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Digitonthophagus ulcerosus G��nier, new species http://zoobank.org/urn:lsid:zoobank.org:act: CEBB 9AB5-F6F4-4BDF-A6C1-037C15 BBBD 24 (Figs. 21, 51���52, 69, 85, 115���116, 133, 177���179; Map 16) Type locality. Ouarsangueli [=Warsangeli Sultanate], Somalia. Diagnosis. Male cephalic horns with coarse granules internally at apex (Fig. 52); males with pronotal anterior angles surface deeply concave, simply round anteriorly (Fig. 51); FLP sclerite ventral portion with a single carinae (Fig. 179); SA sclerite apex interrupted much before apex of apical portion of FLP sclerite (Figs. 177���178). Description. Holotype ♂ (Fig. 21). Measurements. Length 13.5 mm, width 8.0 mm. Head (Figs. 51���52). Anterior clypeal edge sinuous on median seventh in dorsal view; clypeofrontal carina broadly arcuate and interrupted at gena; vertex lacking median tubercle, surface with fine and minute punctures; fine, simple punctures separated by one to three diameters. Horns rather long, arcuate and slightly divergent in frontal view apical third more abruptly curved internally, parallel sided on basal two-thirds, and tapering into a point on apical third; posterointernal edge unmodified, internal surface with numerous coarse granules; genal edge upturned and simply arcuate, almost in line with clypeal edge. Pronotum (Fig. 69). Surface with rather small, granulate punctures restricted to anterior half medially, with smaller scattered simple or slightly scabrous punctures on posterior half of disc; punctures scattered and obsolete posterolaterally, with minute punctures throughout. Anteromedian tubercle well developed, produced into small acute denticle on each side of midline; median longitudinal sulcus deep; surface behind the eyes concave, surface of anterior angles with a deeply concave, anterior angles unmodified; anterior half of lateral edge arcuate in dorsal view and slightly sinuous in lateral view; posterior angles slightly lobate in dorsal view. Anterior hypomeral ridge nearly straight posteriorly and strongly arcuate on anterior third, anterior hypomeral depression surface slightly darker in color medially. Elytra (Fig. 21). Intervals 2 and 4 with odd, fine granules on apical declivity only. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus (Fig. 133). Parameres with dorsal and ventral edges diverging toward apex in lateral view. Internal sac sclerites (Figs. 177���179). Axial sclerite strongly sclerotized, abruptly bent ventrally on apical third. Subaxial sclerite, extending straight and interrupted in line with right lateral fold apical portion, with villi on apical third. Frontolateral peripheral sclerite basoventral apophysis moderately developed; a single medioventral carina present, carina distinctly folded and angulate in ventral view; right lateral fold produced into a large everted and open apically conical process with sinuous apical edge located behind oblique right lateral edge of apical lobe; left lateral lobe well sclerotized; subapicodorsal lobe membranous, narrow, not reaching anterior edge, apex set medially in dorsal view; apical lobe truncate apically and approximately in line with the axis frontolateral peripheral sclerite, ventral portion produced into and additional fold separating two concavities, ventral portion of additional lobe more-or-less parallel to apex of ventral portion of apical lobe, apical villi regular in shape; subapicoventral lobe small, not reaching apical edge of apical lobe, oriented ventrally and in line with ventrally folded left edge of apical lobe. Variation. Measurements (96 ♂♂, 92 ♀♀). Length: male 9.0��� 13.5 mm (11.7 �� 0.9 mm), female 8.0���13.0 mm (11.1 �� 1.0 mm). Female allotype. Cephalic outline in dorsal view as in Fig. 85; vertex with a broadly arcuate, transverse carina, dorsal edge broadly arcuate in frontal view, lateral portion gradually sloping down posteriorly; anterior pronotal tubercles well developed, external lateral edges parallel sided in dorsal view, anterolateral surface simply convex, anterosuperior edge arcuate in dorsal view (Fig. 115), lateral portion of anterosuperior edge slightly upturned (Fig. 116). Protibia short, with external teeth more robust. Primary type data. Holotype ♂ (MNHN): [MUSEUM PARIS / SOMALI / OUARSANGUELI / REVOIL 1881] green card; [1648 / 81] blue disc verso; [WORLD / SCARAB. / DATABASE / WSD00029929] barcode label; [HOLOTYPE ♂ / Digitonthophagus / ulcerosus n.sp. / des. F. G��nier, 2016] red card. MAP 16. Distribution of Digitonthophagus ulcerosus. Material examined (134 ♂♂, 111 ♀♀), distribution (Map 16): ETHIOPIA: Fourri, [no date], [anonymous] ��� 1 ♀ (paratype) (MNHN); Hora Oitu, [no date], [anonymous]���2 ♀♀ (2 paratypes) (MNHN); DIRE DAWA, Dire Dawa (9��36'N, 41��51'E), ix.1935, H. Ulenhuth��� 1 ♂ (paratype) (BMNH); Dire Dawa (9��36'N, 41��51'E), 1919, A. Marchand��� 4 ♀♀, 5 ♂♂ (9 paratypes) (FGIC, MNHN); Dire Dawa (9��36'N, 41��51'E), iv, [anonymous] ��� 3 ♀♀ (3 paratypes) (IRSNB); Dire Dawa (9��36'N, 41��51'E), [no date], [anonymous] ���1 ♀, 5 ♂♂ (6 paratypes) (IRSNB); Dire Dawa (9��36'N, 41��51'E), 1912, G. Kristensen��� 2 ♂♂ (2 paratypes) (MNHN); Dire Dawa (9��36'N, 41��51'E), 1936, [anonymous] ��� 1 ♂ (paratype) (MNHN); Dire Dawa (9��36'N, 41��51'E), iii, [anonymous] ��� 1 ♂ (paratype) (MNHN); Dire Dawa (9��36'N, 41��51'E), 2.x.1903, [anonymous] ��� 1 ♂ (paratype) (MNHN); HARARI, Harrar (9��19'N, 42��7'E), [no date], [anonymous] ���6 ♀♀, 8 ♂♂ (14 paratypes) (MNHN); Harrar (9��19'N, 42��7'E), 1910, [anonymous]���4 ♀♀ (4 paratypes) (MNHN); Harrar (9��19'N, 42��7'E), 1911, [anonymous]���1 ♂ (paratype) (MNHN); OROMIA, 35 km S Yabello (4��38'N, 38��14'E), 10.v.2013, I. Martinu��� 1 ♂ (paratype) (JFJC); 76 km S Agere Maryam (5��1'N, 38��13'E), 8.v.2013, I. Martinu��� 2 ♂♂ (2 paratypes) (JFJC); Bulbulla (7��43'N, 38��39'E), [no date], [anonymous]��� 9 ♀♀, 9 ♂♂ (18 paratypes) (FGIC, MNHN); Environs de Kibre Mengis, 1800���2000 m (5��53'N, 38��59'E), 12���15.xi.1973, P.C. Rougeot��� 1 ♂ (paratype) (MNHN); Lake K���ok���a [= Lake Koka; Harairobisee; Garairobisee] (8��21'N, 39��7'E), ii.1931, Huyn��� 15 ♀♀, 19 ♂♂ (34 paratypes) (FGIC, NMPC); Lake K���ok���a [= Lake Koka; Harairobi-see; Garairobisee] (8��21'N, 39��7'E), [no date], A. Hepp��� 2 ♀♀ (2 paratypes) (SMF); Maleka, 15 km W Kibre Mengist (5��58'N, 38��54'E), 14���16.iv.2002, Josso, Juhel & Legrand��� 1 ♂ (paratype) (JFJC); Yabelo (4��53'N, 38��5'E), iv.1994, Werner��� 1 ♀, 1 ♂ (paratypes) (CMD); SOUTHERN NATIONS, NATIONALITIES, AND PEOPLE���S REGION, Maraquo (8��15'N, 38��0'E), viii.1914, [anonymous] ���4 ♀♀, 10 ♂♂ (14 paratypes) (FGIC, IRSNB); Maraquo (8��15'N, 38��0'E), [no date], [anonymous]���19 ♀♀, 21 ♂♂ (40 paratypes) (IRSNB, MNHN); KENYA: Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 18.x.1973, T.J. Kingston��� 2 ♀♀, 4 ♂♂ (6 paratypes) (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 19.xi.1973, T.J. Kingston��� 1 ♂ (paratype) (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 21.xi.1973, T.J. Kingston��� 1 ♂ (paratype) (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 29.xi.1973, T.J. Kingston��� 1 ♂ (paratype) (OUMNH); ISIOLO, 10 km N Isiolo (0��27'14''N, 37��36'36''E), 16.xii.1990, B.D. Gill��� 2 ♀♀, 3 ♂♂ (5 paratypes) (BDGC); Buffalo Spring National Reserve, 900 m (0��31'N, 37��35'E), 15���16.xii.1990, B.D. Gill��� 1 ♀, 1 ♂ (paratypes) (BDGC); KITUI, S of Mwingi, Kavati Road. (0��56'S, 38��3'E), 27���28.xi.1999, P. Smerz��� 1 ♂ (paratype) (JFJC); LAIKIPIA, Karama [Hunting Camp], Tigania (0��12'25''N, 36��54'15''E), 23.x.1982, R. Mourglia��� 7 ♀♀, 6 ♂♂ (13 paratypes) (FETC); MARSABIT, near Gof Bongole Crater, Marsabit National Park & Reserve (2��12'N, 37��56'E), 18.v.1975, Davis & Dewhurst��� 1 ♀, 1 ♂ (paratypes) (BMNH); RIFT VALLEY, Samburu National Reserve (1��13'N, 36��57'E), 4.xii.1989, Moreno-Mestre��� 2 ♀♀, 2 ♂♂ (4 paratypes) (PMOC); SAMBURU, approximately 35 km N Wamba, Mattews Range, Rift Valley, 1300���1400 m (1��10.707'N, 37��18.962'E), 7���12.xii.2002, C. Hauser, D. Bartsch, & A. Zahm���1 ♀ (paratype) (SMNS); Uaso Nyiro (0��35'S, 37��32'E), [no date], Machulka��� 1 ♀, 2 ♂♂ (3 paratypes) (NMPC); TAITA TAVETA, Entrance Gate Tsavo National Park, Voi (3��21'47''S, 38��35'46''E), 26.xii.1990, B.D. Gill��� 1 ♂ (paratype) (BDGC); Maktau, Pori, 1050 m (3��24'S, 38��7'E), iii.1912, Alluaud & Jeannel (St. 62)��� 1 ♀ (paratype) (MNHN); Mbuyuni, Pori, 1110 m (3��24'S, 37��54'E), iii.1912, Alluaud & Jeannel (St. 63)��� 1 ♀, 1 ♂ (paratypes) (MNHN); Mwatate, 25 km SW Voi (3��30'S, 38��22'E), 26.xii.1990, B.D. Gill��� 5 ♀♀, 3 ♂♂ (8 paratypes) (BDGC); Taita (3��22'S, 38��22'E), [no date], [anonymous]��� 3 ♀♀, 2 ♂♂ (5 paratypes) (BMNH, MNHN); Taita Hills (3��25'S, 38��20'E), 12.xii.1989, Moreno-Mestre��� 2 ♀♀, 3 ♂♂ (5 paratypes) (PMOC); Taita Hills Game Lodge (3��30'42''S, 38��15'5''E), 26.xii.1990, B.D. Gill��� 1 ♀ (paratype) (BDGC); Taveta, 750 m (3��24'S, 37��41'E), iii.1912, Alluaud & Jeannel (St. 65)��� 1 ♀ (paratype) (MNHN); Tsavo, 50 km N Voi (3��0'S, 38��27'E), 25.xii.1990, B.D. Gill��� 1 ♂ (paratype) (BDGC); Voi, Tsavo East National Park (3��23'S, 38��34'E), 31.xii.1990, B.D. Gill��� 4 ♀♀, 2 ♂♂ (6 paratypes) (BDGC); SOMALIA: [unspecified locality], [no date], [anonymous]��� 2 ♂♂ (2 paratypes) (MNHN); [unspecified locality], [no date], R��voil���3 ♂♂ (3 paratypes) (MNHN); SANAAG, Ouarsangueli [=Warsangeli Sultanate] (11��0'N, 48��0'E), 1881, R��voil��� 1 ♀, 2 ♂♂ (holotype, allotype, 1 paratype) (MNHN); TANZANIA: KILIMANDJARO, Rivi��re Himo, zone inf��rieure, versant sudouest, Kilimandjaro, 1000 m (3��20'41''S, 37��31'56''E), iii.1912, Alluaud & Jeannel (St. 66)��� 2 ♀♀ (2 paratypes) (MNHN); Same (4��4'S, 37��45'E), v.1962, [anonymous]��� 1 ♀ (paratype) (CNC); Same (4��4'S, 37��45'E), 12��� 17.v.1999, O. Bužga��� 1 ♂ (paratype) (PMOC); RUKWA, Near Kala, Tanganyika (8��8'S, 30��58'E), [no date], J.P. Lacroix��� 1 ♀ (paratype) (MNHN). Etymology. Ulcerosus is a Latin adjective pertaining to the deeply concave surface of the male anterior pronotal angles of this species. Natural history. No habitat data are known. Some specimens were collected in buffalo and elephant dung., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 68-70, DOI: 10.5281/zenodo.439444

Digitonthophagus gazella species group 7. Digitonthophagus biflagellatus Génier, new species Distribution: Burundi, D.R. Congo, Kenya, Rwanda, Tanzania, Uganda, Zambia 8. Digitonthophagus gazella (Fabricius, 1787), conservation of name in prevailing usage Scarabaeus dorcas Olivier, 1789, proposed new synonymy Distribution: Angola, Botswana. Democratic Republic of the Congo, Kenya, Malawi, Mozambique, Namibia, South Africa, Swaziland, Tanzania, Uganda, Zambia, Zimbabwe; Introduced worldwide in tropical, subtropical, and southern temperate pastures (including the Comoros Islands, Mayotte, and Madagascar) 9. Digitonthophagus petilus Génier, new species Distribution: Ethiopia, Kenya, South Sudan, Sudan, Uganda 10. Digitonthophagus viridicollis Génier, new species Distribution: Angola, Congo, Democratic Republic of the Congo, Namibia

Digitonthophagus biflagellatus G��nier, new species http://zoobank.org/urn:lsid:zoobank.org:act:C93B1EB9-87CE-4E82-9017-F6D5D946 DDFB (Figs. 13, 35���36, 61, 77, 99���100, 125, 153���155; Map 6) Type locality. Lake Ndutu area (02��59���S 35��01���E), Serengeti National Park, Shinyanga, Tanzania. Diagnosis. Axial sclerite as long as or longer than subaxial sclerite (Fig. 153). Description. Holotype ♂ (Fig. 13). Measurements. Length 11.0 mm, width 6.5 mm. Head (Figs. 35���36). 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. Horns rather short, slightly divergent in frontal view, gradually tapering from base to apex; posterointernal edge unmodified basally, apicointernal surface lacking granules; genal edge upturned and distinctly angulate on anterior third, forming a broad angle with clypeal edge. Pronotum (Fig. 61). Surface with granulate punctures restricted to anterior half medially, with distinct scattered umbilicate punctures on posterior half of disc; punctures weaker but distinct posterolaterally, with more-or-less distinct, minute punctures throughout. Anteromedian tubercle atrophied, simply round in lateral view, median longitudinal sulcus well defined; surface behind the eyes with a simple, round depression, surface of anterior angles convex; anterior half of lateral edge arcuate in dorsal and lateral view; posterior angles simply arcuate in dorsal view. Anterior hypomeral ridge arcuate, anterior hypomeral depression surface slightly darker in color medially. Elytra (Fig. 13). Intervals 2 and 4 lacking fine granules from base to apex. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus (Fig. 125). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites (Figs. 153���155). Axial sclerite weekly sclerotized, longer than subaxial sclerite, and almost straight. Subaxial sclerite, large basally, and abruptly reduced, extending straight and interrupted before apex of frontolateral peripheral sclerite apical portion, with fine and scattered villi on apical 10th. Frontolateral peripheral sclerite basoventral apophysis moderately developed; lacking medioventral carinae; right lateral fold produced into a rather large everted and open apically conical process, apical edge deeply emarginate externally and open internally, appearing tong shaped; left lateral lobe absent; subapicodorsal lobe membranous, not reaching anterior edge, apex set on right side in dorsal view; apical lobe round apically, narrow and directed obliquely on left side, apical villi regular in shape; subapicoventral lobe large, round and interrupted in line with apical lobe apically. Variation. Measurements (94 ♂♂, 69 ♀♀). Length: male 7.5���11.5 mm (9.8 �� 0.9 mm), female 8.0���12.0 mm (9.8 �� 0.7 mm). Female allotype. Cephalic outline in dorsal view as in Fig. 77; vertex with a slightly sinuate, transverse carina, dorsal edge broadly arcuate in frontal view, lateral portion gradually sloping down posteriorly; anterior pronotal tubercles moderately developed, external lateral edges widening toward apex, anterolateral surface simply convex, anterosuperior edge slightly arcuate in dorsal view (Fig. 99), lateral portion of anterosuperior edge distinctly upturned (Fig. 100). Protibia short, with external teeth more robust. Primary type data. Holotype ♂ (OUMNH): [18 /05 /91 / L. Ndutu area, / Serengeti National Park / Tanzania, 35��1���E 2��59���S / dung-baited pitfall trap / R. Foster O.U.M. 2-1993] partly handwritten; [WORLD / SCARAB. / DATABASE / WSD00030055] barcode label; HOLOTYPE ♂ / Digitonthophagus / biflagellatus n.sp. / des. F. G��nier, 2016] red card. Material examined (135 ♂♂, 82 ♀♀), distribution (Map 8): BURUNDI: [unspecified locality], [no date], R.P. Giraudin��� 1 ♀, 4 ♂♂ (5 paratypes) (MNHN); BURURI, Bururi (3��54'S, 29��31'E), [no date], R.P. Giraudin��� 1 ♀, 1 ♂ (paratypes) (MNHN); Muyinga, Colline Kobero (2��38'59''S, 30��24'48''E), 10.ix.1940, H.-J. Br��do��� 1 ♂ (paratype) (IRSNB); RUTANA, 03��45���S 30��10���E, 2000 m (3��45'S, 30��10'E), iii.1968, R.P. Giraudin��� 1 ♀, 1 ♂ (paratypes) (OMOC); Mpinga (3��45'S, 30��10'E), [no date], R.P. Giraudin��� 1 ♂ (paratype) (MNHN); DEMOCRATIC REPUBLIC OF THE CONGO: NORTH-KIVU, Parc National Rwindi [= SE Parc National Virunga] (1��23'S, 29��28'E), 26.iii.1979, [anonymous]��� 1 ♂ (paratype) (PHWC); Rutshuru (1��11'S, 29��27'E), 1927, G. Babault��� 2 ♀♀, 1 ♂ (paratypes) (MNHN); SOUTH-KIVU, Bukavu (2��30'S, 28��52'E), 30.i.1952, C. Verbeke���3 ♂♂ (3 paratypes) (IRSNB); KENYA: HOMA BAY, Baie de Kavirondo, Lac Victoria (0��31'S, 34��29'E), ix���x.1903, C. Alluaud��� 2 ♂♂ (2 paratypes) (MNHN); Baie de Kavirondo, Lac Victoria (0��31'S, 34��29'E), xii.1911, Alluaud & Jeannel��� 1 ♂ (paratype) (MNHN); Ile de Lusinga [= Rusinga], Victoria-Nyanza N. - E. (0��24'S, 34��11'E), x.1903, C. Alluaud��� 1 ♂ (paratype) (IRSNB); KAJIADO, 60 km SW Nairobi, 1200 m (1��45'S, 36��21'E), 23.xii.1990, B.D. Gill��� 2 ♀♀, 1 ♂ (paratypes) (BDGC); KAKAMEGA, Ilala, 14 mi. E Mumias, 1372 m (0��17'N, 34��41'E), 18���21.vi.1911, S. A. Neave��� 1 ♂ (paratype) (BMNH); KISUMU, Kisumu (0��6'S, 34��46'E), [no date], P.S. Corbet��� 2 ♀♀, 4 ♂♂ (6 paratypes) (BMNH); MURANG���A, Fort Hall [= Muranga Town] (0��43'S, 37��9'E), i.1912, Alluaud & Jeannel��� 1 ♂ (paratype) (MNHN); NAIROBI, Nairobi (1��17'S, 36��49'E), 24.iv.1957, [anonymous]��� 2 ♀♀, 2 ♂♂ (4 paratypes) (BMNH); Nairobi National Park (1��22'S, 36��51'E), iv.1993, W. Hoek��� 1 ♂ (paratype) (CEMT); NANDI, Kamililo, vi.1903, F.J. Jackson��� 1 ♂ (paratype) (BMNH); NAROK, Lemek Valey (1��6'S, 35��23'E), ii.1913, G.B. ��� 6 ♀♀, 8 ♂♂ (14 paratypes) (FGIC, MNHN); TAITA TAVETA, Bura (3��2'S, 38��19'E), i���iv.1904, C. Alluaud��� 5 ♂♂ (5 paratypes) (IRSNB, MNHN); Mwatate (3��30'S, 38��22'E), i��� iv.1904, C. Alluaud��� 1 ♀, 3 ♂♂ (4 paratypes) (IRSNB, MNHN); Mwatate, 25 km SW Voi (3��30'S, 38��22'E), 26.xii.1990, B.D. Gill��� 2 ♀♀, 2 ♂♂ (4 paratypes) (BDGC); VIHIGA, For��t de Kakamegoes [= Kakamega], Environs de Kaimosi, Mont-Elgon, 2000 m (0��10'N, 34��52'E), 1913, G. Babault��� 6 ♂♂ (6 paratypes) (FGIC, MNHN); RWANDA: EASTERN, Ranch Mpanga (2��3'13''S, 30��47'0''E), iii.1987, [anonymous]��� 2 ♀♀, 3 ♂♂ (5 paratypes) (PSCW); NORTHERN, Bugarula District, [unspecified locality] (1��33'S, 29��42'E), 21.viii.1941, H.-J. Br��do��� 2 ♀♀, 3 ♂♂ (5 paratypes) (IRSNB); WESTERN, Rwaza (1��42'S, 29��16'E), 1927, G. Babault��� 2 ♂♂ (2 paratypes) (MNHN); TANZANIA: Lake Rukwa, iv.1945, H.-J. Br��do��� 1 ♀, 2 ♂♂ (3 paratypes) (FGIC, IRSNB); Lake Rukwa, xii.1948, H.-J. Br��do��� 1 ♀, 3 ♂♂ (4 paratypes) (IRSNB); ARUSHA, Arusha environs (3��22'S, 36��41'E), 3���4.iv.1997, M. Kuboň��� 5 ♀♀, 1 ♂ (paratypes) (PMOC); Karatu Ndutu Safari Lodge, Serengeti (3��1'S, 35��0'E), 15.xi.1990, R. Foster��� 1 ♂ (paratype) (BDGC); Mumba Hills (3��30'S, 35��3'E), 8.iii.1938, Dr. S.G. Kohl- Larsen��� 1 ♀ (paratype) (SMF); Mumba Hills (3��30'S, 35��3'E), 23.iii.1938, Dr. S.G. Kohl-Larsen��� 1 ♂ (paratype) (SMF); Mumba Hills (3��30'S, 35��3'E), 24.iii.1938, Dr. S.G. Kohl-Larsen��� 1 ♂ (paratype) (SMF); Orekeryan, Mount Longido (2��43'47''S, 36��43'26''E), 8.vi���9.viii.2012, Smith, Takano & Garner��� 1 ♂ (paratype) (BMNH); DODOMA, 40 km N Dodoma, 1100 m (5��54'S, 35��45'E), 15.xii.2007, F. Kantner��� 5 ♀♀, 3 ♂♂ (8 paratypes) (SMNS); 70 km N Dodoma, 1330 m (5��40'S, 35��48'E), 17.xii.2006, F. Kantner��� 1 ♂ (paratype) (SMNS); Lulanguru, 17 mi. W Tabora, 1148 m (5��6'S, 32��38'E), x���xii.1917, G.D. H. Carpenter��� 1 ♀, 1 ♂ (paratypes) (BMNH); Tosamaganga, near Iringa (6��40'S, 36��19'E), [no date], W. Ruddock��� 1 ♂ (paratype) (BMNH); IRINGA, Magema, Ulembwe Village, 2048 m (9��18.389'S, 34��37.213'E), 7.i.2011, P. Darge��� 1 ♂ (paratype) (MHNL); KILIMANDJARO, Same (4��4'S, 37��45'E), v.1962, [anonymous]��� 2 ♂♂ (2 paratypes) (CNC); Zone inf��rieure, Kilimandjaro, i���iv.1904, C. Alluaud��� 1 ♀, 3 ♂♂ (4 paratypes) (MNHN); MANYARA, 100 km NNE Singida, 1650 m (4��31'S, 35��21'E), 20.xii.2006, F. Kantner��� 5 ♀♀, 2 ♂♂ (7 paratypes) (SMNS); Giting, Mount Hanang, 1964 m (4��24'14''S, 35��24'56''E), 22���26.xi.2011, R. Smith & H. Takano��� 8 ♀♀, 2 ♂♂ (10 paratypes) (BMNH); MARA, Shirati (1��9'S, 34��2'E), 1909, [anonymous]��� 5 ♀♀, 1 ♂ (paratypes) (SMF); MOROGORO, Llonga (9��2'S, 36��51'E), 21.x.1966, Robertson��� 1 ♀, 2 ♂♂ (3 paratypes) (FETC); Majawanga, 1400 m (6��5'S, 36��50'E), 14���18.iii.2004, S. Tind Nielsen��� 1 ♀, 3 ♂♂ (4 paratypes) (ZMUC); RUKWA, Tumba (7��7'S, 31��10'E), 15.xii.1950, H.-J. Br��do��� 1 ♀, 2 ♂♂ (3 paratypes) (FGIC, IRSNB); SHINYANGA, Lake Ndutu area, Serengeti National Park (2��59'S, 35��1'E), 15.xi.1990, R. Foster��� 3 ♀♀, 2 ♂♂ (allotype, 4 paratypes) (OUMNH); Lake Ndutu area, Serengeti National Park (2��59'S, 35��1'E), 17.xii.1990, R. Foster��� 3 ♀♀, 1 ♂ (paratypes) (OUMNH); Lake Ndutu area, Serengeti National Park (2��59'S, 35��1'E), 18.xii.1990, R. Foster��� 1 ♂ (paratype) (OUMNH); Lake Ndutu area, Serengeti National Park (2��59'S, 35��1'E), 22.xii.1990, R. Foster��� 1 ♂ (paratype) (OUMNH); Lake Ndutu area, Serengeti National Park (2��59'S, 35��1'E), 26.xii.1990, R. Foster��� 1 ♂ (paratype) (OUMNH); Lake Ndutu area, Serengeti National Park (2��59'S, 35��1'E), 18.v.1991, R. Foster��� 2 ♂♂ (holotype, 1 paratype) (OUMNH); Old Shinyanga (3��34'S, 33��25'E), 1.xii.1935, E. Burtt��� 1 ♂ (paratype) (BMNH); Old Shinyanga (3��34'S, 33��25'E), 26.i.1936, E. Burtt��� 1 ♂ (paratype) (BMNH); Old Shinyanga (3��34'S, 33��25'E), 10.iv.1936, E. Burtt��� 1 ♀ (paratype) (BMNH); TABORA, Tabore (5��2'S, 32��49'E), 1886, R��voil��� 1 ♂ (paratype) (MNHN); UGANDA: CENTRAL, Entebbe (0��4'N, 32��28'E), ii���vii.1972, H. Falke��� 10 ♀♀, 9 ♂♂ (19 paratypes) (CNC, FGIC); Entebbe (0��4'N, 32��28'E), ii.1972, H. Falke��� 2 ♂♂ (2 paratypes) (CNC); Entebbe (0��4'N, 32��28'E), 22.vi.1912, C. A. Wiggins��� 1 ♀ (paratype) (OUMNH); Entebbe (0��4'N, 32��28'E), 15.viii.1912, C. A. Wiggins��� 1 ♂ (paratype) (OUMNH); Entebbe (0��4'N, 32��28'E), 9.ii.1914, [anonymous]��� 1 ♂ (paratype) (BMNH); Entebbe (0��4'N, 32��28'E), 11.ii.1914, [anonymous]��� 1 ♀ (paratype) (BMNH); Entebbe (0��4'N, 32��28'E), 20���28.v.1914, C.C. Gowdey��� 2 ♂♂ (2 paratypes) (BMNH); Kampala (0��21'N, 32��35'E), 1���10.i.1918, C.C. Gowdey��� 1 ♀, 5 ♂♂ (6 paratypes) (BMNH); Kampala (0��21'N, 32��35'E), 11.ii.1918, [anonymous]��� 1 ♀, 1 ♂ (paratypes) (BMNH); R��gion du Bouganda (0��27'S, 31��55'E), 1906, [anonymous]��� 1 ♂ (paratype) (IRSNB); [unspecified locality] (0��15'N, 32��15'E), i���ii.1909, C. Alluaud��� 1 ♀, 3 ♂♂ (4 paratypes) (MNHN); EASTERN, Jinja, 1151 m (0��26'N, 33��12'E), 14.ii.1906, C. A. Wiggins��� 1 ♂ (paratype) (OUMNH); WESTERN, Kafu River near Hoima, Kampala Road (1��18'N, 31��27'E), 29���31.xii.1911, S. A. Neave��� 1 ♂ (paratype) (BMNH); Province d���Unyoro, R��gion de l���Albert-Nyanza (1��17'N, 31��10'E), ii.1909, C. Alluaud��� 1 ♂ (paratype) (MNHN); ZAMBIA: NORTHERN PROVINCE, Mbala (8��50'S, 31��28'E), 15.xii.1943, H.-J. Br��do��� 2 ♂♂ (2 paratypes) (IRSNB). MAP 8. Distribution of Digitonthophagus biflagellatus. Etymology. Biflagellatus is a Latin adjective referring to the configuration of the axial and subaxial sclerites of the internal sac. Natural history. The single specimen with habitat data was collected in woodland in Eastern Province, Uganda. The few specimens with collecting data indicate light and dung traps., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 37-39, DOI: 10.5281/zenodo.439444

