IANA 590.5 Fl N.S. no. 105(2005) Zoolog NEW IO. 105 Morphological Phylogeny of the Bat Genus Platyrrhinus Saussure, 1860 (Chiroptera: Phyllostomidae) with the Description of Four New Species Paul M. Velazco November 15, 2005 Publication 1535 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY FIELDIANA Zoology NEW SERIES, NO. 105 Morphological Phylogeny of the Bat Genus Platyrrhinus Saussure, 1860 (Chiroptera: Phyllostomidae) with the Description of Four New Species Paul M. Velazco Division of Mammals Department of Zoology Field Museum of Natural History 1400 South Lake Shore Drive Chicago, Illinois 60605-2496 U.S.A. Department of Biological Sciences University of Illinois at Chicago Chicago, Illinois 60680-4348 U.S.A. BWL06Y LIBRARY Accepted July 22, 2005 101 BURRJLL HAll SSSKST- 1S' 2005 m * 4 ms PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY £ 2005 Field Museum of Natural History ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents Abstract 1 Introduction 1 Background 2 Materials and Methods 4 Taxonomic Sample and Outgroup Deter- mination 4 Sources of Data 5 Definition of the Characters and Ordering of Character States 5 Polarity 5 Methods of Phylogenetic Analysis 5 Character Descriptions 6 Pelage and Integument 6 Skull 10 Dentition 12 Postcranial Skeleton 17 Results 18 Discussion and Conclusions 18 Taxonomic Diagnoses of the Platyrrhinus Species 20 Acknowledgments 42 Literature Cited 42 Appendix 1: Specimens Examined 45 Appendix 2: Data Matrix 51 Appendix 3: Optimization of Characters ... 51 List of Illustrations 1 . Frontal view of Platyrrhinus sp. nov 2 2. Owen's (1987; redrawn from Fig. 17) hypothesis of relationships among Pla- tyrrhinus species, Sturnira erythromos, Uroderma magnirostrum, Chiroderma villosum, and Vampyrodes caraccioli 3 3. Schematic view of the heads of three hypothetical bats 7 4. Ventral view of the chin of five hypo- thetical bats 8 5. Frontal view of the noseleaf of Sturni- ra erythromos, Platyrrhinus infuscus, and P. nigellus 9 6. Detail of the posterior margin of the hard palate of Platyrrhinus brachyceph- alus and P. helleri "Western" 10 7. Detail of the postorbital processes of Platyrrhinus infuscus and P. nigellus .... 11 8. Detail of the mastoid and paraoccipital processes of Platyrrhinus helleri "West- ern," P. nigellus, P. vittatus "South- ern," and Sturnira erythromos 11 9. Detail of the fossa on the squamosal end of the zygomatic arch of Platyr- rhinus brachycephalus, P. dorsalis, and P. infuscus 12 10. Detail of the posterior clinoid process- es of Platyrrhinus infuscus and Uro- derma magnirostrum 13 1 1 . Occlusal view of the maxillary tooth- row of Platyrrhinus infuscus and Stur- nira erythromos 13 1 2. Lateral view of the maxillary toothrow of Platyrrhinus infuscus 14 13. Occlusal view of M1-M2 of Pla- tyrrhinus infuscus and P. dorsalis "Norte" 15 14. Strict consensus tree of the morpholog- ical analysis with the characters unor- dered 19 15. Distributional map of Platyrrhinus al- bericoi sp. nov 21 16. Labial views of the left p2 and p4 of Platyrrhinus vittatus and P. albericoi sp. nov 25 17. Distributional map of Platyrrhinus au- rarius and P. brachycephalus 26 18. Distributional map of Platyrrhinus cho- coensis and P. dorsalis 27 19. Distributional map of Platyrrhinus hel- leri 28 20. Distributional map of Platyrrhinus in- fuscus and P. lineatus 29 2 1 . Frontal view of Platyrrhinus ismaeli sp. nov 30 22. Distributional map of Platyrrhinus is- maeli sp. nov 31 23. Occlusal views of the left Ml and M2 of Platyrrhinus ismaeli sp. nov. and P. dorsalis 33 24. Frontal view of Platyrrhinus masu sp. nov 34 25. Distributional map of Platyrrhinus masu sp. nov 35 26. Distributional map of Platyrrhinus ma- tapalensis sp. nov 38 27. Labial views of the left p2 and p4 of Platyrrhinus matapalensis sp. nov., P. helleri, and P. brachycephalus 40 28. Distributional map of Platyrrhinus ni- gellus, P. recifinus and P. vittatus 41 29. Strict consensus tree with nodes num- bered as referenced in Appendix 3 52 m List of Tables 4. Measurements of the type series of Pla- tyrrhinus ismaeli sp. nov 30 5. Measurements of the type series of Pla- 1. Taxa examined 4 tyrrhinus masu sp. nov 34 2. Measurements of the type series of Pla- 6. Selected measurements of Platyrrhinus tyrrhinus albericoi sp. nov 21 ismaeli sp. nov. and P. masu sp. nov 36 3. Selected measurements of members of 7. Measurements of the type series of Pla- the genus Platyrrhinus 22 tyrrhinus matapalensis sp. nov 37 Morphological Phylogeny of the Bat Genus Platyrrhinus Saussure, 1860 (Chiroptera: Phyllostomidae) with the Description of Four New Species Paul M. Velazco Abstract Platyrrhinus, comprising 10 species of broad-nosed bats, is one of the most diverse genera in the Neotropical family Phyllostomidae. To evaluate the content and limits of this genus, the phylogenetic relationships among these species and other closely related taxa were examined using a data set of 60 morphological characters including external, cranial, dental, and post- cranial characters. Four species (Carollia subrufa, Sturnira erythromos, Uroderma magniros- trum, and Vampyrodes caraccioli) of the family Phyllostomidae were used as outgroups to test the monophyly of Platyrrhinus and to resolve intrageneric relationships. Parsimony analysis resulted in two most parsimonious trees. The results support Platyrrhinus monophyly with a bootstrap value of 84% and a Bremer support of 3. Three synapomorphies are recognized for the genus. The results also support the recognition of P. umbratus as a junior synonym of P. dorsalis, the division of P. dorsalis into three distinct species, and the discovery of two new species previously considered part of P. helleri and P. vittatus. Taxonomic diag- noses of all the species of Platyrrhinus based on character optimizations as well as descriptions of four new species are presented. This revision brings the total number of species of Platyr- rhinus to 14, making it the most speciose genus in the family Phyllostomidae. Introduction The genus Platyrrhinus Saussure, 1860, be- longs to the family Phyllostomidae (Mammalia: Chiroptera). The distribution of this family is re- stricted to the Neotropics. Morphologically, it is characterized by the presence of a noseleaf, de- veloped in most of its members, as well as the presence of two synapomorphic characters that differentiate it from the other Microchiropterans: oviductal folds restricted to the extramural ovi- duct (Hood & Smith, 1982) and friction lock in the digits of feet (Simmons & Quinn, 1994; Sim- mons, 1998). Within the Phyllostomidae, Platyr- rhinus is placed in the subfamily Stenodermati- nae, which is diagnosed by a combination of the presence of short and flattened muzzle, short and wide nose leaf, narrow uropatagium, lack of tail, and frequent presence of facial and/or dorsal stripes (Linares, 1986; Albuja, 1999). This sub- family also has two synapomorphic characters: a single common uterine lumen, without remnants of cornual lumina, and the oviducts enter the fun- die border of the uterine body near the midsagittal line (Hood & Smith, 1982). The species of Platyrrhinus (Fig. 1) are thought to be monophyletic (Owen, 1987; Lim, 1993; Wetterer et al., 2000). Platyrrhinus is diagnosed from other Stenodermatinae by a combination of three characters: two accessory cusps on the pos- terior face of P4, presence of three upper molars, and presence of a fringe of hair on the edge of the uropatagium. Although other genera also have these characters, no other genus possesses all three (Lim, 1993; Albuja, 1999). Ten nominal species of Platyrrhinus are com- monly recognized (Owen, 1987; Koopman, 1993; Nowak, 1999). The distributional range of FIELDIANA: ZOOLOGY, N.S., NO. 105, NOVEMBER 15, 2005, PP. 1-54 Fig. 1. Frontal view of Platyrrhinus sp. nov. (FMNH 172108); adult male photographed at Pillahuata in the Cultural zone of the Manu Biosphere Reserve, Cuzco, Peru, by P. M. Velazco. Platyrrhinus includes North, Central, and South America, from southern Mexico to Bolivia, Par- aguay, Uruguay, and southeastern Brazil (San- born, 1955; Cabrera, 1957; Jones & Carter, 1976; Hall, 1981; Koopman, 1982, 1993). Some spe- cies of this genus are widely distributed — for ex- ample, P. helleri is distributed from southern Mexico to Peru, Bolivia, and Brazil as well as Trinidad and Tobago (Koopman, 1982; Ferrell & Wilson. 1991). The genus Platyrrhinus presents difficulties both in terms of its diagnosis and in how many species it comprises. The present work seeks to elucidate the phylogenetic relationships among the members of Platyrrhinus. To do so, cladistic analyses of discrete morphological characters were conducted. External, cranial, dental, and postcranial skeletal characters were examined. The results from these analyses offer a new inter- pretation of morphological diversity within the genus and a reassessment of the hypotheses of relationships presented by Owen (1987). Lim (1993), Baker et al. (2000, 2003), and Wetterer et al. (2000). Background All the species of Platyrrhinus Saussure, 1860, described before 1990 were recognized under the genus Vampyrops Peters, 1865. Gardner and Fer- rell (1990) reviewed the nomenclatural history of these bats and determined that Platyrrhinus and Vampyrops were synonymous and that Platyr- rhinus had priority over Vampyrops. Koopman (1993) recognized 10 nominal spe- cies of Platyrrhinus: P. aurarius (Handley & Fer- ris, 1972), endemic to the Guianan Shield (south- ern Venezuela. Guyana, and Suriname); P. bra- chycephalus (Rouk & Carter, 1972), distributed in the Amazon and the coasts of northern South America and adjacent islands (northern Brazil, Colombia to Guyana, Ecuador, Peru, and Bolivia); P. chocoensis Alberico and Velasco, 1991, dis- tributed in the Choco region (western Colombia, in lowlands between the western slope of the An- des Cordillera and the Pacific coast); P. dorsalis (Thomas, 1900), distributed on both slopes of the Andes Cordillera and Central America (Panama to Peru and Bolivia); P. helleri (Peters, 1866), dis- FIELDIANA: ZOOLOGY tributed from Central America (Oaxaca and Ve- racruz [Mexico]), south to Peru, Bolivia, the Am- azon of Brazil, and Trinidad and Tobago; P. in- fuscus (Peters, 1880), distributed on the eastern slope of the north part of the Andes Cordillera and in the Amazon (Colombia to Peru, Bolivia, and northwestern Brazil); P. lineatus (E. Geof- frey, 1810), distributed in the Pampean subregion, Amazon, and the eastern slope of the north part of the Andes Cordillera (Colombia to Peru, Bo- livia, Uruguay, northern Argentina, and southern and eastern Brazil); P. recifinus (Thomas, 1901), distributed in the highlands and coast of eastern Brazil and the Amazon (eastern Brazil); P. um- bratus (Lyon, 1902), distributed in the eastern slope of the northern Andes Cordillera, north coast of South America and adjacent islands, and the Pacific coast of Ecuador and Colombia (Pan- ama, Colombia, and northern Venezuela); and P. vittatus (Peters, 1860), distributed on the eastern slope of the north part of the Andean Cordillera, north coast and islands of South America, and Central America (Costa Rica to Venezuela and from Peru to Bolivia) (Koopman 1982, 1993; Al- berico 1990; Lim & Engstrom, 2000). Sanborn (1955), Gardner and Carter (1972b), and Carter and Rouk (1973) considered Platyr- rhinus umbratus to be conspecific with P. dorsa- lis, synonymizing the former with the latter. How- ever, Handley (1976) reported that P. umbratus differed from P. dorsalis, which was followed by Koopman (1993) and Owen (1987). Velazco and Solari (2003) considered P. aquilus, P. oratus, and P. umbratus as junior synonyms of P. dor- salis, and they stated that P. dorsalis as it is now recognized is a complex of three taxa. One of these corresponds to P. dorsalis (Thomas, 1900), with aquilus, oratus, and umbratus as junior syn- onyms; the other two taxa are undescribed spe- cies, denoted P. dorsalis "Norte" and P. dorsalis "Centro-Sur." Platyrrhinus dorsalis "Norte" is distributed on both slopes of the Andes in Ecua- dor and Peru, and P. dorsalis "Centro-Sur" is dis- tributed on the eastern slope of the Andes in Peru (Velazco & Solari, 2003). Tuttle (1970) regarded Platyrrhinus infuscus as a junior synonym of P. vittatus, but Gardner and Carter (1972b) and Koopman (1978) considered P. infuscus distinct from P. vittatus. Gardner and Carter (1972a) described P. nigellus as a species present in Colombia and probably Ecuador and Peru, but Koopman (1978, 1982), Jones and Cart- er (1979), and Willig and Hollander (1987) treated nigellus as a subspecies of P. lineatus. Owen Sturnira erythromos Uroderma magnirostrum Chiroderma villosum Vampyrodes caraccioli Platyrrhinus infuscus Platyrrhinus recifinus Platyrrhinus brachycephalus Platyrrhinus aurarius Platyrrhinus lineatus Platyrrhinus umbratus Platyrrhinus vittatus Platyrrhinus dorsalis Platyrrhinus helleri Platyrrhinus nigellus Fig. 2. Owen's (1987; redrawn from Fig. 17) hy- pothesis of relationships among Platyrrhinus species, Sturnira erythromos, Uroderma magnirostrum, Chirod- erma villosum, and Vampyrodes caraccioli, based on a consensus of continuous and discrete characters. (1987) and Velazco and Solari (2003) recognized P. nigellus as a species distinct from P. lineatus. The first explicit hypothesis of phylogenetic re- lationships within Platyrrhinus was presented by Owen (1987; Fig. 2) in the context of his analysis of the subfamily Stenodermatinae. That analysis has been criticized both generally (Lim, 1993) as well as for specific genera included in his analysis (Pacheco & Patterson, 1991). In that analysis, Owen used 22 discrete and 33 continuous exter- nal, cranial, and dental characters. Owen's preferred phylogenetic hypothesis (Owen, 1987; Fig. 2) placed the monotypic genus Vampyrodes as a sister genus of Platyrrhinus. He considered Platyrrhinus to be a monophyletic group and indicated two clades that were not re- solved within Platyrrhinus, one containing P. ni- gellus + P. helleri + P dorsalis and the other P. vittatus + P. aurarius + P. lineatus + P. umbra- tus. The relationships of the remaining species in the tree (P. brachycephalus, P. infuscus, and P. recifinus) also were unresolved. Lim (1993) sub- sequently argued that Owen (1987) did not judi- ciously select the best characters for his cladistic analysis and used distance methods not suitable for the reconstruction of evolutionary history. Lim (1993) used 20 external, cranial, dental, and internal morphological characters to generate a hypothesis of relationships at the generic level for the Stenodermatinae. In this analysis, he ob- VELAZCO: PLATYRRHINUS PHYLOGENY tained Platyrrhinus + Vampyrodes as sister taxa, supporting Owen's (1987) hypothesis. Two char- acters unite these taxa: one unambiguous syna- pomorphy (fringe of hair on the edge of the uro- patagium) and one parallel synapomorphy, which is also present in Uroderma (accessory cusps on P4). Lim (1993) found that the sister group to this pair includes Ectophylla + Mesophylla + Vam- pyressa + Chiroderma. Wetterer et al. (2000) evaluated the phyloge- netic relationships among the genera of the family Phyllostomidae using 150 morphological, karyo- logical, and molecular characters. In this analysis, they also found Platyrrhinus and Vampyrodes to be sister taxa, supported with a bootstrap value of 46. Their ACCTRAN (accelerated transformation optimization) analysis indicated five synapomor- phies for this grouping: uropatagial fringe pres- ence, II occlusal margin generally straight or slightly rounded, ml paraconid present, pharyn- geal region of tongue completely covered with pa- pillae, cerebral vermis completely covering the longitudinal fissure between inferior colliculi, and inferior colliculi exposed dorsally only along lat- eral edges of cerebelar vermis. Their DELTRAN (delayed transformation optimization) analysis also identified three of these as synapomorphies: uropatagial fringe presence. Ml hypocone pre- sent, and pharyngeal region of tongue completely covered with papillae. In their analysis Uroderma was sister to the clade Platyrrhinus + Vampyro- des, and Ectophylla + Vampyressa + Chiroderma was sister to the clade Uroderma + Platyrrhinus + Vampyrodes. Baker et al. (2000, 2003) evaluated the phylo- genetic relationships of the family Phyllostomidae at the generic level using nuclear and mitochon- drial genes. In those analyses they also found Pla- tyrrhinus and Vampyrodes to be sister taxa. More- over, Mesophylla and Vampyressa were found to be the sister of Platyrrhinus + Vampyrodes. Materials and Methods Taxonomic Sample and Outgroup Determination For this cladistic analysis, the taxonomy of Koopman (1993) was followed with the exception of P. dorsalis, P. lineatus, and P. umbratus. For these species, Velazco and Solari (2003), who rec- ognized three distinct morphs within P. dorsalis Table 1. List of examined taxa. Ingroup Playtrrhinus Platyrrhinus Playtrrhinus Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus aurarius brachycephalus chocoensis dorsalis dorsalis "Norte" dorsalis "Centro-Sur" helleri "Eastern" helleri "Western" infuscus lineatus nigellus recifinus vittatus "Northern" vittatus "Southern" Outgroups Carollia subrufa Sturnira erythromos Uroderma magnirostrum Vampyrodes caraccioli (two not yet described: they are here denoted P. dorsalis, P. dorsalis "Norte," and P. dorsalis "Centro-Sur"), was followed. They also recog- nized P. nigellus as a valid species and considered P. umbratus as a junior synonym of P. dorsalis. After the character analysis, two different popu- lations in P. helleri and P. vittatus were recog- nized. For the purposes of the analysis herein, they are denoted as P. helleri "Eastern," P. hel- leri "Western," P. vittatus "Northern," and P. vittatus "Southern." A list of all 14 ingroup and 4 outgroup taxa is given in Table 1. Only one outgroup is necessary to root a phy- logenetic tree (Nixon & Carpenter, 1993), but to establish the polarity of ambiguous characters and to test Platyrrhinus monophyly, four taxa were used as outgroups: Carollia subrufa (Hahn, 1905), Sturnira erythromos (Tschudii, 1844), Uroderma magnirostrum Davis, 1968, and Vampyrodes car- accioli (Thomas, 1889). Freeman (2000) consid- ered Carollia as the most ancestral and least spe- cialized genus in the tribe Stenodermatini. Wet- terer et al. (2000) found that Carollia + Rhino- phylla is the sister group of the subfamily Stenodermatinae; they placed Sturnira as the most basal taxon in the Stenodermatinae. Owen (1987). Lim (1993), Baker et al. (2000, 2003), and Wet- terer et al. (2000) all agreed that Vampyrodes was the sister genus of Platyrrhinus. With respect to the position of Uroderma, Owen (1987) placed it in a clade together with Ectophylla and Artibeus, Lim (1993) placed Uroderma in a clade together with Artibeus, and Wetterer et al. (2000) placed FIELDIANA: ZOOLOGY Uroderma as the sister taxon of the clade Platyr- rhimis + Vampyrodes. Baker et al. (2003) placed Uroderma as sister to the clade Mesophylla + Ec- tophylla + Platyrrhinus + Vampyrodes. Sources of Data Only adult specimens were examined based on Morris (1972), Anthony (1988), and Pacheco and Patterson (1992), and all the specimens belong to scientific collections of museums (skins, skulls, skeletons, and alcohol-preserved specimens). A complete list of the specimens examined is pro- vided in Appendix 1. A total of 1,314 specimens were examined, be- longing to the scientific collections of the follow- ing museums; acronyms from museums holding holotypes of Platyrrhinus species were also in- cluded: AMNH, American Museum Natural of History, New York, New York, USA; BMNH, Natural History Museum, London, England; EPN, Escue- la Politecnica Nacional, Quito, Ecuador; FMNH, Field Museum of Natural History, Chicago, Illi- nois, USA; IND-M, La Unidad de Investigation "Federico Medem" — Inderena, Bogota, Colom- bia; LSUMZ, Louisiana State University, Muse- um of Natural Science, Baton Rouge, Louisiana, USA; MCZ, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA; MNHN, Museum National d'Histoire Na- turelle, Paris, France; MSB, Museum of South- western Biology, University of New Mexico, Al- buquerque, New Mexico, USA; MUSM, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru; MVZ, Museum of Vertebrate Zoology, University of California, Berkeley, California, USA; QCAZ, Pontificia Universidad Catolica del Ecuador, Museo de Zoologfa — Mamfferos, Quito, Ecuador; ROM, Royal Ontario Museum, Toronto, Ontario, Cana- da; TCWC, Texas Cooperative Wildlife Collec- tion, Texas A&M Collection, College Station, Texas, USA; UMMZ, University of Michigan, Museum of Zoology, Ann Arbor, Michigan, USA; USNM, National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA; UV, Universidad del Valle, Cali, Colombia; ZMB, Museum fur Naturkunde der Humboldt- Universitat zu