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Catalogue
Catalogue of primary types of Neotropical Myotis (Chiroptera, Vespertilionidae)
expand article infoRoberto Leonan M. Novaes, Don E. Wilson§, Ricardo Moratelli
‡ Fundação Oswaldo Cruz, Fiocruz Mata Atlântica, Rio de Janeiro, Brazil
§ National Museum of Natural History, Washington, DC, United States of America
Open Access

Abstract

Myotis comprises a diverse group of vespertilionid bats with worldwide distribution. Neotropical Myotis have an accentuated phenotypic conservatism, which makes species delimitation and identification difficult, hindering our understanding of the diversity, distribution, and phylogenetic relationships of taxa. To encourage new systematic reviews of the genus, a catalogue of the primary types and names is presented, current and in synonymy, for Neotropical Myotis. Currently 33 valid species (and three subspecies) are recognized, and their primary types are deposited in 12 scientific collections in the USA (30 types), Brazil (two types), England (two types), and France (one type). The names of 29 Neotropical Myotis species currently in synonymy were found. However, it is possible that some synonyms represent independent evolutionary lineages, considering recent results provided by taxonomic revisions.

Resumo

Myotis compreende um grupo diverso de morcegos vespertilionídeos amplamente distribuídos ao redor do planeta. Myotis neotropicais têm um acentuado conservatismo fenotípico, o que dificulta a delimitação e identificação de espécies a partir de caracteres morfológicos, criando barreiras para a compreensão da diversidade, distribuição e relações filogenéticas entre os táxons. Visando encorajar novas revisões sistemáticas para o gênero, é apresentado este catálogo de nomes e tipos primários, válidos e sob sinonímia, para Myotis neotropicais. Atualmente, são reconhecidas 33 espécies válidas (e três subespécies) e seus tipos estão depositados em 12 coleções científicas nos EUA (30 tipos), Brasil (2 tipos), Inglaterra (2 tipos) e França (1 tipo). Compilamos 29 nomes de espécies de Myotis atualmente sob sinonímia. Entretanto, é possível que alguns desses sinônimos possam representar linhagens evolutivas independentes, considerando resultados recentes de revisões taxonômicas.

Keywords

Myotinae, taxonomy, type locality, type specimen, vespertilionid bats, zoological nomenclature

Palavras-chave

espécime-tipo, localidade-tipo, morcego vespertilionídeo, Myotinae, nomenclatura zoológica, taxonomia

Introduction

Taxonomy is the discipline of Biology responsible for describing, classifying, and naming organisms, as well as hypothesize about the evolutionary relationships between taxa (Tancoigne et al. 2011). Therefore, understanding and organizing biological diversity is the primary task of the taxonomist. Taxonomic studies have profound implications in virtually all areas of the biological sciences, such as ecology, evolution, genetics, and epidemiology, in addition to directly influencing public policies focused on health and environment (Cracraft 2002; Pearson et al. 2011; Cook et al. 2020). Furthermore, knowing the real diversity of organisms on our planet is critical for the sustainable use of natural resources and for the management and conservation of species (May 1988), especially in the current biodiversity crisis, where the rate of extinction indicates that we are witnessing a sixth mass extinction (Ceballos et al. 2015, 2017).

Species are the central unit of taxonomy and the association between an unambiguous scientific name and a species is of paramount importance for a reliable biological information system (Wheeler 2004). For that, the existence of primary types, which are those specimens designated as the name-bearing representative of a species is essential. In addition to serving as a reference point for the existence of any organism, type-specimens are a particularly important source of information for scientists to track and unravel the taxonomic history of biologically complex groups, such as bats of the genus Myotis Kaup, 1829.

Myotis is the most speciose genus of bats and the second largest genus of mammals, with more than 140 extant species (MDD 2021). It is also the genus with the greatest area of distribution among non-human mammals (Moratelli et al. 2019a). The greatest diversity and abundance of Myotis is reported in temperate and subtropical areas (Nowak 1994; Moratelli et al. 2019a). However, recent systematic reviews have indicated that there is a high diversity of Myotis in the Neotropics (e.g., Larsen et al. 2012; Moratelli et al. 2011a, 2013, 2016, 2017, 2019b; Carrión-Bonilla and Cook 2020; Novaes et al. 2021a, b, c). Nevertheless, our knowledge of species limits, name validity, and distributional boundaries for several Neotropical Myotis species remains incipient.

Part of the taxonomic hurdle is due to the accentuated morphological conservatism and lack of specimen series covering all geographic distributions (Menu 1987; Smith et al. 2012; Moratelli et al. 2019a). On the other hand, molecular studies have revealed the existence of more independent evolutionary lineages than species recognized from morphology-based taxonomy (Larsen et al. 2012; Novaes et al. 2021a, b). This indicates the existence of hidden diversity possibly composed of multiple cryptic species, which challenges the delimitation of species and raises the need for new systematic reviews, especially those based on multiple lines of evidence.

To contribute to the organization of systematic knowledge about this genus, and to support future studies of taxonomy, we present a catalogue of the primary types of Neotropical Myotis. Later, we briefly comment on the validity and distribution of some species.

Materials and methods

The catalogue was mostly compiled by analysis of the specimens deposited in 12 biological collections: American Museum of Natural History (New York, USA), Field Museum of Natural History (Chicago, USA), Louisiana State University Museum of Natural Science (Baton Rouge, USA), Museum of Texas Tech University (Lubbock, USA), Museum of Vertebrate Zoology at University of California (Berkeley, USA), Kansas University Biodiversity Institute and Natural History Museum (Lawrence, USA), Natural History Museum, Los Angeles County (Los Angeles, USA), Smithsonian’s National Museum of Natural History (Washington D.C., USA), Natural History Museum, London (London, UK), Zoologisches Staats-Sammlung München (Munich, Germany), Muséum National D’Histoire Naturelle (Paris, France), Muséum d’Histoire Naturelle (Geneva, Switzerland), Museu de Zoologia da Universidade de São Paulo (São Paulo, Brazil), Universidade Federal Rural do Rio de Janeiro (Seropédica, Brazil). When it was not possible to visit the collection to examine the type specimen, the information was retrieved from the original species descriptions or other available bibliography (e.g., LaVal 1973; Carter and Dolan 1978; Carrión-Bonilla and Cook 2020) and by direct consultation with the curators of the collections. Abbreviations of biological collections cited in the text are available below.

ALP Universidade Federal Rural do Rio de Janeiro, Seropédica, Brazil;

ANSP Academy of Natural Sciences of Drexel University, Philadelphia, USA;

AMNH American Museum of Natural History, New York, USA;

BMNH Natural History Museum, London, UK;

FMNH Field Museum of Natural History, Chicago, USA;

KU Natural History Museum, Kansas University, Lawrence, USA;

LACM Natural History Museum, Los Angeles County, Los Angeles, USA;

LSU Louisiana State University Museum of Natural Sciences, Baton Rouge, USA;

MHNG Muséum d’Histoire Naturelle, Geneva, Switzerland;

MNHN Muséum National D’Histoire Naturelle, Paris, France;

MSB Museum of Southwestern Biology, University of New Mexico, Albuquerque, USA;

MVZ Museum of Vertebrate Zoology, University of California, Berkeley, USA;

MZUSP Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil;

RNH Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands;

TTU Museum of Texas Tech University, Lubbock, USA;

USNM Smithsonian’s National Museum of Natural History, Washington D.C., USA;

ZSM Zoologisches Staats-Sammlung München, Munich, Germany.

The list of Myotis species adopted here is based on systematic reviews conducted for the genus Myotis in the Neotropical region (i.e., LaVal 1973; Bogan 1978; Moratelli et al. 2019a, b; Carrión-Bonilla and Cook 2020; Novaes et al. 2021a, b, c). Following LaVal (1973), we excluded species from the definition of Neotropical Myotis when their distributions extend from the Nearctic into the Neotropics. Geographical coordinates of type localities were retrieved, when available, directly from the original publications or by consulting the museum database and the gazetteer of Gardner (2008). In cases where they were not available, we used proximal coordinates of the locality from the search in the USA’s National Geospatial – Intelligence Agency (https://geonames.nga.mil/namesgaz/). We follow the International Code of Zoological Nomenclature (ICZN 1999) as a reference for the categories of type specimens.

