Biology of Bats of the New World Family Phyllostomatidae. Part I

SPECIAL PUBLICATIONS THE MUSEUM TEXAS TECH UNIVERSITY Biology of Bats of the New World Family Phyllostomatidae. Part I Edited by Robert J. Baker, J. Knox Jones, Jr., and Dilford C. Carter June 1976 f ,,,,.III11"n'IIUllfl"'UIl1l1' ...... ••· l'IUIl'I~'''IIlI'lltl'I''111I1111 11111 11I1 III 1" II 11,,,,,,11 111"'11\"1 \Il/1l1l1lllUl/ll llf II 1I1f1l 1l1l1 I III 1111.1111111111 II/flJ\ 1111111., "I III ~I 1111 /.. II .,\h" .. 1I11ldh 1/ 1111 1 '1111111,1111< It, IIII' •• 'II'I(l"iI,Ulfl lt ~111 No. 10 1"'lllllnUU\l"I"'"\llllll'IIIIIIlIlI"~llmlllUlIlI'lIl1"l TEXAS TECH UNIVERSITY Grover E. Murray, President Glenn E. Barnett, Executive Vice President Regents.-Clint Formby (Chairman), J. Fred Bucy, Jr., Bill E. Collins, John J. Hinchey, A. J. Kemp, Jr., Robert L. Pfluger, Charles G. Scruggs, Judson F. Williams, and Don R. Workman. Academic Publications Policy Committee.-J. Knox Jones, Jr. (Chairman), Dilford C. Carter (Executive Director), C. Leonard Ainsworth, Samuel E. Curl, Harold E. Dregne, Hugh H. Genoways, Ray C. Janeway, William R. Johnson, S. M. Kennedy, Thomas A. Langford, George F. Meenaghan, Harley D. Oberhelman. Robert L. Packard, and Charles W. Sargent. The Museum Special Publications No. 10 218 pp. 25 June 1976 $6.00 Special Publications of The Museum are numbered separately and published on an irregular basis under the auspices of the Dean of the Graduate School and Director of Academic Publications, and in cooperation with the International Center for Arid and Semi-Arid Land Studies. Copies may be obtained on an exchange basis from, or purchased through, the Exchange Librarian, Texas Tech University, Lubbock, Texas 79409. Texas Tech Press, Lubbock, Texas 1976 SPECIAL PUBLICATIONS THE MUSEUM TEXAS TECH UNIVERSITY Biology of Bats of the New World Family Phyllostomatidae. Part I Edited by Robert J. Baker, J. Knox Jones, Jr., and Dillord C. Carter No. 10 June 1976 TEXAS TECH UNIVERSITY Grover E. Murray, President Glenn E. Barnett, Executive Vice President Regents.-Clint Formby (Chairman), J. Fred Bucy, Jr., Bill E. Collins, John J. Hinchey, A. J. Kemp, Jr., Robert L. Pfluger, Charles G. Scruggs, Judson F. Williams, and Don R. Workman. Academic Publications Policy Committee.-J. Knox Jones, Jr. (Chairman), Dilford C. Carter (Executive Director), C. Leonard Ainsworth, Samuel E. Curl, Harold E. Dregne, Hugh H. Genoways, Ray C. Janeway, William R. Johnson, S. M. Kennedy, Thomas A. Langford, George F. Meenaghan, Harley D. Oberhelman, Robert L. Packard, and Charles W. Sargent. The Museum Special Publications No. 10 218 pp. 25 June 1976 $6.00 Special Publications of The Museum are numbered separately and published on an irregular basis under the auspices of the Dean of the Graduate School and Director of Academic Publications, and in cooperation with the International Center for Arid and Semi-Arid Land Studies. Copies may be obtained on an exchange basis from, or purchased through, the Exchange Librarian, Texas Tech University, Lubbock, Texas 79409. Texas Tech Press, Lubbock, Texas 1976 CONTENTS INTRODUCTION 5 ANNOTATED CHECKLIST, WITH KEYS TO SUBFAMILIES AND GENERA. . . . . . . 7 J. Knox Jones, Jr., and Dilford C. Carter, The Museum, Texas Tech University, Lubbock, 79409. ZOOGEOGRAPHY 39 Karl F. Koopman, Department of Mammalogy, The American Museum of Natural History, Central Park West at 79th St., New York, 10024. CHIROPTERAN EVOLUTION 49 James Dale Smith, Department of Biological Sciences, California State University, Fullerton, 92634. COLLECTING TECHNIQUES 71 Merlin D. Tuttle, Vertebrate Division, Milwaukee Public Museum, Milwaukee, Wisconsin 53233. CARE IN CAPTIVITY. ........................................ .. 89 Arthur M. Greenhall, U.S. Fish and Wildlife Service, National Fish and Wildlife Laboratory, National Museum of Natural History, Washington, D.C. 20560. ECONOMICS AND CONSERVATION. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 133 Clyde Jones, U.S. Fish and Wildlife Service, National Fish and Wildlife Laboratory, National Museum of Natural History, Washington, D.C. 20560. BRAIN ANATOMY 147 V. Rick McDaniel, Division of Biological Sciences, Arkansas State University, Jonesboro, 72467. LACTATION AND MILK 201 Robert Jenness and Eugene H. Studier, Department of Biochemistry, University of Minnesota, St. Paul, 55108, and Department of Biology, University of Michigan, Flint, 48503. INTRODUCTION Because of their adaptive diversity and, in many instances, unique morphological attributes, bats of the family Phyllostomatidae have long fascinated biologists. Known only from the New World, most genera of phyllostomatids are strictly limited to tropical environs, but some representatives occur as far north as the southwestern United States and others southward to the northern parts of Argentina and Chile; some species also are distributed in the Bahamas and on the islands of the Greater and Lesser Antilles. With the advent in relatively recent years of improved methods of collecting bats (see Tuttle, this volume), atremendous wealth of information on phyllostomatids has been gathered and it is the purpose of this publication, which ultimately will contain more than 20 individual chapters, to bring these data together in order to assess what now is known about the family and to provide adeparture point for further studies. Owing to the large number of contributions, all of which were solicited by us from persons we felt to be knowledgeable of the subject matter, and the fact that several contributions are necessarily lengthy, the decision was made to group chapters into three parts. Each part will be aseparately numbered Special Publication of The Museum at Texas Tech University. In order to establish aworkable approach by which reference could be made consistently to taxa throughout the series, the annotated checklist by Jones and Carter was circulated to all authors. Each was asked to follow the nomenclature and systematic arrangement in the checklist or, alternatively, to document departures therefrom. This system, it is hoped, will allow readers to relate information from one chapter to the next without the handicap of conflicting names for the same organism. Manuscripts for most contributions first were solicited in 1973. Most manuscripts had been received by the end of 1974. As editorial work progressed, some authors provided up-dated information and all authors of chapters in Part 1had the opportunity to insert limited materials at the time they received galley proofs (in most cases October 1975). Therefore, content is as current as reasonably could be anticipated for aproject of this kind. Organization and editorial style follows that established for the Special Publications of The Museum at Texas Tech University. Otherwise, authors were allowed broad latitude concerning material to be included in their chapters. Accordingly, and for obvious other reasons, some chapters will overlap others in content. Even though some redundacy has reSUlted, we thought it best to have asection on the cited literature with each contribution. Citations to manuscripts in this collected series are carried in text as "this volume," which does not necessarily indicate that the chapter appears in the same part of the series as the one in which it is cited. November 1975 5 Robert J. Baker J. Knox Jones, Jr. Dilford C. Carter ANNOTATED CHECKLIST, WITH KEYS TO SUBFAMILIES AND GENERA J. KNOX JONES, JR., AND DILFORD C. CARTER Leaf-nosed bats of the New World family Phyllostomatidae are primarily limited in distribution to tropical and subtropical regions. A few species reach subtemperate areas. The fanlily has aknown fossil record dating back to Miocene times. Most phyllostomatids are fruit eaters or nectar feeders, but some, primarily species in the subfamily Phyllostonlatinae, are carnivorous or insectivorous, and the unique desmodontines are sanguivorous. The family is unusually diverse from an evolutionary point of view, comprising six subfamilies, 49 currently recognized Recent genera, and 137 Recent nominal species. Twenty-four genera are monotypic. The subfamilies contain the following numbers of genera and species as here recognized: Phyllostomatinae, 11 and 32; Glossophaginae, 13 and 32; Carolliinae, two and seven; Stenoderminae, 17 and 54; Phyllonycterinae, three and seven; and Desmodontinae, three and three. Systematic inquiry in the past decade has tended to reduce the number of recognized genera and species, but the discovery of new taxa continues. Some species are rare in museum collections and their relationships poorly understood. Various new techniques applied in recent years to the study of phyllogenetic relationships have resulted in recognition of new taxonomic alignments--for example, inclusion of the vampire bats as asubfamily of the Phyllostomatidae (Forman et aI., 1968) and exclusion of the Mormoopidae (Smith, 1972), formerly regarded as asubfamily of this group. As standard references for apoint of departure in compilation of this annotated list, we used Hall and Kelson (1959) for North America and Cabrera (1958) for South Anlerica. Avariety of publications has appeared subsequent to these two basic documents in which the distribution and systematics of phyllostomatids are treated. Of these, revisions and reviews are cited at the appropriate places in the accounts. Faunal reports of special interest are noted below. Recourse to the literature we have cited will lead the interested researcher to most of the published sources used in compiling this synopsis. Villa-R. (1967) summarized material on Chiroptera of Mexico. Publications since that time on Chihuahua (Anderson, 1972), Jalisco (Watkins et aI., 1972), Oaxaca (Goodwin, 1969), Sinaloa (Jones et al., 1972), the Yucatan Peninsula (Jones et al., 1973), and Zacatecas (Genoways and Jones, ]968; Matson and Patten, 1975) treat major faunal units as awhole. For Central America, the papers of Jones (1966) on Guatemala, Burt and Stirton (1961) on EI Salvador, Jones et al. (1971 b) and Baker and Jones (1975) on Nicaragua, Starrett and Casebeer (1968) and Gardner et al. (1970) on Costa Rica, and Handley (1966) on Panama are useful, as well as those by Davis et al. (1964) and Carter et al. (1966) 7 8 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY on the region as awhole. Choate and Birney (1968), Koopman (1968) and Jones and Phillips (1970) are useful recent references to bats in the Antillean region. Relatively few major contributions have been published on South America since Cabrera's (op. cit.) compendiunl, those on Trinidad and Tobago (Goodwin and Greenhall, 1961), Surinam (Husson, 1962), Peru (Tuttle, 1970), Colombia (Aellen, 1970; Marinkelle and Cadena, 1972), and Uruguay (Ximenez et ai., 1972) being especially noteworthy. Studies of amore limited scope, such as Hill's (1964) report of asmall collection from Guyana, Brosset's (1965) and Baker's (1974) papers on Ecuador, and the publication by Villa-R. and Cornejo (1969) on northern Argentina also have proved useful (see also the appendix of the contribution on zoogeography by K. F. Koopman in this volume). It is of interest that few reports on the Brazilian fauna have appeared since Cabrera's work, papers by Handley (1967) on the Belem area and by Pine et al. (1970) on acollection fronl Mato Grosso, and Peracchi and de Albuquerque (1971) on the states of Rio de Janeiro and Guanabara being notable exceptions. [See also Gardner's (1976) recent paper on Peru. ] In arecent paper on the mammalian fauna of the Antilles, Varona (1974) incorporated anumber of systematic changes with respect to bats found in that region. For example, he regarded all species of Ardops, Ariteus, and Phyllops as assignable to the subgenus Ariteus of the genus Stenoderma, and placed Monophyllus as asubgenus of Giossophaga. Because Varona presented no evidence supportive of these and other changes, we have not followed his arrangement here. We are indebted to anumber of colleagues, principally Robert J. Baker, Alfred L. Gardner, Hugh H. Genoways, Clyde Jones, Karl F. Koopman, and Don E. Wilson, for scrutinizing an early draft of this manuscript. SUBFAMI LY PHYLLOSTOMATINAE Genus MICRONYCTERIS Gray Micronycteris megalotis (Gray, 1842) Distribution.-Western (Jalisco) and eastern (Tamaulipas) Mexico southeastward through Middle America and much of northern and central South America to Amazonian Peru and Sao Paulo, Brazil; also recorded from Grenada in the Lesser Antilles. Systematics.-Four subspecies currently are recognized: megaiotis (most of South American segment of species distribution); homezi (northwestern Venezuela); mexicana (Mexico south to western Nicaragua and adjacent Costa Rica); microtis (eastern Nicaragua southeastward to Panama and adjacent parts of northwestern South America). Micronycteris schmidtorum Sanborn, 1935 Distribution.- Yucatan Peninsula of Mexico southeastward to northwestern South America. Systematics.-M. schmidtorum is amonotypic species. BIOLOGY OF THE PHYLLOSTOMATIDAE 9 Micronycteris minuta (Gervais, 1855) Distribution.-Nicaragua southeastward to South An}erica (including Trinidad) at least to Brazil and eastern Peru. Systematics.-M. minuta is currently regarded as amonotypic species. Together with megalotis and schmidtorum this species represents the subgenus Micronycteris. Micronycteris hirsuta (Peters, 1869) Distribution.-Honduras southeastward to northern South America (Colon1bia, Venezuela, Guyana, Trinidad, and Peru). Systematics.-M. hirsuta is amonotypic species and the sole representative of the subgenus Xenoctenes. Micronycteris brachyotis (Dobson, 1878) Distribution.-Oaxaca southeastward through Central America to Amazonian Brazil. Systematics.-M. brachyotis is amonotypic species and represents the subgenus Lampronycteris. The specific name platyceps, widely used for this bat for several decades, is asynonym of brachyot is. Micronycteris pusilla Sanborn, 1949 Distribution.-Northern Brazil, eastern Colombia, probably adjacent regions of South America. Systematics.-M. pusilla is amonotypic species and represents the subgenus Neonycteris. Micronycteris nicefori Sanborn, 1949 Distribution.-Nicaragua to northern South America (including Trinidad and south at least to northern Brazil and northern Amazonian Peru). Systematics.-M. nice/ori is amonotypic species and the only representative of the subgenus Trinycteris. Micronycteris sylvestris (Thomas, 1869) Distribution.-Western (Nayarit) and eastern (Veracruz) Mexico southeastward through Central America to Panama and into northern South America at least as far east as Trinidad, northeastern Brazil, and eastern Peru. Systematics.-M. sylvestris is thought to be amonotypic species. Micronycteris behni (Peters, 1865) Distribution.-Known only from central Brazil and Peru. Systematics.- This nominal species is poorly known. Along with M. sylvestris, with which it evidently is closely related, behni constitutes the subgenus Glyphonycteris. Lonchophylla mordax Thomas, 1903 Distribution.-Reported from Ecuador, Bolivia, and Brazil. Systematics.-L. mordax is considered here to be amonotypic species but may include concava as anorthern subspecies. Lonchophylla robusta Miller, 1 91 2 Distribution.-Reported fronl Nicaragua, Costa Rica, Panama, Calombia, Venezuela, and Peru. Systematics.-L. robusta is amonotypic species. Lonchophylla thomasi J. A. Allen, 1904 Distribution.-Known from Panama, Venezuela, Guyana, Surinam, Brazil, Peru, and Bolivia. Systematics.-L. thomasi is amonotypic species. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Genus ANouRA Gray Anoura geoffroyi Gray, 1838 Distribution.-Western (Sinaloa) and eastern (San Luis Potosi) Mexico south to southeastern Brazil and northwestern Argentina. Systematics.- Three nominal subspecies are: geoffroyi (Venezuela, Surinam, Trinidad, Brazil, Argentina, and Bolivia); lasiopyga (Mexico south to northern Colombia); and peruana (Colombian Andes south to Peru). Genus LIONYCTERIS Thomas Lionycteris spurrelli Thomas, 1 91 3 Distribution.-Eastern Panama and northern South America (recorded from Guyana, northern Brazil, Venezuela, Colombia, and Amazonian Peru). Systematics.-L. spurrelli is amonotypic species. Lonchophylla concava Goldman, 1914 Distribution.-Reported from Costa Rica, Panama, Colombia, and Peru. Systematics.-L. concava is recognized provisionally as amonotypic species distinct from mordax, with which it may be conspecific. Handley (1966) considered concava to be anorthern subspecies of mordax, but recent authors have not followed that arrangement. Anoura caudifer (E. Geoffroy St.-Hilaire, 1818) Distribution.-Northern South America south to Peru and Brazil. Systematics.- Two subspecies currently are recognized: caudifer (Colombia east through Venezuela and the Guianas and south in eastern Brazil to Sao Paulo); aequatoris (Ecuador and Peru). 16 BIOLOGY OF THE PHYLLOSTOMATIDAE Anoura cultrata Handley, 1960 Distribution.-Known from Costa Rica, Panama, and Venezuela. Systematics.-A. cultrata is amonotypic species. 17 Anoura werckleae Starrett, 1 969 Distribution.-Known only from Costa Rica. Systematics.-A. werckleae is amonotypic species closely related to A. cultrata. Ano;:;ra brevirostrum Carter, 1 968 Distribution.-Recorded from Colombia (Santander) and eastern Peru. Systematics.-As presently known, A. brevirostrum is amonotypic species. Genus SCLERONYCTERIS Thomas Scleronycteris ega Thomas, 1895 Distribution.- This rare species is known only from Brazil and Venezuela. Systematics.-S. ega is amonotypic species. Genus LICHONYCTERIS Thomas Lichonycteris degener Miller, 1931 Distribution.-Known only from lower Amazon region of Brazil. Systematics.-L. degener is amonotypic species. Lichonycteris obscura Thomas, 1895 Distribution.-Guatemala southeastward to South America at least as far as Surinam and east-central Peru. Systematics.-L. obscura is amonotypic species. Genus HYLONYCTERIS Thomas Hylonycteris underwoodi Thomas, 1903 Distribution.-Western Mexico (north to Jalisco) southeastward to western Panama. Systematics.- Two subspecies (Phillips and Jones, 1971), underwoodi (Veracruz and northern Oaxaca southeastward to Panama) and minor (western Mexico), are recognized. Genus PLATALINA Thomas Platalina genovensium Thomas, 1928 Distribution.- This rare bat is known only from Peru, principally west of the Andes. Systematics.-P. genovensium is amonotypic species. Choeroniscus periosDs Handley, 1966 Distribution.-Known only from Pacific Coast of Colombia. Systematics.-C. periosus is adistinctive, monotypic species. Choeroniscus minor (Peters, 1868) Distribution.-Recorded from Brazil, Colombia, Ecuador, Peru, and Surinam. Systematics.-C. minor is amonotypic species closely related to C. inca and C. intermedius. Choeroniscus intermedius (J. A. Allen and Chapman, 1893) Distribution.- Thought to be restricted to Trinidad, but reported also from Peru by Tuttle (1970). Systematics.-C. intermedius is amonotypic species closely related to C. minor and C. inca. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Genus CHOERONYCTERIS Tschudi Choeronycteris mexicana Tschudi, 1844 Distribution.-Extreme southern parts of California, Arizona, and New Mexico southward to Honduras. Systematics.-C. mexicana is here regarded as amonotypic species. However, Pirlot (1967) described asubspecies (ponsL) from northwestern Venezuela; we are unconvinced by Pirlot's brief description that his two specinlens are referable to the genus Choeronycteris. Choeroniscus inca (Thomas, 1912) Distribution.-Recorded from Guyana, Ecuador, Peru, and Venezuela. Systematics.-C. inca is amonotypic species closely related to the two preceding taxa. Specimens of the genus Choeroniscus are rare in museum collections. Only about two dozen individuals of the minor-intermedius-inca complex have been reported in the literature, and the characteristics of the three species never have been defined in acomparative sense. Clearly, this group is in need of systematic review. It may well b~ that minor, intermedius, and inca represent asingle species. Genus CHOERONISCUS Thomas Choeroniscus godmani (Thomas, 1903) Distribution.-Western Mexico (Sinaloa) southeastward to Colombia and Venezuela. Systematics.-C. godmani is amonotypic species. 18 BIOLOGY OF THE PHYLLOSTOMATIDAE 19 Genus MUSONYCTERIS Schaldach and McLaughlin Musonycteris harrisoni Schaldach and McLaughlin, 1960 Distribution.-Presently known only from the states of Colima and Guerrero in western Mexico. Systematics.-M. harrisoni is amonotypic species. Although some recent authors have regarded Musonycteris as asynonym of Choeronycteris, we follow Phillips (1971) in regarding it as adistinct genus. SUBFAMILY CAROLLIINAE Genus CAROLLIA Gray Carollia castanea H. Allen, 1890 Distribution.-Honduras southeastward through Colombia, Ecuador, and Peru to Bolivia. Systematics.-According to Pine (1972), C. castanea is amonotypic species. Carollia subrufa (Hahn, 1905) Distribution.-Western Mexico (Jalisco) southeastward, mostly in the Pacific versant of Middle America, to Nicaragua. Systematics.-C. subrufa was regarded by Pine (1972) as amonotypic species. Carollia brevicauda (Schinz, 1821) Distribution.-Eastern Mexico (southern San Luis Potosi and adjacent Veracruz) southeastward to northern and western South America (northeastern Brazil, Colombia, Venezuela, Ecuador, Amazonian Peru, and Bolivia). Systematics.-As in the case of the previous two species of Ca ro Ilia, Pine (1972) considered C. brevicauda to be monotypic. Carollia perspicillata (Linnaeus, 1758) Distribution.-Veracruz and Oaxaca southeastward to South America, where the species is widely distributed south to Bolivia, Paraguay, and southern Brazil; also reported from Trinidad, Tobago, and the Antillean island of Grenada (recorded occurrences on Jamaica and Redondo Island, in the northern Lesser Antilles, are questionable). Systematics.- Two subspecies currently are tentatively recognized (Pine, 1972), perspicillata in much of the South American range of the species and azteca in Middle America and adjacent northwestern South America. Pine (op. cit.) also noted that the name C. p. tricolor might apply to specimens from the southern part of the range of the species. Rhinophylla alethina Handley, 1966 Distribution.-Known only from western Colombia. Systematics.-R. alethina is amonotypic species. Rhinophylla fischerae Carter, 1966 Distribution.-Known only from Amazonian parts of Peru and Brazil, and adjacent Colombia and Ecuador. Systenlatics.-R. fischerae is amonotypic species. Genus RHINOPHYLLA Peters Rhinophylla pumilio Peters, 1865 Distribution.-Northern South America in Guyana, Surinam, Venezuela, Colombia, Brazil, and eastern Ecuador and Peru. Systematics.-R. pumilio is amonotypic species. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 20 SUBFAMILY STENODERMINAE Genus STURNIRA Gray The systematics of bats of the genus Sturnira are, for the most part, poorly understood. Several new species have been named in the past decade or so. The list presented here is provisional, pending publication of Luis de la Torre's long awaited revision of the genus. Sturnira thomasi de la Torre and Schwartz, 1966 Distribution.-Known only from Guadeloupe, Lesser Antilles. Systematics.-S. thomasi is tentatively recognized here as avalid, monotypic species because it differs in several ways from other named Antillean populations of Sturnira. It is related to lilium and ultimately may prove best regarded as asubspecies of that species. Sturnira Iilium (E. Geoffroy St.-Hilaire, 1810) Distribution.-Widely distributed from western (Sonora) and eastern (Tamaulipas) Mexico southward through Middle America and throughout most of tropical and subtropical South America to Uruguay, northern Argentina, and possibly Chile; also in southern Lesser Antilles and reported from Jamaica. Systematics.- The following subspecies are tentatively recognized: !ilium (most of South America, including Trinidad); angeli (Dominica in Lesser Antilles); luciae (St. Lucia in Lesser Antilles); paulsoni (St. Vincent in Lesser Antilles); parvidens (Mexico southeastward to Colombia); zygomaticus (Martinique in Lesser Antilles). The taxa angeli and paulsoni, originally named as species, are here listed as subspecies of lilium following Koopman (1968) and Jones and Phillips (1970, 1976). BIOLOGY OF THE PHYLLOSTOMATIDAE 2l Sturllira tildae de la Torre, 1959 Distribution.-Originally named from Trinidad, this species now is known to be widely distributed in northern and central South America, south at least to Mato Grosso, Brazil, and Amazonian Peru. Systematics.-S. tildae is amonotypic species. Sturnira magna de la Torre, 1966 Distribution.-Known only from Amazon drainage in Colombia, Ecuador, and Peru. Systematics.-S. magna is amonotypic species. Sturnira mordax (Goodwin, 1938) Distribution.-Recorded only from Costa Rica. Systematics.-S. mordax is amonotypic species described originally as the sole representative of the genus Sturnirops, possibly valid a~ asubgenus (see Davis et al., 1964). Sturnira bidens (Thomas, 1915) Distribution.-Known only from near Tarrlbo, Colombia, the type locality at Baeza, Ecuador, and cloud forests of eastern Peruvian Andes. Systematics.-S. bidens is amonotypic species and for many years was placed in the genus Corvira. Gardner and O'Neill (1969) reduced Corvira to subgeneric status under Sturnira. Sturnira nana Gardner and O'Neill, 1971 Distribution.-Known only from the type locality, Huanhuachayo, Ayacucho, Peru. Systematics.-S. nana is amonotypic species in the subgenus Corvira (Gardner and O'Neill, 1971). Sturnira aratathomasi Peterson and Tamsitt, 1968 Distribution.-Southwestern Colonlbia and Ecuador west of Andes. Systemat ics.-S. aratathomasi is amonotypic species. Sturnira ludovici Anthony, 1 924 Distribution.-Westem (Sinaloa) and eastern (Tamaulipas) Mexico southeastward through Central America at least to Colombia, Venezuela, Ecuador, and Peru; limits of range poorly understood owing to confusing systematic picture (see below). Systematics.- Two subspecies presently recognized in the literature are ludovici (central Mexico to South America) and occidentalis (western Mexico). However, several named kinds of Sturnira (including hondurensis, bogotensis, and oporophilum) related to ludovici, but not currently recognized in literature, may, in fact, be valid species or subspecies. Vampyrops dorsalis Thomas, 1900 Distribution.-Known from intermediate elevations in Costa Rica, Panama, Colombia, Ecuador, Peru, and questionably from Venezuela. Sturnira erythromos (Tschudi, 1844) Distribution.-Presently recorded only from eastern slope of Andes in Peru, but probably widely distributed in northern South America. Systematics.-S. erythromos is currently regarded as amonotypic species. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Genus VAMPYROPS Peters Vampyrops infuscus Peters, 1881 Distribution.-Colombia south to Peru and Brazil. Systematics.- V. infuscus is amonotypic species. Vampyrops vittatus (Peters, 1860) Distribution.-Known to occur at intermediate elevations (900 to 2600 meters) from Costa Rica south to Peru and east to Venezuela. Systematics.- V. vittatus is amonotypic species. Genus URODERMA Peters Uroderma bilobatum Peters, 1866 Distribution.--Southern Mexico (Veracruz and Oaxaca), Central America, South America as far south as southern Peru and adjacent Bolivia, and southeastern Brazil. Systematics.-Currently recognized subspecies (Davis, 1968; Baker and McDaniel, 1972) include: bilobatum (eastern Bolivia, Brazil, the Guianas, and Venezuela); convexum (Pacific versant of Middle America from Nicaragua southeastward to adjacent South America); davisi (Pacific versant of Middle America from Chiapas to EI Salvador and probably Honduras); molaris (Caribbean versant of Middle America from Veracruz to Costa Rica); thomasi (Ecuador, Peru, northwestern Bolivia); trinitatum (Trinidad). Uroderma magnirostrum Davis, 1968 Distribution.-Chiapas southeastward in Pacific versant of Middle America to Panama, and northern and central South America east of Andes (reported from northern Bolivia, Brazil, Colombia, eastern Peru, eastern Ecuador, and Venezuela). Systematics.-A. magnirostrum is amonotypic species. 22 BIOLOGY OF THE PHYLLOSTOMATIDAE 23 Systematics.- This species is in need of systematic review, but probably is polytypic, with disjunct populations occurring above 900 meters from eastern Panama south to Peru and east into Venezuela. We follow Gardner and Carter (1972) and Carter and Rouk (1973) in our treatment of V. dorsalis. Vampyrops aurarius Handley and Ferris, 1972 Distribut ion.-Known only from the Guiana Highlands of Venezuela. Systematics.- V. aurarius is recognized provisionally as avalid species, but may prove to be asynonym of V. dorsalis. Vampyrops nigellus Gardner and Carter, 1972 Distribution.-Recorded only from Colombia and Peru, but probably occurs also in Ecuador. Systematics.- V. nigellus is considered to be amonotypic species. Vampyrops brachycephalus Rouk and Carter, 1972 Distribution.-Known from ColoITlbia, Venezuela, Guyana, Amazonian Brazil, Ecuador, and Peru. Systematics.- V. brachycephalus is here considered as monotypic and to include V. latus and V. l. saccharus of Handley and Ferris. Vampyrops helleri Peters, 1867 Distribution.--Southern Mexico south through Middle America and northern South America to Peru, Bolivia, and Brazil; also found on Trinidad. Systematics.-As pointed out by Rouk and Carter (] 972), certain differences exist between specimens of helleri from Mexico south through Middle America and those in Peru, but too few specimens are available to interpret these differences. The name incarum Thomas, 1912, would apply to Peruvian specimens and probably other Amazonian material should the differences prove to be of subspecific import. V. zarhinus is considered to be asynonym of V. helleri, and the holotype to have come from Panama. Vampyrops lineatus (E. Geoffroy St.-Hilaire, 1810) Distribution.-Reported from Central (Mato Grosso) and eastern (Bahia) Brazil south to Uruguay, Paraguay, Bolivia, and northern Argentina (Chaco). Although recorded by several authors from localities in western South America, these reports evidently refer to other species of bats. Systematics.- V. lineatus is amonotypic species as presently understood, but appears closely allied to V. recifinus, with which it may be conspecific. Vampyrops recifinus Thomas, 1901 Distribution.-Known from the Brazilian state of Pernambuco and purported to occur in those of Bahia and Sao Paulo. Systematics.-Provisionally recognized as amonotypic species closely related to V. lineatus, from which it may not be distinct even at the subspecific level. Vampyressa bidens (Dobson, 1878) Distribution.-Ecuador, Peru, Colombia, northern Brazil, and Guyana. Vampyressa brocki Peterson, 1968 Distribution.-Presently known only from Guyana and Colombia. Systematics.- V. brocki is amonotypic species. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Genus V AMPYRESSA Thomas Vampyressa pusilla (Wagner, 1843) Distribution.-Southern Mexico (Veracruz), Central America, northern and central South Anlerica south at least to southeastern Brazil and Peru. Systematics.-Goodwin (1963), who reviewed the genus Vampyressa, recognized three subspecies (pusilla, thyone, and venUla). Later, Peterson (1968), in his synopsis of the genus, listed only two (regarding venUla as indistinct from thyone). Handley (1966) did not recognize subspecies in V. pusilla. The species nattereri, named by Goodwin (op. cit.), is here regarded as asynonym of pusilla following Peterson (op. cit.). Genus V AMPYRODES Thomas Vampyrodes caraccioloi (Thomas, 1889) Distribution .. --Southern Mexico (Oaxaca, Veracruz) southeastward through Central America to South America as far south as northern Brazil and Amazonian Peru. Systematics.-Two subspecies are recognized, following Handley (1966): caraccioloi (Trinidad and Tobago, and adjacent regions of northeastern South American mainland); major (Mexico to Peru, ornatus asynonym). Some recent authors have regarded major as aspecies distinct from, but closely related to, caraccioloi. Vampyressa nymphaea Thomas, 1909 Distribution.-Reported from Nicaragua, Costa Rica, Panama, and western Colombia. Systematics.- V. nymphaea is amonotypic species representing, along with V. brocki, the subgenus Metavampyressa. Vampyressa melissa Thomas, 1926 Distribution.-Known only from eastern slope of Andes in Peru. Systematics.- V. melissa is amonotypic species, which together with V. pusilla constitutes the subgenus Vampyressa. 24 BIOLOGY OF THE PHYLLOSTOMATIDAE 25 Systematics.- V. bidens is amonotypic species and the sole representative of the subgenus Vampyriscus, which has been used in the generic sense by some recent authors. Genus CHIRODERMA Peters Chiroderma doriae Thomas, 1891 Distribution.-Eastern Brazil (Minas Gerais). Systematics.-C. doriae is amonotypic species. Chiroderma improvisum Baker and Genoways, 1976 Distribution.-Known only from the Lesser Antillean island of Guadeloupe. Systematics.-C. improvisum is amonotypic species known only from the holotype. Chiroderma villosum Peters, 1860 Distribution.--Southern Mexico (Oaxaca and Veracruz) south to Peru, Bolivia, and Brazil. Systematics.- Two subspecies are recognized, villosum (Trinidad and adjacent Venezuela south to Peru and Brazil) and jesupi (Mexico south through Central America to northern Colombia). Chiroderma salvini Dobson, 1878 Distribution.-Western Mexico (Chihuahua) south to Colombia and Ecuador. Systematics.- Two subspecies currently are recognized, salvini (Puebla, Mexico, south to northern South America) and scopaeum (western Mexico from Chihuahua south to Guerrero). Chiroderma trinitatum Goodwin, 1958 Distribution.-Panama east to Trinidad and south to Peru, Bolivia, and Brazil (Mato Grosso). Systematics.- Two subspecies are recognized, trinitatum (Trinidad and Amazonian South America south to Peru and Brazil) and gorgasi (Darien, Panama, east to Venezuela). Genus ECTOPHy.LLA H. Allen Ectophylla alba H. Allen, 1892 Distribution.-Known only from Nicaragua, Costa Rica, and western Panama. Systematics.-E. alba is amonotypic species. Ectophylla macconnelli Thomas, 1901 Distribution.