Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae) Génier, François Moretto, Philippe Zootaxa 2017 4248 1 1 110 363WV Genier Genier [151,596,570,596] Insecta Scarabaeidae Digitonthophagus Animalia Coleoptera 53 54 Arthropoda species viridicollis sp. nov.     Typelocality. Moanda, Zaïre[= Democratic Republic of the Congo].   Diagnosis. Pronotum rather glossy ( Fig. 64); FLP sclerite subapicodorsal lobe short anteriorly ( Figs. 162– 163); right lateral fold produced into a longitudinal spoon-shape process with apical portion open ventrally, ventral edge with minute villi ( Fig. 186 A), with a well-sclerotized projection on apicoventral edge ( Fig. 186B); axial sclerite shorter than subaxial sclerite, acute apically ( Fig. 163); subaxial sclerite short and robust ( Fig. 163).   Description. Holotype ♂( Fig. 16). Measurements. Length 12.5 mm, width 7.0 mm. Head( Figs. 41–42). Anterior clypeal edge straight on median fourth in dorsal view; clypeofrontal carina broadly arcuate and interrupted at gena; vertex lacking median tubercle, surface with punctures fine to small, separated by one to four diameters. Horns rather short, slightly divergent in frontal view, parallel sided on basal three-fourth and abruptly tapering externally and narrower on apical fifth; posterointernal edge with a distinct low angular projection basally; apicointernal surface lacking granules, with few scabrous punctures; genal edge upturned and bluntly angulate on anterior third, forming a broad angle with clypeal edge. Pronotum( Fig. 64). Surface with granulate punctures extending on a short distance on posterior half with distinct, finely granulate, simple punctures on posterior half of disc; punctures scattered and smaller posterolaterally with distinct, minute punctures throughout. Anteromedian tubercle atrophied, simply round in lateral view; median longitudinal sulcus weakly-defined and shallow; surface behind the eyes with a simple, round, rather deep depression; surface of anterior angles slightly concave; anterior half of lateral edge arcuate in dorsal and lateral view; posterior angles unmodified and simply arcuate in dorsal view. Anterior hypomeral ridge arcuate anteriorly, anterior hypomeral depression surface slightly darker in color medially. Elytra( Fig. 16). Intervals 2 and 4 with the odd fine granules on apical declivity. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus( Fig. 128). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites( Figs. 162–164). Axial sclerite broadly arcuate, with apex acute. Subaxial sclerite, gradually tapering from base to apex, extending straight approximately in line with apex of right lateral fold, with a brush of villi on apical fourth. Frontolateral peripheral sclerite basoventral apophysis moderately developed; lacking medioventral carinae; right lateral fold produced into a rather large everted and open internally conical process with irregular apical edge, always