Distribution and natural history of Myotis lavali (Chiroptera, Vespertilionidae)

Transcription

1 Journal of Mammalogy, 94(3): , 2013 Distribution and natural history of Myotis lavali (Chiroptera, Vespertilionidae) RICARDO MORATELLI* AND DON E. WILSON Pavilhão Olympio da Fonseca Filho, Campus Fiocruz da Mata Atlântica, Fundação Oswaldo Cruz, Rio de Janeiro, Brazil (RM) Present address of RM: Division of Mammals, National Museum of Natural History, Smithsonian Institution, Washington, DC , USA Division of Mammals, National Museum of Natural History, Smithsonian Institution, Washington, DC , USA (DEW) * Correspondent: rmoratelli@fiocruz.br LaVal s myotis (Myotis lavali) was recently described from the M. nigricans complex based on specimens from the Caatinga of northeastern Brazil. We present new distributional records for the Alto Chaco in Paraguay and for the Atlantic Forest of Brazil and Paraguay. These new records extend the distribution of the species approximately 2,000 km southwest and 400 km east, and document its co-occurrence with M. nigricans. Both results have taxonomic and ecological implications for M. lavali. Additionally, we provide comments on its natural history and reproduction. Key words: bionomy, geographic distribution, Gloger s rule, Myotinae, range extension, South America Ó 2013 American Society of Mammalogists DOI: /12-MAMM-A LaVal s myotis (Myotis lavali Moratelli et al., 2011a) was described from the M. nigricans (Schinz, 1821) complex based on museum collections from 3 localities in northeastern Brazil. All records are within the limits of the Brazilian Caatinga ecosystem, without confirmed evidence of sympatry between M. nigricans and M. lavali (Moratelli et al. 2011a). As with many other newly described species, the natural history and distribution limits of M. lavali remain practically unknown. As part of a critical review of South American samples of Myotis we found specimens from outside of the Caatinga that fit the description of M. lavali. Herein, we analyze these potential records from Atlantic Forest and Alto Chaco habitats, and the syntopy of M. lavali and M. nigricans. The results have taxonomic and ecological implications for M. lavali. We also comment on the natural history and reproduction of the species based on literature and museum specimens currently identified as M. lavali. MATERIALS AND METHODS Five Paraguayan specimens deposited in the Smithsonian s National Museum of Natural History (USNM) constitute the 1st records of M. lavali from that country, and document its syntopy with M. nigricans. Three of them, prepared as dry skins and skulls, were collected by the Biological Survey Program of the United States Geological Survey in Parque Nacional Teniente Agripino Enciso, 200 miles east of Del Fortin, Boquerón Department. According to the labels, USNM (male) was caught by D. B. Abrell on 22 July 1982; whereas USNM (female) and USNM (female) were caught by M. Ludlow, respectively, on 21 and 22 July The other 2 specimens (females) were collected by W. T. Foster: USNM , prepared as a fluid-preserved specimen, comprises a skin (in spirit) and skull, and was captured in Villarica, Guairá Department, on 14 September 1900; and USNM , represented only by a skull, was captured in Sapucaí, Paraguarí Department, on 20 April Myotis lavali can be distinguished from the remaining species in South America by the following set of traits: dorsal fur strongly bicolored, with medium-brown bases (two-thirds of the total length) and light-brown tips, fringe of hairs along the trailing edge of uropatagium absent, plagiopatagium attached at toes, forehead steeply sloping with regard to the braincase, and occipital region projecting beyond the line of the occipital condyles (Moratelli et al. 2011a). The species also can be distinguished cranially by the upwardly inclined rostrum, a 650