The list of names was divided in two parts, the first with accounts of name-bearing type specimens of currently recognized species; and the second with accounts of name-bearing type specimens in synonymy. The accounts were arranged chronologically, following the date of taxa description. The format of accounts was inspired by Fisher and Ludwig (2015), but with modifications. Each account reads as follows: (i) Current name (for recognized species) or original published name (for names in synonymy) followed by the author’s or authors’ names; (ii) Original citation, including publication, volume, pages, and year of publication; (iii) Taxonomy, species original published name if different from the currently name, followed by information on subspecies, if any; (iv) Type designation as holotype, lectotype, paralectotype, neotype, or syntype, including collection number, age and sex, date collected and collector(s) name(s), and preparation of specimen; (v) Type locality: Verbatim locality as given in the original description or neotype designation, published restrictions, and supplementary data. Abbreviations are used for miles (mi), kilometers (km), feet (ft), and meters (m); (vi) Remarks, with additional information is provided as needed, but especially to explain types designated subsequent to description.

Results

Name-bearing type specimens of recognized species

For the 33 species (and three subspecies) of Neotropical Myotis currently recognized (Table 1), primary types are deposited in 12 zoological collections in the USA (eight collections), Brazil (two collections), England (one collection) and France (one collection). The USA is home to 30 primary types of Neotropical Myotis, while Brazil and England are home to two primary types each and France to one type specimen. The collections with the largest number of primary types are the Smithsonian’s National Museum of Natural History (11 types), followed by the American Museum of Natural History (eight types), both in the USA. The other collections have 1–4 type specimens each (Fig. 1).

Table 1.

Valid species and subspecies of Neotropical Myotis including information on their primary types.

Species Type specimen Proximal type locality
M. albescens Neotype AMNH 205195 Paraguarí, Paraguay
M. armiensis Holotype MSB 262089 Chiriquí, Panamá
M. atacamensis Neotype USNM 391786 Tarapacá, Chile
M. attenboroughi Holotype USNM 540693 St. John Parish, Tobago Island
M. bakeri Holotype MVZ 136907 Lima, Peru
M. carteri Holotype LACM 36876 Jalisco, Mexico
M. caucensis Holotype AMNH 32787 Valle del Cauca, Colombia
M. chiloensis Neotype FMNH 24029 Chiloé Island, Chile
M. clydejonesi Holotype TTU 109227 Sipaliwini, Suriname
M. cobanensis Holotype AMNH 145017 Alta Verapaz, Guatemala
M. diminutus Holotype USNM528569 Los Ríos, Ecuador
M. dinellii Holotype BMNH 0.7.9.4 Tucumán, Argentina
M. dominicensis Holotype USNM 113564 Dominica
M. elegans Holotype KU 88398 Veracruz, Mexico
M. findleyi Holotype USNM 512417 Islas Tres Marías, Mexico
M. handleyi Holotype USNM 370932 Distrito Federal, Venezuela
M. izecksohni Holotype ALP 6675 Rio de Janeiro, Brazil
M. keaysi Holotype AMNH 15814 Puno, Peru
M. larensis Holotype AMNH 130709 Lara, Venezuela
M. lavali Holotype MZUSP 18762 Pernambuco, Brazil
M. levis Syntype MNHN 1997-1805 Southern Brazil
M. martiniquensis Holotype AMNH 214062 Tartane, Martinique
M. midastactus Holotype AMNH 211156 Beni, Bolívia
M. moratellii Holotype USNM 513482 Los Ríos, Ecuador
M. nesopolus Holotype USNM 101849 Curaçao, Netherlands Antilles
M. n. nigricans Neotype LACN 36877 Rio de Janeiro, Brazil
M. n. extremus Holotype USNM 77670 Chiapas, Mexico
M. n. osculati Not located Eastern Ecuador
M. nyctor Holotype KU 109473 St. Thomas Parish, Barbados
M. o. oxyotus Neotype LACN 36878 Carchi, Ecuador
M. o. gardneri Holotype LSU 12924 San José, Costa Rica
M. pampa Holotype AMNH 205471 Artigas, Uruguay
M. pilosatibialis Holotype LACN 36879 Francisco Morazán, Honduras
M. riparius Holotype USNM 310255 Darién, Panamá
M. ruber Neotype USNM 115097 Paraguarí, Paraguay
M. simus Holotype BMNH 8.5.12.2 Loreto, Peru
Figure 1. 

Number of primary type specimens of Neotropical Myotis deposited in zoological collections. The name of the institution for each acronym shown in the graph is described in the methods.

Of all recognized Neotropical Myotis types, 28 are holotypes, six are neotypes, and one is syntype. Only one taxon lacks a type specimen (M. nigricans osculati), which presumably was destroyed. About 95% of the type specimens are preserved as skin and skull, with mandible; while only 5% are preserved in fluid (usually alcohol 70°GL). Most types (80%) are in a good condition, with complete skulls and untorn skin. The other 20% are damaged, especially the oldest ones. Damages include broken skulls, loss of bone elements, or torn skins (Figs 28).

Figure 2. 

Some type specimens from valid species of Neotropical Myotis: AMNH 205195, neotype of M. albescens; USNM 391786, neotype of M. atacamensis; USNM 540693, holotype of M. attenboroughi; MVZ 136907, holotype of M. bakeri.

Figure 3. 

Some type specimens from valid species of Neotropical Myotis: LACM 36876, holotype of M. carteri; AMNH 32787, holotype of M. caucensis; FMNH 24029, neotype of M. chiloensis; TTU 109227, holotype of M. clydejonesi.

Figure 4. 

Some type specimens from valid species of Neotropical Myotis: USNM528569, holotype of M. diminutus; KU 88398, holotype of M. elegans; USNM 512417, holotype of M. findleyi; USNM 370932, holotype of M. handleyi.

Figure 5. 

Some type specimens from valid species of Neotropical Myotis: ALP 6675, holotype of M. izecksohni; AMNH 15814, holotype of M. keaysi; AMNH 130709, holotype of M. larensis; MZUSP 18762, holotype of M. lavali.

Figure 6. 

Some type specimens from valid species of Neotropical Myotis: MNHN 1997-1805, syntype of M. levis; AMNH 211156, holotype of M. midastactus; USNM 513482, holotype of M. moratellii; LACN 36877, neotype of M. nigricans.

Figure 7. 

Some type specimens from valid species of Neotropical Myotis: LACN 36878, neotype of M. oxyotus; AMNH 205471, holotype of M. pampa; LACN 36879, holotype of M. pilosatibialis; USNM 310255, holotype of M. riparius.

Figure 8. 

Some type specimens from valid species of Neotropical Myotis: USNM 115097, neotype of M. ruber; BMNH 8.5.12.2, holotype of M. simus; USNM 113564, holotype of M. dominicensis.

Below is an annotated list of Neotropical Myotis species (organized in chronological order of description), with information about the primary specimen types and the type locality. We include a map containing the geographical point of all type localities (Fig. 9).

Figure 9. 

Type localities of the currently valid species (red) and subspecies (blue) of Neotropical Myotis.

Myotis ruber (É. Geoffroy, 1806)

Annales du Muséum d’Histoire Naturelle 8: 187–205.

Taxonomy: Described as Vespertilio ruber by Geoffroy Saint-Hilaire (1806: 204). Treated as monotypic (Wilson 2008; Moratelli et al. 2019a).

Neotype: USNM 115097, adult male collected on May 22, 1901 by W. Foster; skull, mandible, and skin.

Type locality: Sapucay (= Sapucai, Paraguarí), Paraguay (25°40'S, 56°57'W; ca. 200 m a.s.l.) by neotype designation (LaVal 1973: 45).

Remarks: The holotype was not specified by the author, who based his description on the Azara’s (1801) “chauve-sourris cannelle”. LaVal (1973) noted that D. C. Carter did not locate the specimen at the Muséum National d’Histoire Naturelle (Paris, France) or in any other European museum, concluding that it was lost or destroyed. The neotype was designated by LaVal (1973: 45), following the same reasoning presented for M. albescens (see below).

Myotis albescens (É. Geoffroy, 1806)

Annales du Muséum d’Histoire Naturelle 8: 187–205.

Taxonomy: Described as Vespertilio albescens by Geoffroy Saint-Hilaire (1806: 204–205). Treated as monotypic (Moratelli and Oliveira 2011; Moratelli et al. 2019a).

