Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae) Génier, François Moretto, Philippe Zootaxa 2017 4248 1 1 110 363WL Genier Genier [151,561,1186,1213] Insecta Scarabaeidae Digitonthophagus Animalia Coleoptera 32 33 Arthropoda species falciger sp. nov.     Typelocality. 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 typedata. Holotype ♂( CMNC): [ BURKINA FASO: LORUM/  Forêt de Boulon,  270m/ 10°16’27”N004°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.     Materialexamined( 292 ♂♂, 159 ♀♀), distribution (Map 7):  BENIN:  MONO, Agoué( 6°15'N, 1°41'E),  vii– ix.1878, AbbéMénager— 2 ♀♀, 2 ♂♂( 4 paratypes) ( FGIC, MNHN);  Agoué( 6°15'N, 1°41'E), 1879, AbbéMénager— 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 kmN 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 kmS Pama( 11°4'37''N, 0°40'50''E),  1–16.vi.1988, Sanborne, Landry& Tou— 2 ♀♀( 2 paratypes) ( CMNC);  Kompienga,  20 kmS 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 kmSW 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, Bringakronear ( 6°25'53''N, 5°4'32''W),  12.vi.2003–   13.vi.2006, Newman et al.— 2 ♀♀, 3 ♂♂( 5 paratypes) ( BMNH);  Bringakronear ( 6°25'53''N, 5°4'32''W), 13.  vi.20 0 3, Newman et al.— 2 ♀♀, 4 ♂♂( 6 paratypes) ( BMNH);  Bringakronear ( 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 WestDistrict, JenoiDEC ( 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,  RioPongo ( 10°10'N, 14°2'W),  1883–1884, P. Lutz— 1 ♂( paratype) ( MNHN);   GUINEA-BISSAU: BAFATÁ, Chime [= Xime],  RioGéba ( 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, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  22.vi.2009,  A. Coache— 1 ♀, 3 ♂♂( 4 paratypes) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  23.vi.2009,  A. Coache— 2 ♀♀, 3 ♂♂( 5 paratypes) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  24.vi.2009,  A. Coache— 3 ♂♂( 3 paratypes) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  16.x.2009,  A. Coache— 1 ♂( paratype) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  28.xi.2009,  A. Coache— 1 ♀( paratype) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  20.iv.2010,  A. Coache— 1 ♂( paratype) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  15.vii.2010,  A. Coache— 1 ♀, 1 ♂( paratypes) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  22.vii.2010,  A. Coache— 1 ♂( paratype) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  13.viii.2010,  A. Coache— 1 ♀, 1 ♂( paratypes) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  14.viii.2010,  A. Coache— 2 ♀♀( 2 paratypes) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  15.viii.2010,  A. Coache— 1 ♂( paratype) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  16.viii.2010,  A. Coache— 1 ♂( paratype) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  17.viii.2010,  A. Coache— 2 ♂♂( 2 paratypes) (CAC);  Réserve de Fathala, Parcnational du delta du Saloum( 13°39'N, 16°27'W),  21.ix.2010,  A. Coache— 1 ♀, 3 ♂♂( 4 paratypes) (CAC);  Réserve de Fathala, Parcnational 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 Nationaldes  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, MissionENS-IFAN aux MontsLoma— 1 ♀, 4 ♂♂( 5 paratypes) ( MNHN);   UGANDA: Nebbi District, Pamora, Padyeresubcounty ( 2°24'25''N, 31°11'40''E),  v.2006, Localcollectors— 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. 