--South America (reported from Anlazonian Ecuador and Peru, Bolivia, Brazil, Colombia, Venezuela, Guyana, and Trinidad) and reported from Costa Rica and Panama in North America. 4. Two lower premolars ......................................... .. Phyllostomus Three lower premolars (second sometimes crowded to lingual side of toothrow) 5 5. Rostrum as long as braincase Vampyrum Rostrum shorter than braincase ............................................. 6 6. Second lower premolar large, subequal in size to first and third premolars 7 Second lower premolar small to minute, much smaller than first and third premolars .. · 8 Phyllostomatinae 1. One lower incisor 2 Two lower incisors 4 2. Two lower premolars Mimon Three lower premolars (second small to minute) 3 3. Second lower premolar crowded to lingual side of toothrow, first and third lower premolars usually in contact Chrotopterlls Second lower premolar not crowded from toothrow, first and third lower premolars not in contact ........................................................ .. Tonatia 2. Noseleaf rudimentary, without distinct upright process; tail present Phyllonycterinae Noseleaf usually well developed; tail absent if noseleaf rudimentary 3 3. Tongue elongate, with conspicuous bristlelike papillae on anterodorsal surface; first upper premolar usually distinctly separated from canine and rarely in contact with second upper premolar (first upper premolar sometimes in contact with canine in MonophY//llS, but distinctly separated from second upper premolar) Glossophaginae Tongue not elongate, lacking conspicuous bristlelike papillae; first upper premolar in contact with canine and usually with second upper premolar 4 4. Zygomatic arch incomplete Caro//iinae Zygomatic arch complete 5 5. Molars dilambdodont (distinct W-shaped pattern of lophs on occlusal surface) . ·....................................................... .. Phyllostof11atinae Molars lacking dilambdodont pattern Stenoderminae SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 32 7. Auditory bullae large, greatest diameter much exceeding distance between them ..... ·.............................................................. .. Macrotis Auditory bullae small, greatest diameter less than distance between them . ·.......................................................... .. Micronycteris 8. Second lower premolar displaced lingually from toothrow; first and second lower premolars in contact or nearly so 9 Second lower premolar not displaced lingually from toothrow; first and second lower premolars usually not in contact 10 9. Greatest length of skull less than 20 mm Macrophyllum Greatest length of skull more than 20 mm. Trachops 10. Dorsal profile of rostrum strongly convex; deep depression present between orbits ..... ·........................................................... .. Lonchorhina Dorsal profile of rostrum not convex; no depression between orbits Phy//oderma BIOLOGY OF THE PHYLLOSTOMATIDAE 33 Glossophaginae 1. Permanent lower incisors lacking 2 Two pairs of permanent lower incisors, usually weJl developed 8 2. Premolars 3/3 Anoura Premolars 2/3 3 3. Molars 2/2 Lichonycteris Molars 3/3 4 4. Pterygoids highly modified, expanded at base and inflated in appearance; pterygoid wings long and in contact, or nearly so, with auditory bullae 5 Pterygoids normal, not expanded at base or inflated in appearance: pterygoid wings short and not in contact with auditory bullae 7 5. First and second upper incisors separated by distinct gap; upper premolars low, barely exceeding height of molars Choeroniscus First and second upper incisors in contact, or nearly so; upper premolars distinctly higher than molars 6 6. Rostrum distinctly longer than postrostral part of cranium: upper molars essentially equal in size, all with adistinct metastyle Musonycteris Rostrum about equal in length to postrostral part of cranium; third upper molar somewhat smaller than first two and lacking adistinct metastyle Choeronycleris 7. Upper molars lacking mesostyle; lower molars long and narrow; known only from Middle America Hylonycleris Mesostyle present on all upper molars; lower molars only moderately compressed: known only from Brazil and Venezuela Scleronycteris 8. Molars 2/2 Leptonycteris Molars 3/3 9 9. Zygomatic arch complete, first upper incisor not markedly enlarged and spatulate 10 Zygomatic arch incomplete, first upper incisor enlarged and spatulate 11 10. Evident gap between upper premolars and between them and adjacent teeth; tail relatively long and extending beyond posterior border of uropatagium Monophyll us Upper premolars usually in contact and filling space between canine and first molar: tail short and not extending beyond posterior border of uropatagium ..... Glossophaga 11. Rostrum elongate, longer than postrostral part of cranium; postcanine maxillary teeth reduced in size and with evident gaps between them Platalina Rostrum not elongate, no longer than postrostral part of cranium; postcanine maxillary teeth of normal size, last premolar and molars in contact (or nearly so) 12 12. First upper premolar smaller than second and laterally compressed Lonchophylla First upper premolar essentially same size as second, not laterally compressed (triangular in outline) Lionycteris Carolliinae 1. Tail present; upper premolars essentially equal in size Carollia Tail absent; first upper premolar much smaller than second . . .. Rhinophylla ]0. Second upper molar noticeably larger than first; upper premolars separated from each other and from adjacent teeth by evident gaps Ecrophylla (part) Second upper molar equal in size to, or smaller than, first; no gaps between anterior upper cheekteeth 11 II. Incisors 2/l or 2/2; height of first incisor greater than height of first premolar; greatest length of skull less than 22 Vampyressa (part) Incisors 2/2; height of first incisor much less than height of first premolar; greatest length of skull more than 24 Vampyrodes 12. Upper dental arcade expanded laterally to form semicircular arc 13 Upper dental arcade not expanded laterally, U-shaped in occlusal view 14 13. Orbital space wider than long; interorbital constriction less than 5 Amerrida Orbital space longer than wide; interorbital constriction more than 5 . · Sphaeronycreris 14. Palate short, posterior palatal emargination reaching level of first upper molar ..... 15 Palate of medium length or long, posterior border variously emarginate but never to level of toothrow ...................................................... .. 17 Stenoderminae ]. Molars 2/2 2 Molars 2/3 or 3/3 7 2. Upper dental arcade semicircular, rostrum less than half as long as braincase . ·.............................................................. Centurio Upper dental arcade not semicircular, rostrum more than half as long as braincase ... 3 3. Rostrum inflated, nearly cuboid in form , Pygoderma Rostrum not inflated or cuboid in form 4 4. Posterior margin of external nares with marked, lyre-shaped emargination . ·............................................................ .. CII iroderl11a Posterior margin of external nares lacking lyre-shaped emargination 5 5. Second upper mol ar markedly larger than first; upper premolars separated from each other and adjacent teeth by evident gaps . . . . . . . . . . . . . . . . . . . . . . .. Ectophylla (part) Second upper molar essentially equal in size to, or smaller than, first; no gaps between anterior upper cheekteeth 6 6. Posterior margin of external nares more or less straight; second upper molar much smaller than first and differing in form Artihells (part) Posterior margin of external nares broaQly V-shaped; second upper molar resembling first in size and form .. ................................... .. Vampyressa (part) 7. Molars 2/3 8 Molars 3/3 12 8. Palate short, posterior border having deep U-shaped emargination that reaches level of first molar Ariteus Palate long, posterior border having shallow emargination that falls far short of level of toothrow ..............................................................9 9. First upper incisor markedly bifid, less than twice size of second incisor . · Artibells (part) First upper incisor not bifid or only weakly so, more than twice size of second incisor. . ....................................................................... ]0 34 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY BIOLOGY OF THE PHYLLOSTOMATIDAE 35 15. Palatal emargination broadly V-shaped Phyl/ops Palatal emargination deeply U-shaped 16 J 6. Well-developed V-shaped ridge from sagittal crest to anterior margin of orbits, forming deep rostral depression . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Stenoderma V-shaped ridge from sagittal crest to anterior margin of orbits lacking, rostrum normal Ard()ps 17. Upper molars distinctly grooved longitudinally, the first two subquadrate in outline and lacking well-developed cusps; first upper incisor approximately half as high as canine Sturnira Upper molars lacking longitudinal groove, the first two not subquadrate in outline and possessing well-developed cusps; first upper incisor much less than half as high as canine 18 18. First upper incisor less than twice size of second and resembling it in shape: upper incisors in contact and fill ing space between canines 19 First upper incisor more than twice size of second and differing from it in shape; evident gaps present between upper incisors .........................................20 19. First upper incisor deeply bifid; m3, if present, minute and peglike· Artihells (part) First upper incisor not bifid; m3 relatively large and well developed Enchisthenes 20. Crowns of first upper incisors parallel, deeply bifid; lower incisors in contact . ·............................................................. .. Ur()derI11a Crowns of first upper incisors converge distally, not deeply bifid; lower incisors separated by distinct gaps. .......................................... .. Vampyrops Phyllonycterinae 1. Tail not extending beyond edge of uropatagium Brachyphylla Tail extendin_g beyond edge of uropatagium 2 2. Zygomatic arch complete; second and third lower molars distinctly cuspidate . · Erophylla Zygomatic arch incomplete; second and third lower molars not distinctly cuspidate .... ·.......................................................... .. Phyllonycteris Desmodontinae 1. First lower incisors in contact; interfemoral membrane with distinct fringe of moderately long hairs Diphylla First lower incisors not in contact; interfemoral membrane without fringe of hair 2 2. Lower incisors not bifid; wing white from middle of proximal phalanx to tip . ·.............................................................. .. Diaelnlls Lower incisors bifid; wing usually pigmertted to tip (if white-tipped, white does not extend proximally to first phalanx) ................................. .. Desl110d us LITERATURE CITED AELLEN, V. 1970. Catalogue raisonne des chiropteres de la Colombie. Rev. Suisse Zool., 77:1-37. ALLEN, G. M. 1911. Mammals of the West Indies. Bull. Mus. Compo Zool., 54: 173-263. ANDERSON, S. 1972. Mammals of Chihuahua: taxonomy and distribution. Bull. Amer. Mus. Nat. Hist., 148: 149-410. ANDERSON, S., AND C. E. NELSON' 1965. A systematic revision of Macrotlls (Chiroptera). Amer. Mus. Novit., 2212:1-39. BAKER, R. H. 1974. Records of mammals from Ecuador. Pubi. Mus., Michigan State Univ .. BioI. Ser., 5: 129-146. BAKER, R. J., AND J. K. JONES, JR. 1975. Additional records of bats from Nicaragua, with a revised checklist ofChiroptera. Occas. Papers Mus., Texas Tech Univ., 32:1-13. BAKER, R. J., AND V. R. McDANIEL. 1972. A new subspecies of Uroderma bilobatum (Chiroptera: Phyllostomatidae) from Middle America. Occas. Papers Mus., Texas Tech Univ., 7:1-4. BROSSET, A. 1965. Contribution al"etude des chiropteres de l'ouest de I'Ecuador. Mammalia, 29:211-227. BURT, W. H., AND R. A. STiRTON. 1961. The mammals of EI Salvador. Misc. Publ. Mus. Zool., Univ. Michigan, 117:1-69. CABRERA, A. 1958. Catalogo de los mamiferos de America del Sur. Rev. Mus. Argentino Cien. Nat. "Bernardino Rivadavia," Cien. Zool., 4: iv +1-307. CARTER, D. C., R. H. PINE, AND W. B. DAVIS. 1966. Notes on Middle American bats. Southwestern Nat., 11:488-499. CARTER, D. C., AND C. S. ROUK. 1973. The status of recently described species of Vampyrops (Chiroptera: Phyllostomatidae). J. Mamm., 54:975-977. CHOATE, J. R., AND E. C. BIRNEY. 1968. Sub-Recent Insectivora and Chiroptera from Puerto Rico, with the description of a new bat of the genus Stenoderma. J. Mamm., 49:400-412. DAVIS, B. L., AND R. J. BAKER. 1974. Morphometries, evolution, and cytotaxonomy of mainland bats of the genus Macrotus (Chiroptera: Phyllostomatidae). Syst. Zool., 23:26-39. DAVIS, W. B. 1968. Review of the genus Urol!erma (Chiroptera). J. Mamm .• 49:676-698. 1969. A review of the small fruit bats (genus Artihells) of Middle America. Southwestern Nat., 14: 15-29. 1970a. A review of the small fruit bats (genus Artihells) of Middle America. Part II. Southwestern Nat., 14:389-402. 1970h. The large fruit bats (genus Artiheus) of Middle America, with a review of the Artihellsjal1laicensis complex. J. Mamm., 51:105-122. DAVIS, W. B., AND D. C. CARTER. 1962. Review of the genus Leptonycteris (Mammalia: Chiroptera). Proc. BioI. Soc. Washington, 75:193-197. DAVIS W. B., D. C. CARTER, AND R. H. PINE. 1964. Noteworthy records of Mexican and Central American bats. J. Mamm., 45:375-387. FORMAN, G. L., R. J. BAKER, AND J. D. GERBER. 1968. Comments on the systematic status of vampire bats (family Desmodontidae). Syst. Zool., 17:417-425. GARDNER, A. L. 1976. The distributional status of some Peruvian mammals. Occas. Papers Mus., Louisiana State Univ., 48: 1-18. GARDNER, A. L., AND D. C. CARTER. 1972. A review of the Peruvian species of Vampyrops (Chiroptera: Phyl]ostomatidae). J. Mamm., 53:72-82. GARDNER, A. L., AND J. P. O'NEILL. 1969. The taxonomic status of Sturnira bidens (Chiroptera: Phyllostomidae) with notes on its karyotype and life history. Occas. Papers Mus. Zool., Louisiana State Univ., 38: 1-8. 1971. A new species of Stllrnira (Chiroptera: Phy]]ostomidae) from Peru. Occas. Papers Mus. Zool., Louisiana State Univ., 42:1-7. GARDNER, A. L., AND J. L. PATTON. 1972. New species of Philander (MarsupiaJia: Didelphidae) and Mimon (Chiroptera: Phy]]ostomidae) from Peru. Occas. Papers Mus. Zool., Louisiana State Univ., 43:1-12. GARDNER. A. L., R. K. LA VAL, AND D. E. WILSON. 1970. The distributional status of some Costa Rican bats. J. Mamm., 51 :712-729. GENOWAYS. H. H., AND J. K. JONES, JR. 1968. Notes on bats from the Mexican state of Zacatecas. J. Mamm., 49:743-745. 36 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY BIOLOGY OF THE PHYLLOSTOMATIDAE 37 GOODWIN, G. G. 1963. American bats of the genus Vampyressa, with the description of a new species. Amer. Mus. Novit., 2125:]-24. ]969. Mammals from the state of Oaxaca, Mexico, in the American Museum of Natural History. Bull. Amer. Mus. Nat. Hist., 141:]-269. GOODWIN, G. G., AND A. M. GREENHALL. 1961. Areview of the bats of Trinidad and Tobago. Bull. Amer. Mus. Nat. Hist., ]22: 187-30 1. HALL, E. R., AND K. R. KELSON. ]959. The mammals of North America. Ronald Press, New York, ]:xxx+ 1-546+ 79. HANDLEY, C. 0., JR. ]960. Descriptions of new bats from Panama. Proc. U.S. Nat. Mus., ]12:459-479. 1966. Checklist of mammals of Panama. Pp. 753-793, in Ectoparasites of Panama (R. L. Wenzel and V. J. Tipton, eds.), Field Mus. Nat. Hist., Chicago, xii+861 pp. ]967. Bats of the canopy of an Amazonian forest. Atas do simposio sabre a biota Amazonica, 5:2 1 1-2 15. HILL, J. E. ]964. Notes on bats from British Guiana, with the description of anew genus and species of Phyllostomidae. Mammalia, 28:553-572. HUSSON, A. M. ]962. The bats of Suriname. Zoo!. Verhand., 58: 1-282. IRWIN, D. W., AND R. J. BAKER. ]967. Additional records of bats from Arizona and Sinaloa. Southwestern Nat., 12: 195. JONES, J. K., JR. 1966. Bats from Guatemala. Univ. Kansas Publ., Mus. Nat. Hist., 16:439-472. JONES, J. K., JR., AND C. J. PHILLIPS. 1970. Comments on systematics and zoogeography of bats in the Lesser Antilles. Studies on the Fauna of Curacao and other Caribbean Islands, 32:131-145. 1976. Bats of the genus Sturnira in the Lesser Antilles. Occas. Papers Mus., Texas Tech Univ., 40:]-16. JONES, J. K., JR., AND A. SCHWARTZ. 1967. Bredin-Archbold-Smithsonian Biological Survey of Dominica. 6. Synopsis of bats of the Antillean genus Ardops. Proc. U.S. Nat. Mus., 124(3634):1-13. JONES, J. K., JR., H. H. GENOWAYS, AND R. H. BAKER. 1971a. Morphological variation in Stenaderma rufum. J. Mamm., 52:244-247. JONES, J. K., JR., J. D. SMITH, AND R. W. TuRNER. 1971 b. Noteworthy records of bats from Nicaragua, with achecklist of the chiropteran fauna of the country. Occas. Papers Mus. Nat. Hist., Univ. Kansas, 2: 1-35. JONES, J. K., JR., J. R. CHOATE, AND A. CADENA. 1972. Mammals from the Mexican state of Sinaloa. II. Chiroptera. Occas. Papers Mus. Nat. Hist., Univ. Kansas, 6: 1-29. JONES, J. K., JR., J. D. SMITH, AND H. H. GENOWAYS. 1973. Annotated checklist of mammals of the Yucatan Peninsula, Mexico. I. Chiroptera. Occas. Papers Mus., Texas Tech Univ., 13:1-31. KOOPMAN, K. F. 1958. Does pygoderma occur in North America? J. Mamm., 39: 584-585. 1968. Taxonomic and distributional notes on Lesser Antillean bats. Amer. Mus. Novit., 2333:1-13. MARINKELLE, C. J., AND A. CADENA. 1972. Notes on bats new to the fauna of Colombia. Mammalia, 36:50-58. MATSON, J. 0., AND D. R. PATTEN. 1975. Notes on some bats from the state of Zacatecas, Mexico. Contrib. Sci., Los Angeles Co. Mus. Nat. Hist., 263:1-12. OJASTl, J., AND O. J. LINARES. 1971. Adiciones de la fauna de murcieJagos de Venezuela con notas sobre las especies del genero Diclidurus (Chiroptera). Acta BioI. Venezolana, 7:421-441. PARADISO, J. L. 1967. Areview of the wrinkle-faced bats (Centuria senex Gray), with description of a new subspecies. Mammalia, 31 :595-604. PERACTHI, A. L., AND S. T. DE ALBUQUERQU E. 1971. Lista provisoria dos quiropteros dos est ados do Rio de Janeiro e Guanabara, Brasil (Mammalia, Chiroptera). Rev. Brasil. BioI., 31 :405-413. PETERSON, R. L. 1965. A review of the bats of the genus Ametrida, family Phyllostomidae. Contrib. Life Sci., Royal Ontario Mus., 65:1-13. 1968. A new bat of the genus Vampyressa from Guyana, South America. Contrib. Life Sci., Royal Ontario Mus., 73:1-17. PHILLIPS, C. J. 1971. The dentition of glossophaginae bats: development, morphological characteristics, variation, pathology, and evolution. Misc. PubJ. Mus. Nat. Hist., Univ. Kansas, 54:1-138. PHILLIPS, C. J., AND J. K. JONES, JR. 1971. A new subspecies of the long-nosed bat, Hylonycteris lInderl\'oodi, from Mexico. J. Mamm., 52:77-80. PINE, R. H. 1972. The bats of the genus Caro//ia. Tech. Monogr., Texas Agric. Exp. Sta., Texas A&M Univ., 8:1-125. PINE, R. H., I. R. BISHOP, AND R. L. JACKSON. 1970. Preliminary list of mammals of the Xavantina/Cachimbo expedition (central Brazil). Trans. Royal Soc. Trop. Med. Hygiene, 64:668-670. PIRLOT, P. 1967. Nouvelle recolte de chiropteres dans rouest du Venezuela. Mammalia, 31 :260-274. POWER, D. M., AND J. R. TAMSITT. 1973. Variation in Phyllostomus discolor (Chiroptera: Phyllostomatidae). Canadian J. Zool., 5] :461-468. ROUK, C. S., AND D. C. CARTER. 1972. A new species of Vampyrops (Chiroptera: PhyJlostomatidae) from South America. Occas. Papers Mus., Texas Tech Univ., 1: 1-7. SCHWARTZ, A., AND J. K. JONES, JR. 1967. Bredin-Archbold-Smithsonian Biological Survey of Dominica. 7. Review of bats of the endemic Antillean genus Monophy//us. Proc. U.S. Nat. Mus., 124(3635):1-20. SILVA TABOADA, G., AND R. H. PINE. 1969. Morphological and behavioral evidence for the relationship between the bat genus Brachyphylla and the Phyllonycterinae. Biotropica, 1:10-19. SMITH, J. D. 1972. Systematics of the chiropteran family Mormoopidae. Misc. Pub I. Mus. Nat. Hist., Univ. Kansas, 56: 1-132. STARRETT, A., AND R. S. CASEBEER. 1968. Records of bats from Costa Rica. Contrib. Sci., Los Angeles Co. Mus., ]48:1-21. TUTTLE, M. D. 1970. Distribution and zoogeography of Peruvian bats, with comments on natural history. Univ. Kansas Sci. Bull., 49:45-86. VARONA, L. S. 1974. Catalogo de los mamiferos vivientes y extinguidos de las Ant ill as. Acad. Cien. Cuba, viii +1-139 pp. VILLA-R., B. 1967. Los murcielagos de Mexico. Inst. BioI., Univ. Nac. Autonoma Mexico, xvi +1-491 pp. VILLA-R., B., AND M. VILLA CORNEJO. 1969. Algunos murcietagos del norte de Argentina. Misc. Publ. Mus. Nat. Hist., Univ. Kansas, 51 :407-428. WATKINS, L. C., J. K. JONES, JR., AND H. H. GENOWAYS. 1972. Bats of Jalisco, Mexico. Spec. Publ. Mus., Texas Tech Univ., 1:1-44. XIMENEZ, A., A. LANGGUTH, AND R. PRADERI. 1972. Lista sistematica de los mamiferos del Uruguay. An. Mus. Nac. Hist. Nat. Montevideo, ser. 2,7(5):1-49. 38 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY ZOOGEOGRAPHY KARL F. KOOPMAN Of the nine families of bats in the Western hemisphere, three (Emballonuridae, Vespertilionidae, Molossidae) are found also in the Old World. Of the six endemic New World (and chiefly Neotropical) families, the Phyllostomatidae is by far the largest grouping. The Noctilionidae, Furipteridae, and Thyropteridae have only two species each, the Natalidae probably only six species, and the Mormoopidae eight species (see Smith, 1972). The Phyllostomatidae, however, have 136 species as recognized in the classification in this volume. Except for a few areas (West Indies, southeastern Brazil, northern Mexico) even the Pleistocene fossil record of bats for the Neotropical region is poor. To my knowledge the only pre-Pleistocene record of bats in the Neotropical region is of Notonycteris (a phyllostomatid) from the Miocene of Colombia. This shows that the Phyllostomatidae were in South America by at least that time. In the Miocene, South America was still an island continent separated from other continents by ocean barriers. Judging by present diversity in South America, it is likely that the Phyllostomatidae was the first chiropteran family to reach South America during its long period of isolation, and may even have originated as a family on that continent. The other five families more or less confined to the Neotropical region have too few species for anything very definite to be said about their area of origin (see Koopman, 1970). While it is thus probable that South America was the primary center of phyllostomatid evolution, it is clear that both Middle America and the West Indies have been important secondary centers. In the following sections, after ashort discussion of distribution of the subfamilies, the various special regions of South America, Middle America, and the West Indies will be taken up in turn with adiscussion of the phyllostomatids found within each of them It should be emphasized that oceanic straits as well as high mountains and cold temperate lowland areas (such as Patagonia) constitute formidable barriers for the Phyllostomatidae. DISTRIBUTION OF MAJOR PHYLLOSTOMATID GROUPS Of the six subfamilies in to which the Phyllostomatidae are currently divided, all but the Phyllonycterinae are widely distributed in South and Middle America. The phyllonycterines are endemic to the West Indies, which also have phyllostonlatines, glossophagines, and stenodermines. Vampires are known in the fossil record of Cuba, but there are no certain records of carolliines in the West Indies as defined here. Several genera of glossophagines and stenodermines are confined to the Antilles. On the mainland, many genera and even species are distributed over alarge part of the total range of the family. Others, particularly in the Glossophaginae (and to alesser extent the Stenoderminae), have restricted ranges (for example, 39 Platalina). It should also be mentioned.that the Sturnirini (all now included in the genus Sturnira) are strongly concentrated in the northern Andean region with relatively few species far away from it. For futher information on distribution of individual species, the reader is referred to the preceding article in this volume and the appendix of this paper. REGIONAL BAT FAUNAS South America Patagonian subregion.- This is that portion of South America south of the tropical forests. It has never been precisely defined but would certainly include, for our purposes, all of Chile and Uruguay, also all of Argentina except small portions of the northeast and northwest. The high Andes and altiplano of western Bolivia and southern Peru also would be included as well as dryer areas of western Paraguay, southern Bolivia, and extreme southeastern Brazil. Although there are several species of bats that are mostly confined to the Patagonian region, all are vespertilionids. A few species of phyllostomatids, the main range of which lies farther north, do reach the Patagonian subregion to alimited degree, but only six species really penetrate the subregion. Of these, only one, Desmodus rotundus, reaches any distance southward-to central Argentina and even central Chile. While it is possible that this may in part reflect the man-made availability of cattle as food, this is by no means certain. Sturnira /ilium also reaches central Argentina, but an old record from Chile is apparently erroneous. Of the other four species, Chrotopterus auritus, Glossophaga soricina, and Artibeus lituratus range no farther than northern Argentina, whereas Vampyrops lineatus has recently been recorded from Uruguay. Eastern Brazilian highlands and coast.- The dry chaco zone of northern Argentina, western Paraguay, and southeastern Bolivia continues in modified form as the caatinga, abelt of scrub forest (really asavanna), which reaches the Atlantic coast alittle to the west of easternmost Brazil. This isolates the mountain and coastal forests of eastern Brazil, eastern Paraguay, and northeastern Argentina from those of the Amazon basin. As aresult, there are anumber of mammals, particularly primates and rodents, that are confined to this eastenl area. However, out of 36 species of phyllostomatids known from the region, only four (Tonatia brasiliense, Vampyrops recifinus, Chiroderma doriae, and possibly Lonchophylla mordax) are, as at presently recognized, endemic to it. In view of the fact that bats are able to fly across short stretches of unfavorable habitat, however, this is not surprising. There are, on the other hand, some 42 species of phyllostomatids in the Amazon basin that are not known from the eastern Brazilian highlands and coastal area, although some may eventually be found there. Anlazon Basin.-This represents perhaps the reaJ heartland of the Neotropical region. It includes the entire Amazon drainage of Brazil and also includes northeastern Bolivia (with an extension along the eastern face of the Andes into northwestern Argentina), the eastern lowlands of Peru and Ecuador, the Amazon and Orinoco drainages of Colombia and Venezuela, and also the Guianas. Some 40 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY BIOLOGY OF THE PHYLLOSTOMATIDAE 41 74 species of phyllostomatids are known from this area, 14 of which apparently are endemic. However, the list of endemics (Micronycteris behni, M. daviesi, Lonchorhina orinocensis, Tonatia carrikeri, Phyllostofnus latifolius, Scleronycteris ega, Choeroniscus inca, Lichonycteris degener, Rhinophylla fischerae, Vampyrops infuscus, V. aurarius, Vampyressa brocki, V. bidens, and Artibeus concolor) includes several species that are poorly known or of dubious validity. Four other species (Mimon bennetti, Phyllostomus elongatus, Rhinophylla pumilio, and Pygoderma bilabiatum) are shared only with the eastern Brazilian area. More taxonomic work undoubtedly will change considerably the figures both for total number and number of endemics. Eastern slopes of the northern Andes.- The upper forested slopes on the eastern side of the Andes from Colombia to Bolivia, while ecologically continuous with the Amazonian lowlands to the east, are environmentally distinctive to the extent that many lowland species extend only to alimited extent up these slopes, whereas other species are confined to higher elevations. Unfortunately the altitudinal distributions of phyllostomatids are not well known in most of this Andean belt and, at present, it is only about Peru that much can be written. I have chosen to ignore species that do not occur above about 1000 meters but include as endemics all species that are confined to elevations above 500 meters. Using these criteria, some 25 species are known from the upper slopes of the eastern Andes, and three of these (Mimon koepkeae, Sturnira nana, and Vampyressa melissa) are endemic. There would be more endemic species (such as Sturnira erythromos and S. bidens) if higher elevations of the internal Andean valleys of Colombia were included. Northern coast and islands.-I would define this area on the mainland as extending from the northern end of the Cordillera Occidental Uust east of the Gulf or Uraba) east along the coast to the Paria Peninsula in northeastern Venezuela. In Venezuela, it would include only arather narrow coastal strip including the mountains directly to the south, but in Colombia would extend up the river valleys between the cordilleras, but not west of the Cordillera Occidental nor east of the Cordillera Oriental. The boundaries are most difficult to draw in Colombia. Here the higher elevations in the internal Andean valleys are perhaps better placed with eastern slope highlands. The lowlands of the Cauca Valley, on the other hand, are almost equally well placed with lowlands of the Pacific coast. However, in the absence of agreat deal more detailed distributional information, Ihave been unable to draw abetter boundary. Anumber of islands are also included, chiefly Aruba, Cura~ao, Bonaire, Margarita, Trinidad, Tobago, and Grenada. Ihave published previously on the bat faunas of these islands (Koopman, 1958), but anumber of species have been recorded since. Ipreviously (Koopman, 1959) treated Grenada and the Grenadines as part of the West Indies, but, as explained in the Lesser Antillean section, Ibelieve these islands are better placed here. Perhaps the greatest significance of this northern coast and island area is that anumber of the species occurring there have affinities with Central America (and sometimes the West Indies) rather than with the Amazon Basin. Some 64 species are known from this area. Although only Leptonycteris CHIROPTERAN EVOLUTION JAMES DALE SMITH One of the most intriguing problems in vertebrate evolution is the evolution and diversification of the mammalian order Chiroptera. Among vertebrates, bats share with birds and possibly the reptilian pterosaurs (the latter may have been ~dapted to gliding rather than true flight) the unique ability of sustained flight. Whereas the pelvic appendages of birds have remained relatively unchanged for terrestrial locomotion and only the pectoral appendages modified for flight, bats have become totally committed, in an anatomical sense, to astrategy for flight. The difference in the mode of adaptation to flight by birds and bats no doubt reflects two quite different selective regimes involving bipedal and quadrupedal ancestry, respectively. Bats apparently became adapted for an aerial existence in order to exploit an aerial insectivorous food source. On the other hand, birds initially may have developed flight to pursue apredacious mode of life, to escape predators, for dispersal, or acombination of these (Ostrom, 1974). Admittedly, aerial insectivory has been important in the adaptation and diversification of birds, but this particular feeding strategy has not been the central focus in their speciation. Although bats are aremarkably successful group and comprise the second largest mammalian order, they remain one of the least known groups in terms of afossil record. The delicacy of the chiropteran skeleton and the cave and forestdwelling habit of bats have apparently contributed to the paucity of fossils. The antiquity of the Chiroptera is confirmed by Icaronycteris from the early Eocene of Wyoming and France (Russell et al., 1973; includes description and comparisons of Icaronycteris? menui); Palaeochiropteryx and Archaeonycteris from the early Eocene of Austria; Cecilionycteris from the middle Eocene of Germany; Ageina from the early Eocene of France; and the extant genus Hipposideros from the middle Eocene of Europe. By the Oligocene and Miocene, six chiropteran families (Pteropodidae, Rhinolophidae, Emballonuridae, Phyllostomatidae, Vespertilionidae, and Molossidae) are represented in the geologic record. Unfortunately, most of the fossilized remains of bats are extremely fragmentary with the exception of Icaronycteris index, which is beautifully preserved. Martin (1972) compiled asynopsis of late Pliocene and Pleistocene bats (including phyllostomatids) from North America and the Antilles. For the most part, his list includes extant species or extinct species that are clearly related to living taxa. Paula Couto (1938) reported numerous Pleistocene fossil bats from Brazil but these, too, were extant species or related thereto. Two vespertilionids, Miomyotis f/oridanus and Suaptenos whitei, were described by Lawrence (1943) from the early Miocene of Florida, and, more recently, Sutton and Genoways (1974) described Ancenycteris rasmusseni from late Miocene deposits in Gallatin County, Montana. Galbreath (1962) described Oligomyotis casementi from Middle Oligocene deposits in Logan County, Colorado. In addition to these fos49 EVOLUTIONARY IMPETUS OF THE CHIROPTERAN GRADE Because of the meager fossil record for bats, students of chiropteran evolution have been forced to extrapolate the past history of the order based on features of living species. For the most part, anatomical features of the flight mechanism and dental morphology have been utilized in this endeavor, whereas the ecological role, in terms of feeding strategy and niche diversity, mostly has been overlooked. The primary emphasis of chiropteran biology has been descriptive and it is only recently that there has been ashift to synthesizing this information in terms of faunal and ecological complexity. The problem is further aggravated by the paucity of information relating to world ecosystems in the late Mesozoic and early Cenozoic. However, to arrive at areasonable interpretation of the evolution of the Phyllostomatidae as well as that of the Chiroptera as awhole, one must at least be aware that their adaptive radiation progressed as an integral part of developing global ecological complexity. Based on known fossils, the chiropteran grade was fully established in the early Eocene. Reasonable conjecture might project the origin of the group back as far as the early Paleocene or perhaps even into the late Cretaceous. At that point in geologic time, the angiosperm radiation was in its initial stages (Axelrod, sils, which clearly are assignable to the Chiroptera, there are anumber of fragmented insectivore renlains that are suggestive of achiropteran grade, but that can only be categorized as "incertae sedis" (Simpson, 1945; Russell and Sige, 1970). The development of flight by bats (which has involved nearly all major organ systems) was primarily concerned with providing adelivery system for the feeding apparatus. Based on dental morphology of extant species as compared with that of early fossils, it is generally assunled that aerial insectivory was the initial impetus for chiropteran evolution. Subsequent diversification has been associated with the further partitioning and specialization of this generalized feeding strategy into carnivory, piscivory, foliage gleaning, frugivory, nectarivory, and sanguivory. [For convenience, Ihave selected the trophic categories described by Wilson (1973) in this discussion; Irealize that these, in themselves, represent generalized strategies that could be further partitioned. ] It is the goal of this chapter to consider the evolution of the Phyllostomatidae. This family has conlmanded the interest of students of chiropteran evolution because it represents one of the most, if not the most, diverse families in terms of feeding strategy-all categories except piscivory being represented within the context of the family. Furthermore, although ranking third in number of species (136, as compared to 285 for vespertilionids and 150 for pteropodids), the phyllostomatids exceed all other families of bats in nunlber of genera (49, Koopman and Jones, 1970; and Jones and Carter, this volume). Lastly, the adaptive radiation of phyllostomatids apparently has been confined to the tropical regions of the New World. Before proceeding further with adiscussion of the phyllostomatids, Ibelieve it is relevant and important to consider some of the overall aspects of chiropteran evolution. