A new species of Myotis (Chiroptera, Vespertilionidae) from Peru

Eight species of Myotis are known to occur west of the Andes, of which four are from western Peru. Based on four vouchers from the arid coast of northern and central Peru, a new species is added to this list. Myotis bakeri species novum externally
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  A N EW  S PECIES   OF    M  YOTIS   (C HIROPTERA , V ESPERTILIONIDAE ) FROM  P ERU  R  ICARDO  M  ORATELLI   , R OBERTO  L  EONAN   M. N  OVAES   , C   ARLOS   C   ARRIÓN   B ONILLA  ,  AND  D ON   E. W   ILSON  A BSTRACT I NTRODUCTION The genus  Myotis  Kaup, 1829 in South America currently comprises 22 species (Moratelli and Wilson 2014; Moratelli et al. 2017). The Andean Cordillera  plays an important role in the distribution of most of this assemblage, acting as a corridor for a few species and a partial or complete barrier for others (LaVal 1973; Moratelli et al. 2013). Among the South American as - semblage,  Myotis albescens  (É. Geoffroy, 1806),  Myotis nigricans  (Schinz, 1821), and  Myotis riparius  Handley, 1960, are widespread on the continent, occurring on  both sides of the Cordillera. Southernmost records on the west side of the Andes are in southwestern Ecuador for  Myotis riparius , and northwestern Peru for  Myotis albescens  and  Myotis   nigricans .  Myotis caucensis  J. A. Allen, 1914,  Myotis keaysi  J. A. Allen, 1914,  Myotis oxyotus  (Peters, 1866), and  Myotis pilosatibialis  LaVal, 1973, occur throughout the Cordillera.  Myotis keaysi  and  M. pilosatibialis  are restricted to mid-elevations (≈ 900–2000 m), whereas  M. oxyotus reaches the lowlands on the west side of the Cordillera, and  M. caucensis  occurs in intermontane valleys and adjacent Amazon lowlands on the east side.  Myotis levis  (I. Geoffroy, 1824),  Myotis ruber   (É. Geoffroy, 1806), and  Myotis izecksohni  Moratelli et al., 2011, are restricted to east - ern South America.  Myotis izecksohni  occurs in the mountains of eastern Brazil and Argentina, whereas  M. levis  and  M. ruber   occur from Paraguay eastward to the lowlands of southern Brazil and Argentina, and northward throughout the mountains of eastern Brazil. Apparently,  Myotis simus  Thomas, 1901, is restricted to the lowlands of the Amazon River. One of the authors (RM) has reanalyzed all vouchered specimens of  Myotis  simus  Thomas, 1901, supporting the occurrence of the species for the west side of the Andes, and found that they correspond to misidentications of other species in the ruber   group (R. Moratelli, unpublished data).  Myotis midastactus  Moratelli and Wilson, 2014, is known from only the Cerrado in Bolivia and Alto Chaco in Paraguay.  Myotis chiloensis  (Waterhouse, 1840) is restricted to localities in southwestern Chile and Argentina.  Myotis dinellii  Thomas, 1902, occurs in Argentina and southern Bolivia.  Myotis atacamensis  (Lataste, 1892) occupies arid and semiarid habitats in western Peru and Chile.  Myotis diminutus  Moratelli and Wilson, 2011, is in western Colombia and Ecuador, where the species appears to be restricted to the Chocó Ecoregion.  Myotis handleyi  Moratelli et al., 2013, is known from two cordilleras in northern Venezuela, and  M. nesopolus  Miller, 1900, occurs in a restricted arid zone in northwestern Venezuela (  M. nesopolus larensis  LaVal, 1973) and on Curaçao (  M. nesopolus nesopolus ).  Myotis attenboroughi  Moratelli et al., 2017, is endemic to the Island of Tobago, Trinidad and Tobago, and  Myotis clydejonesi  Moratelli et al., 2016, is known from a single individual collected in tropi - cal lowland forest in Sipaliwini, Suriname. Finally,  Myotis lavali  Moratelli et al., 2011, occurs throughout the South American dry corridor (LaVal 1973; Wilson 2008; Stevens et al. 2010; Muñoz-Garay and Mantilla-Meluk 2012; Mantilla-Meluk and Muñoz-Garay 2014; Moratelli and Wilson 2014, 2015; Moratelli et al. 2013, 2015a,b, 2016, 2017; Barquez et al. 2017).Eight species of  Myotis  are known to occur west of the Andes, of which four are from western Peru. Based on four vouchers from the arid coast of northern and central Peru, a new species is added to this list.  