2 June 2013 MORATELLI AND WILSON NATURAL HISTORY OF MYOTIS LAVALI 651 character shared by most specimens of M. lavali, and absent in other South American species (Moratelli et al. 2011a). In the course of our review of South American specimens of Myotis spp. deposited at USNM, the 1st indication of the occurrence of M. lavali in Paraguay was the strong contrast between bases and tips of the dorsal fur of USNM , which corresponds exactly with the pattern observed in specimens of M. lavali from northeastern Brazil. The pelage colors of USNM and USNM do not fit this pattern and resemble those of M. nigricans, with bases and tips of dorsal hairs contrasting just slightly (see Moratelli et al. 2011a). However, identifications of skulls of USNM and USNM do not agree with identifications of skins. The skull of USNM resembles that of M. lavali in shape; whereas the skulls of USNM and USNM are identical and resemble that of M. nigricans (see Moratelli et al. 2011a). In summary, USNM resembles M. lavali in pelage color but is more like M. nigricans in the size and shape of skull; USNM resembles M. nigricans in pelage color and M. lavali in the size and shape of skull; and USNM resembles M. nigricans in both pelage color and size and shape of skull. The skull of the USNM fits the description of M. lavali, but the contrast between bases and tips in the dorsal fur is not so evident; and USNM is a skull only that resembles M. lavali in shape, although it is slightly smaller. An additional female specimen (USNM ) collected by A. L. Gardner at Estação Ecológica do Tapacurá, Pernambuco, on 17 March 1978, with a skull resembling that of M. lavali, but with slightly darker pelage, was included in the analyses to test the occurrence of the species in the Atlantic Forest of northeastern Brazil. To test identifications of specimens reported above, each skull was classified with regard to samples representing M. lavali and most South American species currently recognized from Paraguay and adjacent countries based on the frequencies of shortest Mahalanobis distances obtained in 1,000 bootstrap iterations. Because multivariate procedures require complete data sets, missing values (1.9% of total data set) were estimated from the existing raw data using the expectationmaximization algorithm (Little and Rubin 1987; Strauss et al. 2003). All measurements and estimated values were then logtransformed and the covariance matrices were computed considering all variables. Statistical procedures were performed in Matlab (The MathWorks, Inc., Natick, Massachusetts) using functions written by R. Strauss (Strauss 2012). Samples used in the analysis included 218 adult specimens (Appendix I) representing M. albescens (É. Geoffroy, 1806), M. ruber (É. Geoffroy, 1806), M. nigricans, M. levis levis (I. Geoffroy, 1824), M. oxyotus (Peters, 1866), M. simus Thomas, 1901, M. levis dinellii Thomas, 1902, M. keaysi J. A. Allen, 1914, M. riparius Handley, 1960, M. izecksohni Moratelli et al., 2011a, and M. lavali. Except for M. lavali, restricted to northeastern Brazil, this assemblage comprises most species that occur in Paraguay and adjacent localities (see López- González 2005; Wilson 2008; Stevens et al. 2010). Only M. aelleni Baud, 1979, and M. chiloensis (Waterhouse, 1840) were absent in the analysis. A subset of the dimensions (in mm) used by Moratelli et al. (2011a) was used here, as follows: greatest length of skull (GLS), condyloincisive length (CIL), mastoid breadth (MAB), braincase breadth (BCB), interorbital breadth (IOB), postorbital breadth (POB), breadth across canines (BAC), breadth across molars (BAM), maxillary toothrow length (MTL), molariform toothrow length (M13), and mandibular toothrow length (MAN). For a complete description of measurements see Moratelli et al. (2011a). Notes on natural history were obtained from labels and literature reporting museum specimens currently identified as M. lavali. Most of the information on the biology and reproduction of M. lavali was made available from fieldwork conducted by M. R. Willig from 1976 to 1978 in 2 localities in the Caatinga ecosystem (Exu, Pernambuco and Crato, Ceará) in northeastern Brazil. In the original publications specimens currently identified as M. lavali are referred to as M. nigricans (Mares et al. 1981; Willig 1983, 1985a, 1985b, 1986; Willig et al. 1986) or M. riparius (Willig and Moulton 1989; Willig and Hollander 1995). Specimens referred to M. nigricans by Gregorin et al. (2011) from Estação Ecológica Serra Geral do Tocantins, in the Cerrado ecosystem, were analyzed from photographs and probably correspond to M. lavali. Other specimens from northeastern Brazil currently identified as M. nigricans are pending revision (e.g., Astúa and Guerra 2008 [Pernambuco]; Fábian 2008 [Ceará]; and Feijó and Nunes 2010 [Rio Grande do Norte]). RESULTS Specimen identification. In the probability allocations of skulls, specimens USNM and USNM were classified with M. nigricans (73% and 59%, respectively), whereas USNM was classified with M. lavali (100%), confirming previous