Digitonthophagus gazella (Fabricius, 1787), conservation of name in prevailing usage (Figs. 14, 37���38, 62, 78, 101���102, 126, 156���158, 185, 188, 190; Map 9) Scarabaeus Gazella Fabricius, 1787b: 377 (original description) Scarabaeus Dorcas Olivier, 1789: 121 (original description) proposed new synonymy Scarabaeus Gazella ��� Gmelin 1790: 1537 (diagnosis) Scarabaeus gazella ��� Fabricius 1792: 56 (diagnosis) Copris Dorcas ��� Fabricius 1798: 31 (diagnosis) Scarabaeus Dorcas ��� Illiger 1800: 228 (monograph) Scarabaeus Gazella ��� Illiger 1800: 228 (mentioned as synonym) Copris Dorcas ��� Fabricius 1801: 44 (diagnosis) Copris Gazella ��� Fabricius 1801: 47 (diagnosis) Copris Gazella ��� Illiger 1804: 149 (mentioned as synonym) Copris Gazella ��� Sch��nherr 1806: 48 (catalog) Onthophagus Gazella ��� Dejean 1821: 53 (catalog) Onthophagus Gazella ��� Sturm 1826: 178 (catalog) Onthophagus bonasus ��� Latreille 1827: 282 (diagnosis, distribution, comment) Onthophagus Gazella ��� LePeletier & Audinet-Serville 1828: 354 (comment taxonomy) Onthopagus [sic] Gazella ��� Klug 1833: 163 (diagnosis) Scarabaeus Dorcas ��� Klug 1833: 163 (synonymy) Onthophagus Gazella ��� Klug 1835: 33 (distribution) Onthophagus Gazella ��� Dejean 1836: 157 (catalog) Onthophagus Gazella ��� Hope 1837: 33 (catalog) Ontophagus gazella ��� Reiche 1840: 243 (comment) Ontophagus gazella ��� Laporte 1840: 85 (diagnosis, distribution) Onthophagus Gazella ��� Sturm 1843: 107 (catalog) Ontophagus gazella ��� Erichson 1843: 208 (mention) Onthophagus Gazella ��� Bertoloni 1849: 415 (diagnosis, distribution) Onthophagus gazella ��� F��hraeus 1857: 275 (redescription) Onthophagus gazella ��� Motschulsky 1860: 153 (mention) Onthophagus gazella ��� Klug 1862: 232 (comment, distribution) Onthophagus gazella ��� Harold 1867: 49 (comment) Onthophagus gazella ��� Harold 1869: 1029 (catalog) [Scarabaeus] dorcas ��� Harold 1869: 1029 (mentioned as synonym) Copris gazella ��� Gerstaecker 1871: 50 (synonymy) Onthophagus gazella ��� Harold 1871: 119 (comment) Scarabaeus Dorcas ��� Gerstaecker 1873: 130 (mentioned as synonym) Scarabaeus gazella ��� Gerstaecker 1873: 130 (mentioned as synonym) Onthophagus gazella ��� Wallengren 1881: 21 (comment) Onthophagus gazella ��� Lansberge 1882: XXII (mention) Onthophagus gazella ��� Reitter 1892: 183 (identification key, distribution) Onthophagus gazella ��� P��ringuey 1892: 29 (distribution) Onthophagus gazella ��� Fairmaire 1893: 10 (distribution) Onthophagus gazella ��� Brancsik 1893: 223 (distribution) Onthophagus gazella ��� Reitter 1893: 62 (identification key, distribution) Onthophagus gazella ��� Kolbe 1897a: 150 (distribution) Onthophagus gazella ��� Kolbe 1897b: 83 (distribution) Onthophagus gazella ��� Gahan & Arrow 1900: 24 (distribution) Onthophagus (O.) gazella ��� d���Orbigny 1900a: 218 (monograph) Onthophagus gazella ��� d���Orbigny 1900b: 299 (comment, distribution) Onthophagus gazella ��� P��ringuey 1901: 181 (monograph) Onthophagus (O.) gazella ��� d���Orbigny 1904: 304 (distribution) Onthophagus (O.) gazella ��� d���Orbigny 1905a: 494 (diagnosis, distribution) Onthophagus gazella ��� d���Orbigny 1905b: 445 (distribution) Onthophagus gazella ��� Kolbe 1908: 128 (distribution) Onthophagus gazella ��� d���Orbigny 1909: 525 (distribution) Onthophagus gazella ��� d���Orbigny 1911: 20 (distribution) Onthophagus (O.) gazella ��� d���Orbigny 1913a: 249 (monograph) Onthophagus (O.) gazella ��� d���Orbigny 1913b: 96 (comment, distribution) Onthophagus (O.) gazella ��� Kolbe 1914: 297 (distribution) Onthophagus (O.) gazella ��� d���Orbigny 1914: 654 (catalog) Onthophagus (O.) gazella ��� d���Orbigny 1916: 11 (distribution) Onthophagus gazella ��� Boucomont 1923: 96 (distribution) Onthophagus gazella ��� Gillet & Boucomont 1927: 138 (catalog) [Scarabaeus] dorcas ��� Gillet & Boucomont 1927: 138 (mentioned as synonym) Scarabaeus gazella ��� Arrow 1931: 230 (mentioned as synonym) Onthophagus gazella ��� Boucomont 1934: 26 (mentioned as synonym) Onthophagus gazella ��� Paulian 1936: 132 (identification key) Onthophagus gazella ��� Boucomont 1937: 131 (distribution) Onthophagus gazella ��� Balthasar 1941: 110 (distribution) Onthophagus gazella ��� M��ller 1942: 96 (comment, distribution) Onthophagus gazella ��� Gomes Alves 1944: 3 (distribution) Onthophagus gazella ��� Paulian 1948: 151 (mentioned as synonym) Onthophagus (O.) gazella ��� Gomes Alves 1956: 94 (distribution) Onthophagus gazella ��� Frey 1956: 654 (distribution) Onthophagus gazella ��� Ferreira 1958: 512 (faunistic) Onthophagus gazella ��� Frey 1958: 73 (distribution) Onthophagus gazella ��� Ferreira 1959: 84 (distribution) Onthophagus gazella ��� Frey 1961: 90 (distribution, ecology) Onthophagus gazella ��� Petrovitz 1961: 102 (distribution) Onthophagus gazella ��� Ferreira 1962: 153 (faunistic) Onthophagus (Digitonthophagus) gazella ��� Balthasar 1963: 365 (monograph) [Scarabaeus] dorcas ��� Balthasar 1963: 365 (mentioned as synonym) Onthophagus gazella ��� Ferreira 1967: 1169 (catalog) Onthophagus dorcas ��� Ferreira 1968: 593 (mentioned as synonym) Onthophagus gazella ��� Ferreira 1968: 593 (monograph) Onthophagus (Digitonthophagus) gazella ��� Balthasar 1969: 68 (faunistic) Onthophagus gazella ��� Bornemissza 1970: 31 (natural history) Onthophagus (O.) gazella ��� Endr��di 1971: 294 (faunistic) Onthophagus (Digitonthophagus) gazella ��� Cambefort 1972: 246 (distribution, ecology) Onthophagus (Digitonthophagus) gazella ��� Ferreira 1972: 780 (catalog) Onthophagus dorcas ��� Ferreira 1972: 780 (mentioned as synonym) Onthophagus gazella ��� Balthasar 1972: 22 (faunistic) Onthophagus gazella ��� Endr��di 1973a: 125 (faunistic) Onthophagus (O.) gazella ��� Endr��di 1973b: 198 (distribution) Onthophagus gazella ��� Endr��di 1974: 3 (faunistic) Onthophagus gazella ��� Blume & Aga 1975: 735 (natural history) Onthophagus gazella ��� Endr��di 1976: 156 (distribution) Onthophagus gazella ��� Alfieri 1976: 205 (faunistic) Onthophagus gazella ��� Bornemissza 1976: 7 (natural history) Onthophagus gazella ��� Blume & Aga 1978: 190 (distribution, ecology) Onthophagus gazella ��� Rougon & Rougon 1980: 381 (natural history, description of larvae) Digitonthophagus gazella ��� Zunino 1981: 414 (new combination, faunistic) Onthophagus gazella ��� Rougon & Rougon 1981: 6 (description of larvae) Onthophagus gazella ��� Rougon & Rougon 1983: 499 (natural history) Onthophagus gazella ��� Fincher et al. 1986: 2 (natural history) Onthophagus gazella ��� Paulian 1986: 107 (mentioned as synonym) Onthophagus gazella ��� Gutierrez et al 1988: 39 (natural history) Digitonthophagus gazella ��� Zunino & Halffter 1988: 17 (distribution) Digitonthophagus gazella ��� Kohlmann 1994: 35 (distribution) Onthophagus gazella ��� Ripa et al. 1995: 438 (distribution) Digitonthophagus gazella ��� Barbero et al. 1998: 242 (distribution) Onthophagus (Digitonthophagus) gazella ��� Kim & Lumaret, 1989: 336 (description of larvae) Digitonthophagus gazella ��� Deloya 2000: 125 (faunistic) Onthophagus gazella ��� MacRae & Penn 2001: 49 (distribution) Digitonthophagus gazella ��� Noriega 2002: 213 (distribution) Digitonthophagus gazella ���Barbero & Palestrini 2003: 316 (distribution) Digitonthophagus gazella ���Barbero et al. 2003: 333 (phylogeny) Digitonthophagus gazella ��� Cambefort & Bordat 2003: 560 (distribution, ecology) Digitonthophagus gazella ��� Noriega et al. 2006: 379 (distribution) Digitonthophagus gazella ��� Moretto & Bordat 2007: 122 (distribution, ecology) Digitonthophagus gazella ��� Monaghan et al. 2007: 678 (phylogeny) Digitonthophagus gazella ��� Ivie & Philips 2008: 10 (distribution) Digitonthophagus gazella ���Cabral-de-Mello et al. 2008: 1244 (karyotype) Onthophagus gazella ��� Wirta et al. 2008: 1079 (phylogeny) Digitonthophagus gazella ��� Noriega et al. 2010: 451 (distribution) Digitonthophagus gazella ��� Noriega et al. 2011: 35 (distribution) Digitonthophagus gazella ��� Tarasov & Solodovnikov 2011: 9 (phylogeny) Digitonthophagus gazella ��� Whipple et al. 2012: 45 (DNA analysis) Digitonthophagus gazella ��� Mlambo et al. 2015: 318 (phylogeny) Digitonthophagus gazella ��� Roggero et al. 2016a: [2] (phylogeny) Digitonthophagus gazella ��� Breeschoten et al. 2016: 90 (phylogeny) Digitonthophagus gazella ��� G��nier & Davis 2017: 497 (comment taxonomy, distribution) Scarabaeus dorcas ��� G��nier & Davis 2017: 498 (comment taxonomy) Type locality. Unspecified. Diagnosis. Pronotum rather glossy (Fig. 62); FLP sclerite subapicodorsal lobe short anteriorly (Figs. 156��� 157); right lateral fold produced into a longitudinal spoon-shape process with apical portion open ventrally, ventral edge bordered with a conspicuous brush of villi (Fig. 185 A), lacking well-sclerotized projection on apicoventral edge (Fig. 185 B); axial sclerite shorter than subaxial sclerite, acute apically (Fig. 156); subaxial sclerite moderately long and robust (Fig. 153). Description. Male specimen from Central Province, Zambia (Fig. 14). Measurements. Length 12.5 mm, width 7.0 mm. Head (Figs. 37���38). 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. Horns rather short, slightly divergent in frontal view, parallel sided on basal four-fifths and abruptly tapering externally and narrower on apical sixth, posterointernal edge produced into low, angular projection basally, apicointernal surface lacking granules, with few scabrous punctures; genal edge slightly upturned and feebly angulate on anterior third, forming a broad angle with clypeal edge. Pronotum (Fig. 62). Surface with granulate punctures extending on posterior half, with distinct umbilicate punctures on posterior half of disc; punctures smaller posterolaterally, with distinct, minute punctures throughout. Anteromedian tubercle atrophied, simply round in lateral view, median longitudinal sulcus narrow and moderately deep; surface behind the eyes with a simple, round, shallow depression, surface of anterior angles flat; anterior half of lateral edge arcuate in dorsal and lateral view; posterior angles slightly upturned and simply arcuate in dorsal view; anterior hypomeral ridge arcuate anteriorly, anterior hypomeral depression surface slightly darker in color medially. Elytra (Fig. 14). Interval 2 with few scattered 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. 126). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites (Figs. 156���158). Axial sclerite broadly arcuate, with apex acute. Subaxial sclerite, gradually tapering from base to apex, slightly angled on apical third, extending straight approximately in line with apex of frontolateral peripheral sclerite apical portion, with a thick brush of villi on apical fifth. Frontolateral peripheral sclerite basoventral apophysis well developed; lacking medioventral carinae; right lateral fold produced into longitudinal spoon-shape process with apical portion open ventrally, ventral edge bordered with a brush of 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 irregular in shape; subapicoventral lobe round and interrupted in line with apical lobe, with few villi along apical edge only. Variation. Measurements (237 ♂♂, 169 ♀♀). Length: male 7.0���13.0 mm (10.6 �� 1.1), female 7.5���13.0 mm (10.5 �� 1.0 mm). Female specimen. Cephalic outline in dorsal view as in Fig. 78; vertex with a slightly sinuate transverse carina, dorsal edge slightly sinuous in frontal view, lateral portion abruptly sloping down; anterior pronotal tubercles well developed, external lateral edges slightly widening toward apex in dorsal view, anterolateral surface obliquely concave, anterosuperior edge slightly arcuate in dorsal view (Fig. 101), lateral portion of anterosuperior edge slightly upturned (Fig. 102). Protibia short, with external teeth more robust. Primary type data. Scarabaeus gazella Fabricius : Syntype (ZMUK): [] small green square; [ZMUC / 00508509]; [TYPE] red card; [WORLD / SCARAB. / DATABASE / WSD00029479] barcode label; [SYNTYPE ♂ / Scarabaeus / gazella / Fabricius, 1787 / vid. F. G��nier, 2016] red card; [Guinea / Isert / Mus. T. Lund / Copris / gazella / ♂ / F.] handwritten [in error] see Nomenclature and taxonomy below for more details. Note. The first label, a small green square without inscription, confirms this specimen as a Fabrician type (Zimsen 1964). The last label was added after Fabricius���s time and is erroneously cited as African. See nomenclature and taxonomy below section for more details. Scarabaeus dorcas Olivier : Neotype ♂ (MNHN) present designation: [Madagascar / Diego-Suarez 7 / Ch. Alluaud, 1893] partly handwritten; [MUSEUM PARIS / MADAGASCAR / COLL. CH. ALLUAUD 1904] green card; [WORLD / SCARAB. / DATABASE / WSD00030025] barcode label; [NEOTYPE / Scarabaeus / dorcas / Olivier, 1789 / des. F. G��nier, 2016] red card. Note. This neotype designation is made with the express purpose of clarifying the taxonomic status and the type locality of Scarabaeus dorcas Olivier. The specimen designated agrees in morphology to the description presented above. The original type material used for Olivier���s description is lost or destroyed (Olivier Montreuil, MNHN, personal communication). The neotype has been selected to match Olivier���s description and type locality, in addition to possessing the best diagnostic character for the species. The neotype is deposited in the collection of the Museum national d���Histoire naturelle de Paris along with the remains of Olivier���s collection. MAP 9. Distribution of Digitonthophagus gazella (native range in continental Africa only). Material examined (715 ♂♂, 407 ♀♀), distribution (Map 9): ANGOLA: CUANDO CUBANGO, Menongue (14��39'S, 17��41'E), iii.1914, Mission Rohan-Chabot��� 1 ♂ (MNHN); HUILA, 10 km S Caconda (13��50'S, 15��6'E), 5.xi.2011, G. Werner��� 2 ♀♀, 2 ♂♂ (FGIC, GWPC); 10 km S Cacula (14��35'S, 14��3'E), 4.xi.2011, G. Werner��� 1 ♂ (GWPC); 30 km W Quipungo (14��48.02'S, 14��15.32'E), 26.xi.2013, G. Werner��� 3 ♀♀, 2 ♂♂ (FGIC, GWPC); 33 km S Cacula (14��41'S, 13��54'E), 13.xi.2011, G. Werner��� 1 ♀, 1 ♂ (GWPC); Capelongo pr��s Luceque (14��39'S, 15��4'E), xii.1914, Mission Rohan-Chabot��� 1 ♂ (MNHN); Capelongo-Dongo (14��31'S, 16��1'E), 1914, Mission Rohan-Chabot��� 1 ♀ (MNHN); Capelongo-Dongo (14��31'S, 16��1'E), xii.1914, Mission Rohan-Chabot��� 2 ♂♂ (MNHN); AUSTRALIA: NEW SOUTH WALES, Camp Blackman, Warrumbungle National Park (31��16'37''S, 148��59'51''E), 5���6.xii.1988, H. & A. Howden��� 1 ♀, 1 ♂ (CMNC); Gilgandra (31��43'S, 148��40'E), 12.iv.1988, H. & A. Howden��� 1 ♂ (CMNC); Narrabri (30��20'S, 149��47'E), 5.xii.1977, G.G. Burns��� 2 ♀♀ (CMNC); NORTHERN TERRITORY, CSIRO McMillans Road, Darwin (12��24'42''S, 130��55'19''E), 1���10.xii.1993, S. & J. Peck (93-85)��� 1 ♀, 1 ♂ (CMNC); CSIRO McMillans Road, Darwin (12��24'42''S, 130��55'19''E), 5.i.1994, S. & J. Peck (93-166)��� 1 ♀, 1 ♂ (CMNC); CSIRO McMillans Road, Darwin (12��24'42''S, 130��55'19''E), 9.i.1994, S. & J. Peck (93-167)��� 1 ♂ (CMNC); Elsey National Park, Mataranka (14��58'S, 133��11'E), 9.xii.1993, S. & J. Peck (93- 115)��� 1 ♀, 1 ♂ (CMNC); Elsey National Park, Mataranka (14��58'S, 133��11'E), 9.xii.1993, S. & J. Peck (93- 116)��� 1 ♀, 1 ♂ (CMNC); Gungaree Rainforest, South Alligator River, Kakadu National Park (12��40'46''S, 132��29'9''E), 22.xii.1993, S. & J. Peck (93-133)��� 1 ♂ (CMNC); Magela Creek (12��40'S, 133��1'E), 4.xi.1989, C.W. & L.B. O'Brien��� 1 ♂ (CMNC); Tenat Creek (19��39'S, 134��11'E), 22.ix.1988, H. & A. Howden��� 1 ♀, 1 ♂ (CMNC); QUEENSLAND, 12 mi. S Ayr (19��44'S, 147��28'E), 26.ii.1975, H. & A. Howden��� 1 ♂ (CMNC); 16 km W Ravenshoe (17��39'S, 145��21'E), 2.i.1975, M.S. & B.J. Moulds��� 1 ♂ (CMNC); 4���8 km S Atherton (17��20'S, 145��30'E), 26.xii.1988, H. & A. Howden��� 1 ♂ (CMNC); 40 km N Lynd (18��36'S, 144��43'E), ii.1993, H. Demarz&mdas, Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 39-52, DOI: 10.5281/zenodo.439444, {"references": ["Fabricius, J. C. (1787 b) Mantissa insectorum sistens species nuper detectas adiectis synonymis, observationibus, descriptionibus, emendationibus. Tom. II. Christ. Gottl. Proft., Copenhagen, Denmark, [2] + 382 pp.", "Olivier, A. G. (1789) Entomologie, ou histoire naturelle des insectes, avec leurs caracteres generiques et specifiques, leur descriptions, leur synonymie, et leur figure enluminee. Coleopteres. Tome premier. Beaudoin, Paris, France, xx + 497 pp. https: // doi. org / 10.5962 / bhl. title. 61905", "Gmelin, J. F. 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Digitonthophagus namaquensis species group 16. Digitonthophagus namaquensis Génier, new species Distribution: Angola, Namibia, South Africa

Digitonthophagus falciger species group 6. Digitonthophagus falciger G��nier, new species Distribution: Benin, Burkina Faso, Cameroon, Chad, Cote d���Ivoire, Gambia, Ghana, Guinea, Guinea-Bissau, Mauritania, Niger, Nigeria, Senegal, Sierra Leone,? Uganda, Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on page 74, DOI: 10.5281/zenodo.439444

Digitonthophagus gazella species group Diagnosis. Subapicoventral lobe of FLP sclerite round apically (Figs. 155, 158, 161, 164); right lateral fold of FLP sclerite apical portion more-or-less spoon shape (Figs. 153, 156, 159, 162)., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on page 37, DOI: 10.5281/zenodo.439444

Digitonthophagus catta species group Diagnosis. Pronotal punctures scattered and small on posterior half (Figs. 58, 93); surface behind anterior pronotal angles concave; male with two erected cephalic horns (Figs. 30, 32); females with anterior pronotal tubercles well developed and deeply obliquely concave (Figs. 94, 96); subapicodorsal lobe of FLP sclerite enlarged and sclerotized (Figs. 144, 147); FLP left surface with a conspicuous apophysis (Figs. 144, 147)., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on page 24, DOI: 10.5281/zenodo.439444