with a well-sclerotized projection on apicoventral edge, ventral edge bordered with minute villi; left lateral lobe membranous, slightly developed; subapicodorsal lobe membranous, not reaching anterior edge, apex set medially in dorsal view; apical lobe round and directed obliquely on left side, left edge emarginate, apical villi regular in shape; subapicoventral lobe round and interrupted in line with apical lobe, with few villi along apical edge only.  Variation. Measurements ( 170 ♂♂, 112 ♀♀). Length: male 7.0–13.0 mm (9.8 ± 1.0 mm), female 8.0– 11.5 mm(9.7 ± 0.8 mm). Female allotype. Cephalic outline in dorsal view as in Fig. 80; vertex with a straight transverse carina, dorsal edge broadly arcuate in frontal view, lateral portion round and gradually sloping down posteriorly; anterior pronotal tubercles moderately developed, external lateral edges strongly divergent toward apex, anterolateral surface slightly concave, anterosuperior edge slightly arcuate in dorsal view (Fig. 105), lateral portion of anterosuperior edge strongly upturned (Fig. 106). Protibia short, with external teeth more robust.  Primary type data. Holotype ♂ (CMNC): [Moanda / (Zaïre) 30.iv.1979] partly handwritten; [WORLD / SCARAB. / DATABASE / WSD00032092] barcode label; [HOLOTYPE ♂ /  Digitonthophagus/  viridicollis n.sp./ des. F. Génier, 2016] red card.  MAP 11.Distribution of  Digitonthophagus viridicollis.     Materialexamined( 183 ♂♂, 115 ♀♀), distribution (Map 11):  ANGOLA: [unspecified locality], [no date], [anonymous]—2 ♂♂ (2 paratypes) (BMNH); [unspecified locality], [no date], Monteiro— 3 ♂♂( 3 paratypes) ( BMNH); BENGUELA, Benguela( 12°35'S, 13°25'E), iv.1981, L. Hilburger—2 ♀♀, 3 ♂♂ (5 paratypes) (OUMNH);  CABINDA, Cabinda( 5°34'S, 12°11'E), 1885, Hesse— 1 ♂( paratype) ( SMF);  CUANZA SUL, Cossoço[= Coçoço] ( 11°24'S, 14°1'E),  iii.2005, [anonymous]—2 ♀♀, 3 ♂♂ (5 paratypes) (PMOC); Cossoço [=Coçoço] (11°24'S, 14°1'E), 3–13.iii.2005, M. Hansson & Th. Bouyer—1 ♂ (paratype) (PMOC); [unspecified locality], ii–iii.1999, T. Bouyer & M. Hasson—2 ♀♀, 1 ♂ (paratypes) (PMOC);  CUNENE, 10 kmN Humbe( 16°35'49''S, 14°54'24''E),  3.xi.2011, G. Werner— 2 ♀♀, 4 ♂♂( 6 paratypes) ( FGIC, GWPC);  Near Categuero( 17°45.765'S, 14°29.411'E), 11.xii.2012, S. Rojkoff—1 ♂ (paratype) (CMD); Roçadas (16°45'S, 14°59'E),   19– 22.ii.1972, SouthernAfrica Expedition— 3 ♀♀, 3 ♂♂( 6 paratypes) ( BMNH);  HUILA, 26 kmS Cahila( 16°6'18.3''S, 14°14'7.6''E), 2.xii.2015, G. Werner—1 ♂ (paratype) (GWPC); 5 mi. NE Negola (14°8'S, 14°30'E),   25.iii.1972, SouthernAfrica Expedition— 2 ♀♀, 1 ♂( paratypes) ( BMNH); João de Almeida[=Chibia] ( 15°11'S, 13°42'E),   29.iii.1972, [anonymous] ( A41)— 1 ♂( paratype) ( BMNH);  LUANDA, Luanda( 8°50'S, 13°17'E), [no date], Welwitch— 1 ♀, 1 ♂( paratypes) ( BMNH);  Luanda( 8°50'S, 13°17'E), [no date], [anonymous]— 1 ♂( paratype) ( IRSNB);  UIGE, 3 mi. N Santa Clara( 7°22'S, 14°44'E),   3.iii–1.iv.1972, SouthernAfrica