Berlin, Berlin, Germany. The number of specimens per species examined in this study varied according to their availability in these museum collections; each taxon was rep- resented by at least 12 individuals. Definition of the Characters and Ordering of Character States Autapomorphies were included in the phylo- genetic analysis for the species of Platyrrhinus because they are useful in diagnosis. In addition, relevant characters in the relationships among the outgroups were included to provide resolution. In cases where variation inside the species is present, the character was scored as polymorphic. Sim- mons and Conway (2001) were followed for the categorization of missing data. The final data set comprised a total of 60 char- acters: 17 characters of the pelage and integu- ment, 6 of the skull, 35 dental characters, and 2 characters of the postcranial skeleton (Appendix 2). Fifty-eight percent (35) of the characters were binary, and 42% (25) of the characters were mul- tistate. Polarity The "outgroup method" was used to establish the polarity of the character states (Watrous & Wheeler, 1981). On the basis of earlier studies, outgroup species formed the sequence Carollia subrufa, Sturnira erythromos, Uroderma magni- rostrum, and Vampyrodes caraccioli. Carollia subrufa was used to root the tree because it is considered ancestral to the remaining three out- group species (Freeman, 2000). Methods of Phylogenetic Analysis The matrix elaboration and the visualization of the character mapping were performed using MacClade 4.0 (Maddison & Maddison, 2000). The phylogenetic analysis was performed using PAUP version 4.0b6 (Swofford, 2001) on a Mac- intosh PowerBook G4. The branch and bound option of PAUP was used to identify the most parsimonious trees with the smallest number of steps and to calculate tree statistics. The analysis was performed using all characters unordered. A strict consensus was per- formed because more than one parsimonious tree was obtained. To perform the decay analysis (Bre- mer, 1988), the program TreeRot version 2b (So- VELAZCO: PLATYRRHINUS PHYLOGENY renson, 1999) was used. A branch and bound bootstrap analysis was performed, with 10,000 replications, to evaluate the support of the result- ing clades. Character optimizations were calculated using both the ACCTRAN and DELTRAN. The results of these optimizations are presented in Appen- dix 3. Character Descriptions Pelage and Integument Character J — Facial stripes. Absent (0); pre- sent (1). The facial stripes are present as two lines on each side of the face. The dorsomedial stripe begins at the posterolateral edge of the noseleaf, continues above the eye, and terminates close to the pinna. The ventrolateral stripe begins in the corner of the mouth and runs toward the tragus. The facial stripes are present in all the species of Platyrrhinus, Vampyrodes caraccioli, and Urod- errna magnirostrum. In contrast, the facial stripes are absent in Carollia subrufa and Sturnira ery- thromos. Owen (1987: character 4) uses the facial stripes as part of a multistate character that he termed "Pelage patterns." In this character Owen in- cludes facial stripes, dorsal stripe, shoulders patches, and striping on the wing. I agree with Owen on his facial stripes scoring for the species used in the present analysis. Lim (1993: character 1) and Wetterer et al. (2000: character 6) include facial stripes in their phylogenetic analyses; char- acter states and scoring used in this analysis co- incide with theirs. Character 2 — Brightness of facial stripe. Dor- somedial and ventrolateral stripes obviously sub- equal (0); dorsomedial stripe more marked than the ventrolateral stripe (1). In most Platyrrhinus species, both facial stripes have the same tonality, either dark or brilliant (e.g., P. aurarius, P. helleri "Eastern," P. nigellus). In contrast, P. vittatus "Northern" and P. vittatus "Southern" have a more luminous dorsomedial stripe. Carollia sub- rufa and Sturnira erythromos were scored as " — " because they do not have facial stripes. This is the first use of this character in a phylogenetic analysis. Character 3 — Facial stripe coloration. Bril- liant-white (0); dark (1). The facial stripes are brilliant or white in P. brachycephalus, P. helleri "Eastern," P. helleri "Western," P. lineatus, P. recifinus, P. vittatus "Southern," Vampyrodes caraccioli, and Uroderma magnirostrum. In con- trast, they are dark in P. aurarius, P. chocoensis, P. dorsalis, P. dorsalis "Norte," P. dorsalis "Centro-Sur," P. infuscus, P. nigellus, and P. vit- tatus "Northern." Carollia subrufa and Sturnira erythromos were scored as " — " because they lack both facial stripes. This is the first use of this character in a phylogenetic analysis. Character 4 — Basal protuberance where the genal vibrissae are implanted. Absent (0); pre- sent (1). The genal vibrissae are implanted in the cheek, ventral, and/or posterior to the eye. The genal vibrissae are present in all the species of Platyrrhinus. These vibrissae are implanted in a basal protuberance in some Platyrrhinus (e.g., P. brachycephalus, P. helleri "Eastern," P. lineatus) (Fig. 3A). In contrast, the genal vibrissae are im- planted directly on the cheek in Carollia subrufa and some species of Platyrrhinus (e.g., P. aurar- ius, P. chocoensis) (Fig. 3B-C). This character has not been used in previous studies. Character 5 — Number of vibrissae surround- ing the margins of the noseleaf in a single ar- ray. Nine vibrissae present (0); eight vibrissae present (1); seven vibrissae present (2); six vi- brissae present (3). Some species of Platyrrhinus have seven vibrissae surrounding the margins of the noseleaf in a single array (e.g., P. brachyce- phalus, P. chocoensis, P. lineatus) (Fig. 3B). In contrast, six vibrissae are presented to each side of the noseleaf in some Platyrrhinus and Vam- pyrodes (e.g., P. aurarius, P. dorsalis) (Fig. 3A). Platyrrhinus helleri "Eastern" presents intraspe- cific variation (seven and eight vibrissae present; Fig. 3B-C). Sturnira erythromos have nine vi- brissae surrounding the margins of the noseleaf. Character 6 — Number of vibrissae present on the upper lip, ventral to the vibrissae that sur- round the margins of the noseleaf. One vibrissa present on each side of the face (0); two vibrissae present (1). In some species of Platyrrhinus (e.g., P. aurarius, P. chocoensis, P. dorsalis), Sturnira erythromos, Uroderma magnirostrum, and Vam- pyrodes, a single vibrissa is present on the upper lip, ventral to the vibrissae that surround the mar- gins of the noseleaf (Fig. 3A). Only P. brachy- cephalus and Carollia subrufa have two vibrissae on the upper lip (Fig. 3B). Platyrrhinus helleri "Eastern," P. infuscus, and P. nigellus present in- traspecific variation in this character (one vibrissa and two vibrissae present) (Fig. 3A-B). This char- acter has not been used in any other study. FIELDIANA: ZOOLOGY B Fig. 3. Schematic view of the heads of three hypo- thetical bats, showing details of basal protuberance (BP) and pinna folds (PF). Scale bar = 15 mm. Character 7 — Number of submental vibrissae on each side of the chin. Absence of submental vibrissae (0); two submental vibrissae present (1); three submental vibrissae present (2); four sub- mental vibrissae present (3); five submental vi- brissae present (4). The submental vibrissae are on the chin and lower lip and are usually arranged in two notable parallel longitudinal rows (Pocock, 1914; Brown, 1971). Some Platyrrhinus (e.g., P. aurarius, P. lineatus, P. nigellus), Uroderma magnirostrum, and Vampyrodes (Fig. 4C, E) al- ways have four submental vibrissae. Platyrrhinus dorsalis "Centro-Sur" has two submental vibris- sae (Fig. 4B). Platyrrhinus brachycephalus pre- sent five submental vibrissae (Fig. 4D). Intraspe- cific variation was noted (three and four submen- tal vibrissae) in P. chocoensis (Fig. 4A, C, E). Carollia subrufa lacks submental vibrissae. This character has not been used in previous studies. Character 8 — Interramal vibrissae. Absence (0); one interrramal vibrissa present (1); two in- terramal vibrissae present (2). The interramal vi- brissae form a median tuft on the gular region between the two rami of the lower jaws and well behind the mandibular symphysis (Pocock, 1914; Brown, 1971). Platyrrhinus recifinus, Sturnira er- ythromos, and Uroderma magnirostrum lack in- terramal vibrissae (Fig. 4A). An interramal vibris- sa is present in some species of Platyrrhinus (e.g., P. brachycephalus, P. lineatus) and Vampyrodes (Fig. 4B-D). Two interramal vibrissae are present in P. infuscus, P. vittatus "Northern," P. helleri "Western," and Carollia subrufa (Fig. 4E). Intra- specific variation is found in P. vittatus "South- ern" (one and two interramal vibrissae present; Fig. 4B-E). This character was used by Wetterer et al. (2000: character 13). Character 9 — Inferior border of horseshoe. Inferior border with no distinct boundary between horseshoe and lip (0); inferior border partially joined to the upper lip (1); inferior border com- pletely free (2). The inferior border of the horse- shoe is completely free of the upper lip in some Platyrrhinus (e.g., P. brachycephalus, P. linea- tus), Uroderma magnirostrum, and Vampyrodes (Fig. 5C). Platyrrhinus chocoensis, P. dorsalis "Centro-Sur," and P. infuscus have intraspecific variation (inferior border completely free and par- tially joined (Fig. 5B-C). In Carollia subrufa and Sturnira erythromos, there is no distinct boundary between horseshoe and the upper lip (Fig. 5A). According to Wetterer et al. (2000: character 25), the inferior border is completely free in all the species of Platyrrhinus; however, I do not agree with that assessment. Character 10 — Parallel folds located in the pinna. Absent (0); poorly marked but distinguish- able (1); well marked (2). Sturnira erythromos lacks these folds (Fig. 3C). The folds in the pinna are not very well defined in some species of Pla- tyrrhinus (e.g., P. dorsalis, P. lineatus, P. recifi- nus) and Vampyrodes (Fig. 3B). In contrast, P. aurarius, P. chocoensis, P. dorsalis "Centro- Sur," P. nigellus, P. vittatus "Northern," Carollia subrufa, and Uroderma magnirostrum have well- VELAZCO: PLATYRRHINUS PHYLOGENY A B D Fig. 4. Ventral view of the chin of five hypothetical bats, showing states of interramal (IV) and submental vibrissae (SV). Scale bar = 5 mm. marked pinna folds (Fig. 3A). Platyrrhinus helleri "Eastern" and P. infuscus present intraspecific variation, presenting both states (Fig. 3A-B). This is the first use of this character in a phylogenetic analysis. Character II — Dorsal stripe. Absent (0); in- distinct, thin, and obscured (1); definite but nar- row (2); brilliant-white and wide (3). A dorsal midline stripe is absent in Carollia subrufa and Sturnira erythromos. A narrow dorsal stripe is presented in some species of Platyrrhinus (e.g., P. chocoensis, P. dorsalis) and Uroderma mag- nirostrum. Platyrrhinus lineatus, P. recifinus, P. vittatus "Northern," P. vittatus "Southern," and Vampyrodes have the dorsal stripe that is white and wide. In Platyrrhinus infuscus, the dorsal stripe is thin and obscure. Platyrrhinus hrachy- cephalus, P. helleri "Eastern," and P. nigellus all show intraspecific variation in this character, some individuals with a narrow dorsal stripe and others with white and wide stripes. Owen (1987: char- acter 4) used the dorsal stripe as part of a complex multistate character termed "Pelage patterns," in which he included facial stripes, dorsal stripe, and shoulder patches. I agree with Owen (1987) on the scores for dorsal stripe in the species used in the present analysis. Wetterer et al. (2000: char- acter 7) used the dorsal stripe as a character with three states: absent, present, and occasionally pre- sent. FIELDIANA: ZOOLOGY f A B Fig. 5. Frontal view of the noseleaf in (A) Sturnira erythromos (MUSM 53 1 0); (B.) Platyrrhinus infuscus (MUSM 9897); (C) P. nigellus (MUSM 16170). Scale bar = 5 mm. Character 12 — Length of the dorsal fur. Long, >8 mm (0); medium, average, 6.30-7.50 mm (1); short, <6.30 mm (2). Dorsal fur is of average length in some species of Platyrrhinus (e.g., P. helleri "Eastern," P. recifinus), Carollia subrufa, Uroderma magnirostrum, and Vampyro- des. In Platyrrhinus aurarius, P. dorsalis, P. dor- salis "Norte," P. nigellus, P. vittatus "Northern," P. vittatus "Southern," and Sturnira erythromos, the dorsal fur is long. Platyrrhinus infuscus char- acteristically has short dorsal fur. Platyrrhinus chocoensis presents intraspecific variation, in which some individuals have short dorsal fur, while in others it is normal. This character was used by Marques- Aguiar (1994: character 3) in her study of the genus Artibeus. Character 13 — Bands of contrast in the dor- sal fur. Bicolored (0); tricolored (1); tetracolored (2). In most Platyrrhinus (e.g., P. brachycephalus, P. chocoensis, P. dorsalis), Carollia subrufa, and Uroderma magnirostrum, the dorsal fur is dis- tinctly tricolored. In contrast, tetracolored dorsal fur is presented in P. aurarius, P. recifinus, and Sturnira erythromos. Vampyrodes has bicolored dorsal fur. Owen (1987: character 3) and Wetterer et al. (2000: character 5) scored S. erythromos as having tricolored dorsal fur; according to my ob- servations, it has tetracolored dorsal fur. Wetterer et al. (2000) scored all Platyrrhinus as having bi- colored dorsal fur, as did Owen (1987) with the exception of P. dorsalis which had tricolored dor- sal fur. I agree with Owen's score for P. dorsalis but do not agree with the scores of the other Pla- tyrrhinus species. Instead, I observed interspecific variation in dorsal fur patterns among the species of Platyrrhinus. I agree with Owen (1987) and Wetterer et al. (2000) on the score of Vampyrodes. Character 14 — Bands of contrast in the ven- tral fur. Unicolored (0); bicolored (1); tricolored (2); tetracolored (3). A tricolored ventral fur is present in some species of Platyrrhinus (e.g., P. brachycephalus, P. infuscus), Carollia subrufa, and Vampyrodes. In contrast, P. chocoensis, P helleri "Eastern," P. vittatus "Northern," P. vit- tatus "Southern," and Uroderma magnirostrum have a bicolored ventral fur. Platyrrhinus helleri "Western" presents intraspecific variation be- cause some individuals present a bicolored ventral fur and others a unicolored ventral fur. Sturnira erythromos has tetracolored ventral fur. This char- acter has not been used in any other study. Character 15 — Hair on the dorsum of the feet. Sparse and short (0); intermediate in density and length (1); dense and long (2). Dense and long hair is present on the dorsal side of the feet in some Platyrrhinus (e.g., P. lineatus, P. vittatus "Southern") and Sturnira erythromos. In Platyr- rhinus aurarius, P. chocoensis, P. dorsalis, P. helleri "Eastern," and Vampyrodes, the hair on the dorsal side of the feet is intermediate in den- sity and length. Only P. infuscus, P. helleri "Western," and Uroderma magnirostrum have sparse and short hairs on the dorsal side of the feet. Platyrrhinus brachycephalus presents intra- specific variation in this character, some individ- uals with sparse hairs, and others with an inter- mediate distribution of hair on the dorsal side of the feet. This character has not been used in any other study. Character 16 — Posterior edge of the uropa- tagium. "V" shaped (0); "U" shaped (1). The uropatagium is "U" shaped in some Platyrrhinus (e.g., P. brachycephalus, P. recifinus), Carollia subrufa, and Sturnira erythromos. In contrast, VELAZCO: PLATYRRHINUS PHYLOGENY Vampyrodes has "V"-shaped uropatagium. Pla- tyrrhinus dorsalis, P. helleri "Eastern." P. helleri "Western," P. infuscus, P. nigellus, P. vittatus "Northern," and Uroderma magnirostrum have intraspecific variation. This character was evalu- ated only in alcohol-preserved specimens. This character has not been used in any other study. Character 17 — Distribution of the fringe of hair on the edge of the uropatagium. Absent (0); usually hairy, occasionally sparsely (1); densely haired (2). A densely haired uropatagium fringe is present in some Platyrrhinus (e.g.. P. helleri "Eastern." P. nigellus) and Sturnira ery- thromos. In contrast, P. brachycephalus, P. cho- coensis, P. dorsalis, P. infuscus, P. vittatus "Northern." and Vampyrodes have a regular or occasionally sparsely haired uropatagium fringe. Carollia subrufa and Uroderma magnirostrum lack this fringe of hair on the edge of the uropa- tagium. I agree with Lim (1993: character 13) and Wetterer et al. (2000: character 10) on the score of Platyrrhinus, Carollia, Sturnira, Uroderma, and Vampyrodes. Skull Character 18 — Posterior border of the hard palate. "U" shaped (0): "V" shaped (1). A "U"- shaped posterior border of the hard palate is pre- sent in some species of Platyrrhinus (e.g., P. li- neatus, P. recifinus). Carollia subrufa, Sturnira eiythromos, and Uroderma magnirostrum (Fig. 6A). In contrast. P. aurarius, P. dorsalis, and P. dorsalis "Norte" have a "V"-shaped posterior border of the hard palate (Fig. 6B). Platyrrhinus brachycephalus, P. chocoensis, P. dorsalis "Cen- tro-Sur." P. helleri "Eastern." P. helleri "West- ern." P. infuscus, P. nigellus, P. vittatus "North- ern." P. vittatus "Southern." and Vampyrodes have intraspecific variation for this character, with some individuals possessing the "U"-shaped pos- terior border of the hard palate and other individ- uals "V" shaped (Fig. 6A-B). Wetterer et al. (2000: character 45) used this character, but they scored Platyrrhinus, Vampyrodes, Uroderma, and Sturnira as " — " because they considered that there was a continuous range of intra- and inter- specific variation among the shape of the posterior border, making it not a useful character. Accord- ing to my observations, I found some species with intraspecific variation. Owen (1987: character 10) used this character, and I do not agree with his scores of Vampyrodes: he indicated that Vampy- B Fig. 6. Ventral view of the basicranium, showing de- tails of the posterior margin of the hard palate: (A) Pla- tyrrhinus brachycephalus (MUSM 13793); (B) P. helleri "Western" (MUSM 10725). Scale bar = 8 mm. rodes only has a "U" -shaped posterior border of the hard palate, but I also found some individuals having a "V"-shaped posterior border of the hard palate. Within Platyrrhinus, he did not find intra- specific variation, but according to my observa- tions, Platyrrhinus demonstrates intraspecific var- iation. Lim (1993: character 4) scored Platyrrhi- nus, Vampyrodes, and Sturnira as having a hard palate extended into interpterygoidal space, but he did not score the shape of the posterior border of 10 FIELDIANA: ZOOLOGY A B Fig. 7. Dorsal view of two crania, showing details of the postorbital process (PP): (A) Platyrrhinus nigellus (MUSM 16170); (B) P. infuscus (MUSM 9897). Scale bar = 12 mm. PoP B PoP D PoP Fig. 8. Lateral view of the temporal region, showing details of the mastoid process (MP) and paraoccipital process (PoP): (A) Platyrrhinus helleri "Western" (MUSM 10726); (B) P. nigellus (MUSM 16170); (C) P. vittatus "Southern" (MUSM 14584); (D) Sturnira ery- thromos (MUSM 5260). Scale bar = 2 mm. the hard palate in these genera. This character was used by Marques-Aguiar (1994: character 21) and Owen (1991: character 23) in their studies of the systematics of the genus Artibeus and Dermanu- ra, respectively. Character 19 — Postorbital process. Absent or poorly developed (0); moderately developed (1); well developed (2). A poorly developed postor- bital process is present in some species of Platyr- rhinus (e.g., P. chocoensis, P. dorsalis) and Stur- nira erythromos (Fig. 7A). In Platyrrhinus aurar- ius, P. helleri "Eastern," P. infuscus, P. recifinus, P. vittatus "Northern," P. vittatus "Southern," and Vampyrodes, the postorbital process is mod- erately developed (Fig. 7B). Platyrrhinus lineatus, Carollia subrufa, and Uroderma magnirostrum have a well-developed postorbital process. In Pla- tyrrhinus dorsalis "Norte," there is intraspecific variation, with individuals having a poorly devel- oped or moderately developed postorbital process (Fig. 7A-B). This character was used by Marques-Aguiar (1994: character 15) in her study of the genus Artibeus. Character 20 — Paraoccipital process. Poorly developed, almost imperceptible (0); moderately developed (1); well developed (2). The paraoccip- ital process appears moderately developed, about half the size of the mastoid process, in some spe- cies of Platyrrhinus (e.g., P. helleri "Eastern," P. nigellus), Carollia subrufa, Sturnira erythromos, and Vampyrodes (Fig. 8B). In Platyrrhinus au- rarius, P. infuscus, P. lineatus, P. vittatus "South- ern," and Uroderma magnirostrum, the paraoc- cipital process is well developed, almost the same size as the mastoid process (Fig. 8C-D). Platyr- rhinus brachycephalus and P. helleri "Western" have the paraoccipital process poorly developed, VELAZCO: PLATYRRHINUS PHYLOGENY 11 scarcely a third of the size of the mastoid process (Fig. 8A). Intraspecific variation is present in P. chocoensis, with the paraoccipital process poorly to moderately developed (Fig. 8A-B). I do not agree with Owen's (1987: character 12) score for P. brachycephalus, P. helleri, and P. nigellus', he scored them as if they lacked a paraoccipital pro- cess. Owen (1987) also scored P. dorsalis and P. infuscus as having minimal development of the paraoccipital process, but according to personal observations, P. dorsalis has a moderately devel- oped process (Fig. 8B), and P. infuscus has a well-developed paraoccipital process (Fig. 8C). This character has been used by Marques-Aguiar (1994: character 20), Straney (1980: character J- 6), and Owen (1987: character 12) in their phy- logenetic analyses of Artibeus, Phyllostominae. and Dermanura, respectively. Character 21 — Apophysis in the paraoccipi- tal process. Present (0); absent (1). All species of Platyrrhinus, Carollia subrufa, and Vampyrodes caraccioli lack an apophysis in the paraoccipital process (Fig. 8A-C). In contrast, Sturnira ery- thromos and Uroderma magnirostrum have an apophysis in the paraoccipital process (Fig. 8D). This character has not been used in any other study. Character 22 — Fossa on the squamosal end of the zygomatic arch, lateral to the glenoid fossa. Absent or almost imperceptible (0); shallow (1); deep (2). This fossa is absent or almost im- perceptible in some species of Platyrrhinus (e.g.. P. helleri "Eastern." P. lineatus), Carollia sub- rufa, Sturnira erythromos, Uroderma magniros- trum, and Vampyrodes (Fig. 9A). Platyrrhinus dorsalis "Norte." P. infuscus, and P. recifinus have a shallow fossa (Fig. 9B). Platyrrhinus au- rarius and P. dorsalis have a deep fossa (Fig. 9C). Platyrrhinus chocoensis and P. dorsalis "Centro- Sur" present intraspecific variation, some individ- uals have the fossa on the squamosal end of the zygomatic arch absent or almost imperceptible, and others have this fossa shallow. Platyrrhinus nigellus also demonstrate intraspecific variation, with a shallow or deep fossa. This character has not been used in any other study. Character 23 — Posterior clinoid processes. Absent (0); present (1). This process is located on the posterior face of the sella turcica in the sphe- noid bone (Stromsten, 1947). Carollia subrufa, Sturnira erythromos, and Uroderma magniros- trum lack these processes (Fig. 10A). In contrast, all the species of Platyrrhinus and Vampyrodes possess posterior clinoid processes (Fig. 10B). ZA ZA Fossa ZA Fossa B Fig. 9. Posterior view of the temporal region, show- ing details of zygomatic arch (ZA) and fossa on the squamosal end of the zygomatic arch: (A) Platyrrhinus brachycephalus (MUSM 13793): (B) P. 