identifications (Table 1). The skull confirmed as belonging to M. lavali (USNM ) has a longer and upwardly oriented rostrum when compared with the other 2 specimens classified as M. nigricans (USNM and ). Both of these traits are used to distinguish these species (Moratelli et al. 2011a), and although not all specimens currently identified as M. lavali have the rostrum upwardly inclined, this condition is absent in other South American Myotis spp. The sagittal crest is absent in USNM (lavali), and present in USNM (nigricans) and USNM (nigricans), but as shown by previous studies (e.g., Moratelli and Oliveira 2011; Moratelli et al. 2011a, 2011b), this structure is quite variable in other South American Myotis, and should not be regarded as a diagnostic trait in isolation. Regarding skins, the dorsal fur color of USNM and USNM is medium brown, with bases and tips contrasting just slightly. Both are similar to other Paraguayan specimens of M. nigricans in the color of the dorsal fur (e.g., USNM , , , , and ), except for a few slightly darker individuals (e.g., USNM ). These differences may be attributed to time in storage and variation in storage conditions, or different habitat or roost

3 652 JOURNAL OF MAMMALOGY Vol. 94, No. 3 TABLE 1. Frequency distribution of classification of single specimens of Myotis using minimum Mahalanobis distances to centroids of selected samples on 1,000 bootstrap iterations. Boldface values correspond to higher classification values and dashes ( ) correspond to zero. USNM ¼ National Museum of Natural History; NE ¼ northeastern; SE ¼ southeastern; S ¼ southern; N ¼ northern. USNM albescens lavali nigricans l. dinellii l. levis izecksohni oxyotus keaysi riparius simus ruber Paraguay NE Brazil SE Brazil Argentina and Bolivia SE and S Brazil and Argentina S Brazil Peru Peru N Brazil Bolivia and N Brazil SE Brazil conditions. But the slight contrast between bases and tips in the dorsal fur is consistent throughout the distribution of M. nigricans (see Moratelli et al. 2011a). USNM has bases and tips of dorsal hairs strongly contrasting, with mediumbrown bases and light-brown tips, similar to M. lavali from Ceará, northeastern Brazil, in the pelage banding pattern (e.g., USNM , , , and ). The analyses of skins and skulls lead us to speculate that these parts were interchanged in the specimens USNM and USNM The skulls of the other 2 Paraguayan specimens (USNM and ) were mainly classified with M. lavali (Table 1). Both have upwardly oriented rostra, and occipitals projecting beyond the posterior limit of the occipital condyles, but are cranially smaller than northeastern specimens. The sagittal crest is absent in USNM and present but very low in USNM In the latter specimen the 2nd upper premolar (P3) is displaced to the lingual side and not visible in labial view, whereas in the former this tooth is aligned with others. The skin of USNM has less obvious contrast in comparison with other M. lavali, but may have faded as an effect of more than a century preserved in spirit (Simmons and Voss 2009). The female specimen from Pernambuco, northeastern Brazil (USNM ), had its identity confirmed in the classification analysis (Table 1). This specimen was previously identified as M. lavali based on its cranial size and shape, but the dorsal fur is darker and less contrasting than that of other specimens from northeastern Brazil (e.g., USNM , , , and ). It was collected in the Estação Ecológica do Tapacurá, São Lourenço da Mata, Pernambuco, a seasonal lowland forest in the Atlantic Forest ecosystem. We speculate that the difference in color is related to habitat differences. Based on the specimens identified as M. lavali (USNM [skin? and skull], [skull], [skin], [skull], and [skin and skull]) and M. nigricans (USNM [skull] and [skin and skull]), we confirm the occurrence of M. lavali in the Alto Chaco in Paraguay and in the Atlantic Forest of northeastern Brazil and Paraguay. We also report the syntopy of M. lavali and M. nigricans in the Parque Nacional Teniente Agripino Enciso, Boquerón Department, Paraguay. These records from Paraguay are at least 2,000 km southwest from Barra, Bahia, which previously constituted the southernmost limit for the species (see Moratelli et al. 2011a). Including M. lavali, 7 species of Myotis are reported from Paraguay (López-González et al. 2001; López-González 2005; Stevens et al. 2010), and 8 from Brazil (Moratelli et al. 2011a). The rostrum upwardly oriented and the