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Blumenbach, 1779. Handbuch der Naturgeschichte, p. 74

Chiropteriformes Kinman, 1994; Vespertilionidae Gray, 1825.

1474

Acerodon, Aethalops, Afronycteris, Alionycteris, Ametrida, Amorphochilus, Anoura, Anthops, Antrozous, Aproteles, Ardops, Arielulus, Ariteus, Artibeus, Asellia, Aselliscus, Austronomus, Baeodon, Balantiopteryx, Balionycteris, Barbastella, Bauerus, Boneia, Brachyphylla, Cabreramops, Cardioderma, Carollia, Casinycteris, Cassistrellus, Centronycteris, Centurio, Chalinolobus, Cheiromeles, Chilonatalus, Chiroderma, Chironax, Choeroniscus, Choeronycteris, Chrotopterus, Cistugo, Cloeotis, Cnephaeus, Coelops, Coleura, Cormura, Corynorhinus, Craseonycteris, Cynomops, Cynopterus, Cyttarops, Dermanura, Desmalopex, Desmodus, Diaemus, Diclidurus, Diphylla, Dobsonia, Doryrhina, Dryadonycteris, Dyacopterus, Ectophylla, Eidolon, Emballonura, Enchisthenes, Eonycteris, Epomophorus, Epomops, Eptesicus, Erophylla, Euderma, Eudiscoderma, Eudiscopus, Eumops, Falsistrellus, Furipterus, Gardnerycteris, Glauconycteris, Glischropus, Glossophaga, Glyphonycteris, Haplonycteris, Harpiocephalus, Harpiola, Harpyionycteris, Hesperoptenus, Hipposideros, Histiotus, Hsunycteris, Hylonycteris, Hypsignathus, Hypsugo, Ia, Idionycteris, Kerivoula, Laephotis, Lampronycteris, Lasionycteris, Lasiurus, Latidens, Lavia, Leptonycteris, Lichonycteris, Lionycteris, Lonchophylla, Lonchorhina, Lophostoma, Lyroderma, Macroderma, Macroglossus, Macronycteris, Macrophyllum, Macrotus, Megaderma, Megaerops, Megaloglossus, Melonycteris, Mesophylla, Micronomus, Micronycteris, Mimetillus, Mimon, Miniopterus, Mirimiri, Mirostrellus, Molossops, Molossus, Monophyllus, Mops, Mormoops, Mormopterus, Mosia, Murina, Musonycteris, Myonycteris, Myopterus, Myotis, Mystacina, Myzopoda, Nanonycteris, Natalus, Neoeptesicus, Neonycteris, Neoplatymops, Neopteryx, Neoromicia, Nesonycteris, Noctilio, Notopteris, Nyctalus, Nycteris, Nycticeinops, Nycticeius, Nyctiellus, Nyctimene, Nyctinomops, Nyctophilus, Otomops, Otonycteris, Otopteropus, Ozimops, Paranyctimene, Parastrellus, Paratriaenops, Paremballonura, Penthetor, Perimyotis, Peropteryx, Pharotis, Philetor, Phoniscus, Phylloderma, Phyllonycteris, Phyllops, Phyllostomus, Pilonycteris, Pipistrellus, Platalina, Platymops, Platyrrhinus, Plecotus, Plerotes, Promops, Pseudoromicia, Ptenochirus, Pteralopex, Pteronotus, Pteropus, Pygoderma, Rhinolophus, Rhinonicteris, Rhinophylla, Rhinopoma, Rhogeessa, Rhynchonycteris, Rhyneptesicus, Rousettus, Saccolaimus, Saccopteryx, Sauromys, Scleronycteris, Scoteanax, Scotoecus, Scotomanes, Scotonycteris, Scotophilus, Scotorepens, Scotozous, Setirostris, Sphaerias, Sphaeronycteris, Stenoderma, Stenonycteris, Sturnira, Styloctenium, Submyotodon, Syconycteris, Tadarida, Taphozous, Thainycteris, Thoopterus, Thyroptera, Tomopeas, Tonatia, Trachops, Triaenops, Trinycteris, Tylonycteris, Uroderma, Vampyressa, Vampyriscus, Vampyrodes, Vampyrum, Vansonia, Vespadelus, Vespertilio, Xeronycteris.

Africa, Americas, Asia, Europe, Oceania

Chiropteran monophyly is strongly supported by both morphological and molecular data (summarized by Simmons, 1994, 1995; see also Kirsch et al., 1995; Miyamoto, 1996; Murphy et al., 2001; Arnason et al., 2002; Teeling et al., 2002, 2005; Van Den Bussche et al., 2002; Miller-Butterworth et al., 2007). Within Mammalia, some previous morphological work supported the placement of bats in Volitantia (Dermoptera + Chiroptera; see Simmons, 1993; Szalay and Lucas, 1993, 1996) and/or Archonta (with Dermoptera, Primates, and Scandentia; see Wible and Novacek, 1988; Simmons, 1995; Miyamoto, 1996); however, numerous molecular studies based on both nuclear and mitochondrial genes, support the placement of Chiroptera within Laurasiatheria, usually grouping with either the cetferungulates (pholidotans, carnivores, cetaceans, artiodactyls, and perissodactyls) or eulipotyphlans (minimally including soricids and talpids; see e.g., Miyamoto et al., 2000; Murphy et al., 2001; Arnason et al., 2002; Douady et al., 2002; Van Den Bussche et al., 2002; Van Den Bussche and Hoofer, 2004, Meredith et al., 2011, Amador et al. 2016). The recent analysis of O'Leary et al. (2013) using both molecular and morphological data also found strong support for the placement of bats within Laurasiatheria, where they appeared as the sister taxon of a clade that included Euungulates and their extinct relatives.
Within Chiroptera, the substantial disagreements concerning higher-level relationships that emerged over the past two decades now appear to be largely settled. The traditional arrangement of bats into the two suborders Megachiroptera (non-echolocating fruit bats) and Microchiroptera (echolocating bats) that prevailed in classification for more than 150 years (e.g., Gray 1821; Gill, 1872; Dobson, 1875; Miller 1907; Simpson, 1945; McKenna and Bell, 1997) and preceeded the discovery of echolocation has been falsified as a hypothesis. Phylogenetic studies based on morphological data supported this traditional view (e.g., Simmons, 1998; Simmons and Geisler, 1998; Gunnell and Simmons, 2005). Molecular data, however, strongly supported a conflicting hypothesis, accepted both here and widely within the taxonomic community, that identifies Yinpterochiroptera (comprising the non-echolocating Pteropodidae - formerly Megachiroptera - plus the echolocating Rhinolophoidea, see below), while Yangochiroptera includes the remaining echolocators (formerly in Microchiroptera; Hutcheon et al., 1998; Teeling et al., 2000, 2002, 2005; Hulva and Horacek, 2002; Van Den Bussche and Hoofer, 2004; Van Den Bussche et al., 2002, 2003; Miller-Butterworth et al., 2007; Amador et al. 2016). Along with this reorganization at the subordinal level, several additional conflicts between morphological work and molecular data have now been settled, with the acceptance of Nycteridae + Emballonuridae within Yangochiroptera as the superfamily Emballonuroidea and support for the inclusion within Noctilionoidea of Mystacinidae, Furipteridae, and Thyropteridae. The position of Myzopodidae remains problematic (see Amador et al., 2016). These changes are reflected in the classification presented here, which largely follows the supplementary Figure 6 of Simmons et al. (2008) and Amador et al. (2016).
The name Chiroptera has been widely applied to the group including all bats, especially in mammalian classifications or literature pertaining to fossil bats (e.g., McKenna and Bell, 1997; Simmons and Geisler, 1998); however, the vast majority of the biological literature on bats focuses on extant species within the crown clade. Here we restrict the name Chiroptera to the crown clade, including all extant bats and their fossil relations. Stem taxa (e.g., Eocene fossil taxa such as Onchyonycteris) are not included in this taxon. As such, the classification presented here for Chiroptera is incomplete as it includes only extant bat species, but not their fossil relations.

References:

Amador, L.I., R.L. Moyers Arévalo, F.C. Almeida, S.A. Catalano, and N.P. Giannini. 2016. Bat systematics in the light of unconstrained analyses of a comprehensive molecular supermatrix. Journal of Mammalian Evolution 2016: 1-34. Read abstract.

Arnason, U., J. A. Adegoke, K. Bodin, E. W. Born, Y. B. Esa, A. Gullberg, M. Nilsson, R. V. Short, X. Xu, and A. Janke. 2002. Mammalian mitogenomic relationships and the root of the eutherian tree. Proceedings of the National Academy of Sciences of the United States of America 99(12): 8151-8156. Read article.

Blumenbach, J.F. 1779. Handbuch der Naturgeschichte. Johann Christian Dieterich, Gottingen: 1-448. Read book.

Dobson, G.E. 1875. Conspectus of the suborder, families, and genera of Chiroptera arranged according to their natural affinities. Annals and Magazine of Natural History ser. 4, 16(1875): 345-357. Read article.

Douady, C.J., P.I. Chatelier, O. Madsen, W.W. de Jong, F. Catzeflis, M.S. Springer, and M.J. Stanhope 2002. Molecular phylogenetic evidence confirming the Eulipotyphla concept and in support of hedgehogs as the sister group to shrews. Molecular Phylogenetics and Evolution 25(1): 200-209. Read abstract.

Gill. T. 1872. Arrangement of the Families of Mammals. Smithsonian Institution, Washington, D.C.: 1-98. Read book.

Gray, J.E. 1821. On the natural arrangement of vertebrose animals. London Medical Repository 15: 296-310. Read article.

Gunnell, G.F., and N.B.Simmons 2005. Fossil evidence and the origin of bats. Journal of Mammalian Evolution 12: 209-246. Read article.

Hulva, P., and I. Horáĉek. 2002. Craseonycteris thonglongyai (Chiroptera: Craseonycteridae) is a rhinolophoid: molecular evidence from cytochrome b. Acta Chiropterologica 4(2): 107-120. Read article.

Hutcheon, J. M., J. A. W. Kirsch, and J. D. Pettigrew. 1998. Base compositional biases and the bat problem. III. The question of microchiropteran monophyly. Philosophical Transactions of the Royal Society B: Biological Sciences 353: 607-617. Read article.

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Meredith, R. W., J. E. Janečka, J. Gatesy, O. A. Ryder, C. A. Fisher, E. C. Teeling, A. Goodbla, E. Eizirik, T. L. L. Simão, T. Stadler, D. L. Rabosky, R. L. Honeycutt, J. J. Flynn, C. M. Ingram, C. Steiner, T. L. Williams, T. J. Robinson, A. Burk-Herrick, M. Westerman, N. A. Ayoub, M. S. Springer, and W. J. Murphy 2011. Impact of the Cretaceous Terrestrial Revolution and KPg extinction on mammal diversification. Science 344(6055): 521-524. Read article.

Miller, G.S., Jr. 1907. The families and genera of bats. Bulletin of the U.S. National Museum 57: 1-282. Read volume.

Miller-Butterworth, C. M., W. J. Murphy, S.J. O’Brien, D.S. Jacobs, M.S. Springer, and E.C. Teeling. 2007. A family matter: conclusive resolution of the taxonomic position of the long-fingered bats, Miniopterus. Molecular Biology and Evolution 24(7): 1553-1561. Read article.

Miyamoto, M.M. 1996. A congruence study of molecular and morphological data for eutherian mammals. Molecular Phylogenetics and Evolution 6(3): 373-390. Read abstract.

Miyamoto, M.M., C. Porter, and M. Goodman 2000. c-Myc gene sequences and the phylogeny of bats and other eutherian mammals. Systematic Biology 49(3): 501-514. Read abstract.

Murphy, W.J., E. Eizirik, W.E. Johnson, Y.P. Zhang, O. Ryder, and S.J. O'Brien. 2001. Molecular phylogenetics and the origins of placental mammals. Nature 409: 614-618. Read abstract.

O'Leary, M.A. , J. I. Bloch, J. J. Flynn, T. J. Gaudin, A. Giallombardo, N. P. Giannini, S. L. Goldberg, B. P. Kraatz, Z.-X. Luo, J. Meng, X. Ni, M. J. Novacek, F. A. Perini, Z. S. Randall, G. W. Rougier, E. J. Sargis, M. T. Silcox, N. B. Simmons, M. Spaulding, P. M. Velazco, M. Weksler, J. R. Wible, and A. L. Cirranello 2013. The placental mammal ancestor and the post-K-Pg radiation of placentals. Science 339(6120): 662-667. Read abstract.

Simmons, N.B. 1993. The importance of methods: archontan phylogeny and cladistic analysis of morphological data. Pages 1-61 In R.D.E. MacPhee (eds.) Primates and their relatives in phylogenetic perspective. Advances in Primatology Series 67: 384 pp. Read abstract.

Simmons, N.B. 1994. The case for chiropteran monophyly. American Museum Novitates 3103: 1-54. Read article.

Simmons, N.B. 1998. A reappraisal of interfamilial relationships of bats. Pages 3-26 In T.H. Kunz and P.A. Racey (eds.) Bat Biology and Conservation . Washington, D.C.: Smithsonian Institution Press. Not available online.

Simmons, N.B., and J.H. Geisler 1998. Phylogenetic relationships of Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx to extant bat lineages, with comments on the evolution of echolocation and foraging strategies in Microchiroptera. Bulletin of the American Museum of Natural History 235: 1-182. Read volume.

Simmons, N.B., K. L. Seymour, J. Habersetzer, and G. F. Gunnell. 2008. Primitive Early Eocene bat from Wyoming and the evolution of flight and echolocation. Nature 451: 818-821. Read abstract.

Simpson, G.G. 1945. The principles of classification and a classification of mammals. Bulletin of the American Museum of Natural History 85: i-xvi, 1-350. Read volume.

Szalay, F.S., and S.G. Lucas 1993. Cranioskeletal morphology of archontans, and diagnoses of Chiroptera, Volitantia, and Archonta. Pages 187-226 In R.D.E. MacPhee (eds.) Primates and their relatives in phylogenetic perspective. Advances in Primatology Series 67: 384 pp. Read abstract.

Szalay, F.S., and S.G. Lucas 1996. The postcranial morphology of Paleocene Chriacus and Mixodectes and the phylogenetic relationships of archontan mammals. Bulletin of the New Mexico Museum of Natural History and Science 7: 1-47. Read volume.

Teeling, E. C., M. Scully, D. J. Kao, M. L. Romagnoli, M. S. Springer, and M. J. Stanhope. 2000. Molecular evidence regarding the origin of echolocation and flight in bats. Nature 403: 188-192. Read abstract.

Teeling, E.C., M.S. Springer, O. Madsen, P. Bates, S.J. O’Brien, and W.J. Murphy. 2005. A molecular phylogeny for bats illuminates biogeography and the fossil record. Science 307: 580-584. Read article.

Teeling, E.C., O. Madsen, R.A. Van Den Bussche, W.W. de Jong, M.J. Stanhope, and M.S. Springer. 2002. Microbat paraphyly and the convergent evolution of a key innovation in Old World rhinolophid microbats. Proceedings of the National Academy of Sciences of the United States of America 99(3): 1431-1436. Read article.

Van Den Bussche, R. A., S.A. Reeder, E.W. Hansen, and S.R. Hoofer. 2003. Utility of the dentin matrix protein 1 (DMP1) gene for resolving mammalian intraordinal relationships. Molecular Phylogenetics and Evolution 26(1): 89-101. Read abstract.

Van Den Bussche, R.A., and S.R. Hoofer 2004. Phylogenetic relationships among recent chiropteran families and the importance of choosing appropriate out-group taxa. Journal of Mammalogy 85(2): 321-330. Read article.

Van Den Bussche, R.A., S.R. Hoofer, and E.W. Hansen 2002. Characterization and phylogenetic utility of the mammalian protamine P1 gene. Molecular Phylogenetics and Evolution 22(3): 333-341. Read abstract.

Wible, J.R., and M.J. Novacek 1988. Cranial evidence for the monophyletic origin of bats. American Museum Novitates 2911: 1-19. Read article.

Koopman, 1985. Bat Research News, 25 (3/4): 26

Vespertilioniformes Hutcheon & Kirsch, 2006.