Digitonthophagus catta (Fabricius, 1787) (Figs. 10, 29���30, 58, 74, 93���94, 122, 144���146; Map 5) Scarabaeus Catta Fabricius, 1787a: 12 (original description) Scarabaeus Catta ��� Olivier 1789: 125 (redescription) Scarabaeus Catta ��� Gmelin 1790: 1534 (diagnosis) Scarabaeus Catta ��� Fabricius 1792: 44 (diagnosis) Scarabaeus Catta ��� Illiger 1800: 232 (monograph) Copris Catta ��� Fabricius 1801: 35 (diagnosis) Copris Catta ��� Illiger 1804: 149 (comment taxonomy) Copris Catta ��� Sch��nherr 1806: 49 (mentioned as synonym) Onthophagus Catta ��� Dejean 1821: 53 (mentioned as synonym) Copris Catta ��� LePeletier & Audinet-Serville 1828: 354 (comment taxonomy) Copris Catta ��� Klug 1833: 163 (mentioned as synonym) [Onthophagus] Catta ��� Dejean 1836: 157 (mentioned as synonym) Onthophagus Catta ��� Hope 1837: 32 (catalog) Ontophagus Catta ��� Reiche 1840: 243 (comment taxonomy) [Ontophagus] Catta ��� Laporte 1840: 85 (mentioned as synonym) Onthophagus Catta ��� Sturm 1843: 107 (mentioned as synonym) Onthophagus catta ��� Motschulsky 1860: 154 (comment taxonomy) Onthophagus catta ��� Harold 1869: 1029 (mentioned as synonym) Onthophagus catta ��� Gerstaecker 1871: 50 (distribution) Onthophagus catta ��� Gerstaecker 1873: 130 (monograph) Onthophagus catta ��� Quedenfeldt 1884: 272 (distribution) Onthophagus catta ��� Kolbe 1887: 246 (distribution) Onthophagus catta ���Kolbe 1897: 150 (mentioned as synonym) Onthophagus catta ��� P��ringuey 1901: 181 (mentioned as synonym) Onthophagus catta ��� Gillet & Boucomont 1927: 138 (mentioned as synonym) Onthophagus catta ��� Arrow 1931: 230 (monograph) Onthophagus catta ��� Boucomont 1934: 26 (mentioned as valid species) Onthophagus catta ��� Boucomont 1935: 284 (distribution) Onthophagus catta ��� Paulian 1937: 12 (distribution) Onthophagus catta ��� Paulian 1948: 151 (distribution) Onthophagus catta ��� Janssens 1953: 1549 (distribution) Onthophagus catta ��� Frey 1958: 73 (mentioned as synonym) [Scarabaeus] catta ��� Balthasar 1963: 365 (mentioned as synonym) Onthophagus catta ��� Ferreira 1967: 1169 (mentioned as synonym) Onthophagus catta ��� Ferreira 1968: 593 (mentioned as synonym) Onthophagus catta ��� Ferreira 1972: 780 (mentioned as synonym) Onthophagus catta ��� Paulian 1986: 107 (catalog) Onthophagus gazella ��� Veenakumari & Veeresh 1996: 252 (natural history) Onthophagus catta ��� Gaikwad & Bhawane 2015: 1538 (natural history) Digitonthophagus catta ��� G��nier & Davis 2017: 497 (comment taxonomy, distribution) Diagnosis. Species distributed from Pakistan eastward; left lateral apophysis of FLP sclerite moderately developed (Fig. 144). Description. Alloreferent ♂ (Fig. 10): Measurements. Length 11.5 mm, width 6.5 mm. Head (Figs. 29���30). 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. Horns moderately long, arcuate in frontal view, parallel sided on basal four-fifths; distinctly tapering externally before apex and narrower on apical sixth; posterointernal edge unmodified basally; apicointernal surface lacking granules. Genal edge upturned and distinctly angulate on anterior third, forming a broad angle with clypeal edge. Pronotum (Fig. 58). Surface with granulate punctures restricted to anterior half medially, with smaller, irregularlyscattered, simple punctures on posterior half of disc; punctures lacking posterolaterally, with more-or-less distinct, minute punctures throughout. Anteromedian tubercle atrophied, simply round in lateral view; median longitudinal sulcus well defined; surface behind the eyes slightly concave, surface of anterior angles concave, anterior angles unmodified; anterior half of lateral edge arcuate in dorsal and slightly sinuous anteriorly in lateral view; posterior angles simply arcuate in dorsal view. Anterior hypomeral ridge broadly arcuate, anterior hypomeral depression surface slightly darker in color medially. Elytra (Fig. 10). Interval 2 with few fine granules on apical declivity, interval 4 lacking granules from base to apex. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus (Fig. 122). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites (Figs. 144���146). Axial sclerite moderately sclerotized, shorter than subaxial sclerite, almost straight. Subaxial sclerite, larger basally and gradually tapering, extending straight and interrupted at apex of frontolateral peripheral sclerite apical portion, with villi on apical sixth. Frontolateral peripheral sclerite basoventral apophysis moderately developed, angular in lateral view; with a single acute medioventral carinae; right lateral fold produced into a uneven bifurcate process, lateral edge with few indentations; left lateral apophysis present, moderately developed; left lateral lobe absent; subapicodorsal lobe well sclerotized, reaching anterior edge, partly fused to apical lobe, with villi similar to apical lobe, right edge round before apex; apical lobe round apically, longitudinally folded and partly fused to subapicodorsal lobe, apical villi regular in shape; subapicoventral lobe atrophied, oriented ventrally. Variation. Measurements (107 ♂♂, 106 ♀♀). Length: male 7.0���13.0 mm (10.6 �� 1.0 mm), female 7.5���12.5 mm (10.2 �� 1.0 mm). Female specimen. Cephalic outline in dorsal view similar to Fig. 74; vertex with a broadly arcuate transverse carina, dorsal edge broadly arcuate in frontal view, lateral portion gradually sloping down posteriorly; anterior pronotal tubercles well developed, external lateral edges approximately parallel in dorsal view (Fig. 93), anterolateral surface obliquely concave, anterosuperior edge slightly arcuate in dorsal view, lateral portion of anterosuperior edge slightly upturned (Fig. 94). Protibia short, with external teeth more robust. Primary type data. Lectotype ♀ (ZMUK) present designation: [WORLD / SCARAB. / DATABASE / WSD00032221] barcode label; [LECTOTYPE ♀ / Scarabaeus / catta / Fabricius, 1787 / des. F. G��nier, 2016] red card. Material examined (188 ♂♂, 199 ♀♀, 855 sex no recorded), distribution (Map 5): AFGHANISTAN: KHYBER PAKHTUNKHWA, Hazara, Abbottabad (39��10'N, 73��13'E), vi.2002, [anonymous] ���2 ♀♀, 2 ♂♂ (CEMT); BANGLADESH: RANGPUR, Dinajpur (25��45'N, 88��43'E), iv.1970, Barbe��� 1 ♂ (CMNC); INDIA: Coromandel Coast, [no date], Zschuck��� 4 ♀♀ (lectotype, 3 paralectotypes) (ZMUC); ANDAMAN AND NICOBAR ISLANDS, [unspecified locality] (12��30'N, 92��45'E), [no date], Roepstorff��� 1 ♂ (ZMUC); GOA, Calangute (15��32'30''N, 73��45'45''E), 15.xii.1976, [anonymous]��� 4 ♀♀, 3 ♂♂ (OMOC); [unspecified locality] (15��22'N, 74��4'E), 22���23.x.1986, J.L. & A. Nicolas��� 1 ♀, 1 ♂ (CMD); JHARKHAND, Ranchi (23��20'N, 85��19'E), 1970, Dr. Soběslavsk����� 1 ♀ (CVM); Ranchi (23��20'N, 85��19'E), [no date], R.P. Bretaudeau��� 1 ♀, 1 ♂ (MNHN); KARNATAKA, 15 km W Gadag, 670 m (15��25'48''N, 75��28'39''E), 13.viii.1984, B. Gill��� 1 ♀ (BDGC); 20 km SW Hubli, 650 m (15��12'3''N, 74��59'34''E), 11.vii.1984, B. Gill��� 3 ♀♀, 1 ♂ (BDGC); Bangalore, 970 m (12��58'N, 77��34'E), 17���19.vii.1984, B. Gill��� 7 ♀♀, 2 ♂♂ (BDGC); Chikkangalur, pr��s de Bangalore [=Chickmagaluru] (13��18'53''N, 74��46'29''E), vii���ix.1899, R.P. Tabourel��� 1 ♀ (MNHN); Hampi (15��20'6''N, 76��27'36''E), 18.x.1986, J.L. Nicolas��� 2 ♀♀, 2 ♂♂ (CMD); Harihar, 750 m (14��30'30''N, 75��48'30''E), 15.vii.1984, B. Gill��� 2 ♀♀, 5 ♂♂ (BDGC); Hubli, 670 m (15��21'42''N, 75��5'6''E), 7���14.vii.1984, B. Gill��� 5 ♀♀, 8 ♂♂ (BDGC); Sindhanur (15��46'N, 76��45'E), 21.xii.1989, C. Mahalingappa��� 1 ♀, 2 ♂♂ (OUMNH); Mysore District, Mysore (12��18'N, 76��39'E), [no date], [anonymous]��� 1 ♀, 1 ♂ (MNHN); Shimoga District, Bhadravati (13��50'24''N, 75��42'7.2''E), 13.v.1935, [anonymous]��� 1 ♀, 1 ♂ (CMNC); Shimoga (13��55'N, 75��34'E), v.1897, [anonymous]��� 1 ♀, 1 ♂ (MNHN); Shimoga (13��55'N, 75��34'E), vi.1897, [anonymous]��� 1 ♀, 1 ♂ (MNHN); Shimoga (13��55'N, 75��34'E), iv.1936, [anonymous]��� 1 ♂ (PMOC); KERALA, Chembra Peak, 1067 m (11��33'12''N, 76��5'14''E), iv.1970, T.R.S. Nathan��� 1 ♂ (CMNC); Thenmala (8��58'N, 77��4'E), v.1988, T.R.S. Nathan��� 1 ♀, 1 ♂ (CMNC); Thiruvananthapuram District, Poonmudi Range, 914 m (8��45'27''N, 77��6'54''E), v.1989, T.R.S. Nathan��� 1 ♀, 1 ♂ (CMNC); KERALA / TAMIL NADU, Anaimalai Hills, 1067 m (10��22'N, 77��7.5'E), v.1963, P.S. Nathan��� 1 ♀ (CNC); Anaimalai Hills, 1067 m (10��22'N, 77��7.5'E), v.1977, [anonymous]��� 4 ♀♀, 4 ♂♂ (CNC); Anaimalai Hills, 1067 m (10��22'N, 77��7.5'E), iv.1963, P.S. Nathan��� 2 ♂♂ (CNC); Anaimalai Hills, 1067 m (10��22'N, 77��7.5'E), x.1977, T.R.S. Nathan��� 1 ♂ (RMNH); Cinchona, Anaimalai Hills, 1067 m, iv.1964, P.S. Nathan��� 1 ♂ (CNC); MADHYA PRADESH, Jabalpur (23��10'N, 79��56'E), x.1957, Nathan��� 1 ♀, 1 ♂ (CMNC); Maihar (24��16'10''N, 80��45'24''E), x.1972, [anonymous]��� 1 ♀ (ZMUC); Pachmarhi, Satputa Hills, 1067 m (22��28'.12''N, 78��25'59.88''E), x.1970, T. Nathan��� 1 ♀, 1 ♂ (CNC); Ratlam (23��20'N, 75��2'E), [no date], [anonymous]��� 1 ♂ (MNHN); Satna (24��36'N, 80��50'E), [no date], Stloukal��� 1 ♂ (CMNC); MAHARASHTRA, Pune (18��31'N, 73��51'E), viii.1984, S. Pokorn����� 2 ♀♀, 2 ♂♂ (CVM); ODISHA, Brahmapur (19��19'N, 84��47'E), xi.1972, [anonymous]��� 1 ♂ (ZMUC); ORISSA, Berhampur (19��18'30''N, 84��47'26''E), [no date], Atkinson��� 1 ♂ (OUMNH); Balasore District, Balasore (21��30'N, 86��56'E), [no date], R.P. Gengler��� 1 ♀, 1 ♂ (MNHN); PONDICHERRY, Karaikal District, Karaikal (10��56'N, 79��50'E), viii.1991, T.R.S. Nathan��� 1 ♀ (CMNC); Karaikal District, Karaikal (10��56'N, 79��50'E), x.1991, T.R.S. Nathan��� 1 ♂ (CMNC); Karaikal District, Karaikal (10��56'N, 79��50'E), iv.1962, P.S. Nathan��� 1 ♀, 1 ♂ (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), x.1963, P.S. Nathan��� 1 ♀, 1 ♂ (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), ix.1952, P.S. Nathan��� 2 ♀♀ (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), vi.1962, P.S. Nathan��� 1 ♂ (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), vii.1963, P.S. Nathan��� 1 ♂ (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), i.1962, P.S. Nathan��� 11 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), ii.1962, P.S. Nathan��� 36 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), iii.1962, P.S. Nathan��� 34 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), vii.1962, P.S. Nathan��� 40 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), viii.1962, P.S. Nathan��� 74 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), ix.1962, P.S. Nathan��� 146 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), x.1962, P.S. Nathan��� 83 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), xi.1962, P.S. Nathan��� 39 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), xii.1962, P.S. Nathan��� 9 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), i.1963, P.S. Nathan��� 16 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), ii.1963, P.S. Nathan��� 12 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), iii.1963, P.S. Nathan��� 34 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), iv.1963, P.S. Nathan��� 12 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), vi.1963, P.S. Nathan��� 4 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), x.1966, P.S. Nathan��� 13 specimens (CNC); Karaikal District, Karaikal (10��56'N, 79��50'E), viii.1963, T.R.S. Nathan��� 2 ♀♀, 2 ♂♂ (MMUE); Karaikal District, Karaikal (10��56'N, 79��50'E), i���ii.1964, T.R.S. Nathan��� 4 ♀♀, 4 ♂♂ (MMUE); Karaikal District, Karaikal (10��56'N, 79��50'E), vii.1970, P. Susai Nathan��� 1 ♂ (RMNH); Nedungadu (10��58'N, 79��46'E), [no date], [anonymous]��� 1 ♂ (NMPC); RAJASTHAN, Barmer (25��45'N, 71��23'E), ix.1973, T.R.S. Nathan��� 1 ♀ (RMNH); Kota, 250 m (25��10'28''N, 75��50'14''E), x.1981, T.R.S. Nathan��� 2 ♀♀, 2 ♂♂ (OUMNH); Thar desert around Jaisalmer (26��55'N, 70��56'E), 2.viii.2007, I. N��dai��� 4 ♀♀, 7 ♂♂ (PMOC); TAMIL NADU, 20 km SE Sagar, 600 m (14��6.37'N, 75��8.93'E), 12.v.2005, M. Halada��� 1 ♀ (OUMNH); 20 km SW Karamadai, 450 m (11��12.6'N, 76��44.9'E), 27.iv.2005, M. Halada��� 1 ♂ (OUMNH); Cap Comorin [=Kanyakumari] (8��5'N, 77��32'E), 1888, R.P. Castets��� 16 specimens (MNHN); Coimbatore, 427 m (11��0'N, 76��57'E), xi.1969, P.S. Nathan��� 1 ♀, 3 ♂♂ (CMNC, CNC); Coimbatore, 427 m (11��0'N, 76��57'E), iv.1959, P.S. Nathan��� 1 ♀ (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), iv.1962, P.S. Nathan��� 25 specimens (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), v.1962, P.S. Nathan��� 19 specimens (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), vii.1962, P.S. Nathan��� 18 specimens (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), ix.1962, P.S. Nathan��� 14 specimens (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), x.1962, P.S. Nathan��� 3 specimens (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), xi.1962, P.S. Nathan���20 specimens (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), xii.1962, P.S. Nathan��� 5 specimens (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), i.1963, P.S. Nathan��� 6 specimens (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), iv.1963, P.S. Nathan��� 50 specimens (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), xi.1966, P.S. Nathan��� 20 specimens, 2 ♂♂ (CNC, PMOC); Coimbatore, 427 m (11��0'N, 76��57'E), viii.1963, T.R.S. Nathan��� 6 ♀♀, 6 ♂♂ (MMUE); Coimbatore, 427 m (11��0'N, 76��57'E), ix.1964, T.R.S. Nathan��� 1 ♀, 5 ♂♂ (MMUE); Coimbatore, 427 m (11��0'N, 76��57'E), x���xi.1964, T.R.S. Nathan��� 4 ♂♂ (MMUE); Coimbatore, 427 m (11��0'N, 76��57'E), xi.1967, P.S. Nathan��� 1 ♀ (PMOC); Coimbatore, 427 m (11��0'N, 76��57'E), ix.1972, T.R.S. Nathan��� 2 ♀♀, 1 ♂ (PMOC, RMNH); Coimbatore, 427 m (11��0'N, 76��57'E), x.1972, T.R.S. Nathan��� 2 ♀♀, 2 ♂♂ (PMOC); Coimbatore, 427 m (11��0'N, 76��57'E), xii.1976, T.R.S. Nathan��� 1 ♂ (RMNH); Gingee (12��15'N, 79��25'E), [no date], M. Andr����� 1 ♂ (PMOC); Madurai (9��55'N, 78��7'E), x.1975, M. Coe��� 1 ♀, 4 ♂♂ (OUMNH); Maravakkadu (10��38'N, 79��27'E), v.1962, T.R.S. Nathan��� 2 ♀♀ (CNC); NE of Madurai, 200 m (10��8.7'N, 77��46.5'E), 6.v.2005, M. Halada��� 1 ♀ (OUMNH); Palni Hills (10��12'N, 77��30'E), 1898, R.P. Castets��� 1 ♀, 1 ♂ (MNHN); Point Calimere (10��17'35''N, 79��52'6''E), 1972, T.R.S. Nathan��� 1 ♀ (RMNH); Pudukkottai (10��23'N, 78��49'E), x.1984, T.R.S. Nathan��� 1 ♀, 1 ♂ (CMNC); Singara, Nilgiri Hills (11��22'N, 76��46'E), v.1963, T.R.S. Nathan��� 2 ♀♀, 2 ♂♂ (MMUE); Thambikkottai (10��23'N, 79��27'E), vi.1962, P.S. Nathan��� 3 ♀♀ (CNC); Tirunelveli (8��44'N, 77��42'E), x.1986, [anonymous]��� 1 ♀, 1 ♂ (OMOC); Tranquebar [= Tharangambadi] (11��2'N, 79��5'E), [no date], [anonymous]��� 1 ♂ (ZMUC); Vellore (12��55'N, 79��8'E), [no date], L��venthal��� 2 ♂♂ (ZMUC); Chennai District, Madras Airport [= Chennai Airport] (12��59'38''N, 80��9'28''E), v.1962, P.S. Nathan��� 64 ♀♀, 42 ♂♂ (CNC); Chennai District, Madras Airport [= Chennai Airport] (12��59'38''N, 80��9'28''E), viii.1962, P.S. Nathan��� 19 specimens (CNC); Chennai District, Madras Airport [= Chennai Airport] (12��59'38''N, 80��9'28''E), ix.1962, P.S. Nathan��� 9 specimens (CNC); Chennai District, Madras Airport [= Chennai Airport] (12��59'38''N, 80��9'28''E), x.1962, P.S. Nathan��� 14 specimens (CNC); Coimbatore District, Coimbatore (11��0'N, 76��57'E), x.1958, P.S. Nathan��� 1 ♀ (CNC); Coimbatore District, Coimbatore (11��0'N, 76��57'E), vi.1962, P.S. Nathan��� 1 ♀ (CNC); Coimbatore District, Coimbatore (11��0'N, 76��57'E), x.1976, [anonymous]��� 1 ♀, 1 ♂ (CNC); Coimbatore District, Coimbatore (11��0'N, 76��57'E), xi.1976, [anonymous]��� 1 ♂ (CNC); Coimbatore District, Coimbatore (11��0'N, 76��57'E), iv.1976, [anonymous]��� 1 ♂ (CNC); Coimbatore District, Coimbatore (11��0'N, 76��57'E), xi.1957, P.S. Nathan��� 3 specimens (CNC); Coimbatore District, Coimbatore (11��0'N, 76��57'E), xi.1962, P.S. Nathan��� 12 specimens (CNC); Coimbatore District, Coimbatore (11��0'N, 76��57'E), xi.1965, P.S. Nathan��� 10 specimens (CNC); Marudamalai Hills (11��2'46''N, 76��51'7''E), xi.1969, P.S. Nathan��� 29 specimens, 1 ♀, 1 ♂ (CMNC, CNC); Marudamalai Hills (11��2'46''N, 76��51'7''E), x.1969, P.S. Nathan��� 2 ♀♀ (CNC); Kancheepuram District, Manapakkam (13��0'N, 80��10'E), 20.ix.2002, S. Saluk��� 1 ♂ (PMOC); UTTAR PRADESH, Babina, 300 m (25��15'N, 78��28'E), viii.1984, T.R.S. Nathan��� 1 ♀, 2 ♂♂ (CMNC); Babina, 300 m (25��15'N, 78��28'E), viii.1987, T.R.S. Nathan��� 3 ♀♀, 3 ♂&male, Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 24-28, DOI: 10.5281/zenodo.439444, {"references":["Fabricius, J. C. (1787 a) Mantissa insectorum sistens eorum species nuper detectas adiectis characteribus generici, differentiis specificis, emendationibus, observationibus. Tom. I. Christ. Gottl. Proft., Copenhagen, Denmark, xx + 348 pp. https: // doi. org / 10.5962 / bhl. title. 36471","Olivier, A. G. (1789) Entomologie, ou histoire naturelle des insectes, avec leurs caracteres generiques et specifiques, leur descriptions, leur synonymie, et leur figure enluminee. Coleopteres. Tome premier. Beaudoin, Paris, France, xx + 497 pp. https: // doi. org / 10.5962 / bhl. title. 61905","Gmelin, J. F. (1790) Caroli a Linne, systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, loci. Editio decima tertia, aucta, reformata. Tom. I, Pars IV. Classis V. Insecta. G. E. Beer, Leipzig, Germany, pp. 1517 - 2224.","Fabricius, J. C. (1792) Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species adjictis synonimis, locis, observationibus, descriptionibus. Tom. I. Christ. Gottl. Proft., Copenhagen, Denmark, xx + 330 pp.","Illiger, J. C. W. (1800) Olivier's Entomologie oder Naturgeschichte der Insekten mit ihren Gattungs- und ArtMerkmalen, ihrer Beschreibung und Synonymie. Kafer. Uebersetzt und mit Zusatzen und Anmerkungen durchgangig begleitet. Erster Theil. Karl Reichard, Braunschweig, Germany, xvi + 309 pp., 2 plates.","Fabricius, J. C. (1801) Systema Eleutheratorum secundum ordines, genera, species: adiectis synonymis, locis, observationibus, descriptionibus. Tomus I. Bibliopoli academici novi, Kiel, Germany, xxiv + 506 pp.","Illiger, K. (1804) Zusatze, Berichtigungen und Bemerkungen zu Fabricii. Systema Eleutheratorum. Tomus II. Magazin fur Insektenkunde, 3, 146 - 180.","Schonherr, C. J. (1806) Synonymia insectorum, ode: Versuch einer Synonymie aller bisher bekannten Insecten; nach Fabricii Systema Eleutheratorum geordnet, mit Berichtigungen und Anmerkungen, wie auch Beschreibungen neuer Arten und illuminirten Kupfern. Erter Band. Eleutherata oder Kafer. Erster Theil. Lethrus ... Scolytes. A. Nordstrom, Stockholm, Sweden, xxii + 293 pp., 3 plates.","Dejean, P. F. M. A. (1821) Catalogue de la collection de coleopteres de M. le Baron Dejean. Crevot, Paris, France, viii + 136 + [2] pp.","LePeletier de Saint Fargeau, A. L. M. & Audinet-Serville, J. G. de (1828) Scarabe, Scarabaeus. In: Encyclopedie methodique, ou par ordre de matieres; par une societe de gens de lettres, de savant et d'artistes; precede d'un vocabulaire universel, servans de table pour tout l'ouvrage, ornee des portraits de Mm. Diderot & d'Alembert, premiers editeurs de l'Encyclopedie. Tome dixieme. Pars 2. Mme. veuve Agasse, Paris, France, pp. 345 - 832.","Klug, J. C. F. (1833) Bericht uber eine auf Madagascar veranstaltete Sammlung von Insecten aus der Ordnung Coleoptera. Eine in der Konigl. Akademie der Wissenschaften am 29. Marz 1832 gelesene Abhandlung. Mit funf illuminirten Tafeln. Koniglichen Akademie der Wissenschaften, Berlin, Germany. 135 pp, 5 plates.","Dejean, P. F. M. A. (1836) Catalogue des coleopteres de la collection de M. le Comte Dejean. Troisieme edition, revue, corrigee et augmentee. Mequignon-Marvis Pere et Fils, Paris, France, xiv + 503 pp.","Hope, F. W. (1837) The Coleopterist's Manual, containing the lamellicorn insect of Linneus and Fabricius. Henry G. Bohn, London, United Kingdom, xiii + [2] + 15 - 121 + [4] pp., 3 plates.","Reiche, L. J. (1840) Notes sur deux especes d' Ontophages. Revue Zoologique, 1840, 243 - 244.","Laporte, F. (Comte de Castelnau) (1840) Histoire Naturelle des Insectes Coleopteres. Tome deuxieme. Histoire Naturelle des animaux articulees, crustaces, arachnides, myriapode et insectes. Tome troisieme, 2, P. Dumenil, Paris, France, 564 pp.","Sturm, J. (1843) Catalog der Kaefer-Sammlung von Jacob Sturm. Jacob Sturm, Nurenberg, Germany, xii + 386 pp., 6 plates.","Motschulsky, V. de (1860) Sur les Collections Coleopterologiques de Linne et de Fabricius (Continuation.). 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Berliner Entomologische Zeitschrift, 28, 265 - 340. https: // doi. org / 10.1002 / mmnd. 18840280207","Kolbe, H. J. (1887) Beitrage zur Zoogeographie Westafrikas nebst einem Bericht uber die wahrend der Loango-Expedition von Herrn Dr. Falkenstein bei Chinchoxo gesammelten Coleoptera. Nova Acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae Curiosorum, 50, 153 - 364.","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa (Lucanidae and Scarabaeidae). Transactions of the South African Philosophical Society, 12, 1 - 563, 9 plates.","Gillet J. J. E. & Boucomont A. (1927) Scarabaeidae: Coprinae II, Termitotroginae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus. Partes 38 et 90, 38, 90. W. Junk, Berlin, Germany, pp. 101 - 264.","Arrow, G. J. (1931) Coleoptera Lamellicornia Part III. (Coprinae). The Fauna of British India, including Ceylon and Burma. Taylor and Francis, London, United Kingdom, 428 pp.","Boucomont, A. (1934) Mission J. de Lepiney au Soudan francais 1933 - 34 (Quatrieme note). Bulletin de la Societe des Sciences Naturelles du Maroc, 14, 25 - 27.","Boucomont, A. (1935) Coleoptera. VI. Scarabaeidae Scarabaeini. In: Jeannel, R. (Ed.), Mission Scientifique de l'Omo. Tome 2 Zoologie, Fascicule 16. Paul Chevalier & Fils, Paris, France, pp. 279 - 290.","Paulian, R. (1937) Contribution a l'etude de la faune entomologique de L'Angola, Mission scientifique suisse en Angola. Coleopteres Lamellicornes Laparoscticti. Memorias e Estudos do Museu Zoologico da Universidade de Coimbra, 1 (105), 1 - 41.","Paulian, R. (1948) 12. Coleoptera, Scarabaeidae: Troginae, Geotrupinae, Dynamopinae, Hybosoridae, Coprinae, Aphodiinae. British Museum (Natural History) Expedition to South-Est Arabia 1937 - 38, Volume 1 (9 - 15). British Museum, London, United Kingdom, pp. 141 - 155.","Janssens, A. (1953) Contribution a l'etude de la zone d'inondation du Niger (mission G. Remaudiere, 1950). V - Coleopteres Scarabaeinae. Bulletin de l'Institut Francais de l'Afrique Noire, 15, 1548 - 1551.","Frey, G. (1958) Onthophagini (Coleoptera Polyphaga) (Fam. Scarabaeidae). Exploration du Parc National de l'Upemba, I. Mission G. F. de Witte en collaboration avec W. Adam, A. Janssens, L. van Meel et R. Verheyen (1946 - 1949), 54, 69 - 90.","Balthasar, V. (1963) Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region. Coleoptera: Lamellicornia. Band, 2. Coprinae (Onitini, Oniticellini, Onthophagini). Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region, 2. Verlag der Tschechoslowakischen Akademie der Wissenschaften, Prague, Czech Republic, 627 pp., 16 plates.","Ferreira, M. C. (1967) Catalogo dos Coleopteros de Angola. Revista de Entomologia de Mocambique, 8, 417 - 1317.","Ferreira, M. C. (1968) Os escarabideos de Mocambique. I. Revista de Entomologia de Mocambique, 10, 5 - 778.","Ferreira, M. C. (1972) Os escarabideos de Africa (sul do Saara). I. Revista de Entomologia de Mocambique, 11, 5 - 1088.","Paulian, R. (1986) Catalogue des coleopteres Scarabaeidae de Madagascar. Bulletin de l'Academie Malgache, 62, 89 - 111.","Veenakumari, K. & Veeresh, G. K. (1996) Some aspects of the reproductive biology of Onthophagus gazella (F) and Onthophagus rectecornutus (Lansb) (Coleoptera, Scarabaeidae). Journal of Bombay Natural History Society, 93, 252 - 256.","Gaikwad, A. R. & Bhawane, G. P. (2015) Study of nidification behavior in three dung beetle species (Scarabaeidae: Scarabaeinae) from south-western Maharashtra, India. International Journal of Science and Research, 4, 1538 - 1542.","Genier, F. & Davis, A. L. V. (2017) Digitonthophagus gazella auctorum: an unfortunate case of mistaken identity for a widely introduced species (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Zootaxa, 4221 (4), 497 - 500. https: // doi. org / 10.11646 / zootaxa. 4221.4.8","Evenhuis, N. L. (1997) Litteratura taxonomica dipterorum (1758 - 1930); being a selected list of the books and prints of Diptera taxonomy from the beginning of Linnean zoological nomenclature to the end of the year 1930; containing information on the biographies, bibliographies, types, collections, and patronymic genera of the authors listed in this work; including detailed information on publication dates, original and subsequent editions, and other ancillary data concerning the publications listed herein. Volume I: A - K. Backhuys Publishers, Leiden, Netherlands, vii + 426 pp.","Bousquet, Y. (2016) Litteratura Coleopterologica (1758 - 1900): a guide to selected books related to the taxonomy of Coleoptera with publication dates and notes. ZooKeys, 583, 1 - 776.","Zimsen, E. (1964) The Type Material of I. C. Fabricius. Munksgaard, Copenhagen, Denmark, 280 pp."]}