Expedition— 3 ♀♀, 2 ♂♂( 5 paratypes) ( BMNH);   CONGO: BRAZZAVILLE,  Environs de Brazzaville( 4°15'S, 15°16'E), 1907, E. Roubaud& A Weiss— 1 ♂( paratype) ( MNHN);   DEMOCRATIC REPUBLIC OF THE CONGO: BAS- CONGO, Banana( 5°59'53''S, 12°23'56''E), [no date], F. Busschodts— 2 ♂♂( 2 paratypes) ( IRSNB);  Banana( 5°59'53''S, 12°23'56''E), iii.1885, Hesse—3 ♀♀, 8 ♂♂ (11 paratypes) (SMF); Banana (5°59'53''S, 12°23'56''E), v.1886, Hesse—4 ♂♂ (4 paratypes) (SMF);  Banana-Boma( 5°50'S, 12°43'E), 1891, M. Tschoffen— 4 ♂♂( 4 paratypes) ( IRSNB);  Boma( 5°51'S, 13°3'E), [no date], P. Rolin— 1 ♀, 1 ♂( paratypes) ( IRSNB);  Boma( 5°51'S, 13°3'E), [no date], M. Tschoffen— 4 ♂♂( 4 paratypes) ( IRSNB);  Boma( 5°51'S, 13°3'E),  3.i.1977, P. Walter— 8 ♀♀, 6 ♂♂( 14 paratypes) ( PHWC);  Boma( 5°51'S, 13°3'E),  4.i.1977, P. Walter— 16 ♀♀, 14 ♂♂( 30 paratypes) ( PHWC);   Environs de Kangu, Mayumbé( 5°17'S, 12°57'E),  6.i.1977, Yimi-di-Phambu— 18 ♀♀, 21 ♂♂( 39 paratypes) ( PHWC);  Kwilu( 5°30'S, 14°41'E),  iv.1885, Hesse— 1 ♀, 1 ♂( paratypes) ( SMF);  Matadi( 5°49'S, 13°28'E), [no date], [anonymous]—1 ♂ (paratype) (IRSNB); Muanda (5°56'S, 12°21'E), 30.iv.1979, P. Walter—1 ♀, 4 ♂♂ (holotype, allotype, 3 paratypes) (CMNC, PHWC);  Vista[= Nsiamfumu] ( 5°52'S, 12°17'E), [no date], V. Moerenhout— 1 ♂( paratype) ( IRSNB);  Vivi( 5°48'S, 13°27'E), [no date], [anonymous]— 1 ♂( paratype) ( IRSNB);  Zambi( 5°51'S, 12°52'E), [no date], M. Tschoffen— 2 ♀♀, 6 ♂♂( 8 paratypes) ( IRSNB);  Zambi( 5°51'S, 12°52'E), [no date], C. Haas— 1 ♂( paratype) ( IRSNB);   NAMIBIA: ERONGO, Ameib Farm,  19 kmNW Karibib( 21°47'S, 15°38'E),   31.i–2.ii.1972, SouthernAfrica Expedition— 2 ♀♀( 2 paratypes) ( BMNH); Otjikoko SüdFarm, 33 mi.  ENE Omaruru( 21°10'S, 16°22'E),  10–13.ii.1972, SouthernAfrica Expedition— 3 ♀♀, 3 ♂♂( 6 paratypes) ( BMNH);  KUNENE, Kamanjab( 19°38'S, 14°51'E),  9.iv.2005, W. Schawaller— 1 ♀, 1 ♂( paratypes) ( SMNS);  Ongongo( 18°8'S, 13°22'E),  17–18.iv.2005, W. Schawaller— 1 ♂( paratype) ( SMNS);  Otjitambifarm, 27 mi. ESE Kamanjab( 19°49'S, 15°10'E),  13–15.ii.1972, SouthernAfrica Expedition— 1 ♀, 2 ♂♂( 3 paratypes) ( BMNH);  Outjo( 20°7'S, 16°10'E),  i–ii.1998, Forti, Gzeppel& Giannatelli— 1 ♂( paratype) ( FETC);  Purros( Hoaruzsib Valley) ( 18°46'S, 12°57'E),  15–16.iv.2005, W. Schawaller— 1 ♂( paratype) ( SMNS);  OHANGWENA, Oshikango( 17°24'S, 15°53'E),  19.ii.1972,  SouthernAfrica Expedition— 1 ♂( paratype) ( BMNH);  OMAHEKE, Kuzikus Wildlife Reserve( 23°14'17''S, 18°23'28''E),  30.iii–11.iv.2011, J. Constant— 3 ♂♂( 3 paratypes) ( IRSNB);  Kuzikus Wildlife Reserve( 23°14'17''S, 18°23'28''E),  3–10.iv.2011, J. Constant— 1 ♂( paratype) ( IRSNB);  OMUSATI, 2 kmE Renostervlei( 19°9'59''S, 14°33'12''E),  26.xii.1999, Mann& Marais— 1 ♂( paratype) ( OUMNH);  Narawanduroad, Etosha National Park( 18°51'4''S, 15°32'55''E),  24.xii.1999, Mann, Marais& Newman— 2 ♀♀, 1 ♂( paratypes) ( OUMNH);  Narawanduroad, Etosha National