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 50 BIOLOGY OF THE PHYLLOSTOMATIDAE 51 1952, 1970). Also, by the close of the Cretaceous, the anthophilous insect orders Coleoptera, Diptera, and quite probably the Lepidoptera (principal food resources of insectivorous bats) were well established in an evolutionary and ecological sense (Leppik, 1957, 1960; Baker and Hurd, 1968). In addition, eutherian and metatherian mammals were differentiated and both were expanding into numerous terrestrial niches. If tooth morphology is any indication, most of the terrestrial niches, open to mammals, were geared to insectivory or some form of carnivory, although generalized herbivory would certainly have been within the functional potential of these small vertebrates. It is not difficult to visualize the chiropteran ancestry as having taken the form of small arboreal insectivores that nlay have possessed gliding membranes. The argument for an arboreal ancestor as opposed to astrictly terrestrial ancestor seems obvious in light of the fact that all volant mammals normally launch from trees or heights above the ground. The transition from gliding to a movable wing could have progressed by way of elongation .of the digits and interconnected membranes to increase the surface area of the patagium. The patagial arrangement possessed by living Dermoptera may resemble an early stage through which the chiropteran ancestor passed. It seems reasonable to suspect that digital elongation would have reached apoint of diminishing returns in that further progression would have produced an ungainly and clunlsy structure that necessitated movement as awing rather than use as afixed gliding device. [It should be noted here that birds and the recently described giant pterosaur (Lawson, 1975a, 1975b) apparently achieved greatly elongated wings by fusion and elongation of nondigital elements or elongation of asingle digit with broad-based articulations, respectively.] Having successfully traversed this critical point in wing development, apparently by simply utilizing the existing dorsal and ventral thoracic musculature to drive the wing (Vaughan, 1970a, 1970b, 1970c), bats were well on their way to occupying an aerial insectivorous niche. Further refinements of the wing probably related to such parameters as maneuverability and speed. The arguments to support the conjectured insectivory of the chiropteran ancestor may be regarded as open to question. As noted above, this general assumption is based on the morphology of the dental arcade of known fossil bats. The primitive tribosphenic dentition of eutherian-metatherian mammals was modified in the earliest bats to adilambdodont condition with amarked W-shaped ectoloph. This configuration, which allows an increase in the number of shearing facets on the postcanine dentition, is generally associated with insectivory (or carnivory) in living bats as well as in living insectivores such as shrews and moles. The point to be made here is that whereas an arboreal habit seems requisite for the development of chiropteran flight, insectovory, in and of itself, does not. Surely, niche partitioning would have played as integral apart in the various mammalian adaptive radiations in the late Cretaceous or early Paleocene as it has in contemporary ecosystems. With this in mind we might ask the question, why do animals occupy an arboreal niche to begin with? If living forms are any indication, we might con- sider spatial segregation of such parameters as nesting and roosting sites, escape from terrestrial (nonarboreal) predators, and the utilization of such food resources as insects or other small organisms gleaned from branches and foliage, seeds, fruits, flowers, and the like. All of these variables would have been important to the chiropteran ancestor, and, certainly, the utilization of various food items (omnivory) would have been well within the potential of their tribosphenic dentition. The foregoing discussion points out the evolutionary impetus for the chiropteran grade. It is relevant to our consideration of the evolution of the Phyllostomatidae (a group that exploits many feeding strategies) because it establishes the rationale and potential of the arboreal niche with regard to chiropteran evolution. Although insectivory, in the form of foilage gleaning or perhaps aerial insectivory, may have been important to the chiropteran ancestor, certainly opportunistic carnivory or even frugivory (utilization of fruits, seeds, and flowers) would have been possible. The latter is especially important considering the degree of dental specialization and other anatomical departures from the chiropteran norm seen in living pteropodids (apparently exclusive frugivores and the only bats other than phyllostomatids to utilize this food resource). On this basis alone and without agreat deal of conjecture it would be possible to postualte diphyly, or at least an early dichotomy, for bats with respect to the two distinct lineages-Megachiroptera (Pteropodidae) and Microchiroptera (all other living families of bats). PHYLOGENETIC RELATIONSHIPS OF THE CHIROPTERA Judgements as to the phylogeny and evolution of major groups of bats have been mostly intuitive and based on features exhibited by living species. Hill (1974), in his description of the new bat family Craseonycteridae, warned of the inherent difficulties and dangers of this practice. With respect to bats, the problem historically has involved the assessment of the degree of specialization of the flight mechanism and the dental arcade; other systems most certainly could be added in this consideration, but, to date, few have been examined. Whether rightly or wrongly, if we are to proceed with an interpretation of chiroptenin phylogeny based on living taxa and the meager sample of fossil representatives, we must establish an inference as to the nature of the prototype. The recent description of [caronycteris index from the early Eocene of Wyoming (Jepsen, 1966, 1970; Russell and Sige, 1970; Segall, 1971) has provided chiroptologists with atantalizing insight into the prospective chiropteran prototype. [caronycteris along with Paleochiropteryx, Archaeonycteris, Cecilionycreris, and Ageina establishes the nature of aworld-wide paleochiropteran grade in early to middle Eocene times. Cranially, the Paleochiroptera reserrlbled tupaiids in general shape, although the braincase may not have been as inflated. The facial portion of the cranium was long and tapered distally. The premaxillaries of !caronycteris appear to have been fused, although Jepsen (1970) claimed they were not united at the midline. The zygomatic plate was broad and the zygoma were well developed. There is no SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 52 BIOLOGY OF THE PHYLLOSTOMATIDAE 53 indication of apostorbital bar on any of the paleochiropteran fossils. The dental formula was i2-3?/3 c 1/1 P3/3 m3/3. The incisors were well developed and the premolars exhibited areduction in size from posterior to anterior. The molars were typically tribosphenic and dilambdodont, with a W-shaped ectoloph and small hypocone on the upper teeth. The lower molars had well-developed talonids. All molar cusps and associated conlmissures were high and contributed to the complex of shearing facets so characteristic of insectivore dentitions. The wing of paleochiropterans was fully developed, but primitive in most aspects. The globular head of the humerus was the most prominent feature of the proximal portion of this bone and the greater tuberosity was not particularly enlarged and did not extend proximal of the head of the humerus. Contrary to Jepsen (1966), Idoubt that there was a ~'secondary articulation" established between the greater tuberosity and tbe supraglenoid region of the primitive, yet chiropteran, scapula (Smith, 1972). Distally, the humerus was rather primitive, with ashort medial process and generalized trochlear and capitular surfaces. The radius appears to have been typically chiropteran, and, ju~ging from the distal articulation of the humerus, there were no special locking facets present in the elbow region; disjointing stresses developed during flight in this region were probably prevented by muscular and ligamentous binding. The ulna of Icaronycteris was not fused with the proximal portion of the radius and, albeit reduced in size, it appears to have been nearly complete, resernbling the condition found in megachiropterans (Pteropodidae). The first digit (thumb) of Icaronycteris was large and apparently free of the patagium, and the second digit terminated with aclaw, again resembling the Megachiroptera. Although some phalangial elements are missing from all available fossils, the wing apparently was broad, of low aspect, and without any special tip modifications (Findley et al., 1972). The degree of sacral fusion to the pelvic region in paleochiropterans was somewhat less than that which occurs in either the Megachiroptera or Microchiroptera. The head of the femur was large and globular and set between the flangelike greater and lesser trochanters. Paleochiropteryx had awelJ-developed calcar, whereas the remaining paleochiropterans apparently did not, although this absence may be an artifact of preservation. The tail was long and slender in Icaronycferis and Paleochiropteryx. Isuspect that Archaeonycleris also possessed along tail. The paleochiropteran grade, as exemplified by the Eocene fossils, was primitive and generalized in most respects. There is little doubt that these bats were insectivorous, but their capacity for acoustic orientation remains questionable (Segall, 1971). With further refinements in the wing, for speed and maneuverability, and specialization of the cochlear region, for acoustic orientation, the Microchiroptera are easily derived from the paleochiropteran grade as characterized above. Isuspect that such divergence occurred in the Paleocene. The question concerning the relationship of the Megachiroptera and the paleochiropteran grade is not so easily resolved. Meschinelli (1903) described Archaeopteropus transiens from the early Oligocene of Italy. Meschinelli, along with Andersen (1912), Revilliod (1922), and Dal Paiz (1937), regarded Arch- aeopteropus as representing the oldest member of the Megachiroptera. This assignment is based primarily upon similarities of wing morphology, because RevilHod (1922), Russell and Sige (1970), and Slaughter (1970) have pointed out that the dentition of Archaeopteropus, which is badly fragmented, more closely resembles that of the Microchiroptera in appearance. Isuspect that Archaeopteropus represents afurther differentiation of the paleochiropteran grade and perhaps is not at all related to the Megachiroptera. The distinctness and marked departure of megachiropteran dentition from that of the Microchiroptera, as well as from known Tertiary fossils, suggests to me that this group of bats had their origin much earlier in the paleochiropteran grade or perhaps, as noted above, separately from an insectivorous ancestral stock. It is important to point out the rationale for weighting dental morphology, in deference to wing morphology, at this level of interpretation of chiropteran evolution. Since the Oligocene, there appears to have been relatively little variation in the dental morphology of the Microchiroptera; the greatest departure from the basic dilambdodont condition is seen in the phyllonycterine and desmodontine phyllostomatids. The reduction and modification of the dentition in these two subfamilies seems to be the predictable consequence of ahighly specialized feeding strategy and in both cases the dental pattern is traceable to the "primitive" dilambdodont condition (Slaughter, 1970). The degree and consistency of difference of the megachiropteran dentition as well as the apparent total absence in living or fossil taxa of any dentition renlotely similar to the dilambdodont condition, further suggests arather lengthy separation from the paleochiropteran ancestor. [Slaughter's (1970:77, fig. 1) argument for asingle divergence of the Microchiroptera and Megachiroptera from apaleochiropteran prototype based on supposed similarity of the dentitions of Archaeopteropus-- see comment above--and Harpyionycteris is weak.] On the other hand, marked differences in wing morphology might not be expected. The retention of a"primitive" wing by megachiropterans may simply reflect the adequacy of this structure to the habit of these bats; whereas, the wing of microchiropterans has been modified to provide greater maneuverability or speed, thereby facilitating further partitioning of the insectivorous niche. Therefore, with regard to the Microchiroptera, the departure from the paleochiropteran prototype had to do more with refining the wing for maneuverability and speed and with less emphasis on modifying the dental morphology. Among the Microchiroptera, the phyllostomatids illustrate the greatest diversity in dental modification and this appears to have taken place since the Oligocene as will be discussed beyond. The Microchiroptera, no doubt, have their origin within the paleochiropteran grade. Twenty-one genera (age and geographic location in parenthesis) occur as Tertiary fossils and are assignable to the following six living families (Revilliod, 1922; Stirton, 1931; Lawrence, 1943; Simpson, 1945; Galbreath, 1962; Russell and Sige, 1970; SJaughter, 1970; Smith, 1972; Sutton and Genoways, 1974): EmballonuridaeVespertiliavus (Eocene-Oligocene, Europe); RhinoJophidaePalaeophyllophora (Eocene-Oligocene, Europe), Paraphyllophora (EoceneMiocene, Europe), Palaeonycteris (Oligocene, Europe), Rhinolophus (EoceneRecent, Europe), Pseudorhinolophus (Eocene-Oligocene, Europe), Hipposideros 54 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY BIOLOGY OF THE PHYLLOSTOMATIDAE 55 (Eocene-Recent, Europe); Megadernlatidae-Necromantis (Eocene, Europe), Miomegaderma (Miocene, Europe), Provampyrus (Eocene-Oligocene, Africa); Phyllostomatidae-Notonycteris (Miocene, South America); VespertilionidaeStehlinia (Eocene-Oligocene, Europe), Nycterobius (= Revilliodia) (EoceneOligocene, Europe), Samonycteris (Miocene? Pliocene, western Asia), Suaptenos (Miocene, North America), Miomyotis (Miocene, North America), Ancenycteris (Miocene, North America), Oligomyotis (Oligocene, North America), Myotis (Oligocene-Recent, Europe), Simonycteris (Pliocene, North America); MolossidaeTadarida (= Nyctinomus) (Oligocene-Recent, Europe). In addition, there are several genera of such fragmentary remains that familial assignment is not possible at this time. From this, it is evident that most of the major families of Microchiroptera were well established at least by middle Oligocene or Miocene. It seems to be generally agreed upon that the majority of the Microchiroptera originated in the Old World. This thesis is supported by the middle Tertiary occurrence of the Emballonuridae, Rhinolophidae, Megadermatidae, Vespertilionidae, and Molossidae in European deposits. With the exception of the Megadermatidae, which apparently has no living representative in the Palearctic (temperate Eurasia), all of the above-listed families have differentiated in, and presently occur in, all major zoogeographic regions of the Old World. It is noteworthy that the differentiation of these five families, as well as the remaining five in the Old World (Rhinopomatidae, Craseonycteridae, Nycteridae, Myzopodidae, and Mystacinidae), has proceeded along the theme of insectivory or, in several cases, carnivory or piscivory (for example, Megaderma, Macroderma, and, perhaps, Cardioderma). Of these 10 Old World families, only three (Emballonuridae, Vespertilionidae, and Molossidae) also have adaptively radiated in the New World, the emballonurids being confined there to the Neotropics. Although it has not been precisely stated, an emballonuroidlike ancestry generally has been accepted as the base for the Microchiroptera (Fig. 1A). Seemingly, this hypothesis is based more on the apparent antiquity of the group rather than any particular set of primitive characteristics. Certainly, the long and slender free tail of Rhinopoma is reminiscent of acondition noted in several paleochiropterans. However, the trend toward facial shortening by reduction in size and number of premolars, rostral inflation, unfused and unique form of the premax ill aries, trend toward complicated osseous processes on and posterior to the dorsonasal plate, complicated basisphenoidal pits, and specializations of the humerus (both proximally and distally) suggest asomewhat more specialized state for these bats than would be expected for an ancestral group. Afurther corollary of the hypothesis is that the New World noctilionids, mormoopids, and phyllostomatids evolved from an Old World emballonuroid migrant. The Neotropically endemic natalids, thyropterids, and furipterids apparently were derived somewhat later from avespertilionoid stock, which in turn had evolved from the emballonuroid complex. Until the discovery of lcaronycteris index (Jepsen, 1966), which verified the existence of aworld-wide paleochiropteran grade, the foregoing hypothesis would not have been totally untenable. With !caronycteris, more light is shed, albeit dim, on the question of early chiropteran phylogeny, and it now becomes B Pteropodidae FIG. J .-A, cladogram of the generally accepted view of chiropteran phylogeny with the M icrochiroptera derived from a common emballonuroidlike ancestry. B, an alternative proposal for chiropteran evolution with several microchiropteran lineages being derived, independently, from aworld-wide paleochiropteran grade and the Megachiroptera (Pteropodidae) derived either separately from an insectivorous stock or early from the paleochiropteran grade. a, Emballonuroidea; b, Rhinolophoidea; c, Phyllostomatoidea; d, Vespertilionoidea. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Rhinopomatidae }a Craseonycteridae Emballonuridae Megadermatidae }b Nycteridae Rhinolophidae Phyllostomatidae }c Mormoopidae Noctilionidae Molossidae Mystacinidae Natalidae Thyropteridae d Furipteridae Vespertilionidae Myzapodidae A Pteropod idae Rhinopomatidae }a Craseonycteridae Emballonuridae ~b Megadermatidae }b ~ Nycteridae ~o ~ :"t. Rhinolophidae ~ ':.--.;; ~ Phyllostomatidae }c §Mormoopidae \ <a Noctilionidae \0 \ ~ Molossidae \\ Mystacinidae 1\ Natalidae '-......1 \Thyropteridae d \Furipteridae Vespertilionidae Myzapodidae 56 BIOLOGY OF THE PHYLLOSTOMATIDAE 57 possible to suggest an alternative to an exclusively emballonuroid ancestry of the Microchiroptera, at least in the New World. This alternative hypothesis would be to recognize apaleochiropteran grade as asource from which several lineages of Microchiroptera adaptively radiated (Fig. IB). In the consideration of the phylogeny of the New World bats, especially noctilionids, mormoopids, and phyllostomatids, the above proposition is of paramount importance. The discovery of Notonycteris magdalenesis (Savage, 1951, from the late Miocene of Colombia) indicates that phyllostomatids were weJ] established in an evolutionary sense in the late Tertiary. The oldest emballonuroids from the Quercy fauna (Eocene-Oligocene of Europe) were not markedly dissimilar from modern species of that family. The amount of time involved to account for the magnitude of anatomical differentiation of the phyllostomatids from an emballonuroidlike ancestor, amigrant from the Old World, seenlS to shift unduly the whole evolutionary sequence of the Microchiroptera to amuch earlier and as yet undocumented age. It is equally plausible to suggest aseparate and independent radiation of these three unique New World families from the paleochiropteran grade present in the early to middle Tertiary of the New World. In further support of the alternative hypothesis, aconsideration of the overaH anatomical adaptation to aparticular feeding strategy is relevant. With regard to the Microchiroptera, some varied modes of insectivory (or carnivory) apparently were the initial impetus for the differentiation of the various families. Ithink it is important to note that it is only in the New World tropics that alternative microchiropteran feeding strategies such as frugivory, nectarivory, and sanguivory developed. It could be argued that, in the Old World, the various frugivorous, nectarivorous, and other similar niches were already occupied by the Megachiroptera. This, of course, is entirely possible, but it is not consistent in an evolutionary sense to invoke an adaptive potential for an emballonuroidlike ancestor in the New World (where alternative feeding niches apparently were available) and not to consider the same potential as likely in the Old World tropics. Therefore, it seems reasonable to expect that alternative feeding strategies would have been expressed, even in aminor way, in the adaptive radiation of Old World microchiropterans. Yet, the oldest microchiropteran fossils from the Old World now available for interpretation as well as the entire Old World microchiropteran complex, are specialized for insectivory or a relC;lted feeding strategy. The argument for ecological competitive exclusion of the Microchiroptera by the Megachiroptera for frugivorous and nectarivorous feeding niches in the Old World also seems weak. If behavior of living representatives of both groups and the mode by which food resources are partitioned is any indication of past history, then, indeed, the availability of alternative feeding niches to Microchiroptera in the Old World tropics is to be expected. This thesis is proposed on the basis that the nonacousticalJy orienting megachiropterans, facultatively, utilize the food resources (fruits, flowers, nectar, and pollen) during the twilight (crepuscular) period, thereby leaving these resources available during the nocturnal hours for acoustically orienting microchiropterans. Smith (1972), in considering the phylogenetic relationships of the Mormoopidae, suggested that the Phyllostomatidae, Mormoopidae, and Noctilionidae were intimately related to the extent that they might have been derived from common ancestry. This relationship is supported on the basis of similar cranial and postcranial skeletal morphology as well as on similarities of the soft anatomy. The anatomical similarities of the phyllostomatids with these two families is especially pronounced if one considers the subfamily Phyllostomatinae. With regard to the three families, the phyllostomatids are the most divergent, with the mormoopids being somewhat intermediate between noctilionids and phyllostomatids in this regard (Fig. 1). This divergence simply may be areflection of the diversity in feeding strategies utilized by the latter. RELATIONSHIPS WITHIN THE PHYLLOSTOMATIDAE The adaptive radiation of the Phyllostomatidae apparently was aresponse to exploit the various frugivorous niches in the New World tropics. One subfamily, the Desmodontinae, developed the unique feeding strategy of sanguivory. In addition to these specialized strategies, some members of the family, particularly the phyllostomatines, pursue the more typical chiropteran feeding strategy of insectivory and, in several instances, carnivory and omnivory. The family as currently understood is divided into six (perhaps seven) subfamilies: Phyllostomatinae, Glossophaginae, Carolliinae, Phyllonycterinae, Stenoderminae (here including the Sturnirinae), and Desmodontinae. This classification is traditionally based mostly on dental morphology. The phylogenetic relationships within the family are complex and are not well understood at this time. Part of the confusion may be due to similar, but unrelated, adaptations to similar feeding strategies. The phyllostomatines are generally considered to represent the most primitive of phyllostomatid subfamilies. The dental arcade of these bats shows the least amount of modification when compared to other subfamilies. Slaughter (1970) suggested the prototypic dentition of the phyllostomatids would have had aformula of i2/2, c1/1, P2/3, m 3/3, which is found in mo~ living members of the subfamily. The upper molars had well-pronounced and W-shaped ectolophs, acharacteristic of insectivorous, piscivorous, and carnivorous bats. As in noctilionids and mormoopids, the Wshaped ectoloph on the upper molars of phyllostomatines extends at least half the width of the tooth and the protocone-hypoconal shelf is not particularly broadened. In addition, P3 and p3 probably were not much reduced in size in the prototyptic dentition. In comparing Notonycteris magdalenensis with other phyllostomatines, Savage (1951) recognized several groups of genera within the subfamily. He distinguished these primarily on the configuration of the cusps on the cheekteeth. In one group, he included Notonycteris, Chrotopterus, and Vampyrum, which he found to differ strikingly from Phyllostomus. Walton and Walton (1968) and Smith (1972) also noted severa] groups of genera within the phyllostomatine bats based on various postcranial characteristics. In their Macrotus type, Walton and Walton (1968) included Savage's (1951) Notonycteris-ChrotopterusSPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 58 BIOLOGY OF THE PHYLLOSTOMATIDAE 65 Chiroderma salvini Dobson VENEZUELA (Aragua): Cueva de Quebrada Honda (SR), Linares, 1968. Artibeus jamaicensis Leach MEXICO (Yucauin): Lara's Cave (SR), Has Cave (SR), Loltun Cave (SR), Coyok Cave (SR), Spikul Cave (SR), Chacaljas Cave (SR), Hatt et ai., 1953. CUBA: Daiquiri Cave (LP), Anthony, 1919; Camaguey Cave (LP), Koopman and RUibal, 1955. HAITI: Cave near St. Michel (SR), Diquini (SR), Miller, 1929. PUERTO RIco: Cueva Monte Grande (LP), Anthony, 1918; Cueva de Clara (LP), Cueva del Perro (LP), Choate and Birney, 1968; Reynolds et al., 1953. VENEZUELA (Aragua): Cueva de Quebrada Honda (SR), Linares, 1968. Artibeus cinereus Miller MEXICO (Yucatan): Coyok Cave (SR), Hatt et ai., 1953. VENEZUELA (Aragua): Cueva de Quebrada Honda (SR), Linares, 1968. Enchisthenes harti (Thomas) VENEZUELA (Aragua): Cueva de Quebrada Honda (SR), Linares, 1968. Sphaeronycteris toxophyllum Peters VENEZUELA (Aragua): Cueva de Quebrada Honda (SR), Linares, 1968. Phyllopsfalcatus (Gray) CUBA: Daiquiri Cave (LP), Anthony, 1919; Camaguey Cave (LP), Koopman and Ruibal, 1955. Phyllops haitiensis (J. A. Allen) HAl TI: Cave near St. Michel (SR), Cave near Atalaye (SR), Diquini (SR), Miller, 1929; Cave near EnCafe (SR), Miller, 1930. tPhyllops vetus Anthony CUBA: Daiquiri Cave (LP), Anthony, 1919. Ariteus flavescens (Gray) JAMAICA: Dairy Cave (LP), Williams, 1952. tStenoderma rufum anthonyi Choate and Birney PUERTO RICO: Cueva de Clara (LP), Cueva del Perro (LP), Choate and Birney, 1968. Subfamily Desmodontinae Desmodus rOtltndus (Wagner) UNITED STATES (Texas): Cave in Terlingua district (LP), Cockerell, 1930. MEXICO (Yucatan): Loltun Cave (SR), Hatt et al., 1953. CUBA: Cueva Lamas (LP), Koopman, 1958. VENEZUELA (Aragua): Cueva de Quebrada Honda (SR), Linares, 1968. t Desmodus rotundus puntajudensis Woloszyn and Mayo CUBA: Centenario de Lenin, Lorna del Medio, Punta Judas, NE coast of Las Villas (SR), Woloszyn and Mayo, 1974 (these authors were uncertain about assigning Koopman's, 1958, specimen from Cueva Lamas to this taxon). tDesmodus stocki Jones UNITED STATES (California): Potter Creek Cave (LP), Hutchinson, 1967; (Florida): Reddick (LP), Gut, 1959, and Olsen, 1960; Arredondo (LP), Martin, 1972. MEXICO (Nuevo Leon): San Josecito Cave (LP), Jones, 1958; (Mexico): Tlapacoya (LP), Alvarez, 1972. tDesmodus sp. VENEZUELA (Monagas): Cueva del Guacharo (LP), (Clayton Ray and Omar Linares, personal communication) [Paula Couto (1938) reported Schizostoma (= Micronycteris), Lophostoma (= Tonatia), Vampyrus (= Chrotopterus, Tonatia, or Vampyrum),Phyllostoma ( = Phyllostomus), Tylostoma (= Mimon crenulatum), Carollia, Lonchoglossa ( = Anoura), Glossophaga, Chiroderma, Sturnira, Vampyrops, Artibeus, Desmodus, "etc.," from Pleistocene cave deposits of Brazil. Ihave not included these in the above listing because he did not designate species and their determination would be difficult from the generic list that he presented. No locality information other than Brazil was given. In addition, there is avague reference to phyllostomatid genera cited by Peter Lund and Herluf Winge from Brazilian Pleistocene caves.] SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY LITERATURE CITED ALLEN, G. M. 1939. Bats. Harvard Univ. Press, Cambridge, x+369 pp. ALLEN, H. 1898. On the Glossophaginae. Trans. Amer. Philos. Soc., 237-266. ALVAREZ, T. ]972. Nuevo registro para el vampiro del Pleistoceno. Desmodus stoki [sic] de Tlapacoya, Mexico. An. Esc. Nac. Cienc. BioI., Mexico, 19: 163-165. ANDERSEN. K. ]912. Catalogue of the Chiroptera in the collection of the British Museum. British Mus. (Nat. HisL), London, 2nd ed .• ]:ci +1-854 +30. ANTHONY, H. E. ]917. Two new fossil bats from Puerto Rico. Bull. Amer. Mus. Nat. Hist., 37:565-568. 1918. The indigenous land mammals of Puerto Rico, living and extinct. Mem. Amer. Mus. Nat. Hist.. 2:33]-435. 1919. Mammals collected in eastern Cuba in 1917. with descriptions of two new races. Bull. Amer. Mus. Nat. Hist., 41 :625-643. ?Phyllostomatidae incertae sedis Univ. California Mus. Paleo. No. 54572, ant. two-thirds of right p2 from UCMP loco V-5847 in Big Cat Quarry, Cuyanla Valley Badlands, Santa Barbara Co., California. Age.--early CI arendonian. James (1963) noted that this tooth was in the size range of Phyllostomus, Chrotopterus, Vampyrum, and Notonycteris, but admitted that generic assignment was impossible and familial assignment was conjectural. UCMP No. 82144, left lower canine; UCMP No. 80324, edentulous and incomplete right dentary, from UCMP loco V-6761 Branch Canyon Formation, Santa Barbara Co., California. Age.-Hemingfordian. Hutchison and Lindsay (1974) noted that these specimens resembled Pteronotus (family Mormoopidae), Lonchorhina, and Macrophyllum, but deferred generic assignment. 66 AXELROD, D. I. 1952. Atheory of angiosperm evolution. Evolution, 6:29-60. 1970. Mesozoic paleogeography and early angiosperm history. Bot. Rev., 36:277-319. BAKER, R. J. 1967. Karyotypes of bats of the family phy]]ostomatidae and their taxonomic implications. Southwestern Nat., 12:407-428. 1970. The role of karyotypes in phylogenetic studies of bats. Pp. 303-312, in About bats (B. H. Slaughter and D. W. 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APleistocene vampire bat from Florida. J. Mamm., 40:534-538. HATT, R. T., H. I. FISHER, D. A. LANGEBARTEL, AND G. W. BRAINERD. 1953. Faunal and archeological researches in Yucatan caves. Cranbrook Inst. Sci. BuH., 33:1-119. HILL, J. E. 1974. Anew family, genus and species of bat (Mammalia: Chiroptera) from Thailand. Bull. British Mus. (Nat. Hist.), Zoology, 27:304-336. HUTCHINSON, J. H. 1967. APleistocene vampire bat (Desmodus stocki) from Potter Creek Cave, Shasta County, California. Paliobios, 3: 1-16. HUTCHINSON, J. H., AND E. H. LINDSAY. 1974. The Hemingford mammal fauna of the Vedder locality, Branch Canyon Formation, Santa Barbara County, California. Part 1: lnsectivora, Chiroptera, Lagomorpha, and Rodenta (Sciuridae). Paleobios, 15:1-19. JAMES, G. T. 1963. Paleontology and nonmarine stratigraphy of the Cuyama VaHey Badlands, California. Part I. Geology, faunal interpretations and systematic descriptions of Chiroptera, Insectivora, and Rodentia. Univ. California Publ. Geol., 45:1-154. JEPSEN, G. L. 1966. Early Eocene bat from Wyoming. Science, 154:1333-1339. BIOLOGY OF THE PHYLLOSTOMATIDAE 67 1970. Bat origins and evolution. Pp. 1-64, in Biology of bats (W. A. Wimsatt, ed.), Academic Press, New York. 1: xii+ 406 pp. JONES, J. K., JR. 1958. Pleistocene bats from San Josecito Cave, Nuevo Leon, Mexico. Univ. Kansas Publ., Mus. Nat. Hist., 9:389-396. KOOPMAN, K. F. 1951. Fossil bats from the Bahamas. J. Mamm., 32:229. 