Digitonthophagus bonasus (Fabricius, 1775) (Figs. 7, 23���24, 55, 71, 87���88, 119, 135���137, 183; Map 2) Scarabaeus Bonasus Fabricius, 1775: 23 (original description) Scarabaeus Bonasus ��� Goeze 1777: 69 (diagnosis) Scarabaeus Bonasus ��� Fabricius 1781: 26 (redescription) Scarabaeus bonasus ��� Pallas 1781: 5 (redescription) Scarabaeus Bonasus ��� Herbst 1786: 153 (diagnosis, distribution) Scarabaeus Bonasus ��� Fabricius 1787a: 14 (diagnosis) Scarabaeus Bonasus ��� Olivier 1789: 121 (redescription) Scarabaeus Bonasus ��� Herbst 1789 2: 180 (monograph) Scarabaeus Bonasus ��� Gmelin 1790: 1541 (diagnosis) Scarabaeus Bonasus ��� Fabricius 1792: 50 (diagnosis) Scarabaeus Bonasus ��� Illiger 1800: 228 (monograph) Copris Bonasus ��� Fabricius 1801: 40 (diagnosis) Copris Bonasus ��� Illiger 1804: 149 (comment taxonomy) Copris Bonasus ��� Sch��nherr 1806: 42 (catalog) Ontophagus Bonasus ��� Sturm 1826: 177 (catalog) Onthophagus bonasus ��� Latreille 1827: 282 (misidentification) Onthophagus bonasus ��� LePeletier & Audinet-Serville 1828: 353 (identification key, comment) Onthophagus Bonasus ��� Hope 1837: 32 (catalog) Ontophagus bonasus ��� Laporte 1840: 85 (diagnosis, distribution) Onthophagus Bonasus ��� Sturm 1843: 107 (catalog) Onthophagus bonasus ��� Motschulsky 1860: 154 (comment taxonomy) Onthophagus bonasus ��� Motschulsky 1863: 457 (checklist) Onthophagus bonasus ��� Harold 1869: 1026 (catalog) Onthophagus (O.) bonasus ��� d���Orbigny 1900a: 218 (monograph) Onthophagus bonasus ��� Boucomont 1914: 241 (distribution) Onthophagus bonasus ��� Boucomont 1921: 29 (identification key, distribution) Onthophagus bonasus ��� Gillet & Boucomont 1927: 133 (catalog) Onthophagus bonasus ��� Arrow 1931: 231 (monograph) Onthophagus (O.) bonasus ��� Paulian 1945: 101 (monograph) Onthophagus bonasus ��� Petrovitz 1955: 277 (distribution) Onthophagus bonasus ��� Balthasar 1956: 435 (distribution) Onthophagus (Digitonthophagus) bonasus ��� Balthasar 1959: 464 (genotype designation) Onthophagus (Digitonthophagus) bonasus ��� Balthasar 1963: 296 (monograph) Digitonthophagus bonasus ��� Zunino 1981: 413 (new combination) Onthophagus (Matashia) bonasus ��� Masumoto 1988: 140 (distribution) Digitonthophagus bonasus ��� Zunino & Halffter 1988: 20 (comment taxonomy) Digitonthophagus bonasus ���Barbero & Palestrini 2003: 333 (phylogeny) Digitonthophagus bonasus ���Cabral-de-Mello et al. 2008: 1248 (chromosomal formula) Digitonthophagus bonasus ��� Tarasov & Solodovnikov 2011: 4 (phylogeny) Digitonthophagus bonasus ��� G��nier 2013: 2 (comment taxonomy) Digitonthophagus bonasus ��� Roggero et al. 2016a: [2] (phylogeny) Digitonthophagus bonasus ��� Roggero et al. 2016b: 2 (phylogeny) Digitonthophagus bonasus ��� Breeschoten et al. 2016: 90 (phylogeny) Type locality. Tranquebariae [= Tharangambadi, Tamil Nadu State, India]. Diagnosis. Male and female pronotal anteromedian tubercle similar, well developed, forming an angular projection in lateral view (Fig. 88); pronotal anterior median longitudinal sulcus wide and deep (Figs. 55, 87). Description. Alloreferent ♂ (Fig. 7). Measurements. Length 14.5 mm, width 9.0 mm. Head (Figs. 23���24). Anterior clypeal edge slightly sinuous on median seventh in dorsal view; clypeofrontal carina broadly arcuate and interrupted at gena; vertex with an acute median conical process higher than wide basally, surface with dense irregular granules. Horns long, obliquely divergent in frontal view, gradually tapering from base to apex, posterointernal edge unmodified basally, internal surface coarsely and densely granulated; genal edge slightly upturned and arcuate on anterior third, forming a broad angle with clypeal edge. Pronotum (Fig. 55). Surface with granules restricted to median third, lacking distinct umbilicate punctures, punctures weaker but distinct posterolaterally with distinct, minute punctures throughout. Anteromedian tubercle well developed, forming an angular projection in lateral view and forming two short blunt longitudinal carinae in dorsal view; median longitudinal sulcus wide and deep; surface behind the eyes with a small round depression; surface of anterior angles sloping laterally; anterior half of lateral edge strongly sinuous in dorsal and lateral view; posterior angles produced into a obliquely-oriented tooth in dorsal view. Anterior hypomeral ridge broadly arcuate anteriorly with anterior most portion crenulate, anterior hypomeral depression surface slightly darker in color. Elytra (Fig. 7). Intervals 2 and 4 with few fine granules on apical fourth. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus (Fig. 119). Parameres with dorsal and ventral edges diverging toward apex in lateral view. Internal sac sclerites (Figs. 135���137). Axial sclerite strongly sclerotized, crescent shaped. Subaxial sclerite extending straight approximately in line with apex right lateral fold apical portion, with fine villi. Frontolateral peripheral sclerite basoventral apophysis moderately developed; two medioventral carinae present; right lateral fold produced into a rather large everted and open apically conical process with extremely irregular apical edge; left lateral lobe absent; subapicodorsal lobe membranous, narrow, not reaching anterior edge, apex set medially in dorsal view; apical lobe round and directed obliquely on left side, apical villi regular in shape; subapicoventral lobe short not reaching apical edge of apical lobe, oriented obliquely. Variation. Measurements (111 ♂♂, 103 ♀♀). Length: male 11.0��� 16.5 mm (13.9 �� 1.3 mm), female 10.0���16.0 mm (13.4 �� 1.2 mm). Female lectotype. Cephalic outline in dorsal view similar to Fig. 71; vertex with horns wide basally and regularly tapering to apex, posterointernal edge with an acute tooth basally; anterior pronotal configuration (Figs. 87���88) similar to males. Protibia short, with external teeth more robust. A number of females from various localities in Asia east of India present much shorter cephalic horns. The horns in these females (Fig. 183) are wide and taper abruptly just before the apex, which is internally diverging in frontal view. Males collected with these females are identical to typical male D. bonasus and we consider this variation as intraspecific. Primary type data. Lectotype ♀ (ZMUK) present designation: [ZMUC / 00021685]; WORLD / SCARAB. / DATABASE / WSD00032218] barcode label; [LECTOTYPE ♀ / Scarabaeus / bonasus / Fabricius, 1775 / d��s. F. G��nier, 2016] red card. MAP 2. Distribution of Digitonthophagus bonasus. Material examined (169 ♂♂, 164 ♀♀, 377 sex not recorded), distribution (Map 2): BANGLADESH: CHITTAGONG, Chittagong Hill Tracts (22��20'N, 92��13'E), 1883, H.M. Parish��� 1 ♀, 1 ♂ (BMNH); DHAKA, Dacca [= Dhaka] (23��42'N, 90��22'30''E), 5.v.1945, D. Leston��� 1 ♀ (BMNH); CAMBODIA: KOH KONG, 20 km SE Koh Kong, Tatai river, 50���300 m (11��34'N, 103��7'E), 17.x���15.xi.2011, Z. Linek���2 ♀♀, 2 ♂♂ (FGIC); PHNOM PENH, Kompong Luong (11��50'N, 104��48'E), [no date], [anonymous]��� 1 ♀ (MNHN); SIEM REAP, Kompong Chikreng (13��5'N, 104��18'E), 1912, G. Thomas��� 1 ♂ (MNHN); INDIA: C��te de Malabar, 1900, T. Deschamps��� 4 ♀♀, 6 ♂♂ (MNHN); HIMACHAL PRADESH, Near Kullu, Pirpandzhal range, 1500 m (31��57'30''N, 77��6'30''E), 21.vii.2003, A. Gorodinski��� 1 ♂ (CMD); Yol, Kangra Valley, 1300 m (32��10'N, 76��12'E), viii���ix.1943, C. Lomi���1 ♀, 1 ♂ (CMNC); KARNATAKA, Kanara [=Coastal Karnataka], [no date], [anonymous]��� 2 ♂♂ (BMNH); Belgaum District, Belgaum (15��51'N, 74��33'0''E), [no date], [anonymous]��� 1 ♀, 3 ♂♂ (BMNH); Kodagu District, Mercara [= Madikeri] (12��25'14''N, 75��44'23''E), v.1973, T.R.S. Nathan��� 1 ♂ (RMNH); Mysore District, Mysore (12��18'N, 76��39'E), [no date], [anonymous]��� 1 ♀, 1 ♂ (BMNH); Shimoga District, Bhadravati (13��50'24''N, 75��42'7.2''E), vii.1938, [anonymous]��� 1 ♀ (FETC); Shimoga District, Bhadravati (13��50'24''N, 75��42'7.2''E), 13.v.1935, [anonymous]��� 1 ♀ (CMNC); Shimoga (13��55'N, 75��34'E), vi.1897, [anonymous]��� 1 ♀, 1 ♂ (MNHN); Shimoga (13��55'N, 75��34'E), [no date], [anonymous]��� 1 ♂ (NMPC); Thirthahalli, 700 m (13��42'N, 75��13'48''E), 14.xii.1989, A. Kumar��� 2 ♀♀ (OUMNH); KERALA, Kumily (9��37'6''N, 77��9'29''E), v.1986, T.R.S. Nathan��� 1 ♀, 1 ♂ (CMNC); Kumily (9��37'6''N, 77��9'29''E), vi.1986, T.R.S. Nathan���1 ♂ (CMNC); KERALA / TAMIL NADU, Anaimalai Hills, 1067 m (10��22'N, 77��7.5'E), iv.1963, P.S. Nathan��� 3 ♀♀, 3 ♂♂ (CNC); ORISSA, Balasore District, Balasore (21��30'N, 86��56'E), [no date], R.P. Gengler��� 1 ♀, 2 ♂♂ (MNHN); PONDICHERRY, Karaikal District, Karaikal (10��56'N, 79��50'E), iii.1963, P.S. Nathan��� 91 specimens, 1 ♀ (BDGC, CNC); PONDICHERRY, Karaikal District, Karaikal (10��56'N, 79��50'E), iv.1963, P.S. Nathan��� 20 specimens, 1 ♀ (BDGC, CNC); Karaikal (10��56'N, 79��50'E), ix.1969, P.S. Nathan��� 1 ♀ (CMNC); Karaikal (10��56'N, 79��50'E), iii.1968, T.R.S. Nathan��� 1 ♀ (CMNC); Karaikal (10��56'N, 79��50'E), v.1988, T.R.S. Nathan��� 1 ♀ (CMNC); Karaikal (10��56'N, 79��50'E), viii.1991, T.R.S. Nathan��� 1 ♀, 1 ♂ (CMNC); Karaikal (10��56'N, 79��50'E), xii.1961, P.S. Nathan��� 3 ♀♀, 2 ♂♂ (CNC); Karaikal (10��56'N, 79��50'E), i.1962, P.S. Nathan��� 5 ♀♀, 5 ♂♂ (CNC); Karaikal (10��56'N, 79��50'E), ii.1962, P.S. Nathan��� 5 ♀♀, 5 ♂♂ (CNC); Karaikal (10��56'N, 79��50'E), iii.1962, P.S. Nathan��� 5 ♀♀, 5 ♂♂ (CNC); Karaikal (10��56'N, 79��50'E), vi.1962, P.S. Nathan��� 1 ♀ (CNC); Karaikal (10��56'N, 79��50'E), iv.1962, P.S. Nathan��� 14 specimens (CNC); Karaikal (10��56'N, 79��50'E), v.1962, P.S. Nathan��� 2 specimens (CNC); Karaikal (10��56'N, 79��50'E), vii.1962, P.S. Nathan��� 45 specimens (CNC); Karaikal (10��56'N, 79��50'E), viii.1962, P.S. Nathan��� 21 specimens (CNC); Karaikal (10��56'N, 79��50'E), ix.1962, P.S. Nathan��� 63 specimens (CNC); Karaikal (10��56'N, 79��50'E), x.1962, P.S. Nathan��� 42 specimens (CNC); Karaikal (10��56'N, 79��50'E), xii.1962, P.S. Nathan��� 5 specimens (CNC); Karaikal (10��56'N, 79��50'E), i.1963, P.S. Nathan��� 36 specimens (CNC); Karaikal (10��56'N, 79��50'E), ii.1963, P.S. Nathan��� 18 specimens (CNC); Karaikal (10��56'N, 79��50'E), vi.1963, P.S. Nathan��� 8 specimens (CNC); Karaikal (10��56'N, 79��50'E), x.1963, P.S. Nathan��� 4 specimens (CNC); Karaikal (10��56'N, 79��50'E), x.1966, P.S. Nathan��� 4 specimens (CNC); Karaikal (10��56'N, 79��50'E), iii.1967, P.S. Nathan��� 4 specimens (CNC); Karaikal (10��56'N, 79��50'E), [no date], [anonymous]��� 1 ♀, 2 ♂♂ (FETC); Karaikal (10��56'N, 79��50'E), i���ii.1964, T.R.S. Nathan��� 4 ♀♀, 1 ♂ (MMUE); Karaikal (10��56'N, 79��50'E), viii.1963, T.R.S. Nathan��� 1 ♂ (MMUE); Karaikal (10��56'N, 79��50'E), v.2005, Nathan��� 1 ♀, 1 ♂ (PMOC); Karaikal (10��56'N, 79��50'E), v.1973, T.R.S. Nathan��� 1 ♂ (RMNH); Kurumbagaram (10��59'N, 79��46'E), xii.1950, P.S. Nathan��� 1 ♂ (CMNC); Nedungadu (10��58'N, 79��46'E), v.1938, [anonymous]��� 1 ♂ (FETC); Nedungadu (10��58'N, 79��46'E), [no date], [anonymous]��� 1 ♂ (FETC); Nedungadu (10��58'N, 79��46'E), 1936, P.S. Nathan��� 14 ♀♀, 10 ♂♂ (MNHN); Nedungadu (10��58'N, 79��46'E), 1.vi.1931, P.S. Nathan��� 1 ♂ (BMNH); PUNJAB, Rormazara, Hoshiarpur (31��11'50.7''N, 76��12'58''E), 6���9.vii.2004, M. Uhler��� 2 ♀♀ (CVM); TAMIL NADU, Arcot (12��54'N, 79��19'E), ix.2012, [anonymous]��� 1 ♀ (CMD); Coimbatore, 427 m (11��0'N, 76��57'E), iv.1962, P.S. Nathan��� 10 ♀♀, 12 ♂♂ (BDGC, CNC); Coimbatore, 427 m (11��0'N, 76��57'E), ix.1962, P.S. Nathan��� 1 ♀ (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), x.1962, P.S. Nathan��� 1 ♀, 4 ♂♂ (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), xi.1962, P.S. Nathan��� 1 ♀, 2 ♂♂ (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), xii.1962, P.S. Nathan��� 1 ♀, 3 ♂♂ (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), i.1963, P.S. Nathan��� 1 ♀, 1 ♂ (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), iv.1963, P.S. Nathan��� 10 ♀♀, 10 ♂♂ (CNC); Coimbatore, 427 m (11��0'N, 76��57'E), viii.1963, T.R.S. Nathan��� 1 ♀, 3 ♂♂ (MMUE); Coimbatore, 427 m (11��0'N, 76��57'E), ix.1964, T.R.S. Nathan��� 4 ♀♀, 2 ♂♂ (MMUE); Coimbatore, 427 m (11��0'N, 76��57'E), x���xi.1964, T.R.S. Nathan��� 1 ♀ (MMUE); Pudukkottai (10��23'N, 78��49'E), x.1984, T.R.S. Nathan��� 1 ♀ (CMNC); Thambikkottai (10��23'N, 79��27'E), vi.1962, P.S. Nathan��� 1 ♀ (CNC); Tranquebar [= Tharangambadi] (11��2'N, 79��5'E), [no date], [anonymous]��� 3 ♀♀, 1 ♂ (lectotype, paralectotypes, alloreferent) (ZMUC, ZMUK); Chennai District, Madras [= Chennai] (13��5'2''N, 80��16'12''E), xi.1907, Tomlin��� 1 ♀ (BMNH); Madras Airport [= Chennai Airport] (12��59'38''N, 80��9'28''E), x.1962, P.S. Nathan��� 3 ♀♀, 2 ♂♂ (CNC); Coimbatore District, Coimbatore (11��0'N, 76��57'E), iii.1962, P.S. Nathan��� 3 ♀♀ (CNC); Coimbatore District, Coimbatore (11��0'N, 76��57'E), vii.1939, [anonymous]��� 1 ♂ (FETC); Coimbatore District, Coimbatore (11��0'N, 76��57'E), [no date], P.S. Nathan��� 1 ♀, 2 ♂♂ (MNHN); Coimbatore District, Coimbatore (11��0'N, 76��57'E), xii.1971, T.R.S. Nathan��� 1 ♂ (RMNH); Coimbatore District, Coimbatore (11��0'N, 76��57'E), 15.ix.1947, P.S. Nathan��� 2 ♂♂ (CMNC); Marudamalai Hills (11��2'46''N, 76��51'7''E), xi.2004, Nathan��� 1 ♂ (PMOC); Tiruchirappalli District, Trichinopoly [= Tiruchirappalli] (10��48'18''N, 78��41'8''E), iii.1907, [anonymous]��� 1 ♀ (BMNH); Tiruchirappalli District, Trichinopoly [= Tiruchirappalli] (10��48'18''N, 78��41'8''E), x.1919, C. Leigh��� 1 ♂ (BMNH); Tiruchirappalli District, Trichinopoly [= Tiruchirappalli] (10��48'18''N, 78��41'8''E), [no date], R.P. Castets��� 1 ♀, 1 ♂ (MNHN); UTTAR PRADESH, Kheri Reserved Forest (25��21'11''N, 79��17'9''E), ii.1916, H.G. Champion��� 1 ♂ (BMNH); UTTARAKHAND, Haldwani District, Kumaon [= Haldwani] (29��13'N, 79��31'E), [no date], H.G. Champion��� 2 ♀♀ (BMNH); WEST BENGAL, Howrah District, Sarda (22��20'N, 88��8'E), [no date], F.W. Champion��� 2 ♂♂ (BMNH); Kalkata District, Calcutta [= Kolkata] (22��34'N, 88��22'E), [no date], [anonymous]��� 1 ♂ (BMNH); Kalkata District, Calcutta [= Kolkata] (22��34'N, 88��22'E), [no date], Waagen��� 1 ♀ (SMF); LAOS: CHAMPASAK, Ban Sepian (15��9'N, 106��15'E), 1���7.iv.2003, R. N.��� 1 ♂ (CEMT); PHONGSALY, Gnot-Ou District, Mount Phu Nam Kh�� (22��5'N, 102��27'E), 15���20.vi.2006, Local collectors��� 12 ♀♀, 7 ♂♂ (FGIC, OUMNH); VIENTIANE, Phou Khao Khouay National Bio-Biversity Conservation Area, 700��� 800 m (18��20.369'N, 102��48.523'E), 25���30.v.2008, A. Solodovnikov & J. Pedersen��� 6 ♀♀, 7 ♂♂ (FGIC, ZMUC); MALAYSIA: MALACCA, Malacca (2��12'N, 102��15'E), [no date], [anonymous]��� 1 ♂ (SMF); MYANMAR: KAYIN, Hpapun (18��3'30''N, 97��26'30''E), 1932, R.P. Loizeau��� 1 ♂ (MNHN); YANGON, Hlegu- Goygon (17��5'43''N, 96��13'16''E), iii.1997, M. Klicha��� 2 ♂♂ (OUMNH); Htauk Kyant (17��2'40''N, 96��8'12''E), ii.1997, M. Klicha��� 1 ♀, 1 ♂ (OUMNH); Rangoon [= Yangon] (16��48'N, 96��9'E), v.1934, F.J. Meggitt��� 3 ♀♀, 2 ♂♂ (BMNH); Rangoon [= Yangon] (16��48'N, 96��9'E), 1933���1934, F.J. Meggitt��� 5 ♀♀, 4 ♂♂ (BMNH); NEPAL: CENTRAL, Chitwan District, Sauraha, Chitwan National Park, 213 m (27��31'57''N, 84��30'20''E), 4.vi.1983, M.J.D. Brendell��� 1 ♂ (BMNH); Chitwan District, Sauraha, Chitwan National Park, 213 m (27��31'57''N, 84��30'20''E), 1���4.vi.1999, M. Pejcha��� 1 ♀ (FETC); Chitwan District, Sauraha, Chitwan National Park, 213 m (27��31'57''N, 84��30'20''E), 4���6.vi.1999, A. Kudrna��� 1 ♂ (FETC); PAKISTAN: ISLAMABAD CAPITAL TERRITORY, Islamabad (33��43'N, 73��4'E), 6.vi.1991, S. Prepsl��� 1 ♀ (CMD); PENJAB, Changa Manga Forest, 70 km S Lahore (31��5'N, 74��0'E), 19���21.viii.1998, L. Cern����� 3 ♀♀ (CVM); Kallar Kahar (32��47'32''N, 72��43'34''E), 26.vi.2008, A. Zubair��� 1 ♂ (PMOC); SRI LANKA: EASTERN, Pottuvil (6��52'N, 81��50'E), 1��� 12.vii.1983, O. Mehl��� 1 ♂ (ZMUC); NORTH CENTRAL, Girithale (8��0'N, 80��56'E), 16.viii.1979, G. de Rougemont��� 1 ♀ (OUMNH); Polonnaruwa (7��56'N, 81��0'E), 24.vii.1982, P. Moretto��� 2 ♂♂ (PMOC); NORTHERN, Jaffa (9��20.5'N, 80��24.3'E), 1.iii.2010, Kuduvidanage et al.��� 1 ♂ (OUMNH); SOUTHERN, Hambantota District, Tissamaharama (6��17'N, 81��17'E), 1.viii.1982, P. Moretto��� 2 ♀♀, 2 ♂♂ (PMOC); THAILAND: BANGKOK, Bangkok (13��45'N, 100��30'E), [no date], S.S. Flower��� 1 ♀ (BMNH); CHIANG MAI, Chiang Dao (19��25'N, 98��52'E), 17���24.v.1991, V. Kub��ň��� 1 ♀, 1 ♂ (CVM); Chiang Mai (18��48'N, 99��0'E), v.1986, N. Koyama��� 2 ♂♂ (CEMT); Chiang Mai (18��48'N, 99��0'E), 28.vi.1990, R. Minetti��� 1 ♀ (FETC), Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 15-19, DOI: 10.5281/zenodo.439444, {"references":["Fabricius, J. C. (1775) Systema Entomologiae sistens Insectorum Classes, Ordines, Genera, Species adiectis Synonymis, Locis Descriptionibus, Observationibus. Officina Libraria Kortii, Flensburg & Leipzig, Germany, xxxii + 832 pp.","Goeze, J. A. E. (1777) Entomologische Beytrage zu des Ritter Linne zwolften Ausgabe des Natursystems. Erster Teil. Weidmanns Erben und Reich, Leipzig, Germany, xvi + 736 pp.","Fabricius, J. C. (1781) Species insectorum exhibens eorum differentias specificas, synonyma auctorum, loca natalia, metamorphosis, adiecitis observationibus, descriptionibus. Tomus I. Carol Ernest Bohnii, Hamburg & Kiel, Germany, viii + 552 pp.","Pallas, P. S. (1781) Icones Insectorum praesertim Rossiae Sibiriaeque peculiarium quae collegit et descriptionibus illustravit. Wolfgangi Waltheri, Erlangen, Germany, [6] + 104 pp, 8 plates.","Herbst, J. F. W. (1786) Erste Mantisse zum Verzeichniss der ersten Klasse meiner Insektensammlung. Archiv der Insectengeschichte, 7 - 8, 153 - 182, plates 43 - 48.","Fabricius, J. C. (1787 a) Mantissa insectorum sistens eorum species nuper detectas adiectis characteribus generici, differentiis specificis, emendationibus, observationibus. Tom. I. Christ. Gottl. Proft., Copenhagen, Denmark, xx + 348 pp. https: // doi. org / 10.5962 / bhl. title. 36471","Olivier, A. G. (1789) Entomologie, ou histoire naturelle des insectes, avec leurs caracteres generiques et specifiques, leur descriptions, leur synonymie, et leur figure enluminee. Coleopteres. Tome premier. Beaudoin, Paris, France, xx + 497 pp. https: // doi. org / 10.5962 / bhl. title. 61905","Herbst, J. F. W. (1789) Natursystem aller bekannten in- und auslandischen Insekten, als eine Fortsetzung der von Buffonschen Naturgeschichte. Nach dem System des Ritters von Linne und Fabricius zu bearbeiten angefangen von Carl Gustav Jablonsky von Johan Friedrich Wilhem Herbst. Der Kafer zweyter Theil. Joachim Pauli, Berlin, Germany, lxiv + 330 + [6] pp., plates 7 - 20.","Gmelin, J. F. (1790) Caroli a Linne, systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, loci. Editio decima tertia, aucta, reformata. Tom. I, Pars IV. Classis V. Insecta. G. E. Beer, Leipzig, Germany, pp. 1517 - 2224.","Fabricius, J. C. (1792) Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species adjictis synonimis, locis, observationibus, descriptionibus. Tom. I. Christ. Gottl. Proft., Copenhagen, Denmark, xx + 330 pp.","Illiger, J. C. W. (1800) Olivier's Entomologie oder Naturgeschichte der Insekten mit ihren Gattungs- und ArtMerkmalen, ihrer Beschreibung und Synonymie. Kafer. Uebersetzt und mit Zusatzen und Anmerkungen durchgangig begleitet. Erster Theil. Karl Reichard, Braunschweig, Germany, xvi + 309 pp., 2 plates.","Fabricius, J. C. (1801) Systema Eleutheratorum secundum ordines, genera, species: adiectis synonymis, locis, observationibus, descriptionibus. Tomus I. Bibliopoli academici novi, Kiel, Germany, xxiv + 506 pp.","Illiger, K. (1804) Zusatze, Berichtigungen und Bemerkungen zu Fabricii. Systema Eleutheratorum. Tomus II. Magazin fur Insektenkunde, 3, 146 - 180.","Schonherr, C. J. (1806) Synonymia insectorum, ode: Versuch einer Synonymie aller bisher bekannten Insecten; nach Fabricii Systema Eleutheratorum geordnet, mit Berichtigungen und Anmerkungen, wie auch Beschreibungen neuer Arten und illuminirten Kupfern. Erter Band. Eleutherata oder Kafer. Erster Theil. Lethrus ... Scolytes. A. Nordstrom, Stockholm, Sweden, xxii + 293 pp., 3 plates.","Sturm, J. (1826) Catalog meiner Insecten-Sammlung. Erster Theil. Kafer. J. Sturm, Nuremberg, Germany, viii + 207 pp., 4 plates.","Latreille, P. A. (1827) Description des Insectes Figures sur la pl. LVIII du tom. II, recueillis par M. Cailliaud, et decrits par M. Latreille, membre de l'Academie royale des sciences, Etc. In: Voyage a Meroe, au Fleuve Blanc, au-dela de Fazoql dans le midi du royaume de Sennar, a Syouah et dans cinq autres oasis; fait dans les annees 1819, 1820, 1821, et 1822, par M. Frederic Cailliaud de Nantes, 4. Imprimerie Royale, Deleure, Paris, France, pp. 271 - 292.","LePeletier de Saint Fargeau, A. L. M. & Audinet-Serville, J. G. de (1828) Scarabe, Scarabaeus. In: Encyclopedie methodique, ou par ordre de matieres; par une societe de gens de lettres, de savant et d'artistes; precede d'un vocabulaire universel, servans de table pour tout l'ouvrage, ornee des portraits de Mm. Diderot & d'Alembert, premiers editeurs de l'Encyclopedie. Tome dixieme. Pars 2. Mme. veuve Agasse, Paris, France, pp. 345 - 832.","Hope, F. W. (1837) The Coleopterist's Manual, containing the lamellicorn insect of Linneus and Fabricius. Henry G. Bohn, London, United Kingdom, xiii + [2] + 15 - 121 + [4] pp., 3 plates.","Laporte, F. (Comte de Castelnau) (1840) Histoire Naturelle des Insectes Coleopteres. Tome deuxieme. Histoire Naturelle des animaux articulees, crustaces, arachnides, myriapode et insectes. Tome troisieme, 2, P. Dumenil, Paris, France, 564 pp.","Sturm, J. (1843) Catalog der Kaefer-Sammlung von Jacob Sturm. Jacob Sturm, Nurenberg, Germany, xii + 386 pp., 6 plates.","Motschulsky, V. de (1860) Sur les Collections Coleopterologiques de Linne et de Fabricius (Continuation.). Etudes Entomologiques, redigees par Victor de Motschulsky, 8, 147 - 168.","Motschulsky, V. de (1863) Essaie d'un catalogue des insectes de l'Ile Ceylan. Bulletin de la Societe Imperiale des Naturalistes de Moscou, 36, 421 - 532.","Harold, E. von (1869) Tom. IV. Scarabaeidae. In: Gemminger. M. & Harold, E. v. (Eds.), Catalogus Coleopterorum hucusque descriptorum synonymicus et systematicus autoribus Dr. Gemminger et B. de Harold. E. H. Gummi, Munich, Germany, pp. 979 - 1346, [1347 - 1353].","d'Orbigny, H. (1900 a) Synopsis des Onthophagides palearctiques. L'Abeille, 29, 117 - 254.","Boucomont, A. (1914) Onthophagus asiatiques nouveaux ou peu connus. Annali del Museo civico di storia naturale di Genova Serie 3. a, Volune 6, 46, 210 - 243.","Boucomont, A. (1921) [species treatment] In: Boucomont, A. & Gillet, J. J. E. (Eds.), Fam. Scarabaeidae, Laparosticti (Coleopteres). Faune entomologique de l'Indochine francaise. Fascicule 4, 1 - 76.","Gillet J. J. E. & Boucomont A. (1927) Scarabaeidae: Coprinae II, Termitotroginae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus. Partes 38 et 90, 38, 90. W. Junk, Berlin, Germany, pp. 101 - 264.","Arrow, G. J. (1931) Coleoptera Lamellicornia Part III. (Coprinae). The Fauna of British India, including Ceylon and Burma. Taylor and Francis, London, United Kingdom, 428 pp.","Paulian, R. (1945) Coleopteres Scarabeides de l'Indochine. Faune de l'empire Francais, 3. Librairie Larose, Paris, France, 227 pp.","Petrovitz, R. (1955) The 3 rd Danish expedition to central Asia. Zoological resuts 18. Scarabaeidae (Insecta) aus Afganistan. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i KObenhavn, 117, 275 - 287.","Balthasar, V. (1956) Scarabaeidae (Coleoptera) der Afghanistan-Expedition (1952 u. 1953) J. Klapperichs. 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(1988) Analisis taxonomico, ecologico y biogeografico de un grupo americana de Onthophagus (Coleoptera, Scarabaeidae). Monographie, Museo Regionale di Scienze Naturali, Torino, 9, 1 - 211.","Tarasov, S. I. & Solodovnikov, A. Y. (2011) Phylogenetic analyses reveal reliable morphological markers to classify megadiversity in Onthophagini dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Cladistics, 27, 1 - 39. https: // doi. org / 10.1111 / j. 1096 - 0031.2011.00351. x","Genier, F. (2013) Transfert d' Onthophagus bubalus Harold, 1867, dans le genre Phalops Erichson, 1847, et notes sur sa position phylogenetique (Coleoptera, Scarabaeidae, Scarabaeinae). Catharsius, La Revue, 7, 1 - 4.","Roggero, A., Barbero, E. & Palestrini, C. (2016 a) Revised classification and phylogeny of an Afrotropical species group based on molecular and morphological data, with the description of a new genus (Coleoptera: Scarabaeidae: Onthophagini). 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Digitonthophagus lusinganus (d���Orbigny, 1905) (Figs. 20, 49���50, 68, 84, 113���114, 132, 174���176; Map 15) Onthophagus (O.) gazella var. lusinganus d���Orbigny, 1905: 495 (original description) Onthophagus (O.) gazella var. lusinganus ��� d���Orbigny 1913b: 249 (monograph) Onthophagus (O.) gazella race lusinganus ��� Kolbe 1914: 297 (distribution) Onthophagus gazella var. lusinganus ��� Gillet & Boucomont 1927: 138 (mentioned as synonym) Onthophagus lusinganus ��� M��ller 1942: 96 (new combination, distribution) Onthophagus (Digitonthophagus) gazella var. lusinganus ��� Balthasar 1963: 365 (mentioned as synonym) Onthophagus (O.) lusinganus ��� Ferreira 1972: 680 (catalog) Onthophagus (Digitonthophagus) lusinganus ��� Kim & Lumaret, 1989: 336 (description of larvae) Type locality. Ile de Lusinga (Victoria-Nyanza N. -E.), Afrique Orientale Anglaise [Kenya]. Diagnosis. Male cephalic horns lacking granules internally at apex (Fig. 50); males with pronotal anterior angles surface distinctly concave, simply round anteriorly (Fig. 49); pronotal surface with dense, small punctures throughout (Fig. 70); FLP sclerite ventral portion with a single carinae (Fig. 176); SA sclerite apex interrupted much before apex of apical portion of FLP sclerite (Figs. 174���175). Description. Lectotype ♂ (Fig. 20). Measurements. Length 13.5 mm, width 8.0 mm. Head (Figs. 49���50). Anterior clypeal edge straight on median fifth in dorsal view; clypeofrontal carina broadly arcuate and interrupted at gena; vertex lacking median tubercle, surface with fine and minute punctures; fine punctures separated by one to three diameters. Horns rather long, arcuate, divergent in frontal view, gradually tapering from base to apex; posterointernal edge with a low, angular projection; apicointernal surface lacking granules. Punctures slightly scabrous; genal edge strongly upturned and distinctly angulate on anterior third, forming a broad angle with clypeal edge. Pronotum (Fig. 68). Surface with large, weakly-defined, strongly granulate punctures restricted to anterior half medially, with smaller scattered scabrous punctures on posterior half of disc; punctures weaker but distinct posterolaterally, with dense small punctures throughout; anteromedian tubercle well developed, produced into a small, transverse, sharp ridge on each side of midline, median longitudinal sulcus deep; surface behind the eyes slightly concave, surface of anterior angles concave, anterior angles unmodified; anterior half of lateral edge arcuate in dorsal and lateral view; posterior angles simply arcuate in dorsal view. Anterior hypomeral ridge arcuate throughout, anterior hypomeral depression surface slightly darker in color medially. Elytra (Fig. 20). Intervals 2 and 4 lacking fine granules from base to apical declivity. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus (Fig. 132). Parameres with dorsal and ventral edges diverging toward apex in lateral view. Internal sac sclerites (Figs. 174���176). Axial sclerite weekly sclerotized, short, semicircular. Subaxial sclerite wide basally and abruptly narrowed at mid distance, apical half short, extending straight, interrupted before apex of frontolateral peripheral sclerite apical portion, with villi on apical half. Frontolateral peripheral sclerite basoventral apophysis well developed, forming an acute angle in lateral view; with a more-or-less rectangular medioventral carina; right lateral fold produced into a spoon-like process apically; left lateral lobe moderately sclerotized and well developed; subapicodorsal lobe membranous, small, extending in line with apex of right lateral fold, apex set on left side in dorsal view; right preapical edge entire; apex of apical lobe round apically, directed obliquely on left side, apical villi regular in shape, ventral portion with a small additional lobate projection above subapicoventral lobe; subapicoventral lobe rather small, round and interrupted before apex of apical lobe. Variation. Measurements (64 ♂♂, 73 ♀♀). Length: male 9.5���15.0 mm (12.6 �� 1.2 mm), female 9.0���14.0 mm (12.1 �� 1.1 mm). Female paralectotype. Cephalic outline in dorsal view as in Fig. 84; vertex with a broadly arcuate, transverse carina, dorsal edge broadly arcuate in frontal view, lateral low and portion shortly gradually sloping down posteriorly; anterior pronotal tubercles well developed, external lateral edges rather short and slightly widening toward apex in dorsal view, anterolateral surface slightly obliquely concave, anterosuperior edge slightly arcuate in dorsal view (Fig. 113), lateral portion of anterosuperior edge slightly upturned (Fig. 114). Protibia short, with external teeth more robust. Primary type data. Lectotype ♂ (MNHN) present designation: [AFRIQUE OR le ANGLAISE / Ile de LUSINGA / (VICTORIA-NYANZA N.-E) / CH. ALLUAUD X. 1903]; [MUSEUM PARIS / Coll. H. d��� ORBIGNY / 1913] blue card; [WORLD / SCARAB. / DATABASE / WSD00030000] barcode label; [gazella Fabr. / v. lusinganus / var. nov. d���Orb.] handwritten on card; [LECTOTYPE ♂ / Onthophagus / gazella v. lusinganus / d���Orbigny, 1905 / d��s. F. G��nier, 2016] red card; [Digitonthophagus / lusinganus ♂ / (d���Orbigny, 1905) / d��t. F. G��nier, 2016]. MAP 15. Distribution of Digitonthophagus lusinganus. Material examined (91 ♂♂, 116 ♀♀), distribution (Map 15): ETHIOPIA: SOUTHERN NATIONS, NATIONALITIES, AND PEOPLE���S REGION, Arba Minch (6��2'N, 37��33'E), iv.1993, Werner��� 1 ♂ (PMOC); Arba Minch (6��2'N, 37��33'E), 6.ii.1978, P.-C. Rougeot��� 1 ♀ (MNHN); Arba Minch (6��2'N, 37��33'E), 6.ii.1978, P.-C. Rougeot��� 1 ♀ (MNHN); Arba Minch (6��2'N, 37��33'E), 11���17.ix.2000, P. L��onard��� 19 ♀♀, 12 ♂♂ (FGIC, PMOC); Lac Awassa (7��2'N, 38��28'E), 10.xi.1973, P.C. Rougeot��� 1 ♀ (MNHN); Nanoropus, bord du Rodolphe, 565 m (4��51'N, 36��5'E), 1932���1933, C. Arambour, P.- A. Chapuis, & R. Jeannel��� 6 ♀♀, 6 ♂♂ (FGIC, MNHN); Soddu [= Sodo] (6��51'N, 37��45'30''E), 20.vii.1971, [anonymous]��� 1 ♂ (FETC); KENYA: Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 14.xi.1972, T.J. Kingston��� 1 ♀, 1 ♂ (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 18.x.1973, T.J. Kingston��� 8 ♀♀, 4 ♂♂ (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 21.xi.1973, T.J. Kingston��� 4 ♀♀ (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 29.xi.1973, T.J. Kingston��� 1 ♂ (OUMNH); HOMA BAY, Baie de Kavirondo, Lac Victoria (0��31'S, 34��29'E), ix���x.1903, C. Alluaud��� 1 ♂ (MNHN); Ile de Lusinga [= Rusinga], Victoria-Nyanza N. -E. (0��24'S, 34��11'E), x.1903, C. Alluaud��� 12 ♀♀, 11 ♂♂ (lectotype, 22 paralectotypes) (IRSNB, MNHN); ISIOLO, Isiola Samburni (0��21'23''N, 37��34'56''E), 18.xi.1975, H. A.��� 1 ♀ (BMNH); Meru National Park, Tana River near Post 96, 275 m (0��10'N, 38��18'E), 13.xi.1975, C.F. Dewhurst��� 1 ♂ (BMNH); KISUMU, 3 km SE Kisumu, 1120 m (0��9'S, 34��51'E), 17.iv.1975, Davis & Dewhurst��� 1 ♀ (MNHN); 6 km S Ahero, 1200 m (0��13'16''S, 34��57'24''E), 22.xi.1975, C.F. Dewhurst��� 1 ♂ (MNHN); Kisumu (0��6'S, 34��46'E), [no date], P.S. Corbet��� 1 ♀ (BMNH); MAKUENI, Kibwezi (2��24'38''S, 37��58'4''E), 20.xi.1989, P. Smerz��� 1 ♂ (JFJC); MERU, Leopard Rock Swamp - Post 12���33, Meru National Park, 600 m (0��12'N, 38��14'E), 14.xi.1975, C.F. Dewhurst��� 1 ♂ (MNHN); NAROK, Massai Mara National Reserve (1��29'S, 35��7'E), ii.1994, Czeppel��� 1 ♂ (FETC); Massai Mara National Reserve (1��29'S, 35��7'E), iii.1995, Cursi��� 1 ♂ (FETC); RIFT VALLEY, Amboseli National Park (2��38'S, 37��14'E), 22.i.1986, R. Minetti��� 2 ♀♀, 1 ♂ (PMOC); Headquarters area, Lake Amboseli basin, Amboseli National Park, 1050 m (2��39'14''S, 37��15'22''E), 5.v.1975, Davies & Dewhurst��� 1 ♀ (BMNH); Talek River, Massai Mara, 1300 m (1��26'S, 35��10'E), xii.1997, P. Bleuzen��� 1 ♀, 1 ♂ (JFJC); SAMBURU, approximately 35 km N Wamba, Mattews Range, Rift Valley, 1300���1400 m (1��10.707'N, 37��18.962'E), 7���12.xii.2002, C. Hauser, D. Bartsch, & A. Zahm��� 1 ♂ (SMNS); TAITA TAVETA, Entrance Gate Tsavo National Park, Voi (3��21'47''S, 38��35'46''E), 26.xii.1990, B.D. Gill��� 1 ♀ (BDGC); Sagala Hills (3��29'S, 38��35'E), xii.1993, Werner��� 1 ♀ (CMD); Taita (3��22'S, 38��22'E), [no date], [anonymous]��� 2 ♀♀, 1 ♂ (BMNH); Taveta (3��24'S, 37��41'E), i���iv.1904, C. Alluaud��� 1 ♂ (MNHN); Taveta (3��24'S, 37��41'E), iii.1912, Alluaud & Jeannel (St. 65)��� 1 ♂ (MNHN); Tsavo, 60 km N Voi (2��59'42''S, 38��27'41''E), 25.xii.1990, B.D. Gill��� 2 ♂♂ (BDGC); Voi (3��23'S, 38��34'E), xi.1997, M. Snižek��� 1 ♂ (CEMT); Voi (3��23'S, 38��34'E), vi/1997, O. Bužga��� 1 ♂ (PMOC); Voi (3��23'S, 38��34'E), 10.xii.1999, P. Smerz��� 1 ♀ (JFJC); Voi, Tsavo East National Park (3��23'S, 38��34'E), 31.xii.1990, B.D. Gill��� 20 ♀♀, 12 ♂♂ (BDGC); SOMALIA: JUBBADA HOOSE, Kismaayo, Mareri (0��21'37''S, 42��32'56''E), 29.x.1986, M. J.W. Cock��� 1 ♀, 1 ♂ (BMNH); SHABEELLAHA HOOSE, Genale (1��48'N, 44��42'0''E), v.1935, F. Bigi��� 1 ♀ (FETC); Genale (1��48'N, 44��42'0''E), [no date], [anonymous]��� 1 ♀ (NMPC); SOUTH SUDAN: UPPER NILE, Malakal (9��32'N, 31��39'E), 30.xi.1960, B. Hocking��� 1 ♀ (BMNH); SUDAN: KHARTOUM, Kawa, 157 mi. S Khartoum (13��45'N, 32��30'E), 2���3.xii.1961, J.L. Cloudsley- Thompson��� 2 ♀♀, 1 ♂ (BMNH); SENNAR, [unspecified locality], 1906, C. Alluaud���2 ♀♀, 2 ♂♂ (IRSNB); WHITE NIL, El Jebelain (12��35'N, 32��50'E), 3���4.xii.1961, J.L. Cloudsley-Thompson��� 1 ♀ (BMNH); TANZANIA: ARUSHA, Arusha environs (3��22'S, 36��41'E), 3���4.iv.1997, M. Kuboň��� 5 ♀♀, 1 ♂ (PMOC); Orekeryan, Mount Longido (2��43'47''S, 36��43'26''E), 8.xi.2011, R. Smith & H. Takano��� 1 ♂ (BMNH); DODOMA, 40 km N Dodoma, 1100 m (5��54'S, 35��45'E), 15.xii.2007, F. Kantner��� 1 ♂ (SMNS); Babati-Dodoma (5��15'S, 35��52'E), 6.xii.1997, Werner & Lizler��� 1 ♂ (CMD); KILIMANDJARO, Ibaya Camp, Mkomazi Game Reserve (4��5'11''S, 38��4'55''E), 13.iv.1995, J.G. Davies��� 1 ♀ (BMNH); Ibaya Camp, Mkomazi Game Reserve (4��5'11''S, 38��4'55''E), 29.iii.1996, G.C. McGavin��� 1 ♂ (OUMNH); Same (4��4'S, 37��45'E), 12���17.v.1999, O. Bužga��� 3 ♂♂ (PMOC); MANYARA, Kibaone, 1348 m (3��20'11''S, 35��46'36''E), 8���10.v.2012, R. Smith & H. Takano��� 1 ♂ (BMNH); Mto Wa MBu, Lake Manyara National Park, 963 m (3��23'12''S, 35��52'36''E), 21.xi.2011, R. Smith & H. Takano��� 14 ♀♀, 11 ♂♂ (BMNH); MARA, Seronera Wildlife Lodge (2��26'50''S, 34��48'24''E), 14��� 17.ii.1989, G. Bassi & Scaramozzino��� 1 ♂ (CMNC); MOROGO, Monts Uluguru (7��14'S, 37��33'E), 20.xi.2005, Local collectors��� 1 ♀ (PMOC); SHINYANGA, Lake Ndutu area, Serengeti National Park (2��59'S, 35��1'E), 20.i.1991, R. Foster��� 1 ♀ (OUMNH). Natural history. No habitat data, some specimens collected in buffalo and elephant dung. This species is attracted to mercury vapor and ultraviolet light traps. Nomenclature and taxonomy. Digitonthophagus lusinganus was first described as a variety of Onthophagus gazella by d���Orbigny (1905). In a note on the Coleoptera of Benadir (Somalia) M��ller (1942) concluded that the coexistence of two forms of ��� O. gazella ��� and the ��� O. gazella form lusinganus ��� in several East African localities was not possible and considered this form as a good species. We concur with M��ller (1942) and Ferreira (1972) that this revised status is correct. The original syntype series comprises 23 specimens deposited in the Mus��um national d���Histoire naturelle (Paris) and Royal Belgian Institute of Natural Sciences (Brussels). In order to stabilize nomenclature a male specimen possessing the best diagnostic characters is selected as lectotype, (present designation). This lectotype is designated in order to choose a single specimen as name bearing type in the event that the other specimens of the syntype series could belong to a different cryptic species in this genus., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 65-67, DOI: 10.5281/zenodo.439444, {"references":["d'Orbigny, H. (1913 b) Insectes Coleopteres II. Scarabaeidae Onthophagini. In: Voyage de Ch. Alluaud et R. Jeannel en Afrique Orientale (1911 - 1912). Resultats scientifiques. Librairie Albert Schulz, Paris, pp. 65 - 128.","Kolbe, H. J. (1914) Lieferung 3: Scarabaeiden. In: Shubotz, H. (Ed.), Wissenschaftliches Ergebnisse der Deutschen Zentral- Afrika-Expedition 1907 unter Fuhrung Adolf Friedrichs, Herzog zu Mecklenburg. Zoologie III, 5. Klinkhardt & Biermann, Leipzig, Germany, pp. 261 - 416.","Gillet J. J. E. & Boucomont A. (1927) Scarabaeidae: Coprinae II, Termitotroginae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus. Partes 38 et 90, 38, 90. W. Junk, Berlin, Germany, pp. 101 - 264.","Muller, G. (1942) Coleotterie del Benadir (Somalia) raccolti dal Prof. Giuseppe Russo. Bollettino del Laboratorio di Zoologia generale e agraria della Facolta Agraria in Portici, 32, 70 - 103.","Balthasar, V. (1963) Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region. Coleoptera: Lamellicornia. Band, 2. Coprinae (Onitini, Oniticellini, Onthophagini). Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region, 2. Verlag der Tschechoslowakischen Akademie der Wissenschaften, Prague, Czech Republic, 627 pp., 16 plates.","Ferreira, M. C. (1972) Os escarabideos de Africa (sul do Saara). I. Revista de Entomologia de Mocambique, 11, 5 - 1088.","Kim, J. I. & Lumaret, J. - P. (1989) Les larves du genre Onthophagus Latreille, 1802. 5 - Les especes des sous-genres Digitonthophagus, Trichonthophagus et \" Incetae Sedis \" (Coleoptera Scarabaeidae). Bulletin Mensuel de la Societe Linneenne de Lyon, 58, 335 - 344."]}