Park( 18°51'4''S, 15°32'55''E),  24–25.xii.1999, Mann& Marais— 1 ♂( paratype) ( OUMNH);  Narawanduroad, Etosha National Park( 18°51'4''S, 15°32'55''E),  24.xii.1999, Mann& Marais— 1 ♂( paratype) ( OUMNH);  Narawanduroad, Etosha National Park( 18°51'4''S, 15°32'55''E),  25– 26.xii.1999, Mann& Marais— 1 ♀, 1 ♂( paratypes) ( OUMNH);  OSHANA, Oshakati( 17°52'30''S, 15°22'30''E),  15.vii.1975, R. Venter— 1 ♂( paratype) ( BMNH);  OSHIKOTO, Andoniroad ( 18°45'37''S, 16°54'24''E),  20.xii.1999, Mann& Marais— 6 ♀♀, 6 ♂♂( 12 paratypes) ( OUMNH);  Etosha National Park( 19°2'S, 16°28'E),  i– ii.1998, Czeppel, Forti, & Giannatelli— 1 ♀( paratype) ( FETC);  Etosha National Park( 19°2'S, 16°28'E),  13.ii.1998, Forti— 1 ♂( paratype) ( SMNS);  Mushara area, sandy road ( 18°36'24''S, 16°53'23''E),  21.xii.1999, Mann& Marais— 5 ♀♀, 4 ♂♂( 9 paratypes) ( OUMNH);  Mushara Waterhole( 18°35'19''S, 16°55'11''E),  22–23.xii.1999, Mann& Marais— 2 ♂♂( 2 paratypes) ( OUMNH);  Mushara Waterhole( 18°35'19''S, 16°55'11''E),  23.xii.1999, Mann& Marais— 3 ♀♀, 2 ♂♂( 5 paratypes) ( OUMNH);  NorthTsintsabis ( 18°46'S, 17°58'E),  1.i.2007, R. Perissinoto& L. Clennell— 1 ♂( paratype) ( PMOC);  Okaukuejo, Etosha National Park,  1113 m( 19°10'S, 15°55'E),  i.2007, M. Forti— 1 ♂( paratype) ( SMNS);  Oluconda( 17°59'7''S, 16°1'24''E), 1901, Schinz— 1 ♀, 1 ♂( paratypes) ( SMF);  Onguma Farm, 55 mi. NW Tsumeb( 18°44'S, 17°3'E),  17–19.ii.1972, SouthernAfrica Expedition— 7 ♀♀, 4 ♂♂( 11 paratypes) ( BMNH);  Otjikata[= Otjikoto] Lake( 19°11'S, 17°33'E),  16.xi.1933, K. Jordan— 7 ♀♀, 4 ♂♂( 11 paratypes) ( BMNH);  OTJOZONDJUPA, 12 mi. E Otavi( 19°39'S, 17°31'E),  17.ii.1972,  SouthernAfrica Expedition— 3 ♀♀, 5 ♂♂( 8 paratypes) ( BMNH);  Epupa Falls,  660 m( 17°0'3''S, 13°14'36''E),  11–12.iv.2005, W. Schawaller— 5 ♂♂( 5 paratypes) ( SMNS);  Hohenfels( 20°41.928'S, 16°50.498'E),  7.i.2011, G. Werner— 1 ♂( paratype) ( FGIC);  Okahandja( 21°59'S, 16°55'E),  11.xi.1909, G. Fock— 2 ♀♀, 3 ♂♂( 5 paratypes) ( IRSNB);  Okahandja( 21°59'S, 16°55'E),  2–18.iii.1922, R.E. Turner— 2 ♂♂( 2 paratypes) ( BMNH);  Okahandja( 21°59'S, 16°55'E),  1–12.i.1928, R.E. Turner— 1 ♂( paratype) ( BMNH);  Okahandja( 21°59'S, 16°55'E),  3–9.ii.1928, R.E. Turner— 2 ♂♂( 2 paratypes) ( BMNH);  Otavi( 19°39'S, 17°20'E),  8.i.1934, J. Ogilvie— 1 ♀( paratype) ( BMNH);  Otavifontein( 19°39'S, 17°24'E),  21.xi.1933, K. Jordan— 1 ♂( paratype) ( BMNH);  Otjawarongo( 20°27'S, 16°40'E), 1909, Külz— 1 ♀, 4 ♂♂( 5 paratypes) ( SMF);  Otjawarongo( 20°27'S, 16°40'E),  13.i.1934, J. Ogilvie— 1 ♀, 1 ♂( paratypes) ( BMNH);  Otjawarongo( 20°27'S, 16°40'E),  4.i.2004, R. Perissinoto& L. Clennell— 1 ♂( paratype) ( PMOC); Swakop River, 3 mi. S Okahandja( 22°2'S, 16°56'E),  7.iv.1972,  SouthernAfrica Expedition— 1 ♂( paratype) ( BMNH);  Grootfontein District, Okaputa334 at: campsite ( 20°7'26''S, 16°57'54''E),  12.xii.1999, Mann, Marais& Newman— 1 ♂( paratype) ( OUMNH).   Etymology.  