Myotis

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 1829P04 (see below for synonymy)
    |--M. alcathoeFS15
    `--+--+--M. hajastanicusFS15
       |  `--+--+--M. brandtiiFS15
       |     |  `--M. insularumFS15 [=M. (Selysius) insularumF72]
       |     `--+--+--+--M. sodalis Miller & Allen 1928FS15, B75 [=M. (Selysius) sodalisF72]
       |        |  |  `--M. volansFS15 [=M. (Leuconoe) volansF72]
       |        |  `--+--+--M. vivesiFS15 [=M. (Leuconoe) vivesiF72, *Pizonyx vivesiM07]
       |        |     |  `--+--M. federatus [=M. (Leuconoe) federatus]F72
       |        |     |     |--M. fortidensFS15 [=M. (Leuconoe) fortidensF72]
       |        |     |     `--M. peytoni [=M. (Leuconoe) peytoni]F72
       |        |     `--+--+--M. auriculusFS15
       |        |        |  |    |--M. a. auriculusMB86
       |        |        |  |    `--M. a. apache Hoffmeister & Krutsch 1955MB86
       |        |        |  `--+--M. csorbaiFS15
       |        |        |     `--M. septentrionalis (Trouessart 1897)FS15, B75 (see below for synonymy)
       |        |        `--+---+--+--M. californicus (Audubon & Bachman 1842)FS15, MB86 [=M. (Selysius) californicusF72]
       |        |           |   |  |    |--M. c. californicusMB86
       |        |           |   |  |    `--M. c. mexicanus (Saussure 1860)MB86
       |        |           |   |  `--M. leibii (Audubon & Bachman 1842)F72, FS15, B75 [=M. (Selysius) leibiF72]
       |        |           |   `--+--M. planiceps Baker 1955FS15, MB86
       |        |           |      `--+--M. ciliolabrum (Merriam 1866)FS15, B75 [=M. leibii ciliolabriumB75]
       |        |           |         `--M. melanorhinus (Merriam 1890)FS15, MB86 [=M. leibii melanorhinusMB86]
       |        |           `--+--+--M. aelleniFS15
       |        |              |  `--+--M. lucifugus (Le Conte in McMurtrie 1831)FS15, B75 [=Vespertilio lucifugusB75]
       |        |              |     `--M. occultusFS15 [=M. (Leuconoe) lucifugus occultusF72]
       |        |              `--+--M. keeniiFS15
       |        |                 `--+--M. evotisFS15
       |        |                    `--M. thysanodes Miller 1897FS15, MB86
       |        |                         |--M. t. thysanodesG69
       |        |                         `--M. t. aztecusG69
       |        `--+--+--M. keaysi Allen 1914FS15, C57 [=Myotis ruber keaysiC57]
       |           |  `--+--M. ruber (Geoffroy 1806)FS15, C57 (see below for synonymy)
       |           |     `--+--M. simus Thomas 1901FS15, C57 [=M. (*Hesperomyotis) simusC57]
       |           |        `--+--M. elegansFS15
       |           |           `--M. ripariusFS15 [=M. (Leuconoe) ripariusF72]
       |           `--+--+--M. austroripariusFS15 [=M. (Leuconoe) austroripariusF72]
       |              |  `--M. grisescens Howell 1909FS15, B75 [=M. (Leuconoe) grisescensF72]
       |              `--+--+--M. velifer (Allen 1890)FS15, K92 [=M. (Leuconoe) veliferF72]
       |                 |  `--+--M. cobanensisFS15
       |                 |     `--M. yumanensisFS15 [=M. (Leuconoe) yumanensisF72]
       |                 |          |--M. y. yumanensisMB86
       |                 |          `--M. y. lutosus Miller & Allen 1928MB86
       |                 `--+--M. chiloensis (Waterhouse 1838)FS15, C57 (see below for synonymy)
       |                    |    |--M. c. chiloensis [incl. Vespertilio gayi Lataste 1892]C57
       |                    |    |--M. c. alter Miller & Allen 1928C57
       |                    |    `--M. c. arescens Osgood 1943C57
       |                    `--+--+--M. oxyotus (Peters 1866)FS15, C57 (see below for synonymy)
       |                       |  `--+--M. levisFS15