Digitonthophagus aksumensis G��nier, new species http://zoobank.org/urn:lsid:zoobank.org:act:2316941C-3BDF-4900-9212-25C83F5AC88F (Figs. 17, 43���44, 65, 81, 107���108, 129, 165���167, 184; Map 12) Type locality. Obock, Colonie fran��aise des Somalies [= Djibouti]. Diagnosis. Male cephalic horns lacking granules internally at apex (Fig. 44); FLP sclerite apical portion rather large, obliquely oriented and tapering to apex (Fig. 165���166); males with pronotal anterior angles surface convex, simply round anteriorly (Fig. 43); FLP sclerite ventral portion with two carinae (Fig. 167); SA sclerite apex approximately in line with apex of apical portion of FLP sclerite (Figs. 165���166). Description. Holotype ♂ (Fig. 17). Measurements. Length 11.0 mm, width 6.5 mm. Head (Figs. 43���44). 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. Horns rather short, slightly divergent in frontal view, gradually tapering from base to apex; posterointernal edge produced into an low, angular projection basally; apicointernal surface lacking granules; genal edge upturned and distinctly angulate on anterior third, forming a distinct angle with clypeal edge. Pronotum (Fig. 65). Surface with granulate punctures restricted to anterior half medially, with distinct umbilicate punctures on posterior half of disc; punctures weaker but distinct posterolaterally with distinct, minute punctures throughout. Anteromedian tubercle atrophied, forming an obtuse angle in lateral view, median longitudinal sulcus weakly-defined; surface behind the eyes with a simple round depression, surface of anterior angles convex; anterior half of lateral edge arcuate in dorsal and lateral view; posterior angles slightly upturned and simply arcuate in dorsal view. Anterior hypomeral ridge arcuate, anterior hypomeral depression slightly darker in color medially. Elytra (Fig. 17). Intervals 2 and 4 lacking fine granules from base to apex. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus (Fig. 129). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites (Figs. 165���167). Axial sclerite weekly sclerotized, reduced in size and almost straight. Subaxial sclerite, large basally and abruptly reduced, extending straight approximately in line with apex of frontolateral peripheral sclerite apical portion, with fine and scattered villi. Frontolateral peripheral sclerite basoventral apophysis well developed; two medioventral carinae present; right lateral fold produced into a rather large everted and open apically conical process with irregular apical edge; left lateral lobe membranous, slightly developed; subapicodorsal lobe membranous not reaching anterior edge, apex set on left side in dorsal view; apical lobe tapering into a point and directed obliquely on left side, apical villi regular in shape; subapicoventral lobe acute and in line with apical lobe apically. Variation. Measurements (86 ♂♂, 69 ♀♀). Length: male 6.5���11.5 mm (10.1 �� 1.0 mm), female 7.0���11.0 mm (9.6 �� 0.9 mm). Female allotype. Cephalic outline in dorsal view as in Fig. 81; vertex with a straight, transverse carina, dorsal edge broadly arcuate in frontal view, lateral portion gradually sloping down; anterior pronotal tubercles atrophied, external lateral edges parallel sided, anterolateral surface simply convex, anterosuperior edge slightly arcuate in dorsal view (Fig. 107), lateral portions of anterosuperior edge slightly upturned (Fig. 108). Protibia short, with external teeth more robust. Primary type data. Holotype ♂ (MNHN): [C. FR. Des SOMALIES / Obock] handwritten; [MUS��UM PARIS / 1918 / Dr. Yousseaume] partly handwritten on blue card; [WORLD / SCARAB. / DATABASE / WSD00029901]; [HOLOTYPE ♂ / Digitonthophagus / aksumensis n.sp. / des. F. G��nier, 2016] red card. MAP 12. Distribution of Digitonthophagus aksumensis. Material examined (92 ♂♂, 74 ♀♀), distribution (Map 12): DJIBOUTI: ARTA, Grand Bara (11��16'N, 42��39'E), 22.xii.1989, [anonymous] ���1 ♂ (paratype) (CMD); Grand Bara (11��16'N, 42��39'E), 24.v.1991, G. Nazaret��� 4 ♀♀, 2 ♂♂ (6 paratypes) (CMD); DJIBOUTI, Ambouli (11��33'13''N, 43��9'31''E), v.1990, G. Nazaret��� 3 ♀♀, 3 ♂♂ (6 paratypes) (CMD); Ambouli (11��33'13''N, 43��9'31''E), 9.xi.1972, Mission Balachowsky-Menier��� 1 ♀, 3 ♂♂ (4 paratypes) (MNHN); Ambouli (11��33'13''N, 43��9'31''E), 15.ii.1990, G. Nazaret��� 4 ♀♀, 4 ♂♂ (8 paratypes) (PMOC); Ambouli (11��33'13''N, 43��9'31''E), 21.ii.1990, G. Nazaret��� 2 ♀♀, 2 ♂♂ (4 paratypes) (CMD); Ambouli (11��33'13''N, 43��9'31''E), 12.i.1992, G. Nazaret��� 2 ♂♂ (2 paratypes) (CMD); Ambouli (11��33'13''N, 43��9'31''E), 12.i.1992, [anonymous]���1 ♂ (paratype) (PCSR); Djibouti (11��35'N, 43��9'E), vii.1897, Jousseaume��� 6 ♀♀, 7 ♂♂ (13 paratypes) (MNHN); Djibouti (11��35'N, 43��9'E), 1897, H. Couti��re��� 2 ♀♀, 1 ♂ (paratypes) (MNHN); Djibouti (11��35'N, 43��9'E), [no date], [anonymous]��� 2 ♀♀, 1 ♂ (paratypes) (MNHN); OBOCK, Obock (11��58'N, 43��17'E), 1918, Dr. Yousseaume��� 3 ♀♀, 4 ♂♂ (holotype, allotype, 5 paratypes) (MNHN); Obock (11��58'N, 43��17'E), [no date], Faurot��� 1 ♀ (paratype) (MNHN); Obock (11��58'N, 43��17'E), 1893, Maindron��� 1 ♀, 1 ♂ (paratypes) (MNHN); ERITREA: SEMENAWI-KEYIH-BAHRI, Massawa (15��36'35''N, 39��27'0''E), [no date], [anonymous]��� 1 ♂ (paratype) (BMNH); ETHIOPIA: DIRE DAWA, Dire Dawa (9��36'N, 41��51'E), ix.1935, H. Ulenhuth��� 1 ♀, 2 ♂♂ (3 paratypes) (BMNH); Dire Dawa (9��36'N, 41��51'E), 1936, [anonymous] ��� 1 ♀, 1 ♂ (paratypes) (MNHN); Dire Dawa (9��36'N, 41��51'E), 1935, [anonymous] ��� 1 ♀ (paratype) (MNHN); OROMIA, 23 km NEE Welenchiti (8��46'N, 39��36'E), 29.iv.2013, [anonymous] ��� 1 ♂ (paratype) (CMD); 30 km SW Awash (8��54'N, 39��56'E), 29���30.iv.2013, I. Martinu���1 ♂ (paratype) (JFJC); SOMALI, Adigala (10��25'35''N, 42��14'25''E), [no date], Dr. Martin��� 2 ♂♂ (2 paratypes) (MNHN); Dewele (11��2'5''N, 42��37'53''E), [no date], Dr. Martin��� 1 ♂ (paratype) (MNHN); Lassarat (10��39'52''N, 42��27'21''E), [no date], Dr. Martin��� 2 ♀♀, 7 ♂♂ (9 paratypes) (MNHN); SAUDI ARABIA: Hedjaz (24��30'N, 38��30'E), [no date], [anonymous]��� 1 ♀, 2 ♂♂ (3 paratypes) (MNHN); [unspecified locality], [no date], [anonymous] ���1 ♀, 1 ♂ (paratypes) (MNHN); ASEER, 35 km N Mahayl [=Muhayil] (18��50'N, 41��58'E), 7.xi.1982, A.R. Pittaway��� 1 ♂ (paratype) (OUMNH); Muhayl [=Muhayil] (18��32'40''N, 42��3'3''E), 8.i.1983, A.R. Pittaway��� 1 ♀, 1 ♂ (paratypes) (OUMNH); MAKKAH, Lith, 10 mi. inland (20��9'N, 40��16'E), i.1945, B.P. Uvarov��� 1 ♂ (paratype) (BMNH); Wadi Jizan (21��37'N, 39��10'E), 18.iii.1978, Filipponi��� 1 ♂ (paratype) (FETC); Wadi Jizan (21��37'N, 39��10'E), 26.iii.1978, Filipponi��� 1 ♀, 1 ♂ (paratypes) (FETC); Wadi Jizan (21��37'N, 39��10'E), 6.viii.1978, Filipponi��� 1 ♂ (paratype) (FETC); Wadi Jizan (21��37'N, 39��10'E), 11.viii.1978, Filipponi��� 2 ♀♀, 6 ♂♂ (8 paratypes) (FETC); Wadi Jizan (21��37'N, 39��10'E), 14.viii.1978, Filipponi��� 7 ♀♀, 7 ♂♂ (14 paratypes) (FETC); Wadi Jizan (21��37'N, 39��10'E), 18.viii.1978, Filipponi��� 7 ♀♀, 2 ♂♂ (9 paratypes) (FETC); Wadi Jizan (21��37'N, 39��10'E), 24.viii.1978, Filipponi��� 1 ♂ (paratype) (FETC); Wadi Jizan (21��37'N, 39��10'E), 30.xi.1978, Filipponi��� 1 ♀, 2 ♂♂ (3 paratypes) (FETC); MINTAQAT MAKKAH, 12 mi. N Jidda [=Jeddah] (21��38'15''N, 39��13'35''E), 4.xii.1944, A.R. Waterston��� 1 ♂ (paratype) (BMNH); SOMALIA: Dolphin Bay, 22.iii.1903, M. Cameron��� 2 ♀♀, 1 ♂ (paratypes) (BMNH); SUDAN: KHARTOUM, Environs de Khartoum (15��35'N, 32��32'E), [no date], [anonymous]��� 2 ♀♀, 1 ♂ (paratypes) (MNHN); Khartoum (15��35'N, 32��32'E), 18.i.1904, Zaphiro��� 1 ♀, 1 ♂ (paratypes) (BMNH); RED SEA, Khor Arbaat Delta (20��1'N, 37��8'E), iv���v.1926, H.B. Johnston���1 ♂ (paratype) (BMNH); Tokar (18��26'N, 37��44'E), 1914, H.W. Bedford��� 1 ♂ (paratype) (BMNH); YEMEN: [unspecified locality] (14��14'N, 44��29'E), [no date], Mellingen��� 3 ♀♀, 2 ♂♂ (5 paratypes) (BMNH); [unspecified locality] (14��14'N, 44��29'E), [no date], [anonymous]���2 ♀♀, 1 ♂ (paratypes) (MNHN); ADEN, Aden (12��45'56''N, 45��1'1''E), [no date], [anonymous] ���2 ♀♀, 1 ♂ (paratypes) (BMNH); Aden (12��45'56''N, 45��1'1''E), [no date], C. P��rez��� 4 ♀♀, 4 ♂♂ (8 paratypes) (MNHN); AL HUDAYDAH, Kamaran Island (15��20'N, 42��35'E), 24.xii.1902, M. Cameron��� 1 ♀, 1 ♂ (paratypes) (BMNH); Lahij, Lahij (13��3'N, 44��53'E), [no date], Dodson��� 2 ♀♀, 1 ♂ (paratypes) (MNHN); TA���IZZ, Taizz (13��34'N, 44��2'E), xi.1975, N. Hadi��� 1 ♂ (paratype) (BMNH). Etymology. Derived from Aksum, a historical kingdom approximately corresponding to the distribution of the species. Natural history. Unknown except for two specimens collected from camel dung and five specimens collected at light. Remarks. The Egyptian records for D. gazella from Quena in the Nil Valley (Alfieri 1976) presumably refer to this species., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 57-59, DOI: 10.5281/zenodo.439444, {"references":["Alfieri, A. (1976) The Coleoptera of Egypt. Memoires de la Societe Entomologique d'Egypte, 5. Atlas Press, Cairo, Egypt, xvi + 362 pp."]}

Digitonthophagus lusinganus species group 11. Digitonthophagus aksumensis G��nier, new species Distribution: Djibouti, Eritrea, Ethiopia, Saudi Arabia, Somalia, Sudan, Yemen 12. Digitonthophagus dilatatus G��nier, new species Distribution: Burkina Faso, Cameroon, Chad, C��te d���Ivoire, Ghana, Nigeria 13. Digitonthophagus fimator G��nier, new species Distribution: Benin, Burkina Faso, Cameroon, Central Africa republic, Chad, C��te d'Ivoire, Democratic Republic of the Congo, Ethiopia, Ghana, Guinea, Guinea-Bissau, Nigeria, Senegal, Sierra Leone, Togo, Uganda 14. Digitonthophagus lusinganus (d���Orbigny, 1905), valid species Distribution: Ethiopia, Kenya, Somalia, South Sudan, Sudan, Tanzania 15. Digitonthophagus ulcerosus G��nier, new species Distribution: Ethiopia, Kenya, Somalia, Tanzania, Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on page 74, DOI: 10.5281/zenodo.439444

Digitonthophagus bonasus species group 1. Digitonthophagus bonasus (Fabricius, 1775), valid species Distribution: Bangladesh, Cambodia, India, Laos, Malaysia, Myanmar, Nepal, Pakistan, Sri Lanka, Thailand, Vietnam 2. Digitonthophagus uks G��nier, new species Distribution: Afghanistan, India, Iran, Pakistan 3. Digitonthophagus sahelicus Moretto, new species Distribution: Burkina Faso, Cameroon, Chad, Djibouti, Eritrea, Mali, Mauritania, Niger, Senegal, Sudan, Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on page 74, DOI: 10.5281/zenodo.439444, {"references":["Fabricius, J. C. (1775) Systema Entomologiae sistens Insectorum Classes, Ordines, Genera, Species adiectis Synonymis, Locis Descriptionibus, Observationibus. Officina Libraria Kortii, Flensburg & Leipzig, Germany, xxxii + 832 pp."]}

