Digitonthophagus falciger Génier, 2017

Génier, François & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), Zootaxa 4248 (1), pp. 1-110 : 33-36

Digitonthophagus falciger Génier , new species

http://zoobank.org/urn:lsid:zoobank.org:act:84E0D6CA-8B72-407C-8012-024F76BCCBE0 ( Figs. 12 , 33–34 , 60 , 76 , 97–98, 124, 150–152; Map 7)

Type locality. Forêt de Boulon (10°16’27”N 04°27’15”W), 270 m, Comoé, Burkina Faso.

Description. Holotype ♂ ( Fig. 12 ). Measurements. Length 11.0 mm, width 7.0 mm. Head ( Figs. 33–34 ). 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, scabrous anteriorly, separated by one to four diameters. Horns moderately long, parallel on apical half 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 very broad angle with clypeal edge. Pronotum ( Fig. 60 ). 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 flat; anterior half of lateral edge arcuate in dorsal and lateral view; posterior angles simply arcuate in dorsal view. Anterior hypomeral ridge arcuate anteriorly, nearly straight posteriorly, anterior hypomeral depression surface slightly darker in color medially. Elytra. ( Fig. 12 ). Intervals 2 and 4 with few scattered fine granules from base to apex. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus ( Fig. 124 ). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites ( Figs. 150–152 ). Axial sclerite weekly sclerotized, as long as subaxial sclerite and almost straight. Subaxial sclerite, more-or-less even in width, extending straight and interrupted before apex of frontolateral peripheral sclerite apical portion, with short to long villi on apical sixth. Frontolateral peripheral sclerite basoventral apophysis moderately developed; with a spiniform medioventral carina; right lateral fold produced into a bifurcate process apically; left lateral lobe moderately sclerotized and developed; subapicodorsal lobe membranous, small, not reaching anterior edge, apex set on right side in dorsal view; right preapical edge deeply notched, basal edge of emargination produced into a sickle-shape projection; apex of apical lobe round apically, narrow and directed at right angle on the left side, apical villi regular in shape; subapicoventral lobe large, round, and interrupted in line with apical lobe.

Variation. Measurements (117 ♂♂, 123 ♀♀). Length: male 7.5–11.5 mm (9.9 ± 1.0 mm), female 7.5–12.0 mm (10.0 ± 0.8 mm). Female allotype. Cephalic outline in dorsal view as in Fig. 76 ; vertex with a broadly arcuate and slightly sinuous medially transverse carina, dorsal edge slightly sinuous in frontal view (Fig. 97), lateral portion subangular and sloping down posteriorly; anterior pronotal tubercles well developed, external lateral edges nearly straight and slightly widening toward apex in dorsal view (Fig. 98), anterolateral surface obliquely concave, anterosuperior edge slightly arcuate in dorsal view, lateral portion of anterosuperior edge slightly upturned. Protibia short, with external teeth robust.

Primary type data. Holotype ♂ ( CMNC): [ BURKINA FASO: LORUM / Forêt de Boulon, 270m / 10°16’27”N 004°27’15”O / 6.VII.2006, zone soudanienne / savane boisée, piège lumineux / F. & S. Génier, 2006- 05]; [COLÉOPTÈRES / DU / BURKINA FASO / BF015489] barcode label; [HOLOTYPE ♂ / Digitonthophagus / falciger n.sp. / des. F. Génier, 2016] red card.

MAP 7. Distribution of Digitonthophagus falciger .

