Updated checklist of bats (Mammalia: Chiroptera) from Brazil

ABSTRACT

We present an updated checklist of Brazilian bats, commenting on the endemic and threatened status of the species listed and providing information on recent taxonomic and nomenclatural changes. The bats of Brazil comprise 186 species, 68 genera, and nine families, with 13 species exclusive to the country. From the previous checklists, we add eight species to Brazil: Artibeus amplus, Choeroniscus godmani, Glossophaga bakeri, Lichonycteris obscura, Platyrrhinus guianensis, Trachops ehrhardti, Molossus melini and Myotis pampa. The latter is reported for the country for the first time in this study. The Brazilian list of threatened species includes three species as vulnerable (Furipterus horrens, Lonchophylla bokermanni, and Natalus macrourus) and Lonchophylla dekeyseri as endangered. The International Union for the Conservation of Nature (IUCN) lists Natalus macrourus as “near threatened”, and Lonchophylla bokermanni and L. dekeyseri as “endangered”. Twenty-two additional species are probable for the country. Compared to previous lists, five species are now considered doubtful records and 19 are considered erroneous records. We reinforce the importance of a continuous update of the bat checklists in all Brazilian states as well as taxonomic revisions and bat inventories in unsampled areas.

KEY WORDS:
Neotropics; nomenclature; species list; taxonomy

INTRODUCTION

Lists of species are essential for the scientific community and general users, forming a base on the richness and diversity of studies and projects in conservation, local livelihoods, zoonoses surveillance, and scientific research (Garnett et al. 2020). With that in mind, there has been a recent push to create a single, authoritative list of the Earth’s biota, which has been met with a series of publications both for and against the proposal (Raposo et al. 2017, Thomson et al. 2018). Irrespective of the merits associated with the global list proposal, a specific set of 10 principles has been recommended to form the foundation of its governance (Garnett et al. 2020). Among these principles, it is important to underscore the imperative for transparent decision-making, unrestricted taxonomic autonomy, and the active pursuit of community support and utilization.

Brazil is one of the megadiverse countries in the world and ranks fourth amongst South American countries with the largest number of bat species, behind Colombia, Ecuador, and Peru (Díaz et al. 2021, Pacheco et al. 2021, Ramírez-Chaves et al. 2022). The first effort to list the bat species occurring in Brazil was made by August von Pelzeln (von Pelzeln 1883), who described the mammals collected by Johann Natterer during the Austrian Expedition to Brazil from 1817 to 1835 and listed 48 species of bats. In the following years, there have been several subsequent efforts to list the bat richness that occurs in Brazil, especially the catalogs that include bats for the entire country (Vieira 1942, 1955), and more frequently after the 1990s (for historical reviews see Vieira 1942, Nogueira et al. 2014).

In 2011, the Brazilian Bat Research Society (Sociedade Brasileira para o Estudo dos Quirópteros - SBEQ) created the Committee of the List of Brazilian Bats (Comitê da Lista de Morcegos do Brasil - CLMB). Since then, the CLMB has been in charge of compiling and maintaining a list of Brazilian bat species, with the initial lists comprising 178 species (Nogueira et al. 2014), 182 species (Nogueira et al. 2018), and 181 species (Garbino et al. 2020a). From the initial workgroup of six, the CLMB now has 13 members as of 2024, all of whom study taxonomic aspects of bat species occurring in Brazil. One goal of the CLMB bat checklist is to create a list that both the scientific community and other important users, such as government stakeholders in charge of threatened species legislation, can use.

Simultaneously with the publication of the CLMB 2020 list, two checklists of Brazilian mammals, including bats, were published by the Brazilian Society of Mammalogy (Sociedade Brasileira de Mastozoologia - SBMz; Abreu et al. 2021) and independent researchers (Quintela et al. 2020). Added to these three recent lists of Brazilian bats, there is an ongoing project to produce a list of the country’s animal species that has united more than 800 zoologists to produce an online catalog (Brazilian Zoology Group (2024). This effort, named Taxonomic Catalog of the Brazilian Fauna (Catálogo Taxonômico da Fauna do Brasil - CTFB) is constantly being updated by experts in the field, and includes members of both SBEQ and SBMz committees.

Considering that four checklists of Brazilian bats have been published in the last five years, the goals of this paper are to (1) produce an updated checklist of Brazilian bats to serve as a taxonomic and nomenclatural reference to be used in further research and by stakeholders, such as governmental agencies; and (2) compare the similarities and differences between the new list and the three previous lists.

MATERIAL AND METHODS

To update the checklist of bats native to Brazil, we have considered the latest taxonomic and nomenclatural studies available in the literature (e.g., Velazco and Patterson 2019, Loureiro et al. 2020, Basantes et al. 2020, Fonseca et al. 2024). We have also examined articles that report on range extensions for Brazil (e.g., Velazco et al. 2017, Garbino et al. 2022, Zortéa et al. 2023). Family, genus, and species-level taxonomy follow Wilson and Mittermeier (2019) except where noted.

We considered only formal records of bats for the checklist, which include records from peer-reviewed publi cations (e.g., articles, books, and short communications) that mention at least one voucher deposited in a zoological collection (Nogueira et al. 2014). Also following Nogueira et al. (2014), three lists are presented herein: a list of the formal records of bats from Brazil, a list of doubtful records, and a list of erroneous records. A formal record was considered doubtful if it was not supported by a voucher, if the taxon had an uncertain taxonomic status, or if the occurrence of the taxon in Brazil was uncertain after a taxonomic revision. An erroneous record was considered as such if the evidence supporting its occurrence in Brazil was refuted or if a taxonomic revision redefined the taxon concept. For example, Pavan and Marroig (2016) restricted Pteronotus parnellii to Cuba and Jamaica, and this taxon was considered an erroneous record for Brazil.

