R. Camacho Lara, Zumel- za y Cia., La Paz; or, Mapa de la Reputblica de Bolivia (1: 1,000,000), 1980, third edi- tion, Instituto Geografico Mil- itar, La Paz. Middle America Research Unit (now closed), Balboa Heights, Canal Zone, Panama Museo Nacional de Ciencias Naturales, Madrid Museum of Comparative Zo- ology, Harvard University, Cambridge, Massachusetts Map of Hispanic America (1: 1,000,000), American Geo- graphical Society, New York (on one of the nine sheets covering Bolivia) Mapa del Departamento de La Paz, 1:500,000, 1981 Museo de Historia Natural MNLP MSB MSU MVZ N.L. ONC PASB PRBN RAP RON SNCFHB TIPNIS UAF UAGRM UCD UCONN UMMZ UMSA UNM USBGN USNM or US "Noel Kempff Mercado," Santa Cruz Museo Nacional de Historia Natural, La Paz Museum of Southwestern Bi- ology, Albuquerque Michigan State University, East Lansing Museum of Vertebrate Zool- ogy, Berkeley Not located Operational Navigational Chart (1: 1,000,000), Aero- nautical Chart and Informa- tion Center, U.S. Air Force, St. Louis, Mo. (reference to either sheet ONC N-26, re- vised 1967, or to sheet ONC P-26, 1965). Referred to as WAC (World Aeronautical Chart) in field journals of AMNH 1963, etc. Pan American Sanitary Bu- reau Proyecto Rfos Blanco y Ne- gro Rapid Assessment Program, Conservation International, Washington, D.C. Territorio de Rondonia (1: 1,000,000), Instituto Brasilei- ro de Geografia e Estatftica, Divisao de Cartografia, 1961 Servicio Nacional de Control de Fiebre Hemorragia Bolivi- ana Territorio Indigena Nacional Isiboro-Secure University of Alaska, Fair- banks Universidad Autonoma "Ga- briel Rene Moreno," Santa Cruz University of California at Davis University of Connecticut University of Michigan Mu- seum of Zoology, Ann Arbor Universidad Mayor de San Andres, La Paz University of New Mexico, Albuquerque U.S. Board of Geographic Names, Dept. of the Interior, Washington, D.C. (Gazetteer No. 4, Bolivia, 1955) United States National Muse- um, now the National Muse- 81 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY um of Natural History, Wash- ington, D.C. World Aeronautical Chart, see Operational Navigational Chart Zoologisches Forschungsin- stitut und Museum Alexander Koenig, Bonn 1618 /6753 etc., this example is for 160181 south latitude and 67053' west longitude. The coordinates of rivers are given for their mouths, unless oth- erwise noted. MAMMALOGICAL GAZETTEER OF BOLIVIA Paynter et al. (1975) published an ornitho- logical gazetteer of Bolivia after which the following gazetteer is patterned. Because many collectors obtained both birds and mammals, there is some overlap. References to all type localities for Recent Mammalia in Bolivia are included here. The intent here is to document every lo- cality from which a specimen of Recent mammal has been preserved in a museum collection. A few localities referred to in the literature but not represented by specimens are also included. The original version of January 1980 was compiled with the help of Ted Danforth, Jr., and was based on the lit- erature and on specimens in the American Museum of Natural History. Other localities represented in this and other collections and localities visited since 1979 have been added as I have had opportunity to obtain this in- formation. By using this alphabetical list in conjunc- tion with the "List of Collectors" and the chronological "History of Collecting," lo- cations of origin of specimens can usually be resolved. Since more than one place may have the same name, it is necessary to know collector and date to be certain. The adequa- cy of maps available to collectors through the years and the care used by collectors in as- certaining and designating localities have varied; therefore, some imprecision and some inaccuracy persist. For each locality, the name is followed by the name of the department (or country, if outside Bolivia). The latitude and longitude are then given (abbreviated as noted in list of abbreviations), along with the map or oth- er source of those coordinates. Then, listed in chronological order are names of collec- tors, dates, and names of authors (with dates of publications in parentheses), notes on type localities, and variant spellings that have been used. Variant spellings may be found on different maps published within Bolivia and in the absence of an official arbiter I do not indicate which is correct. Occasionally there is a clear misspelling, but even if there is one correct spelling, it is useful to know the variants that appear on specimen labels or in the published literature. For rivers, the coordinates are for the mouths, unless oth- erwise indicated. Collections often have been made at one or more places designated as being some distance from the named refer- ence locality rather than at that locality. Co- ordinates are given for each actual source of specimens. Coordinates not followed by a map source in parentheses were estimated in some way by me and are sometimes less ac- curate than those from maps. For example, a site designated as "20 km SE by road" might be estimated to be 15 km SE in a straight line and coordinates calculated accordingly. Different maps may also show somewhat dif- ferent coordinates for a named place. Cross-references are provided when some specimens have one designation and others from the same place have a different desig- nation on specimen labels or in published re- ports. In some cases, the province as well as the larger department is given. This is done be- cause some original labels show both prov- ince name and local place name and these names have been confused in museum re- cords or in publications. Consulting the gaz- etteer should help reduce this confusion. Oth- erwise, provincial names have not been in- serted. Recently, Bolivian collectors have in- cluded province names in designating localities more often than collectors did in earlier years. WAC ZFMK NO. 231 82 ANDERSON: MAMMALS OF BOLIVIA The sequence of localities is alphabetical in conventional English form, thus LI, N, and Ch, used in Spanish as letters different from L, N, and C and placed in Spanish diction- aries after all entries for those letters, are not separated here. Locality designations are sometimes complex, e.g., a locality is des- ignated in two different ways, and one is cit- ed one time and the other at other times. Some designations begin with an article such as el or las or with a general term such as no, serrania, or laguna. For the convenience of readers these are cited under the specific term rather than under the general term and many of these are cross-referenced. LIST OF LOCALITIES ABAIXO DA LANTERNA, Brazil N.L. On rio Guapore, probably on the Brazilian side, Natterer (Pelzeln, 1883: 94). ABAPO; Santa Cruz 1850/6328 (USBGN) Pine (specimen from Gene Harris of Santa Cruz), Aug. 1980, from 3 km N of (and 7 km S of rio Grande; 1848/6328). ABRA, LA; Santa Cruz 1722/6302 Hibbs, Aug. 1965, 18 km NE of Warnes; and 2 km N of La Abra (1721/6302). ABUNA; Pando 0942/6523 Felis pardalis (L. Salles, personal commun.); RAP team, June and July 1992, Puesto Abufia (0947/6532), 140 m, Bolivian naval post on S bank of rio Abufia in Provincia General Federico Roman, opposite Fortaleza, Brasil; same RAP team, "Cachuela Camp" on N bank of rio Abufia at first cachuela about 3 km upstream from Puesto Abufia (0945/6533); on some maps as Abuna or Abufia. ACACIO; Potosi 1800/6605 Kessler, Aug. 1991, 31 km from, on road to Uncia (1806/6608). ACAPULCO, see San Joaquin. ACERAMARCA, RIO, see Aceromarca. ACEROMARCA, RfO; La Paz 1618/6753 (USBGN) Tate, May 1926, 3275 m, tributary to rio Un- duavi, Beni drainage. "The stream in the great glaciated valley that is visible directly across the Unduavi valley from the point of rails at Ichulo- ma. Camp half an hour up valley where the pro- jected railbed crosses the stream." Creighton, Nov. 1979, 2600 m, 1 km S of Yerbani. Anthony (1926, pl. 1), type locality of Thomasomys ladewi; Tate (1931b: 12), type locality of Marmosa ac- eramarcae (= Gracilinanus aceramarcae); AMNH-MSB party, Aug. 1992, 2990 m, (1619/6753 GPS); see also Yerbani. "Cabezilla de" Hippocamelus antisensis, obtained by Jorge Condon, Apr. 1992 (1618/6753). AMNH-MSB party, Aug. 1992, 2990 m (1619/6753); CBF party, Sept. 1992, 2080 (1619/6753). ACHACACHI; La Paz 1603/6843 (USBGN) Simons, Oct. 1900, Feb. 1901, 3827 m, SE of Titicaca; Thomas (191 1b: 256), as Achacoche; Gilmore, Nov. 1942, 5 km W of (1603/6846), 15 km N of (1555/6843), and Hacienda Poccata (or Pocoata) to the north (ca. 1600/6845), in province of Omasuyos; Baudoin, May 1968, 10 km E of (1603/6838); also spelled Achacach. ACHACHAIRU; Beni 1324/6404 (MARU) Webb, 20 Apr. 1970, on rio Itonamas. ACHACHICOLA; La