Large-footed bat Myotis adversus, from here.

Belongs within: Vespertilionidae.

Myotis is a genus of more than eighty species of bat found in most parts of the world within the tree line. Species of Myotis have been divided between the subgenera Myotis, Leuconoe and Selysius, which differ in both morphology and feeding behaviour (Findley 1972). Species of Myotis are long-winged and -legged forms whose slow but maneuverable flight suits them for feeding by gleaning from the ground or vegetation. Leuconoe species have shorter wings and legs and a more rapid, direct flight, and feed near the surface of water (with several species at least partially piscivorous). Selysius species are specialised for aerial capture of prey, and may use their broad uropatagia in doing so. A separate subgenus Hesperomyotis has also been recognised for the velvety myotis Myotis simus from South America, distinguished from other species in the genus by its particularly short velvety fur and largely hairless wing membranes. However, this subgenus is doubtfully distinct from Leuconoe.

Characters (from Miller 1907): Dental formula I2/3, C1/1, P3/3, M3/3. Upper incisors well developed, subequal, closely crowded, crowns higher than long, subterete ; inner incisor with distinct posterior secondary cusp, outer incisor with well-developed concave surface directed toward canine, from which it is separated by space not quite equal to diameter of both incisors together. Lower incisors with crowns about equal in length, forming continuous, strongly convex row between canines, first and second with rather narrow, trilobed cutting edge, third much wider, subterete, with three or four tubercles. Canines well developed, simple, with distinct though rather small cingulum and no secondary cusps. Cheek teeth both above and below strictly normal, m1 and m2 rather narrow on inner edge and with hypocone absent or very imperfectly developed, m3 with three commissures and four or five cusps, according to varying condition of metacone, m3, with second triangle smaller than first, but with all elements of tooth complete. Skull slender and lightly built, without special peculiarities of form, rostrum nearly as long as braincase, depth of braincase, including audital bulla, about equal to mastoid breadth, sagittal crest low, but usually distinct, palate deeply emarginate in front, abruptly contracted behind, the sides of its posterior extension parallel. Audital bullae well developed and covering more than half surface of cochleae, but simple in form and not very large, their diameter about equal to the distance between them. Ear well developed, slender, occasionally rather large; tragus slender and nearly or quite straight. Foot very variable in size, but never peculiar in form. Tail about as long as outstretched leg. Interfemoral membrane large, its surface furred at extreme base above.