Neotype: AMNH 205195, adult female collected on June 2, 1963 by M. D. Tuttle; skull, mandible, complete post-cranial skeleton and skin.

Type locality: Yaguarón, Paraguarí, Paraguay (25°33'S, 57°17'W; ca. 200 m a.s.l.) based on neotype designation (LaVal 1973: 26).

Remarks: The holotype was not specified by the author, who based his description on the Azara’s (1801) “chauve-souris donzième”. LaVal (1973) noted that D. C. Carter was unable to locate a specimen in European zoological collections from Azara’s expedition. According to Cabrera (1958), É. Geoffroy based his description of Phyllostoma lineatum (currently Platyrrhinus lineatus) on specimens collected in Paraguay and listed by Azara (1801) in the same publication in which he listed the specimen that Geoffroy described as Vespertilio albescens. Cabrera stated that the specimen of P. lineatum was destroyed. LaVal (1973: 26) considered that the type specimen of V. albescens met the same fate and designated a neotype.

Myotis nigricans (Schinz, 1821)

Taxonomy: Originally Vespertilio nigricans Schinz, 1821. We follow Bogan (1978) and Moratelli et al. (2019a) in recognizing three subspecies, and in treating M. carteri as a distinct species, instead of a subspecies of M. nigricans.

Myotis nigricans nigricans (Schinz, 1821)

In “Das tierreich eingetheilt nach dem Bau der thiere als Grundlage ihrer Naturgeschichte und der vergleichenden Anatomie von dem Herrn Ritter von Cuvier, volume 1”. Saugethiere und Vögel, Stuttgart and Tübingen, 894 pp.

Taxonomy: Described as Vespertilio nigricans by Schinz (1821: 179).

Neotype: LACM 36877, adult female collected on October 14, 1968 by A. L. Peracchi; skull, mandible, and skin.

Type locality: Seropédica, 42 km S Rio de Janeiro, Brazil (22°45'S, 43°41'W; 33 m a.s.l.).

Remarks: The species was described based on a specimen collected by Prinz Maxililian zu Wied-Neuwied at Fazenda do Agá, near the Rio Iritiba, Espírito Santo, Brazil. Miller and Allen (1928) were not able to confirm the existence of this specimen. From a personal communication of D. C. Carter, that he could not locate it in among the important collection of Wied’s Brazilian specimens at the American Museum of Natural History, LaVal (1973: 9) presumed it has been lost and designated a neotype.

Myotis nigricans osculati (Cornalia, 1849)

In “Vertebratorum synopsis in Museo Mediolanense extantium que per novam orbem Cajetanus Osculati collegit annis 1846–47–1848 (…)”. Typographia Corbetta, Modoetiae, 16 pp.

Taxonomy: Described as Vespertilio osculati by Cornalia (1849: 11).

Type specimen: None. The specimen used in the original description was collected by G. Osculati between 1846 and 1848. Osculati’s collection deposited in the Museo di Storia Naturale di Milano, Italy, in 1848, was destroyed during World War II (Cagnolaro and Violani 1988; Moratelli et al. 2013).

Type locality: Eastern Ecuador.

Myotis nigricans extremus Miller & Allen, 1928

Bulletin of the United States National Museum 144: 1–218.

Taxonomy: Described as Myotis nigricans extremus by Miller and Allen (1928: 181).

Holotype: USNM 77670, adult female collected by the E. W. Nelson and E. A. Goldman on March 1, 1896; skull, mandible, and skin.

Type locality: Huehuetán, Chiapas, Mexico (15°01'N, 92°22'W; 91 m a.s.l.).

Myotis levis (I. Geoffroy, 1824)

Annales de Sciences Naturelles de Paris 3: 440–447.

Taxonomy: Described as Vespertilio levis by Geoffroy Saint-Hilaire (1824: 444); currently monotypic (Moratelli et al. 2019a).

Syntype: MNHN type 203 (also referred to as MNHN 1997-1805), adult, sex unknown, collected by A. Geoffroy Saint-Hilaire (date not specified); mounted specimen, with skull removed and severely damaged.

Type locality: Southern Brazil.

Myotis chiloensis (Waterhouse, 1840)

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”. Smith, Elder and Co, London, 97 pp.

Taxonomy: Described as Vespertilio chiloensis by Waterhouse (1840: 5); currently monotypic (Novaes et al. 2018; Moratelli et al. 2019a).

Neotype: FMNH 24029, adult female collected by J. Vera in 1923; skull partially damaged, mandible, and skin.

Type locality: Cucao, Chiloé Island, Los Lagos, Chile (42°38'S, 74°06'W; sea level).

Remarks: The original description was based on a specimen collected in January 1836 by Lieutenant Sullivan and given to C. R. Darwin during the H. M. S. Beagle voyage. Miller and Allen (1928) were unable to locate the specimen. LaVal (1973: 43) presumed it lost and designated a neotype.

Myotis oxyotus (Peters, 1866)

Taxonomy: Originally Vespertilio oxyotus Peters, 1866. We follow LaVal (1973) and Moratelli et al. (2019a) in recognizing two subspecies.

Myotis oxyotus oxyotus (Peters, 1866)

Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin 1867: 16–25.

Taxonomy: Originally Vespertilio oxyotus as described by Peters (1866: 19).

Neotype: LACM 36878, adult female collected by D. C. Carter on July 4, 1964; skull, mandible, and skin.

Type locality: Gruta Rumichaca, 2 mi E La Paz, Carchi, Ecuador (00°29'N, 77°50'W; ca. 2,600 m a.s.l.).

Remarks: The original description was based on an adult female preserved in spirit at “Zoologischen Cabinet zu München” (Peters 1866). Presumably this “Zoologischen Cabinet” is the same as the current Zoologische Sammlung des Bayerischen Staates (= Zoologische Staatssammlung München) in Munich, Germany. LaVal (1973) noted that D. C. Carter was unable to locate the holotype there in 1966, and he pointed out that many specimens in the museum in Munich were destroyed during World War II, and must be presumed lost. Therefore, LaVal (1973: 41) designated a neotype.

Myotis oxyotus gardneri LaVal, 1973

Bulletin of the Natural History Museum of Los Angeles County 15: 1–54.

Taxonomy: Described as Myotis oxyotus gardneri by LaVal (1973: 42).

Holotype: LSU 12924, adult male collected by A. L. Gardner on May 8, 1967; skull, mandible, baculum, and skin.

Type locality: Fila La Maquina, ca. 7.5 km E Canaan, San José, Costa Rica (09°27'N, 83°32'W; 2,610 m a.s.l.).

Myotis atacamensis (Lataste, 1892)

Actes de la Société Scientifique du Chili 1: 70–91.

Taxonomy: Described as Vespertilio atacamensis Lataste (1892: 79); currently monotypic (Moratelli et al. 2019a).

Neotype: USNM 391786, adult female collected by W. Mann and S. Mann in January 1944; skull partially damaged, mandible, and skin.

Type locality: Near Minimini, Tarapacá, Chile (19°10'S, 69°41'W; 1,800 m a.s.l.).

Remarks: The original description was based on three syntypes, probably collected in February 1885 in San Pedro de Atacama, Antofagasta, Chile, and deposited at Museo Nacional de Historia Natural (Santiago, Chile), including a mounted specimen (number 277), a skull (number 1007), and a fluid preserved specimen (number 276). LaVal (1973) argued these specimens are lost or, more probable, were destroyed. Novaes et al. (2022: 3) designated a neotype.

Myotis nesopolus Miller, 1900

Proceedings of the Biological Society of Washington 13: 123–127.

Taxonomy: Described as Myotis nesopolus by Miller (1900: 123). Treated as monotypic (Novaes et al. 2021a).

Holotype: USNM 101849, adult male collected by L. J. Guthrie on November 4, 1899; complete specimen preserved in alcohol.

Type locality: Near Willemstad, Curaçao, Netherlands Antilles (12°07'N, 68°52'W, ca. 35 m a.s.l.).

Myotis simus Thomas, 1901

Annals and Magazine of Natural History (ser. 7) 7: 189–193.

Taxonomy: Described as Myotis simus by Thomas (1901: 541). Currently monotypic (Moratelli et al. 2011b, 2019a).

