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 363WQ Genier Genier [151,628,1230,1256] Insecta Scarabaeidae Digitonthophagus Animalia Coleoptera 69 70 Arthropoda species namaquensis sp. nov.     Typelocality. Farm Klipkoppies (6) ( 28.87137 S 21.30944 E),  957 m,  NorthernCape, South Africa.   Description. Holotype ♂( Fig. 22). Measurements. Length 10.5 mm, width 6.0 mm. Head( Figs. 53–54). Anterior clypeal edge straight on median fifth 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, scabrous. Horns rather short, slightly arcuate in frontal view, gradually tapering from base to apex; posterointernal edge unmodified basally, apicointernal surface with few scabrous punctures; genal edge slightly upturned and broadly arcuate, forming a broad angle with clypeal edge. Pronotum( Fig. 70). Surface with weakly-defined granulate punctures on anterior half only, with scattered punctures on posterior half of disc; punctures scattered and small posterolaterally, with distinct minute punctures throughout; anteromedian tubercle atrophied, forming an obtuse angle in lateral view, median longitudinal sulcus rather wide and moderately deep; surface behind the eyes with a transverse oval and shallow depression, surface of anterior angles slightly concave; anterior half of lateral edge arcuate in dorsal and lateral view; posterior angles slightly upturned and simply arcuate in dorsal view; anterior hypomeral ridge broadly arcuate anteriorly, anterior hypomeral depression surface light in color. Elytra( Fig. 22). Interval 2 with one to three fine granules on apical declivity, interval 4 lacking fine granules from base to apex. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus( Fig. 134). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites( Figs. 180–182). Axial sclerite arcuate, with apex acute. Subaxial sclerite, gradually tapering from base to apex, extending straight and interrupted before apex of frontolateral peripheral sclerite apical portion, with a brush of villi on apical fourth. Frontolateral peripheral sclerite basoventral apophysis large; lacking medioventral carinae; right lateral fold produced into a rather small everted and open apically conical process with irregular apical edge located behind oblique right lateral edge of apical lobe; left lateral lobe membranous, slightly developed; subapicodorsal lobe membranous, small and obliquely projecting dorsally, interrupted well before anterior edge of apical lobe, apex set on left side in dorsal view; apical lobe truncate and slightly directed obliquely on left side, lateral edges folded ventrally, with a small additional longitudinal fold in line with subapicoventral lobe; apical villi becoming longer toward apex; subapicoventral lobe acute in lateral view, interrupted before apex of apical lobe.  