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