bat, Blumenbach, 1779Megadermatidae, H. Allen, 1864

Cakenberghe, Victor Van, Tungaluna, Guy-Crispin Gembu, Akawa, Prescott Musaba, Seamark, Ernest & Verheyen, Erik, 2017, The bats of the Congo and of Rwanda and Burundi revisited (Mammalia: Chiroptera), European Journal of Taxonomy 382 (382), pp. 1-327 : 112-125

Key to the bat species of the CRB region

This key is primarily modified from Hayman et al. (1966), with additional information from Hayman & Hill (1971), Bergmans (1997), Cotterill (2001b), Happold & Happold (2013), Kerbis Peterhans et al. (2013) and Foley et al. (2014).

1. Cheekteeth without W-pattern of cusps; second finger with claw; tail absent or rudimentary; interfemoral membrane reduced; ears very short, well separated, no tragus; margin of ear forming a complete ring (family Pteropidae )..................................................................................................... 9

– Cheekteeth with W-pattern of cusps; second finger entirely included in patagium and without claw; tail generally well developed; interfemoral membrane developed; ears very short to very long, widely separed or joined; tragus present in most cases (except for Rhinolophidae , Rhinonycteridae and Hipposideridae )........................................................................................................................... 2

2. Tail more or less completely included in uropatagium (in certain forms the last vertebra can be free) ................................................................................................................................................... 3

– Tail not completely included in uropatagium .................................................................................... 8

3. No nasal appendages; tragus conspicuous; ears very short to very long, widely separated or joined, but not fused in the mediosagittal plane .......................................................................................... 64

– Nasal appendages present.................................................................................................................. 4

4. No tragus or tragus inconspicuous; ears very short to medium, not fused mediosagitally ............... 5

– Tragus present (sometimes small); ears fused or very close to each other mediosagitally ............... 6

5. Posterior leaflet of nose-leaf low, rounded or with three pointed projections (tridentate); two pedal phalanges; tragus inconspicuous...................................................................................................... 43

– Posterior leaflet of nose-leaf high, erect, pointed; three pedal phalanges; no tragus (family Rhinolophidae , genus Rhinolophus )................................................................................................ 53

6. Tail shorter than uropatagium; no postorbital processes; no premaxillae present; noseleaf very large, comparatively simple; tragus very conspicuous, bifid (family Megadermatidae )............................. 7

– Tail as long as uropatagium; last vertebra Y- or T-shaped; postorbital process and premaxillae present; noseleaf is formed by a longitudinal slit bordered by fleshy outgrowths; tragus conspicuous, not bifid (family Nycteridae ) ........................................................................................................... 35

7. Noseleaf less than 15 mm; pelage grey without yellow tinge; frontal shield of skull strongly concave, minimum breadth of shield less than 6.5 mm................................... Cardioderma cor (Peters, 1872)

– Noseleaf over 18 mm; pelage bluish-grey, generally with yellowish tinge; frontal shield of skull comparatively flat, minimum breadth of shield over 6.5 mm ............................................................. ........................................................................................ Lavia frons (E. Geoffroy St.-Hilaire, 1810)

8. Basal part of tail included in uropatagium; distal part of tail resting free on the dorsal side of the uropatagium; ears very short, widely separated, not fused, backwards-pointing; tragus conspicuous (family Emballonuridae ).................................................................................................................. 31

– Distal part of tail (ca half the width of the interfemoral membrane) not included in uropatagium and not resting on the dorsal side of the uropatagium; ears short, mostly united to each other by a cutaneous wall (family Molossidae ).............................................................................................. 108

9. Tongue simple, fixed to floor of mouth by posterior half, and without unfringed filiform papillae at tip; muzzle not elongated................................................................................................................. 10

– Tongue more extensible, fixed to floor of mouth by its posterior third, its terminal fourth or fifth covered above with unfringed filiform papillae (“brush-tipped”), muzzle elongated.................................................................. Megaloglossus woermanni Pagenstecher, 1885

10. Facial pattern of white nose and eye patches....................................................................................11

– No facial pattern .............................................................................................................................. 12

11. White tufts at base of ears inconspicuous or absent; bony palate extending well beyond teeth; rostrum, when viewed laterally, not upturned........... Scotonycteris bergmansi Hassanin et al., 2015

– White tufts at base of ears present, although sometimes inconspicuous; no postdental bony palate; rostrum, when viewed laterally, upturned................................. Casinycteris argynnis Thomas, 1910

12. No white tufts present at base of ears, or tufts inconspicuous......................................................... 13

– White tufts at base of ears present, although sometimes inconspicuous ......................................... 17

13. Facial axis of skull not noticeably deflected against basicranial axis; forearm 54–90 mm, length second digit 71–81 % of forearm; toes partially webbed; cheek teeth squarish or oblong; 6 + 2 or 7 + 2 palatal ridges (genus Myonycteris )......................................................................................... 14

– Facial axis of skull more or less strongly deflected against basicranial axis; forearm 75–130 mm, length of second digit 64–71 % of forearm; toes not webbed; cheek teeth oblong; 7 + 1 palatal ridges................................................................................................................................................ 16

14. Tibia dorsally furred; P 4 and M 1 squarish in outline; two lower incisors; M 3 reduced but present; forearm 68–88 mm (♂♂) and 67–90 mm (♀♀); weight 60–97 g (♂♂) and 60–100 g (♀♀); third metacarpal 73–75 % of forearm length; tibia 42–46 % of forearm length .......................................... ............................................................................................. Myonycteris angolensis (Bocage, 1898)

– Tibia dorsally practically naked over distal quarter to third; P 4 and M 1 oblong in outline; one or two lower incisors; M 3 reduced or absent; forearm 54–76 mm; third metacarpal 67–73 % of forearm length; tibia 36–43 % of forearm length.......................................................................................... 15

15. Interfemoral membrane only furred near legs; palatal ridge pattern 6 + 2 (sometimes 7 + 2); two lower incisors; M 3 absent; forearm 65–76 mm .......................... Myonycteris relicta Bergmans, 1980

– Interfemoral membrane wholly furred; palatal ridge pattern 7 + 2; one or two lower incisors; M 3 reduced but normally present; forearm 54–68 mm ........... Myonycteris torquata (Dobson, 1878)