dorsal fur strongly contrasting can be used to identify most specimens of M. lavali. But in specimens with the rostrum not upwardly oriented and with the dorsal fur less contrasting, other qualitative and quantitative traits are useful. For instance, M. lavali can be distinguished from M. albescens and M. levis by the fringe of hairs along the trailing edge of uropatagium absent; from M. simus, M. riparius, and M. ruber by the woolly and bicolor dorsal fur, and occipital projecting well beyond the posterior limit of the occipital condyles; also distinguished from M. simus by the plagiopatagium attached at the toes and longer dorsal fur; and from M. nigricans and M. izecksohni by the sagittal crest generally present and bicolor dorsal fur; also distinguished from M. nigricans by the longer rostrum, and from M. izecksohni by smaller forearm, thumb, and tragus. Distribution and natural history. Myotis lavali apparently occurs in a diagonal corridor composed of the Brazilian Caatinga and Cerrado and the Paraguayan Alto Chaco, encompassing localities predominantly composed of semiarid (Caatinga and Alto Chaco) and savanna (Cerrado) formations, with peripheral records in adjacent Atlantic Forest localities in northeastern Brazil and Paraguay (Fig. 1). We expect the occurrence of the species in other localities inside the Cerrado, filling the gap between the Brazilian savanna and the Alto Chaco. Elevational records range from 15 m in Russo, Ceará, to 900 m in the Floresta Nacional do Araripe, Pernambuco, but the species appears to be more frequent between approximately 350 and 550 m. Specimens from Caatinga (Figs. 2 and 3) occur in deciduous, drought-adapted formations (localities 1 and 4 in the map) and open-forest savanna formations (locality 2). In most of these localities M. lavali is a common and abundant species, with sexes occurring in similar proportions (Willig 1983, 1985a, 1985b). The species frequently was found roosting in roofs of abandoned buildings, but without forming aggregations of any size (Willig 1983). The record for the Atlantic Forest of northeastern Brazil consists of 1 specimen from Estação

4 June 2013 MORATELLI AND WILSON NATURAL HISTORY OF MYOTIS LAVALI 653 FIG. 1. Map of part of South America showing previous (spheres [1 4]) and new (stars [5 8]) localities for Myotis lavali: 1) Exu, Pernambuco, Brazil; 2) Crato, Ceará, Brazil; 3) Russo, Ceará, Brazil; 4) Barra, Bahia, Brazil; 5) São Lourenço da Mata, Pernambuco, Brazil; 6) Sapucaí, Paraguarí, Paraguay; 7) Villarica, Guairá, Paraguay; and 8) Parque Nacional Teniente Agripino Enciso, Boquerón, Paraguay. Localities 1 4 are in the Caatinga ecosystem, localities 5 7 are in the Atlantic Forest, and locality 8 is in the Alto Chaco. Ecológica do Tapacurá, São Lourenço da Mata, Pernambuco, a seasonal lowland forest (locality 5). In the Cerrado ecosystem the species apparently occurs in the Estação Ecológica Serra Geral do Tocantins, Tocantins State, where specimens were captured in habitats of vereda bordered by humid grassland, FIG. 2. Myotis lavali (male) from the Caatinga of Pernambuco. Photograph courtesy of the collector, Roberto L. M. Novaes. and riparian forest bordered by deforested area (Gregorin et al. 2011). In Paraguay the species occurs in 2 distinct phytogeographic zones, Alto Chaco (west of Paraguay River) and central Paraguay (east of Paraguay River [see Presley et al. 2009]). On the west side of the Paraguay River, the species is reported from Parque Nacional Teniente Agripino Enciso, Boquerón Department. In that region the landscape is characterized by xerophytic thorn-scrub forest (locality 8). In central Paraguay the species occurs in lowland humid forests (approximately m; Villarica, Guairá, and Sapucaí, Paraguarí), in a region that constitutes the westernmost limit of the Atlantic Forest ecosystem (localities 6 and 7). In samples from Caatinga males and females are similar in external and cranial linear measurements (Willig 1983; Willig et al. 1986; Moratelli et al. 2011a). The only significant difference among sexes found by Willig (1983) was for body mass, with females heavier than males in both Pernambuco (means; males ¼ 4.50 g, females ¼ 4.95 g) and Ceará (males ¼ 4.34 g, females ¼ 4.88 g). Regarding the reproductive cycle, Willig (1985a) found evidence of continuous breeding year-round, with no peaks of pregnancy or lactation detected. He found the following numbers (in parentheses) for pregnant, lactating, pregnant and lactating, and inactive females: Pernambuco: January (2, 1,

