Zoosyst. Evol. 96 (2) 2020, 411-429 | DOI 10.3897/zse.96.51059 Et iba a0 Pg ga Pg SS _________ae SPENSOFT. pp a NATURKUNDE BERLIN A systematic revision of the bats (Chiroptera) of Honduras: an updated checklist with corroboration of historical specimens and new records Manfredo Alejandro Turcios-Casco!2, Hefer Daniel Avila-Palma?, Richard K. LaVal*, Richard D. Stevens°, Eduardo Javier Ordofiez-Trejo°, José Alejandro Soler-Orellana’, Diego Ivan Ordofiez-Mazier? 1 Departamento de Vida Silvestre, Instituto Nacional de Conservacion y Desarrollo Forestal, Areas Protegidas y Vida Silvestre (ICF), Brisas de Olancho, Francisco Morazan, Honduras 2 Biological Institute, Tomsk State University (TSU), 36 Lenin Ave, Tomsk, Russia 3 Escuela de Biologia, Facultad de Ciencias, Universidad Nacional Autonoma de Honduras (UNAH), Boulevard Suyapa, Francisco Morazadn, Honduras 4 Monteverde Bat Jungle, Monteverde, Costa Rica 5 Department of Natural Resources Management and Natural Science Research Laboratory, Museum of Texas Tech University, Texas, USA http://zoobank.org/695 DASCE-S8 178-45 17-9A83-FABD64AD398C Corresponding author: Manfredo Alejandro Turcios-Casco (manturcios21@gmail.com) Academic editor: Thomas von Rintelen # Received 12 February 2020 # Accepted 13 May 2020 @ Published 22 June 2020 Abstract 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 7rinycteris 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, Chi- roderma gorgasi, Eumops ferox, Gardnerycteris keenani, Lasiurus frantzii, Myotis pilosatibialis, Molossus and Pteronotus species, and Jonatia 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 re- quires 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. Key Words Artibeus intermedius, Central America, Cynomops greenhalli, Diaemus youngii, Natalus lanatus, taxonomy Introduction et al. (2014), 139 species were reported; this is the high- est number of bat species for any Mesoamerican country. Rodriguez Herrera and Sanchez (2015) reported 98 spe- Recent efforts to study bats in the Mesoamerican region cies of bats for Guatemala, 109 for Honduras, 68 for El have increased the number of bat species known. For ex- Salvador, 102 for Nicaragua and 114 for Costa Rica. Inthe ample, Kraker-Castafieda et al. (2016) listed 100 species recent list of bats of Mexico presented by Ramirez-Pulido for Guatemala, Mora et al. (2018) and Turcios-Casco et Copyright Mantredo Alejandro Turcios-Casco et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 412 al. (2020a) increased the list for Honduras to 111 plus four expected species and recently, Medina-Fitoria and Martinez-Fonseca (2019) and Saldafia Tapia et al. (2020) increased the list for Nicaragua to 111 species. In addi- tion, 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 (Rodriguez Herrera and Sanchez 2015), of which over 65.29% occur in Hon- duras. Not only have researchers recorded new species for each country, but they have also carried out systemat- ic and taxonomic studies that have defined new lineages, new species and new taxonomic arrangements. Frequent taxonomic changes, molecular vs. morpho- logical 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 conserva- tion assessments of bat species have been based on his- torical records (Turcios-Casco and Medina-Fitoria 2019). Mora (2016) provided the first taxonomic keys for iden- tification of Honduran bats, so it was no longer neces- sary to rely on keys from other countries [e.g. Costa Rica (Timm et al. 1999) and Mexico (Medellin et al. 2008)]. Such practices led to misidentifications of species endem- ic 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-Castafieda et al. 2016), Honduras has had a questionable number of known bat species since the 1900s. Unlike other megad- iverse countries such as Mexico (Medellin et al. 2008; Ramirez-Pulido et al. 2014) and Costa Rica (LaVal and Rodriguez-H 2002; Rodriguez-Herrera et al. 2014; York et al. 2019), Honduras has had relatively few bat studies and distribution and ecology of most species are still un- known (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) pre- sented the first checklist of mammals of Honduras with 69 bat species including Macrotus macrotus (= M. water- housii Gray, 1843). More records were added to the bat list of Honduras by Davis and Carter (1964), Davis et al. (1964), 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 zse.pensoft.net Turcios-Casco, M.A. et al.: Taxonomy of the bats of Honduras 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). Hernandez (2015) mentioned 112 species based on the efforts done by the Program for Bat Conservation in Honduras (PCMH), but in the final table in Rodriguez Herrera and Sanchez (2015), there were only 109 spe- cies 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 Good- win, 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 Mufioz-Garay (2014) for Myotis pilosatibialis LaVal, 1973 and Pavan and Marroig (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. (2018, 2020) for Molossus E. Geoffroy, 1805. Finally, new re- cords are given based on the verification of museum spec- imens 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 deposit- ed in museums over the world, we re-examined certain vouchers that were misidentified and that we could cor- rectly identify, based on cranial and external measure- ments. These were measured to the nearest 0.01 mm with digital calipers. Measurements followed Srinivasu- lu 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 (Smt); third metacarpal length (3mt); length of the first phalanx Zoosyst. Evol. 96 (2) 2020, 411-429 of digit III (1ph); length of the second phalanx of dig- it If] (2ph); and the length of the third phalanx of digit Il (3ph). Skull measurements obtained from vouchers followed Simmons and Voss (1998), Tejedor (2011) and Giménez and Giannini (2016). These included condyloin- cisive length (CIL), condylocanine length (CCL), zygo- matic 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). We accept the changes of Gardnerycteris Hurtado & Pacheco, 2014 for G. keenani (Handley, 1960) (Hurtado and D’Elia 2018) and Mimon Gray, 1847 for M. cozu- melae Goldman, 1914 (Hurtado and Pacheco 2014). We recognise Lophostoma evotis (Davis & Carter, 1978) as a separate species from L. si/vicolum d Orbigny, 1836 (Da- vis and Carter 1978; Medellin and Arita 1989) and accept the changes proposed by Pavan and Marroig (2016) for Mormoopidae. The two species of Sturnira Gray, 1842 that occur in Honduras are S. hondurensis and S. parv- idens Goldman, 1917 (Velazco and Patterson 2013). We also accept the taxonomic changes of Uroderma biloba- tum Peters, 1866 to U. convexum Lyon, 1902 and U. da- visi Baker and McDaniel, 1972 (Mantilla-Meluk 2014). We recognise the distinction of the two species of Rho- geessa H. Allen, 1866 proposed by Baird et al. (2008, 2012). We treat Aeorestes Fitzinger, 1870 and Dasypterus W. Peters, 1870 proposed by Baird et al. (2015) as sub- genera of Lasiurus Gray, 1831. We recognise Dermanura Gervais, 1856 as a genus separate from Artibeus Leach, 1821, following Hoofer et al. (2008) and York et al. (2019). In addition, we agree with York et al. (2019) in the recognition of A. intermedius supported by Larsen et al. (2013) and currently accepted by York et al. (2019) and Simmons and Cirranello (2020). We also consider Eumops ferox (Gundlach, 1861) as the species that occurs in Central America (McDonough et al. 2008; Baker et al. 2009; Gregorin et al. 2016) and we followed Simmons and Voss (1998) in the recognition of Eptesicus andinus J. A. Allen, 1914, E. brasiliensis (Desmarest, 1819) and E. chiriquinus Thomas, 1920 as different species [see York et al. (2019)]. For the other species, we follow Wilson and Mittermeier (2019). Finally, we reviewed the new records for Honduras following McCarthy et al. (1993) that could be verified as vouchers with museum catalogue numbers, records based on vocalisations found in published articles or new information based on the authors’ data. Acronyms used for the museums are the following: EAPZ Biodiversity Collection, Escuela Agricola Panamericana, Honduras LACM _ Vertebrate Collection, Natural History Muse- um of Los Angeles County, USA MVB Museum of Vertebrate Zoology, Portland State University, USA PSM Vertebrates Collection, Museum of Natural History, University of Puget Sound, USA 413 TCWC _ The Biodiversity Research and Teaching Col- lections (Texas A&M University), USA TTU Museum of Texas Tech University, USA UCR Universidad de Costa Rica, Costa Rica UNAH — Universidad Nacional Autonoma de Hondu- ras, Ciudad Universitaria, Honduras UVS Vertebrate Collection, Universidad Nacional Autonoma de Honduras, Valle de Sula, Hon- duras Results Annotated list of species of the species that occur in Hon- duras, authors and years follow Simmons and Cirranello (2020): Emballonuridae Gervais, 1856 Emballonurinae Gervais, 1856 Balantiopteryx Peters, 1867 1. Balantiopteryx io Thomas, 1904 2. Balantiopteryx plicata Peters, 1867 Centronycteris Gray, 1838 3. Centronycteris centralis Thomas, 1912 Diclidurus Wied-Neuwied, 1819 4. Diclidurus albus Wied-Neuwied, 1819 Peropteryx Peters, 1867 5. Peropteryx kappleri Peters, 1867 6. Peropteryx macrotis (Wagner, 1843) Rhynchonycteris Peters, 1867 7. Rhynchonycteris naso (Wied-Neuwied, 1820) Saccopteryx Mliger, 1811 8. Saccopteryx bilineata (Temminck, 