Myotis   bakeri species novum externally resembles  M. atacamensis   but it can be distinguished from this species and all other South American congeners by a unique set of external and craniodental qualitative and quantitative characters. Herein, this new spe - cies that inhabits the lowland arid regions on the northern and central coasts of western Peru is named and described. As a result, 22 species of  Myotis  are now known from South America, nine of which occur in Peru. Key words: desert, Myotinae,  Myotis bakeri  species novum, neotropics, South America 239  240 S PECIAL  P UBLICATIONS , M USEUM   OF  T EXAS  T ECH  U NIVERSITY Moratelli and Wilson (2014) and Moratelli et al. (2013) arranged this assemblage into the albescens  and ruber   groups based on the oldest available names. Spe - cies in each of these groups share external and cranial features, and have been partially retrieved as closely related in the molecular phylogenies of Stadelmann et al. (2007) and Larsen et al. (2012a), but several species are still pending positioning in phylogenies. Species in the albescens  group generally have silky fur, rounded occipital region, and either lack a sagittal crest or the crest is low and weakly developed. This species group includes  M. albescens ,  M. atacamensis ,  M. caucensis ,  M. chiloensis ,  M. diminutus ,  M. dinellii ,  M. izecksohni ,  M. lavali ,  M. levis ,  M. nesopolus ,  M. nigricans , and  M. oxyotus . Species in the ruber   group are characterized  by woolly fur, generally attened occipital region, and the presence of a sagittal crest, which usually varies from moderately developed to high. The ruber   group includes  M. keaysi ,  M. midastactus ,  M. pilosatibialis ,  M. riparius ,  M. ruber  , and  M. simus .Seven species are found west of the Andes—  al-bescens , atacamensis , chiloensis , diminutus , nigricans , oxyotus , and riparius  —with all but  M. riparius  in the albescens  group (LaVal 1973; Wilson 2008; Moratelli et al. 2013). Based on four vouchers from desert localities in western Peru (Lambayeque and Lima), a new species of  Myotis, named and described herein, is added to this list. The new species is morphologically a member of the albescens  group, and particularly allied with  M. atacamensis  —the name under which the vouchers were misidentied. Although similar in external appearance, this new species shows a particular set of external and cranial traits that distinguish it from  M. atacamensis  and all other South American congeners. M ETHODS This research is part of a critical review of col - lections of Neotropical  Myotis , and more than 3,800 specimens from different localities in South America, covering all species currently recognized, have been examined. The new species was compared with 286 vouchers representing all species currently recog - nized from Peru. These specimens are preserved in the collections of the American Museum of Natural History (AMNH, New York), Carnegie Museum of  Natural History (CM, Pittsburgh, Pennsylvania), The Field Museum (FMNH, Chicago, Illinois), Muséum d’histoire naturelle (MHNG, Geneva, Switzerland), Museum of Natural Science, Louisiana State Univer  - sity (LSUMZ, Baton Rouge), Museum of Texas Tech University (TTU, Lubbock), Museum of Vertebrate Zoology (MVZ, Berkeley, California), Natural History Museum, University of Kansas (KU, Lawrence), and  National Museum of Natural History, Smithsonian Institution (USNM, Washington, DC).Four vouchers, three adult males and one adult female, constitute the source material for the descrip - tion of this new species. They were collected in the departments of Lambayeque (LSUMZ 21306 ♂, 21307 ♀) and Lima (MVZ 137906 ♂, 137907 ♂), western Peru, between 1969 and 1981. The complete list of specimens from Peru analyzed here is presented in Appendix I, including those that could not be identi -ed. Measurements are in millimeters (mm) and the  body mass in grams (g), and are from adults and one subadult (the holotype of  M. diminutus ). The total length (TL), tail, hind foot, ear, and body mass were recorded from skin labels, and reported to the nearest millimeter or nearest gram. Other measurements were taken using digital callipers accurate to 0.02 mm. Cra - niodental measurements were taken under a binocular microscope at low magnication (usually 6x). These dimensions were recorded and analyzed to the near  - est 0.01 mm, but values were rounded off to 0.1 mm throughout the text because this is the smallest unit that allows accurate repeatability with calipers (Voss et al. 2013). Measurements include forearm length (FA), third metacarpal length (3MC), length of dorsal hair (LDH), length of ventral hair (LVH), greatest length of skull (GLS), condylocanine length (CCL), condylobasal length (CBL), condylo-incisive length (CIL), basal length (BAL), zygomatic breadth (ZB), mastoid breadth (MAB), braincase breadth (BCB), interorbital breadth (IOB), postorbital breadth (POB),  breadth across canines (BAC), breadth across molars (BAM), maxillary toothrow length (MTL), length of the upper molars (M1–3), mandibular length (MAL), and mandibular toothrow length (MAN). These mea -surements are dened in Moratelli et al. (2013). The cranial index—CRI = ((IOB + BCB) x GLS)/2—used  by Moratelli and Wilson (2011, 2014), was applied  M ORATELLI   ET   AL .