A systematic revision of the bats (Chiroptera) of Honduras: an updated checklist with corroboration of historical specimens and new records

During the last century, survey efforts for mammals in Honduras have been few and most distributional and conservation assessments of bats have been based on historical records. Taxonomy of many records has changed. Moreover, a number of supposed Honduran occurrences are based on records from bordering countries without confirmation by a Honduran voucher. Therefore, the list of bats of Honduras lacks precision. Here, we update the number of species in the country, including taxonomic changes not reflected in recent works and new records based on museum specimens. The known number of species for Honduras is 113 with seven expected (Cormura brevirostris, Lampronycteris brachyotis, Mesophylla macconnelli, Molossus coibensis, M. pretiosus, Thyroptera discifera and Trinycteris nicefori), based on records in adjoining countries. We provide a new record for Honduras of Natalus lanatus. We confirm the presence of Cynomops greenhalli and Diaemus youngii and clarify the taxonomic status of Artibeus intermedius, Chiroderma gorgasi, Eumops ferox, Gardnerycteris keenani, Lasiurus frantzii, Myotis pilosatibialis, Molossus and Pteronotus species, and Tonatia bakeri. We recommend a reassessment of the conservation status of the bats of Honduras considering recent changes and that a number of species (e.g. Choeronycteris mexicana) have not been observed since their reports in historical records. This requires an update of the taxonomic identification keys for Honduras. The updated checklist below demonstrates the high biodiversity of Honduran bats but is also an example of how poorly many groups have been studied since they were first recorded in the country.


Introduction
Rodríguez Herrera and Sánchez (2015) reported 98 species of bats for Guatemala, 109 for Honduras, 68 for El Salvador, 102 for Nicaragua and 114 for Costa Rica. In the recent list of bats of Mexico presented by Ramírez-Pulido al. (2020a) increased the list for Honduras to 111 plus four expected species and recently, Medina-Fitoria and Martínez-Fonseca (2019) and Saldaña Tapia et al. (2020) increased the list for Nicaragua to 111 species. In addition, York et al. (2019) reported 120 for Costa Rica. This demonstrates that Central America is one of the regions in the world with the highest number of genera (66) of bats with more than 170 recorded species (Rodríguez Herrera and Sánchez 2015), of which over 65.29% occur in Honduras. Not only have researchers recorded new species for each country, but they have also carried out systematic and taxonomic studies that have defined new lineages, new species and new taxonomic arrangements.
Frequent taxonomic changes, molecular vs. morphological methods of delineating species and historical vouchers whose identifications have not been updated, are amongst the factors that affect the number of species known for each country. For this reason, our study aims to update species distributions, species checklists and corroborate museum vouchers for the bats of Honduras.
During the last century, sampling effort in Honduras has been very low and most distributional and conservation assessments of bat species have been based on historical records (Turcios-Casco and Medina-Fitoria 2019). Mora (2016) provided the first taxonomic keys for identification of Honduran bats, so it was no longer necessary to rely on keys from other countries [e.g. Costa Rica (Timm et al. 1999) and Mexico (Medellín et al. 2008)]. Such practices led to misidentifications of species endemic to northern Central America, but not found in Costa Rica or Mexico [e.g. Artibeus inopinatus Davis & Carter, 1964 (Fig. 1A, B)]. Like Guatemala (Kraker-Castañeda et al. 2016), Honduras has had a questionable number of known bat species since the 1900s. Unlike other megadiverse countries such as Mexico (Medellín et al. 2008;Ramírez-Pulido et al. 2014) and Costa Rica (LaVal and Rodríguez-H 2002;Rodríguez-Herrera et al. 2014;York et al. 2019), Honduras has had relatively few bat studies and distribution and ecology of most species are still unknown (Turcios-Casco et al. 2019b).