We also produced a fourth list, including species that could potentially occur in Brazil. For this, we considered a buffer area of 200 km around the boundaries of the Brazilian territory and verified extralimital records in the literature that fall within this area. The criteria to consider these records as valid followed the same for the formal records. To establish the distribution of marginal taxa, we used the compilations of Rojas et al. (2018), Marsh et al. (2022), and the IUCN shapefile data (https://www.iucnredlist.org/resources/spatial-data-download). After identifying the potential species, we removed taxa that could not occur in Brazil due to geographical barriers (e.g., montane species that fell in the 200 km buffer zone) and also disregarded erroneous records that are present in older shapefiles (e.g., the IUCN distribution polygon of Molossus sinaloae). The used distribution polygons overlaid with the 200 km buffer zone can be found in ^{Supplementary Material S1}.

RESULTS

Species diversity and endemismTable 1 Table 2 Table 3 Table 1 Table 3 ^{Supplementary Material S1}

The checklist of bats from Brazil includes 186 species, 68 genera, and nine families (^{Appendix 1}, Fig. 1). Consider ing that the order Chiroptera now contains 1,466 extant species, Brazil harbors ca. 13% of the world diversity of bats (MDD 2023). Thirteen species are endemic to Brazil: Dryadonycteris capixaba, Glyphonycteris behnii, Histiotus alienus, Lasiurus ebenus, Lonchophylla bokermanni, L. inexpectata, L. mordax, L. peracchii, Neoeptesicus taddeii, Neonycteris pusilla, Platyrrhinus recifinus, Trachops ehrhardti, and Xeronycteris vieirai (^{Appendix 1}). The most diverse group is the family Phyllostomidae (96 species), followed by Molossidae (34 species), Vespertilionidae (27 species), Emballonuridae (17 species), Thyropteridae (five species), Mormoopidae (four species), Noctilionidae (two species), Natalidae (one species), and Furipteridae (one species).

Figure 1
Number of bat species recorded for Brazil according to different checklists produced between 1995 and 2023. We fit a loess regression in the scatterplot. 1: Aguiar and Taddei (1995), 2: Fonseca et al. (1996), 3: Taddei (1996), 4: Marinho-Filho and Sazima (1998), 5: Tavares et al. (2008- list produced in 2005), 6: Peracchi et al. (2006), 7: Reis et al. (2007), 8: Peracchi et al. (2010), 9: Peracchi et al. (2011), 10: Paglia et al. (2012), 11: Nogueira et al. (2014), 12: Nogueira et al. (2018), 13: Garbino et al. (2020a), 14: Quintela et al. (2020), 15: Abreu et al. (2022), 16: Abreu et al. (2023),17: This study.

Table 1
List of bat species with doubtful records for Brazil. In the Reference column, we cite the source for the doubtful records.

Family	Taxon	Reference
Phyllostomidae	Artibeus glaucus Thomas, 1893	Koopman (1993)
Phyllostomidae	Artibeus phaeotis (Miller, 1902)	Taddei (1996)
Phyllostomidae	Carollia castanea H. Allen, 1890	Uieda (1980)
Phyllostomidae	Lonchorhina orinocensis Linares & Ojasti, 1971	Taddei (1996)
Phyllostomidae	Lonchorhina marinkellei Hernández-Camacho & Cadena, 1978	Taddei (1996)

Table 2
List of erroneous records of bat species in Brazil, considering Nogueira et al. (2014) as a starting point.

Family	Taxon	Reason for removal	Reference
Phyllostomidae	Carollia subrufa (Hahn, 1905)	Specimens reidentified as Carollia brevicauda	Nogueira et al. (2014)
Phyllostomidae	Chiroderma salvini Dobson, 1878	Specimens reidentified as Chiroderma villosum	Garbino et al. (2020b)
Phyllostomidae	Chiroderma vizottoi Taddei & Lim, 2010	Considered valid as subspecies of C. doriae	Garbino et al. (2020b)
Phyllostomidae	Enchisthenes hartii (Thomas, 1892)	Erroneous identification	Nogueira et al. (2014)
Phyllostomidae	Lichonycteris degener Miller, 1931	Junior synonym of Lichonycteris obscura Thomas, 1895	Zamora-Gutierrez and Ortega (2020)
Phyllostomidae	Micronycteris brosseti Simmons & Voss, 1998	Specimen reidentified as Micronycteris sp.	Garbino (2016)
Phyllostomidae	Micronycteris homezorum Pirlot, 1967	Junior synonym of Micronycteris minuta (P. Gervais, 1856)	Ochoa and Sánchez (2005)
Phyllostomidae	Platyrrhinus helleri (Peters, 1866)	Specimens reidentified as Platyrrhinus incarum	Garbino et al. (2024)
Phyllostomidae	Sturnira bidens (Thomas, 1915)	Specimen reidentified as Stunira lilium	Nogueira et al. (2014)
Phyllostomidae	Tonatia saurophila Koopman & Williams, 1951	Restricted to a fossil from Jamaica	Basantes et al. (2020)
Mormoopidae	Pteronotus davyi Gray, 1838	Specimen reidentified as Pteronotus gymnonotus	Nogueira et al. (2014)
Mormoopidae	Pteronotus parnellii (Gray, 1843)	Restricted to the Antilles	Pavan and Marroig (2016)
Vespertilionidae	Neoeptesicus andinus J.A. Allen, 1914	Specimen reidentified as Neoeptesicus sp.	This study
Vespertilionidae	Eptesicus fuscus (Palisot de Beauvois, 1796)	Specimens reidentified as Neoeptesicus furinalis	Nogueira et al. (2014)
Vespertilionidae	Lasiurus atratus Handley, 1996	Specimens reidentified as Lasiurus castaneus	Nogueira et al. (2014)
Vespertilionidae	Lasiurus cinereus (Palisot de Beauvois, 1796)	Restricted to North America	Baird et al. (2015)
Vespertilionidae	Lasiurus salinae Thomas, 1902	Considered valid as subspecies of L. blossevillii	Baird et al. (2015)
Vespertilionidae	Myotis alter Miller & G.M. Allen, 1928	Junior synonym of Myotis levis (I. Geoffroy, 1824)	Wilson (2008)
Vespertilionidae	Myotis dinellii Thomas, 1902	Specimens reidentified as Myotis albescens	Moratelli et al. (2019)