Paz 1621/6803 (ACB) Pacheco, 1980s, 2 km NE of Alto Achachicola, sometimes spelled Achachicala. ACHIRI; La Paz 1712/6900 Alvarado, June 1989, in province of Pacajes. ACHOCALLA; La Paz 1634/68 10 (ACB) Mercado, 1987. ACRE, PUERTO; Beni 1219/6428 (RON) AMNH, July 1964. AGUADULCE, see San Joaquin. AGUA DULCE; Pando 1101/6612 AMNH-MSB party, July 1986; 160 m. AGUA HEDIONDA; Santa Cruz 1810/6344 de la Barrera, Nov. 1954, 1955 (Fonseca, 1959; Hopkins and Rothschild, 1966: 120); also see Florida and Floripondio. AGUAIRENDA MISSION; Tarija 2151/6340 (USBGN) ca. 850 m, (MHA); Thomas (1898b). AGUA RICA; Oruro 1820/6836 An estancia, 40 km E and 22 km S of the vil- lage of Sajama; AMNH-MSB party, Sept. 1986, 3850 m. AGUIRRE N.L. F B. Steinbach, Mar. 1940. AIQUILE; Cochabamba 1807/6509 (USBGN) F. B. Steinbach, July 1936; de la Barrera, 2225 m, Sept. 1955 (Fonseca, 1959: 130); also spelled Aiguile or Aquile. ALALAY, LAGUNA; Cochabamba 1725/6609 EBD party, Sept. 1982, or Alalai, at S side of city of Cochabamba. ALASKA MINE; La Paz 1617/6802 (Tate field map) Tate, Mar. 1926; "A ... tin mine lying at the head of a glaciated valley immediately to the north of and above Pongo" (1620/6757, USBGN), ca. 4300 m. 1997 83 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY ALCOCHE; La Paz 1542/6740 (USBGN) MARU, 1963; Creighton, May 1978, 4 km (by road) NW of (1540/6742) 1400 ft (= 425 m); cit- ed as Aleoche by Pine (1972: 22). ALEJANDRIA; Beni 1205/6509 (IGM) AMNH, May 1965, on rio Mamor6, 5 km NW of, on left bank (1203/6510). ALISUNI; Cochabamba 1646/6637 (MAPA as Alisani) Simons, July 1901; Thomas (1902a), 2600 m, "on the high paramos north and northwest of Co- chabamba." ALMACEN, PUERTO; Beni 1447/6451 (MAPA) Pilleri (1969b), on rio Ibare west of Trinidad; Anderson, Aug. 1983, 230 m. ALMACEN, PUERTO; Santa Cruz 1530/6225 EBD party, 1986; Pajuelo, Jan. 1988; Tapia, Dec. 1989, on nro Negro in province of Nuflo de Chavez. ALOTA; Potosi Salazar, May 1989, 3 km NE of (2118/6720). ALOTA, VILLA; Potosi 2124/6741 EBD party, Nov. 1982; 2 km N of (2123/6741); 2 km W of (2124/6741); 9 km W of (2124/6747). ALTAMACHI; Cochabamba 1702/6626 (USBGN) Gilmore, Mar. 1943. ALTIPLANO; this, like other terms such as cha- co, yungas, and valles, can be used either in a general sense or as a proper noun or part of the name of a specific place or area. When used in the specific sense to refer to the Bolivian Altipla- no or the larger Andean Altiplano, Altiplano is capitalized. If used to refer to high plains in gen- eral, lower case altiplano is used. ALTO MADIDI, see Moire. ALTO PALMAR, see Palmar. ALTWA SAN JUAN; Beni 1308/6444 (MARU) MARU 1963, 4 km S of San Juan. AMBOR6, see also San Rafael de Amboro. AMBOR6, CERRO; Santa Cruz 1744/6339 (USBGN) J. Steinbach, Mar. 1916 (Crespo, 1974: 16); F. Steinbach, Sept. 1947; Riddle, Sept. 1985, rio Pi- tasama, 4.5 km N and 1.5 km E of Cerro Amboro, 620 m (1745/6340). AMULLAJTA; La Paz 1702/6748 (USBGN) Baudoin, 1970, 5 km down valley from Cara- cato, also spelled Amullachta. ANDES OF BOLIVIA, see La Paz. ANDRJES IBANEZ; Santa Cruz 1745/6315 (USBGN) F B. and J. Steinbach, Apr.-July 1937, 430 m; a province later known as Cercado. APERE, RIO; Beni 1344/6518 (USBGN) AMNH, 1965, 4 or 5 km above mouth (1345/6519); Tello, July 1986, rio Apere (1400/6515), and Aug. 1986, rio Apere flood plain about 40 km W of San Ignacio de Moxos (1448- 6555). APOLO; La Paz 1443/6830 (MAPA) 1440 m, on rio Beni headwater; now province of Franz Tamayo; Weddell, 1853; Deville (1849: 55) and Weddell (1853), type locality of Midas Weddellii (= Saguinus fuscicollis weddellii) as Province of Apolobamba; Abel Fornes, 1969; Emmons, June 1990, 50 km W of, at 1500 m, 5 km beyond Correo on new road to La Paz (1455/6820); Emmons, June 1990, 17 km N of, at 1000 m, on trail to Tuichi, on rio Machariapo, a tributary of rio Tuichi (1434/6828). APOLOBAMBA, see Apolo. ARANI; Cochabamba 1734/6546 (USBGN) 9000 ft (= 2770 m), d'Orbigny, Oct. 1830, d'Orbigny (1834-1847); F B. Steinbach, Jan. and Feb. 1927, Sept. 1928, Feb. 1939; Baptista and Salazar, June 1987, 17 km by road W of (1731/6555); Kessler, Aug. 1991, 3350 m, 25 km S of, on road to Mizque area (1738/6539). ARAONAS; La Paz Harada, 1984; Iseki et al. (1985). 1230/6830 ARIRUMA; Santa Cruz AMNH-MSB party, June 1991, 7 km (by road) SE of (1824/6419), 1750 m. AROMA; La Paz No other data. 1717/6734 ARQUE; Cochabamba 1748/6623 Collins-Day Expedition (Osgood, 1916: 199); perhaps this locality. ARROYO CANADA; Pando AMNH, July 1986, mouth of. 1123/6658 ARROYO MERCEDES, see Buena Hora. ARRUDA; Beni MARU, 1963-1964. N.L. ASCENSI6N DE GUARAYOS; Santa Cruz 1542/6305 (USBGN) MARU, June 1964, in province of Nuflo de Chaivez; AMNH-MSB Aug. 1985, 6 km by road W of Ascension (1543/6309). ASTILLERO; La Paz ca. 1616/6733 Simons, Nov. 1900, Feb. 1901, Thomas (1902a: 129), 2700 m; Thomas (1914: 362), type locality of Sciurus cuscinus ochrescens (= S. ignitus ig- nitus); Hershkovitz (1962) as Astilleras. ASUNTA; Beni 1307/6523 (MARU) MacKenzi, Nov. 1971. But also see San Joa- quin. 84 NO. 231 ANDERSON: MAMMALS OF BOLIVIA ATACHEBA; Cochabamba N.L. J. Steinbach, Aug. 1920 (FMNH) ATOCHA; Potosi AMNH-MSB party, Sept. 1986, 40 km by road SE of, 4000 m (2107/6602). ATOLLADAR; Chuquisaca N.L. Arduz, Aug. 1951, in valley of rio Bafiado. AVAP6, see Abap6. AXOJO; Beni N.L. Boca Axojo (Inia). AYACUCHO; Santa Cruz 1700/6355 Estaci6n on railroad, AMNH and CENETROP, Oct. 1987, 250 m. AYACUCHO; Santa Cruz 1751/6320 (USBGN) In province of Ibafiez; Acasigue, Becerra, Kerr, Ridell, Villalobos (MARU), Aug. and Oct. 1966; also 1 km E of Ayacucho. AYANE, see Yani. AYATA; La Paz Yoneda, June 1982. 1521/6845 (USBGN) AYDAYO 17??/6822 Felis concolor at USNM, 20 km W of Aydayo; minutes of latitude unknown. AYOPAYA; Cochabamba 1630/6635 (USGBN) Province in western Cochabamba; d'Orbigny, Sept. 1830; as Ayupayo, F B. Steinbach, June, July, and Aug. 1953, some recorded as Ayopaca, 3500 m. AZERO, RIO; Chuquisaca 1912/6357 (USBGN) Carriker, Nov., 1936, 1220 m; Azuero (USBGN) AZUNTA, see San Joaquin. BAHIA CHAPLIN, LAGUNA; Santa Cruz 1428/6102 EBD party, Aug. 1986, in Velasco province. BAHIA DAS ONCAS; Beni 1205/6445 (MAPA) On rio Itenez ca. 50 km NW of Principe da Beira. AMNH, July 1964; between Bahia das On- cas and Casa Alta. BAHIA DE LAS PIEDRAS N.L. Hydrochaeris specimen. BAHIA DE LOS CASARA; Beni 1313/6220 AMNH, Aug. 1964; see Larangeiras. BAJO PALMAR, see Palmar. BALLIVIAN; PROVINCE OF; Beni BALNEARIO LAS POZAS; Cochabamba 1657/6523 EBD party, Sept. 1982, see Villa Tunari. BALZON, LAKE; E. Bolivia N.L. Tate (1933: 198). BANANEIRA, CACHUELA: Beni 1036/6525 (USBGN) Natterer, 3 Sept. 1829 (Pelzeln, 1883: 127) as Cachoeira do Bananeira. BANARES, RIO; Beni (?) N.L. Natterer, Aug. 1829 (Pelzeln, 1883: 127); pos- sibly rio Baures? BARADOR, PUERTO, see Puerto Varador. BARRANQUITA; Beni 1313/6448 MARU, 1963, or 10 km S of (1318/6448); also Estacion Barranquita; probably this is Bauricito (MAPA, USBGN, 1313/6448). BARRERAS, LAS; Santa Cruz 1726/6305 (MARU) Hibbs, July 1965; 11 km NE of Warnes. BASILIO; Santa Cruz 1808/6312 (USBGN) Becerra, Kerr, Villalobos, Acasigue, Oct. 1966, 45 km SSW of Santa Cruz, in province of Cor- dillera; at Basilio and 3.4 km S of Basilio (1810/6312); MNK party, Aug. 1992, 2 km W of (1807/6313). BATO; Beni, see rio Negro. BAURES; Beni 1335/6335 (USBGN) MARU, 1963; Ranck, Nov. 1966. BAURES, RIO; Beni 1230/6418 (IGM) AMNH party, October 1964. BELEM; Beni 1230/6340 AMNH, July 1964, island in rio Itenez between Santa Rosa and Isla de Bispa. BELGICA, LA; Santa Cruz 1733/6313 MSB party, June 1988, 4 km SW of (1735/6315); MNK party, Feb. and June 1992, 2 km S of (1731/6312). BELLA ESPERANZA; Santa Cruz N.L. Silva, June 1948, in province of Cercado, now Iba'nez. BELLA VISTA; Beni 1340/6550 Estancia between Santa Ana de Yacuma and Santa Rosa, photo of Chrysocyon skin by Jaime Peralta T, 23 Oct. 1992; see also Boa Vista. BELLAVISTA, SERRANIA; La Paz 1540/6735 Schmitt and Remsen, June 1979, a tea planta- tion, 35 km by road N of Caranavi, 1650 m (1540/6735); Webster, 1979 (Webster and Jones, 1980); 47 km by road N of Caranavi, 1350 m (1538/6732); AMNH-MSB party, as Bella Vista, July 1992, at 1300 m (1542/6729 GPS), and at 1525 m (1541/6730 GPS). BELLA VISTA; La Paz 1520/6813 (USBGN) Simons, Aug. 1900, 1400 m; Andersen (1906: 419), as Bellavista, type locality of Uroderma Thomasi (= Uroderma bilobatum thomasi). BELLA VISTA; Pando 1123/6712 AMNH-MSB party, July 1986, 170 m. 1997 85 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY BENI, RIO; 1023/6524 (USBGN) Gray (1866b: 826), type locality of Cebus pal- lidus (= C. apella pallidus), by restriction (Ca- brera, 1958: 166); Dr. Heath, prior to 1901, exact location unknown; Mann, June and Sept. 1922; Scherrer, Apr.