1958. Afossil vampire bat from Cuba. Breviora, 90:1-4. KOOPMAN, K. F., AND E. E. WILLIAMS. 1951. Fossil Chiroptera collected by H. E. Anthony in Jamaica, 1919-1920. Amer. Mus. Novit., 1519:1-29. KOOPMAN, K. F., AND R. RUIBAL. 1955. Cave-fossil vertebrates from Camaguay, Cuba. Breviora, 46: 1-8. KOOPMAN, K. F., M. K. HECHT, AND E. LEDECKy-JANECEK. 1957. Notes on the mammals of the Bahamas with special references to the bats. J. Mamm., 38:164-174. KOOPMAN, K. F., AND J. K. JONES, JR. 1970. Classification of bats. Pp. 22-28, in About bats (B. H. Slaughter and D. W. Walton, eds.), Southern Methodist Univ. Press, Dallas, vii+339 pp. LAWSON, D. A. 1975a. Pterosaur from the latest Cretaceous of west Texas: discovery of the largest flying creature. Science, 187:947-948. 1975h. Could pterosaurs fly? Science, 188:676-678. LAWRENCE, B. 1943. Miocene bat remains from Florida, with notes on the generic characters of the humerus of bats. J. Mamm., 24:356-369. LEPPIK, E. E. 1957. Evolutionary relationship between entomophilous plants and anthophilous insects. Evolution, 11 :466-481. 1960. Early evolution of flower types. Lloydia, 23:72-92. LINARES, O. J. 1968. Quiropteros subfosiles encontrados en las cuevas venezolanas, Parte 1. Boi. Soc. Venezolana Espeleologia, 1: 119-145. MACHADO-ALLISON, C. E. 1967. The systematic position of the bats Desmodus and Chilonycteris, based on host-parasite reI ationships (Mammalia: Chiroptera). Proc. BioI. Soc. Washington, 80:223-226. MARTIN, R. A. 1972. Synopsis of late Pliocene and Pleistocene bats of North America and the Antilles. Amer. Midland Nat., 87:326-335. MESHINELLI, L. 1903. Un nuovo chiroptero fossile (Archaeopterus transiens Mesch.) deBe Iiquiti di Monteviale. Atti. reale Istit. veneto Sci. Lett. Arti., 62(2): 1329-1344. MILLER, G. S., JR. 1907. The families and genera of bats. Bull. U.S. Nat. Mus., 57: xvii +1-282. 1929. A second collection of mammals from caves near St. Michel, Haiti. Smithsonian Misc. CoiL, 81:1-30. 1930. Three small collections of mammals from Hispaniola. Smithsonian Misc. CoIl., 82:1-10. OLSEN, S. J. 1960. Additional remains of Florida's Pleistocene vampire. J. Mamm., 41 :458-462. OSTROM, J. H. 1974. Archaeopteryx and the origin of flight. Quart. Rev. BioI., 49:2747. PAULA COUTO, C. DE. 1938. Os chiropteIos fosseis do Brasil. Rev. Acad. Colombiana Cien., 2:464-465. PHILLIPS, C. J. 1971. The dentition of glossophagine bats: development, morphological characteristics, variation, pathology, and evolution. Misc. Pub!. Mus. Nat. Hist., Univ. Kansas. 54:1-138. REVILLIOD, P. 1922. Contribution a l'etude des chiropteres des terrains Tertiaires 2. Mem. Soc. Pal. Suisse, 45:] 33-195. REYNOLDS, T. E., K. F. KOOPMAN, AND E. E. WILLIAMS. 1953. A cave faunule from western Puerto Rico with discussion of the genus lsolohodon. Breviora, 12: 1-8. RUSSELL, D. E., AND B. SIG·E. 1970. Revision des chiropteres lutetien de Messel (Hesse, AJlemagne). Palaeovertebrata, MontpeBier, 3:83-182. RUSSELL, D. E., P. LOUIS, AND D. E. SAVAGE. 1973. Chiroptera and Dermoptera of the French early Eocene. Univ. California Publ. GeoJ., 95: I-57. 68 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY BIOLOGY OF THE PHYLLOSTOMATIDAE 69 SAVAGE, D. E. 1951. A Miocene phyllostomatid bat from Colombia, South America. Univ. California Publ. Geol., 28:357-366. SEGALL, W. 1971. Auditory region of bats including lc{/ronycteris index. Fieldiana Zool., 58:]03-108. SILVA-TABOADA, G. 1974. Fossil Chiroptera from cave deposits in Central Cuba, with descriptions of two new species (genera Pteronotlls and Mormoops) and the first West Indian record of Mormoops megalophyllll. Acta Zool. Cracoviensia. 19:33-73. SILVA-ToBOADA, G., AND R. H. PINE. 1969. Morphological and behavioral evidence for the relationship between the bat genus Brachyphyl/{/ and the PhylJonycterinae. Biotropica, 1: 10-19. SIMPSON, G. G. ]945. The principles of classification and a classification of mammals. Bull. Amer. Mus. Nat. Hist., 85:1-350. SLAUGHTER, B. H. 1970. Evolutionary trends of chiropteran dentitions. Pp. 51-83, ill About bats (B. H. Slaughter and D. W. Walton. eds.). Southern Methodist Univ. Press, Dallas, vii +339 pp. SMITH, J. D. 1972. Systematics of the chiropteran family Mormoopidae. Misc. Publ. Mus. Nat. Hist., Univ. Kansas, 56: 1-132. ST ARRETT, A., AND R. S. CASEBEER. 1968. Records of bats from Costa Rica. Contrib. Sci., Los Angeles Co. Mus., ]48: ] -2 l. STIRTON, R. A. 1931. Anew genus of the family Vespertilionidae from San Pedro Pliocene of Arizona. Univ. California Publ., Bull. Dept. Geosci., 20:27-30. SUTTON, J. F., AND H. H. GENOWAYS. 1974. Anew Vespertilionine bat from the Barstovian deposits of Montana. Occas. Papers Mus., Texas Tech. Univ., 20:]-8. VAUGHAN, T. A. ]970a. The skeletal system. Pp. 97-138. in Biology of bats (W. A. Wimsatt, ed.), Academic Press, New York, ]:xii+406. 1970h. The muscular system. Pp. 139-194. in Biology of bats (W. A. Wimsatt. ed.), Academic Press, New York, l:xii +406. 1970c. Flight patterns and aerodynamics. Pp. 195-216, in Biology of bats (W. A. Wimsatt, ed.), Academic Press, New York, l:xii +406. WALTON, D. W., AND G. W. WALTON. 1968. Comparative osteology of the pelvic and pectoral girdles of the Phyllostomatidae (Chiroptera; Mammalia), J. Grad. Res. Center, Southern Methodist Univ .. 37:]-35. WENZEL, R. L., V. J. TIPTON, AND A. KIEWLICZ. 1966. The streblid batflies of Panama (Diptera: Calypterae: Streblidae). Pp. 405-575, in Ectoparasites of Panama (R. L. Wenzel and V. J. Tipton, eds.), Field Mus. Nat. Hist., Chicago, xii+ 861 pp. WILLIAMS, E. E. 1952. Additional notes on fossil and subfossil bats from Jamaica. J. Mamm., 33: 171-] 79. WILSON, D. E. 1973. Bat faunas: a trophic comparison. Syst. Zool., 22: 14-29. WOLOSZYN, B. W., AND N. A. MAYO. 1974. Postglacial remains of a vampire bat (Chiroptera: Desmodlls) from Cuba. Acta Zool. Cracoviensia. 19:253-265. COLLECTING TECHNIQUES MERLIN D. TUTTLE Phyllostomatids exhibit an unusual diversity in roosting and foraging behavior. Hence, while several collecting methods (such as mist-netting and trapping) are exceptionally versatile, even these fall far short of capturing all species under alJ circumstances. Because each technique results in selective capture of certain species while practically excluding others, faunal analyses should be based upon the widest possible variety .of collecting methods. By contrast, ecological and behavioral studies of one or a few species should employ only those techniques best adapted to obtaining desired data while, at the same time, minimizing disturbance to the population. This chapter provides information on means of locating phyllostomatids and their roost sites, and discusses those collecting techniques that have proven to be effective. For asummary of other methods not mentioned here see Greenhall and Paradiso (1968:8-15). MATERIALS Many materials for collecting bats may be placed in one of two categoriesthose employed at roosts and those used along flyways or at places where bats forage. However, other materials are useful in both kinds of situations and will be discussed first; equipment used primarily for specialized collecting will be dealt with later. An electric headlight is essential for most types of collecting. The best light Ihave been able to find is the Justrite Headlight (obtainable from Justrite Manufacturing Co., 2061 NSouthport Avenue, Chicago, Illinois 60614). This light has an adjustable beam and space to store spare bulbs behind the headlight reflector. Agood power source is the alkaline Eveready battery, no. 520. Acanvas battery holder can be carried on an army pistol belt. Several kinds of holding cages have been described (Greenhall and Paradiso, 1968:20-21), but Ihave found it more convenient to hold captured bats in bags made of nylon army mosquito netting with tie strings near the top. Muslin bags may also be used. However, muslin is bulky and much heavier to carry, rots easily in tropical environments, and bats cannot be seen without opening the top of the bag. Mist nets are the most versatile devices for collecting bats. They can be purchased from the following suppliers: Bleitz Wildlife Foundation, 5334 Hollywood Boulevard, Hollywood, California 90027; Eastern Bird Banding Association, Biology Department, Indiana University of Pennsylvania, Indiana, Pennsylvania 15701; Northeastern Bird Banding Association, 37 Old Brook Road, West Hartford, Connecticut 06117; and W. B. Davis, P. O. Box 3522, Bryan, Texas 77801. They are available in widths of 6, 9, 12, and 18 meters by 1.2 or 71 HANDLING OF NETS AND TRAPS Mist-netting and trapping are the two most effective methods known for collecting avariety of bats. Much of their success, however, is dependent upon knowledge of how and where to use them. Operational details are dealt with here, whereas factors influencing where and when to use nets will be discussed later. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 72 2.4 meters high, and are constructed of 30 to 70-denier thread in 25 to 36 millimeters or longer mesh. Various colors are available, but black seems to be most efficient for night use. The most versatile nets for catching bats have four shelves, are 6 or 12 meters wide and 2.4 meters high, and are constructed of 50 or 70-denier thread with 36-millimeter mesh (Handley, 1968:15-16). One should always sample one or afew nets from agiven supplier before ordering more. It is wise to check each one for the following possible defects: 1) improperly threaded shelf strings, which cause uneven distribution of netting; 2) shelf strings that have become untied and must be rethreaded; 3) shelf strings of unequal length; 4) inadequate amount of netting between shelf strings; and 5) netting that is not soft and pliable. Also, the loops at the ends of each shelf string should be made of cotton because nylon loops tend to become untied easily. Arecently developed double-framed trap has been used under awide range of conditions and has proven successful in capturing many temperate and tropical bats (Tuttle, 1974a). The bats collide with fine vertical wires and fall unharmed into alarge canvas receptacle from which they are unable to escape. This trap is easily carried by one man, can be assembled or broken down in 45 minutes, and is particularly useful in studies that require rapid handling of large samples. At present these traps are not produced commercially, but acomplete description with specifications for construction has been published (Tuttle, 1974c). Several earlier and less versatile traps also have been described (Constantine, 1958, 1962, 1969). Other important items are amachete and gloves. In tropical areas, amachete with a 15 to 18-inch blade (and abelt sheath) is an efficient tool for clearing netting and trapping sites, for preparing poles for nets, and for chopping into roosts in small holes. Apair of leather gloves should be used when catching roosting bats by hand and for handling captured specimens. Mist Nets Preparatory to setting amist net, appropriate poles must be obtained. In tropical rain forests, one rarely experiences difficulty in finding adequate saplings that can be cut into lengths of two and ahalf to three meters. These should be straight, stiff, and about five centimeters in diameter at the base. All twigs should be removed. Amachete may be used to sharpen the larger end of each pole. Work in relatively dry areas may require carrying asupply of poles; telescoping aluminum poles and adjustable metal pole clamps are convenient. When asuitable netting site has been found, amachete is used to remove sufficient vegetation and debris so that the net will not become tangled. At the same time, it is important not to remove too much vegetation, thus leaving the BIOLOGY OF THE PHYLLOSTOMATIDAE 73 net exposed and conspicuous. The goal is to allow barely enough space for the collector to pass freely along both sides without endangering the net, but no more. At windy sites, extra space should be allowed for billowing of the net. In most tropical areas, poles can be driven into the ground by repeated jabbing and twisting. When several have been cut, it is best to select the heaviest pole with the hardest, sharpest point to make holes for the rest. Depending on the rigidity of the poles, holes should be angled so that erected poles lean slightly outward. This allows for bending from the inward pull of the net. When apole is limber or inadequately anchored in the ground, guy lines can be used to hold it in place. In rocky river beds, it may be necessary to provide additional support by piling large rocks against the bases of the poles. Before the first pole is finally secured, the loops of one end of the net are placed in proper sequence over the upper end. When anew net is first unpacked, the main loops, which fit over the poles, usually will be gathered in the center in two bunches in the correct order. It is important that these be separated carefully to avoid initial confusion. Next, an outside loop is found and the rest of the loops on that end are arranged in appropriate sequence over afinger, from which all are slipped over the end of the pole. The pole is firmly secured, and the net is unfolded until it is pulled tight to mark the spot where the second pole is to be placed. At this point, one person may hold the net off the ground while another prepares the hole. If alone, refold the net before making the hole, unfolding it only when ready to secure the second pole. Before slipping the loops over the pole, it is necessary to check the top shelf string to be sure that the net is not twisted, with the loops in reverse sequence. When possible, nets should be set and adjusted before dark, but they should not be opened on the poles until it is time to use them. In the evening, when collectors tend their nets, they should be equipped with aheadlight, agood battery, spare bulbs, holding bags, apair of gloves, insect repellent, and spare string to repair broken shelf strings and to guy leaning poles. In some areas they may also wish to carry agun; poisonous snakes, caiman, crocodiles, and large cats may be attracted to the squeals of trapped bats. Nets should not be left unattended for long, and must be guarded almost constantly when set over trails and around villages where domestic animals and humans are likely to pass. Even in the absence of other problems, bats themselves will soon destroy an unattended net. Such large species as Chrotopterus auritus, Phyllostomus hastatus, or Vampyrum spectrum can completely ruin anet and escape in as little time as aminute. Also, the longer abat struggles in the net the more difficult it is to remove. As aresult, anetter should be careful not to set too many nets, as more bats may be caught than can be removed, leading to loss of both bats and nets. As soon as possible after abat strikes the net, the collector should grasp it with agloved hand and determine from which side it entered. It should be held firmly in the gloved left hand (for aright-handed person) while the ungloved right hand is used to extricate the bat from the open side of the pocket, starting with the head. Iusually try to remove netting from the bat's mouth first to pre- 74 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY vent further damage to the net, and then work back, freeing the wings and finally the feet. However, the reverse is sometimes more convenient. Each net must be readjusted at regular intervals so that the bag does not move along the sh~lf strands, leaving tight places where bats bounce off or bunched places that they more easily detect. If rain or fog causes water droplets to collect, these should be shaken from the net as soon as possible. Also, leaves, twigs, and insects should be removed quickly before they tangle anet. Badly tangled sticks can be removed most easily if first they are broken into small pieces; large beetles should be disarticulated. Even when one is careful, an occasional large bat will succeed in chewing through ashelf string and, as aresult, alarge section of net ceases to function. If possible, broken shelf strands should be rethreaded in daylight, but two experienced netters can accomplish the job in afew minutes at night. The net loops immediately above and below the broken string should be spread as far apart as possible and raised until the track of the broken string is roughly at eye level. A piece of woven nylon fishline of about the same diameter as the original string and ameter long should be tied to the end of the longest string so that this leader can be threaded easily without tension. The shelf string then is carefully threaded through about every fourth mesh along the original track. When the ends meet, one person holds the two strings to release tension (or an end loop is removed from its pole) while the other ties the splice in place (including asmall piece of leader) so that the repaired shelf string is the same length as before. Holes in the mesh are not easily repaired, and after enough accumulate the net should be discarded or cut into smaller pieces. Anet should not be closed until all insects, twigs and other debris have been removed, after which all of the loops are pushed together near the upper ends of the poles. In areas where human interference is not aproblem, nets may be left "closed" on the poles until the following night, but when they are removed it is important to keep the loops in order. Iusually follow Handley's (1968:17) method of tying apiece of white string about 40 centimeters long to the top loop of each end. Before removing anet from its poles, the string at each end is threaded through the rest of the loops and tied. Next the loops are removed from one pole, and the net is folded by reaching out about ameter at atime to grasp the net, folding it back to the first hand again and again as one walks toward the other pole. Finally, the loops are removed from the other pole, tied, and the net is folded and stored in asmall bag. Traps Traps generally are not broken down between settings, but if this is necessary, as for shipping, they are reasserrlbled in the following manner. Each frame is assembled separately and bolted to the other using four threaded rods. The legs are bolted in place and the trap righted before the angled rods and wires are bolted to the top of each frame and carefully unrolled. The threaded rods for adjusting wire tension are extended as far as possible, and the botton angled rods are bolted to the bottom on each frame. Finally the threaded rods at the top are tightened to adjust wire tension. BIOLOGY OF THE PHYLLOSTOMATIDAE 75 Proper adjustment of trap frames and wires is essential. Much depends on the speed and angle of approach by bats. The trap should be vertical and perpendicular to the flight path. Traps are generally most effective when adjusted so that the two parallel sets of vertical wires are roughly seven and ahalf centimeters apart, although this distance may need to be varied for different conditions and kinds of bats. During initial testing of crude trap designs, Isucceeded in capturing an impressive number of phyllostomatids (Tuttle, 1974a). However, all subsequent trap modifications were designed to increase vespertilionid captures, without consideration of phyllostomatids, and the wire spacing was increased from two centimeters to two and ahalf centimeters. Aspacing of two centimeters or less, combined with increased elasticity of the springs, might prove advantageous for phyllostomatids. Regardless of spacing, the tautness of the wires should be proportional to the speed of the bats. Normally, wires should be adjusted so that they are barely tight. When bats escape by bouncing off the trap, wires should be loosened; when bats pass completely through the trap, both frames should be tightened. Captured bats are easily removed with gloved hands, and should be sorted into separate bags to avoid placing carnivorous species with other bats. Several thousand can be handled in an hour. Even though large numbers of bats do not damage traps, one must be constantly vigilant lest bats rapidly accumulate and suffocate before removal. Whenever atrap begins to catch more bats than can be removed conveniently, it can be turned sideways, carried out of the flight path, or covered on one side with asmall canvas. CAPTURE TECHNIQUES Roosts Little is known about the ecological requirements of phyllostomatid bats, and there is apaucity of information available on roosting behavior. Iselected 28 sources from which information pertaining to roosting habits was taken (Table 1). Pine (] 972) was used as the sole source of material on Carollia due to prior confusion in identification. Walker (1964) is cited only when original observations could not be found. Sources are numbered (see parenthetical numbers in Literature Cited), and numbers of references cited appear in the appropriate places in Table 1. Species for which I was unable to find information on roosting habits are not included. Early literature emphasized discovery of new species and seldom mentioned how or where bats were collected. Recently, the use of mist nets has enhanced knowledge of overall distribution and provided much ecological data. Nevertheless, netting has been so convenient that few researchers have been forced to look for roosting bats. Searches for roosts have been limited to a few obvious types of places. As aresult, roosts in caves, houses, hollow trees, or culverts are often reported whereas those in foliage and other less evident places are not, leaving the roosting habits of even some common species unknown. With this bias in mind, Iwill provide suggestions for finding the types of roosting sites that have proven most productive. Caves.-Caves may provide roosting sites for more different species of phyllostomatids than any other kind of shelter. Most caves are located in limestone, How to Collect at Roosts In addition to the more general equipment already discussed, abee smoker, hand net, and .22-caliber pistol are essential for collecting at many roosts. In the discussion that follows, it is assumed that at least two persons will be working together. In all collecting at roosts, it is vital to avoid alarming bats with unnecessary noise, vibration, or light. Hand netting.-A hand net ideally should have asturdy hoop about 40 centimeters in diameter attached to a1.2-meter aluminum handle. If needed, additional sections of telescoping aluminunl can be purchased for extending the handle to five meters. The bag should be made of nylon army mosquito netting, at least 75 centimeters deep, rounded at the bottom, and sewn at the top to heavy cloth fitting over the hoop. Apiece of heavy plastic 18 centimeters wide should hang freely around the inside of the hoop, preventing climbing bats from escaping. Hand nets are most frequently used at roosts in hollow trees, animal burrows, rock crevices, or caves. At ahollow tree, careful inspection should be made to determine the number and size of openings fronl which bats could escape. Each potential exit then should be covered with anet or somehow blocked. Many bats can be frightened into attempting to leave by pounding on the tree trunk with a rock. If that fails, alimber stick of appropriate length may be Gut, leaving small branches and foliage intact at one end. This can be carefully inserted and twirled near the bats. If bats remain stubborn in their refusal to come out, abee smoker can be used. Alength of flexible tubing may be attached, if needed, to direct the smoke to aspecific place. Emerging bats are caught in the hand nets, from which they are transferred to holding bags. Hand-netting in caves is much more difficult. Especially near the entrance, each depression or crevice should be approached cautiously, with the headlight not aimed beyond the reach of the net. Frequently, an extra section of handle is required so that bats can be reached quickly before they become alarmed. When acolony is heard, the roosting bats should be approached by sound rather than by sight, with the headlight aimed at the floor just ahead. Speed is crucial inasmuch as many bats will fly as soon as possible after sighting an approaching light. Most collectors find it easier to wait until evening when emerging bats can be trapped or netted at the cave entrance. Shoot ing.-Bats can be shot at roosts with a.22-caliber pistol and long rifle dust shot. Although such pistols frequently are bored smooth to improve the shot pattern, Ihave never found that to be necessary. The acceptable collecting range is roughly four to nine meters. For greater range, one can use a.32 or .410-caliber auxilIary barrel and dust shot in a16-gauge shotgun. The most frequent problem is that of shooting at too short arange, thus damaging specimens. Shooting is best employed when collecting at sites that are easy to see from a distance and difficult to approach without alarming bats. Examples of such places are foliage roosts, cavities in cliff faces, overhanging roots, large caves, culverts, and bridges. Apistol also can be used in large hollow trees and in small caves, but there is danger of damaging the ears of the collector. Whenever 82 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY BIOLOGY OF THE PHYLLOSTOMATIDAE 83 other options are available, shooting should be avoided, as some specimens may be damaged while others, which escape, may be needlessly injured. Netting.-Nets can be effective in capturing bats emerging from roosts not easily covered by ahand net, and are especially inlportant at bridges where small colonies may be difficult to approach. At aculvert, for example, one person tends the net, which is set to block one end, while another frightens the bats from the opposite end. Damaged nets can be cut to make one or more small nets, two to three meters long, which are handy at culverts. When not in use, short nets are easily rolled onto an aluminum pole. Mist nets also may be employed in front of such roosting sites as buildings and caves during evening emergence. Frequently, bats occupying these places cannot be reached or forced to exit before their natural departure. When colonies are small, mist nets can be quite convenient; when large numbers are involved, however, nets often entangle hundreds of bats at atime and are ruined long before bats can be extricated. Trapping.- Traps are particularly useful at entrances to caves containing large colonies. They may be set anywhere along the flight path of emerging bats, but the best place is often some distance from aroost entrance-for example, where bats normally enter foliage that can assist in obscuring the trap from detection. Prior observation of emergence patterns permits optimal trap placement. Frequently, however, traps simply can be set directly in front of a point of emergence or in acave entrance; the area around the trap may be partially blocked with brush or netting. Foraging Sites At least a few foraging bats can be found almost anywhere at night in the tropics; however, some places are far more productive than others. The few examples presented here may be intuitively obvious; many additional possibilities become apparent only with experience. Often the best places are discovered only by careful observation at twilight or at night while searching with aheadlight. Searches at night should include frequent pauses with the light turned off, listening for the sounds of falling fruit, flying bats, and the squabbling that occurs at major feeding sites. Trails.-Most forest trails at least ameter in width are likely to be used by bats, particularly when the surrounding vegetation is both tall and dense. In the tropics, the best trails are those that lead from villages to gardens or plantations. While these trails are especially productive places for collecting frugivorous species, others connecting pastures or leading to livestock sheds are more likely to be used by vampires. The widest trails, especially short sections between clearings, are better for foraging insectivores. Forest edges.-Bats forage and fly along the edges of most forested areas, but edges of small clearings within forests are best for collecting phyllostomatids, except when there are feeding or watering places in larger open areas nearby. Streams.--Streams provide natural flyways, especially where surrounding forest is dense. Slow-flowing streams, three to 10 meters in width, seem to have How to CollecT at Foraging Sites Shooting.- Though shooting at dusk or later with ashotgun and number 12 shot is an excellent method for collecting many emballonurids, noctilionids, vespertilionids, and nlolossids, this method seldom works well for obtaining the most traffic. Swift mountain streams and large rivers frequently are less productive of phyllostomatids and, in the latter case, pose difficulties in collecting. Ponds.-Isolated ponds in areas where there are no other sources of available water often attract bats in spectacular numbers and variety in the dry season. Other watering places, sometimes only afew centimeters in diameter, appear to be highly attractive to some stenodermines, even in the rainy season and in places where other water is abundant (Tuttle, 1974b). These sites are often well known to local native hunters who visit them in order to hunt tapirs (Tapirus terrestris), which also are attracted in unusual numbers. Feeding site s.-Many phyllostomatids are best collected in proximity to their feeding places. Glossophagines visit many flowering trees and shrubs, the best of which may be found by watching hummingbirds; certain plants that attract these birds during the day are equally attractive to bats at night. Flowering banana and cashew trees are well worth checking. Bats of the subfamilies Carolliinae and Stenoderminae are most commonly collected near fruiting trees or shrubs. Wild figs attract avariety of these bats in large numbers, and gardens containing fruiting bananas, guavas, papayas, or mangos also are excellent attractions. Especially in virgin forest, fruit-eating birds and monkeys often provide clues to additional food sources. Vampires frequently are numerous around the borders of villages when chickens, dogs, or pigs are present. Desmodus is encountered most frequently near cattle or horses, whereas Diaemus and Diphylla are more likely to be found near poultry. The presence of vampires is easily confirmed by the presence of dried blood on the head and shoulder regions of livestock, or on perches where poultry roost at night. Feeding areas of phyllostomatines seem to be more generalized and unpredictable, but Ihave nearly always succeeded in collecting Phyllostomus discolor, P. elongatus, and P. hastatus on small banana plantations in forested areas. Goodwin and Greenhall (1961 :240) reported P. hastatus flying in groups of up to 100 to feed on the seeds of spacaia nut trees (Lecythis zabucajo),and that Micronycteris megalotis was attracted to fruiting guava trees (op. cit., 228). I have nl0st frequently collected other phyllostomatines such as Chrotopterus, Mimon, Phylloderma, Tonatia, and Trachops in natural clearings beneath adense canopy of virgin lowland forest, where they appeared to be foraging. Such areas are found where undergrowth has been eliminated by wet-season flooding. Highland passes.-A surprising number and variety of phyllostomatids can be collected as they fly through low places along ridges. These are most easily found along roads that follow ridges. At elevations between 1400 and 2800 meters, Ihave commonly collected such interesting genera as Chiroderma, Enchisthenes, and Vampyrops. 84 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY BIOLOGY OF THE PHYLLOSTOMATIDAE 85 phyllostomatids. For shooting to be practical, bats must fly high enough to be seen against the horizon while there is stilJ adequate light, and there should be relatively bare ground or water below so that downed specimens can be found. The foraging habits of only a few phyllostomatids fit these requirements. Some of the larger species, such as Phyllostomus hastatus, can be shot at dusk from asmall boat as they attempt to cross rivers enroute to their feeding sites. They float and can be scooped from the river in ahand net. Large frugivorous species sometimes can be spotted with aheadlight and shot while feeding. Netting and trapping.-Many phyllostomatids have been collected efficiently only in mist nets and traps. Whereas nets have been used almost exclusively since the late 1950's, the potential of traps has received widespread attention only recently. Either nets or traps can be set at almost any place where bats are expected to fly, although they are not equally practical under alJ circumstances. Traps are especially convenient whenever large numbers of bats must be handled rapidly. They are not easily damaged by bats or other animals and do not require frequent attention unless exceptionally large numbers are being caught. Nets can be raised into the forest canopy, but the procedure is difficult and costly (Humphrey et aI., 1968). Traps, however, can be set easily in dense foliage on the ground or hoisted into the canopy without danger of becoming tangled. They are unaffected by wind, whereas anet must be set exactly perpendicular to even alight breeze or the netting quickly blows to one end, making the net virtually inoperable. The main disadvantages of traps are that they are much heavier than nets, and cover asmaller area. Either nets or traps may be set at any of the previously discussed kinds of places, but much of the success in using these devices depends on the collector's ability to camouflage them. Many feeding sites involve flowering or fruiting trees where nets or traps are set as close in front of atree as possible, or immediately beneath the lowest branches. Sometimes, however, nearby openings or trails used by approaching bats provide easier collecting sites. Along trails and streams, nets and traps should be set in the narrowest places, preferably where there are natural obstructions, such as fallen trees, that block all but asmall space. Traps are particularly effective at such sites. Hanging vines, overhanging limbs, and sharp turns provide additional concealment. At ponds and small clearings, where larger areas must be covered, nets are more easily used and should be set around the edges parallel to the vegetation. Around native gardens and other similar sites, Ifrequently have strung as many as 10 12-meter nets end to end, alternating the loops from two nets on each pole, but such an array of nets must be manned by several people. If traps are to be used at these places, they must be set either where vegetation forces natural funneling of the bats or where artificial blocking at the sides can achieve the same end. Often the sides can be blocked by tying astrong line to the top of the trap on each side, running the lines to nearby trees. Leafy vegetation is then cut and hung from the lines. This is especially effective at the approaches to ponds or where traps are set over streams. At low passes along mountain ridges, nets are preferable, and several may be set end-to-end just below the crest where they blend with the steep hillside. ACKNOWLEDGMENTS Iwish to thank Dr. Robert S. Hoffmann and Ms. Diane E. Stevenson for critically reading the manuscript and otherwise assisting in its preparation. Dr. Charles O. Handley, Jr., generously provided assistance during my early collecting experience, which was in part supported by U.S. Army contract DA-49193-MD-2788. LITERATURE CITED ALLEN, G. M. 1939. Bats. Harvard Univ. Press, Cambridge, Massachusetts, x+368 pp. ALLEN, J. A. 1911. Mammals from Venezuela collected by Mr. M. A. Carriker, Jr., 1909- ] 9] I. Bull. Amer. Mus. Nat. Hist., 30:239-273. (l) ANTHONY, H. F. 19]8. The indigenous land mammals of Porto Rico, living and extinct. Mem. Amer. Mus. Nat. Hist., n.s., 2:333-435. (2) BOND, R. M., AND G. A. SEAMAN. ]958. Notes on a colony of Brachyphylla cavernarul11. J. Mamm., 39: 150-151. (3) BURT, W. H., AND R. A. STiRTON. 