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Nowitate MUSEUM ovitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3426, 14 pp., 3 figures, 3 tables February 27, 2004 A New Chocoan Species of Lonchophylla (Chiroptera: Phyllostomidae) LILIANA M. DAVALOS! ABSTRACT Lonchophylla is a diverse genus of glossophagines characterized by large, forwardly projecting inner upper incisors and the absence of zygomatic arches. Seven species are currently recognized, including the large-bodied (greatest length of skull >24.5 mm) robusta, handleyi, hesperia, and bokermanni and the small-bodied (greatest length of skull <24.5 mm) thomasi, dekeyseri, and mordax. Lonchophylla species range throughout the Neotropics and include endemics in Amazonia, the Cerrado, and the arid regions of coastal Peru and Ecuador. In this paper I describe a new large-bodied species, Lonchophylla chocoana, from the subtropical rainforests of the Choc6 in southwestern Colombia and northwestern Ecuador. I also document the diagnostic external, craniodental, and mitochondrial characters of the new species and summarize morphological characteristics for the new species and its sympatric congeners. INTRODUCTION three genera constitute the phyllostomid tribe Lonchophyllini. Recent analyses of morpho- The genus Lonchophylla Thomas includes logical and mitochondrial DNA-sequence seven species of glossophagines character- characters from lonchophyllines have recov- ized by large, forwardly projecting inner up- ered a paraphyletic Lonchophylla, although per incisors and the absence of zygomatic low support values indicate that new evi- arches (Taddei et al., 1983; Thomas, 1903). dence might easily overturn this conclusion Two closely related monotypic genera distin- (Davalos and Jansa, 2004). guished primarily by larger size and the mor- As members of the phyllostomid subfam- phology of premolars are Platalina Thomas ily Glossophaginae, species of Lonchophylla and Lionycteris Thomas. Together, these have a rostrum roughly equal in length to the ' Division of Vertebrate Zoology (Mammalogy), American Museum of Natural History; and Department of Ecology, Evolution, and Environmental Biology, Columbia University. e-mail: [email protected] Copyright © American Museum of Natural History 2004 ISSN 0003-0082 a AMERICAN MUSEUM NOVITATES NO. 3426 braincase and a small but well-developed METHODS nose leaf (Koopman, 1994). Lonchophylla STUDY SITE AND COLLECTION METHODS species range from Nicaragua to Peru, Boliv- ia and southeastern Brazil, and west of the The known specimens of this new species Andes through the Choc6é south to the arid were collected 2 km south of Alto Tambo regions of Ecuador and northwestern Peru. (00°54'N, 78°33'W; 700 m) in Provincia Es- Large-bodied Lonchophylla species (greatest meraldas, Ecuador, and at La Guaraperia, ca. length of skull >24.5 mm) include robusta, 3 km NW of Junin (1°21'N, 78°08'W; ca. handleyi, hesperia, and bokermanni. Loncho- 900 m) in Departamento Narifio, Colombia Phylla thomasi, dekeyseri, and mordax are (fig. 1; Burton Lim, personal commun.; Cad- comparatively smaller (greatest length of ena et al., 1998). The average annual rainfall skull <24.5 mm, Taddei et al., 1983). Lon- at the elevation belt encompassing both Alto chophylla species are typically found in for- Tambo and La Guaraperia (fig. 1) is estimat- ests, gardens, and plantations, with L. hes- ed to be around 2800 mm (ESRI, 1997), and peria restricted to deciduous forest, L. bok- the average annual temperature is estimated ermanni to the Cerrado, and L. handleyi, L. to be around 25°C (ESRI, 1999). The Alto robusta, and L. mordax ranging to cloud-for- Tambo locality comprised humid disturbed est habitat (Cadena et al., 1998; Handley, secondary forest (Burton Lim, personal com- 1976; Koopman, 1994; Sazima et al. 1978). mun.). The vegetation at La Guaraperia had Lonchophylla feed predominantly on nectar, been selectively logged without ever being although insects, fruit, and