16. Forearm 115–130 mm; M 1 equal in length to M 2 and M 3 combined; tympanic forming a short tubular bony auditory meatus; premaxillae spaced in front; colour generally tawny ..................................... …………… ......................................................................................... Eidolon helvum (Kerr, 1792)

– Forearm 79–107 mm; M 1 shorter than M 2 and M 3 combined; tympanic not forming a bony auditory meatus; premaxillae in contact or co-ossified in front; colour generally not tawny nor yellowish .... ......................................................................................................................................................... 22

17. Cheekteeth 4 / 5, 4 / 6 or 5 / 6; unusual weak dentition; interfemoral membrane greatly reduced; calcar absent or rudimentary; forearm 47–53 mm ................................... Plerotes anchietae (Seabra, 1900)

– Cheekteeth 3 / 5; interfemoral membrane well developed; calcar present; forearm generally larger (except for Micropteropus ) .............................................................................................................. 18

18. Size very large, forearm 118–133 mm; rostrum very deep, end of muzzle truncated, ending in fleshy plate; no white shoulder tuft in ♀ .................................... Hypsignathus monstrosus H. Allen, 1862

– Size smaller, forearm 45–100 mm; rostrum shallow, end of muzzle simple; white shoulder tuft in ♀ .... ......................................................................................................................................................... 19

19. Forearm 60–100 mm ........................................................................................................................ 20

– Forearm 45–55 mm .......................................................................................................................... 21

20. Six prominent palatal ridges, not more than two being post-dental; the latter not differentiated from inter-dental ridges; post-dental palate strongly concave posteriorly (genus Epomophorus )........... 24

– Five, or 9–11, palatal ridges, with sometimes from 5 to 7 post-dental; latter strongly differentiated from inter-dental ridges; post-dental palate flattened posteriorly (genus Epomops ) ....................... 29

21. Rostrum broad; 5–6 palatal ridges, obviously divided by median groove (genus Micropteropus ). 30

– Rostrum slender; 12–13 palatal ridges, only posterior group narrowly divided ................................ ........................................................................................... Nanonycteris veldkampii (Jentink, 1888)

22. Pm 1 subequal in bulk to a lower incisor; premaxillae co-ossified; braincase slightly deflected; wings from second toe; fur long and silky; forearm 79–83.5 mm ................................................................. ............................................................................................. Myonycteris angolensis (Bocage, 1898)

– Pm 1 much larger in bulk than a lower incisor; premaxillae in contact; braincase conspicuously deflected; forearm at least 82.0 mm; tibia furred or naked.............................................................. 23

23. Braincase moderately deflected; width of last upper premolar about ⅓ of palate between fronts of last upper premolars, clearly larger than half their lengths; wings from first toe (occasionally between first and second); antitragal lobe distinct; fur short; tibiae dorsally practically naked; predominantly in lowland areas; forearm 85–107 mm (♂♂) and 82–107 mm (♀♀); weight 111–168 g (♂♂) and 100–155 g (♀♀) ........................................... Rousettus aegyptiacus (E. Geoffroy St.-Hilaire, 1810)

– Braincase strongly deflected; cheekteeth excessively narrow; width of last upper premolar about 1 / 5 of that of the palate between fronts of last upper premolars, about half their lengths or slightly more; wings from second toe (occasionally between first and second); antitragal lobe obsolete; fur long and coarse; tibiae dorsally furred; mostly above 1000 m; forearm 85–94 mm (♂♂) and 85–95 (♀♀); weight 102–140 g (♂♂) and 94–162 g (♀♀) ........................... Stenonycteris lanosus Thomas, 1906

24. One conspicuous, thick postdental palatal ridge; total length of skull 44.4–57.3 mm (♂♂), 41.0– 55.5 mm (♀♀); forearm 72–95 mm (♂♂), 67–89 mm (♀♀) …………………………………………………… ……………………………………………………………………………… Epomophorus wahlbergi (Sundevall, 1846)

– Two postdental palatal ridges .......................................................................................................... 25

25. Size small to medium; total length of skull <50 mm (♂♂) and <46 mm (♀♀); forearm <80 mm (♂♂) and <76 mm (♀♀)................................................................................................................. 26

– Size larger; total length of skull> 51.8 mm (♂♂) and> 44 mm (♀♀); forearm> 80 mm (♂♂) and> 75 mm (♀♀); fourth palal ridge mid-way between third and fifth................................................... ............................................................................................... Epomophorus crypturus Peters, 1852

26. Two thick postdental palatal ridges in ♂♂, one postdental and one partially postdental in ♀♀; total length of skull 47.1 mm (♂), 38.7 and>41.2 mm (♀♀); forearm 77 mm (♂♂), 68–74 mm (♀♀) (otherwise known from Malawi and SW Tanzania only) ………………………………………………………… ………………………………………………………………… Epomophorus anselli Bergmans & Van Strien, 2004

– Two thick postdental palatal ridges in both sexes; total length of skull <48 mm (♂♂) and <46 mm (♀♀); forearm <80 mm (♂♂) and <76 mm (♀♀)......................................................................... 27

27. Size smaller; total length of skull <40 mm (♂♂) and <36.5 mm (♀♀) ........................................... ........................................................................... Epomophorus minimus Claessen & De Vree, 1991

– Size medium; total length of skull> 40 mm (♂♂) and> 36.5 mm (♀♀)....................................... 28

28. Size larger; forearm 66.7–80.3 mm (♂♂) and 64.8–78.3 mm (♀♀); weight 54–99 g (♂♂) and 51–81 g (♀♀); mainly woodlands. Distributed from NE Nigeria over S Sudan to Eritrea and to NE Tanzania; furthermore known from few localities in DRC, Malawi and SE Kenya......................................................................... Epomophorus labiatus (Temminck, 1837)

– Size smaller; forearm 57.0–68.9 mm (♂♂) and 54.1–67.6 mm (♀♀); weight 32–58 g (♂♂) and 25–62 g (♀♀); mainly woodlands. Distributed in E Sudan, Ethiopia, Somalia, lower parts of Kenya to S Malawi................................................................................. Epomophorus minor Dobson, 1880

29. Four thick, prominent interdental and about 5–7 thinner postdental palate ridges ............................ ...................................................................................................... Epomops franqueti (Tomes, 1860)

– Three thick and prominent interdental palate ridges; postdental palate with two pairs of prominent, triangular ridges at middle and one or a few thin and serrate ridges at palation border........................................................................................... Epomops dobsonii ( Bocage, 1889)