-May 1925, exact location un- known; AMNH, MSB, and IBBA party, Sept. 1985, (1304/6711, 1309/6712, 1315/6718, 1316/6717, 1401/6731, 1416/6732, all in La Paz, and 1315/6717 and 1327/6721, in Beni); Dicker- man and Ruedas, July 1986, on W bank opposite Hamburgo, 160 m, in Pando (1101/6606); AMNH-MSB party, July 1986, left bank, 160 m (1057/6603); see also Consuelo. BENI RESERVE; Beni Salazar, Sept. 1987, at rio Curiraba, "BIOLAT ZONA 2" (1450/6623), see Estacion Biologica de Beni and Porvenir. BERMEJO; Santa Cruz 1810/6336 (MAPA) Miller and Boyle, Oct. 1915, 3500 ft (= 1100 m). BERMEJO, RIO; Tarija 2651/5827 (USBGN) Carriker, Sept. 1936, 410 m, at Bermejo (2210/6442); Carrizo, Daneri, and Vinas, July 1989 (Vaccaro, 1990), left bank opposite mouth of rio Lipeo (2227/6430). BETHANIA; Beni N.L. MARU, 1963, Estancia Bethania, 35 km E of. BIATA, RIO; Beni 1144/6648 AMNH-MSB party, Aug. 1986, 170 m, at San Jos6 about 1 km above mouth of rio, designated as boca del rio Biata (1144/6647). BIBOSI; Pando ca. 1150/6840 An estancia; CBF, year unknown. BLANCO, RIO; Beni N.L. MNLP, Sept. 1983. BLANCO, RIO; Cochabamba 1636/6637 Simons, June 1901; see Charuplaya, coordi- nates erroneously given as 1550/6748 by Smit (1987: 215). BOA VISTA; BENI 1306/6348 (MAPA) Also Bella Vista or Buena Vista; Webb, Apr. 1970; Remsik (specimen from Harris), Mar. 1980, about 1000 ft elevation, see San Joaquin also; which location uncertain. BOBICA N.L. Wesmael (1841: 59), type locality of Maston- otus popelairi (= Myocastor coypus popelairi). BOLIVAR; Tarija 2138/6234 An estancia; AMNH-MSB party, Aug. 1991, 400 m; 4 km N of (2136/6234), 400 m; 5 km W of (2138/6237), 400 m. BOLIVIA, LAGUNA; Beni N.L. CBF party, June 1992, TIPNIS; 1 km N of; at mouth of Arroyo Negro; W bank of and N of. BOLIVIA Gray (1846: 212), type locality of Jacchus leu- comerus (= Callithrix argentata melanura); Gray (1850b: 380), by inference from the name, type locality of Coendou boliviensis (= Coendou pre- hensilis boliviensis); Gray (1873b: 16), by infer- ence from the name, type locality of Tatusia bo- liviensis (= Dasypus novemcinctus novemcinc- tus); Brass (1911: 613), by inference from the name, type locality of Chinchilla boliviana (= C. lanigera). BOLIVIAN CHACO Thomas (1898b: 2). BOLIVIAN CORDILLERA d'Orbigny. BOLPEBRA; Beni 1330/6426 Estancia; Yates and Salazar, Mar. 1993. BOROICA; Beni 1313/6409 (USBGN) P. A. Webb, Apr. 1970, in Itonama Province. BOYUIBE; Santa Cruz 2025/6317 (USBGN) de la Barrera, 1955, spelled Boyuiba; Bromley, Aug. 1957, spelled Boyuibi; AMNH-MSB party, July 1991, 53 km E of (2027/6250), 600 m; 26 km E of (2026/6302), 800 m; 1 km S and 4.5 km E of (2027/6314), 815 m. BRESTA; Beni ca. 1433/6720 (MHA) Olalla, 1938; Lonnberg (1942: 55). BUENA HORA; Brazil 1133/6512 (RON) Used to designate places in Beni; AMNH, May 1965, Arroyo Mercedes, 6 km S of Buena Hora (1136/6512); 5 km SW of (1134/6513); See RON, there are two places named Boa Hora in Rondon- ia, fieldnotes indicate the northern one. BUENA VISTA; Beni 1306/6348 (MAPA) MARU, 1963, on rio Blanco, 25 mi E of Mag- dalena, also termed Bella Vista or Boa Vista. BUENA VISTA; Santa Cruz 1727/6340 (USBGN) J. Steinbach, at various times from 1914 through 1938, 500 m; F. Steinbach, at various times from 1926 through 1952; Miller and Boyle, Aug. 1915, 1300 ft (= 400 m); Thomas (1928b: 293), type locality of Cyclopes didactylus catel- lus; Nelson and Goldman (1929), type locality of Felis concolor osgoodi; Tate (1931b), type local- ity of Marmosa agilis buenavistae (p. 10, = Gra- cilinanus agilis buenavistae); Marmosa ocellata (p. 7, = Marmosops dorothea); Pocock (1941b), type localities of Leopardus pardalis steinbachi (p. 235, = Felis pardalis steinbachi) and of Leo- pardus wiedii boliviae (p. 237, = Felis wiedii bo- liviae); Cabrera (1958), restricted type locality of Arctopithecus gularis Gray (1850c) (= Bradypus variegatus) to here; Nelson and Goldman (1933: 229), type locality of Felis onca boliviensis (= 86 NO. 231 ANDERSON: MAMMALS OF BOLIVIA Panthera onca); G. L. Ranck, Nov. 1966; Schmitt and Cole, Dec. 1978 and Jan. 1979, 7 km N and 17 W of Buena Vista (1724/6344), at km 119 on road to Yapacani, Propiedad Nuevo Mundo, 353 m; Creighton, Feb. 1980, 25 km (by road) W of Buena Vista, on W bank of rio Yapacani (1724/6346), 400 m; Harada, Aug. 1984; MSB party, Aug. 1987, 2 km SW of (1728/6341), 2 km WSW of (1728/6341), 4 km SW of (1728/6342), and 6 km S of (1730/6340); MSB party, July 1993, 16 km by road E of (1723/6333). BUEN RETIRO; Santa Cruz 1717/6338 (MAPA) Delgadillo, 1950, province of Ichilo; de la Bar- rera, Oct. and Nov. 1954 (Hopkins and Roths- child, 1966: 120, gave coordinates as 1716/6345); AMNH-MSB, Sept. 1984, 4.5 km N of, 300 m (1714/6338); Anderson et al. (1987); 6 km N of, 300 m (1713/6338, restricted type locality of Ctenomys steinbachi Thomas, 1907b); MSB party, Aug. 1987, 3 km N of (1716/6338). BUSURUCUCU; Beni 1439/6617 Campamento in Estaci6n Biologica de Beni; Hinojosa and Salazar, June, Aug., and Oct. 1988. CABALLO, see Cabayu Nambi. CABALLO, PUERTO; Beni 1343/6521 (USBGN) AMNH, Sept. 1965. CABANA; Beni 1450/6621 An estancia; Flores, Nov. 1985, CBF, 40 km [E of] San Borja, between estancias Tajibos and Flor- ida; see also Ulla Ulla and Tabana. CABAYU NAMBI; Tarija 2138/6301 (MAPA) An estancia; Eisentraut, Oct. 1979, for ZFMK; 90 km SE Villa Montes, as Caballo Nambia; as El Cabayo Nambi (2138/6258) in USBGN. CABEZAS; Santa Cruz 1846/6324 de la Barrera, July 1955 (Fonseca, 1959: 93). CABRADA, POSTA LA; Chuquisaca 1905/6505 (Chubb, 1919: 5) Simons, Sept. 1901, 3500 m, as El Cabrado; Thomas (1902c: 116), type locality of Andinomys edax. CABRERA; Beni 1323/6440 (MARU) Webb, Feb. 1969. CACERES; Santa Cruz 1857/5747 Daveron, prior to 1941, to W of [Laguna] Cal- ceres near Bolivian-Brazilian border; also Sao Luis de Caceres. CACHIMAYO, RIO; Potosil918/6612 (USBGN) Tributary on the left bank of the rio Pilcomayo; Miller and Boyle, 1915; Miller (1918). CACHOEIRA DO BANANEIRA; Beni 1036/6525 (MAPA) Natterer, 1829; Wagner (1842b), type locality of Callithrix brunea (= Callicebus brunneus), by revisor's designation (Hershkovitz, 1963a); on rio Mamore. CACHUELA CAMP, see rio Abuna. CACHUELA ESPERANZA; Santa Cruz 1647/6314 An estancia; AMNH-MSB, Aug. 1984, 300 m. CACHUELITA; Beni 1312/6410 (MARU) Webb, 23-24 Apr. 1970, on rio Itonamas. CAFECES, see Montegrande. CAFETAL; Beni Barba (USNM), July 1966, San Ramon. CAIGUA; Tarija Eisentraut (1986) reported near here. CAIZA, see Villa Ingavi. CAJUATA; La Paz MNLP, 1982, 1817 m. CALABATEA; La Paz Carriker, Nov. 1934, 1981. CALABATEA; La Paz RAP, May and June 1991). N.L. 1325/6435 about 20 km SE 2110/6326 Catagonus from 1649/6715 ca. 1600/6750 1400 m; MNLP, Mar. 1458/6830 1990, camp (Emmons, CALACOTO, see La Paz. CALAHUASI; Cochabamba 1740/6446 Olrog, Aug. 1959, "250 km E of Cochabam- ba," also Carahuasi. CALIFORNIA; Santa Cruz N.L. Miller, Oct. 1915; a valley near Comarapa; Mil- ler (1918: 331). CALLIPAMPA; Oruro ca. 1835/6657 Carriker, June 1936, 12,000 ft (= 3660 m). CALUYO, see Kaluyo. CAMACHO, PUERTO; Pando 1131/6742 CBF party, Oct. 1991, in Manuripi province, 15 km NO de [= NW of] (1128/6750); 25 km de Comunidad Camacho at Madrid (coordinates un- known). CAMANDUI; Beni M. Ayala, May 1988, SW of rio Matos (prob- ably SW of 1451/6621); also see Katmandu. CAMARGO; Chuquisaca 2039/6513 (USBGN) 0. P. Pearson, Oct. 1955, 8500 f (= 2615 m); Hershkovitz (1962: 62); AMNH-MSB party, Sept. 1986, 68 km by road E of, 3400 m (2009/6517). CAMATAQUI, see Villa Abecia. CAMATINDI; Chuquisaca AMNH-MSB party, July 1991, 1 km S of (and in Tarija; 2100/6323), 650 m. 1997 87 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY CAMIACO; Beni 1519/6444 (AMNH field map) AMNH, Aug. 1965, = 16 km N Limoquije; "1515/6446" (Eger, 1974: 4); 6 km N of (1516/6444). CAMIARE; Beni 1415/6550 Cabot and Castroviejo, Apr. 1982, Yacuma province, near Estancia Venecia. CAMINO CUPESI; Santa Cruz N.L. Silva, Apr. 1938, in province of Cercado, now Ibaniez. CAMINO VILCHES; Beni 1304/6448 (MARU) MARU, 1963 and Feb. 1966, on rio Machupo near San Joaquin. CAMIRI; Santa Cruz ca. 2005/6334 (MAPA) Moll, prior to 1939, "Camiri and Choreti Regions, Chaco Frontier"; Cors Medina, 1941, 5 km S of Choreti in Azero province; Trusta, near Camiri, Aug. 1964; Acasigue and Becerra, Sept. 1966; Kerr, Sept. 1966, at Cordillera Yuti, 10 km S Camiri, (2010/6330); Cabot, Nov. 1984, 15 km [N of] (1955/6334); Cook and Gardner, July 1985, 17 km N of (1955/6334). CAMPAMENTO ENDE, see Colorada, Laguna. CAMPAMENTO I; see Sajama. CAMPAMENTO II; Cochabamba N.L. Tapia, Mar. 1990; in province of