Table 3
List of bat species potentially occurring in Brazil. See ^{Supplementary Material S1} for their distribution maps. In the Reference column, we cite the source for the closest formal records to the Brazilian border.

Family	Taxon	Reference
Emballonuridae	Centronycteris centralis Thomas, 1912	Hice and Solari (2002)
Phyllostomidae	Artibeus glaucus Thomas, 1893	Handley (1987)
Phyllostomidae	Artibeus phaeotis (Miller, 1902)	Handley (1987)
Phyllostomidae	Carollia castanea H. Allen, 1890	Shapley et al. (2005)
Phyllostomidae	Enchisthenes hartii (Thomas, 1892)	Solari et al. (2006)
Phyllostomidae	Hsunycteris dashe Velazco et al., 2017	Velazco et al. (2017)
Phyllostomidae	Lonchorhina fernandezi Ochoa & Ibáñez, 1984	Ochoa and Ibáñez (1984)
Phyllostomidae	Lonchorhina orinocensis Linares & Ojasti, 1971	Hernández-Camacho and Cadena (1978)
Phyllostomidae	Lonchorhina marinkellei Hernández-Camacho & Cadena, 1978	Hernández-Camacho and Cadena (1978)
Phyllostomidae	Micronycteris brosseti Simmons & Voss, 1998	Simmons and Voss (1998)
Phyllostomidae	Micronycteris matses Simmons, Voss & Fleck, 2002	Simmons et al. (2002)
Phyllostomidae	Micronycteris yatesi Siles & Brooks, 2013	Siles et al. (2013)
Molossidae	Eumops nanus (Miller, 1900)	Eger (1977)
Molossidae	Molossus alvarezi González-Ruiz, Ramírez-Pulido & Arroyo-Cabrales, 2011	Loureiro et al. (2019)
Molossidae	Molossus fentoni Loureiro, Lim & Engstrom, 2018	Loureiro et al. (2018)
Molossidae	Nyctinomops mbopicuare Barquez et al., 2023	Barquez et al. (2023)
Vespertilionidae	Histiotus humboldti Handley, 1996	Handley (1996)
Vespertilionidae	Lasiurus atratus Handley, 1996	Handley (1996)
Vespertilionidae	Myotis clydejonesi Moratelli et al., 2016	Moratelli et al. (2016)
Vespertilionidae	Myotis midastactus Moratelli & Wilson, 2014	Moratelli and Wilson (2014)
Vespertilionidae	Neoeptesicus orinocensis (Ramírez-Chaves et al., 2021)	Ramírez-Chaves et al. (2021)
Natalidae	Natalus tumidirostris Miller, 1900	Williams et al. (1983)

Threatened and near threatened species

The list of threatened species produced by the Ministry of the Environment, Brazilian government, lists three species as vulnerable (Furipterus horrens, Lonchophylla bokermanni, and Natalus macrourus) and one, Lonchophylla dekeyseri, as endangered (MMA 2014, 2022, 2023). The International Union for the Conservation of Nature’s (IUCN) Red List includes Natalus macrourus (named “Natalus espiritosantensis”) as “near threatened” (Tejedor and Dávalos 2016) and two species of Lonchophylla, L. bokermanni and L. dekeyseri, as “endangered” (Aguiar 2016, Aguiar and Bernard 2016). Additionally, Lochophylla mordax, Vampyrum spectrum, and Myotis ruber are classified as “near threatened” in the IUCN.

DISCUSSION

Comparisons among the lists

While it is desirable to maintain species lists as stable as possible, taxonomic revisions and consequent changes are an integral aspect of scientific progress and facilitate communication within the scientific community. In this context, we briefly discuss here the main differences between the most recent lists of Brazilian bats.

From the latest checklist developed by the Committee of the List of Brazilian Bats (CLMB; Garbino et al. 2020a), eight species were added (Artibeus amplus, Choeroniscus godmani, Glossophaga bakeri, Lichonycteris obscura, Platyrrhinus guianensis, Molossus melini, Trachops ehrhardti, and Myotis pampa), and three species were removed (Glossophaga commissarisi, Micronycteris homezorum and Lichonycteris degener).