Variation. Measurements ( 97 ♂♂, 74 ♀♀). Length: male 7.0–12.0 mm (10.0 ± 0.9 mm), female 8.0– 11.5 mm(9.9 ± 0.8 mm). Females allotype. Cephalic outline in dorsal view as in Fig. 86; vertex with a high and broadly arcuate, transverse carina, dorsal edge subangular medially in frontal view, lateral portion gradually sloping down posteriorly; anterior pronotal tubercles well developed, external lateral edges distinctly widening toward apex in dorsal view, anterolateral surface obliquely excavated, anterosuperior edge slightly arcuate in dorsal view (Fig. 117), lateral portion of anterosuperior edge slightly upturned (Fig. 118). Anterior tibia short, with external teeth more robust.  Primary type data. Holotype ♂ (CMNC): [SOUTH AFRICA: N. CAPE / Farm Klipkoppies (6) / S28.87137° E21°30.944’ [ sic] / 957 m; 12–14 March 2004/ A.L.V. Davis & C. Deschodt]; [WORLD / SCARAB. / DATABASE / WSD00029107] barcode label; [HOLOTYPE ♂ /  Digitonthophagus/  namaquensis n.sp./ des. F. Génier, 2016] red card.  MAP 17.Distribution of  Digitonthophagus namaquensis.     Materialexamined( 100 ♂♂, 75 ♀♀), distribution (Map 17):  ANGOLA: BENGUELA,  Benguela( 12°35'S, 13°25'E), [no date], [illegible]— 1 ♀, 1 ♂( paratypes) ( IRSNB);   NAMIBIA: ERONGO, Ameib Farm, 19 kmNW Karibib( 21°47'S, 15°38'E),  31.i–2.ii.1972,  SouthernAfrica Expedition— 1 ♂( paratype) ( BMNH);  Gobabeb( 23°34'S, 15°2'E),  10.ii.1978, O. Lomhodlt— 1 ♂( paratype) ( ZMUC); Kahn River, 5 mi. N Usakos( 22°0'S, 15°36'E),  30–31.i.1972,  SouthernAfrica Expedition— 1 ♀, 1 ♂( paratypes) ( BMNH); Kuiseb Canyon( 23°18'S, 15°45'E),  22–23.i.1972,  SouthernAfrica Expedition— 1 ♀, 1 ♂( paratypes) ( BMNH);  HARDAP, Farm Lovedale, N Tiras Mountains,  1600 m( 25°32'24''S, 16°23'24''E),  16–17.ii.2010, W. Schawaller— 1 ♀, 5 ♂♂( 6 paratypes) ( SMNS);  Sesriem Farm, Maltahoe District( 24°11'S, 15°59'E),  19–20.i.1972, SouthernAfrica Expedition— 1 ♀( paratype) ( BMNH);  Voigtgrund( 24°49'S, 17°26'E), [no date], K. Jordan— 1 ♀, 2 ♂♂( 3 paratypes) ( BMNH);  Malthahöhe district, Zais6 ( 24°2'58''S, 16°9'14''E),  16.i.2000, Mann& Marais— 1 ♀( paratype) ( OUMNH);  Rehoboth district, on road verge, Kambes498 ( 24°4'32''S, 16°28'21''E),  17.i.2000, Mann& Marais— 1 ♀, 3 ♂♂( 4 paratypes) ( OUMNH);  KARAS, Barby Farm, 25 mi. W Helmeringhausen( 25°53'S, 16°49'E),  17–18.i.1972,  SouthernAfrica Expedition— 11 ♀♀, 16 ♂♂( 27 paratypes) ( BMNH);  Keetmannshoop( 26°34'S, 18°8'E), 1913, S. Hardt& G. Lotz— 3 ♀♀, 2 ♂♂( 5 paratypes) ( SMF); Mesosaurus Fossil Farm, NE Keetmanshoop, 1100 m(28°14'24''S, 18°16'48''E), 18–19.ii.2010, W. Schawaller—1 ♂ (paratype) (SMNS);  Noachabeb, 27 mi. NNE Grunau( 27°23'S, 18°28'E),  10–12.i.1972, SouthernAfrica Expedition— 3 ♀♀, 5 ♂♂( 8 paratypes) ( BMNH); near Onseepkans, Orange RIVERbanks ( 28°44'S, 19°18'E),  8–10.i.1972,  SouthernAfrica Expedition— 1 ♂( paratype) ( BMNH);  PlateauFarm, 22 mi. E Aus( 26°38'S, 16°34'E),  14–17.i.1972,  SouthernAfrica Expedition— 1 ♀, 1 ♂( paratypes) ( BMNH);  Seeheim( 26°49'S, 17°47'E),  16.ii.1934, J. Ogilvie— 1 ♀( paratype) ( BMNH);  KAVANGO, Mayana Lodge, Rundu