1046

Afronycteris, Ametrida, Amorphochilus, Anoura, Antrozous, Ardops, Arielulus, Ariteus, Artibeus, Austronomus, Baeodon, Balantiopteryx, Barbastella, Bauerus, Brachyphylla, Cabreramops, Carollia, Cassistrellus, Centronycteris, Centurio, Chalinolobus, Cheiromeles, Chilonatalus, Chiroderma, Choeroniscus, Choeronycteris, Chrotopterus, Cistugo, Cnephaeus, Coleura, Cormura, Corynorhinus, Cynomops, Cyttarops, Dermanura, Desmodus, Diaemus, Diclidurus, Diphylla, Dryadonycteris, Ectophylla, Emballonura, Enchisthenes, Eptesicus, Erophylla, Euderma, Eudiscopus, Eumops, Falsistrellus, Furipterus, Gardnerycteris, Glauconycteris, Glischropus, Glossophaga, Glyphonycteris, Harpiocephalus, Harpiola, Hesperoptenus, Histiotus, Hsunycteris, Hylonycteris, Hypsugo, Ia, Idionycteris, Kerivoula, Laephotis, Lampronycteris, Lasionycteris, Lasiurus, Leptonycteris, Lichonycteris, Lionycteris, Lonchophylla, Lonchorhina, Lophostoma, Macrophyllum, Macrotus, Mesophylla, Micronomus, Micronycteris, Mimetillus, Mimon, Miniopterus, Mirostrellus, Molossops, Molossus, Monophyllus, Mops, Mormoops, Mormopterus, Mosia, Murina, Musonycteris, Myopterus, Myotis, Mystacina, Myzopoda, Natalus, Neoeptesicus, Neonycteris, Neoplatymops, Neoromicia, Noctilio, Nyctalus, Nycteris, Nycticeinops, Nycticeius, Nyctiellus, Nyctinomops, Nyctophilus, Otomops, Otonycteris, Ozimops, Parastrellus, Paremballonura, Perimyotis, Peropteryx, Pharotis, Philetor, Phoniscus, Phylloderma, Phyllonycteris, Phyllops, Phyllostomus, Pipistrellus, Platalina, Platymops, Platyrrhinus, Plecotus, Promops, Pseudoromicia, Pteronotus, Pygoderma, Rhinophylla, Rhogeessa, Rhynchonycteris, Rhyneptesicus, Saccolaimus, Saccopteryx, Sauromys, Scleronycteris, Scoteanax, Scotoecus, Scotomanes, Scotophilus, Scotorepens, Scotozous, Setirostris, Sphaeronycteris, Stenoderma, Sturnira, Submyotodon, Tadarida, Taphozous, Thainycteris, Thyroptera, Tomopeas, Tonatia, Trachops, Trinycteris, Tylonycteris, Uroderma, Vampyressa, Vampyriscus, Vampyrodes, Vampyrum, Vansonia, Vespadelus, Vespertilio, Xeronycteris.

Africa, Americas, Asia, Europe, Oceania

Includes Emballonuroidea (Emballonuridae + Nycteridae; see Teeling et al., 2002). In addition to DNA hybridization data that supports this group (see Hutcheon et al., 1998), two base pair deletions (one in the BRCA1 gene, and one in PLCB4) are found in all yangochiropteran bats but are absent in all yinpterochiropterans and outgroup taxa (Teeling et al. 2005). Studies of molecular data have collectively analyzed more than 13 kilobases of DNA sequence from 20 nuclear and 4 mitochondrial genes (Teeling et al. 2000, 2002, 2003, 2005; Springer et al., 2001; Hulva and Horacek, 2002; Van Den Bussche and Hoofer, 2004; Van Den Bussche et al., 2002; Miller-Butterworth et al., 2007; Agnarsson et al. 2011, Meredith et al. 2011, Amador et al., 2016), and all have recovered Yangochiroptera, often with very high support values (e.g., bootstrap, Bremer values, posterior probabilities). Finally, two combined analyses of morphological and molecular data (Hermsen and Hendricks, 2008; O'Leary et al., 2013) also recovered Yangochiroptera including Emballonuroidea, and the O'Leary et al. (2013) study reported very high support values (jacknife: 100%). The inference of this clade from two combined data analyses is noteworthy. Morphological analyses strongly support monophyly of Noctilionoidea + Vespertilionoidea (Yangochiroptera sensu Koopman, 1985; see Simmons, 1998; Simmons and Geisler, 1998; Gunnell and Simmons 2005), but do not provide support for this group plus Emballonuroidea.
We use Yangochiroptera (Koopman, 1985) instead of Vespertilioniformes (Hutcheon and Kirsch, 2006), because the former has been more widely used to refer to this clade.

References:

Gunnell, G.F., and N.B.Simmons 2005. Fossil evidence and the origin of bats. Journal of Mammalian Evolution 12: 209-246. Read article.

Koopman, K.F. 1985. A synopsis of the families of bats, Part VII. Bat Research News 25(3/4): 25-27. Not available online.

Miller, G.S., Jr. 1907. The families and genera of bats. Bulletin of the U.S. National Museum 57: 1-282. Read volume.

Thomas, 1904. Proc. Zool. Soc. Lond., 1904(2): 5

Myzopoda.

Africa

Monogeneric. Myzopodidae was long considered a close relative of vespertilionids (Miller, 1907; Simpson, 1945; Smith, 1976). However, recent phylogenetic analyses using molecular data now indicate that the only concrete thing we can say about myzopodid relationships is that this family does not belong to Yinpterochiroptera. Current possibilities for the placement of Myzopodidae include all three yangochiropteran superfamilies: Emballonuroidea (Agnarrson et al., 2011; Amador et al., 2016), Vespertilionoidea (Van Den Bussche and Hoofer, 2004; Eick et al., 2005; Meredith et al., 2011), and Noctilionoidea (Teeling et al., 2005; Miller-Butterworth et al., 2007), or as a basal taxon within Yangochiroptera (Hoofer et al., 2003). We have chosen to place this taxon incertae sedis within Yangochiroptera until there is more clarity concerning its superfamilial affinities. See Gunnell et al. (2014) for an emended diagnosis of the family.

References:

Agnarsson, I., C.M. Zambrana-Torrelio, N.P. Flores-Saldana, L.J. May-Collado. 2011. A time-calibrated species-level phylogeny of bats (Chiroptera, Mammalia). PLOS Currents Tree of Life 2011(1):RRN1212: . Read article.

Eick, G.N., D.S. Jacobs, and C.A. Matthee 2005. A nuclear DNA phylogenetic perspective on the evolution of echolocation and historical biogeography of extant bats (Chiroptera) . Molecular Biology and Evolution 22(9): 1869-1886. Read article.

Gunnell, G.F., N.B. Simmons, and E.R. Seiffert 2014. New Myzopodidae (Chiroptera) from the Late Paleogene of Egypt: emended family diagnosis and biogeographic origins of Noctilionoidea. PLoS One 9(2): 1-11. Read article.

Hoofer, S.R., S.A Reeder, E.W. Hansen, and R.A. Van Den Bussche. 2003. Molecular phylogenetics and taxonomic review of noctilionoid and vespertilionoid bats (Chiroptera, Yangochiroptera). Journal of Mammalogy 84(3): 809-821. Read article.

Miller, G.S., Jr. 1907. The families and genera of bats. Bulletin of the U.S. National Museum 57: 1-282. Read volume.

Simpson, G.G. 1945. The principles of classification and a classification of mammals. Bulletin of the American Museum of Natural History 85: i-xvi, 1-350. Read volume.

Smith, J.D. 1976. Chiropteran evolution. Pages 49-69 In R.J. Baker, J.K.Jones Jr., and D.C. Carter (eds.) Biology of the Bats of the New World family Phyllostomatidae. Part I. Special Publications, The Museum, Texas Tech University 10: 218 pp. Not available online.

Thomas, O. 1904. On the osteology and systematic position of the rare Malagasy bat Myzopoda aurita. Proceedings of the Zoological Society of London 1904(2): 2-6. Read article.

Gervais, 1856. In Comte de Castelnau, Exped. Parties Cen. Am. Sud., Zool. (Sec. 7), Vol. 1, pt. 2, (Mammifères), p. 62 footnote

Nycteroidea Van der Hoeven, 1855.

Balantiopteryx, Centronycteris, Coleura, Cormura, Cyttarops, Diclidurus, Emballonura, Mosia, Nycteris, Paremballonura, Peropteryx, Rhynchonycteris, Saccolaimus, Saccopteryx, Taphozous.

Africa, Americas, Asia, Oceania

Formerly included in Yinochiroptera (see e.g., Koopman, 1985) but clearly part of Yangochiroptera; see Teeling et al. (2002), Teeling et al. (2005), Eick et al. (2005), Miller-Butterworth et al. (2007), and Amador et al. (2016). Includes Nycteridae; does not include Rhinopomatidae or Craseonycteridae (see e.g., Teeling et al. 2002, 2005). Relationships of Myzopodidae to this superfamily remain unclear; see that entry and Van Den Bussche and Hoofer (2004), Eick et al. (2005), Teeling et al. (2005), Miller-Butterworth et al. (2007), Agnarsson et al. (2011), Meredith et al. (2011), and Amador et al. (2016). ACR (2007-2018; see 2018: 329) has applied the name Nycteroidea van der Hoeven, 1855 to this superfamily, arguing that the "oldest, typified name is Nycteris G. Cuvier and E. Geoffroy, 1795, rather than Emballonura Temminck, 1838 (or even Emballonuridae, Gervais, 1856)." The priority of a family-group name is independent of the priority of genus-group names, and, following Article 64 of the ICZN (1999), "An author who wishes to establish a new nominal family-group taxon may choose as type genus any included nominal genus the name of which he or she regards as valid [Art. 11.7.1], not necessarily that having the oldest name." ACR (2007-2018) is correct that, within the family-group, the name Nycteridae van der Hoeven, 1855 has priority over Emballonuridae Gervais 1856 [dated 1855]; however, since 1999, the ICZN (1999: Article 35.5) has conserved family-group names in widespread usage, regardless of priority. Therefore, we continue to apply the the name Emballonuroidea Gervais, 1856, first used by Weber (1928), and applied by a succession of subsequent authors, to this superfamily.

References:

ACR 2018. African Chiroptera Report 2018. AfricanBats NPC, Praetoria: i-xvi + 1-8028 pp. Read report.

Gervais, P. 1856 [dated 1855]. Chéiroptères sud-américains. Pages 25-88 In F. de Castelnau (eds.) Expédition dans les parties centrales de l'Amérique du Sud: de Rio de Janeiro à Lima, et de Lima au Para, (Sec. 7), Vol. 1, pt. 2: (Mammifères, ed. P. Gervais). Paris: P. Bertrand. Read volume.

International Commission on Zoological Nomenclature 1999. International Code of Zoological Nomenclature. Fourth Edition. [Including Declaration 44, amendments of Article 74.7.3, and the Amendment on e-publication]. The International Trust for Zoological Nomenclature, London: 306 pp. Read book.

Koopman, K.F. 1985. A synopsis of the families of bats, Part VII. Bat Research News 25(3/4): 25-27. Not available online.

Weber, M.W.C. 1928. Die säugetiere: Einführung in die anatomie und systematik der recenten und fossilen Mammalia, Volume 2. Gustav Fisher, Jena: 898 pp. Not available online.

Gervais, 1856. In Comte de Castelnau, Exped. Parties Cen. Am. Sud., Zool. (Sec. 7), Vol. 1, pt. 2, (Mammifères), p. 62 footnote

Balantiopteryx, Centronycteris, Coleura, Cormura, Cyttarops, Diclidurus, Emballonura, Mosia, Paremballonura, Peropteryx, Rhynchonycteris, Saccolaimus, Saccopteryx, Taphozous.

Africa, Americas, Asia, Oceania

Two subfamilies are recognized: Taphozoinae, consisting of Old World genera, and Emballonurinae, which includes the Old World tribe Emballonurini and the New World tribe Diclidurini; see Lim (2007), Lim et al. (2008), and Ruedi et al. (2012). For additional phylogenies see Barghoorn (1977), Robbins and Sarich (1988), Griffiths and Smith (1991), and Dunlop (1998). South American species reviewed by Jones and Hood (1993).

References:

Barghorn, S.F. 1977. New material of Vespertiliavus Schlosser (Mammalia: Chiroptera) and suggested relationships. American Museum Novitates 2618: 1-29. Read article.

Dunlop, J. 1998. The evolution of behavior and ecology in the Emballonuridae (Unpublished Doctoral Dissertation). York University North York, Ontario, Canada. Read thesis.

Griffiths, T.A., and A.L. Smith 1991. Systematics of emballonuroid bats (Chiroptera: Emballonuridae and Rhinopomatidae) based on hyoid morphology. Pages 62-83 In T.A. Griffiths and D. Klingener (eds.) Contributions to Mammalogy in Honor of Karl F. Koopman. Bulletin of the American Museum of Natural History 206: 432 pp. Read article.

Jones, J.K., Jr., and C.S. Hood. 1993. Synopsis of South American bats of the family Emballonuridae. Occasional Papers, The Museum, Texas Tech University 155: 1-32. Read article.

Lim, B.K. 2007. Divergence times and origin of Neotropical sheath-tailed bats (tribe Diclidurini) in South America. Molecular Phylogenetics and Evolution 45(2007): 777-791. Read abstract.

Lim, B.K., M.D. Engstrom, J.W. Bickham, and J.C. Patton 2007. Molecular phylogeny of New World sheath-tailed bats (Emballonuridae: Diclidurini) based on loci from the four genetic transmission systems in mammals. Biological Journal of the Linnean Society 93(1): 189-209. Read article.

Robbins, L.W., and V.M. Sarich. 1988. Evolutionary relationships in the family Emballonuridae (Chiroptera). Journal of Mammalogy 69(1): 1-13. Read abstract.

Ruedi, M., N. Friedli-Weyeneth,E.C. Teeling, S.J. Puechmaille, S.M. Goodman. 2012. Biogeography of Old World emballonurine bats (Chiroptera: Emballonuridae) inferred with mitochondrial and nuclear DNA. Molecular Phylogenetics and Evolution 64(1): 204-211. Read abstract.

Gervais, 1856. In F. Comte de Castelnau, Exped. Partes Cen. Am. Sud. Zool. (Sec. 7), Vol. 1, pt. 2 (Mammifères), p. 62 footnote

Balantiopteryx, Centronycteris, Coleura, Cormura, Cyttarops, Diclidurus, Emballonura, Mosia, Paremballonura, Peropteryx, Rhynchonycteris, Saccopteryx.

Africa, Americas, Asia, Oceania

McKenna and Bell (1997) divided this subfamily into two tribes, Emballonurini Gervais, 1856 (Mosia, Emballonura, Coleura) and Diclidurini Gray, 1866 (Neotropical emballonurids), which are here recognized; see Lim (2007), Lim et al. (2008), and Ruedi et al. (2012); but see Dunlop (1998). For a key to Neotropical species see Jones and Hood (1993).

References:

Dunlop, J. 1998. The evolution of behavior and ecology in the Emballonuridae (Unpublished Doctoral Dissertation). York University North York, Ontario, Canada. Read thesis.

Gray, J.E. 1866. Synopsis of the genera of Vespertilionidae and Noctilionidae. Annals and Magazine of Natural History ser. 3, 17(1866): 89-93. Read article.

Lim, B.K. 2007. Divergence times and origin of Neotropical sheath-tailed bats (tribe Diclidurini) in South America. Molecular Phylogenetics and Evolution 45(2007): 777-791. Read abstract.

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Gray, 1866. Ann. Mag. Nat. Hist., ser. 3, 17: 92

Balantiopteryx, Centronycteris, Cormura, Cyttarops, Diclidurus, Peropteryx, Rhynchonycteris, Saccopteryx.

Americas

See Lim (2007), Lim et al. (2008), and Ruedi et al. (2012); but see Dunlop (1998).

References:

Dunlop, J. 1998. The evolution of behavior and ecology in the Emballonuridae (Unpublished Doctoral Dissertation). York University North York, Ontario, Canada. Read thesis.

Gray, J.E. 1866. Synopsis of the genera of Vespertilionidae and Noctilionidae. Annals and Magazine of Natural History ser. 3, 17(1866): 89-93. Read article.

Lim, B.K. 2007. Divergence times and origin of Neotropical sheath-tailed bats (tribe Diclidurini) in South America. Molecular Phylogenetics and Evolution 45(2007): 777-791. Read abstract.

Gray, 1866. Ann. Mag. Nat. Hist., ser. 3, 17: 92

Balantiopteryx, Cormura, Cyttarops, Diclidurus, Peropteryx.

Americas

See Lim (2007) and Lim et al. (2007).

References:

Gray, J.E. 1866. Synopsis of the genera of Vespertilionidae and Noctilionidae. Annals and Magazine of Natural History ser. 3, 17(1866): 89-93. Read article.

Lim, B.K. 2007. Divergence times and origin of Neotropical sheath-tailed bats (tribe Diclidurini) in South America. Molecular Phylogenetics and Evolution 45(2007): 777-791. Read abstract.

Lim, 2007. Mol. Phy. Evol., 45(2007): 789

Centronycteris, Rhynchonycteris, Saccopteryx.

Americas

See also Lim et al. (2007).

References:

Lim, B.K. 2007. Divergence times and origin of Neotropical sheath-tailed bats (tribe Diclidurini) in South America. Molecular Phylogenetics and Evolution 45(2007): 777-791. Read abstract.