5 654 JOURNAL OF MAMMALOGY Vol. 94, No. 3 FIG. 3. Caatinga of Pernambuco in the A) rainy and B) dry seasons where specimens of Myotis lavali were captured. Photographs courtesy of Luiz A. M. da Silva. 0, 8), February (0, 3, 1, 6), March (1, 2, 0, 5), April (3, 12, 0, 12), May (2, 1, 0, 7), June (0, 1, 0, 5), July (0, 0, 0, 1), August (1, 1, 0, 2), September (1, 1, 0, 1), October (1, 1, 0, 3), November (1, 1, 0, 0), December (0, 1, 0, 0); Ceará: January (4, 0, 0, 1), February (2, 1, 0, 2), March (1, 0, 0, 7), April (0, 1, 0, 1), June (0, 0, 0, 2), September (1, 0, 0, 1), October (1, 0, 0, 0), December (1, 0, 0, 1). DISCUSSION We found that M. lavali occurs in ecosystems other than Caatinga, and in syntopy with M. nigricans in part of its distribution. These findings have the following taxonomic and ecological implications for M. lavali: 1) reinforce its specific status by demonstrating that M. lavali is not a geographic form of M. nigricans; and 2) give us clues on the plasticity of the species, which appears to occur predominantly in semiarid and savanna ecosystems in South America, but also in adjacent humid forests. Among South American Myotis, apparently only M. nesopolus larensis LaVal, 1973, M. atacamensis (Lataste, 1892), and M. chiloensis occur in semiarid habitats, with the first 2 restricted to those environments (LaVal 1973; Wilson 2008). Records of M. lavali must be examined in a more detailed and accurate geographic scale within these ecosystems to determine habitat preferences. Specimens from the corridor Caatinga Cerrado Alto Chaco, currently identified as M. nigricans, must be critically reviewed to determine their identity and possible sympatry of M. nigricans and M. lavali in that region. In an ecological generalization known as Gloger s rule, animals from humid habitats tend to be darker than those from dry habitats (Mayr 1999; Caro 2005; Kamilar and Bradley 2011). This pattern was 1st revealed for birds (Gloger 1833), and subsequently extended to terrestrial mammals (Dice and Blossom 1937). Although hypotheses to explain it are difficult to test, microorganism resistance, camouflage, and thermoregulation have been proposed (Dice and Blossom 1937; Burtt and Ichida 2004; Caro 2005; Kamilar and Bradley 2011). Pelage color of the only available specimen from the Atlantic Forest prepared as a dry skin (USNM ) is darker than others from Caatinga and Alto Chaco, suggesting that animals from humid habitats are darker than animals from dry habitats. But to support this hypothesis, large samples from the Atlantic Forest must be examined, and records within dry habitats must be evaluated accurately. Despite being a newly described species, a reasonable amount of information is available for M. lavali under the names M. nigricans and M. riparius. We believe that our understanding about the biology of the species may increase further with the review of specimens deposited in additional biological collections, particularly those from midwestern and northeastern Brazil, and the analyses of accompanying field notes. We strongly recommend dry skin preparations for vouchers because of the importance of pelage color and color banding patterns in identifications of South American Myotis. During the preparation of dry skins special care must be taken to adequately stretch the uropatagium (to verify the occurrence of fringe of hairs along the trailing edge) and the plagiopatagium (to verify the site of its attachment), remembering to tie the tags around the leg midway between the ankle and the knee, never on the foot (Simmons and Voss 2009). RESUMO Myotis lavali Moratelli et al., 2011a, foi recentemente descrita a partir do complexo M. nigricans com base em amostras da Caatinga do nordeste do Brasil. Assim como muitas espécies recém-descritas, seus limites de distribuição e sua biologia ainda são pouco conhecidos. Fornecemos novos registros de distribuição de M. lavali para o Alto Chaco no Paraguai e para a Mata Atlântica no Brasil e Paraguai. Esses registros ampliam a distribuição da espécie na América do Sul em aproximadamente 2,000 km para o sudoeste e 400 km para o leste, revelando ainda a sintopia de M. lavali e M. nigricans no Alto Chaco paraguaio. Esses resultados têm implicações taxonômicas e ecológicas para M. lavali. Fornecemos ainda comentários sobre a história natural e reprodução dessa espécie na Caatinga do nordeste do Brasil.