District( 17°51'55''S, 19°55'48''E),  19.xii.1999, Mann& Marais— 6 ♀♀, 7 ♂♂( 13 paratypes) ( OUMNH);  Mile46 Agricultutal Station, Sovo area( 18°19'14''S, 19°15'28''E),  23.iii.2002, Mann, Marais, & Kasch— 1 ♀, 1 ♂( paratypes) ( OUMNH);  KHOMAS, Farm Namibgrens, Spreetshoogte,  1800 m( 23°22'12''S, 16°8'24''E),  3–6.ii.2010, W. Schawaller— 1 ♀, 2 ♂♂( 3 paratypes) ( SMNS);  Solitaire( 23°53'S, 16°0'E),  22.i.1972, SouthernAfrica Expedition— 2 ♀♀, 8 ♂♂( 10 paratypes) ( BMNH);  Windhoek( 22°34'12''S, 17°5'1''E), 1909, Külz— 2 ♀♀( 2 paratypes) ( SMF);  KUNENE, 87 kmW Kamanjab( 19°49'S, 14°10'E),  2.iv.1977, J. Boomker— 1 ♂( paratype) ( BMNH);  Khorixas( 20°22'S, 14°58'E),  iii.2004, O. Aschenborn— 3 ♀♀, 2 ♂♂( 5 paratypes) (CMD); Orupembe( 18°10'S, 12°34'E),  14.iv.2005, W. Schawaller— 1 ♀, 2 ♂♂( 3 paratypes) ( SMNS);  Palmwag Lodge( 19°53'10''S, 13°56'13''E),  18.iv.2005, W. Schawaller— 1 ♀, 2 ♂♂( 3 paratypes) ( SMNS);  Purros( Hoaruzsib Valley) ( 18°46'S, 12°57'E),  15–16.iv.2005, W. Schawaller— 1 ♀( paratype) ( SMNS);  Khorixas District, Ae-ams/ Khoanibconfluence ( 19°14'44''S, 13°20'34''E),  28.xii.1999, Mann, Marais, & Newman— 1 ♀, 1 ♂( paratypes) ( OUMNH);  Hoanib rivernear Dubis( 19°13'36''S, 13°22'E),  28.xii.1999, Mann& Marais— 1 ♂( paratype) ( OUMNH);  Palm708, adjacent to spring ( 19°52'6''S, 14°1'13''E),  4.i.2000, Mann& Marais— 1 ♂( paratype) ( OUMNH);  Spaarwater711, near water hole ( 20°3'24''S, 14°2'55''E),  5.i.2000, Mann& Marais— 3 ♀♀, 1 ♂( paratypes) ( OUMNH);  Opuwo District, Lower Hoanib( 19°23'7''S, 13°4'11''E),  30.xii.1999, Mann& Marais— 1 ♀, 1 ♂( paratypes) ( OUMNH);  OMUSATI, Narawanduroad, Etosha National Park( 18°51'4''S, 15°32'55''E),  24–25.xii.1999, Mann& Marais— 1 ♀( paratype) ( OUMNH);  OSHIKOTO, Mushara area, sandy road ( 18°36'24''S, 16°53'23''E),  21.xii.1999, Mann& Marais— 1 ♂( paratype) ( OUMNH);  Mushara Waterhole( 18°35'19''S, 16°55'11''E),  23.xii.1999, Mann& Marais— 1 ♂( paratype) ( OUMNH);  OTJOZONDJUPA, Epupa Falls,  660 m( 17°0'3''S, 13°14'36''E),  11–12.iv.2005, W. Schawaller— 3 ♀♀, 4 ♂♂( 7 paratypes) ( SMNS); SOUTH AFRICA:   EASTERNCAPE, Willowmore( 33°18'S, 23°29'E),  ii.1916, Dr. Brauns— 1 ♀, 1 ♂( paratypes) ( NMPC);   NORTHERNCAPE, Farm Klipkoppies,  957 m( 28°52'16.93''S, 21°18'33.98''E),  12–14.iii.2004,  A.L.V. Davis & C. Deschodt— 11 ♀♀, 12 ♂♂( holotype, allotype, 21 paratypes) ( BDGC, CMNC, FGIC, OUMNH, PMOC); Groblershoop, Farm Brulpan( 28°53'56''S, 21°46'49''E),  23–26.iii.2001,  A. Davis— 2 ♂♂( 2 paratypes) ( BMNH); Keograbie Farm(1),  1038 m( 29°4'33.6''S, 21°46'5.63''E),  6–8.iii.2004,  A.L.V. Davis & C. Deschodt— 8 ♀♀, 7 ♂♂( 15 paratypes) ( FGIC, PMOC); Rooiplato( 31°26'S, 19°24'E),  14.i.1980, [anonymous]—1 ♂ (paratype) (BMNH);  Upington( 28°27'S, 21°15'E),  7.xii.1933, L. Ogilvie— 1 ♀( paratype) ( BMNH).   Etymology.  Namaquensis(from Namaqualand) a Latin adjective pertaining to the distribution of this species.  