1455385315 [151,1162,1329,1354] Foret de Boulon Burkina Faso 270 10.274166 Comoe 21 -4.4541664 32 33 1 Comoe holotype 1455385302 2006-07-06 CMNC, LORUM Foret de Boulon & F. & S. Genier & F. Genier Burkina Faso 270 10.274166 Primary Foret de Boulon 33 34 2 2 holotype 1455385309 [199,995,1436,1461] Benin Material 33 34 451 159 292 1455385289 1878-07 1878-09-31 1878-07 FGIC, MNHN Abbe Benin 6.25 Agoue 1305 1.6833334 33 34 8 2 2 MONO 1455385346 FGIC, MNHN Abbe Burkina Faso 6.25 Agoue 1305 1.6833334 33 34 44 4 18 1455385338 1977-08 FETC Burkina Faso 12.366667 Ouagadougou 1293 -1.5166667 33 34 1 1 CENTRE paratype 1455385355 PMOC Burkina Faso 12.366667 Ouagadougou 1293 -1.5166667 33 34 1 1 CENTRE paratype 1455385312 2012-08-16 PMOC Burkina Faso 314 13.146167 Kaya 1 -0.8783611 33 34 1 1 CENTRE-NORD paratype 1455385326 2005-08-10 2005-08-15 2005-08-10 PMOC Foret de Sorobouli & Foret Classee de Baporo Burkina Faso 270 11.795556 Foret Classee de Baporo 21 -2.8902776 Foret de Sorobouli 33 34 18 6 3 CENTRE-OUEST paratype 1455385347 2005-06-09 2005-06-12 2005-06-09 PCSR Reserve de Nazinga Burkina Faso 11.15 Reserve de Nazinga 1296 -1.6 33 34 1 1 CENTRE-SUD paratype 1455385322 1988-09-03 1988-09-24 1988-09-03 CMNC F. Genier Burkina Faso 11.073889 Kompienga 21 0.7022222 33 34 132 19 47 EST paratype 1455385330 1988-06-01 1988-06-16 1988-06-01 CMNC Sanborne & Landry Burkina Faso 11.076944 Kompienga 21 0.6805556 33 34 4 2 EST paratype 1455385358 1988-09-03 1988-09-24 1988-09-03 BDGC F. Genier Burkina Faso 11.076944 Kompienga 21 0.6805556 33 34 8 2 2 EST paratype 1455385331 1975-01-11 BMNH Burkina Faso 11.2 Bobo-Dioulasso 1295 -4.3 33 34 1 1 HAUTS-BASSINS paratype 1455385336 1976-11-14 CMNC Burkina Faso 11.2 Bobo-Dioulasso 1295 -4.3 33 34 6 3 HAUTS-BASSINS paratype 1455385349 [267,1155,1904,1929] 2005-06-15 PMOC Burkina Faso 11.083333 Matourkou 1296 -4.366667 33 34 1 1 HAUTS-BASSINS paratype 1455385327 2005-08-07 PMOC Burkina Faso 320 12.862223 Passore 21 -2.3097222 33 34 1 1 NORD paratype 1455385318 2012-05 PMOC Burkina Faso 324 10.899028 Bondigui 15 -3.5708332 33 34 1 1 SUD-OUEST paratype 1455385316 2012-08-20 PMOC Burkina Faso 324 10.899028 Bondigui 15 -3.5708332 33 34 3 2 1 SUD-OUEST paratype 1455385344 MNHN Burkina Faso 34 35 10.316667 Gaoua 1298 -3.1666667 33 34 1 1 SUD-OUEST paratype 1455385337 2006-07-06 CMNC Foret de Boulon & F. & S. Genier Burkina Faso 270 10.274166 Foret de Boulon 21 -4.4541664 34 35 2 1 1 COMOE holotype 1455385345 2006 2017 2006 FGIC Foret de Boulon & F. & S. Genier Burkina Faso 270 10.274166 Foret de Boulon 21 -4.4541664 34 35 4 2 COMOE holotype 1455385343 2006 2024 2006 FGIC Foret de Boulon & F. & S. Genier Burkina Faso 270 10.274166 Foret de Boulon 21 -4.4541664 34 35 1 1 COMOE paratype 1455385329 2005 2039 2005 FGIC F. Genier Burkina Faso 160 11.084722 Kompienga 21 0.71166664 34 35 14 2 5 COMOE paratype 1455385350 2005 2044 2005 FGIC F. Genier Burkina Faso 230 11.283334 Pama 925 0.7163889 34 35 1 1 COMOE paratype 1455385341 2006 2047 2006 FGIC Foret de Nazinga & F. & S. Genier Burkina Faso 285 11.154722 Kouzougou 21 -1.5361111 Foret de Nazinga 34 