1838) 9. Saccopteryx leptura (Schreber, 1774) Phyllostomidae Gray, 1825 Carolliinae Miller, 1924 Carollia Gray, 1838 10. Carollia castanea H. Allen, 1890 11. Carollia perspicillata (Linnaeus, 1758) 12. Carollia sowelli Baker, Solari, & Hoffmann, 2002 13. Carollia subrufa (Hahn, 1905) Desmodontinae J.A. Wagner, 1840 Desmodus Wied-Neuwied, 1826 14. Desmodus rotundus (E. Geoffroy, 1810) Diaemus Miller, 1906 15. Diaemus youngii (Jentink, 1893) Diphylla Spix, 1823 16. Diphylla ecaudata Spix, 1823 Glossophaginae Bonaparte, 1845 Anoura Gray, 1838 17. Anoura geoffroyi Gray, 1838 Choeroniscus Thomas, 1928 18. Choeroniscus godmani (Thomas, 1903) Choeronycteris Tschudi, 1844 19. Choeronycteris mexicana Tschudi, 1844 Glossophaga E. Geoffroy, 1818 20. Glossophaga commissarisi Gardner, 1962 zse.pensoft.net 414 21. Glossophaga leachii Gray, 1844 22. Glossophaga soricina (Pallas, 1766) HAylonycteris Thomas, 1903 23. Hylonycteris underwoodi Thomas, 1903 Leptonycteris Lydekker, 1891 24. Leptonycteris yerbabuenae Martinez & Villa-R, 1940 Lichonycteris Thomas, 1895 25. Lichonycteris obscura Thomas, 1895 Glyphonycterinae Baker, Solari, Cirranello & Sim- mons, 2016 Glyphonycteris Thomas, 1896 26. Glyphonycteris daviesi (Hill, 1964) 27. Glyphonycteris sylvestris Thomas, 1896 Lonchorhininae Gray, 1866 Lonchorhina Tomes, 1863 28. Lonchorhina aurita Tomes, 1863 Micronycterinae Van Den Bussche, 1992 Micronycteris Gray, 1866 29. Micronycteris hirsuta (Peters, 1869) 30. Micronycteris microtis Miller, 1898 31. Micronycteris minuta (Gervais, 1856) 32. Micronycteris schmidtorum Sanborn, 1935 Phyllostominae Gray, 1825 Chrotopterus Peters, 1865 33. Chrotopterus auritus (Peters, 1856) Gardnerycteris Hurtado & Pacheco, 2014 34. Gardnerycteris keenani (Handley, 1960) Lophostoma @ Orbigny, 1836 35. Lophostoma brasiliense Peters, 1866 36. Lophostoma evotis (Davis & Carter, 1978) 37. Lophostoma silvicolum @’ Orbigny, 1836 Macrophyllum Gray, 1838 38. Macrophyllum macrophyllum (Schinz, 1821) Mimon Gray, 1847 39. Mimon cozumelae Goldman, 1914 Phylloderma Peters, 1865 40. Phylloderma stenops Peters, 1865 Phyllostomus Lacépéde, 1799 41. Phyllostomus discolor Wagner, 1843 42. Phyllostomus hastatus (Pallas, 1767) Tonatia Gray, 1827 43. Tonatia bakeri Williams, Willig & Reid, 1995 Trachops Gray, 1847 44, Trachops cirrhosus (Spix, 1823) Vampyrum Rafinesque, 1815 45. Vampyrum spectrum (Linnaeus, 1758) Stenodermatinae Gervais, 1856 Artibeus Leach, 1821 46. Artibeus inopinatus Davis & Carter, 1964 47. Artibeus intermedius J.A. Allen, 1897 48. Artibeus jamaicensis Leach, 1821 49. Artibeus lituratus (Olfers, 1818) Centurio Gray, 1842 50. Centurio senex Gray, 1842 Chiroderma Peters, 1860 51. Chiroderma gorgasi Handley, 1960 52. Chiroderma salvini Dobson, 1878 zse.pensoft.net Turcios-Casco, M.A. et al.: Taxonomy of the bats of Honduras 53. Chiroderma villosum Peters, 1860 Dermanura Gervais, 1856 54. Dermanura azteca K. Andersen, 1906 55. Dermanura phaeotis (Miller, 1902) 56. Dermanura tolteca (Saussure, 1860) 57. Dermanura watsoni Thomas, 1901 Ectophylla H. Allen, 1892 58. Ectophylla alba H. Allen, 1892 Enchisthenes K. Andersen, 1906 59. Enchisthenes hartii (Thomas, 1892) Platyrrhinus Saussure, 1860 60. Platyrrhinus helleri (Peters, 1866) Sturnira Gray, 1842 61. Sturnira hondurensis Goodwin, 1940 62. Sturnira parvidens Goldman, 1917 Uroderma Peters, 1865 63. Uroderma convexum Lyon, 1902 64. Uroderma davisi Baker & McDaniel, 1972 65. Uroderma magnirostrum Davis, 1968 Vampyressa Thomas, 1900 66. Vampyressa thyone Thomas, 1909 Vampyriscus Thomas, 1900 67. Vampyriscus nymphaea (Thomas, 1909) Vampyrodes Thomas, 1900 68. Vampyrodes major Allen, 1908 Mormoopidae Saussure, 1860 Mormoops Leach, 1821 69. Mormoops megalophylla (Peters, 1864) Pteronotus Gray, 1838 70. Pteronotus fulvus (Thomas, 1892) 71. Pteronotus gymnonotus (J.A. Wagner, 1843) 72. Pteronotus mesoamericanus Smith, 1972 73. Pteronotus psilotis (Dobson, 1878) Noctilionidae Gray, 1821 Noctilio Linnaeus, 1766 74. Noctilio albiventris Desmarest, 1818 75. Noctilio leporinus (Linnaeus, 1758) Thyropteridae Miller, 1907 Thyroptera Spix, 1823 76. Thyroptera tricolor Spix, 1823 Natalidae Gray, 1866 Natalus Gray, 1838 77. Natalus lanatus Tejedor, 2005 78. Natalus mexicanus Miller, 1902 Molossidae Gervais, 1856 Molossinae Gervais, 1856 Cynomops Thomas, 1920 79. Cynomops greenhalli Goodwin, 1958 80. Cynomops mexicanus (Jones & Genoways, 1967) Eumops Miller, 1906 81. Eumops auripendulus (Shaw, 1800) 82. Eumops ferox (Gundlach, 1861) 83. Eumops hansae Sanborn, 1932 84. Eumops nanus (Miller, 1900) 85. Eumops underwoodi Goodwin, 1940 Molossus ¥. Geoffroy, 1805 86. Molossus alvarezi Gonzalez-Ruiz, Ramirez-Pu- lido & Arroyo-Cabrales, 2011 Zoosyst. Evol. 96 (2) 2020, 411-429 87. Molossus aztecus Saussure, 1860 88. Molossus bondae J.A. Allen, 1904 89. Molossus molossus (Pallas, 1766) 90. Molossus nigricans Miller, 1902 Nyctinomops Miller, 1902 91. Nyctinomops aurispinosus (Peale, 1848) 92. Nyctinomops laticaudatus (E. Geoffroy, 1805) 93. Nyctinomops macrotis (Gray, 1840) Promops Gervais, 1855 94, Promops centralis Thomas, 1915 Tadarida Rafinesque, 1814 95. Tadarida brasiliensis (1. Geoffroy, 1824) Vespertilionidae Gray, 1821 Myotinae Tate, 1942 Myotis Kaup, 1829 96. Myotis albescens (E. Geoffroy, 1806) 97. Myotis elegans Hall, 1962 98. Myotis nigricans (Schinz, 1821) 99. Myotis pilosatibialis LaVal, 1973 100. Myotis riparius Handley, 1960 101. Myotis velifer (J. A. Allen, 1890) Vespertilioninae Gray, 1821 Bauerus Van Gelder, 1959 102. Bauerus dubiaquercus (Van Gelder, 1959) Eptesicus Rafinesque, 1820 103. Eptesicus brasiliensis (Desmarest, 1819) 104. Eptesicus furinalis (d’ Orbigny, 1847) 105. Eptesicus fuscus (Beauvois, 1796) Lasiurus Gray, 1831 106. Lasiurus cinereus (Palisot de Beauvois, 1796) 107. Lasiurus ega (Gervais, 1856) 108. Lasiurus egregius (Peters, 1870) 109. Lasiurus frantzii Peters, 1870 110. Lasiurus intermedius H. Allen, 1862 Perimyotis Menu, 1984 111. Perimyotis subflavus (F. Cuvier, 1832) Rhogeessa H. Allen, 1866 112. Rhogeessa bickhami Baird, Marchan-Rivade- neira, Perez & Baker, 2012 113. Rhogeessa menchuae Baird, Marchan-Rivade- neira, Perez & Baker, 2012 Specific remarks Artibeus Leach, 1821. Recently, Portillo-Reyes et al. (2019) reported a presumed roosting site of approximate- ly 50 individuals which they identified as A. inopinatus (Fig. 1A, B) in southern Honduras, but no criteria for identification were provided. These individuals might have been misidentified considering the difficulty of iden- tification of the four species of large Artibeus in Hondu- ras. Moreover, Davis (1970) mentioned that the subspe- cies (e.g. Artibeus jamaicensis paulus Davis, 1970) that occurs on the Pacific versant was smaller than those from the Atlantic versant in Honduras, further complicating identification. However, Portillo-Reyes et al. (2019) not- ed that the establishment of an Area de Importancia para 415 la Conservacion de Murcielagos (AICOM) is warranted on Isla del Tigre for the conservation of A. inopinatus. This is based on three unofficial records in 2016 by the PCMH (Programa de Conservacion de Murciélagos en Honduras). Hernandez (2015) suggested A. inopinatus be considered an endangered species in Honduras, spe- cifically 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 south- ern 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 (Medellin 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 Nic- aragua. 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. in- opinatus has been considered a rare species (see Turci- os-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 respec- tively 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 Morazan (Carboneras, Saba- nagrande). To our knowledge, the individual captured in San Buenaventura in Francisco Morazan by Turcios-Cas- co et al. (2020c) is the highest elevational record for the species which was captured at 1435 ma.s.1. The case of Artibeus (sensu lato) has been controver- sial and York et al. (2019) mentioned an easy distinguish- ing 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. in- termedius represent individuals of A. /. 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. 1. 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 Marchan-Rivadeneira et al. (2012) provided indirect evidence of genetic isolation of A. intermedius by iden- tifying sympatric A. intermedius and A. I. 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. /ituratus 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 zse.pensoft.net 416 Turcios-Casco, M.A. et al.: Taxonomy of the bats of Honduras Figure 1. Artibeus inopinatus. A — Adult male captured in Carboneras, Sabanagrande in the department of Francisco Morazan (center of Honduras). B— Adult female captured in Nagarejo, Nacaome in the department of Valle (southern Honduras). size-class (referring to the formerly A. /ituratus) and a small size-class (referring to what he hypothesised to be A. intermedius) from Santa Barbara (western Honduras) and Brus Laguna (eastern Honduras). Davis (1984) iden- tified specimens of A. intermedius from the following de- partments of Honduras: Choluteca and Valle in southern Honduras; Copan, Intibuca, La Paz, Ocotepeque and San- ta Barbara in western Honduras; Francisco Morazan and Comayagua in Central Honduras; and Gracias a Dios in eastern Honduras. Additionally, in the GBIF.org (2019), there are preserved specimens from Atlantida, Colon 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 Paraiso, Yoro, Olancho and Islas de la Bahia. Finally, Simmons and Cirranello (2020) recognised A. intermedi- us as a distinct species from A. /ituratus. 