Holotype: BMNH 8.5.12.2, adult female collected in 1876 by W. Davis; body preserved in alcohol (skin is faded and the dorsum and venter have blocks of hair losses) with skull and mandible removed, being partially damaged.

Type locality: Thomas (1901) indicates Sarayacu (06°44'S, 75°06'W; Carter and Dolan 1978), Peru, as type-locality. Later, LaVal (1973) added Rio Ucayali, Loreto, Peru, 100 m a.s.l.

Myotis dominicensis Miller, 1902

Proceedings of the Biological Society of Washington 15: 243–244.

Taxonomy: Described as Myotis dominicensis by Miller (1902: 243). Currently monotypic (Moratelli et al. 2019a).

Holotype: USNM 113564, adult male collected by H. S. Branch on July 18, 1901; body preserved in alcohol, skull and mandible removed.

Type locality: Island of Dominica.

Myotis dinellii Thomas, 1902

Annals and Magazine of Natural History (ser. 7) 10: 493–494.

Taxonomy: Described as Myotis dinellii by Thomas (1902: 493), treated as subspecies by LaVal (1973), and as species by Miranda et al. (2013) and Moratelli et al. (2019a). Monotypic (Moratelli et al. 2019a).

Holotype: BMNH 0.7.9.4, adult female collected by L. Dinelli on April 7, 1899; skull severely damaged, mandible missing, and skin.

Type locality: Tucumán, Argentina.

Myotis keaysi J.A. Allen, 1914

Bulletin of the American Museum of Natural History 33(29): 381–389.

Taxonomy: Described as Myotis ruber keaysi by Allen (1914: 383). Monotypic (Mantilla-Meluk and Muñoz-Garay 2014; Moratelli et al. 2019a).

Holotype: AMNH 15814, adult male collected by H. H. Keays on December 2, 1899; skull severely damaged and skin.

Type locality: Inca Mines, Puno, Peru (13°30'S, 70°00'W, 1,830 m a.s.l.).

Myotis caucensis J.A. Allen, 1914

Bulletin of the American Museum of Natural History 33(29): 381–389.

Taxonomy: Described as Myotis caucensis by Allen (1914: 386). Currently monotypic (Moratelli et al. 2013, 2019a).

Holotype: AMNH 32787, adult male collected by L. E. Miller on November 29, 1911; skull, mandible, and skin

Type locality: Rio Frío, Cauca River, Valle del Cauca, Colombia (04°09'N, 76°17'W; 1,066 m a.s.l.).

Myotis cobanensis Goodwin, 1955

American Museum Novitates 1744: 1–5.

Taxonomy: Described as Myotis velifer cobanensis by Goodwin (1955: 2), but considered as a full species by de la Torre (1958) and Hall (1981). Monotypic (Moratelli et al. 2019a).

Holotype: AMNH 145017, adult male collected by T. Larson on June 21, 1946; skull, mandible, and skin.

Type locality: Cathedral at Cobán, Alta Verapaz, Guatemala (15°28'S, 90°22'W; 1,305 m a.s.l.).

Myotis riparius Handley, 1960

Proceedings of the United States National Museum 112: 459–479.

Taxonomy: Described as Myotis simus riparius by Handley (1960: 466), raised to the species level by LaVal (1973). Treated as monotypic (Moratelli et al. 2019a).

Holotype: USNM 310255, an adult female (USNM 310255) with one embryo (7 mm crown-rump) collected by C.O. Handley and B.R. Feinstein on February 9, 1959; skull, mandible, and skin.

Type locality: Tacarcuna Village, Río Pucro, Darién, Panamá (07°51'N, 77°43'W, 945 m.a.s.l.).

Myotis elegans Hall, 1962

University of Kansas Publications, Museum of Natural History 14(13): 161–164.

Taxonomy: Described as Myotis elegans by Hall (1962: 163). Currently monotypic (Moratelli et al. 2019a).

Holotype: KU 88398, adult female collected by P.L. Clifton on September 24, 1961; skull severely damaged, mandible missing, and skin.

Type locality: 12.5 mi N of Tihuatlán, Veracruz, Mexico (20°41'N, 97°30'W; 90 m a.s.l.).

Myotis carteri LaVal, 1973

Bulletin of the Natural History Museum of Los Angeles County 15: 1–54.

Taxonomy: Described as Myotis nigricans carteri by LaVal (1973: 13), and elevated to the species level by Bogan (1978). Monotypic (LaVal 1973; Bogan 1978).

Holotype: LACM 36876, adult male collected by D. C. Carter on January 19, 1960; skull, mandible, baculum, and skin.

Type locality: 16 mi NE of Tamazula, Jalisco, Mexico (19°41'N, 103°14'W; 1,500 m a.s.l.).

Myotis larensis LaVal, 1973

Bulletin of the Natural History Museum of Los Angeles County 15: 1–54.

Taxonomy: Described as full species by LaVal (1973: 44), but posteriorly treated as subspecies of M. nesopolus by Genoways and Williams (1979). Novaes et al. (2021a) raised M. larensis to species level. Monotypic (LaVal 1973; Novaes et al. 2021a).

Holotype: AMNH 130709, adult female collected by G. H. H. Tate on March 23, 1938; skull, mandible, and skin.

Type locality: Rio Tocuyo, Lara, Venezuela (10°16'N, 69°56'W; 500 m a.s.l.).

Myotis martiniquensis LaVal, 1973

Bulletin of the Natural History Museum of Los Angeles County 15: 1–54.

Taxonomy: Described as Myotis martiniquensis by LaVal (1973: 35). Monotypic (Moratelli et al. 2019a).

Holotype: AMNH 214062, adult female collected by H. Beatty on March 15, 1967; body in alcohol, skull and mandible removed.

Type locality: Ca, 6 km E La Trinité, Tartane, Martinique (14°45'N, 60°54'W; ca. 65 m a.s.l.).

Myotis pilosatibialis LaVal, 1973

Bulletin of the Natural History Museum of Los Angeles County 15: 1–54.

Taxonomy: Described as Myotis keaysi pilosatibialis by LaVal (1973: 24), and raised to the species level by Mantilla-Meluk and Muñoz-Gray (2014). Treated as monotypic (Moratelli et al. 2019a).

Holotype: LACM 36879, adult male collected by R.K. LaVal and R. Valdez on July 26, 1969; skull, mandible, and skin partially damaged.

Type locality: 1 km W Talanga, Francisco Morazán, Honduras (14°24'N, 87°05'W; 750 m a.s.l.).

Myotis nyctor LaVal & Schwartz, 1974

Caribbean Journal of Science 14: 189–192.

Taxonomy: Described as Myotis nyctor by LaVal and Schwartz (1974: 190). Currently monotypic (Moratelli et al. 2019a).

Holotype: KU 109473, adult male collected by D. C. Leber and A. Schwartz on February 16, 1961; skull, mandible, and skin.

Type locality: Cole’s Cave, Saint Thomas Parish, Barbados (13°11'N, 59°34'W; 270 m).

Myotis findleyi Bogan, 1978

Journal of Mammalogy 59(3): 519–530.

Taxonomy: Described as Myotis findleyi by Bogan (1978: 524). Currently monotypic (Bogan 1978).

Holotype: USNM 512417, adult male collected by C. B. Robbins on March 14, 1976; skull, mandible, and skin.

Type locality: Isla Maria Magdalena, Islas Tres Marias, Nayarit, Mexico (21°27'N, 106°25'W; ca. 300 m).

Myotis diminutus Moratelli & Wilson, 2011

Mammalian Biology 76: 608–614.

Taxonomy: Described as Myotis diminutus by Moratelli and Wilson (2011a: 609). Monotypic (Moratelli and Wilson 2011a; Moratelli et al. 2019a).

Holotype: USNM 58569, sub-adult male collected by D. E. Wilson on February 11, 1979; skull, mandible, and skin.

Type locality: Río Palenque Science Center, 47 km S (by road) from Santo Domingo, Los Rios, Ecuador (00°35'S, 79°21'W; ca. 150 m).

Myotis izecksohni Moratelli, Peracchi, Dias & Oliveira, 2011

Mammalian Biology 76: 592–607.

Taxonomy: Described as Myotis izecksohni by Moratelli et al. (2011a: 597). Currently monotypic (Moratelli et al. 2011a, 2019a).