We summarise the history of bat research in Honduras: the description of Ectophylla alba H. Allen, 1892 (see McCarthy et al. 1993) was based on Honduran specimens in 1892 and then Allen (1898) further described the skull and teeth of this species. Goodwin (1940) described three new bats from Honduras (Phylloderma septentrionalis Goodwin, 1940, Sturnira hondurensis Goodwin, 1940 and Eumops underwoodi Goodwin, 1940) and provided the first record of Enchisthenes hartii (Thomas, 1892) for North America, based on a mammal collection made by C. F. Underwood. Subsequently, Goodwin (1942) presented the first checklist of mammals of Honduras with 69 bat species including Macrotus macrotus (= M. waterhousii Gray, 1843). More records were added to the bat list of Honduras by , , LaVal (1969) and Valdez and LaVal (1971). Jones et al. (1977) included at least 96 species for Honduras in their "Annotated checklist of the bats of Mexico and Central America", including Micronycteris brachyotis Dobson, 1879 [= Lampronycteris brachyotis (Dobson, 1878)]. Benshoof et al. (1984) added more records and Jones et al. (1988) reported that at least 100 species of bats included Honduras in their known distributions (e.g. Artibeus intermedius J.A. Allen, 1897). Consequently, McCarthy et al. (1993) noted 99 species for Honduras in his checklist (but only listed 98 in their final table). Reid (2009) included the distribution of some species that still remained uncertain in Honduras (e.g. Diaemus youngii (Jentink, 1893) and Hylonycteris underwoodi Thomas, 1903). Hernández (2015) mentioned 112 species based on the efforts done by the Program for Bat Conservation in Honduras (PCMH), but in the final table in Rodríguez Herrera and Sánchez (2015), there were only 109 species listed. Mora (2016) reported 110 species and four expected species and McCranie et al. (2018) and Mora et al. (2018) upheld the same number of species. The only new record for Honduras after the report by Mora et al. (2018) is by Turcios-Casco et al. (2020a), who updated the number of Honduran species to 111 with Chiroderma gorgasi Handley, 1960(referring to C. trinitatum Goodwin, 1958. In response to the uncertain number of bat species in Honduras, we provide an update, including taxonomic proposals not considered by Mora et al. (2018). These include: Mantilla-Meluk and Muñoz-Garay (2014) for Myotis pilosatibialis LaVal, 1973 andPavan andMarroig (2016) for Mormoopidae. Mora et al. (2018) included the changes of Pteronotus mesoamericanus Smith, 1972, but not P. fulvus (Thomas, 1892) and P. psilotis (Dobson, 1878).
They also failed to consider Gregorin et al. (2016) for Eumops Miller, 1906, Moras et al. (2018 for Cynomops Thomas, 1920 and some works of Loureiro et al. (2018Loureiro et al. ( , 2020 for Molossus É. Geoffroy, 1805. Finally, new records are given based on the verification of museum specimens examined by us, with remarks on their taxonomy and systematics.

Materials and methods
To update the number of bat species in Honduras, we first reviewed the database of GBIF.org (2019). Amongst the approximately 9000 Honduran records of bats deposited in museums over the world, we re-examined certain vouchers that were misidentified and that we could correctly identify, based on cranial and external measurements. These were measured to the nearest 0.01 mm with digital calipers. Measurements followed Srinivasulu et al. (2010), except for tragus length (Tr) and width (TrW) that followed Dietz and von Helversen (2004): forearm length (FA); tibia length (Tib); ear length (E); ear width (EW); thumb length (Th); length of the calcar (Ca); tail length (T); body length (BH); hindfoot length (HF); wingspan (WS); fifth metacarpal length (5mt); third metacarpal length (3mt); length of the first phalanx of digit III (1ph); length of the second phalanx of digit III (2ph); and the length of the third phalanx of digit III (3ph). Skull measurements obtained from vouchers followed Simmons and Voss (1998), Tejedor (2011) and Giménez and Giannini (2016). These included condyloincisive length (CIL), condylocanine length (CCL), zygomatic breadth (ZB), height of braincase (HB), mastoid breadth (MB), postorbital breadth (PB), mandibular tooth row length (MBL), maxillary tooth row length (MTL), depth of braincase (DB), breadth across molars (BAM), and breadth across canines (BAC).