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 repre- sent, 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 Rodriguez Herrera and Sanchez (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 assump- tions. The only unofficial record was made by Hernandez et al. (2016) in which vocalisations of C. centralis were recorded in the Cuyamel-Omoa National Park in the de- zse.pensoft.net partment of Cortés, but without a precise location (coor- dinates and elevation). No further information 1s known of the species in the country. Chiroderma gorgasi Handley, 1960. The first record of C. gorgasi (UVS—V—02529) for Honduras was collect- ed 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 Comaya- gua as the first record of Molossops greenhalli Good- win & 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. Cur- rently, Simmons and Cirranello (2020) recognised C. greenhalli to be distinct from C. mexicanus. Erroneously, the species that had been previously rec- ognised in Honduras was C. mexicanus [e.g. Hernandez (2015), Mora (2016), Mora et al. (2018)], but to date, the only specimen of Cynomops 1s the female captured in the Zoosyst. Evol. 96 (2) 2020, 411-429 417 Table 1. Comparison of the new record of C. mexicanus with the record of 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 mea- surements are in mm. Cynomops mexicanus (TTU 104070), al. (2018) new record Sex 1 female 1 male 5 females FA 36.71 37.05 35.08 (33.02-36.3) GLS 18.90 18.78 17.02 (15.87-17.57) CIL 17.02 19.39 16.87 (16.05-17.19) PB 4.07 4.73 4.55 (4.23-4.80) ZB 10.75 13.20 11.76 (11.32-12.00) BB 7.51 9.18 8.75 (8.65-8.92) MB 8.47 34-6 11.21 (10.71-11.41) MTL 5.80 7.62 6.79 (6.38-6.92) 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 mas.l., in the department of Atlantida (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, 1n the SAG, La Ceiba, in the department of Atlantida in northern Hondu- ras (not Colon, 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 (in mm) of specimen TTU 104070 are TL = 93.0, T = 26.0, HF = 6.0, Th = 4.11, CCL = 16.54 and MTL = 7.93. Diaemus youngii (Jentink, 1893). Amongst the three haematophagous species, D. youngii (Fig. 3A—C) was the only species never officially recorded from Honduras. We now document its occurrence in the country. During a survey in November 2018, in Finca Don Richard in Tru- jillo, in the department of Colon (15.916N, 85.934W, Fig. 7), we captured an adult female at 22:00 h along with individuals of A. jamaicensis, A. lituratus, Desmodus ro- tundus (E. Geoffroy, 1810), Micronycteris microtis (Mill- er, 1898) and S. parvidens. Unfortunately, the individu- al escaped after chewing the bag in which it was held. However, the individual was already identified and its morphometric data and photographs had been taken: FA = 55.40; E = 15.75; Tr = 7.20; TrW = 2.60; Th = 11.70; Tib = 24.60; HF = 15.95; HB = 73.80; Hu = 53.10; 1ph = 11.50; 2ph = 28.65; 3ph = 22.05; WS = 450.2 mm. The individual was identified as D. youngii following Timm et al. (1999), Reid (2009) and Mora (2016): 1) ears tri- angular; facial and dorsal colour grey brown and lightly frosted; ventrum whitish (Fig. 3A), uropatagium reduced and hairy; and submandibular glands present as men- tioned by Medina-Fitoria (2014); 2) wingtips white (Fig. Cynomops mexicanus by Moras et Cynomops greenhalli Cynomops greenhalli by Moras et al. (TCWC 22123) by LaVal ( (1969) 1 female 11 males 22 females 35.7 37.41 (35-39.7) 35.36 (33.40-38.28) LOF3 18.41 (17.37-19.23) 17.12 (15.91-17.82) - 18.68 (17.92-19.5) 16.93 (16.06-17.68) 4.8 4.78 (4.53-5.02) 4.63 (4.29-4.97) 12.6 12.77 (12.06-13.65) 11.85 (11.23-12.51) - 9.24 (8.68-9.65) 8.86 (8.20-9.25) 12.1 12.26 (11.9-13.01) 11.24 (10.49-11.86) Fi2 7.18 (6.79-7.67) 6.61 (6.22-7.05) 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) re- viewed the E. brasiliensis complex, he mentioned that there is a population in the highlands of Middle America that appears to be identical to £. andinus from the high- lands 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 FE. 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 EF. andinus and Koopman (1978, 1993, 1994) considered E. andinus as a subspe- cies to E. brasiliensis and E. chiralensis H. E. Anthony, 1926 and E. montosus to E. furinalis (d’Orbigny, 1847). Simmons and Voss (1998), in accordance with Davis (1966), concluded that FE. inca and E. chiriquinus are conspecific and selected chiriquinus as the senior syno- nym 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 Bo- livia northwards at upper elevations along the Andes of Peru, Ecuador and Colombia, into north-western Vene- zuela 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 east- ern Ecuador, Peru and Bolivia and elsewhere in Central America north-westwards into the Chiapan Highlands of Mexico. While reviewing the systematics of the E. andi- nus 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 Atlantida (not Colon, as is incorrectly recorded on the original label) in Honduras. We corroborate the identifi- cation 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), zse.pensoft.net 418 2 3 PNT TTT Figure 2. A ventral, B lateral and C dorsal views of the skull of C. mexicanus (TTU 104070) captured in the SAG, La Ceiba in the deparment of Atlantida in northern Honduras. the key to Costa Rican and Nicaraguan bats (York et al. 2019) and the keys for Eptesicus in South America by Davis and Gardner (2008). There is no clarification that the specimen of E. andinus [collected at approximately 1.8 miles (ca. 1.6 km) west in Ixhuatan, Chiapas, Mexi- co by M. D. Tuttle in 1962 (AMNH 203916)] mentioned by Davis (1966) and available in the GBIF.org database (Trombone 2016) corresponds to that species [see Ar- royo-Cabrales et al. (2008)]. This individual supports the supposed distribution of E. chiriquinus to Chiapas, Mex- ico. Arroyo-Cabrales et al. (2008) mentioned a personal communication of R. A. Medellin in 1998 stating that one individual of E. furinalis gaumeri (J. A. Allen, 1897) from Jalcocotan, captured in the field in January of 1977, zse.pensoft.net Turcios-Casco, M.A. et al.: Taxonomy of the bats of Honduras belongs to E. chiriquinus according to the shape of the interorbital region. Recently, there 1s personal comment by A. Gardner in Reid (2009) regarding the distribution of E. chiriquinus, in which the species may occur in the highlands of Chiapas. However, the specimen AMNH 203916 must be verified to determine which species 1s the one that occurs in Mexico. We identified the specimen TTU 104074 as E. furinalis and not E. andinus following Davis and Gardner (2008) and York et al. (2019), because 1) the FA was 39.27 mm; 2) dorsal fur was 7.0 mm with a pale greyish-brown with black basal band; 3) it was collected at low elevations; 4) GSL > 15.00 mm; and 5) MDTL > 5.40 mm. E. brasiliensis was not included by McCarthy et al. (1993) for Central America and there is no voucher in the GBIF.org (2019) that supports its occurrence in Hondu- ras. Nevertheless, the species is included on the bat list for Honduras by Rodriguez Herrera and Sanchez (2015) and Mora et al. (2018). Espinal and Mora (2012b) record- ed a female of EF. brasiliensis captured in El Corpus, de- partment of Choluteca (southern Honduras), which was preserved, but unfortunately, it is part of a personal col- lection and is not deposited as a voucher in any museum. Identification was supported by vocalisations recorded to individuals of £. 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. In conclusion, there are only three species of Eptesi- cus (E. brasiliensis, E. furinalis and E. fuscus Beauvois, 1796) reported for Honduras and, even though Davis and Gardner (2008) stated that EF. 