Gervais, 1856. In F. Comte de Castelnau, Exped. Partes Cen. Am. Sud. Zool. (Sec. 7), Vol. 1, pt. 2 (Mammifères), p. 62 footnote

Coleura, Emballonura, Mosia, Paremballonura.

Africa, Asia, Oceania

See Lim (2007), Lim et al. (2008), and Ruedi et al. (2012); but see Dunlop (1998).

References:

Dunlop, J. 1998. The evolution of behavior and ecology in the Emballonuridae (Unpublished Doctoral Dissertation). York University North York, Ontario, Canada. Read thesis.

Lim, B.K. 2007. Divergence times and origin of Neotropical sheath-tailed bats (tribe Diclidurini) in South America. Molecular Phylogenetics and Evolution 45(2007): 777-791. Read abstract.

Jerdon, 1867. Mammals of India, p. 30

Saccolaimus, Taphozous.

Africa, Asia, Oceania

Equivalent to tribe Taphozoini of McKenna and Bell (1997).

References:

Jerdon, T.C. 1867. The Mammals of India. Thomason College Press, Roorkee: 1-319. Read volume.

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Van der Hoeven, 1855. Handb. Dierkunde, 2nd ed., 2: 1028

Nycteris.

Africa, Asia

Monogeneric; see Griffiths (1994) for a phylogeny of the family. The correct name for the family is "Nycteridae" not "Nycterididae" as has been suggested by some authors (e.g., Russell and Sigé 1970; Habersetzer and Storch, 1987; Kock et al., 2002). The stem for forming a family group name based on Nycteris would be Nycterid-; however, following Article 29.3.1.1 of the ICZN: "If the stem so formed ends in -id, those letters may be elided before adding the family-group suffixes." Furthermore, Article 29.5 adds: "If a spelling of a family-group name was not formed in accordance with Article 29.3 but is in prevailing usage, that spelling is to be maintained, whether or not it is the original spelling and whether or not its derivation from the name of the type genus is in accordance with the grammatical procedures in Articles 29.3.1 and 29.3.2." Consequently, we maintain the correct and commonly accepted spelling (Nycteridae) for family-group names based on the Greek Nycteris (see discussion in Simmons and Geisler [1998: footnote 13]).

References:

Griffiths, T.A 1994. Phylogenetic systematics of slit-faced bats (Chiroptera, Nycteridae), based on hyoid and other morphology. American Museum Novitates 3090: 1-17. Read article.

Habersetzer, J., and G. Storch. 1987. Klassifikation und funktionelle flügelmorphologie paläogener fledermäuse (Mammalia, Chiroptera). Courier Forschungsinstitut Senckenberg 91: 11-150. Not available online.

Kock, D., L. Barnett, J. Fahr, and C. Emms. 2002. On a collection of bats from Gambia (Mammalia: Chiroptera). Acta Chiropterologica 4(1): 77-97. Read article.

Russell, D.E., and B. Sigé. 1970. Révision des chiroptères lutétiens de Messel (Hesse, Allemagne). Palaeovertebrata 3: 83-182. Read abstract.

van der Hoeven, J. 1855. Handboek der Dierkunde. J.C.A. Sulpke, Amsterdam: 1-1068. Read volume.

Gray, 1821. London Med. Repos., 15: 299

708

Afronycteris, Antrozous, Arielulus, Austronomus, Baeodon, Barbastella, Bauerus, Cabreramops, Cassistrellus, Chalinolobus, Cheiromeles, Cistugo, Cnephaeus, Corynorhinus, Cynomops, Eptesicus, Euderma, Eudiscopus, Eumops, Falsistrellus, Glauconycteris, Glischropus, Harpiocephalus, Harpiola, Hesperoptenus, Histiotus, Hypsugo, Ia, Idionycteris, Kerivoula, Laephotis, Lasionycteris, Lasiurus, Micronomus, Mimetillus, Miniopterus, Mirostrellus, Molossops, Molossus, Mops, Mormopterus, Murina, Myopterus, Myotis, Neoeptesicus, Neoplatymops, Neoromicia, Nyctalus, Nycticeinops, Nycticeius, Nyctinomops, Nyctophilus, Otomops, Otonycteris, Ozimops, Parastrellus, Perimyotis, Pharotis, Philetor, Phoniscus, Pipistrellus, Platymops, Plecotus, Promops, Pseudoromicia, Rhogeessa, Rhyneptesicus, Sauromys, Scoteanax, Scotoecus, Scotomanes, Scotophilus, Scotorepens, Scotozous, Setirostris, Submyotodon, Tadarida, Thainycteris, Tomopeas, Tylonycteris, Vansonia, Vespadelus, Vespertilio.

Africa, Americas, Asia, Europe, Oceania

Includes Natalidae; see Van Den Bussche et al. (2003), Hoofer et al. (2003), and Van Den Bussche and Hoofer (2004). Does not include Thyropteridae, Furipteridae, or Mystacinidae; see Pierson (1986), Pierson et al. (1986), Van Den Bussche and Hoofer (2004), Agnarsson et al., (2011), and Amador et al. (2016). Relationships of Myzopodidae to this superfamily are unclear. See that entry and Van Den Bussche and Hoofer (2004), Eick et al. (2005), Teeling et al., (2005), Miller-Butterworth et al. (2007), Agnarsson et al. (2011), Meredith et al. (2011), and Amador et al. (2016). Two newly recognized families, formerly included within Vespertilionidae, are included in Vespertilionoidea: Cistugidae (Lack et al., 2010) and Miniopteridae (Van Den Bussche and Hoofer, 2004; Miller-Butterworth et al., 2007).

References:

Gray, J.E. 1821. On the natural arrangement of vertebrose animals. London Medical Repository 15: 296-310. Read article.

Lack, J.B., Z.P. Roehrs, C.E. Stanley, Jr., M. Ruedi,and R.A. Van Den Bussche 2010. Molecular phylogenetics of Myotis indicate familial-level divergence for the genus Cistugo (Chiroptera). Journal of Mammalogy 91(4): 976-992. Read article.

Pierson, E.D. 1986. Molecular systematics of the Microchiroptera: higher taxon relationships and biogeography (Unpublished Doctoral Dissertation). University of California, Berkeley Berkeley, CA. Not available online.

Pierson, E.D., V.M. Sarich, J.M. Lowenstien, M.J. Daniel, and W.E. Rainey 1986. A molecular link between the bats of New Zealand and South America. Nature 323: 60-63. Read abstract.

Lack, Roehrs, Stanley, Ruedi & Van Den Bussche, 2010. J. Mammal., 91(4): 980

Cistugo.

Africa

Previously included in Vespertilionidae as either a subgenus of Myotis (e.g., Corbet and Hill, 1991; Koopman, 1993, 1994) or a distinct genus within Vespertilioninae (e.g., Eick et al. 2005; Stadelmann et al., 2004), but clearly distinct at the family level; see Lack et al. (2010). Although Benda (2010) and the African Chiroptera Report (2017) have used the name Cistugonidae as the correct form of the family level name, the etymology of Cistugo was not clarified by Thomas (1912) and the name is not clearly Greek or Latin; therefore, as in Article 29.3.3 of the Code (ICZN, 1999), the stem for the formation of the family-group name is chosen by the author of the family level name.

References:

ACR 2017. African Chiroptera Report 2017. AfricanBats NPC, Praetoria: i-xxi, 1-7635 pp. Read volume.

Benda, P. 2010. České vernakulární jmenosloví netopýrů. I. Návrh úplného jmenosloví. Vespertilio 13-14: 263-308. Read article.

Corbet, G.B., and J.E. Hill 1980. A World List of Mammalian Species, second edition. British Museum (Natural History) Publications, London: 1-226. Not available online.

Koopman, K.F. 1993. Chiroptera. Pages 137-241 In D.E. Wilson and D.M. Reeder (eds.) Mammal Species of the World. A Taxonomic and Geographic Reference. Washington, D.C.: Smithsonian Institution Press. Not available online.

Koopman, K.F. 1994. Chiroptera: Systematics. Handbuch der Zoologie, vol. VIII, Mammalia, Part 60. de Gruyter, Berlin: 224 pp. Not available online.

Stadelmann, B., D.S. Jacobs, C. Schoeman, and M. Ruedi 2004. Phylogeny of African Myotis bats (Chiroptera, Vespertilionidae) inferred from cytochrome b sequences. Acta Chiropterologica 6(2): 177-192. Read article.

Thomas, O. 1912. A new vespertilionine bat from Angola. Annals and Magazine of Natural History ser. 8, 10(56): 204-206. Read article.

Gervais, 1856. In Comte de Castelnau, Exped. Partes Cen. Am. Sud., Zool. (Sec. 7), Vol. 1, pt. 2 (Mammiferes), p. 53 footnote

134

Africa, Americas, Asia, Europe, Oceania

Includes Tomopeatinae; see Barkley (1984), Sudman et al. (1994), Simmons (1998), and Simmons and Geisler (1998). Revised by Freeman (1981) and Legendre (1984). For recent phylogenies that cover much of the family see Ammerman et al. (2012), Gregorin and Cirranello (2015), and Amador et al. (2018). Afro-Malagasy species reviewed by Lamb et al. (2011).

References:

Ammerman, L.K., D.N. Lee, and T.M. Tipps. 2012. First molecular phylogenetic insights into the evolution of free-tailed bats in the subfamily Molossinae (Molossidae, Chiroptera). Journal of Mammalogy 93(1): 12-28. Read article.

Barkley, L.J. 1984. Evolutionary relationships and natural history of Tomopeas ravus (Mammalia: Chiroptera) (Unpublished Masters Thesis). Louisiana State University Baton Rouge. Not available online.

Freeman, P.W 1981. A multivariate study of the family Molossidae (Mammalia: Chiroptera): morphology, ecology, evolution. Fieldiana: Zoology, n.s. 7: 1-173. Read volume.

Gregorin, R., and A.L. Cirranello 2016. Phylogeny of Molossidae Gervais (Mammalia: Chiroptera) inferred by morphological data. Cladistics 32(1): 2-35. Read abstract.

Lamb, J.M., T. Ralph, T. Naidoo, P.J. Taylor, F. Ratrimomanarivo, W.T. Stanley, and S.M. Goodman. 2011. Toward a molecular phylogeny for the Molossidae (Chiroptera) of the Afro-Malagasy region. Acta Chiropterologica 13(1): 1-16. Read abstract.

Legendre, S. 1984. Étude odotologíque des représentants actuels du groupe Tadarida (Chiroptera, Molossidae). Implications phylogéniques, systématiques et zoogéographiques. Revue Suisse de Zoologie 91: 399-442. Read article.

Sudman, P, D., L.J. Barkley, and M.S. Hafner. 1994. Familial affinity of Tomopeas ravus (Chiroptera) based on protein electophoretic and cytochrome B sequence data. Journal of Mammalogy 75: 365-377. Read abstract.

Miller, 1907. Bull. U.S. Natl. Mus. , 57: 237

Tomopeas.

Americas

Formerly included in Vespertilionidae; see Barkley (1984), Sudman et al. (1994).

References:

Gervais, 1856. In Comte de Castelnau, Exped. Partes Cen. Am. Sud., Zool. (Sec. 7), Vol. 1, pt. 2 (Mammifères), p. 53 footnote

Cheiromelinae Legendre, 1984; Tadaridinae Legendre, 1984; Tadarinae Peterson, Eger & Mitchell, 1995.

133

Africa, Americas, Asia, Europe, Oceania

Equivalent to Molossidae sensu Freeman (1981), Legendre (1984), Koopman (1993, 1994), and Peterson et al. (1995). While this group has been subdivided into two (Gregorin, 2000) or three (Legendre, 1984) subgroups, continuing confusion concerning intergeneric relationships (see e.g., Ammerman et al., 2012; Gregorin and Cirranello, 2015; and Amador et al., 2016) leads us to await further studies prior to recognizing tribes within this subfamily. See Gregorin and Taddei (2002) for a key to Brazilian species.

References:

Freeman, P.W 1981. A multivariate study of the family Molossidae (Mammalia: Chiroptera): morphology, ecology, evolution. Fieldiana: Zoology, n.s. 7: 1-173. Read volume.

Gregorin, R. 2000. Filogenia de Molossidae Gervais, 1856 (Mammalia: Chiroptera) (Unpublished Doctoral Dissertation). Universidade de São Paulo Sao Paulo, Brazil. Read abstract.

Gregorin, R., and A.L. Cirranello 2016. Phylogeny of Molossidae Gervais (Mammalia: Chiroptera) inferred by morphological data. Cladistics 32(1): 2-35. Read abstract.

Gregorin, R., and V.A. Taddei 2002. Chave artificial para a identificãço de molossídeos Brasileiros (Mammalia, Chiroptera). Mastozoologia Neotropical 9: 13-32. Read article.

Koopman, K.F. 1994. Chiroptera: Systematics. Handbuch der Zoologie, vol. VIII, Mammalia, Part 60. de Gruyter, Berlin: 224 pp. Not available online.

Peterson, R.L., J.L. Eger, and L. Mitchell. 1995. Chiroptères. Vol. 84. Faune de Madagascar. Muséum National d'Histoire Naturelle, Paris: 204. Not available online.

Dobson, 1875. Ann. Mag. Nat. Hist., ser. 4, 16: 349

Miniopterus.

Africa, Asia, Europe, Oceania

Previously included in Vespertilionidae as a subfamily, but clearly distinct; see Van Den Bussche and Hoofer (2004) and Miller-Butterworth et al. (2007).

References:

Gray, 1866. Ann. Mag. Nat. Hist., ser. 3, 17: 90

Chilonatalus, Natalus, Nyctiellus.

Americas

Once considered monogeneric, the family is now comprised of three genera, with Nyctiellus and Chilonatalus raised to generic rank by Morgan (1989) and Morgan and Czaplewski (2003). Dávalos (2005) reviewed phylogenetic relationships and biogeography. Tejedor (2011) revised the family.

References:

Dávalos, L.M. 2005. Molecular phylogeny of funnel-eared bats (Chiroptera: Natalidae), with notes on biogeography and conservation. Molecular Phylogenetics and Evolution 37(1): 91-103. Read abstract.

Gray, J.E. 1866. Synopsis of the genera of Vespertilionidae and Noctilionidae. Annals and Magazine of Natural History ser. 3, 17(1866): 89-93. Read article.

Morgan, G.S. 1989. Fossil Chiroptera and Rodentia from the Bahamas, and the historical biogeography of the Bahamian mammal fauna. Pages 685-740 In C.A. Woods (eds.) Biogeography of the West Indies: Past, Present, and Future. Gainesville: Sandhill Crane Press. Not available online.

Morgan, G.S., and N.J. Czaplewski. 2003. A new bat (Chiroptera: Natalidae) from the early Miocene of Florida, with comments on natalid phylogeny. Journal of Mammalogy 84: 729-752. Read article.

Tejedor, A. 2011. Systematics of funnel-eared bats (Chiroptera: Natalidae). Bulletin of the American Museum of Natural History 353(): 1-140. Read article.

Gray, 1821. London Med. Repos., 15: 299

531

Africa, Americas, Asia, Europe, Oceania

Does not include Tomopeatinae; see Barkley (1984), Sudman et al. (1994), Simmons (1998), and Simmons and Geisler (1998). Does not include Cistugo; see Lack et al. (2010). Does not include Miniopterinae; see Hoofer and Van Den Bussche (2003), Van Den Bussche and Hoofer (2004), and Miller-Butterworth et al. (2007). Includes Antrozoinae; see discussion under that subfamily. For a phylogeny including representatives of most genera, see Volleth and Heller (1994), and see Kawai et al. (2002), Hoofer and Van Den Bussche (2003), and Amador et al. (2016) for relationships among subfamilies.

References:

Gray, J.E. 1821. On the natural arrangement of vertebrose animals. London Medical Repository 15: 296-310. Read article.

Hoofer, S.R., and R.A. Van Den Bussche. 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica 5(suppl.): 1-63. Read abstract.

Kawai, K., M. Nikaido, M. Harada, S. Matsumura, L.-K. Lin, Y. Wu, M. Hasegawa, and N. Okada. 2002. Intra- and interfamilial relationships of Vespertilionidae inferred by various molecular markers including SINE insertion data. Journal of Molecular Evolution 55: 284-301. Read abstract.

Volleth, M., and K.-G. Heller. 1994. Phylogenetic relationships of vespertilionid genera (Mammalia: Chiroptera) as revealed by karyological analysis. Zeitschrift für Zoologische Systematik und Evolutionsforschung 32: 11-34. Read abstract.

Miller, 1907. Bull. U.S. Natl. Mus., 57: 232

Kerivoula, Phoniscus.

Africa, Asia, Oceania

References:

Miller, G.S., Jr. 1907. The families and genera of bats. Bulletin of the U.S. National Museum 57: 1-282. Read volume.

Miller, 1907. Bull. U.S. Natl. Mus., 57: 229

Harpiocephalus, Harpiola, Murina.

