Digitonthophagus ulcerosus 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 : 68-70

Digitonthophagus ulcerosus Génier , new species

http://zoobank.org/urn:lsid:zoobank.org:act:CEBB9AB5-F6F4-4BDF-A6C1-037C15BBBD24 ( 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. Génier, 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], Révoil—3 ♂♂ (3 paratypes) (MNHN); SANAAG, Ouarsangueli [=Warsangeli Sultanate] (11°0'N, 48°0'E), 1881, Révoil— 1 ♀, 2 ♂♂ (holotype, allotype, 1 paratype) ( MNHN) ; TANZANIA: KILIMANDJARO , Rivière Himo, zone inférieure, 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.

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.