Natural history. Davis & Scholtz (2004), collected this species at the Brulpan Farm near Groblershoop, in Bushmanland Nama Karoo, in the south-west arid late summer rainfall region of  NorthernCape. Specimens were collected in cow dung baited traps and the species is recorded in this work as a soil generalist. The current data indicates a Namibian savanna woodlands and a Nama karoo centered distribution. Some specimens collected in cow and elephant dung. Asingle specimen collected under old elephant dung. This species is attracted to mercury vapor and ultraviolet light traps. 1455384277 [151,1278,1373,1398] South Africa 957 -28.87137 Northern Cape 1 21.30944 69 70 1 Northern Cape holotype 1455384287 [199,1206,1506,1531] Angola Material 70 71 175 75 100 BENGUELA 1455384279 IRSNB Angola -12.583333 Benguela 1293 13.416667 70 71 2 1 1 Benguela paratype 1455384286 1972-01-31 1972-02-02 1972-01-31 Ameib Farm Namibia -21.783333 Karibib 1262 15.633333 70 71 1 ERONGO paratype 1455384283 [666,1317,1578,1603] BMNH Namibia Southern 70 71 1 1 Southern paratype 1455384291 1978-02-10 ZMUC Namibia -23.566668 Gobabeb 1255 15.033334 70 71 1 1 Southern paratype 1455384295 1972-01-30 1972-01-31 1972-01-30 N Usakos Namibia -22.0 River 1262 15.6 70 71 1 River paratype 1455384292 1972-01-22 1972-01-23 1972-01-22 BMNH Kuiseb Canyon Namibia -23.3 Southern 1256 15.75 70 71 2 1 1 Southern paratype 1455384273 [428,1118,1686,1711] BMNH Namibia Southern 70 71 2 1 1 Southern paratype 1455384275 2010-02-16 2010-02-17 2010-02-16 SMNS Farm Lovedale, N Namibia 1600 -25.539999 Tiras Mountains 21 16.39 70 71 12 1 5 HARDAP paratype 1455384284 1972-01-19 1972-01-20 1972-01-19 BMNH Namibia -24.183332 Maltahoe District 1252 15.983334 Sesriem Farm 70 71 1 1 Southern paratype 1455384294 [397,1432,1794,1819] BMNH Namibia -24.816668 Voigtgrund 1249 17.433332 70 71 6 1 2 Southern paratype 1455384297 [151,1432,1830,1855] 2000-01-16 OUMNH Namibia Malthahohe district -24.049444 Mann 21 16.153889 Zais 70 71 1 1 Southern paratype 1455384280 2000-01-17 OUMNH Namibia Rehoboth district -24.075556 Mann 21 16.4725 Kambes 70 71 8 1 3 Southern paratype 1455384281 [426,1431,1902,1927] 1972-01-17 1972-01-18 1972-01-17 Barby Farm & W Helmeringhausen Namibia -25.883333 KARAS 1244 16.816668 70 71 1 KARAS paratype 1455384276 [151,935,1938,1963] BMNH Namibia Southern 70 71 54 11 16 Southern paratype 1455384290 SMF S. Hardt Namibia -26.566668 Keetmannshoop 1241 18.133333 70 71 10 3 2 Southern paratype 1455384293 1972-01-10 1972-01-12 1972-01-10 BMNH Namibia 71 72 -27.383333 Grunau 1237 18.466667 Noachabeb 70 71 16 3 5 Southern paratype 1455384282 1972-01-08 1972-01-10 1972-01-08 Onseepkans Namibia -28.733334 RIVER 1230 19.3 71 72 1 RIVER paratype 1455384278 BMNH Namibia Southern 71 72 1 1 Southern paratype 1455384288 [270,977,223,248] 1972-01-14 1972-01-17 1972-01-14 E Aus Namibia -26.633333 Plateau 1241 16.566668 71 72 1 Plateau paratype 1455384289 BMNH Namibia Southern 71 72 2 1 1 