Digitonthophagus fimator 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 : 61-65

Digitonthophagus fimator Génier , 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. Forêt de Sorobouli (11°47’44”N 02°53’25”W), 270 m, Sanguié , 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: SANGUIÉ / Forêt de Sorobouli , 270m / 11°47’44”N 002°53’25”W / 13.VIII.2005, F. Génier / zone soudanienne sud / piège excr. Humain, 2005-14]; [COLÉOPTÈRES / DU / BURKINA FASO / BF015498] barcode label; [HOLOTYPE ♂ / Digitonthophagus / fimator n.sp. / des. F. Génier, 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, Forêt 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, Forêt de Wari-Maro (WM2) (9°1'37''N, 2°24'13''E), 14.iv.2007, S. Tchibozo— 1 ♂ (paratype) ( PMOC) ; DONGA, Forêt de Pénélan, Pénéssoulou , 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, Forêt de Houéyogbé, Houéyogbé , 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); OUÉMÉ, 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, Forêt de Sorobouli [= Forêt Classée 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, Réserve de Nazinga (11°9'N, 1°36'W), 9– 12.vi.2005, S. Rojkoff— 17 ♀♀, 16 ♂♂ (33 paratypes) ( PCSR, PMOC) ; BALÉ, Boromo, 250 m (11°45'6''N, 2°51'58''W), 10.viii.2005, F. Génier (2005-10)—1 ♂ (paratype) (FGIC); COMOÉ, Forêt de Boulon (savane boisée) , 270 m (10°16'27''N, 4°27'15''W), 5.vii.2006, F. & S. Génier (2006-02)—1 ♂ (paratype) (FGIC); Forêt de Boulon (savane boisée), 270 m (10°16'27''N, 4°27'15''W), 6.vii.2006, F. & S. Génier (2006-05)— 1 ♂ (paratype) ( FGIC) ; Forêt de Boulon (savane boisée), 270 m (10°16'27''N, 4°27'15''W), 9.vii.2006, F. & S. Génier (2006-17)— 3 ♂♂ (3 paratypes) ( FGIC) ; Koflandé ( Village ), 290 m (10°14'42''N, 4°27'50''W), 4.vii.2006, F. & S. Génier (2006- 01)— 4 ♂♂ (4 paratypes) ( FGIC) ; NAHOURI, Forêt de Nazinga , 275 m (11°9'24''N, 1°36'44''W), 20.vii.2006, F. & S. Génier (2006-46)— 8 ♀♀, 9 ♂♂ (17 paratypes) ( FGIC); Forêt de Nazinga, Akwazena , 275 m (11°9'24''N, 1°36'44''W), 21.vii.2006, F. & S. Génier (2006-53)—1 ♂ (paratype) (FGIC); Forêt de Nazinga, Akwazena , 275 m (11°9'24''N, 1°36'44''W), 26.vii.2006, F. & S. Génier (2006-73)— 1 ♀, 2 ♂♂ (3 paratypes) ( FGIC) ; Forêt de Nazinga, Akwazena , 275 m (11°9'24''N, 1°36'44''W), 26.vii.2006, F. & S. Génier (2006-75)— 2 ♀♀, 3 ♂♂ (5 paratypes) ( FGIC) ; Forêt de Nazinga, Barka , 265 m (11°8'30''N, 1°36'35''W), 22.vii.2006, F. & S. Génier (2006- 57)— 7 ♀♀, 10 ♂♂ (17 paratypes) ( FGIC); Forêt de Nazinga, Barka , 265 m (11°8'30''N, 1°36'35''W), 24.vii.2006, F. & S. Génier (2006-62)—1 ♂ (paratype) (FGIC); Forêt de Nazinga, Boulieselo , 310 m (11°11'50''N, 1°35'9''W), 27.vii.2006, F. & S. Génier (2006-82)— 2 ♂♂ (2 paratypes) ( FGIC); Forêt de Nazinga, Kalie Boulou , 275 m (11°11'29''N, 1°30'26''W), 22.vii.2006, F. & S. Génier (2006-54)—1 ♂ (paratype) (FGIC); Forêt de Nazinga, Kalie Boulou , 275 m (11°11'29''N, 1°30'26''W), 25.vii.2006, F. & S. Génier (2006-67)— 1 ♀ (paratype) ( FGIC) ; Forêt de Nazinga, Kouzougou , 285 m (11°9'17''N, 1°32'10''W), 21.vii.2006, F. & S. Génier (2006-47)— 8 ♀♀, 2 ♂♂ (10 paratypes) ( FGIC); Forêt de Nazinga, Naguio , 270 m (11°7'52''N, 1°34'38''W), 24.vii.2006, F. & S. Génier (2006- 66)—1 ♂ (paratype) (FGIC); SANGUIÉ, Forêt de Sorobouli , 270 m (11°47'44''N, 2°53'25''W), 13.viii.2005, F. Génier (2005-14)— 1 ♀, 1 ♂ (holotype, allotype) ( CMNC) ; Forêt de Sorobouli , 270 m (11°47'44''N, 2°53'25''W), 14.viii.2005, F. Génier (2005-18)— 2 ♀♀, 1 ♂ (paratypes) ( FGIC) ; TAPOA, Kaabougou, 280 m (11°57'22''N, 2°0'40''E), 12.vii.2006, F. & S. Génier (2006-30)—1 ♂ (paratype) (FGIC); Kaabougou , 280 m (11°57'22''N, 2°0'40''E), 12.vii.2006, F. & S. Génier (2006-31)— 1 ♂ (paratype) ( FGIC) ; CAMEROON: Yaoundé-Bertoua , v.2003, M. Ondoa— 55 ♀♀, 60 ♂♂ (115 paratypes) ( CMD, PMOC) ; ADAMAWA, Ngaoundéré (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, Yaoundé (3°52'16''N, 11°27'14''E), 18.x.1973, B. de Miré— 1 ♀, 2 ♂♂ (3 paratypes) ( MNHN) ; Nkolbisson, Yaoundé (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 Miré— 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 Maboké (3°53'N, 17°58'E), 20.vi.1965, [anonymous]— 1 ♀ (paratype) ( MNHN) ; Station de La Maboké (3°53'N, 17°58'E), 23.ix.1965, R. Pujol— 1 ♀ (paratype) ( MNHN) ; Station de La Maboké (3°53'N, 17°58'E), 24.ix.1965, R. Pujol— 2 ♀♀, 2 ♂♂ (4 paratypes) ( MNHN) ; Station de La Maboké (3°53'N, 17°58'E), 31.xii.1965, [anonymous]— 2 ♀♀ (2 paratypes) ( MNHN) ; Station de La Maboké (3°53'N, 17°58'E), 22.iii.1966, R. Pujol— 1 ♂ (paratype) ( MNHN) ; Station de La Maboké (3°53'N, 17°58'E), 27.iv.1968, R. Pujol— 97 specimens (paratypes) ( MNHN) ; Station de La Maboké (3°53'N, 17°58'E), 19.ii.1969, P. Teocchi— 2 ♀♀, 2 ♂♂ (4 paratypes) ( MNHN) ; MAMBERE-KADEI, Berbérati (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. Hervé— 1 ♀, 3 ♂♂ (4 paratypes) ( PMOC) ; CHAD: LOGONE ORIENTAL, Komé, 80 km Moundou (8°30'N, 16°41'E), vii.2002, T. Garnier— 2 ♀♀, 2 ♂♂ (4 paratypes) ( PMOC) ; CÔTE D’IVOIRE: BAFING, Ranch de Dolla, Biémasso , 450 m (8°4'N, 7°33'W), iii–iv.2002, P. Moretto— 43 ♀♀, 36 ♂♂ (79 paratypes) ( FGIC, PMOC) ; Ranch de Dolla, Biémasso , 450 m (8°4'N, 7°33'W), 1–5.vi.2000, P. Moretto— 6 ♀♀, 6 ♂♂ (12 paratypes) ( PMOC) ; DENGUELE, hotel des frontières, Odienné , 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, Forêt 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 Marahoué (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) ; VALLÉE DU BANDAMA, Bouaké (7°41'N, 5°2'W), 5.iv.1974, E. & I. Munroe— 1 ♀, 1 ♂ (paratypes) ( CMNC) ; Bouaké (7°41'N, 5°2'W), 7–28.v.1980, O. Kukal— 3 ♀♀, 2 ♂♂ (5 paratypes) ( CMNC) ; Bouaké (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); Forêt de Kawa (1°33'N, 30°32'E), 15.iv.1929 , A. Collart— 1 ♂ (paratype) ( IRSNB); Forêt 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. Léonard— 3 ♂♂ (3 paratypes) ( FGIC, PMOC) ; Dila (6°24'30''N, 38°18'30''E), ix.2000, P. Léonard— 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) ; NZÉRÉKORÉ, Thio , Mont Nimba (7°35'10''N, 8°30'7''W), ii–vi.1942, M. Lamotte— 2 ♀♀, 3 ♂♂ (5 paratypes) ( IRSNB) ; GUINEA- BISSAU : BAFATÁ, Chime [= Xime] , Rio Géba (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. Léonard & P. Vingerhoedt— 3 ♀♀, 5 ♂♂ (8 paratypes) ( PMOC) ; [unspecified locality], vi.1955, A.M. Robertson— 2 ♀♀ (2 paratypes) ( BMNH) ; CROSS RIVER, Calabar (4°57'N, 8°19'E), 11.v.1984, [anonymous]— 1 ♀ (paratype) ( BMNH) ; Calabar (4°57'N, 8°19'E), 2–3.vii.1984, [anonymous]— 1 ♂ (paratype) ( BMNH); KADUNA , Kaduna (10°31'23''N, 7°26'25''E), 9.vi.1970, P.H. Ward— 2 ♂♂ (2 paratypes) ( BMNH) ; Samaru (11°10'N, 7°38'E), 18–25.v.1970, P.H. Ward— 5 ♀♀, 3 ♂♂ (8 paratypes) ( BMNH) ; KWARA, Offa (8°8'25''N, 4°43'32''E), [no date], R.C. Hiscock— 1 ♂ (paratype) ( BMNH) ; OYO, Ibadan (7°22'N, 3°56'E), 2.v.1957, J.L. Gregory— 1 ♀, 3 ♂♂ (4 paratypes) ( BMNH) ; PLATEAU, Jos (9°55'N, 8°54'E), vi.1976, [anonymous]— 