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 Rodriguez Herrera and Sanchez (2015) and Mora et al. (2018) in the bat lists of Honduras. McCarthy et al. (1993) mentioned that D. C. Carter cap- tured 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, ob- served it as 1t escaped after chewing a hole in the bag in which it was held. Fortunately, the bat was already ten- tatively identified by D. C. Carter and photographed by LaVal [stored in the Mammal Slide Library (no. 378) of the American Society of Mammalogists]. McCarthy et al. (1993) also mentioned six individuals [one male and five females: FA = 41.9 mm (41.1-43.3 mm)] of G. sy/vestris Thomas, 1896 (previously referred as Micronycteris syl- vestris) captured in Laguna de Bacalar in the department of Gracias a Dios by B. H. Gaskell on 7 October 1982. Unfortunately, specimens were not retained and besides the locality, measurements taken of individuals are the Zoosyst. Evol. 96 (2) 2020, 411-429 419 Figure 3. Diaemus youngii. A— Frosted brownish venter; B — Whitish tip of the wings; C — One pad on the base of the thumb. only information we have for the species in Honduras. These sparse records are all from the Caribbean Low- lands in the north-eastern part of the country. HAylonycteris underwoodi Thomas, 1903. The occur- rence of H. underwoodi has been debated, because there are no vouchers that can confirm presence in Honduras. Turcios-Casco and Medina-Fitoria (2019) confirmed oc- currence of the species in Honduras, based on an adult female (UVS—V—02527) captured in the department of Gracias a Dios in eastern Honduras. Lasiurus Gray, 1831. Mora and Lopez (2013) de- scribed the first record of L. cinereus (Palisot de Beau- vois, 1796) (EAPZ—06), based on a male specimen found dead in Cerro de Hula in Santa Ana municipality in south- ern Francisco Morazan 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 Mar- cos de Colon, in the department of Choluteca in south- ern 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 Cata- camas, department of Olancho (north-eastern Honduras). These records cover a gap between Guatemala and Pana- ma (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 and not L. blossevillii (Lesson & Garnot, 1826), as is mentioned by Mora et al. (2018). Leptonycteris yerbabuenae Martinez and Villa-R, 1940. We followed Cole and Wilson (2006a, 2006b), when recognising occurrence in Honduras of L. yerb- abuenae. This distinction is well supported by published data (Wilkinson and Fleming 1996), in which L. curas- oae Miller, 1900 and L. yerbabuenae separated approx- imately 540,000 years ago. In 1991, R. D. Bradley re- ported two specimens (TTU 61087, TTU 61088) from Nacaome, in the department of Valle. Lee and Bradley (1992) reported three additional individuals [(TCWC 49747-49749) (GBIF.org 2019, Fig. 7). More recently in 1995, M. Sandiford reported one more individual (ZD 1999.194) from Yusguare, in the department of Cholute- ca (GBIF.org 2019, Fig. 7). Even though there are two records in the GBIF.org database, in the catalogue pre- sented 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. Hernandez (2015) proposed the creation of the Area de Importancia para la Conservacion de Murcielagos (AI- COM) Golfo de Fonseca for the conservation of L. yerb- abuenae. Its rarity in Central America is one of the rea- sons that this species is considered as Near Threatened (NT) by the IUCN. In Table 2, we give cranial measure- ments for Bradley’s records. Lonchorhina Tomes, 1863, Mimon Gray, 1847, Phyl- loderma Peters, 1865 and Tonatia Gray, 1827. A sev- enth locality of L. aurita Tomes, 1863 (UVS—V—02067, UVS-V-02075) was reported by Avila-Palma et al. (2020) in Sabanagrande in the department of Francisco Morazan. Avila-Palma et al. (2019) reported MM. cozume- lae (UVS—V—02059) at two sites in the core of the Rio Platano Biosphere Reserve in the department of Colon. These records represent the fifth and sixth localities of this species in Honduras and the first record since 2001. A new record of Phylloderma stenops (UVS—V-—02526) was obtained after 46 years by Turcios-Casco et al. (2020b) in the Caribbean Lowlands of the department of Gracias a Dios in eastern Honduras. Regarding 7onatia, we follow Basantes et al. (2020). The species that occurs in Hon- duras 1s T. bakeri Williams, Willig & Reid, 1995 and 7 saurophila Koopman & Williams, 1951 is only known, based on subfossil remains found in caves of the type lo- cality in Jamaica. zse.pensoft.net 420 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 de- partment 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 coun- try for the occurrence of M. bondae and consequently Burnett et al. (2001), Reid (2009), Hernandez (2015), Mora (2016) and Mora et al. (2018) all listed MZ. bondae as occurring in Honduras. Additionally, Lopez-Gonzalez 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 suffi- ciently similar that they considered them to be conspecif- ic, based on morphometric data. Eger (2008) believed that M@ bondae and M. curren- tium were different species due to differences in colour- ation and fur length. However, we follow Loureiro et al. (2020) as the most recent taxonomy for Mo/ossus, based on Dolan (1989) and Gonzalez-Ruiz et al. (2011). We rec- ognise 14 species of Mo/lossus, 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 La- guna in the department of Gracias a Dios in north-eastern Honduras (Dolan and Carter 1979; Dolan 1989; Lopez- Gonzalez 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 Rio Coco, in the department of El Paraiso 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. alva- rezi Gonzalez-Ruiz, Ramirez-Pulido & Arroyo-Cabrales, Table 2. Cranial measurements (in mm) of the two individu- als of L. yerbabuenae recorded in 1991 by R. D. Bradley et al. in the department of Valle, southern Honduras (Garner 2016b, 2016c). The records of Cole and Wilson (2006a) are from south- ern Arizona and Mexico (means before parentheses and ranges are included on them). TCWC TCWC Cole and Wilson Cole and Wilson 49749 49748 (2006a) (2006a) Sex Male Male 10 males 10 females GLS 26.68 DTS 26.8 (26.0-27.3) 26.8 (26.4-27.3) CIL 26.14 27.04 - - PB 5.35 5:25 - - ZB 10.75 11.16 11.1 (10.7-11.5) 10.8 (10.6-10.9) BB 9.66 10.39 - - MB 10.85 11.06 10.3 (10.1-10.6) 10.3 (10.1-10.5) MBL 8.86 8.72 9.6 (9.2-9.8) 9.5 (9.0-9.9) CGL 25.09 25.69 - - MDTL 9.46 9.86 - - zse.pensoft.net Turcios-Casco, M.A. et al.: Taxonomy of the bats of Honduras 2011. There has not been any doubt about the occurrence of M. molossus (Pallas, 1766) in the country. On the oth- er 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 west- ern Honduras. Additionally, Loureiro et al. (2020) and Simmons and Cirranello (2020) consider its occurrence in Jalisco and Cozumel in Mexico to Nicaragua, south- ern Venezuela, and south-eastern Brazil. Finally, M. ni- gricans Miller, 1902 is the species that occur in Mexico and Central America and M. rufus E. Geoffroy, 1805 is restricted to South America including Guianas, Ecuador, Peru, to central and northern Brazil and Bolivia, and Trin- idad and Tobago (Loureiro et al. 2020; Simmons and Cir- ranello 2020). Myotis pilosatibialis LaVal, 1973. Mantilla-Meluk and Mufioz-Garay (2014) elevated MM. keaysi pilosatibi- alis to species and they included new discrete characters distinguishing M. pilosatibialis from M. keaysi J. A. Al- len, 1914 which is considered a similar species. In agree- ment with York et al. (2019), we consider M. pilosatibial- is (Holotype: LACM 36879) to be the species that occurs in Central America. The type locality is a small cave 1 km west of Talanga, in the department of Francisco Morazan (750 m a.s.I.) in central Honduras (LaVal 1973). Accord- ing to LaVal (1973), he collected the holotype and the large series of topotypes from a cluster of several hundred bats in a small domed room not far from the entrance of the cave. This species is known from few Honduran lo- calities besides the type locality, based on the GBIF.org (2019): 8.5 miles (ca. 12.9 km) south-west San Lorenzo in the department of Valle (southern Honduras); 49.89 km north in the department of Santa Barbara (western Honduras); and at 31.8 miles (ca. 50 km) east southeast of Cuyamel, Santo Domingo in the department of Cortés (northern Honduras). Notably, LaVal (1973) mentioned that almost all specimens from Honduras were from ele- vations of 1000 to 2000 m a.s.l. Natalus lanatus Tejedor, 2005 and N. mexicanus Miller, 1902. Tejedor (2011) examined two specimens from Honduras and identified them as N. mexicanus, one captured in La Tigra National Park, in the department of Francisco Morazan (TTU 83664) in Central Hondu- ras and the other at 12 km north of Santa Barbara (TTU 13418) in western Honduras [this record was considered as a complex by Turcios-Casco et al. (2019a), because N. mexicanus was previously included in N. stramineus Gray, 1838]. Recently, Miller (2014) reported individuals of N. mexicanus from Honduras, but no vouchers were collected. N. /Janatus was considered an expected species in Honduras (Mora 2016; Mora et al. 2018), based on re- cords in Mexico (Tejedor 2011), Nicaragua (Medina-Fi- toria et al. 2015) and Costa Rica (Rodriguez Herrera et al. 2011). The occurrence of N. /anatus in Honduras is based on two historical specimens that we re-examined and identified as N. /anatus (TCWC 10992 and TCWC 11008), respectively, captured by J. R. Meyer in 1963 at 6 miles (ca. 9.6 km) north of Zamorano, Francisco Morazan Zoosyst. Evol. 96 (2) 2020, 411-429 , and by J. V. Mankins, at 2 mi (ca. 3.2 km) W Cueva del Viejo, La Paz in 1963 (Fig. 7). We identified the two individuals (TCWC 10992 and TCWC 11008) as N. /anatus following Tejedor (2011): (1) bicoloured ventral pelage (Fig. 4C); (2) hairy legs and feet with ungual tufts [Fig. 5A, see comparison with feet of N. mexicanus (Fig. 5B of individual TCWC 19698)]; (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) mar- gins of the ears were straight. See Table 3 for comparison with individuals mentioned by Tejedor (2011), Rodriguez Herrera et al. (2011) and Medina-Fitoria et al. (2015). These two specimens confirm the presence of N. /anatus in Honduras and help fill the gap between Mexico and Nicaragua. N. /anatus is the second species of the genus known for the country, although it was expected by Mora et al. (2018). Nyctinomops Miller 1902. McCarthy et al. (1993) in- cluded only WN. laticaudatus (E. Geoffroy, 1805) for Hon- duras, based on a record of a female [TCWC 19759 (Pre- stridge 2019a] captured by LaVal on the Aguan River in the department of Yoro (northern Honduras) and consti- tutes 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 Colon and El Corpus in the department of Choluteca (southern Honduras). Addition- A Figure 4. A Lateral and B ventral views of the skull of Natalus lanatus (TCWC 10992) captured in Zamorano in department of Francisco Morazan, August 1963. C Bicolor ventral pelage of N. /anatus of the same specimen. 