Viridicollis, a Latin adjective pertaining to the metallic green coloration of the pronotum typical for this species.   Natural history. The material examined shows a Kalahari xeric savanna and Angolan scarp savanna and woodland centered distribution. The only specimens with habitat data were collected in pastures. Some specimens collected at light traps and cow, elephant, and zebra dung. Afew specimens collected in pitfall traps baited with giraffe, zebra, or human dung.   Remarks. This species is very closely related to  D. gazella.Despite the relative high number of individuals examined, no intermediates were found for the character of the shape of the right lateral fold (see diagnosis). Large females of  D. viridicollisalso have very characteristic strongly upturned anterosuperior edges of the anterior pronotal tubercles. The green metallic tinge of the dorsum is also characteristic of most specimens. 1455384669 [151,928,710,735] Democratic Republic of the Congo Zaire Moanda 53 54 1 holotype 1455384650 Angola Material 54 55 298 115 183 1455384679 BMNH Angola -12.583333 BENGUELA 1293 13.416667 54 55 6 3 BENGUELA 1455384653 [298,1225,1030,1055] SMF Angola -5.5666666 Cabinda 1305 12.183333 54 55 1 1 Cabinda paratype 1455384643 2005-03 Angola -11.4 Cococo 1295 14.016666 Cossoco 54 55 1 CUANZA SUL paratype 1455384651 2011-11-03 FGIC, GWPC Angola -16.596945 Humbe 21 14.906667 54 55 12 2 4 CUNENE paratype 1455384667 Angola -17.76275 Near Categuero 1 14.490183 54 55 1 CUNENE paratype 1455384670 1972-02-19 1972-02-22 1972-02-19 BMNH Angola Southern 54 55 12 3 3 Southern paratype 1455384668 Angola -16.105083 Cahila 1 14.235445 54 55 1 HUILA paratype 1455384644 1972-03-25 BMNH Joao de Almeida Angola -15.183333 Southern 1286 13.7 54 55 3 2 1 Southern paratype 1455384661 [266,941,1354,1379] 1972-03-29 BMNH Angola A 54 55 1 1 A paratype 1455384672 BMNH Angola -8.833333 LUANDA 1300 13.283334 54 55 2 1 1 LUANDA paratype 1455384659 IRSNB Angola -8.833333 Luanda 1300 13.283334 54 55 1 1 Luanda paratype 1455384657 [389,897,1426,1451] N Santa Clara Angola -7.366667 UIGE 1303 14.733334 54 55 1 UIGE paratype 1455384652 1972-03-03 1972-04-01 1972-03-03 BMNH Angola Southern 54 55 10 3 2 Southern paratype 1455384663 MNHN Environs de Brazzaville & E. Roubaud Democratic Republic of the Congo -4.25 Environs de Brazzaville 1306 15.266666 54 55 1 1 paratype 1455384664 IRSNB Democratic Republic of the Congo -5.998056 Banana 21 12.398889 54 55 4 2 CONGO paratype 1455384656 Democratic Republic of the Congo -5.998056 Banana 21 12.398889 54 55 1 CONGO paratype 1455384658 IRSNB Democratic Republic of the Congo -5.8333335 Banana-Boma 1305 12.716666 54 55 8 4 CONGO paratype 1455384660 [394,1257,1642,1667] IRSNB Democratic Republic of the Congo -5.85 Boma 1305 13.05 54 55 2 1 1 CONGO paratype 1455384684 IRSNB Democratic Republic of the Congo -5.85 Boma 1305 13.05 54 55 8 4 CONGO paratype 1455384662 1977-01-03 PHWC Democratic Republic