<==Myotis Kaup 1829 P04 (see below for synonymy)
    |--M. (Myotis) F72
    |    |--+--M. (M.) emarginatus (Geoffroy 1806) F72, P04 [=Isotus emarginatus M07]
    |    |  |--M. (M.) goudoti (Smith 1834) F72, K92
    |    |  `--M. (M.) tricolor F72
    |    |--+--M. (M.) formosus [incl. M. flavus, M. hermani, M. rufoniger, M. rufopictus] F72
    |    |  |    |--M. f. formosus I92
    |    |  |    `--M. f. tsuensis Kuroda 1922 I92
    |    |  `--M. (M.) welwitschii F72
    |    |--+--M. (M.) nattereri (Kuhl 1818) F72, P04 [=Isotus nattereri M07]
    |    |  |--M. (M.) pequinius F72
    |    |  `--M. (M.) thysanodes Miller 1897 F72, MB86
    |    |       |--M. t. thysanodes G69
    |    |       `--M. t. aztecus G69
    |    |--+--M. (M.) auriculus F72
    |    |  |    |--M. a. auriculus MB86
    |    |  |    `--M. a. apache Hoffmeister & Krutsch 1955 MB86
    |    |  |--M. (M.) evotis F72
    |    |  `--M. (M.) keeni F72
    |    |       |--M. k. keeni B75
    |    |       `--M. k. septentrionalis (Trouessart 1897) [=Vespertilio gryphus var. septentrionalis] B75
    |    `--+--*M. (M.) myotis (Borkhausen 1797) C57, F72, P04 [=Vespertilio myotis C57]
    |       |--M. (M.) bechsteini (Kuhl 1818) F72, P04
    |       |    |--M. b. bechsteini P04
    |       |    `--M. b. robustus Topál 1963 P04
    |       |--M. (M.) blythi (Tomes 1857) F72, P04
    |       |    |--M. b. blythii P04
    |       |    `--M. b. longicaninus Popov 2004 P04
    |       |--M. (M.) chinensis F72
    |       `--M. (M.) sicarius F72
    |--M. (Hesperomyotis Cabrera 1957) C57
    |    `--M. (*H.) simus Thomas 1901 C57
    |--M. (Leuconoe Boie 1830) C57 (see below for synonymy)
    |    |--M. (L.) dasycneme F72 [=Brachyotus dasycneme M07, Comastes dasycneme M07]
    |    |--+--M. (L.) montivagus F72
    |    |  `--M. (L.) riparius F72
    |    |--+--‘M. (Rickettia) pilosus Peters 1869 [=Vespertilio (Leuconoe) pilosus] C57
    |    |  `--M. (L.) ricketti (Thomas 1894) F72, C57 (see below for synonymy)
    |    |--+--M. (L.) federatus F72
    |    |  |--M. (L.) fortidens F72
    |    |  `--M. (L.) peytoni F72
    |    |--+--M. (L.) macrotarsus F72 [=Exochurus macrotarsus M07]
    |    |  |--M. (L.) stalkeri F72
    |    |  `--M. (L.) vivesi F72 [=*Pizonyx vivesi M07]
    |    |--+--M. (*L.) daubentoni (Kuhl 1819) C57, F72, K92 [=Vespertilio daubentonii C57, Brachyotus daubentonii M07]
    |    |  |--M. (L.) capaccini (Bonaparte 1837) F72, P04 [=*Capaccinius capaccinii M07, Comastes capaccinii M07]
    |    |  |--M. (L.) fimbriatus F72
    |    |  |--M. (L.) longipes F72
    |    |  |--M. (L.) macrodactylus F72 [=Exochurus macrodactylus M07]
    |    |  `--M. (L.) taiwanensis F72
    |    |--+--M. (L.) albescens (Geoffroy 1806) F72, C57 (see below for synonymy)
    |    |  |--M. (L.) austroriparius F72
    |    |  |--M. (L.) carissima [=M. lucifugus carissima] F72
    |    |  |--M. (L.) petax F72
    |    |  |--M. (L.) volans F72
    |    |  `--M. (L.) yumanensis F72
    |    |       |--M. y. yumanensis MB86
    |    |       `--M. y. lutosus Miller & Allen 1928 MB86
    |    |--+--M. (L.) chiloensis (Waterhouse 1838) F72, C57 (see below for synonymy)
    |    |  |    |--M. c. chiloensis [incl. Vespertilio gayi Lataste 1892] C57
    |    |  |    |--M. c. alter Miller & Allen 1928 C57
    |    |  |    |--M. c. arescens Osgood 1943 C57
    |    |  |    |--M. c. atacamensis (Lataste 1892) (see below for synonymy) C57
    |    |  |    `--M. c. oxyotus (Peters 1866) [=Vespertilio oxyotus; incl. M. thomasi Cabrera 1901] C57
    |    |  |--M. (L.) grisescens Howell 1909 F72, B75
    |    |  |--M. (L.) incautus F72 [=M. velifer incautus MB86]
    |    |  |--M. (L.) occultus [=M. lucifugus occultus] F72
    |    |  `--M. (L.) velifer (Allen 1890) F72, K92
    |    `--+--M. (L.) abboti F72
    |       |--M. (L.) adversus F72 [incl. Trilatitus macellus M07]
    |       |    |--M. a. adversus M72
    |       |    `--M. a. continentis F72
    |       |--M. (L.) bocagei [incl. M. dogalensis] F72
    |       |--M. (L.) carimatae F72
    |       |--M. (L.) deignani F72
    |       |--M. (L.) dryas F72
    |       |--M. (L.) horsefieldi F72 [=Exochurus horsfieldii M07]
    |       |--M. (L.) jeannei F72
    |       |--M. (L.) lepidus F72
    |       |--M. (L.) macropus F72 [=M. adversus macropus M72]
    |       |--M. (L.) moluccarum F72 [=M. adversus moluccarum M72]
    |       `--M. (L.) peshwa F72
    `--M. (Selysius Bonaparte 1841) C57
         |  i. s.: M. (S.) ruber (Geoffroy 1806) (see below for synonymy) C57
         |--M. (S.) altarium F72
         |--M. (S.) scotti F72
         |--M. (S.) siligorensis F72
         |--+--M. (S.) nigricans (Schinz 1821) F72, C57 [=Vespertilio nigricans C57, Aeorestes nigricans C57]
         |  |    |--M. n. nigricans (see below for synonymy) C57
         |  |    |--M. n. dalquesti Hall & Alvarez 1961 G69
         |  |    `--M. n. nicholsoni Sanborn 1941 C57
         |  `--M. (S.) sodalis Miller & Allen 1928 F72, B75
         |--+--M. (*S.) mystacinus C57, F72 [=Vespertilio mystacinus C57, Brachyotus mystacinus M07]
         |  |--M. (S.) davidii F72
         |  |--M. (S.) frater F72
         |  |--M. (S.) insularum F72
         |  `--M. (S.) meinertzhageni F72
         |--+--M. (S.) californicus (Audubon & Bachman 1842) F72, MB86
         |  |    |--M. c. californicus MB86
         |  |    `--M. c. mexicanus (Saussure 1860) MB86
         |  `--M. (S.) leibi (Audubon & Bachman 1842) F72, B75
         |       |--M. l. leibi MB86
         |       |--M. l. ciliolabrium (Merriam 1866) B75
         |       `--M. l. melanorhinus (Merriam 1890) MB86
         `--+--M. (S.) ater F72
            |--M. (S.) blanfordi F72
            |--M. (S.) browni F72
            |--M. (S.) caliginosus F72
            |--M. (S.) latirostris Kishida 1932 F72, TYM08
            |--M. (S.) moupinensis F72
            |--M. (S.) muricola F72 [=Vespertilio muricola M07; incl. V. (*Pternopterus) lobipes M07]
            |--M. (S.) niasensis F72
            `--M. (S.) primula F72