421 ally, Espinal et al. (2016) reported a dead male of N. au- rispinosus (Peale, 1848) also from San Marcos de Colon. Unfortunately, even though they gave morphometric in- formation 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 de- posited 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 Intibuca, which is over 1000 ma.s.1. Following Velazco and Patter- son (2013), the only two species that occur in Honduras are S. parvidens and S. hondurensis. Thyroptera Spix, 1823. Goodwin (1942) considered the type locality of 7. discifera (Lichtenstein & Peters, 1854) as Puerto Caballos, department of Cortés, Hon- duras. However, Wilson (2008) clarified that, although many researchers assumed the type locality of T dis- cifera (Fig. 6A, B) to be in Honduras, the correct type locality is Puerto Cabello, Carabobo, Venezuela. See Turcios-Casco and Medina-Fitoria (2019) for a detailed occurrence of 7: tricolor Spix, 1823 (UVS—V—02525, UVS-V—02532, UVS—V—02533) in Honduras, which re- zse.pensoft.net 422 Turcios-Casco, M.A. et al.: Taxonomy of the bats of Honduras LQ LJ Figure 6. A — Roost of Thyroptera discifera in a (Musaceae) from the Caribbean lowlands of Costa Rica. B— View of the colony 7: discifera inside the leaves of Musa x acuminata. Photos by Bernal Rodriguez. cently is only known in the department of Gracias a Dios _ exploitation of resources, changes in land use and live- (eastern Honduras). There are historical records in the — stock grazing (Turcios-Casco and Medina-Fitoria 2019; departments of Francisco Morazan (Goodwin 1942) and __ Larsen 2019). As York et al. (2019) included 7° discifera Cortés (Hall 1981). The distribution of 7’ tricolor in Hon- — in Nicaragua and Costa Rica, we can expect to encounter duras has likely been affected by deforestation, excessive _ this species in Honduras. More sampling effort is needed zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 411-429 A423 Table 3. Comparison of N. /anatus specimens that comprise new records for Honduras with individuals captured by Tejedor (2011) in Mexico, Nicaragua (Medina-Fitoria et al. 2015), and Costa Rica (Rodriguez Herrera et al. 2011). Means (ranges in parentheses); all the measurements are in mm. TCWC 10992 TCWC 11008 Individuals mentioned by Tejedor (2011) Rodriguez Herrera et al. Medina-Fitoria et al. (2011) (2015) Sex M F females males male undetermined FA 38 40 36.8 (35.4-38.6) 37.0 (35.4-38.3) 39.0 36.5-37 Ti 19 17 16.5 (15.9-17.3) 16.8 (15.5-18.4) - - 3mt 34 35 32.6 (31.2-33.9) 33.0 (32.0-33.8) - - 5mt 36 37 34.2 (33.2-35.5) 34.2 (33.2-34.9) - - F 15 14 13.9 (13.0-15.3) 14.1 (12.0-15.6) 16.0 14.5-14.5 GSL 16.5 16.2 15.8 (15.3-16.2) 16.0 (15.4-16.4) - - ZB 8.4 8.3 8.1 (7.9-8.3) 8.2 (7.8-8.7) - - BB 7.9 8.0 7.7 (7.6-7.9) 7.9 (7.5-8.2) - - BAM 5,7 5.5 5.4 (5.2-5.5) 5.4 (5.2-5.6) - - BAC 3.9 4.2 3.5 (3.4-3.6) 3.6 (3.5-3.8) - - MTL 6.9 6.7 6.5 (6.3-6.8) 6.7 (6.4-6.9) - - MBL 7.0 6.8 6.9 (6.7-7.2) 7.1 (6.9-7.4) - - PB 3.4 3.4 3.2 (3.1-3.3) 3.2 (3.1-3.3) - - Legend Species @ Diaemus youngii HS Natalus lanatus GS Leptonycteris yerbabuenae Caribbean Sea 16°N Nob Political Divisions (_] Departments of Honduras @ Central America Elevation Model (m a.s.|.) 14°N Nob Datum WGS84 Source: SINIT Pacific Ocean QGIS 3.4.1 "Madeira" November, 2019 0 88°W 86°W 84°W Figure 7. Map with new records of D. youngii and N. lanatus for Honduras. None of the previous records (except D. youngii) have exact coordinates, however, we shaded the municipality on each department in which they were recorded. The blue shaded area indicates the localities of N. /anatus captured in the municipality of Zamorano (municipality of San Antonio de Oriente) in the department of Francisco Morazan in central Honduras, and in La Cueva del Viejo in the department of La Paz in western Hon- duras. The big yellow-shaded area refers to two localities in which L. yerbabuenae was recorded in the municipality of Nacaome in the department of Valle in southern Honduras by Lee and Bradley (1992) and R. D. Bradley in 1991 (GBIF.org 2019). The small brown-shaded areas in southern Honduras refer to the localities in which M. Sandiford reported one L. yerbabuenae from the municipality of Santa Ana de Yusguare in the department of Choluteca (GBIF.org 2019). Finally, the purple point refers to the exact location of D. youngii recorded in the department of Colon in northeastern Honduras. Abbreviations as follows: La Paz (LP), Nacaome (NC), San Antonio de Oriente (SAO), and Santa Ana de Yusguare (SAY). There is no specific area highlighted in La Paz because there are several caves known as Cueva del Viejo. to study roosting sites of bats in general. For example, 7. tricolor probably uses banana leaves, as they do in Cos- ta Rica, as well as Heliconia leaves, which are abundant plants in wet forests and disturbed and secondary forests (Rodriguez et al. 2007). Vampyriscus nymphaea (Thomas, 1909). Mora et al. (2014) confirmed the presence of this species, based on a pregnant female (EAPZ 72) from the Coco River in Nue- va Esperanza, in the Patuca National Park, in the depart- ment of Olancho (north-eastern Honduras). zse.pensoft.net 424 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 stud- ied (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 spe- cies (York et al. 2019) and above Nicaragua, with 111 species considering Medina-Fitoria and Martinez-Fonse- ca (2019) and Saldafia Tapia et al. (2020). We augmented the most recent bat list of Hondu- ras (Mora et al. 2018) by confirming the presence of D. youngii and C. greenhalli and with the new record of N. lanatus. The species that occur in Honduras, based on new taxonomic arrangements in addition to Mora et al. (2018), are A. intermedius, C. gorgasi, C. greenhalli, E. ferox, G. keenani, L. frantzii, M. pilosatibialis, M. alva- rezi, M. nigricans, P. fulvus, P. psilotis and T: bakeri. The seven expected species, based on occurrence in neigh- bouring countries and consideration of previous authors, are Cormura brevirostris (Wagner, 1843), L. brachyotis, Mesophylla macconnelli Thomas, 1901, M. coibensis J. A. Allen, 1904, M. pretiosus Miller, 1902, 7: discifera and Trinycteris nicefori (Sanborn, 1949). The occurrence of E. bonariensis 1n 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 Ameri- ca. Although Jones et al. (1977) and Simmons (2005) have suggested that FE. perotis (Schinz, 1821) occurs in Hon- duras, we remain unconvinced, because there are no ver- ified records from any Central American country. We did not consider the occurrence of MZ. coibensis in Honduras, Mora et al. (2018) considered it in southern Honduras, be- cause 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 Hon- duras, 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 Hon- duras, 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. zse.pensoft.net Turcios-Casco, M.A. et al.: Taxonomy of the bats of Honduras 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. macrophyl- Jum and V. major. We strongly recommend a revision of the threatened species list proposed by Hernandez (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. syl- vestris 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, 2017) or of using Dermanura for some species for- merly in the genus Artibeus (Cirranello et al. 2016; Sim- mons 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. /ituratus. Based on the re- vision of Davis (1984), the occurrence of A. intermedius in Honduras is well supported. Another species, formerly considered rare, was A. inopinatus (Reid 2009; Medina and Reid 2016; Portillo-Reyes et al. 2019). Nonetheless, between 1966 and 2001, there were 422 records of A. in- opinatus for Honduras [not 454 records as erroneously mentioned by Portillo-Reyes et al. (2019)] (Turcios-Cas- co et al. 2020c); in fact, there are 193 just at TTU. In ad- dition, there are 21 for El Salvador and 11 for Nicaragua in the database of GBIF.org (2019). This suggests that the Species is relatively common in several areas of Hondu- ras, although there has been little sampling reported for Honduras since 2001. For a recent example, in the depart- ment of Francisco Morazan, Sabanagrande [also an im- portant site for the conservation of L. aurita (Avila-Palma et al. 2020)], A. inopinatus comprised 64% of one night’s catch in August 2018. However, studies are needed to de- termine the conservation status of the species and mor- phometric and systematic studies are necessary to solve the identification problem amongst species of Artibeus. 