pollen are also fully cleared (Cadena et al., 1998). part of their diet (Gardner, 1977; Sazima et Both specimens were captured using mist al., 1978). nets. At Alto Tambo, nets were set along a The lowland Lonchophylla species are eas- trail and along a river surrounded by dis- ily captured by mistnetting (Handley, 1976; turbed secondary growth, and there was rain Sazima et al., 1978; Simmons and Voss, before the capture of the specimen (B. Lim, 1998), and L. thomasi is widespread through- personal commun.). At La Guaraperia nets out its range and appears to be the most com- were set across a stream, a trail, and in a mon glossophagine found in Amazonia (Em- cleared area (Cadena et al., 1998). Mist nets mons, 1997). Lonchophylla congeners, how- were opened at 1700 hours and stayed open ever, have not been found in sympatry at overnight until 0500 hours at Alto Tambo, well-sampled lowland localities (Simmons while at La Guaraperia nets were open at and Voss, 1998; Voss and Emmons, 1996), dusk and closed around 2200 hours (B. Lim, although mordax and robusta were recently personal commun.; Cadena et al., 1998). collected together at 1400 m above sea level Nets varied from 10 to 18 m in length and in Altaquer, southwestern Choc6é of Colom- were set to reach 3 m above ground at both bia (Cadena et al., 1998). localities. In 1999 Sharon A. Jansa and I initiated a collaboration to sequence cytochrome b from MEASUREMENTS AND ABBREVIATIONS lonchophyllines to resolve relationships All measurements reported here are from among the bats in this tribe. Among others, adult individuals with closed epiphyses. The we obtained samples from 11 individuals first five measurements listed below were representing five species of Lonchophylla taken from skin tags or other records made (for details and results see Davalos and Jan- by the collector, but other dimensions were sa, 2004). One of those species is an unde- measured using digital calipers. Linear mea- scribed large Lonchophylla resembling L. ro- surements of external and craniodental di- busta and L. handleyi in size, but differing mensions are reported in millimeters (mm); from these and other known congeners by a weights are reported in grams (g). Measure- unique combination of traits. I describe this ments follow Simmons et al. (2002), and are new species below, document its diagnostic described below: morphological and molecular attributes, and summarize available observations in a com- Total length: Distance from the tip of the snout to parative context. the tip of the last caudal vertebra. 2004 DAVALOS: NEW SPECIES OF LONCHOPHYLLA 3 7 : ; || a y . ‘7 f4 * ‘Le j — Sin" “| ‘o (| i. ; PN “| ff fr 7 \ ~_ B P f o Hy SS a ’ J / = ¥ |\ be # | F, él XY ‘ ~ Yo, ae a i 1 A —— i's 4 ! p a ag x wh VY I \ | : en Junin* / pmmsfi/ f my ‘is é/‘ x — : Alt aquear Ny - “he\ \ Cer ~ Bieta 3 Ss Ricaurte \ y i" i ( .— x i S o—~, 1 \ < a \ 7 ae 2 { ey \ \ . _ 7 ta SS oF Ae io oa f % | \ ue “oe, Ay He ie ~. | ~ 4 \ | ie a Colombia { KY i“ Alto Tambo#—. ~~ 5 / te SY : ( \ ve ow Ecuador Tulcan 5 \’ ‘ ian. =, ; ( My } hy \ \ Af y iSae e A PSee sao . as f _ é | Py es \ gon 4 : ‘. Weg \ 7 f oya } ey s h 8| ()() PT)f e | se. =~ [ A Lo \ | {1 a J | / | ali ty \ \ i d \ a { ~ ioe \ \ , } [ a sn, vo 2 ee se ee eee Fa Fig. 1. Southern Chocé of Colombia (Departamento Narifio) and Ecuador (Provincia Esmeraldas), showing the towns closest to the localities at Alto Tambo (Alto Tambo) and La Guaraperia (Junin). Gray outlines indicate major roads, black outlines correspond to rivers. See text for map coordinates. Tail length: Measured from the point of dorsal Palatal length: From the anteriormost point be- flexure of the tail with the sacrum to the tip of hind the incisors to the edge of the bony pala- the last caudal vertebra. tion. Hindfoot length: From the anterior edge of the Condylo-incisive length: From the posteriormost base of the calcar to the tip of the claw of the point on the occipital condyles to the