Digitonthophagus sahelicus Moretto, new species http://zoobank.org/urn:lsid:zoobank.org:act:659D2617-892E-4CA3-803B-A727B64D643C (Figs. 9, 27���28, 57, 73, 91���92, 121, 141���143; Map 4) Type locality. Toulf�� (13��53���43���N 01��52���25���W), 300 m, Lorum, Burkina Faso. Diagnosis. Africa species; Male pronotal anterior angles surface sloping laterally and usually with a dark spot (Figs. 57); female vertex with horns (Fig. 73). Description. Holotype ♂ (Fig. 9). Measurements. Length 9.5 mm, width 5.5 mm. Head (Figs. 27���28). Anterior clypeal edge emarginate on median fifth in dorsal view; clypeofrontal carina nearly straight and extending to clypeal edge; vertex convex medially, surface with scabrous punctures. Horns long, divergent in frontal view, gradually tapering from base to apex, posterointernal edge unmodified basally, internal surface with numerous coarse granules. Genal edge slightly upturned and arcuate on anterior third, forming a broad angle with clypeal edge. Pronotum (Fig. 57). Surface with granulate punctures extending slightly beyond posterior half, with a few scattered simple punctures on posterior portion of disc, larger punctures absent posterolaterally, with distinct, minute punctures throughout. Anteromedian tubercle atrophied, forming a simply convex projection in lateral view, surface of tubercles covered with rather regular and fine squamiform granules; median longitudinal sulcus absent; surface behind the eyes with a small and shallow oval depression, surface of anterior angles irregularly sloping laterally; anterior half of lateral edge slightly sinuous in dorsal and lateral view; posterior angles simply arcuate in dorsal view. Anterior hypomeral ridge nearly straight posteriorly and abruptly arcuate anteriorly with anterior most portion crenulate, anterior hypomeral depression surface light in color. Elytra (Fig. 9). Interval 2 and 4 lacking fine granules from base to apex. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge interrupted before apex. Aedeagus (Fig. 121). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites (Figs. 141���143). Axial sclerite strongly sclerotized, abruptly bent ventrally on apical third. Subaxial sclerite, extending straight past apex of right lateral fold apical portion, with villi on apical half. Frontolateral peripheral sclerite basoventral apophysis moderately developed; a single medioventral carinae present; right lateral fold short, produced into a rather large everted and open apically conical process with extremely irregular apical edge; left lateral lobe absent; subapicodorsal lobe membranous, narrow, not reaching anterior edge, apex set on left side in dorsal view; apical lobe elongate and round apically, directed obliquely on left side, apical villi regular in shape; subapicoventral lobe short not reaching apical edge of apical lobe, oriented obliquely and in line with ventrally folded left edge of apical lobe. Variation. Measurements (103 ♂♂, 111 ♀♀). Length: male 6.5���11.0 mm (9.5 �� 0.8 mm), female 7.5���11.0 mm (9.4 �� 0.7 mm). Female lectotype. Cephalic outline in dorsal view as in Fig. 73; vertex with horns low, very wide basally and tapering to apex, posterointernal edge lacking tooth basally; anterior pronotal surface (Figs. 91���92) with a single callus approximately half interocular distance in width, dorsal edge of callus weakly defined. Protibia short, with external teeth more robust. Primary type data. Holotype ♂ (CMNC): [BURKINA FASO: LORUM / Toulf��, 300m / 13��53���43���N 01��52���25���O / 18.VII.2006, zone sah��lienne / steppe arbor��e, collecte g��n��- / rale, F. & S. G��nier, 2006-43]; [COL��OPT��RES / DU / BURKINA FASO / BF008585] barcode label; [HOLOTYPE ♂ / Digitonthophagus / sahelicus n.sp. / des. P. Moretto, 2016] red card. Map 4. Distribution of Digitonthophagus sahelicus. Material examined (266 ♂♂, 193 ♀♀, 496 sex not recorded), distribution (Map 4): BURKINA FASO: HAUTS-BASSINS, Bobo-Dioulasso (11��12'N, 4��18'W), viii.2002, [anonymous] ���1 ♀ (paratype) (PMOC); SAHEL, apr��s le pont, route Essakane-Dori, Gorum-Gorum, 271 m (14��9'32.8''N, 0��1'20.4''E), 7.viii.2012, P. Moretto��� 1 ♀, 1 ♂ (paratypes) (PMOC); Conseil r��gional, Dori, 283 m (14��2'4.5''N, 0��3'10.5''W), 23��� 25.viii.2013, P. Moretto (2013-50)��� 9 ♀♀, 9 ♂♂ (18 paratypes) (PMOC); Conseil r��gional, Dori, 283 m (14��2'4.5''N, 0��3'10.5''W), 30.viii.2013, P. Moretto��� 1 ♀ (paratype) (PMOC); Dori (14��2'N, 0��1'W), iv.2007, [anonymous]��� 8 ♀♀, 3 ♂♂ (11 paratypes) (JFJC); Essakane, Gorum-Gorum, 264 m (14��22'40.2''N, 0��5'47.7''E), 13���15.viii.2012, P. Moretto��� 1 ♀, 1 ♂ (paratypes) (PMOC); N���djomga, Dori (14��3'55.7''N, 0��3'3.4''W), 24��� 27.viii.2013, P. Moretto��� 3 ♂♂ (3 paratypes) (PMOC); LOROUM, Toulf��, 300 m (13��53'43''N, 1��52'25''W), 16.vii.2006, F. & S. G��nier (2006-38)��� 24 ♀♀, 28 ♂♂ (52 paratypes) (FGIC); Toulf��, 300 m (13��53'43''N, 1��52'25''W), 16.vii.2006, F. & S. G��nier (2006-40)��� 2 ♀♀, 13 ♂♂ (15 paratypes) (FGIC); Toulf��, 300 m (13��53'43''N, 1��52'25''W), 17.vii.2006, F. & S. G��nier (2006-41)��� 2 ♀♀, 7 ♂♂ (9 paratypes) (FGIC); Toulf��, 300 m (13��53'43''N, 1��52'25''W), 18.vii.2006, F. & S. G��nier (2006-43)��� 23 ♀♀, 24 ♂♂ (holotype, allotype, 45 paratypes) (CMNC, FGIC); Toulf��, 300 m (13��53'43''N, 1��52'25''W), 18.vii.2006, F. & S. G��nier (2006-42)��� 4 ♂♂ (4 paratypes) (FGIC); NAHOURI, For��t de Nazinga, Boulieselo, 310 m (11��11'50''N, 1��35'9''W), 27.vii.2006, F. & S. G��nier (2006-82)��� 2 ♂♂ (2 paratypes) (FGIC); CAMEROON: EXTREME NORTH, Waza National Park (11��15'N, 14��39'E), 18.viii.2006, C. Vanderbergh��� 1 ♂ (paratype) (PMOC); CHAD: ENNEDI, Fada (17��11'N, 21��35'E), 9/4/1935, [anonymous]��� 1 ♂ (paratype) (MNHN); HADJER-LAMIS, Sitje, rive sud Lac Tchad (12��53'N, 14��52'E), 11.viii.2006, C. Vanderbergh��� 1 ♀, 2 ♂♂ (3 paratypes) (PMOC); KANEM, Zigue�� (14��43'N, 15��47'E), 19���21.vii.1935, [anonymous] ���1 ♀ (paratype) (MNHN); N���DJAMENA, Farcha (12��7'N, 14��59'E), vi.1968, [anonymous]��� 1 ♀, 1 ♂ (paratypes) (MNHN); DJIBOUTI: DJIBOUTI, Djibouti (11��35'N, 43��9'E), [no date], [anonymous]��� 4 ♀♀, 3 ♂♂ (7 paratypes) (IRSNB); ERITREA: ANSEBA, Hagaz (15��42'N, 38��16'E), vii.2003, M. Forti��� 1 ♂ (paratype) (PMOC); GASH-BARKA, 18 km E Akordat (15��33'N, 38��1'E), vii.2003, M. Forti��� 2 ♀♀, 3 ♂♂ (5 paratypes) (FGIC, PMOC); NORTHERN RED SEA, Nefasit (15��20'N, 39��4'E), vii.2002, Czeppel & Forti��� 3 ♀♀, 3 ♂♂ (6 paratypes) (FETC); MALI: SEGOU, Sansanding (13��43'18''N, 6��0'11''W), [no date], R. Demange��� 1 ♀ (paratype) (MNHN); MAURITANIA: Trarza Desert, [no date], [anonymous]��� 2 ♀♀, 2 ♂♂ (4 paratypes) (IRSNB); ADRAR, Atar (20��31'N, 13��3'W), 24.ix.1968, P. Tauzin��� 1 ♂ (paratype) (PMOC); INCHIRI, Oued Akjoujt (19��45'N, 14��23'W), x.1937, P. Malzy��� 1 ♂ (paratype) (MNHN); NOUAKCHOOT, Nouakchoot (18��5'N, 15��59'W), 15.viii���30.ix.2006, Famille Stalmanns��� 5 ♀♀, 1 ♂ (paratypes) (PMOC); S��LIBABY, Tachoot [=Tassota Barene] (15��26'N, 12��16'W), 20.x.1941, [anonymous]��� 1 ♂ (paratype) (OMOC); NIGER: AGADEZ, Agadez (16��58'N, 7��59'E), 1909, Cortier��� 2 ♂♂ (2 paratypes) (MNHN); Agadez (16��58'N, 7��59'E), 2.viii.1947, L. Chopard & A. Villiers��� 1 ♀ (paratype) (MNHN); Agadez (16��58'N, 7��59'E), 20.viii.1947, L. Chopard & A. Villiers��� 1 ♂ (paratype) (MNHN); Agadez (16��58'N, 7��59'E), 25.viii.1947, L. Chopard & A. Villiers��� 1 ♀ (paratype) (MNHN); Agadez (16��58'N, 7��59'E), 20.viii.1989, [anonymous]��� 3 ♀♀, 1 ♂ (paratypes) (OMOC); Arlit (18��44'N, 7��23'E), 15.ix.1977, P. Tauzin��� 1 ♀ (paratype) (PMOC); Dabaga, A��r sud, 600 m (17��18'N, 8��10'E), 13���16.viii.1947, L. Chopard & A. Villiers��� 2 ♂♂ (2 paratypes) (MNHN); Taquei, A��r Massif (18��25'48''N, 8��24'36''E), 26.viii.1983, P.C. Matteson��� 1 ♂ (paratype) (BMNH); DIFFA, N���Guigmi (14��15'N, 13��7'E), viii.1919, Dr. Noel��� 7 ♀♀, 5 ♂♂ (12 paratypes) (MNHN); N���Guigmi (14��15'N, 13��7'E), ix.1919, Dr. Noel��� 12 ♀♀, 13 ♂♂ (25 paratypes) (MNHN); N���Guigmi (14��15'N, 13��7'E), vi���vii.1919, Dr. Noel��� 1 ♂ (paratype) (MNHN); N���Guigmi (14��15'N, 13��7'E), x.1919, Dr. Noel��� 1 ♂ (paratype) (MNHN); DOSSO, Toudou (14��7'N, 3��54'E), 10.viii.1989, [anonymous] ���1 ♀ (paratype) (OMOC); Toudou (14��7'N, 3��54'E), 13.viii.1989, [anonymous]��� 2 ♀♀, 1 ♂ (paratypes) (OMOC); SENEGAL: [unspecified locality], [no date], [anonymous] ���1 ♂ (paratype) (BMNH); DAKAR, Keur Ndiaye L�� (14��45'25''N, 17��14'11''W), 20.viii.2013, I. de Dinechin��� 1 ♂ (paratype) (CMD); DIOURBEL, Bambey (14��42'N, 16��28'W), viii.1939, J. Risbec��� 2 ♂♂ (2 paratypes) (BMNH); Bambey (14��42'N, 16��28'W), 1945, J. Risbec��� 1 ♂ (paratype) (BMNH); FATICK, Diouroup (14��21'N, 16��32'W), 14���18.viii.2007, P. Moretto��� 4 ♀♀, 9 ♂♂ (13 paratypes) (FGIC, PMOC); KAOLAK, Missirah, 45 m (13��59'26''N, 15��7'4''W), 16.vii.2007, F. G��nier (2007-01)��� 12 ♀♀, 9 ♂♂ (21 paratypes) (FGIC); LOUGA, Lingu��re (15��24'N, 15��7'W), ix.1967, A. Descarpentries, T. Leye & A. Villiers��� 496 specimens (paratype) (MNHN); Ndam-Dam (15��36'22''N, 16��23'35''W), xi.1967, A. Descarpentries, T. Leye & A. Villiers��� 2 ♀♀, 7 ♂♂ (9 paratypes) (MNHN); Ndilla (15��20'N, 15��3'W), ix.1967, A. Descarpentries, T. Leye & A. Villiers��� 1 ♂ (paratype) (MNHN); SAINT-LOUIS, Commune de Saint-Louis, 7 m (16��4'34''N, 16��22'40''W), 27.viii.2009, F. G��nier (2009-33)��� 1 ♀ (paratype) (FGIC); Commune de Saint-Louis, 7 m (16��4'34''N, 16��22'40''W), 27.viii.2009, P. Moretto��� 2 ♀♀, 2 ♂♂ (4 paratypes) (PMOC); Podor (16��39'N, 14��57'30''W), viii���ix.1911, R. Chudeau��� 1 ♀ (paratype) (MNHN); Richard-Toll (16��28'N, 15��41'W), 7���10.viii.2008, P. Moretto��� 1 ♀ (paratype) (PMOC); Richard-Toll (16��28'N, 15��41'W), 28���31.viii.2009, F. G��nier (2009-38)��� 8 ♀♀, 20 ♂♂ (28 paratypes) (FGIC); Richard-Toll (16��28'N, 15��41'W), 28���30.viii.2009, F. G��nier (2009-40)��� 4 ♀♀, 1 ♂ (paratypes) (FGIC); Richard- Toll (16��28'N, 15��41'W), 28.viii.2009, F. G��nier (2009-36)��� 1 ♂ (paratype) (FGIC); Richard-Toll (16��28'N, 15��41'W), 28.viii���1.ix.2009, P. Moretto & F. G��nier��� 1 ♀, 2 ♂♂ (3 paratypes) (PMOC); Richard-Toll (16��28'N, 15��41'W), 30.viii.2009, F. G��nier (2009-43)��� 2 ♀♀, 3 ♂♂ (5 paratypes) (FGIC); Richard-Toll (16��28'N, 15��41'W), 30.viii.2009, F. G��nier (2009-44)��� 3 ♀♀, 17 ♂♂ (20 paratypes) (FGIC); Richard-Toll (16��28'N, 15��41'W), 31.viii.2009, F. G��nier (2009-48)��� 6 ♂♂ (6 paratypes) (FGIC); Richard-Toll (16��28'N, 15��41'W), 1.ix.2009, F. G��nier (2009-49)��� 1 ♀, 4 ♂♂ (5 paratypes) (FGIC); Richard-Toll (16��28'N, 15��41'W), 1.ix.2009, F. G��nier (2009-50)��� 3 ♂♂ (3 paratypes) (FGIC); Richard-Toll (16��28'N, 15��41'W), 3.ix.2009, F. G��nier (2009- 52)��� 1 ♀, 1 ♂ (paratypes) (FGIC); Saint-Louis (16��0'30''N, 16��29'30''W), 1899, V. Planchat��� 13 ♀♀, 19 ♂♂ (32 paratypes) (MNHN); THI��S, Kayar (14��55'N, 17��7'E), viii.1971, A. Villiers��� 1 ♀, 1 ♂ (paratypes) (OMOC); Mbour (14��25'N, 16��58'W), viii.1995, P. Moretto��� 13 ♀♀, 8 ♂♂ (21 paratypes) (PMOC); Meckh�� (15��7'N, 16��38'W), [no date], [anonymous]��� 1 ♂ (paratype) (MNHN); Nianing, 8 m (14��19'N, 16��55'43''W), 23.vii.2006, A. Coache��� 1 ♀, 1 ♂ (paratypes) (JFJC); SUDAN: KHARTOUM, Environs de Khartoum (15��35'N, 32��32'E), [no date], [anonymous]��� 2 ♀♀, 1 ♂ (paratypes) (MNHN). Etymology. Sahelicus is a Latin adjective pertaining to the distribution of this species. Natural history. A mostly Sahelian species associated with grassland and wooded and acacia steppes habitats and grassland enclave within degraded sudanese vegetation and along dried sandy riverbeds, on deep sandy soils. Individuals are common at light traps and also collected from, donkey, cow, and human dung. Some individuals collected in pitfall traps baited with rumen content, recently killed toads and monitor lizards., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 21-24, DOI: 10.5281/zenodo.439444

Digitonthophagus fimator G��nier, new species http://zoobank.org/urn:lsid:zoobank.org:act:534 A 9261-6403-4272-93EA-24D061542886 (Figs. 19, 47���48, 67, 83, 111���112, 131, 171���173; Map 14) Type locality. For��t de Sorobouli (11��47���44���N 02��53���25���W), 270 m, Sangui��, Burkina Faso. Diagnosis. Male cephalic horns lacking granules internally at apex (Fig. 48); males with pronotal anterior angles surface flat, at most slightly produced anteriorly (Fig. 47); FLP sclerite ventral portion with a single carina (Fig. 173); SA sclerite apex interrupted much before apex of apical portion of FLP sclerite (Figs. 171���172). Description. Holotype ♂ (Fig. 19). Measurements. Length 11.5 mm, width 6.5 mm. Head (Figs. 47���48). Anterior clypeal edge slightly sinuous on median fifth and appearing arcuate in dorsal view; clypeofrontal carina broadly arcuate and interrupted at gena; lacking median tubercle, surface with punctures fine to small, scabrous anteriorly, separated by one to four diameters. Horns rather long and thick, parallel on apical half in frontal view, parallel sided on apical two-thirds, posterointernal edge unmodified basally, apicointernal surface with few, fine, scabrous punctures; genal edge slightly upturned and simply arcuate on anterior third, forming a broad angle with clypeal edge. Pronotum (Fig. 67). Surface with coarse granulate punctures present on posterior half medially and laterally, with more-or-less distinct, minute punctures throughout. Anteromedian tubercle atrophied, round in lateral view; median longitudinal sulcus rather deep but narrow; surface behind the eyes slightly concave, surface of anterior angles flat with anterior angles unmodified; anterior half of lateral edge slightly sinuous in dorsal and lateral view; posterior angles unmodified and simply arcuate in dorsal view. Anterior hypomeral ridge regularly arcuate, anterior hypomeral depression surface completely dark. Elytra (Fig. 19). Intervals 2 and 4 with fine granules on disc and apical declivity. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus (Fig. 131). Parameres with dorsal and ventral edges diverging toward apex in lateral view. Internal sac sclerites (Figs. 171���173). Axial sclerite short, angular and extending into a point apically. Subaxial sclerite, large basally and abruptly reduced, extending straight and approximately in line with apex of right lateral fold of frontolateral peripheral sclerite, with fine and scattered villi. Frontolateral peripheral sclerite basoventral apophysis moderately developed; a single large medioventral carina present, carina semicircular in ventral view; 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 sclerotized, well developed; subapicodorsal lobe membranous, not reaching anterior edge, apex set on right side in dorsal view; apical lobe truncated apically and directed obliquely on left side, apical villi regular in shape, with a small additional fold ventrally in line with subapicoventral lobe; subapicoventral lobe small, acute apically and obliquely oriented ventrally, short not reaching apical edge of apical lobe. Variation. Measurements (115 ♂♂, 106 ♀♀). Length: male 7.0���13.0 mm (10.8 �� 1.2 mm), female 9.0���12.0 mm (10.6 �� 0.7 mm). Female allotype. Cephalic outline in dorsal view as in Fig. 83; vertex with a broadly arcuate, transverse carina, dorsal edge broadly arcuate in frontal view, lateral portion obtusely angulated, sloping down posteriorly; anterior pronotal tubercles well developed, external lateral edges slightly widening toward apex in dorsal view, anterolateral surface simply convex, anterosuperior edge slightly arcuate in dorsal view (Fig. 111), lateral portion of anterosuperior edge slightly upturned (Fig. 112). Protibia short, with external teeth more robust. Primary type data. Holotype ♂ (CMNC): [BURKINA FASO: SANGUI�� / For��t de Sorobouli, 270m / 11��47���44���N 002��53���25���W / 13.VIII.2005, F. G��nier / zone soudanienne sud / pi��ge excr. Humain, 2005-14]; [COL��OPT��RES / DU / BURKINA FASO / BF015498] barcode label; [HOLOTYPE ♂ / Digitonthophagus / fimator n.sp. / des. F. G��nier, 2016] red card. MAP 14. Distribution of Digitonthophagus fimator. Material examined (415 ♂♂, 345 ♀♀, 279 sex no recorded), distribution (Map 14): BENIN: ATAKORA, Batia (10��53'N, 1��29'E), 27.vii.2009, B. Rainon��� 1 ♂ (paratype) (CMD); ATLANTIQUE, For��t de Niaouli, Attogon, 117 m (6��44'7''N, 2��8'27''E), 1.v.2013, B. Rainon��� 3 ♀♀, 2 ♂♂ (5 paratypes) (CBR); BORGOU, Alafiarou, For��t de Wari-Maro (WM2) (9��1'37''N, 2��24'13''E), 14.iv.2007, S. Tchibozo��� 1 ♂ (paratype) (PMOC); DONGA, For��t de P��n��lan, P��n��ssoulou, 392 m (9��14'50''N, 1��31'45''E), 23.v.2013, B. Rainon��� 12 ♀♀, 9 ♂♂ (21 paratypes) (CBR); Saramanga, 390 m (9��13'9''N, 1��46'29''E), 24.v.2013, B. Rainon��� 1 ♂ (paratype) (CBR); MONO, For��t de Hou��yogb��, Hou��yogb��, 82 m (6��33'43''N, 1��51'2''E), 8.v.2013, B. Rainon���2 ♀♀, 2 ♂♂ (4 paratypes) (CBR); Lokossa (6��38'N, 1��43'E), 24.iii.2006, B. Rainon���4 ♀♀, 8 ♂♂ (12 paratypes) (CBR); Lokossa (6��38'N, 1��43'E), 12.viii.2007, B. Rainon���1 ♀, 1 ♂ (paratypes) (CMD); OU��M��, environs de Porto Novo (6��30'N, 3��30'E), 1912, Waterlot��� 1 ♀, 2 ♂♂ (3 paratypes) (MNHN); ZOU, Quartier Soglogon, Bohicon, 200 m (7��10'54''N, 2��1'54''E), 14.iv.2012, B. Rainon��� 3 ♀♀, 4 ♂♂ (7 paratypes) (CBR); Quartier Soglogon, Bohicon, 200 m (7��10'54''N, 2��1'54''E), 18.iv.2012, B. Rainon���2 ♀♀, 1 ♂ (paratypes) (CBR); BURKINA FASO: CENTRE, Ouagadougou (12��22'N, 1��31'W), viii.1977, P. Ardoin��� 2 ♀♀, 8 ♂♂ (10 paratypes) (FETC); CENTRE- OUEST, For��t de Sorobouli [= For��t Class��e de Baporo], 270 m (11��47'44''N, 2��53'25''W), 10���15.viii.2005, P. Moretto���4 ♀♀, 2 ♂♂ (6 paratypes) (FETC, PMOC); CENTRE-SUD, R��serve de Nazinga (11��9'N, 1��36'W), 9��� 12.vi.2005, S. Rojkoff��� 17 ♀♀, 16 ♂♂ (33 paratypes) (PCSR, PMOC); BAL��, Boromo, 250 m (11��45'6''N, 2��51'58''W), 10.viii.2005, F. G��nier (2005-10)���1 ♂ (paratype) (FGIC); COMO��, For��t de Boulon (savane bois��e), 270 m (10��16'27''N, 4��27'15''W), 5.vii.2006, F. & S. G��nier (2006-02)���1 ♂ (paratype) (FGIC); For��t de Boulon (savane bois��e), 270 m (10��16'27''N, 4��27'15''W), 6.vii.2006, F. & S. G��nier (2006-05)��� 1 ♂ (paratype) (FGIC); For��t de Boulon (savane bois��e), 270 m (10��16'27''N, 4��27'15''W), 9.vii.2006, F. & S. G��nier (2006-17)��� 3 ♂♂ (3 paratypes) (FGIC); Kofland�� (Village), 290 m (10��14'42''N, 4��27'50''W), 4.vii.2006, F. & S. G��nier (2006- 01)��� 4 ♂♂ (4 paratypes) (FGIC); NAHOURI, For��t de Nazinga, 275 m (11��9'24''N, 1��36'44''W), 20.vii.2006, F. & S. G��nier (2006-46)��� 8 ♀♀, 9 ♂♂ (17 paratypes) (FGIC); For��t de Nazinga, Akwazena, 275 m (11��9'24''N, 1��36'44''W), 21.vii.2006, F. & S. G��nier (2006-53)���1 ♂ (paratype) (FGIC); For��t de Nazinga, Akwazena, 275 m (11��9'24''N, 1��36'44''W), 26.vii.2006, F. & S. G��nier (2006-73)��� 1 ♀, 2 ♂♂ (3 paratypes) (FGIC); For��t de Nazinga, Akwazena, 275 m (11��9'24''N, 1��36'44''W), 26.vii.2006, F. & S. G��nier (2006-75)��� 2 ♀♀, 3 ♂♂ (5 paratypes) (FGIC); For��t de Nazinga, Barka, 265 m (11��8'30''N, 1��36'35''W), 22.vii.2006, F. & S. G��nier (2006- 57)��� 7 ♀♀, 10 ♂♂ (17 paratypes) (FGIC); For��t de Nazinga, Barka, 265 m (11��8'30''N, 1��36'35''W), 24.vii.2006, F. & S. G��nier (2006-62)���1 ♂ (paratype) (FGIC); For��t de Nazinga, Boulieselo, 310 m (11��11'50''N, 1��35'9''W), 27.vii.2006, F. & S. G��nier (2006-82)��� 2 ♂♂ (2 paratypes) (FGIC); For��t de Nazinga, Kalie Boulou, 275 m (11��11'29''N, 1��30'26''W), 22.vii.2006, F. & S. G��nier (2006-54)���1 ♂ (paratype) (FGIC); For��t de Nazinga, Kalie Boulou, 275 m (11��11'29''N, 1��30'26''W), 25.vii.2006, F. & S. G��nier (2006-67)��� 1 ♀ (paratype) (FGIC); For��t de Nazinga, Kouzougou, 285 m (11��9'17''N, 1��32'10''W), 21.vii.2006, F. & S. G��nier (2006-47)��� 8 ♀♀, 2 ♂♂ (10 paratypes) (FGIC); For��t de Nazinga, Naguio, 270 m (11��7'52''N, 1��34'38''W), 24.vii.2006, F. & S. G��nier (2006- 66)���1 ♂ (paratype) (FGIC); SANGUI��, For��t de Sorobouli, 270 m (11��47'44''N, 2��53'25''W), 13.viii.2005, F. G��nier (2005-14)��� 1 ♀, 1 ♂ (holotype, allotype) (CMNC); For��t de Sorobouli, 270 m (11��47'44''N, 2��53'25''W), 14.viii.2005, F. G��nier (2005-18)��� 2 ♀♀, 1 ♂ (paratypes) (FGIC); TAPOA, Kaabougou, 280 m (11��57'22''N, 2��0'40''E), 12.vii.2006, F. & S. G��nier (2006-30)���1 ♂ (paratype) (FGIC); Kaabougou, 280 m (11��57'22''N, 2��0'40''E), 12.vii.2006, F. & S. G��nier (2006-31)��� 1 ♂ (paratype) (FGIC); CAMEROON: Yaound��-Bertoua, v.2003, M. Ondoa��� 55 ♀♀, 60 ♂♂ (115 paratypes) (CMD, PMOC); ADAMAWA, Ngaound��r�� (7��19'N, 13��35'E), iii���v.2003, M. Dongmo��� 1 ♂ (paratype) (CMD); Ranch de Ngaoundaba (7��8'N, 13��41'45''E), iv.1976, J.L. & A. Nicolas��� 2 ♀♀, 3 ♂♂ (5 paratypes) (CMD); CENTRAL, Nanga-Eboko (4��41'N, 12��22'E), 1960, Dr. Lenczy��� 1 ♀, 1 ♂ (paratypes) (CNC); Nkolbisson, Yaound�� (3��52'16''N, 11��27'14''E), 18.x.1973, B. de Mir����� 1 ♀, 2 ♂♂ (3 paratypes) (MNHN); Nkolbisson, Yaound�� (3��52'16''N, 11��27'14''E), 24.iv.1979, J. Carayon��� 1 ♂ (paratype) (MNHN); Soa (3��59'N, 11��36'E), i���iii.2003, M. Ondoa��� 1 ♀, 1 ♂ (paratypes) (CMD); EAST, Batouri (4��26'N, 14��22'E), 1939, P. Lepesme, R. Paulian, & A. Villiers��� 18 ♀♀, 8 ♂♂ (26 paratypes) (FGIC, MNHN); EXTREME NORTH, Waza (11��24'N, 14��34'E), v.2008, D. Moore��� 2 ♀♀, 2 ♂♂ (4 paratypes) (PMOC); NORTHWEST, Bankim (6��2'25.8''N, 10��16'25.32''E), 8���13.vi.2009, P. LeGall��� 11 ♀♀, 8 ♂♂ (19 paratypes) (PMOC); Santa, Monts Bamboutos (5��48'N, 10��10'E), 26.v.1967, [anonymous]��� 1 ♂ (paratype) (MNHN); WEST, Foumbot (5��30'N, 10��38'E), iv.1966, B. de Mir����� 1 ♂ (paratype) (MNHN); CENTRAL AFRICAN REPUBLIC: KEMO, Sibut (5��44'N, 19��5'E), [no date], [anonymous]��� 1 ♂ (paratype) (IRSNB); LOBAYE, Boukoko (3��54'N, 17��55'E), iv.1964, R. Pujol��� 142 specimens (paratype) (MNHN); Boukoko (3��54'N, 17��55'E), 5.vi.1965, M. Boulard��� 40 specimens (paratype) (MNHN); Kapou (4��2'28''N, 18��19'28''E), 2.v.1970, M. Boulard��� 1 ♀, 2 ♂♂ (3 paratypes) (MNHN); Station de La Mabok�� (3��53'N, 17��58'E), 20.vi.1965, [anonymous]��� 1 ♀ (paratype) (MNHN); Station de La Mabok�� (3��53'N, 17��58'E), 23.ix.1965, R. Pujol��� 1 ♀ (paratype) (MNHN); Station de La Mabok�� (3��53'N, 17��58'E), 24.ix.1965, R. Pujol��� 2 ♀♀, 2 ♂♂ (4 paratypes) (MNHN); Station de La Mabok�� (3��53'N, 17��58'E), 31.xii.1965, [anonymous]��� 2 ♀♀ (2 paratypes) (MNHN); Station de La Mabok�� (3��53'N, 17��58'E), 22.iii.1966, R. Pujol��� 1 ♂ (paratype) (MNHN); Station de La Mabok�� (3��53'N, 17��58'E), 27.iv.1968, R. Pujol��� 97 specimens (paratypes) (MNHN); Station de La Mabok�� (3��53'N, 17��58'E), 19.ii.1969, P. Teocchi��� 2 ♀♀, 2 ♂♂ (4 paratypes) (MNHN); MAMBERE-KADEI, Berb��rati (4��15'41''N, 15��47'22''E), 10.iii.1996, Poirier & Ducrocq��� 2 ♀♀, 6 ♂♂ (8 paratypes) (PMOC); OMBELLA-M���POKO, Bozo (5��8'N, 18��29'E), ix.1975, L. Herv����� 1 ♀, 3 ♂♂ (4 paratypes) (PMOC); CHAD: LOGONE ORIENTAL, Kom��, 80 km Moundou (8��30'N, 16��41'E), vii.2002, T. Garnier��� 2 ♀♀, 2 ♂♂ (4 paratypes) (PMOC); C��TE D���IVOIRE: BAFING, Ranch de Dolla, Bi��masso, 450 m (8��4'N, 7��33'W), iii���iv.2002, P. Moretto��� 43 ♀♀, 36 ♂♂ (79 paratypes) (FGIC, PMOC); Ranch de Dolla, Bi��masso, 450 m (8��4'N, 7��33'W), 1���5.vi.2000, P. Moretto��� 6 ♀♀, 6 ♂♂ (12 paratypes) (PMOC); DENGUELE, hotel des fronti��res, Odienn��, 416 m (9��30'20.3''N, 7��34'23.6''W), 9���10.vii.2013, P. Moretto��� 2 ♂♂ (2 paratypes) (PMOC); LACS, Bringakro near (6��25'53''N, 5��4'32''W), 13.vi.2003, Newman et al.��� 1 ♂ (paratype) (BMNH); Bringakro near (6��25'53''N, 5��4'32''W), 16���17.vi.2003, Newman et al.��� 1 ♂ (paratype) (BMNH); Bringakro near (6��25'53''N, 5��4'32''W), 19.vi.2003, Newman et al.��� 1 ♂ (paratype) (BMNH); LAGUNES, For��t du Banco (5��23'N, 4��3'W), ii.1989, [anonymous]��� 2 ♀♀, 1 ♂ (paratypes) (CEMT); Lamto (6��13'25''N, 5��1'36''W), 10.iv.1964, [anonymous]��� 2 ♂♂ (2 paratypes) (OMOC); Lamto (6��13'25''N, 5��1'36''W), 20.ii���28.iii.1968, C. Girard��� 1 ♀, 1 ♂ (paratypes) (MNHN); Lamto (6��13'25''N, 5��1'36''W), 20���21.iii.1968, C. Girard��� 1 ♀ (paratype) (MNHN); Lamto (6��13'25''N, 5��1'36''W), 20���24.iii.1968, C. Girard��� 1 ♀, 4 ♂♂ (5 paratypes) (MNHN); MARAHOUE, Parque National Marahou�� (7��5'N, 6��2'W), 17.v.1975, G. & V. Halffter��� 1 ♀, 1 ♂ (paratypes) (CEMT); NZI-COMOE, Dimbokro (6��39'N, 4��42'W), [no date], [anonymous]��� 1 ♀, 4 ♂♂ (5 paratypes) (IRSNB); SAVANES, M���banto, Boundiali, 395 m (9��34'10.4''N, 6��40'56.5''W), 6���8.vii.2013, P. Moretto���1 ♂ (paratype) (PMOC); M���banto, Boundiali, 395 m (9��34'10.4''N, 6��40'56.5''W), 13���15.vii.2013, P. Moretto��� 1 ♀, 7 ♂♂ (8 paratypes) (PMOC); VALL��E DU BANDAMA, Bouak�� (7��41'N, 5��2'W), 5.iv.1974, E. & I. Munroe��� 1 ♀, 1 ♂ (paratypes) (CMNC); Bouak�� (7��41'N, 5��2'W), 7���28.v.1980, O. Kukal��� 3 ♀♀, 2 ♂♂ (5 paratypes) (CMNC); Bouak�� (7��41'N, 5��2'W), 8���17.v.1980, O. Kukal��� 1 ♀ (paratype) (CMNC); DEMOCRATIC REPUBLIC OF THE CONGO: EQUATEUR, Libenge (3��39'N, 18��38'E), 14.xi.1947, R. Cremer & M. Neuman��� 1 ♂ (paratype) (IRSNB); Savane Liki-Bembe, Libenge (3��39'N, 18��38'E), iii.1948, R. Cremer & M. Neuman��� 1 ♀, 1 ♂ (paratypes) (IRSNB); Savane Liki-Bembe, Libenge (3��39'N, 18��38'E), iii.1948, R. Cremer & M. Neuman��� 1 ♂ (paratype) (IRSNB); ORIENTALE, Bambesa (3��28'N, 25��42'E), 13.ii.1940, J. Vrydagh��� 1 ♂ (paratype) (IRSNB); Cellule II, Parc National de la Garamba (3��53'N, 29��24'E), 28.ii.1951, H. de Saeger (1391)��� 1 ♀, 2 ♂♂ (3 paratypes) (IRSNB); Cellule II, Parc National de la Garamba (3��53'N, 29��24'E), 5.iii.1951, H. de Saeger (1402)��� 2 ♀♀, 3 ♂♂ (5 paratypes) (IRSNB); Cellule II, Parc National de la Garamba (3��53'N, 29��24'E), 5.iii.1951, H. de Saeger (1405)��� 1 ♀, 2 ♂♂ (3 paratypes) (IRSNB); Cellule II, Parc National de la Garamba (3��53'N, 29��24'E), 20.iii.1951, H. de Saeger (1406)��� 1 ♂ (paratype) (IRSNB); Cellule II, Parc National de la Garamba (3��53'N, 29��24'E), 20.iii.1951, H. de Saeger (1499)��� 1 ♂ (paratype) (IRSNB); Cellule II, Parc National de la Garamba (3��53'N, 29��24'E), 7.iv.1952, H. de Saeger (3530)��� 2 ♀♀, 1 ♂ (paratypes) (IRSNB); Faradje (3��44'N, 29��42'E), 13.iv.1930, A. Collart��� 2 ♀♀, 3 ♂♂ (5 paratypes) (IRSNB); For��t de Kawa (1��33'N, 30��32'E), 15.iv.1929, A. Collart��� 1 ♂ (paratype) (IRSNB); For��t de Kawa (1��33'N, 30��32'E), 20.iv.1929, A. Collart��� 1 ♀, 3 ♂♂ (4 paratypes) (IRSNB); ETHIOPIA: OROMIA, 12 km N Ch���ida (7��15'25''N, 36��48'5''E), 15.v.2013, I. Martinu��� 1 ♀, 2 ♂♂ (3 paratypes) (JFJC); Gimbi (9��10'N, 35��50'E), 26.vii.2002, Werner��� 1 ♂ (paratype) (CMD); SOUTHERN NATIONS, NATIONALITIES, AND PEOPLE���S REGION, Arba Minch (6��2'N, 37��33'E), 11��� 17.ix.2000, P. L��onard��� 3 ♂♂ (3 paratypes) (FGIC, PMOC); Dila (6��24'30''N, 38��18'30''E), ix.2000, P. L��onard��� 1 ♂ (paratype) (PMOC); GHANA: EASTERN, Hwapa (5��56'N, 0��15'E), 3.iv.1930, [anonymous]��� 1 ♂ (paratype) (BMNH); GREATER ACCRA, Accra (5��36'N, 0��11'W), ii���iii.1989, Strada��� 2 ♂♂ (2 paratypes) (FETC); north end, Shai Hills, 125 m (5��56.376'N, 0��3.406'E), 23���25.vi.2006, Bell, Davis, Paemka & Philips���1 ♀, 2 ♂♂ (3 paratypes) (PMOC); GUINEA: KINDIA, Kindia (10��4'N, 12��51'W), 20.v.1983, S. Murzin��� 1 ♀, 1 ♂ (paratypes) (FETC); Kindia (10��4'N, 12��51'W), 23.vi.1983, [anonymous]��� 1 ♂ (paratype) (OUMNH); NZ��R��KOR��, Thio, Mont Nimba (7��35'10''N, 8��30'7''W), ii���vi.1942, M. Lamotte��� 2 ♀♀, 3 ♂♂ (5 paratypes) (IRSNB); GUINEA- BISSAU: BAFAT��, Chime [= Xime], Rio G��ba (11��58'N, 14��56'W), 1906, G. Favarel��� 1 ♀, 2 ♂♂ (3 paratypes) (IRSNB, MNHN); NIGERIA: ABIA, Umuahia (5��32'N, 7��29'E), 12.vii���2.ix.1960, J.L. Gregory��� 1 ♂ (paratype) (BMNH); Umudike (5��29'N, 7��33'E), 23���31.iii.1960, J.L. Gregory��� 8 ♀♀, 6 ♂♂ (14 paratypes) (BMNH); ADAMAWA, Zungera (9��59'N, 12��28'E), iv.1911, J.W. Scott-Macfie��� 1 ♀, 1 ♂ (paratypes) (BMNH); BAUCHI, Lame Burra Game Reserve, 630 m (11��4'57''N, 8��37'26''E), 21���25.vii.2006, P. L��onard & P. Vingerhoedt��� 3 ♀♀, 5 ♂♂ (8 paratypes) (PMOC); [unspecified loca, Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 61-65, DOI: 10.5281/zenodo.439444, {"references":["Moretto, P. & Bordat, P. (2007) Les scarabeides coprophages du Parc National du Niokolo-Koba au Senegal (Coleoptera, Scarabaeoidea). Nouvelle Revue d'Entomologie (N. S.), 23, 115 - 145."]}