2 ♀♀, 1 ♂ (paratypes) ( PHWC) ; Jos (9°55'N, 8°54'E), v.1976, R. Raoux— 1 ♂ (paratype) ( PHWC) ; Jos (9°55'N, 8°54'E), 13.iii.1976, R. Raoux— 3 ♀♀ (3 paratypes) ( PHWC) ; SENEGAL: DAKAR, Sangalkam (14°47'N, 17°14'W), viii.1971 , A. Villiers— 2 ♂♂ (2 paratypes) ( OMOC) ; FATICK, Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 25.iii.2009 , A. Coache— 1 ♀, 2 ♂♂ (3 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 26.iii.2009 , A. Coache— 2 ♀♀, 1 ♂ (paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 27.iii.2009 , A. Coache— 1 ♂ (paratype) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 28.iv.2009 , A. Coache— 2 ♀♀, 1 ♂ (paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 22.vi.2009 , A. Coache— 4 ♀♀, 7 ♂♂ (11 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 23.vi.2009 , A. Coache— 2 ♀♀, 4 ♂♂ (6 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 24.vi.2009 , A. Coache— 7 ♀♀, 4 ♂♂ (11 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 21.iv.2010 , A. Coache— 1 ♂ (paratype) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 12.vii.2010 , A. Coache— 4 ♀♀, 2 ♂♂ (6 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 13.vii.2010 , A. Coache— 2 ♀♀ (2 paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 14.vii.2010 , A. Coache— 2 ♀♀, 1 ♂ (paratypes) (CAC); Réserve de Fathala, Parc national du delta du Saloum (13°39'N, 16°27'W), 15.vii.2010 , A. Coache— 2 ♂♂ (2 paratypes) (CAC) ; KAFFRINE, Tamba (13°53'49''N, 15°15'33''W), 2.vii.1948, [anonymous]— 1 ♂ (paratype) ( CNC) ; KOLDA, Sare Gnako , 31 m (12°44'16''N, 15°9'56''W), 31.vii.2007, F. Génier & P. Moretto (2007 -49)—1 ♀ (paratype) (FGIC); SAINT-LOUIS, Richard-Toll (16°28'N, 15°41'W), xi.1967 , A. Descarpentries, T. Leye & A. Villiers— 8 ♀♀, 4 ♂♂ (12 paratypes) ( MNHN) ; Richard-Toll (16°28'N, 15°41'W), 28.viii–1.ix.2009, P. Moretto & F. Génier— 4 ♀♀, 4 ♂♂ (8 paratypes) ( PMOC) ; Richard-Toll (16°28'N, 15°41'W), 1.ix.2009, F. Génier (2009-49)— 1 ♂ (paratype) ( FGIC) ; TAMBACOUNDA, Ancien Poste, Parc National Niokolo-Koba , 144 m (12°53'19''N, 12°43'10''W), 24– 26.vii.2007, F. Génier & P. Moretto (2007-27)— 1 ♀, 2 ♂♂ (3 paratypes) ( FGIC) ; Ancien Poste, Parc National Niokolo-Koba , 144 m (12°53'19''N, 12°43'10''W), 24.vii.2007, F. Génier & P. Moretto (2007-23)— 3 ♂♂ (3 paratypes) ( FGIC); Ancien Poste, Parc National Niokolo-Koba , 144 m (12°53'19''N, 12°43'10''W), 24–25.vii.2007, F. Génier & P. Moretto (2007 -26)—1 ♂ (paratype) (FGIC); ZIGUINCHOR, Boukitimgo , 18 m (12°26'41''N, 16°35'53''W), 4.viii.2007, P. Moretto & F. Génier (2007-58) — 1 ♂ (paratype) ( FGIC) ; Ziguinchor (12°35'N, 16°16'W), 28–31.iii.1977, J.L. & A. Nicolas— 2 ♀♀, 2 ♂♂ (4 paratypes) ( CMD) ; SIERRA LEONE: NORTHERN, Kanike (9°18'N, 11°52'W), 22.v.1912, J.J. Simson— 1 ♂ (paratype) ( BMNH) ; Koinadugu (9°32'9''N, 11°22'6''W), 23.v.1963, Mission ENS-IFAN aux Monts Loma— 7 ♀♀, 10 ♂♂ (17 paratypes) ( MNHN) ; Musaia [= Musala ] (9°45'N, 11°34'W), 26.v.1950, F. A.S.— 1 ♀, 2 ♂♂ (3 paratypes) ( BMNH) ; WESTERN, Freetown (8°29'N, 13°14'W), 1889 , A. Mocquerys— 1 ♀ (paratype) ( MNHN) ; TOGO: CENTRALE, Sokode (8°59'N, 1°8'E), iii.1957, P. Jauffret— 2 ♂♂ (2 paratypes) ( PMOC) ; UGANDA: BUSIA , Busia (0°28'N, 34°5'E), i.1990, B. Wandera— 2 ♀♀, 1 ♂ (paratypes) ( FETC) ; Busia (0°28'N, 34°5'E), vii.1990, [anonymous]— 1 ♂ (paratype) ( FETC) ; NORTHERN , Nebbi District, Pamora, Padyere subcounty (2°24'25''N, 31°11'40''E), v.2006, Local collectors— 2 ♀♀, 3 ♂♂ (5 paratypes) ( PMOC) .