anterior- longest toe. most point on the upper incisors. Ear length: From the notch to the fleshy tip of the Postorbital breadth: Least breadth across frontals pinna. posterior to the postorbital bulges. Forearm length: From the elbow (tip of oleocra- Braincase breadth: Greatest breadth of the glob- non process) to the wrist (including the car- ular part of the braincase. pals), measured with the wing partially folded. Mastoid breadth: Greatest cranial breadth across Thumb length: From the metacarpal-phalangeal the mastoid region. joint to the tip of the claw of the thumb. Maxillary toothrow length: From the anteriormost Tibia length: From the proximal end of the tibia edge of the canine crown to the posteriormost to the posterior base of the calcar. edge of the crown of M3. Greatest length of skull: From the posteriormost Breadth across molars: Greatest breadth across point on the occiput to the anteriormost point the outer edges of the crowns of the upper mo- on the premaxillae (excluding the incisors). lars. 4 AMERICAN MUSEUM NOVITATES NO. 3426 Height of upper canine: Dorsoventral height from exhaustive mammal inventories in neotropi- the anterior base of the crown of the upper ca- cal forests west of the Andes have been re- nine to the tip. stricted to Barro Colorado, Panama, and La Breadth at the base of upper canine: Greatest an- Selva, Costa Rica (Voss and Emmons, 1996), teroposterior breadth at the base of the upper it is likely that the known distribution is just canine. an artifact of inadequate collecting, com- In attempting to make taxonomic identifi- pounded with superficial similarities to L. ro- cations I compared the vouchers from Alto busta and L. handleyi. In particular, speci- Tambo and La Guaraperia with original de- mens of L. handleyi reported from the west- scriptions, notes on distribution, and com- ern slopes of the Cordillera Occidental of parative series of specimens of all other Lon- Departamentos of Valle and Narifio, Colom- chophylla species. The following institution- bia, and specimens of Lonchophylla sp. from al acronyms correspond to museums in the Pacific lowlands of Ecuador (Alberico which vouchers and other specimens exam- and Orejuela, 1982; Alberico, 1987; Albuja, ined are preserved: AMNH, American Mu- 1999) deserve investigation. seum of Natural History (New York); ICN, ETYMOLOoGy: For the Choc6, an area of en- Instituto de Ciencias Naturales (Bogota); demism comprising tropical and subtropical MHN, Museo de Historia Natural (Popayan); humid forests west of the Andes from west- ROM, Royal Ontario Museum (Toronto); ern Panama through Colombia to northern TTU, The Museum, Texas Tech University Ecuador. (Lubbock); USNM, National Museum of DIAGNOsIs: A large-sized species of Lon- Natural History (Washington, DC). chophylla (forearm 45—48 mm; weight 19— 23 g) with chocolate-brown to chestnut-col- SYSTEMATICS ored dorsal fur and brown ventral fur; dorsal FAMILY PHYLLOSTOMIDAE GRAY, 1825 and ventral hairs bicolored (the former more distinctly banded than the latter), 7-8 mm SUBFAMILY GLOSSOPHAGINAE BONAPARTE, 1845 long in shoulder region; pinnae short with rounded tips; thumb large (7.5—8.3 mm); GENUS LONCHOPHYLLA THOMAS, 1903 fringe of the uropatagium sparse; calcar Lonchophylla chocoana, new species shorter than foot; palate long; with postpa- Figures 2, 3 latine torus; right and left upper I1 meet at distal third of the crown; large gap present TYPE MATERIAL: The holotype (ROM 105786), an adult female preserved as skin, between I] and I2, and smaller gap between skull, skeleton, and tissue, was collected in I2 and C; outer upper incisors (12) pointed the vicinity of Alto Tambo by Mark Engs- ventrally more than ventromedially; upper trom, Burton Lim, and Francisco Sornoza canines large; height of P3 slightly less than (original number F40079) on 3 March 1996. P4; gap present between C and P3; smaller One paratype is an additional female (CN gap present between P3 and P4; P4 with 13649) collected