'infuscus (MUSM 9897): (C) P. dorsalis (FMNH 128141). Scale bar = 4 mm. Dentition Character 24 — Upper median incisors. In contact (0): without contact ( 1 ). The upper median incisors are in contact in some species of Platyr- rhinus (e.g., P. chocoensis, P. lineatus), Carollia subrufa, Sturnira erythromos, and Vampyrodes. In contrast, Uroderma magnirostrum has the upper median incisors not in contact. Platyrrhinus bra- chycephalus, P. helleri "Eastern," P. helleri "Western," P. infuscus, and P. recifinus have in- traspecific variation. Character 25 — Lobulation of the upper lat- eral incisors. Monolobed (0): bilobed (1). Mon- olobed upper lateral incisors are present in some species of Platyrrhinus (e.g., P. chocoensis, P. helleri "Eastern"). Carollia subrufa, and Vam- pyrodes. In contrast. P. dorsalis "Centro-Sur" 12 FIELDIANA: ZOOLOGY A Lingual B PCP Fig. 10. Detail of the posterior clinoid processes (PCP), in a posterior view of cranium: (A) Absence of PCP in Urodenna magnirostrum (MUSM 14029); (B) Presence of the PCP in Platyrrhinus infuscus (MUSM 9897). Scale bar = 8 mm. and Urodenna magnirostrum have bilobed upper lateral incisors. Platyrrhinus brachycephalus, P. dorsalis "Norte," P. helleri "Western," P. infus- cus, P. lineatus, P. vittatus "Southern," and Stur- nira erythromos have intraspecific variation. Character 26— Alignment of P3 in the tooth- row. P3 aligned along the axis of the toothrow (0); P3 aligned transversally to the toothrow (1). Carollia subrufa and Sturnira erythromos have the P3 aligned along the axis of the toothrow (Fig. 11 A). In contrast, all the species of Platyrrhinus, Urodenna magnirostrum, and Vampyrodes have the P3 aligned transversally to the toothrow (Fig. 11B). This character has not been used in any other study. Fig. 11. Occlusal view of the maxillary toothrow: (A) Sturnira erythromos (MUSM 5310); (B) Platyrrhin- us infuscus (MUSM 9897). Scale bar = 8 mm. Character 27 — Size of the P3 with regard to P4. P3 slightly smaller than P4 (0); P3 about half the size of P4 (1). All the species of Platyrrhinus, Uroderma magnirostrum, and Vampyrodes pos- sess a P3 slightly bigger than half the size of P4. Carollia subrufa and Sturnira erythromos have a P3 almost the same size as P4. This character has not been used in any other study. Character 28 — Stylar cusps on the posterior face of P4. P4 lacking stylar cusps or with one stylar cusp present (0); two stylar cusps present (1). All the species of Platyrrhinus, Uroderma magnirostrum, and Vampyrodes have two stylar cusps on the posterior face of P4 (Fig. 12). In contrast, Carollia subrufa and Sturnira erythro- mos do not possess two stylar cusps on P4, al- though occasionally they exhibit one stylar cusp. I agree with Lim (1993: character 6) in his scores of Platyrrhinus species. I do not agree with Wet- terer et al. (2000: character 57) on their scores for Platyrrhinus because they indicated that this ge- nus only occasionally has the second stylar cusp. According to my observations, all the individuals of Platyrrhinus always have two stylar cusps. They also stated that Lim (1993: character 6) did VELAZCO: PLATYRRHINUS PHYLOGENY 13 Fig. 12. Lateral view of the maxillary toothrow: Platyrrhinus infuscus (MUSM 9897). Scale bar = 8 mm. not report the presence of the second stylar cusp for Vampyrodes, when in fact he did report it. Character 29 — Sulcus on the posterior face of P4. Absent (0); present (1). All the species of Pla- tyrrhinus, Uroderma magni rostrum, and Vampy- rodes have a sulcus on the posterior face of P4, starting near the accessory cusps and ending at the tip of P4. In contrast, Carollia subrufa and Sturnira erythromos lack this sulcus. This char- acter has not been used in any previous study. Character 30 — Sulcus on the anterior face of P4. Absent (0); present (1). All the species of Pla- tyrrhinus have a sulcus on the anterior face of P4, starting near the cingulum and ending at the tip of P4. In contrast, Carollia subrufa, Sturnira er- ythromos, Uroderma magnirostrum, and Vampy- rodes lack this sulcus on the anterior face of P4. This character has not been used in any previous study. Character 31 — Fossa on the hypoconal basin of P4. Shallow (0); deep (1). The fossa of the hypoconal basin of P4 is deep in some species of Platyrrhinus (e.g., P. recifinus, P. vittatus "South- ern"). In contrast, P. aurarius, P. chocoensis, P. helleri "Eastern," P. lineatus, Carollia subrufa, Sturnira erythromos, and Uroderma magniros- trum have a shallow fossa. Platyrrhinus brachy- cephalus, P. dorsalis, P. dorsalis "Norte," P. dorsalis "Centro-Sur," P. helleri "Western," P. ni gel I us, and Vampyrodes have intraspecific var- iation. This character has not been used by other researchers. Character 32 — Ml Parastyle. Absent (0); pre- sent (1). The Ml parastyle is present in some spe- cies of Platyrrhinus (e.g., P. helleri "Eastern," P. recifinus). In contrast, other species of Playtrrhin- us (e.g., P. aurarius, P. chocoensis), Carollia sub- rufa, Sturnira erythromos, Uroderma magniros- trum, and Vampyrodes lack the parastyle on Ml. Platyrrhinus lineatus has intraspecific variation. Straney (1980: character: K 27) used this char- acter in his phylogeny of the Phyllostominae. Character 33 — Ml Mesostyle. Absent (0); pre- sent (1). The mesostyle on Ml is absent in some species of Platyrrhinus (e.g., P. chocoensis, P. dorsalis), Carollia subrufa, and Sturnira erythro- mos. In contrast, P. vittatus "Southern," Urod- erma magnirostrum, and Vampyrodes possess a mesostyle on M 1 . Platyrrhinus brachycephalus, P. helleri "Eastern," P. helleri "Western," P. in- fuscus, P. dorsalis "Norte," and P. nigellus have intraspecific variation. Character 34 — Labial cingulum at the base of the Ml metacone. Absent (0); present (1); sty- lar cusp on the labial cingulum of the Ml meta- cone (2). Carollia subrufa, Sturnira erythromos, and Uroderma magnirostrum lack the labial cin- gulum on the base of the Ml metacone. Platyr- rhinus brachycephalus, P. helleri "Eastern," P. helleri "Western," P. recifinus, and Uroderma magnirostrum have the labial cingulum without a stylar cusp. In contrast, a stylar cusp on the labial cingulum of the Ml metacone is present in some species of Platyrrhinus (e.g., P. chocoensis, P. in- fuscus) and Vampyrodes. Platyrrhinus lineatus and P. nigellus have intraspecific variation, as some individuals have a stylar cusp on the labial cingulum of the Ml metacone and others lack this stylar cusp. This character has not been used by other researchers. Character 35 — Sulcus on the posterior face of the Ml paracone. Absent (0); present (1). All the species of Platyrrhinus, Uroderma magnirostrum, and Vampyrodes have a sulcus on the posterior face of the Ml paracone. In contrast, Carollia subrufa and Sturnira erythromos lack this sulcus on Ml. Character 36 — Lingual cingulum on the base of the Ml metacone. Absent (0); lingual cingu- lum present at the base of the Ml metacone (1); stylar cusp on the lingual cingulum of the Ml metacone (2). Carollia subrufa, Sturnira erythro- mos, and Vampyrodes lack the lingual cingulum on the M 1 metacone. A stylar cusp is present on the cingulum of the lingual face of the Ml meta- cone in some species of Platyrrhinus (e.g., P. in- fuscus, P. lineatus). In contrast, P. helleri "East- ern," P. helleri "Western," P. recifinus, and Uroderma magnirostrum lack this stylar cusp. Platyrrhinus brachycephalus, P. chocoensis, P. dorsalis, and P. nigellus have intraspecific varia- tion, as some individuals have a stylar cusp on the lingual cingulum of the Ml metacone and others FIELDIANA: ZOOLOGY Labial Cingulum Lingual Cingulum Labial Interruption B ingua Cingulum restricted to Metacone Fig. 13. Occlusal view of M1-M2: (A) Platyrrhinus infuscus (MUSM 9897); (B) P. dorsalis "Norte" (MUSM 4946). Scale bar = 2.5 mm. lack this stylar cusp. This is the first use of this character in a phylogenetic analysis. Character 37 — Sulcus on the posterior face of the paracone not joined to the cingulum of the lingual face of the metacone on Ml. Absent (0); present (1). The majority of the Platyrrhinus spe- cies (e.g., P. aurarius, P. brachycephalus, P. cho- coensis) have this sulcus on the posterior face of the paracone joined to the cingulum of the lingual face of the metacone on Ml (Fig. 13 A). In con- trast, P. dorsalis "Norte" and Uroderma magni- rostrum have the sulcus on the posterior face of the paracone not joined to the cingulum of the lingual face of the Ml metacone (Fig. 13B). Car- ollia subrufa, Sturnira erythromos, and Vampy- rodes caraccioli were scored as " — " because they lack the cingulum on the lingual face of the Ml metacone. This character has not been used by other researchers. Character 38 — Ml metastyle. Absent (0); pre- sent (1). The metastyle on Ml is present in the majority of Platyrrhinus species (e.g., P. dorsalis, P. dorsalis "Norte"), Carollia subrufa, and Vam- pyrodes. In contrast, Sturnira erythromos and Uroderma magnirostrum lack the metastyle on M 1 . Platyrrhinus brachycephalus, P. chocoensis, P. helleri "Eastern," and P. helleri "Western" have intraspecific variation. Character 39 — Ml protocone. Well-devel- oped, large (0); moderately developed (1); small and blunt (2). A well-developed Ml protocone is present in Platyrrhinus brachycephalus. In con- trast, P. chocoensis, P. dorsalis, P. dorsalis "Norte," and P. helleri "Eastern" possess an Ml protocone that is small and blunt. Some species of Platyrrhinus (e.g., P. aurarius, P. infuscus), Carollia subrufa, Sturnira erythromos, Uroderma magnirostrum, and Vampyrodes have a moderate- ly developed protocone on Ml. Platyrrhinus vit- tatus "Northern" has intraspecific variation: some individuals have the protocone with a moderate development and others with a well-developed protocone on Ml. This character has not been used by other researchers. Character 40 — M2 parastyle. Absent (0); present (1). The parastyle on M2 is present in all Platyrrhinus species and Vampyrodes. In contrast, Carollia subrufa, Sturnira erythromos, and Uro- derma magnirostrum lack a parastyle on M2. This character has not been used in any other study. Character 41 — Labial cingulum of the M2 paracone. Absent (0); present ( 1 ). The labial cin- gulum on the M2 paracone is present in some spe- cies of Platyrrhinus (e.g., P. brachycephalus, P. chocoensis) and Vampyrodes. In contrast, Carol- lia subrufa, Sturnira erythromos, and Uroderma magnirostrum lack the labial cingulum on the M2 paracone. Platyrrhinus dorsalis, P. dorsalis "Norte," and P. dorsalis "Centro-Sur" have in- traspecific variation. This character has not been used in any other study. Character 42 — Stylar cusp on the lingual face of the M2 paracone. Absent (0); present (1). A stylar cusp on the lingual face of the M2 paracone is present in some species of Platyrrhinus (e.g., P. infuscus, P. vittatus "Southern"). In contrast, other species of Platyrrhinus (e.g., P. aurarius, P. brachycephalus), Carollia subrufa, Sturnira ery- thromos, Uroderma magnirostrum, and Vampy- rodes lack this stylar cusp on the lingual face of the M2 paracone. Platyrrhinus chocoensis and P. dorsalis have intraspecific variation. This char- acter has not been used in any other study. Character 43 — M2 metastyle. Absent (0); pre- sent (1). A metastyle on M2 is present in some species of Platyrrhinus (e.g., P. brachycephalus, P. chocoensis) and Uroderma magnirostrum. In contrast, P. vittatus "Southern," Carollia subrufa, Sturnira erythromos, and Vampyrodes lack a me- tastyle on M2. Platyrrhinus nigellus has intraspe- VELAZCO: PLATYRRHINUS PHYLOGENY 15 cific variation. This character has not been used in any other study. Character 44 — Stylar cusp on the lingual face of the M2 metacone. Absent (0); present (1). A stylar cusp on the lingual face of the M2 meta- cone is present in some species of Platyrrhinus (e.g., P. infuscus, P. vittatus "Southern") and Vampyrodes. In contrast, other species of Platyr- rhinus (e.g., P. aurarius, P. chocoensis), Carollia subrufa, Sturnira erythromos, and Uroderma ma gni rostrum lack a stylar cusp on the lingual face of the M2 metacone. Platyrrhinus dorsalis "Norte," P. dorsalis "Centro-Sur," and P. linea- tus have intraspecific variation. This character has not been used in any other study. Character 45 — Lingual cingulum of the M2 metacone. Absent (0); lingual cingulum of the M2 metacone restricted to the metacone (1); lin- gual cingulum of the M2 metacone continuous to the paracone (2). Carollia subrufa, Sturnira ery- thromos, and Vampyrodes lack the lingual cin- gulum of the M2 metacone. The lingual cingulum of the M2 metacone is continuous to the paracone in Platyrrhinus chocoensis, P. dorsalis, and P. re- cifinus (Fig. 13 A). In contrast, the lingual cingu- lum of the M2 metacone is restricted to the meta- cone in some species of Platyrrhinus (e.g., P. hra- chycephalus, P. vittatus "Southern") and Uro- derma magnirostrum (Fig. 13B). Platyrrhinus helleri "Western," P. infuscus, P. lineatus, and P. nigellus have intraspecific variation in this char- acter: some individuals have the lingual cingulum of the M2 metacone restricted to the metacone, whereas others have lingual cingulum of the M2 metacone continuous to the paracone. This char- acter has not been used by other researchers. Character 46 — M2 hypoconal basin. Devel- oped (0); rudimentary or small (1). A developed hypoconal basin on M2 is present in all the spe- cies of Platyrrhinus and Sturnira erythromos. In contrast, in Carollia subrufa, Uroderma magni- rostrum, and Vampyrodes, the M2 hypoconal ba- sin is rudimentary or small. Character 47 — Two sulci on the lingual face of the main cone of p4. Absent (0); present (1). Two sulci on the lingual face of the main cone of p4 are present in all the Platyrrhinus species, Uroderma magnirostrum, and Vampyrodes. These sulci originate at the base of the main cone and continue to the tip of the main cone. In contrast, Carollia subrufa and Sturnira erythromos lack these sulci on p4. This character has not been used in any other study. Character 48 — Cones on the posterior face of p4 (one lingual and the other one labial). Two cones present (0); one cone present on either the labial or the lingual face (1); no cones present (2). Two cones on the posterior face of p4 are present in all the species of Platyrrhinus and Vampyrodes. Sturnira erythromos and Uroderma magnirostrum have only one cone. Carollia subrufa lack both cones on the posterior face of p4. This character has not been used by other researchers. Character 49 — Labial and lingual cingulum of p4. Both cingula absent (0); only one cingulum present, either the lingual or the labial (1); both cingula present (2). Both cingula on p4 are pre- sent in the majority of Platyrrhinus species (e.g., P. dorsalis, P. vittatus "Southern"). In Platyr- rhinus aurarius, P. chocoensis, Sturnira erythro- mos, Uroderma magnirostrum, and Vampyrodes, only one cingulum is present on p4. Carollia sub- rufa lack cingula on p4. This character has not been used by other researchers. Character 50 — Stylid cusps on the anterior or anterolingual face of the main cone of p4. Absent (0); one stylid cusp present (1); two stylid cusps present (2). Two stylid cusps on the anterior face of the main cone of p4 are present in Pla- tyrrhinus brachycephalus. In Platyrrhinus vittatus "Northern" and Vampyrodes, one and two stylid cusps, respectively, are present on the anterolin- gual face. Platyrrhinus helleri "Eastern" has a single stylar cusp on the anterior face of p4. In contrast, other Platyrrhinus species (e.g., P. au- rarius, P. chocoensis), Carollia subrufa, Sturnira erythromos, and Uroderma magnirostrum lack stylid cusps on the anterior face of the main cone of p4. This is the first use of this character in a phylogenetic study. Character 51 — Stylid cusp on the posterior face of the main cone of p4. Absent (0); present (1). One or two stylid cusps are present on the posterior face of the main cone of p4 in some species of Platyrrhinus (e.g., P. helleri "Eastern," P. helleri "Western"). In contrast, other species of Platyrrhinus (e.g., P. aurarius, P. chocoensis), Carollia subrufa, Sturnira erythromos, Uroderma magnirostrum, and Vampyrodes lack stylid cusps on the posterior face of the main cone of p4. This character has not been used by other researchers. Character 52 — ml paraconid. Present (0); ab- sent (1). Most Platyrrhinus species (e.g., P. au- rarius, P. chocoensis), Carollia subrufa, Uroder- ma magnirostrum, and Vampyrodes lack a para- conid on ml. In contrast, Sturnira erythromos has a paraconid on ml. Platyrrhinus nigellus has in- traspecific variation. Wetterer et al. (2000: char- 16 FIELDIANA: ZOOLOGY acter 68) used this character in their phylogeny of the Phyllostomidae, but I do not agree on their score for Platyrrhinus. They scored it as a poly- morphism (with the paraconid occasionally pre- sent). However, my observations indicate that the majority of Platyrrhinus lack the paraconid on ml, as does Vampyrodes. Character 53 — ml cingulum. Absent (0); only the labial cingulum present (1); both labial and lingual cingula present (2). Both cingula on ml are present in all the species of Platyrrhinus. In contrast, Carollia subrufa and Sturnira erythro- mos lack both cingula on ml. Uroderma magni- rostrum and Vampyrodes have only the labial cin- gulum. This character has not been used in any other study. Character 54 — Stylid cusps on the anterior face of the ml protoconid. Absent (0); present (1). A stylid cusp on the anterior face of the ml protoconid is present in all species of Platyrrhin- us. In contrast, Carollia subrufa, Sturnira ery- thromos, Uroderma magnirostrum, and Vampy- rodes lack this stylid cusp on the anterior face of the ml protoconid. This character has not been used in any other study. Character 55 — ml metaconid. Well developed (0); poorly developed (1); absent (2). A well-de- veloped ml metaconid is present in Platyrrhinus infuscus, P. lineatus, and Sturnira erythromos. However, other species of Platyrrhinus (e.g., P. aurarius, P. recifinus), Uroderma magnirostrum, and Vampyrodes lack a metaconid on