Acronyms used for the museums are the following:  Fig. 1A, B) in southern Honduras, but no criteria for identification were provided. These individuals might have been misidentified considering the difficulty of identification of the four species of large Artibeus in Honduras. Moreover, Davis (1970) mentioned that the subspecies (e.g. Artibeus jamaicensis paulus Davis, 1970 (2015) suggested A. inopinatus be considered an endangered species in Honduras, specifically for the loss of its habitat due to anthropogenic causes (e.g. extensive clearing for agriculture, livestock and the general reduction of tropical dry forests in southern Honduras). Before Mora (2016) and Mora et al. (2018), there were no identification keys for bats in Honduras and most researchers used the keys of other countries, especially those of Mexico (Medellín et al. 2008) and Costa Rica (Timm et al. 1999), which do not include A. inopinatus, because it is endemic to Honduras, El Salvador and Nicaragua. Considering the controversial identification of A. inopinatus, many researchers could have confused it with certain subspecies of A. jamaicensis Leach, 1821 or even with juvenile A. jamaicensis or A. lituratus (Olfers, 1818). This could be one of the main reasons that A. inopinatus has been considered a rare species (see Turcios-Casco et al. 2020c) by Reid (2009) or categorised with deficient data by Reid and Medina (2016). Only in TTU and TCWC we know of 214 and 221 specimens respectively and there are two museum specimen in Honduras, one (CZB-2019-10) in the department of Comayagua (La Carbonera, El Rosario) and one (UVS-V-02063) in the department of Francisco Morazán (Carboneras, Sabanagrande). To our knowledge, the individual captured in San Buenaventura in Francisco Morazán by Turcios-Casco et al. (2020c) is the highest elevational record for the species which was captured at 1435 m a.s.l.

EAPZ
The case of Artibeus (sensu lato) has been controversial and York et al. (2019) mentioned an easy distinguishing characteristic between A. intermedius and A. lituratus (e.g. A. lituratus has both pairs of facial stripes distinct and A. intermedius only the stripes above eye) in Costa Rica. Simmons (2005) did not recognise A. intermedius, because she believed that individuals identified as A. intermedius represent individuals of A. l. palmarum J. A. Allen & Chapman, 1897 that fall at the lower end of the normal range of size variation for that species. However, we followed Larsen et al. (2013) who suggest that that A. l. intermedius in Central America is the product of a recent ecologically-driven Neotropical expansion by A. lituratus. In addition, they noted that Davis (1984) and Marchán-Rivadeneira et al. (2012) provided indirect evidence of genetic isolation of A. intermedius by identifying sympatric A. intermedius and A. l. palmarum as morphotypes in Middle America. In conclusion, Larsen et al. (2013) gave concrete information, based on review of previous works plus molecular evidence, that support that A. intermedius must be treated as a separate species from A. lituratus reinforcing the hypothesis of Davis (1984). Evidence in Honduras supporting occurrence of A. intermedius (Davis 1984) is based on the existence of two size classes of Artibeus lituratus (sensu lato), a large size-class (referring to the formerly A. lituratus) and a small size-class (referring to what he hypothesised to be A. intermedius) from Santa Bárbara (western Honduras) and Brus Laguna (eastern Honduras). Davis (1984) identified specimens of A. intermedius from the following departments of Honduras: Choluteca and Valle in southern Honduras; Copán, Intibucá, La Paz, Ocotepeque and Santa Bárbara in western Honduras; Francisco Morazán and Comayagua in Central Honduras; and Gracias a Dios in eastern Honduras. Additionally, in the GBIF.org (2019), there are preserved specimens from Atlántida, Colón and Cortés deposited at TTU. These specimens from TTU complement the wide distribution of the species in almost all the country, excluding the departments of Lempira, El Paraíso, Yoro, Olancho and Islas de la Bahía. Finally, Simmons and Cirranello (2020) recognised A. intermedius as a distinct species from A. lituratus.