by Alberto Cadena, Pilar small but well-developed lingual cusp; M1 Rivas, and Robert P. Anderson (original and M2 roughly the same width; lower in- number ACG 2765) at La Guaraperia on 12 cisors small and trilobed, with crown height March 1995. approximately equal to crown width. DISTRIBUTION: Currently known only from Of the characters listed above, five are par- northwestern Ecuador and southwestern Co- ticularly useful for field identification of Lon- lombia (fig. 1), but perhaps widespread chophylla chocoana: size (forearm 44—48 throughout the lower to middle elevations of mm; weight 19—23 g); thumb length (7.5-8.3 the southern biogeographic Choc6. Because mm); dorsal fur color; ventral fur bicolored; -> Fig. 2. Dorsal (A), ventral (B), and lateral (C) views of the skull of the holotype of Lonchophylla chocoana (ROM 105768). Scale bar = 5 mm. 2004 DAVALOS: NEW SPECIES OF LONCHOPHYLLA 6 AMERICAN MUSEUM NOVITATES NO. 3426 TABLE 1 probably larger than their male conspecifics. Measurements of the Specimens of Although this might produce some overlap Lonchophylla chocoana in measurements between female robusta and male chocoana, other nonmetric char- Holotype Paratype acters remain diagnostic. ROM 105786 ICN 13649 From all other large Lonchophylla, cho- Sex Female Female coana can be unambiguously distinguished Weight 23.0 19.0 on the basis of fur coloration, forearm length, Total length 78.0 84.5 greatest length of skull, and palatal length. Tail length 7.0 10.5 Additionally, chocoana can be distinguished Hind foot length 15.0 13.2 from robusta on the basis of size of the Ear length 17.0 14.0 thumb, greatest length of skull, palatal Forearm length 48.0 45.0 length, orientation and size of the outer upper Thumb length dd 8.3 incisors, and the relative size of gaps be- Tibia length 16.5 18.2 tween I1, I2 and C, which are roughly equal Greatest length of skull 27.7 28.3 in robusta. L. chocoana can be distinguished Palatal length 7.0 6.9 from handleyi on the basis of the furry fringe Condyloincisive length 20.9 27.4 Postorbital breadth 5.3 5.3 along the uropatagium, size of the thumb, the Braincase breadth 10.2 10.5 relative size of the upper canines, presence Mastoid breadth 6.9 7.0 of a ridge along the posterior edge of the Maxillary toothrow length 9.8 11.7 palate (postpalatine torus), and the well-de- Breadth across molars 7.3 7.3 veloped lingual cusp on P4. Lonchophylla Height of upper canine 2.6 2.7 chocoana is larger than all remaining con- Breadth at base of upper canine 1.7 1.7 geners in most anatomical dimensions. L. hesperia and bokermanni may reach similar length of skull, but they have greater skull and trilobed lower incisors. Within the genus length-to-width ratios and are absent from Lonchophylla this combination of traits is the known range of chocoana (see measure- unique to L. chocoana. Appendix 1 summa- ments in Taddei et al., 1983). rizes the molecular data that prompted the As with robusta and handleyi, the dorsal investigation of morphological differentia- pelage of chocoana is composed of bicolored tion in L. chocoana. The average uncorrected hairs with cream-white bases and brown tips. pairwise sequence divergence of the mito- The length of the dorsal fur along the upper chondrial cytochrome b gene of L. chocoana back in chocoana is approximately 7—8 mm, with respect to L. robusta from Colombia is slightly longer than in handleyi and robusta 11.6%, to L. robusta from Panama is 12.3%, (4.0-7.5 mm). The ventral pelage of cho- and to L. handleyi is 11.6%. These genetic coana contrasts with both handleyi and ro- distances are indicative of species-level dif- busta in being bicolored from neck to genital ferentiation, according to one recently pro- region, whereas in handleyi the ventral fur is posed criterion (Bradley and Baker, 2001). exclusively beige-brown unicolored, while MEASUREMENTS: Dimensions of both spec- some robusta individuals (particularly fe- imens of Lonchophylla chocoana are provid- males) show