Mammalian Biology 76 (2011) Contents lists available at ScienceDirect. Mammalian Biology. journal homepage:

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1 Mammalian Biology 76 (2011) Contents lists available at ScienceDirect Mammalian Biology journal homepage: Original Investigation A new species of Myotis Kaup, 1829 (Chiroptera, Vespertilionidae) from Ecuador Ricardo Moratelli a,, Don E. Wilson b a Campus Fiocruz da Mata Atlântica, Fundação Oswaldo Cruz, Rio de Janeiro, Brazil b National Museum of Natural History, Smithsonian Institution, Washington, DC, USA article info abstract Article history: Received 1 July 2010 Accepted 12 October 2010 Keywords: Myotis South America Chocó ecoregion Morphology Taxonomy A new species of bat in the genus Myotis (Vespertilionidae, Myotinae) is described from the Chocó ecoregion on the western slope of the Andes in northwestern Ecuador. The genus Myotis comprises a diverse group of small to large-sized vespertilionid bats distributed worldwide. Twelve South American species are recognized currently, 6 of which occur in Ecuador. Morphological relationships among the new species and the other 12 South American species of Myotis were examined using 15 cranial and 5 external characters. Both qualitative and quantitative analyses found the new species to be distinct. As an aid to future identifications, we provide a key to the Ecuadorian species of Myotis Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved. Introduction The genus Myotis Kaup, 1829 comprises a diverse and widely distributed group of small to large-sized vespertilionid bats with 103 nominal species recognized (Simmons, 2005). Based primarily on LaVal s (1973) revisionary work on Neotropical Myotis, Wilson (2008) recently recognized 12 South American species, 6 of them reported to occur in Ecuador: M. albescens (É. Geoffroy, 1806), M. nigricans (Schinz, 1821), M. oxyotus (Peters, 1867), M. simus Thomas, 1901, M. keaysi J.A. Allen, 1914, and M. riparius Handley, Two subspecies of M. nigricans occur in Ecuador: the nominate form, restricted to lowland localities on the west side of the Andes; and M. n. osculatii (Cornalia, 1849), in the Andean highland area, intermontane valleys, and Amazon lowlands (LaVal, 1973; Wilson and LaVal, 1974; Wilson, 2008). LaVal (1973) used the names M. n. punensis J.A. Allen, 1914 and M. n. caucensis J.A. Allen, 1914 for these subspecies, respectively, but subsequently they have been regarded as junior synonyms (Bogan, 1978; Wilson, 2008). The rainforests of the coastal and lower slopes of the western Ecuadorian and Colombian Andes have a long history of isolation from the Amazon forests on the eastern slopes, and the region is home to a highly endemic and poorly known fauna (Albuja Corresponding author at: Campus Fiocruz da Mata Atlântica, Presidência, Fundação Oswaldo Cruz, Estrada Rodrigues Caldas, 3400, Taquara, Jacarepaguá, Pavilhão Agrícola, CEP , Rio de Janeiro, RJ, Brazil. Tel.: ; fax: addresses: rmoratelli@fiocruz.br, rimoratelli@yahoo.com (R. Moratelli). and Gardner, 2005). The recent description of some new species, including 3 sympatric nectarivorous bat species of the genus Lonchophylla Thomas, 1903 (Dávalos, 2004; Albuja and Gardner, 2005; Woodman, 2007) and 3 frugivorous bat species of Platyrrhinus Saussure, 1860 and Sturnira Gray, 1842 (Velazco, 2005; McCarthy et al., 2006; Velazco and Gardner, 2009), is a reflection of that poorly known, endemic fauna of the tropical and subtropical humid forests on the western slope of the Ecuadorian and Colombian Andes. During fieldwork at the Río Palenque Scientific Center (RPSC) in 1979, Don E. Wilson collected a single specimen of Myotis with a unique set of qualitative and quantitative characters separating it from South American congeners. Despite superficial resemblance to M. nigricans, it is easily distinguished by color, smaller overall size, and shape of the skull. The RPSC is a private reserve, with ha of humid lowland rainforest and very humid premontane forest surrounded by agricultural land, with elevations ranging from 130 to 190 m, at the extreme southern limit of the Chocó ecoregion. The reserve is located in central coastal Ecuador, along the Río Baba, in the northern part of the province of Los Ríos (Dodson and Gentry, 1978; Rachowiecki and Palmerlee, 2003; Pearson et al., 2010). We delayed description of this new species in anticipation of additional specimens becoming available. However, the continuing disappearance of forest habitat on the western slope of the Andes over the past 3 decades argues against further delay. Herein, as part of an ongoing systematic and biogeographic review of South American Myotis, we name and describe a new species of Myotis based on that single available specimen, documenting its diagnostic external and cranial characters by comparing it to other South American /$ see front matter 2010 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved. doi: /j.mambio