Balantiopteryx io Thomas, 1904. This species is known from Honduras only by the records of Divoll and Buck (2013) of six individuals (four females and two males) captured in a harp trap on the Masca River, Piedra Chaca in the department of Cortés. It appears there are no Honduran specimens of this species in any museum. The caves, in which these individuals were captured represent, the only known locality for the species in Honduras. B. io is classed as Vulnerable (Lim 2015) by the IUCN (International Union for the Conservation of Nature).
Centronycteris centralis Thomas, 1912. This species is included by Rodríguez Herrera and Sánchez (2015) and Mora et al. (2018) for Honduras, even though there is no record of C. centralis in the GBIF.org (2019). Goodwin (1942) and McCarthy et al. (1993) placed this species [then referred to as C. maximilliani (J. Fischer, 1829)] in the bat list of Honduras with no further details. Although Simmons (2005) and Hood and Gardner (2008) suggest this species occurs in Honduras, there are no published records or museum specimens to back up these assumptions. The only unofficial record was made by Hernández et al. (2016) in which vocalisations of C. centralis were recorded in the Cuyamel-Omoa National Park in the de-partment of Cortés, but without a precise location (coordinates and elevation). No further information is known of the species in the country.
Chiroderma gorgasi Handley, 1960. The first record of C. gorgasi (UVS-V-02529) for Honduras was collected in 2017 [= referring to C. trinitatum by Turcios-Casco et al. (2020a)]. This specimen came from the Caribbean lowlands of the Honduran Mosquitia in the historical city of Ciudad Blanca, in the department of Gracias a Dios. This record is the only one known from northern Central America and represents a range extension of 512 km from Tortuguero, Costa Rica (Turcios-Casco et al. 2020a). However, we followed Lim et al. (2020) and the species that occurs in north-western Ecuador, western Colombia, Panama, Costa Rica and Honduras is C. gorgasi.
Cynomops Thomas, 1920. LaVal (1969 reported a lactating female captured in the department of Comayagua as the first record of Molossops greenhalli Goodwin & Greenhall, 1958 in Central America. Although the systematics of Cynomops (= Molossops) have been somewhat problematic, we decided to follow Peters et al. (2002) who stated that C. greenhalli Goodwin, 1958 is the species that occurs in Honduras. C. mexicanus (Jones & Genoways, 1967) had been previously reported from Honduras, but Peters et al. (2002) restricted C. mexicanus to Mexico. Additionally, Peters et al. (2002) suggested that C. paranus (Thomas, 1901), C. planirostris (Peters, 1865), C. g. greenhalli and C. g. mexicanus cannot be definitely separated based on size, and this has been one of the main causes of taxonomic confusion within this group. Simmons (2005), amongst others, does not agree with this treatment, but more recent authors [Eger (2008), Moras et al. (2016) and Moras et al. (2018)] recognised C. greenhalli and C. mexicanus as distinct species. Currently, Simmons and Cirranello (2020) recognised C. greenhalli to be distinct from C. mexicanus.
Erroneously, the species that had been previously recognised in Honduras was C. mexicanus [e.g. Hernández (2015), Mora (2016), Mora et al. (2018)], but to date, the only specimen of Cynomops is the female captured in the department of Comayagua by LaVal (1969) which is C. greenhalli and not C. mexicanus. Additionally, there is a record of C. greenhalli (TCWC 24178) from Lancetilla, 40 m a.s.l., in the department of Atlántida (Valdez and LaVal 1969;Prestridge 2019b). Table 1 shows a morphometric comparison amongst other Central American specimens plus that of the female misidentified as C. greenhalli (TTU 104070) that was captured by R. D. Bradley et al. in 2004, in the SAG, La Ceiba, in the department of Atlántida in northern Honduras (not Colón, as is incorrectly recorded on the original label). The specimen (TTU 104070) was identified as C. mexicanus ( Fig. 2A-C) following Moras et al. (2018), for the following reasons: 1) forearm shorter than 40.0 mm; 2) the rostrum was relatively low; 3) the anterior face of the lacrimal ridges sloping smoothly to the forehead; 4) incisive and accessory foramina arranged in the shape of an equilateral triangle instead of an isosceles triangle when viewed from above. Other measurements ( Timm et al. (1999), Reid (2009) and Mora (2016): 1) ears triangular; facial and dorsal colour grey brown and lightly frosted; ventrum whitish ( Fig. 3A), uropatagium reduced and hairy; and submandibular glands present as mentioned by Medina-Fitoria (2014); 2) wingtips white (Fig.   3B); 3) only one pad at the base of the thumb (Fig. 3C); 4) obvious large glands in the mouth. See Simmons and Cirranello (2020) for a discussion of the correct epithet, which is youngii.