196 J. The mammals of EI Salvador. Misc. PubL Mus. Zoo)., Univ. Michigan. I] 7: 1-69. (4) CONSTANTINE, D. G. 1958. An automatic bat-collecting device. J. Wildlife Mgt., 22: 17-22. 1961. Locality records and notes on western bats. J. Mamm., 42:404405. (30) SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Effectiveness at all places will be increased by frequent changes of position to counter learning behavior of bats. When choices are available, nets and traps should be set in the darkest places and at times when there is little or no n100n. Usually, the first and last two hours of the night are most productive, but afew species are more likely to be caught at other times. Techniques for luring bats.- There are two methods that have considerable potential for attracting bats to collecting sites. Many bats can be attracted to nets or traps when other bats are induced to call in distress. In general the smallest species make the best "callers." Often, the distress cries of asmall bat will attract large species in addition to others of its own size, whereas alarge species, such as Phylloslomus hastatus, calls its own kind while frightening most others away. On several occasions Ihave achieved excellent results by hanging abag containing 20 to 30 quarreling bats of several species on the side of abat trap in aplace where no bats had been caught previously. Ipredict that taped recordings may someday prove invaluable for luring bats into nets and traps. The less tested of the two techniques is the use of bait. At one locality in Peru, Carollia was so persistent in searching out my rat traps baited with banana that even those set beneath dense vegetation or fallen logs caught them. Although this was unusual, Carollia often was attracted to ripening stalks of bananas in native huts. Ialso have taken Rhinophylla pumilio in abanana-baited rat trap. Fruit, caged animals, or even caged insects could be hung behind traps set in places where bats would be forced to approach from the opposite side. It is quite possible that some glossophagines could be attracted to hummingbird feeders. Many phyllostomatids probably could be lured into baited bat traps. 86 BIOLOGY OF THE PHYLLOSTOMATIDAE 87 1962. Methodos de lucha contra los vampiros transmisores de la rabia. BoI. Ofic. Sanitaria Panamericana, 53:7-12. 1969. Trampa portatil para vampiros usada en programas de campana antirabica. BoI. Ofic. Sanitaria Panamericana, 67:39-42. DAVIS, R., AND E. L. COCKRUM. 1963. Bridges utilized as day-roosts by bats. J. Mamm., 44:428-430. DAVIS, W. B., AND R. J. RUSSELL. 1954. Mammals of the Mexican state of Morelos. J. Mamm., 35:63-80. (5) DAVIS, W. B., D. C. CARTER, AND R. H. PINE. 1964. Noteworthy records of Mexican and Central American bats. J. Mamm., 45:375-387. (6) DAVIS, W. B., AND D. C. CARTER. 1964. A new species of fruit-eating bat (genus Artiheus) from Central America. Proc. Biol. Soc. Washington, 77: 119-122. (7) GOODWIN, G. G., AND A. M. GREENHALL. 1961. Areview of the bats of Trinidad and Tobago. Descriptions, rabies infection, and ecology. Bull. Amer. Mus. Nat. Hist., 122:187-302 (8) GOODWIN, R. E. 1970. The ecology of Jamaican bats. J. Mamm., 51:571-579. (9) GREENHALL, A. M., AND J. L. PARADISO. 1968. Bats and bat banding. Resource PubI. Bur. Sport Fisheries and Wildlife, 72:iv +1-48. HALL, E. R., AND W. W. DALQUEST. 1963. The mammals of Veracruz. Univ. Kansas Publ., Mus. Nat. Hist., 14:165-362. (10) HALL, E. R., AND W. B. JACKSON. 1953. Seventeen species of bats recorded from Barro Colorado Island, Panama Canal Zone. Univ. Kansas PubI., Mus. Nat. Hist., 5:641-646. (11) HANDLEY, C. 0., JR. 1966. Checklist of the mammals of Panama. Pp. 753-795, in Ectoparasites of Panama (R. L. Wenzel and V. J. Tipton, eds.), Field Mus. Nat. Hist., Chicago, xii +861 pp. (12) 1968. Capturing bats with mist nets. Pp. 15-19, in Bats and bat band ing, Resource Publ., Bur. Sport Fisheries and Wildlife, 72:iv+1-48. HUSSON, A. M. 1954. On Vampyrodes caracciolae (Thomas) and some other bats from the Island of Tobago (British West Indies). ZooI. Medad. Mus. Leiden, 33:6367. (13) 1962. The bats of Suriname. Zool. Verhand. Rijksmus. Nat. Hist. Leiden, 58:1-282, 30 pIs. (14) INGLES, L. G. 1953. Observations on Barro Colorado Island mammals. J. Mamm., 34:266-268. (15) JONES, J. K .• JR. 1966. Bats from Guatemala. Univ. Kansas PubI., Mus. Nat. Hist.. 16:439-472. (16) JONES, J. K .• JR., AND A. SCHWARTZ. 1967. Synopsis of bats of the Antillean genus Ardops. Proc. U.S. Nat. Mus., 124(3634):1-13. (l7) JONES. J. K .• JR., J. R. CHOATE, AND A. CADENA. 1972. Mammals from the Mexican State of Sinaloa. II. Chiroptera. Occas. Papers, Mus. Nat. Hist., Univ. Kansas, 6: 129. (18) MITCHELL, H. A. 1963. Ammonia tolerance of the California leaf-nosed bat. J. Mamm., 44:543-551. (31) PHILLIPS, C. J., AND J. K. JONES, JR. 1971. A new subspecies of the long-nosed bat, Hylonycteris underwoodi, from Mexico. J. Mamm., 52:77-80. (19) PINE, R. H. 1972. The bats of the genus Carollia. Tech. Monogr., Agr. Exp. Sta., Texas A&M Univ., 8:1-125. (20) SANBORN, C. C. 1936. Records and measurements of neotropical bats. Publ. Field Mus. Nat. Hist., Zool. Ser., 20:93-106. (21) 1951. Mammals from Marcapata, Southeastern Peru. PubI. Mus. Hist. Nat. "Javier Prado," Ser. A, Zool., 6: 1-26. (22) 1954. Bats from Chimanta-Tepui, Venezuela with remarks on Choeroniscus. Fieldiana Zool., 34:289-293. (23) Mexico, Inst. (28) Johns Hopk ins Press. (29) SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 1955. Remarks on the bats of the genus Vllmpyrops. Fieldiana Zool., 37: 403-413. (24) SCHWARTZ, A., AND J. K. JONES, JR. ]967. Review of bats of the endemic Antillean genus MOf1ophylllls. Proc. U.S. Nat. Mus., 124(3635):1-20. (25) STARRETT, A., AND L. DE LA TORRE. 1964. Notes on acoJ)ection of bats from Central America, with the third record for Cy((arops alecto Thomas. Zoologia,49:53-63. TABOADA, G. S., AND R. H. PINE. ]969. Morphological and behavioral evidence for the relationship between the bat genus Brachyphyl/a and the Phyllonycterinae. Biotropica, 1:10-]9. (26) TUTTLE, M. D. 1970. Distribution and zoogeography of Peruvian bats, with comments on natural history. Univ. Kansas Sci. Bull., 49:45-86. (27) 1974a. Bat trapping: results and suggestions. Bat Banding News, ]5:4-7. 1974/7. Unusual drinking behavior in some stenodermine bats. Mammalia. 38: 141 -I 44. 1974('. An improved trap for bats. J. Mamm., 55:475-477. VILLA-R., B. ]966. Los murcielagos de Mexico. Univ. Nac. Autonoma BioI., xvi +491 pp. WALKER, E. P., AND OTHERS. 1964. Mammals of the world. Baltimore, 1:xlviii + ]-644. 88 CARE IN CAPTIVITY ARTHUR M. GREENHALL In the past 50 years only about adozen articles have appeared that dealt solely with care of bats in captivity. With the exception of the spectacular flying foxes and the vampire bats, maintenance of bats either as experimental animals or as zoological park exhibits has been neglected. Interest in bats as laboratory animals increased after 1953, mostly stimulated by discovery in the United States of rabies in insectivorous bats (Constantine, 1970). As it became evident that public health relationships existed between bats and man, attempts increased to study live bats under control1ed conditions. Valuable sources of the information included in this chapter have been unpublished manuscripts or papers in press. Ihave also included some personal observations where appropriate. Ihave reviewed most of the available literature, but despite active research interest, published information is woefully lacking for captive Chiroptera, particularly the Phyllostomatidae. Little or nothing is known about the care of insectivorous phyllostomatids. However, because anumber of vespertilionids and molossids have been raised successfully on artificial diets, I have described diets, cage systems, and techniques for their husbandry, as apotential guide for the care of insect-eating phyllostomatids. Phyllostomatid bats present a nUITlber of unusual maintenance problems. What these problems are, and how they have been or may be solved, is the topic of this chapter. This information is intended primarily for those who maintain bats in captivity for research or educational purposes and not for someone simply interested in keeping bats as pets. TRANSPORTATION Transportation of bats to the laboratory or zoo should be carefully planned. Because phyllostomatids are mainly tropical, transport time should be minimal, and the shipper must be aware that weather conditions may change rapidly from the hot lowlands to the cool uplands or from tropical to temperate latitudes. It is often possible to arrange for commercial carriers to take special intransit precautions with the animals being shipped. Ihave had excellent cooperation from airlines and shipping companies in keeping my bats away from extreme heat or cold or other potentially stressful situations-. Bats may be transported in metal cans, wire cages, light-weight wooden boxes, or in cardboard or plastic cartons. Most bats, unlike many other mammals, will not attempt to gnaw out of containers, but they can squeeze through incredibly small holes and cracks. Bats generally travel better individually than in groups, with each animal placed in asingle compartment or in alight cloth bag within a rigid container. Vampire bats and carnivorous species should not be grouped with bats of other species. Care must be taken to avoid exposing bats to the sun, to provide proper ventilation, and to control temperature and humidity. Food 89 and water must be provided for long trips. All persons wishing to transport bats should be aware of the rules and regulations governing the national and international shipment of live animals. On arrival, the shipping containers in which the animals are received should be either incinerated or thoroughly cleaned and sterilized to minimize contamination by disease organisms or parasites. If permanent living quarters are unavailable, cleaned temporary cages, such as those used for shipping, could suffice. The physical condition of every bat should be assessed upon arrival. THE LABORATORY ENVIRONMENT Temperature and Relative Humidity Among the more important factors influencing the successful maintenance of bats are temperature and relative humidity. If possible, these should be controlled automatically. Under natural conditions bats are exposed to daily fluctuations of temperature and relative humidity; however, little is 'known about the optimum conditions for captive tropical bats. Nevertheless, atemperature of 20 to 25°C and arelative humidity of 70 to 75 per cent seems satisfactory for many species. Low humidities can be injurious to the wing membranes (Racey, 1972). Uwe Schmidt (personal communication) maintains his animal room at aconstant temperature of 27°C and arelative humidity between 65 and 75 per cent. This is satisfactory for Phyllostomus discolor, Carollia perspicillata, Artibeus lituratus, and Desmodus rotundus. Rasweiler and de Bonilla (1972:659) and Rasweiler and Ishiyama (1973:56-57) maintained their laboratories at tenlperatures between 21 and 28°C and arelative humidity between 55 and 92 per cent, which proved satisfactory for Glossophaga soricina, Anoura caudifer, Phyllostomus discolor, Carollia perspicillata, Artibeus lituratus, and Sturnira lilium. My bat laboratory in Trinidad was not air-conditioned. The daily temperature ranged between 21.1 and 29.4°C, and the relative humidity between 55 and 95 per cent, conditions undoubtedly suitable for Glossophaga soricina, Phyllostomus discolor, P. hastatus, Vampyrum spectrum, Carollia perspicillata, Vampyrops helleri, Artibeus jamaicensis, A. lituratus, and Desmodus rotundus in that many of these bats lived in the laboratory for several years. The Desmodontinae do well under avariety of laboratory conditions. Wimsatt and Guerriere (1961 :450) maintained Desmodus rotundus in an air-conditioned laboratory at temperatures between 20 and 25 °C and arelative humidity between 30 and 65 per cent, which approximated temperatures and humidities previously recorded in Mexico. They also observed that vampires in the laboratory tolerated higher relative humidities and short exposures to lower temperature, but that their tolerance of temperature above 25°C was poor. In Trinidad, Greenhall (1965b:442) kept Desmodus and Diaemus in alaboratory at temperatures between 21 and 29°C and an average daily relative humidity of 75 per cent. The colony of Desmodus and Diaemus studied by Dickson and Green (1970:38) in London was kept in quarters held at aconstant temperature of 240Cduring the SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 90 BIOLOGY OF THE PHYLLOSTOMATIDAE 91 day but which cooled naturally to 21 °during the night. The relative humidity was fairly constant at about 50 per cent. At an elevation of 2545 meters, where vampires do not normally occur in Mexico, Schmidt and Greenhall (1972:243) attempted to maintain alaboratory temperature of between 25 °and 30°C and arelative humidity above 55 per cent for Desmodus. At this elevation there were problems in controlling an acceptable temperature and relative humidity. Power failures, aggravated by the lack of astandby generator for emergency use, complicated matters. We normally used thernl0statically controlled electric heaters and infrared heat lamps to maintain the temperature and cool mist electric hunlidifiers to control relative humidity. Uwe Schmidt (personal communication) reported Desmodus thrived in his animal room in Germany, which was maintained at aconstant temperature of 27°C and arelative humidity of between 65 and 75 per cent. Daily fluctuation in temperature and relative humidity can be monitored with ahygrothermograph. The thermograph readings should be calibrated with an accurate nlinimum-maxinlum thermonleter; the hydrograph with awet and drybulb hygrometer. Ventilation.- There is practically no information on the importance of ventilation and circulation ofair in laboratories housing bats. Pye (I967) cautions that many bat species are sensitive to draughts and overventilation should be avoided. In my experience, bats in poorly ventilated laboratories appear restless. Light Light appears to be an important factor in regulating the daily activities of bats (DeCoursey and DeCoursey, 1964). Uwe Schmidt (personal communication) claimed that acolleague investigating daily bat activity found that Phyllostomus discolor, Carollia perspicillata, and Artibeus lituratus died when kept continuously in total darkness for 10 days. Illumination is automatically controlled in many laboratories and 13 hours of light and II hours of darkness has been found to be satisfactory for Phyllostomus discolor, Sturnira lilium, and Artibeus lituratus (Rasweiler and Ishiyama, 1973), Glossophaga soricina, Anoura caudifer, and Carollia perspicillata (Rasweiler and de Bonilla, 1972), Desmodus rotundus and Diaemus youngii (Dickson and Green, 1970), and Desmodus rotundus (Schmidt and Greenhall, 1972). Many of the same phyllostomatids, except Diaemus, have been displayed successfully in alarge simulated South American cave and tropical rain forest exhibit at the New York Zoological Park where, by varying the intensities of white, blue, green, and occasionally red light, the activity patterns of the bats were reversed. Vampire bats kept by Wimsatt and Guerriere (1961) were subjected to low-intensity illumination from light entering through two glass-brick windows and glass panel in adoor. The bat cage itself further reduced the light because only the front was made of transparent material. Electric lights were turned on in the bat room only briefly when the cages were cleaned or the animals were being attended. No effort was made by the investigators to control the light regime because wild vampires are found in roosts receiving varying intensities of light. Insectivores disinfectant with aresidual available iodine of at least 1: 10,000, or 1%concentration of soapy water or detergent can be used. The solution should be autoclaved and discarded after each use. Hot soapy water or detergent can be used for swabbing floors and tables. "Glassware, plasticware and instruments ... should be discarded into plastic or glass receptacles containing one of the disinfectants mentioned above. They should be autoclaved. "Carcasses and animal tissue ... are best disposed of in plastic bags and incinerated." SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY In discussing the feeding habits of the Phyllostomatidae in another chapter, Gardner noted that the family's only true insectivore may be Macrophyllum macrophyllum. He cited examples of insect remains having been found in the stomachs or fecal remains of all subfamilies, suggesting that, regardless of the basic food preferences of each group, insects are probably an important food component of the diet. However, in the case of the Desmodontinae, insects probably were ingested accidentally when preparing abite site on some animal selected as prey, or during grooming activity. Racey (1972:297-299), in his review of the care and management of bats, listed 33 genera and 54 species of insectivorous bats that have been kept in captivity, but the only phyllostomatid mentioned was Macrotus. Micronycteris mega/otis was kept by Ruschi (1953a), who indicated that the bats lived and reproduced, but he did not provide further information. With the exception of Macrotus and Micronycteris, species of which also eat fruit, it is not surprising that the literature is wanting on the care of DIET Of all bats, the Phyllostomatidae probably have the most varied food preferences. Their natural feeding habits are discussed by Gardner (this volume), and his chapter on food habits will be indispensable to anyone who must prepare adiet for any phyllostomatid not yet successfully kept in captivity. Except for a few species, the literature is meager about the diets used by laboratories or zoological parks when keeping New World leaf-nosed bats in captivity. For this discussion Ihave grouped the phyllostomatids as insectivorous, nectarivorous, frugivorous, omnivorous, carnivorous, and sanguivorous. No bat in captivity can eat exactly as it would under natural conditions and specific natural food items may be impossible to supply, making substitutions essential. Adaily intake of protein appears necessary, but it is impossible to state what the required amounts should be in formulating abalanced diet. Hopefully, however, with vitamins, minerals, and other food additives, the nutritional requirements of acaptive bat may be resolved. Various vitamin preparations are available and many investigators have their own preferences. One, Stuart Formula Liquid, seems almost to be apanacea for diet deficiencies, not only for bats but for other captive small mammals (see Appendix] 8for sources of products mentioned in this chapter). 98 insectivorous phyllostomatid bats. Ibelieve, therefore, that it will be of value to describe diets used successfully to maintain insectivorous bats. Insect diets.-Vespertilionid and molossid bats have been fed avariety of insects such as greenbottle flies (both adults and maggots), house flies, instars of grasshoppers, locusts, and crickets in addition to bees, June beetles, termites, waxworms, and waxmoths (Gates, 1936:270~ Ramage, 1947:61). Most captive insectivorous bats thrive on adiet of mealworms (Tenebrio molitor)~ although the larvae are preferred, the pupae also are eaten. Pye (1967) cautioned that dietary deficiencies may occur if bats are fed mealworms that have had apurely farinaceous diet. This deficiency may be overcome by the addition of agood quality commercial animal feed to the mealworm's diet. Also, mealworms may be dusted with vitamin and mineral mixes (Rasweiler, 1975) or coated with vitamin drops such as those used for children (Gardner, personal communication). Concerning adiet of mealworms, Racey (] 972) and Ladische et al. (1967) advised that there may be some toxic quinones in some mealworm imagoes. Gates (1938b) added pieces of honey bees to the diets of Eptesicus, Myotis, Lasiurus, Plecotus, and Tadarida. The bees seemed to improve the consistency of the feces and added to the palatability of the food. He tried other insects, such as grasshoppers and June beetles, but the bats preferred bees. Ramage (1 947: 61) had no success in persuading various species of Myotis and Eptesicus to eat the foods suggested by Gates (1936) until she provided the larvae, pupae, and adults of greenbottle flies, "which can be easily reared in enormous quantities." AMyotis californicus she kept refused to eat flies but was raised successfully on termites. Ramage (1947:61) commented, however: "Termites have the dual disadvantage that they cannot be cultured rapidly enough to feed the bats and must be chloroformed or killed to keep thenl from crawling away before the bats have achance to eat them." Racey (1972:302), in his discussion on insects as food for insectivorous bats, mentioned that the larger bat species also will take early instars of many Orthoptera, "the most commonly cultured of which are locusts and cockroaches." Orr (1954:168) mostly fed mealworms to Antrozous pallidus during the early phases of his study, but later he used aprepared diet recommended to him by Ernest P. Walker. However, he offered (p. 234) alisting of other kinds of animal foods such as avariety of flies, moths, and even snails. Elsewhere in his study, Orr (op. cit., 232-233) cited, "records of captive pallid bats which were observed to eat western skinks (Eumeces skiltonianus), aSonoran desert gecko (Coleonyx variegatus), and were suspected of eating the head and neck of aMexican freetailed bat (Tadarida mexicana). It seems likely that starvation was responsible for such deviation from an insectivorous diet, although ... it is possible that small night lizards may be preyed upon locally by pallid bats." Artificial diets.-l have used insect traps to catch insects for bat food. At times, however, insects may be either scarce or not available. Consequently, investigators have had to devise substitute diets. These are mashes or mixtures comprised of anumber of items that are readily taken by the bats and usually inBIOLOGY OF THE PHYLLOSTOMATIDAE 99 100 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY clude such things as banana, cottage cheese, hard-boiled egg, and vitamins. Such mixtures are commonly called Hgl op "by bat biologists. Gates (1936, 1938b) found that various species of Myotis, Pipistrellus, Eptesicus, and Plecotus do well in captivity on artificial diets that may be completely different from their normal diets. He (1938b: 157) noted: ~'Under captive conditions they have been known to eat practically everything, unless it is too highly seasoned. This includes all cereals, breads, crackers, cakes, meats, eggs, vegetables of all kinds, both fresh and cooked, lettuce, celery, and all of the not too acid fruits, apples, pears, peaches, prunes, pineapples, and figs. All milk products, both fresh and sour milk, buttermilk, cheese, and even butter are acceptable. In fact, the author has hardly found any food which they will not eat. They of course have their preferences, apparently preferring the milder cheeses to anything else. However, bread crumbs moistened with buttermilk are also greatly enjoyed." Gates (1936: 270) first suggested that chitin was essential for the proper formation of fecal pellets and the prevention of intestinal obstruction in insect-eating bats. The ease of obtaining bananas and cottage cheese tempted Racey (1970) and others to feed amash lacking insects. The bats did poorly, however, and their pelage deteriorated. Empirically, it was discovered that mealworms or other insects added to the mixture corrected the condition. There are anumber of recipes for glop. Walker (1966: 138) developed afood mixture relished by many small mammals at the National Zoological Park (Appendix 9). Davis and Luckens (1966) used banana, cream cheese, canned dog food, and multivitamins to feed Epte sicus (Appendix 10). Mohos (1961) used the Walker (1966) formula to feed Myotis, Pipistrellus, and Eptesicus. However (p. 371), Hoccasionally equal parts of beef and beef liver were substituted for the cottage cheese, since it was found that, after an initial adaptation period, the bats fare equally well on this diet." J. Frederick Bell (personal communication) fed Myotis lucifugus homogenized whole baby mice, which were readily available in his laboratory. Krutzsch and Sulkin (1958:262-264) tried a number of feeding techniques and food combinations to induce their captive Tadarida brasiliensis to feed. Live mealworm larvae were unsatisfactory, and a nutrient fluid containing amino acids, simple sugars, and vitamins caused the bats to develop dysentery with fatal results. They were finally successful in maintaining Tadarida on glop. Food storage.-Diet preparation may be simplified in that the various food ingredients, including insects, can be mixed in an electric blender, preweighed in wax paper or plastic bags, and then stored frozen at 4°C until used (Mohos, 1961 :37] ;Davis and Luckens, 1966:226; Barbour and Davis, 1969:246; Rasweiler, 1975). The size of the food packets should depend on the number of bats to be fed at anyone time. Before feeding the bats, frozen food should be removed from the freezer and allowed to thaw. Once thawed, food will last about a week under ordinary refrigeration. It is important not to serve wet mashes and other liquid diets too early in the day because they may begin to spoil before all bats have fed. This can lead to diarrhea and malnutrition. BIOLOGY OF THE PHYLLOSTOMATIDAE Nectarivores 101 Until recently the standard diet for such captive nectar-feeding bats as G/ossophaga and Anoura has been sugar or honey in water, fruit juice, succulent fruits, and vitamins (Ruschi, 1953c, 1953d, 1953e; Goodwin and Greenhall, 1961 ;McNab, 1969). Rasweiler (1975) claimed that ~'laboratory diets for frugivorous or nectarivorous species based solely upon fruit pUlp andlor fruit juices may be grossly inadequate from anutritional standpoint." He believed that addition of insects and pollen could significantly increase the protein, fat, mineral, and vitamin levels in diets of fruit-eating bats, as well as supply essential amino acids that are inadequately represented in the fruit component of the diet. Rasweiler and de Bonilla (1972) and Rasweiler (1973) have formulated diets (Appendix 12) that have been successful for the long-term maintenance of large numbers of G/ossophaga soricina, Anoura geoffroyi, A. caudifer, and Carol/ia perspicil/ala. The New York Zoological Park prepares an artificial nectar (Appendix 11) dispensed from large watering bottles hidden among the plastic plants of the exhibit. In these exhibits, the nectar-feeding bats also have access to the solid diet (Appendix 15) offered to the frugivorous phyllostomatids (House and Doherty, 1975). The bats housed in this exhibit have included G/ossophaga soricina, Anoura geoffroyi, Phyllostomus discolor, and Carol/ia perspicil/ata. Donna J. Howell (personal communication) successfully raised nectarivorous bats on adifferent diet (Appendix 13). She also maintained Phyllostomus discolor and Carol/ia perspicil/ata, but treated them as fruit eaters. Howell wrote: HI've kept Leptonycleris, Choeronycleris, Glossophaga, Anoura, and Hylonycleris on the nectar diet for periods exceeding ayear. All the diet ingredients seemed necessary to duplicate the very nutritious contents of 'bat-adapted' pollen and nectar from chiropterophilous plants. One must be careful to give the bats enough protein, yet not over protein-load or over sugar-load their kidneys. Concentrating ability of the glossophagines is very poor. Protein should stay about 9to 11 per cent, sugar 14 to 20 per cent." Frugivores The diet of nectarivorous and frugivorous bats is influenced mostly by the seasonal abundance of flowers and fruits (Greenhall, 1956, 1957; Goodwin and Greenhall, 1961; Fleming el ai., 1972). The diets of captive frugivores will be determined by the availability of fruits. In temperate regions, tropical fruits, with the exception of bananas, may not always be available. Captive fruit-eating bats generally prefer sweet fruits such as bananas, mangoes, peaches, plums, melons, grapes, and papayas (Pye, 1967; Racey, 1972). Although citrus fruits are not preferred, sweet oranges and grapefruit occasionally may be accepted (Greenhall, 1966). Pye (1967) mentioned that apples can be substituted if exotic fruits are in short supply. The banana-based diet formulated by Rasweiler (1975) and Rasweiler and de Bonilla (1972) is given as Appendix 14. 