30. Forearm 50–53 mm; total skull length 29–29.8 mm .............. Micropteropus pusillus (Peters, 1868)

– Forearm 59–63 mm; total skull length 33.2–33.4 mm ... Micropteropus intermedius Hayman, 1963

31. Six lower incisors; two or four upper incisors; forearm normally less than 52 mm ........................... ................................................................................................................. Coleura afra (Peters, 1852)

– Four lower incisors; two upper incisors (very minute, often absent); forearm more than 57 mm .. 32

32. Forearm more than 80 mm (83–96 mm); total skull length 30–33 mm; skull with shallow frontal depression; no radio-metacarpal pouch; bullae complete; fur dark at base......................................... ................................................................................................... Saccolaimus peli (Temminck, 1853)

– Forearm less than 80 mm; total skull length 20–28 mm; skull with shallow ( T. nudiventris ) or deep frontal depression; radio-metacarpal pouch present; bullae incomplete; fur light at base .............. 33

33. Forearm more than 70 mm, posterior part of body naked above and below ....................................... ......................................................................................... Taphozous nudiventris Cretzschmar, 1830

– Forearm less than 70 mm; body fully haired ................................................................................... 34

34. Dorsally light brown, ventrally pale grey to dark greyish-brown; throat usually sepia-brown, darker than belly; no gular patch in either sex; forearm 60–67 mm ............................................................... .......................................................................... Taphozous perforatus E. Geoffroy St. -Hilaire, 1818

– Dorsally grizzled grey, ventrally pure white or stained yellowish; throat white; large gular sac in male; forearm 58–64 mm ............................... Taphozous mauritianus E. Geoffroy St. -Hilaire, 1818

35. Second premolar of lower jaw large (equal to or larger than half of the first lower premolar); I 1–2 bifid (I 2 exceptionally trifid); marked constriction halfway along the height of the tragus ( arge -group) 38

– Second lower premolar small (smaller than half of the first lower premolar); I 1–2 bi- or trifid; no constriction in the tragus.................................................................................................................. 36

36. I 1–2 bifid ............................................................................................................................................ 37

– I 1–2 trifid ( hispida -group).................................................................................................................. 41

37. Tragus semilunate ( macrotis -group)................................................................................................ 42

– Tragus pyriform (free portion narrowest at base; its outer and inner margin evenly convex); forearm length 42–52 mm; ear length 28–37 mm ( thebaica -group) ……………………………………………… ……………………………………………………… ............................ Nycteris thebaica E. Geoffroy St. -Hilaire, 1818

38. Forearm length more than 39 mm; skull length more than 19 mm ................................................. 39

– Forearm length less than 38 mm; skull length less than 19 mm ...................................................... 40

39. Forearm length 47.3– 50 mm; skull length 20.9–22.4 mm; ear length measuring 50–65 % of forearm length ......................................................................................... Nycteris major (K. Andersen, 1912)

– Forearm length 39.5– 46 mm; skull length 19.3–20.9 mm; ear length measuring 63–77 % of forearm length ...................................................................................................... Nycteris arge Thomas, 1903

40. Forearm length 36.5– 37 mm; skull length 18–18.7 mm; tibia longer than half the forearm length ... ........................................................................................................ Nycteris intermedia Aellen, 1959

– Forearm length 32–36.2 mm; skull length 15.9–16.6 mm; tibia shorter than half the forearm length ..................................................................................................... Nycteris nana (K. Andersen, 1912)

41. Forearm length 39–43 mm ............................................................. Nycteris hispida (Schreber, 1774)

– Forearm length 57–66 mm .................................................................... Nycteris grandis Peters, 1865

42. Forearm length 45–52 mm; ear length 22–34 mm .......................... Nycteris macrotis Dobson, 1876

– Forearm length 37–42 mm; ear length 29–34 mm ......................... Nycteris parisii (de Beaux, 1924)

43. Posterior leaflet of noseleaf simple in outline, elliptical or rounded, not tridentate; ears either separate or united by low band; rostrum at least half as long as braincase; small upper premolar present; forearm over 40 mm (family Hipposideridae , genera Doryrhina , Hipposideros , Macronycteris ).. 45

– Posterior leaflet of noseleaf tridentate; ears always separate; rostrum sometimes less than half as long as braincase; small upper premolar present or absent; forearm 31–54 mm ............................ 44

44. Rostrum less than half as long as braincase; ears low, 8 mm, rounded, rim-like; tridentate nose-leaf; small, forearm 31–35 mm; tail not projecting beyond end of interfemoral membrane; two upper premolars on each side; M 3 not reduced (family Rhinonycteridae ) … Cloeotis percivali Thomas, 1901

– Rostrum at least half as long as braincase; ears large, 18–20 mm with pointed tip; larger, forearm 44– 54 mm; tail projecting up to 7 mm beyond end of interfemoral membrane; one upper premolar on each side; M 3 greatly reduced (family Hipposideridae )..... Asellia tridens (E. Geoffroy St.-Hilaire, 1813)

45. Ears long and narrow, pointed; cochleae greatly expanded, their width equal to at least 4 times the distance between them..................................................................................................................... 49

– Ears short, broad, rounded or triangular; cochleae not expanded, their width approximately equal to the distance between them............................................................................................................... 46

46. Ears rounded or broadly triangular; bluntly pointed upper incisors weak, the outer lobe obsolescent or obsolete; crown area of outer lower incisors equal to or only slightly greater than that of the inner lower incisors................................................................................................................................... 50

– Ears triangular; pointed upper incisors strong, usually retaining much of the outer lobe; crown area of the outer lower incisors much greater than that of the inner lower incisors................................ 47

47. Ears with a small projection at the antitragal fold; sphenoidal bridge narrow, not concealing lateral apertures; pterygoid wings underdeveloped; forearm 58–66 mm ……………………………………… ……… ……………………………………………………………………………………… Hipposideros abae J.A. Allen, 1917

– Ears without antitragal modification; sphenoidal bridge wide, partially concealing lateral apertures; pterygoid wings expanded; forearm 78–114 mm ............................................................................ 48

48. Number of lateral leaflets 3; smaller on average, forearm 93–106 mm (♂♂) and 84–101 mm (♀♀); pelage shorter, not slightly wooly; predominantly in savannas........................................................... ................................................................................................. Macronycteris vittatus (Peters, 1852)