35 1 1 COMOE paratype 1455385328 2006 2070 2006 FGIC Foret de Nazinga & F. & S. Genier Burkina Faso 285 11.154722 Kouzougou 21 -1.5361111 Foret de Nazinga 34 35 1 1 COMOE paratype 1455385335 2005 2023 2005 FGIC F. Genier Burkina Faso 340 11.184167 Tiakane 21 -1.2077779 34 35 8 1 3 COMOE paratype 1455385314 2005 2034 2005 FGIC F. Genier Burkina Faso 340 11.184167 Tiakane 21 -1.2077779 34 35 1 1 COMOE paratype 1455385372 2006 2064 2006 FGIC F. & S. Genier Burkina Faso 340 11.184167 Tiakane 21 -1.2077779 34 35 1 1 COMOE paratype 1455385363 1979-07 EXTREME, PHWC Cameroon 10.583333 Maroua 1297 14.316667 34 35 6 1 2 NORTH paratype 1455385380 2006-08-18 PMOC Cameroon 11.25 Waza National Park 1295 14.65 34 35 4 2 NORTH paratype 1455385334 1973-10 MNHN Cameroon 9.3 Garoua 1300 13.4 34 35 6 1 2 NORTH paratype 1455385352 OMOC Chad 9.3 Garoua 1300 13.4 34 35 2 1 1 paratype 1455385371 2006-08-11 PMOC Chad 12.883333 Sitje 1292 14.866667 34 35 2 1 1 HADJER-LAMIS paratype 1455385348 1968-06 MNHN Chad 12.116667 Farcha 1293 14.983334 34 35 6 1 2 HADJER-LAMIS paratype 1455385353 1967-06-28 MNHN Chad 12.116667 Farcha 1293 14.983334 34 35 2 1 1 HADJER-LAMIS paratype 1455385382 [402,1432,799,824] PMOC Chad 12.116667 N'Djamena 1293 15.05 Fort Lamy 34 35 1 1 HADJER-LAMIS paratype 1455385356 [151,1337,836,861] 1958-07-17 FETC Chad 12.116667 N'Djamena 1293 15.05 Fort Lamy 34 35 4 2 HADJER-LAMIS paratype 1455385360 2000-06-01 2000-06-05 2000-06-01 PMOC Ranch de Dolla Ivory Coast 8.066667 Biemasso 1302 -7.55 Ranch de Dolla 34 35 32 2 14 paratype 1455385375 [396,1070,908,933] 2003-06-12 Ivory Coast 6.431389 Bringakro 21 -5.0755553 34 35 1 LACS paratype 1455385351 2006-06-13 BMNH Ivory Coast Newman 34 35 10 2 3 LACS paratype 1455385342 vi. 2 BMNH Newman Ivory Coast 6.431389 Bringakro 21 -5.0755553 34 35 12 2 4 LACS paratype 1455385340 2003-06-17 BMNH Newman Ivory Coast 6.431389 Bringakro 21 -5.0755553 34 35 1 1 LACS paratype 1455385368 [270,1432,1015,1040] 1968-08 MNHN Ivory Coast 6.2166667 Lamto 1305 -5.016667 34 35 10 2 3 LAGUNES paratype 1455385365 [151,1125,1052,1077] 1989-02 CEMT Ivory Coast 5.383333 Banco 1305 -4.05 34 35 1 1 Banco paratype 1455385370 IRSNB Ivory Coast 6.65 Dimbroko 1304 -4.7 34 35 1 1 SAVANES paratype 1455385369 2000-06-24 2000-06-25 2000-06-24 PMOC Ivory Coast 9.433611 Korhogo 21 -4.7191668 34 35 1 1 Koko paratype 1455385357 1976-03 PMOC Ivory Coast 7.6833334 Bouake 1303 -5.0333333 34 35 3 2 1 VALLEE DU BANDAMA paratype 1455385374 1976-04 PMOC Ivory Coast 7.6833334 Bouake 1303 -5.0333333 34 35 18 2 7 VALLEE DU BANDAMA paratype 1455385364 1977-07-13 PMOC Ivory Coast 7.6833334 Bouake 1303 -5.0333333 34 35 1 1 VALLEE DU BANDAMA paratype 1455385354 [394,759,1231,1256] Gambia CENTRAL RIVER 34 35 1 CENTRAL RIVER paratype 1455385362 1997-08-23 OUMNH Gambia 13.65 Ballangharr 1290 -15.416667 34 35 28 7 7 Lower Saloum District paratype 1455385339 1997-09-09 OUMNH Gambia 13.666667 Kuntaur Agricultural Station 1290 -14.883333 34 35 1 1 Naini District paratype 1455385396 [999,1197,1304,1328] Gambia LOWER RIVER 34 35 1 LOWER RIVER paratype 1455385386 1997-08-29 OUMNH Mann Gambia 13.480833 Jenoi 21 -15.561945 34 35 8 2 2 Jarra West District paratype 1455385403 1997-08-26 OUMNH D. J. Mann Gambia 13.281388 