2 R. Moratelli, D.E. Wilson / Mammalian Biology 76 (2011) species of Myotis. In addition, we provide a key to the Ecuadorian species of Myotis. Material and methods Specimen identification We confirmed the taxonomic status of the new species by comparing it to currently recognized South American species of Myotis, including examination of type specimens, original descriptions, and comparative series from different localities. A list of specimens examined with their localities is in Appendix A. The museum specimens were identified on the basis of a set of qualitative and quantitative characters reported as diagnostic by previous authors (Thomas, 1901, 1902; Miller and Allen, 1928; Handley, 1960; LaVal, 1973; Baud and Menu, 1993; López-González et al., 2001; López-González, 2005; Rocha, 2008; Wilson, 2008). The color nomenclature used to describe pelage follows Ridgway (1912). Measurements Fifteen cranial and 5 external dimensions were measured using a digital caliper accurate to 0.02 mm. The measurements, reported in millimeters (mm), and their abbreviations are defined as follows (lengths were measured from the anteriormost point of the first structure to the posteriormost point of the second structure mentioned below): greatest length of skull (GLS), from the premaxillae, including the incisors, to the occiput; condylo-canine length (CCL), from the occipital condyles to the anterior surface of the upper canines; condylo-incisive length (CIL), from a line connecting the posterior surfaces of the occipital condyles to the upper incisors; basal length (BL), from the foramen magnum to the upper incisors; zygomatic breadth (ZB), greatest breadth across the outer edges of the zygomatic arches; mastoid breadth (MAB), greatest breadth across the mastoid region; braincase breadth (BCB), greatest breadth of the globular part of the braincase; interorbital breadth (IOB), least breadth dorsally between the orbits; postorbital breadth (POB), least breadth across frontals posterior to the postorbital bulges; breadth across canines (BAC), greatest breadth across outer edges of the crowns of upper canines including cingulae; breadth across molars (BAM), greatest breadth across outer edges of the crowns of upper molars; maxillary toothrow length (MTL), from the upper canine crown including cingulum to the crown of M3; molariform toothrow length (M13), from the crown of M1 to the crown of M3; mandibular length (MAL), from the line of internal incisors, to the angular process; mandibular toothrow length (MAN), from the lower canine to m3; ear length, from the ear notch to the tip of the ear; forearm length, from the elbow to the distal end of the forearm including carpals; third metacarpal length, from the distal end of the forearm to the distal end of the third metacarpal; and length of the dorsal (LDH) and ventral hairs (LVH), from the base to the tip of the hair, measured between scapulas. The length of the ear and weight were obtained from the skin labels. The cranial index (CRI = (((IOB + BCB) GLS)/2)) and a modification of the maxillary index (MXI = (((BAC + BAM) MTL)/2)), as proposed by Baud and Menu (1993) and López-González et al. (2001), were applied to summarize the shape of skull. Results Family Vespertilionidae Gray, 1821 Subfamily Myotinae Tate, 1942 Genus Myotis Kaup, 1829 Myotis diminutus, new species Holotype: A sub-adult male (USNM ) skin and skull collected by Don E. Wilson (field number DEW 4724) on 11 February The skin and skull, including mandible, are complete, with the exception of the zygomatic arches, which are both broken. Type locality: Río Palenque Science Center ( S, W), 47 Km S (by road) from Santo Domingo, Los Rios, Ecuador, elevation ca. 150 m. Distribution: Currently known only from the type locality, but to be expected in other areas of similar habitat on the western slope of the Ecuadorian and Colombian Andes (Fig. 1). Etymology: This