Myotis incertae sedis:
  M. abei IT07
  M. aelleni IT07
  M. annectans IT07
  M. argentatus G69
  M. australis IT07
  M. bartelsii BP87
  M. bombinus IT07
  M. brandti IT07
  M. cobanensis IT07
  M. danutae Kowalski 1956 P04
  M. delicatus Heller 1936 [incl. M. dasycneme subtilis Kowalki 1956] P04
  M. dominicensis IT07
  M. elegans F72
  M. exilis Heller 1936 P04
  M. farater I92
    |--M. f. farater I92
    `--M. f. kaguyae Imaizumi 1956 I92
  M. findleyi IT07
  M. ghardalamensis Storch 1974 P04
  M. gundersheimensis Heller 1936 P04
  M. hasseltii IT07 [=Trilatitus hasseltii M07]
  M. hosonoi Imaizumi 1954 I92
  M. ikonnikovi Ognev 1912 I92
  M. janossyi Topál 1983 P04
  M. kretzoii Topál 1981 P04
  M. laniger F72
  *Nystactes’ leisleri M07
  M. lesueuri IT07
  M. levis IT07
  ‘Comastes’ limnophilus M07
  M. lucifugus (Le Conte in McMurtrie 1831) [=Vespertilio lucifugus] B75
  M. martiniquensis IT07
  ‘Comastes’ megapodius M07
  M. milleri IT07
  M. morrisi IT07
  M. nathalinae BP87
  M. nesopolus IT07
  M. oreias IT07
  M. oxygnathus KK54
  M. ozensis Imaizumi 1954 I92
  M. peninsularis IT07
  M. planiceps Baker 1955 MB86
  M. pruinosus Yoshiyuki 1971 I92
  M. ridleyi IT07
  M. rosseti IT07
  M. schaubi IT07
  M. seabrai F72
  M. sowerbyi F72
  M. subulatus F68
  ‘Aeorestes’ villosissimus M07
  M. yesoensis IT07

Myotis Kaup 1829 P04 [incl. Aeorestes Fitzinger 1870 M07, Brachyotus Kolenati 1856 non Gould 1837 M07, Exochurus Fitzinger 1870 M07, Isotus Kolenati 1856 M07, Nystactes Kaup 1829 M07, Pternopterus Peters 1867 M07, Trilatitus Gray 1842 M07, Vespertilio Keyserling & Blasius 1839 non Linnaeus 1758 M07]