ml. Platyr- rhinus dorsalis "Norte," P. vittatus "Northern," P. vittatus "Southern," and Carollia subrufa have a poorly developed ml metaconid. Platyrrhinus nigellus has intraspecific variation, with some specimens lacking the ml metaconid and others with it poorly developed. I do not agree with the score of present that Wetterer et al. (2000: char- acter 69) assigned to Vampyrodes because, ac- cording to my observations, Vampyrodes lacks a metaconid on m 1 . Character 56 — m2 hypoconid. Present (0); ab- sent (1). The m2 hypoconid is present in Carollia subrufa and Sturnira erythromos. In contrast, the majority of the species of Platyrrhinus (e.g., P. brachycephalus, P. nigellus), Uroderma magni- rostrum, and Vampyrodes lack the hypoconid on m2. Platyrrhinus helleri "Eastern" has intraspe- cific variation. This character has not been used in other studies. Character 57 — Stylid cusp between the meta- conid and protoconid on m2. Absent (0); present (1). A stylid cusp between the metaconid and pro- toconid on m2 is present in some species of Pla- tyrrhinus (e.g., P. brachycephalus, P. vittatus "Southern") and Vampyrodes. In contrast, P. cho- coensis, P. infuscus, P. recifinus, Carollia subru- fa, Sturnira erythromos, and Uroderma magniros- trum lack this stylid cusp between the metaconid and protoconid on m2. This character has not been used in any other study. Character 58 — Cingula on m2. Both cingula labial and lingual absent (0); labial cingulum pre- sent (1); both cingula present (2). Both cingula on m2 are present in the majority of species of Pla- tyrrhinus (e.g., P. brachycephalus, P. infuscus) and Uroderma magnirostrum. In contrast, Vam- pyrodes has only the labial cingulum, and Car- ollia subrufa and Sturnira erythromos lack both cingula on m2. Platyrrhinus helleri "Western" has intraspecific variation, with some individuals with both cingula and others with only the labial cingulum. This character has not been used by other researchers. Postcranial Skeleton Character 59 — Third and fifth metacarpal arrangement. Length of M III > M V (0); length of M III = M V (1); length of M III < M V (2). Platyrrhinus aurarius, P. dorsalis "Norte," P. li- neatus, P. nigellus, P. vittatus "Northern," Stur- nira erythromos, and Vampyrodes have the third metacarpal shorter that the fifth metacarpal. In Platyrrhinus helleri "Eastern," P. helleri "West- ern," P. recifinus, and Uroderma magnirostrum, the third metacarpal is longer than the fifth meta- carpal. The third metacarpal is subequal to the fifth in P. vittatus "Southern." Platyrrhinus cho- coensis, P. dorsalis, P. dorsalis "Centro-Sur," and P. infuscus exhibit intraspecific variation: the third metacarpal can be subequal to or shorter than the fifth metacarpal. Platyrrhinus brachyce- phalus and Carollia subrufa also have intraspe- cific variation: the third metacarpal can be sube- qual to or longer than the fifth metacarpal. Wet- terer et al. (2000: character 84) used this character in their phylogeny of the Phyllostomidae; they in- dicated that in all species of Platyrrhinus, the third metacarpal was subequal in length to the fifth metacarpal. Simmons (1996: character 8) used this character in her phylogeny of the genus Micronycteris. Character 60 — Insertion of the posterior edge of the plagiopatagium. Onto the ankle re- gion (0); onto the first metatarsal (1). The pos- VELAZCO: PLATYRRHINUS PHYLOGENY 17 terior edge of the plagiopatagium inserts at the first metatarsal in all the species of Platyrrhinus (e.g.. P. helleri "Eastern."" P. recifinus), Urod- erma magnirostrum, and Vampyrodes. The pos- terior edge of the plagiopatagium inserts at the ankle region in Carollia subrufa and Sturnira er- ythromos. Owen (1987: character 6) and Wetter- er et al. (2000: character 84) used this character, and I agree with the scores assigned to Platyr- rhinus and the outgroup. Marques- Aguiar (1994: character 7) used this character in her phylogeny of the genus Artibeus, as did Straney (1980: characters G 9-13) in his phylogeny of the sub- family Phyllostominae. Results Parsimony analysis of the data resulted in two most parsimonious trees of 162 steps each (CI = 0.54: CI excluding uninformative characters = 0.52; RI = 0.58). A strict consensus of these trees with the bootstrap and decay values is shown in Figure 14. Monophyly of Platyrrhimis was sup- ported with bootstrap and decay values of 84% and 3%. respectively. Two main clades were observed within Platyr- rhinus. One is comprised of the smaller species + P. lineatus and P. recifinus and is supported weakly by a bootstrap value of 33% and a decay value of 1 . In this clade. the basal species P. line- atus and P. recifinus were unresolved, whereas the clade formed by P. helleri "Eastern" + P. brachycephalus + P. helleri "Western"' was sup- ported by a bootstrap value of 72% and a decay value of 1. The clade formed by P. brachyceph- alus + P. helleri "Western"" also was supported by a bootstrap value of 72% and a decay value of 1. The second main clade comprised of the re- maining nine species of Platyrrhinus was com- pletely resolved and was supported by a bootstrap value of 43% and a decay value of 2. Platyrrhinus chocoensis, P. dorsalis, and P. aurarius occupied the three most basal branches, followed by P. ni- gellus + P. dorsalis "Centro-Sur."* P. dorsalis "Norte." P. infuscus, P. vittatus "Northern. " and P. vittatus "Southern." There was strong support (bootstrap = 98% and decay = 6) for Vampyrodes caraccioli as sister taxon of Platyrrhinus. The analysis also supported (bootstrap = 100% and decay = 10) Uroderma magnirostrum as sister taxa of the clade com- prised of Platyrrhinus + Vampyrodes. The most basal branches of the tree were occupied by Car- ollia subrufa (subfamily Carolliinae) and Sturnira erythromos (most basal genus of the Stenoder- matinae). Discussion and Conclusions The present phylogenetic analysis strongly sup- ports Platyrrhinus monophyly but only weakly supports the interspecific relationships (Fig. 14). Owen"s (1987) phylogenetic hypothesis also sup- ported Platyrrhinus monophyly but did not in- clude all the species, nor did it provide support values. Three synapomorphies are described for the first time for Platyrrhinus: sulcus on the an- terior face of the P4. both labial and lingual cin- gula present on the ml. and stylid cusps present on the anterior face of the ml protoconid. Two main divisions of Platyrrhinus were evi- dent in both analyses. However, they were weak- ly supported, so I do not suggest recognizing them as separate subgenera. The first clade in- cludes all the smaller species of Platyrrhinus (P. brachycephalus + P. helleri "Eastern"" + P. hel- leri "Western"*). P. lineatus, and P. recifinus and is supported by six characters: poorly marked but distinguishable pinna folds (character 10). dense and long hair on the dorsum of the feet (15). a densely haired uropatagium edge (17). Ml par- astyle present (32). labial cingulum at the base of the Ml metacone present (34). and both labial and lingual cingula present on p4 (49). The pre- sent hypothesis strongly contradicts with Owen's interspecific phylogenetic hypothesis. Owen's (1987) preferred hypothesis placed P. brachy- cephalus and P. helleri (the only "small" spe- cies of Platyrrhinus recognized at that time) in different clades. In the present study. P. brachy- cephalus and the two forms of P. helleri were placed in the same clade. The second clade. containing the nine larger species of Platyrrhinus. is supported by seven synapomorphies: dark facial stripes (character 3). absence of the basal protuberance where the genal vibrissae are implanted (4). folds in the pinna well marked (10). dorsal stripe definite but narrow (11). posterior border of the hard palate "V"- shaped (18). postorbital process absent or poorly developed (19). and Ml protocone small and blunt (39). The present analysis supports the hypothesis of Velazco and Solari (2003). recognizing P. 18 FIELDIANA: ZOOLOGY 10 100 43 13 13 Carollia subrufa Sturnira erythromos Uroderma magnirostrum Vampyrodes caraccioli Platyrrhinus chocoensis Platyrrhinus dorsalis Platyrrhinus aurarius Platyrrhinus nigellus Platyrrhinus dorsalis "Centro-Sur" Platyrrhinus dorsalis "Norte" Platyrrhinus infuscus Platyrrhinus vittatus "Southern" • — Platyrrhinus vittatus "Northern" — Platyrrhinus lineatus — Platyrrhinus recifinus — Platyrrhinus helleri "Eastern" I — Platyrrhinus brachycephalus ■ Platyrrhinus helleri "Western" Fig. 14. Strict consensus tree of the analysis with the characters unordered. Decay values are given above each branch; bootstrap values are below. dorsalis as a complex of three different species (two not yet described). The analysis placed P. dorsalis close to P. chocoensis, which also is a species that occurs on the western slope of the Andes. Platyrrhinus dorsalis "Norte" and P. dorsalis "Centro-Sur" (the two undescribed spe- cies) were placed closer to P. nigellus and P. infuscus, species that are distributed on the east- ern slope of the Andes. No differences were found between the paratypes of P. umhratus and the specimens of P. dorsalis. Therefore, I treat P. umhratus as a junior synonym of P. dorsalis. Platyrrhinus dorsalis "Norte" and P. dorsalis "Centro-Sur" exhibit several autopomorphies that differentiate them from P. dorsalis and each other. Consequently, these two taxa are named and described here. Platyrrhinus lineatus and P. nigellus demonstrate clear morphological differ- ences and are located in different clades, sup- porting their validity as species. The branches representing P. helleri "Eastern" and P. helleri "Western" have sufficient distin- guishing characteristics to be considered different species. The distribution of P. helleri "Eastern" is the widest of the genus, comprising most of Central and South America and encompassing VELAZCO: PLATYRRHINUS PHYLOGENY 19 type localities of P. helleri (Peters, 1866), P. za- rhinus (Allen, 1891), and P. zarhinus incarum (Thomas, 1912). Comparisons between the de- scriptions of these species and P. helleri "East- ern" suggest that P. helleri is the appropriate bi- nomial name for P. helleri "Eastern," with P. zarhinus and P. zarhinus incarum representing ju- nior synonyms. Platyrrhinus helleri "Western," which is found in the Choco region in Colombia and Ecuador, does not have a name available and is described here. Platyrrhinus vittatus "Northern" and P. vitta- tus "Southern" are the largest forms in this genus. Platyrrhinus vittatus "Northern" is restricted to the lowlands in Costa Rica, Panama, Atlantic and Pacific Colombia, and Venezuela. Platyrrhinus vittatus "Southern" is distributed mainly in the highlands of Ecuador, Peru, and Bolivia. The ho- lotype of P. vittatus is from Puerto Cabello, Vene- zuela, and is distinguishable both qualitatively and morphometrically from P. vittatus "South- ern." Platyrrhinus vittatus "Southern" is larger than P. vittatus "Northern" and has qualitative morphological and morphometric differences. Platyrrhinus vittatus "Southern" is a taxon with- out an available name and therefore is described here. Like previous phylogenetic hypotheses for the family Phyllostomidae that placed Platyrrhinus and the monotypic genus Vampyrodes as sister taxa (Baker et al., 2000, 2003; Lim, 1993; Owen, 1987 [only discrete characters]; Smith, 1976; Van Den Bussche, 1992; Wetterer et al., 2000), the present study places Vampyrodes as a sister taxa of Platyrrhinus. Taxonomic Diagnoses of the Platyrrhinus Species A taxonomic diagnosis for 14 species of Pla- tyrrhinus used in the phylogenetic analysis is pre- sented. This includes descriptions of four new species. The diagnoses are based on the ACCT- RAN and DELTRAN optimizations of the data. Unambigously derived conditions are in italics. States with ambiguous optimizations are not ital- icized. All observations are based on adult indi- viduals, external and craniodental measurements are in millimeters, and body mass (weight) is in grams. The first five measurements listed here were obtained from skin tags or field notes made by the collector of each specimen; the other di- mensions were measured with a digital caliper to the nearest 0.01 mm. Measurements are defined as follows: Weight (W): Body mass in grams. Total length (TL): Distance from the snout to the tip of the last caudal vertebra. Hind foot length (HL): Distance from the an- terior edge of the base of the calcar to the tip of the claw of the longest toe. Ear length (E): Distance from the notch to the fleshy tip of the pinna. Forearm length (FA): Distance from the olec- ranon process to the carpals. Tibia length (Tibia): Distance from the prox- imal end of the tibia to the posterior base of the calcar. Greatest length of skull (GLS): Distance from the most posterior point of the occiput to the an- teriormost point of the premaxilla (excluding in- cisors). Condiloincisive length (CIL): Distance be- tween the posterior-most point of the occipital condyles and the anterior-most point on the upper incisors. Condilocanine length (CCL): Distance be- tween the posterior-most point of the occipital condyles and the anterior-most point on the upper canines. Postorbital breadth (PB): Least breadth across the frontals posterior to the postorbital processes or bulges. Zygomatic breadth (ZB): Greatest breadth across zygomatic arches. Braincase breadth (BB): Greatest breadth of the braincase, excluding the mastoid and paraoc- cipital processes. Mastoid breadth (MB): Greatest breadth across the mastoid region. Maxillary toothrow length (MTRL): From the anterior-most edge of the canine crown to the posterior-most edge of the crown of M3. Breadth across molars (BAM): Greatest breadth across the outer edges of the crowns of the upper molars. Measurements of male and females were com- bined in species comparisons because no differ- ences in size were apparent between males and females in any species of Platyrrhinus. Velazco and Solari (2003) evaluated sexual dimorphism in the Peruvian populations of P. dorsalis "Norte," P. dorsalis "Centro-Sur," and P. nigellus and concluded that these three species did not dem- onstrate significant sexual dimorphism. 20 FIELDIANA: ZOOLOGY Table 2. Measurements of the type series of Pla- tyrrhinus albericoi. Holotype MUSM 19149 Paratype FMNH 170145 Sex Female Female (pregnant) Weight 55 68 Total length 100 100 Hind foot length 16 15 Ear length 25 24 Forearm length 63 62 Tibia length 23.93 23.71 Greatest length of skull 32.84 32.51 Condyloincisive length 32.63 31.63 Condylocanine length 31.89 30.82 Postorbital beadth 7.65 7.39 Zygomatic breadth 20.85 20.68 Braincase breadth 13.38 13.58 Mastoid breadth 15.85 16.12 Maxillary toothrow length 13.89 13.44 Breadth across molars 15.23 15.57 Systematics Family Phyllostomidae Gray, 1825 Subfamily Phyllostominae Gray, 1825 Genus Platyrrhinus Saussure, 1866 Platyrrhinus albericoi, new species. Figures 1,16 (bottom) Vampyrops vittatus Gardner and Carter, 1972b: 74- 75 (part) Vampyrops vittatus Koopman, 1978: 11 Vampyrops vittatus Swanepoel and Genoways. 1979: 66, 104 (part) Vampyrops vittatus Barquez and Olrog, 1980: 53-54 Vampyrops vittatus Anderson et al., 1982: 7-8 Vampyrops vittatus Anderson, 1985: 7 Vampyrops vittatus Alberico, 1990: 349 (part) Platyrrhinus vittatus Anderson, 1993: 8, 24, 69 Platyrrhinus vittatus Anderson, 1997: 36, 244, 589 (part) Platyrrhinus vittatus Albuja, 1999: 134-135 (part) Platyrrhinus vittatus Emmons et al., 2001: 255 Platyrrhinus vittatus Romo, 2001: 258 Platyrrhinus vittatus Solari et al., 2001: 112, 263 Platyrrhinus vittatus Salazar-Bravo et al., 2003: 15 Platyrrhinus vittatus "Southern" (this paper) Type Material— The holotype, MUSM 19149, is an adult female collected by Sergio Solari (orig- inal field number SS 2067) on 14 April 2001 at San Pedro, Paucartambo-Pilcopata road, 1480 m in elevation, Province of Paucartambo, Depart- ment of Cuzco, Peru, approximately 13°0TS, 71°32'46'W. The type locality is in the Cultural ^x? • Platyrrtiinus albericoi 0 262.5 52: Fig. 15. Distributional map of Platyrrhinus albericoi. Zone of the Manu Biosphere Reserve. The skin, skull, and alcohol preserved carcass were in good condition. Frozen tissues are deposited at the Field Museum of Natural History (FMNH 172107). The paratype is an adult female (FMNH 170145) from Suecia, km 138.5 Carretera Shin- tuya, Province of Paucartambo, Department of Cuzco, Peru. The holotype and paratype as well as other specimens from the known distributional range are listed in Appendix 1. Dimensions of each specimen of the type series of Platyrrhinus albericoi are provided in Table 2. Distribution — Platyrrhinus albericoi is cur- rently known from the Eastern slope of the Andes in Ecuador, Peru, and Bolivia. Currently, the northern limit of its distribution is the Province of Pichincha (Ecuador), and the southern limit is the Department of Cochabamba (Bolivia). However, this taxon is expected to occur in southern Colom- bia (Fig. 15). The elevational range comprise from 1480 to 2500 m. Etymology — The specific epithet is chosen to honor Michael Alberico, who devoted his scien- tific career to the study of Colombian mammals. Diagnosis — Size large (FA = 62-63 mm; CIL = 31.6-32.6 mm; W = 55-68 g); facial stripes brilliant-white; folds in the pinna poorly marked but distinguishable: fringe of hair on the edge of the uropatagium dense; paraoccipital process well developed; Ml mesostyle present; M2 me- VELAZCO: PLATYRRHINUS PHYLOGENY 21 Table 3. Selected measurements^ of members of the genus Platyrrhinus. Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus aurarius brachycephalus chocoensis dorsalis helleri w 30.5 ± 0.7 13.6 ± 2.8 30.5 ± 2.1 14.5 ± 2.7 30-31 (2) 10-20 (20) 29-32 (2) 11-19 (10) TL 78.5 ± 4.9 58.3 ± 3.8 76.5 ± 2.1 75.5 ± 6.4 57.7 ± 3.9 75-82 (2) 52-68 (33) 75-78 (2) 71-80(2) 52-65 (15) HL 15.0 ± 0.8 11.0 ± 1.0 14.5 ± 0.7 13.2 ± 0.7 10.5 ± 0.9 14-17 (12) 9-13 (37) 14-15 (2) 12-15 (11) 8-12 (21) E 20.7 ± 0.6 16.3 ± 1.2 19.0 ± 1.4 18.8 ± 0.7 15.7 ± 1.4 20-22 (12) 13-19 (33) 18-20 (2) 18-20(11) 12-17 (14) FA 52.7 ± 1.1 37.6 ± 1.8 48.5 ± 0.7 48.9 ± 1.3 37.8 ± 1.5 51-54 (12) 33-42 (37) 48-49 (2) 46-50(11) 35-40 (21) Tibia 21.30 ± 0.76 15.36 ± 0.93 19.65 ± 1.72 20.32 ± 1.34 15.04 ± 0.69 20.00-22.00 (12) 13.18-17.60(33) 18.43-20.86 (2) 17.39-21.88 (11) 13.60-16.00 (21) GLS 27.23 ± 0.38 20.44 ± 0.50 27.33-0.49 26.42 ± 0.47 21.21 ± 0.75 26.53-27.87 (12) 19.50-21.53 (47) 26.77-27.69 (3) 25.46-27.40 (13) 19.04-22.50 (67) CIL 26.18 ± 0.39 19.08 ± 0.53 25.59 ± 0.84 25.06 ± 0.48 19.95 ± 0.80 25.62-27.01 (12) 18.30-20.26(47) 24.64-26.22 (3) 24.33-25.75 (13) 18.03-21.28 (66) CCL 25.55 ± 0.36 18.64 ± 0.50 25.18 ± 0.76 24.59 ± 0.48 19.49 ± 0.79 25.12-26.35 (12) 17.75-19.86 (47) 24.33-25.80 (3) 23.82-26.43 (13) 17.65-20.80(67) PB 6.65 ±0.12 5.39 ± 0.22 6.55 ± 0.22 6.32 ±0.12 5.44 ± 0.15 6.39-6.83 (12) 4.92-6.15 (48) 6.33-6.77 (3) 6.13-6.59 (13) 5.17-5.88 (67) ZB 16.89 ± 0.33 12.29 ± 0.55 16.91 ± 0.34 16.13 ± 0.67 12.56 ± 0.49 16.38-17.57 (9) 11.46-13.64 (38) 16.57-17.26(3) 14.88-17.10 (12) 11.49-13.54 (55) BB 11.85 ± 0.19 9.31 ± 0.30 11.82 ± 0.08 1 1 .52 ± 0.33 9.38 ± 0.27 11.55-12.28 (12) 8.76-9.94 (47) 11.74-11.89 (3) 10.74-11.93 (13) 8.62-9.89 (66) MB 14.01 ± 0.34 10.66 ± 0.32 13.75 ± 0.43 13.34 ± 0.33 10.88 ± 0.38 13.42-14.58 (12) 10.04-11.44(47) 13.26-14.06(3) 12.76-13.91 (13) 10.04-11.61 (66) MTRL 11.07 ± 0.26 7.37 ± 0.27 (41) 10.94 ± 0.29 10.68 ± 0.33 7.88 ± 0.40 10.71-11.55 (12) 6.82-7.94 (41) 10.70-11.25 (3) 10.17-11.34 (13) 6.96-8.59 (63) BAM 12.45 ± 0.33 8.75 ± 0.37 11.55 ± 0.12 1 1 .49 ± 0.49 9.02 ± 0.45 11.88-12.90(12) 8.11-9.72 (48) 11.45-11.69 (3) 10.88-12.60(13) 8.14-9.80(65) tastyle absent: length of the third metacarpal sub- equal to the fifth metacarpal. Description — The largest known species of Platyrrhinus (FA = 62-63 mm; GLS = 32.5-32.8 mm; Tables 2-3). Dorsal fur dark brown, ventral fur light brown, dorsal and ventral hair tricolored: dorsomedial facial stripes more marked than the ventrolateral: dorsal hairs > 8 mm long on the back: narrow dorsal stripe brighter than the facial ones; folds in the pinna poorly marked but distin- guishable: six vibrissae surrounding the margins of the noseleaf in a single array; four submental vibrissae present: one or two interramal vibrissae present: noseleaf longer than wide, inferior border of the horseshoe completely free of the upper lip: dense and long hair on the dorsum of the feet; "U" -shaped posterior edge of the uropatagium with a densely haired fringe, uropatagium extends along the midline 4.5-7 mm long: insertion of the posterior edge of the plagiopatagium onto the first metatarsal; third metacarpal subequeal in length with fifth metacarpal. Almost imperceptible fossa on the squamosal end of the zygomatic arch; postorbital process moderately developed; paraoccipital process well developed: upper median incisors convergent and in contact; sulcus on the posterior face of the up- per canines; two stylar cusps present on the pos- terior face of the P4; stylar cusp present on the cingulum of the lingual face of the Ml metacone; Ml metastyle and mesostyle present; Ml proto- cone moderately developed: stylar cusp present on the lingual face of the M2 paracone: M2 metastyle absent; M2 parastyle present: stylar cusp present on the lingual face of the M2 metacone: lingual cingulum of the M2 metacone restricted to the metacone; two sulci present on the lingual face of the main cone of p4; both labial and lingual cin- gula present on p4: stylid cusps absent on the an- terior or anterolingual face of the main cone of p4; ml paraconid absent; stylid cusp present on the anterior face of the ml protoconid: poorly de- veloped ml metaconid: m2 hypoconid absent; stylid cusp between the metaconid and protoconid 22 FIELDIANA: ZOOLOGY Table 3. Continued. Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus Platyrrhinus infuscus lineatus nigellus recifinus vittatus w 44.5 ± 5.1 21.9 ± 3.1 21.6 ± 3.1 17.7 ± 0.8 62.3 ± 2.5 36-59 (23) 18-28 (20) 19-30(13) 17-19 (6) 60-65 (3) TL 88.4 ± 6.4 70.1 ± 4.7 65.9 ± 3.3 91.3 ± 1.5 92.6 ± 7.3 77-105 (36) 60-78 (22) 60-74 (32) 89-93 (6) 83-100 (5) HL 15.5 ± 0.9 12.3 ± 0.9 12.1 ± 0.9 12.2 ± 0.7 14.9 ± 1.5 14-18 (57) 10-1 (28) 10-15 (33) 11-13 (6) 13-17 (7) E 22.4 ± 1.1 19.4 ± 1.4 18.5 ± 1.1 19.2 ± 0.7 24.0 ± 1 .0 20-24 (36) 15-2 (22) 16-21 (31) 18-20 (6) 23-25 (5) FA 57.9 ± 1.6 47.1 ± 0.8 43.0 ± 1.6 42.5 ± 0.5 59.8 ± 1.2 54-62 (52) 46-48 (22) 40-47 (33) 42-43 (6) 57-61 (7) Tibia 23.75 ± 1.13 19.51 ± 0.82 16.94 ± 0.73 16.28 ± 0.76 21.74 ± 0.90 20.80-26.00 (49) 18.12-21.00 (20) 15.62-18.46(30) 15.20-17.50 (6) 20.43-22.82 (7) GLS 29.74 ± 0.73 24.01 ± 0.46 24.43 ± 0.5 1 23.32 ± 0.29 30.57 ± 0.82 27.96-31.22 (63) 23.08-24.94 (29) 23.51-25.88 (35) 23.03-23.76 (6) 28.90-31.39 (7) CIL 28.32 ± 0.65 22.52 ± 0.39 23.04 ± 0.54 22.26 ± 0.26 29.63 ± 1.09 26.39-29.89 (61) 21.98-23.45 (24) 21.94-24.50(34) 21.99-22.61 (6) 28.12-31.29 (8) CCL 27.70 ± 0.63 21.89 ± 0.39 22.56 ± 0.54 21.73 ± 0.21 28.91 ± 1.03 25.93-29.01 (63) 21.36-22.79 (24) 21.57-23.98 (35) 21.55-22.06(6) 27.48-30.41 (8) PB 6.88 ± 0.20 6.31 ± 0.18 6.12 ± 0.17 5.74 ± 0.21 7.49 ± 0.24 6.32-7.34 (65) 5.98-6.61 (30) 5.61-6.41 (36) 5.46-6.10(6) 7.23-7.89 (8) ZB 18.39 ± 0.48 14.42 ± 0.35 14.30 ± 0.35 14.20 ± 0.33 19.07 ± 0.43 17.42-19.33 (60) 13.80-15.28 (26) 13.45-15.48 (35) 13.55-14.45 (6) 18.41-19.67 (8) BB 12.69 ± 0.28 10.77 ± 0.24 10.72 ±0.19 10.44 ± 0.21 13.17 ± 0.26 11.96-13.35 (62) 10.77-11.25 (29) 10.37-11.20(36) 10.18-10.80(6) 12.81-13.50(8) MB 15.15 ± 0.38 12.22 ± 0.33 12.32 ± 0.32 11.94 ± 0.30 15.43 ± 0.38 14.16-15.97 (63) 11.76-12.89 (25) 11.66-13.18 (35) 11.57-12.41 (6) 14.68-15.96 (7) MTRL 12.17 ± 0.28 8.91 ± 0.19 9.54 ± 0.30 8.96 ± 0.26 12.72 ± 0.45 11.53-12.79 (63) 8.57-9.30 (30) 8.96-10.16(31) 8.62-9.39 (6) 12.02-13.30(7) BAM 13.79 ± 0.39 10.13 ± 0.21 10.55 ± 0.29 10.41 ± 0.30 14.43 ± 0.57 12.88-14.52 (63) 9.62-10.83 (31) 10.04-11.09 (36) 9.90-10.71 (6) 13.59-15.37 (8) ■ Weight is in grams, all other measurements are in millimeters. b Summary statistics (mean and standard deviation [above], observed range and sample size [below]) of measure- ments for each species (see Appendix 1 for a list of the specimens measured). of m2 present; both labial and lingual cingula pre- sent on m2. Comparisons — Platyrrhinus albericoi is found sympatrically with P. dorsalis "Norte," P. dor- salis "Centro-Sur," and P. nigellus. Platyrrhinus albericoi is easily distinguishable from these three species by its larger forearm length and greatest length of skull: P. albericoi (FA = 62-63 mm; GLS = 32.5-32.8 mm), P. dorsalis "Centro-Sur" (FA = 45-51 mm; GLS = 25.0-26.8 mm), and P. nigellus (FA = 40-47 mm; GLS = 23.5-25.9 mm). Platyrrhinus albericoi has been traditionally confused with P. vittatus, so subsequent compar- isons are restricted to these two large species. Externally, Platyrrhinus albericoi can be distin- guished from P. vittatus by darker dorsal fur and brighter and wider facial stripes in P. albericoi. Moreover, folds in the pinna are poorly marked but distinguishable in P. albericoi, whereas in P. vittatus the folds in the pinna are well marked. The dorsal stripe is white and narrow in P. alber- icoi, whereas in P. vittatus the dorsal stripe is white and wide. The posterior edge of the uro- patagium always is "U" shaped in P. albericoi, whereas it can be "U" or "V" shaped in P. vit- tatus. A densely haired uropatagium fringe is pre- sent in P. albericoi, whereas the uropatagium fringe is usually hairy, occasionally sparsely in P. vittatus. The third metacarpal is greater in length than the fifth metacarpal in P. albericoi but is sub- equal in length in P. vittatus. Cranially, Platyrrhinus albericoi and P. vittatus are very similar. The only apparent differences are that the cranium of P. albericoi is more robust than that of P. vittatus, and the paraoccipital pro- cess is better developed in P. albericoi. Dentally, Platyrrhinus albericoi and P. vittatus differ as follows: Ml mesostyle present in P. al- bericoi and absent in P. vittatus; moderately de- veloped Ml protocone in P. albericoi, whereas in VELAZCO: PLATYRRHINUS PHYLOGENY 23 P. vittatus the protocone on M 1 tends to be better developed; M2 metastyle absent in P. albericoi and present in P. vittatus; stylid cusps absent on the anterior or anterolingual face of the main cone of p4 in P. albericoi, whereas one stylid cusp on the anterolingual face of the main cone of p4 is present in P. vittatus (Fig. 16). Platyrrhinus aurarius (Handley and Ferris, 1972) Vampyrops aurarius Handley and Ferris, 1972. Type locality: km 125, 85 km SSE El Dorado, State of Bolivar, Venezuela. Holotype — Vampyrops aurarius USNM 387163, an adult male preserved as skin and skull, was collected by Merlin D. Tuttle and Arden L. Tuttle on 18 May 1966 at km 125, 85 km SSE El Dorado, State of Bolivar, Venezuela, elevation 1000 m. Distribution — Endemic to the Guianan Shield (southern Venezuela, Guyana, and Suriname), from 140 to 1250 m (Fig. 17). Diagnosis — Size medium (FA = 51-54 mm; CIL = 25.6-27.0 mm; W = 30-31 g; Table 3); dorsal fur tetracolored; postorbital process mod- erately developed; paraoccipital process well de- veloped; only the labial cingulum present on p4. Platyrrhinus brachycephalus (Rouk and Carter, 1972) Vampyrops brachycephalus Rouk and Carter, 1972. Type locality: 3 mi S Tingo Maria, Department of Huanuco, Peru. Vampyrops latus Handley and Ferris, 1972. Type lo- cality: San Juan, Province of Oxapampa, Depart- ment of Pasco, Peru (USNM 364408). Vampyrops latus saccharus Handley and Ferris, 1972. Type locality: Manacal, 5 km S and 25 km E Ca- riipano, State of Sucre, Venezuela (USNM 408411). Holotype — Vampyrops brachycephalus TCWC 12193, adult male preserved as skin and skull, was collected by D. C. Carter (original field num- ber 5513) on 28 August 1964 at 3 mi S Tingo Maria, Department of Huanuco, Peru, elevation 2400 ft (731 m). Distribution — Known from the Amazon Basin (Brazil, Bolivia, Colombia, Ecuador, French Gui- ana, Guyana, Peru, Suriname, and Venezuela) and the mountain forests in Ecuador and Peru, from 90 to 1295 m (Fig. 17). Diagnosis — Size small (FA = 33-42 mm; CIL = 18.3-20.3 mm; W = 10-20 g) (Table 3); one vibrissa present on the upper lip, ventral to the vibissae that surround the margin of the noseleaf; five submental vibrissae present on each side of the chin; ventral fur tricolored; fringe of hair on the edge of the uropatagium usually hairy, occa- sionally sparsely; a well-developed Ml proto- cone; two stylid cusps present on the anterior face of the main cone of p4. Platyrrhinus chocoensis Alberico and Velasco, 1991 Platyrrhinus chocoensis Alberico and Velasco, 1991. Type locality: Quebrada El Platinero, 12 km W Ist- mina (by road), Department of Choco, Colombia. Holotype — Platyrrhinus chocoensis UV 3817, adult male preserved as skin and skull, was col- lected by M. S. Alberico (original field number, MSA 1316) on 16 April 1984 at Quebrada El Pla- tinero, 12 km W Istmina (by road), Department of Choco, Colombia, elevation 100 m. Distribution — Found in the lowlands of the Choco biogeographic region of Colombia, to northwestern Ecuador, from 35 to 305 m (Fig. 1 8). Diagnosis — Size medium (FA = 48-49 mm; CIL = 24.6-26.2 mm; W = 29-32 g; Table 3); ventral fur bicolored; Ml protocone small and blunt; only the labial cingulum present on the p4; stylid cusp between the metaconid and protoconid of the ml present. Platyrrhinus dorsalis (Thomas, 1900) Vampyrops dorsalis Thomas, 1900. Type locality: Paramba, Province of Imbabura, Ecuador. Vampyrops umbratus Lyon, 1902. Type locality: San Miguel, Department of La Guajira, Colombia (MCZ B8180). Vampyrops oratus Thomas, 1914. Type locality: "Galifari" Picacho de Galipan, Cerro del Avila, Distrito Federal, Venezuela (BMNH 14.7.27.1). Vampyrops aquilus Handley and Ferris, 1972. Type locality: On the head of the Rio Pucro, Cerro Mali, Province of Darien, Panama (USNM 338025). Holotype — Vampyrops dorsalis BMNH 99.12.5.1, juvenile male preserved as skin and skull, was collected by R. Miketta (original field number 61) on 14 April 1899 at Paramba, Prov- ince of Imbabura, Ecuador, elevation 1100 m. Distribution — From southern Panama, along both slopes of the Andes in Colombia, and only the western slope in Ecuador, and northern Vene- zuela, from 230 to 2012 m (Fig. 18). Diagnosis — Size medium (FA = 46-50 mm; CIL = 24.3-25.7 mm; Table 3); folds in the pinna 24 FIELDIANA: ZOOLOGY Fig. 16. Labial views of the left p2 and p4 illustrating taxonomic differences in the presence or absence of accessory cusps on the anterolingual face of the main cone of p4. (Top) Platyrrhinus vittatus (LSUMZ 25465) with one accessory cusp (arrow). (Bottom) Platyrrhinus albericoi (FMNH 170145) without accessory cusp (arrow). absent; Ml protocone small and blunt; only the labial cingulum present on p4; stylar cusp present on the lingual face of the M2 metacone; both la- bial and lingual cingula present on p4. Platyrrhinus helleri (Peters, 1866) Vampyrops helleri Peters, 1866. Type locality: Mex- ico. Vampyrops zarhinus Allen, 1891. Type locality: Obis- po,'Canal Zone, Panama (MCZ 321 1). Vampyrops zarhinus incarum Thomas, 1912. Type lo- cality: Pozuzo, Department of Pasco, Peru (BMNH 12.1.15.1). Syntype — Vampyrops helleri ZMB 3276, adult, preserved as skin with the skull not removed, was collected by Heller in 1850 in Mexico. Distribution — From Mexico (Oaxaca and Ve- racruz) to Peru, Bolivia, and Amazonian Brazil, plus Trinidad and Tobago, from 160 to 1295 m (Fig. 19). Diagnosis — Size small (FA = 35-40 mm; CIL = 18.0-21.3 mm; W = 1 1-19 g; Table 3); ventral fur bicolored; hair on the dorsum of the feet in- termediate in density and length; Ml protocone small and blunt; one stylid cusp present on the anterior face of the main cone of p4. VELAZCO: PLATYRRHINUS PHYLOGENY 25 A Platyrrhinus aurarius # Platyrrhinus brachycephalus ^> ^ Fig. 17. Distributional map of Platyrrhinus aurarius and P. brachycephalus. Platyrrhinus infuscus (Peters, 1880) Vampyrops infuscus Peters. 1880. Type locality: Gru- ta de Ninabamba. Hacienda Ninabamba. Province of Hualgayoc. Department of Cajamarca. Peru. Vampyrops fumosus Miller. 1902. Type locality: Hui- tanaa. upper Purus River, State of Amazonas. Bra- zil (USNM 105530). Vampyrops intermedins Marinkelle. 1970. Type lo- cality: Mina de Upin. near Restrepo. Department of Meta. Colombia (Universidad de los Andes, Co- lombia 14885). Holotype — The holotype collected by L. Tac- zanowski from Gruta de Ninabamba, Hacienda Ninabamba. Province of Hualgayoc, Department of Cajamarca, Peru, and deposited in the Warsaw Museum (Polska Akademia Nauk, Instytut Zool- ogiczny) was apparently destroyed by fire, per- haps as early as 1936 (Gardner & Carter. 1972b). In 1972, Gardner and Carter designated a neo- type: TCWC 12199. adult male, preserved as skin and skull, collected by D. C. Carter (original field number 5477) on 26 August 1964 at 2 mi N Tingo Maria, Province of Leoncio Prado, De- partment of Huanuco. Peru, elevation 2000 ft (610 m). Distribution — Colombia to Peru, Bolivia, and NW Brazil, from 183 to 1900 m (Fig. 20). Diagnosis — Size large (FA = 54-62 mm; CIL = 26.4-29.9 mm: W = 36-59 g: Table 3); dorsal stripe indistinct, thin and obscured; dorsal fur short, <6.30 mm; sparse and short hair on the dorsum of the feet: uropatagium fringe usually hairy, occasionally sparsely; paraoccipital process well developed; Ml parastyle present; ml meta- 26 FIELDIANA: ZOOLOGY • Platyrrhinus chocoensis ▲ Platyrrhinus dorsalis Fig. 18. Distributional map of Platyrrhinus chocoensis and P. dorsalis. conid developed; stylid cusp absent between the metaconid and protoconid of m2. Platyrrhinus ismaeli, new species Figure 21 (top) 23 cf. Vampyrops aurarius Lemke et al., 1982: 230-231 Vampyrops sp. C Alberico, 1990: 349-352 Platyrrhinus dorsalis Pacheco and Patterson, 1991: 101, 104, 111, 112, 113, 114, 121 Platyrrhinus dorsalis Albuja, 1999: 135-136 (part) Platyrrhinus "dorsalis norte" Solari et al., 2001: 263 Platyrrhinus dorsalis "Norte" Velazco and Solari, 2003 Platyrrhinus dorsalis "Norte" (this paper) Type Material — The holotype, MUSM 4946, is an adult collected by Bruce D. Patterson (orig- inal field number BDP 2484) on 2 May 1987 at 19 km E of Balsas, 1945 m (6380 ft) in elevation, Province of Chachapoyas, Department of Ama- zonas, Peru. The skin, skull, and skeleton were in good condition. Frozen tissues are deposited at the Field Museum of Natural History (FMNH 129138). Paratypes include five adult specimens: one fe- male (FMNH 129134) and one male (FMNH 129136) from the type locality; and two females (FMNH 129139, 129146) and one male (FMNH VELAZCO: PLATYRRHINUS PHYLOGENY 27 Fig. 19. Distributional map of Platyrrhinus helleri. 129143) from Hacienda Limon, W of Balsas, Province of Celendin, Department of Cajamarca, Peru. The holotype and paratypes along with other specimens from the known distributional range are listed in Appendix 1 . Measurements of each specimen of the type series of Platyrrhinus is- maeli are provided in Table 4. Distribution — Platyrrhinus ismaeli (Fig. 21) is currently known from both slopes of the Andes in Colombia, Ecuador, and Peru. The northern limit of the distribution is the Department of Huila (Co- lombia), and the southern limit is the Department of San Martin (Peru). The elevation is from 1230 to 2950 m (Fig. 22). Etymology — Platyrrhinus ismaeli is named in honor of Ismael Ceballos Bendezii, an interna- tionally well-known naturalist from Cuzco, Peru, in recognition of his important contributions to the study of Peruvian bats. Diagnosis — Size medium (FA = 50-56 mm; CIL = 26.6-28.2 mm; W = 30-51 g); folds in the pinna absent; sulcus on the posterior face of the paracone not joined to the cingulum of the lingual face of the metacone on Ml; ml meta- conid poorly developed. Description — A medium-sized species of Pla- tyrrhinus (FA = 50-56 mm; GLS = 27.8-30.1 mm) with dark brown dorsal fur, grayish ventral 28 FIELDIANA: ZOOLOGY • Platyrrhinus infuscus A Platyrrhinus lineatus Fig. 20. Distributional map of Platyrrhinus infuscus and P. lineatus. fur, and tricolored dorsal and ventral hair. Dor- somedial and ventrolateral facial stripes obviously subequal and darkish, dorsal hairs 8-10 mm long on the back; narrow dorsal stripe brighter than the facial ones; folds in the pinna poorly marked but distinguishable; six vibrissae surrounding the margins of the noseleaf in a single array; four sub- mental vibrissae present; one interramal vibrissa present; noseleaf longer than wide, inferior border of the horseshoe completely free of the upper lip; dense and long hair on the dorsum of the feet; "U"-shaped posterior edge of the uropatagium with a densely haired fringe, uropatagium extends along the midline 4.5-7 mm long; insertion of the posterior edge of the plagiopatagium onto the first metatarsal; third metacarpal shorter than fifth metacarpal; fossa on the squamosal end of the zy- gomatic arch shallow; upper median incisors con- vergent and in contact; sulcus present on the pos- terior face of the upper canines; two stylar cusps present on the posterior face of P4; labial cingu- lum present at the base of the metacone of Ml; stylar cusp present on the cingulum of the lingual face of the Ml metacone; sulcus on the posterior face of the paracone not joined to the cingulum of the lingual face of the metacone on Ml; Ml metastyle present; Ml protocone small and blunt; stylar cusp present on the lingual face of the M2 paracone; M2 metastyle present; two sulci present on the lingual face of the main cone of p4; ml paraconid absent; ml entoconid present; stylid cusp present on the anterior face of the ml pro- VELAZCO: PLATYRRHINUS PHYLOGENY 29 Table 4. Measurements of the type series of Platyrrhinus ismaeli. Holotvpe Paratvpe Paratvpe Paratvpe Paratvpe Paratvpe MUSM FMNH FMNH FMNH FMNH FMNH 4946 129134 129136 129139 129143 129146 Sex Male Female Male Female Male Female Weight 35 40 38 35 30 31 Total length 84 82 87 82 83 79 Hind foot length 18 13 18 15 16 17 Ear length 20 21 22 20 21 21 Forearm length 52 53 54 54 52 56 Tibia length 21.68 19.52 21.68 21.27 20.56 21.31 Greatest length of skull 27.94 27.77 28.15 28.14 28.47 28.17 Condyloincisive length 27.10 26.57 27.47 27.06 26.83 26.62 Condylocanine length 26.22 26.07 26.88 26.49 26.14 26.06 Postorbital breadth 6.46 6.27 6.57 6.43 6.36 6.65 Zygomatic breadth 17.13 16.93 18.21 17.31 17.33 16.41 Braincase breadth 12.00 11.57 12.29 12.03 12.02 11.70 Mastoid breadth 13.99 13.63 14.72 14.04 13.98 14.06 Maxillary toothrow length 1 1 .35 11.43 11.66 11.75 11.33 11.48 Breadth across molars 12.63 12.65 12.81 13.08 12.45 12.41 toconid: poorly developed ml metaconid: m2 hy- poconid absent: stylid cusp present between the metaconid and protoconid of m2: both labial and lingual cingula present on m2. Velazco (2001) and Velazco and Solari (2003) did not find secondary sexual dimorphism in populations of P. ismaeli from Peru in either qualitative morphologic or morphometric characters. Fig. 21. Frontal view of Platyrrhinus ismaeli: adult male photographed at Hacienda Limon. Cajamarca. Peru, by B. D. Patterson. 30 FIELDIANA: ZOOLOGY Fig. 22. Distributional map of Platyrrhinus ismaeli. Comparisons — In several localities Platyr- rhinus ismaeli is found sympatrically with P. al- bericoi and P. nigellus, and it occurs with P. dor- salis in one locality at El Parque Nacional Nat- ural de la Cueva de Los Guacharos, Department of Huila, in Colombia. Platyrrhinus ismaeli is easily distinguished from P. albericoi and P. ni- gellus by its forearm length and greatest length of skull: P. albericoi (FA = 62-63 mm; GLS = 32.5-32.8 mm) and P. nigellus (FA = 40-47 mm; GLS - 23.5-25.9 mm; Tables 2-4). It dif- fers from P. dorsalis in its GLS, condiloincisive length, condilocanine length, and maxillary toothrow length (Tables 3-4). Platyrrhinus is- maeli has traditionally been confused with P. dorsalis, so subsequent comparisons are restrict- ed to these two species. Externally, Platyrrhinus ismaeli can be distin- guished from P. dorsalis by two characters. Pla- tyrrhinus ismaeli has dense and long hair on the dorsal side of the feet, whereas the hair in the dorsum of the feet is sparsely distributed and with some long hairs in P. dorsalis. The hairs on the edge of the uropatagium are densely haired and longer than 2. 1 mm in P. ismaeli, whereas P. dor- salis presents a regular or occasionally sparsely VELAZCO: PLATYRRHINUS PHYLOGENY 31 haired uropatagium fringe, with hairs shorter than 1.5 mm. Cranially, Platyrrhinus ismaeli and P. dorsalis are very similar. Platyrrhinus ismaeli has a more robust cranium with a wider rostrum. Moreover, P. ismaeli has a shallow fossa on the squamosal end of the zygomatic arch, lateral to the glenoid fossa, whereas P. dorsalis has a deep fossa. Dentally, Platyrrhinus ismaeli and P. dorsalis exhibit some differences. Whereas in P. ismaeli the sulcus on the posterior face of the paracone not joined to the cingulum of the lingual face of the metacone on Ml, this discontinuity is absent in P. dorsalis (Fig. 23). The lingual cingulum of the M2 metacone is restricted to the metacone in P. ismaeli, whereas in P. dorsalis the lingual cin- gulum of the M2 metacone continuous to the M2 paracone. Finally, a poorly developed metaconid on m 1 is present in P. ismaeli, whereas P. dorsalis lacks the ml metaconid. Comments — In his systematic review of Pla- tyrrhinus from the northwestern South America, Alberico (1990) identified Platyrrhinus sp. C as a potentially new species from Colombia. Platyr- rhinus sp. C (FA = 50.9-53.2 mm) was collected at several localities in the Cueva de los Guacharos National Park near the headwaters of the Mag- dalena River at midelevations (ca. 1500 m) in Co- lombia. Platyrrhinus ismaeli has been collected at Cueva de los Guacharos National Park, and the range of variation of the FA size in Platyrrhinus sp. C falls within the range of P. ismaeli (FA = 50-56 mm). Therefore, it is likely that Platyr- rhinus sp. C is conspecific with P. ismaeli. Platyrrhinus lineatus (E. Geoffroy, 1810) Phyllostoma lineatum E. Geoffroy St.