Holotype: ALP 6675, adult male collected by D. Dias on June 25, 2005; skull, mandible, complete post-cranial skeleton, and skin.

Type locality: Fazenda Maria Brandina, Tinguá Biological Reserve, Rio de Janeiro, Brazil, (22°36'S, 43°27'W; 760 m).

Myotis lavali Moratelli, Peracchi, Dias & Oliveira, 2011

Mammalian Biology 76: 592–607.

Taxonomy: Described as Myotis lavali by Moratelli et al. (2011a: 602). Currently monotypic (Moratelli et al. 2011a, 2019a).

Holotype: MZUSP 18762, adult male collected by M. R. Willig on April 3, 1977; skull, mandible, and skin.

Type locality: 6 km S of Exu, Pernambuco State, Brazil (07°30'S, 39°43'W; 523 m).

Myotis handleyi Moratelli, Gardner, Oliveira & Wilson, 2013

American Museum Novitates 3780: 1–36.

Taxonomy: Described as Myotis handleyi by Moratelli et al. (2013: 11) Currently monotypic (Moratelli et al. 2013, 2019a).

Holotype: USNM 370932, adult male collected by the Smithsonian Venezuela Project team on August 19, 1965; skull, mandible, and skin.

Type locality: Pico Ávila, 5 km northeast of Caracas, Distrito Federal, Venezuela (10°33'N, 66°52'W; 2,092 m).

Myotis midastactus Moratelli & Wilson, 2014

Journal of Mammalogy 95: E17–E25.

Taxonomy: Described as Myotis midastactus by Moratelli and Wilson (2014: E19). Currently monotypic (Moratelli and Wilson 2014; Moratelli et al. 2019a).

Holotype: AMNH 211156, adult male collected by S. Anderson on September 9, 1965; skull, mandible, complete post-cranial skeleton, and skin.

Type locality: Cercado, Río Mamoré, Beni, Bolívia, ca. 23 km W of San Javier (14°34'S, 64°55'W, 160 m).

Myotis clydejonesi Moratelli, Wilson, Gardner, Fisher & Gutiérrez, 2016

Special Publications, Museum of Texas Tech University 65: 49–66.

Taxonomy: Described as Myotis clydejonesi by Moratelli et al. (2016: 56). Currently monotypic (Moratelli et al. 2016, 2019a).

Holotype: TTU 109227, adult female collected by H. H. Genoways on January 23, 2008; skull, mandible, skin, and tissue (TK 151465).

Type locality: Raleigh Falls, Sipaliwini, Suriname (04°43'N, 56°12'W; 55 m).

Myotis attenboroughi Moratelli, Wilson, Novaes, Helgen & Gutiérrez, 2017

Journal of Mammalogy 98: 994–1008.

Taxonomy: Described as Myotis attenboroughi by Moratelli et al. (2017: 997). Currently monotypic (Moratelli et al. 2017, 2019a).

Holotype: USNM 540693, adult male collected on April 4, 1981 by G. S. Morgan, L. K. Gordon and F. A. Harrington; skull, mandible, and skin.

Type locality: Charlottesville, 1 km N of Pirate’s Bay, Saint John Parish, Tobago Island, Republic of Trinidad and Tobago (ca. 11°19'N, 60°33'W; sea level).

Myotis bakeri Moratelli, Novaes, Carrión & Wilson, 2019

Special Publications, Museum of Texas Tech University 71: 239–256.

Taxonomy: Described as Myotis bakeri by Moratelli et al. (2019b: 241). Currently monotypic (Moratelli et al. 2019b).

Holotype: MVZ 137909, adult male collected by M. L. Hawes on July 30, 1969; skull, mandible, and skin.

Type locality: 7 km SE of Chilca, Lima, Peru (12°33'S, 76°41'W; ca. 250 m).

Myotis armiensis Carrión-Bonilla & Cook, 2020

Therya 11: 508–532.

Taxonomy: Described as Myotis armiensis by Carrión-Bonilla and Cook (2020: 515). Currently monotypic (Carrión-Bonilla and Cook 2020).

Holotype: MSB 262089, adult male collected by J.A. Cook and collaborators on March 20, 2012; skull, mandible, complete post-cranial skeleton, and skin.

Type locality: Las Nubes Ranger Station, Parque Internacional La Amistad, District of Bugaba, Province of Chiriquí, Panamá (08°53'N, 82°36'W; 2,214 m).

Myotis pampa Novaes, Wilson & Moratelli, 2021

Vertebrate Zoology 71: 711–722.

Taxonomy: Described as Myotis pampa by Novaes et al. (2021b: 716), who considered it monotypic.

Holotype: AMNH 205471, adult female collected by M. D. Tuttle in January, 1963; skull, mandible, and skin.

Type locality: Ca. 6 km NW from Belén, Artigas, Uruguay (30°37'S, 57°50'W; 32 m elevation).

Myotis moratellii Novaes, Cláudio, Carrión, Abreu, Wilson, Maldonado & Weksler, 2021

Journal of Mammalogy 103: 1–20.

Taxonomy: Described as Myotis moratellii by Novaes et al. (2021c: 10), who considered it monotypic.

Holotype: USNM 513482, adult male collected by A. L. Gardner on July 22, 1976; skull, mandible, and skin, all well-preserved.

Type locality: Vinces Canton, 3 km NE of Puerto Nuevo, Los Ríos, Ecuador (01°15'S, 78°31'W; 15 m elevation).

Name-bearing type specimens of species in synonymy

There are at least 29 names currently in synonymy of recognized species (Table 2). Fourteen names are junior synonyms of M. nigricans, three are under M. albescens, three under M. chiloensis, two under M. ruber, one under M. atacamensis, one under M. oxyotus, and one under M. simus. Below is an annotated list of these names (in chronological order), with information about the primary specimen types, their preservation, and the type localities.

Table 2.

Names under synonymy of valid species of Neotropical Myotis, including information on their primary types.

Nomenclatural types Type specimen Synonymy Proximal type locality
argentatus Holotype KU 19228 M. albescens Veracruz, Mexico
isidori Holotype? MNHN 1997-1806 M. albescens Corrientes, Argentina
leucogaster Lectotype AMNH 385 M. albescens Bahia, Brazil
punensis Holotype AMNH 36263 M. albescens Guayas, Ecuador
nicholsoni Holotype FMNH 50783 M. atacamensis Arequipa, Peru
aelleni Holotype MHNG 1486.76 M. chiloensis Chubut, Argentina
arescens Holotype FMNH 24396 M. chiloensis Valparaiso, Chile
gayi Not located M. chiloensis Los Lagos, Chile
alter Holotype BMNH 0.6.29.23 M. levis Paraná, Brazil
nubilus Holotype? ZSM 121 M. levis Southern Brazil
polythrix Syntypes MNHN 842, 843 M. levis Rio Grande do Sul, Brazil
arsinoe Holotype RNH 17635 M. nigricans Suriname
bondae Holotype AMNH 14587 M. nigricans Santa Marta, Colombia
brasiliensis Not located M. nigricans Brazil
chiriquensis Holotype AMNH 18736 M. nigricans Chiriquí, Panama
concinnus Syntypes ANSP 1114, 1115 M. nigricans San Salvador, El Salvador
dalquesti Holotype KU 23839 M. nigricans Veracruz, Mexico
esmeraldae Holotype AMNH 33239 M. nigricans Esmeraldas, Ecuador
exiguus Holotype ANSP 5626 M. nigricans Panamá, Panama
hypothrix Holotype? MNHN 1903-41 M. nigricans Beni, Bolivia
maripensis Holotype AMNH 17069 M. nigricans Bolívar, Venezuela
mundus Holotype ANSP 1829 M. nigricans Zulia, Venezuela
parvulus Lectotype RNH 17621 M. nigricans Brazil
spixi Not located M. nigricans Brazil
splendidus Holotype? ZSM 142 M. nigricans US Virgin Islands
thomasi Not located M. oxyotus Napo, Ecuador
cinnamomeus Not located M. ruber Paraguay
kinnamon Holotype? MNHN 1997-2056 M. ruber Minas Gerais, Brazil
guaycuru Holotype ALP 9277 M. simus Mato Grosso do Sul, Brazil

Vespertilio leucogaster Schinz, 1821

In “Das tierreich eingetheilt nach dem Bau der thiere als Grundlage ihrer Naturgeschichte und der vergleichenden Anatomie von dem Herrn Ritter von Cuvier, volume 1”. Saugethiere und Vögel, Stuttgart and Tübingen, 894 pp.

