Digitonthophagus biflagellatus 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 : 37-39

Digitonthophagus biflagellatus Génier , new species

http://zoobank.org/urn:lsid:zoobank.org:act:C93B1EB9-87CE-4E82-9017-F6D5D946DDFB ( Figs. 13 , 35–36 , 61 , 77 , 99–100, 125, 153–155; Map 6)

Type locality. Lake Ndutu area (02°59’S 35°01’E), Serengeti National Park, Shinyanga, Tanzania.

Diagnosis. Axial sclerite as long as or longer than subaxial sclerite ( Fig. 153 ).

Description. Holotype ♂ ( Fig. 13 ). Measurements. Length 11.0 mm, width 6.5 mm. Head ( Figs. 35–36 ). Anterior clypeal edge straight on median fifth 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, 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 broad angle with clypeal edge. Pronotum ( Fig. 61 ). 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 convex; anterior half of lateral edge arcuate in dorsal and lateral view; posterior angles simply arcuate in dorsal view. Anterior hypomeral ridge arcuate, anterior hypomeral depression surface slightly darker in color medially. Elytra ( Fig. 13 ). Intervals 2 and 4 lacking fine granules from base to apex. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus ( Fig. 125 ). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites ( Figs. 153–155 ). Axial sclerite weekly sclerotized, longer than subaxial sclerite, and almost straight. Subaxial sclerite, large basally, and abruptly reduced, extending straight and interrupted before apex of frontolateral peripheral sclerite apical portion, with fine and scattered villi on apical 10th. Frontolateral peripheral sclerite basoventral apophysis moderately developed; lacking medioventral carinae; right lateral fold produced into a rather large everted and open apically conical process, apical edge deeply emarginate externally and open internally, appearing tong shaped; left lateral lobe absent; subapicodorsal lobe membranous, not reaching anterior edge, apex set on right side in dorsal view; apical lobe round apically, narrow and directed obliquely on left side, apical villi regular in shape; subapicoventral lobe large, round and interrupted in line with apical lobe apically.

Variation. Measurements (94 ♂♂, 69 ♀♀). Length: male 7.5–11.5 mm (9.8 ± 0.9 mm), female 8.0–12.0 mm (9.8 ± 0.7 mm). Female allotype. Cephalic outline in dorsal view as in Fig. 77 ; vertex with a slightly sinuate, transverse carina, dorsal edge broadly arcuate in frontal view, lateral portion gradually sloping down posteriorly; anterior pronotal tubercles moderately developed, external lateral edges widening toward apex, anterolateral surface simply convex, anterosuperior edge slightly arcuate in dorsal view (Fig. 99), lateral portion of anterosuperior edge distinctly upturned (Fig. 100). Protibia short, with external teeth more robust.

Primary type data. Holotype ♂ ( OUMNH): [18 /05 /91 / L. Ndutu area, / Serengeti National Park / Tanzania, 35°1’E 2°59’S / dung-baited pitfall trap / R. Foster O.U.M. 2-1993] partly handwritten; [ WORLD / SCARAB. / DATABASE / WSD00030055] barcode label ; HOLOTYPE ♂ / Digitonthophagus / biflagellatus n.sp. / des. F. Génier, 2016] red card.

Material examined (135 ♂♂, 82 ♀♀), distribution (Map 8): BURUNDI: [unspecified locality], [no date], R.P. Giraudin— 1 ♀, 4 ♂♂ (5 paratypes) ( MNHN) ; BURURI, Bururi (3°54'S, 29°31'E), [no date], R.P. Giraudin— 1 ♀, 1 ♂ (paratypes) ( MNHN) ; Muyinga, Colline Kobero (2°38'59''S, 30°24'48''E), 10.ix.1940, H.-J. Brédo— 1 ♂ (paratype) ( IRSNB) ; RUTANA, 03°45’S 30°10’E, 2000 m (3°45'S, 30°10'E), iii.1968, R.P. Giraudin— 1 ♀, 1 ♂ (paratypes) ( OMOC) ; Mpinga (3°45'S, 30°10'E), [no date], R.P. Giraudin— 1 ♂ (paratype) ( MNHN) ; DEMOCRATIC REPUBLIC OF THE CONGO: NORTH-KIVU, Parc National Rwindi [= SE Parc National Virunga] (1°23'S, 29°28'E), 26.iii.1979, [anonymous]— 1 ♂ (paratype) ( PHWC) ; Rutshuru (1°11'S, 29°27'E), 1927, G. Babault— 2 ♀♀, 1 ♂ (paratypes) ( MNHN) ; SOUTH-KIVU, Bukavu (2°30'S, 28°52'E), 30.i.1952, C.