       |                       |     `--+--M. findleyiFS15
       |                       |        `--+--M. nesopolusFS15
       |                       |           `--M. nigricans (Schinz 1821)FS15, C57 (see below for synonymy)
       |                       |                |--M. n. nigricans (see below for synonymy)C57
       |                       |                |--M. n. dalquesti Hall & Alvarez 1961G69
       |                       |                `--M. n. nicholsoni Sanborn 1941C57
       |                       `--+--+--M. martiniquensisFS15
       |                          |  `--+--M. atacamensis (Lataste 1892)FS15, C57 (see below for synonymy)
       |                          |     `--M. dominicensisFS15
       |                          `--+--M. peninsularisFS15
       |                             `--+--M. albescens (Geoffroy 1806)FS15, C57 (see below for synonymy)
       |                                `--+--M. pilosus Peters 1869FS15, C57 (see below for synonymy)
       |                                   `--M. ricketti (Thomas 1894)F72, C57 (see below for synonymy)
       `--+--M. moluccarumFS15 [=M. adversus moluccarumM72, M. (Leuconoe) moluccarumF72]
          `--+--+--+--M. tricolorFS15
             |  |  `--+--M. emarginatus (Geoffroy 1806)FS15, P04 [=Isotus emarginatusM07]
             |  |     `--M. morrisiFS15
             |  `--+--+--M. scottiFS15 [=M. (Selysius) scottiF72]
             |     |  `--+--M. bocagiiFS15 [=M. (Leuconoe) bocageiF72; incl. M. dogalensisF72]
             |     |     `--M. welwitschiiFS15
             |     `--+--+--M. formosusFS15 [incl. M. flavusF72, M. rufonigerF72]
             |        |  |    |--M. f. formosusI92
             |        |  |    `--M. f. tsuensis Kuroda 1922I92
             |        |  `--M. hermaniFS15
             |        `--+--M. anjouanensisFS15
             |           `--+--M. goudoti (Smith 1834)FS15, K92
             |              `--+--M. dinellii Thomas 1902FS15, C57 [=M. chiloensis dinelliiC57]
             |                 `--M. rufopictusFS15
             `--+--M. dasycnemeFS15 [=Brachyotus dasycnemeM07, Comastes dasycnemeM07, Myotis (Leuconoe) dasycnemeF72]
                `--+--+--+--+--M. aterFS15 [=M. (Selysius) aterF72]
                   |  |  |  `--M. muricolaFS15 (see below for synonymy)
                   |  |  `--+--+--M. annectansFS15
                   |  |     |  `--M. gomantongensisFS15
                   |  |     `--+--M. pruinosus Yoshiyuki 1971FS15, I92
                   |  |        `--+--M. montivagusFS15 [=M. (Leuconoe) montivagusF72]
                   |  |           `--M. yanbarensisFS15
                   |  `--+--M. nipalensisFS15
                   |     `--+--+--M. aurascensFS15
                   |        |  `--M. mystacinusFS15 (see below for synonymy)
                   |        `--+--+--M. altariumFS15 [=M. (Selysius) altariumF72]
                   |           |  `--M. oreiasFS15
                   |           `--+--M. dieteriFS15
                   |              `--M. ikonnikovi Ognev 1912FS15, I92
                   `--+--+--+--M. bechsteinii (Kuhl 1818)FS15, P04
                      |  |  |    |--M. b. bechsteiniiP04
                      |  |  |    `--M. b. robustus Topál 1963P04
                      |  |  `--M. sicariusFS15
                      |  `--+--M. daubentonii (Kuhl 1819)FS15, K92 (see below for synonymy)
                      |     `--+--M. bucharensisFS15
                      |        `--M. fraterFS15 [=M. (Selysius) fraterF72]
                      `--+--+--+--M. punicusFS15
                         |  |  `--+--*M. myotis (Borkhausen 1797)C57, FS15, P04 [=Vespertilio myotisC57]
                         |  |     `--M. blythi (Tomes 1857)FS15, P04