102 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY When offering food, it is advisable to use small pieces, about I-centimeter cubes, well mixed. This will prevent dominant individuals from selecting all the choice fruits or obtaining more food than their cage mates. Small pieces can be easily managed by bats while eating or flying. The smalJer fruit-eating bats can handle whole fruits only when they are overripe or soft enough that asmall hole can be bitten in the skin through which the fruit pulp and juice can be extracted. Some biologists have fed phyllostomatids some unusual dietary items. For example, Ruschi (1953j) fed Artibeus lituratus blood as well as fruit and insects, because he believed that this species feeds upon blood in the wild. Goodwin and Greenhall (1961) stated that Phyllostomus discolor will not eat flesh in captivity and prefers fruit such as bananas, mangoes, and papayas. McNab (] 969) reported, however, that captive P. discolor require asmall, but regular, intake of meat. Uwe Schmidt (personal communication) mentioned having successfulJy reared this species on sliced bananas, mealworms, and neonatal laboratory mice. McNab (1969) reported that he kept anumber of frugivorous phyllostomatids healthy for extended periods of time, but, unfortunately, provided no information on diets other than the fact that Rhinophylla pumilio, Uroderma bilobatum, Artibeus cinereus, and A. concolor were fed fruit. His Phyllostomus elongatus and Vampyressa nymphaea would not eat. Polyphagous Phyllostomatids Agood example of an omnivorous phyllostomatid is Phyllostomus hastatus, which readily adjusts to captivity. Dunn (J 933) fed P. hastatus mice, bats, birds, defibrinated blood, fruit, and raw meat, including liver. Ruschi (1953b) and McNab (] 969) provided similar fare, the former adding cockroaches when available. Goodwin and Greenhall (J 96]) found that this species, in addition to accepting awide variety of fruit, thrived on mice and young birds. It did not hesitate to kill and eat other bats placed in its cage although it appeared to be uneasy in the presence of Desmodus. The diet used by the New York Zoological Park for their colony of P. hastatus is given in Appendix 16. Donna J. Howell (personal communication) fed both Phyllostomus hastatus and P. discolor aspecial diet for frugivorous bats (Appendix 17) and commented that these bats also got a dish of mealworms at each feeding. Carnivores The best known carnivorous phyllostomatid is Vampyrum spectrum. It has been raised successfully in captivity on raw meat as welJ as dead whole chicks and pigeons (Ditmars, 1935, J 936; Goodwin and Greenhall, 1961; Crandall, 1964; Bradbury, 1970). Greenhall (1968) described the care of Vampyrum, which successfully raised young during the five years bats were maintained in captivity. They were fed pigeons, chicks, wild birds, and dead laboratory rats and mice as weI) as raw meat cut into 2-centimeter chunks. Although these Vampyrum never fought over food, there was always the risk of injury to their wings when they stalked live prey because the cage was too small. Therefore, all BIOLOGY OF THE PHYLLOSTOMATIDAE 103 food was killed and usually presented to the bats by forceps or placed on the cage floor. Food was thoroughly masticated by the bats; feathers and rodent tails usually were discarded. Other carnivorous phyllostomatids kept in captivity include Trachops cirrhosus and Phylloderma stenops, which ate lizards (Pye, 1967), Chrotopterus auritus, which was kept on adiet of white mice by Villa-R. and Villa-C. (1969), and meat and bats by McNab (1969). Vampire Bats The three vampire bats, Desmodus rotundus, Diaemus youngii, and Diphylla ecaudata have been maintained in captivity with varying degrees of success. Desmodus adapts easily to captive conditions, whereas Diphylla is most difficult to maintain. Diaemus does fairly well once its basic requirements are recognized and met. Presently there are breeding colonies of Desmodus in many laboratories and zoological parks. Ditmars and Greenhall (1935) were the first to describe keeping vampire bats in captivity. Trapido (1946) demonstrated that Desmodus easily adapted to laboratory conditions and reported alongevity record of 12 years. Wimsatt and Guerriere (1961) detailed the care of Desmodus in temperate zone laboratories, whereas Greenhall (1965b) described its maintenance in the tropics. Wild Desmodus may consume amounts of blood equalling or exceeding their body weight (Wimsatt, 1969). The average weight of Desmodus is about 30 grams, and captive bats daily may drink up to 50 milliliters of blood, although 15 to 20 is usually sufficient (Pye, 1967). Wimsatt and Guerriere (1962) cited the unusual capacity of one captive 28-gram nonpregnant female they maintained in isolation for 17 days, which consumed blood in excess of her body weight on 13 days. On two days the amounts ingested were 47 and 52 milliliters, respectively. Dickson and Green (1970), in order to reduce the time and labor required in the maintenance of vampires, utilized trisodium citrate (1 milliliter 3.8 per cent per 10 milliliters of blood) to prevent blood coagulation and dispensed the blood meal in plastic hoppers. The use of citrated blood instead of defibrinated blood reduced the preparation time and dispensing meals in plastic hoppers prevented splattering and spillage thereby reducing the time for cage cleaning. According to Dickson and Green (1970), defibrination results in the loss of blood volume and the removal of factors essential for the vampire's welfare. Blood can be stored frozen at -20°C for up to six weeks without becoming unpalatable to vampires. Dickson and Green (1970) presented the daily ration warmed to 37°C at 1630 hours. Each food container was filled with 100 milliliters of bloodsufficient for five bats. These hoppers or glass-tubed drinking bottles prevented the wastage and contamination of the blood by urine and feces that usually occurs when vampires are fed from open dishes. Radiological examination revealed no evidence of bone decalcification following 15 months of citrate in the diet. 104 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Frozen blood, whether defibrinated or citrated, has atendency to spoil rapidly after thawing (Wimsatt and Guerriere, 1961; Greenhall, 1965 b). Therefore, the blood meal should be offered when the bats normally commence feeding. In the field, where fresh blood is usually difficult to obtain, an ingenious emergency supply was devised by Wimsatt and Guerriere (1961). This "instant blood" is shell-frozen and lyophilized defibrinated blood and is reconstituted with water, as needed. Diaemus has been kept successfully in the laboratory on citrated bovine blood -the same diet fed to Desmodus (Dickson and Green, 1970). Aweekly supplement of chicken blood was also provided by permitting the vampires to feed on the toes of alive chicken placed on the top of their cage. Nonetheless, Goodwin and Greenhall (1961) reported that Diaemus refused to drink defibrinated cattle blood even when mixed with chicken blood. Their bats, however, would accept chicken blood. At alater date, Greenhall (1970) reported observing Diaemus feeding on cattle. Ibelieve that Diaemus should be fed citrated whole blood as individuals apparently refuse to drink defibrinated blood. Diphylla has not been successfully kept in captivity for any length of time. Ruschi (195 1) stated that he maintained Diphylla and Desmodus on both citrated and defibrinated blood, and that the latter was preferred. However, VillaR. (1967) reported that Diphylla would not accept cattle blood, and although defibrinated chicken blood was consumed, the bats died within 48 hours. Perhaps captive Diphylla would accept citrated rather than defibrinated blood. Irecommend that liquid multivitamins be added to all blood meals. House and Doherty (1975) used 2.4 cubic centimeters of "Pet Drops" per pint of blood (Appendix 18). An unusual observation about vampire bat diet was reported by Kumm (1932), who noted that Desmodus fed readily in captivity on bananas and live animals. Ican find no confirmation that vampires feed on fruit. Vampire fasting.~chmidt et af. (1971) observed that wild Desmodus may not forage every night. If true, one must assume that such fasts are not deleterious. Wimsatt and Guerriere (1961) routinely skipped feeding their Desmodus on Sundays. On the few occasions that their bats were neglected accidentally for as long as three days, the effect was marked emaciation but not death. Greenhall (l965b:443) found that Desmodus could go without food for at least 62 and a half hours without apparent ill effects. Vampire cannibalism.- The only reference to cannibalism among Desmodus in captivity was described by Win1satt (1959). Wimsatt's colony consisted of four cages, each containing 15 to 20 bats. In one cage the daily ration of blood was always consumed, whereas in the other three there was always some blood left over. After several weeks, the bats in the first cage developed hairless patches on the shoulders, interscapular region, and back of the head. Associated lacerations were obviously caused by bites. This did not occur with the occupants of the other three cages, suggesting that this situation reflected unusual behavior. This abnormal behavior ceased and the injured animals subsequently recovered when blood was supplied in adequate amounts. The feeding on cage mates was BIOLOGY OF THE PHYLLOSTOMATIDAE 105 interpreted to be an attempt to secure blood in any way possible whenever the amount of blood available was deficient. Greenhall (1965b:442) observed that cannibalism might occur anl0ng newly captured vampires, but could easily be prevented by supplying the bats with .a greater quantity of blood than could be consumed in one night. Preparation of defibrinated and citrated blood.-A major obstacle to keeping avampire bat colony in the laboratory is the uncertainty of acontinuous fresh supply of blood. Defibrinated cattle blood has been used successfully for many years, but citrated blood may be preferable inasmuch as nothing is removed. Blood for defibrinating should first be collected in aclean vessel as it 1l0ws from the slaughtered animal. The blood must then be whipped or agitated with either wooden applicators, cocoa beaters, swizzle sticks, large wire beaters, whisks, or roughened glass beads. After several minutes of agitation, the fibrin will adhere to the beating instrument or glass beads and then may be discarded. The remaining blood may then be poured into another clean vessel for storage. Defibrinated blood will remain as aliquid and will keep under refrigeration for afew weeks. The color, usually bright red, might slowly turn purplish. The blood will be accepted by the bats if not spoiled. Greenhall (1965b) froze defibrinated blood in ice cube trays and other containers. Frozen defibrinated blood may be maintained safely for several weeks. Inasmuch as it may spoil rapidly, only single rations of blood should be thawed each day, and this should be used as quickly as possible. Iknow of some laboratories and zoos that have used outdated human blood and plasma from blood banks. The consensus is that vampires apparently do not do well on either fare. Dickson and Green (1970:40) described their method of citrating blood as follows: HBovine blood is collected from anearby slaughterhouse in 5liter containers, each containing 500 ml of 3.8 per cent trisodium citrate to prevent coagulation. This blood is strained through muslin into 500 ml polythene bottles; it can be stored frozen at -200Cfor up to 6weeks without becoming unpalatable to the bats." DRINKING WATER In their reviews of the care and management of bats, Pye (1967:498) and Racey (1972:303) stressed that all captive bats should have aplentiful supply of drinking water available. Most information about water requirements pertains to vespertilionid and molossid bats. Perhaps only afew phyllostomatids, such as Desmodus, are able to live for extended periods without drinking water. Most bats will learn to use drinking tubes, nozzles of inverted bottles (Racey, 1972: 303), or plastic hoppers of the types used for caged birds. Pye (1967), however, asserted that bats do not readily use drinking tubes and that water is best supplied in shallow dishes. These water dishes should be cleaned daily. According to Pye (1967:498), there is adanger that sick animals may drown in water dishes and the depth of the water, for smaller bats, should not exceed 3to 5millimeters. Some bats are reluctant to crawl on the floor so that food and water dishes should be placed on ashelf, fastened to the side, or suspended from the top of the cage. GENERAL CARE OF CAPTIVE BATS Acclimation There is little information about the acclimation of bats to captive conditions. Rasweiler (1973) initiated acclimation of his phyllostomatids immediately SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 106 Racey (1972) observed that vespertilionids will lap water from saturated cotton and this is auseful way of providing water during transport. Rasweiler (1975) daily provided water for his captive Phyllostomus discolor, P. hastatus, Artibeus lituratus, and Sturnira lilium. There are divergent opinions as to whether captive Desmodus rotundus require water to drink. King and Saphir (1 937) and Trapido (1 946) provided water ad libitum to these vampires. Wimsatt and Guerriere (1961) expressed surprise that their Desmodus drank little if any water; bowls of fresh water were kept in the cages at all times, but were not used even when the bats missed aday's feeding. They finally discontinued the practice and, after nearly two years without water, the bats showed no ill effects. Greenhall (l965b) observed that his vampire colony required aconstant supply of water for the bats' well-being. In addition to water supplied during the day, another 150 milliliters was given with the daily meal. During atwo-month check period (May and June), 113 milliliters was the greatest amount of water consumed in one night, whereas 10 nlilliliters was the least. Water evaporation was not afactor owing to the constant high humidity. At high elevations in Mexico, Greenhall et al. (1971) noted that the daily blood intake of Desmodus kept in glass jars increased when asupply of drinking water was added. However, when large numbers of vampires (about 200) were kept at one time, water was not provided in order to reduce the maintenance required to clean up to 100 extra water hoppers. Over aperiod of months the bats suffered no ill effects as aresult of withholding water. Water was always provided for the small group of vampires in the flight cage. Uwe Schmidt (personal communication) provided his Desmodus with water ad libitum day and night, and blood only during the night. He wrote: HI don't know whether water is essential for keeping vampires, but my feeling is they need it for their well-being, especially if they have eaten spoiled blood." While maintaining acolony of Desmodus in a flight cage, Rexford D. Lord (personal communication) observed: "The other day Isaw avampire come down and drink water even before drinking blood which was also available. He drank about 5milliliters of water, then flew up into the roost box." Lord described afood preference test where water was offered to ]9 bats. Each drank an average of 23.4 milliliters of blood and 2.5 milliliters of water nightly, over aperiod of nine nights. On one night, no water was consumed but during another night the bats drank atotal of I32 milliliters. Lord conducted another test where vampires were kept individually in oil can cages placed in an outdoor enclosure. Water consumption varied with the ambient temperature. On extremely hot days, the bats consumed 20 to 30 milliliters of water, drinking even during the day. On cool days the bats did not drink. Lord concluded: "I think water is necessary in hot climates or when there is no control of cage tenlperature." BIOLOGY OF THE PHYLLOSTOMATIDAE 107 following capture. In the field, food is placed in the transport containers. Upon arrival at his laboratory, all bats are hand-fed to provide some nourishment. This quick introduction to the new diet probably assists the bats to cope with the trauma associated with capture, transit, and the initial period of adjustment to captivity. Recalcitrant, weakened, or torpid Glossophaga soricina, for example, usually can be induced to consume some of their new diet, if at first a small amount is placed over the bat's nostrils (Rasweiler, 1975). Asecond method of feeding bats that are reluctant to accept hand-offered food is to force carefully an eyedropper containing the liquid diet into the bat's mouth (Rasweiler and Ishiyama, 1973:57). Upon tasting asmall sample of food, many animals will readily consume more. The bats are then fed from dishes. The same regimen was applied to Carollia perspicillata, Phyllostomus dicolor, Artibeus lituratus, and Sturnira lilium. Rasweiler and Ishiyama (1973:58) stated: "The hand-feedings were given in the hope that they would facilitate the transition of the animals to alaboratory existence." Training Bats to Eat Mealworms In the wild, most insect-eating bats catch their prey in flight, whereas a few may glean their prey from foliage or other substrates. In captivity, where food is placed in containers, many insectivorous bats require training before they wilJ feed themselves. Racey (1972), who has had much experience with captive insectivorous bats, emphasized that this training is the nlost crucial and timeconsuming stage in the acclimation of these bats to captivity. This is also true of weanling bats born in captivity. Novick (1963:51) stated that captive Macrotus must be hand-fed mealworms before they adapt to taking their own food from a dish. Although anumber of other phyllostomatid bats feed on insects in the wild, there is no information as to how these bats were trained to feed on insects in the laboratory. Lacking this information on insect-eating phyllostomatids, it may be valuable to describe how some investigators have trained vespertilionid and molossid bats to feed. The success of training bats to do what the investigator wants them to do depends in large measure on the patience and skill of the trainer, but also on the inclination of the bat to learn. Some bats require less training to eat mealworms than others. There are several methods, ranging from holding abat in agloved hand and offering food to simply allowing bats to learn to feed themselves from apile of nlealworms in adish. Constantine (1952:397) placed mealworms on the cage floor hoping that specimens of Tadarida brasiliensis would recognize their future diet after they ate their first few hand-offered worms. Racey (1970: 178) held the bat in his hand, decapitated the mealworm, and then applied the viscera to the bat's lips. The bat's jaws closed and the bat usually would chew the insect. If the bat did not chew in half aminute, then adrop of water was placed in the corner of the bat's mouth. If there was still no response, the mealwornJ was squeezed so that its contents entered the bat's mouth. After the first mealworm was swallowed, the bat's nose was brought in contact with the insects moving in adish or on the cage Food Poisoning Uwe Schmidt (personal communication) described some deaths in his Desmodus colony that he attributed to food poisoning. The condition may appear suddenly, even after bats have been in captivity for severa] years. Schmidt observed that "bats hung with stretched legs from the cage top and regorged blood (stomach was always completely filled) while urinating bloody urine." The cause, Schmidt believed, was ingested spoiled blood. Spoiled blood does not appear to be distasteful to vampires. Some bats died; others recovered. The animals that survived drank great amounts of water. The dead bats tested negative for rabies. Sore Limbs Swelling of the wrist joints was observed by Orr (1958), who thought that the high protein content of food, after ayear or two, may have caused this condition. Some bats will develop sores on the bottom of their feet from resting on a horizontal surface such as the bottom of abox or bottle. To correct this condition Barbour and Davis (1969:246) suggested fixing ascreen to permit the bats to hang head downward. Racey (1970) believed that swelling of joints in noctules might be associated with the lack of exercise. Some pteropodids and possibly other large bats such as Vampyrum, which normally hang pendant and free, may suffer from sore feet if not supplied with proper roosting surfaces such as tree branches. Excessively curved claws may result from continued hanging on wire (Pye, 1967). An excessively dry atmosphere in alaboratory may result in dry and brittle wing membranes. Raising the relative humidity and the application of amoistening skin lotion or baby oil may remedy the ailment. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY chiroptera and Microchiroptera. This condition is unsightly, but there do not seem to be any other adverse effects. After some time in captivity on unsupplemented diets, many bats are prone to weakening of the jaws and limbs, and general listlessness. Stuart Fornlula Liquid effects acomplete cure in some cases." Others have noted that bats fed on artificial diets sometimes lose their hair and that multiple vitamins added to the diet corrected the condition (Mohos, 1961; Davis and Luckens, 1966; Barbour and Davis, 1969). Rasweiler (1975) mentioned correspondence from J. Frederick Bell, who found that food containers must be kept clean to prevent the loss of hair on bats' bellies. Rasweiler (1975) also observed that individuals of Artibeus lituratus, unable to fly in small cages, were forced to crawl and rub against the cage wire, which resulted in the loss of ventral hair. Uwe Schmidt (personal communication) discovered that large numbers of ticks on abat will cause lose of hair through constant scratching by the bat in an attempt to remove the parasites. Temporary relief nlay be provided by washing the affected area of the bat with soap and lukewarm water. 114 MISCELLANEOUS LABORATORY TECHNIQUES Anesthesia and Euthanasia The use of anesthetics and euthanasics in laboratory animals is often necessary for humane and technical reasons. Muscle relaxants or paralytics are not anesthetics and should not be used alone for surgical restraint. They may be used for surgery in conjunction with drugs known to produce adequate analgesia. Ether is an anesthetic used by many biologists. Mohos (1961) administered ether from aregular dripping bottle onto asmall, gauze-lined, wire-mesh basket covering the bat's nose and mouth. Precautions must be taken to avoid heavily soaking the gauze inasmuch as the bat may swallow the ether and die. The depth of anesthesia can be regulated and recovery is usually fast and uneventful. Excessive ether may be used for euthanasia because bats have anarrow tolerance range for this anesthetic. Mohos (1961) found that nembutal, although easier to apply than ether, gave less satisfactory results. Pye (1967) preferred pentobarbitone sodium (nembutal) instead of ether and used one volume of commercial solution to nine volumes of 10 per cent ethyl alcohol injected intraperitoneally to induce surgical anesthesia, Exercise and ObesiTY There are various opinions on the exercise requirements of captive bats. Orr (1958) found that Antrozous, like many captive animals, tends to overeat once accustomed to captivity. If more than one bat is in acage, it is unwise to limit the amount of food offered because aggressive bats may consume more than their share. Overeating will lead to obesity and one solution is regular exercise. This may be accomplished by permitting the bats to fly in aroom. Mohos (J96 J) provided alarge room, 5 by 9meters, for bats to fly unhindered on the assumption that exercise would keep them in good condition. The exercise flight was satisfactory for small numbers of bats. The practice was later discontinued due to accidents and high mortality when hundreds of bats were exercised at once. However, when the exercise was stopped, single and multiple pyogenic infections developed in many of the animals. Racey (1970) believed that flying is not apractical exercise for large-scale bat husbandry, but did mention conditions he thought to be caused by the lack of exercise. His serotines often developed sore wrist joints and occasionally fatal wrist joint infections of Pseudomonas aeruginosa. The wrist joints of his noctulus also became stiff and infected. Wimsatt and Guerriere (1961) observed that, although exercise for vampire bats might be desirable for physiological reasons, their bats appeared healthy and vigorous without exercise. Glossophaga and Carollia are able to exercise within small cages because they can hover. Aldo M. Voute (personal communication) reported that the best way he found to keep captive bats in good condition was to place their cage in alarge room and allow them to leave the cage and fly about in search of food as they wished. Food was located at aspecific spot on a she1f~ the bats learned to find the food and always returned to their cage to roost. BIOLOGY OF THE PHYLLOSTOMATIDAE 1] 5 Cesarian Section Adams and Baer (1966) described cesarian sections used on Tadarida brasiliensis. Ether was used. The hair of the abdomen was clipped and the area dampened with one per cent bensalkonium chloride. Amidline incision was made through the skin and abdominal musculature with ascalpel, and the right hom of the uterus withdrawn. The uterine wall was carefully incised and the infant bat quickly withdrawn; fluids immediately were sucked out of its mouth at 30 to 50 nlilligrams per kilogram of body weight. There is considerable individual variation in response and the bat should be fully aroused before injection, and aconstant body temperature of 37° to 40°C must be maintained afterwards to·.ensure successful anesthesia and recovery. Overdosage with nembutal may be used to kill bats humanely (Pye, 1967). To anesthetize safely vampire bats, Dickson and Green (1970:43) introduced amixture of oxygen (2 liters per minute), nitrous oxide (1 liter per minute), and fluothane (halothane, 2.5 units Fluotec scale) into the cages. The bats were anesthetized in four to five nlinutes and recovered completely in three to four minutes. No bats died as aresult of this method. Racey (1972) preferred the halothane/oxygen method of anesthesia for vespertilionid bats. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 116 Marking Bats Various methods are employed to mark bats for individual recognItIon. Numbered metal bands attached to forearms have been widely used. Another method utilizing differently colored plastic bands, such as used for cage birds, is satisfactory for asmall number of bats. Some phyllostomatids, such as Glossophaga soricina, have apeculiar structure of the antebrachial membrane that makes attachment of bands to the forearm difficult. Neonates, the forearms of which are too small to take bands, may be numbered on the wing membrane with atatooing forceps (Racey, 1970). This method is useful only for short-term marking, because the small holes outlining the nurnbers heal over and become obliterated in afew days. However, the holes can be reopened with asuitable needle without danger of infection. When scar tissue forms, it is often unpigmented and the number can be recognized for several weeks. Rasweiler (1975) marked bats by means of spots bleached onto their fur. "The bleaching solution consists of six per cent H202,Lady Clairol Protinator and Lady Clairol Cremogenized Hair Lightener [Appendix 18] in the proportions of 10:2:5 respectively. The H202and protinator are mixed vigorously for about five seconds in asmall vial. The lightener is then added, and the mixture agitated for another 20 seconds. The working solution retains its activity for at least an hour after preparation. Immediately after the application of small amounts of the bleaching solution with abrush, the bats may be released back into their cages." Up to 30 different numbers are possible by varying the combination and position of the spots on the back and head of abat. Several hundred Glossophaga and Carol/ia have been so marked, with only afew minor cases of hair loss in adults and skin damage in juvenile Carol/ia. BIOLOGY OF THE PHYLLOSTOMATIDAE 1]7 with apipette or wiped out with acotton swab. The umbilical cord was clamped with ahemostat, ligatures tied on either side of the hemostat, and the cord cut. It was necessary to perform the entire procedure rapidly because any delay meant increased mortality in the young. AJ] young, after drying, were placed in an incubator (37°C) on aslightly moistened towe1. No mention was made of whether any females survived the operation. Extraction of Bat Milk To determine some of the chemical and physical properties of bat milk, Huibregtse (l966:551) related atechnique for removing milk from lactating Leptonycteris sanborni and Tadarida brasiliensis as foHows: "The animals were lightly anesthetized with sodium pentobarbital (60 mg 20 mI) .... Adose of 0.03 to 0.10 ml of the anesthetic was found to be satisfactory. This was administered prior to an injection of oxytocic hormone (Pitochin, Parke-Davis) of less than 0.1 m1. The mammae were bathed with warm water, and the milk expressed manually with thumb and index finger. The extruded droplets were collected in a smalJ pipette on arubber tube (a hemocytometer pipette served well)." A small drop of 10 per cent formalin was added to preserve refrigerated samples. Bleeding Bats Basic extraction techniques have been devised to obtain blood from bats, and two are generally used--one from the heart, the other from the eye. Cardiac puncture is the technique most widely used. Disposable needles and syringes are recomnlended. The appropriate needle size depends on the size of the bat. For bats of average size, Sudia et al. (1970) recommended a25-guage, 3/8 to 5/8inch needle. A2-milliliter syringe is satisfactory. Asuitable supply of needles are heparinized by drawing aone per cent solution of heparin through them and then allowing the needles to air-dry. The puncture site should be cleaned and disinfected before inserting the needle into the heart. Some investigators prefer to take blood from the orbital sinus. Sudia et al. (1970) have described this method. The anesthetized bat is held firmly in the left hand, the thunlb exerting sufficient pressure just behind the eye to cause it to bulge slightly. Amicrosampling pipette (hematocrit) of either a50 or 100 microliters is inserted into the posterior orbit of the eye (carefu11y pushing the eyeball to one side to avoid damaging it) and gently rotated so that the capillaries are ruptured against the bone and thus initiate the flow of blood. Once the flow is started it is necessary to draw the tube back slightly and incline it downward. Usually the blood flows freely into the tube, at times so profusely that two tubes can be filled easily; sometimes, however, it is necessary to repeat the rotation a few times. The pipette is then discharged into atube containing ameasured volume of diluent. The hematocrit tube or microliter tube may be heparinized. A50-microliter pipette wil1 take up to 0.1 ml. of blood and a100-microliter pipette will take up to 0.2 ml. blood (the amount usually taken from each bat). Larger amounts of blood may be obtained by holding the bat between the thumb and first and second fingers to apply pressure to the thoracic area. Apparently, this raises the blood pressure and increases the yield of blood from the orbital sinus. Bats bled in this manner do not appear to be harmed seriously and such bleeding may be continued daily for up to 10 to 15 days if necessary. Saliva Collection Dickson and Green (1970:41-43) required the collection of saliva from Desmodus and Diaemus. These biologists developed an ingenious plastic box for the safe restraint of these bats. The vampires first are anesthetized as described under "Anesthesia." They then are placed in the salivation units so that pilocarpine can be administered to the buccal mucosa and the saliva collected as described in detail by the investigators. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Urine Collection Bladder catheterization has been the technique used to obtain urine samples from small animals. The usual techniques require either continuous anesthesia, restraint, or extensive surgery. Kanthor (1965) devised amethod of repeated and accurate urine collection from unrestrained Myotis lucifugus (body weight about 7grams). He wrote (p. 326): "We developed acatheter which could be passed through their extremely well-developed urethral spincter with aminimum of subsequent irritation. Moreover, it prevented leakage and retention of urine in the bladder, while ensuring free passage to permit collection of serial samples over extended periods. Females were animals of choice as the use of these catheters in males involved extensive surgery." The bats were anesthetized with ether. More than 50 bats tolerated the procedure and none tried to remove the catheter. The catheters permitted freedom of movement under all conditions and remained functional for up to three days. Operant Conditioning for Experiments Bats, unlike most mammals, cannot operate the conventional gadgetry normally used in experimental situations with either of their specialized limbs. Beecher (1971) demonstrated that Phyllostomus hastatus could operate aconventional pigeon key by either nosing or licking it. Schmidt and Greenhall (1972) found that Desmodus rotundus would respond quickly to training. Their bats were conditioned to feed at an observation table from I100 to 1300 hours. After adish of blood was placed on the table, the cage was opened and aconditioning noise was made by scraping two forceps together. Conditioning could take up to aweek, but, when bats were trained, they usually flew to the experimental table within five minutes after the noise in anticipation of ameal. After the bats were trained, avariety of small animals were introduced into the cage so that the investigators could observe and photograph the vampires as they stalked, attacked and fed on their prey. Thus, Schmidt and Greenhall had to wait approximately five minutes before they were able to observe the feeding behavior that otherwise might have required several hours. 118 Personnel Precautions All persons handling captive bats or in contact with them should be aware of the possible health risks in their routine work. Simple hygiene precautions should be observed, such as washing the hands after handling animals or immediately on leaving the animal facilities. In my laboratories, Ihave used wall-mounted dispensers containing either tincture of green soap, rubbing alcohol, or other general antiseptics. Laboratory coats should be worn to protect clothing. Handling bats with bare hands should be avoided if possible and leather gloves should be worn as aprotection against bites. Rubber or disposable plastic gloves should be used while performing operations and autopsies. All cuts and abrasions 119 BIOLOGY OF THE PHYLLOSTOMATIDAE Bat Brain Removal Bats that have bitten people should be killed and sent to adiagnostic laboratory for examination. For the purposes of accurate taxonomic identification, the head probably is the most important part of the animal to the mamnlalogist, whereas brain and other body tissues are of paramount importance to the epidemiologist. The conventional laboratory techniques used for brain tissue removal for rabies diagnosis frequently mutilates the skin and skull, not only making accurate identification difficult but often completely ruining the specimen for museum purposes. To solve this problem, Greenhall (1965a) devised methods for tissue removal, with little or no head and body damage, that proved satisfactory to both mammalogist and epidemiologist. In the case of extremely small bats, or those with specialized attachments between the ears or unusual glandular structures on the head, it is unnecessary to open the brain case because sufficient brain tissue may be hypodermically withdrawn through aneedle inserted into the foramen magnum without damage to the skull. HUMAN HEALTH PROBLEMS The World Health Organization (1973) reported that increasing numbers and kinds of animals are now used in biomedical studies. Zoologists should be aware that bats may carry diseases and, therefore, may be ahazard to human and animal health. Jones in his chapter on economics and conservation (this volume) has discussed the diseases occurring in wild phyllostomatids that may be transmitted to humans and livestock. Acomprehensive review of zoonoses and bats was presented by Constantine (1970). There is always the risk that biologists and technicians working with captive bats may contract an infection from the parasites, urine, feces, skin, blood, and other tissues from their own laboratory animals. Yunker (1964) has reviewed some of the common arthropod associates of laboratory animals that are hazardous to man. The public health importance of Neotropical bats has been discussed in detail by Greenhall (1964), Acha (1967), Chalmers and Scott (1969), and Tamsitt and Valdivieso (1970). Because little is known about the health of captive bats, any information on the causes of death in the laboratory is of value and autopsies should be performed when possible. 