Asia, Europe, Oceania

References:

Miller, G.S., Jr. 1907. The families and genera of bats. Bulletin of the U.S. National Museum 57: 1-282. Read volume.

Tate, 1942. Bull. Amer. Mus. Nat. Hist., 80: 229

143

Eudiscopus, Myotis, Submyotodon.

Africa, Americas, Asia, Europe, Oceania

Originally named as a tribe within Vespertilioninae; raised to subfamily level by Simmons (1998), following the suggestion of Volleth and Heller (1994). Subsequent work by Hoofer and Van Den Bussche (2003), Kawaii et al. 2002; Lack et al. (2010), and Stadelmann et al. (2004) provides support for subfamilial status for this clade. Includes Eudiscopus, following Amador et al. (2016), although this placement is provisional (see also Borisenko et al., 2008). Includes Submyotodon (Ruedi et al., 2013, 2015; Amador et al., 2016).

References:

Borisenko, A.V., S.V. Kruskop, and N.V. Ivanova. 2008. A new mouse-eared bat (Mammalia: Chiroptera: Vespertilionidae) from Vietnam. Russian Journal of Theriology 7(2): 57-69. Read article.

Hoofer, S.R., and R.A. Van Den Bussche. 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica 5(suppl.): 1-63. Read abstract.

Novaes, R.L.M., D.E. Wilson, and R. Moratelli. 2022. Catalogue of primary types of Neotropical Myotis (Chiroptera, Vespertilionidae). Zootaxa 1105(): 127-164. Read article.

Ruedi, M., B.Stadelmann, Y. Gager, E.J.P. Douzery, C.M. Francis, L. Lin, A. Guillén-Servent, and A. Cibois. 2013. Molecular phylogenetic reconstructions identify East Asia as the cradle for the evolution of the cosmopolitan genus Myotis (Mammalia, Chiroptera). Molecular Phylogenetics and Evolution 69(3): 437-449. Read abstract.

Ruedi, M., G. Csorba, L. Lin, and C. Chou. 2015. Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920(1): 301-342. Read abstract.

Tate, G.H.H. 1942. Review of the vespertilionine bats: with special attention to genera and species of the Archbold collections. Bulletin of the American Museum of Natural History 8(7): 1-297. Read article.

Gray, 1821. London Med. Repos., 15: 299

314

Africa, Americas, Asia, Europe, Oceania

Does not include Myotis; see Volleth and Heller (1994), Hoofer and Van Den Bussche (2003), Kawaii et al. (2002), Lack et al. (2010), and Stadelmann et al. (2004). We place Rhyneptesicus, Parastrellus, and Perimyotis incertae sedis within the subfamily (see Juste et al., 2013; Koubínová et al. (2013), Amador et al. 2016), although the latter two should be placed into their own tribe (see Roehrs et al., 2010; Amador et al., 2016), which has not been formally named. See Lack and Van Den Bussche (2010) for a discussion of problems associated with resolving relationships especially within this subfamily. Roehrs et al. (2010) summarized previous classifications. See Hoofer and Van Den Bussche (2003), Roehrs et al. (2010), and Amador et al. (2016) for recent phylogenies. Hill and Harrison (1987) reviewed bacular morphology, and Volleth (1992), Volleth and Heller (1994), and Volleth et al. (2001, 2006) reviewed karyotypes.

References:

Hill, J.E., and D.L. Harrison. 1987. The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a systematic review, a synopsis of Pipistrellus and Eptesicus, and the description of a new genus and subgenus. Bulletin of the British Museum (Natural History), Zoology Series 52(7): 225-305. Read article.

Hoofer, S.R., and R.A. Van Den Bussche. 2001. Phylogenetic relationships of plecotine bats and allies based on mitochondrial ribosomal sequences. Journal of Mammalogy 82: 131-137. Read article.

Hoofer, S.R., and R.A. Van Den Bussche. 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica 5(suppl.): 1-63. Read abstract.

Juste, J., P. Benda, J.L. Garcia‐Mudarra, and C. Ibáñez 2013. Phylogeny and systematics of Old World serotine bats (genus Eptesicus, Vespertilionidae, Chiroptera): an integrative approach.. Zoologica Scripta 42(5): 441-457. Read abstract.

Koubínová, D., N. Irwin, P. Hulva, P. Koubek and J. Zima. 2013. Hidden diversity in Senegalese bats and associated findings in the systematics of the family Vespertilionidae. Frontiers in Zoology 10(48): 1-16. Read article.

Lack, J.B., and R.A. Van Den Bussche 2010. Identifying the confounding factors in resolving phylogenetic relationships in Vespertilionidae. Journal of Mammalogy 91(6): 1435-1448. Read article.

Roehrs, Z.P., J.B. Lack, and R.A. Van Den Bussche. 2010. Tribal phylogenetic relationships within Vespertilioninae (Chiroptera: Vespertilionidae) based on mitochondrial and nuclear sequence data. Journal of Mammalogy 91(5): 1073-1092. Read article.

Volleth, M. 1992. Comparative analysis of the banded karyotypes of the European Nyctalus species (Vespertilionidae; Chiroptera). Pages 221-226 In I. Horácek and V. Vohralík (eds.) Prague Studies in Mammalogy. Prague: Charles University Press. Not available online.

Volleth, M., G. Bonner, M.C. Göpfert, K.-G. Heller, O. von Helversen, and H.-S. Yang. 2001. Karyotype comparison and phylogenetic relationships of Pipistrellus-like bats (Vespertilionidae; Chiroptera; Mammalia). Chromosome Research 9: 25-46. Read abstract.

Volleth, M., K-G. Heller, and J.Fahr 2006. Phylogenetic relationships of three “Nycticeiini” genera (Vespertilionidae, Chiroptera, Mammalia) as revealed by karyological analysis. Mammalian Biology 71(1): 1-12. Read abstract.

Bianchi, 1917. Annu. Mus. Zool., 21(1916): lxxvi

Rhineptesicus Horáček and Hanák, 1986.

Rhyneptesicus.

Asia

Previously considered part of Eptesicus, but clearly merits recognition at the generic level; see Horáček and Hanák (1986), Horáček et al. (2000), Juste et al. (2013), and Amador et al. (2016).

References:

Bianchi, V. 1917. Predvaritel’nyja zametki o letučih’ myšah’ (Chiroptera) Rossi [Notes préliminares sur les chauve-souris ou Chiroptères de la Russie]. Annu. Mus. Zool. [Petrograd] XXI: 1916: LXXII-LXXXII. Read article.

Horáček, I., and V. Hanák. 1985-1986. Generic status of Pipistrellus savii and comments on classification of the genus Pipistrellus (Chiroptera, Vespertilionidae). Myotis 23-24: 9-16. Not available online.

Horáček, I., V.Hanák, and J. Gaisler. 2000. Bats of the Palearctic region: a taxonomic and biogeographic review. Pages 11-157 In B.W. Woloszyn (eds.) Proceedings of the VIIIth European Bat Research Symposium. Vol I. Approaches to Biogeography and Ecology of Bats. Krakow, Poland: Chiropterological Information Center, Institute of Systematics and Evolution of Animals PAS. Not available online.

Hoofer, Van Den Bussche & Horáček, 2006. J. Mammal., 87: 985

Parastrellus Horáček and Hanák, 1985 [nomen nudum]; Parastrellus Horáček and Hanák, 1985-1986 [nomen nudum].

Parastrellus.

Americas

Originally part of Pipistrellus. Horáček and Hanák (1985, 1985-1986) found that hesperus differed substantially morphologically from the other American pipistrelle (subflavus) as well as European forms to warrant recognition as a new genus ("Parastrellus"), which they did not formally describe. Consequently, the name Parastrellus Horáček and Hanák (1985, 1985-1986) is a nomen nudum. Molecular data also support recognition of Parastrellus; see Hoofer and Van Den Bussche (2003), Hoofer, Van Den Bussche and Horáček (2006, who formally described the genus), Roehrs et al. (2011), and Amador et al. (2016). This genus belongs, with Perimyotis, in a tribe within Vespertilioninae (e.g., the "Perimyotine group" of Roehrs et al. 2010; and see Amador et al., 2016); however, such a taxon has not yet been formally described and we place both Parastrellus and Perimyotis incertae sedis within Vespertilioninae at present.

References:

Hoofer, S.R., and R.A. Van Den Bussche. 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica 5(suppl.): 1-63. Read abstract.

Hoofer, S.R., R.A. Van Den Bussche, and I. Horáček. 2006. Generic status of the American pipistrelles (Vespertilionidae) with description of a new genus. Journal of Mammalogy 87(5): 981-992. Read article.

Horáček, I., and V. Hanák. 1985. Generic status of Pipistrellus savii (Bonaparte, 1837) and remarks on systematics of the genus Pipistrellus. Bat Research News 26: 62 (abstract). Not available online.

Menu, 1984. Mammalia, 48(3): 415

Perimyotis.

Americas

Originally part of Pipistrellus. Menu (1984, 1987) suggested that Perimyotis had a close relationship with species in the myotine subgenus Leuconoe and differed substantially enough from the other American pipstrelle (hesperus) as well as European forms to warrant recognition at the generic level. Molecular data support recognition of Perimyotis at the generic level; see Horáček and Hanák (1985-1986), Hoofer and Van Den Bussche (2003), Stadelmann et al. (2004), Roehrs et al. (2011), and Amador et al. (2016), but do not support a close relationship of Perimyotis with either Pipistrellus or Myotis. Rather, this genus belongs, with Parastrellus, in a tribe within Vespertilioninae (e.g., the "Perimyotine group" of Roehrs et al. 2010; and see Amador et al., 2016); however, such a taxon has not yet been formally described and we place both Parastrellus and Perimyotis incertae sedis within Vespertilioninae at present.

References:

Hoofer, S.R., and R.A. Van Den Bussche. 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica 5(suppl.): 1-63. Read abstract.

Menu, H. 1984. Révision du statut de Pipistrellus subflavus (F. Cuvier, 1832). Proposition d'un taxon générique nouveau: Perimyotis nov. gen.. Mammalia 48: 409-416. Read abstract.

Menu, H. 1987. Morphotypes dentaires actuels et fossiles des chiroptères. Palaeovertebrata 17: 77-150. .

Miller, 1897. North American Fauna, 13: 41

Antrozous, Baeodon, Bauerus, Rhogeessa.

Americas

Includes Rhogeessa and Baeodon (Hoofer and Van Den Bussche, 2001; Roehrs et al., 2010, 2011; Amador et al., 2016). Hoofer and Van Den Bussche (2001) found that Otonycteris was sister taxon to Antrozoini, but this result has not been supported by subsequent studies (e.g., Roehrs et al. 2010, 2011). Simmons (1998) raised this group to family level and moved it to Molossoidea, but subsequent studies based on DNA sequence data (e.g., Hoofer and Van Den Bussche, 2001, 2003; Roehrs et al. 2010; Amador et al. 2016) indicate that Antrozoini belongs in Vespertilionidae and nests within Vespertilioninae, a placement more in line with traditional classifications (e.g., Simpson, 1945; Koopman, 1993, 1994; McKenna and Bell, 1997).

References:

Hoofer, S.R., and R.A. Van Den Bussche. 2001. Phylogenetic relationships of plecotine bats and allies based on mitochondrial ribosomal sequences. Journal of Mammalogy 82: 131-137. Read article.

Hoofer, S.R., and R.A. Van Den Bussche. 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica 5(suppl.): 1-63. Read abstract.

Koopman, K.F. 1994. Chiroptera: Systematics. Handbuch der Zoologie, vol. VIII, Mammalia, Part 60. de Gruyter, Berlin: 224 pp. Not available online.

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Miller, G.S., Jr. 1897. Revision of the North American bats of the family Vespertilionidae. North American Fauna 13: 5-135. Read article.

Roehrs, Z.P., J.B. Lack, and R.A. Van Den Bussche 2011. A molecular phylogenetic reevaluation of the tribe Nycticeiini (Chiroptera: Vespertilionidae). Acta Chiropterologica 13(1): 17-31. Read abstract.

Simpson, G.G. 1945. The principles of classification and a classification of mammals. Bulletin of the American Museum of Natural History 85: i-xvi, 1-350. Read volume.

Tate, 1942. Bull. Am. Mus. Nat. Hist., 80: 290

Lasiurus.

Americas

Although Amador et al. (2016) found that Euderma was weakly associated with Lasiurini, this appears to be a novel result. Consequently, we only include Lasiurus in this tribe; see e.g. Hoofer and Van Den Bussche (2001) and Roehrs et al. (2010).

References:

Hoofer, S.R., and R.A. Van Den Bussche. 2001. Phylogenetic relationships of plecotine bats and allies based on mitochondrial ribosomal sequences. Journal of Mammalogy 82: 131-137. Read article.

Gervais, 1856. In F. Comte de Castelanu, Exped. Partes Cen. Am. Sud., Zool. (Sec. 7), Vol. 1, pt. 2, Mammifères, p. 71

Arielulus, Cnephaeus, Eptesicus, Glauconycteris, Hesperoptenus, Histiotus, Ia, Lasionycteris, Neoeptesicus, Nycticeius, Scoteanax, Scotomanes, Scotorepens, Thainycteris.

Africa, Americas, Asia, Europe, Oceania

Nycticeiini sensu Tate (1942) and subsequent classifications such as Koopman (1994), McKenna and Bell (1997), and Simmons (2005) is not monophyletic; see Roehrs et al. (2011) for a review and Volleth and Heller (1994), Hoofer and Van Den Bussche, (2001, 2003), Volleth et al., 2006; Roehrs et al. (2010, 2011); Amador et al. (2016). Does not include Nycticeinops, Rhogeessa, Scotoecus, or Scotophilus. Includes Arielulus, Eptesicus, Glauconycteris, Hesperoptenus, Ia, and Lasionycteris. Relationships of Nycticeius remain unclear and in some recent phylogenies (e.g., Roehrs et al., 2011; Amador et al., 2016) this taxon does not form a clade with the other taxa in this tribe. However, until additional research is undertaken to resolve this issue, we retain Nycticeius in this group and apply the name Nycticeiini, rather than Eptescini, which would be more appropriate should relationships of Nycticieus fall outside this group.

References:

Hoofer, S.R., and R.A. Van Den Bussche. 2001. Phylogenetic relationships of plecotine bats and allies based on mitochondrial ribosomal sequences. Journal of Mammalogy 82: 131-137. Read article.

Hoofer, S.R., and R.A. Van Den Bussche. 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica 5(suppl.): 1-63. Read abstract.

Koopman, K.F. 1994. Chiroptera: Systematics. Handbuch der Zoologie, vol. VIII, Mammalia, Part 60. de Gruyter, Berlin: 224 pp. Not available online.

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Simmons, N.B. 2005. Chiroptera. Pages 312-529 In D.E. Wilson and D.M. Reeder (eds.) Mammal Species of the World: A Taxonomic and Geographic Reference, 3rd edition, Volume 1. Baltimore: Johns Hopkins University Press. Read chapter.

Gray, 1866. Ann. Mag. Nat. Hist., ser. 3, 17: 90

Barbastella, Corynorhinus, Euderma, Idionycteris, Otonycteris, Plecotus.

Africa, Americas, Asia, Europe

Provisionally accepted as monophyletic (Hoofer and Van Den Bussche, 2001; Amador et al. 2016) despite weak support in some recent analyses and some results that show this clade is paraphyletic with respect to Lasiurini (Roehrs et al. 2010). Apparently includes Otonycteris; see Qumsiyeh and Bickham (1993), Bogdanowicz et al. (1998), Roehrs et al. (2010), Amador et al. (2016), but see also Pine et al. (1971) and Hoofer and Van Den Bussche (2001, 2003). Although in two recent phylogenies Idionycteris does not appear as part of Plecotini (see Roehrs et al. 2010; Amador et al., 2016), given the long association between Idionycteris and Euderma, we have retained Idionycteris in this clade pending additional investigation of this unusual result. For phylogenies see Tumilson and Douglas (1992), and Frost and Timm (1992) as well as the aforementioned molecular studies. Frost and Timm (1992) provided generic diagnoses, but note that they included Idionycteris in Euderma.

References:

Bogdanowicz, W., S. Kasper, and R.D. Owen. 1998. Phylogeny of plecotine bats: reevaluation of morphological and chromosomal data. Journal of Mammalogy 79: 78-90. Read article.

Frost, D.R., and R.M. Timm 1992. Phylogeny of plecotine bats (Chiroptera: "Vespertilionidae"): summary of the evidence and proposal of a logically consistent taxonomy. American Museum Novitates 3034: 1-16. Read article.

Gray, J.E. 1866. Synopsis of the genera of Vespertilionidae and Noctilionidae. Annals and Magazine of Natural History ser. 3, 17(1866): 89-93. Read article.

Hoofer, S.R., and R.A. Van Den Bussche. 2001. Phylogenetic relationships of plecotine bats and allies based on mitochondrial ribosomal sequences. Journal of Mammalogy 82: 131-137. Read article.