Etymology. Fimator (from fimus) a Latin noun in apposition meaning dung remover. Pertaining to the natural history of this species.

Natural history. The distribution of this species in the soudanian zone and forest/savanna mosaic areas, with intrusions in the northern portion of the central forest block, suggests that it can colonize wetter environments when used as pastures. All Cameroonian and Central African records are from pastures in forest/savanna mosaic. Few individuals were collected from the Sahelian region. Specimens were collected from ruminants and non-ruminants such as cow, elephant, roan antelope ( Hippotragus equinus ), pig, rhinoceros, warthog, and zebra dung. This species is attracted to mercury vapor and ultraviolet light.

Figures

FIGURES 19 – 22. Male dorsal habitus. 19. Digitonthophagus fimator, holotype; 20. D. lusinganus, lectotype; 21. D. ulcerosus, holotype; 22. D. namaquensis, holotype.

FIGURES 47 – 54. 47, 49, 51, 53. Male head, dorsal view; 48, 50, 52, 54. Male head, frontal view. 47 – 48. Digitonthophagus fimator, holotype; 49 – 50. D. lusinganus, lectotype; 51 – 52. D. ulcerosus, holotype; 53 – 54. D. namaquensis, holotype.

FIGURES 63 – 70. Male pronotum, dorsal view. 63. D. petilus, holotype; 64. Digitonthophagus viridicollis, holotype; 65. D. aksumensis, holotype; 66. D. dilatatus, holotype; 67. D. fimator, holotype; 68. D. lusinganus, lectotype; 69. D. ulcerosus, holotype 70. D. namaquensis, holotype.

FIGURES 79 – 86. Female head, dorsal view. 79. Digitonthophagus petilus, allotype; 80. D. viridicollis, allotype; 81. D. aksumensis, allotype; 82. D. dilatatus, allotype; 83. D. fimator, allotype; 84. D. lusinganus, paralectotype; 85. D. ulcerosus, allotype 86. D. namaquensis, allotype.

FIGURES 171 – 182. Internal sac sclerites (FLP, A + SA). 171, 174, 177, 180. Dorsal view; 172, 175, 178, 181. Ventral view. 173, 176, 179, 182. Right lateral view. 171 – 173. Digitonthophagus fimator, holotype; 174 – 176. D. lusinganus, lectotype; 177 – 179. D. ulcerosus, holotype; 180 – 182. D. namaquensis, holotype.

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.