Eptesicus Rafinesque, 1820. When Davis (1965) reviewed the E. brasiliensis complex, he mentioned that there is a population in the highlands of Middle America that appears to be identical to E. andinus from the highlands of Colombia and described the E. andinus group (E. chiriquinus, E. inca O. Thomas, 1920, E. montosus O. Thomas, 1920 as having soft pelage and long forearm (43-48 mm). Davis (1966) mentioned that E. andinus occurs from the Chiapas Highlands to the highlands of Colombia and Ecuador. Simmons and Voss (1998) noted that Davis (1966) considered E. chiriquinus and E. inca to be strict junior synonyms of E. andinus and Koopman (1978and Koopman ( , 1993and Koopman ( , 1994) considered E. andinus as a subspecies to E. brasiliensis and E. chiralensis H. E. Anthony, 1926 andE. montosus to E. furinalis (d'Orbigny, 1847). Simmons and Voss (1998), in accordance with Davis (1966), concluded that E. inca and E. chiriquinus are conspecific and selected chiriquinus as the senior synonym and provided a new diagnosis for the species. Davis and Gardner (2008) revised the genus Eptesicus for South America, restricted E. andinus from the highlands of Bolivia northwards at upper elevations along the Andes of Peru, Ecuador and Colombia, into north-western Venezuela with no details of distributions in Central America. On the other hand, Davis and Gardner (2008) stated that E. chiriquinus occurred at moderate to lower elevation in Colombia, Venezuela, the Guianas, Brazil and eastern Ecuador, Peru and Bolivia and elsewhere in Central America north-westwards into the Chiapan Highlands of Mexico. While reviewing the systematics of the E. andinus group, we encountered no mention of an adult female (TTU 104074) misidentified as Eptesicus andinus by R. D. Bradley et al. (Garner 2016a) which was captured in 2004 at S.A.G. Laboratorio, La Ceiba, in the department of Atlántida (not Colon, as is incorrectly recorded on the original label) in Honduras. We corroborate the identification of this specimen with the identification keys for Eptesicus in Central America of Davis (1966), the new diagnosis of E. chiriquinus by Simmons and Voss (1998),  LaVal (1969) in the department of Comayagua of C. greenhalli, and the measurements presented for both species by Moras et al. (2018). Means (ranges in parentheses); all the measurements are in mm.  (1993) for Central America and there is no voucher in the GBIF.org (2019) that supports its occurrence in Honduras. Nevertheless, the species is included on the bat list for Honduras by Rodríguez Herrera and Sánchez (2015) and Mora et al. (2018). Espinal and Mora (2012b) recorded a female of E. brasiliensis captured in El Corpus, department of Choluteca (southern Honduras), which was preserved, but unfortunately, it is part of a personal collection and is not deposited as a voucher in any museum. Identification was supported by vocalisations recorded to individuals of E. brasiliensis during the same survey. For comparison, one of us (LaVal) has recorded echolocation calls of this species at 27 of 59 localities in the Costa Rican Highlands and also in the Pacific and Caribbean Lowlands. This may well be a widespread species in Honduras, but more echolocation recordings are needed.

Cynomops greenhalli by Moras et al. (2018)
In conclusion, there are only three species of Eptesicus (E. brasiliensis, E. furinalis and E. fuscus Beauvois, 1796) reported for Honduras and, even though Davis and Gardner (2008) stated that E. chiriquinus may occur in Honduras, there is no evidence in the country.