Hoofer, S.R., and R.A. Van Den Bussche. 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica 5(suppl.): 1-63. Read abstract.

Pine, R.H., D.C. Carter, and R.K. LaVal. 1971. Status of Bauerus Van Gelder and its relationships to other nyctophiline bats. Journal of Mammalogy 52: 663-669. Read abstract.

Qumsiyeh, M.B., and J.W. Bickham. 1993. Chromosomes and relationships of long-eared bats of the genera Plecotus and Otonycteris. Journal of Mammalogy 74: 376-382. Read abstract.

Tumlison, R., and M.E. Douglas. 1992. Parsimony analysis and the phylogeny of plecotine bats (Chiroptera: Vespertilionidae). Journal of Mammalogy 73: 276-285. Read abstract.

Tate, 1942. Bull. Am. Mus. Nat. Hist., 80: 232

Glischropus, Nyctalus, Nycticeinops, Pipistrellus, Scotoecus, Scotozous, Vansonia.

Africa, Asia, Europe, Oceania

Includes Pipistrellus, Glischropus, Nyctalus, Scotozous (Volleth, 1992; Volleth and Heller, 1994; Volleth et al., 2001; Mayer and von Helversen, 2001, Koubínová et al., 2013) and Scotoecus (Amador et al., 2016; Roehrs et al., 2010, 2011; Volleth et al., 2006).

References:

Mayer, F., and O. von Helversen. 2001. Sympatric distribution of two cryptic bat species across Europe. Biological Journal of the Linnean Society 74: 365-374. Read article.

Hill & Harrison, 1987. Bull. Br. Mus. Nat. Hist. Zool., 52: 261

Scotophilus.

Africa, Asia

Only includes Scotophilus; see e.g., Amador et al. (2016); Hoofer and Van Den Bussche (2003); Roehrs et al. (2010).

References:

Hoofer, S.R., and R.A. Van Den Bussche. 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica 5(suppl.): 1-63. Read abstract.

Gray, 1821. London Med. Repos., 15: 299

Afronycteris, Cassistrellus, Chalinolobus, Falsistrellus, Hypsugo, Laephotis, Mimetillus, Mirostrellus, Neoromicia, Nycticeinops, Nyctophilus, Pharotis, Philetor, Pseudoromicia, Tylonycteris, Vespadelus, Vespertilio.

Africa, Asia, Europe, Oceania

Does not include Arielulus, Eptesicus, Hesperoptenus, Glischropus, Nyctalus, Pipistrellus, and Scotozous (Volleth, 1992; Volleth and Heller 1994; Volleth et al., 2001; Roehrs et al., 2010, 2011; Amador et al., 2016). Includes Hypsugo, Cassistrellus, Falsistrellus, Neoromicia, Nyctophilus, Tylonycteris, and Vespadelus (Volleth and Tidemann, 1991; Volleth and Heller, 1994, Volleth et al., 2001; Roehrs et al., 2010, 2011; Amador et al., 2016; Ruedi et al., 2018). Pharotis has not been included in recent analyses and is provisionally placed here. Roehrs et al. (2010) found that Vespertilio was more closely related to our tribe Pipistrellini and consequently named this the "Hypsugine group".

References:

Gray, J.E. 1821. On the natural arrangement of vertebrose animals. London Medical Repository 15: 296-310. Read article.

Ruedi, M., J.L. Eger, B.K. Lim, and G. Csorba. 2015. A new genus and species of vespertilionid bat from the Indomalayan region. Journal of Mammalogy 99(1): 209-222. Read article.

Volleth, M., and C.R. Tidemann. 1991. The origin of the Australian Vespertilioninae bats, as indicated by chromosomal studies. Zeitschrift für Säugetierkunde 56: 321-330. Read article.

Gray, 1821. London Med. Reposit., 15: 299

267

Ametrida, Amorphochilus, Anoura, Ardops, Ariteus, Artibeus, Brachyphylla, Carollia, Centurio, Chilonatalus, Chiroderma, Choeroniscus, Choeronycteris, Chrotopterus, Dermanura, Desmodus, Diaemus, Diphylla, Dryadonycteris, Ectophylla, Enchisthenes, Erophylla, Furipterus, Gardnerycteris, Glossophaga, Glyphonycteris, Hsunycteris, Hylonycteris, Lampronycteris, Leptonycteris, Lichonycteris, Lionycteris, Lonchophylla, Lonchorhina, Lophostoma, Macrophyllum, Macrotus, Mesophylla, Micronycteris, Mimon, Monophyllus, Mormoops, Musonycteris, Mystacina, Natalus, Neonycteris, Noctilio, Nyctiellus, Phylloderma, Phyllonycteris, Phyllops, Phyllostomus, Platalina, Platyrrhinus, Pteronotus, Pygoderma, Rhinophylla, Scleronycteris, Sphaeronycteris, Stenoderma, Sturnira, Thyroptera, Tonatia, Trachops, Trinycteris, Uroderma, Vampyressa, Vampyriscus, Vampyrodes, Vampyrum, Xeronycteris.

Americas, Oceania

Includes Mystacinidae, Furipteridae, and Thyropteridae. See Pierson (1986), Pierson et al. (1986), Kennedy et al. (1999), Van Den Bussche and Hoofer (2000, 2004), Agnarsson et al. (2011), and Amador et al. (2016). May include Myzopodidae; see that entry and Van Den Bussche and Hoofer (2004), Eick et al. (2005), Teeling et al. (2005), Miller-Butterworth et al. (2007), Agnarsson et al. (2011), Meredith et al. (2011), and Amador et al. (2016). For a phylogeny of the superfamily see Rojas et al. (2016). Rojas et al. (2018) updated distributional maps for species within Noctilionoidea. Teeling et al. (2005), Lim (2009), and Gunnell et al. (2014) discuss biogeographic patterns, but note that all these studies include Myzopodidae within Noctilionoidea.

References:

Gray, J.E. 1821. On the natural arrangement of vertebrose animals. London Medical Repository 15: 296-310. Read article.

Kennedy, M., A.M. Paterson, J.C. Morales, S. Parsons, A.M. Winnington, and H.G. Spencer. 1999. The long and the short of it: branch lengths and the problem of placing the New Zealand short-tailed bat, Mystacina. Molecular Phylogenetics and Evolution 13: 405-416. Read abstract.

Lim, B.K. 2009. Review of the origins and biogeography of bats in South America. Chiroptera Neotropical 15(1): 391-410. Read abstract.

Pierson, E.D., V.M. Sarich, J.M. Lowenstien, M.J. Daniel, and W.E. Rainey 1986. A molecular link between the bats of New Zealand and South America. Nature 323: 60-63. Read abstract.

Rojas, D., M. Moreira, M.J.R. Pereira, C. Fonseca, and L.M. Dávalos. 2018. Updated distribution maps for neotropical bats in the superfamily Noctilionoidea.. Ecology 99(9): 2131-2131. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Van Den Bussche, R.A., and S.R. Hoofer. 2000. Further evidence for inclusion of the New Zealand short-tailed bat (Mystacina tuberculata) within Noctilionoidea. Journal of Mammalogy 81: 865-874. Read article.

Gray, 1866. Ann. Mag. Nat. Hist., ser. 3, 17: 91

Amorphochilus, Furipterus.

Americas

References:

Gray, J.E. 1866. Synopsis of the genera of Vespertilionidae and Noctilionidae. Annals and Magazine of Natural History ser. 3, 17(1866): 89-93. Read article.

Saussure, 1860. Revue et Mag. Zool., 2: 286

Mormoops, Pteronotus.

Americas

Revised by Smith (1972). For phylogenies, see Lewis-Oritt et al. (2001), Simmons and Conway (2001), Van Den Bussche et al. (2002), Van Den Bussche and Weyandt (2003), Dávalos (2006), Pavan and Marroig (2016), and Morgan et al. (2019). See Smith (1972) for a discussion of authorship and priority of the name Mormoopidae.

References:

Dávalos, L.M. 2006. The geography of diversification in the mormoopids (Chiroptera: Mormoopidae). Biological Journal of the Linnean Society 88(1): 101-118. Read article.

Lewis-Oritt, N., C.A. Porter, and R.J. Baker 2001. Molecular systematics of the family Mormoopidae (Chiroptera) based on cytochrome b and recombination activating gene 2 sequences. Molecular Phylogenetic and Evolution 20(3): 426-436. Read article.

Morgan, G.S., N.J. Czaplewski, and N.B. Simmons 2019. A new mormoopid bat from the Oligocene (Whitneyan and Early Arikareean) of Florida, and phylogenetic relationships of the major clades of Mormoopidae (Mammalia: Chiroptera).. Bulletin of the American Museum of Natural History 434: 1-146. Read article.

Pavan, A.C., and G. Marroig. 2016. Integrating multiple evidences in taxonomy: Species diversity and phylogeny of mustached bats (Mormoopidae: Pteronotus). Molecular Phylogenetics and Evolution 103(2016): 184-198. Read article.

Saussure, M.H., de 1860. Notes sur quelques mammifères du Mexique. Revue et Magasin de Zoology Pure et Appliquée ser 2, v. 12: 281-293. Read article.

Simmons, N.B., and T.M. Conway 2001. Phylogenetic relationships of mormoopid bats (Chiroptera, Mormoopidae) based on morphological data. Bulletin of the American Museum of Natural History 258: 1-97. Read volume.

Smith, J.D. 1972. Systematics of the chiropteran family Mormoopidae. Miscellaneous Publication - University of Kansas, Museum of Natural History 56: 1-132. Read article.

Van Den Bussche, R., and S.E. Weyandt 2003. Mitochondrial and nuclear DNA sequence data provide resolution to sister-group relationships within Pteronotus (Chiroptera: Mormoopidae). Acta Chiropterologica 5(1): 1-13. Read abstract.

Van Den Bussche, R.A., S.R. Hoofer, and N.B. Simmons. 2002. Phylogenetic relationships of mormoopid bats using mitochondrial gene sequences and morphology. Journal of Mammalogy 83: 40-48. Read article.

Dobson, 1875. Ann. Mag. Nat. Hist., ser. 4, 16: 349

Mystacopidae Miller, 1907.

Mystacina.

Oceania

Monogeneric. See Dwyer (1962) for a discussion of the name Mystacopidae Miller, 1907.

References:

Dwyer, P.D. 1962. Studies on the two New Zealand bats. Zoology Publications from Victoria University of Wellington 28: 1-28. Read article.

Gray, 1821. London Med. Reposit., 15: 299

Noctilio.

Americas

Monogeneric.

References:

Gray, J.E. 1821. On the natural arrangement of vertebrose animals. London Medical Repository 15: 296-310. Read article.

Gray, 1825. Zool. Journ., 2(6): 242

227

Americas

Includes Desmodontidae; see Jones and Carter (1976). For use of Phyllostomidae rather than Phyllostomatidae, see Handley (1980). The classification used here generally follows that of Baker et al. (2016); see also Cirranello et al. (2016). See Baker et al. (1989), Van Den Bussche (1992), Wetterer et al. (2000), Baker et al. (2003), Datzmann et al. (2010), Botero-Casto etal. (2014), Dávalos et al. (2012, 2014), Amador et al. (2016), and Rojas et al. (2012, 2016) for phylogenies of the family. Wetterer et al. (2000) summarized historical background prior to 2000. Ferrarezi and Gimenez (1996), Wetterer et al. (2000), Schondube et al. (2001), Casotti et al. (2006), Rojas et al. (2011, 2016), Cruz-Neto et al. (2011), Baker et al. (2012), Rossoni et al. (2017), and Kries et al. (2018) discussed morphology and evolutionary patterns. Villalobos and Arita (2010) and Villalobos et al. (2014) examined biogeography.

References:

Baker, R.J., C.S. Hood, and R.L. Honeycutt. 1989. Phylogenetic relationships and classification of the higher categories of the New World bat family Phyllostomidae. Systematic Zoology 38: 228-238. Read article.

Baker, R.J., O.R.P. Bininda-Emonds, H. Mantilla-Meluk, C.A. Porter, and R.A. Van Den Bussche. 2012. Molecular timescale of diversification of feeding strategy and morphology in New World leaf-nosed bats (Phyllostomidae): a phylogenetic perspective.. Pages 385-409 In G.F. Gunnell and N.B. Simmons (eds.) Evolutionary history of bats: fossils, molecules and morphology. New York: Cambridge University Press. Read abstract.

Baker, R.J., S. Solari, A. Cirranello, and N.B. Simmons. 2016. Higher level classification of phyllostomid bats with a summary of DNA synapomorphies. Acta Chiropterologica 18(1): 1-38. Read abstract.

Baker, R.J., S.R. Hoofer, C.A. Porter, and R.A. Van Den Bussche. 2003. Diversification among New World Leaf-Nosed Bats: an evolutionary hypothesis and classification inferred from digenomic congruence of DNA sequence. Occasional Papers, Museum of Texas Tech University 230: 1-32. Read article.

Botero-Castro, F., M. Tilak, F. Justy, F. Catzeflis, F. Delsuc, and E.J.P. Douzery. . Next-generation sequencing and phylogenetic signal of complete mitochondrial genomes for resolving the evolutionary history of leaf-nosed bats (Phyllostomidae). Molecular Phylogenetics and Evolution 69(3): 728-739. Read abstract.

Casotti, G., L.G. Herrera M., J.J. Flores M., C.A. Mancina, E.J. Braun 2006. Relationships between renal morphology and diet in 26 species of New World bats (suborder Microchiroptera). Zoology 109(2006): 196-207. Read abstract.

Cirranello, A., N.B. Simmons, S. Solari, and R.J. Baker. 2016. Morphological diagnoses of higher-level phyllostomid taxa (Chiroptera: Phyllostomidae). Acta Chiropterologica 18(1): 39-71. Read abstract.

Cruz-Neto, A.P., T. Garland, Jr., and A.S. Abe. 2001. Diet, phylogeny, and basal metabolic rate in phyllostomid bats. Zoology 104(2001): 49-58. Read abstract.

Datzmann, T., O. von Helversen, and F. Mayer. 2010. Evolution of nectarivory in phyllostomid bats (Phyllostomidae Gray, 1825, Chiroptera: Mammalia). BMC Evolutionary Biology 10(1): 1-14. Read article.

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Dávalos, L.M., P.M. Velazco, O.M. Warsi, P.D. Smits, and N.B. Simmons. 2014. Integrating incomplete fossils by isolating conflicting signal in saturated and non-independent morphological characters. Systematic Biology 63(4): 582-600. Read article.

Ferrarezi H., and E.A. Gimenez. 1996. Systematic patterns and the evolution of feeding habits in Chiroptera (Archonta: Mammalia).. Journal of Comparative Biology 1(): 75-94. Not available online.

Gray, J.E. 1825. An attempt at a division of the family Vespertilionidae into groups. The Zoological Journal 2(6): 242-243. Read article.

Handley, C.O., Jr. 1980. Inconsistencies in formation of family-group and subfamily-group names in Chiroptera. Pages 9-13 In D.E. Wilson and A.L. Gardner (eds.) Proceedings of the Fifth International Bat Research Conference. Lubbock, Texas: Texas Tech University Press. Not available online.

Jones, J.K., Jr., and D.C. Carter 1976. Annotated checklist, with keys to subfamilies and genera. Pages 7-38 In R.J. Baker, J.K. Jones, Jr., and D.C. Carter (eds.) Biology of the bats of the New World Family Phyllostomatidae. Part I. Special Publications, The Museum, Texas Tech University 10: 218 pp. Read article.

Rojas, D., A. Vale, V. Ferrero, and L. Navarro. 2011. When did plants become important to leaf‐nosed bats? Diversification of feeding habits in the family Phyllostomidae. Molecular Ecology 20(10): 2217-2228. Read abstract.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Schondube, J.E., L.G. Herrera-M, and C.M. del Rio. 2001. Diet and the evolution of digestion and renal function in phyllostomid bats. Zoology 104(1): 59-73. Read abstract.

Van Den Bussche, R. 1992. Restriction site variation and molecular systematics of New World Leaf-Nosed Bats. Journal of Mammalogy 73(1): 29-42. Read abstract.

Villalobos, F., A. Lira-Noriega, J. Soberón, and H.T. Arita. . Co-diversity and co-distribution in phyllostomid bats: evaluating the relative roles of climate and niche conservatism. 15(1): 85-91. Read abstract.

Villalobos, F., and H.T. Arita. 2010. The diversity field of New World leaf‐nosed bats (Phyllostomidae). Global Ecology and Biogeography 19(2): 200-211. Read abstract.

Wetterer, A.L., M.V. Rockman, and N.B. Simmons. 2000. Phylogeny of phyllostomid bats (Mammalia: Chiroptera): data from diverse morphological systems, sex chromosomes, and restriction sites. Bulletin of the American Museum of Natural History 248: 1-200. Read article.

Van Den Bussche, 1992. J. Mammal., 73(1): 36

Macrotus.