Carrasco; a field station of the University of Cochabamba. CAMPAMENTO 6 DE AGOSTO; Beni 1517/6705 Cabot, Nov. 1984 (Cabot, 1989; Salazar et al., 1994: 126). CAMPAMENTO 08; Beni 1444/6615 In Estacion Biologica de Beni, Gomez, Hino- josa, and Salazar, Aug. and Oct. 1988; or Cam- pamento Chima&n 08. CAMPO DE GUANACOS; Santa Cruz ca. 1900/6300 (Doutt, 1938: 100) J. Steinbach, Aug. 1909; Doutt (1938), 450 m, type locality of Galea spixi campicola (= Galea spixii campicola); in the western part of the prov- ince of Cordillera; also spelled Guanakos. CANDELARIA, PUERTO; Pando 1102/6613 Flores, Aug. 1985. CANUMA; La Paz Villalba, 1989, Cabania, rio Caniuma, province of Franz Tamayo. CAPAPARI N.L. Maddren, 1924, 10 mi S of, perhaps equals Carapari. CAPIRENDA, RIO; Chuquisaca ca. 2028/6404 Hershkovitz (1960: 561); see Ticucha. CAPIRENDO; Tarija 2107/6300 (MAPA) Stager and Bromley, Aug. to Nov. 1957, 15 km NE of (2101/ 6254), 7 km N of (2103/6300), 8 km NE of (2104/6257), 5 km S of (2110/6300), 7 km S of (2111/6300), 12 km SW of (2112/ 6305), 10 km S of (2113/6300), 13 km SW of (2113/6305), 20 km SW of (2115/6308); also spelled Capirenda; as Caipipendi (USBGN and MAPA) in Chuquisaca. CAPIVARA; Beni AMNH, July 1964. CARACATO; La Paz Baudoin, Aug. 1970; 1982, 2900 m. 1242/6308 (USBGN) 1659/6749 (USBGN) Anderson party, Apr. CARACOLLO; Oruro 1739/6710 (USBGN) 3840 m, (MHA); Simons, Mar. 1901; Thomas (1902a: 143) as Caracolto at 4000 m; Instituto de Ecologia, May 1979; Cabot and Yoneda, 1982. CARAHUARANI; La Paz ca. 1530/6758 Road from Carahuarani, via Tipuani Gold Mine, to Chuguini, 3000 m down to 2000 m and back to 3000 m, Tate, Apr. 1926; Tipuani is at 1533/6800. CARAHUASI; Cochabamba IML, 1959. 1740/6446 CARANAVI; La Paz 1546/6736 (USBGN) In province of Nor Yungas; Webster, Feb. 1978, 8 km SE of (1548/6734), in June 1978 at 606 m, July and Aug. 1979; 10 km W of (1546/6741), in June 1978; 20 km N of (1542/6736), in Jan. 1978; 5 km E of (1546/6734); Creighton, May 1978, 20 km (by road) NNE of (1542/6735), 2000 ft; Schmitt and Cole, two localities-Mar.-Apr. 1979, 6.6 km (by road) downstream from Caran- avi (1538/6739), in rio Coroico Valley, east bank, 653 m; and, June 1979, 35 km (by road) N of Caranavi at Serranfa Bella Vista (1540/6735), 1650 m; Webster, 1979 (Webster and Jones, 1980), 606 m at the town; Remsen and Cardiff, July 1980, 47 km (by road) N of Caranavi at Ser- rania Bella Vista (1538/6732), 1350 m; Hinojosa, Mar. 1988, 5 km N of (1544/6735), and 8 km N of (1541/6731); EBD party, date uncertain, 2 km E of (1546/6735). CARANDA; Santa Cruz 1732/6331 MSB party, Aug. 1987, 1.5 km SW of (1733/6332), 2 km S of (1733/6332), 3 km SE of (on rio Metillas, 1734/6331). CARANDAYTI; Chuquisaca 2045/6304 (USBGN) Bromley, Aug. to Oct. 1957, also spelled Car- andaita or Carandaiti; Stager, July 1957, 10 SE of (2049/6308); 18 km SE of (2053/6312); 30 km SE of (2058/6251); 35 km S of (2106/6304); AMNH-MSB party, July 1985, 4.5 km by road W of (2045/6306), 3.8 km by road E of (2046/6303), 9 km by road E of (2046/6300), 9.7 km by road E of (2046/6300). 88 NO. 231 ANDERSON: MAMMALS OF BOLIVIA CARAPARI; Tarija 2149/6346 (USBGN) Budin, Apr., July, and Aug. 1924, 1000 m; Tho- mas (1925), type localities of Akodon sylvanus pervalens (p. 579, = A. pervalens); Mazama sar- ae (p. 581, = Mazamna americana sarae); Ory- zomys legatus (p. 575); Thomas (1925: 580) as Carapiri; type locality of Oxymycterus paramensis jacentior; F Steinbach, field number 653, year un- known; AMNH-MSB party, Aug. 1991, 3 km WNW of (2148/6347), 850 m. CARAVANI, see San Joaquin. CARICARI, see Kari Kari. CARLAZO; Tarija 2128/6432 (USBGN) Budin, Mar 1924, Feb., Mar., and May 1925; Thomas (1926b: 322), 2200 m and 2300 m, "40 km to the east of" (2128/6410); and type locality of Marmosa janetta (p. 327, = Thylamys elegans venusta), see also Tarija. CARMEN; Beni 1356/6340 (MAPA) C. Gans, Feb. 1954; MARU, 1963, or El Car- men, on rio Blanco. CARNAVAL; Beni, see Espiritu. CARRASCO, PARQUE NACIONAL; see Villa Tunari. CASABINDO, CERRO; N.L. Cabrera (1961: 558). CASARA, see Bahia de los Casara. CASARABE; Beni 1448/6414 (USBGN) Anderson, Aug. 1983, 230 m, 6 km W of (1454/6422); Ruedas and Cuenca, Aug. 1985. CASCAJAL; Beni 1213/6513 (USBGN) AMNH, May 1965, collected at Cascajal and opposite Cascajal on E side of rio Mamore. CASTEDO, see San Ramon. CASTOR, see Khastor. CATAGAITA; Potosi 2050/6540 Pearson, Sept. 1955, 30 mi WNW of (2043/6557), 11,000 ft (= 3385 m); also spelled Cotagoita or Cotagaita. CATAVI; Potosi 1824/6636 (USBGN) Carriker, May 1936, 12,600 ft (= 3840 m). CATMANDUJ, see Katmandu. CAVINAS; La Paz 1236/6701 (USBGN) Dasyprocta from zoo, in USNM, died 1927. CAYOBA; Beni 1310/6413 (MARU) Webb, Feb. 1969 and Apr. 1970, or La Cayoba; SNCFHB, Jan. and Feb. 1985 (Torres et al., 1988). CEDRITO, EL; Santa Cruz 1748/6311 Tapia, Sept. 1992, cave near rio Pirai at west edge of Santa Cruz de la Sierra. CENTINELA; Beni N.L. MARU, 1963, on rio Machupo, also spelled Centenela. CENTRO DIECIOCHO; Pando 1036/6647 RAP team in June and July 1992, 18 km NNW of San Juan de Nuevo Mundo, a seasonally used castania camp. CENTRO GRANDE (Centro Branda); Pando 1120/6908 (USBGN) Izawa, Nov. and Dec. 1979. CERCADO, see Andr6s Ibaiiez. CERDAS: Potosi 2048/6629 (USBGN) Carriker, Feb. 1938, 3900 m. CERRO, see names such as Amboro, Itahuaticua, etc. CERRO COLORADO; Santa Cruz 1927/6221 An estancia; EBD party, Sept. 1986; 10 km S of (1933/6221); 50 km S of, at Perforaci6n (1955/6233); 58 km S of (2000/6233). CHACALTAYA, MT; La Paz 1620/6808 (USBGN) Barker. CHACO; when used as a proper noun as in Gran Chaco, the word is capitalized. When used in the general sense of any clearing or open area, it is not capitalized. The same dichotomy of usage arises with the words yungas and valles. CHACO, EL; Beni, see Espiritu. CHACO LEJOS; Beni 1324/6442 (MARU) Barba, July 1966, ca. 20 km SE of San Ram6n; also as Chaco Lejo. CHACO MAIZ; Beni 1333/6429 An estancia; Yates and Salazar, Mar. 1993, 15 km NW of El Valle. CHALLANA, RiO, see Guanay. CHALLAPATA; Oruro 1854/6647 (USBGN) Simons, Oct. 1901, 3800 m, east of Lake Poop6 (Thomas, 1902b: 225), type locality of Phyllotis hirtipes (= Eligmodontia puerulus); Thomas (1902e: 161), type locality of Marmosa elegans pallidior (= Thylamys pallidior); Anderson, Nov. 1979, 45 km and 123° [= compass bearing] from Challapata (1910/6625); Galictis at CBF (Yensen et al., 1994: 408). CHAPARE; Cochabamba 1709/6530 F B. Steinbach, June and July 1927, at 2000 m (the old road from Cochabamba into the Chapare valley passed the 2000 m elevation just above El Palmar at about 1709/6530), Sept. 1939, at 2700 m, Dec. 1941, Nov. 1948; Zischka, 1945, 1954, 1962 (Munich, R. M. Wetzel, personal commun.); mouth of rio at 1558/6841; Yoneda, Sept. 1982. A large area is known as the Chapare. 89 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY CHAPARE, RIO; Cochabamba 1558/6442 Carriker, Aug. 1937, 250 m.; Arce Pereira et al. (1963) reported various mammals from the rio Chapare, without specimens. See also rio Ichilo. CHAQUECAMATA, see Choquecamata. CHARAL, EL; Beni 1523/6706 CBF, 1989, 6 km on road to Yacumo from Qui- quibey. CHARAQUA; Santa Cruz 1948/6313 (USBGN) Wavrin, (Myotis), BM; de la Berrera, July 1955 (Fonseca, 1959: 93), or Charagua. CHARAZANI; La Paz 1512/6902 (MAPA) Anderson, Sept. 1980, 3450 m, also known as General Juan Jose Perez; also 2 km W of (1512/6903). CHAROPLAYA; Cochabamba ca. 1636/6637 Simons, May and June 1901; probably on the rio Santa Elena rather than the headwaters of the rfo Secure, at 1300 to 1400 m, northward of La- gunillas (Chubb, 1919); Thomas (1902a), type lo- calities of Coendou simonsi (p. 141, = C. bicolor simonsi), Oryzomys yunganus (p. 130), and Proe- chimys securus (p. 140, = P. longicaudatus se- curus); Allen (1915a: 205) as Charumplaya; Tho- mas (1916h: 478), type locality of Oxymycterus doris (= Oxymycterus inca doris); usually spelled Charuplaya. CHASQUIPAMPA; La Paz 1631/6800 (Yoneda, 1984) Yoneda, Apr. 1982, 3750 m. CHATARONA; Beni 1422/6728 (USBGN) Carriker, Sept. 1934, 180 m, as Chaterona. CHIANTA; Oruro N.L. A Phyllotis presented to the British Museum in 1912 by Frank A. Moss; 12,000 ft (= 3690 m), Hershkovitz (1962: 247). According to Hershkov- itz's map (p. 245), Chianta is very near Challa- pata, Oruro; also spelled Chayanta. CHICO, RANCHO; Santa Cruz 2015/6234 EBD party, Sept. 1986. CHIJCHIPA; La Paz 1608/6744 (USBGN) IBBA, Feb. to Apr. 1985, in Nor Yungas prov- ince, ca. 1400 m; or Chijchipani, Ruedas, Aug. 1986; AMNH-MSB party, as Chijchijpa, at 1114 m (1609/6745 GPS) and 1224 m (1609/6744 GPS). CHILCANI; La Paz 1544/6840 Olalla, July and Aug. 1938, near Mt. Illampu, 3700 m (Patterson, 1992: 