Mansanjang Kunda 21 -14.203611 Bassa Government Rest Home 34 35 1 1 UPPER RIVER paratype 1455385359 [738,914,1412,1436] Gambia WEST COAST 34 35 1 WEST COAST paratype 1455385377 1997-07 OUMNH Gambia 13.361945 Yundum Agricultural Station 21 -16.661943 34 35 1 1 Kombo District paratype 1455385397 1997-08-19 D. J. Mann Gambia 13.361945 Yundum Agricultural Station 21 -16.661943 34 35 1 Kombo District paratype 1455385373 OUMNH Gambia A 34 35 4 2 A paratype 1455385379 [290,1203,1519,1544] ZMUC Ghana Danish Guinea 34 35 2 1 1 paratype 1455385402 [151,1065,1556,1581] 1883 1884 1883 MNHN Ghana 10.166667 Rio 1298 -14.033334 34 35 1 1 Rio paratype 1455385378 Guinea-Bissau BAFATA 34 35 1 BAFATA paratype 1455385387 [372,1432,1591,1616] MNHN Guinea-Bissau 11.966666 Rio 1294 -14.933333 34 35 8 2 2 Rio paratype 1455385400 [151,895,1628,1653] 1995-06-25 1995-07-17 1995-06-25 Guinea-Bissau 11.533334 Quebo 1295 -14.766666 34 35 1 TOMBALI paratype 1455385376 [927,1432,1628,1652] CEMT Guinea-Bissau A 34 35 1 1 A paratype 1455385367 FGIC, MNHN Mauritania 15.366667 GUIDIMAKA 1285 -12.1 34 35 18 1 8 GUIDIMAKA paratype 1455385366 1976-01-22 PHWC Mauritania 13.426111 Gorou Kirey 21 2.1422224 34 35 1 1 NIAMEY paratype 1455385393 1972-03-15 PHWC Mauritania 12.922778 Kolo 21 1.9566667 34 35 1 1 TILLABERI paratype 1455385390 [266,529,1772,1797] Nigeria KADUNA 34 35 1 KADUNA paratype 1455385394 [542,1432,1772,1797] 1962-05-04 CNC Nigeria 10.523055 Kaduna 21 7.440278 34 35 1 1 Kaduna paratype 1455385395 [151,1213,1807,1832] 1970-05-18 1970-05-25 1970-05-18 BMNH Nigeria 11.166667 Samaru 1295 7.633333 34 35 8 1 3 Kaduna paratype 1455385383 1970-06-15 1970-06-22 1970-06-15 BMNH Nigeria 11.166667 Samaru 1295 7.633333 34 35 1 1 Kaduna paratype 1455385398 1952-05 BMNH W.D. Edmonds Nigeria 11.996389 KANO 21 8.516666 34 35 1 1 KANO paratype 1455385388 [391,1357,1988,2013] 1972-03-19 BMNH Niger 9.05 Badeggi 1300 6.15 34 35 1 1 paratype 1455385381 1957-04-02 BMNH J.-P. Lumaret Niger 35 36 7.366667 Ibadan 1303 3.9333334 34 35 2 1 1 OYO paratype 1455385384 [1101,1355,151,176] Senegal DAKAR 35 36 1 DAKAR paratype 1455385361 MNHN P. Lepesme & R. Paulian Senegal 14.766666 Dakar 1286 -17.366667 35 36 1 1 Dakar paratype 1455385419 MNHN P. Lepesme & R. Paulian Senegal 14.720555 Hann 21 -17.436666 35 36 60 15 15 A paratype 1455385411 2013-08-18 Senegal 14.756945 Keur Ndiaye Lo 21 -17.23639 35 36 1 1 A paratype 1455385430 2009-06-22 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 1 FATICK paratype 1455385385 2009-06-23 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 8 1 3 A paratype 1455385407 2009-06-24 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 10 2 3 A paratype 1455385420 2009-10-16 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 6 3 A paratype 1455385399 2009-11-28 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 1 1 A paratype 1455385405 2010-04-20 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 1 1 A paratype 1455385429 2010-07-15 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 1 1 A paratype 1455385401 2010-07-22 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 2 1 1 A paratype 1455385410 2010-08-13 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 1 1 A paratype 1455385424 