Glyphonycteris Thomas, 1896. We did not find any specimens in GBIF.org (2019) of the genus but both species are included by Rodríguez Herrera and Sánchez (2015) and Mora et al. (2018) in the bat lists of Honduras. McCarthy et al. (1993) mentioned that D. C. Carter captured an individual of unknown sex of G. daviesi (Hill, 1964) (originally described as Barticonycteris daviesi) on the Perlas River where a trail to Valencia on the Patuca River crosses the river near Catacamas, in the department of Olancho. In April 1967, one of us (LaVal), who had actually captured the bat while netting with Carter, observed it as it escaped after chewing a hole in the bag in which it was held. Fortunately, the bat was already ten-  Lasiurus Gray, 1831. Mora and López (2013) described the first record of L. cinereus (Palisot de Beauvois, 1796) (EAPZ-06), based on a male specimen found dead in Cerro de Hula in Santa Ana municipality in southern Francisco Morazán at 1658 m a.s.l. There is only one more record of the species in Honduras by Espinal and Mora (2012a), based on audio recordings from San Marcos de Colón, in the department of Choluteca in southern Honduras. Mora (2012) recorded Lasiurus egregius (Peters, 1870) for the first time in Honduras, based on two specimens, a male (UCR 2067) and a female (UNAH 1456) captured during the night of 9 May 1998 in Catacamas, department of Olancho (north-eastern Honduras). These records cover a gap between Guatemala and Panama (see Mora (2012) for more details). In addition, based on Baird et al. (2015), the species that occur in Central America (see York et al. 2019) is L. frantzii Peters, 1870 andnot L. blossevillii (Lesson &Garnot, 1826), as is mentioned by Mora et al. (2018).
Leptonycteris yerbabuenae Martínez and Villa-R, 1940. We followed Wilson (2006a, 2006b), when recognising occurrence in Honduras of L. yerbabuenae. This distinction is well supported by published data (Wilkinson and Fleming 1996), in which L. curasoae Miller, 1900 and L. yerbabuenae (ZD 1999.194) from Yusguare, in the department of Choluteca (GBIF.org 2019, Fig. 7). Even though there are two records in the GBIF.org database, in the catalogue presented by the Natural History Museum (London) (2019), there is only the one individual captured by M. Sandiford in 1995. L. yerbabuenae is known only from southern Honduras in the departments of Choluteca and Valle. Hernández (2015) proposed the creation of the Área de Importancia para la Conservacion de Murcielagos (AI-COM) Golfo de Fonseca for the conservation of L. yerbabuenae. Its rarity in Central America is one of the reasons that this species is considered as Near Threatened (NT) by the IUCN. In Table 2 Basantes et al. (2020). The species that occurs in Honduras is T. bakeri Williams, Willig &Reid, 1995 andT. saurophila Koopman &Williams, 1951 is only known, based on subfossil remains found in caves of the type locality in Jamaica. Molossus. Goodwin (1942) gave the first records of Molossus bondae J.A. Allen in the department of La Paz, north-western Honduras. Dolan (1989) considered M. bondae to occur in Honduras, specifying that the species was known with certainty from Brus Laguna, in the department of Gracias a Dios in north-eastern Honduras, but LaVal (1977) stated that the individuals mentioned by Goodwin (1942) were not M. bondae, based on his personal examination of the specimens. McCarthy et al. (1993) considered Honduras as the northernmost country for the occurrence of M. bondae and consequently Burnett et al. (2001), Reid (2009), Hernández (2015, Mora (2016) and Mora et al. (2018) all listed M. bondae as occurring in Honduras. Additionally, López-González and Presley (2001) indicated that M. currentium Thomas, 1901 has priority over M. bondae. Although differences exist in cranial and external measurements in South and Central American individuals, the populations are sufficiently similar that they considered them to be conspecific, based on morphometric data. Eger (2008) believed that M. bondae and M. currentium were different species due to differences in colouration and fur length. However, we follow Loureiro et al. (2020) as the most recent taxonomy for Molossus, based on Dolan (1989) and González-Ruiz et al. (2011). We recognise 14 species of Molossus, based on Loureiro et al. (2020), including M. bondae, the species that occurs in Central America, and M. currentium restricted to South America. This species has its northern limit in Brus Laguna in the department of Gracias a Dios in north-eastern Honduras (Dolan and Carter 1979;Dolan 1989;López-González and Presley 2001); these reports are the only ones that are known for Honduras, although there is an old record by R. W. Adams of 1963, from Río Coco, in the department of El Paraíso in south-eastern Honduras (GBIF.org 2019).