– Number of lateral leaflets 3 or 4; larger on average, forearm 100–124 mm (♂♂) and 95–116 (♀♀); pelage longer, slightly wooly; predominantly in forests.............. Macronycteris gigas Wagner, 1845

49. Smaller, length of forearm less than 74.0 mm; anteorbital foramen relatively large .......................... ................................................................................................ Doryrhina cyclops (Temminck, 1853)

– Larger, length of forearm exceeding 74.0 mm; anteorbital foramen relatively small ......................... ..................................................................................... Hipposideros camerunensis Eisentraut, 1956

50. Posterior nose-leaf without a transverse supplementary structure developed from its posterior face or with such a structure low, undeveloped and lacking a serrated upper edge .................................... ...................................................................................... Hipposideros fuliginosus (Temminck, 1853)

– Posterior nose-leaf having a transverse supplementary structure with a serrated upper edge developed from its posterior face...................................................................................................................... 51

51. Anterior upper premolar small, slightly extruded from toothrow, or compressed between canine and second upper premolar..................................................................................................................... 52

– Anterior upper premolar minute, extruded from toothrow, canine and second upper premolar in contact or nearly so........................................................... Hipposideros beatus (K. Andersen, 1906)

52. Frontal sac opening in both sexes; forearm generally less than 48 mm (42–52 mm); tibia 18–22 mm; dorsal pelage greyish or pale orange; median posterior narial compartiment of nasal swelling relatively narrow, with wide lateral inflations ....................... Hipposideros caffer (Sundevall, 1846)

– Frontal sac opening in ♂♂ only; forearm generally over 48 mm (47–55 mm); tibia 18–24 mm; dorsal pelage brownish or rufous; median posterior narial compartment of nasal swelling relatively wide, with narrow lateral inflations ............................................. Hipposideros ruber (Noack, 1893)

53. First upper premolar placed between the upper canine and second premolar or slightly reduced and partially displaced labially............................................................................................................... 54

– First upper premolar missing or very reduced and external to toothrow; the canine and second premolar touching each other or nearly so ...................................................................................... 62

54. Connecting process of nose-leaf rises to a distinct point................................................................. 55

– Connecting process of nose-leaf low, subtriangular or obsolete ..................................................... 56

55. First phalange of fourth finger long, more than half as long as second, 24–28 % of metacarpal length; sella wedge-shaped, top narrow; lancet subtriangular or hastate, tip rounded .................................... ......................................................................................................... Rhinolophus blasii Peters, 1867

– First phalange of fourth finger shorter, 19–23 % of metacarpal length; sella sides slightly concave; lancet hastate, tip bluntly pointed ................................................ Rhinolophus landeri Martin, 1838

56. Connecting process of nose-leaf obsolete or poorly developed; ears large; forearm 50 mm or more ................................................................................................................................................. 57

– Connecting process of nose-leaf low and rounded or subtriangular; forearm 55 mm or less ......... 60

57. Eight or nine internal ear folds ........................................................................................................ 58

– 11 or 12 internal ear folds ................................................................................................................ 59

58. Lancet tip truncated, only moderately exceeding height of sella; ear with eight folds; larger, forearm 55–62 mm; skull with braincase conspicuously constricted behind the mastoid process; infraorbital bridge longer and more slender; lower molars comparatively small; ear length 32– 40 mm .............................................................................. Rhinolophus ruwenzorii J. Eric Hill, 1942

– Lancet tip bluntly pointed, conspicuously exceeding height of sella; ear with nine folds; smaller, forearm 54 mm; skull with braincase not conspicuously constricted behind the mastoid process; infraorbital bridge shorter and stouter; lower molars comparatively large; ear length 28.5 mm ............................................................................................. Rhinolophus hilli Aellen, 1973

59. 11 internal ear folds; forearm length 54.5 mm; ear length 34.5 mm ................................................... ......................................................................... Rhinolophus kahuzi Fahr & Kerbis Peterhans, 2013

– 12 internal ear folds; forearm length 49.7–51.5 mm; ear length 24.2– 29 mm .................................... ........................................................................ Rhinolophus willardi Kerbis Peterhans & Fahr, 2013

60. Forearm length 49–54 mm; skull length 22.4–23.3 mm ...... Rhinolophus alcyone Temminck, 1853

– Forearm length 40–45 mm; skull length 17.6 mm or less ............................................................... 61

61. Sella comparatively narrow (1.2–1.3 mm prior to preservation); lancet subtriangular, almost hastate, sides concave, tip bluntly pointed; horseshoe 6.0–7.4 mm ......... Rhinolophus swinnyi Gough, 1908

– Sella comparatively broad (1.5–1.7 mm after preservation); lancet subtriangular or hastate, tip somewhat rounded; horseshoe 6.7–9.0 mm.................... Rhinolophus simulator K. Andersen, 1904

62. Face and edges of sella naked; forearm length 50–56 mm; lancet hastate, tip rounded; horseshoe 6.6– 9.6 mm, narrower as muzzle; nasal swellings very low; frontal depression very shallow; supraorbital ridges weak; interpterygoid groove absent or very indistinct.............................................................. ........................................................................................... Rhinolophus clivosus Cretzschmar, 1828

– Face and edges of sella with long hairs ........................................................................................... 63

63. Forearm length 60–67 mm; tibia 26–31 mm; ear length 26–36 mm; baculum> 3 mm; anterior medial nasal swellings longer than broad............................ Rhinolophus hildebrandtii Peters, 1878

– Forearm length 53–63 mm; tibia 22–25 mm; ear length 21–38 mm; baculum <3 mm ...................... ......................................................................................... Rhinolophus eloquens K. Andersen, 1905

– Forearm length 47–60 mm; tibia 19–24 mm; ear length 19–28 mm; anterior medial nasal swellings broader than long ................................................................... Rhinolophus fumigatus Rüppell, 1842

64. Second phalange of third digit nearly 3 times as long as the first (family Miniopteridae ; genus Miniopterus ) .................................................................................................................................... 75

– Second phalange of the third digit shorter (family Vespertilionidae ).............................................. 65

65. Ear slightly but evidently funnel-formed; very high braincase (subfamily Kerivoulinae ; genus Kerivoula )........................................................................................................................................ 76

– Ears largely open at their base and not funnel-formed; normal braincase....................................... 66

66. Six Pm and M in upper and lower jaw (subfamily Myotinae , genus Myotis ) ................................. 80

– Less than 6 Pm and M ..................................................................................................................... 67