is the smallest Myotis yet known from South America and the name diminutus is Latin for diminutive. Common name: Diminutive Myotis. Diagnosis: A small species of Myotis (forearm length 33.3 mm; weight 3.5 g); small ears (11.0 mm); long and silky pelage (dorsal fur 6.5 mm; ventral fur 5.2 mm); dorsal hairs cinnamon-brown, bicolored, with bases and tips only slightly contrasting; ventral fur strongly bicolored with mummy-brown bases (2/3 of total length) and cinnamon-buff tips (1/3), contrasting bases and tips; fringe of hairs along the trailing edge of the uropatagium absent; plagiopatagium attached to feet at the level at toes by a broad band of membrane (Fig. 2); skull small (greatest length mm; mastoid breadth 6.01 mm) and narrow (CRI: 59.2, MXI: 17.2); supraoccipital and posteriormost part of parietals rounded in lateral view, projecting beyond the limit of the occipital condyles; rostrum slightly sloping with regard to the braincase in lateral view; sagittal crest absent; occipital crests low; P3 in toothrow, smaller than P2 (50% of the total length of P2) and visible in lateral view (Fig. 3). Dimensions of the holotype are provided in Table 1. Of the characters listed above, five are useful for field identification: general size (forearm length 33.3 mm; ear length 11.0 mm; weight 3.5 g); texture (silky) and color of fur (dorsal hairs cinnamon-brown, with slight contrast between bases and tips; ventral hairs with mummy-brown bases and cinnamon-buff tips); lacks a fringe of hairs along trailing edge of uropatagium; and the plagiopatagium attached at base of toes. Description and comparisons: Myotis diminutus is the smallest Ecuadorian form of the genus, overlapping in external size only with smaller specimens of M. nigricans. Myotis diminutus has a narrower skull than M. nigricans and differs in the color of fur. Myotis diminutus differs qualitatively from other Ecuadorian species as follows: from M. keaysi and M. riparius by the absence of a sagittal crest; from M. albescens by the absence of a fringe of hairs along the trailing edge of the uropatagium; from M. simus by its silky pelage and plagiopatagium attached at toes by a broad band of membrane; and from M. oxyotus by the paler coloration. Compared with its other South American congeners, M. diminutus overlaps in size with M. atacamensis (Lataste, 1891), but differs in having the dorsal surface of the uropatagium almost naked and only slightly bicolored dorsal fur that appears almost uniformly cinnamon-brown. In M. atacamensis the fur extends distally on the dorsal surface of the uropatagium to a level halfway between the knee and ankle, and the dorsal fur is soft and tricolored, with black bases (LaVal 1973). Myotis diminutus differs qualitatively from other South American species as follows: from M. ruber (É. Geoffroy, 1806) by the absence of a sagittal crest; from M. levis (I. Geoffroy, 1824) by the absence of a fringe of hairs along the trailing edge of the uropatagium; from M. aelleni Baud, 1979 by the slightly bicolored fur (as opposed to tricolored); and from M. chiloensis (Waterhouse, 1840) by the almost uniform-appearing dorsal fur with bases and tips contrasting only slightly and a more rounded braincase. As posted above, with the exception of the color of the fur, it is impossible to distinguish M. nigricans and M. oxyotus from M. diminutus qualitatively. However, those species are measurably larger in both external and cranial measurements (Fig. 4), with little or no overlap, when compared with the holotype of M. diminutus (Table 1). Furthermore, the cranial and maxillary indexes distinguish the holotype of M. diminutus (CRI: 59.2, MXI:

3 610 R. Moratelli, D.E. Wilson / Mammalian Biology 76 (2011) Fig. 1. Map of Ecuador showing the Río Palenque Science Center, type locality of Myotis diminutus. See text for coordinates. 17.2) from the Ecuadorian samples of M. nigricans (CRI: [N = 8], MXI: [N = 7]), which include specimens assigned to M. n. punensis by LaVal (1973), and M. oxyotus (CRI: , MXI: [N = 7]) (Table 1), indicating a narrower skull. These values reported for M. diminutus are too very smaller than the values verified for young/subadult specimens of M. nigricans (CRI: , MXI: [N = 4]), as well as for adult specimens of M. nigricans (CRI: , MXI: [N = 271]) and M. Fig. 2. Dorsal and ventral views of the skin of Myotis diminutus (USNM ). Forearm length 33.3 mm. See Table 1 for other measurements.