6 June 2013 MORATELLI AND WILSON NATURAL HISTORY OF MYOTIS LAVALI 655 ACKNOWLEDGMENTS The following curators and collection staff provided access to specimens under their care: A. L. Peracchi (Universidade Federal Rural do Rio de Janeiro, Brazil); F. de C. Passos (Universidade Federal do Paraná, Brazil); M. de Vivo and J. G. Barros (Museu de Zoologia da Universidade de São Paulo, Brazil); N. Simmons and E. Westwig (American Museum of Natural History, United States); and K. Helgen, D. Lunde, and L. Gordon (National Museum of Natural History, United States). A. L. Gardner (United States Geological Survey Patuxent Wildlife Research Center, United States) provided information on specimens he collected in northeastern Brazil. R. Gregorin (Universidade Federal de Lavras, Brazil) and R. L. M. Novaes (Universidade Federal do Estado do Rio de Janeiro, Brazil) provided photographs of specimens from Tocantins and Pernambuco, respectively. L. A. M. da Silva (Universidade Federal de Pernambuco, Brazil) provided photographs of the Caatinga of Pernambuco where specimens of M. lavali were captured. M. A. Mares and an anonymous reviewer provided comments that improved the manuscript. This work was partially supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico, Brazil (CNPq /2011-2). LITERATURE CITED ALLEN, J. A New South American bats and a new octodont. Bulletin of the American Museum of Natural History 33: ASTÚA, D. M., AND D. Q. GUERRA Caatinga bats in the mammal collection of the Universidade Federal de Pernambuco. Chiroptera Neotropical 14: BAUD, F. J Myotis aelleni, nov. sp., chauve-souris nouvelle d Argentine (Chiroptera: Vespertilionidae). Revue Suisse de Zoologie 86: BURTT, E. H., JR., AND J. M. ICHIDA Gloger s rule, featherdegrading bacteria, and color variation among song sparrows. Condor 106: CARO, T The adaptative significance of coloration in mammals. BioScience 55: DICE, L. R., AND P. M. 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7 656 JOURNAL OF MAMMALOGY Vol. 94, No. 3 THOMAS, O On Azara s Chauve-souris onzieme (Myotis ruber, Geoff.) and a new species allied to it. Annals and Magazine of Natural History, Series 7, 10: WATERHOUSE, G. R Mammalia. Pp in The zoology of the voyage of the H.M.S. Beagle, under the command of Captain Fitzroy, R. N., during the years 1832 to 1836 (G. R. Waterhouse and C. Darwin, eds.). Smith, Elder and Co., London, United Kingdom. WILLIG, M. R Composition, microgeographic variation, and sexual dimorphism in Caatingas and Cerrado bat communities from northeast Brazil. Bulletin of Carnegie Museum of Natural History 23: WILLIG, M. R. 1985a. Reproductive activity of female bats from northeast Brazil. Bat Research News 26: WILLIG, M. R. 1985b. Reproductive patterns of bats from Caatingas and Cerrado biomes in northeast Brazil. Journal of Mammalogy 66: WILLIG, M. R Bat community structure in South America: a tenacious chimera. Revista Chilena de Historia Natural 59: WILLIG, M. R., AND R. R. HOLLANDER Secondary sexual dimorphism and phylogenetic constraints in bats: a multivariate approach. Journal of Mammalogy 76: WILLIG, M. R., AND M. P. MOULTON The role of stochastic and deterministic processes in structuring Neotropical bat communities. Journal of