Taxonomy: Described as Vespertilio leucogaster (currently allocated to Myotis) by Schinz (1821: 180). Currently a junior synonym of Myotis albescens (Miller and Allen 1928; LaVal 1973; Wilson 2008).

Lectotype: AMNH 385, adult (undetermined sex) collected by Maximilian, Prinz zu Wied-Neuwied (date not specified); taxidermized skin and skull not removed (see Avila-Pires 1965).

Type locality: Mucurí, Bahia, Brazil.

Vespertilio brasiliensis Spix, 1823

In “Simiarum et Vespertilionum brasiliensium species novae (…)”. Typis Francisci Serephici Hübschmanni, Monaco, xvi + 72 pp.

Taxonomy: Described as Vespertilio brasiliensis (currently allocated to Myotis) by Spix (1823: 63). Currently a junior synonym of Myotis nigricans (Miller and Allen 1928; Wilson 2008).

Type specimen: No specimen was designated by the author. Just like Carter and Dolan (1978), we have not found any reference specimens deposited in European collections.

Type locality: Brazil.

Remarks: The original name combination is preoccupied by Vespertilio brasiliensis Desmarest, 1822 (currently Eptesicus brasiliensis), hence, Fischer (1829) replaced it by Vespertilio spixii.

Vespertilio polythrix I. Geoffroy, 1824

Annales de Sciences Naturelles de Paris 3: 440–447.

Taxonomy: Described as Vespertilio polythrix (currently allocated to Myotis) by Geoffroy Saint-Hilaire (1824: 443). Currently a junior synonym of Myotis levis (LaVal 1973; Wilson 2008).

Syntypes: MNHN 842 (adult, undetermined sex), MNHN 843 (adult male), ZMB 3911 (adult, undetermined sex) collected by A. Geoffroy Saint-Hilaire, date not specified. All specimens are skins taxidermized (faded) with skull not removed.

Type locality: Rio Grande do Sul or Minas Gerais, Brazil.

Remarks: According to Turni and Kock (2008), the name polythrix is a nomen oblitum, due to page priority. This is the first available name (p. 443), whereas levis (nomen protectum) is on page 444 in Geoffroy’s publication (1824).

Vespertilio spixii Fischer, 1829

In “Synopsis mammalium”. Stuttgardtiae: J. G. Cottae, xlii + 752 pp.

Taxonomy: This name was proposed in replacement for Vespertilio brasiliensis Spix, 1823, considering that this name was preoccupied by Vespertilio brasiliensis Desmarest, 1822 (= Eptesicus brasiliensis). Currently a junior synonym of Myotis nigricans (Miller and Allen 1928; Cabrera 1958; Wilson 2008).

Vespertilio parvulus Temminck, 1840

In “Monographies de mammalogie ou description de quelques genres de mammifères dont les espèces ont été observées dans les différens musées de l’Europe”. E. d’Ocagne et A. Bertrand, Paris, 141–272.

Taxonomy: Described as Vespertilio parvulus (currently allocated to Myotis) by Temminck (1840: 246). Currently a junior synonym of Myotis nigricans (Miller and Allen 1928; LaVal 1973; Wilson 2008).

Lectotype: RNH 17621, adult, sex undetermined, collected by J. Natterer (date not specified); skull severely damaged and skin faded.

Type locality: Brazil.

Vespertilio arsinoe Temminck, 1840

In “Monographies de mammalogie ou description de quelques genres de mammifères dont les espèces ont été observées dans les différens musées de l’Europe”. E. d’Ocagne et A. Bertrand, Paris, 141–272.

Taxonomy: Described as Vespertilio arsinoe (currently allocated to Myotis) by Temminck (1840: 247). Currently a junior synonym of Myotis nigricans (LaVal 1973; Wilson 2008).

Holotype: RNH 17635, adult female (collector and date of capture are unknown); skull partially damaged and skin faded.

Type locality: Surinam.

Vespertilio hypothrix d’Orbigny & Gervais, 1847

In “Voyage dans l’Amérique méridionale (…). P. Bertrand/Strasbourg: V. Levrault, Paris 4: 1–32.

Taxonomy: Described as Vespertilio hypothrix (currently allocated to Myotis) by d’Orbigny and Gervais (1847: 14). Currently a junior synonym of Myotis nigricans (Miller and Allen 1928; Cabrera 1958; Wilson 2008).

Holotype: MNHN AC 1903-41, sex, age, collector, and date undetermined; stretched skin only.

Type locality: Moxos [Beni], Bolivia.

Vespertilio isidori d’Orbigny & Gervais, 1847

In “Voyage dans l’Amérique méridionale (…)”. P. Bertrand/Strasbourg: V. Levrault, Paris 4: 1–32.

Taxonomy: Described as Vespertilio isidori (currently allocated to Myotis) by d’Orbigny and Gervais (1847: 16). Currently a junior synonym of Myotis albescens (Miller and Allen 1928). However, based on observations made by Carter and Dolan (1978), Wilson (2008) did not include this name in the synonym list for M. albescens (see discussion in Remarks section)

Holotype: Probably MNHN 1997-1806, adult, sex undetermined; skull (damaged), mandible, and skin.

Type locality: Corrientes, Argentina.

Remarks: Rode (1941) indicated the specimen MNHN 865 as the holotype. However, Carter and Dolan (1978) show that there was confusion when interpreting a Cadre number, used to guide visitors about a specimen on display in the museum, with the catalog number. Thus, Carter and Dolan indicate that this is not the type specimen of this name and have not found any other specimen in collections in Europe that could be. One of us (RM) found the supposed specimen used for the description by d’Orbigny and Gervais deposited in the mammal collection of the Muséum National D’Histoire Naturelle, Paris, France. The presumable holotype (MNHN 1997-1806) is an adult (sex undetermined). The skull reassembles M. albescens, but the color pattern of the skin is not a Neotropical Myotis.

Vespertilio splendidus Wagner, 1855

In “Die säugthiere in abbildungen nach der natur mit beschreibungen von Dr. Johann Christian Daniel von Schreber (…)”. T.O. Weigel, Leipzig, xxvi + 810 pp.

Taxonomy: Described as Vespertilio splendidus (currently allocated to Myotis) by Wagner (1855: 148). Currently a junior synonym of Myotis nigricans (see Carter and Dolan 1978; Wilson 2008).

Holotype: ZSM 142, adult of undetermined sex, probably collected by A. F. W. Schimper (date not specified); skin only, slightly faded.

Type locality: St. Thomas [American Virgin Islands (Carter and Dolan 1978)].

Vespertilio nubilus Wagner, 1855

In “Die säugthiere in abbildungen nach der natur mit beschreibungen von Dr. Johann Christian Daniel von Schreber (…)”. T. O. Weigel, Leipzig, xxvi + 810 pp.

Taxonomy: Described as Vespertilio nubilus (currently allocated to Myotis) by Wagner (1855: 752). Currently a junior synonym of Myotis levis (see LaVal 1973; Wilson 2008).

Holotype: ZSM 121, subadult, sex undetermined; collector and date of capture are unknown; skin taxidermized with skull not removed.

Type locality: Brazil.

Vespertilio cinnamomeus Wagner, 1855

In “Die säugthiere in abbildungen nach der natur mit beschreibungen von Dr. Johann Christian Daniel von Schreber (…)”. T. O. Weigel, Leipzig, xxvi + 810 pp.

Taxonomy: Wagner (1855: 755) proposed the name Vespertilio cinnamomeus as a substitute for Vespertilio ruber É. Geoffroy, 1806 believing that “chauve-sourris cannelle” from Azara (1801) was a Noctilio Linnaeus, 1766. However, Miller and Allen (1928) resolved Wagner’s misunderstanding, indicating that both the name ruber and cinnamomeus were based on the same specimen. Currently a junior synonym of Myotis ruber (see Miller and Allen 1928; LaVal 1973; Wilson 2008).