—N EW  S PECIES   OF    M  YOTIS    FROM  P ERU  241 to compare skull shape. Descriptive statistics (mean and range) were calculated for all dimensions. The statistical significance of craniometric differences among samples was assessed by single analyses of variance (one-way ANOVA). Based on a subset of the craniodental dimensions (GLS, CCL, BCB, POB, BAM, MTL, M1–3, MAL, MAN, BAC), a discriminant function analysis (DFA) was performed to compare the new species with  M. atacamensis ,  M. diminutus , and  M. nigricans. Myotis atacamensis  and  M. nigricans  were included because they are the most similar species in external and craniodental features, respectively.  Myotis   diminutus  was included due to its geographic proximity, although it is less similar as compared to the other taxa. DFA was performed using MASS and Lattice pack  - ages in R software (R Development Core Team 2008). Multivariate procedures require complete datasets, thus missing data (approximately 5% of total dataset) were estimated from the existing raw data using the Amelia II packpage (Honaker et al. 2011) implemented in the R software. The complete list of specimens used in the statistical analyses is in Appendix II. Capitalized color nomenclature is from Ridgway (1912). R  ESULTS  Myotis bakeri  , species novum Baker’s Myotis,  Myotis de Baker  Figs. 1, 2, and 4; Tables 1 and 2  Myotis atacamensis  Wilson 2008:472; part, not  M. atacamensis  (Lataste, 1892).  Myotis atacamensis  Rodríguez-San Pedro et al. 2014;  part, not  M. atacamensis  (Lataste, 1892).  Holotype and type locality .—The holotype (MVZ 137907) consists of the skin and skull of an adult male (Figs. 1 and 2) collected by M. L. Hawes (eld number 232) on 30 July 1969 at about 7 km SE of Chilca (≈ 12°33′S, 76°41′W; ca. 200–250 m altitude, obtained from Google Earth), department of Lima, Peru.  Paratypes.  —One adult female (LSUMZ 21307) and two adult males (LSUMZ 21306, MVZ 137906). MVZ 137906 is from the type locality. LSUMZ 21306 and 21307 are from the department of Lambayeque, Peru, about 12 km N of Olmos. See Table 1 for external and craniodental measurements and body mass for the type series. These vouchers include skin and skull, and those from Lambayeque (LSUMZ 21306, 21307) include parts of the post-cranial skeleton.  Distribution and habitat.  —   Myotis bakeri  is known from single localities in the departments of Lima and Lambayeque, Peru (Fig. 3). Specimens from Lima are from ca. 7 km SE of Chilca (≈ 12°33′S, 76°41′W; Google Earth), and those from Lambayeque are from ca. 12 km N of Olmos (05°55′S, 79°47′W; Gardner’s Gazetteer of marginal localities [Gardner 2008]). Information about elevation is available only for specimens from Lambayeque, 12 km N of Olmos (ca. 335 m [1100 ft] above sea level). The elevation obtained from Google Earth for Lima, 7 km SE of Chilca, is ca. 200–250 m above sea level. Based on these records,  M. bakeri  is restricted to western Peru, where it inhabits the lowland arid formations on the northern and central Pacic coast. Localities are in the extreme north of the South Ameri - can Transition Zone, Desert province (see Morrone 2014:24, g. 12). Morrone’s (2014) Desert province corresponds to Cabrera and Willink’s Desert province (Cabrera and Willink 1973). In this region, vegetation is scarce and usually found along margins of rivers and near the ocean (Morrone 2001).  Diagnosis.  —Similar to most species of  Myotis , a single character was not found that could distinguish  M. bakeri  from all other American congeners. However,  M. bakeri  can be distinguished from other Neotropi - cal species in the genus by the unique combination of the following set of traits: silky fur; dorsal pelage strongly bicolored, with Mummy Brown bases (2/3), and Buckthorn Brown tips (1/3 of the total fur length); absence of a fringe of hairs along the trailing edge of uropatagium (although a few sparse hairs may be pres - ent); plagiopatagium broadly attached to the side of the foot at the level of the toes; rounded occipital region; low, weakly developed sagittal crest; inated, but not rounded braincase.  242 S PECIAL  P UBLICATIONS , M USEUM   OF  T EXAS  T ECH  U NIVERSITY Figure 1. Dorsal (A) and ventral (B) views of the skin of the holotype of  Myotis bakeri  (MVZ 137907). Scale bar = 10 mm. See Table 1 for measurements.Figure 2. Dorsal (A), ventral (B), and lateral (C) views of the skull, and lateral (D) and dorsal (E) views of the mandible of the holotype of  Myotis bakeri  (MVZ 137907). Scale bar = 10 mm. See Table 1 for measurements.  M ORATELLI   ET   AL .—N EW  S PECIES   OF    M  YOTIS    FROM  P ERU  243 Table 1. Selected measurements (mm) and body mass (g) of the holotype (MVZ 137907) and paratypes of  Myotis bakeri  (LSU 21306, 21307, MVZ 137906). See Methods for variable abbreviations and Appendix I for localities of specimens .CharactersMVZ 137907 ♂ MVZ 137906 ♂ LSU 21306 ♂ LSU 21307 ♀ TL79747372Tail30263031Hind foot7677 Ear 14121212Body mass4. LDH7.– 6.46.1GLS13.813.513.413.1CCL12.211.811.711.7 CBL12.712.412.312.3CIL12.912.612.512.5BAL11.411.311.211.1ZB–  ––– MAB7. BAC3.–
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