4 R. Moratelli, D.E. Wilson / Mammalian Biology 76 (2011) Table 1 Measurements of the holotype of M. diminutus and summary descriptive statistics for Ecuadorian samples of M. nigricans and M. oxyotus. Myotis diminutus Myotis nigricans Myotis oxyotus Character Holotype Mean (Range) N Mean (Range) N Ear length Forearm length ( ) ( ) 7 Third metacarpal length ( ) ( ) 7 Dorsal fur ( ) ( ) 7 Ventral fur ( ) ( ) 7 Weight Greatest length of skull ( ) ( ) 7 Condylo-canine length ( ) ( ) 7 Condylo-incisive length ( ) ( ) 7 Basal length ( ) ( ) 7 Zygomatic breadth 8.40 ( ) ( ) 5 Mastoid breadth ( ) ( ) 7 Braincase breadth ( ) ( ) 7 Interorbital breadth ( ) ( ) 7 Postorbital breadth ( ) ( ) 7 Breadth across canines ( ) ( ) 7 Breadth across molars ( ) ( ) 7 Maxillary toothrow length ( ) ( ) 7 Molariform toothrow length ( ) ( ) 7 Mandibular length ( ) ( ) 7 Mandibular toothrow length ( ) ( ) 7 Cranial index ( ) ( ) 7 Maxillary index ( ) ( ) 7 Summary statistics: N = sample size (adults only). Measurements, with the exception of the indexes, are given in millimeters and weight is given in grams. See text for a description of measurement methods. oxyotus (CRI: , MXI: [N = 18]) across South America. Natural history: As with many other newly described species, we know nothing about the natural history of this bat. Unfortu- Fig. 3. Dorsal, ventral and lateral views of the cranium and dorsal and lateral views of the mandible of Myotis diminutus (USNM ). The picture of the lateral view of the mandible was inverted to allow alignment with remaining skull pictures in the plate. See Table 1 for measurements. nately, the prospects for learning more about it are bleak. The moist forests of western Ecuador at one time covered a large part of the Pacific Coastal area, blanketing the coastal plain and extending up the relatively steep slopes of the western side of the Andes. These forests undoubtedly harbored an enormous wealth and diversity of species, but exist today as a disrupted series of tiny fragments under continuing threat and facing an uncertain future. The RPSC exists as an isolated remnant of vegetation dominated by trees with a dense canopy over 30 m high, rich in lianas and epiphytes, as well as mosses, lichens, ferns, and palms, many of which are endemic. Dissected by steep-sided drainages, a single square kilometer of this forest housed 1250 plant species in 136 families, and of these species 43 were endemic (Dodson and Gentry, 1978). Myotis diminutus is at least the fifth new species of mammal described from the area in recent decades (Dodson and Gentry, 1993). Key to the Ecuadorian species of Myotis 1a. Dorsal fur extremely short (ca. 3 4 mm) and woolly Myotis simus 1b. Dorsal fur longer (> 4 mm) and silky 2 2a. Fringe of hairs on the trailing edge of the uropatagium Myotis albescens present 