-Hilaire, 1810. Type locality: Asuncion, Paraguay. Vampyrops lineatus sacrillus Thomas, 1 924. Type lo- cality: Rio Doce, State of Espfrito Santo, Brazil (BMNH 23.12.12.9). Holotype — Phyllostoma lineatum MNHN 953, adult, preserved as skin and skull, was collected by Felix de Azara in Asuncion, Paraguay. Distribution — Lowlands from eastern and southeastern Brazil, northern Uruguay, northeast- ern Argentina, Paraguay, and Bolivia, from 122 to 971 m (Fig. 20). Diagnosis — Size medium (FA = 46-48 mm; CIL = 22.0-23.4 mm; W = 18-28 g; Table 3); dense and long hair on the dorsum of the feet; postorbital process well developed; paraoccipital process well developed; ml metaconid developed. Platyrrhinus masu, new species. Figure 24 Vampyrops dorsalis Sanborn, 1951: 10 Vampyrops dorsalis Barquez and Orlog, 1980: 54 Vampyrops dorsalis Anderson et al., 1982: 7 Vampyrops dorsalis Anderson, 1985: 7 Platyrrhinus dorsalis Anderson, 1993: 8, 23, 69 Platyrrhinus dorsalis Anderson et al., 1993: 14, 24 Platyrrhinus dorsalis Anderson, 1997: 36, 37, 239, 240 (part) Platyrrhinus dorsalis Emmons et al., 2001: 255 Platyrrhinus dorsalis Romo, 2001: 258 Platyrrhinus cf. dorsalis Solari et al., 2001: 1 12 Platyrrhinus "dorsalis sur" Solari et al., 2001: 263 Platyrrhinus dorsalis Salazar-Bravo et al., 2003: 15 Platyrrhinus dorsalis "Centro-Sur" Velazco and So- lari (2003) Platyrrhinus dorsalis "Centro-Sur" (this paper) Type Material — The holotype, FMNH 123917, is an adult female collected by David E. Willard (original field number DEW 425) on 18 October 1981 at Consuelo, km 165, 17 km by road west of Pilcopata, Province of Paucartambo, Department of Cuzco, Peru, approximately 13°07'59'S, 71°15'W The type locality is in the Cultural Zone of the Manu Biosphere Reserve. The holotype is a specimen preserved in alcohol with the skull removed and cleaned. The skull and body were in good condition. Paratypes include six adult specimens: one fe- male (FMNH 172100) from San Pedro, Paucar- tambo-Pilcopata road, Province of Paucartambo, Department of Cuzco, Peru; one male (MVZ 166595) from Kiteni, Rio Urubamba, Province of La Convencion, Department of Cuzco, Peru; two females (MUSM 14562, 14566) from Campamen- to Llactahuaman, Kimbiri, Province of La Con- vencion, Department of Cuzco, Peru; and two males (MUSM 18265, 18266) from Cordillera de Carpish, Chinchao, Province of Huanuco, Depart- ment of Huanuco, Peru. The holotype, paratypes, and other specimens from the known distribution- al range are listed in Appendix 1. Measurements of each specimen of the type series of Platyrrhi- nus masu are provided in Table 5. Distribution — Platyrrhinus masu (Fig. 24) is currently known from the eastern slope of the An- des in Bolivia and Peru. The northern limit of the distribution is the Department of Huanuco (Peru) and the southern limit is the Department of La Paz (Bolivia). The elevational range extends from 650 to 3350 m (Fig. 25). Etymology — From the Quechua word "masu," meaning bat. Most of the collection lo- calities of Platyrrhinus masu have Quechua as the main language. 32 FIELDIANA: ZOOLOGY Fig. 23. Occlusal views of the left Ml and M2 illustrating taxonomic differences in the presence or absence of a connection between the sulcus on the posterior face of the paracone on Ml and the cingulum of the lingual face of the metacone on Ml. (Top) Platyrrhinus ismaeli (MUSM 4946) not connected (arrow). (Bottom) Platyrrhinus dorsalis (FMNH 128141) connected (arrow). Diagnosis — Size medium (FA = 45-51 mm; CIL = 24.0-25.5 mm; W = 23-33 g; Table 6); two submental vibrissae on each side of the chin; dorsal stripe brilliant-white and wide; dorsal hairs on the back 6.3-7.5 mm; upper lateral incisors bilobed. Description — A medium-sized species of Pla- tyrrhinus (FA = 45-51 mm; GLS = 25.0-26.8 mm; Table 6). Dorsal fur dark brown, ventral fur grayish, dorsal and ventral hair tricolored; dorso- medial and ventrolateral facial stripes obviously subequal and darker; dorsal hairs 6.3-7.5 mm long on the back; narrow dorsal stripe brighter that the facial ones, starting at the end of the dor- somedial facial stripes and ending on the rump; dorsal stripe definite but narrow; folds in the pin- na well marked; six vibrissae surrounding the margins of the noseleaf in a single array; two sub- mental vibrissae present on each side of the chin; one interramal vibrissae present; noseleaf longer than wide; dense and long hair on the dorsum of the feet; "U"-shaped posterior edge of the uro- patagium with a densely haired fringe, uropata- gium extends along the midline 4.5-7 mm long; insertion of the posterior edge of the plagiopata- gium onto the first metatarsal; third metacarpal VELAZCO: PLATYRRHINUS PHYLOGENY 33 Table 5. Measurements of the type series of Platyrrhinus masu. Holotype Paratvpe Paratvpe Paratvpe Paratype Paratvpe Paratvpe FMNH FMNH MVZ MUSM MUSM MUSM MUSM 123917 172100 166595 14562 14566 18265 18266 Sex Female Female Male Female Female Male Male Weight — 28 30 26 28 28 33 Total length 73 75 82 75 70 73 77 Hind foot length 12 14 15 14 13 13 14 Ear length 18 18 18 19 20 17 19 Forearm length 47 50 48 50 50 49 51 Tibia length 19.58 19.03 20.41 19.13 19.28 20.15 19.63 Greatest length of skull 26.03 26.42 26.14 26.70 26.84 26.53 26.41 Condyloincisive length 24.87 24.77 25.52 25.40 25.13 25.35 25.51 Condylocanine length 24.39 24.19 24.87 24.69 24.54 24.87 25.04 Postorbital breadth 6.19 6.50 6.57 6.30 6.39 6.14 6.10 Zygomatic breadth 16.02 16.52 16.11 16.78 15.93 16.31 16.96 Braincase breadth 1 1 .33 11.62 11.93 11.74 11.56 11.70 11.76 Mastoid breadth 13.19 13.09 13.71 13.60 13.50 13.81 13.91 Maxillary toothrow length 10.37 10.33 10.53 10.69 10.59 10.96 11.11 Breadth across molars 11.82 12.11 11.55 12.58 12.02 12.17 12.78 shorter than or subequal to fifth metacarpal; post- orbital process absent or poorly developed: zy- gomatic arches slightly divergent; paraoccipital process well developed; upper median incisors convergent and in contact; upper lateral incisors bilobed; sulcus present on the posterior face of the upper canines; two stylar cusps present on the posterior face of P4; labial cingulum present at Fig. 24. Frontal view of Platyrrhinus masu (FMNH 174760): adult male photographed at La Esperanza in the Cultural zone of the Manu Biosphere Reserve. Cuzco. Peru, by B. D. Patterson. 34 FIELDIANA: ZOOLOGY Fig. 25. Distributional map of Platyrrhinus masu. the base of the metacone on M 1 ; sulcus present on the posterior face of the M 1 paracone; stylar cusp present on the cingulum of the lingual and labial face of the Ml metacone; Ml metastyle present; moderately developed Ml protocone; sty- lar cusp present on the lingual face of the M2 paracone; M2 metastyle present; lingual cingulum of the M2 metacone restricted to the metacone; M2 hypoconal basin developed; two sulci present on the lingual face of the main cone of p4; two cones present on the posterior face of p4; both labial and lingual cingula present on p4; both la- bial and lingual cingula present on m2; ml para- conid absent; ml entoconid present; stylid cusp on the anterior face of the ml protoconid; ml metaconid absent; m2 hypoconid present; stylid cusp present between the metaconid and proto- conid of m2; both labial and lingual cingula pre- sent on m2. Velazco (2001) and Velazco and So- lari (2003) did not find secondary sexual dimor- phism in either qualitative morphologic or mor- phometric characters. Comparisons — Platyrrhinus masu is found sympatrically with P. albericoi and P. nigellus in several localities. Morphometrically, P. masu is easily distinguished from P. albericoi, but there is overlap in some measurements of masu and ni- gellus: P. albericoi (FA = 62-63 mm; CIL = 31.6-32.6 mm; W = 55-68 g), P. nigellus (FA = 40-47 mm; CIL = 21.9-24.5 mm; W = 19-30 g), and P. masu (FA = 45-51 mm; CIL = 23.0- 25.5 mm; W = 23-33 g). Platyrrhinus masu has been traditionally confused with P. dorsal is; how- ever, it also exhibits overlap of measurements VELAZCO: PLATYRRHINUS PHYLOGENY 35 Table 6. Selected measurements"11 of Platyrrhinus ismaeli and P. masu. Platyrrhinus ismaeli Platyrrhinus masu Weight Total length Hind foot length Ear length Forearm length Tibia length Greatest length of skull Condyloincisive length Condylocanine length Postorbital breadth Zygomatic breadth Braincase breadth Mastoid breadth Maxillary toothrow length Breadth across molars 37.1 30-51 86.7 78-98 15.5 13-18 21.2 20-22 52.7 50-56 21.97 18.34- 28.53 27.77- 27.19 26.57- 26.57 26.06- 6.48 6.17 17.21 16.25 11.97 11.57 14.07 13.43 11.84 11.33 13.08 12.40 ± 5.2 (19) ± 5.3 (20) ± 1.6 (20) ± 0.7 (19) ± 1.4 (20) ± 0.87 -21.96 (20) ± 0.56 30.08 (20) 0.43 28.33 (20) ± 0.44 27.59 (20) ± 0.20 6.86 (20) ± 0.49 18.21 (20) ± 0.23 12.29 (20) ± 0.30 ■14.72 (20) ± 0.34 12.48 (20) ± 0.46 -13.83 (20) 27.0 23-33 74.3 70-82 13.6 12-16 19.2 16-21 48.4 45-51 19.44 17.35- 26.02 24.96- 24.87 23.98- 24.36 23.45- 6.33 5.99 16.13 15.62 11.54 11.19 13.46 13.09 10.57 10.17 12.05 11.55 ± 2.7 (21) ± 3.2 (25) ± 0.9 (26) ± 1.4 (25) ± 1.5 (26) ± 0.80 -20.86 (26) ± 0.53 -26.84 (26) ± 0.44 25.52 (26) ± 0.41 25.04 (26) ± 0.18 -6.61 (26) ± 0.34 16.96 (25) ± 0.21 -11.93 (26) ± 0.22 -13.91 (26) ± 0.22 11.11 (25) ± 0.28 12.78 (25) ■ Weight is in grams, all other measurements are in millimeters. b Summary statistics (mean and standard deviation [above], observed range and sample size [below]) of measure- ments for each species (see Appendix 1 for a list of the specimens measured). with P. ismaeli and P. nigellus (Tables 3, 6). Therefore, comparisons will focus on differentia- tion of these four species. Externally, Platyrrhinus masu can be distin- guished from these species by three submental vi- brissae on each side of the chin in P. masu, whereas P. dorsalis, P. ismaeli, and P. nigellus have four submental vibrissae on each side of the chin; inferior border of the noseleaf partially joined to the upper lip in some individuals and completely free in others in P. masu, whereas in P. dorsalis, P. ismaeli, and P. nigellus, the infe- rior border of the noseleaf is completely free in all individuals; parallel folds in the pinna well marked in P. masu and P. nigellus, whereas in P. dorsalis and P. ismaeli, the parallel folds are poorly marked but distinguishable; dorsal stripe white and wide in P. masu and P. nigellus, where- as in P. dorsalis and P. ismaeli, the dorsal stripe is definite but narrow; dorsal fur length 6.3-7.5 mm in P. masu, whereas P. dorsalis, P. ismaeli, and P. nigellus have the dorsal fur length >8 mm; dense and long hair on the dorsum of the feet in P. ismaeli, P. masu, and P. nigellus, whereas P. dorsalis has the hair on the dorsum of the feet intermediate in density and length; and fringe of hair on the edge of the uropatagium densely haired in P. ismaeli, P. masu, and P. nigellus, whereas P. dorsalis has the uropatagium fringe usually hairy, occasionally sparsely. Dentally, Platyrrhinus masu can be distin- guished from these species by the following char- acters: upper lateral incisors bilobed in P. masu, whereas P. dorsalis and P. nigellus have the upper lateral incisors monolobed; P. dorsalis, P. masu, and P. nigellus present the sulcus on the posterior face of the paracone of Ml joined to the cingulum of the lingual face of the metacone on Ml, where- as P. ismaeli present these two structures not joined; Ml protocone moderately developed in P. masu and P. nigellus, whereas P. dorsalis and P. ismaeli have the Ml protocone small and blunt; 36 FIELDIANA: ZOOLOGY Table 7. Measurements of the type species of Platyrrhinus matapalensis. Holotype Paratype Paratype Paratype Paratype FNH 81079 FMNH 81080 FMNH 81081 MUSM 10725 MUSM 10726 Sex Male Male Male Male Female (pregnant) Weight — — — 16 20 Total length 58 58 56 65 65 Hind foot length 13 13 13 10 12 Ear length 17 17 17 16 19 Forearm length 39 37 39 37 38 Tibia length 17.34 15.50 17.09 14.69 14.43 Greatest length of skull 22.04 22.00 21.46 22.24 21.07 Condyloincisive length 20.77 20.92 20.22 21.34 20.12 Condylocanine length 20.30 20.40 19.82 20.82 19.65 Postorbital breadth 5.43 5.54 5.40 5.67 5.47 Zygomatic breadth 12.99 13.06 — — 12.57 Braincase breadth 9.41 9.49 9.47 9.75 9.35 Mastoid breadth 10.99 11.31 10.54 11.11 10.61 Maxillary toothrow length 8.31 8.54 8.22 8.56 8.13 Breadth across molar 9.14 9.46 9.06 9.39 9.01 stylar cusp present on the lingual face of the M2 paracone in P. ismaeli and P. masu, whereas in P. nigellus, that stylar cusp is lacking; lingual cin- gulum of the M2 metacone restricted to the meta- cone in P. ismaeli and P. masu, whereas in P. dorsalis, the lingual cingulum of the M2 meta- cone continues to the paracone; and lack of the ml metaconid in P. dorsalis and P. masu, where- as in P. ismaeli, the ml metaconid is poorly de- veloped. Platyrrhinus matapalensis new species. Figure 27 (top) Vampyrops helleri Sanborn, 1955: 412 (part) Vampyrops helleri Tuttle, 1970: 73 (part) Vampyrops helleri Koopman, 1978: 12 (part) Platyrrhinus helleri Rodriguez, 1998: 73 Platyrrhinus helleri Albuja, 1999: 139-140 (part) Platyrrhinus helleri "Western" (this paper) Type Material— The holotype, FMNH 81079, is an adult male collected by Celestino Kalinows- ki (original field number 1524) on 3 July 1954 at Matapalo, 54 m in elevation, Province of Zaru- milla, Department of Tumbes, Peru, approximate- ly 3°40'59'S, 80°12'W. The skin and skull were in good condition. Paratypes include four adult specimens: two males (FMNH 81080, 81081) from the type lo- cality; and one female (MUSM 10726) and one male (MUSM 10725) from Quebrada Naranjal- Campo Verde, Z.R. Tumbes, 620-680 m in ele- vation, Province of Zarumilla, Department of Tumbes, Peru. All these specimens are adult and are listed in Appendix 1 with other specimens from the known distributional range. Measure- ments of each specimen of the type series of Pla- tyrrhinus matapalensis are provided in Table 7. Distribution — Platyrrhinus matapalensis is currently known from the western slope of the Andes in Ecuador and Peru. The northern limit of the distribution is the province of Esmeraldas (Ec- uador) and the southern limit is the Department of Tumbes (Peru). The elevational range extends from 54 to 680 m (Fig. 26). Etymology — The specific name matapalensis refers to the type locality Matapalo (Tumbes, Peru), where the taxon was first collected. Diagnosis — Size small (FA = 37-39 mm; CIL = 20.1-21.3 mm; W = 16-20 g); two interramal vibrissae; dorsal stripe definite but narrow, ven- tral fur hicolored, occasionally unicolored; dense and long hair on the dorsum of the feet. Description — A small-sized species of Platyr- rhinus (FA = 37-39 mm; GLS = 21.1-22.2 mm). Dorsal fur light brown, ventral fur brownish, dor- sal hair tricolored, ventral hair bicolored in some individuals and in others unicolored; dorsomedial and ventrolateral facial stripes obviously subequal and brighter, dorsal hairs 6.3-7.5 mm long on the back; narrow dorsal stripe with the same colora- tion as the facial ones; folds in the pinna poorly marked but distinguishable; seven vibrissae sur- rounding the margins of the noseleaf in a single array; four submental vibrissae present; two inter- ramal vibrissae present; noseleaf longer than wide, inferior border of the horseshoe completely VELAZCO: PLATYRRHINUS PHYLOGENY 37 Fig. 26. Distributional map of Platyrrhinus matapalensis. free of the upper lip; hairs on the dorsum of the feet sparse and short; "U" or "V" shaped pos- terior edge of the uropatagium with a fringe densely haired, uropatagium extends along the midline <4.5 mm long; insertion of the posterior edge of the plagiopatagium onto the first metatar- sal; fifth metacarpal shorter than third metacarpal: postorbital and paraoccipital processes poorly de- veloped; fossa on the squamosal end of the zygo- matic arch almost imperceptible; sulcus present on the posterior face of the upper canines: two stylar cusps present on the posterior face of P4; both la- bial and lingual cingulum present at the base of the metacone of Ml: sulcus on the posterior face of the paracone joined to the cingulum of the lingual face of the Ml metacone; Ml parastyle present: Ml metastyle occasionally present; Ml protocone moderately developed: stylar cusp absent on the lingual face of the M2 paracone: labial cingulum of the M2 paracone present; M2 metastyle present; stylar cusp absent on the lingual face of the M2 paracone: both labial and lingual cingula present on p4: stylid cusps absent on the anterior or anter- olingual face of the main cone of p4; stylid cusp present on the posterior face of the main cone of p4; ml paraconid absent: ml metaconid absent; m2 hypoconid absent: stylid cusp between the meta- conid and protoconid of m2 present; both labial and lingual cingula present on m2, but occasionally only the labial cingulum is present. Comparisons — Platyrrhinus matapalensis is found sympatrically with P. chocoensis and P. dorsalis. Morphometrically. P. matapalensis is easily distinguished from both as follows: P. cho- coensis (FA = 48-49 mm; CIL = 24.6-26.2 mm; W = 29-32 g). P. dorsalis (FA = 46-50 mm; CIL = 24.3-25.7 mm), and P. matapalensis (FA = 37-39 mm; CIL = 20.1-21.3 mm; W = 16- 20 g). Platyrrhinus matapalensis has been con- fused with P. helleri (Sanborn. 1955; Tuttle, 1970; 38 FIELDIANA: ZOOLOGY Koopman, 1978); however, P. matapalensis could also be mistaken for P. brachycephalus because there is overlap in the measurements (Table 3, 7). Therefore, the comparisons focus on P. matapa- lensis, P. brachycephalus, and P. helleri because these are the smaller species in the genus and are subject to potential confusion. Externally, Platyrrhinus matapalensis can be dis- tinguished from the other two species by the follow- ing: one vibrissa present on the upper lip of P. ma- tapalensis, ventral to the vibrissae that surround the margins of the noseleaf in, whereas P. brachyce- phalus has two vibrissae on the upper lip; four sub- mental vibrissae on each side of the chin in P. hel- leri and P. matapalensis, whereas P. brachycephal- us has five submental vibrissae on each side of the chin; two interramal vibrissae present in P. mata- palensis, whereas P. brachycephalus and P. helleri have one interramal vibrissa; ventral fur unicolored or bicolored in P. matapalensis, whereas P. brachy- cephalus has the ventral fur tricolored and P. helleri bicolored; sparse and short hair on the dorsum of the feet in P. matapalensis, whereas in P. helleri, the hair on the dorsum of the feet is intermediate in density and length; and fringe of hair on the edge of the uropatagium densely haired in P. helleri and P. matapalensis, whereas P. brachycephalus has the fringe of hair on the edge of the uropatagium usually hairy, occasionally sparsely. Cranially, Platyrrhinus matapalensis can be distinguished from the other two species by the following characters: a poorly developed postor- bital process in P. brachycephalus and P. mata- palensis, whereas in P. helleri, the postorbital pro- cess is moderately developed; and paraoccipital process almost imperceptible in P. brachycephal- us and P. matapalensis, whereas in P. helleri, the paraoccipital process is moderately developed. Dentally, Platyrrhinus matapalensis can be dis- tinguished from the other two species by the fol- lowing: M 1 protocone moderately developed in P. matapalensis, whereas in P. brachycephalus, the Ml protocone is well developed, and in P. helleri, the Ml protocone is small and blunt, and lack of stylid cusps on the anterior face of the main cone of p4 in P. matapalensis, whereas P. brachyce- phalus has two stylid cusps on the anterior face of the main cone of p4, and P. helleri has only one stylid cusp (Fig. 27). Platyrrhinus nigellus (Gardner and Carter, 1972) Vampyrops nigellus Gardner and Carter, 1972a. Type locality: Huanhuachayo, Department of Ayacucho, Peru. Holotype — Vampyrops nigellus LSUMZ 16415, adult male, preserved as skin and skull, was collected by A. L. Gardner (original field number ALG 11684) on 6 May 1971 at Huan- huachayo, Department of Ayacucho, Peru, eleva- tion 1660 m. Distribution — Along the Andes from western Venezuela south to Colombia, Ecuador, Peru, and Bolivia, from 620 to 2757 m (Fig. 28). Diagnosis — Size medium (FA = 40-47 mm; CIL = 21.9-24.5 mm; W = 19-30 g; Table 3); facial stripes darker; absence of a basal protuber- ance, where the genal vibrissae are implanted; postorbital process moderately developed; paraoc- cipital process moderately developed; a deep fos- sa on the squamosal end of the zygomatic arch, lateral to the glenoid fossa; ml metaconid poorly developed. Platyrrhinus recifinus (Thomas, 1901) Vampyrops recifinus Thomas, 1901. Type locality: Recife, State of Pernambuco, Brazil. Holotype — Vampyrops recifinus BMNH 81.2.16.4, adult male, preserved in alcohol with the skull removed and cleaned, was collected by W. A. Forbes at Recife, State of Pernambuco, Bra- zil. Distribution — Eastern Brazil, elevation 200 m (Fig. 28). Diagnosis — Size small to medium (FA = 42- 43 mm; CIL = 22.0-22.6 mm; W = 17-19 g; Table 3); interramal vibrissae absent; dorsal fur tetracolored; dense and long hair on the dorsum of the feet; an almost imperceptible fossa on the squamosal end of the zygomatic arch, lateral to the glenoid fossa; a deep fossa on the hypoconal basin of P4; stylid cusp absent between the meta- conid and the protoconid of m.2. Platyrrhinus vittatus (Peters, 1860) Artibeus vittatus Peters, 1860. Type locality: Puerto Cabello, State of Carabobo, Venezuela. Holotype — Artibeus vittatus ZMB 568, adult male, preserved in alcohol with the skull removed and cleaned, was collected by von Appun at Puerto Cabello, State of Carabobo, Venezuela. Distribution — Costa Rica, Panama, western and northern Colombia, and northern Venezuela, from 640 to 1400 m (Fig. 28). Diagnosis — Size large (FA = 57-61 mm; CIL = 28.1-31.3 mm; W - 60-65 g; Table 3); par- VELAZCO: PLATYRRHINUS PHYLOGENY 39 Fig. 27. Labial views of the left p2 and p4 illustrating taxonomic differences in the number of stylid cusps on the anterior face of the main cone of p4. (Top) Platyrrhinus matapalensis (FMNH 81079) lack stylid cusps (arrow). (Center) Platyrrhinus helleri (FMNH 127127) with one stylid cusp present (arrow). (Bottom) Platyrrhinus brachy- cephalus (FMNH 139584) with two stylid cusps present (arrow). 