                         |  |          |--M. b. blythiiP04
                         |  |          `--M. b. longicaninus Popov 2004P04
                         |  `--+--M. chinensisFS15
                         |     `--+--M. schaubiFS15
                         |        `--+--M. pequiniusFS15
                         |           `--+--M. bombinusFS15
                         |              `--M. nattereri (Kuhl 1818)FS15, P04 [=Isotus nattereriM07]
                         `--+--+--+--M. adversusFS15 [=M. (Leuconoe) adversusF72; incl. Trilatitus macellusM07]
                            |  |  |    |--M. a. adversusM72
                            |  |  |    `--M. a. continentisF72
                            |  |  `--M. fimbriatusFS15 [=M. (Leuconoe) fimbriatusF72]
                            |  `--+--+--M. davidiiFS15 [=M. (Selysius) davidiiF72]
                            |     |  `--M. siligorensisFS15 [=M. (Selysius) siligorensisF72]
                            |     `--+--M. lanigerFS15
                            |        `--+--M. annamiticusFS15
                            |           `--M. longipesFS15 [=M. (Leuconoe) longipesF72]
                            `--+--M. capaccinii (Bonaparte 1837)FS15, P04 (see below for synonymy)
                               `--+--+--M. australisFS15
                                  |  `--M. rossetiFS15
                                  `--+--M. ridleyiFS15
                                     `--+--+--M. hasseltiiFS15 [=Trilatitus hasseltiiM07]
                                        |  `--M. macropus Gould 1855FS15, MB12 (see below for synonymy)
                                        `--+--M. horsfieldiiFS15 [=Exochurus horsfieldiiM07, M. (Leuconoe) horsefieldiF72]
                                           `--+--M. macrodactylusFS15 (see below for synonymy)
                                              `--+--M. macrotarsusFS15 [=Exochurus macrotarsusM07, M. (Leuconoe) macrotarsusF72]
                                                 `--M. stalkeriFS15 [=M. (Leuconoe) stalkeriF72]
Myotis incertae sedis:
  M. abeiIT07
  M. abboti [=M. (Leuconoe) abboti]F72
  M. argentatusG69
  M. bartelsiiBP87
  M. blanfordi [=M. (Selysius) blanfordi]F72
  M. browni [=M. (Selysius) browni]F72
  M. caliginosus [=M. (Selysius) caliginosus]F72
  M. carimatae [=M. (Leuconoe) carimatae]F72
  M. carissima [=M. lucifugus carissima, M. (Leuconoe) carissima]F72
  M. danutae Kowalski 1956P04
  M. deignani [=M. (Leuconoe) deignani]F72
  M. delicatus Heller 1936 [incl. M. dasycneme subtilis Kowalki 1956]P04
  M. dryas [=M. (Leuconoe) dryas]F72
  M. exilis Heller 1936P04
  M. faraterI92
    |--M. f. faraterI92
    `--M. f. kaguyae Imaizumi 1956I92
  M. ghardalamensis Storch 1974P04
  M. gundersheimensis Heller 1936P04
  M. hosonoi Imaizumi 1954I92
  M. incautusF72 [=M. velifer incautusMB86, M. (Leuconoe) incautusF72]
  M. janossyi Topál 1983P04
  M. jeannei [=M. (Leuconoe) jeannei]F72
  M. kretzoii Topál 1981P04
  M. latirostris Kishida 1932TYM08 [=M. (Selysius) latirostrisF72]
  ‘*Nystactes’ leisleriM07
  M. lepidus [=M. (Leuconoe) lepidus]F72
  M. lesueuriIT07
  ‘Comastes’ limnophilusM07
  ‘Comastes’ megapodiusM07
  M. meinertzhageni [=M. (Selysius) meinertzhageni]F72
  M. milleriIT07
  M. moupinensis [=M. (Selysius) moupinensis]F72
  M. nathalinaeBP87
  M. niasensis [=M. (Selysius) niasensis]F72
  M. oxygnathusKK54
  M. ozensis Imaizumi 1954I92
  M. peshwa [=M. (Leuconoe) peshwa]F72
  M. petax [=M. (Leuconoe) petax]F72
  M. primula [=M. (Selysius) primula]F72
  M. seabraiF72
  M. sowerbyiF72
  M. subulatusF68
  M. taiwanensis [=M. (Leuconoe) taiwanensis]F72
  ‘Aeorestes’ villosissimusM07
  M. yesoensisIT07