5). CHILEAN BORDER; La Paz 1730/6930 Specimens in Frankfurt, "at border to Bolivia and Peru." CHILILAYA; La Paz 1613/6827 (Rusby, 1933) 0. and G. Garlepp, Aug. 1893, on Lago Titi- caca; Simons, Sept. and Oct. 1900, 4000 m; also spelled Chililya. CHIMASI; La Paz 1530/6753 (USBGN) Simons, Jan. and Feb. 1901, June 1903, 1500 m, Yungas, as Chimosi. CHIMATE; La Paz ca. 1525/6800 Simons, Sept. 1900; Thomas (1901b: 369), at 700 m, type locality of Rhipidomys benevolens (= Oecomys bicolor bicolor); Thomas (1903b: 488), type locality of Sciurus castus (= S. spadiceus); spelled Chirimote by Gyldenstolpe (1932: 40), and Chimote by Tate (1932f: 20). CHIMATE, RIO; La Paz ca. 1525/6800 (Tate, 1926) Tate, Apr. 1926, 1900 ft (= 585 m); tributary of rio Mapiri about 4 hours below the rio Velique on the same side. CHIMATI; La Paz N.L. Pusch (1941: 209), type locality of Cebus apel- la sagitta (= C. apella pallidus). CHIMOREI, RIO; Cochabamba 1643/6449 (USBGN) Miller, Aug. 1915; Miller (1918: 311); AMNH, July 1965, 2 km N of mouth of the river (1642/6449); Stearman, Mar. and Apr. 1991, in Carrasco province; see also rio Ichilo; also spelled Chimori. CHIMORI; La Paz ANSP. N.L. CHIMOSI; La Paz 1530/6753(USBGN) 1700 m; Thomas (1910c: 505), type locality of Dasyprocta variegata yungarum (= D. punctata yungarum); see Chimasi. CHINIRI; La Paz 1516/6745 (USBGN) Carriker, Aug. 1934, 335 m, on or near rio Kaka. CHIPIRIRI; Cochabamba 1611/6518 Rio; Arce Pereira, 1962, (Arce Pereira et al., 1963). CHIPIRIRI; Cochabamba 1635/6525 Cabot, Aug. 1988, 450 m, cercanias de San Pe- dro. CHIQUITOS; Santa Cruz 1750/6048 300-1000 m, d'Orbigny; province in llanos of E Santa Cruz; San Jose de Chiquitos (1749/6045); Kreig, 1925, 1926; Olrog, ca. 1960; San Jose de Chiquitos, Becarra, Oct. 1966; Heltne, Freese, and Whitesides, primate census, Aug. 1975 (Heltne et al., 1976: appendix I; near Esperanza at 1730/6052 some 40 km N and 12 km W of San Jose de Chiquitos; near Natividad at 1755/6055 some 20 km W and 10 km S of San Jose de Chi- quitos; and at 1750/6035 some 20 km E of San NO. 231 90 ANDERSON: MAMMALS OF BOLIVIA Jose de Chiquitos on road to Robore); Cook and Moore, Oct. 1984, 4 km S and 24 km E of (1752/6031); Olrog, Aug.-Sept. 1959. CHIVE; Pando 1223/6835 (USBGN) Yoneda, July 1982, as "1250/6840"; Harada, Sept. 1984; AMNH-MSB party, July 1986. CHIYIJTE N.L. E B. Steinbach, Feb. 1939. CHOCAYA; Potosi 2115/6545 (Paynter et al., 1975) Carriker, June 1936, 3660 m. CHONTA, LA; Santa Cruz N.L. MNK party, Oct. 1992, on rio Negro. CHOQUECAMATA; Cochabamba 1655/6637 (USBGN) Simons, July 1901; Thomas (1902a), type lo- calities of Oxymycterus paramensis (p. 139, = 0. paramensis paramensis) and of Akodon puer (p. 136, = A. lutescens puer), 4000 m on high para- mos northwest of Cochabamba; usually Chuqui- camata; Reed, 1939, mountains near, as Chaque- camata. CHORETI; Santa Cruz 2003/6335 (MARU) MARU, Sept. 1966; see also Camiri. CHORO, El; Cochabamba 1656/6642 (USBGN) Simons, May, July 1901, 3200-3500 m, en el Alto S6cure, province of Ayopaya; Thomas (1902a), type localities of Akodon bacchante (p. 138, = Chroeomys jelskii bacchante), Akodon fu- meus (p. 137); Phyllotis lutescens (p. 131, = Phyl- lotis osilae osilae); Conepatus chorensis (p. 126, = Conepatus chinga rex), Eligmodontia carilla (p. 133, = Calomys lepidus carillus); Thomas (1920b: 363), type locality of Eptesicus montosus (= E. furinalis montosus); F B. Steinbach, June, July, and Aug. 1953, 3500 m. CHORO, EL; Oruro 1821/6708 AMNH-MSB, Oct. 1986, Chaetophractus ob- tained by Carminia Miranda Moreira and Freddy Caceres Vega. CHUHUAYACO, see Chuyayacu. CHULUMANI; La Paz 1624/6731 (USBGN) d'Orbigny, 1830, 2000 m; Kalinowski, 1896; Simons, Nov. and Dec. 1900, Jan. and Feb. 1901, at 1600, 2000, and 2200 m; Allen (1901c: 411), type locality of Metachirus nudicaudatus bolivi- anus; (Thomas, 1907b: 163) type locality of Gri- son furax luteola (= Galictis vittata luteola); as 1800 m, Thomas (1917a: 158), type locality of Cavia nana (= C. aperea sodalis); Olalla, Sept. 1938, 1740 m (Patterson, 1992: 5); Lonnberg (1941: 42), type locality of Aotus roberti (= Aotus azarai boliviensis); capital of Yungas Province; 1734 m. CHUQUINI; La Paz 1534/6804 (USBGN) Tate, Apr. 1926, ca. 3000 m, "A house on the Gritado River which is tributary to the Tipuani on its right-hand side," also "road from Chuquini to Tora, 3000-3400 m." CHUQUISACA, see Sucre. CHURO NEGRO, see Comarapa. CHUSMAS, RIO; N.L. Olrog, Aug. 1957, 5 km W of Las Madrejones. CHUSPIPATA; La Paz 1618/6748 (IGM) Remsen party, July 1981, 1 km S of, 3050 m (also see Sacramento Alto and Cotapata); Mer- cado, Nov. 1990; CBF, Apr. 1992, Cerro Chuspi- pata. CHUYAYACU; Chuquisaca 1943/6352 Arduz, Aug. 1951; AMNH-MSB party, Aug. 1990, 2 km E of Chuhuayaco, 1200 m (1943/6351). CIENEGA, see Vallegrande. CINCO; Beni 1249/6503 MARU, 1963, designation for km 5 on road from Puerto Siles to San Joaquin, coordinates are as measured from Puerto Siles. COBIJA; Pando 1102/6844 (USBGN) On "Rio Madeira"; the alleged origin of a specimen of Saguinus imperator (Hershkovitz, 1977: 684); Hill (1957: 237), as Colija; Freese, 1975, as 1110/6858; Heltne et al. (1976: appendix I; 20 to 25 km west of Cobija on rfo Acre, 1100/6845; on foot between Espirito Santo, 1100/6858, to rio Nareuda, at 1120/6900; and on rio Acre from Espirito Santo to Buenos Aires, at 1120/6900); Swing, Feb. 1982; D. Lopez for MNLP, no date; see also Tahuamanu; 10 km S of Cobija (1108/6844); Harada, Aug. 1984; Iseki et al. (1985). COCABOMBA N.L. J. Steinbach, June 1920, Aug. 1921, Dec. 1928; on rio Tajajos. COCAPATA; Cochabamba 1731/6517 Kessler, Aug. 1991, 3200 m, above Cocapata on Cochabamba to Santa Cruz road (1730/6516). COCAPUNCO; La Paz 1530/6829 (Tate fieldno- tes) Tate, Mar. 1926, 3100 m, "on trail from Sorata to Mapiri via Ingenio," also "road from Coco- punco to Pararani"; Anthony (1926: 2), type lo- cality of Thomasomys oreas. Hershkovitz (1959b: 45), type locality of Mazama chunyi (= M. bri- cenii chunyi). COCHABAMBA; Cochabamba 1724/6609 (USBGN) Simons, Apr. 1901, at 2400 m and at 2600 m; J. Steinbach, at 2700 m (Hershkovitz, 1962: 248); Thomas (1902a), type locality of Akodon varius 1997 91 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY (p. 134), at 2400 m, and of Conepatus porcinus (p. 128, = Conepatus chinga rex); Collins-Day Expedition, 1915 (Osgood, 1916: 199); F Stein- bach, Dec. 1928, Aug. 1929, July 1936; EBD party, Sept. 1985, 15 km E of (1724/6559); Cabot, Sept. 1988, at Country Club, 2560 m, near Co- chabamba; Szwagrzak, Nov. 1985, owl pellets; Serrano, 1986, owl pellets; Kessler, Aug. 1991, km 79 on Cochabamba to Oruro road (coordinates unknown); capital city of department. COIMBRA; Brazil 1129/6518 (RON) Used to designate places in Beni opposite Coimbra (1129/ 6519) and 5 km SE of Coimbra (1 131/6517) AMNH, July 1965. COLCHANI, EL; Potosi 2020/6656 EBD party, Nov. 1982, entre Colchani y Uyuni (2024/6653). COLINA CACHAPA; La Paz 1640/6800 Mercado, Feb. 1987, Feb. 1989. COLLANA; La Paz 1638/6858 (Yoneda, 1984b) Baudoin, Aug. 1968; Yoneda, Apr. 1983, 3750- 3800 m. COLOMI; Cochabamba 1721/6552 (USBGN) F B. Steinbach, Jan. and Aug. 1928, 3800 and 4000 m, June, July, and Dec. 1936, 3075 m, Dec. 1941; 5 km N of (1718/6552); 1993 expedition; 13 km N of, 3125 m ; Poseidon Resort, 12.5 km N of, 3200 m (1713/6553, GPS data); 16.5 km NW of, 3500 m (1714/6557). COLONIA PIRAY; Santa Cruz 1620/6339 Minezawa et al. (1985), Alouatta. COLORADA, LAGUNA; Potosi 2217/6747 (USBGN) Walcott, Feb. 1924, 4196 m.; EBD party, Sept. and Nov. 1982; AMNH-MSB, Sept. 1986, 21 km by road SE of ENDE camp, 4500 m (2218/6744), 7 km E of ENDE camp, 4280 m (2210/6745), 2 km E of ENDE camp (2210/6742); Salazar, Oct. 1989, 1 km E of Campamento Sol de Manana II, (2210/6743), 22.5 km E of ENDE camp (2210/6730). COLORADO, CERRO; Santa Cruz 1927/6221 Iba'niez, Sept. 1986, hacienda on rio Parapeti. COLORADO, EL; Chuquisaca EBD party, Nov. 1982, near Villa Abecia (2100/6523). COLORADO, RIO; La Paz 1547/6717 Mann, Sept. 1921. COMARAPA; Santa Cruz 1754/6429 (USBGN) F. B. Steinbach, Sept. and Oct. 1926, at 2500 m, as Comparapa (USNM); Pearson, Sept. 1955, 5 mi. W of (1754/6434), 7500 ft (= 2310 m); Crossin, Mar. 1973, 28 km W of (1751/6440), Dec. 1972, 30 km W of (1751/6440), in Cocha- bamba; Schmitt and Cole, Feb. 1979, 6 km S and 10 km E of (1757/6424), in rio Pulquina Valley, Propiedad Bado Hondo, 1527 m; Anderson, Nov. 1979, 13 km SE of, by road (1758/6425); Creigh- ton, Feb. 1980, in the Siberia cloud forest, 25 km (by road) W of (1751/6440), 2800 m, in Cocha- bamba, originally judged to be in Santa Cruz; Cabot, July 1982, Mar. 1983, 11 km [by road] W of, at Churo Negro (1755/6433), 2180 