The Taxonomic Catalog of the Brazilian Fauna (TCBF) differs from the present list in the following aspects: (1) the TCBF considers Micronycteris brosseti (Phyllostomidae) as occurring in Brazil, which we do not (see Garbino 2016); (2) it treats M. homezorum as valid, while we consider it as a synonym of M. minuta; (3) the TCBF considers the species Chiroderma vizottoi as valid, while we treat it as a subspecies of C. doriae (Garbino et al. 2020b); (4) the mormoopid Pteronotus parnellii is still present in the TCBF, but now it has been considered endemic to the Antilles, whereas the name availa ble for the Brazilian populations is P. rubiginosus (Pavan and Marroig 2016; Pavan 2019); (5) in the family Molossidae, the TCBF considers Cynomops paranus of Thomas, 1901 as valid, while we treat it as a synonym of C. planirostris (Moras et al. 2016, 2018); (6) we included Eumops chimaera, which has not been included in the TCBF; (7) the Vespertilionidae in the TCBF differs in not including the valid taxon Histiotus diapha nopterus, and by including Lasiurus cinereus and L. salinae, which are not considered valid or occur elsewhere (Baird et al. 2015). These inconsistencies in the TCBF stem from the fact that it is a multi-author list that can be updated at any time. As a result, some taxa were last updated in 2015, and others were altered in the last couple of years.

The main difference between the list presented here and Quintela et al. (2020), is that the latter authors did not include Lasiurus villosissimus, included L. cinereus, which does not occur in Brazil, and considered L. salinae valid, despite the considerations of a previous study (Baird et al. 2015). The former authors also consider Cynomops paranus as valid, despite previous studies treating it as a junior synonym of C. planirostris (Moras et al. 2016, 2018). Additionally, Quintela et al. (2020) included Myotis dinellii in their checklist, which is not considered to occur in Brazil, as the records are misidentifications of Myotis albescens (see Nogueira et al. 2018).

The 2022 checklist from the Brazilian Society of Mammalogy (SBMz) listed 182 bat species in Brazil (Abreu et al. 2022), differing solely in terms of recent post-2022 changes that we have incorporated into the current list. The most recent checklist from the SBMz includes the same species presented here, except for Molossus melini and Trachops ehrhardti (Abreu et al. 2023). The SBMz committee shares members with the SBEQ list and both societies aim to have the same list for Chiroptera. Therefore, as the SBMz is updated, it should be congruent with the list presented here.

We note that Artibeus amplus and C. godmani were previously considered doubtful records for Brazil (Nogueira et al. 2014), and their removal from this category is the main change in our second list (Table 1). We have also, in this study, removed Myotis alter from the doubtful record list and assigned it as an erroneous record (Table 2). The 22 species with potential occurrence in Brazil (Table 3) represent ca. 10% of current number reported for the country, and include mostly phyllostomids (10), but also vespertilionids (4), and molossids (3). These three families also account for most records in our main list (^{Appendix 1}). In 1996, Taddei estimated the number of bat species that could potentially be recorded in Brazil based on geographic proximity, accounting for 22 species (Taddei 1996). Nearly three decades later, it has been confirmed that eight of these species do indeed occur in the country. One species predicted by him, Natalus tumidirostris, is also present in our list or potential species, but the other 13 he assigned were not considered here because their closest records either fall outside our 200 km buffer, or, if within it, they were restricted to higher altitudes along the eastern Andes.

Taxonomic and nomenclatural comments

Micronycteris homezorum - In our first assessment, we retained M. homezorum Pirlot, 1967 as valid (Nogueira et al. 2014), contra Ochoa and Sánchez (2005). However, because species limits in the subgenus Schizonycteris are still unclear and M. minuta might represent a species complex (Morales-Martínez et al. 2021), we opted for following Ochoa and Sánchez (2005) and several recent authors in treating this species as junior synonym of M. minuta (Siles and Baker 2020, Díaz et al. 2021, Morales-Martínez et al. 2021, Simmons and Cirranello 2023), until additional data are available.

Micronycteris sanborni - Solari et al. (2019) have conside red the records of M. sanborni from Bolivia to be referable to M. yatesi. However, a recent revision of Bolivian bats mentions M. sanborni for the country (Poma-Urey et al. 2020, 2023). Therefore, we do not consider M. sanborni a Brazilian endemic.

Gardnerycteris crenulata - Brandão et al. (2019) were the first to correct the spelling of Gardnerycteris crenulatum to G. crenulata to agree with the feminine -nycteris. We followed their suggestion, as did the Mammal Diversity Database and batnames lists (MDD 2023, Simmons and Cirranello 2023).

Trachops ehrhardti - Williams and Genoways (2008) recognize a polytypic Trachops with two subspecies occurring in Brazil: T. c. cirrhosus (Spix, 1823) and T. c. ehrhardti Felten, 1956. An unpublished PhD dissertation suggested treating ehrhardti as a full species based on DNA sequence data and morphology (Fonseca 2019). The Mammal Diversity Database includes ehrhardti “tentatively” in cirrhosus, citing the unpublished dissertation of Fonseca (2019). The 2022B version of Batnames (https://doi.org/10.5281/zenodo.6857865) lists T. ehrhardti as a valid species, but the 2023 update (Version 1.3) treats it as a junior synonym of T. cirrhosus. A recent study provided morphological and molecular evidence for splitting T. cirrhosus in three species, suggesting that T. ehrhardti is endemic to southern and southeastern Brazil (Fonseca et al. 2024).

Choeroniscus godmani - We include the species in the list, following Garbino et al. (2022), who recorded the species in Brazil based on museum specimens, including material collected in 1977 in the state of Mato Grosso.

Glossophaga commissarisi and G. bakeri - Webster and Jones (1987) originally described bakeri as a subspecies of Glossophaga commissarisi from the western Amazonia of Brazil, Colombia, and Peru. However, Velazco et al. (2021) showed that bakeri is morphologically distinct from G. commissarisi and should be recognized as a distinct species, endemic to the western Amazonia. Following the latter authors, we consider the records of G. commissarisi from Brazil to represent G. bakeri.