67. One upper incisor and M 1 and M 2 with W-pattern distorted or nearly absent (subfamily Scotophilinae ; genus Scotophilus )........................................................................................................................... 82

– Two upper incisors or 1 upper incisor and M 1 and M 2 with W-pattern normal (subfamily Vespertilioninae ).............................................................................................................................. 68

68. One upper incisor............................................................................................................................. 85

– Two upper incisors........................................................................................................................... 69

69. Five Pm and M in upper and lower jaw (genera Neoromicia / Pipistrellus / Hypsugo ).................. 99

– Four Pm and M in upper and 5 in lower jaw ................................................................................... 70

70. Ears very enlarged, about 15 mm and more (genus Laephotis )....................................................... 74

– Ears normally much less enlarged ................................................................................................... 71

71. Very short and broad rostrum; the height of the braincase (with the bullae) more than ⅔ of the condylo-basal length; lower lip with fleshy lobes at angle of mouth (genus Glauconycteris )........ 87

– Height of the braincase (bullae included) much less than the distance from incisors to posterior zygoma root ..................................................................................................................................... 72

72. Very shortened wings; the third digit not longer than head and body length; skull much flattened (genus Mimetillus ) ........................................................................................................................... 73

– Normal wings; skull not especially flattened (genus Neoromicia ) .................................................. 94

73. Pelage dark brown; wing membranes dark; occurring in forest habitats............................................. ................................................................................................. Mimetillus moloneyi (Thomas, 1891)

– Pelage pale brown; wing membranes lighter; occurring in savanna habitats...................................... ....................................................................................................... Mimetillus thomasi Hinton, 1920

74. On average larger; greatest skull length 14.1–15.5 mm; ear length 17–22 mm; forearm length 30– 39 mm; palatal length 6.2–7.3 mm ............................................... Laephotis botswanae Setzer, 1971

– On average smaller; greatest skull length 13.7–13.8 mm; ear length 15–18 mm; forearm length 32–36 mm; palatal length 6.1–6.2 mm ...................................... Laephotis angolensis Monard, 1935

75. Forearm length 45–52 mm; greatest skull length 16.4–17.8 mm; hind foot (including claw) 7–11 mm; total length 92–128 mm; weight 8–19 g ..................................... Miniopterus inflatus Thomas, 1903

– Forearm length 35–42 mm; greatest skull length 11.1–14.5 mm; hind foot (including claw) 5–10 mm; total length 82–97 mm; weight 5–10 g ............................................ Miniopterus minor Peters, 1867

– Forearm length 42–47 mm; greatest skull length 13.6–15.5 mm; hind foot (including claw) 6–11 mm; total length 104–115 mm; weight 6–11 g .......................... Miniopterus natalensis (A. Smith, 1833)

– Forearm length 41–45 mm; greatest skull length 13.3–15.5 mm; hind foot (including claw) 7–11 mm; total length 88–103 mm; weight 7–11 g ........... Miniopterus fraterculus Thomas & Schwann, 1906

76. Comb-like fringe of hairs on interfemoral membrane well developed............................................ 77

– Comb-like fringe of hairs on interfemoral membrane absent or scarcely developed...................... 78

77. Dorsal pelage bright reddish-brown with conspicuous frosting (highlighted tips); ventral pelage cream, dirty-white or white; size on average larger, forearm length 28–40 mm; skull longer and more robust, greatest length 14.7–16.5 mm; outer lower incisors usually with three cusps, sometimes four; inner upper incisors uni-, bi- or tricuspid ...................................... Kerivoula argentata Tomes, 1861

– Dorsal pelage dark brown to pale grey with (in)conspicuous frosting, but not bright reddish-brown with conspicuous frosting; ventral pelage sepia brown, grey to whitish; size on average smaller, forearm length 26–34 mm; skull shorter and more delicate, greatest length 11.2–13.5 mm; outer lower incisors with two or three cusps; inner upper incisors uni- or bicuspid.... Kerivoula lanosa (A. Smith, 1847)

78. Size smaller; forearm length 25–30 mm; dorsal pelage pale reddish-brown, fawn-brown or greyishbrown, usually darker at base, without frosting; outer lower incisors unicuspid; inner upper incisors uni- or bicuspid ............................................................................ Kerivoula phalaena Thomas, 1912

– Size larger; forearm length 30 mm or more; dorsal pelage dark, frosted; inner upper incisors bicuspid............................................................................................................................................ 79

79. Dorsal pelage blackish or dark brown; forearm length 32–36 mm; lower incisors unicuspid.......................................................................................... Kerivoula smithii Thomas, 1880

– Dorsal pelage dark brown with terminal third bright rusty-brown or chocolate brown; forearm length 30–33 mm; lower incisors tricuspid .............................................. Kerivoula cuprosa Thomas, 1912

80. With dichromatic (black and orange) wing membranes; forearm length 52–60 mm; sparse fringe of bristle-like hairs along margin of interfemoral membrane between tips of calcars and tip of tail ..................................................................................................... Myotis welwitschii (Gray, 1866)

– Wing membranes uniformly dark; size smaller; no fringe of bristle-like hairs along margin of interfemoral membrane.................................................................................................................... 81

81. Larger, forearm length 47–53 mm; wing membrane uniformly dark or inconspicuously particoloured, black on most of the membrane and orange to dark red along the fingerbones; ventral and dorsal hairs tricoloured............................................................................ Myotis tricolor (Temminck, 1832)

– Smaller, forearm length 33–42 mm; wing membrane uniformly blackish-brown; ventral hairs bicoloured; dorsal hairs bi- or tricoloured ............................................ Myotis bocagii (Peters, 1870)

82. Very large, forearm length 78–88 mm ...................................... Scotophilus nigrita (Schreber, 1774)

– Smaller, forearm not over 61 mm .................................................................................................... 83

83. Ventral pelage dark brown to reddish-orange; habitat rainforest....... Scotophilus nux Thomas, 1904

– Ventral pelage pure white, yellowish, yellow, orange-yellow; habitat savannas and rainforestsavanna mosaics............................................................................................................................... 84

84. Ventral pelage pure white to medium brown, never yellowish; dorsal pelage pale to medium brown, never tinged with yellow or green; forearm length 44–58 mm; skull total length 16– 18 mm .......................................................................... Scotophilus leucogaster (Cretzschmar, 1826)