Vespertilio kinnamon Gervais, 1856

In “Animaux nouveaux ou rares recueillis pendant l’expédition dans les parties centrales de l’Amérique du Sud (…)”. P. Bertrand, Paris, 25–88.

Taxonomy: Described as Vespertilio kinnamon (currently allocated to Myotis) by Gervais (1856: 84). Currently a junior synonym of Myotis ruber (Miller and Allen 1928; Cabrera 1958).

Holotype: MNHN 1997-2056, adult male collected on 1844 (collector not specified); skin only.

Type locality: Capela Nova, Minas Gerais, Brazil.

Vespertilio mundus H. Allen, 1866

Proceedings of the Academy of Natural Sciences of Philadelphia 18: 279–288.

Taxonomy: Described as Vespertilio mundus (currently allocated to Myotis) by Allen (1866: 280). Currently a junior synonym of Myotis nigricans (Miller and Allen 1928; LaVal 1973; Wilson 2008).

Holotype: ANSP 1829 (=USNM 5547), subadult female collected by S. Hayes (date not specified), currently deposited in the mammal collection of the Academy of Natural Sciences of Drexel University (Philadelphia, USA); complete specimen preserved in alcohol with skin faded.

Type locality: Maracaibo, Venezuela.

Vespertilio concinnus H. Allen, 1866

Proceedings of the Academy of Natural Sciences of Philadelphia 18: 279–288.

Taxonomy: Described as Vespertilio concinnus (currently allocated to Myotis) by Allen (1866: 281). Currently a junior synonym of Myotis nigricans (Miller and Allen 1928; Cabrera 1958; LaVal 1973; Wilson 2008).

Syntypes: ANSP 1114 and ANSP 1115, are adult females, collected by J. Leidy (date not specified); Body preserved in alcohol with skin faded, skull and mandible removed.

Type locality: San Salvador, El Salvador.

Vespertilio exiguus H. Allen, 1866

Proceedings of the Academy of Natural Sciences of Philadelphia 18: 279–288.

Taxonomy: Described as Vespertilio exiguus (currently allocated to Myotis) by Allen (1866: 281). Currently a junior synonym of Myotis nigricans (Miller and Allen 1928; Cabrera 1958).

Holotype: ANSP 5626 (= USNM 5373), adult female collected by S. Hayes (date not specified) is currently deposited in the mammal collection of the Academy of Natural Sciences of Drexel University (Philadelphia, USA); complete specimen preserved in alcohol with skin faded.

Type locality: Aspinwall, NG. (= Colón, Panama).

Vespertilio gayi Lataste, 1892

Actes de la Société Scientifique du Chili 1: 70–91.

Taxonomy: Described as Vespertilio gayi (currently allocated to Myotis) by Lataste (1892: 79), currently considered a junior synonym of Myotis chiloensis (Miller and Allen 1928; Cabrera 1958; Wilson 2008).

Type specimen: None. We did not access the original publication describing the species. However, no specimens from Lataste that could match the description of M. gayi are available in collections in Europe or South America. Probably, these specimens are lost.

Type locality: Valdivia, Chile.

Myotis thomasi Cabrera, 1901

Boletín de la Sociedad Española de Historia Natural 1: 367–373.

Taxonomy: Described as Myotis thomasi by Cabrera (1901: 370). Currently a junior synonym of Myotis oxyotus (Miller and Allen 1928; LaVal 1973; Wilson 2008).

Type specimen: None. The original description was based on an adult female preserved in alcohol that, according to the author, was deposited in the mammal collection of the Museo Nacional de Ciencias Naturales (Madrid, Spain). However, a voucher number for the specimen was not listed by Cabrera (1901). Nevertheless, Carter and Dolan (1978) did not find the representative specimen in the MNCN collection and suspect that when moving to Argentina, A. Cabrera would have taken the type specimens. There are no specimens of Myotis that can represent the type of M. thomasi in the Museo de La Plata (Itatí Olivares, pers. comm.). We presume that probably this type specimen is lost.

Type locality: In the original description, Cabrera (1901) argued that he did not have reliable data on the geographical origin of the specimen, but that it was probably from southern Brazil. Later, Cabrera (1902) corrected this to “Archidona [sobre el citado río], Napo, Ecuador”.

Myotis chiriquensis J.A. Allen, 1904

Bulletin of the American Museum of Natural History 20: 29–80.

Taxonomy: Described as Myotis chiriquensis by Allen (1904: 77). Currently a junior synonym of Myotis nigricans (Miller and Allen 1928; Cabrera 1958; LaVal 1973; Wilson 2008).

Holotype: AMNH 18736, adult female, collected by J. H. Batty on October 16, 1901; skull, mandible, and skin.

Type locality: Boquerón, Chiriquí, Panama.

Myotis punensis J.A. Allen, 1914

Bulletin of the American Museum of Natural History 33(29): 381–389.

Taxonomy: Described as Myotis punensis by Allen (1914: 383). Currently a junior synonym of Myotis albescens (see Moratelli and Wilson 2011b).

Holotype: AMNH 36263, sub-adult male collected by W.B. Richardson on May 8, 1913; skull (partially damaged), mandible, and skin.

Type locality: Isla Puna, Guayaquil, Guayas, Ecuador.

Myotis bondae J.A. Allen, 1914

Bulletin of the American Museum of Natural History 33(29): 381–389.

Taxonomy: Described as Myotis bondae by Allen (1914: 384). Currently a junior synonym of Myotis nigricans (Miller and Allen 1928; Cabrera 1958; LaVal 1973; Wilson 2008).

Holotype: AMNH 14587, adult of undetermined sex, collected by H. H. Smith in June 1898; skull, mandible, and skin.

Type locality: Bonda, Santa Marta, Colombia.

Myotis maripensis J.A. Allen, 1914

Bulletin of the American Museum of Natural History 33(29): 381–389.

Taxonomy: Described as Myotis maripensis by Allen (1914: 385). Currently a junior synonym of Myotis nigricans (Miller and Allen 1928; Cabrera 1958; LaVal 1973; Wilson 2008).

Holotype: AMNH 17069, adult female collected by S. M. Klages on December 13, 1909; skull, mandible, and skin.

Type locality: Maripa, Venezuela.

Myotis esmeraldae J.A. Allen, 1914

Bulletin of the American Museum of Natural History 33(29): 381–389.

Taxonomy: Described as Myotis esmeraldae by Allen (1914: 385). Currently a junior synonym of Myotis nigricans (Miller and Allen 1928; Cabrera 1958; Wilson 2008).

Holotype: AMNH 33239, adult male, collected by W. B. Richardson on November 5, 1912; skull, mandible, and skin.

Type locality: Esmeraldas, Ecuador.

Myotis chiloensis alter Miller & Allen, 1928

Bulletin of the United States National Museum 144: 1–218.

Taxonomy: Described as a subspecies of Myotis chiloensis by Miller and Allen (1928). Currently a junior synonym of Myotis levis (LaVal 1973; Wilson 2008).

Holotype: BMNH 0.6.29.23, adult female collected by G. Grillo (date not specified); body in alcohol, skull and mandible removed.

Type locality: Palmeira, Paraná, Brazil.

Myotis nigricans nicholsoni Sanborn, 1941

Field Museum of Natural History, Zoological Series 27: 371–387.

Taxonomy: Described as a subspecies of Myotis nigricans by Sanborn (1941: 382). Currently a junior synonym of Myotis atacamensis (LaVal 1973; Wilson 2008).

Holotype: FMNH 50783, adult male collected by C. C. Sanborn on October 17, 1939; skull, mandible, and skin, all well-preserved.

Type locality: Hacienda Chucarapi, Tambo Valley, Arequipa, Peru.

Myotis chiloensis arescens Osgood, 1943

Field Museum of Natural History, Zoological Series 30: 1–268.

Taxonomy: Described as a subspecies (Osgood 1943: 55), but currently considered a junior synonym of Myotis chiloensis (LaVal 1973; Wilson 2008).

Holotype: FMNH 24396, adult male collected by J. A. Wolffsohn on January 1, 1925; skin only.

Type locality: Hacienda Limache, Valparaíso, Chile.

Myotis guaycuru Proença, 1943

Revista Brasileira de Biologia 3: 313–315.

Taxonomy: Described as Myotis guaycuru by Proença (1943: 314), but currently considered a junior synonym of Myotis simus (Wilson 2008; Moratelli et al. 2011b).