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 identifica- tion needs to be verified. More sampling is needed in cer- tain areas of the country, especially in western Honduras (e.g. La Paz, Intibuca) 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 1s espe- cially true for many species that have not been record- ed 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 Zoosyst. Evol. 96 (2) 2020, 411-429 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. Acknowledgements We are grateful to Gary Shugart for reviewing the Sturni- ra of the PSM Vertebrate Collection; to Eric Van Den Berghe for receiving specimens of the Escuela Agricola Panamaericana (EAP); to Alex Vallejo for authorising the examination of specimens in the Vertebrate Collection, Universidad Nacional Autonoma de Honduras, Valle de Sula, Honduras; to the private collection and rescue cen- tre El Ocotal for all their support for bat conservation; to the PCMH for their efforts in the conservation of bats in Honduras. To the Instituto Nacional de Conservacion y Desarrollo Forestal, Areas Protegidas y Vida Silvestre (ICF) for investigation and research permits (Resolucion DE-—MP-020-17, Resoluci6n—DE—MP-64—2017) and to the Protected Areas department, because the record of the D. youngii was caught during a survey funded by them and coordinated by Felix E. Mancia. Our gratitude also to Silvia Rodriguez and José Luis Ramos for their field assistance. We want to thank Don Israel and Don Richard for their great hospitality and support during the fieldwork in Don Richard’s Finca. Finally, to Nancy B. Simmons, Heather York, Thomas von Rintelen and Mike Skinner who revised our manuscript and improved it with import- ant comments and observations. References Allen H (1892) Description of a new genus of phyllostome bats. Pro- ceedings of the United States National Museum 15(913): 441-442. https://doi.org/10.5479/si.00963801.15-913.441 Allen H (1898) The skull and teeth of Ectophylla alba. Transactions of the American Philosophical Society, New Series 19(2): 267—272. https://doi.org/10.2307/1005455 Arroyo-Cabrales J, Polaco OJ, Wilson DE, Gardner AL (2008) Nue- vos registros de murci¢lagos para el estado de Nayarit, México. Revista Mexicana de Mastozoologia 12(1): 141-162. https://doi. org/10.22201/1e.20074484e.2008.12.1.50 Avila-Palma HD, Turcios-Casco MA, Ordofiez Bautista DJ, Martinez M, Ordofiez-Mazier DI (2019) First records of Mimon cozumelae Goldman, 1914 (Chiroptera, Phyllostomidae) in the Rio Platano Biosphere Reserve in northeastern Honduras. Check List 15(6): 1113-1118. https://doi.org/10.15560/15.6.1113 Avila-Palma HD, Turcios-Casco MA, Velasquez A (2020) The Tomes’s Sword-nosed bat (Lonchorhina aurita) in Honduras, with new re- cords in Sabanagrande, Francisco Morazan. Bat Research News 61(1): 1-6. Baird AB, Braun JK, Engstrom MD, Holbert AC, Huerta MG, Lim BK, Mares MA, Patton JC, Bickham JW (2017) Nuclear and mtDNA phylogenetic analyses clarify the evolutionary history of two species of native Hawaiian bats and the taxonomy of Lasiurini (Mammalia: 425 Chiroptera). PLoS ONE 12(10): 1—27. https://doi.org/10.1371/jour- nal.pone.0186085 Baird AB, Braun JK, Mares MA, Morales JC, Patton JC, Tran CQ, Bickham JW (2015) Molecular systematic revision of tree bats (Lasiurini): doubling the native mammals of the Hawaiian Islands. Journal of Mammalogy 96(6): 1255-1274. https://doi.org/10.1093/ jmammal/gyv135 Baird AB, Hillis DM, Patton JC, Bickham JW (2008) Evolutionary history of the genus Rhogeessa (Chiroptera: Vespertilionidae) as revealed by mitochondrial DNA sequences. Journal of Mammalogy 89(3): 744-754. https://doi.org/10.1644/07-MAMM-A-135R2.1 Baird AB, Marchan-Rivadeneira MR, Pérez SG, Baker RJ (2012) Mor- phological analysis and description of two new species of Rhogees- sa (Chiroptera: Vespertilionidae) from the Neotropics. Occasional Papers Museum of Texas Tech University 307: 1—25. https://www. biodiversitylibrary.org/page/57756021 Baker RJ, McDonough MM, Swier VJ, Larse PA, Carrera JP, Am- merman LK (2009) New species of bonneted bat, genus Eumops (Chiroptera: Molossidae) from the lowlands of western Ecua- dor and Peru. Acta Chiropterologica 11(1): 1-13. https://doi. org/10.3161/150811009X465659 Basantes M, Tinoco N, Velazco PM, Hofmann MJ, Rodriguez-Posa- da ME, Camacho MA (2020) Systematics and taxonomy of Tonatia saurophila Koopman & Williams, 1951 (Chiroptera, Phyllostomidae). ZooKeys 915: 59-86. https://doi.org/10.3897/ zookeys.915.46995 Benshoof L, Yates TL, Froehlich JW (1984) Noteworthy records of mammals from eastern Honduras. The Southwestern Naturalist 29(4): 511-514. https://doi.org/10.2307/3671014 Burnett SE, Jennings JB, Rainey JC, Best TL (2001) Molossus bondae. Mammalian Species 668: 1-3. https://doi.org/10.1644/1545-1410(2 001)668%3C0001:MB%3E2.0.CO;2 Cirranello A, Simmons NB, Solari S, Baker RJ (2016) Morphological diagnoses of higher-level phyllostomid taxa (Chiroptera: Phyllosto- midae). Acta Chiropterologica 18(1): 39-71. https://doi.org/10.3161 /15081109ACC2016.18.1.002 Cole FR, Wilson DE (2006a) Leptonycteris yerbabuenae. Mammalian Species 797: 1-7. https://doi.org/10.1644/797.1 Cole FR, Wilson DE (2006b) Leptonycteris curasoae. Mammalian Spe- cies: 796: 1-3. https://doi.org/10.1644/796. 1 Davis WB (1965) Review of the Eptesicus brasiliensis complex in Middle America with the description of a new subspecies from Costa Rica. Journal of Mammalogy 46(2): 229-240. https://doi. org/10.2307/1377842 Davis WB (1966) Review of South American bats of the genus Ept- Southwestern Naturalist 11(2): 245-274. https://doi. org/10.2307/3669648 Davis WB (1970) The large fruit bats (genus Artibeus) of Middle Amer- ica, with a review of the Artibeus jamaicensis complex. Journal of Mammalogy 51(1): 105-122. https://doi.org/10.2307/1378537 Davis WB (1984) Review of the large fruit-eating bats of the Artibeus “lituratus”’ complex (Chiroptera: Phyllostomidae) in Middle Amer- eSICus. ica. Occasional Papers Museum Texas Tech University 93: 1-16. https://doi.org/10.5962/bh1.title. 156552 Davis WB, Carter DC (1964) A new species of fruit-eating bat (genus Artibeus) from Central America. Proceedings of the Biological So- ciety of Washington 77: 119-121. https://www.biodiversitylibrary. org/page/34605354#page/495/mode/1up zse.pensoft.net 426 Davis WB, Carter DC (1978) A review of the round-eared bats the 7o- natia silvicola complex, with descriptions of three new taxa. Occa- sional Papers Museum Texas Tech University 53: 1-12. https://doi. org/10.5962/bhI title. 15652 Davis WB, Carter DC, Pine RH (1964) Noteworthy records of Mexican and Central American bats. Journal of Mammalogy 45(3): 375-387. https://doi.org/10.2307/1377410 Davis WB, Gardner AL (2008) Family Vespertilionidae. In Gard- ner AL (Ed) Mammals of South America marsupials, xenar- thrans, shrews, and bats. Vol. 1. University of Chicago Press, Chicago and London, 399-440. _https://doi.org/10.7208/chica- g0/9780226282428.001.0001 Dietz C, von Helversen O (2004) Illustrated identification key to the bats of Europe. Tuebingen & Erlangen. https://auvergne-rhone-alpes.|po. fr/images/chiroptere/telecharger/dietz_von_helversen_2004_1.pdf [December 04, 2019] Divoll TJ, Buck DG (2013) Noteworthy field observations of cave roosting bats in Honduras. Mastozoologia Neotropical 20(1): 149-— 151. https://www.redalyc.org/articulo.oa?id=45728549012 Dolan PG (1989) Systematics of Middle American mastiff bats of the genus Molossus. Special Publications Museum Texas Tech Univer- sity 29: 1-80. https://doi.org/10.5962/bh1 title. 142636 Dolan PG, Carter DC (1979) Distributional notes and records for Mid- dle American Chiroptera. Journal of Mammalogy 60(3): 644-649. https://doi.org/10.2307/1380115 Eger JL (2008) Family Molossidae. In Gardner AL (Ed.) Mammals of South America marsupials, xenarthrans, shrews, and bats. Vol. 1. University of Chicago Press, Chicago and London, 399-440. Espinal M, Mora JM (2012a) Ampliacion en la distribucion de cinco especies de murciélagos en Honduras basada en deteccién por me- dios acusticos. Ceiba 53(1): 30-37. https://doi.org/10.5377/ceiba. v5311.2014 Espinal M, Mora JM (2012b) Noteworthy record of Eptesicus brasilien- sis (Vespertilionidae) in Honduras. Ceiba 53(2): 77—80. https://doi. org/10.5377/ceiba.v53i2.2433 Espinal M, Mora JM, OReilly CM (2016) The occurrence of the Peale’s free-tailed bat (Nyctinomops aurispinosus, Molossidae). Caribbean Journal of Science 49(1): 79-82. https://doi.org/10.18475/cjos.v4911.a8 Garner H (2016a) TTU Mammals collection. Version 9.1. Museum of Texas Tech University (TTU). https://www.gbif.org/occur- rence/911734780 [November 30, 2019] Garner H (2016b) TTU Mammals collection. Version 9.1. Museum of Texas Tech University (TTU). https://www.gbif.org/occur- rence/911692088 [July 04, 2019] Garner H (2016c) TTU Mammals collection. Version 9.1. Museum of Texas Tech University (TTU). https://www.gbif.org/occur- rence/911692134 [September 09, 2019] GBIF.org (2019) GBIF occurrence. https:/Awww.gbif.org/occurrence/ search [September 02, 2019] Giménez AL, Giannini NP (2016) Morphofunctional segregation in mo- lossid bats species (Chiroptera: Molossidae) from the South Ameri- can Southern Cone. Hystrix 27(2): 1-11. Gonzalez-Ruiz N, Ramirez-Pulido J, Arroyo-Cabrales J (2011) A new species of mastiff bat (Chiroptera: Molossidae: Molossus). Mam- malian Biology 76(4): 461-469. https://doi.org/10.1016/j.mam- bio.2010.06.004 Goodwin GG (1940) Three new bats from Honduras and the first re- cord of Enchisthenes harti (Thomas) for North America. American zse.pensoft.net Turcios-Casco, M.A. et al.: Taxonomy of the bats of Honduras Museum Novitates 1075: 1-3. http://digitallibrary.amnh.org/han- dle/2246/4806 Goodwin GG (1942) Mammals of Honduras. Bulletin of the American Museum of Natural History 79(2): 107-195. http://digitallibrary. amnh.org/handle/2246/986 Gregorin R, Moras LM, Acosta LH, Vasconcellos KL, Poma JL, Ro- drigues dos Santos F, Paca RC (2016) A new species of Eumops (Chiroptera: Molossidae) from southeastern Brazil and Bolivia. Mammalian Biology 81(3): 235-246. https://doi.org/10.1016/). mambio.2016.01.002 Hall ER (1981) The mammals of North America. 