of the Congo -5.85 Boma 1305 13.05 54 55 28 8 6 CONGO paratype 1455384647 1977-01-04 PHWC Democratic Republic of the Congo -5.85 Boma 1305 13.05 54 55 60 16 14 CONGO paratype 1455384666 1977-01-06 PHWC Environs de Kangu Democratic Republic of the Congo -5.2833333 Mayumbe 1305 12.95 Environs de Kangu 54 55 78 18 21 CONGO paratype 1455384641 [399,1238,1786,1811] 1885-04 SMF Democratic Republic of the Congo -5.5 Kwilu 1305 14.683333 54 55 2 1 1 CONGO paratype 1455384694 Democratic Republic of the Congo -5.8166666 Matadi 1305 13.466666 54 55 1 CONGO paratype 1455384675 IRSNB Democratic Republic of the Congo -5.866667 Nsiamfumu 1305 12.283334 Vista 54 55 1 1 CONGO paratype 1455384708 [602,1432,1894,1919] IRSNB Democratic Republic of the Congo -5.8 Vivi 1305 13.45 54 55 1 1 CONGO paratype 1455384674 [151,1141,1930,1955] IRSNB Democratic Republic of the Congo -5.85 Zambi 1305 12.866667 54 55 16 2 6 CONGO paratype 1455384677 IRSNB Democratic Republic of the Congo -5.85 Zambi 1305 12.866667 54 55 1 1 CONGO paratype 1455384671 Namibia -21.783333 Ameib Farm 1262 15.633333 54 55 1 ERONGO paratype 1455384693 [265,1115,2003,2028] 1972-01-31 1972-02-02 1972-01-31 BMNH Namibia Southern 54 55 4 2 Southern paratype 1455384695 [1127,1436,2003,2028] Namibia Sud 54 55 1 Sud paratype 1455384692 1972-02-10 1972-02-13 1972-02-10 BMNH -21.166666 Omaruru 1265 16.366667 55 56 12 3 3 Southern 1455384676 [272,1431,188,213] 2005-04-09 SMNS -19.633333 Kamanjab 1271 14.85 55 56 2 1 1 KUNENE paratype 1455384685 [151,1111,223,248] 2005-04-17 2005-04-18 2005-04-17 SMNS -18.133333 Ongongo 1276 13.366667 55 56 1 1 KUNENE paratype 1455384697 1972-02-13 1972-02-15 1972-02-13 BMNH -19.816668 Kamanjab 1270 15.166667 Otjitambi 55 56 6 1 2 Southern paratype 1455384680 [151,1210,295,320] 1998-01 1998-02-31 1998-01 FETC Forti & Gzeppel -20.116667 Outjo 1269 16.166666 55 56 1 1 Southern paratype 1455384683 2005-04-15 2005-04-16 2005-04-15 SMNS -18.766666 Hoaruzsib Valley 1274 12.95 Purros 55 56 1 1 Southern paratype 1455384678 1972-02-19 -17.4 Oshikango 1279 15.883333 55 56 1 OHANGWENA paratype 1455384687 [520,1165,367,392] BMNH Southern 55 56 1 1 Southern paratype 1455384689 2011-03-30 2011-04-11 2011-03-30 IRSNB -23.238056 Kuzikus Wildlife Reserve 21 18.391111 55 56 6 3 OMAHEKE paratype 1455384682 2011-04-03 2011-04-10 2011-04-03 IRSNB -23.238056 Kuzikus Wildlife Reserve 21 18.391111 55 56 1 1 OMAHEKE paratype 1455384681 1999-12-26 OUMNH Mann -19.166388 Renostervlei 21 14.553333 55 56 1 1 OMUSATI paratype 1455384686 1999-12-24 OUMNH Mann & Marais -18.851112 Etosha National Park 21 15.548612 Narawandu 55 56 3 2 1 OMUSATI paratype 1455384701 1999-12-24 1999-12-25 1999-12-24 OUMNH Mann -18.851112 Etosha National Park 21 15.548612 Narawandu 55 56 1 1 OMUSATI paratype 1455384688 1999-12-24 OUMNH Mann -18.851112 Etosha National Park 21 15.548612 Narawandu 55 56 1 1 OMUSATI paratype 1455384673 1999-12-25 1999-12-26 1999-12-25 OUMNH Mann -18.851112 Etosha National Park 21 15.548612 Narawandu 55 56 2 1 1 OMUSATI paratype 1455384690 1975-07-15 BMNH -17.875 Oshakati 21 15.375 55 56 1 1 OSHANA paratype 1455384665 1999-12-20 OUMNH Mann -18.760279 Andoni 21 16.906666 55 56 24 6 6 OSHIKOTO paratype 1455384719 1998-01 1998-02-31 1998-01 FETC Czeppel & Forti -19.033333 Etosha National Park 1273 16.466667 55 56 1 1 OSHIKOTO paratype 1455384700 1998-02-13 SMNS -19.033333 Etosha National Park 1273 16.466667 55 56 1 1 OSHIKOTO paratype 1455384730 1999-12-21 OUMNH Mann -18.606667 Mushara area 21 16.889723 55 56 18 5 4 OSHIKOTO paratype 1455384696 1999-12-22 1999-12-23 1999-12-22 OUMNH Mann -18.588612 Mushara Waterhole 21 16.919722 55 56 4 2 OSHIKOTO paratype 1455384712 1999-12-23 OUMNH Mann -18.588612 Mushara Waterhole 21 16.919722 55 56 10 3 2 OSHIKOTO paratype 1455384705 2007-01-01 PMOC R. Perissinoto -18.766666 North 1274 17.966667 55 56 1 1 North paratype 1455384717 2007-01 SMNS 1113 -19.166666 Etosha National Park 1273 15.916667 Okaukuejo 55 56 1 1 North paratype 1455384720 SMF -17.985277 Oluconda 21 16.023333 55 56 2 1 1 North paratype 1455384718 1972-02-17 1972-02-19 1972-02-17 BMNH NW Tsumeb -18.733334 Onguma Farm 1274 17.05 55 56 22 7 4 Southern paratype 1455384710 1933-11-16 BMNH -19.183332 Otjikoto 1273 17.55 Otjikata 55 56 22 7 4 Lake paratype 1455384713 [453,1233,1087,1112] 1972-02-17 E Otavi -19.65 OTJOZONDJUPA 1271 17.516666 55 56 1 OTJOZONDJUPA paratype 1455384725 BMNH Southern 55 56 16 3 5 Southern paratype 1455384707 2005-04-11 2005-04-12 2005-04-11 SMNS 660 -17.000834 Epupa Falls 21 13.243334 55 56 10 5 Southern paratype 1455384709 2011-01-07 FGIC -20.6988 Hohenfels 1 16.841633 55 56 1 1 Southern paratype 1455384699 [379,1432,1196,1221] 1909-11-11 IRSNB -21.983334 Okahandja 1262 16.916666 55 56 10 2 3 Southern paratype 1455384703 [151,1186,1231,1256] 1922-03-02 1922-03-18 1922-03-02 BMNH -21.983334 Okahandja 1262 16.916666 55 56 4 2 Southern paratype 1455384715 1928-01-01 1928-01-12 1928-01-01 BMNH -21.983334 Okahandja 1262 16.916666 55 56 1 1 Southern paratype 1455384706 1928-02-03 1928-02-09 1928-02-03 BMNH -21.983334 Okahandja 1262 16.916666 55 56 4 2 Southern paratype 1455384711 [604,1432,1303,1328] 1934-01-08 BMNH -19.65 Otavi 1271 17.333334 55 56 1 1 Southern paratype 1455384704 [151,1148,1340,1365] 1933-11-21 BMNH -19.65 Otavifontein 1271 17.4 55 56 1 1 Southern paratype 1455384726 SMF -20.45 Otjawarongo 1268 16.666666 55 56 10 1 4 Southern paratype 1455384702 1934-01-13 BMNH -20.45 Otjawarongo 1268 16.666666 55 56 2 1 1 Southern paratype 1455384698 2004-01-04 PMOC R. Perissinoto -20.45 Otjawarongo 1268 16.666666 55 56 1 1 Southern paratype 1455384716 [385,1088,1447,1472] 1972-04-07 S Okahandja -22.033333 River 1261 16.933332 55 56 1 River paratype 1455384691 BMNH Southern 55 56 1 1 Southern paratype 1455384746 1999-12-12 OUMNH Mann & Marais -20.12389 Okaputa 21 16.965 Grootfontein District 55 56 1 1 Southern paratype