2b. Fringe of hairs on the trailing edge of the uropatagium 3 absent 3a. Sagittal crest present; supraoccipital and posteriormost 4 part of parietals flattened in lateral view 3b. Sagittal crest absent; supraoccipital and posteriormost 5 part of parietals rounded in lateral view 4a. Dorsal fur on uropatagium covering the legs at least Myotis keaysi halfway from knee to foot along tibia; P3 in toothrow, not crowded to the lingual side 4b. Dorsal fur on uropatagium not reaching to the knees; Myotis riparius P3 may be displaced to the lingual side 5a. Length of forearm more than 35 mm; greatest length of Myotis oxyotus skull more than 14.2 mm; length of ears more than 13 mm; cranial index more than 81 5b. Length of forearm less than 35 mm; greatest length of 6 skull less than 14.2; length ears less than 13 mm; cranial index less than 81 6a. Greatest length of skull more than 13 mm; cranial Myotis nigricans index more than 68; maxillary index more than 19 6b. Greatest length of skull less than 13 mm; cranial index Myotis diminutus ca. 59; maxillary index ca. 17

5 612 R. Moratelli, D.E. Wilson / Mammalian Biology 76 (2011) Fig. 4. Dorsal, ventral and lateral views of the skulls of (A) M. diminutus, (B) M. nigricans and (C) M. oxyotus. (A) Specimen from Los Rios, Ecuador (USNM , GLS: mm); (B) specimen from northeastern Brazil (USNM , GLS: mm); and (C) specimen from Amazonas, Venezuela (USNM , GLS: mm). Acknowledgments We are grateful to Cal Dodson and Roy McDiarmid for their help in the field, and for facilitating our work in Ecuador. The following curators and collection staff provided access to specimens under their care: Adriano L. Peracchi (Universidade Federal Rural do Rio de Janeiro, Brazil), João Alves de Oliveira, Stella Franco (Museu Nacional, Brazil), Fernando de Camargo Passos (Universidade Federal do Paraná, Brazil), Teresa Cristina C. Margarido (Museu de História Natural Capão da Imbuia, Brazil), Mario de Vivo, Juliana G. Barros (Museu de Zoologia da Universidade de São Paulo, Brazil), Eliana Morielle-Versute (Universidade Estadual Paulista Julio de Mesquita Filho, Brazil), Nancy Simmons, Eileen Westwig (American Museum of Natural History, USA), Alfred L. Gardner (USGS Patuxent Wildlife Research Center), Linda K. Gordon (National Museum of Natural History, USA) and Patrick Boussès (Muséum national d Histoire naturelle, France). José Luís Passos Cordeiro (Fundação Oswaldo Cruz, Brazil) composed the map. This research was partially supported by the Coordenação de Aperfeiçoamento de Pessoal de Ensino Superior (CAPES, Ministério da Educação, Brazil), the National Museum of Natural History (Short-Term Visitor Program Grant) and the American Museum of Natural History (Collection Study Grant). We are grateful to Alfred L. Gardner for helpfully reviewing a draft of the manuscript. Appendix