– Ventral pelage pale yellow, bright yellow or orange-yellow, not tinged with brown; dorsal pelage sepia brown, greenish-brown, greyish-brown or reddish-brown; forearm 47–60 mm; skull total length 18–21 mm ................................................................... Scotophilus dinganii (A. Smith, 1833)

85. Penis short (5.6–6.2 mm); upper canine with anterior surface rounded, not grooved; anterior lower premolar ca half height of posterior premolar ...................... Nycticeinops schlieffenii (Peters, 1859)

– Penis extremely long (ca 9–16 mm);upper canine with anterior surface flat and grooved; anterior lower premolar at least two-thirds height of posterior premolar, usually subequal (genus Scotoecus )..... 86

86. Wing membranes dark-brown, slightly translucent.......................... Scotoecus hindei Thomas, 1901

– Wing membranes light, semi-translucent, becoming brownish around edges and near body............. ................................................................................................ Scotoecus albofuscus (Thomas, 1890)

87. Wing and interfemoral membranes conspicuously veined; wings pale; colour of dorsal hairs creamybuff to yellowish-fawn (tricoloured or inconspicuously bicoloured); forearm length 38–45 mm ...... ............................................................................................ Glauconycteris variegata (Tomes, 1861)

– Wing and interfemoral membranes not conspicuously veined ........................................................ 88

88. Dorsal body colour black and white; five to seven dorsal markings (spots, patches and stripes); ears subquadrangular; inner upper incisors unicuspid; forearm length 45–48 mm .................................... .............................................................................................. Glauconycteris superba Hayman, 1939

– Dorsal body colour not black and white; ears rounded ................................................................... 89

89. No body pattern; dorsal body colour umber-, sepia-, or reddish-brown; dorsal hairs tricoloured; inner upper incisors strongly bicuspid, with unequal cusps; forearm length 34–38 mm ............................. .................................................................................. Glauconycteris curryae Eger & Schlitter, 2001

– Markings restricted to flank stripes or shoulder spots; dorsal hairs bi- or tricoloured; inner upper incisors generally bicuspid (except for argentata ), with subequal or unequal cusps; forearm length 44 mm or less................................................................................................................................... 90

90. Wing and interfemoral membranes pale brown; dorsal hairs pale golden-fawn or darker and greyer ................................................................................ Glauconycteris argentata (Dobson, 1875)

– Wing and interfemoral membranes dark brown; dorsal hairs medium to dark greyish or dark sepia brown ............................................................................................................................................... 91

91. Inner upper incisors strongly bicuspid, with subequal or unequal cusps; dorsal hairs tricoloured; no body patterns or at most a shoulder spot ......................................................................................... 92

– Inner upper incisors weakly bicuspid, with unequal cusps; dorsal hairs bicoloured; body pattern includes flank stripe and/or shoulder spot ....................................................................................... 93

92. No light lateral markings on body (perhaps rarely a white shoulder spot); forehead region of skull weakly concave....................................................................... Glauconycteris beatrix Thomas, 1901

– White shoulder spot is generally present; forehead region of skull moderately concave.................... ........................................................................................ Glauconycteris humeralis J.A. Allen, 1917

93. Dorsal hairs sepia brown; wings not reticulated........... Glauconycteris alboguttata J.A. Allen, 1917

– Dorsal hairs medium to dark greyish or brownish; wings faintly reticulated...................................... .................................................................................................. Glauconycteris poensis (Gray, 1842)

94. Wings pale or white translucent; mid-ventral pelage cream, whitish or pale grey.......................... 98

– Wings dark; mid-ventral pelage whitish or pale greyish-brown to medium brown ........................ 95

95. Mid-ventral pelage white, whitish, cream or buff; forehead region of skull straight to weakly concave; inner upper incisors weakly bicuspid (unicuspid when worn); forearm length 28–38 mm; occipital helmet usually well developed............................................... Neoromicia capensis (A. Smith, 1829)

– Mid-ventral pelage pale (greyish-)brown to (medium-)brown; forehead region of skull weakly to moderately concave (exceptionally almost straight); occipital helmet absent ................................ 96

96. Size large; forearm length 33–38 mm; greatest length of skull 13.0–14.1 mm; dorsal hairs unicoloured; mid-ventral pelage greyish-brown to medium brown; inner upper incisors unicuspid; no anterior upper premolar; forehead region of skull weakly concave...... Neoromicia brunnea (Thomas, 1880)

– Size smaller; forearm length less than 33 mm; dorsal hairs not unicoloured; inner upper incisors usually unicuspid; anterior upper premolar extremely rarely present, in which case minute ......... 97

97. Dorsal and ventral hairs faintly bicoloured; mid-ventral colour pale brown or brown; forehead region of skull weakly concave to almost straight; forearm length 22–32 mm .............................................. ................................................................................................ Neoromicia somalica (Thomas, 1901)

– Dorsal hairs bi- or tricoloured;mid-ventral colour pale greyish-brown or brown;forehead region of skull weakly to moderately concave; forearm length 27–33 mm .... Neoromicia zuluensis (Roberts, 1924)

98. Dorsal colour dark brown, in strong contrast to pale wings; dorsal hairs usually unicoloured; ventral hairs bicoloured; inner upper incisors weakly bicuspid; forearm length 28–33 mm .......................... ............................................................................................... Neoromicia tenuipinnis (Peters, 1872)

– Dorsal colour light brown, not in strong contrast to pale wings; dorsal hairs bi- or tricoloured; ventral hairs bicoloured; inner upper incisors unicuspid; forearm length 29–38 mm ..................................... .................................................................................................. Neoromicia rendalli (Thomas, 1889)

99. Outer upper incisor greatly reduced; entire underside pure white; dorsal pelage grey to sepia brown [lower molars nyctalodont; forearm length 29–37 mm; anterior upper premolar medium sized, conspicuous, within toothrow or slightly displaced; dorsal and ventral hairs bicoloured] ................. ................................................................................................. Pipistrellus rueppellii (Fischer, 1829)

– Outer upper incisor not greatly reduced; underside light, dark or grey-tipped.............................. 100

100. Inner upper incisors unicuspid; both dorsal and ventral hairs bicoloured; dorsal pelage greyishbrown, reddish-brown, dark brown or blackish; anterior upper premolar medium-size to minute, usually visible, within toothrow or fully displaced lingually; no white border on hind border of wing; forearm length 29–38 mm ................................... Pipistrellus hesperidus (Temminck, 1840)