bicolored hairs around the neck ed in table 1, and metrical comparisons with but never in the abdominal region. representative series of other large congeners The cranial morphology of chocoana is within its range are summarized in table 2. similar to that of other members of the genus. DESCRIPTION AND COMPARISON: Loncho- L. chocoana has a relatively long rostrum, a phylla chocoana is the largest member of the small but noticeable anteorbital inflation, and genus, as large or slightly larger than han- a large braincase. Zygomatic arches are ab- dleyi, and larger than robusta, hesperia, and sent as in all other lonchophyllines, and the bokermanni (tables 1—3; Taddei et al., 1983). palate is the longest of any congener. Like Females are generally larger than males in all Lonchophylla, L. chocoana has a dental robusta and handleyi (table 2), and the two formula 12/2, C1/1, P2/3, M3/3 X 2 = 34. chocoana females (table 1) are therefore The inner upper incisors are large compared 2004 DA, VALOS NEW SPECIES OF LONCHOPHYLLA ) ) 9 L 9 0 ' ' 7 SoyeUl OZ (I1Z ) O'EL (0'Z8 8-0$7 (O'ZS -0'0) O7I (OPZv I-O (Zt€ 9-4"'S) Z( ELIS) Z( O'L7-7'9Z) Z (VO0I-L'6) ~ (69-99) Z (101-66) Zt (Pl-7Z) ZOI-SD F BS ol 9°92 66 LO OO! EZ 9 S I OeU091 08 09 Or | g's 891 0'L7Z vol 39 66 77 9'| Z| So[PUl GE! 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In the two disjunct subspecific populations). the latter respect chocoana also resembles all In chocoana P4 is longer than P3 (antero- other species of Lonchophylla. posterior dimension), as it is in robusta, han- Differences in the dentition are subtle, but dleyi, mordax, and thomasi. The height (dor- they provide the means to distinguish species soventral dimension) of P3 is either equal to of Lonchophylla (see Hill, 1980). In cho- or very slightly less than P4. Both robusta coana the inner upper incisors have a small and handleyi have shorter (dorsoventral di- triangular gap between them, meeting at the mension) P3 than P4, while mordax and tho- distal third of the crown. This is unlike ro- masi show upper premolars that are subequal busta, handleyi, mordax, and thomasi, all of in height. The basal lingual cusp on P4 is which show a taller gap between the inner present and well developed in chocoana, ro- upper incisors, meeting at the distal quarter busta, and thomasi, less developed in mor- of the crown. The upper canines of chocoana dax, and poorly developed or absent in han- are relatively and absolutely larger than those dleyi. The first molar of chocoana is com- of robusta and handleyi. The posterior cusp paratively wide, and it is larger (both in an- of the upper canines of chocoana is blunt, teroposterior and lateral dimensions) than similar to that of handleyi, while the poste- M2, which in turn is larger than M3. Both rior cusp is sharp in robusta, thomasi, and robusta and handleyi have similar widths of Fig. 3. Anterior views of the lower dentition of Lonchophylla chocoana (left, ROM 105786) and L. robusta (right, ICN 13648, male). Note the shape and height of the gap between inner upper incisors and the trilobed lower incisors in chocoana. These morphological traits can be used in conjunction with size and other characters for field identification. 2004 DAVALOS: NEW SPECIES OF LONCHOPHYLLA 9 M1 and M2, although M1 is longer (antero- Tambo indicate that it was captured along posterior dimension) than M2, with overall with other glossophagines (Anoura spp.) in a smaller M3. The first two molars of thomasi disturbed secondary forest in nets left open and mordax are similar in length and height. overnight due to continuous rain (Burton The lower outer incisors of chocoana are Lim, personal commun.). The specimen from trilobed, as they are in thomasi. Lower inci- La Guaraperia was also captured in second- sors are rarely trilobed in robusta; m. mordax ary forest, between dusk and 2200 hours, has trilobed incisors, but m. concava does along with Anoura caudifera, A. cultrata, not; and incisors