Americas

Includes only Macrotus. Previously included in Phyllostominae (see Wetterer et al., 2000), but distinct; see Baker et al. (1989, 2000, 2003), Van Den Bussche (1992), and e.g., Rojas et al. (2016). For additional discussion see Cirranello et al. (2016) and Baker et al. (2016).

References:

Baker, R.J., C.A. Porter, J.C. Patton, and R.A. Van Den Bussche. 2000. Systematics of bats of the family Phyllostomidae based on RAG2 DNA sequences. Occasional Papers, The Museum, Texas Tech University 201: 1-16. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Van Den Bussche, R. 1992. Restriction site variation and molecular systematics of New World Leaf-Nosed Bats. Journal of Mammalogy 73(1): 29-42. Read abstract.

Van Den Bussche, 1992. J. Mammal., 73(1): 36

Lampronycteris, Micronycteris.

Americas

Does not include Glyphonycteris, Trinycteris, and Neonycteris, previously considered part of Micronycteris; see Simmons and Voss (1998) and Wetterer et al. (2000). Previously considered part of Phyllostominae (see Wetterer et al., 2000), but distinct, see Baker et al. (1989, 2000, 2003), Van Den Bussche (1992), and e.g., Rojas et al. (2016). For additional discussion see Cirranello et al. (2016) and Baker et al. (2016).

References:

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Simmons, N.B., and R.S. Voss 1998. The mammals of Paracou, French Guiana, a Neotropical lowland rainforest fauna. Part 1, Bats. Bulletin of the American Museum of Natural History 237: 1-219. Read article.

Van Den Bussche, R. 1992. Restriction site variation and molecular systematics of New World Leaf-Nosed Bats. Journal of Mammalogy 73(1): 29-42. Read abstract.

J.A. Wagner, 1840. Die Säugthiere in Abbildungen, p. 375

Desmodus, Diaemus, Diphylla.

Americas

See Handley (1980) for the correct derivation of the stem of this name (Desmodont-, not Desmod-). Although Handley attributed authorship of the family group name to Gill (1884), this use is predated by Wagner (1840). For additional discussion see Cirranello et al. (2016) and Baker et al. (2016).

References:

Gill. T. 1884. The Standard Natural History. S.E. Cassino, Boston: i-viii, 535. Read volume.

Wagner, J.A. 1840. Die Säugthiere in Abbildungen nach der Natur, mit Beschreibungen von Dr. Johann Christian Daniel von Schreber. Supplementband. Erste Abtheilung: Die Affen und Flederthiere. Expedition das Schreber'schen Säugthier-und des Esper'sschen Schmetterlingswerkes, und in Commission der Voss'schen Buchhandlung in Leipzig, Erlangen: i-xiv, i-vi + 551. Read volume.

Gray, 1866. Proc. Zool. Soc. Lond., 1866: 113

Lonchorhina.

Americas

Previously considered part of Phyllostominae (see e.g., Baker et al. 1989; Wetterer et al., 2000), but possibly distinct; see Baker et al. (2000, 2003), Rojas et al. (2016), but see Amador et al. (2016). For additional discussion see Cirranello et al. (2016) and Baker et al. (2016). For a discussion of relationships based on chromosome painting see Farias et al. (2021).

References:

Farias, J.C., N. Santos, D.P. Bezerra, and C.G. Sotero-Caio. 2012. Chromosome painting in Lonchorhina aurita sheds light onto the controversial phylogenetic position of sword-nosed bats (Chiroptera, Phyllostomidae). Cytogenetic and Genome Research 2021(161): 569-577. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Gray, 1825. Zool. Journ., 2(6): 242

Chrotopterus, Gardnerycteris, Lophostoma, Macrophyllum, Mimon, Phylloderma, Phyllostomus, Tonatia, Trachops, Vampyrum.

Americas

Does not include Macrotus, Micronycteris, Glyphonycteris, Lampronycteris, Lonchorhina, Neonycteris, or Trinycteris; see Baker et al. (2003), Dávalos et al. (2012, 2014). Baker et al. (1989) applied the name Phyllostominae broadly to a clade that included most of the family, but this usage has not been widely followed. We recognize three tribes within this subfamily: Macrophyllini, Phyllostomini, and Vampyrini. See Cirranello et al. (2016) and Baker et al. (2016) for additional discussion.

References:

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Gray, J.E. 1825. An attempt at a division of the family Vespertilionidae into groups. The Zoological Journal 2(6): 242-243. Read article.

Gray, 1866. Proc. Zool. Soc. Lond., 1866: 113

Macrophyllum, Trachops.

Americas

Originally comprised of Macrophyllum only. Expanded by Baker et al. (2003) to include Trachops; see also Rojas et al., (2016); Dávalos et al., (2012), and Amador et al. (2016). See Cirranello et al. (2016) and Baker et al. (2016) for additional discussion.

References:

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Gray, J.E. 1866. Revision of the genera of Phyllostomidae, or leaf-nosed bats. Proceedings of the Zoological Society of London 1866: 111-118. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Gray, 1825. Zool. Journ., 2(6): 242

Gardnerycteris, Lophostoma, Phylloderma, Phyllostomus, Tonatia.

Americas

Does not include Mimon, Lonchorhina, or Macrophyllum; see Rojas et al. (2016), Dávalos et al., (2012), and Amador et al. (2016). For additional discussion see Cirranello et al. (2016) and Baker et al. (2016).

References:

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Gray, J.E. 1825. An attempt at a division of the family Vespertilionidae into groups. The Zoological Journal 2(6): 242-243. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Bonaparte, 1838. Nuovi Ann. Sci. Nat. Bologna, 1(2): 112

Chrotopterus, Mimon, Vampyrum.

Americas

Used at the subfamily level by Baker et al. (1989) and as a tribe by Wetterer et al. (2000). Includes Mimon s.s.; does not include Trachops or Tonatia s.l.; see Rojas et al. (2016) and Dávalos et al., (2014), as these studies are the most taxonomically comprehensive to date. See Cirranello et al. (2016) and Baker et al. (2016) for additional discussion.

References:

Bonaparte, C.L. 1838. Synopsis Vertebratorum Systematis. Nuovi Annali delle Scienze Naturali, Bologna 1(2): 1-390. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Bonaparte, 1845. Cat. Met. Mamm. Europe, 5

Anoura, Brachyphylla, Choeroniscus, Choeronycteris, Dryadonycteris, Erophylla, Glossophaga, Hylonycteris, Leptonycteris, Lichonycteris, Monophyllus, Musonycteris, Phyllonycteris, Scleronycteris.

Americas

Includes Brachyphylla, Phyllonycteris, and Erophylla. Does not include lonchophyllines. See Griffiths (1982), Baker et al. (2003), Datzmann et al. (2010), Rojas et al. (2016), but see also Wetterer et al. (2000), Carstens et al. (2002) and Dávalos et al. (2012). We recognize three tribes within this subfamily: Brachyphyllini, Choeronycterini, and Glossophagini. Morphology, biology, and evolution reviewed by Solmsen (1998). See Dávalos et al. (2012), Cirranello et al. (2016), and Baker et al. (2016) for additional discussion.

References:

Bonaparte, C.L. 1845. Catalogo metodico dei mammiferi Europei. L. di Giacomo Pirola, Milano: 36 pp. Read book.

Carstens, B.C., B.L. Lundrigan, and P. Myers. 2002. A phylogeny of the Neotropical nectar-feeding bats (Chiroptera: Phyllostomidae) based on morphological and molecular data. Journal of Mammalian Evolution 9: 23-53. Read abstract.

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Griffiths, T.A 1982. Systematics of the New World nectar-feeding bats (Mammalia, Phyllostomidae), based on the morphology of the hyoid and lingual regions. American Museum Novitates 2742: 1-45. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Solmsen, E.H. 1998. New World nectar-feeding bats: biology, morphology, and craniometric approach to systematics. Bonner Zoologische Mongraphien 44: 1-118. Read article.

Solmsen, 1998. Bonn . Zool. Monogr., 44: 97

Anoura, Choeroniscus, Choeronycteris, Dryadonycteris, Hylonycteris, Lichonycteris, Musonycteris, Scleronycteris.

Americas

"Choeronycterine" was first used informally by Allen (1898). See Cirranello et al. (2016) for a full discussion of the authorship of this name. Solmsen (1998) included Choeroniscus, Choeronycteris (with Musonycteris as a subgenus), and Hylonycteris in Choeronycterini. Carstens et al. (2002) used the name infomally, but applied the name to the taxa we have included here. See Wetterer et al. (2000), Baker et al., (2003); Datzmann et al., (2010); Dávalos et al., (2012); and Rojas et al., (2016).

References:

Allen, H.A. 1898. On the Glossophaginae. Transactions of the American Philosophical Society 19(1898): 237-266. Read abstract.

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Solmsen, E.H. 1998. New World nectar-feeding bats: biology, morphology, and craniometric approach to systematics. Bonner Zoologische Mongraphien 44: 1-118. Read article.

Gray, 1866. Proc. Zool. Soc. Lond., 1866: 115

Brachyphylla, Erophylla, Phyllonycteris.

Americas

Includes Phyllonycteris and Erophylla; see Taboada and Pine (1969), Baker and Lopez (1970), Baker and Bass (1979), Baker et al. (1981, 2003), Carstens et al. (2002), Datzmann et al. (2010), Dávalos et al. (2012, 2014), and Rojas et al. (2016); but see Wetterer et al. (2000). Appears most closely related to Glossophagini; see e.g., Amador et al. (2016). See Cirranello et al. (2016) and Baker et al. (2016) for additional discussion.

References:

Baker, R. J., and R. A. Bass. 1979. Evolutionary relationship of the Brachyphyllinae to the glossophagine genera Glossophaga and Monophyllus. Journal of Mammalogy 60(2): 364-372. Read abstract.

Baker, R.J., and G. Lopez. 1970. Karyotypic studies of the insular populations of bats on Puerto Rico. Caryologia 23: 465-472. Read article.

Baker, R.J., H.H. Genoways, and P.A. Seyfarth. 1981. Results of the Alcoa Foundation-Suriname expeditions. VI. Additional chromosomal data for bats (Mammalia: Chiroptera) from Suriname. Mammalogy Papers, University of Nebraska State Museum 230: 333–344. Read article.

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Gray, J.E. 1866. Revision of the genera of Phyllostomidae, or leaf-nosed bats. Proceedings of the Zoological Society of London 1866: 111-118. Read article.

Peters, W.C.H. 1861. Hrn. Dr. Gundlach beschriebene Gattung von Flederthieren aus Cuba, Phyllonycteris. Monatsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin 1860: 817-819. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Taboada, G.S., and R.H. Pine. 1969. Morphological and behavioral evidence for the relationship between the bat genus Brachyphylla and the Phyllonycterinae. Biotropica 1: 10-19. Read abstract.

Bonaparte, 1845. Cat. Met. Mamm. Europe, 5

Glossophaga, Leptonycteris, Monophyllus.

Americas

Recognized informally by Carstens et al. (2002). Named as a tribe by Baker et al. (2003). See also Datzmann et al., (2010); Dávalos et al., (2012, 2014); Rojas et al., (2016); but see Wetterer et al. (2000). Apparently more closely related to Brachyphyllini than to Choeronycterini; see e.g., Amador et al. (2016).

References:

Bonaparte, C.L. 1845. Catalogo metodico dei mammiferi Europei. L. di Giacomo Pirola, Milano: 36 pp. Read book.

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Griffiths, 1982. Am. Mus. Novitates, 2742: 43

Hsunycteris, Lionycteris, Lonchophylla, Platalina, Xeronycteris.

Americas

Equivalent to the tribe Lonchophyllini of McKenna and Bell (1997) and Simmons (2005). Previously included in Glossophaginae, but distinct; see Griffiths (1982), Baker et al. (2003), Datzmann et al. (2010), and Rojas et al. (2016). Morphology, biology, and evolution reviewed by Solmsen (1998). For phylogenies see Dávalos and Jansa (2004), Woodman and Timm (2006), and Parlos et al. (2014). See Dávalos et al. (2012), Cirranello et al. (2016), and Baker et al. (2016) for additional discussion.

References:

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Dávalos, L.M., and S.A. Jansa 2004. Phylogeny of the Lonchophyllini (Chiroptera: Phyllostomidae). Journal of Mammalogy 85(3): 404-413. Read article.

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Parlos, J.A., R.M. Timm, V.J. Swier, H. Zeballos, and R.J. Baker 2014. Evaluation of the paraphyletic assemblages within Lonchophyllinae, with description of a new tribe and genus. Occasional Papers, Museum of Texas Tech University 320: 1-23. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Solmsen, E.H. 1998. New World nectar-feeding bats: biology, morphology, and craniometric approach to systematics. Bonner Zoologische Mongraphien 44: 1-118. Read article.

Woodman, N., and R.M. Timm 2006. Characters and phylogenetic relationships of nectar-feeding bats, with descriptions of new Lonchophylla from western South America (Mammalia: Chiroptera: Phyllostomidae: Lonchophyllini). Proceedings of the Biological Society of Washington 119(4): 437-477. Read article.

Baker, Solari, Cirranello & Simmons, 2016. Acta Chiropterol., 18(1): 16

Glyphonycterinae Baker, Hoofer, Porter & Van Den Bussche, 2003 [nomen nudum].

Glyphonycteris, Neonycteris, Trinycteris.

Americas

Previously included within Micronycteris (sensu Sanborn, 1949) and Phyllostominae, but distinct as both genera (see Simmons and Voss, 1998; Wetterer et al., 2000) and as a subfamily (Baker et al., 2003). See Dávalos et al. (2014) who confirmed the inclusion of Neonycteris within this subfamily. For additional discussion see Cirranello et al. (2016) and Baker et al. (2016).

References:

Sanborn, C.C. 1949. Bats of the genus Micronycteris and its subgenera. Fieldiana: Zoology 31: 215-233. Read article.

Miller, 1924. Bull. U.S. Natl. Mus., 128: 53

Hemiderminae Miller, 1907.

Carollia.

Americas

Does not include Rhinophylla. Restricted to the genus Carollia by Baker et al. (2003). See Cirranello et al. (2016) and Baker et al. (2016) for additional discussion.

References:

Miller, G.S., Jr. 1907. The families and genera of bats. Bulletin of the U.S. National Museum 57: 1-282. Read volume.

Miller, G.S., Jr. 1924. List of North American Recent mammals 1923. Bulletin of the U.S. National Museum 128: 1-673. Read article.

Baker, Solari, Cirranello & Simmons, 2016. Acta Chiropterol., 18(1): 17

Rhinophyllinae Baker, Hoofer, Porter & Van Den Bussche, 2003 [nomen nudum].

Rhinophylla.

Americas

Previously included in Carolliinae, but distinct; see Baker et al. (2003). For additional discussion see Cirranello et al. (2016) and Baker et al. (2016).

References:

Gervais, 1856. In Comte de Castelnau, Exped. Partes Cen. Am. Sud., Zool. (Sec. 7), Vol. 1, pt. 2 (Mammiferes), p. 32n

101

Americas

Equivalent to tribe Stenodermatini of McKenna and Bell (1997) and Baker et al. (2003). McKenna and Bell (1997) included Carollia and Rhinophylla as a tribe (Carolliini) within their subfamily Stenodermatinae. Sturnira has occasionally been considered as a distinct subfamily (Sturnirinae Miller, 1907; Baker, 1967), but we continue to recognize it at the tribal level within Stenodermatinae. See Cirranello et al. (2016) and Baker et al. (2016) for additional discussion. See Baker (1967), Gerber, (1968), Gerber and Leone (1971), Wetterer et al. (2000), Baker et al. (2003), Dávalos et al. (2012), Amador et al. (2016), and Rojas et al. (2016). Phylogenetic relationships have been discussed by Owen (1987, 1991), Lim (1993), Van Den Bussche et al. (1993), Wetterer et al. (2000), and Baker et al. (2000); roosting ecology by Garbino and Tavares (2018).

References:

Baker, R.J. 1967. Karyotypes of bats of the family Phyllostomidae and their phylogenetic implications. Southwestern Naturalist 12: 407-428. Read abstract.

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Garbino, G.S.T., and V. da C. Tavares. 2018. Roosting ecology of Stenodermatinae bats (Phyllostomidae): evolution of foliage roosting and correlated phenotypes. Mammal Review 48(2): 75-89. Read abstract.

Gerber, J.D. 1968. Electrophoretic and immunological comparisons of the serum proteins of bats (Unpublished Doctoral Dissertation). University of Kansas Lawrence. Not available online.

Gerber, J.D., and C.A. Leone. 1971. Immunologic comparisons of the sera of certain phyllostomatid bats. Systematic Zoology 20(2): 160-166. Read abstract.

Lim, B.K. 1993. Cladistic reappraisal of Neotropical stenodermatine bat phylogeny. Cladistics 9: 147-165. Read abstract.