40 FIELDIANA: ZOOLOGY • Platyrrhinus nigellus * Platyrrhinus recifmus ▲ Platyrrhinus vittatus 1,000 Fig. 28. Distributional map of Platyrrhinus nigellus, P. recifinus, and P. vittatus. allel folds in the pinna well marked; dorsal stripe brilliant-white and wide; fringe of hair on the edge of the uropatagium usually hairy, occasion- ally sparsely; paraoccipital process moderately developed; one stylid cusp present on the antero- lingual face of the main cone of p4. In older in- dividuals (AMNH 15100-15101, 34232; ZMB 568 [Holotype]) with a moderate degree of tooth wear, this stylid cusp appears as a nub on the an- terolingual face of the main cone. Comments — In his systematic review of Pla- tyrrhinus from northwestern South America, Al- berico (1990) identified Platyrrhinus sp. B as a potentially new species from Colombia. Platyr- rhinus sp. B (FA = 56.9-59.4 mm) was collected only at the northern end of the Andes in Colom- bia, in the Department of Antioquia at 500 m; this taxon shows the same range of variation in FA size as P. vittatus (sensu this paper) and was found within the geographic range of P. vittatus. Some populations of P. vittatus (sensu Alberico, 1 990) belong to P. albericoi, suggesting that Pla- tyrrhinus sp. B is conspecific with P. vittatus. From the three potentially new species identi- fied by Alberico (1990), two are conspecific with Platyrrhinus ismaeli and P. vittatus; the third probably constitutes a new species. Platyrrhinus sp. A (FA = 53.0-57.0 mm) is found along the western base of the western Andes of Colombia from the department of Choco, south almost to the Department of Narino, from 230 to 1000 m in elevation. Based on both geographical range and FA size, this taxon does not overlap with any of the other 14 Platyrrhinus taxa. VELAZCO: PLATYRRHINUS PHYLOGENY 41 Commonly, between 8 and 10 nominal species are recognized for Platyrrhinus (Alberico. 1990: Ferrell & Wilson. 1991: Jones & Carter. 1976. 1979: Koopman. 1993. 1994: Nowak. 1999: Owen. 1987: Swanepoel & Genoways. 1979). Based on the results obtained in this work, the genus Platyrrhinus contains at least 14 species. Therefore, it currently ranks as the most speciose genus in the family Phyllostomidae. Acknowledgments For the loan of specimens or for their hospital- ity while visiting their respective institutions, I thank Luis Albuja (EPN). Michael D. Carleton (USNM), Judy Chupasko (MCZ), Luis Coloma (QCAZ). Joseph A. Cook and William L. Gannon (MSB). Judith Eger and Burton Lim (ROM). Mark S. Hafner (LSUMZ). Philip Myers (UMMZ). Victor Pacheco (MUSM). Bruce D. Patterson (FMNH). James L. Patton (MVZ). and Nancy B. Simmons and Robert S. Voss (AMNH). The visits to the AMNH and FMNH were sup- ported by the AMNH Collection Study Grant (AMNH) and the Barbara E. Brown Fund for Mammal Research (FMNH). I am also very grate- ful to Robert Asher (ZMB), who kindly took some pictures of the holotype of Platyrrhinus vittatus. Betty Strack (FMNH) assisted with the SEM pho- tography. Sean Bober (FMNH) shared his exper- tise in GIS. Useful comments on various drafts of this man- uscript were provided by Christine L. Hice. Burton K. Lim. Victor Pacheco. Bruce D. Patter- son, and two anonymous reviewers. I am very grateful to the Departamento de Mas- tozoologfa at the Museo de Historia Natural (UNMSM) and the Division of Mammals of the Field Museum of Natural History for providing space, technical, and scientific support. During the preparation of this manuscript. I was supported by a curatorial assistantship funded by a NSF grant (DEB-9870191) to Bruce D. Patter- son. D. F. Stotz. and J. W. O. Ballard and by the Barbara E. 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Llactahuaman (FMNH 177477, MUSM 14583-14584, 14586, 14588-14589); Cuzco, Paucartambo, San Pedro (MUSM 19149*); Cuz- co, Paucartambo, Suecia, km 138.5 Carretera Shintuya (FMNH 170145*); Cuzco, Paucartambo, Pillahuata (FMNH 172108); Pasco, Oxapampa, Pozuzo, Palmira (MUSM 10973). Platyrrhinus aurarius: Guyana: Potaro-Sipa- runi, Mount Ayanganna, Toe Slope Camp (ROM 1 14679*); Potaro-Siparuni, Mount Ayanganna, First Plateau Camp (ROM 1 14702*); Venezuela: Amazonas, Cerro de la Neblina, Base del Pico Maguire (FMNH 137294-137298*, 137299- 137303, 137304-137307*, 137308-137311); Amazonas, Cerro de la Neblina, Left Bank del Rio Baria (FMNH 137312, 137313*); Amazonas, 3Vi km W Pico Zuloaga (AMNH 261225- 261229); Bolivar, Cerro Guaiquinima (AMNH 235339-235342); Bolivar, La Escalera (AMNH 265572-265584). Platyrrhinus brachycephalus: Bolivia: Cocha- bamba, 4 km SE Villa Tunari (UMMZ 126755); El Beni, Itenez, Buenavista (FMNH 1 15012*); El Beni, Mamore, San Joaquin (FMNH 96092*, 1 15016-1 15018*); El Beni, Vaca Diez, La Esper- anza (FMNH 115003*, 115005*); La Paz, Santa Ana de Madidi (AMNH 262353); Pando, Santa Rosa (AMNH 262520); Brazil: Amazonas, Ta- batinga (AMNH 170623); Colombia: Arauca, Rio Arauca (FMNH 92302-92303, 92326*, 92327); Meta, La Macarena Parque, Refugio (FMNH 58739*); Meta, Villavicencio (AMNH 207872-207873, 207875); Meta, Villavicencio, Finca El Buque (FMNH 121261*); Putumayo, Es- tacion de Bombeo, Guamues (FMNH 113657); Putumayo, San Antonio, Rio Guamues (FMNH 114018-114020); Ecuador: Napo, Marian (FMNH 124990*); Napo, San Jose (AMNH 64033); Napo, San Jose de Payamino (FMNH 124991-124992*); Napo, Zancudo (FMNH 124985-124986*); Orellana, Canton Aguarico, Pozo Exploratorio PCSA-2 (EPN 985199, 985201); Pastaza, Sarayacu (AMNH 67649); French Guiana: Cayenne (FMNH 21745- 21746); Peru: Amazonas, Condorcanqui, Santa Rosa de Nieva (MUSM 12789); Amazonas, San Juan (Bagua Grande) (MUSM 1010); Cuzco, La Convencion, Camisea, Armihuari (MUSM 13770, 13774-13775, 13778-13779, 13786, 13808); Cuzco, La Convencion, Camisea, Cashiriari (MUSM 13788-13790, 13821, 13833); Cuzco, La Convencion, Kiteni (MUSM 1012; UMMZ 160637); Cuzco, La Convencion, Camisea, Kon- kariari (MUSM 14788); Cuzco, La Convencion, Camisea, Pagoreni (MUSM 13791-13792); Cuz- co, La Convencion, Camisea, San Martin (MUSM 13793-13794, 13844); Cuzco, La Convencion, Camisea, Segakiato (MUSM 14793); Cuzco, Pau- cartambo, Consuelo, km 165, 17 km by road W Pilcopata (FMNH 123920*, 123923*); Cuzco, Quispicanchi, Collpa de San Lorenzo (FMNH 93580*, 93582-93585*); Cuzco, Quispicanchi, Huajyumbe (FMNH 84381*); Huanuco, Leoncio Prado, Tingo Maria, Rio Azul (FMNH 98008*); Loreto, Alto Amazonas, Rio Morona, Quebrada Pushaga (FMNH 89091*); Loreto, Alto Amazon- as, Yurimaguas (FMNH 19651*); Loreto, Quebra- da Sucusari, Camp. Explornapo (MUSM 6750); Loreto, Requena, Jenaro Herrera (MUSM 5927); Loreto, Loreto, Rfo Samiria, Base Atun (FMNH 122874-122876*, 122878*); Loreto, Loreto, Rfo Tigre, 1 km above Rfo Tigrillo (FMNH 122880*); Loreto, Mariscal Ramon Castilla, Rio Amazonas, mouth of Rfo Peruate (FMNH 89096*); Loreto, Mariscal Ramon Castilla, Rfo Yavari Mirim, Que- brada Esperanza (FMNH 89095*); Loreto, May- nas, Rfo Nanay, Santa Luisa (FMNH 87074*); Madre de Dios, Albergue Rfo Madre de Dios, 12 km E Puerto Maldonado (MUSM 1013); Madre de Dios, Manu, Alto Rfo Madre de Dios, Haci- VELAZCO: PLATYRRHINUS PHYLOGENY 45 enda Amazonia (FMNH 139639*, FMNH 125930*, 125938*; MUSM 8850): Madre de Dios, Hacienda Erika, Rio Alto Madre de Dios. opposite Salvacion (UMMZ 160636, 160638); Madre de Dios, Reserva Cuzco Amazonico, 15 km NE Puerto Maldonado (MUSM 6268-6270, 7150); Madre de Dios. Manu, Pakitza (MUSM 6769, 12558, 12560, 12562, 12564); Madre de Dios, Manu, Rio Palotoa (MUSM 9844); Madre de Dios, Manu. Rio Palotoa, left bank. 1 2 km up- stream from mouth (FMNH 139581*. 139584- 139585*, 139587*); Madre de Dios, Rio Tambo- pata (MUSM 191); Madre de Dios, Rio Tambo- pata, Explorer's Inn (MUSM 1002, 1006-1007); Pasco, Oxapampa, Huancabamba, Comunidad Nativa Castillo (MUSM 394-396, 398, 400, 430. 496-499); Puno, Carabaya, Coasa. mouth of Que- brada Ursulinda with the Rio Candamo (MUSM 15882); Ucayali, Coronel Portillo, Pucallpa (FMNH 64316-64317*); Ucayali, Coronel Porti- llo, Yarinacocha (FMNH 98009*); Ucayali, Padre Abad, B. N. von Humboldt (MUSM 8527); Su- riname: Nickerie, Wageningen (UMMZ 175688); Paramaribo, Peu Et Contant (UMMZ 175768); Venezuela: Tachira, San Juan de Colon (FMNH 21117*). Platyrrhinus chocoensis: Colombia: Cauca. Alto Micay, Betania (FMNH 113745*, 113822- 113823, 113825-113927, 113830-1 13832); Cau- ca, Quebrada Guangui (AMNH 235774-235779); Choco, Quebrada Docordo (AMNH 233186- 233187); Valle del Cauca, Concesion Bajo Cali- ma, Cuartel BV83 (FMNH 140696-140697*); Valle del Cauca, Zabaletas (FMNH 85838; UMMZ 169039-169040, 169048); Ecuador: Es- meraldas, Borbon, Rio Cayapas. Community of Zapote (QCAZ 2063); Esmeraldas, Borbon, Rio Cayapas, Community of Viruela (QCAZ 2175); Esmeraldas, San Lorenzo, Estacion La Chiquita (QCAZ 2375. 2378, 3281-2382); Esmeraldas, 3 km S San Miguel (EPN 84361). Platyrrhinus dorsalis: Colombia: Cauca, Alto Micay,' Betania (FMNH 1 13372, 1 13380, 1 13385, 113394-113396, 113821*, 113824*, 113828- 113829*, 113831*, 113833-113835*); Cauca, Charguayaco (FMNH 113538-113539*); Cauca, Popayan (FMNH 90327*); Huila, Las Cuevas Parque. Upper Cabana (FMNH 58740*); La Gua- jira, San Antonio (MCZ B-8300); Magdalena, Palomino (MCZ B-8301); Valle del Cauca, El Sil- encio (UMMZ 169038); Ecuador: Carchi, Pailon, NW Parroquia de Chical (EPN 871732, 871748, 871778, 871843-871848. 871868); Carchi, Par- roquia Tobar Donoso, El Pailon (EPN 871747, 871765); Esmeraldas, Borbon, Rio Cayapas, Rio Chimbocal, Comuna Corriente Grande (QCAZ 2157); Esmeraldas, Eloy Alfaro, Pichiyacu, Rio Cayapas, upstream Borbon (QCAZ 1468); Pichin- cha. Nanegal (QCAZ 1975); Pichincha, Tambillo, 25 km S Quito (EPN 7465, 7467); Panama: Da- rien, Parque Nacional Darien, Rancho Frio (FMNH 128141*). Platyrrhinus helleri: Belize: Cayo district, Ba- nana Bank (FMNH 58264*); Cayo district. Ba- nana Bank Ranch (FMNH 58265-58268*); Cayo district, Macaw Bank (FMNH 106804*); Toledo district, Columbia Forest Preserve, Forestry Camp (FMNH 58270*); Toledo district, Columbia For- est Preserve, Mahogany Plant, N Forestry Camp (FMNH 58269*); Toledo district, Forest Home (FMNH 128073*); Bolivia: Beni, 6 km W Cas- arabe (AMNH 255919, 255921-255922, 255927); Beni, Gral. Jose Ballivian, Serrania Pilon, 27 km N Rio Quiquibey by road to San Borja (AMNH 261058, 261062, 261064-261069, 261071- 261074); Beni. Itenes, Buenavista (FMNH 115001*, 115009-115011*, 115012, 115013*); Beni, Mamore, San Joaqufn (FMNH 96092, 115002, 115006. 115007-115008*, 115016- 1 1 50 1 8, 1 1 50 1 9*); Beni, opposite Costa Marques, Brazil (AMNH 209502); Beni, Puerto Almacen (AMNH 255909, 255912); Beni, Vaca Diez, La Esperanza (FMNH 115003-115005, 115014*); Cochabamba, 3 km SW Villa Tunari (AMNH 244629-244630); Cochabamba, 50 km NW Villa Tunari (UMMZ 126754, 126756); La Paz, Santa Ana de Madidi (AMNH 261628-261631, 261634-261639, 261642-261647, 262351, 262354-262355); La Paz, Sararia, 2 hr (river) NW Puerto Linares (UMMZ 126757-126758); Pando, La Cruz (AMNH 262507); Pando, Las Piedras (AMNH 262514); Pando, Rio Nareuda (AMNH 248880, 249062); Pando. Santa Rosa (AMNH 262518, 262522-262523); Pando, W Bank Rio Beni, opposite Hamburgo (AMNH 262510) ; Santa Cruz, 23 km S Camp Los Fierros, Parque Nacional Noel Kempff Mercado (AMNH 264068-264070); Santa Cruz, 4.5 km N Buen Re- tiro (AMNH 260228); Santa Cruz, Camp Los Fierros (AMNH 263616); Santa Cruz, Estancia Cachuela Esperanza (AMNH 260219-260221, 260223-260224, 260226-260227); Santa Cruz, San Rafael de Amboro (AMNH 261663, 262350); Brazil: Para. Belem, Mocambo/Embrapa (FMNH 126601*. 126602, 126603*. 126604-126605, 126606*); Rondonia, Pedras Negras (AMNH 209517); Colombia: Cauca, Alto Micay, Betania (FMNH 113321, 113330, 113344, 113348, 46 FIELDIANA: ZOOLOGY 113883-113884, 113885*, 113886, 113887- 113889*, 114023-114026); Cauca, Quebrada Guangui (AMNH 235780-235788); Cauca, Rio Saija, Quebrada Huanqui (FMNH 104841, 104843); Choco, Quebrada Docordo (AMNH 233188); Cundinamarca, Mesitas de Colegio (AMNH 207863, 207871); Cundinamarca, Sasai- ma (FMNH 49156*); Magdalena, Cacagualito (FMNH 13202*); Meta, Villavicencio (FMNH 121262*); Valle del Cauca, Rio Zabaletas (UMMZ 168982-168984, 168987, 168993- 168995, 168999); Costa Rica: Alajuela, Guayabo (FMNH 18045-18046*); Guanacaste, Guanacaste 2 km S, 12 km E Bolson (FMNH 123152); Pun- tarenas, 2 km S San Vito, Finca Las Cruces (FMNH 123153); Puntarenas, Dominical (UMMZ 112024, 112031); Ecuador: Napo, Estacion La Selva, 85 km SE Coca (QCAZ 2518-2519); Napo, Orellana, Comuna Indillana, mouth of Rio Indillana and Rio Napo, P. N. Yasuni (QCAZ 969); Pastaza, Rio Cushueme, Cushueme, 150 km SE Puyo (FMNH 104753, 104759); Guatemala: Alta Vera Paz, Los Rapidos (AMNH 214242); Pe- ten, Tikal National Park (FMNH 58584*); Santa Rosa, Avellana (AMNH 243774, 243923); Santa Rosa, 13 km N Avellana, Finca Camalote (AMNH 243775); Santa Rosa, 2 km ESE Ixpaco (AMNH 244337); Santa Rosa, 5 km N Avellana, Chiquihuitan (AMNH 245325); Honduras: Cor- tes, Santo Domingo, aprox. 5.5 km ESE Cuyamel (AMNH 265119-265120); Lempira, Tepusuna (FMNH 47615-47616*); Mexico: Oaxaca, Tutla (FMNH 51856*); Veracruz, Tuxtla Mts (FMNH 127127-127130*); Veracruz, Tuxtla Mts, 0.5 km E cerro Balzapote (FMNH 127115-127120*, 127125-127126*, 127131-127132*, 127134- 127136*); Veracruz, Tuxtla Mts, 0.5 km W cerro Balzapote (FMNH 127121-127124*); Panama: Darien, Cana (LSUMZ 25485-25486*); Peru: Amazonas, Bongara, Rio Utcubamba, between Churuja and Pedro Ruiz (FMNH 129149- 129150*; MUSM 4950-4952); Amazonas, Luya, Rio Utcubamba, 1 1 km by road NW Pedro Ruiz (FMNH 129152-129153*); Cuzco, La Conven- tion, Camisea, Armihuari (MUSM 13799, 13803, 13807); Cuzco, La Convencion, Camisea, Cashi- riari (MUSM 13811, 13824-13825); Cuzco, La Convencion, Camisea, Las Malvinas (MUSM 14794); Cuzco, La Convencion, San Martin (MUSM 13846-13847, 13849); Cuzco, Quispi- canchi, Collpa de San Lorenzo (FMNH 93586- 93587*); Junfn, Alto Yurinaqui (MUSM 1009); Loreto, Mariscal Ramon Castilla, Rib Yavari Mir- im, Quebrada Esperanza (FMNH 89094*); Lore- to, Maynas, Rio Nanay, Santa Luisa (FMNH 87080*); Loreto, Requena, Jenaro Herrera (MUSM 871, 4217, 5526, 5598, 5928); Loreto, Requena, Jenaro Herrera, 1 .4 km N Centro de In- vestigation Jenaro Herrera (MUSM 5500); Lore- to, Rib Yarapa, Albergue Yacumana, SE Iquitos (MUSM 9434); Madre de Dios, Albergue, Rio Madre de Dios, 12 km E Puerto Maldonado (MUSM 1008); Madre de Dios, Manu, Aguas Calientes, Rio Alto Madre de Dios, 1 km below Shintuya (UMMZ 160635); Madre de Dios, Manu, Hacienda Amazonia (FMNH 125951*, 139631*; MUSM 8855, 9863-9864); Madre de Dios, Manu, Pakitza (MUSM 6768, 12555- 12557, 12559, 12563, 12565-12575); Madre de Dios, Manu, Rio Palotoa (MUSM 9884); Madre de Dios, Manu, Rio Palotoa, left bank, 1 2 km up- stream from mouth (FMNH 139648*); Madre de Dios, Rio Tambopata, Explorer's Inn (MUSM 1003); Pasco, Oxapampa, Pozuzo (MUSM 10942, 10946); Pasco, Villa America (MUSM 1005); Ucayali, Padre Abad, Bosque Nacional Alexander von Humboldt (MUSM 8633-8636, 8528-8531); Trinidad and Tobago: Saint George, Arima Val- ley (AMNH 149624); Saint George, Chaguara- mas, U.S. Naval Station (AMNH 183164); Saint George, Maracas Valley, Water Fall road (AMNH 176283); Saint George, Port of Spain, Saint Clair, 2 Scott (AMNH 176284); Saint George, 4 mi by road N Arima (AMNH 246222-246224); Saint Patrick, Point Fortin (AMNH 183858-183859); Venezuela: Aragua, Rancho Grande (AMNH 144386); Guarico, 8 km N 13 km W San Jose de Guaribe (AMNH 247644); Tachira, San Juan de Colon (FMNH 20539-20540*); Zulia, 9 km N Rio Catatumbo (AMNH 244039); Zulia, Puerto Delicias (AMNH 244038). Platyrrhinus infuscus: Bolivia: Beni, Gral. Jose Ballivian, Serrania Pilon, 27 km by road N Rio Quiribay (AMNH 261076); Cochabamba, 4 km SE Villa Tunari (UMMZ 126761); La Paz, 6.6 km downstream Canavari, Valle del Rio Coroico (AMNH 246621); La Paz, Serrania Bellavista, 35 km N Canavari (AMNH 246622-246623); Pando, Rio Nareuda (AMNH 248882-248883); Santa Cruz, 3 km N and 13.5 km W San Rafael de Am- boro, Rio Saguayo (AMNH 261623); Santa Cruz, San Rafael de Amboro (AMNH 261664); Colom- bia: Meta, La Macarena Parque, Camp. Izawa (FMNH 58741*, 58745*); Meta, La Macarena Parque, Cano, Refugio Cabana (FMNH 58742- 58743*, 58744); Meta, La Macarena Parque, Cano Cabana, Cabana Duda (FMNH 58746*); Meta, La Macarena Parque, Refugio (FMNH VELAZCO: PLATYRRHINUS PHYLOGENY 47 58747*); Meta, Villavicencio, Villavicencio (FMNH 51733*); Meta, Villavicencio, Small Cano (FMNH 58748); Putumayo, Estacion de Bombeo, Guamues (FMNH 113699, 113752, 113898); Putumayo, San Antonio. Rio Guamues (FMNH 113413, 113415, 113417, 113899- 113900, 114128-114129); Putumayo, Rio Me- caya (FMNH 71525. 72123*); Ecuador: Napo. Loreto, P. N. Napo-Galeras, W side, line 28 (QCAZ 1533-1534); Napo, Loreto, P. N. Napo- Galeras, W side, line 30 (QCAZ 1406-1407) Napo, Rio Cotapino, Oriente (FMNH 47587*) Napo, Rio Suno (AMNH 67929); Napo, San Jose (AMNH 67907-67912, 67918-67919, 67922, 67925-67926); Napo, San Jose de Payamino (FMNH 124989*); Pastaza, Montalvo, (FMNH 41429*); Pastaza, Rio Copotaza (FMNH 53503*) Pastaza, Rio Pindo Yaco (FMNH 43137-43139*) Sucumbios. Gonzalo Pizarro, Los Cedros Bosque, 10 km S Lumbaqui (QCAZ 521-523) Tungurahua, Banos, Oriente (FMNH 47588*) Tungurahua, Palmera (AMNH 67658-67664) Peru: Amazonas, 43 km NE Chiriaco (MUSM 143); Amazonas, Luya, Rio Utcubamba, 15 km by road NW Pedro Ruiz (FMNH 129158- 129159*, 129161*; MUSM 4953-4954); Cuzco, La Convencion, Camisea, Armihuari (MUSM 13850); Cuzco, La Convencion, Camisea, Pago- reni (MUSM 13858-13859); Cuzco, La Conven- cion, Camisea, San Martin (MUSM 13862); Cuz- co, La Convencion Camisea, Segakiato (MUSM 14798); Cuzco, Paucartambo, Consuelo, km 165, 17 km by road W Pilcopata (FMNH 123931- 123932*; MUSM 9923-9924); Cuzco, Paucar- tambo, Tono. 5 km S Rio Tono and 18 km W Patria (MUSM 9950); Cuzco, Quispicanchi, Coil- pa de San Lorenzo (FMNH 93545*); Cuzco. Quispicanchi, Hacienda Cadena (FMNH 68447*. 68449*. 78691-78692*. 78694*); Cuzco, Quis- picanchi, Huajyumbe (FMNH 68450-68454*, 75153-75154*. 78693*, 84406*); Cuzco, Quis- picanchi, San Juan Grande (FMNH 75155- 75156*); Junin, Alto Yurinaqui (MUSM 994); Ju- nin, Chanchamayo, Tarma, Vitoc Valley (FMNH 51525*); Loreto, Mouth of Rio Curaray (AMNH 71696-71701, 71708); Loreto, Loreto, Rio Tigre, 1 km above Rio Tigrillo (FMNH 122902*); Ma- dre de Dios, Hacienda Erika, Rio Alto Madre de Dios opposite Salvacion (UMMZ 160628- 160629); Madre de Dios, Manu, Rio Alto Madre de Dios, Hacienda Amazonia (FMNH 125990*. 126010*, 126019*, 139663-139665*, 139674*, 139676*, 139685*, 139695-139696*, 139700- 139702*, 139710*; MUSM 994. 9847, 9897, 9902-9904. 9910, 9928); Madre de Dios, Manu, Aguas Calientes, Rio Alto Madre de Dios, 1 km below Shintuya (UMMZ 160630-160632); Madre de Dios, Manu, Cerro de Pantiacolla (FMNH 122134*, 139737*, 139739-139742*, 139745*, 139748-139749*; MUSM 9918, 9920-9921); Madre de Dios, Manu, Itahuania (FMNH 84405*); Madre de Dios, Manu, Pakitza (MUSM 12576); Madre de Dios, Reserva Cuzco Amazon- ico, 15 km NE Puerto Maldonado (MUSM 6272- 6273); Pasco, Oxapampa, Pozuzo (MUSM 10960, 10969); Pasco, Oxapampa, Pozuzo. Palmira (MUSM 10941. 