m; EBD party, Mar. 1983, 3 km NW of Churo Negro (1751/6440); EBD party, Nov. 1983, 30 km W of (1751/6441); AMNH-MSB party, Sept. 1984, 31 km (by road) W of, 2800 m (1751/6442), in Co- chabamba; 28 km by road W of, 2800 m (1751/6440), in Cochabamba; Myers and Patton (1989a: 4), type locality of Akodon siberiae; Hi- nojosa et al. (1987: 15), type locality of Oxymyc- terus hucucha; 21 km by road W of, 2900 m (1751/6437); 15 km by road W of, 2700 m (1753/6435); 8.5 km by road W of, 2300 m (1754/6432); 1 km N and 8 km W of, 2450 m (1755/6434); 3 km by road SE of, 1700 m (1757/6430); 5 km by road SE of, 1695 m (1758/6429); AMNH and MNK, Nov. 1987, 5 km SW of, 1750 m (1757/6432); 8 km SW of, 1700 m (1758/6433); Glanz and Anderson (1990; 23), type locality of Abrocoma boliviensis; AMNH- MSB party, July 1992, 1800 m, 4 km E of (1754/6427). COMAUCHI; La Paz 1658/6829 BM 13.3.18.1, Ctenomys leucodon; EBD party, Felis concolor, 1982, on map as Comanche. CONCEPCION; Santa Cruz 1615/6204 (USBGN) d'Orbigny, 1831; Mision, 490 m; d'Orbigny and Gervais (1847), type locality of Noctilio af- finis (= N. albiventris). Goodfellow, July 1919; Thomas (1921a: 136), type locality of Ctenomys goodfellowi (= C. boliviensis goodfellowi) is Es- peranza (1628/6116), near Concepci6n. Davis, Mar. 1986 (she gave coordinates as 1608/6202); Nov. 1985, 80 km by road SE of, at Zapocoz (1633/6140); Oct. 1985, 35 km S of, at El Carmen (1625/6200); Szwagrzak, 1980s, year unknown, owl pellets (1615/6204). CONDO; Oruro 1903/6644 EBD party, Nov. 1982. CONQUISTA; Beni N.L. A hacienda; Miranda, 1990 and 1991, in prov- ince of Ballivian. CONSUELO, EL; Beni 1420/6715 (Gyldenstolpe, 1945, map) A. M. Olalla, 1937, 1938; Lonnberg (1939: 17), type locality of Callicebus modestus; Lonnberg (1942: 30), estancia on pampas about 12 km E of Reyes; (Patterson, 1992: 5). COPACABANA; La Paz 1610/6905 (USBGN) Yepes and Crespo, Mar. 1941. NO. 231 92 ANDERSON: MAMMALS OF BOLIVIA COPACABANA, see also Montero. CORANI; Cochabamba 1713/6552 (GPS) June and July 1993, MSB expedition, 2630 m, hydroelectric plant; July, Laguna de Corani, 12.5 km N of Colomi, 3200 m (1714/6553, GPS data). CORDILLERA [RIO GRANDE]; Santa Cruz J. Steinbach, 1909 (FMNH). COROICO; La Paz 1610/6744 (USBGN) In province of Nor Yungas; Creighton, May 1978, 31 km (1618/6748, on road to La Paz) southward of river crossing near Coroico, 2900 m; S. and R. J. Anderson, May 1982, 1715 m; EBD party, Oct. and Nov. 1982, 4 km from, on road to Caranavi and on right bank of rio Coroico (1611/6743); Hinojosa, 1988, 3 km N of, at co- munidad Marka (1608/6744). COSMINI; La Paz 1655/6812 (USBGN) Simons, Mar. 1901, as 4300 m; Thomas (1902a: 134). COSTA MARQUES; Brazil 1228/6417 (RON) Used to designate localities in Beni across the no Itenez, opposite bank (1229/6417), 500 m aba- jo de (1229/6417), and 40 km by river above town (1232/6403); 4 km above (1229/6415), 1 km above (1228/ 6417); 2 km above (1229/6416); 1.5 km below (1229/6418); 0.5 km S of river and 4 km SW of town (1230/6418); AMNH, 1964; also spelled Costa Marquez. COTACAJES, RIO; Cochabamba 1630/6650 CBF party, Aug. 1992, San Antonio, in prov- ince of Ayopaya; Seque Rancho (1635/6645). COTA COTA; La Paz 1633/6805 MNLP, Nov. 1982; see also La Paz. COTAGAITA, see Catagaita. COTAPATA; La Paz ca. 1613/6754 Remsen and Cardiff, June 1980, 4.5 km by road WNW of Chuspipata, at 3300 m; Hinojosa, Feb. 1991, at Santa Barbara (coordinates uncertain); Apr. 1992, 4 km N of (1611/6754). COTOCA; Santa Cruz 1745/6257 (MAPA) Becerra, Sept. 1966; Flores, May 1985, and Szwagrzak, Oct. 1985, owl pellets; in Province of Iba'nfez; MSB party, May 1988 (1746/6258). CRETA, ISLA DE, see Estaci6n Biologia de Beni. CRUCE VENTILLA; Oruro AMNH-MSB party, Sept. 1986, 7 km S and 4 km E of, 3450 m (1908/6607). CRUCES, LAS; Santa Cruz 1746/6321 UAGRM party, June and July 1990; 2 km SW of (1747/6322); 3 km SW of (1747/6322). CRUS, LA; Beni 1320/6340 An estancia, Flores, Sept. 1983. CRUZ, LA; Pando AMNH/MSB, Aug. 1986, 170 m. 1124/6713 CUCHARCA, RIO; Beni Bats reported by Barquez (1984a) from Santa Rosa, rio Cucharca, 70 km N of Trinidad, date and collector not given. Cocharcas on some maps. CUCHICANCHA; Cochabamba ca. 1721/6543 (MHA) Miller and Boyle, 1915, 3400 m; or Cuchacan- cha. CUESTA CUCHO; Cochabamba 1715/6545 Olalla, May 1938, 2300 m (Patterson, 1992). CUEVAS; Santa Cruz 1807/6343 An estancia; de la Barrera, November 1954; AMNH-MSB party, May 1991, 1 km NE of (1811/6344), 1300 m. CUEVO; Chuquisaca 2027/6332 de la Barrera, July 1955 (Fonseca, 1959). CUMBRE, LA; La Paz 1622/6803 (USBGN) Tate, Feb. 1926, 15,200 ft (= 4675 m); the highest point reached by the former La Paz-Yun- gas railroad; Carriker, Jan. 1935; Sanborn (1950), type locality of Hesperomys lepidus montanus (= Calomys 1. carillus); Mercado and Miralles, 1987. CUPESI, PUESTO DE; Santa Cruz 2012/6233 EBD party, 1986, donated skins, 98 km S of Cerro Colorado; 40 km [S] of, at Rancho Chico, in Cordillera province. CURICHE; Santa Cruz 1844/6316 (USBGN) Rio Grande, J. Steinbach, Aug. 1909 (FMNH); de la Barrera, 1955. CURICHI GRANDE; Santa Cruz ca. 1530/6016 (MAPA) As "about 150 S., a few miles from Brazilian border," Daveron, May 1932, USNM, about 15° 30' on Brazilian border, Daveron, Dec. 1935; name means simply 'big swamp.' MAPA uses "Bahia Grande." CURICHE, RIO; Beni ca. 1236/6354 (MAPA) AMNH, July 1964, at mouth of W end of Isla Capim. CURIRABA, RIO; Beni 1450/6623 Wilson and Salazar, Sept. 1987 (Wilson and Sa- lazar, 1990: 47); Salazar, July 1988 (1451/6621); also spelled Curiaba or Cureraba; see Beni Re- serve. CURUYUQUI; Santa Cruz 1846/6214 Emmons, Oct. 1991 (Parker et al., 1993). CUSI, LAS; Beni Tello, Sept. 1986. 1355/6450 CUTIKHUCHU; La Paz 1608/6807 (GPS) MSB party, June 1993, 2697 m, or Cuticucho. 93 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY CUYAMBUYO; Tarija 2215/6436 (USBGN) Creighton, Jan. 1980; 4 km (by road) N of (2213/6436), 980 m; 8 km (by road) N of (2212/6436), Hacienda Conzelmann, 1000 m; AMNH party, Aug. 1991, 3 km SE of (2216/6433), 900 m. Ch, see in list with names beginning with C. DESAGUADERO, RIO; La Paz 1634/6902 (USBGN) Pentland, 1850; Thomas (1911: 256). DESIERTO; Beni 1342/6732 (Gyldenstolpe, 1945) Olalla, 1937 (Lonnberg, 1940c); on rio Beni, also termed El Desierto. DOLORES, RIO; Santa Cruz 1722/6341 (USBGN) Province of Sara; headwaters of Yapacani, near Buenavista; J. and F Steinbach (Sanbom, 1932a: 175). ECUADOR; Beni MARU, Mar. 1971. 1412/6526 (MARU) EL, used with names such as Astillero, Choro, Consuelo, etc. ELE ELE, see Mizque, Rio. EL PUENTE; department uncertain N.L. Tello, July 1986. EMBOCADA, LA; Beni 1503/6658 Creighton, Nov. 1979, 1 km E, Estancia La Ca- bania, 600 m, on some maps spelled as Embos- cada. ENCANTO, EL; Beni, see Espiritu. ENCANTO, EL; Santa Cruz 1436/6042 Cascades; MSB party, July 1990; Emmons, Nov. 1992. ENGENHO DO CAP. GAMA; Brazil N.L. "Du rio Guapori." Natterer, 8 Sept. 1826 (Pel- zeln, 1883: 18, 104). ENTRE RIOS; La Paz AMNH-MSB party, July 1992, 1000 m, 2 km W of (1539/6726). ENTRE RIOS; Tarija 2132/6412 (USBGN) Carriker, Oct. 1936; Gilmore, July 1943, in Province O'Connor, 1200 to 1300 m; Crossin, Jan. 1973, 25 km NW of (2123/6421); AMNH- MSB party, Aug. 1991, 5 km NNW of (2129/6412), 1600 m. EPIZANA, Cochabamba, see Siberia Cloud For- est. ERQUIS; Tarija 2128/6448 AMNH-MSB party, July 1991, 2100 m. ESCOMA; La Paz 1540/6908 EBD party, Nov. 1984, at Lago Titicaca. ESPEJILLOS; Santa Cruz 1758/6327 Ibainez, 1986, 10 km W of San Jose, in province of Andres Ibainez. ESPERANZA; Beni 1401/6446 Pilleri (1977b: 21), Puesto La Esperanza on rio Ipurupuru. ESPERANZA; Beni 1425/6532 (MARU) MARU, 1963; Barba, Sept. 1966, 42 km NE of San Joaquin (1251/6436); also La Esperanza, see San Joaquin for another place of this name. ESPERANZA; La Paz 1749/6847 (USBGN) 4000 m, northeast of Mt. Sahama in province of Pacajes, G. Garlepp, Apr., May, June 1897; Thomas (1898a), type localities of Conepatus rex (p. 278, = C. chinga rex), Phyllotis (?) garleppii (p. 279, = Galenomys garleppii), Chinchillula sa- hamae (p. 280), Akodon berlepschii (p. 281, = Akodon albiventer berlepschii, Cavia niata (p. 282, = Microcavia niata niata); Thomas (1907a: 443), Viscaccia lutea (= Lagidium viscacia cu- vieri); Thomas (1914: 357), Pseudalopex cul- paeus andina; F Steinbach, Nov. 1941, at 4200 m (Hershkovitz, 1962: 247); Cabrera (1958: 230) cited as "Esperanza, departamento de Oruro" and (1961a: 496) as department of La Paz. ESPERANZA; Santa Cruz 1628/6116 Thomas (1921a: 136), type locality of Cteno- mys goodfellowi. ESPERANZA; Santa Cruz 1954/6259 Zalles, May 1990, Rancho La Esperanze or Ha- cienda Nueva Esperanza, 40 km from Charagua. ESPERANZA; "Tarija," see