Lichonycteris obscura and L. degener - We follow Zamora-Gutierrez and Ortega (2020) in considering L. degener Miller, 1931 as a junior synonym of L. obscura Thomas, 1895 pending a taxonomic revision of the genus. Although Griffiths and Gardner (2008) recognize two species in Lichonycteris, the authors point out that Miller compared degener with a specimen of “obscura” that probably was collected in Brazil. Other authors treat Lichonycteris as monotypic (Gardner 1976, Hill 1985, Zamora-Gutierrez and Ortega 2020).

Glyphonycteris behnii - Here, we consider the species to be endemic to Brazil, due to re-identification of the Bolivian specimen, which is a Microncyteris hirsuta (Poma-Urey et al. 2023).

Carollia brevicauda - Solari et al. (2019) suggested using the spelling brevicaudum for C. brevicauda, arguing that the original description as Phyllostoma brevicaudum Schinz, 1821 should be followed and that an emendation would be unjustified. Considering that the author of the genus Carollia, and the first subsequent authors that used it, considered the name as feminine (Gray 1838, 1866, Peters 1865), we assume the gender as feminine. We also follow Article 30.2 of The International Code of Zoological Nomenclature, which states that “If no gender was specified or indicated, the name is to be treated as masculine, except that, if the name ends in -a the gender is feminine.” The genitive of the feminine cauda is caudae and the stem is caud-. Therefore, considering that Carollia is a feminine noun, the species name would be Carollia brevicauda (Handley 1980).

Artibeus amplus - We include this species of subgenus Artibeus in the list, following Zortéa et al. (2023).

Platyrrhinus guianensis - We include the species in the list, following Lopes et al. (2023).

Platyrrhinus helleri - Despite appearing in recent papers on Brazilian bats (Fischer et al. 2015, 2022, Acero-Murcia et al. 2023), the occurrence of P. helleri has been restricted to northern South America west of the Andes and to Mexico/Central America (Velazco et al. 2010). Garbino et al. (2024) have sequenced mitochondrial DNA from specimens of small-sized Platyrrhinus from the Brazilian Cerrado and Pantanal and confirmed that the taxon occurring in the region is P. incarum.

Molossus melini - This recently described species was previously known only for Argentina and has now been recorded for Curitiba, in the Brazilian state of Paraná (Montani et al. 2021, Olímpio et al. 2024)

Genus Lasiurus - Baird et al. (2015) classified hoary, yellow, and red bats into three genera, respectively, Aeorestes, Dasypterus, and Lasiurus based on molecular and phenotypic divergences. As Lasiurus, the generic name used to encompass these bats, is still monophyletic and the morphological differences among the clades are not sharply contrasting, we retain the use of a single genus Lasiurus for red, yellow, and hoary bats. In this case, the use of subgenera is recommended to maintain binomen stability and still convey the idea of three distinct clades (Garbino 2015, Novaes et al. 2018, Teta 2019, Burgin 2023).

Neoeptesicus - We use here the genus name Neoeptesicus Cláudio et al., 2023, for the Brazilian taxa formerly included in Eptesicus. Over the last decades, Eptesicus (sensu lato) has been recovered as non-monophyletic in molecular phylogenies (Hoofer and Bussche 2003, Juste et al. 2013, Yi and Latch 2022), and different taxonomic arrangements were proposed and used to solve this. Cláudio et al. (2023) reevaluated the taxonomic status of Eptesicus and restricted the name to the type species Eptesicus fuscus (Palisot de Beauvois, 1796), as well as E. guadeloupensis Genoways & Baker, 1975, and E. miradorensis (H. Allen, 1866), all of which do not occur in Brazil (Ramírez-Chaves et al. 2023). All other New World species were allocated to Neoeptesicus. As consequence of this arrangement, we use Histiotus as genus, instead as a subgenus of Eptesicus, which was proposed by some authors (e.g., Giménez et al. 2019).

Neoeptesicus andinus - The occurrence of N. andinus in Brazil is based on a specimen from Anápolis, Goiás (AMNH 134910). This specimen was named by Davis (1966) as Eptesicus montosus montosus in his broad review of South American Eptesicus. Later, Simmons and Voss (1998) partially reviewed the taxonomy of Eptesicus andinus and E. chiriquinus, and listed E. montosus as conspecific with E. andinus (Davis and Gardner 2008). We compared the specimen AMNH 134910 with the holotype of Eptesicus andinus (AMNH 33807) and it does not share the diagnostic traits of E. andinus. Consider ing this, we opted to remove E. andinus from the list, and the identity of specimen AMNH 134910 remains uncertain pending additional morphological and molecular analyses. Pine et al. (1970) mention E. m. montosus (USNM 393769) for Mato Grosso state; this specimen was examined by us and does not represent Neoeptesicus andinus.

Myotis pampa - We add here the first record of M. pampa for Brazil, based on specimen AMNH 235919 from Candelária, Rio Grande do Sul. This species was described based on specimens from Uruguayan grasslands, very close to the Brazil border. Although its occurrence in Brazil was considered probable by Novaes et al. (2021), the present record is the first to confirm its occurrence in Brazil.

Final remarks

Knowledge of bat diversity in Brazil seems to be close to an asymptote (Fig. 1). However, this may be result of having several lists published in a short time interval. Despite having a more stable species list of Brazilian bats, there are still knowledge gaps on the specific occurrence records of the species in the country. For example, less than half of the 27 Brazilian Federative Units have updated bat checklists (Tavares et al. 2024). Even in states that have bat checklists, dramatic changes can be seen as these lists are updated (Ferreira et al. 2024). Also, a vast area of Brazil is undersampled for bats (Bernard et al. 2011), increasing the potential of obtaining new records for the country. We suggest that future efforts focus on cataloging bat diversity in each Federative Unit of Brazil, refining our knowledge of species occurring in each ecosystem. A more detailed species-level list, including subspecies, is also important to refine our understanding of Brazilian bat diversity.