Southern paratype 1455384285 [404,1282,260,285] 1934-02-16 BMNH Namibia -26.816668 Seeheim 1240 17.783333 71 72 1 1 Southern paratype 1455384311 1999-12-19 OUMNH Namibia Mayana Lodge -17.865278 Mann 21 19.93 Rundu District 71 72 26 6 7 KAVANGO paratype 1455384300 2002-03-23 OUMNH Namibia Mile -18.320557 Sovo area 21 19.257778 Agricultutal Station 71 72 2 1 1 KAVANGO paratype 1455384305 2010-02-03 2010-02-06 2010-02-03 SMNS Namibia 1800 -23.37 Spreetshoogte 21 16.14 Farm Namibgrens 71 72 6 1 2 KHOMAS paratype 1455384308 1972-01-22 BMNH Namibia -23.883333 Solitaire 1254 16.0 71 72 20 2 8 Southern paratype 1455384304 SMF Namibia -22.570002 Windhoek 21 17.083612 71 72 4 2 Southern paratype 1455384318 1977-04-02 BMNH Namibia -19.816668 Kamanjab 1270 14.166667 71 72 1 1 KUNENE paratype 1455384314 2004-03 2005-04-14 2004-03 SMNS Namibia -18.166666 Orupembe 1276 12.566667 Khorixas 71 72 16 4 4 KUNENE paratype 1455384312 [151,1351,583,608] 2005-04-18 SMNS Namibia -19.88611 Palmwag Lodge 21 13.936945 71 72 6 1 2 KUNENE paratype 1455384313 2005-04-15 2005-04-16 2005-04-15 SMNS Namibia -18.766666 Hoaruzsib Valley 1274 12.95 Purros 71 72 1 1 KUNENE paratype 1455384299 1999-12-28 OUMNH Namibia Khorixas District -19.245556 Khoanib 21 13.342777 Ae-ams 71 72 2 1 1 KUNENE paratype 1455384321 1999-12-28 OUMNH Namibia Hoanib river -19.226667 Mann 873 13.366667 Dubis 71 72 1 1 KUNENE paratype 1455384315 2000-01-04 OUMNH Namibia -19.868334 Mann 21 14.020278 Palm 71 72 1 1 KUNENE paratype 1455384303 2000-01-05 OUMNH Namibia -20.056665 Mann 21 14.048612 Spaarwater 71 72 4 3 1 KUNENE paratype 1455384316 1999-12-30 OUMNH Namibia Opuwo District -19.385277 Mann 21 13.069722 Lower Hoanib 71 72 2 1 1 KUNENE paratype 1455384306 1999-12-24 1999-12-25 1999-12-24 OUMNH Namibia Narawandu -18.851112 Mann 21 15.548612 Etosha National Park 71 72 1 1 OMUSATI paratype 1455384307 1999-12-21 OUMNH Namibia -18.606667 Mann 21 16.889723 Mushara area 71 72 1 1 OSHIKOTO paratype 1455384317 1999-12-23 OUMNH Namibia -18.588612 Mann 21 16.919722 Mushara Waterhole 71 72 1 1 OSHIKOTO paratype 1455384310 2005-04-11 2005-04-12 2005-04-11 SMNS Namibia 660 -17.000834 Epupa Falls 21 13.243334 71 72 14 3 4 OTJOZONDJUPA paratype 1455384322 1916-02 NMPC Namibia -33.3 Willowmore 1205 23.483334 71 72 2 1 1 EASTERN CAPE paratype 1455384301 [265,1348,1052,1077] 2004-03-12 2004-03-14 2004-03-12 Farm Klipkoppies Namibia 957 -28.87137 NORTHERN CAPE 1 21.309439 71 72 1 NORTHERN CAPE paratype 1455384309 2001-03-23 2001-03-26 2001-03-23 BDGC, CMNC, FGIC, OUMNH, PMOC Groblershoop & Farm Brulpan Namibia -28.89889 A 20 21.780277 71 72 44 11 12 A holotype 1455384319 2004-03-06 2004-03-08 2004-03-06 BMNH Keograbie Farm Namibia 1038 -29.076 A 1 21.76823 71 72 4 2 A holotype 1455384320 1980-01-14 FGIC, PMOC Rooiplato Namibia -31.433332 A 1216 19.4 71 72 30 8 7 A holotype 1455384302 [268,1153,1231,1256] 1933-12-07 BMNH Namibia -28.45 Upington 1232 21.25 71 72 1 1 A paratype