Concepci6n, Santa Cruz. ESPIRITU; Beni 1413/6640 La Estancia Elsner Hermanos "Yacuma" (Han- agarth, 1993: 4); coordinates differ as given by different sources (e.g. Hanagarth, 1993: 4, 1408/6624; Instituto Nacional de Estadistica, 1982: 255, 1413/6640; USBGN, 1412/6640; MAPA, 1413/6640; Aguirre, 1994: 6, 1413/6624; Aguirre and Urioste, 1994: 71, 1413/6624); Han- agarth, Instituto de Ecologia, Sept. 1980, May and Aug. 1981; Mendez, Jan. 1986; Salazar and Sar- miento, June 1986; Aguirre, June into Oct. 1992. Thirteen collecting sites within 6 km of the estan- cia, as follows, vicinity of the ranch, tusecal de Espiritu, islas de bosque Peru, El Encanto, and Don Herbert (all near 1413/6640); tajibal de Peru (1413/6641); isla de bosque Vacia (1413/6639); isla de bosque and tajibal de Espiritu Viejo (1411/6638); islas de bosque El Encanto and Don Herbert (1413/6640); Arroyo Carnaval y Puerto Senda and isla de bosque El Chaco (1410/6640); rio Yacuma (1410/6639); "Pastizal" (1413/6640). ESTACI6N BIOL6GICA DEL BENI; Beni 1451/6621 EBD parties, Oct. 1983 to Nov. 1985; 6 km N of Isla de Creta (1445/6621); 5 km N of rio Cu- NO. 231 94 ANDERSON: MAMMALS OF BOLIVIA riraba, Isla de Creta (1449/6621); Isla de Creta, 3 km N of EBB (1449/6621); 60 km E of San Borja on road (1451/6621); Arroyo Aguas Negras (co- ordinates unknown); AMNH, MSB and IBBA, Aug. 1985; 1 km SW of (at Totaisal, 1451/6621), 6 km E of (at rio Matos, 1451/6617); Sarmiento, 1986; also known as Estancia El Porvenir; Cabot et al. (1986); Wilson and Salazar (1989); Merca- do, 1987, 15 km N of Porvenir (1443/6621); Mir- anda, Aug. 1990; Paynter, Oct. 1990; other des- ignations are Estacion Biologica Beni and Esta- cion Biologica de Beni; see also Beni Reserve, Florida, Trapiche, Pascana, Campamento 08, Porvenir, Villa Dorita, and Busurucucu. ESTANCIA, see name of the estancia, such as Estancia Bethania. EUCALIPTUS; Oruro 1735/6731 (MAPA) Schmitt and Cole, Dec. 1968; 11.2 km S and 0.8 km W of (1741/6731), at 3365 m; 12 km S and 1.6 km E of (1742/ 6730); and 9 km S of (1740/6731); 9 km S and 0.8 km W of (1740/6731); see also Huancaroma. EVA EVA, SERRANIA; Beni 1536/6638 Hinojosa, Oct. 1990. EXALTACION; Beni 1316/6515 (USBGN) MARU, 1963; AMNH, Sept. 1965, 8 km N of Exaltacion (1312/ 6515), on rio Mamore; 2 km S of (1317/6515); 7 km S of (1319/6516). FIERROS, LOS; Santa Cruz 1430/6110 EBD party, Aug. 1986, 450 m, 2, 3, 3.5, 4, or 5 km N of, at Campamento Precimbrico (1432/6054); MSB party, June 1990, 3 km E de Campamento "Los Fierros" (1433/6052); MSB party, July 1991, 3 km S of (1434/6053); 4 km S of (1434/6053); 6 km S of (1434/6053); 7 km S of (1435/6052); 17 km S of (1433/6049); 23 km S of (1438/6045); 27.5 km S of (1438/6044); 52 km S of (1445/6035); see also Florida and Moira. FILADELFIA; Beni MARU, 1963. 1259/6453 (MARU) FLOR DE ORO; Santa Cruz 1535/6034 MNK party, Sept. 1991, 187 m, Parque Na- cional "Noel Kempff Mercado." FLORIDA, see Pitiguaya. FLORIDA; Beni 1451/6621 Cabot, June 1985; Miranda, Aug. 1990, an es- tancia or hacienda near EBB. FLORIDA; Santa Cruz N.L. Delgadillo, Aug. 1950, near Floripondio, or near Agua Hedionda. FLORIDA, LA; in La Paz, see Pitiguaya. FLORIDA, LA; Santa Cruz 1436/6111 Village on rio Paragua; Iba'nez, Aug. 1986; 50 km E of (1436/6047); 38 km E of (1436/6054 = Los Fierros); Braza and Garcia (1988) observa- tions of primates at various localities from La Florida to 60 km E thereof. FLORIPONDIO; Cochabamba 1804/6445 de la Barrera, Feb. 1955 (Smit, 1987: 350). FORESTAL, LA; Santa Cruz N.L. MNK party, Apr. 1990, Florida province [or Andres Ibainez]. FORTALEZA; Beni 1412/6528 (MARU) Webb, Feb. 1971 and Nov. 1972, 95 km S of Santa Ana, in Yacuma province. FORTIN CAMPERO; Tarija 2255/6418 (MAPA) Carriker, Sept. 1936, 350 m. GENERAL PEREZ, see Perez. GLAVIR; Beni 1451/6610 An estancia, 65 km on road from San Borja to Trinidad; EBD party, Oct. 1985; adjacent to EBB. GRANDE, RIO; Santa Cruz 1551/6439 (USBGN) d'Orbigny, June 1831, at ca. 1745/6245 and ca. 1845/6415 (d'Orbigny, 1846: 162); J. Steinbach, Sep. 1909, ca. 1855/6320; Miller, Oct.-Nov. 1915 (perhaps near 1850/6416; Miller, 1918: 344); Steinbach, Oct. 1942, at Bajo Rio Grande, 1000 ft AMNH, Aug. 1965, 5 km NW (some specimens were erroneously labeled 4 km NE and some as 5 km NE) mouth of rio Grande, on rio Mamore (1550/6441), in Beni; Humberto Montero, Dec. 1988 (1740/6245). GRANJA, LA; Beni 1318/6409 Webb, Apr. 1970; Kunz, July 1971; Pine (1975: 321), type locality of Monodelphis kunsi, W bank of rio Itonamas, 4 km N of Magdalena. GUADALOUPE; Santa Cruz 1833/6405 (USBGN), 1827/6406 (MAPA) Marzana and Macchiavello, May 1949, 1800 m; as Guadalupe, 10 km S of Vallegrande. GUALEVA; Beni Kuns, Feb. 1972; in province of Itenez. N.L. GUANACOS; Santa Cruz 1849/6309 (USBGN) J. Steinbach, Sept. 1915, in Cordillera province; MCZ no. 26948, see also Campo de Guanacos. GUANAY; La Paz 1528/6752 (USBGN) Tate, Apr. 1926, 1800 m, "small, neatly laid- out village at junction of R. Tipuani and the Ma- piri," also "road from Guanay to Carahuarani, 1800-3000 m"; Carriker, Aug. 1934, 460 m; Creighton, June 1978, 5 km (by road) SE of, on rio Challana (1530/6750), 1100 ft; 20 km NNE of (1518/6748); also spelled Huanay. GUAPORJt, RIO; see Itenez, rio. GUARAYITOS; Santa Cruz1735/6146 (USBGN) Kreig, 1926. 95 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY GUARAYOS; Santa Cruz 1614/6037 (Hershkov- itz, in litt.) "Province in Santa Cruz"; d'Orbigny (1834: 89), type locality by restriction (Elliot, 1913) of Callithrix boliviensis (= Saimiri sciureus boli- viensis), as Sierras Guarayas; d'Orbigny (1837: plate 9), type locality of Noctilio rufipes (= Noc- tilio leporinus rufescens), as rio San Miguel, Lla- nos de Guarayos; at present there is no province of Guarayos in Santa Cruz. See also Ascension de Guarayos. Pilleri and Arvy (1977) mapped San Miguel at 1619/6026. GUARINA, see Huarina. GUARIPEMBI, see Guirapembi. GUAYARAMERIN (Puerto Sucre); Beni 1049/6525 (MAPA) AMNH, 1964, 1965; 2 km N of, on island (1048/6525); 1.5 km N, on island in rio Mamore (1048/6525); 1.5 km NW of (1048/6526); 4 kLm S of (1051/6525); 5 km S of (1052/6525); Fugler and Swing, Feb. 1982; Tello, July 1986. GUAYARAMERIN AND GUAYARA ASSU, CACHUELAS; Beni 1045/6525 (RON) Natterer, 1829 (Pelzeln, 1883: 127) as Cach- oeiras Guayara mirim and guacu. GUIRAPEMBI; Santa Cruz 1926/6231 EBD party, Sept. 1986, on rio Parapeti, in prov- ince of Cordillera, as 1930/6730. GUTIERREZ; Santa Cruz 1925/6334 (USBGN) de la Barrera, July 1955 (Smit, 1987: 350). Be- cerra and Acasigue, Sept. 1966, in province of Cordillera; Schmitt and Cole, Jan. 1979 and Feb. and Apr. 1984, 10 km E of, at Laguna Caucaya (1925/6329), 875 m. HABANA; Beni 1410/6449 An estancia, Pilleri, 1976 (Pilleri, 1977b: 21). HACHA, see Jhacha. HAMACAS; Santa Cruz 1744/6311 (USBGN) Silva, July 1938, also spelled Hamecas. HAMBURGO, see rio Beni. HARDEMAN; Santa Cruz 1620/6340 Harada, Sept. 1984. HENEY, RIO N.L. Pine and Wetzel (1975). HERBERT, DON; Beni, see Espiritu. HORCUS; Chuquisaca 1929/6433 (USBGN) Prince, March 1965, 80 km SE Sucre, or Hor- cas (USBGN, MAPA). HORQUILLA; Beni 1234/6425 (MAPA) MARU, 1963; AMNH, Sept. 1964, 15 km above Horquilla, on rio Machupo (1241/6432); 20 km above Horquilla, on rio Machupo (1243/6435). HOSANE, CERRO; Santa Cruz N.L. 1200 m, J. Steinbach, Aug. 1917; Tate (1933: 75) as Cerro Hosana. HOYADA, see Vallegrande. HUACARAJE; Beni 1333/6345 (USBGN) MARU, 1963; Acasigue, Nov. 1966, in Itenez Province; also spelled Huancaraje. HUACHI; La Paz 1540/6715 (USBGN) Mann, "1921-2" (USNM); also known as San Miguel de Huachi, on rio "Boki" [= Bopi]. HUACULLANI; La Paz 1627/6845 or 1658/6850? Instituto de Ecologia, Feb. 1979. HUAJCHILLA; La Paz 1637/6803 Yoneda, 1984 Instituto de Ecologia, Yoneda, Feb. 1982 to Feb. 1983, 3050 m; Mercado, Feb. and May 1982, 4 km SW of (1639/6805); EBD parties, Sept. and Oct. 1982, Oct. 1984. HUAKARANI: perhaps in Oruro N.L. Pujol, April 1969, "Departamento d'Urumo"; specimens in Paris, see Microcavia. HUANAY, see Guanay. HUANCARAJE, see Huacaraje. HUANCAROMA; Oruro 1739/6731 (MAPA 1980) Baudoin, Aug. 1968, Mar. and Apr. 1971, ranch near Eucaliptus; AMNH-MSB, Aug. 1984, 3720 m (1740/6729); 1 km W of, 3730 m (1740/6730); 2.5 km NE of, 3720 m (1739/6728); 3.5 km E of, 3720 m (1740/6727); 1 km E and 3 km S of, 3720 m (1742/6728); AMNH-MSB party, Oct. 1986, (1740/6729), 3.5 km E of (1740/6727, 1 km W of (1740/6730); 3 km W of, rio Desaguadero (1740/6732). HUANCASAYA; La Paz N.L. Villalba, Feb. 1990, prov. Franz Tamayo. HUANCHACA, SERRANIA; Santa Cruz 1425/6050 EBD party, Aug. 1986, or Serrania Caparus. HUANCUNI; La Paz 1544/6835 Olalla, July 1938, 3050 m (Patterson, 1992). HUANDA; Santa Cruz N.L. MNK party, 1990, in province of Andr6s Iba- fnez; see Wenda. HUANUNI; Oruro 1816/6651 Anderson and Olds, July 1983, at 4150 m; 3 km E [or NE] of (1815/6648). HUARACO; La Paz 1710/6755 (Yoneda's notes) Instituto de Ecologia, June, Nov., and Dec. 1980; Jan., May, Aug., Sept., and Nov., 1981; Feb. 1982; Pefiaranda, Jan. 1991 (as 1722/6738) and June 1991, as Huaraco-Antipampa (1727/6737); Kessler, Aug. 1991, 3700 m (1710/6755). 