120 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY should be cleansed immediately and agood first aid kit should be easily accessible. Particular care should be given to eye protection and glasses or goggles should be worn when conducting postmortem dissection on suspected rabies cases. Disposable paper face masks serve as some protection against the inhalation of dust and spores and are recommended for persons suffering from allergies and respiratory ailments. Whether these masks are effective against infectious aerosols is not known. Strict adherence to aU laboratory safety rules should be compulsory for staff as well as visitors. Clinical Symptoms of Infections Many people working with captive animals are not aware of the clinical signals that may be indicative of an infection acquired through exposure to miscellaneous animal nlaterial. If the symptoms fail to be resolved quickly, medical advice should be obtained. Irvin et al. (1972) listed the following symptoms: 1) allergies such as asthmalike symptoms of running, itching, and burning eyes, and skin hypersensitivity and irritation; 2) skin infections; 3) respiratory symptoms, especial1y apersistent cough, sore throat, or running nose; 4) influenza-type symptoms; 5) local inflanlation and infection, expecially on the hand or exposed parts of the body; 6) swelling of lymph nodes; 7) generalized symptoms such as fever, headache, vertigo, diarrhea, nausea, vomiting, and malaise. Rabies Rabies is perhaps the greatest threat to the chiroptologist because any bat may contract rabies. Anumber of phyl1ostomatids (notably Phyllostomus, Glossophaga, Carollia, Artibeus, Desmodus, Diaemus, and Diphylla) have been found to be positive for rabies and these bats should be considered as potentially dangerous laboratory animals (Acha, )967). Dickson and Green (1970) enlphasized that vampires from areas that are endemic for rabies should be kept only in laboratories especially equipped for the protection of the staff. The precautions observed in their laboratory are as follows (pp. 37-38): "The bats are housed in aquarantine room, which may be entered only by persons immunised against rabies; gowns, masks and gloves must be worn at all times; al1 waste materials are sealed in sacks before removal from the animal room, and autoclaved prior to disposal; the brains of al1 bats which die are removed and examined for rabies virus." Rabies in alaboratory colony of vampire bats.- The unexpected appearance of rabies in alaboratory colony of vampire bats demonstrates the importance of quarantine measures and the practical value of using cages in which the bats are visible at all times. "The following incident is of interest because it was contrary to the usual pattern of behavior of vampires in captivity (Horst and Langworthy, (1972:903). "In January )970, 30 vampire bats (Desmodus rotundus murinus) were collected from ... Mexico. The acclimation of these bats to laboratory conditions followed the pattern described by Wimsatt and Guerriere (1961) and 20 bats survived this adjustment. "On 29 January 1971, 30 additional vampire bats were obtained in ... Mexico and added to the 15 remaining animals in the laboratory colony. After aweek of adjustment during which five of the new animals succumbed, the colony was stable and no unusual events occurred until the first week of April, two months later. Up to this time the bats, when disturbed, normally would bunch together, hiding in the darkest corner of the cage. However, beginning about 1April 1971, there was intense fighting at the slightest disruptive stimulus, such as switching on the lights, moving the cage, or sudden loud noises. These fights were so intense that the entangled pairs would fall ... onto the floor, still screaming and viciously biting each other." Mortality occurred at arate of about one bat per day for about two weeks. Those bats checked by the fluorescent antibody method were positive for rabies. Horst and Langworthy (1972:904) correctly recommended that "individuals who maintain these animals in captive colonies are well advised to take proper quarantine precautions with recently captured vampires, lest they suffer asimilar loss of valuable animals." Although Wimsatt and Guerriere (1961) described their routine care and maintenance of vampires, Icould not find one mention of the pattern of acclimation of bats referred to by Horst and Langworthy (1972:903). Nonbite rabies in laboratory animals and technicians.-Winkler et al. (1972, 1973) reported an unusual outbreak of nonbite transmitted rabies in alaboratory colony of wild carnivores and afatal case of nonbite rabies in alaboratory worker possibly caused by astrain of bat rabies virus. Sixty-four animals died, including 39 that had no known exposure history. The human victim had been vaccinated against rabies 13 years earlier, but had not developed demonstrable serum antibodies. Investigation confirmed that direct contact transmission did not occur and suggested that airborne bat virus may have been responsible. The human case emphasizes the necessity for biologists and technicians working with potentially rabid animals to be immunized, followed by verification of demonstrable serum antibodies. Preexposure immunization.-The best protection against rabies an individual can have is preexposure vaccination, as recommended by the United States Public Health Service (1974:16) and the World Health Organization (WHO) Expert Committee on Rabies (1973:30): Preexposure immunization consists of three injections of duck embryo vaccine (DEY) spaced over aperiod of several weeks, followed by abooster injection of vaccine one month later, and lastly by the confirmation of antibodies (that is, immunization) in the serum of the vaccinated individual. If negative, booster doses should be repeated until antibodies become demonstrable. Further booster injections should be given at intervals of one to three years as long as the person remains exposed. Some people have been reluctant to accept the preexposure immunication because of the misconception that the regimen required alarge number of daily injections. First aid treatment for bite wounds.-According to the WHO Expert Committee on Rabies (1973:28) and Kaplan (1973:15-16), the n10st important first aid treatment for all bite wounds and scratches in preventing possible rabies infecBIOLOGY OF THE PHYLLOSTOMATIDAE 121 ACKNOWLEDGMENTS I am sincerely grateful to the following colleagues for providing either manuscripts that are in press or permitting the inclusion of their unpublished obtion is the gentle washing and flushing of the wounds with soap and water, detergent, or water alone. Next apply either 40 to 70 per cent alcohol, or a 5 to 7per cent tincture or aqueous solution of iodine, or 0.1 per cent quaternary ammonium compounds, which can kill rabies virus on contact within one minute. Alcoholic beverages of 86 proof or greater can be used in emergencies. When soap has been used to clean wounds, all traces of it should be removed before the application of quaternary ammonium compounds because soap neutralizes their activity. The WHO Expert Committee on Rabies (1973:28) stated: HAlthough judicious use of concentrated nitric acid in puncture wounds has its advocates, there is no evidence that this product is more effective than quaternary ammonium compounds or 20%liquid soap solution." Postexposure treatment.- The United States Public Health Service (1974: 1) has recommended the following postexposure treatment against rabies. "If an immunized person is bitten by arabid animal [bat], the rabies virus stimulates rapid production of antibodies because the individual has already been sensitized by his preexposure vaccination. Therefore, an immunized person needs only 1to 6doses of DEV [Duck Embryo Vaccine] even after being bitten by a known rabid animal, instead of the regimen of antiserum and up to 23 doses of DEY recommended for an unimmunized person in the same situation. But, most importantly, aperson who has received pre-exposure vaccination and receives 6doses of vaccine after exposure is considered significantly better protected than someone who receives only the full post-exposure regimen of antiserum and vaccine." The physician attending abite exposure must decide whether antirabies treatment is indicated and, if so, must administer the most effective treatment available to him. If serum is indicated, the physician must first test for allergies and check the patient's history for allergenic reaction. At present the duck embryo vaccine (DEV-Eli Lilly &Co., Indianapolis, Indiana) is the most widely used in the United States. Newer and safer vaccines such as the Wistar Vaccine, are being investigated. Physicians are strongly urged to check for the most recent recommendations at either of the following two World Health Organization Rabies Reference Centers located in the United States: United States Public Health Service Center for Disease Control Bureau of Epidemiology, Viral Zoonoses Section Atlanta, Georgia 30333 Telephone (404) 633-3311, Extensions 3415 or 3683 After 5p.m. (404) 633-2176 or The Wistar Institute Rabies Division 36th Street at Spruce Philadelphia, Pennsylvania 19104 Telephone (215) EV 7-6700. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 122 servations: James G. Doherty, H. Bradford House, Donna J. Howell, Rexford D. Lord, John J. Rasweiler, IV, Uwe Schmidt, and Aldo M. Voute. Ialso extend my appreciation to Alfred L. Gardner for his valuable criticism. Iwish to thank the World Health Organization for granting permission to cite parts of the Sixth Report of the WHO Expert Committee on Rabies. LITERATURE CITED ACHA, P. N. 1967. Epidemiology of paralytic bovine rabies and bat rabies. Bol. Off. Int. Epizol., 67:343-382. ADAMS, D. B., AND G. M. BAER. 1966. Cesarian section and artificial feeding device for suckling bats. J. Mamm., 47:524-525. BARBOUR, R. W., AND W. H. DAVIS. 1969. Bats of America. Univ. Press Kentucky, Lexington, 286 pp. BEECHER, M. D. 1971. Operant conditioning in the bat Phyllostomus hastatus. J. Exper. Animal Behav., 16:219-223. BRADBURY, J. W. 1970. Target discrimination by the echolocating bat, Vampyrlll11 spectrum. Exp. Zool., 173:23-46. BUCKLAND-WRIGHT, J. C., AND J. D. PYE. 1973. Dietary deficiency in fruit bats. Internal. Zoo Yearbook, 13:271-277. BULLARD, R. W., AND S. A. SHUMAKE. 1973. Food temperature preference response of Desmodus rotundus. J. Mamm., 54:299-302. CHALMERS, A. W., AND G. R. SCOTT. 1969. Ecology of rabies. Trop. Animal Health Production, 1:33-55. CONSTANTINE, D. G. 1952. A program for maintaining the freetail bat in captivity. J. Mamm.,33:395-397. 1970. Bats in relation to health, welfare and economy of man. Pp. 319-449, in Biology of bats (W. A. Wimsatt, ed.), Academic Press, New York, 2:xv+ 1-477. CRANDALL, L. S. 1964. The care and management of wild animals in captivity. Univ. Chicago Press, Chicago, xv +761 pp. DAVIS, W. H., AND M. M. LUCKENS, 1966. Use of big brown bats (Eptesicus [uscus) in biomedical research. Lab. Animal Care, 16:224-227. DECOURSEY, G., AND P. G. DECOU~SEY. 1964. Adaptive aspects of activity rhythms in bats. BioI. Bull., 126:14-27. DICKSON, J. M., AND D. G. GREEN. 1970. The vampire bat (Desmodus rotundus): improved methods of laboratory care and handl ing. Laboratory Animals (London), 4:37-44. DITMARS, R. L. 1935. Collecting bats in Trinidad. Bull. New York Zool. Soc., 38:213218. 1936. A Vampyrum spectrum is born. Bull. New York Zool. Soc., 39:162-163. DITMARS, R. L., AND A. M. GREENHALL. 1935. The vampire bat. Zoologica, 19:53-76. DUNN, L. H. 1933. Observations on the carnivorous habits of the spear-nosed bat, PhyllostomushastatuspanamensisAlIen, in Panama. J. Mamm., 14:188-199. FLEMING, T. H., E. T. HOOPER, AND D. E. WILSON. 1972. Three Central American bat communities: structure, reproductive cycles, and movement patterns. Ecology, 53:555-569. FLORES CRESPO, R., R. J. BURNS, AND S. B. LINHART. 1971. Comportamiento del vampiro (Desmodus roTundus) durante su alimentaci6n en ganado bovino en cautiverio. Tecnica Pecuaria Mexico, 18:40-44. GATES, W. H. 1936. Keeping bats in captivity. J. Mamm., 17:269-273. 1938a. Raising the young of red bats on an artificial diet. J. Mamm., 19:461464. 1938h. On the keeping of bats in captivity. Proc. Louisiana Acad. Sci., 4:157158. 123 BIOLOGY OF THE PHYLLOSTOMATIDAE centimeters multivitamin drops (Vi-Penta); six teaspoons Marine Protein Concentrate; three teaspoons beef extract. Add water to make one quart and blend. Then add two quarts of water and stir. Amount sufficient for approximately 50 bats. 16. Bronx Zoo Diet for Phyllostomus hastatus House (1968: 141) used this formula to maintain Phyllostomlls hastatus: 50 per cent bananas; 30 per cent grapes; 20 per cent chopped meat; vitamin supplement; one teaspoon Mellin's Food (per day). The daily amount offered each bat was approximately 35 grams ad libitllm. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 13. Artificial Diet for Nectarivorous Bats This formula was provided by Donna J. Howell (personal correspondence): 10 eggs; nine tablespoons Brewer's yeast; nine tablespoons instant protein (Alpine Marine); one tablespoon bone meal; 18 capsules amino acids (Wolin's); 18 cubic centimeters multivitamin drops (Poly-Vite Wolin's); 18 ounces (2 1 Acups) condensed (not sweetened) milk; three pounds honey or strawberry jam; two small jars "high-meat diet" baby food (strained beef or chicken); three-fourths beef extract (Difco Culture Medium); 18 tablespoons Super Hydramin (Nion). Mix the ingredients well in blender, add six quarts water, and mix again. Makes one and a half gallons. The mixture is frozen in pint containers and used as needed. Each jar must be shaken very thoroughly before it is poured into bat feeders. The beefextract provides salt. 15. Bronx Zoo Diet for Frugivorous Phyllostomatid Bats House and Doherty (1975) used this diet for their frugivores: 24 bananas; one apple; onefourth pound grapes; 5ounces Zu/Preem Primate Diet (canned); four ounces Zu/Preem Feline Diet (canned); six tablespoons Mellin's Food; six tablespoons Super Hydramin. The daily quantity fed was approximately 25 to 30 grams per bat ad libitum. 14. Banana-based Did for Frllgivorolls Phyllostomatid Bats This diet was taken from Rasweiler (1976) and Rasweiler and Ishiyama (1973:59): 700 grams banana; 9.87 grams wheat germ (pulverized in acoffee mill); 12.90 grams whole milk powder; 24.00 grams calcium caseinate; 24.48 grams sugar; 4.04 grams protein (Gevral); 1.71 grams mineral supplement; 0.70 grams vitamin mix; 8.68 milliliters corn oil mixture. The bananas are cut into slices I-centimeter thick and gently mixed with the other ingredients. The slices are kept intact so that the bats can hold and carry away the pieces. An excess of food is provided. The daily amount offered each bat is approximately 32 grams for Artibells litllratlls and Phyllosotmlls discolor, and about 20 grams for Swrnira /ilium. 12. Artificial Diet for Nectarivorolls Phyllostomatid Bats This formula was given by Rasweiler and de Bonilla (1972:661) and Rasweiler (1973: 394): 700 milliliters fruit base (peach nectar or guava nectar); 29.52 grams cereal (high protein); 4.92 grams wheat germ; 12.81 grams milk powder; 20.39 grams calcium caseinate; 68.00 grams sugar; 3.99 grams protein (Gevral); 1.68 grams mineral supplement; 0.69 grams vitamin mix; 9.00 milliliters corn oil mixture; 300 milliliters water. To facilitate suspension in the liquid diet, the cereal and wheat germ are reduced to a fine powder by passage through acoffee mill prior to their mixture with the other dry components. The dry mixture can be stored at 4°C until use. The composition of the corn oil mixture (polyoxyethylene sorbitan mono-oleate as an emulsifier and isopentylacetate as a flavoring agent) can be stored at 4°C until use. The final diet consisting of peach nectar, the powdered premix, and the corn oil mixture is prepared freshly each day in an electric blender and served to the bats in shallow dishes. The amount offered by Rasweiler and de Bonilla was approximately 24 milliliters per bat per day. 130 BIOLOGY OF THE PHYLLOSTOMATIOAE 131 J 7. Diet for Frugivorous and Omnivorous Plzyl/ostomatid Bars Donna J. Howell (personal communication) recommended this formula for maintaining frugivorous and omnivorous phyllostomatids: 10 bananas (very ripe, chopped); three to four cups melon pieces (cantaloupe or honeydew); eight ounces cottage cheese: one pound hamburger or one can dog food; one-fourth tube beef extract; one-half Pervinal powder (available in pet stores); two eggs. This should be tossed to mix. or else the first bat at the dish will eat all the choice pieces. However, it should not be mixed into amush or paste because bats will get the mush on themselves in their enthusiasm and competition for the food. If bats are obviously pregnant, condensed milk and bone meal should be added. The formula has proved satisfactory for Phyllostomus hastatus, P. discolor, Carol/hi, Artibeus, and Sturnira. 18. Products Mentioned in the Text* Beef Extract-Difco Laboratories, Detroit, Michigan (Difco Culture Medium); GevralProtein--Lederle Laboratories, Pearl River, New York; Instant Protein--Alpine Marine Industries, New Bedford, Massachusetts 02742; Jeculin--Upjohn Company, Kalamazoo. Michigan 49001 (liver extract and iron); Lady Clairol Protinator-Clairol Incorporated. Stamford, Connecticut; Lady Clairol Cremenogenized Hair Lightener-CI airol Incorporated, Stamford, Connecticut; Marine Protein Concentrate--Alpine Marine Industries, New Bedford, Massachusetts 02742; Mellin's Food-Consolidated Royal Chemical Company, Chicago, Illinois 60610 (maltose-dextrin mixture with added thiamin ferric glycerophosphate and potassium bicarbonate); Ostermilk--Glaxo laboratories, Ltd., Greenford, Middlesex. England (reconstituted milk); Pet Drops--Upjohn Company. Kalamazoo, Michigan 49001 (multivitamins); Poly Vite Drops--Wolins Company, Farmingdale, New York (multivitamins); Stuart Formula Liquid-Stuart Company, Pasadena, California (multivitamins); Super Hydramin Powder-N ion Corporation. Los Angeles, California 90038; Tego Solution--Hough Hoseanson Ltd., Chapel Street, Manchester J 9, England; Theragran--E. R. Squibb and Sons, New York, New York 10022 (multivitamins); ViPenta-Roche Laboratories, Nutley, New Jersey 07110; Zu/Preem Diets--Riviana Food. Inc., Hills Division, Topeka, Kansas 66601. *Mention of trade names or products does not imply endorsement by the U.S. Government. ECONOMICS AND CONSERVATION CLYDE JONES Historically, bats have been considered by man as objects of mystery, superstition, fear, and basically as indicators of some unknown or evil significance. The role of bats in the writings of early naturalists, as subjects of early artists, as objects of superstition and even worship, and as ingredients of concoctions for varied purposes was summarized in detail by G. M. Allen (1939), who prepared the first general summary on the biology of bats. In modern times, there has been an increasing awareness of bats for several reasons, mostly because of the development of tools with which to obtain numerous species for study, as well as an increase in knowledge of some diseases of bats important to man. As aresult of these and other factors, there has been a great surge of interest and activity in various studies of bats during the past two decades. A few of the major general works on bats within this period of time include the summaries provided by Brosset (1966), Barbour and Davis (1969), Leen and Novick (1969), Slaughter and Walton (1970), and Wimsatt (1970). As an additional indication of current interest in studies of bats, four international conferences have been held since 1968, and annual conferences have been held in the United States since 1970. Newsletters about bats have appeared during recent years in Australia, Europe, and the United States. Adetailed review of the proliferation of literature on Chiroptera in the past few years was provided by Anderson and Van Gelder (1970). In spite of the aforementioned interests and activities, much remains unknown with regard to the biology of bats. It seems that certain groups, such as the Phyllostomatidae and some other tropical taxa, as well as special topics, such as the status of populations and the need for conservation' have been relatively neglected. This report attempts to summarize and discuss some of the broad problems concerning the economic importance and conservation of the Phyllostomatidae, and to review briefly some of the specific needs for conservation of these bats. PROCEDURES It is not within the scope of this report to present acomplete summary of literature containing information and notes on the aspects of the biology of phyllostomatids. References are provided that contain either additional information or examples pertinent to the purposes of this paper. Information was obtained from the literature, from newsletters and various reports on bats, and from correspondence with researchers who either worked on phyllostomatids or conducted field studies in areas where these bats occur. Data were taken also from the files of the bat-banding program of the National Fish and Wildlife Laboratory. The scientific names used herein are in accordance with the nomenclature provided by Jones and Carter (this volume). 133 134 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY ECONOMICS It seems impossible to attempt to determine meaningful economic values with regard to most bats; at least, no precise determinations will be made herein. However, it is relevant to review briefly the role of bats within ecosystems, especially with regard to relationships between phyllostomatid bats and certain things that seem important to man. Relationships with Plants Bats playa role in the natural dispersal of plants. Members of the family Phyllostomatidae are the principle agents of chiropterochory in the New World tropics. In general, most dissemination of plants by bats results from seeds that are dropped from the mouths of animals either in flight or in roosts. Relatively few plant seeds are passed through the digestive tracts of bats, although seeds of some plants, such as Ficus, probably are scattered in this manner. Chiropterochory is highly developed in certain families of plants, especially the Moraceae, Palmae, Anacardiaceae, Sapotaceae, and Meliaceae. However, this condition also is present in many other families. For lists of chiropterochorous plants, characteristics of bat fruits, and discussions of the syndrome of dispersal of plants by bats, see the works by Van der Pijl (1957, 1968, 1969, and others). Sonle specific examples of synzooic relationships between plants and Phyllostomus discolor, P. hastatus, Artibeus jamaicensis, A. lituratus, and Carollia perspicillata are given by Van der Pijl (1957) and Greenhall (1956, 1965, 1966). Phyllostomatids are the agents of chiropterophily in the Neotropical region. According to Van der Pijl (1969), bat flowers occur mostly in the genera Musa, Parkia, Sonneratia, Agave, and Carnegiea, but are present also in various stages of evolutionary development in some other taxa. Flowers pollinated by bats show some specialized traits; they have an abundance of nectar and pollen and tend to open at night. There is some evidence that pollen of chiropterophilous plants have more amino acids than do related species pollinated in other ways (Howell, 1970). Because flower bats rely on flowers for asupply of protein in the form of pollen, mutualistic adaptations are exhibited by some pollinating bats. For example, projections on the cuticular scales of hairs of some glossophagines seemingly are correlated with chiropterophily (Howell, 1971). Other morphological adaptations related to feeding and flight, as well as seasonal movements of bats that coincide with flowering of certain plants, are better known than the aforementioned example. There is no doubt that phyllostomatid bats are important as agents of dispersal of seeds and pollen, at least in alimited range. Some botanists are of the opinion that, among mammals, bats are the most important dispersers of seeds (Van der Pijl, 1957). Recent analyses of trophic roles of bats demonstrate the richness of the Neotropical region with regard to frugivory and nectarivory of the chiropteran fauna (Wilson, 1973). There are similarities between the distributions of certain phyllostomatids and chiropterochorous and chiropterophilous plants; in fact, the ranges of some plants may result from the actions of bats on the re- BIOLOGY OF THE PHYLLOSTOMATIDAE 135 production of the plants. Numerous plants with typical bat flowers and bat fruits are economically important to man either as sources of food or for ornamental purposes. Relationships with Insects Phyllostomatid bats are associated ecologically with an array of insects. Although usually referred to as fruit bats or fruit-eating bats, members of this family consume considerable quantities of insects. For example, 33 of 143 stomachs from atotal of 11 taxa of phyllostomatids examined by Arata ef al. (1967) contained insect materials. Information provided by Wilson (1973) revealed the relative importance of insects in the diets of the genera of phyllostomatids. Wilson also indicated the need for additional information on the food habits of bats. Few data are available for evaluating the role of phyllostomatids with regard to the consumption of insects within an ecosystem. For some limited information on the impact of large colonies of insect-eating bats on insect populations, see the articles by Cockrum (1969, 1970). Although almost impossible to quantify on the basis of current knowledge, it is obvious that the predator-prey relationships between phyllostomatid bats and insects are important. Relationships with Man Phyllostomatid bats are considered occasionally as pests of fruit trees. The possibilities of these bats damaging fruit trees were implied in the discussion of the relationships between bats and plant dispersal and reproduction. There are, however, few reports of serious damage to fruit trees by bats in the New World tropics. Some data on disturbances to crop plants by phyllostomatids were presented by GreenhalJ (1956, 1966). All of the available data indicate that whatever harm phyllostomatids do to the fruit industry is of little or no consequence, except for some isolated instances where only limited damage is done. Phyllostomatids create some annoyances for man because of fruit consumption in gardens and homes and corresponding deposition of fruit pulp and other dropped materials. Phyllostomatids also create anuisance by roosting in man-made structures. Bats commonly occur in attics, walls, and between layers of thatch of buildings throughout the Neotropical area. People usually object to noises made by either vocalizations or movements of the animals, odors and stains from urine and fecal materials, and rejected food particles that frequently accumulate below roosting sites. In addition, ammonia gas may accumulate in guano deposits and cause unpleasantness to man. For an indication of the relationship between ammonia and Macrotus californicus, see the report by Mitchell (1963). Human food may become contaminated by droppings from bats that inhabit buildings or from insects and arthropods that live in guano deposits. Guano deposited by bats has been used as fertilizer, especially in the early stages of the development of the fertilizer industry. Except for some figures for guano production and mining in northern Mexico and the southwestern United States (Hutchinson, 1950), good data on modern production and use are not 136 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY TABLE I.-Occurrence of bacterial diseases in phyllostomatids. Bacteria Disease Host species Country Salmonella Salmonellosis Glossophaga soricina Panama Artibeus litllratus Colombia Sturnira lilium Colombia Bartonella and Bartonellos is Carollia perspicillatll Brazil Grahamella Desmodus rotundus Peru available. Guano is still mined, especially in some areas in Mexico, and some of the larger caves apparently yield worthwhile amounts. However, the importance of bat guano as fertilizer is not what it was previously because of the depletion of supplies and sources, as well as the development of other commercial fertilizers in recent years. The role of phyllostomatids as producers of guano in sufficient quantities for commercial exploitation is poorly known. Members of the Phyllostomatidae also are considered pests because they harbor ectoparasites that cause some concern to humans. However, no further discussion is warranted herein because this topic is dealt with in detail by Webb and Loomis (this volume). Phyllostomatids, like other bats, are often considered as pests by man in connection with diseases that may be carried by the animals and possibly transmitted to man. Alist of the bacterial, mycotic, and protozoan diseases known to occur in wild phyllostomatids is presented in Tables 1to 3. Known occurrences of viruses in these bats are summarized in Tables 4and 5. Most of these data are from Constantine (1970), who also gave detailed discussions of each disease and virus, including information on how these diseases are manifested in humans. For additional information on pathogens in the Neotropical area, with special emphasis on Puerto Rico, see the paper by Tamsitt and Valdivieso (1970). The reported occurrences of rabies virus infections in wild phyllostomatids are summarized in Table 5. Rabies in vampire bats is more common than in all other members of the family. According to Constantine (1970), reported deaths of humans from rabies transmitted by vampire bats are relatively insignificant causes of human mortality (Table 6). However, rabies transmitted by vampire bats to livestock is an important concern because this disease is amajor cause of mortality in cattle in Latin America. Rabies in cattle has been reported in all Latin American countries except Chile and Uruguay. Estimates of annual cattle mortality vary considerably. For example, Constantine (] 970) provided asummary of data on estimated cattle losses that totaled half amillion head (47.6 million dollars) in ]966 and two million head (100 million dollars) in 1969. The magnitude of this problem is reflected in the rash of investigations in recent years concerned with the control of either rabies or vampire bats. "Vaccination has been the most effectively applied method of combatting the problem in livestock" (Constantine, ]970). Some methods for controlling vampire bats have resulted from studies of certain biological aspects of these animals in programs operated by the Food and Agriculture Organization of the United Nations and the BIOLOGY OF THE PHYLLOSTOMATIDAE TABLE 2.-0ccllrrence of mycoTic diseases in phyllostonwtids. 137 Fungus Histoplasma Blastomyces Scvplliariopsis Cryptococcus Candida Torulopsis TrichophYTon, Microsporum, Tricho~poron Disease Histoplasmosis Blastomycosis Scopulariopsos is Cryptococcosis Candidiasis Torulopsosis Superficial mycoses Host species Leptonycteris sanborni Desmodus rOfllndus Phyllostomus discolor Artibeusjamaicensis Carollia perspicillata Glvssophaga soricina Lvnchophylla rvhusta Lonchorhina aurita Micronycleris megalotis Tonatia bidens Phyllostomlls hastatus Anvura geoffroyi Artibeus lituratus Glvssophaga soricina Caro" ia perspicillata Artibeus lilUratus Desmodus rotundus Carvllia perspicillata Desmodus rotundus Leplonycteris sanbvrni Carollia perspicillata Desmodus rotundus LeptonYCleris sanborni LeptonycTeris sanborni Glvssophaga soricina Country United States Mexico, Panama, Colombia, Trinidad El Salvador, Panama El Salvador, Panama Panama, Colombia, Trinidad Panama, Colombia. Trinidad Panama Panama Panama Panama Trinidad Trinidad Colombia M~xico Mexico Mexico Mexico Colombia Colombia United States Colombia Colombia United States United States Colombia Denver Wildlife Research Center of the United States Fish and Wildlife Service. The attempt to reduce rabies in cattle by the control ofvampire bats is acomplex and sometimes controversial issue. For additional information, consult the reports by Constantine (1970), Schmidt et ale (1970), Rosenthal (1972), Yosti et al. (1971); Mendez (1971), Greenhall (1971, 1972), Tamsitt and Valdivieso (1970), Thompson et al. (1972), and Turner (1975). Bats are extremely useful to man as objects of research. Some indications of the increasing awareness of this importance are implicit in the introductory remarks with regard to the proliferation of literature on these mammals in recent years. Bats are used in several aspects of medical research, especially vaccine development, epidemiological studies, mechanisms of disease resistance, aging, and thermoregulation. Transillumination of bat wings is aconvenient way to make gross and microscopic observations of natural and experimental physiological phenomena, as weI) as for pathological studies and culturing of organisms in animal tissues. 138 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY TABLE 3.-0ccllrrence of protozoan diseases in phyllostomatids. Protozoa Disease Hosl species Country Bats are important in space biology for studies dealing with awide array of tolerances of environmental extremes and stresses. They also are useful in numerous investigations of aerodynamics and related topics. Studies in bat echolocation are useful for many purposes, as documented in some detail by Griffin (1958). Some interesting and sophisticated studies of ultrasonic orientation in bats have been conducted in recent years. For some exTABLE 4.-0ccurrence of viruses in phyllostomatids. Host species Country Colombia Panama Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Trinidad Trinidad Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Colombia Colombia Colombia Colombia. Mexico Colombia Colombia Colombia Carollia perspicillata Phyllostomus discolor Phyllostomus hastatus Desmodlls rotundus Glossophaga soricina Artibeus lituratus Artibells lituratus Group AArboviruses Artibellsjamaice/lsis Carol/ia perspicil/llta Artibeus lituratus Carol/ia perspicil/a({[ Artibellsjamaicensis Group BArboviruses Rhinophyl/a pumilio Glossophaga soricina Miscellaneous Arboviruses Artibeus lituratlls Artibeus lituratus Artibeus jamaicensi,\ Caroll ia perspicil/ata Artibeus lituraflls Artibeus liturlltliS Artihellsjll/l1llil'ensi.\ Art ibe liS flliginosiis Artibeus liWratlls Arfibeus jamaicensis Artibells lituratus Artibells jamaicensis Phyllostomus hastlltus Vampyrops hel/eri Artibellsjamaicensis Toxoplasmosis Trypanosomiasis Toxoplasma Trypanosoma Eastern equine encephalitis Venezuelan equine encephalitis Mucambo / Virus Itaporanga Carapara Jurona Utinga Tacaribe Saint Louis encephalitis Yellow fever Catu Tacaiuma BIOLOGY OF THE PHYLLOSTOMATIDAE TABLE 5.-0ccurrence of rabies in phyllostomatids. 