Material examined (292 ♂♂, 159 ♀♀), distribution (Map 7): BENIN : MONO, Agoué (6°15'N, 1°41'E), vii– ix.1878, Abbé Ménager— 2 ♀♀, 2 ♂♂ (4 paratypes) ( FGIC, MNHN) ; Agoué (6°15'N, 1°41'E), 1879, Abbé Ménager— 4 ♀♀, 18 ♂♂ (22 paratypes) ( FGIC, MNHN); BURKINA FASO : CENTRE, Ouagadougou (12°22'N, 1°31'W), viii.1977, P. Ardoin— 1 ♂ (paratype) ( FETC) ; Ouagadougou (12°22'N, 1°31'W), 1977, J. Remy— 1 ♂ (paratype) ( PMOC) ; 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, Forêt de Sorobouli [= Forêt Classée de Baporo ], 270 m (11°47'44''N, 2°53'25''W), 10–15.viii.2005, P. Moretto— 6 ♀♀, 3 ♂♂ (9 paratypes) ( PMOC) ; CENTRE-SUD, Réserve de Nazinga (11°9'N, 1°36'W), 9–12.vi.2005, S. Rojkoff— 1 ♂ (paratype) ( PCSR) ; EST, Kompienga (11°4'26''N, 0°42'8''E), 3–24.ix.1988, F. Génier & M. Sanborne— 19 ♀♀, 47 ♂♂ (66 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. Génier & M. Sanborne— 2 ♀♀, 2 ♂♂ (4 paratypes) ( BDGC) ; HAUTS-BASSINS, Bobo-Dioulasso (11°12'N, 4°18'W), 11.i.1975, H. Pollitzar— 1 ♂ (paratype) ( BMNH) ; Bobo-Dioulasso (11°12'N, 4°18'W), 14.xi.1976, R.H. Gooding— 3 ♂♂ (3 paratypes) ( CMNC) ; Matourkou (11°5'N, 4°22'W), 15.vi.2005, S. Rojkoff— 1 ♂ (paratype) ( PMOC) ; NORD, 8 km SW Yako, Passoré , 320 m (12°51'44''N, 2°18'35''W), 7.viii.2005, P. Moretto— 1 ♂ (paratype) ( PMOC) ; SUD-OUEST, Bondigui , 324 m (10°53'56.5''N, 3°34'15''W), v.2012, F. Rousset— 1 ♂ (paratype) ( PMOC) ; Bondigui , 324 m (10°53'56.5''N, 3°34'15''W), 20.viii.2012, P. Moretto— 2 ♀♀, 1 ♂ (paratypes) ( PMOC) ; Gaoua (10°19'N, 3°10'W), 1924, H. Labouret— 1 ♂ (paratype) ( MNHN) ; COMOÉ, Forêt de Boulon (savane boisée), 270 m (10°16'27''N, 4°27'15''W), 6.vii.2006, F. & S. Génier (2006-05)— 1 ♀, 1 ♂ (holotype, allotype) ( CMNC) ; Forêt de Boulon (savane boisée), 270 m (10°16'27''N, 4°27'15''W), 9.vii.2006, F. & S. Génier (2006-17)— 2 ♂♂ (2 paratypes) ( FGIC) ; Forêt de Boulon (savane boisée), 270 m (10°16'27''N, 4°27'15''W), 10.vii.2006, F. & S. Génier (2006- 24)— 1 ♂ (paratype) ( FGIC) ; KOMPIENGA, Kompienga , 160 m (11°5'5''N, 0°42'42''E), 21.viii.2005, F. Génier (2005-39)— 2 ♀♀, 5 ♂♂ (7 paratypes) ( FGIC) ; Pama , 230 m (11°17'N, 0°42'59''E), 24.viii.2005, F. Génier (2005- 44)— 1 ♂ (paratype) ( FGIC) ; NAHOURI, Forêt de Nazinga, Kouzougou , 285 m (11°9'17''N, 1°32'10''W), 21.vii.2006, F. & S. Génier (2006-47)— 1 ♂ (paratype) ( FGIC) ; Forêt de Nazinga, Kouzougou , 285 m (11°9'17''N, 1°32'10''W), 25.vii.2006, F. & S. Génier (2006-70)— 1 ♂ (paratype) ( FGIC) ; Tiakané , 340 m (11°11'3''N, 1°12'28''W), 18.viii.2005, F. Génier (2005-23)— 1 ♀, 3 ♂♂ (4 paratypes) ( FGIC) ; Tiakané , 340 m (11°11'3''N, 1°12'28''W), 20.viii.2005, F. Génier (2005-34)— 1 ♂ (paratype) ( FGIC) ; Tiakané , 340 m (11°11'3''N, 1°12'28''W), 24.vii.2006, F. & S. Génier (2006-64)— 1 ♂ (paratype) ( FGIC) ; CAMEROON: EXTREME NORTH, Maroua (10°35'N, 14°19'E), vii.1979, G. Onore— 1 ♀, 2 ♂♂ (3 paratypes) ( PHWC) ; Waza National Park (11°15'N, 14°39'E), 18.viii.2006, C. Vanderbergh— 2 ♂♂ (2 paratypes) ( PMOC) ; NORTH, Garoua (9°18'N, 13°24'E), x.1973, B. de Miré— 1 ♀, 2 ♂♂ (3 paratypes) ( MNHN) ; Garoua (9°18'N, 13°24'E), [no date], [anonymous]— 1 ♀, 1 ♂ (paratypes) ( OMOC); CHAD : HADJER-LAMIS, Sitje , rive sud Lac Tchad (12°53'N, 14°52'E), 11.viii.2006, C. Vanderbergh— 1 ♀, 1 ♂ (paratypes) ( PMOC) ; N’DJAMENA, Farcha (12°7'N, 14°59'E), vi.1968, [anonymous]— 1 ♀, 2 ♂♂ (3 paratypes) ( MNHN) ; Farcha (12°7'N, 14°59'E), 28.vi.1967, [anonymous]— 1 ♀, 1 ♂ (paratypes) ( MNHN) ; Fort Lamy [= N’Djamena _] (12°7'N, 15°3'E), [no date], M. André— 1 ♀ (paratype) ( PMOC) ; Fort Lamy [= N’Djamena _] (12°7'N, 15°3'E), 17.vii.1958, J. Mateu— 2 ♂♂ (2 paratypes) ( FETC) ; CÔTE D’IVOIRE: BAFING, Ranch de Dolla, Biémasso (8°4'N, 7°33'W), 1–5.vi.2000, P. Moretto— 2 ♀♀, 14 ♂♂ (16 paratypes) ( PMOC) ; LACS, Bringakro near (6°25'53''N, 5°4'32''W), 12.vi.2003 – 13.vi.2006, Newman et al.— 2 ♀♀, 3 ♂♂ (5 paratypes) ( BMNH) ; Bringakro near (6°25'53''N, 5°4'32''W), 13. vi.2 0 0 3, Newman et al. — 2 ♀♀, 4 ♂♂ (6 paratypes) ( BMNH) ; Bringakro near (6°25'53''N, 5°4'32''W), 17.vi.2003, Newman et al.— 1 ♂ (paratype) ( BMNH) ; LAGUNES, 5 km Lamto (6°13'N, 5°1'W), viii.1968, C. Girard— 2 ♀♀, 3 ♂♂ (5 paratypes) ( MNHN) ; Forêt du Banco (5°23'N, 4°3'W), ii.1989, [anonymous]— 1 ♂ (paratype) ( CEMT) ; NZI-COMOE, Dimbroko (6°39'N, 4°42'W), [no date], [anonymous]— 1 ♂ (paratype) ( IRSNB); SAVANES , Koko, Korhogo (9°26'1''N, 4°43'9''W), 24–25.vi.2000, P. Moretto— 1 ♂ (paratype) ( PMOC) ; VALLÉE DU BANDAMA, Bouaké (7°41'N, 5°2'W), iii.1976, R. Couilloud— 2 ♀♀, 1 ♂ (paratypes) ( PMOC) ; Bouaké (7°41'N, 5°2'W), iv.1976, R. Couilloud— 2 ♀♀, 7 ♂♂ (9 paratypes) ( PMOC) ; Bouaké (7°41'N, 5°2'W), 13.vii.1977, [anonymous]— 1 ♀ (paratype) ( PMOC) ; GAMBIA: CENTRAL RIVER , Lower Saloum District, Ballangharr (13°39'N, 15°25'W), 23.viii.1997, D.J. Mann— 7 ♀♀, 7 ♂♂ (14 paratypes) ( OUMNH) ; Naini District, Kuntaur Agricultural Station (13°40'N, 14°53'W), 9.ix.1997, D.J. Mann— 1 ♂ (paratype) ( OUMNH) ; LOWER RIVER , Jarra West District, Jenoi DEC (13°28'51''N, 15°33'43''W), 29.viii.1997, Mann & Woodcock— 2 ♀♀, 2 ♂♂ (4 paratypes) ( OUMNH) ; UPPER RIVER, Bassa Government Rest Home, Mansanjang Kunda (13°16'53''N, 14°12'13''W), 26.viii.1997, D.J. Mann & B. Woodcock— 1 ♂ (paratype) ( OUMNH) ; WEST COAST , Kombo District, Yundum Agricultural Station (13°21'43''N, 16°39'43''W), vii.1997, D.J. Mann— 1 ♂ (paratype) ( OUMNH) ; Kombo District, Yundum Agricultural Station (13°21'43''N, 16°39'43''W), 19.viii.1997, D.J. Mann & B . A. Woodcock— 2 ♂♂ (2 paratypes) ( OUMNH) ; GHANA: “ Danish Guinea ”, [no date], P. E. Isert— 1 ♀, 