Digitonthophagus gazella species group 7. Digitonthophagus biflagellatus G��nier, new species Distribution: Burundi, D.R. Congo, Kenya, Rwanda, Tanzania, Uganda, Zambia 8. Digitonthophagus gazella (Fabricius, 1787), conservation of name in prevailing usage Scarabaeus dorcas Olivier, 1789, proposed new synonymy Distribution: Angola, Botswana. Democratic Republic of the Congo, Kenya, Malawi, Mozambique, Namibia, South Africa, Swaziland, Tanzania, Uganda, Zambia, Zimbabwe; Introduced worldwide in tropical, subtropical, and southern temperate pastures (including the Comoros Islands, Mayotte, and Madagascar) 9. Digitonthophagus petilus G��nier, new species Distribution: Ethiopia, Kenya, South Sudan, Sudan, Uganda 10. Digitonthophagus viridicollis G��nier, new species Distribution: Angola, Congo, Democratic Republic of the Congo, Namibia, Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on page 74, DOI: 10.5281/zenodo.439444, {"references":["Olivier, A. G. (1789) Entomologie, ou histoire naturelle des insectes, avec leurs caracteres generiques et specifiques, leur descriptions, leur synonymie, et leur figure enluminee. Coleopteres. Tome premier. Beaudoin, Paris, France, xx + 497 pp. https: // doi. org / 10.5962 / bhl. title. 61905"]}

Digitonthophagus petilus G��nier, new species http://zoobank.org/urn:lsid:zoobank.org:act:4B8CB6CC-46E9-49CC-9 A 3D-5C8C74 A 4F878 (Figs. 15, 39���40, 63, 79, 103���104, 127, 159���161; Map 10) Type locality. El Wak, North Eastern Province, Kenya. Diagnosis. Pronotum rather opaque (Fig. 63); FLP sclerite subapicodorsal lobe strongly produced anteriorly (Figs. 159���160); right lateral fold produced into a longitudinal spoon-shape process with apical portion open ventrally, ventral edge with a brush of minute villi, lacking well-sclerotized projection on apicoventral edge (Fig. 160); axial sclerite shorter than subaxial sclerite, acute apically (Figs. 159���160); subaxial sclerite long and thin (Fig. 160). Description. Holotype ♂ (Fig. 15). Measurements. Length 11.5 mm, width 6.0 mm. Head (Figs. 39���40). 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. Horns rather short, divergent from base to apex in frontal view, gradually tapering from base to apex; posterointernal edge with a low angular projection; apicointernal surface lacking granules, with few feebly scabrous punctures; genal edge slightly upturned and feebly angulate on anterior third, forming a broad angle with clypeal edge. Pronotum (Fig. 63). Surface with granulate punctures restricted to anterior half, with moderately large punctures on posterior half of disc; punctures smaller posterolaterally, with minute punctures distinct and rather dense throughout. Anteromedian tubercle atrophied, simply round in lateral view; median longitudinal sulcus narrow and shallow; surface behind the eyes with a simple round shallow depression, surface of anterior angles convex; 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. 15). Intervals 2 and 4 with few scattered, fine granules on apical declivity. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus (Fig. 127). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites (Figs. 159���161). Axial sclerite broadly arcuate, with apex acute. Subaxial sclerite, gradually tapering from base to mid-distance, fine and straight on apical half, extending straight beyond apex of right lateral fold of frontolateral peripheral sclerite apical portion, with villi on apical third. Frontolateral peripheral sclerite basoventral apophysis well developed; lacking medioventral carinae; right lateral fold produced into longitudinal spoon-shape process with apical portion open internally, ventral edge bordered with a brush of villi; left lateral lobe membranous, slightly developed; subapicodorsal lobe membranous, reaching anterior edge, apex set on right side in dorsal view; apical lobe round and directed obliquely on left side and bent ventrally, left edge emarginate, apical villi similarly shaped throughout; subapicoventral lobe apical portion acute and in line with apical lobe apically, with distinct villi on apical third. Variation. Measurements (50 ♂♂, 40 ♀♀). Length: male 8.0���12.0 mm (10.1 �� 1.0 mm), female 8.0���11.0 mm (9.7 �� 0.7 mm). Female allotype. Cephalic outline in dorsal view as in Fig. 79; vertex with a straight, transverse carina, dorsal edge broadly arcuate in frontal view, lateral portion gradually sloping down; anterior pronotal tubercles atrophied, external lateral edges parallel sided, anterolateral surface simply convex, anterosuperior edge slightly arcuate in dorsal view (Fig, 103), lateral portion of anterosuperior edge slightly upturned (Fig. 104). Protibia short, with external teeth more robust. Primary type data. Holotype ♂ (CMNC): [KENYA: NE PROV. / El Wak / 1-3.V.2001 / Werner & Smrz leg.]; [WORLD / SCARAB. / DATABASE / WSD00029750] barcode label; [HOLOTYPE ♂ / Digitonthophagus / petilus n.sp. / des. F. G��nier, 2016] red card. MAP 10. Distribution of Digitonthophagus petilus. Material examined (69 ♂♂, 63 ♀♀), distribution (Map 10): ETHIOPIA: OROMIA, Yabelo (4��53'N, 38��5'E), iv.1994, Werner��� 1 ♂ (paratype) (CMD); SOUTHERN NATIONS, NATIONALITIES, AND PEOPLE���S REGION, Arba Minch (6��2'N, 37��33'E), 11���17.ix.2000, P. L��onard��� 23 ♀♀, 22 ♂♂ (45 paratypes) (FGIC, PMOC); Dila (6��24'30''N, 38��18'30''E), ix.2000, P. L��onard��� 1 ♂ (paratype) (PMOC); KENYA: Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 18.x.1973, T.J. Kingston��� 1 ♀, 3 ♂♂ (4 paratypes) (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 9.xi.1973, T.J. Kingston��� 1 ♀, 1 ♂ (paratypes) (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 13.xi.1973, T.J. Kingston��� 1 ♀ (paratype) (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 18.xi.1973, T.J. Kingston��� 1 ♀ (paratype) (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 21.xi.1973, T.J. Kingston��� 3 ♀♀, 1 ♂ (paratypes) (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 10.xii.1973, T.J. Kingston��� 1 ♀ (paratype) (OUMNH); Tsavo National Park [=Tsavo East National Park] (2��45'S, 38��49'E), 15.xii.1973, T.J. Kingston��� 1 ♂ (paratype) (OUMNH); BARINGO, Loiminange, 980 m (0��31'N, 36��7'E), 24.x.1988, M.N. Mungal & R.K. Butlin��� 6 ♀♀, 5 ♂♂ (11 paratypes) (OUMNH); ISIOLO, 10 km N Isiolo (0��27'14''N, 37��36'36''E), 16. xii.1 990, B.D. Gill��� 7 ♀♀, 11 ♂♂ (18 paratypes) (BDGC); MANDERA, El Wak (2��49'N, 40��56'E), 1���3.v.2001, Werner & Smrz��� 10 ♀♀, 9 ♂♂ (holotype, allotype, 17 paratypes) (CMD, CMNC); RIFT VALLEY, Samburu National Reserve (1��13'N, 36��57'E), 4.xii.1989, Moreno-Mestre��� 1 ♀, 3 ♂♂ (4 paratypes) (PMOC); TAITA TAVETA, Tsavo, 50 km N Voi (3��0'S, 38��27'E), 25.xii.1990, B.D. Gill��� 1 ♀, 4 ♂♂ (5 paratypes) (BDGC); Voi, Tsavo East National Park (3��23'S, 38��34'E), 31.xii.1990, B.D. Gill��� 3 ♀♀, 2 ♂♂ (5 paratypes) (BDGC); SOUTH SUDAN: CENTRAL EQUATORIA, Rejaf (4��45'N, 31��34'E), 29.xii.1911, [anonymous]��� 4 ♀♀, 2 ♂♂ (6 paratypes) (MNHN); SUDAN: SENNAR, Sennar (13��34'N, 33��34'E), [no date], Tritsch��� 1 ♂ (paratype) (SMF); UGANDA: CENTRAL, Entebbe (0��4'N, 32��28'E), [no date], H. Rolle��� 2 ♂♂ (2 paratypes) (IRSNB). Etymology. Petilus is a Latin adjective (frail, slight, puny) pertaining to the aspect of the apical portion of the internal sac subaxial sclerite (SA). Natural history. Some specimens collected in buffalo, cow, and elephant dung. Remarks. We first regarded D. petilus as a northern population of D. gazella. The very similar configuration of the right lateral fold and lacking the ventral sclerotized projection was nearly identical to typical D. gazella. But after careful observations of the FLP sclerite, several additional congruous characters were found; the much more developed subapicodorsal lobe and the much narrower subapicoventral lobe with its apex covered with distinct villi (Figs. 159���161) combined with the external characters such as the overall dorsum dull aspect and smaller average size are consistent. The possibility that D. petilus could be a subspecies of D. gazella needs to be tested. However, the fact that its distribution is parapatric with D. gazella seems to preclude this possibility., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 52-54, DOI: 10.5281/zenodo.439444

Digitonthophagus lusinganus species group Diagnosis. FLP sclerite ventral portion with one or two medioventral carinae (Fig. 184 A); right lateral fold produced into a rather large everted and open apically conical process with irregular apical edge (Figs. 165, 168, 171, 174, 177); FLP sclerite subapicoventral lobe acute apically (Figs. 167, 176); axial and subaxial sclerite rather short, with a distinct transverse uniting sclerotized area medially (Fig. 171, 174, 177)., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on page 57, DOI: 10.5281/zenodo.439444

Digitonthophagus lusinganus species group 11. Digitonthophagus aksumensis Génier, new species Distribution: Djibouti, Eritrea, Ethiopia, Saudi Arabia, Somalia, Sudan, Yemen 12. Digitonthophagus dilatatus Génier, new species Distribution: Burkina Faso, Cameroon, Chad, Côte d’Ivoire, Ghana, Nigeria 13. Digitonthophagus fimator Génier, new species Distribution: Benin, Burkina Faso, Cameroon, Central Africa republic, Chad, Côte d'Ivoire, Democratic Republic of the Congo, Ethiopia, Ghana, Guinea, Guinea-Bissau, Nigeria, Senegal, Sierra Leone, Togo, Uganda 14. Digitonthophagus lusinganus (d’Orbigny, 1905), valid species Distribution: Ethiopia, Kenya, Somalia, South Sudan, Sudan, Tanzania 15. Digitonthophagus ulcerosus Génier, new species Distribution: Ethiopia, Kenya, Somalia, Tanzania

Digitonthophagus namaquensis species group Diagnosis. A single species with subapicodorsal lobe of FLP sclerite set on left side (Fig. 180); right lateral fold of FLP sclerite apical portion atrophied, set behind oblique right lateral edge of apical lobe (Fig. 180); basoventral apophysis of FLP sclerite large (Fig. 182)., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on page 70, DOI: 10.5281/zenodo.439444

Digitonthophagus eucatta G��nier, new species http://zoobank.org/urn:lsid:zoobank.org:act: D66876 EF-FA95-493B-B87B-1 D88858 C8406 (Figs. 11, 31���32, 59, 75, 95���96, 123, 147���149, 187; Map 6) Type locality. Toulf�� (13��53���33���N 01��53���39���W), 325 m, Loroum, Burkina Faso. Diagnosis. Species distributed from Oman westward; left lateral apophysis of FLP sclerite large (Fig. 147). Description. Holotype ♂ (Fig. 11). Measurements. Length 11.5 mm, width 6.5 mm. Head (Figs. 31���32). 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 moderate, scabrous anteriorly, separated by less than a diameter. Horns moderately long, parallel on apical half in frontal view, parallel sided on basal four-fifths, slightly tapering externally before apex; posterointernal edge produced into a low, angular projection; apicointernal surface lacking granules, with a few small scabrous punctures; genal edge upturned and distinctly angulate on anterior third, forming a broad angle with clypeal edge. Pronotum (Fig. 59). Surface with granulate punctures restricted to anterior half medially, with smaller, irregularly-scattered, simple punctures on posterior half of disc; punctures lacking posterolaterally, with more-or-less distinct minute punctures throughout; anteromedian tubercle atrophied, simply round in lateral view, median longitudinal sulcus well defined; surface behind the eyes slightly concave, surface of anterior angles concave, anterior angles unmodified; anterior half of lateral edge arcuate in dorsal and slightly sinuous anteriorly in lateral view; posterior angles simply arcuate in dorsal view. Anterior hypomeral ridge straight, abruptly curved before anterior angle, anterior hypomeral depression surface light in color medially. Elytra (Fig. 11). Intervals 2 and 4 with few fine granules on apical declivity only. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus (Fig. 123). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites (Figs. 147���149). Axial sclerite moderately sclerotized, shorter than subaxial sclerite and almost straight. Subaxial sclerite, larger basally and abruptly tapering in line with apex of axial sclerite, extending straight and interrupted approximately at apex of frontolateral peripheral sclerite apical portion, with fine villi on apical sixth. Frontolateral peripheral sclerite basoventral apophysis moderately developed, angular in lateral view; with a single acute medioventral carinae; right lateral fold produced into a uneven bifurcate process, lateral edge straight; left lateral apophysis present, large; left lateral lobe absent; subapicodorsal lobe well sclerotized, entirely covering apical lobe, with villi much shorter than apical lobe villi, right edge obliquely truncated before apex; apical lobe truncated apically, apical villi regular in shape; subapicoventral lobe atrophied, oriented ventrally Variation. Measurements (107 ♂♂, 118 ♀♀). Length: male 7.0��� 12.5 mm (10.4 �� 1.0 mm), female 8.0��� 12.5 mm (10.6 �� 1.0 mm). Female allotype. Cephalic outline in dorsal view as in Fig. 75; vertex with a broadly arcuate transverse carina, dorsal edge straight in frontal view, lateral portion subangular and sloping down posteriorly; anterior pronotal tubercles well developed, external lateral edges rather strongly widening in dorsal view (Fig. 95), anterolateral surface deeply, obliquely concave, anterosuperior edge slightly arcuate in dorsal view, lateral portion of anterosuperior edge slightly upturned (Fig. 96). Protibia short, with external teeth more robust. Primary type data. Holotype ♂ (CMNC): [BURKINA FASO: LORUM / Toulf��, 325m / 13��53���33���N 001��53���39���O / 17.VII.2006, zone sah��lienne / steppe arbor��e, pi��ge lumineux / F. & S. G��nier, 2006-41]; [COL��OPT��RES / DU / BURKINA FASO / BF015417] barcode label; [HOLOTYPE ♂ / Digitonthophagus / eucatta n.sp. / des. F. G��nier, 2016] red card. MAP 6. Distribution of Digitonthophagus eucatta Material examined (403 ♂♂, 237 ♀♀), distribution (Map 6): BENIN: ATAKORA, Batia (10��53'N, 1��29'E), 24.vii.2009, B. Rainon��� 1 ♂ (paratype) (CMD); BURKINA FASO: CENTRE, Ouagadougou (12��22'N, 1��31'W), viii.1977, P. Ardoin��� 2 ♀♀, 7 ♂♂ (9 paratypes) (FETC); CENTRE-NORD, 19 km N Kaya, 314 m (13��8'46.2''N, 0��52'42.1''W), 16.viii.2012, P. Moretto��� 1 ♂ (paratype) (PMOC); CENTRE-OUEST, For��t de Sorobouli [= For��t Class��e de Baporo], 270 m (11��47'44''N, 2��53'25''W), 10���15.viii.2005, P. Moretto��� 31 ♀♀, 41 ♂♂ (72 paratypes) (FETC, PMOC); Koudougou (12��15'N, 2��22'W), vii.1998, J.B. Yam��ogo��� 1 ♂ (paratype) (PMOC); CENTRE- SUD, R��serve de Nazinga (11��9'N, 1��36'W), 9���12.vi.2005, S. Rojkoff��� 2 ♀♀, 1 ♂ (paratypes) (PCSR); EST, 25 km SE Pama (11��7'49''N, 0��52'38''E), 3���24.ix.1988, F. G��nier & M. Sanborne��� 2 ♀♀, 6 ♂♂ (8 paratypes) (CMNC); Kompienga, 20 km S Pama (11��4'37''N, 0��40'50''E), 1���16.vi.1988, Sanborne, Landry & Tou��� 2 ♂♂ (2 paratypes) (CMNC); Kompienga, 20 km S Pama (11��4'37''N, 0��40'50''E), 3���24.ix.1988, F. G��nier & M. Sanborne��� 2 ♀♀, 4 ♂♂ (6 paratypes) (CMNC); HAUTS-BASSINS, A��roport, Bobo-Dioulasso (11��9'47''N, 4��19'12''W), 27.ix.2012, P. Moretto��� 1 ♀, 2 ♂♂ (3 paratypes) (PMOC); Bobo-Dioulasso (11��12'N, 4��18'W), 29.iv.1953, Hamon��� 7 ♀♀, 7 ♂♂ (14 paratypes) (MNHN); Bobo-Dioulasso (11��12'N, 4��18'W), 7.v.1953, Hamon��� 1 ♀, 2 ♂♂ (3 paratypes) (MNHN); Matourkou (11��5'N, 4��22'W), 15.vi.2005, S. Rojkoff��� 3 ♂♂ (3 paratypes) (CMD, PCSR); site de Samand��ni, Bama, 330 m (11��22'24.5''N, 4��33'57''W), vii.2012, F. Rousset��� 1 ♂ (paratype) (PMOC); NORD, 8 km SW Yako, Passor��, 320 m (12��51'44''N, 2��18'35''W), 7.viii.2005, P. Moretto��� 1 ♀, 1 ♂ (paratypes) (PMOC); Toulf��, Lorum, 330 m (13��54'13''N, 1��54'19''W), 9.viii.2005, P. Moretto��� 5 ♀♀, 6 ♂♂ (11 paratypes) (PMOC); PLATEAU-CENTRAL, Piste de Bombor�� (12��15'N, 0��53'W), 11.viii.2012, P. Moretto��� 1 ♂ (paratype) (PMOC); Ziniare (12��35'N, 1��18'W), 19.vi.2005, S. Rojkoff��� 1 ♀, 4 ♂♂ (5 paratypes) (PCSR, PMOC); SAHEL, 3,7 km NW piste vers Essakane, N���djomga, Dori, 287 m (14��5'20.9''N, 0��1'36.3''W), 1��� 5.ix.2013, P. Moretto��� 2 ♂♂ (2 paratypes) (PMOC); apr��s le pont, route Essakane-Dori, Gorum-Gorum, 271 m (14��9'32.8''N, 0��1'20.4''E), 7.viii.2012, P. Moretto��� 2 ♂♂ (2 paratypes) (PMOC); Conseil r��gional, Dori, 283 m (14��2'4.5''N, 0��3'10.5''W), 30.viii.2013, P. Moretto��� 1 ♂ (paratype) (PMOC); Dori (14��2'N, 0��1'W), iv.2007, [anonymous]��� 1 ♂ (paratype) (JFJC); Dori (14��2'N, 0��1'W), 24.viii.2012, P. Moretto��� 1 ♂ (paratype) (PMOC); Essakane, Gorum-Gorum, 264 m (14��22'40.2''N, 0��5'47.7''E), 13���15.viii.2012, P. Moretto��� 1 ♂ (paratype) (PMOC); Essakane, Gorum-Gorum, 264 m (14��22'40.2''N, 0��5'47.7''E), 22���24.viii.2012, P. Moretto��� 2 ♀♀, 2 ♂♂ (4 paratypes) (PMOC); Essakane, Gorum-Gorum, 264 m (14��22'40.2''N, 0��5'47.7''E), 10���13.ix.2012, P. Moretto��� 1 ♀, 2 ♂♂ (3 paratypes) (PMOC); Essakane, Gorum-Gorum, 264 m (14��22'40.2''N, 0��5'47.7''E), 2���4.x.2012, P. Moretto��� 1 ♀, 1 ♂ (paratypes) (PMOC); N���djomga, Dori (14��3'55.7''N, 0��3'3.4''W), 24���27.viii.2013, P. Moretto��� 4 ♂♂ (4 paratypes) (PMOC); SUD-OUEST, Bondigui, 324 m (10��53'56.5''N, 3��34'15''W), v.2012, F. Rousset��� 1 ♀, 1 ♂ (paratypes) (PMOC); BAL��, Boromo, 250 m (11��45'6''N, 2��51'58''W), 10.viii.2005, F. G��nier (2005-10)��� 1 ♀, 4 ♂♂ (5 paratypes) (FGIC); GOURMA, Fada N���Gourma, 300 m (12��3'5''N, 0��21'27''E), 14.vii.2006, F. & S. G��nier (2006-35)��� 1 ♂ (paratype) (FGIC); KOMPIENGA, 15 km E Nadiagou, 155 m (11��4'31''N, 0��56'8''E), 22.viii.2005, F. G��nier (2005-40)��� 1 ♂ (paratype) (FGIC); Kompienga, 160 m (11��5'5''N, 0��42'42''E), 21.viii.2005, F. G��nier (2005-39)��� 2 ♂♂ (2 paratypes) (FGIC); Pama, 230 m (11��17'N, 0��42'59''E), 24.viii.2005, F. G��nier (2005-44)��� 2 ♂♂ (2 paratypes) (FGIC); Pama, 230 m (11��17'N, 0��42'59''E), 25.viii.2005, F. G��nier (2005-45)��� 1 ♂ (paratype) (FGIC); Pama, 230 m (11��17'N, 0��42'59''E), 26���27.viii.2005, F. G��nier (2005-49)��� 2 ♂♂ (2 paratypes) (FGIC); LOROUM, Toulf��, 330 m (13��54'16''N, 1��54'19''W), 9.viii.2005, F. G��nier (2005-08)��� 1 ♀, 2 ♂♂ (3 paratypes) (FGIC); Toulf��, 330 m (13��54'16''N, 1��54'19''W), 16.vii.2006, F. & S. G��nier (2006-38)��� 1 ♀, 3 ♂♂ (4 paratypes) (FGIC); Toulf��, 330 m (13��54'16''N, 1��54'19''W), 16.vii.2006, F. & S. G��nier (2006-40)��� 1 ♀, 1 ♂ (paratypes) (FGIC); Toulf��, 330 m (13��54'16''N, 1��54'19''W), 17.vii.2006, F. & S. G��nier (2006-41)��� 22 ♀♀, 21 ♂♂ (holotype, allotype, 41 paratypes) (CMNC, FGIC); Toulf��, 330 m (13��54'16''N, 1��54'19''W), 18.vii.2006, F. & S. G��nier (2006-42)��� 2 ♂♂ (2 paratypes) (FGIC); Toulf��, 330 m (13��54'16''N, 1��54'19''W), 18.vii.2006, F. & S. G��nier (2006-43)��� 2 ♂♂ (2 paratypes) (FGIC); NAHOURI, For��t de Nazinga, 275 m (11��9'24''N, 1��36'44''W), 20.vii.2006, F. & S. G��nier (2006-46)��� 1 ♂ (paratype) (FGIC); For��t de Nazinga, Akwazena, 275 m (11��9'24''N, 1��36'44''W), 26.vii.2006, F. & S. G��nier (2006-76)��� 3 ♂♂ (3 paratypes) (FGIC); For��t de Nazinga, Boulieselo, 310 m (11��11'50''N, 1��35'9''W), 27.vii.2006, F. & S. G��nier (2006-82)��� 2 ♂♂ (2 paratypes) (FGIC); For��t de Nazinga, Kouzougou, 285 m (11��9'17''N, 1��32'10''W), 21.vii.2006, F. & S. G��nier (2006-47)��� 1 ♀, 4 ♂♂ (5 paratypes) (FGIC); For��t de Nazinga, Naguio, 270 m (11��7'52''N, 1��34'38''W), 24.vii.2006, F. & S. G��nier (2006-66)��� 1 ♂ (paratype) (FGIC); Tiakan��, 320 m (11��11'37''N, 1��14'10''W), 20.viii.2005, F. G��nier (2005-29)��� 2 ♀♀ (2 paratypes) (FGIC); PASSOR��, 8 km SW Yako, 320 m (12��51'44''N, 2��18'35''W), 7.viii.2005, F. G��nier (2005-03)��� 4 ♀♀, 2 ♂♂ (6 paratypes) (FGIC); 8 km S.O. Yako, 320 m (12��51'44''N, 2��18'35''W), 7.viii.2005, F. G��nier (2005-04)���1 ♀ (paratype) (FGIC); 8 km S.O. Yako, 320 m (12��51'44''N, 2��18'35''W), 7.viii.2005, F. G��nier (2005-02)���7 ♂♂ (7 paratypes) (FGIC); SANGUI��, For��t de Sorobouli, 270 m (11��47'44''N, 2��53'25''W), 13.viii.2005, F. G��nier (2005-14)��� 1 ♀, 1 ♂ (paratypes) (FGIC); For��t de Sorobouli, 270 m (11��47'44''N, 2��53'25''W), 14.viii.2005, F. G��nier (2005-18)��� 2 ♂♂ (2 paratypes) (FGIC); For��t de Sorobouli, 270 m (11��47'44''N, 2��53'25''W), 15.viii.2005, F. G��nier (2005-19)��� 2 ♀♀, 2 ♂♂ (4 paratypes) (FGIC); CAMEROON: EXTREME NORTH, Gu��tal��, Koza (10��53'N, 13��54'E), 13.vi.1967, [anonymous]��� 1 ♀ (paratype) (MNHN); Maroua (10��35'N, 14��19'E), vii.1989, D. Bernaud��� 3 ♀♀, 5 ♂♂ (8 paratypes) (CMD); Maroua (10��35'N, 14��19'E), vii.1979, G. Onore��� 2 ♀♀, 2 ♂♂ (4 paratypes) (PHWC); Maroua (10��35'N, 14��19'E), 5.vii.1978, [anonymous]��� 1 ♂ (paratype) (MNHN); Waza (11��24'N, 14��34'E), 18.viii.2006, C. Vanderbergh��� 4 ♀♀, 2 ♂♂ (6 paratypes) (PMOC); Waza National Park (11��15'N, 14��39'E), 18.viii.2006, C. Vanderbergh��� 3 ♀♀, 4 ♂♂ (7 paratypes) (PMOC); NORTH, Garoua (9��18'N, 13��24'E), x.1973, B. de Mir����� 1 ♀ (paratype) (MNHN); CHAD: HADJER-LAMIS, Sitje, rive sud Lac Tchad (12��53'N, 14��52'E), 11.viii.2006, C. Vanderbergh��� 1 ♂ (paratype) (PMOC); LOGONE ORIENTAL, Kom��, 80 km Moundou (8��30'N, 16��41'E), vii.2002, T. Garnier��� 2 ♂♂ (2 paratypes) (PMOC); N���DJAMENA, Farcha (12��7'N, 14��59'E), 28.vi.1967, [anonymous]��� 4 ♀♀, 8 ♂♂ (12 paratypes) (MNHN); Fort Lamy [= N���Djamena _] (12��7'N, 15��3'E), [no date], M. Andr����� 3 ♀♀, 5 ♂♂ (8 paratypes) (PMOC); OUADAI, Massif de Kabla (13��25'N, 21��9'W), 5.viii.1949, P. de Mir����� 1 ♂ (paratype) (MNHN); ERITREA: ANSEBA, Hagaz (15��42'N, 38��16'E), vii.2003, M. Forti��� 1 ♀, 1 ♂ (paratypes) (PMOC); GASH-BARKA, 18 km E Akordat (15��33'N, 38��1'E), vii.2003, M. Forti��� 1 ♀, 1 ♂ (paratypes) (PMOC); Tesseney, 1200 m (15��6'36''N, 36��39'27''E), 30.viii.2001, L. & M. Stalmans��� 2 ♀♀, 4 ♂♂ (6 paratypes) (CADB, PMOC); NORTHERN RED SEA, Nefasit (15��20'N, 39��4'E), vii.2002, Czeppel & Forti��� 2 ♀♀, 5 ♂♂ (7 paratypes) (FETC); ETHIOPIA: AFAR, Yangudi Rassa National Park (10��33'N, 40��59'E), 9/2/2006, S. Rojkoff��� 1 ♀, 1 ♂ (paratypes) (CMD); GAMBIA: CENTRAL RIVER, Lower Saloum District, Ballangharr (13��39'N, 15��25'W), 23.viii.1997, D.J. Mann��� 2 ♀♀, 5 ♂♂ (7 paratypes) (OUMNH); MALI: KIDAL, Adrar des Ifoghas (19��32'N, 1��35'E), viii.1941, Volkonsky��� 2 ♂♂ (2 paratypes) (MNHN); Etambar, Kidal (18��26'N, 1��25'E), ix.1941, M. Volkoneky��� 1 ♀ (paratype) (MNHN); SEGOU, [Station centrale de] Kogoni (14��44'N, 6��2'W), 19.ix.1955, J.-G. Pointel��� 2 ♀♀, 1 ♂ (paratypes) (MNHN); [Station centrale de] Kogoni (14��44'N, 6��2'W), 7.x.1955, J.-G. Pointel��� 2 ♀♀ (2 paratypes) (MNHN); [Station centrale de] Kogoni (14��44'N, 6��2'W), 11.x.1955, J.-G. Pointel��� 1 ♂ (paratype) (MNHN); [Station centrale de] Kogoni (14��44'N, 6��2'W), 24.x.1955, J.-G. Pointel��� 1 ♀, 1 ♂ (paratypes) (MNHN); [Station centrale de] Kogoni (14��44'N, 6��2'W), 27.x.1955, J.-G. Pointel��� 1 ♂ (paratype) (MNHN); [Station centrale de] Kogoni (14��44'N, 6��2'W), 2.xi.1955, J.-G. Pointel��� 1 ♀, 2 ♂♂ (3 paratypes) (MNHN); [Station centrale de] Kogoni (14��44'N, 6��2'W), 21.ix.1959, J.-G. Pointel��� 1 ♂ (paratype) (MNHN); MAURITANIA: GUIDIMAKA, [unspecified locality] (15��22'N, 12��6'W), 1908, G. Audan��� 4 ♀♀, 7 ♂♂ (11 paratypes) (FGIC, MNHN); HODH EL GHARBI, Tamchekett (17��14'44''N, 10��40'11''W), x.1947, B. de Mir����� 1 ♂ (paratype) (MNHN); S��LIBABY, Tachoot [= Tassota Barene] (15��26'N, 12��16'W), 20.x.1941, [anonymous]��� 1 ♀ (paratype) (OMOC); NIGER: AGADEZ, Agadez (16��58'N, 7��59'E), 20.viii.1989, [anonymous]��� 1 ♀, 1 ♂ (paratypes) (OMOC); DOSSO, Toudou (14��7'N, 3��54'E), 10.viii.1989, [anonymous]��� 1 ♀, 1 ♂ (paratypes) (OMOC); MARADI, Maradi (13��29'N, 7��6'E), [no date], J. Roggeman��� 2 ♂♂ (2 paratypes) (BMNH); TAHOUA, Entre Madaoua et Bouza (14��15'N, 6��0'E), 3.vii.1980, P. Leblanc��� 2 ♂♂ (2 paratypes) (FETC); TILLABERI, Koure (13��20'N, 2��35'E), 19���21.ix.2003, Josso, Juhel & Monfort��� 1 ♂ (paratype) (JFJC); NIGERIA: KANO, [unspecified locality] (11��59'47''N, 8��31'0''E), v.1952, W.E. S. Merrett��� 4 ♂♂ (4 paratypes) (BMNH); [unspecified locality] (11��59'47''N, 8��31'0''E), vii.1954, W.E.S. Merrett���2 ♂♂ (2 paratypes) (BMNH); [unspecified locality] (11��59'47''N, 8��31'0''E), v.1953, W.E.S. Merrett���4 ♂♂ (4 paratypes) (BMNH); [unspecified locality] (11��59'47''N, 8��31'0''E), vi.1953, W.E.S. Merrett���5 ♂♂ (5 paratypes) (BMNH); [unspecified locality] (11��59'47''N, 8��31'0''E), vi.1954, W.E. S. Merrett��� 1 ♂ (paratype) (BMNH); OMAN: MINTAQAT MASQAT, Mascat (23��36'48''N, 58��35'36''E), ix���x, Maindron��� 2 ♂♂ (2 paratypes) (BMNH); SENEGAL: DAKAR, Cit�� Gendarmerie, Keur Massar (14��45'51''N, 17��16'45''W), 29.viii.2012, I. de Dinechin��� 1 ♂ (paratype) (CMD); Dakar (14��46'N, 17��22'W), 1914, Chissadon��� 3 ♀♀, 4 ♂♂ (7 paratypes) (MNHN); Dakar (14��46'N, 17��22'W), 1902, Waterlot��� 1 ♂ (paratype) (MNHN); Entr��e du parc Marine, Keur Massar, Dakar (14��47'11''N, 17��18'43''W), viii.2015, I. de Dinechin��� 1 ♀, 1 ♂ (paratypes) (CMD); Keur Ndiaye L�� (14��45'25''N, 17��14'11''W), ix.2013, Local collectors��� 1 ♂ (paratype) (CMD); Keur Ndiaye L�� (14��45'25''N, 17��14'11''W), 18.viii.2013, I. de Dinechin��� 1 ♂ (paratype) (CMD); Sebikotane (14��45'N, 17��8'W), viii.1971, A. Villiers��� 1 ♂ (paratype) (OMOC); DIOURBEL, Bambey (14��42'N, 16��28'W), 1945, J. Risbec��� 4 ♀♀, 5 ♂♂ (9 paratypes) (BMNH); Bambey (14��42'N, 16��28'W), viii.1939, J. Risbec��� 3 ♂♂ (3 paratypes) (BMNH); FATICK, Diouroup (14��21'N, 16��32'W), viii.2008, [anonymous]��� 1 ♀, 1 ♂ (paratypes) (OMOC); Diouroup (14��21'N, 16��32'W), 14��� 18.viii.2007, P. Moretto��� 11 ♀♀, 17 ♂♂ (28 paratypes) (FGIC, PMOC); Diouroup (14��21'N, 16��32'W), 19��� 20.xi.2009, P. Moretto��� 1 ♀, 1 ♂ (paratypes) (PMOC); KAFFRINE, Tamba (13��53'49''N, 15��15'33''W), 2.vii.1948, [anonymous]��� 1 ♀, 1 ♂ (paratypes) (CNC); KAOLAK, Missirah, 45 m (13��59'26''N, 15��7'4''W), 16.vii.2007, F. G��nier (2007-01)��� 3 ♀♀, 9 ♂♂ (12 paratypes) (FGIC); Missirah, 45 m (13��59'26''N, 15��7'4''W), 17.vii.2007, F. G��nier (2007-02)��� 2 ♂♂ (2 paratypes) (FGIC); Nioro du Rip (13��44'43''N, 15��46'23''W), 10��� 11.viii.2009, P. Moretto��� 1 ♀, 1 ♂ (paratypes) (PMOC); LOUGA, Lingu��re (15��24'N, 15��7'W), ix.1967, A. Descarpentries, T. Leye &, Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 29-33, DOI: 10.5281/zenodo.439444, {"references":["Moretto, P. & Bordat, P. (2007) Les scarabeides coprophages du Parc National du Niokolo-Koba au Senegal (Coleoptera, Scarabaeoidea). Nouvelle Revue d'Entomologie (N. S.), 23, 115 - 145."]}