2™ ed. John Wiley and Sons, New York, 1271 pp. Hernandez DJ (2015) Programa para la conservacion de los murciélagos de Honduras (PCMH). In: Rodriguez Herrera B, Sanchez R (Eds) Estrategia centroamericana para la conservacion de los murciélagos. Universidad de Costa Rica, San José, 41-55. Hernandez J, Flores R, Cardenas S, Cabrera G (2016) Los murciélagos del area propuesta como Parque Nacional Cuyamel/Omoa Golfo de Honduras, un atractivo de biodiversidad. Programa de Conservacion de Murciélagos de Honduras (PCMH), Tegucigalpa, 14 pp. Hoofer SR, Solari S, Larsen PA, Bradley RD, Baker RJ (2008) Phylo- genetics of the fruit-eating bats (Phyllostomidae: Artibeina) inferred from mitochondrial DNA sequences. Occasional Papers Museum Tex- as Tech University 277: 1-15. https://doi.org/10.5962/bh1.title. 156929 Hood C, Gardner AL (2008) Family Emballonuridae. In: Gard- ner AL (Ed.) Mammals of South America marsupials, xenar- thrans, shrews, and bats. Vol. 1. University of Chicago Press, Chicago and London, 188-203. _https://doi.org/10.7208/chica- g0/9780226282428.001.0001 Hurtado N, D’Elia G (2018) Taxonomy of the genus Gardnerycteris (Chiroptera: Phyllostomidae). Acta Chiropterologica 20: 99-115. https://doi.org/10.3161/15081109ACC2018.20.1.007 Hurtado N, Pacheco V (2014) Analisis filogenético del género Mimon Gray, 1847 (Mammalia, Chiroptera, Phyllostomidae) con la de- scripcion de un nuevo género. Therya 5(3): 75-1791. https://doi. org/10.12933/therya- 14-230 Jones Jr. JK, Arroyo-Cabrales J, Owen RD (1988) Revised checklist of bats (Chiroptera) of Mexico and Central America. Occasion- al Papers Museum Texas Tech University 120: 1-34. https://doi. org/10.5962/bh1 title. 142940 Jones Jr. JK, Swanepoel P, Carter DC (1977) Annotated checklist of the bats of Mexico and Central America. Occasional Papers The Mu- seum Texas Tech University 47: 1-40. https://doi.org/10.5962/bhl. title. 156521 Koopman KF (1978) Zoogeography of Peruvian bats with special em- phasis on the role of the Andes. American Museum Novitates 2651: 1-33. http://digitallibrary.amnh.org/handle/2246/2965 Koopman KF (1993) Order Chiroptera. In: Wilson DM, Reeder DM (Eds) Mammal species of the World, a taxonomic and geographic reference (2™ edn). Smithsonian Institute Press, Washington DC, 137-241. Koopman KF (1994) Chiroptera: systematics. In Niethammer J, Schlie- mann H, Starck D (Eds) Handbook of Zoology. Vol. 8. (Mammalia). Walter de Gruyter, Berlin, 1-217. Kraker-Castafieda C, Pérez SG, Cajas-Castillo JO, Echeverria-Tello JL (2016) Lista actualizada de los murciélagos (Mammalia, Chiroptera) de Guatemala. Revista Mexicana de Biodiversidad 87(2): 409-416. https://doi.org/10.1016/j.rmb.2015.10.005 Zoosyst. Evol. 96 (2) 2020, 411-429 Larsen PA, Marchan-Rivadeneira R, Baker RJ (2013) Speciation dy- namics of the fruit-eating bats (genus Artibeus): with evidence of ecological divergence in Central American populations. In: Adams RA, Pedersen SC (Eds) Bat Evolution, Ecology, and Conservation. Springer, New York, 315-339. https://doi.org/10.1007/978-1-4614- 7397-8 16 Larsen TH (2019) Evaluacion bioldgica rapida en Ciudad del Jaguar, Ciudad Blanca, La Mosquitia, Honduras. RAP Bulletin of Biolog- ical Assessment 72 Conservation International, Arlington, 204 pp. LaVal RK (1969) Records of bats from Honduras and EI Salvador. Jour- nal of Mammalogy 50(4): 819-822. https://doi.org/10.2307/1378267 LaVal RK (1973) A revision of Neotropical bats of the genus Myotis. Science Bulletin Los Angeles County 15: 1-54. LaVal RK (1977) Notes on some Costa Rican bats. Brenesia 10/11: 77-83. LaVal RK, Rodriguez-H B (2002) Murciélagos de Costa Rica. Institu- to Nacional de Biodiversidad (INBio), Santo Domingo de Heredia, 320 pp. Lee Jr. TE, Bradley RD (1992) New distribution records of some mam- mals from Honduras. Texas Journal of Science 44: 109-111. Lim B (2015) Balantiopteryx io. The IUCN Red List of Threatened Species, 8 pp. https://dx.doi.org/10.2305/IUCN.UK.2015-4.RLTS. T2532A22030080.en. [November 20, 2019] https://doi.org/10.2305/ IUCN.UK.2015-4.RLTS.T2532A22030080.en Lim BK, Loureiro LO, Garbino GST (2020) Cryptic diversity and range extension in the big-eyed bat genus Chiroderma (Chiroptera, Phyl- lostomidae). ZooKeys 918: 41-63. https://doi.org/10.3897/zook- eys.918.48786 Lopez-Gonzalez C, Presley SJ (2001) Taxonomic status of Molossus bondae J. A. Allen 1904 (Chiroptera: Molossidae), with descrip- tion of a new subspecies. Journal of Mammalogy 82(3): 760-774. https://doi.org/10.1644/1545-1542(2001)082%3C0760:TSOMB- J%3E2.0.CO;2 Loureiro LO, Lim BK, Engstrom MD (2018) A new species of mastiff bat (Chiroptera, Molossidae, Molossus) from Guyana and Ecuador. Mammalian Biology 90: 10-21. https://doi.org/10.1016/).mam- bio.2018.01.008 Loureiro LO, Engstrom MD, Lim BK (2020) Single nucleotide poly- morphisms (SNPs) provide unprecedented resolution of species boundaries, phylogenetic relationships, and genetic diversity in the mastiff bats (Mol/ossus). Molecular Phylogenetics and Evolution 143: 1-13. https://doi.org/10.1016/j.dib.2020. 105276 Mantilla-Meluk H (2014) Defining species and species boundaries in Uroderma (Chiroptera: Phyllostomidae) with a description of a new species. Occasional Papers Museum of Texas Tech University 325: 1-29. https://www. biodiversitylibrary.org/page/57786029 Mantilla-Meluk H, Mufioz-Garay J (2014) Biogeography and taxonom- ic status of Myotis keaysi pilosatibialis LaVal 1973 (Chiroptera: Vespertilionidae). Zootaxa 3793: 60-70. https://doi.org/10.11646/ zootaxa.3793.1.2 Marchan-Rivadeneira MR, Larsen PA, Phillips CJ, Strauss RE, Bak- er RJ (2012) On the association between environmental gradi- ents and skull size variation in the great fruit-eating bat, Artibeus lituratus (Chiroptera: Phyllostomidae). Biological Journal of the Linnean Society 105(3): 623-634. https://doi.org/10.1111/j.1095- 8312.2011.01804.x McCarthy TJ, Davis WB, Hill JE, Jones Jr. JK, Cruz GA (1993) Bat (Mammalia: Chiroptera) records, early collectors, and faunal lists 427 for Northern Central America. Annals of Carnegie Museum 62(3): 191-228. https://www.biodiversitylibrary.org/page/52454529 McCranie JR, Castafieda FE, Estrada N, Ferrufino L, Germer D, Mat- amoros W, Sagastume-Espinoza KO (2019) Biodiversity in Hondu- ras: The environment, flora, bats, medium and large-sized mammals, birds, freshwater fishes, and the amphibians and reptiles. In Pullaiah T (Ed) Global biodiversity: selected countries in the Americas and Australia. Vol. 4. Apple Academic Press Inc., Oakville, 72. https:// doi.org/10.1201/9780429433634-7 McDonough MM, Ammerman LK, Timm RM, Genoways HH, Larsen PA, Baker RJ (2008) Speciation within bonneted bats (genus Eu- mops). The complexity of morphological, mitochondrial, and nu- clear data sets in systematics. Journal of Mammalogy 89(5): 1306— 1315. https://doi.org/10.1644/07-MAMM-A-349. | Medellin RA, Arita HT (1989) Tonatia evotis and Tonatia silvicola. Mammalian Species 334: 1-5. https://doi.org/10.2307/3504111 Medellin RA, Arita HT, Sanchez O (2008) Identificacion de los mur- ciélagos de México, clave de campo. 2“ ed. Universidad Nacional Autonoma de México, Distrito Federal, 79 pp. Medina-Fitoria A (2014) Murciélagos de Nicaragua, guia de campo. Ministerio del Ambiente y los Recursos Naturales (MARENA), Ma- nagua, 275 pp. Medina-Fitoria A, Martinez-Fonseca JG (2019) Cronologia historica de la quiropterologia en Nicaragua. Revista Mexicana de Mas- tozoologia, Nueva Epoca 9(2): 1-28. https://doi.org/10.22201/ ie.20074484e.2019.9.2.286 Medina-Fitoria A, Saldafia O, Martinez JG, Aguirre Y, Silva W, Chavez M, Salazar M, Carballo N, Jarquin O, Gonzalez RA, Diaz L, Cham- bers C, Reid F, Mies R, Williams K, Zolotoff JM, Molina C, Pérez T, Rodriguez J, Gutiérrez L, Fernandez M, Mendieta R, Pérez J (2015) Nuevos reportes sobre los murciélagos (Mammalia: Chiroptera) de Nicaragua, América Central, con la adicion de siete nuevos registros de especies. Mastozoologia Neotropical 22(1): 43-54. http://www. sarem.org.ar/wp-content/uploads/2015/06/SAREM_MastNeo- trop_22-1_05_Medina.pdf Miller C (2014) Host specificity and ectoparasite load of bat flies in Utila, Honduras. Senior Honors Theses. University of New Orle- ans, 63 pp. https://scholarworks.uno.edu/cgi/viewcontent.cgi?arti- cle=1063&context=honors_theses Mora JM (2012) Big Red Bat Lasiurus egregius (Vespertilionidae) in Honduras. The Southwestern Naturalist 57(1): 104-105. https://doi. org/10.1894/0038-4909-57.1.104 Mora JM (2016) Clave para la Identificacion de las Especies de Mur- ciélagos de Honduras. Ceiba 54(2): 93-117. https://doi.org/10.5377/ ceiba.v5412.3283 Mora JM, Lopez LI (2013) First Record of the Hoary Bat (Lasiurus cinereus, Vespertilionidae) for Honduras. Ceiba 51: 89-90. https:// doi.org/10.5377/ceiba.v5 112.1188 Mora JM, Espinal MR, Ruedas LA, Lopez LI (2016) The big Free-tailed bat, Nyctinomops macrotis (Gray, 1839), in Central America. Masto- zoologia Neotropical 23(2): 551-556. Mora JM, Lopez LI, Espinal MR, Marineros L, Ruedas LA (2018) Di- versidad y conservacion de los murciélagos de Honduras. Master Print S. de R.L, Tegucigalpa, 284 pp. Mora JM, Marineros L, Lopez LI (2014) First record of the striped