2010-08-14 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 2 1 1 A paratype 1455385406 2010-08-15 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 4 2 A paratype 1455385392 2010-08-16 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 1 1 A paratype 1455385391 2010-08-17 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 1 1 A paratype 1455385421 2010-09-21 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 4 2 A paratype 1455385416 2010-09-22 Reserve de Fathala & du Saloum Senegal 13.65 Parc 1290 -16.45 Reserve de Fathala 35 36 8 1 3 A paratype 1455385422 [868,1278,871,896] Senegal A 35 36 1 1 A paratype 1455385418 1948-07-02 CMNC, CNC Senegal 13.896944 Tamba 21 -15.259167 35 36 4 2 KAFFRINE paratype 1455385408 2007-07-16 FGIC F. Genier Senegal 45 13.990556 Missirah 21 -15.117778 35 36 4 2 KAFFRINE paratype 1455385425 2007-07-17 FGIC F. Genier Senegal 45 13.990556 Missirah 21 -15.117778 35 36 1 1 KAFFRINE paratype 1455385414 2008-08-03 PMOC Nioro du Rip Senegal 13.745278 Nioro du Rip 21 -15.773055 35 36 1 1 KAFFRINE paratype 1455385404 2009-08-10 2009-08-11 2009-08-10 PMOC Nioro du Rip Senegal 13.745278 Nioro du Rip 21 -15.773055 35 36 2 1 1 KAFFRINE paratype 1455385409 2008-07-13 2008-07-14 2008-07-13 PMOC Senegal 12.378612 Dinndefelou 21 -12.325 35 36 1 1 KEDOUGOU paratype 1455385389 2008-07-31 2008-08-02 2008-07-31 PMOC Foret de Bakor Senegal 12.85 Mahon 1292 -14.816667 Foret de Bakor 35 36 1 1 KOLDA paratype 1455385440 2009-11-22 2009-11-23 2009-11-22 PMOC Senegal 12.856112 Mahon 21 -14.8116665 35 36 10 2 3 KOLDA paratype 1455385433 2007 2049 2007 FGIC F. Genier & P. Moretto Senegal 31 12.737778 Sare Gnako 21 -15.165555 35 36 4 2 KOLDA paratype 1455385455 2007 2047 2007 FGIC Genier & Moretto Senegal 23 12.6652775 Sothoto 21 -15.400833 35 36 4 2 KOLDA paratype 1455385413 1967-09 Senegal 15.4 Linguere 1285 -15.116667 35 36 1 LOUGA paratype 1455385436 1967-11 MNHN Descarpentries, T. Senegal 15.606112 Ndam-Dam 21 -16.393055 35 36 32 10 6 A paratype 1455385449 MNHN Descarpentries, T. Senegal A 35 36 5 4 1 A paratype 1455385423 1996-10 PSCW Oiseaux du Djoudj Senegal 16.4 Oiseaux du Djoudj 1282 -16.233334 Parc National 35 36 8 1 3 SAINT-LOUIS paratype 1455385428 1967-11 MNHN Descarpentries, T. Senegal 16.466667 Richard-Toll 1282 -15.683333 35 36 106 30 23 A paratype 1455385453 [853,979,1412,1436] Senegal SEDHIOU 35 36 1 SEDHIOU paratype 1455385426 MNHN Senegal 12.7 Sedhiou 1292 -15.566667 35 36 1 1 Sedhiou paratype 1455385434 1947-07-14 MNHN Senegal 12.7 Sedhiou 1292 -15.566667 35 36 1 1 Sedhiou paratype 1455385450 2007-07-20 FGIC F. Genier Senegal 52 13.804444 Boutou 21 -13.579722 35 36 1 1 TAMBACOUNDA paratype 1455385435 1995-07-29 PMOC Senegal 13.213056 Dienoun Diala 21 -13.112223 35 36 24 8 4 TAMBACOUNDA paratype 1455385417 2007 2056 2007 FGIC F. Genier Senegal 18 12.444722 Boukitimgo 21 -16.598057 35 36 1 1 paratype 1455385415 2007 2058 2007 FGIC P. Moretto & F. Genier Senegal 18 12.444722 Boukitimgo 21 -16.598057 35 36 1 1 paratype 1455385445 1963-05-23 MNHN Mission & Monts Sierra Leone 9.535833 Koinadugu 21 -11.368334 35 36 10 1 4 NORTHERN paratype 1455385439 2006-05 PMOC Local Uganda 2.4069445 Padyere 21 31.194443 Pamora 35 36 2 1 1 Nebbi District paratype