139 Host species Macrotus californicus Phyllostomus discolor Phyllostomus hastatus Glossophaga soricina Leptonycteris nivalis Carollia perspicillata Uroderma bilobatum Artibeus sp. Artibeusjamaicensis Artibeus lituratus Desmodlls rotllndus Diaemus youngii Diphylla ecalldata Country United States. Mexico British Honduras. Guatemala Brazil Mexico Mexico Trinidad. Colombia Panama Mexico Panama, Trinidad, Brazil British Honduras, Guatemala, Mexico, Trinidad, Brazil British Honduras. Guatemala. Mexico. Trinidad. Brazil Trinidad Brazil amples, note the studies by Pollak et ale (1972), Simmons (1970), Simmons and Howell (1971), Howell and Pylka (1972), and Simmons et al. (1972). Bats may serve as important bioholographic models in the future (Greguss, 1968). In addition to the aforementioned usefulness in research, bats are important research objects for other biologists. For example, there still is need for, and interest in, basic studies of ecology, life history, distribution, morphology, and the like. The taxonomy of bats is still an intriguing topic, and considerable interest has developed in studies of their behavior and population dynamics. This may become amore active research area in the future, especially with regard to studies of mechanisms permitting bats to survive in extremes of population congestion. CONSERVATION Needs for conservation of American bats in general were recognized by Manville (1962), Davis (1967), and Cockrum (1969, 1970), and presented in their pleas for conservation. Additional comments on needs for bat conservation were presented by Barbour and Davis (1969), Gould (1970), Greenhall (1973), and Findley (1973). Phyllostomatids, as well as some other bats, have been given TABLE 6.-Some reported incidences of rabies transmitted to humans by vampire bats. Deaths Years Country 89 1925-1937 Trinidad 31 1951-196] Mexico 17 1953-1961 Guyana ]1960 Bolivia 81960 Brazil 51965 Argentina 140 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY TABLE 7.-1njormation on phyllostomlltidsjrom surveys oj researchers. Country Dates Genus Status of population some attention in the popular and conservation oriented works by Rood (1971 ), Curry-Lindahl (1972), and Novick and Dale (1973). Awareness and concern for bat conservation has been expressed by numerous spelunkers and the National Speleological Society. For additional information and recommendations of th~s organization, see the paper by Mohr (1972). Some concerns for bat populations in the New World have been expressed by bat researchers. For example, resolutions were developed during the Third Symposium on Bat Research, held in San Diego, California, on 24-25 November 1972 with some guidelines for bat conservation. These actions were based in part on arecent survey that revealed reduced populations of 22 species of bats in the United States, including three species of phyllostomatids (Jones, 1971). In connection with the aforementioned survey, limited data were accumulated on the population status of phyllostomatids at several places in Latin America. Of more than 100 requests for information sent to bat researchers, only 16 of those returned included information on phyllostomatids. Pertinent information from these replies is summarized in Table 7. All reports on phyllostomatids in the United States indicate declining populations; reports for other areas mostly reveal increasing populations, except for one report of decreasing populations and two reports of stable populations. However, most respondents stated that populations of Desmodus were actually reduced or absent in local areas due to eradication or control measures. For awealth of information on populations of vampire bats, see the work by Turner (1975). AU reports included lists of reasons for population changes. Increased populations of bats were associated with increased cultivation of fruit crops and the Declining Declining Declining Declining Declining Increasing Increasing Increasing Increasing Increasing Increasing Decreasing Stable Increasing Stable Increasing Desmodlls Leptonycteris, Choeronycteris Leptonycteris Leptonycteris, Choeronycteris Macrotus Mlicrotu.\ Glossophllga Desmodlls Desmod/l.\ Desmodlls Desmodlls, Phyllostomlls, Artibells, VlImpyrops, Stllrnira, Uroderma, Glossophaga Glossophaga, Carollia Desmodlls Artibells, Carol/ill, Phyl/ostomlls Glossoplwga 1966-1967 1963-1970 1963-] 970 J965-1970 1969-1970 ]953-1963 1965-1971 1967-1971 1962-1969 1960-1970 1965-]970 1961-] 965 J 96] -1971 1968-1971 1969-197] 1957-1971 Costa Rica Colombia Colombia Colombia Argentina Trinidad United States United States United States United States United States Mexico Mexico Mexico Mexico Costa Rica External Internal Nature of study SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY X X X X X X X X x X X X X X X X X X X X X X X X X X X X X X X X X X X X X x X X X X X X X X X X X X X X X X X X X X X X X X X X X X TABLE I.-Brains of phyl!osTomaTid species sTudied. 148 Species Phyllostomatinae LoncllOrhilla aurita Tomes Macrophyl!u11l l1U1crophyl!um (Schinz) MacroTus californicus Baird MicrollycTeris hirsl/Ta (Peters) MicronycTeris mega/oTis (Gray) MicronycTeris minllfa (Gervais) MicronycTeris nicefori Sanborn MicrollVCTeris schmidTorum Sanborn Mimoll'crellll/atllm (E. Geoffroy St.-Hilaire) Phylloderma stellops Peters Phyllostom liS disc%r (W,agner) Phyl!ostomlls e10ngatlls (E. Geoffroy St.-Hilaire) Phyl/oswml/s hastmlls (Pallas) TOllatia hidens (Spix) TOllaTia nicaragl/lle Goodwin Trachops cirr/lOslls (Spix) Vampyrllm spectrum (Linnaeus) Glossophaginae AI/oura geoffroyi Gray ChoeronisclIs godmalli (Thomas) ChoeronisclIs intermedills (Allen and Chapman) C/1OcrollycTeris mexiClInli Tschudi G/ossop/wgll a/tico/a Davis G/ossophaga commissarisi Gardner G/ossop/lliga soricina (Pallas) Hy/ollycteris IInderwoodi Thomas LepTonycteris .\(lIlhOrlli Hoffmeister Lic!1onycteris ohscura Thomas Lonchophyl!lI rohllsfa Miller MOIlOphyllll.\ red mlllli Leach Carol Ii inae Carol! ia perspicillaTa (Linnaeus) Carol/ill sl/hrllfa (Hahn) Rhinophylla pllmi/io Peters Stenoderminae Amctrida celltllrio Gray ArTiheus (/~TCCUS Andersen Artiheus cincreus (Gervais) ArTiheus hirsl/TIIS Andersen Artiheus inopill({[lIS Davis and Carter ArTihellsjamaiccnsis Leach Artihells /iTl/rmliS (Olfers) Artiheus p/llIeotis (Miller) Artihi'us To/tecu~' (Saussure) Artihells Iratsoni Thomas Cellturio senex Gray Chiroderma sa/vini Dobson Chiroderma 1.'il/os//m Peters BIOLOGY OF THE PHYLLOSTOMATIDAE TABLE I.-Continued. Ectophy//a l7U1cconne//i Thomas Enchisthenes hartii (Thomas) Stenoderma rLljwn Desmarest Stllrnira lilillm (E. Geoffroy St.-Hilaire) Sfllrnira Illdovici Anthony Stllrnira mordax (Goodwin) Uroderma hilohatllm Peters Uroderl7ll1 magnirostrllm Davis Vampyressa nymphaea Thomas VlImpyressa pllsil/a (Wagner) Val11pyrodes caraccioloi (Thomas) Vampyrops dorsalis Thomas Vampyrops he//eri Peters Val11pyrops injilsclis Peters Vampyrops vi1tatlls (Peters) Phyllonycterinae Brachyphy//a cavernarllm Gray Erophy//a homhifrons (Miller) Desmodontinae Desl110dllS rvtllndliS (E. Geoffroy St.-Hilaire) Diaemlls yOlillgii (Jentink) Diphy//a ecalldaf({ Spix x X X X X X X X X X X X X X X x X X X X x X X X x X 149 METHODS AND MATERIALS Bats were mist-netted from naturaJ populations or were collected by hand from roosting sites. Brains were prepared by removing the head immediately after the specimen was killed and chipping away the parietal region of the skull case to expose the brain. Fixation with 10 per cent formalin was allowed to proceed for several weeks. To facilitate handling, fixed brains were stored in 70 per cent ethanol. Afew species are represented by specimens preserved originally in alcohol that were collected by other workers. Brains from the latter often showed varying degrees of internaJ deterioration, and in some cases their value was limited to external study only. The skull and meninges were removed carefully from each brain. Then, the available series of brains from each species was examined to determine subjectively a"typical" specimen for each species. These brains were photographed from dorsal, ventral, and lateral views. Such photographs served as aconvenient basis for comparison; however, final judgments were made from observations on the specimen. Certain species were selected for detailed examination of internal anatomy. For these histological preparations, species were selected that were: 1) representative of each major type of external anatomy; 2) representative of each major taxonomic unit; and 3) of questionable phylogenetic position (such as Brachyphylla, which has been assigned by past workers to one of several different subfamilies). Brains for histological study were infiltrated with agum arabic solution (Humason, 1967) and frozen sections were made in the cross-sectional plane at 18 microns. Every second section (third section on some large species) was collected and mounted in step fashion on microscope slides. Sections were stained with "LuxoJ" fast blue MBSN (Matheson Coleman and Bell) to demonstrate myelinated areas and counterstained with cresyl violet acetate (Matheson Coleman and Bell) to outline concentrations of cells in discrete nuclei. Staining was by amodification of techniques described by Drury and Wallington (1967). Specifics for preparation of sections was as follows: 1) sections were dehydrated in 95 per cent ethanol; 2) stained in Luxol fast blue (see McDaniel, 1973, appendix A) for four to eight hours, at 50 degrees Centigrade; 3) hydrated through an ethanol gradient to distilled water; 4) dipped into 0.05 per cent lithium carbonate at one to three degrees Centigrade for 15 to 20 seconds (solution must be freshly prepared daily); 5) differentiated in 70 per cent ethanol until there was aclear distinction between myelinated (blue) and nonmyelinated tissues (clear); 6) hydrated through an ethanol gradient to distilled water; 7) counterstained in cresyl violet acetate (see McDaniel, 1973 appendix A) for five. to 10 minutes; 8) counterstain differentiated in 95 per cent ethanol to which afew drops of acetic acid had been added; 9) sections dehydrated in 100 per cent ethanol, cleared in xylene, and mounted in Permount. Steps 4and 5may be repeated if necessary. This staining procedure yields sections with deeply blue-stained myelinated areas (ranging from large tracts to individual axons) and purplish-red cell bodies embedded in apale violet matrix. EXTERNAL MORPHOLOGY OF THE BRAIN General Description The cerebral hemispheres of phyllostomatid bats are relatively smooth and without convolutions (Fig. 1). A few depressions, which are usually shallow, are found in most species. In the past, these were conveniently regarded as true sulci, but more recently, they have been termed fossae or fissurelike depressions resulting from the presence of adjacent osseous and vascular features (Schneider, 1957; Henson, 1970). My comparative histological examination of aseries of phyllostomatid brains (McDaniel, 1973) revealed that in some cases these depressions are not shallow, and apparently represent more than aconformity of brain to skull (for an extreme example, see the cingulate sulcus of Phyllostomus hastatus). There are no cortical maps published for chiropteran brains; therefore, there are no data concerning functional interpretation of the sulci or pseudosulci, and gyri or pseudogyri. In the absence of afunctional interpretation, the following terminology is based on morphological similarity rather than on actual, or even assumed, homology with brains of higher mammals. Homology within the family Phyllostomatidae is assumed, and hopefully will be substantiated experimentally in the future. Within the Phyllostomatidae, three sulci are consistently well developed: the interhemispheric sulcus, which separates the right and left cerebral hemispheres; the anterior rhinal fissure, which separates the olfactory bulbs from the main mass of the cerebrum; and the hippocampal sulcus, which is ventrally located. Of the remaining sulci, the most consistently present is one that divides the cereSPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 150 BIOLOGY OF THE PHYLLOSTOMATIDAE ] 51 bral surface into anterior and posterior portions. This fissure is similar to, but not homologous with, the central sulcus (Sylvian sulcus) of higher mammals. In this paper it will be referred to as the pseudocentral sulcus. In some cases, another fissure is developed somewhat anterior to the pseudocentral sulcus. The various lobes of the cerebral cortex are relatively unobtrusive and in most cases do not show atendency to bulge in the manner typical of higher mammals. All phyllostomatids have well-developed olfactory bulbs on the rostral end of the cerebral hemispheres. In addition, caudad to the pseudocentral sulcus, apseudotemporal lobe projects ventrally or somewhat posteriorly from the body of the cerebrum. The diencephalon (Fig. 1) is exposed only along its ventral surface. Aportion of the base of the thalamus is exposed anterior to the optic chiasma, and the hypothalamus is exposed posterior to the optic chiasma. The cerebellum (Fig. 1) occurs as adorsal foliated body posterior to the cerebral hemispheres. The cerebellar surface is relatively simple, and rarely has more than primitive sulci developed. The cerebellar body is composed of amedial vermiform body flanked by apair of lateral lobes. As in other mammals, the cerebellum is attached to the brain stem by the inferior, medial, and superior peduncles. The mesencephalon of phyllostomatid bats is rarely seen unless the hypophysis has been removed. The cerebral peduncles are never visible externally, and of the tectal structures, the enlarged inferior colliculi are generally the only structures visible (but even they are not always so). The pons is almost completely covered ventrally by the hypophysis and laterally by the trigeminal nerve. The medulla oblongata (Fig. 1) occurs as arather broad structure in phyllostomatids. Ventrally, the anterior trapezoid body, posterior olives, and medially located pyramidal decussation are visible. As in other vertebrates, the medulla grades posteriorly into the spinal cord. Aspects of Variation in External Anatomy Within the Phyllostomatidae, asurprising array of variation in features of external brain anatomy is encountered. Data concerning inter and intrasubfamilial variation are available for 65 species representing 38 genera and six subfamilies (McDaniel, 1973). Subfamily Phyllostomatinae Data are available for 10 of the 11 genera listed in this subfamily by Jones and Carter in the systematic account in this volume. Pronounced variation of several morphological features creates difficulty in characterizing ageneralized brain for this subfamily. The brain of Mimon (Fig. 2) is characterized by the presence of extremely short, deep cerebral hemispheres, pseudotemporallobes that project ventrally in a rounded rather than angular fashion, and aslight indication of apseudocentral sulcus. The caudal termination of the cerebral henlispheres is dorsally anterior to the inferior colliculi (which are contiguous to one another), resulting in dorsal exposure of precollicular tectunl. Mimon is the only phyllostomatid bat in which the tectum is broadly exposed anterior to the inferior colliculi. The cerebellum of Mimon is simple, having shallow foliations. The brain of Lonchorhina (not figured) resembles that of Mimon in having a short, deep cerebrum and asimple cerebellum. In Lonchorhina, the pseudocentral sulci cut more deeply into the cerebral hemispheres than in Mimon, and the posterior margin of the cerebrum extends almost to the anterior margin of the inferior colliculi. The brains of five species of Micronycteris (Figs. 3-7) vary only slightly among species. The cerebrum of Micronycteris is relatively longer than that of Mimon. The cerebrum is shortest in M. nicefori (Fig. 3) and M. minuta (Fig. 4), longer in M. schmidtorum (Fig. 5) and M. megalotis (Fig. 6), and longest in M. hirsuta (Fig. 7). The cerebral pseudocentral sulci are well developed in all species except M. nicefori. There is shallow development of asulcus anterior to the pseudocentral sulcus. The cerebellum is simple in all species. Consistently within the genus Micronycteris, but in no other phyllostomatine genus, the inferior colliculi are exposed dorsally and are not contiguous dorsally with one another. In Micronycteris, the inferior colliculi are separated by the anterior lobe of the vermiform body of the cerebellum. The brain of Macrotus (Fig. 8) resembles that of Micronycteris megalotis in most details. However, the dorsally exposed inferior colliculi are contiguous. The degree of contiguity is reduced and approaches the condition found in Micronycteris because the anterior lobe of the cerebellar vermiform body projects anteriorly to cover the most posterior portion of the inferior colliculi. In Macrophyllum (Fig. 9), the cerebral hemispheres are elongate relative to the condition in Mimon, and are exceptionally smooth, with only ashallow interhemispheric sulcus. The inferior colliculi are exposed dorsally and are contiguous with one another. The cerebellar tissue is exceptionally nondescript and lightly fissured. The brains of Trachops (Fig. 10) and two species of Tonatia (Figs. 11-12) are similar in external anatomy. The brain of Tonatia nicaraguae (Fig. 11) is the smallest and least ornamented of the three. It is characterized by the presence of deep cerebral hemispheres that are relatively longer than those of Macrophylfum. The pseudocentral sulci of the cerebrum are well developed, and there is shallow development of the sulci anterior to the pseudocentral sulci. The inferior colliculi are exposed dorsally, and are contiguous middorsally. As in Macrotus, the anterior edge of the vermiform body of the cerebellum protrudes forward to cover the posterior portions of the inferior colliculi. The cerebellum is simple in appearance. The brain of Tonatia bidens (Fig. 12) is similar to that of T. nicaraguae except that the cerebral hemispheres are relatively longer and the pseudocentral sulci of the cerebrum are extremely deep. The brain of Trachops (Fig. 10) differs from that of Tonatia bidens by the presence of shallower pseudocentral sulci, but somewhat deeper sulci anterior to the pseudocentral sulci. The brain of Trachops has some secondary foliation at the lateral edge of the vermiform body of the cerebellum. Brains of the genera Phyllostomus (Figs. 13-15) and Phylloderma (Fig. 16) reveal another subgroup within the Phyllostomatinae. The brains of both genera 152 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY BIOLOGY OF THE PHYLLOSTOMATIDAE 153 are characterized by massive cerebral hemispheres that are elongate and anteriorly blunt. Sulcation of the cerebrum is pronounced, and the pseudocentral sulci and sulci anterior to them are well developed. The cerebral hemispheres are well provided with small secondary fissures radiating from the larger sulci. In Phyllostomus elongatus (Fig. 13), the inferior colliculi are exposed dorsally; but in Phyllostomus hastatus (Fig. 14), P. discolor (Fig. 15), and Phylloderma (Fig. 16), the inferior colliculi are completely covered by cerebral and cerebellar tissues. In all four species, the cerebellum has ornamentation in the form of secondary foliation at the lateral edges of the vermiform body, which is itself enlarged to form apronounced medial crest to the cerebellum. The brain of Vampyrum spectrum (Fig. 17) is extremely large, but is not as massive in appearance as that of Phyllostomus because of more pronounced elongation. The cerebral hemispheres are well convoluted and sulcated and have secondary fissures radiating from the major sulci. The cerebellum achieves its maximum ornamentation in Vampyrum. The vermiform body is raised to form an extraordinary medial ridge, and there is considerable secondary foliation at the lateral edges of the vermiform body. Subfamily Glossophaginae Brains of nine genera and 12 species were examined from the Glossophaginae. Brains from this subfamily have relatively smooth and shallow cerebral hemispheres, shallow development of the nlajor sulci, and dorsally unexposed inferior colliculi. The cerebellum is simple and without secondary ornamentation. The brains of Choeroniscus godmani (Fig. 18) and C. intermedius (not figured) are probably impossible to differentiate externally. The brain of Choeroniscus is characterized by the presence of ashort cerebrum having asmooth surface and small olfactory bulbs. The pseudocentral sulcus is extremely shallow, and the pseudotemporal lobes are smoothly rounded on the ventral side. The cerebellum is simple and has no secondary'lobation. Hylonycteris (Fig. 19) and Lichonycteris (Fig. 20) have brains similar to that of Choeroniscus. The cerebrum is short and smooth, the olfactory bulbs are small, and the pseudocentral sulci are shallow. The pseudotemporal lobes of Hylonycteris are ventrally rounded as in Choeroniscus, but those of Lichonycteris are ventrally angular. The cerebellum is simple in both genera. The brains of Glossophaga alticola (Fig. 21), G. commissarisi (Fig. 22), and G. soricina (not figured) are virtually indistinguishable externally. In Glossophaga, the cerebral hemispheres are short, smooth, and almost lacking sulci. The olfactory bulbs are small. The pseudotemporal lobes are somewhat angular and the cerebellar foliations are simple. The brains of Choeronycteris (Fig. 23) and Monophyllus (not figured) are characterized by relatively elongated cerebral hemispheres that are vertically shallow. The pseudotemporal lobes are the shallowest within the Glossophaginae and the cerebellum has only primary foliations. Leptonycteris (Fig. 1) has abrain much like that of Glossophaga alticola. However, the cerebral sulci tend to cut deeper into the mass of the cerebrum in Leptonycteris than in any of the species of Glossophaga. Anoura (Fig. 24) has abrain similar to that of Choeronycteris. The cerebrum is elongate and smooth, and the pseudocentral sulci are shallow. The olfactory bulbs are relatively large, and the pseudotemporal lobes are shallow. In Anoura, the cerebral hemispheres are somewhat more massive than in Choeronycteris, and they reach the greatest relative length within this subfamily. The brain of Lonchophylla (Fig. 25) has the most massive cerebrum within the Glossophaginae. The cerebrum is elongated and has relatively well-developed pseudocentral sulci and large olfactory bulbs. The pseudotemporal lobes project ventrally, and the cerebellum achieves its maximum degree of foliation within the Glossophaginae. Subfamily Carolliinae Brains of both genera in this subfamily have been examined. The brains of Carollia (Fig. 26) and Rhinophylla (Fig. 27) are similar in almost every detail. The brains of Carollia perspicillata (Fig. 26) and C. subrufa' (not figured) are virtually identical externally. In Carollia, the cerebrum is similar to that of Micronycteris in having relatively short and smooth hemispheres. The pseudocentral sulci are well developed, as are the sulci anterior to the pseudocentral sulci. The pseudotemporal lobes are rounded ventrally, and the inferior colliculi are not exposed dorsally. The cerebellum is simple and has only primary lobes. The brain of Rhinophylla (Fig. 27) is similar to that of Carollia, but in RhinophYlla, the pseudotemporallobes project ventrally in an angular fashion. Subfamily Stenoderminae This large subfamily is represented by specimens from 28 species, representing 12 genera. Brains from this subfamily normally have adeep cerebrum with pseudotemporallobes that project ventrally in adistinctive fashion. The genus Artibeus (Figs. 28-35) is represented herein by nine species. The brain of A. aztecus (Fig. 29) is characterized by the most shallow cerebral hemispheres within the Stenoderminae. The pseudotemporal lobes are angular and project ashort distance ventrally. The pseudocentral sulci are shallow, and there are no sulci anterior to the pseudocentral sulci. The inferior colliculi are covered dorsally, and the vermiform body of the cerebellum forms alow crest. In A. phaeotis (Fig. 30), the cerebral hemispheres are relatively deeper than in A. aztecus. The pseudotemporal lobes are angular and project ventrally farther than in A. aztecus. The prepseudocentral gyrus is enlarged and protrudes dorsally. The posterior portions of the inferior colliculi are exposed dorsally, and the cerebellum resembles that of A. aztecus. The brain of A. toltecus (Fig. 31) is anatomically intermediate between the brains of A. aztecus and A. phaeotis. In A. toltecus, the cerebral hemispheres are deep and the pseudotemporallobes project ventrally in an angular fashion. The pseudocentral sulci are shallow, as are the sulci anterior to the pseudocentral sulci. The posterior edges of the cerebral hemispheres cover all but the posterior most edges of the inferior colliculi. The cerebellum has amedial crest and small secondary foliations at the lateral edges of the vermiform body. The brains of A. watsoni (Fig. 32) and A. cinereus (Fig. SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 154 BIOLOGY OF THE PHYLLOSTOMATIDAE J55 28) are indistinguishable except that A. cinereus has alarger brain than does A. watsoni. The brains of these two species are characterized by deep and relatively smooth cerebral hemispheres with shallow sulci. In "both species, the inferior colliculi are not exposed, and the cerebellum is crested and without secondary foliation. The brain of A. inopinatus (Fig. 33) is characterized by deep cerebral hemispheres having angular and ventrally projecting pseudotemporal lobes. The pseudocentral sulci and sulci anterior to the pseudocentral sulci are well developed. The inferior colliculi are not exposed dorsally, and the cerebellum is simple and has ahigh medial crest. A. hirsutus (not figured), A. jamaicensis (Fig. 34), and A. lituratus (Fig. 35) have brains that are similar in morphology. In these species, the brain has relatively well-convoluted cerebral hemispheres and well-developed major sulci. The pseudotemporal lobes project ventrally in an angular fashion, and the inferior colliculi are not dorsally exposed. The cerebellum is well crested and has small secondary foliations at the lateral edges of the vermiform body. The brain of Enchisthenes (Fig. 36) is known to me through examination of one poorly preserved specimen. The relatively smooth cerebrum with deep and angularly projecting pseudotemporal lobes resembles that of Artibeus watson;. The inferior colliculi are not visible from above, and the cerebellum is simple and media.lly crested. The genus Vampyrodes (Fig. 37) is characterized by abrain with large and anteriorly blunt cerebral hemispheres having poorly developed sulci. The pseudotemporal lobes project ventrally in an angular fashion. The posterior portions of the inferior colliculi are dorsally exposed and the cerebellum is simple. The vermiform body forms amedial crest to the cerebellum. The brains of Uroderma bilobatum (Fig. 38) and U. magnirostrum (Fig. 39) are similar in most features. These brains have deep cerebral hemispheres with angular pseudotemporal lobes that protrude ventrally to alesser extent than in Artibeus. The pseudocentral sulci and sulci anterior to the pseudocentral sulci are well developed. The inferior colliculi are not exposed dorsally, and the vermiform body of the cerebellum forms alow medial crest. There are secondary foliations at the lateral edges of the vermiform body. Brains of Sturnira [ilium (not figured), S. mordax (Fig. 40), and S. ludovici (Fig. 41) closely resemble each other. These brains are characterized by deep and extremely smooth cerebral hemispheres. The pseudocentral sulci and sulci anterior to the pseudocentraJ sulci are more poorly developed than in other stenodermine bats. The pseudotemporaJ lobes are angular and project ventrally. The inferior colliculi are completely covered in S. lilium and S. ludovici, but in S. mordax, the posterior portions of the colliculi are dorsally exposed. In all three species, the cerebellum is simple and has amedial crest. The brain of Ectophylla macconnelli (Fig. 42) is similar to that of Artibeus phaeotis. In E. macconnelli, the cerebral hemispheres are deep and relatively smooth. The major sulci are well developed and the prepseudocentral gyrus protrudes dorsally. The pseudotemporal lobes are angular and protrude ventrally. The inferior colliculi are exposed dorsally, and the cerebellum is simple and crested. The brains of Vampyressa nymphaea (Fig. 43) and V. pusilla (Fig. 44) are not alike. The brain of V. nymphaea is characterized by deep and somewhat domed cerebral hemispheres having well-developed major sulci. The pseudotemporal lobes project ventrally in an angular fashion. The inferior colliculi are not exposed dorsally. The cerebellum is simple and has alow medial crest. In V. pusilla, the cerebrum is smooth and has well-developed sulci, but it is not domed as in V. nymphaea. In V. pusilla, the inferior colliculi are exposed dorsa11y. The pseudotemporal lobes and cerebellum of V. pusilla resemble those of V. nymphaea. The brains of Chiroderma salvini (Fig. 45) and C. villosum (Fig. 46) are similar and are characterized by massive cerebral hemispheres that are well sulcated, anteriorly blunt, and somewhat convoluted. The nlajor cerebral sulci are deeper in C. salvini, but have small secondary fissures radiating from them in both species. The pseudotemporal lobes are massive and project ventrally in the typically angular fashion. The inferior colliculi are not exposed dorsally, and the cerebellum is crested medially and has small secondary foliations along the lateral edges of the vermiform body. Brains of species of Vampyrops (Figs. 47-49) have massive cerebral hemispheres, deep sulci and high convolutions in most species, and arelatively complex pattern of foliation to the cerebellum. Among the species exanlined, V. helleri (Fig. 47) has the least convoluted cerebrum and the shallowest cerebral sulci. In V. villatus (Fig. 48) and Vampyrops infuscus (Fig. 49), the cerebral hemispheres are deeply sulcated and well convoluted. The pseudotemporal lobes are relatively large in all three species and project ventrally in an angular fashion. In V. helleri, the posterior portions of the inferior colliculi are exposed dorsally, but in the other two species, the inferior colliculi are completely covered dorsally by the cerebral hemispheres. In all three, the cerebellum is crested medially, and there are secondary foliations at the lateral edges of the vermiform body. The brain of Cenlurio (Fig. 50) is characterized by pronounced anteroposterior compression resulting in adomed brain. The cerebral hemispheres are relatively smooth and have shallow sulci. The pseudotemporal lobes are also somewhat compressed and project ventrally in adifferent fashion than in other members of the subfamily. The inferior colliculi are exposed dorsally, and the cerebellum is simple and slightly crested. Ametrida (Fig. 51) has abrain similar to that of Cenlurio. The cerebrum is compressed and quite smooth, with almost no trace of the major sulci. The pseudotemporal lobes are relatively shallow, angular, and project ventrally. The cerebellum is simple and has alow crest. The brain of Stenoderma (Fig. 52) also resembles that of Centurio. Stenoderma is characterized by massive cerebral hemispheres that are relatively smooth. The major cerebral sulci are shallow, and the pseudotemporallobes are large, angular, and ventrally projecting. The inferior colliculi are exposed dorsally. The cerebellum is simple and has alow crest. 156 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY BIOLOGY OF THE PHYLLOSTOMATIDAE 157 Subfamily Phyllonycterinae Two species (Erophylla bombi/rons and Brachyphylla cavernarum) were examined from this subfamily. Phyllonycteris is 'the only genus not represented. The brain of Erophylla (Fig. 53) has arelatively short and smooth cerebrum, with ashallow pseudocentral sulcus and aslightly developed sulcus anterior to the pseudocentral sulcus. The pseudotemporal lobes are rounded ventrally, and there is asimple pattern of foliation of the cerebellum. The inferior colliculi are dorsally exposed and are not contiguous with each other. In Erophylla, the vermiform body of the cerebellum constitutes about athird of the total dorsal expression of the cerebellum. The brain of Brachyphylla (Fig. 54) is characterized by relatively smooth and massive cerebral hemispheres. The major cerebral sulci are wel1 developed, including the sulcus anterior to the pseudocentral sulcus. The pseudotemporal lobes are ventrally angular, but do not protrude ventrally. The inferior colliculi are not visible from above. The vermiform body of the cerebellum is laterally enlarged and constitutes half of the dorsal exposure of the cerebellar tissues. In addition, Brachyphylla has one characteristic not found in any other phyllostomatid brain in that the uvular portion of the cerebellum is greatly enlarged and forms aprominent lobe at the posterior edge of the vermiform body along the dorsal surface of the medulla. Subfamily Desmodontinae Brains were examined from all three genera of this subfamily. The brains of Desmodus (Fig. 55), Diaemus (Fig. 56), and Diphylla (not figured) are similar in that they al1 have large cerebral hemispheres that are deeply sulcated and well convoluted. The cerebellum is variously ornamented. The brain of Desmodus (Fig. 55) is characterized by elongate and convoluted cerebral hemispheres that are deeply cut by the pseudocentral sulci and the sulci anterior to them. The pseudotemporal lobes project ventrally in an angular fashion (as in the Stenoderminae), and the inferior colliculi are not dorsally exposed. The vermiform body of the cerebellum forms amedial crest, and there are small secondary foliations along its lateral edges. In Diaemus (Fig. 56), the cerebral hemispheres are less elongate than in Desmodus but are well convoluted and deeply sulcated. The pseudotemporal lobes are angular as in Desmodus and the inferior colliculi are not dorsally exposed. The cerebellum has alow medial crest and secondary foliations along the lateral edges of the vermiform body. Diphylla has acerebrum similar to that of Diaemus. The pseudotemporal lobes are angular and project ventrally. The posterior portions of the inferior colliculi are dorsally exposed, and the cerebellum is large and has amedial crest.