A. Specimens examined: Abbreviations of institutions are as follows: Museu Nacional, Rio de Janeiro, Brazil (MN); Universidade Federal Rural do Rio de Janeiro, Seropédica, Brazil (ALP); Museu de Zoologia, São Paulo, Brazil (MZUSP); Universidade do Estado de São Paulo, São José do Rio Preto, Brazil (DZSJRP); Museu de História Natural Capão da Imbuia, Curitiba, Brazil (MHNCI); Universidade Federal do Paraná, Curitiba, Brazil (CCMZ-DZUP); American Museum of Natural History, New York, USA (AMNH); National Museum of Natural History, Washington, DC, USA (USNM); and Museum National d Histoire Naturelle, Paris, France (MNHN). The abbreviation JAO corresponds to field numbers of João Alves de Oliveira, and those specimens will be deposited in mammal collection at Museu Nacional. Myotis albescens: Brazil: Paraná: Paiçandu: DZSJRP 10458, 10459, 10460, 10524, 10525, 10550, 10551, 10552, 10553, 10554, 10555, 10556, 10557, 10558, 10559, 10560, 10561, 10562, 10563, 10564, 10565, 10566, 10567, 10568, 10569, 10570, 10571, 10572, 10573, 10574, 10575, Rio de Janeiro: Piraí: ALP 4659, São Paulo: Avanhandava: MZUSP 1292, São Paulo: Fernando Prestes: DZSJRP 13609, 13612, São Paulo: Itatiba: DZSJRP São Paulo: Vale do Ribeira: DZSJRP Paraguay: Paraguarí: Yaguarón: AMNH Myotis chiloensis: Chile: Coastal Town N of Valparaíso: Zapallar: USNM Myotis keaysi keaysi: Bolivia: Santa Cruz: Vallegrande: AMNH , Peru: Cuzco: Cordillera Vilcabamba: AMNH , , , , , , Huánuco: AMNH Puno: Inca Mines: AMNH Venezuela: Aragua: Rancho Grande Biological Station: USNM Bolívar: Gran Sabana: AMNH , Myotis keaysi pilosatibialis: Venezuela: Aragua: Rancho Grande Biological Station: USNM , , Carabobo: Montalbán: USNM , Miranda: Curupao: USNM , , , Myotis levis dinellii: Argentina: Córdoba: USNM , , Tucumán: La Rocha: AMNH Bolivia: Cochabamba: AMNH Chuquisaca: Tomina: AMNH Santa Cruz: Caballero: AMNH Potosí: Unknown locality: AMNH Myotis levis levis: Argentina: Buenos Aires: La Valle: USNM , Córdoba: USNM Tucumán: Los Vásquez: MZUSP Entre Ríos: Puerto Constanza: USNM Brazil: Minas Gerais: Mariana: MZUSP Minas Gerais: Ouro Preto: MZUSP 15344, Paraná: Palmas: CCMZ-DZUP 380. Paraná: Porto Rico: CCMZ-DZUP 216, 217, 218, 369, 371, 372, 373, 376, 377, 381, 382, 384, 385, 386, 387, 388, 389, 391, 392, 393, 394, 395, 396. Rio de Janeiro: Nova Friburgo: MZUSP Rio de Janeiro: Teresópolis: ALP 6481, Rio Grande do Sul: Camapuã: AMNH , , , , , , , , , , , , , , , , , , Rio Grande do Sul: Harmonia: AMNH Rio Grande do Sul: São Leopoldo: DZSJRP Rio Grande do Sul: Taim: DZSJRP 14548, 14549, 14550, 14552, 14553, 14554, Santa Catarina: Passos Maia:

6 R. Moratelli, D.E. Wilson / Mammalian Biology 76 (2011) CCMZ-DZUP 333, 334, 335, 336. São Paulo: Boracéia: MZUSP 15224, 15225, 15273, 15274, 15304, 15305, 15306, 15307, 15308, 15309, 15358, São Paulo: Cacequí: MZUSP São Paulo: Cananéia: MZUSP 27680, São Paulo: Casa Grande: MZUSP 16473, 16474, 16475, 16476, 16477, 16478, 16479, 16480, 16481, 16482, 16483, 16484, 16485, 16486, 16487, 16488, 16489, 16490, 16491, 16492, 16493, 16494, 16495, 16496, 16497, 16498, 16499, 16500, 16501, 16502, 16503, 16504, 16505, 16506, 16507, 16508, 16509, 16510, São Paulo: Itapeva: DZSJRP 11364, Unknown locality: MNHN type 203. Uruguay: Lavalleja: Piraraja: AMNH , , , , , , , , , , , , Tacuarembó: Yaguari: MZUSP 28979, Colón: USNM Unknown locality: MZUSP 10084, USNM , , Myotis nesopolus: Venezuela: Lara: Rio Tucuyo: USNM Falcón: Capatárida: USNM , , , , , Zulia: Cojoro: USNM Myotis nigricans nigricans: Bolivia: Cochabamba: Chapare: AMNH , , , , , , , , , , , , , , , , , Santa Cruz: El Refugio: USNM , Brazil: Amazonas: MZUSP Mato Grosso do Sul: Rio Brilhante: DZSJRP Mato Grosso do Sul: Pantanal: CCMZ-DZUP 170, 172, 174, 176, 177. Minas Gerais: Caxambu: ALP Minas Gerais: Pains: USNM Minas Gerais: Sete Lagoas: USNM Minas Gerais: Viçosa: USNM , , Paraná: Balsa Nova: CCMZ-DZUP 196, 197, 198, 199, 200, 410, 419, 420, 421, 422. Paraná: Matinhos: CCMZ-DZUP 141, 142, 144, 148. Paraná: São José dos Pinhais: CCMZ-DZUP 428. Rio de Janeiro: Tinguá: ALP 2505, 2870, 6262, 6619, 6620, 6624, 6625, 6679, Rio de Janeiro: Itaguaí: ALP 2284, Rio de Janeiro: Piraí: ALP 4587, Rio de Janeiro: Rio de Janeiro: ALP 3047, 3048, 3049, Rio de Janeiro: Seropédica: ALP 585, 588, 589, 625, 626, 627, 628, 629, 630, 631, 636, 639, 640, 655, 658, 904, 1396, 1826, 2862, 3427, 3428, 3498, 3499, 3581, 3697, 3703, 3705, 3715, 3716, 3718, 3723, 3740, 3741, 3742, 3744, 4815, 4816, 4817, 4819, 4825, 4827, 4839, 4844, 4906, 4930, 4931, 4938, 4939, 4940, 4943, 4944, 4947, 4949, 4950, 5012, 5017, 5020, 5032, 5033, 5034, 5036, 5037, 5040, 5043, 5044, 5045, 5068, 5074, 5075, 5089, 5090, 5092, 5094, 5095, 5097, 5098, 5102, 5103, 5105, 5130, 5131, 5132, 5134, 5171, 5172, 5173, 5174, 5175, 5176, 5179, 5180, 5181, 5185, 5186, 5187, 5188, 5235, 5327, 5331, 5332, 5338, 5340, 5341, 5342, 5344, 5346, 5347, 5348, 5349, 5350, 5499, 5501, 5504, 5510, 5511, 5520, 5521, 5523, 5525, 5543, 5592, 5593, Rio de Janeiro: Teresópolis: ALP 6479, Rio Grande do Sul: Frederico Westphalen: CCMZ-DZUP 338. Santa Catarina: Passos Maia: CCMZ-DZUP 400. São Paulo: Botucatu: ALP 2270, São Paulo: Fernando Prestes: DZSJRP 13616, São Paulo: São José do Rio Preto: DZSJRP São Paulo: São Sebastião: USNM , , , , , , , , , , , , , Colombia: Magdalena: Bonda: AMNH Nariño: Ricuarte: USNM La Nariño: Guayacana: USNM , Ecuador: Esmeraldas: AMNH Pichincha: Santo Domingo: USNM Zamora-Chinchipe: Cumbaratza: USNM , Zamora-Chinchipe: Los Encuentros: USNM San Javier: USNM Pambilar: USNM , Paramba: Guyana: Upper Demerara-Bernice: USNM , Panamá: Canal Zone: Barro Colorado Island: USNM , , , , Darién: Boca de Cupe: USNM