Digitonthophagus viridicollis 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 : 54-56

Digitonthophagus viridicollis Génier , new species

http://zoobank.org/urn:lsid:zoobank.org:act:11622669-B240-4DF1-AF7B-0757 A 587DCEF ( Figs. 16 , 41–42 , 64 , 80 , 105–106, 128, 162–164, 186; Map 11)

Type locality. Moanda, Zaïre [= 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 / (Zaïre) 30.iv.1979] partly handwritten; [WORLD / SCARAB. / DATABASE / WSD00032092] barcode label; [HOLOTYPE ♂ / Digitonthophagus / viridicollis n.sp. / des. F. Génier, 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, Cossoço [= Coçoço ] (11°24'S, 14°1'E), iii.2005, [anonymous] —2 ♀♀, 3 ♂♂ (5 paratypes) (PMOC); Cossoço [=Coçoço] (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); Roçadas (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); João 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, Mayumbé (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 Süd 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, Külz— 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.

Figures

FIGURES 15 – 18. Male dorsal habitus. 15. D. petilus, holotype; 16. D. viridicollis, holotype; 17. D. aksumensis, holotype; 18. D. dilatatus, holotype.

FIGURES 39 – 46. 39, 41, 43, 45. Male head, dorsal view; 40, 42, 44, 46. Male head, frontal view. 39 – 40. Digitonthophagus petilus, holotype; 41 – 42. D. viridicollis, holotype; 43 – 44. D. aksumensis, holotype; 45 – 46. D. dilatatus, 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 159 – 170. Internal sac sclerites (FLP, A + SA). 159, 162, 165, 168. Dorsal view; 160, 163, 166, 169. Ventral view. 161, 164, 167, 170. Right lateral view. 159 – 161. Digitonthophagus petilus, holotype; 162 – 164. D. viridicollis, holotype; 165 – 167. D. aksumensis, holotype; 168 – 170. D. dilatatus, holotype.

FIGURES 183 – 190. Digitonthophagus bonasus, female head, frontal view (Phongsaly Province, Laos); 184. D. aksumensis, FLP sclerite, right lateral view; 185. D. gazella, right lateral fold, ventral view; 186. D. viridicollis, right lateral fold, ventral view; 187. D. eucatta, hypomeron, ventral view, A. anterior hypomeral carina, B. anterior hypomeral depression; D. gazella, hypomeron, ventral view, A. anterior hypomeral carina, B. anterior hypomeral depression; 189. D. namaquensis, vertex dorsal view; 190. D. gazella, vertex dorsal view.

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.