Myotis Kaup 1829^P04 [incl. Aeorestes Fitzinger 1870^M07, Brachyotus Kolenati 1856 non Gould 1837^M07, Capaccinius Bonaparte 1841^C57, Comastes Fitzinger 1870^C57, Exochurus Fitzinger 1870^M07, Hesperomyotis Cabrera 1957^C57, Isotus Kolenati 1856^M07, Leuconoe Boie 1830^C57, Nystactes Kaup 1829^M07, Pizonyx Miller 1906^M07, Pternopterus Peters 1867^M07, Selysius Bonaparte 1841^C57, Tralatitius Gray 1866^C57, Tralatitus Gervais 1849^C57, Trilatitus Gray 1842^M07, Vespertilio Keyserling & Blasius 1839 non Linnaeus 1758^M07]

Myotis albescens (Geoffroy 1806)^{FS15, C57} [=Vespertilio albescens^C57, Aeorestes albescens^C57, Myotis (Leuconoe) albescens^F72; incl. Vespertilio aenobarbus Temminck 1940^C57, V. arsinoe Temminck 1840^C57, Myotis arsinoe^C57, V. isidori D’Orbigny & Gervais 1847^C57, V. leucogaster Wied 1826^C57, V. mundus Allen 1866^C57]

Myotis atacamensis (Lataste 1892)^{FS15, C57} [=Vespertilio atacamensis^C57, M. (Selysius) chiloensis atacamensis^C57]

Myotis capaccinii (Bonaparte 1837)^{FS15, P04} [=*Capaccinius capaccinii^M07, Comastes capaccinii^M07, Myotis (Leuconoe) capaccini^F72]

Myotis chiloensis (Waterhouse 1838)^{FS15, C57} [=Vespertilio chiloensis^C57, M. (Leuconoe) chiloensis^F72, M. (Selysius) chiloensis^C57]

Myotis daubentonii (Kuhl 1819)^{FS15, K92} [=Vespertilio daubentonii^C57, Brachyotus daubentonii^M07, Myotis (*Leuconoe) daubentoni^C57]

Myotis macrodactylus^FS15 [=Exochurus macrodactylus^M07, M. (Leuconoe) macrodactylus^F72]

Myotis macropus Gould 1855^{FS15, MB12} [=M. adversus macropus^M72, M. (Leuconoe) macropus^F72]

Myotis muricola^FS15 [=Vespertilio muricola^M07, M. (Selysius) muricola^F72; incl. V. (*Pternopterus) lobipes^M07]

Myotis mystacinus^FS15 [=Vespertilio mystacinus^C57, Brachyotus mystacinus^M07, Myotis (*Selysius) mystacinus^C57]

Myotis nigricans (Schinz 1821)^{FS15, C57} [=Vespertilio nigricans^C57, Aeorestes nigricans^C57, Myotis (Selysius) nigricans^F72]

Myotis 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 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 oxyotus (Peters 1866)^{FS15, C57} [=Vespertilio oxyotus^C57, M. (Selysius) chiloensis oxyotus^C57; incl. M. thomasi Cabrera 1901^C57]

Myotis pilosus Peters 1869^{FS15, C57} [=Vespertilio (Leuconoe) pilosus^C57, M. (Rickettia) pilosus^C57]

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

Myotis ruber (Geoffroy 1806)^{FS15, C57} [=Vespertilio ruber^C57, Nycticeus ruber^C57, M. (Selysius) ruber^C57; incl. Vespertilio cinnamomeus Wagner 1855^C57, Vespertilio (Myotis) kinnamon Gervais 1856^C57, Vespertilio nubilus Wagner 1844^C57, Nyctophylax nubilus^C57, Vespertilio polythrix Geoffroy 1924^C57, Myotis polythrix^C57, Scotophilus (Pachyotus) polythrix^C57, Vesperus polythrix^C57]

Myotis septentrionalis (Trouessart 1897)^{FS15, B75} [=Vespertilio gryphus var. septentrionalis^B75, M. keeni septentrionalis^B75]

*Type species of generic name indicated

References

[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.

[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.

[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.

[MB12] McKenzie, N. L., & R. D. Bullen. 2012. An acoustic survey of zoophagic bats on islands in the Kimberley, Western Australia, including data on the echolocation ecology, organisation and habitat relationships of regional communities. Records of the Western Australian Museum Supplement 81: 67–108.

[M07] Miller, G. S., Jr. 1907. The families and genera of bats. Smithsonian Institution, United States National Museum, Bulletin 57: i–xvii, 1–282, pls 1–14.

[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.