– Inner upper incisors bicuspid....................................................................................................... 101

101. Dorsal and ventral hairs bicoloured............................................................................................. 102

– Dorsal hairs unicoloured; ventral hairs usually unicoloured ....................................................... 105

102. Outer edge of tragus angular (hatchet-shaped); dorsal pelage chocolate brown or sepia brown, sometimes with gold sheen; anterior upper premolar medium-sized, slightly displaced lingually; no white border on hind border of wing; forearm length 25–35 mm ................................................ ....................................................................................................... Neoromicia nana (Peters, 1852)

– Outer edge of tragus curved (no sharp angle).............................................................................. 103

103. Skull larger, very weakly concave, greatest length 13.7–14.1 mm; wings without white hind border; forearm length 32–36 mm; mid-ventral pelage slightly reddish-brown tinged with beige; anterior upper premolar minute to medium-sized, usually visible, occasionally absent ................................ ...................................................................................................... Hypsugo eisentrauti (Hill, 1968)

– Skull smaller, weakly concave, greatest length less than 13.7 mm; wings with or without white hind border........................................................................................................................................... 104

104. Mid-ventral pelage pale brown, cream or white; anterior upper premolar minute, not visible above gum; greatest length of skull 11.2–13.7 mm; forearm length 29–34 mm ......................................... ................................................................................................... Hypsugo anchietae (Seabra, 1900)

– Mid-ventral pelage dark to medium rust-brown, sometimes speckled; anterior upper premolar absent; greatest length of skull 10.7–11.8 mm; forearm length 23–35 mm ...................................... ............................................................................................ Neoromicia guineensis ( Bocage, 1889)

105. Size larger, forearm 33–38 mm; greatest length of skull 13.8–14.7 mm; mid-ventral pelage pale brown, usually yellowish; anterior upper premolar minute, sometimes not visible, sometimes missing; forehead region of skull weakly concave, sometimes almost straight................................ ........................................................................................... Pipistrellus grandidieri (Dobson, 1876)

– Size smaller, forearm 33 mm or less; greatest length of skull 13.6 mm or less .......................... 106

106. Anterior upper premolar medium-sized to well-developed, visible, in toothrow or slightly lingually displaced; dorsal pelage reddish-brown; white border on wing edge usually missing; forearm length 24–29 mm ................................................................................. Pipistrellus nanulus Thomas, 1904

– Anterior upper premolar minute, barely visible above gum or not at all .................................... 107

107. Size larger, forearm length 28–33 mm; greatest length of skull 12.1–13.6 mm; mid-ventral pelage pale to dark brown or reddish-brown........................................ Hypsugo crassulus (Thomas, 1904)

– Size smaller, forearm length 23–26 mm; greatest length of skull 10.3–11.4 mm; mid-ventral pelage dark umber-brown ................................................................. Hypsugo musciculus (Thomas, 1913)

108. A prominent vertical projection on the zygomatic arch; ears very large, 38–40 mm, conjoined on extended snout; size very large, forearm length 62–73 mm; conspicuous bicolored dorsal pelage pattern, with pale collar separating dark anterior and posterior regions of body .............................. .......................................................................................... Otomops martiensseni (Matschie, 1897)

– Not combining these characters................................................................................................... 109

109. Ears widely separated; ears and wing membranes translucent or whitish; underside of body entirely white; 4 post-canine upper cheek teeth, M 3 much reduced; one lower incisor on each side, with concave crowns lengthened antero-posteriorly; upper incisors in contact with canines; no palatal emargination (genus Myopterus ) ..................................................................................................113

– Ears not widely separated, sometimes conjoined, never translucent; 4 or 5 post-canine upper cheek teeth (first upper premolar often minute, sometimes deciduous); one to three (generally two) lower incisors; upper incisors not in contact with canines .....................................................................110

110. Inner margins of ears meeting or almost meeting to form a V-shaped valley; third ridge of posterior upper molar (M 3) more than half length of second or equal in length; two lower incisors on each side; anterior palatal emargination wide (genus Tadarida ) ......................................................... 133

– Inner margins of ears generally joined by band of skin (uncertain in niangarae , where they might be well separated); anterior palatal emargination narrow or closed .............................................111

111. Third ridge of posterior upper molar (M 3) more than half length of second or equal in length (genus Chaerephon ) .................................................................................................................................114

– Third ridge of posterior upper molar (M 3) less than half length of second or absent (sometimes equal in condylurus ) (genus Mops )...............................................................................................112

112. Anterior palatal emargination closed (subgenus Mops ) .............................................................. 124

– Anterior palatal emargination narrow (subgenus Xiphonycteris )................................................ 130

113. Size smaller, forearm length 33–37 mm; skull length 17–19 mm; pelage dark brown above, without lateral bands ............................................................................... Myopterus whitleyi (Scharff, 1900)

– Size larger, forearm length 48–56 mm; skull length 21.3–25.7 mm; dorsal pelage slaty-grey with two contrasting lateral bands of dirty white .................... Myopterus daubentonii Desmarest, 1820

114. Ears separated by a lobe or lappet ................................................................................................115

– Ears joined by a band....................................................................................................................116

115. Ears long, separated by a large bulbous lobe projecting well over the forehead; ventral flank stripe absent; wing membrane greyish-black; basisphenoid pits deep and large; rostrum uniquely with prominent nasal swellings; forearm length 38 mm ......... Chaerephon gallagheri ( Harrison, 1975)

– Ears separated by a flat lappet over a small, backward-opening pocket; white or whitish ventral flank stripe present; wing membrane dark brown to blackish; depth of basisphenoid pits shallow to moderate, size small to medium; forearm length 39–46 mm ........................................................... .............................................................................................. Chaerephon major (Trouessart, 1897)

116. Size smaller; forearm length less than or equal to 41 mm ............................................................117

– Size larger; forearm length larger than 41 mm .............................................................................119

117. Dorsal pelage pale grey, pale rusty-brown or medium greyish-brown; wing membrane usually white, sometimes pale brown or greyish-brown; ♂♂ with long bicoloured interaural crest....................................................................................... Chaerephon chapini J.A. Allen, 1917

– Dorsal pelage almost black, brown, greyish-brown or reddish-brown; wing membrane white or blackish-brown; ♂♂ with short unicoloured interaural crest .......................................................118

