Abstract

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. nigricansx. Both results have taxonomic and ecological implications for M. lavali. Additionally, we provide comments on its natural history and reproduction.

LaVal's myotis (Myotis lavali [Moratelli et al., 2011]) 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 555672 (male) was caught by D. B. Abrell on 22 July 1982; whereas USNM 555673 (female) and USNM 555674 (female) were caught by M. Ludlow, respectively, on 21 and 22 July 1982. The other 2 specimens (females) were collected by W. T. Foster: USNM 104932, 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 115074, represented only by a skull, was captured in Sapucaí, Paraguarí Department, on 20 April 1901.

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 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 555672, which corresponds exactly with the pattern observed in specimens of M. lavali from northeastern Brazil. The pelage colors of USNM 555673 and USNM 555674 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 555672 and USNM 555673 do not agree with identifications of skins. The skull of USNM 555673 resembles that of M. lavali in shape; whereas the skulls of USNM 555672 and USNM 555674 are identical and resemble that of M. nigricans (see Moratelli et al. 2011a). In summary, USNM 555672 resembles M. lavali in pelage color but is more like M. nigricans in the size and shape of skull; USNM 555673 resembles M. nigricans in pelage color and M. lavali in the size and shape of skull; and USNM 555674 resembles M. nigricans in both pelage color and size and shape of skull. The skull of the USNM 104932 fits the description of M. lavali, but the contrast between bases and tips in the dorsal fur is not so evident; and USNM 115074 is a skull only that resembles M. lavali in shape, although it is slightly smaller. An additional female specimen (USNM 555712) 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 expectation-maximization algorithm (Little and Rubin 1987; Strauss et al. 2003). All measurements and estimated values were then log-transformed 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., 2011), 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 555672 and USNM 555674 were classified with M. nigricans (73% and 59%, respectively), whereas USNM 555673 was classified with M. lavali (100%), confirming previous identifications (Table 1). The skull confirmed as belonging to M. lavali (USNM 555673) has a longer and upwardly oriented rostrum when compared with the other 2 specimens classified as M. nigricans (USNM 555672 and 555674). 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 555673 (lavali), and present in USNM 555672 (nigricans) and USNM 555674 (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 555673 and USNM 555674 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 115070, 115076, 115077, 115079, and 115089), except for a few slightly darker individuals (e.g., USNM 531197). These differences may be attributed to time in storage and variation in storage conditions, or different habitat or roost 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. 201 la). USNM 555672 has bases and tips of dorsal hairs strongly contrasting, with medium-brown bases and light-brown tips, similar to M. lavali from Ceará, northeastern Brazil, in the pelage banding pattern (e.g., USNM 555713–555715, 555717, 555718, and 555720–555722). The analyses of skins and skulls lead us to speculate that these parts were interchanged in the specimens USNM 555672 and USNM 555673.

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 = National Museum of Natural History; NE = northeastern; SE = southeastern; S = southern; N = northern.

 albescens lavali nigricans l. dinellii l. levis izecksohni oxyotus keaysi riparias simus ruber 
USNM 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 
104932 — 0.91 0.09 — — — — — — — — 
115074 — 1.00 — — — — — — — — — 
555672 — 0.01 0.73 — — — 0.16 — — — — 
555673 — 1.00 — — — — — — — — — 
555674 — 0.41 0.59 — — — — — — — — 
555712 — 1.00 — — — — — — — — — 
 albescens lavali nigricans l. dinellii l. levis izecksohni oxyotus keaysi riparias simus ruber 
USNM 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 
104932 — 0.91 0.09 — — — — — — — — 
115074 — 1.00 — — — — — — — — — 
555672 — 0.01 0.73 — — — 0.16 — — — — 
555673 — 1.00 — — — — — — — — — 
555674 — 0.41 0.59 — — — — — — — — 
555712 — 1.00 — — — — — — — — — 

The skulls of the other 2 Paraguayan specimens (USNM 104932 and 115074) 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 104932 and present but very low in USNM 115074. 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 104932 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 555712), 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 555713–555715, 555717, 555718, and 555720–555722). 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 104932 [skin? and skull], 115074 [skull], 555672 [skin], 555673 [skull], and 555712 [skin and skull]) and M. nigricans (USNM 555672 [skull] and 555674 [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, 1 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. 201 la). 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.

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.

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.

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

Fig. 2

Myotis lavali (male) from the Caatinga of Pernambuco. Photograph courtesy of the collector, Roberto L. M. Novaes.