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Miller, G.S., Jr. 1907. The families and genera of bats. Bulletin of the U.S. National Museum 57: 1-282. Read volume.

Owen, R.D. 1987. Phylogenetic analyses of the bat subfamily Stenodermatinae (Mammalia: Chiroptera). Special Publications, The Museum, Texas Tech University 26: 1-65. Read article.

Owen, R.D. 1991. The systematic status of Dermanura concolor (Peters, 1865)(Chiroptera: Phyllostomidae), with description of a new genus. Bulletin of the American Museum of Natural History 206: 18-25. .

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Van Den Bussche, R.A., R.J. Baker, H.A. Wichman, and M.J. Hamilton. 1993. Molecular phylogenetics of stenodermatine bat genera: congruence of data from nuclear and mitochondrial DNA. Molecular Biology and Evolution 10: 944-959. .

Miller, 1907. Bull. U.S. Natl. Mus., 57: 38

Sturnira.

Americas

Monogeneric. Equivalent to subtribe Sturnirina of McKenna and Bell (1997). Owen (1987) recognized Corvirini for the genus Corvira and Sturnirini for Sturnira sensu stricto, but Corvira does not appear to be monophyletic; see Pacheco and Patterson (1991), Velazco and Patterson (2013), Amador et al. (2016), and Rojas et al. (2016). See Cirranello et al. (2016) and Baker et al. (2016) for additional discussion.

References:

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Miller, G.S., Jr. 1907. The families and genera of bats. Bulletin of the U.S. National Museum 57: 1-282. Read volume.

Owen, R.D. 1987. Phylogenetic analyses of the bat subfamily Stenodermatinae (Mammalia: Chiroptera). Special Publications, The Museum, Texas Tech University 26: 1-65. Read article.

Pacheco, V., and B.D. Patterson. 1991. Phylogenetic relationships of the New World bat genus Sturnira (Chiroptera, Phyllostomidae). Pages 101-121 In T.A. Griffiths and D. Klingener (eds.) Contributions to Mammalogy in Honor of Karl F. Koopman. Bulletin of the American Museum of Natural History 206: 432 pp. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Velazco, P.M., and B.D. Patterson. 2013. Diversification of the yellow-shouldered bats, genus Sturnira (Chiroptera, Phyllostomidae), in the New World tropics. Molecular Phylogenetics and Evolution 68(3): 683-698. Read abstract.

Gervais, 1856. In Comte de Castelnau, Exped. Partes Cen. Am. Sud., Zool. (Sec. 7), Vol. 1, pt. 2 (Mammiferes), p. 32n

Ametrida, Ardops, Ariteus, Artibeus, Centurio, Chiroderma, Dermanura, Ectophylla, Enchisthenes, Mesophylla, Phyllops, Platyrrhinus, Pygoderma, Sphaeronycteris, Stenoderma, Uroderma, Vampyressa, Vampyriscus, Vampyrodes.

Americas

Equivalent to the subtribe Stenodermatina of McKenna and Bell (1997). See Owen (1987), Wetterer et al. (2000), Baker et al. (2003), Dávalos et al. (2012), Amador et al. (2016), and Rojas et al. (2016); but see Lim (1993) and Van Den Bussche et al. (1993). The subtribal classification used here follows Cirranello et al. (2016) and Baker et al. (2016).

References:

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Lim, B.K. 1993. Cladistic reappraisal of Neotropical stenodermatine bat phylogeny. Cladistics 9: 147-165. Read abstract.

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Owen, R.D. 1987. Phylogenetic analyses of the bat subfamily Stenodermatinae (Mammalia: Chiroptera). Special Publications, The Museum, Texas Tech University 26: 1-65. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Baker, Solari, Cirranello & Simmons, 2016. Acta Chiropterol., 18(1): 29

Vampyressatini Owen, 1987 [nomen nudum]; Vampyressina Baker, Hoofer, Porter & Van Den Bussche, 2003 [nomen nudum].

Chiroderma, Mesophylla, Platyrrhinus, Uroderma, Vampyressa, Vampyriscus, Vampyrodes.

Americas

Does not include Ectophylla; see Baker et al. (2003), Hoofer and Baker (2006), Dávalos et al. (2012), Amador et al. (2016), and Rojas et al. (2016); but see Lim (1993), Lim et al. (2003), and Wetterer et al. (2000). Owen (1987) applied the name 'Vampyressatini' to a group of Vampyressa pusilla, V. brocki, and V. bidens (the latter two species now belong in the genus Vampyriscus). Baker et al. (2003) later emended the name to Vampyressina and applied it to the group as here defined; however neither usage of the name meets the requirements of the ICZN for availability (ICZN, 1999). See Cirranello et al. (2016) and Baker et al. (2016) for additional discussion. For phylogenies of the group see the aforementioned studies and Hoofer and Baker (2006), Hoofer et al. (2008), and Gomes et al. (2016).

References:

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Gomes, A.J.B., C.Y. Nagamachi, L.R.R. Rodrigues, T.C.M. Benathar, T.F.A. Ribas, P.C.M. O’Brien, F. Yang, M.A. Ferguson-Smith, and J.C. Pieczarka 2016. Chromosomal phylogeny of vampyressine bats (Chiroptera, Phyllostomidae) with description of two new sex chromosome systems. BMC Evolutionary Biology 16(119): 1-11. Read article.

Hoofer, S.R., and R.J. Baker. 2006. Molecular systematics of vampyressine bats (Phyllostomidae: Stenodermatinae) with comparison of direct and indirect surveys of mitochondrial DNA variation. Molecular Phylogenetics and Evolution 39(2): 424–438. Read abstract.

Hoofer, S.R., W.E. Flanary, R.J. Bull, and R.J. Baker. 2008. Phylogenetic relationships of vampyressine bats and allies (Phyllostomidae: Stenodermatinae) based on DNA sequences of a nuclear intron (TSHB-I2). Molecular Phylogenetics and Evolution 47(2): 870-876. Read abstract.

Lim, B.K. 1993. Cladistic reappraisal of Neotropical stenodermatine bat phylogeny. Cladistics 9: 147-165. Read abstract.

Lim, B.K., W.A. Pedro, and F.C. Passos. 2003. Differentiation and species status of the Neotropical yellow-eared bats Vampyressa pusilla and V. thyone (Phyllostomidae) with a molecular phylogeny of the genus. Acta Chiropterologica 5: 15-29. Read abstract.

Owen, R.D. 1987. Phylogenetic analyses of the bat subfamily Stenodermatinae (Mammalia: Chiroptera). Special Publications, The Museum, Texas Tech University 26: 1-65. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Baker, Solari, Cirranello & Simmons, 2016. Acta Chiropterol., 18(1): 29

Enchistheneini Owen, 1987 [nomen nudum]; Enchisthenina Baker, Hoofer, Porter & Van Den Bussche, 2003 [nomen nudum].

Enchisthenes.

Americas

Formerly considered part of Artibeus or Ectophyllina sensu Wetterer et al. (2000); but distinct; see Owen (1987), Baker et al. (2003), Dávalos et al. (2012), and Rojas et al. (2016). Owen (1987) first applied the name "Enchistheneini" at the subtribal level for Enchisthenes only, a usage followed by Baker et al. (2003); however neither name meets the requirements of the ICZN for availability (ICZN, 1999). See Cirranello et al. (2016) and Baker et al. (2016) for additional discussion.

References:

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Owen, R.D. 1987. Phylogenetic analyses of the bat subfamily Stenodermatinae (Mammalia: Chiroptera). Special Publications, The Museum, Texas Tech University 26: 1-65. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Baker, Solari, Cirranello & Simmons, 2016. Acta Chiropterol., 18(1): 30

Ectophyllina Wetterer, Rockman & Simmons, 2000 [nomen nudum]; Ectophyllina Baker, Hoofer, Porter & Van Den Bussche, 2003 [nomen nudum].

Ectophylla.

Americas

Restricted to Ectophylla by Baker et al. (2003), but the name was not made available until Baker et al. (2016). See Cirranello et al. (2016) and Baker et al. (2016) for additional discussion. See Baker et al. (2003), Dávalos et al. (2012), and Rojas et al. (2016), but see also Wetterer et al. (2000). Wetterer et al. (2000) applied the name Ectophyllina at the subtribal level to the group colloquially known as the "long-faced" stenodermatines (and see Gardner [2007] for a similar usage at the tribal level), applications of this name that are not equivalent to our usage here.

References:

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Gardner, A.L. 2008 [dated 2007]. Subfamily Stenodermatinae. Pages 300-301 In A.L. Gardner (eds.) Mammals of South America, Volume I. Chicago: University of Chicago Press. Read abstract.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Allen, 1898. Trans. Am. Philos. Soc., 19(1898): 269

Artibeus, Dermanura.

Americas

Restricted to Artibeus (Artibeus) (including Koopmania - a synonym) and Artibeus (Dermanura), by Baker et al. (2003); see Cirranello et al. (2016) and Baker et al. (2016) for additional discussion. Does not include Enchisthenes; see Wetterer et al. (2000), Baker et al. (2003), Dávalos et al. (2012), and Rojas et al. (2016). In addition to the original usage by Allen (1898), Owen (1987) used the name "Artibeini" at both the tribal and subtribal level, but none of these groups are equivalent to our usage here.

References:

Allen, H. 1898. The skull and teeth of Ectophylla alba. Transactions of the American Philosophical Society 19(1898): 267–272. Read abstract.

Owen, R.D. 1987. Phylogenetic analyses of the bat subfamily Stenodermatinae (Mammalia: Chiroptera). Special Publications, The Museum, Texas Tech University 26: 1-65. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Gervais, 1856. In Comte de Castelnau, Exped. Partes Cen. Am. Sud., Zool. (Sec. 7), Vol. 1, pt. 2 (Mammiferes), p. 32n

Ametrida, Ardops, Ariteus, Centurio, Phyllops, Pygoderma, Sphaeronycteris, Stenoderma.

Americas

Equivalent to tribe Stenodermatini of Owen (1987). See Lim (1993), Wetterer et al. (2000), Baker et al. (2003), Dávalos et al. (2012), Amador et al. (2016), and Rojas et al. (2016). See Cirranello et al. (2016) and Baker et al. (2016) for additional discussion. Dávalos (2007) and Tavares et al. (2018) reviewed the biogeography of this group.

References:

Dávalos, L.M. 2007. Short-faced bats (Phyllostomidae: Stenodermatina): a Caribbean radiation of strict frugivores. Journal of Biogeography 34(): 364-375. Read abstract.

Dávalos, L.M., A.L. Cirranello, J.H. Geisler, and N.B. Simmons. 2012. Understanding phylogenetic incongruence: lessons from phyllostomid bats. Biological Reviews 87(): 991-1024. Read article.

Lim, B.K. 1993. Cladistic reappraisal of Neotropical stenodermatine bat phylogeny. Cladistics 9: 147-165. Read abstract.

Owen, R.D. 1987. Phylogenetic analyses of the bat subfamily Stenodermatinae (Mammalia: Chiroptera). Special Publications, The Museum, Texas Tech University 26: 1-65. Read article.

Rojas, D., O.M. Warsi, and L.M. Dávalos. 2016. Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant neotropical diversity. Systematic Biology 65(3): 432-448. Read article.

Tavares, V. da C., O.M. Warsi, F. Balseiro, C.A. Mancina, L.M. Dávalos 2018. Out of the Antilles: fossil phylogenies support reverse colonization of bats to South America. Journal of Biogeography 45(4): 859-873. Read abstract.

Miller, 1907. Bull. U.S. Natl. Mus., 57: 16, 190

Thyroptera.

Americas

Monogeneric.

References:

Miller, G.S., Jr. 1907. The families and genera of bats. Bulletin of the U.S. National Museum 57: 1-282. Read volume.

Springer, Teeling, Madsen, Stanhope & de Jong, 2001. Proc. Natl. Acad. Sci. USA, 98: 6243

428

Africa, Asia, Europe, Oceania

The name Yinpterochiroptera is a combination of Yinochiroptera (coined by Koopman [1985] for a group of microchiropteran bats that included rhinolophoids and nycterids) and Pteropodidae. In addition to DNA hybridization data (see Hutcheon et al., 1998; Kirsch and Pettigrew, 1998), a unique amino acid apomorphy (Springer et al., 2001) and a karyotypic inversion that appears to be unique to Yinpterochiroptera (Ao et al., 2007) analyses of more than 13 kilobases of DNA sequence from 20 nuclear and 4 mitochondrial genes (Porter et al., 1996; Teeling et al. 2000, 2002, 2003, 2005; Springer et al., 2001; Hulva and Horacek, 2002; Van Den Bussche and Hoofer, 2004; Van Den Bussche et al., 2002; Miller-Butterworth et al., 2007; Agnarsson et al. 2011, Meredith et al. 2011, Amador et al. 2016), have all recovered Yinpterochiroptera, often with very high support values (e.g., bootstrap, Bremer values, posterior probabilities). The sole exception, a recent phylogeny using only the cytochrome b gene (Agnarsson et al. 2011), weakly rejects Yinpterochiroptera when the full data set is used, but supports Yinpterochiroptera monophyly under a pruned data set. In contrast, morphological data (see e.g., Simmons 1998; Simmons and Geisler, 1998; Gunnell and Simmons, 2005) support the traditional split between non-echolocating Pteropodidae (Megachiroptera) and echolocating Microchiroptera, including the members of Rhinolophoidea. Intriguingly, two combined analyses of morphological and molecular data (Hermsen and Hendricks, 2008; O'Leary et al., 2013) found support for Megachiroptera and Microchiroptera.
We use Yinpterochiroptera rather than Pteropodiformes (Hutcheon and Kirsch, 2006) because the former has been in use has been more widely used to refer to this clade.

References:

Ao, L., X. Mao, W. Nie, X. Gu, Q. Feng, J. Wang, W. Su, Y. Wang, M. Volleth, and F. Yang. 2007. Karyotypic evolution and phylogenetic relationships in the order Chiroptera as revealed by G-banding comparison and chromosome painting. Chromosome Research 15 (3): 257–267. Read abstract.

Gunnell, G.F., and N.B.Simmons 2005. Fossil evidence and the origin of bats. Journal of Mammalian Evolution 12: 209-246. Read article.

Hutcheon, J. M., and J. A. W. Kirsch. 2006. A moveable face: deconstructing the Microchiroptera and a new classification of extant bats. Acta Chiropterologica 8(1): 1-10. Read abstract.

Kirsch, J. A. W., and J. D. Pettigrew. 1998. Base-compositional biases and the bat problem. II. DNA hybridization trees based on AT and GC- enriched tracers. Philosophical Transactions of the Royal Society B Biological Sciences 353(1367): 381-388. . Read abstract.

Koopman, K.F. 1985. A synopsis of the families of bats, Part VII. Bat Research News 25(3/4): 25-27. Not available online.

Porter, C. A., M. Goodman, and M. J. Stanhope. 1996. Evidence on mammalian phylogeny from sequences of exon 28 of the von Willebrand factor gene. Molecular Phylogenetics and Evolution 5(1): 89-101. Read abstract.

Springer, M.S., E.C. Teeling, O. Madsen, M.J. Stanhope, and W.W. de Jong. 2001. Integrated fossil and molecular data reconstruct bat echolocation. Proceedings National Academy of Sciences 98: 6241-6246. .

Teeling, E.C., O.Madsen, W.J.Murphy, M.S.Springer, and S.J. O'Brien. 2003. Nuclear gene sequences confirm an ancient link between New Zealand's short-tailed bat and South American Noctilionoid bats. Molecular Phylogenetics and Evolution 28(2): 308-319. .

Gray, 1821. London Med. Repos., 15: 299

Cephalotidae Gray, 1821; Harpyidae C.H. Smith, 1842.

202

Africa, Asia, Europe, Oceania

THIS ENTRY IS UNDER REVIEW. Various workers have recognized between two and six subfamilies of Pteropodidae including: Cynopterinae Andersen, 1912, Epomophorinae K. Andersen, 1912, Harpionycterinae Miller, 1907, Nyctimeninae Miller, 1907, Macroglossinae Gray, 1866, Rousettinae Andersen, 1912, and Pteropodinae Gray, 1821 (Bergmans, 1997; Corbet and Hill, 1980, 1992; Hill and Smith, 1984; Koopman, 1993, 1994; McKenna and Bell, 1997). Recent phylogenetic studies agree that Macroglossinae and Pteropodinae sensu Koopman (1993, 1994) and McKenna and Bell (1997) are not monophyletic (Alvarez et al., 1999; Giannini and Simmons, 2003; Hollar and Springer, 1997; Hood, 1989; Juste et al., 1997; Kirsch et al., 1995; Romagnoli and Springer, 2000; Springer et al., 1995). Monophyly of cynopterines and empomophorines has also been questioned (Alvarez et al., 1999; Hollar and Springer, 1997; Kirsch et al., 1995; Romagnoli and Springer, 2000). Instead of supporting traditional taxonomic groupings, phylogenetic studies based on DNA hybridization and DNA sequences have found support for a large clade of endemic African taxa including genera previously placed in several different subfamilies/tribes (Alvarez et al., 1999; Giannini and Simmons, 2003; Hollar and Springer, 1997; Kirsch et al., 1995; Romagnoli and Springer, 2000). Relationships among pteropodid genera are not yet fully resolved, however, and questions remain concerning the position of Nyctimene, Paranyctimene, Eidolon, and several SE Asian endemic genera. Existing subfamilial and tribal classifications are not adequately congruent with recent phylogenies. Accordingly, no subfamilial or tribal groups are recognized here pending a thorough reevaluation of pteropodid classification.