Darién: Jaque: USNM , , , , Unknown locality: AMNH Paraguay: Paraguarí: USNM , Boquerón: Parque Nacional Teniente Enciso: USNM , Paraguarí: Sapucay: USNM , , Paraguarí: Ibicuy National Park: USNM Peru: Amazonas: Cordillera Del Condor: USNM , Venezuela: Amazonas: Boca Mavaca: USNM Amazonas: Ayacucho: USNM , Apure: Nulita: USNM Bolívar: Maripa: USNM Carabobo: Urama: USNM , , , , , , , , , , , , , , Monagas: San Agustín: USNM , , , , , , , Yaracuy: Urama: USNM Myotis nigricans punensis: Colombia: Chocó: Novita: AMNH Nariño: Ricuarte: USNM Nariño: La Guayacana: USNM , Ecuador: Esmeraldas: Malimpia: USNM , , , Esmeraldas: San Javier: USNM , , Guayas: Puna Island: AMNH Myotis oxyotus: Colombia: Nariño: El Guabo: USNM Ecuador: Chimborazo: Pallatanga: USNM Pastaza: Mera: USNM , Pastaza: Mirador: USNM , , , Peru: Junín: Río Palca: USNM Cuzco: Iquente: USNM Cuzco: Santa Ana: USNM , , , , Venezuela: Amazonas: Cerro Neblina: USNM , , Km SSE Bolívar: El Dorado: USNM Mérida: Tabay: USNM Myotis riparius: Brazil: Mato Grosso: Cachoeira São Simão: MN Mato Grosso do Sul: Paranaíba: DZSJRP Pará: Altamira: USNM , Pará: Belém: USNM , , , , , Pará: Mocambo: ALP 1915, 2002, 2003, 2004, 2554, 2557, 2562, 2568, 2587, 2610, Rio de Janeiro: Tinguá: ALP 4356, 4357, 5421, 6622, 6680, Rio de Janeiro: Piraí: ALP São Paulo: Buri: MZUSP 32968, 32969, São Paulo: Juquitiba: MZUSP 32963, 32964, São Paulo: São Paulo: MZUSP Myotis ruber: Brazil: Minas Gerais: Viçosa: USNM , , Rio de Janeiro: Tinguá: ALP 6621, 6623, Rio de Janeiro: Macaé de Cima: JAO 1751, 1756, Rio de Janeiro: Teresópolis: ALP 6452, 6457, 6458, 6497, 6499, 6506, 6512, MN Rio Grande do Sul: São Lourenço do Sul: MZUSP 1298, São Paulo: Boracéia: MZUSP 28359, 28367, São Paulo: Buri: MZUSP 32971, 32972, 32973, São Paulo: Cananéia: MZUSP São Paulo: São Paulo: MZUSP 15254, 31470, 31471, 31472, 31473, Unknown locality: DZSJRP Myotis simus: Bolivia: Beni: Cercado: AMNH , , , , , , , , , , , , , , , , , , , , , , , Santa Cruz: El Refugio: USNM Brazil: Amazonas: Borba: AMNH 91886, 91887, 91888, 91889, 91890, 91891, 91892, 94224, 94225, 94227, 94230, 94231, 94232, 94233, Amazonas: Itacoatiara: MZUSP Amazonas: Manaus: AMNH 79534, 91472, 91473, 91474, 91475, 91476, 91477, 91478, Amazonas: Parintins: 92983, 93489, 93490, 93491, 93492, 93493, 93494, 93495, 93496, 93497, 93922, 93923, 93924, Amazonas: Rio Juruá: MZUSP 638, Mato Grosso: Taiamã: MZUSP Peru: Loreto: Maynas: AMNH 71483, 71485, 71486, 71487, 71488, 71490, 71491, 71492, 71493, 71494, 74105, 74109, 74110, 74378, 74379, 74380, Loreto: Ucayali: AMNH 76240, 76241, 76242, 76243, 76244, 76245, 76246, 76247, 76248, 76249, 76252, Pasco: San Juan: USNM , References Albuja, V.L., Gardner, A.L., A new species of Lonchophylla Thomas (Chiroptera: Phyllostomidae) from Ecuador. 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