Simmons and Cirranello (2020) mentioned that Loureiro et al. (2020) considered some specimens of M. sinaloae J. A. Allen, 1906 from Honduras to be M. alvarezi Gonzalez-Ruiz, Ramirez-Pulido & Arroyo-Cabrales, 2011. There has not been any doubt about the occurrence of M. molossus (Pallas, 1766) in the country. On the other hand, we consider M. aztecus Saussure, 1860 to occur in Honduras, because McCarthy et al. (1993) referred to two males captured in the department of La Paz in western Honduras. Additionally, Loureiro et al. (2020) and Simmons and Cirranello (2020) consider its occurrence in Jalisco and Cozumel in Mexico to Nicaragua, southern Venezuela, and south-eastern Brazil. Finally, M. nigricans Miller, 1902 is the species that occur in Mexico and Central America and M. rufus É. Geoffroy, 1805 is restricted to South America including Guianas, Ecuador, Peru, to central and northern Brazil and Bolivia, and Trinidad and Tobago Simmons and Cirranello 2020).   (Garner 2016b(Garner , 2016c. The records of Cole and Wilson (2006a) are from southern Arizona and Mexico (means before parentheses and ranges are included on them).  (Fig. 7). We identified the two individuals (TCWC 10992 and TCWC 11008) as N. lanatus following Tejedor (2011) (3) GSL = 16.2-16.5; (4) BB = 7.9-8.0; (5) MTL = 6.7-6.9; (6) the caudal margins of the maxilla have a shallow perpendicular to longitudinal axis of the skull; (7) margins of the ears were straight. See Table 3 for comparison with individuals mentioned by Tejedor (2011), Rodríguez Herrera et al. (2011) and Medina-Fitoria et al. (2015. These two specimens confirm the presence of N. lanatus in Honduras and help fill the gap between Mexico and Nicaragua. N. lanatus is the second species of the genus known for the country, although it was expected by Mora et al. (2018).

Myotis pilosatibialis
Nyctinomops Miller 1902. McCarthy et al. (1993 included only N. laticaudatus (E. Geoffroy, 1805) for Honduras, based on a record of a female [TCWC 19759 (Prestridge 2019a] captured by LaVal on the Aguán River in the department of Yoro (northern Honduras) and constitutes the only record known for the species in Honduras. Mora et al. (2016) confirmed the presence of N. macrotis (Gray, 1840) with two males (MVB 4962, MVB 4963) captured in San Marcos de Colón and El Corpus in the department of Choluteca (southern Honduras). Addition-ally, Espinal et al. (2016) reported a dead male of N. aurispinosus (Peale, 1848) also from San Marcos de Colón. Unfortunately, even though they gave morphometric information and stated that the specimen was preserved in fluids, no catalogue number was listed, and we cannot find the individual (to our knowledge it has not been deposited in any museum collection).
Sturnira Gray, 1842. We re-examined the specimens PSM 13753 and PSM 13754 that were misidentified as S. bogotensis Shamel, 1927. After reviewing individuals and analysing the mensural and cranial characteristics, we corroborate the specimens as S. hondurensis. Both individuals had bilobed lower incisors, forearm lengths greater than 43 mm, dark brown colouration and were captured in La Esperanza, in the department of Intibucá, which is over 1000 m a.s.l. Following Velazco and Patterson (2013), the only two species that occur in Honduras are S. parvidens and S. hondurensis.