yel- low-eared bat, Vampyriscus nvmphaea (Stenodermatinae, Phyllosto- midae) in Honduras. Caribbean Journal of Science 48(1): 49-51. https://doi.org/10.18475/cjos.v48i1.a7 zse.pensoft.net 428 Moras LM, Gregorin R, Sattler T, Tavares V da C (2018) Uncovering the diversity of dog-faced bats of the genus Cynomops (Chiroptera: Molossidae), with the redescription of C. milleri and the descrip- tion of two new species. Mammalian Biology 89: 37-51. https://doi. org/10.1016/j.mambio.2017.12.005 Moras LM, Tavares V da C, Pepato AR, Santos FR, Gregorin R (2016) Reassessment of the evolutionary relationships within the dog-faced bats, genus Cynomops (Chiroptera: Molossidae). Zoologica Scripta 45(5): 465-480. https://doi.org/10.1111/zsc.12169 Natural History Museum (London) (2019) Collection specimens. https://www. gbif.org/occurrence/1919696413 [November 20, 2019] Pavan AC, Marroig G (2016) Integrating multiple evidences in taxono- my: species diversity and phylogeny of mustached bats (Mormoop- idae: Pteronotus). Molecular Phylogenetics and Evolution 103: 184-198. https://doi.org/10.1016/j.ympev.2016.07.011 Peters S, Lim B, Engstrom M (2002) Systematics of Dog-faced bats (Cynomops) based on molecular and morphometric data. Journal of Mammalogy 83(4): 1097-1110. https://doi.org/10.1644/1545-1542( 2002)083%3C1097:SODFBC%3E2.0.CO;2 Portillo-Reyes HO, Aguirre Y, Hernandez J (2019) Registro de una po- blacion del murciélago frutero hondurefio (Artibeus inopinatus), en Nacaome, Valle, Honduras. Scientia hondurensis 2(1): 71—75. Prestridge H (2019a) Biodiversity Research and Teaching Collections — TCWC Vertebrates. Version 9.3. Texas A&M University Biodi- versity Research and Teaching Collections. https://www.gbif.org/ occurrence/675805658 [September 09, 2019] Prestridge H (2019b) Biodiversity Research and Teaching Collections — TCWC Vertebrates. Version 9.3. Texas A&M University Biodi- versity Research and Teaching Collections. https://www.gbif.org/ occurrence/675825411 [September 09, 2019] Ramirez-Pulido J, Gonzalez-Ruiz N, Gardner AL, Arroyo-Cabrales J (2014) List of recent land mammals of Mexico. Special Publica- tions of the Museum of Texas Tech University 63: 1—71. https://doi. org/10.5962/bhL title. 142891 Reid FA (2009) A field guide to the mammals of Central America and southeast Mexico. 2"¢ed. Oxford University Press, New York, 384 pp. Reid F, Medina A (2016) Artibeus inopinatus. The IUCN Red List of Threatened Species, 7 pp. https://doi.org/10.2305/IUCN.UK.2016- 2.RLTS.T2132A21996586.en [November 20, 2019] Rodriguez-Herrera B, Medellin RA, Timm RM (2007) Murciélagos neotropicales que acampan en hojas. Instituto Nacional de Biodiver- sidad (INBio), Santo Domingo de Heredia, 168 pp. Rodriguez Herrera B, Sanchez R (2015) Estrategia centroamericana para la conservacion de los murciélagos. Universidad de Costa Rica, San José, 85 pp. Rodriguez Herrera B, Sanchez R, Pineda W (2011) First record of Natalus lanatus (Chiroptera: Natalidae) in Costa Rica, and current distribution of Natalus in the country. Ecotropica 17: 113-117. https://www.soc- tropecol.eu/PDF/Ecotropica_2011/Rodriguez_et_al_2011.pdf Rodriguez-Herrera B, Ramirez-Fernandez JD, Villalobos-Chaves D, Sanchez R (2014) Actualizacion de la lista de especies de mamiferos vivientes de Costa Rica. Mastozoologia Neotropical 21(2): 275-289. http://www.sarem.org.ar/wp-content/uploads/2014/12/SAREM _ MastNeotrop_21-2 08 Rodriguez.pdf Saldafia Tapia OA, Namendy M, Martinez-Fonseca JG (2020) First record of the Lesser Long-nosed Bat, Leptonycteris yerb- abuenae Martinez & Villa-R., 1940 (Chiroptera, Phyllosto- zse.pensoft.net Turcios-Casco, M.A. et al.: Taxonomy of the bats of Honduras midae), in Nicaragua. Check List 16(2): 451-456. https://doi. org/10.15560/16.2.451 Simmons NB (2005) Order Chiroptera. In: Wilson DE, Reder DM (Eds) Mammal species of the world: a taxonomic and geographic reference, 3"! ed. The Johns Hopkins University Press, Baltimore, 312-529. Simmons NB, Cirranello AL (2020) Bat species of the world: A taxo- nomic and geographic database. https://batnames. org [25 April 2020] Simmons NB, Voss RS (1998) The mammals of Paracou, French Gui- ana: a neotropical lowland rainforest fauna. Part 1 Bats. Bulletin of the American Museum of Natural History 237: 1-219. http://digi- tallibrary.amnh.org/handle/2246/1634 Srinivasulu C, Racey PA, Mistry S (2010) A key to the bats (Mammalia: Chiroptera) of South Asia. Journal of Threatened Taxa 2(7): 1001-— 1076. https://doi.org/10.11609/JoTT.02352.1001-76 Tejedor A (2011) Systematics of funnel-eared bats (Chiroptera: Natal- idae). Bulletin of the American Museum of Natural History 353: 1-141. https://doi.org/10.1206/636.1 Timm RM, LaVal RK, Rodriguez-H B (1999) Clave de campo para los murciélagos de Costa Rica. Brenesia 52: 1-32. https://hdl.handle. net/1808/4521 Trombone T (2016) AMNH Mammal collections. American Museum of Natural History. https://www. gbif.org/occurrence/8593 19403 [26 September 2019] Turcios-Casco MA, Medina-Fitoria A (2019) Occurrence of Hylonyc- teris underwoodi (Chiroptera, Phyllostomidae) and Thyroptera tri- color (Chiroptera, Thyropteridae) in Honduras. Studies on Neotrop- ical Fauna and Environment 54(1): 69-72. https://doi.org/10.1080/0 1650521.2018.1544205 Turcios-Casco MA, Avila-Palma HD, Ordofiez Trejo EJ, Soler Orellana JA, Ordofiez Mazier DI (2019b) Comments on the diet of phyllosto- mid bats (Chiroptera) in a subtropical dry forest in central Hondu- ras. Studies on Neotropical Fauna and Environment 54(3): 239-244. https://doi.org/10.1080/01650521.2019.1663114 Turcios-Casco MA, Ordofiez Mazier DI, Soler Orellana JA, Avila-Pal- ma HD, Ordofiez Trejo EJ (2019a) Two caves in western Honduras are important for bat conservation: first checklist of bats in Santa Barbara. Subterranean Biology 30: 41-55. https://doi.org/10.3897/ subtbiol.30.35420 Turcios-Casco MA, Medina-Fitoria A, Estrada-Andino N (2020a) North- ernmost record of Chiroderma trinitatum (Chiroptera, Phyllostomi- dae) in Latin America, with distributional comments. Caribbean Jour- nal of Science 50(1): 9-15. https://doi.org/10.18475/cjos.v50i1.a2 Turcios-Casco MA, Medina-Fitoria A, Portillo-Alvarez P (2020b) First record of the Pale-face bat (Phylloderma stenops, Phyllostomidae) in the Caribbean lowlands of Gracias a Dios in eastern Honduras. Ca- ribbean Journal of Science 50(1): 16—22. https://doi.org/10.18475/ cjos.v5011.a3 Turcios-Casco MA, Avila-Palma HD, Ordofiez Trejo EJ, Soler Orella- na JA, Ordofiez Mazier DI, Meza-Flores DE, Velasquez A (2020c) Rare or misidentified? On the external identification of the neglected Artibeus inopinatus Davis & Carter, 1964 (Chiroptera, Phyllostomi- dae) in Honduras. Evolutionary Systematics 4(1): 35-43. https://do1. org/10.3897/evolsyst.4.49377 Valdez R, LaVal RK (1971) Records of bats from Honduras and Nicaragua. Journal of Mammalogy 52(1): 247-250. https://doi. org/10.2307/1378465 Zoosyst. Evol. 96 (2) 2020, 411-429 Velazco PM, Patterson BD (2013) Diversification of the yellow-shoul- dered bats genus Sturnira (Phyllostomidae), in the new world trop- ics. Molecular Phylogenetics and Evolution 68(3): 683-698. https:// doi.org/10.1016/j.ympev.2013.04.016 Wilkinson GS, Fleming TH (1996) Migration routes and evolution of lesser long-nosed bats, Leptonycteris curasoae, inferred from mi- tochondrial DNA. Molecular Ecology 5(3): 329-339. https://doi. org/10.1046/j.1365-294X.1996.00081.x Appendix 1 The following list includes all the specimens examined in this study with their respective localities as were written in the labels: Artibeus inopinatus (2) Honduras: Comayagua: La Carbonera (CZB—2019- 10); Francisco Morazadn: Carboneras (UVS—V—02063) Chiroderma gorgasi (1) Honduras: Gracias (UVS—V—02529) a Dios. Ctudad _ Blanca Cynomops greenhalli (1) Honduras: Comayagua: 580 m (TCWC 22123) Cynomops mexicanus (1) Honduras: At/dntida. La Ceiba, S.A.G. Laboratorio (TTU 104070) Eptesicus furinalis (as Eptesicus andinus) (1) Honduras: At/dntida. La Ceiba, S.A.G. Laboratorio (TTU 104074) Hylonycteris underwoodi (1) Honduras: Gracias a_ Dios: (UVS—V-02527). Ciudad _ Blanca Leptonycteris yerbabuenae (2) Honduras: Valle: 1.7 MIS, 8 MI W Nacaome (TCWC 49748, TCWC 49749) 429 Wilson DE (2008) Family Thyropteridae. In: Gardner AL (Ed.) Mammals of South America marsupials, xenarthrans, shrews, and bats. Vol. 1. The University of Chicago Press, Chicago and London, 392-396. Wilson DE, Mittermeier RA (2019) Handbook of the mammals of the world. Vol. 9 Bats. Lynx Ediciones, Barcelona, 1008 pp. York HA, Rodriguez-Herrera B, LaVal RK, Timm RM (2019) Field key to the bats of Costa Rica and Nicaragua. Journal of Mammalogy 100(6): 1726-1749. https://doi.org/10.1093/jmammal/gyz150 Lonchorhina aurita (2) Honduras: Francisco. Morazan: (UVS—V—02067, UVS—V—02075) Carboneras Mimon cozumelae (1) Honduras: Co/o6n: Wuarska (UVS—V—02059) Myotis pilosatibialis (1) Honduras: Francisco Morazan: 1 km W Talanga (LACM 36879) Natalus lanatus (as Natalus mexicanus) (2) Honduras: Francisco Morazan:. 6 mi N Zamorano (TCWC 10992; La Paz: at 2 mi W Cueva del Viejo (TCWC 11008) Natalus mexicanus (1) Honduras: Copdn: 6 km ESE Copan, 900 m (TCWC 19698) Sturnira hondurensis (as Sturnira bogotensis) (2) Honduras: /ntibuca: La Esperanza (PSM 13753, PSM 13754) Thyroptera tricolor (3) Honduras: Gracias a Dios: Ciudad _ Blanca (UVS—V-02525, UVS—V—02532, UVS—V-02533) zse.pensoft.net