Mammalogy 70: WILLIG, M. R., R. D. OWEN, AND R. L. COLBERT Assessment of morphometric variation in natural populations: the inadequacy of the univariate approach. Systematic Zoology 35: WILSON, D. E Genus Myotis Kaup, Pp in Mammals of South America, Volume 1: Marsupials, xenarthrans, shrews, and bats (A. L. Gardner, ed.). University of Chicago Press, Chicago, Illinois. Submitted 2 October Accepted 19 October Associate Editor was Ryan W. Norris. APPENDIX I Listed below are specimens examined and included in the classification analysis, organized according to the taxa herein recognized. Specimens examined consist of skins and skulls that are deposited in the following institutions: Universidade Federal Rural do Rio de Janeiro, Seropédica, Brazil (ALP); American Museum of Natural History, New York, United States (AMNH); Universidade Federal do Paraná, Paraná, Brazil (CCMZ-DZUP); Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZUSP); and National Museum of Natural History, Washington, D.C., United States (USNM). Myotis albescens. PARAGUAY: Yaguaron, Paraguarí (AMNH [neotype]); Curuguaty, Canindeyu (AMNH , , , , , , ); Tacuaral, Cordillera (USNM , ). Myotis izecksohni. BRAZIL: Campinhos, Paraná (CCMZ-DZUP 56 59, 61 67, 85 88, 92, 93, 96, 97, 99, 105, , 112, ). Myotis keaysi. PERU: Cordillera Vilcabamba, Cusco (AMNH , , , , ); Paso Carpish, Huánuco (AMNH ); unknown locality, Cusco (AMNH ). Myotis lavali. BRAZIL: Crato, Ceará (USNM , , , [not included in the classificatory analysis]); 6 km South of Exu, Pernambuco (MZUSP [paratype], [paratype], [paratype], [holotype], [paratypes], [paratype], [paratype], [paratype], [paratypes], [paratypes], [paratype], [paratypes]). PARAGUAY: Parque Nacional Teniente Agripino Enciso, Boquerón (USNM [skin], [skull]); Sapucaí, Paraguarí (USNM ). Myotis levis dinellii. ARGENTINA: La Cocha, Tucumán (AMNH ); Córdoba, Córdoba (USNM , ). BOLIVIA: Caballero, Santa Cruz (AMNH ); Tomina, Chuquisaca (AMNH ). Myotis levis levis. ARGENTINA: La Valle, Buenos Aires (USNM , ); unknown locality, Córdoba (USNM ); Puerto Constanza, Entre Rios (USNM ); Los Vasquez, Tucumán (MZUSP 2055). BRAZIL: Porto Rico, Paraná (CCMZ-DZUP 369, , 376, 377, 381, 382, , 391, 393); Casa Grande, São Paulo (MZUSP 16473, , , 16504, 16506, 16510). Myotis nigricans. BRAZIL: Seropédica, Rio de Janeiro (ALP 5132, 5134, , , , 5235, 5327, 5331, 5332, 5338, ). PARAGUAY: Parque Nacional Teniente Agripino Enciso, Boquerón (USNM ); Asunción, Central (USNM , ); Paraguarí, Paraguarí (USNM , , ); Parque Nacional Ybycuí (USNM ); Sapucaí, Sapucaí (USNM , , ). Myotis oxyotus. PERU: Santa Ana, Cusco (USNM , , , ). Myotis riparius. BRAZIL: Utinga, Belém (USNM ); Fazenda Velha, Belém (USNM , ); Mocambo, Pará (ALP 1915, 2002, 2003, 2554, 2557, 2562, 2568, 2587, 2610, 2710). PARAGUAY: Parque Nacional Cerro Corá, Amambay (USNM , ); Sapucaí, Sapucaí (USNM , ). Myotis ruber. BRAZIL: Boracéia, São Paulo (MZUSP 28359, 28367, 28368); Buri, São Paulo (MZUSP , 32975); Cananéia, São Paulo (MZUSP 27595); São Paulo, São Paulo (MZUSP ). PARAGUAY: Sapucaí, Sapucaí (USNM [neotype]). Myotis simus. BOLIVIA: Cercado, Beni (AMNH , , , , ). BRA- ZIL: Borba, Amazonas (AMNH 91886, , 94224); Parintins, Amazonas (AMNH 92983, 93490, , 93922, 93923).