96 NO. 231 ANDERSON: MAMMALS OF BOLIVIA HUARI; Oruro AMNH-MSB party, Sept. 1986. HUARINA; La Paz 1612/6838 Simons, Aug. 1900, at 4000 m (191 lb: 255), type locality of Kerodon Galea musteloides auceps); as Guarina HUARINILLA, RIO; La Paz CBF, early 1990s. HUARMECHI; Tarija Skin of Felis jacobita from hur 3300 m. HUATAJATA; La Paz Niethammer, Nov. 1951. HUCHULA; Beni 1252/642 USNM no. 461038, Apr. 1970. HUCUCHA Specimen of Akodon albiventer in (Vanzolini, in litt.); hucucha is the word for mouse, so this may not be name at all. HUERRASCA; Beni MARU, Mar. 1965, Feb. 1967, 10 kl Ram6n. HUESOS, LOS; Santa Cruz Marzana and Macchiavello, Apr. 19 dillera Province, 900 m. HUMAHUACA; Argentina Cornalia (1865: 3), type locality of I ita "circa Potosi et Humacuaca in m elevatis"; this may refer to Humahuac Argentina, in which case the type spec have come from either Argentina or E HUMAITA; Pando 1203/6817 (M Flores, Sept. 1985. IBANES, see Santa Cruz de la Sierra. IBARE, RfO; Beni 1437/6457 AMNH, June 1965; 1 km abc (1438/6457); 24 km from mouth (1444 km from mouth (1447/645 1); 27 km f (1447/6451). IBIETO; Beni Townsend, Mateo, Mendoza, and o ous months from Mar. 1991 through also spelled Ibiato or Eviato. IBOPAITI; Chuquisaca 2001/635C Kreig, 1926, as Ibopeite. ICHILO, RIO 1557/6442 Delgadillo, Nov. 1950, near Buen boundary between Cochabamba and at the point given by Paynter et al. AMNH, July 1965, 13 localities on most measured as distances below (no] er is the rio Mamore if the Chapare an regarded as merging to form the TV 1900/6645 above (south) mouth of rio Chapare, two mea- sured below (north) mouth of rio Chimore, those (USBGN) on left bank in Cochabamba, those on right bank GThomas in Santa Cruz; right bank, 5 km N of mouth of auceps (= Chapare (1555/6442); right bank, 2 km N of 3 auceps mouth of Chapare (1557/6441); left bank, 2 km N of mouth of Chapare (1557/6442); left bank, at 1611/6744 mouth of Chapare (1558/6442); right bank, 9 km S of Chapare (1603/6442); right bank, 34 km S 2127/6505 of mouth of Chapare (1617/6442); right bank, 51 Titer, 1991, km S of mouth of Chapare (1627/6444); left bank, 51 km S of mouth of Chapare (1627/ 6445); right 1612/6842 bank, 52 km S of mouth of Chapare (1628/6444); left bank, 52 km S of mouth of Chapare 2 (MARU) (1628/6445); right bank, 54 km S of mouth of Chapare (1629/6444); right bank, 56 km S of mouth of Chapare (1630/6445); right bank, 70 km N.L. S of mouth of Chapare (1636/6448); right bank, SQo Paulo 27 km N of mouth of Chimore (1627/6444); left Quechuan bank, 27 km N of mouth of Chimore (1627/6445); a locality right bank, 2 km N of mouth of Chimor6 (1642/6449); see rio Chapare; Pilleri, Nov. 1976 1317/6438 (1977: 16), at Vita Lei, a farm near Puerto Vil- m E of San larroel (1648/6447). ICHILO PROVINCE; Santa Cruz N.L. F Steinbach, June 1924, June 1945. 149, in Cor- ILLATACO; Cochabamba 1720/6607 Olalla, Dec. 1938, 2600 m (Patterson, 1992). ILLIMANI; La Paz Fnelisjacob- Hartmann, 1978, south slope of Nevado Illi- ontibus sat mani at 1637/6748. imen muight INCACHACA; Cochabamba 1714/6541 imen might ~~~~(USBGN) ol*via. Miller and Boyle, May 1915, 7700 ft (= 2370 APA 1980) m); J. and F B. Steinbach, Sept. 1921, May, June, and Sept. 1927, 2225 m, 2600 m and 3000 m; Anthony (1925: 2), type locality of Thomasomys 7 (USBGN) daphne australis; Carriker, May and June 1937, )ve mouth 2130 to 2440 m, Doutt (1938: 100), type locality 6/645 1); 26 of Monodelphis peruvianus osgoodi (= Monodel- from mouth phis osgoodi); Cabot, Aug. 1988, near Locotal, spelled as Inkachaka (1700/6610), 2250 m. 1448/6418 INDEPENDENCIA; Pando 1126/6734 thers, vari- AMNH-MSB, July and Aug. 1986; 170 m; at Apr. 1992; recent settlement on left bank; also right bank of no Madre de Dios, opposite Independencia. ) (USBGN) INGAVI; Pando 1057/6650 RAP team, June and July 1992, 150 m, barraca 2 (USBGN) (= small settlement) on N bank of nro Orton, in (USBGN) on Provincia Abuna. Retiro; on Santa Cruz INGAVI, see Villa Ingavi, Tarija. (1975: 28); INGENIERO MORA; Santa Cruz 1810/6316 rio Ichilo, Straney, Dec. 1979 and Jan. 1980, Los Tabijos rth; the riv- [a misspelling of Tajibos], 2 km N and 7 km E of d Ichilo are (1809/6312); 10 km E of (1810/6311); Pine and 4amore) or Remsik, August 1980, localities 15 km E of 97 1997 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY (1810/6308), at 1600-1900 ft (= 490-580 m) and 7 km E and 3 km N of (1808/6312), same elevation. INQUISIVI; La Paz 1652/6708 CBF party, Aug. 1991, 2500 m. IPATI; Chuquisaca 2033/6326 de la Barrera, 1955 (Fonseca, 1959: 93); EBD party, Nov. 1984, 81 km from (1954/6346), at Mullupampa or Vaca Guzmain, "hacia Montea- gudo, Pasado Mullupampa, junto al rio Ifiao o Parapeti, Prov. Luis Calvo." IPUNO; department uncertain N.L. A. Fornes, Mar. 1969, specimen at TTU. IPUROPURO, RIO; Beni 1347/6438 (USBGN) Pilleri, Dec. 1976, at Estancia La Havana (1410/6449; Pilleri, 1977b: 21); (Pilleri and Pil- leri, 1984), as Ipurupuru; on some maps as Iru- rupuro. IRUPANA; La Paz 1628/6728 (USBGN) Mercado, July 1986, 2120 m; Mendez, Apr. 1987; Hinojosa, Salazar, and other students, Jan. 1988, and 2 km S of and 3 km S of (1630/6728), 2135 m; in province of Sud Yungas. IRUPLUMO, RIO; Beni CBF party, May 1992, tributary of Secure, at mouth of Iruplumo (1601/6615). ISAMA, RIO; Santa Cruz 1727/6351 (USBGN) Crespo (1959: 10), F Steinbach, May-June 1943 at 420 m, in province of Ichilo (FMNH; also in Buenos Aires, R. M. Wetzel, personal com- mun., as Isamo or Isame). ISCAYACHI; Tarija 2129/6458 (MAPA) Pearson, Dec. 1971, 12 mi NW of (2121/6506), 10,000 ft (= 3075 m); AMNH-MSB, Sept. 1986, 1 km E of, rfo Tomayapo, 3450 m (2129/6457); AMNH-MSB party, July 1991, 1 km E of (2129/6457), at rio Tomayapo, 3416 m; 4.5 km E of (2129/6455) 3750 m. ISIAMAS, see Ixiamas. ISIBOBO; Santa Cruz An estancia; AMNH-MSB party, July 1991, 1 km S and 3 km W of (1931/6336), 930 m. ISIBORO, RIO 1528/6505 (USBGN) d'Orbigny, 1835, as Isibara. ISLA DE LA LUNA: La Paz 1603/6904 CBF party, Apr. 1992. ISLA DEL SOL; La Paz 1601/6910 (MAPA 1980) Libermann, 1986; Bernal, Apr. 1992 ISLA GARGANTUA; Pando 1223/6835 Maps are inconsistent, some show it as in La Paz department. Harada, 1984; Iseki et al. (1985); AMNH-MSB, July 1986, across the north channel of rio Madre de Dios from town of Chive. ITACHEBA; Cochabamba CM 5080. N.L. ITAHUATICUA, CERRO; Santa Cruz 1948/6331 AMNH-MSB party, July 1990. ITAN, see Itau. ITAU; Tarija 2142/6354 (USBGN) Budin, Sept. 1924, at the foot of a range of hills, the Sierra Santa Rosa, near Itau, at about 1600 m; Thomas (1925), type localities of Rhip- idomys collinus (p. 578, = R. leucodactylus col- linus) and of Cavia tschudii pallidior (p. 580, = C. t. sodalis); Cabrera (1961a: 421) as Itan. ITENEZ; see Magdalena. ITENEZ, RIO; Beni 1155/6504 (USBGN) AMNH, Oct. 1965, about 20 km above mouth (1200/6502); Guapore is the Brazilian name for this river. ITONAMA, RIO; Beni 1228/6424 (USBGN) Natterer, 1829 (Pelzeln, 1883: 127); d'Orbigny, Mar. 1832. IXIAMAS, La Paz 1345/6809 (USBGN) Mann, 1922; Eger (1974: 4) cited for Eumops auripendulus, USNM; sometimes spelled Isiamas, or Xiamas; Freese, 1975, as 1340/6810, 221 m; Heltne et al. (1976: appendix I; on foot from 1333/6805 to approximately 35 km N of Ixiamas at 1345/6807; and from 1345/6810 to 5 to 10 km W of Ixiamas); 35 km N of (Hershkovitz, 1984), 1340/6810; Yoneda and Sarmiento, July and Aug. 1990, Aserradero San Francisco (1333/6800); Emmons, June 1990, about 13 km SW of, 400 m, on rio Satariapo (1353/6815); Smith, May 1993, 25 km W of, on road to Alto Madidi (1353/6821). IZOZOG; Santa Cruz 1924/6245 (USBGN) Wavrin, May 1921; Davis (1966: 267); 550 m; at Gamachindy (= Tamachindi, 1927/6232). JACHA KHACA; Oruro 1809/6859 CBF party, Nov. 1992, 4800 m, S of Sajama. JAMACHUMA; Cochabamba MSB expedition, July 1993, 1.3 km W of, 2800 m (1732/6607, GPS). JAPACANI, RIO; see Yapacani. JARIMAYO, RIO; see Totora. JERUSALEN; Beni 1543/6444 (USBGN) AMNH, July 1965, as Jerusaleum; 2 km N of (1542/6444). JESUS DE MACHACA; La Paz AMNH-MSB, Oct. 1986, 3850 m, 12 km by road SW of (1648/6852). JHACHA TOLOKO; La Paz 1621/6802 Miralles and Mercado, May and June 1987, 4740 m, several nearby localities at different el- evations. 98 NO. 231 [Document text truncated for crawler view.]