References:

Gray, J.E. 1821. On the natural arrangement of vertebrose animals. London Medical Repository 15: 296-310. Read article.

Gray, 1866. Proc. Zool. Soc. Lond., 1866: 64

Aethalops, Alionycteris, Balionycteris, Chironax, Cynopterus, Dyacopterus, Haplonycteris, Latidens, Megaerops, Otopteropus, Penthetor, Ptenochirus, Sphaerias, Thoopterus.

Asia

THIS ENTRY IS BEING REVISED. Two tribes are recognized: Cynopterini and Balionycterini. See Almeida et al. (2009) for phylogenetic and biogeographic analyses.

References:

Gray, J.E. 1866. A revision of the genera of pteropine bats (Pteropidae) and the description of some apparently undescribed species. Proceedings of the Zoological Society of London 1866: 62-67. Read article.

Gray, 1866. ,

Cynopterus, Megaerops, Ptenochirus.

Asia

References:

, 0. ,

Aethalops, Alionycteris, Balionycteris, Chironax, Dyacopterus, Haplonycteris, Latidens, Otopteropus, Penthetor, Sphaerias, Thoopterus.

Asia

References:

Almeida, Giannini & Simmons, 2016. Acta Chiropterol., 18(1): 84

Eidolon.

Africa, Asia

THIS ENTRY IS BEING REVISED. Previously included in Rousettinae (Bergmans, 1997; or its equivalent e.g., McKenna and Bell, 1997), but distinct. See also Juste et al. (1999), Romagnoli and Springer (2000), Colgan and da Costa (2002), Giannini and Simmons (2005), and Almeida et al. (2011).

References:

Almeida, F.C., N.P. Giannini, and N.B. Simmons. 2016. The evolutionary history of the African fruit bats (Chiroptera: Pteropodidae). Acta Chiropterologica 18(1): 73-90. Read abstract.

Almeida, F.C., N.P. Giannini, R. DeSalle, and N.B. Simmons. 2011. Evolutionary relationships of the Old World fruit bats (Chiroptera, Pteropodidae): another star phylogeny?. BMC Evolutionary Biology 11(1): 281. Read article.

Bergmans, W. 1997. Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera).5. The genera Lissonycteris Andersen, 1912, Myonycteris Matschie, 1899, and general remarks and conclusions; annex: key to all species. Beaufortia 47: 11-90. Read article.

Colgan, D.J., and P. da Costa 2002. Megachiropteran evolution studied with 12S rDNA and c-mos DNA sequences. Journal of Mammalian Evolution 9(1-2): 3-22. Read abstract.

Giannini, N.P., and N.B. Simmons 2005. Conflict and congruence in a combined DNA–morphology analysis of megachiropteran bat relationships (Mammalia: Chiroptera: Pteropodidae). Cladistics 21: 411-437. Read abstract.

Juste, J., Y. Álvarez, E. Tabarés, A. Garrido-Pertierra, C. Ibáñez, and J.M. Bautista. 1999. Phylogeography of African fruitbats (Megachiroptera). Molecular Phylogenetics and Evolution 13(3): 596-604. Read abstract.

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Romagnoli, M.L., and M.S. Springer. 2000. Evolutionary relationships among Old World fruitbats (Megachiroptera: Pteropodidae) based on 12S rRNA, tRNA Valine, and 16S rRNA gene sequences. Journal of Mammalian Evolution 7: 259-284. Read abstract.

Miller, 1907. ,

Aproteles, Boneia, Dobsonia, Harpyionycteris.

Asia, Oceania

THIS ENTRY IS UNDER REVIEW.

References:

, . ,

References:

, . ,

References:

Almeida, Simmons & Giannini, 2020. Amer. Mus. Novitates, 3950: 13

Macroglossus, Syconycteris.

Asia, Oceania

THIS ENTRY IS UNDER REVIEW.

References:

Miller, 1907. ,

Nyctimene, Paranyctimene.

Asia, Oceania

THIS ENTRY IS UNDER REVIEW.

References:

Almeida, Simmons & Giannini, 2020. Amer. Mus. Novitates, 3950: 12

Notopteris.

Oceania

References:

Gray, 1821. ,

Acerodon, Desmalopex, Melonycteris, Mirimiri, Neopteryx, Nesonycteris, Pteralopex, Pteropus, Styloctenium.

Africa, Asia, Oceania

THIS ENTRY IS UNDER REVIEW.

References:

, . ,

References:

, . ,

References:

, . ,

References:

Andersen, 1912. ,

Eonycteris, Epomophorus, Epomops, Hypsignathus, Megaloglossus, Myonycteris, Nanonycteris, Pilonycteris, Plerotes, Rousettus, Stenonycteris.

Africa, Asia, Europe, Oceania

THIS ENTRY IS BEING REVISED. Does not include Eidolon; see Juste et al. (1999), Romagnoli and Springer (2000), Colgan and da Costa (2002), Giannini and Simmons (2005), Almeida et al (2011, 2016).

References:

Andersen, 1912. ,

Pilonycteris, Rousettus.

Africa, Asia, Europe, Oceania

References:

Almeida, Giannini & Simmons, 2015. ,

Eonycteris.

Asia

References:

Gray, 1866. Proc. Zool. Soc. Lond., 1866: 65

Epomophorus, Epomops, Hypsignathus, Nanonycteris.

Africa

THIS ENTRY IS UNDER REVIEW.

References:

Lawrence & Novick, 1963. ,

Megaloglossus, Myonycteris.

Africa

References:

Nesi, Kadjo, Porrut, Pongombo Shongo, Cruaud & Hassanin, 2012. ,

Stenonycteris.

Africa

References:

Bergmans, 1997. ,

Casinycteris, Scotonycteris.

Africa

References:

Bergmans, 1997. ,

Plerotes.

Africa

References:

Gray, 1825. Zool. Journ., 2(6): 242

226

Africa, Asia, Europe, Oceania

Does not include Nycteridae. Includes Craseonycteridae and Rhinopomatidae. See Hulva and Horacek (2002), Van Den Bussche and Hoofer (2004), and Teeling et al. (2005). Panyutina et al. (2011) examined morphological patterns of the flight apparatus across some of the superfamily, but note that this study includes Nycteridae within Rhinolophoidea.

References:

Gray, J.E. 1825. An attempt at a division of the family Vespertilionidae into groups. The Zoological Journal 2(6): 242-243. Read article.

Panyutinaa, A.A., A.Y. Puzachenkob, and I.B. Soldatova. 2011. Morphological diversity of wing structure in rhinolophoid bats (Chiroptera, Rhinolophoidea). Biology Bulletin 38(7): 679-694. Read abstract.

Hill, 1974. Bull. Brit. Mus. (Nat. Hist.) Zool., 27: 303

Craseonycteris.

Asia

Monotypic.

References:

Hill, J.E. 1974. A new family, genus, and species of bat (Mammalia: Chiroptera) from Thailand. Bull. Brit. Mus. (Nat. Hist.) 27(1974): 301-336. Read article.

H. Allen, 1864. Monogr. Bats N. Am., pp. xxiii, 1

Cardioderma, Eudiscoderma, Lavia, Lyroderma, Macroderma, Megaderma.

Africa, Asia, Oceania

For discussion of the correct formation of the family name, see Handley (1980). Hand (1985, 1996), Griffiths et al. (1992), and Kaňuch et al. (2015) have provided alternative phylogenies for the group. No subfamilies are presently recognized.

References:

Allen, H. 1864. Monograph of the bats of North America. Smithsonian Miscellaneous Collections 7(1): 1-85. Read volume.

Griffiths, T.A., A. Truckenbrod, and P.J. Sponholtz 1992. Systematics of megadermatid bats (Chiroptera, Megadermatidae), based on hyoid morphology. American Museum Novitates 3041: 1-21. Read article.

Hand, S.J. 1985. New Miocene megadermatids (Chiroptera: Megadermatidae) from Australia with comments on megadermatid systematics. Australian Mammalogy 8: 5-43. Read article.

Hand, S.J. 1996. New Miocene and Pliocene megadermatids (Mammalia, Microchiroptera) from Australia, with comments on broader aspects of megadermatid evolution. Geobios 29: 365-377. Read abstract.

Kaňuch, P., T. Aghová, Y. Meheretu, R. Šumbera, and J. Bryja. 1997. New discoveries on the ecology and echolocation of the heart-nosed bat Cardioderma cor with a contribution to the phylogeny of Megadermatidae. African Zoology 50(1): 53-57. Read abstract.

Gray, 1866. Proc. Zool. Soc. Lond., 1866: 81

Triaenopini Benda & Vallo, 2009.

Cloeotis, Paratriaenops, Rhinonicteris, Triaenops.

Africa, Asia, Oceania

Formerly included within Hipposideridae at the tribal or subtribal level (e.g. Gray, 1866; Koopman, 1994; Hand and Kirsch, 2003; Hand and Archer, 2005), raised to familial rank by Foley et al. (2014). McKenna and Bell (1997) used the name Rhinonycterinae Gray, 1866 for the group that includes (as recognized here) Hipposideridae + Rhinonycteridae, but this usage was not accepted by other authors, even though Rhinonycteridae (= Rhinonycterina Gray, 1866) had priority over Hipposideridae Lydekker, 1891 as a family-group name.

References:

Benda, P., and P. Vallo 2009. Taxonomic revision of the genus Triaenops (Chiroptera: Hipposideridae) with description of a new species from southern Arabia and definitions of a new genus and tribe. Folio Zoologica 58(1): 1-45. Read abstract.

Foley, N.M, V.D. Thong, P. Soisook, S.M. Goodman, K.N. Armstrong, D.S. Jacobs, S.J. Puechmaille, and E.C. Teeling 2014. How and why overcome the impediments to resolution: Lessons from rhinolophid and hipposiderid bats. Molecular Biology and Evolution 32(2): 313-333. Read article.

Gray, J.E. 1866. A revision of the genera of Rhinolophidae, or horseshoe bats. Proceedings of the Zoological Society of London 1866: 81-83. Read article.

Hand, S.J., and J.A.W. Kirsch 2003. Archerops, a new annectent hipposiderid genus (Mammalia: Microchiroptera) from the Australian Miocene. Journal of Paleontology 77(6): 1139-1151. Read abstract.

Hand, S.J., and M. Archer 2005. A new hipposiderid genus (Microchiroptera) from an Early Miocene bat community in Australia. Paleontology 48(2): 371-383. Read article.

Koopman, K.F. 1994. Chiroptera: Systematics. Handbuch der Zoologie, vol. VIII, Mammalia, Part 60. de Gruyter, Berlin: 224 pp. Not available online.

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Lydekker, 1891. In Flower and Lydekker, Mamm. Living and Extinct, p. 657

Coelopsinae Tate, 1941.

Anthops, Asellia, Aselliscus, Coelops, Doryrhina, Hipposideros, Macronycteris.

Africa, Asia, Oceania

Does not include Cloeotis, Paratriaenops, Rhinonicteris, and Triaenops. See Murray et al. (2012), Foley et al. (2015), and Amador et al. (2016). Hipposideridae - including rhinonycterids - was treated as a subfamily of Rhinolophidae by Koopman (1993, 1994), McKenna and Bell (1997), Simmons (1998), Simmons and Geisler (1998), and Teeling et al. (2002), but is returned to family rank here following Corbet and Hill (1992), Bates and Harrison (1997), Bogdanowicz and Owen (1998), Hand and Kirsch (1998), and numerous other authors. Equivalent to subtribe Hipposiderina of McKenna and Bell (1997). McKenna and Bell (1997) used the name Rhinonycterinae Gray, 1866 for Hipposideridae sensu Simmons (2005), although the name Hipposideridae had been in widespread use since Miller (1907). There is currently disagreement concerning relationships within this family (see Bogdanowicz and Owen, 1998; Hand and Kirsch, 1998; Murray et al. 2012; Foley et al., 2015, 2017; and Amador et al., 2016), therefore, we do not recognize subfamilies or tribes within Hipposideridae at this time.

References:

Bates, P.J.J., and D.L. Harrison. 1997. Bats of the Indian Subcontinent. Harrison Zoological Museum, Sevenoaks: 258. Not available online.

Bogdanowicz, W., and R.D. Owen. 1998. In the Minotaur's labyrinth: the phylogeny of the bat family Hipposideridae. Pages 27-42 In T.H. Kunz and P.A. Racey (eds.) Bat Biology and Conservation. Washington, D.C.: Smithsonian Institution Press. Not available online.

Corbet, G.B., and J.E. Hill 1992. Mammals of the Indomalayan Region. A Systematic Review. Oxford University Press, Oxford: 1-488. Not available online.

Flower, W.H., and R. Lydekker 1891. An Introduction to the Study of Mammals, Living and Extinct. Adam and Charles Black, London: 763 pp. Read volume.

Foley, N.M, S.M. Goodman, C.V. Whelan, S.J. Puechmaille, and E.C. Teeling 2017. Towards navigating the Minotaur's labyrinth: cryptic diversity and taxonomic revision within the speciose genus Hipposideros (Hipposideridae). Acta Chiropterologica 19(1): 1-18. Read abstract.

Hand, S.J., and J.A.W. Kirsch. 1998. A southern origin for the Hipposideridae (Microchiroptera). Evidence from the Australian fossil record. Pages 72-90 In T.H. Kunz and P.A. Racey (eds.) Bat Biology and Conservation . Washington, D.C.: Smithsonian Institution Press. Not available online.

Koopman, K.F. 1994. Chiroptera: Systematics. Handbuch der Zoologie, vol. VIII, Mammalia, Part 60. de Gruyter, Berlin: 224 pp. Not available online.

McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. Columbia University Press, New York: 1-640. Not available online.

Miller, G.S., Jr. 1907. The families and genera of bats. Bulletin of the U.S. National Museum 57: 1-282. Read volume.

Murray S.W., P. Campbell, T. Kingston, A. Zubaid, C.M. Francis, and T.H. Kunz. 2012. Molecular phylogeny of hipposiderid bats from Southeast Asia and evidence of cryptic diversity. Molecular Phylogenetics and Evolution 62(2): 597-611. Read abstract.

Gray, 1825. Zool. Journ., 2(6): 242

Histiorhina Van der Hoeven, 1855.

112

Rhinolophus.

Africa, Asia, Europe, Oceania

Monogeneric. Does not include Hipposideridae; see discussion under that taxon. See the generic account for additional discussion of relationships within this group.

References:

Gray, J.E. 1825. An attempt at a division of the family Vespertilionidae into groups. The Zoological Journal 2(6): 242-243. Read article.

Bonaparte, 1838. Syn. Vert. Syst. in Nuovi Ann. Sci. Nat., Bologna, 2: 112

Rhinopoma.

Africa, Asia

Monogeneric. Authorship of the family-group name should be attributed to Bonaparte (1838), not Dobson (1872; see e.g., Corbet and Hill, 1992; Van Cakenberghe and De Vree, 1994), who used the correct stem (see Handley, 1980) and suffix, but was not the first to apply the name at the family-group level.

References:

Bonaparte, C.L. 1838. Synopsis Vertebratorum Systematis. Nuovi Annali delle Scienze Naturali, Bologna 1(2): 1-390. Read article.

Corbet, G.B., and J.E. Hill 1992. Mammals of the Indomalayan Region. A Systematic Review. Oxford University Press, Oxford: 1-488. Not available online.

Dobson, G.E. 1872. Chiroptera in Notice of the mammals and birds inhabiting Kachh by Dr. F. Stoliczka. Journal of the Asiatic Society of Bengal 41(3): 220-223. Read article.

Gray, J.E. 1866. A revision of the genera of Rhinolophidae, or horseshoe bats. Proceedings of the Zoological Society of London 1866: 81-83. Read article.

Miller, G.S., Jr. 1907. The families and genera of bats. Bulletin of the U.S. National Museum 57: 1-282. Read volume.

Van Cakenberghe, V., and F. De Vree. 1994. A revision of the Rhinopomatidae Dobson 1872, with the description of a new subspecies. Senkenbergiana Biologica 73(1-2): 1-24. Read article.

Bats of the World

A Taxonomic and Geographic Database

Compiled and Edited by:

Nancy B. Simmons and Andrea L. Cirranello

American Museum of Natural History