10947); Puno. Carabaya, Coasa, mouth of Quebrada Ursulinda with Rio Candamo (MUSM 15824); Ucayali, Affluent of Mamanchi- ta (MUSM 370). Platyrrhinus ismaeli: Colombia: Huila. Las Cuevas Parque, Cueva India (FMNH 58733- 58735*, 58737-58738*); Huila, Cuevas de los Guacharos National Park, 225 m W Upper Ca- bana (IND 1993*); Huila, Cuevas de los Guacha- ros National Park, Entrance to Indian Cave (IND 2244*); Huila, Cueva de Los Guacharos National Park, on top Guacharos Cave (IND 2308*); Huila, Cuevas de los Guacharos National Park, Upper Bridge on Rfo Suaza (FMNH 58732*, 58736*); Ecuador: Azuay. Hacienda Sector de Challtapac. close to Rio Giron (QCAZ 2237); Loja, Loja. San Pedro de Vilcabamba. 3 km N Vilcabamba (QCAZ 1171-1174. 1177, 1180); Morona Santi- ago, San Jose Alto, Canton Paute, Rfo San Vi- cente (EPN 912940); Napo, Loreto, P. N. Napo- Galeras, W side, line 28 (QCAZ 1291); Napo, San Rafael Cascada (FMNH 124988*): Peru: Ama- zonas, 19 km E Balsas (FMNH 129133-129134*, 129136-129137*; MUSM 4944-4945, 4946*); Cajamarca. 12 km SSW by road to San Miguel (MUSM 4949); Cajamarca, Celendin, Hacienda Limon, W Balsas (FMNH 129139*, 129143*, 129145-129146*; MUSM 4947-4948); Cajamar- ca, San Ignacio (MUSM 12884-12889); Lamba- yeque, Ferrenafe, Bosque Chinama (MUSM 925- 926); Piura. Ayabaca, ca. 44 km ESE by road from Ayabaca, ladera Cerro Mayordomo (MUSM 996-998); San Martin, Huallaga (MUSM 16167- 16169); San Martin, La Playa, 28 km NE Pataz (MUSM 7283); San Martin, Las Palmas, 32 km NE Pataz (MUSM 7285-7293); San Martin, Las Papayas (MUSM 7294). Platyrrhinus lineatus: Bolivia: Beni, Rio Ma- more (AMNH 210805-210807. 210809); Beni, Rio Tijamuchi (AMNH 262352); Santa Cruz, 4 km N and 1 km W Santiago de Chiquitos (AMNH 260231); Santa Cruz, 12 km S and 8 km E Santa 48 FIELDIANA: ZOOLOGY Cruz (AMNH 255931-255932); Santa Cruz, Cer- cado, Cercado (FMNH 50990*); Santa Cruz, Chi- quitos, Santiago (FMNH 105844-105862*, 105899-105908, 105909*, 105910-105913); Santa Cruz, Chiquitos, 3 km S and 3 km W San- tiago de Chiquitos (AMNH 261042-261052, 261054-261056); Santa Cruz, Robore (AMNH 260232-260233); Santa Cruz, Santa Rosa de la Roca (AMNH 263618); Brazil: Bahia, Municipio de Valenca, Guaibim (MVZ 185601); Ceara, Serra de Ibiapaba (FMNH 19516); Espifito Santo, Mun- icipio de Santa Teresa, Estacao Biologica Santa Lucia (MVZ 185602); Mato Grosso do Sul, Bel- vedere de Urucum de Corumba (AMNH 37019- 37021); Mato Grosso do Sul, Maracaju (AMNH 134923-134926); Mato Grosso do Sul, Rio Va- caria, Fazenda Capao Bonito (FMNH 47963- 47966, 47968-47977, 47980, 47983-47994, 48008); Mato Grosso do Sul, Urucum (AMNH 36993-37002, 37004-37005, 37007-37018, 39017-39019, 39026-39029; FMNH 26775, 26777-26781*, 26783-26784*, 30032-30048); Parafba, Joao Pessoa, Mata do Buraquinho (MVZ 185596); Pernambuco, Municipio de Tamandare, Coqueiral just outside S border Ibama headquar- ters at Tamandare (MVZ 185598); Rio de Janeiro, Municipio Duas Barras, Town Duas Barras, Gar- den at house of Mario & Tereza Habib (MVZ 185603); Sao Paulo, Sao Paulo, (AMNH 207091); Sao Paulo, Valparaiso (FMNH 41645*); Para- guay: Alto Paraguay, Fuerte Olimpo (FMNH 145258*); Alto Paraguay, W Bank Rio Paraguay, Estancia Puerto Ramos, 5 km SSE Bahia Negra (AMNH 265411-265413); Canendiyu, Igatimi, (AMNH 234285); Central, Asuncion, District Ca- tedral (AMNH 205183-205185); Central, Asun- cion, Recoleta (AMNH 248311-248325, 248503; UMMZ 124321-124332, 125422-125443, 125876-125877, 125879-125901, 133734); Guai- ra, villarrica (AMNH 148662-148666); Itapua, Trinidad (AMNH 36523); La Cordillera, 1.6 km by road S Tobatf (UMMZ 125878); Paraguari, Parque Nacional Ybycui (UMMZ 133732); Para- guari, Saltos de Pirareta (UMMZ 133733); Para- guari, Sapucai (AMNH 23770-23773; FMNH 48791*). Platyrrhinus masu: Bolivia: Cochabamba, 50 km NW Villa Tunari (UMMZ 126759-126760); La Paz, Serrania Bellavista (AMNH 246610- 246614); La Paz, Serrania Bellavista, 47 km by road N Caranavi (UMMZ 158068); Peru: Cuzco, 3 km E Amaybamba (MUSM 989-992); Cuzco, La Convencion, Kimbiri, Camp. Llactahuaman (MUSM 14559-14560, 14562*, 14565, 14566*); Cuzco, La Convencion, Kimbiri, Camp. Wayra- pata (MUSM 14568, 14570, 14572); Cuzco, La Convencion, Rio Urubamba, Kiteni (MVZ 166595*); Cuzco, Paucartambo, 72 km by road NE Paucartambo (UMMZ 160627, 160633); Cuz- co, Paucartambo, Challabamba, P. V Acjanaco (FMNH 170112*; MUSM 8851); Cuzco, Paucar- tambo, Consuelo, km 165, 17 km by road W Pil- copata (FMNH 123917*; MUSM 9854); Cuzco, Paucartambo, La Esperanza (FMNH 174759*, 174760); Cuzco, Paucartambo, Pillahuata (MUSM 11793, 11794, 11795*); Cuzco, Paucar- tambo, Quitacalzon, Carretera Paucartambo-Pil- copata km 163 (MUSM 8852); Cuzco, Paucar- tambo, San Pedro (FMNH 172100* ; MUSM 8853-8854, 11792); Cuzco, Paucartambo, Suecia, km 138.5 road to Shintuya (FMNH 170113*); Cuzco, Quispicanchi, Camante (FMNH 68455*); Cuzco, Quispicanchi, Collpa de San Lorenzo (FMNH 93594*); Cuzco, Quispicanchi, Hacienda Cadena (FMNH 93588*); Madre de Dios, Manu, Cerro de Pantiacolla (FMNH 122136*); Madre de Dios, Manu, Cerro de Pantiacolla, upstream Rib Palotoa (FMNH 139597-139599*, 139601- 129603*, 139606-139607*; MUSM 9849-9853, 9868, 9966); Madre de Dios, Manu, Rio Alto Ma- dre de Dios, Hacienda Amazonia (FMNH 139590-139591*, 139593*; MUSM 9848); Huanuco, Huanuco, Chinchao, Cordillera de Car- pish (MUSM 18265*, 18266*); Pasco, Oxapam- pa, San Alberto (MUSM 10272-10273); Pasco, Pasco, Paucartambo, Auquimarca, Anexo Santa Isabel (MUSM 15881-15884). Platyrrhinus matapalensis: Ecuador: Bolivar, Barraganete, 3 km SW Echeandia (EPN 80.4.1, 80458); Esmeraldas, Borbon, Rio Cayapas, Rio Chimbocal, Comuna Corriente Grande (QCAZ 2155); Esmeraldas, Borbon, Rio Cayapas, Com- munity of Zapote (QCAZ 2066); Esmeraldas, Co- munidad Valle del Sade, 8 km E mouth of the Rio Sade in the Rio Esmeraldas (EPN 85904); Es- meraldas, Eloy Alfaro, Borbon, Rio Santiago, Es- tero Maria, Comuna Selva Alegre (QCAZ 1921); Esmeraldas, Eloy Alfaro, San Miguel, Rio Caya- pas, R. E. Cotacachi-Cayapas (QCAZ 508); Guay- as, Naranjal, La Union (QCAZ 2124); Guayas, El Triunfo, El Piedrero (QCAZ 2624); Los Rios, Quevedo, Rio Palenque (QCAZ 509); Pichincha, Nanegal, Chacapata, Recinto Playa Rica (QCAZ 1900); Peru: Tumbes, Zarumilla, Z.R. Tumbes, Quebrada Los Naranjos, Campo Verde (MUSM 10725-10726*); Tumbes, Zarumilla, Matapalo (FMNH 81079-81081*). Platyrrhinus nigellus: Bolivia: La Paz, 20 km VELAZCO: PLATYRRHINUS PHYLOGENY 49 NNE Caranavi (UMMZ 127174); La Paz, Serran- fa Bellavista, 35 km N Canavari (AMNH 246616- 246620); Santa Cruz, 4.5 km N and 1.5 km E Cerro Amboro, Rib Pitasama (AMNH 261665); Colombia: Cesar, San Sebastian (FMNH 69484*); Narifio, El Carmen (FMNH 113891- 113892, 113894-113897); Ecuador: Azuay, Val- le Yunguilla (FMNH 53504*); Loja, Loja, Mas- anamaca, 12 km S Vilcabamba (QCAZ 1210); Morona Santiago, P. N. Sangay Sardinayacu (EPN 964718); Napo, Loreto, P. N. Napo-Galeras, W side, line 28 (QCAZ 1525); Napo, Loreto. P. N. Napo-Galeras, W side, line 30 (QCAZ 1388- 1389); Napo, San Rafael Cascada (FMNH 124987*); Napo, Tena, Atacapi, Cordillera de Los Guacamayos (QCAZ 1264); Pichincha, Nanegal, Gavilan de Orongo (QCAZ 1974); Pichincha, Es- tacion Forestal La Favorita, close to Chiriboga (EPN 799-10); Pichincha, Reserva Rio Guajalito (QCAZ 3159); Peru: Amazonas, Cordillera del Condor (MUSM 10628); Ayacucho, Huanhu- achayo (AMNH 233644-233684); Cajamarca, San Ignacio (MUSM 12890-12892); Cuzco, La Convencion, Kimbiri, Camp. Wayrapata (MUSM 14579*); Cuzco, La Convencion, Kimbiri, Camp. Llactahuaman (MUSM 14561, 14575*, 14576- 14578); Cuzco, Paucartambo, Bosque de las Nu- bes, km 150 by road Paucartambo-Pilcopata (MUSM 8857); Cuzco, Paucartambo, Consuelo, km 165 by road Paucartambo-Shintuya (FMNH 123947*, 174777*, 174779*, 174781*, 174783*; MUSM 9970-9971, 9975); Cuzco, Paucartambo, Quitacalzon, road Paucartambo-Pilcopata km 163 (MUSM 8858, 8860); Cuzco, Paucartambo, San Pedro (MUSM 11796); Cuzco, Quispicanchi, Collpa de San Lorenzo (FMNH 93589-93593*, 93595-93597*, 93607*); Cuzco, Quispicanchi, Hacienda Cadena (FMNH 93598-93606*); Madre de Dios, Manu. Cerro de Pantiacolla, headwaters of Rio Palotoa (FMNH 139577-139578*, 139580*; MUSM 9969); Madre de Dios, Manu, Hacienda Amazonia (FMNH 126031-126032*; MUSM 9955); Huanuco, Huanuco, Chinchao, Cordillera de Carpish (MUSM 18263*, 18264*, 18273*); Pasco, Oxapampa, Pozuzo, Palmira (MUSM 10943-10944, 10971); San Martin, Huallaga, La Morada (MUSM 16170-16172, 16176); San Martin, Las Palmas, 32 km NE Pataz (MUSM 7295-7296). Platyrrhinus recifinus: Brazil: Sao Paulo (USNM 545002); Sao Paulo, Guaratuba (USNM 542612); Sao Paulo, Iguape (USNM 542613); Sao Paulo, MunicTpio Ilhabela, Ilha de Sao Sebastiao, Parque Estadual Ilhabela (MVZ 185604*, 185605, 185606*, 185607-185608, 185609- 18561 1*); Sao Paulo, Municipio Salesopolis, Mu- seu de Zoologia USP, Esta?ao Biologica Boraceia (MVZ 185901*). Platyrrhinus vittatus: Colombia: Magdalena, Valparaiso (AMNH 15100-15101*); Narino, Ri- caurte (AMNH 34232*); Costa Rica: Puntarenas, Monteverde (UMMZ 116681-116682*); Pana- ma: Darien, ca. 6 km NW Cana, E slope Cerro Pirre (LSUMZ 25464-25466*); Darien, Cerro Mali (AMNH 238203-238209); Darien, Cerro Ta- carcuna, Rio Pucuro (AMNH 238210-238218); Venezuela: Carabobo, Puerto Cabello (ZMB 568). Carol I ia subrufa: Mexico: Chiapas, 7.5 mi SW by road Ixtapa (AMNH 249087); Oaxaca, Juchi- tan, San Jeronimo (AMNH 36037-36038); Oa- xaca, Juchitan. 10 mi S Tapanatepec (AMNH 177625-177626); Oaxaca, Tehuantepec, Limon (AMNH 208238-208240. 167028, 171642- 171644, 171648); Oaxaca, Tehuantepec, Cerro San Pedro (AMNH 146150); Oaxaca, 60 mi Te- huantepec (AMNH 189729). Sturnira erythromos: Peru: Amazonas, ca. 20 km by road W Leymebamba (MUSM 4908- 4910); Ayacucho, Yuraccyacu (MUSM 5269) Cuzco, Paucartambo, Morro Leguia, km 135 road Paucartambo-Shintuya (MUSM 8866-8868) Cuzco, Paucartambo, Challabamba, P. V Acjana- co (MUSM 8876-8877); Cuzco, Paucartambo Pillahuata (MUSM 9980); Cuzco, Paucartambo San Pedro (MUSM 11799); Huanuco, Huanuco Cordillera Carpish (MUSM 172); Huanuco. Un- chog pass between Chunchubamba and Hacienda Paty, NNW Acomayo (MUSM 5260); La Liber- tad, Mashua, E of Tayabamba, road to Ongon (MUSM 5285); La Libertad, Utcubamba, road to Ongon (MUSM 5310). Uroderma magnirostrum: Peru: Cuzco, La Convencion, Camisea, Armihuari (MUSM 14019); Cuzco, La Convencion. Camisea, Kon- kariari (MUSM 14841-14842); Cuzco, La Con- vencion, Camisea, Pagoreni (MUSM 14026); Cuzco, La Convencion, Camisea, San Martin (14027-14037); Cuzco, La Convencion, Camisea, Segakiato (14843-14844); Ucayali, Padre Abad, Bosque Nacional Alexander von Humboldt (MUSM 8630-8631). Vampyrodes caraccioli: Costa Rica: Puntar- enas, 2 km SW Rincon de Osa (MSB 26965- 26969); Ecuador: Pichincha, Estacion Forestal La Favorita, close to Chiriboga (EPN 799-12, 799-15); Sucumbios, Shushurindi. Limoncocha (QCAZ 514); Peru: Cuzco, La Convencion, Cam- 50 FIELDIANA: ZOOLOGY isea, Armihuari (MUSM 14070-14072); Cuzco, La Convencion, Camisea, Pagoreni (MUSM 14073); Cuzco, La Convencion, Camisea, San Martin (MUSM 14075-14078); Loreto, Rio Sa- miria, Tacshacocha (MUSM 1017); Madre de Dios, Manu, Cerro de Pantiacolla (MUSM 10102); Madre de Dios, Manu, Pakitza (MUSM 739, 12625-12624); Madre de Dios, Manu, Ha- cienda Amazonia (MUSM 10100-10101, 10106); Pasco, Oxapampa, Pozuzo (MUSM 10997); Pas- co, Oxapampa, Pozuzo, Rio Negro (MUSM 10996); Ucayali, Atalaya, Sepahua (MUSM 12809). Appendix 2: Data Matrix This data matrix includes all the taxa examined (ingroup and outgroup) and all the characters used in the phylogenetic analysis. Polymorphisms are shown between brackets. Carollia subrufa 0-02 10202 01 120 10021 10000 00000 00000 0- 1 10 00000 10200 01001 000{01 }0 Sturnira erythromos 0-10 02000 00232 12001 0000{01 } 00000 00000 0-010 00000 001 10 00000 00020 Uroderma magnirostrum 10012 03022 21110 {01)0022 00011 11110 00101 11010 00101 11110 01102 10201 Vampyrodes caraccioli 10013 03121 31021 01 {01 } 1 1 10100 11110 {01)0121 0-111 10010 11011 01102 11121 Platyrrhinus aurarius 10103 03122 20221 12112 12100 11111 00021 20111 10101 01010 01212 11221 Platyrrhinus brachycephalus 10012 14121 {23}112{01} 1 1 {01 }00 101 {01 }{01 } 11111 {01 } 1 {01 } 1 1 {12}0{01}01 10101 01022 11212 112{01 }1 Platyrrhinus chocoensis 10102 0{23}1{ 12)2 2{12}111 1 1 {01 }0{01 } 1(01} 100 11111 00021 {12}0{01}21 1{01 } 102 0101001212 102{ 12} 1 Platyrrhinus dorsalis 10103 03121 20121 {01)1101 12100 11111 {01)0021 {12)0121 (01 }(01 } 1 12 01020 01212 112(12)1 Platyrrhinus dorsalis "Norte" — Platyrrhinus is- maeli sp. nov. 10103 03121 20122 121{01}1 1110(01} 11111 {01)0(01)21 21121 (01 }11{01}1 01020 01211 11221 Platyrrhinus dorsalis "Centro-Sur" — Platyrrhin- us masu sp. nov. 10103 011(12)2 31122 12(01)01 1(01)101 11111 (01)0021 20111 {01)11(01)1 01020 01212 112(12)1 Platyrrhinus helleri "Eastern" — Platyrrhinus hel- leri 1001(12} {01)312(12} {23)1111 {01 }2{01 } 1 1 101(01)0 11111 01(01)11 10(01)21 10101 01021 11212 (01)1201 Platyrrhinus helleri "Western" — Platyrrhinus matapalensis sp. nov. 10012 03221 211(01)0 (01}2{01}00 101(01 }{01 } 11111 (01 }1 (01 } 11 10(01}11 1010(12} 01020 11212 11(12)01 Platyrrhinus infuscus 10103 {01 }32{ 12}{ 12} 12120 {01 } 1 {01 } 12 111 (01 }(01 } 11111 11(01)21 20111 1111(12} 01020 01210 102(12)1 Platyrrhinus lineatus 10012 03121 31122 12022 1010(01} 11111 0{01}0{12}1 20111 101{01 }{ 12} 01020 01210 11221 Platyrrhinus nigellus 10103 {01)3122 {23)0122 (01}2{01}01 1{12}100 11111 {01 }0{01 }{ 12}1 (12)0111 10{01 }0{ 12} 01020 0(01}21{12) 11221 Platyrrhinus recifinus 10012 03021 31222 12011 111{01}0 11111 11011 10111 10102 01020 01212 10201 Platyrrhinus vittatus "Northern" — Platyrrhinus vittatus 11103 03222 30112 (01 } 1(01 } 1 1 10100 11111 10021 201{01}1 11111 01021 01211 11221 Platyrrhinus vittatus "Southern" — Platyrrhinus albericoi sp. nov. 11003 03(12)21 20112 12(01)12 1010(01} 11111 10121 20111 11011 0102001211 11211 Appendix 3: Optimization of Characters The following apomorphies support the mono- phyletic clades shown in Figure 29. Both ACCT- RAN and DELTRAN optimizations are given for each clade. Diagnosis for each character is given as follow: (character number; consistency index) character state description, state number -» state number. Unambiguous transformations are indi- cated by a double arrow "=>." transformations VELAZCO: PLATYRRHINUS PHYLOGENY Outgroup Platyrrhinus chocoensis Platyrrhinus dorsalis Platyrrhinus aurarius Platyrrhinus nigellus Platyrrhinus masu sp. nov. Platyrrhinus ismaeli sp. nov. Platyrrhinus infuscus Platyrrhinus albericoi sp. nov. Platyrrhinus vittatus Platyrrhinus lineatus Platyrrhinus recifinus Platyrrhinus helleri Platyrrhinus brachycephalus Platyrrhinus matapalensis sp. nov. Fig. 29. Strict consensus tree with nodes numbered as a reference for Appendix 3, which presents apomorphies of the clades. 10 12 that occur in only one optimization are indicated by a single arrow "— >." Platyrrhinus (Node 1)— ACCTRAN: (30; 1 .000) Sulcus present on the anterior face of P4, 0 => 1; (33; 0.333) Ml mesostyle absent, 1 -> 0; (36; 0.667) Stylar cusp present on the lingual cingulum of the Ml metacone, 0 => 2; (45; 0.500) lingual cingulum of the M2 metacone continuous to the paracone, 0 => 2; (46; 0.500) M2 hypoconal basin developed, 1 => 0; (49; 0.500) both labial and lingual cingula present on p4, 1 => 2; (53; 1.000) both labial and lingual cingula present on ml, 1 => 2; (54; 1.000) stylid cusps present on the anterior face of the ml pro- toconid, 0 => 1. DELTRAN: (30; 1.000) Sulcus on the anterior face of P4, 0 => 1; (36; 0.667) Stylar cusp present on the lingual cingulum of the Ml metacone, 0 => 2; (43: 0.333) M2 metas- tyle present, 0 — > 1; (45; 0.500) lingual cingulum of the M2 metacone continuous to the paracone, 0 => 2; (46; 0.500) M2 hypoconal basin devel- oped, 1 => 0; (53; 1.000) both labial and lingual cingula present on ml, 1 => 2; (54; 1.000) stylid cusps on the anterior face of the ml protoconid, 0=> 1. Node 2— ACCTRAN: (3; 0.500) dark facial stripes, 0 => 1 ; (4; 0.500) Absence of a basal pro- tuberance, where the genal vibrissae are implant- ed, 1 => 0; (10; 0.333) folds in the pinna well marked, 1 -> 2; (1 1; 0.429) dorsal stripe definite, but narrow, 3 — > 2; (18; 1.000) posterior border of the hard palate "V"-shaped, 0 -> 1; (19; 0.286) postorbital process absent or poorly developed, 1 => 0; (39; 0.400) Ml protocone small and blunt, 1 -> 2. DELTRAN: (3; 0.500) dark facial stripes, 0 => 1; (4; 0.500) Absence of a basal protuber- ance, where the genal vibrissae are implanted, 1 => 0; (11; 0.429) dorsal stripe definite, but narrow, 52 FIELDIANA: ZOOLOGY 3 — > 2; (19; 0.286) postorbital process absent or poorly developed, 2 0. Node 3— ACCTRAN: (5; 0.667) six vibrissae surrounding the margins of the noseleaf in a sin- gle array, 2 => 3; (12; 0.500) dorsal fur > 8 mm, 1 => 0; (22; 0.500) A deep fossa on the squamosal end of the zygomatic arch, lateral to the glenoid fossa, 0 => 2. DELTRAN: (5; 0.667) six vibrissae surrounding the margins of the noseleaf in a sin- gle array, 2 => 3; (12; 0.500) dorsal fur > 8 mm, 1 => 0; (18; 1.000) posterior border of the hard palate "V" -shaped, 0 -> 1; (22; 0.500) A deep fossa on the squamosal end of the zygomatic arch, lateral to the glenoid fossa, 0 => 2. Node 4— ACCTRAN: (17; 0.286) A densely haired uropatagium fringe, 1 => 2; (39; 0.400) Ml protocone moderately developed, 2 — > 1; (45; 0.500) lingual cingulum of the M2 metacone re- stricted to the metacone, 2 => 1. DELTRAN: (17; 0.286) A densely haired uropatagium fringe, 1 => 2; (45; 0.500) lingual cingulum of the M2 meta- cone restricted to the metacone, 2 => 1. Node 5— ACCTRAN: (15; 0.286) dense and long hair on the dorsum of the feet, 1 => 2; (22; 0.500) A shallow fossa on the squamosal end of the zygomatic arch, lateral to the glenoid fossa, 2 *-» 1; (31; 0.500) A deep fossa on the hypoconal basin of P4, 0 -> 1. DELTRAN: (15; 0.286) dense and long hair on the dorsum of the feet, 1 => 2; (49; 0.500) both labial and lingual cingula of p4 present, 1 — > 2. Node 6— ACCTRAN: (42; 1.000) stylar cusp present on the lingual face of the M2 paracone, 0 => 1; (44; 0.333) stylar cusp present on the lingual face of the M2 metacone, 0 -> 1 . DELTRAN: (22; 0.500) A shallow fossa on the squamosal end of the zygomatic arch, lateral to the glenoid fossa, 2 — > 1; (42; 1.000) stylar cusp present on the lingual face of the M2 paracone, 0 => 1 . Node 7— ACCTRAN: (10; 0.333) folds in the pinna poorly marked but distinguishable, 2 — > 1; (19; 0.286) postorbital process moderately devel- oped, 0 -h> 1; (55; 0.400) poorly developed ml metaconid, 2 => 1. DELTRAN: (55; 0.400) poorly developed ml metaconid, 2 => 1. Node 8— ACCTRAN: (8; 0.400) two interra- mal vibrissae present, 1 => 2; (17; 0.286) edge of the uropatagium, usually hairy, 2 -> 1; (20; 0.333) A well-developed paraoccipital process, 1 -» 2. DELTRAN: (8; 0.400) two interramal vibrissae present, 1 => 2; (19; 0.286) postorbital process moderately developed, 0 -» 1 ; (3 1 ; 0.500) A deep fossa on the hypoconal basin of P4, 0 — > 1; (44; 0.333) stylar cusp present on the lingual face of the M2 metacone, 0 — » 1. Node 9— ACCTRAN: (2; 1.000) dorsomedial facial stripe more marked than the ventrolateral stripe, 0 => 1; (14; 0.333) ventral fur bicolored, 2 => 1; (22; 0.500) An almost imperceptible fossa on the squamosal end of the zygomatic arch, lat- eral to the glenoid fossa, 1 => 0. DELTRAN: (2; 1.000) dorsomedial facial stripe more marked than the ventrolateral stripe, 0 => 1; (14; 0.333) ventral fur bicolored, 2 => 1 ; (22; 0.500) An al- most imperceptible fossa on the squamosal end of the zygomatic arch, lateral to the glenoid fossa, 1 =* 0. Node 10— ACCTRAN: (15; 0.286) dense and long hair on the dorsum of the feet, 1 — > 2; (17; 0.286) A densely haired uropatagium edge, 1 =» 2; (32; 0.500) Ml parastyle present, 0 -» 1; (34; 1 .000) presence of a labial cingulum at the base of the Ml metacone, 2 -> 1. DELTRAN: (10; 0.333) folds in the pinna poorly marked but dis- tinguishable, 2 -> 1; (17; 0.286) A densely haired uropatagium edge, 1 => 2; (49; 0.500) both labial and lingual cingula on p4 present, 1 — > 2. Node 11— ACCTRAN: (45; 0.500) lingual cin- gulum of the M2 metacone restricted to the meta- cone, 2 => 1; (51; 1.000) stylid cusp present on the posterior face of the main cone of p4, 0 => 1 . DELTRAN: (45; 0.500) lingual cingulum of the M2 metacone restricted to the metacone, 2 =* 1; (51; 1.000) stylid cusp present on the posterior face of the main cone of p4, 0 => 1 . Node 12— ACCTRAN: (19; 0.286) postorbital process, absent or poorly developed, 1 => 0; (20; 0.333) paraoccipital process, poorly developed, 1 => 0. DELTRAN: (19; 0.286) postorbital process, absent or poorly developed, 1 => 0; (20; 0.333) paraoccipital process, poorly developed, 1 => 0. VELAZCO: PLATYRRHINUS PHYLOGENY 53 Field Museum of Natural History 1400 South Lake Shore Drive Chicago, Illinois 60605-2496 Telephone: (312) 665-7055