Fig. 2

Myotis lavali (male) from the Caatinga of Pernambuco. Photograph courtesy of the collector, Roberto L. M. Novaes.

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.

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.

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, 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 200–300 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 = 4.50 g, females = 4.95 g) and Ceará (males = 4.34 g, females = 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, 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 555712) 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).

Acknowledgements

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 202612/2011-2).

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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 205195 [neotype]); Curuguaty, Canindeyu (AMNH 234317–234320, 234323, 234324, 234326, 234329, 234332, 234333); Tacuaral, Cordillera (USNM 105662, 105664).

Myotis izecksohni.—BRAZIL: Campinhos, Paraná (CCMZ-DZUP 56–59, 61–67, 85–88, 92, 93, 96, 97, 99, 105, 107–110, 112, 196–199). Myotis keaysi.—PERU: Cordillera Vilcabamba, Cusco (AMNH 233850, 233851, 233854, 233857, 236134); Paso Carpish, Huánuco (AMNH 216117); unknown locality, Cusco (AMNH 214371).

Myotis lavali.—BRAZIL: Crato, Ceará (USNM 555713-555715, 555717, 555718, 555720–555722 [not included in the classificatory analysis]); 6 km South of Exu, Pernambuco (MZUSP 18753 [paratype], 18755 [paratype], 18759 [paratype], 18762 [holotype], 18783–18785 [paratypes], 18792 [paratype], 18793 [paratype], 18807 [paratype], 18813–18815 [paratypes], 18819–18821 [paratypes], 18823 [paratype], 18846–18849 [paratypes]). PARAGUAY: Parque Nacional Teniente Agripino Enciso, Boquerón (USNM 555672 [skin], 555673 [skull]); Sapucaí, Paraguarí (USNM 115074).

Myotis levis dinellii.—ARGENTINA: La Cocha, Tucumán (AMNH 256987); Córdoba, Córdoba (USNM 142560, 142561). BOLIVIA: Caballero, Santa Cruz (AMNH 260253); Tomina, Chuquisaca (AMNH 263629).

Myotis levis levis.—ARGENTINA: La Valle, Buenos Aires (USNM 236236, 236237); unknown locality, Córdoba (USNM 252766); Puerto Constanza, Entre Rios (USNM 582461); Los Vasquez, Tucumán (MZUSP 2055). BRAZIL: Porto Rico, Paraná (CCMZ-DZUP 369, 371–373, 376, 377, 381, 382, 384–388, 391, 393); Casa Grande, São Paulo (MZUSP 16473, 16476–16478, 16481–16488, 16504, 16506, 16510).

Myotis nigricans.—BRAZIL: Seropédica, Rio de Janeiro (ALP 5132, 5134, 5171–5176, 5179–5181, 5185–5188, 5235, 5327, 5331, 5332, 5338, 5340–5342). PARAGUAY: Parque Nacional Teniente Agripino Enciso, Boquerón (USNM 555674); Asunción, Central (USNM 105601, 115070); Paraguarí, Paraguarí (USNM 115075–115077, 115079, 115081); Parque Nacional Ybycuí (USNM 531197); Sapucaí, Sapucaí (USNM 102948, 105621, 115089).

Myotis oxyotus.—PERU: Santa Ana, Cusco (USNM 194452, 194453, 195147, 195149).

Myotis riparius.—BRAZIL: Utinga, Belém (USNM 361782); Fazenda Velha, Belém (USNM 361786, 361788–361790); Mocambo, Pará (ALP 1915, 2002, 2003, 2554, 2557, 2562, 2568, 2587, 2610, 2710). PARAGUAY: Parque Nacional Cerro Corá, Amambay (USNM 554538, 554539); Sapucaí, Sapucaí (USNM 115071–115073, 115095).

Myotis ruber.—BRAZIL: Boracéia, São Paulo (MZUSP 28359, 28367, 28368); Buri, São Paulo (MZUSP 32971–32973, 32975); Cananéia, São Paulo (MZUSP 27595); São Paulo, São Paulo (MZUSP 31470–31473). PARAGUAY: Sapucaí, Sapucaí (USNM 115097 [neotype]).

Myotis simus.—BOLIVIA: Cercado, Beni (AMNH 211156, 211167–211169, 211171–211174, 211178–211183, 211190). BRAZIL: Borba, Amazonas (AMNH 91886, 91888–91891, 94224); Parintins, Amazonas (AMNH 92983, 93490, 93492–93496, 93922, 93923).

Author notes

Present address of RM: Division of Mammals, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA
Associate Editor was Ryan W. Norris.