Myotis (Leuconoe Boie 1830) C57 [incl. Capaccinius Bonaparte 1841 C57, Comastes Fitzinger 1870 C57, Pizonyx Miller 1906 M07, Tralatitius Gray 1866 C57, Tralatitus Gervais 1849 C57]

Myotis (Leuconoe) albescens (Geoffroy 1806) [=Vespertilio albescens, Aeorestes albescens; incl. V. aenobarbus Temminck 1940, V. arsinoe Temminck 1840, Myotis arsinoe, V. isidori D’Orbigny & Gervais 1847, V. leucogaster Wied 1826, V. mundus Allen 1866] C57

Myotis (Leuconoe) chiloensis (Waterhouse 1838) F72, C57 [=Vespertilio chiloensis C57, M. (Selysius) chiloensis C57]

Myotis (Leuconoe) chiloensis atacamensis (Lataste 1892) [=Vespertilio atacamensis; incl. M. chiloensis dinellii Thomas 1902] C57

Myotis (Leuconoe) ricketti (Thomas 1894) F72, C57 [=Vespertilio (Leuconoe) ricketti C57, M. (*Rickettia) ricketti C57]

Myotis (Selysius) nigricans nigricans (Schinz 1821) [incl. Myotis bondae Allen 1914, M. caucensis Allen 1914, M. chiriquensis Allen 1904, Vespertilio concinnus Allen 1866, M. nigricans concinnus, M. esmeraldae Allen 1914, V. exiguus Allen 1866, V. hypothrix D’Orbigny & Gervais 1847, Myotis ruber keaysi Allen 1914, M. maripensis Allen 1914, Vespertilio osculatii Cornalia 1854, M. nigricans osculatii, V. parvulus Temminck 1837, Nyctophylax parvulus, Myotis punensis Allen 1914, V. quixensis Cornalis 1854, V. spixii Fischer 1829] C57

Myotis (Selysius) ruber (Geoffroy 1806) [=Vespertilio ruber, Nycticeus ruber; incl. Vespertilio cinnamomeus Wagner 1855, Vespertilio (Myotis) kinnamon Gervais 1856, Vespertilio nubilus Wagner 1844, Nyctophylax nubilus, Vespertilio polythrix Geoffroy 1924, Myotis polythrix, Scotophilus (Pachyotus) polythrix, Vesperus polythrix] C57

*Type species of generic name indicated


[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1–184.

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[C57] Cabrera, A. 1957. Catalogo de los mamiferos de America del Sur. I (Metatheria—Unguiculata—Carnivora). Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” e Instituto Nacional de Investigacion de Las Ciencias Naturales, Ciencias Zoológicas 4 (1): 1–307.

[F72] Findley, J. S. 1972. Phenetic relationships among bats of the genus Myotis. Systematic Zoology 21 (1): 31–52.

[F68] Fisher, D. R. 1968. A study of faunal resemblance using numerical taxonomy and factor analysis. Systematic Zoology 18 (4): 48–63.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[KK54] Klemmer, K., & H. E. Krampitz. 1954. Zur Kenntnis der Säugetierfauna Siziliens. Senckenbergiana Biologica 35 (3-4): 121–135.

[K92] Klompen, J. S. H. 1992. Phylogenetic relationships in the mite family Sarcoptidae (Acari: Astigmata). Miscellaneous Publications, Museum of Zoology, University of Michigan 180: i–vi, 1–154.

[MB86] Matson, J. O., & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1–88.

[M72] McKean, J. L. 1972. Notes on some collections of bats (order Chiroptera) from Papua-New Guinea and Bougainville Island. CSIRO Division of Wildlife Research Technical Paper 26: 1–35.

[M07] Miller, G. S., Jr. 1907. The families and genera of bats. Smithsonian Institution, United States National Museum, Bulletin 57: i–xvii, 1–282, pl. I–XIV.

[P04] Popov, V. V. 2004. Pliocene small mammals (Mammalia, Lipotyphla, Chiroptera, Lagomorpha, Rodentia) from Muselievo (north Bulgaria). Geodiversitas 26 (3): 403–491.

[TYM08] Tennent, W. J., M. Yasuda & K. Morimoto. 2008. Lansania Journal of arachnology and zoology—a rare and obscure Japanese natural history journal. Archives of Natural History 35 (2): 252–280.

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