are bilobed in handleyi. The and A. geoffroyi (Cadena et al., 1998). The height of the lower incisors is roughly the survey in Narifio additionally captured Lon- same as the width of these teeth in all species chophylla robusta and L. mordax within the of Lonchophylla. The lower premolar denti- same general region at the Altaquer locality. tion of chocoana resembles that of handleyi. Both specimens were captured in early In robusta the maximum height (dorsoven- March: at Alto Tambo on 3 March 1996, and tral dimension) of the series corresponds to at La Guaraperia on 12 March 1995. p3, and the maximum length (anteroposterior Although these data are scant, some as- dimension) corresponds to p4. In handleyi, pects are noteworthy. First, L. chocoana cur- p4 is the highest and longest tooth of the se- rently seems rare in its range; one specimen ries, and in comparison the p4 of chocoana from Alto Tambo prompted this investiga- is narrow (lateral dimension). In contrast, the tion, and the search for conspecifics only longest lower premolar of thomasi and mor- found 1 more among 11 specimens of large- dax is p2. The height of the premolars in bodied Lonchophylla from west of the Cor- thomasi is roughly the same throughout the dillera Occidental of Colombia. As noted premolar series, whereas in mordax p4 is the above, a few large Lonchophylla from west- highest premolar (dorsoventral dimension). ern Colombia and Ecuador reported in the The first molar of chocoana is the widest and literature might correspond to chocoana. longest of the molar series. This is the same Only further sampling can document the condition as in handleyi, although m1 of complete range and the rarity of chocoana. handleyi is slender in comparison. In robusta Second, as with other Lonchophylla, cho- the m1 is slightly wider (lateral dimension) coana can be captured using ground-level than m2, which in turn is wider than m3. In mist netting. At Paracou, French Guiana, and robusta m1 is the longest molar (anteropos- in southern Venezuela, most L. thomasi were terior dimension). The differences among the captured using ground-level mistnets (Han- molars are less marked in mordax and tho- dley, 1976; Simmons and Voss, 1998), as masi, particularly the latter. Although m1 is were all L. robusta individuals of the Smith- the longest molar (anteroposterior dimen- sonian Venezuelan Project (Handley, 1976) sion) in both species, height and width vary and all L. robusta from Tambito, Colombia only slightly along the series. (Davalos and Guerrero, 1999). Third, the The coronoid process in Lonchophylla habitat associated with the capture of L. cho- chocoana is high and oriented at an angle of coana has been characterized as disturbed, about 110° with respect to the toothrow, as and the simultaneous capture of other glos- in robusta. In handleyi the coronoid process sophagines probably indicates the presence projects at an angle of around 130°. Both tho- of local food resources, regardless of the masi and mordax show even greater angles, habitat condition. as well as a longish (anteroposterior dimen- Little is known about the ecological niche sion) coronoid process in comparison to their of species in this genus, but observed diver- larger congeners. sity in body size and habitat choice (from dry scrub to lowland rainforest) indicates that there is some sort of partitioning in feeding NATURAL HISTORY and roosting ecology. These ecological char- Both known specimens of Lonchophylla acteristics become more intriguing now that chocoana were captured in ground-level mist three species of Lonchophylla have been nets. The notes on the specimen from Alto found in similar habitats, as they were in the 10 AMERICAN MUSEUM NOVITATES NO. 3426 western slopes of the Cordillera Occidental American Museum of Natural History and of Narifio, Colombia (Cadena et al., 1998). Columbia University. ACKNOWLEDGMENTS REFERENCES Alberico, M. 1987. Notes on distribution of some This paper is a contribution from the Mo- bats from southwestern Colombia. Fieldiana nell Molecular Laboratory and the Cullman Zoology 39: 133-135. Research Facility in the Department of Or- Alberico, M., and J. Orejuela. 