In this context, collections-based research is essential, as well as the collection and proper curation of specimens from poorly sampled regions. There has been an increase in local and regional mammal collections in Brazil, which must be followed by investment in research and infrastructure (Chiquito et al. 2021). It is also important to reiterate that several ongoing systematic studies involving bat taxa occurring in Brazil (e.g., Anoura, Furipterus, Lonchophylla, Myotis, Neoeptesicus, Nyctinomops, and Histiotus) are likely to lead to taxonomic rearrangements in the near future. Likewise, frequent faunal inventories and more refined studies using molecular methods for taxonomic identification may yield new records for Brazil. Consequently, the publication of reliable, comprehensive, and regularly updated listings, along with notes on range extensions, should always be prioritized.

ACKNOWLEDGEMENTS

This paper is dedicated to the memory of Dr. Nelio Roberto dos Reis, for his long contribution to the understanding of Brazilian bats, especially through his “Mamíferos do Brasil” and “Morcegos do Brasil” book series, aimed both at mammalogists and at a broader audience. We express our gratitude to the Sociedade Brasileira para o Estudo de Quirópteros (SBEQ) for their unwavering support and for hosting earlier versions of the Brazilian bat list. We also extend our thanks to the curators and staff at the collections we visited. GSTG received support from the Latin American Fellowship Committee of the American Society of Mammalo gists. RLMN has received support from Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ, Brazil; E-26/204.243/2021; E26/200.631/2022 and E26/200.395/2022). RG receives support from Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq process 301946/2022-1) and Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG process PPM 00203/18). RM has received financial support from Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, Brazil; 313963/2018-5), FAPERJ (E-26/200.967/2021), and Coordenação de Vigilância em Saúde e Laboratórios de Referência (CVSLR/Fiocruz). MCN has received support from Conselho Nacional de Desenvolvimento Científico e Tecnológico (PROTAX N.º 22/2020 - process 441714/2020-0) and TCCE ICMBio/Vale II (contrato 005/2021). VCC has received support from Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ, Brazil; E-26/205.820/2022 and E-26/205.821/2022); and research grants from the Field Museum of Natural History and American Museum of Natural History. Two anonymous reviewers provided very important advice and suggestions on an earlier version of this manuscript.

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ADDITIONAL NOTES

• ZooBank register

  https://zoobank.org/BBE43751-D48C-40A0-A198-4A8A700E7C75

• How to cite this article

  Garbino GST, Cláudio VC, Gregorin R, Lima IP, Loureiro LO, Moras LM, Moratelli R, Nascimento MC, Nogueira MR, Novaes RLM, Pavan AC, Tavares VC, Peracchi AL (2024) Updated checklist of bats (Mammalia: Chiroptera) from Brazil. Zoologia 41: e23073, https://doi.org/10.1590/S1984-4689.v41.e23073

• Published by

  Sociedade Brasileira de Zoologia at Scientific Electronic Library Online (https://www.scielo.br/zool)

Supplementary material 1

Supplementary S1. Maps of bat species potentially occurring in Brazil.

Authors: GST Garbino, VC Cláudio, R Gregorin, IP Lima, L Loureiro, L Moras, et al.

Data type: Range maps.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Link: https://doi.org/10.3897/zoologia.41. e23073

Appendix 1

List of families, subfamilies, genera, and species of bats that occur in Brazil. Subfamilies of Phyllostomidae are arranged in phylogenetic order, following Solari et al. (2019). Endemic species are denoted with an asterisk.

Chiroptera Blumenbach, 1779 (9 families, 68 genera, 186 species)

Emballonuridae Gervais, 1856 (7 genera, 17 species)

Emballonurinae Gervais, 1856

Centronycteris maximiliani (Fischer, 1829)

Cormura brevirostris (Wagner, 1843)

Cyttarops alecto Thomas, 1913

Diclidurus albus Wied, 1820

Diclidurus ingens Hernández-Camacho, 1955

Diclidurus isabella (Thomas, 1920)

Diclidurus scutatus Peters, 1869

Peropteryx kappleri Peters, 1867

Peropteryx leucoptera Peters, 1867

Peropteryx macrotis (Wagner, 1843)

Peropteryx pallidoptera Lim et al., 2010

Peropteryx trinitatis Miller, 1899

Rhynchonycteris naso (Wied, 1820)

Saccopteryx bilineata (Temminck, 1838)

Saccopteryx canescens Thomas, 1901

Saccopteryx gymnura Thomas, 1901

Saccopteryx leptura (Schreber, 1774)

Phyllostomidae Gray, 1825 (43 genera, 96 species)

Micronycterinae Van Den Bussche, 1992

Lampronycteris brachyotis (Dobson, 1879)

Micronycteris hirsuta (Peters, 1869)

Micronycteris megalotis (Gray, 1842)

Micronycteris microtis Miller, 1898

Micronycteris minuta (Gervais, 1856)

Micronycteris sanborni Simmons, 1996

Micronycteris schmidtorum Sanborn, 1935

Desmodontinae Wagner, 1840

Desmodus rotundus (É. Geoffroy, 1810)

Diaemus youngii (Jentnik, 1893)