1 ♂ (paratypes) ( ZMUC) ; GUINEA: BOFFA, Rio Pongo (10°10'N, 14°2'W), 1883–1884, P. Lutz— 1 ♂ (paratype) ( MNHN) ; GUINEA-BISSAU: BAFATÁ, Chime [= Xime] , Rio Géba (11°58'N, 14°56'W), 1906, G. Favarel— 2 ♀♀, 2 ♂♂ (4 paratypes) ( MNHN) ; TOMBALI, Quebo (11°32'N, 14°46'W), 25.vi–17.vii.1995 , A. Serrano— 1 ♂ (paratype) ( CEMT) ; MAURITANIA: GUIDIMAKA, [unspecified locality] (15°22'N, 12°6'W), 1908, G. Audan— 1 ♀, 8 ♂♂ (9 paratypes) ( FGIC, MNHN) ; NIGER: NIAMEY, Gorou Kirey (13°25'34''N, 2°8'32''E), 22.i.1976, [anonymous]— 1 ♂ (paratype) ( PHWC) ; TILLABERI, Kolo (12°55'22''N, 1°57'24''E), 15.iii.1972, D. Rougon— 1 ♂ (paratype) ( PHWC) ; NIGERIA: KADUNA , Kaduna (10°31'23''N, 7°26'25''E), 4.v.1962, D.C. Eidt— 1 ♂ (paratype) ( CNC) ; Samaru (11°10'N, 7°38'E), 18–25.v.1970, P.H. Ward— 1 ♀, 3 ♂♂ (4 paratypes) ( BMNH) ; Samaru (11°10'N, 7°38'E), 15–22.vi.1970, P.H. Ward— 1 ♂ (paratype) ( BMNH) ; KANO, [unspecified locality] (11°59'47''N, 8°31'0''E), v.1952, W.E. S. Merrett— 1 ♂ (paratype) ( BMNH) ; [unspecified locality] (11°59'47''N, 8°31'0''E), xi.1951, W.E.S. Merrett—1 ♂ (paratype) (BMNH); [unspecified locality] (11°59'47''N, 8°31'0''E), v.1953, W.E.S. Merrett—1 ♂ (paratype) (BMNH); [unspecified locality] (11°59'47''N, 8°31'0''E), vi.1954, W.E.S. Merrett—1 ♂ (paratype) (BMNH); NIGER, Badeggi (9°3'N, 6°9'E), 19.iii.1972, E.W. Classey— 1 ♂ (paratype) ( BMNH) ; OYO, Ibadan (7°22'N, 3°56'E), 2.iv.1957, J.L. Gregory— 1 ♀, 1 ♂ (paratypes) ( BMNH) ; SENEGAL: DAKAR , Dakar (14°46'N, 17°22'W), 1939, P. Lepesme, R. Paulian, & A. Villiers— 1 ♂ (paratype) ( MNHN) ; Hann (14°43'14''N, 17°26'12''W), 1939, P. Lepesme, R. Paulian, & A. Villiers— 15 ♀♀, 15 ♂♂ (30 paratypes) ( MNHN) ; Keur Ndiaye Lô (14°45'25''N, 17°14'11''W), 18.viii.2013, I. de Dinechin— 1 ♂ (paratype) (CMD) ; FATICK, Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 22.vi.2009 , A. Coache— 1 ♀, 3 ♂♂ (4 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 23.vi.2009 , A. Coache— 2 ♀♀, 3 ♂♂ (5 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 24.vi.2009 , A. Coache— 3 ♂♂ (3 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 16.x.2009 , A. Coache— 1 ♂ (paratype) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 28.xi.2009 , A. Coache— 1 ♀ (paratype) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 20.iv.2010 , A. Coache— 1 ♂ (paratype) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 15.vii.2010 , A. Coache— 1 ♀, 1 ♂ (paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 22.vii.2010 , A. Coache— 1 ♂ (paratype) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 13.viii.2010 , A. Coache— 1 ♀, 1 ♂ (paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 14.viii.2010 , A. Coache— 2 ♀♀ (2 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 15.viii.2010 , A. Coache— 1 ♂ (paratype) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 16.viii.2010 , A. Coache— 1 ♂ (paratype) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 17.viii.2010 , A. Coache— 2 ♂♂ (2 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 21.ix.2010 , A. Coache— 1 ♀, 3 ♂♂ (4 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 22.ix.2010 , A. Coache— 1 ♀ (paratype) (CAC) ; KAFFRINE, Tamba (13°53'49''N, 15°15'33''W), 2.vii.1948, [anonymous]— 2 ♂♂ (2 paratypes) ( CMNC, CNC) ; KAOLAK, Missirah , 45 m (13°59'26''N, 15°7'4''W), 16.vii.2007, F. Génier (2007-01)— 2 ♂♂ (2 paratypes) ( FGIC) ; Missirah , 45 m (13°59'26''N, 15°7'4''W), 17.vii.2007, F. Génier (2007-03)— 1 ♂ (paratype) ( FGIC) ; Nioro du Rip (13°44'43''N, 15°46'23''W), 3.viii.2008, P. Moretto— 1 ♂ (paratype) ( PMOC) ; Nioro du Rip (13°44'43''N, 15°46'23''W), 10–11.viii.2009, P. Moretto— 1 ♀, 1 ♂ (paratypes) ( PMOC) ; KÉDOUGOU, Dinndefelou (12°22'43''N, 12°19'30''W), 13–14.vii.2008, P. Moretto— 1 ♂ (paratype) ( PMOC) ; KOLDA, Forêt de Bakor, Mahon (12°51'N, 14°49'W), 31.vii–2.viii.2008, P. Moretto— 1 ♂ (paratype) ( PMOC) ; Mahon (12°51'22''N, 14°48'42''W), 22–23.xi.2009, P. Moretto— 2 ♀♀, 3 ♂♂ (5 paratypes) ( PMOC) ; Sare Gnako , 31 m (12°44'16''N, 15°9'56''W), 31.vii.2007, F. Génier & P. Moretto (2007-49)— 2 ♂♂ (2 paratypes) ( FGIC) ; Sothoto , 23 m (12°39'55''N, 15°24'3''W), 31.vii.2007, Génier & Moretto (2007 -47)— 2 ♂♂ (2 paratypes) ( FGIC) ; LOUGA, Linguère (15°24'N, 15°7'W), ix.1967 , A. Descarpentries, T. Leye, & A. Villiers— 10 ♀♀, 6 ♂♂ (16 paratypes) ( MNHN); Ndam-Dam (15°36'22''N, 16°23'35''W), xi.1967 , A. Descarpentries, T. Leye, & A. Villiers— 4 ♀♀, 1 ♂ (paratypes) ( MNHN) ; SAINT-LOUIS, Parc National des Oiseaux du Djoudj (16°24'N, 16°14'W), x.1996, [anonymous]— 1 ♀, 3 ♂♂ (4 paratypes) ( PSCW) ; Richard-Toll (16°28'N, 15°41'W), xi.1967, A. Descarpentries, T. Leye & A. Villiers— 30 ♀♀, 23 ♂♂ (53 paratypes) ( MNHN) ; SEDHIOU , Sedhiou (12°42'N, 15°34'W), [no date], [anonymous]— 1 ♂ (paratype) ( MNHN) ; Sedhiou (12°42'N, 15°34'W), 14.vii.1947, [anonymous]— 1 ♂ (paratype) ( MNHN) ; TAMBACOUNDA, Boutou , 52 m (13°48'16''N, 13°34'47''W), 20.vii.2007, F. Génier (2007-06)— 1 ♂ (paratype) ( FGIC) ; Dienoun Diala (13°12'47''N, 13°6'44''W), 29.vii.1995, P. Moretto— 8 ♀♀, 4 ♂♂ (12 paratypes) ( PMOC) ; ZIGUINCHOR, Boukitimgo , 18 m (12°26'41''N, 16°35'53''W), 4.viii.2007, F. Génier (2007-56)— 1 ♂ (paratype) ( FGIC) ; Boukitimgo , 18 m (12°26'41''N, 16°35'53''W), 4.viii.2007, P. Moretto & F. Génier (2007-58) — 1 ♂ (paratype) ( FGIC) ; SIERRA LEONE: NORTHERN, Koinadugu (9°32'9''N, 11°22'6''W), 23.v.1963, Mission ENS-IFAN aux Monts Loma— 1 ♀, 4 ♂♂ (5 paratypes) ( MNHN) ; UGANDA: Nebbi District, Pamora, Padyere subcounty (2°24'25''N, 31°11'40''E), v.2006, Local collectors— 1 ♀, 1 ♂ (paratypes) ( PMOC) .