Discussion
Although Honduras is a region in Mesoamerica of high biodiversity (McCranie et al. 2018;Larsen 2019), there are many mammalian groups that have been poorly studied (e.g. Chiroptera, Didelphimorphia, Rodentia and Xenarthra). Since the 1900s, the number of bat species in Honduras has been debated. After a careful review of recent taxonomic proposals and museum specimens, we confirm that there are 113 species in Honduras, plus seven expected. This makes Honduras the second most diverse country in Central America in number of bat species, just below Costa Rica which has, as of now, 120 recorded species (York et al. 2019) and above Nicaragua, with 111 species considering Medina-Fitoria and Martínez-Fonseca (2019) and Saldaña Tapia et al. (2020).
We augmented the most recent bat list of Honduras (Mora et al. 2018) Allen, 1904, M. pretiosus Miller, 1902, T. discifera and Trinycteris nicefori (Sanborn, 1949. The occurrence of E. bonariensis in Honduras has been controversial, because Jones et al. (1977) included it for Central America and Simmons (2005) listed the occurrence of the species as from Mexico to Peru, Argentina, Paraguay, Uruguay and Brazil. However, the species is currently restricted to South America (Wilson and Mittermeier 2019). In addition, we considered M. pretiosus of expected occurrence, because it was included by Jones et al. (1988) and more recently by Wilson and Mittermeier (2019) to occur in Central America. Although Jones et al. (1977) and Simmons (2005) have suggested that E. perotis (Schinz, 1821) occurs in Honduras, we remain unconvinced, because there are no verified records from any Central American country. We did not consider the occurrence of M. coibensis in Honduras, Mora et al. (2018) considered it in southern Honduras, because there is no veridical evidence of its occurrence in the country. Additionally, Lourerio et al. (2020) and Simmons and Cirranello (2020) mentioned that the distribution of M. coibensis is from México to South America.
Although there is no voucher of D. youngii from Honduras, we presented external measurements, ecological data and photos of a Honduran specimen, so now all three haematophagous species in America are confirmed for Honduras. This is the only record of D. youngii in Honduras, indicating that more sampling effort is needed to determine the distribution of the species in the country, but based on its frequency of occurrence elsewhere, D. youngii is the rarest of the desmodontinae species.
The following species are based on historical records only; in spite of our substantial effort since the 2000s, we do not know of any recent records (even unofficial) of the following species: C. mexicana, G. keenani, G. daviesi, G. sylvestris, L. yerbabuenae, L. obscura, M. macrophyllum and V. major. We strongly recommend a revision of the threatened species list proposed by Hernández (2015) and that it consider some of the previous species which are only known from the historical record in Honduras.
The cases of C. centralis, B. io, G. daviesi and G. sylvestris are also controversial and very similar to the case of D. youngii, because their occurrence in Honduras has been upheld with no museum specimens.
Our acceptance of not dividing Lasiurus into three genera: Lasiurus, Aeorestes and Dasypterus (Baird et al. 2015(Baird et al. , 2017 or of using Dermanura for some species formerly in the genus Artibeus (Cirranello et al. 2016; Simmons and Cirranello 2020) will not affect the real number of bat species in Honduras. York et al. (2019) reported the occurrence of A. intermedius in Nicaragua and Costa Rica. Larsen et al. (2013) gave evidence to support the hypothesis of Davis (1984) and recognised A. intermedius as a different species from A. lituratus. Based on the revision of Davis (1984), the occurrence of A. intermedius in Honduras is well supported. Another species, formerly considered rare, was A. inopinatus (Reid 2009 The keys for Honduras must be updated using the latest taxonomic arrangements and new records for the country. Furthermore, there are more than 9000 vouchers in different museums all over the world whose identification needs to be verified. More sampling is needed in certain areas of the country, especially in western Honduras (e.g. La Paz, Intibucá) or the most eastern region of the department of Gracias a Dios that borders Nicaragua. A reassessment of the conservation status for many species must be done considering these changes and this is especially true for many species that have not been recorded recently (e.g. M. macrophyllum has not been known since 1969). This updated checklist documents the high biodiversity of Honduran bats and is also an example of how poorly many groups have been studied since first being recorded in the country. We hope to encourage the existing and future generations of researchers to not only report new records and update checklists, but to engage in ecological bat research urgently needed in Honduras.