Diphylla ecaudata Spix, 1823

Lonchorhininae Gray, 1866

Lonchorhina aurita Tomes, 1863

Lonchorhina inusitata Handley & Ochoa, 1997

Phyllostominae Gray, 1825

Chrotopterus auritus (Peters, 1856)

Gardnerycteris crenulata (É. Geoffroy, 1803)

Lophostoma brasiliense Peters, 1867

Lophostoma carrikeri (Allen, 1910)

Lophostoma schulzi (Genoways & Williams, 1980)

Lophostoma silvicola d’Orbigny, 1836

Macrophyllum macrophyllum (Schinz, 1821)

Mimon bennettii (Gray, 1838)

Phylloderma stenops Peters, 1865

Phyllostomus discolor Wagner, 1843

Phyllostomus elongatus (É. Geoffroy, 1810)

Phyllostomus hastatus (Pallas, 1767)

Phyllostomus latifolius (Thomas, 1901)

Tonatia bidens (Spix, 1823)

Tonatia maresi Williams, Willig & Reid, 1995

Trachops cirrhosus (Spix, 1823)

Trachops ehrhardti Felten, 1956 *

Vampyrum spectrum (Linnaeus, 1758)

Glossophaginae Bonaparte, 1845

Anoura caudifer (É. Geoffroy, 1818)

Anoura geoffroyi Gray, 1838

Choeroniscus godmani (Thomas, 1903)

Choeroniscus minor (Peters, 1868)

Dryadonycteris capixaba Nogueira et al., 2012 *

Glossophaga bakeri Webster & Jones, 1987

Glossophaga longirostris Miller, 1898

Glossophaga soricina (Pallas, 1766)

Lichonycteris obscura Thomas, 1895

Scleronycteris ega Thomas, 1912

Lonchophyllinae Griffiths, 1982

Hsunycteris pattoni (Woodman & Timm, 2006)

Hsunycteris thomasi (Allen, 1904)

Lionycteris spurrelli Thomas, 1913

Lonchophylla bokermanni Sazima, Vizotto & Taddei, 1978 *

Lonchophylla dekeyseri Taddei, Vizotto & Sazima, 1983

Lonchophylla inexpectata Moratelli & Dias, 2015 *

Lonchophylla mordax Thomas, 1903 *

Lonchophylla peracchii Dias, Esbérard & Moratelli, 2013 *

Xeronycteris vieirai Gregorin & Ditchfield, 2005 *

Carolliinae Miller, 1924

Carollia benkeithi Solari & Baker, 2006

Carollia brevicauda (Schinz, 1821)

Carollia perspicillata (Linnaeus, 1758)

Glyphonycterinae Baker et al., 2016

Glyphonycteris behnii (Peters, 1865) *

Glyphonycteris daviesi (Hill, 1964)

Glyphonycteris sylvestris Thomas, 1896

Neonycteris pusilla (Sanborn, 1949) *

Trinycteris nicefori (Sanborn, 1949)

Rhinophyllinae Baker et al., 2016

Rhinophylla fischerae Carter, 1966

Rhinophylla pumilio Peters, 1865

Stenodermatinae Gervais, 1856

Ametrida centurio Gray, 1846

Artibeus amplus Handley, 1987

Artibeus anderseni Osgood, 1916

Artibeus bogotensis Andersen, 1906

Artibeus cinereus (Gervais, 1856)

Artibeus concolor Peters, 1865

Artibeus fimbriatus Gray, 1838

Artibeus gnomus Handley, 1987

Artibeus lituratus (Olfers, 1818)

Artibeus obscurus (Schinz, 1821)

Artibeus planirostris (Spix, 1823)

Chiroderma doriae Thomas, 1891

Chiroderma trinitatum Goodwin, 1958

Chiroderma villosum Peters, 1860

Mesophylla macconnelli Thomas, 1901

Platyrrhinus angustirostris Velazco, Gardner & Patterson, 2010

Platyrrhinus aurarius (Handley & Ferris, 1972)

Platyrrhinus brachycephalus (Rouk & Carter, 1972)

Platyrrhinus fusciventris Velazco, Gardner & Patterson, 2010

Platyrrhinus guianensis Velazco & Lim, 2014

Platyrrhinus incarum (Thomas, 1912)

Platyrrhinus infuscus (Peters, 1880)

Platyrrhinus lineatus (É. Geoffroy, 1810)

Platyrrhinus recifinus (Thomas, 1901) *

Pygoderma bilabiatum (Wagner, 1843)

Sphaeronycteris toxophyllum Peters, 1882

Sturnira giannae Velazco & Patterson, 2019

Sturnira lilium (É. Geoffroy, 1810)

Sturnira magna de la Torre, 1966

Sturnira tildae de la Torre, 1959

Uroderma bilobatum Peters, 1866

Uroderma magnirostrum Davis, 1968

Vampyressa pusilla (Wagner, 1843)

Vampyressa thyone Thomas, 1909

Vampyriscus bidens (Dobson, 1878)

Vampyriscus brocki (Peterson, 1968)

Vampyrodes caraccioli (Thomas, 1889)

Mormoopidae Saussure, 1860 (1 genus, 4 species)

Pteronotus alitonus Pavan, Bobrowiec & Percequillo, 2018

Pteronotus gymnonotus (Wagner, 1843)

Pteronotus personatus (Wagner, 1843)

Pteronotus rubiginosus (Wagner, 1843)

Noctilionidae Gray, 1821 (1 genus, 2 species)

Noctilio albiventris Desmarest, 1818

Noctilio leporinus (Linnaeus, 1758)

Furipteridae Gray, 1866 (1 genus, 1 species)

Furipterus horrens (Cuvier, 1828)