Etymology. Falciger (sickle + bear) is a Latin adjective in apposition referring to the distinctive configuration of the left edge of the frontolateral peripheral sclerite of this species.

Natural history. The few specimens with data were collected from savanna, disturbed areas, and pastures in soudanian zone and from degraded Guinean Forest. Attracted to mercury vapor and ultraviolet light, some specimens were collected using pitfall traps baited with human or western red colobus ( Procolobus badius ) dung, some specimens retrieved from cow, donkey, elephant, rhinoceros, zebra, and zebu dung.

Remarks. The Ugandan record based on two specimens need confirmation.

Figures

FIGURES 11 – 14. Male dorsal habitus. 11. D. eucatta, holotype; 12. D. falciger, holotype; 13. Digitonthophagus biflagellatus, holotype; 14. D. gazella, non type (Central Province, Zambia).

FIGURES 31 – 38. 31, 33, 35, 37. Male head, dorsal view; 32, 34, 36, 38. Male head, frontal view. 31 – 32. Digitonthophagus eucatta, alloreferent; 33 – 34. D. falciger, holotype; 35 – 36. D. biflagellatus, holotype; 37 – 38. D. gazella, non type (Central Province, Zambia).

FIGURES 55 – 62. Male pronotum, dorsal view. 55. Digitonthophagus bonasus, alloreferent (Tamil Nadu State, India); 56. D. uks, holotype; 57. D. sahelicus, holotype; 58. D. catta, alloreferent (Tamil Nadu State, India); 59. D. eucatta, holotype; 60. D. falciger, holotype; 61. D. biflagellatus, holotype; 62. D. gazella, non type (Central Province, Zambia).

FIGURES 71 – 78. Female head, dorsal view. 71. Digitonthophagus bonasus, non type (Tamil Nadu State, India); 72. D. uks, allotype; 73. D. sahelicus, allotype; 74. D. catta, non type (India); 75. D. eucatta, allotype; 76. D. falciger, allotype; 77. D. biflagellatus, allotype; 78. D. gazella, non type (Antisiranana Province, Madagascar).

FIGURES 119 – 134. Parameres, dorsal view. 119. D. bonasus, alloreferent (Tamil Nadu State, India); 120. Digitonthophagus uks, holotype; 121. D. sahelicus, holotype; 122. D. catta, alloreferent (West Bengal, India); 123. D. eucatta, holotype; 124. D. falciger, holotype; 125. D. biflagellatus, holotype; 126. D. gazella, non type (Antisiranana Province, Madagascar); 127. D. petillus, holotype; 128. D. viridicollis, holotype; 129. D. aksumensis, holotype; 130. D. dilatatus, holotype; 131. D. fimator, holotype; 132. D. lusinganus, lectotype; 133. D. ulcerosus, holotype 134. D. namaquensis, holotype.

FIGURES 147 – 158. Internal sac sclerites (FLP, A + SA). 147, 150, 153, 156. Dorsal view; 148, 151, 154, 157. Ventral view. 149, 152, 155, 158. Right lateral view. 147 – 149. Digitonthophagus eucatta, alloreferent (Tamil Nadu State, India); 150 – 152. D. falciger, holotype; 153 – 155. D. biflagellatus, holotype; 156 – 158. D. gazella, non type (Antisiranana Province, Madagascar).