118. Size smaller; forearm length 32.8–37.8 mm; condylobasal length 14.4–15.9 mm; zygomatic width 9.3–10.3 mm .......................................................... Chaerephon leucogaster (A. Grandidier, 1869)

– Size larger; forearm length 38.3–41.0 mm; condylobasal length 15.3–16.4 mm; zygomatic width 10.4–10.9 mm ................................................................ Chaerephon pumilus (Cretzschmar, 1826) 119. Ventral flank stripe present .......................................................................................................... 120

– Ventral flank stripe absent............................................................................................................ 121

120. Flank stripe whitish; wing membranes pale grey; shallow basisphenoid pits of medium size; anterior palatal emargination narrow; forearm length 41–48 mm; uniquely with pair of tail-glands opening ventrally as a slit on each side of base of tail....... Chaerephon bemmeleni (Jentink, 1879)

– Flank stripe white; wing membranes dorsally whitish or blackish, ventrally white; shallow to medium basisphenoid pits of small to medium size; anterior palatal emargination closed; forearm length 41–51 mm; no such glands ........................................... Chaerephon nigeriae Thomas, 1913

121. Anterior palatal emargination closed; size of basisphenoid pits large......................................... 122

– Anterior palatal emarganation narrow; size of basisphenoid pits small to medium.................... 123

122. Size larger; forearm length 48–53 mm; basisphenoid pits deep; wing membrane blackish-brown; sometimes with greyish spots dorsally ........................ Chaerephon aloysiisabaudiae (Festa, 1907)

– Size smaller; forearm length 42–46 mm; basisphenoid pits moderately deep; wing membrane mostly dark brown, lighter towards tip; no spots or stripes. Chaerephon russatus J.A. Allen, 1917

123. Wing membrane light reddish-brown or light grey; forearm length 43–48 mm; dorsal pelage somewhat frosted, occasionally with white spots................ Chaerephon ansorgei (Thomas, 1913)

– Wing membrane dark brown or light reddish-brown; forearm length 46–51 mm; white or whitish stripes and/or rows of spots on crown of most individuals, and sometimes on shoulders and flanks................................................................................... Chaerephon bivittatus (Heuglin, 1861)

124. Size smaller; forearm length 51 mm or less ................................................................................ 125

– Size larger; forearm length 51 mm or more................................................................................. 127

125. ♂♂ with paired glands between penis and anus; forearm length 41–46 mm; phalanges of 3rd (16– 19 mm) and 4th (13–16 mm) fingers short; ventral pelage white or very pale; crown darker than back; whitish or pale grey flank stripe present, but faint; basisphenoid pits deep and large............. .................................................................................... Mops (Mops) demonstrator ( Thomas, 1903)

– ♂♂ without paired glands between penis and anus; phalanges of 3 rd and 4 th fingers medium to long .............................................................................................................................................. 126

126. Forearm length 45–51 mm; phalanges of 3rd (20–24 mm) and 4th (17–20 mm) fingers long; ventral pelage with little or no white; crown same colour as back; white ventral flank stripe present or absent; basisphenoid pits shallow and small ................ Mops (Mops) condylurus (A. Smith, 1833)

– Forearm length 44–48 mm; phalanges of 3rd (19–21 mm) and 4th (15–17 mm) fingers of medium length; ventral pelage predominantly white or cream; crown darker than back; ventral flank stripe absent; basisphenoid pits moderately deep and medium to large in size........................................... .......................................................................... Mops (Mops) niveiventer Cabrera & Ruxton, 1926

127. Size largest; forearm length 59–67 mm; white or pale ventral flank stripe present; dorsal pelage colour variable; frosted and flecked; basisphenoid pits moderately deep and medium to large in size ....................................................................................... Mops (Mops) midas (Sundevall, 1843)

– Size smaller; forearm 58 mm or less ........................................................................................... 128

128. Inner margin of ears uncertain, possibly well separated; ventral flank stripe probably absent; wing membrane dark brown; forearm length approximately 52 mm ......................................................... ......................................................................................... Mops (Mops) niangarae J.A. Allen, 1917

– Inner margin of ears joined by a band ......................................................................................... 129

129. Forearm length 51–55 mm; wing membrane dark brown; dorsal pelage medium sepia or yellowishbrown, or pale rusty-brown.................................................... Mops (Mops) trevori J.A. Allen, 1917

– Forearm length 54–58 mm; wing membrane blackish-brown; dorsal pelage dark brown to almost black; not frosted or flecked .............................................. Mops (Mops) congicus J.A. Allen, 1917

130. Size smaller, forearm length 31 mm or less................................................................................. 131

– Size larger, forearm length 34 mm or more................................................................................. 132

131. Two lower incisors on each side; lower canines without greatly enlarged cingula; upper incisors procumbent (inclined towards the lips); wing membrane blackish-brown, pale brown to whitish, ventral side paler; forearm length 27–31 mm ....... Mops (Xiphonycteris) nanulus J.A. Allen, 1917

– Usually one lower incisor on each side, rarely two; lower canines with greatly enlarged cingula, especially in ♂♂; wing membrane blackish; forearm length 27–30 mm .......................................... ............................................................................. Mops (Xiphonycteris) spurrelli (Dollman, 1911)

132. Ventral flank stripe present, but contrast weak; wing membrane blackish; ventral pelage dark; occipital helmet prominent; wing insertion higher on body; basisphenoid pits of moderately depth and small to medium in size; forearm length 35–42 mm …………………………………………………………… …………………………………………………………………… Mops (Xiphonycteris) brachypterus (Peters, 1852)

– Ventral flank stripe absent; wing membrane various shades of brown; ventral pelage pale; occipital helmet moderately prominent; wing insertion lower on body; basisphenoid pits moderate or shallow deep and medium to large in size; forearm length 34–41 mm ........................................................... .............................................................................. Mops (Xiphonycteris) thersites ( Thomas, 1903)

133. Little white on ventral surface; forearm length 60–67 mm; both sides of wing membrane dark brown to almost black; basisphenoid pits medium in size; plantar pad absent ................................. ................................................................................................... Tadarida ventralis (Heuglin, 1861)

– Clear white, cream or yellowish ventral flank band present; forearm length 56–61 mm; dorsal side of wing membrane brown, ventral side whitish; basisphenoid pits large in size; plantar pad present..................................................................................... Tadarida fulminans ( Thomas, 1903)