Holotype: ALP 9277, an adult female collected in 1940 by Scientific Committee of the Oswaldo Cruz Institute, headed by L. Travassos; body preserved in fluid (severely damaged), with the skull (including mandible) removed and complete.

Type locality: Rio Miranda, Salobra, Mato Grosso do Sul, Brazil.

Myotis argentatus Dalquest & Hall, 1947

University of Kansas Publications, Museum of Natural History 1(12): 237–244.

Taxonomy: Described as a full species by Dalquest and Hall (1947: 239). Currently a junior synonym of Myotis albescens (see LaVal 1973).

Holotype: KU 19228, adult male collected by W. W. Dalquest on February 2, 1947; skull, mandible, and skin.

Type locality: 14 km SW of Coatzocoalcos, Veracruz, Mexico (30 m elevation).

Myotis nigricans dalquesti Hall & Alvarez, 1961

University of Kansas Publications, Museum of Natural History 14(4): 69–72.

Taxonomy: Described as a subspecies of Myotis nigricans by Hall and Alvarez (1961: 71), but currently considered a junior synonym of Myotis nigricans (LaVal 1973).

Holotype: KU 23839, adult male collected by W. W. Dalquest on January 5, 1948; skull, mandible, and skin.

Type locality: 3 km E of San Andrés Tuxtla, Veracruz, Mexico (304 m elevation).

Myotis aelleni Baud, 1979

Revue Suisse de Zoologie 86(1): 267–278.

Taxonomy: Described as a full species by Baud (1979: 268), but currently considered a junior synonym of Myotis chiloensis (Novaes et al. 2018).

Holotype: MHNG 1486.76, adult male collected by A. Kovacs on December 19, 1975; body preserved in alcohol, skull and mandible removed.

Type locality: El Hoyo de Epuyen, 42°10'S, 71°21'W (230 m elevation), Provincia de Chubut, Argentina.

Discussion

Myotis is the most speciose bat genus in the Neotropics, with 33 species recognized currently (Bogan 1978; Moratelli et al. 2019a, b; Novaes et al. 2021a, b, c). Several species’ descriptions and revalidations have been recently proposed (e.g., LaVal 1973; Moratelli et al. 2011a, 2013, 2016, 2017, 2019b; Novaes et al. 2018, 2021a, b, c), and the evidence available points in the direction of hidden diversity (Clare et al. 2007; Larsen et al. 2012; Novaes et al. 2018; Carrión-Bonilla and Cook 2020). In this scenario of intense taxonomic change, a careful assessment of all name-bearing types is essential to the correct application of names to newly identified lineages and other nomenclatural acts. Below, we point out some nomenclatural issues still associated with name-bearing types of Neotropical Myotis.

Since its description, Myotis nigricans has been treated as a widely distributed species, and several subspecies have been recognized by different authors. However, recent studies have merged evidence indicating that M. nigricans is composite, as currently recognized, representing a complex of allopatric species (Moratelli and Wilson 2011a; Moratelli et al. 2011a, 2016, 2017, 2019b; Novaes et al. 2021b). The name nigricans seems to apply to Atlantic Forest populations from southeastern Brazil and southern South America, considering the type locality (Moratelli and Wilson 2011a; Moratelli et al. 2013, 2017). Therefore, it is necessary to reassess the taxonomic status of populations from tropical Mexico, Central America, and northern South America currently recognized as M. nigricans (and its subspecies). In this case, names currently treated under synonymy might apply to these potential new taxa.

Currently, 14 names are under synonymy of M. nigricans and can be available to use after a careful taxonomic review that considers the examination of type specimens. An example is the name Vespertilio splendidus Wagner, 1855 (= Myotis splendidus), described based on a specimen from “St. Thomas” (Wagner 1855). Carter and Dolan (1978) indicated the type locality as “St. Thomas [American Virgin Islands]”, which was followed by subsequent authors (e.g., Wilson 2008). However, Myotis apparently does not occur either on the US Virgin Islands (Bacle et al. 2008) or on the nearest Caribbean islands (Puerto Rico, British Virgin Islands, Anguilla; Timm and Genoways 2003; Genoways et al. 2007). On the other hand, “St. Thomas” is a locality on the Caribbean Island of Barbados, where M. nyctor is the only species known to occur (Novaes et al. 2021a). Based on this scenario, Myotis splendidus is a very rare (or extinct) species (and unique representative of the genus) on the US Virgin Islands; or the geographical origin of the holotype of Myotis splendidus is Barbados, not US Virgin Islands, and the name is the senior synonym of Myotis nyctor. In any case, Myotis splendidus is unlikely to be a synonym for Myotis nigricans, considering the biogeographical history of colonization of the Caribbean, where each island has its own unique species of Myotis, and there is no evidence of the occurrence of M. nigricans as recognized by Moratelli et al. (2017) and Novaes et al. (2021a).

Another important issue is the validity of some names occasionally found in the literature on Myotis taxonomy. In their catalogue of type specimens of neotropical bats deposited in selected European museums, Carter and Dolan (1978) listed “Vespertilio carbonarius Wagner” based on a specimen (ZSM 124) from Brazil obtained by J.F. Brandt, whose label reads “Vespertilio carbonarius Wagn. / 1843 / Brandt / Brasil”. This specimen (taxidermized skin with skull not removed) was examined by us and it resembles M. riparius in size and the fur texture, length, and coloration. However, as with Carter and Dolan (op cit.), we were also unable to locate the publication with the species description. It is not impossible that this name was formally published (considering the vast, and sometimes rare, production of Wagner). However, if a publication containing the species description is found, we suggest that Vespertilio carbonarius should be treated as a nomen oblitum, following article 23.9.1 from ICZN (1999).

For another example, Cornalia (1849) assigned the name “Vespertilio quixensis Osculati” to the synonymy of Vespertilio osculati (= Myotis nigricans osculati). In an introduction to the facsimile reprint of Cornalia’s (1849) publication, Cagnolaro and Violani (1988) recommended treating V. quixensis as a nomen nudum, but the name became available in the combination Phyllostomus quixensis Osculati, 1854: 53. It is possible that the description of quixensis appeared in the first edition of the Osculati’s publication; however, we have not been able to examine that publication due to its rarity.

Myotis comprises a diverse group in number of species compared to other neotropical bat genera. However, its species richness does not reflect its phenotypic diversity, characterized by a low morphological differentiation (Ghazali et al. 2017; Moratelli et al. 2019a). Due to the large number of species names proposed, this catalogue puts together information on name-bearing types of species treated as valid or under synonymy as an aid for future taxonomic works.

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); 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, USA); B. Patterson, W. Stanley and R. Banasiak (Field Museum of Natural History, USA); J.A. Esselstyn (Museum of Natural Sciences at Louisiana State University, USA); R. Baker and H. Garner (Museum of Texas Tech University, USA); C. Conroy (Museum of Vertebrate Zoology at University of California, USA); R.M. Timm and M. Eifler (Natural History Museum of the Kansas University, USA); J. Dines (Natural History Museum of Los Angeles County, USA); K. Helgen, D. Lunde, and L. Gordon (National Museum of Natural History, USA); R.P. Miguez (Natural History Museum, London, UK); M. Hiermeier (Zoologisches Staats Sammlung München, Germany); C. Callou, C. Denys, J.-M. Pons (Muséum National D’Histoire Naturelle, France). G.S.T. Garbino (Universidade Federal de Viçosa, Brazil) provided access to information about Ángel Cabrera’s publications. N.S. Gilmore (Museum of the Academy of Natural Sciences of Drexel University, USA) and A.L.G. Rodríguez (Museo Nacional de Ciencias Naturales, Spain), Itatí Olivares provided access to information about specimens deposited in Museo de La Plata, (Buenos Aires, Argentina). G. Bardelli (Sezione di Zoologia dei Vertebrati, Museo di Storia Naturale di Milano, Italy) provided information about Gaetano Osculati’s collection. This work was partially supported by the Smithsonian Institution, USA, Conselho Nacional de Desenvolvimento Científico e Tecnológico, Brazil (CNPq 313963/2018-5) and Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro, Brazil (FAPERJ; E-26/200.967/2021) through grants to RM and DEW. RLMN has received support from FAPERJ (E-26/204.243/2021).

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