1982. Diversidad nithology, American Museum of Natural especifica de dos comunidades de murciélagos History, and has received generous support en Narino, Colombia. Cespedesia, Suplemento from the Lewis B. and Dorothy Cullman Pro- 3: 31-40. gram for Molecular Systematics Studies, a Albuja, L. 1999. Murciélagos del Ecuador, 2nd joint initiative of The New York Botanical ed. Quito: Cicetronic Cia. Ltda. Bradley, R.D., and R.J. Baker. 2001. A test of the Garden and The American Museum of Nat- genetic species concept: cytochrome-b se- ural History. This material is based on work quences and mammals. Journal of Mammalogy supported by NASA under grant no. NAGS- 82: 960-973. 8543 and the Center for Biodiversity and Cadena, A., R.P Anderson, and P. Rivas-Pava. Conservation at the American Museum of 1998. Colombian mammals from the Chocoan Natural History. I thank the collectors from slopes of Narifio. Occasional Papers Museum the Instituto de Ciencias Naturales, A. Cad- of Texas Tech University 180: 1-15. ena, P. Rivas, and R.P. Anderson, as well as Davalos, L.M., and J.A. Guerrero. 1999. The bat M. Engstrom, B. Lim, and FE Sornoza from fauna of Tambito, Colombia. Chiroptera Neo- tropical 5: 112-115. the Royal Ontario Museum, for acquiring the Davalos, L.M., and S.A. Jansa. 2004. Phylogeny specimens. A. Cadena, M. Gomez Laverde, of the Lonchophyllini (Chiroptera: Phyllostom- and Y. Mufioz from the Instituto de Ciencias idae). Journal of Mammalogy 85. Naturales made available the specimens un- Emmons, L.H. 1997. Neotropical rainforest mam- der their care, as did M. Engstrom and B. mals: a field guide, 2nd ed. Chicago: University Lim from the Royal Ontario Museum, M. of Chicago Press. Carleton from the United States National ESRI. 1997. World precipitation zones scale: 1: Museum, and N.B. Simmons from the Amer- 15000000. Redlands: Environmental Systems Research Institute. ican Museum of Natural History. R.P. An- ESRI. 1999. World temperature zones (annual) derson, S.A. Jansa, and N.B. Simmons pro- scale: 1:15000000. Redlands: Environmental vided invaluable advice in the course of this Systems Research Institute. project, and B. Carstens and R. Voss com- Gardner, A.L. 1977. Feeding habits. Jn R.J. Baker, mented on the manuscript. J.A. Guerrero as- J.K. Jones, and D.C. Carter (editors), Biology sisted in the 1999 field expedition to Tambito of bats of the New World family Phyllostomi- that obtained all Colombian tissues of robus- dae, part 2: 293-350. Lubbock: Texas Tech ta. That expedition was funded by Bat Con- Press. Handley, C.O. 1976. Mammals of the Smithson- servation International, the Royal Geograph- ian Venezuelan Project. Brigham Young Uni- ic Society (London), The Explorers Club versity Science Bulletin-Biological Series 20: (New York), and the Institute of Latin Amer- 1-89. ican Studies and the Center for Environmen- Hill, J. 1980. A note on Lonchophylla (Chirop- tal Research and Conservation at Columbia tera: Phyllostomatidae) from Ecuador and Peru, University. I extracted and sequenced Col- with the description of a new species. Bulletin ombian specimens of robusta at the molec- of the British Museum of Natural History (Zo- ular laboratory of Instituto de Investigacién ology) 38: 233-236. de Recursos Biolé6gicos Alejandro von Hum- Koopman, K. 1994. Chiroptera: systematics. Handbuch der Zoologie 8: 1—217. boldt at the Biotechnology Research Unit of Sazima, I., L.D. Vizotto, and V.A. Taddei. 1978. the Centro Internacional de Agricultura Trop- Uma nova especie de Lonchophylla do Serra do ical thanks to E. Gaitan, J.D. Palacio, and J. Cip6, Minas Gerais, Brasil (Mammalia, Chirop- Tohme. The author is supported by an inter- tera, Phyllostomidae). Revista Brasileira de national graduate student grant from the Biologia 38: 81-89.