Digitonthophagus namaquensis species group 16. Digitonthophagus namaquensis G��nier, new species Distribution: Angola, Namibia, South Africa, Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on page 75, DOI: 10.5281/zenodo.439444

Digitonthophagus namaquensis G��nier, new species http://zoobank.org/urn:lsid:zoobank.org:act:48DC7E17-6D54-4345-9746-183D5D580AB8 (Figs. 22, 53���54, 70, 86, 117���118, 134, 180���182, 189; Map 17) Type locality. Farm Klipkoppies (6) (28.87137 S 21.30944 E), 957 m, Northern Cape, 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. G��nier, 2016] red card. MAP 17. Distribution of Digitonthophagus namaquensis. Material examined (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 km NW Karibib (21��47'S, 15��38'E), 31.i���2.ii.1972, Southern Africa 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, Southern Africa Expedition��� 1 ♀, 1 ♂ (paratypes) (BMNH); Kuiseb Canyon (23��18'S, 15��45'E), 22���23.i.1972, Southern Africa 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, Southern Africa Expedition��� 1 ♀ (paratype) (BMNH); Voigtgrund (24��49'S, 17��26'E), [no date], K. Jordan��� 1 ♀, 2 ♂♂ (3 paratypes) (BMNH); Malthah��he district, Zais 6 (24��2'58''S, 16��9'14''E), 16.i.2000, Mann & Marais��� 1 ♀ (paratype) (OUMNH); Rehoboth district, on road verge, Kambes 498 (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, Southern Africa 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, Southern Africa Expedition��� 3 ♀♀, 5 ♂♂ (8 paratypes) (BMNH); near Onseepkans, Orange RIVER banks (28��44'S, 19��18'E), 8���10.i.1972, Southern Africa Expedition��� 1 ♂ (paratype) (BMNH); Plateau Farm, 22 mi. E Aus (26��38'S, 16��34'E), 14���17.i.1972, Southern Africa 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); Mile 46 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, Southern Africa Expedition��� 2 ♀♀, 8 ♂♂ (10 paratypes) (BMNH); Windhoek (22��34'12''S, 17��5'1''E), 1909, K��lz��� 2 ♀♀ (2 paratypes) (SMF); KUNENE, 87 km W 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 / Khoanib confluence (19��14'44''S, 13��20'34''E), 28.xii.1999, Mann, Marais, & Newman��� 1 ♀, 1 ♂ (paratypes) (OUMNH); Hoanib river near Dubis (19��13'36''S, 13��22'E), 28.xii.1999, Mann & Marais��� 1 ♂ (paratype) (OUMNH); Palm 708, adjacent to spring (19��52'6''S, 14��1'13''E), 4.i.2000, Mann & Marais��� 1 ♂ (paratype) (OUMNH); Spaarwater 711, 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, Narawandu road, 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: EASTERN CAPE, Willowmore (33��18'S, 23��29'E), ii.1916, Dr. Brauns��� 1 ♀, 1 ♂ (paratypes) (NMPC); NORTHERN CAPE, 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 Northern Cape. 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. A single specimen collected under old elephant dung. This species is attracted to mercury vapor and ultraviolet light traps., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 70-72, DOI: 10.5281/zenodo.439444, {"references":["Davis, A. L. V. & Scholtz, C. H. (2004) Local and regional species ranges of a dung beetle assemblage from the semi-arid Karoo / Kalahari margins, South Africa. Journal of Arid Environments 57, 61 - 85."]}

Digitonthophagus dilatatus G��nier, new species http://zoobank.org/urn:lsid:zoobank.org:act:28F9FCFD-2012-4B76-B546-B444 F54327 D9 (Figs. 18, 45���46, 66, 82, 109���110, 130, 168���170; Map 13) Type locality. For��t de Sorobouli (11��47���44���N 02��53���25���W), 270 m, Sangui��, Burkina Faso. Diagnosis. Male cephalic horns granulate internally at apex (Fig. 46); males with pronotal anterior angles surface concave, produced anteriorly (Fig. 45); FLP sclerite ventral portion with a single carina (Fig. 170); SA sclerite apex interrupted much before apex of apical portion of FLP sclerite (Figs. 168���169). Description. Holotype ♂ (Fig. 18). Measurements. Length 12.0 mm, width 7.0 mm. Head (Figs. 45���46). Anterior clypeal edge straight on median seventh and appearing semicircular 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 long, distinctly arcuate and divergent in frontal view, parallel sided on basal four-fifths and abruptly tapering externally and narrower on apical sixth; posterointernal edge unmodified basally; apicointernal surface with few, coarse granules; genal edge slightly upturned and simply arcuate on anterior third, in line with clypeal edge. Pronotum (Fig. 66). Surface with granulate punctures restricted to anterior half medially, with distinct umbilicate punctures on posterior half of disc; punctures distinct posterolaterally, with distinct minute punctures throughout. Anteromedian tubercle atrophied, forming an obtuse angle in lateral view; median longitudinal sulcus well-defined but narrow; surface behind the eyes slightly concave, surface of anterior angles concave with anterior angles produced forwardly; anterior half of lateral edge sinuous in dorsal and lateral view; posterior angles unmodified and simply arcuate in dorsal view. Anterior hypomeral ridge straight on most of its length and abruptly arcuate on anterior third, anterior hypomeral depression surface darker in color. Elytra (Fig. 18). Intervals 2 and 4 lacking fine granules from base to apex. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus (Fig. 130). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites (Figs. 168���170). Axial sclerite short, angular and extending into a point apically. Subaxial sclerite, large basally and abruptly reduced, extending straight and approximately in line with apex of right lateral fold of frontolateral peripheral sclerite, with fine and scattered villi. Frontolateral peripheral sclerite basoventral apophysis well developed; a single rather large medioventral carina present; 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, not reaching anterior edge, apex set on left side in dorsal view; apical lobe truncate apically and directed obliquely on left side, apical villi regular in shape; subapicoventral lobe small, acute apically, short not reaching apical edge of apical lobe. Variation. Measurements (42 ♂♂, 83 ♀♀). Length: male 8.5���12.0 mm (10.5 �� 0.9 mm), female 7.5���12.0 mm (10.1 �� 0.8 mm). Female allotype. Cephalic outline in dorsal view as in Fig. 82; vertex with a broadly arcuate transverse carina, dorsal edge broadly arcuate in frontal view, lateral portion gradually sloping down posteriorly; anterior pronotal tubercles well developed, external lateral edges slightly widening toward apex in dorsal view, anterolateral surface simply convex, anterosuperior edge slightly arcuate in dorsal view (Fig. 109), lateral portion of anterosuperior edge slightly upturned (Fig. 110). Protibia short, with external teeth more robust. Primary type data. Holotype ♂ (CMNC): [BURKINA FASO: SANGUI�� / For��t de Sorobouli, 270m / 11��47���44���N 0 0 2��53���25���O / 13.VIII.2005, F. G��nier / zone soudanienne sud / pi��ge excr. humain, 2005-14]; [COL��OPT��RES / DU / BURKINA FASO / BF015361] barcode label; [HOLOTYPE ♂ / Digitonthophagus / dilatatus n.sp. / des. F. G��nier, 2016] red card. MAP 13. Distribution of Digitonthophagus dilatatus. Material examined (57 ♂♂, 93 ♀♀), distribution (Map 13): BURKINA FASO: CENTRE-OUEST, For��t de Sorobouli [= For��t Class��e de Baporo], 270 m (11��47'44''N, 2��53'25''W), 10���15.viii.2005, P. Moretto��� 78 ♀♀, 30 ♂♂ (108 paratypes) (FGIC, PMOC); BAL��, Boromo, 250 m (11��45'6''N, 2��51'58''W), 10.viii.2005, F. G��nier (2005-10)���1 ♀, 1 ♂ (paratypes) (FGIC); KOMPIENGA, Pama, 230 m (11��17'N, 0��42'59''E), 24.viii.2005, F. G��nier (2005-44)���3 ♂♂ (3 paratypes) (FGIC); SANGUI��, For��t de Sorobouli, 270 m (11��47'44''N, 2��53'25''W), 13.viii.2005, F. G��nier (2005-14)���1 ♀, 1 ♂ (holotype, allotype) (CMNC); For��t de Sorobouli, 270 m (11��47'44''N, 2��53'25''W), 13.viii.2005, F. G��nier (2005-15)���2 ♀♀, 1 ♂ (paratypes) (FGIC); For��t de Sorobouli, 270 m (11��47'44''N, 2��53'25''W), 28.vii.2006, F. & S. G��nier (2006-83)���1 ♂ (paratype) (FGIC).; CAMEROON: EXTREME NORTH, Maroua (10��35'N, 14��19'E), 4.vii.1978, [anonymous] ���1 ♂ (paratype) (MNHN); Waza (11��24'N, 14��34'E), 18.viii.2006, C. Vanderbergh��� 2 ♂♂ (2 paratypes) (FGIC, PMOC); Waza National Park (11��15'N, 14��39'E), 18.viii.2006, C. Vanderbergh��� 5 ♀♀, 5 ♂♂ (10 paratypes) (FGIC, PMOC); CHAD: N���DJAMENA, Farcha (12��7'N, 14��59'E), vi.1968, [anonymous]��� 1 ♀, 1 ♂ (paratypes) (MNHN); Farcha (12��7'N, 14��59'E), 28.vi.1967, [anonymous]��� 1 ♀, 2 ♂♂ (3 paratypes) (MNHN); Fort Lamy [= N���Djamena] (12��7'N, 15��3'E), 17.vii.1958, J. Mateu��� 1 ♂ (paratype) (FETC); C��TE D���IVOIRE: SAVANES, M���banto, Boundiali, 395 m (9��34'10.4''N, 6��40'56.5''W), 13���15.vii.2013, P. Moretto��� 2 ♀♀, 5 ♂♂ (7 paratypes) (FGIC, PMOC); GHANA: UPPER EAST, Bolgatanga (10��47'N, 0��5'W), 20.xi.2002, Bowku��� 2 ♀♀, 2 ♂♂ (4 paratypes) (OMOC); NIGERIA: KWARA, Bussa (8��51'24''N, 5��13'9''E), 4.v.1962, D.C. Eidt��� 1 ♂ (paratype) (CNC). Etymology. Dilatatus is a Latin adjective pertaining to the development of the anterior pronotal angles of welldeveloped males of this species. This development is unique in the genus. Natural history. The known distribution pattern of this species suggests that it is endemic to the Sudanese savanna. Interestingly, all specimens with data were collected at light or in pitfall traps baited with human dung, no specimens were collected in ruminant dung. Additional observations are required to confirm that it is a nonruminant dung specialist. Remarks. Externally, this species can be confused with D. fimator in West Africa. However, in addition to the characters given, the parameres are smaller and approximately parallel sided in lateral view as opposed to being wider and divergent in lateral view in D. fimator., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 59-61, DOI: 10.5281/zenodo.439444

Digitonthophagus Balthasar, 1959 Digitonthophagus Balthasar, 1959: 464 (original description) Digitonthophagus ��� Balthasar 1963: 159 (monograph) Digitonthophagus ��� Ferreira 1968: 593 (monograph) Digitonthophagus ��� Ferreira 1972: 779 (diagnosis) Digitonthophagus ��� Kabakov 1979: 94 (diagnosis) Digitonthophagus ��� Zunino 1981: 413 (new combination, comment taxonomy) Digitonthophagus ��� Zunino & Halffter 1988: 17 (comment taxonomy) Digitonthophagus ��� Ochi 2003: 260 (diagnosis, comment) Digitonthophagus ��� Monaghan et al. 2007: 678 (phylogeny) Digitonthophagus ��� Tarasov & Kabakov 2010: 24 (comment) Digitonthophagus ��� Tarasov & Solodovnikov 2011: 2 (phylogeny) Digitonthophagus ��� G��nier 2013: 2 (comment taxonomy) Digitonthophagus ��� Mlambo et al. 2015: 318 (phylogeny) Digitonthophagus ��� Breeschoten et al, 2016: 87 (phylogeny) Digitonthophagus ��� Roggero et al. 2016a: [1] (phylogeny) Digitonthophagus ��� Roggero et al. 2016b: 2 (phylogeny) Type species. Scarabaeus bonasus Fabricius, 1775 (original designation) Diagnosis. Externally, the genus Digitonthophagus is almost indistinguishable from the rest of the Onthophagini. However, all Digitonthophagus species have the anterior portion of the hypomeron produced anteriorly and visible from above; therefore, setae that would be located along the anterior edge in ventral portion are moved to a dorsal position. The only other external characters found separating the genus Digitonthophagus and the other closely related genera, is the presence of a distinctly shaped internoapical tooth of the male protibia (Figs. 7���22). This tooth is usually elongate, ventrally flat, and obliquely oriented downward. This character must be combined with females completely lacking this protibial internoapical tooth. In Onthophagus species where a similar tooth is present, the shape of this tooth is spiniform and forwardly directed in males and present in females as well but reduced in size (in African species examined). Internally, the parameres are rather short and the basolateral paramerites always present with the apex of the later adjacent to the lateral portion of the apical carinae. The internal sac of the aedeagus is lacking the lamella copulatrix and associated small sclerites and bristle pads. The frontolateral peripheral sclerite (Figs. 2���5) is located at the apex of the internal sac and a subapicoventral lobe oriented perpendicularly to the main plan of FLP is present. Description. Size. Small to moderately large (6.5���17.0 mm). Color. Overall color fulvous to light brown with structure edges and sutures more-or-less darker; pronotal disc and head darker and with distinct metallic sheen; elytral margins brown to green, disc with a more-or-less distinct darker irregular macula; metasternum and femoral ventral surface usually with much darker maculae. Head. With a sharply defined clypeofrontal carina in both sexes; antenna with 9 antennomeres, with deep pits on distal surface of antennomeres 7���8. Pronotum. With distinct squamiform granules anteriorly posterior to the anterior declivity that is always simply punctate, granules larger anteriorly and extending more-or-less posteriorly; anterior discal punctures weakly-defined and becoming well-defined posteriorly, when present. Elytra. Juxtasutural interval with strong metallic sheen along sutures; usually with a dark oval macula on the apical declivity on intervals 4���5. Pygidium. Surface with scattered, fine granulate punctures; each puncture with an erect, yellow seta. Ventrites. Hypomeron slightly convex; anterior hypomeral carina well-defined throughout; mesosternum with a bisinuate, glossy surface anteriorly and a more-orless developed longitudinal glossy, keel-shaped structure medially; posterior portion completely rugopunctate and setose; metasternal median lobe usually with a distinct longitudinal ridge behind the anterior suture. Aedeagus. Parameres rather short and basolateral paramerites always present, with the apex of the later adjacent to the lateral portion of the apical carinae. Female genitalia. Infundibulum thin, simple with a median portion reduced; receptaculum seminis elongate, simple, lacking annulate ridges internally (female genitalia characters from Zunino 1981). Remarks. Balthasar���s (1959) original description was restricted to external morphology and included a number of unrelated species (O. actaeon Balthasar, O. anguliceps Boucomont, O. avocetta Arrow, O. blumei van Lansberge, O. bonasus Fabricius, O. cameloides d'Orbigny, O. curvicarinatus Boucomont, O. diabolicus Harold, O. digitatus Arrow, O. diversiformis Boucomont, O. fossor Arrow, O. gazella Fabricius, O. kuluensis Bates, O. manipurensis Arrow, O. ohbayashii Nomura, O. ribbei Boucomont, O. rubricollis Hope, O. solivagus Harold, O. troniceki Balthasar, and O. wallacei Harold). The main characteristic used by Balthasar to define his subgenus was the elongate and thin protibia of the male. This character state is highly homoplasious in Scarabaeinae and evolved many times in the Onthophagini. The subgenus, as originally defined, was clearly paraphyletic (Zunino 1981). Remarks for species descriptions and identification key. Diagnoses based on external morphology refer to welldeveloped individuals. As in many scarabaeine species, strong allometric variation in the development of external secondary sexual characters is present. Smallest males will show completely feminized head and thoracic ornamentation, while largest males present two horns on the vertex. Pronotal and elytral granulations and punctation are usually diagnostic but variable, and extremes of one species can be similar to individuals of another species, which makes identification difficult with external characters for some specimens. Examination of the FLP sclerite shape will always provide a definite identification for males. Diagnoses for species groups must be used in conjunction with species diagnoses. Species are arranged into six groups based on the current phylogeny in order to facilitate comparison of closely-related species. Species are listed in alphabetical order within each group except for the Digitonthophagus bonasus group, where D. uks is treated next to D. bonasus since it is the most closely related species from the same biogeographical region., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 14-15, DOI: 10.5281/zenodo.439444, {"references":["Balthasar, V. (1959) Beitrag zur kenntnis der Gattung Onthophagus Latr. Sbornik entomologickeho Oddeleni Narodniho Musea v Praze, 33, 461 - 471.","Balthasar, V. (1963) Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region. Coleoptera: Lamellicornia. Band, 2. Coprinae (Onitini, Oniticellini, Onthophagini). Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region, 2. Verlag der Tschechoslowakischen Akademie der Wissenschaften, Prague, Czech Republic, 627 pp., 16 plates.","Ferreira, M. C. (1968) Os escarabideos de Mocambique. I. Revista de Entomologia de Mocambique, 10, 5 - 778.","Ferreira, M. C. (1972) Os escarabideos de Africa (sul do Saara). I. Revista de Entomologia de Mocambique, 11, 5 - 1088.","Kabakov, O. N. (1979) Obzor plastichnatoystykh zhukov podsemeystva Coprinae (Scarabaeidae, Coleoptera) Dalnego Vostoka i Sopredelnykh territor. In: Krivolutskaya, G. O. (Ed.), Zhuki Dal'nego Vostoka i vostochnoi Sibirii. Akademiya Nauk SSSR, Vladivostok, Russia, pp. 58 - 98.","Zunino, M. (1981) Insect of Saudi Arabia, Coleoptera: Fam. Scarabaeidae, Tribe Onthophagini. Fauna of Saudi Arabia, 3, 408 - 416.","Zunino, M. & Halffter, G. (1988) Analisis taxonomico, ecologico y biogeografico de un grupo americana de Onthophagus (Coleoptera, Scarabaeidae). Monographie, Museo Regionale di Scienze Naturali, Torino, 9, 1 - 211.","Ochi, T. (2003) Studies on the coprophagous scarab beetles from East Asia. VII Descriptions of two new subgenera of the genus Onthophagus (Co / eoptera, Scarabaeidae). Giornale Italiano di Entomologia, 10, 259 - 274.","Monaghan, M. T., Inward, D. J. G., Hunt, T. & Vogler, A. (2007) A molecular phylogenetic analysis of the Scarabaeinae (dung beetles). Molecular Phylogenetics and Evolution, 45, 674 - 692.","Tarasov, S. I. & Kabakov, O. N. (2010) Two new species of Onthophagus (Coleoptera: Scarabaeidae) from Indochina, with a discussion of some problems with the classification of Serrophorus and similar subgenera. Zootaxa, 2344, 17 - 28.","Tarasov, S. I. & Solodovnikov, A. Y. (2011) Phylogenetic analyses reveal reliable morphological markers to classify megadiversity in Onthophagini dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Cladistics, 27, 1 - 39. https: // doi. org / 10.1111 / j. 1096 - 0031.2011.00351. x","Genier, F. (2013) Transfert d' Onthophagus bubalus Harold, 1867, dans le genre Phalops Erichson, 1847, et notes sur sa position phylogenetique (Coleoptera, Scarabaeidae, Scarabaeinae). Catharsius, La Revue, 7, 1 - 4.","Mlambo, S., Sole, C. L. & Scholtz, C. H. (2015) A molecular phylogeny of the African Scarabaeinae (Coleoptera: Scarabaeidae). Arthropod Systematics & Phylogeny, 73, 303 - 321.","Breeschoten, T., Doorenweerd, C., Tarasov, S. & Vogler, A. P. (2016) Phylogenetics and biogeography of the dung beetle genus Onthophagus inferred from mitochondrial genomes. Molecular Phylogenetics and Evolution, 105, 86 - 95. https: // doi. org / 10.1016 / j. ympev. 2016.08.016","Roggero, A., Barbero, E. & Palestrini, C. (2016 a) Revised classification and phylogeny of an Afrotropical species group based on molecular and morphological data, with the description of a new genus (Coleoptera: Scarabaeidae: Onthophagini). Organisms Diversity & Evolution, 2016, [1 - 18].","Roggero, A., Dierkens, M., Barbero, E. & Palestrini, C. (2016 b) Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society, 2016, 1 - 23. https: // doi. org / 10.1111 / zoj. 12498","Fabricius, J. C. (1775) Systema Entomologiae sistens Insectorum Classes, Ordines, Genera, Species adiectis Synonymis, Locis Descriptionibus, Observationibus. Officina Libraria Kortii, Flensburg & Leipzig, Germany, xxxii + 832 pp."]}

Digitonthophagus falciger species group 6. Digitonthophagus falciger Génier, new species Distribution: Benin, Burkina Faso, Cameroon, Chad, Cote d’Ivoire, Gambia, Ghana, Guinea, Guinea-Bissau, Mauritania, Niger, Nigeria, Senegal, Sierra Leone,? Uganda

Digitonthophagus falciger species group Diagnosis. A single species with FLP sclerite right preapical edge deeply notched, basal edge of emargination produced into a sickle-shape projection (Fig. 150); axial sclerite apical portion thong shape, longer than subaxial sclerite., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on page 33, DOI: 10.5281/zenodo.439444

Digitonthophagus viridicollis G��nier, new species http://zoobank.org/urn:lsid:zoobank.org:act:11622669-B240-4DF1-AF7B-0757 A 587 DCEF (Figs. 16, 41���42, 64, 80, 105���106, 128, 162���164, 186; Map 11) Type locality. Moanda, Za��re [= 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. 186 B); 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 / (Za��re) 30.iv.1979] partly handwritten; [WORLD / SCARAB. / DATABASE / WSD00032092] barcode label; [HOLOTYPE ♂ / Digitonthophagus / viridicollis n.sp. / des. F. G��nier, 2016] red card. MAP 11. Distribution of Digitonthophagus viridicollis. Material examined (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, Cosso��o [= Co��o��o] (11��24'S, 14��1'E), iii.2005, [anonymous] ���2 ♀♀, 3 ♂♂ (5 paratypes) (PMOC); Cosso��o [=Co��o��o] (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 km N 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); Ro��adas (16��45'S, 14��59'E), 19��� 22.ii.1972, Southern Africa Expedition��� 3 ♀♀, 3 ♂♂ (6 paratypes) (BMNH); HUILA, 26 km S 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, Southern Africa Expedition��� 2 ♀♀, 1 ♂ (paratypes) (BMNH); Jo��o de Almeida [=Chibia] (15��11'S, 13��42'E), 29.iii.1972, [anonymous] (A 41)��� 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, Southern Africa 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, Mayumb�� (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 km NW Karibib (21��47'S, 15��38'E), 31.i���2.ii.1972, Southern Africa Expedition��� 2 ♀♀ (2 paratypes) (BMNH); Otjikoko S��d Farm, 33 mi. ENE Omaruru (21��10'S, 16��22'E), 10���13.ii.1972, Southern Africa 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); Otjitambi farm, 27 mi. ESE Kamanjab (19��49'S, 15��10'E), 13���15.ii.1972, Southern Africa 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, Southern Africa 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 km E Renostervlei (19��9'59''S, 14��33'12''E), 26.xii.1999, Mann & Marais��� 1 ♂ (paratype) (OUMNH); Narawandu road, Etosha National Park (18��51'4''S, 15��32'55''E), 24.xii.1999, Mann, Marais & Newman��� 2 ♀♀, 1 ♂ (paratypes) (OUMNH); Narawandu road, Etosha National Park (18��51'4''S, 15��32'55''E), 24���25.xii.1999, Mann & Marais��� 1 ♂ (paratype) (OUMNH); Narawandu road, Etosha National Park (18��51'4''S, 15��32'55''E), 24.xii.1999, Mann & Marais��� 1 ♂ (paratype) (OUMNH); Narawandu road, 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, Andoni road (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); North Tsintsabis (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, Southern Africa 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, Southern Africa 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, K��lz��� 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, Southern Africa Expedition��� 1 ♂ (paratype) (BMNH); Grootfontein District, Okaputa 334 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. A few 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. viridicollis also 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., Published as part of G��nier, Fran��ois & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 1-110 in Zootaxa 4248 (1) on pages 54-56, DOI: 10.5281/zenodo.439444