Thyropteridae Miller, 1907 (1 genus, 5 species)

Thyroptera devivoi Gregorin et al., 2006

Thyroptera discifera (Lichtenstein & Peters, 1854)

Thyroptera lavali Pine, 1993

Thyroptera tricolor (Spix, 1823)

Thyroptera wynneae Velazco et al., 2014

Natalidae Gray, 1866 (1 genus, 1 species)

Natalus macrourus (Gervais, 1856)

Molossidae Gervais, 1856 (8 genera, 33 species)

Molossinae Gervais, 1856

Cynomops abrasus (Temminck, 1826)

Cynomops greenhalli Goodwin, 1958

Cynomops mastivus (Thomas, 1911)

Cynomops milleri (Osgood, 1914)

Cynomops planirostris (Peters, 1866)

Eumops auripendulus (Shaw, 1800)

Eumops bonariensis (Peters, 1874)

Eumops chimaera Gregorin et al., 2016

Eumops dabbenei Thomas, 1914

Eumops delticus Thomas, 1923

Eumops glaucinus (Wagner, 1843)

Eumops hansae Sanborn, 1932

Eumops maurus (Thomas, 1901)

Eumops patagonicus Thomas, 1924

Eumops perotis (Schinz, 1821)

Eumops trumbulli (Thomas, 1901)

Molossops neglectus Williams & Genoways, 1980

Molossops temminckii (Burmeister, 1854)

Molossus aztecus Saussure, 1860

Molossus coibensis Allen, 1904

Molossus currentium Thomas, 1901

Molossus fluminensis Lataste, 1891

Molossus melini Montani et al., 2021

Molossus molossus (Pallas, 1766)

Molossus pretiosus Miller, 1902

Molossus rufus É. Geoffroy, 1805

Neoplatymops mattogrossensis (Vieira, 1942)

Nyctinomops aurispinosus (Peale, 1848)

Nyctinomops laticaudatus (É. Geoffroy, 1805)

Nyctinomops macrotis (Gray, 1840)

Promops centralis Thomas, 1915

Promops nasutus (Spix, 1823)

Tadarida brasiliensis (I. Geoffroy, 1824)

Vespertilionidae Gray, 1821 (5 genera, 27 species)

Vespertilioninae Gray, 1821

Histiotus alienus Thomas, 1916 *

Histiotus diaphanopterus Feijó, Rocha & Althoff, 2015

Histiotus laephotis Thomas, 1916

Histiotus montanus (Philippi & Landbeck, 1861)

Histiotus velatus (I. Geoffroy, 1824)

Lasiurus blossevillii ([Lesson, 1826])

Lasiurus castaneus Handley, 1960

Lasiurus ebenus Fazzolari-Corrêa, 1994 *

Lasiurus ega (Gervais, 1856)

Lasiurus egregius (Peters, 1870)

Lasiurus villosissimus (É. Geoffroy, 1806)

Neoeptesicus brasiliensis (Desmarest, 1819)

Neoeptesicus chiriquinus (Thomas, 1920)

Neoeptesicus diminutus (Osgood, 1915)

Neoeptesicus furinalis (d’Orbigny & Gervais, 1847)

Neoeptesicus taddeii (Miranda, Bernardi & Passos, 2006) *

Rhogeessa hussoni Genoways & Baker, 1966

Rhogeessa io Thomas, 1903

Myotinae Tate, 1942

Myotis albescens (É. Geoffroy, 1806)

Myotis izecksohni Moratelli et al., 2011

Myotis lavali Moratelli et al., 2011

Myotis levis (I. Geoffroy, 1824)

Myotis nigricans (Schinz, 1821)

Myotis pampa Novaes, Wilson & Moratelli, 2021

Myotis riparius Handley, 1960

Myotis ruber (É. Geoffroy, 1806)

Myotis simus Thomas, 1901

Edited by

Editorial responsibility

José Anderson Feijó

Data availability

Data citations

Abreu EF, Casali D, Costa-Araújo R, Garbino GST, Libardi GS, Loretto D, et al. (2021) Lista de Mamíferos do Brasil. Zenodo. https://doi.org/10.5281/zenodo.5802047

Abreu EF, Casali D, Costa-Araújo R, Garbino GST, Libardi GS, Loretto D, et al. (2022) Lista de Mamíferos do Brasil. Zenodo. https://doi.org/10.5281/zenodo.7469767

Abreu EF, Casali D, Costa-Araújo R, Garbino GST, Libardi GS, Loretto D, et al. (2023) Lista de Mamíferos do Brasil (2023-1). Zenodo. https://doi.org/10.5281/zenodo.10428436

Aguiar LMS (2016) Lonchophylla bokermanni In: The IUCN Red List of Threatened Species. https://doi.org/10.2305/IUCN.UK.2016-3.RLTS.T12263A22038287.en [Accessed: 22/08/2023]

Aguiar L, Bernard E (2016) Lonchophylla dekeyseri In: The IUCN Red List of Threatened Species. https://doi.org/10.2305/IUCN.UK.2016-2.RLTS.T12264A22038149.en [Accessed: 22/08/2023]

MDD (2023) Mammal Diversity Database. Zenodo. https://doi.org/10.5281/zenodo.7830771

Tejedor A, Dávalos L (2016) Natalus espiritosantensis In: The IUCN Red List of Threatened Species. https://doi.org/10.2305/IUCN.UK.2016-2.RLTS.T136448A21983924.en [Accessed: 21/08/2023]

Publication Dates

• Publication in this collection
  16 Aug 2024
• Date of issue
  2024

History

• Received
  10 Oct 2023
• Accepted
  29 June 2024