Verbeke—3 ♂♂ (3 paratypes) (IRSNB); KENYA: HOMA BAY, Baie de Kavirondo , Lac Victoria (0°31'S, 34°29'E), ix–x.1903, C. Alluaud— 2 ♂♂ (2 paratypes) ( MNHN); Baie de Kavirondo , Lac Victoria (0°31'S, 34°29'E), xii.1911, Alluaud & Jeannel— 1 ♂ (paratype) ( MNHN) ; Ile de Lusinga [= Rusinga ], Victoria-Nyanza N. - E. (0°24'S, 34°11'E), x.1903, C. Alluaud— 1 ♂ (paratype) ( IRSNB) ; KAJIADO, 60 km SW Nairobi , 1200 m (1°45'S, 36°21'E), 23.xii.1990, B.D. Gill— 2 ♀♀, 1 ♂ (paratypes) ( BDGC) ; KAKAMEGA, Ilala , 14 mi. E Mumias, 1372 m (0°17'N, 34°41'E), 18–21.vi.1911, S . A. Neave— 1 ♂ (paratype) ( BMNH) ; KISUMU , Kisumu (0°6'S, 34°46'E), [no date], P.S. Corbet— 2 ♀♀, 4 ♂♂ (6 paratypes) ( BMNH) ; MURANG’A, Fort Hall [= Muranga Town] (0°43'S, 37°9'E), i.1912, Alluaud & Jeannel— 1 ♂ (paratype) ( MNHN) ; NAIROBI, Nairobi (1°17'S, 36°49'E), 24.iv.1957, [anonymous]— 2 ♀♀, 2 ♂♂ (4 paratypes) ( BMNH) ; Nairobi National Park (1°22'S, 36°51'E), iv.1993, W. Hoek— 1 ♂ (paratype) ( CEMT) ; NANDI, Kamililo , vi.1903, F.J. Jackson— 1 ♂ (paratype) ( BMNH) ; NAROK, Lemek Valey (1°6'S, 35°23'E), ii.1913, G.B. — 6 ♀♀, 8 ♂♂ (14 paratypes) ( FGIC, MNHN) ; TAITA TAVETA, Bura (3°2'S, 38°19'E), i–iv.1904, C. Alluaud— 5 ♂♂ (5 paratypes) ( IRSNB, MNHN) ; Mwatate (3°30'S, 38°22'E), i– iv.1904, C. Alluaud— 1 ♀, 3 ♂♂ (4 paratypes) ( IRSNB, MNHN) ; Mwatate , 25 km SW Voi (3°30'S, 38°22'E), 26.xii.1990, B.D. Gill— 2 ♀♀, 2 ♂♂ (4 paratypes) ( BDGC) ; VIHIGA, Forêt de Kakamegoes [= Kakamega], Environs de Kaimosi, Mont-Elgon , 2000 m (0°10'N, 34°52'E), 1913, G. Babault— 6 ♂♂ (6 paratypes) ( FGIC, MNHN) ; RWANDA: EASTERN, Ranch Mpanga (2°3'13''S, 30°47'0''E), iii.1987, [anonymous]— 2 ♀♀, 3 ♂♂ (5 paratypes) ( PSCW) ; NORTHERN, Bugarula District , [unspecified locality] (1°33'S, 29°42'E), 21.viii.1941, H.-J. Brédo— 2 ♀♀, 3 ♂♂ (5 paratypes) ( IRSNB) ; WESTERN, Rwaza (1°42'S, 29°16'E), 1927, G. Babault— 2 ♂♂ (2 paratypes) ( MNHN) ; TANZANIA: Lake Rukwa, iv.1945, H.-J. Brédo— 1 ♀, 2 ♂♂ (3 paratypes) ( FGIC, IRSNB) ; Lake Rukwa, xii.1948, H.-J. Brédo— 1 ♀, 3 ♂♂ (4 paratypes) ( IRSNB) ; ARUSHA , Arusha environs (3°22'S, 36°41'E), 3–4.iv.1997, M. Kuboň— 5 ♀♀, 1 ♂ (paratypes) ( PMOC) ; Karatu Ndutu Safari Lodge, Serengeti (3°1'S, 35°0'E), 15.xi.1990, R. Foster— 1 ♂ (paratype) ( BDGC) ; Mumba Hills (3°30'S, 35°3'E), 8.iii.1938, Dr. S.G. Kohl- Larsen— 1 ♀ (paratype) ( SMF) ; Mumba Hills (3°30'S, 35°3'E), 23.iii.1938, Dr. S.G. Kohl-Larsen— 1 ♂ (paratype) ( SMF) ; Mumba Hills (3°30'S, 35°3'E), 24.iii.1938, Dr. S.G. Kohl-Larsen— 1 ♂ (paratype) ( SMF) ; Orekeryan, Mount Longido (2°43'47''S, 36°43'26''E), 8.vi–9.viii.2012, Smith, Takano & Garner— 1 ♂ (paratype) ( BMNH) ; DODOMA, 40 km N Dodoma, 1100 m (5°54'S, 35°45'E), 15.xii.2007, F. Kantner— 5 ♀♀, 3 ♂♂ (8 paratypes) ( SMNS) ; 70 km N Dodoma, 1330 m (5°40'S, 35°48'E), 17.xii.2006, F. Kantner— 1 ♂ (paratype) ( SMNS) ; Lulanguru , 17 mi. W Tabora, 1148 m (5°6'S, 32°38'E), x–xii.1917, G.D. H. Carpenter— 1 ♀, 1 ♂ (paratypes) ( BMNH) ; Tosamaganga , near Iringa (6°40'S, 36°19'E), [no date], W. Ruddock— 1 ♂ (paratype) ( BMNH) ; IRINGA, Magema, Ulembwe Village , 2048 m (9°18.389'S, 34°37.213'E), 7.i.2011, P. Darge— 1 ♂ (paratype) ( MHNL) ; KILIMANDJARO, Same (4°4'S, 37°45'E), v.1962, [anonymous]— 2 ♂♂ (2 paratypes) ( CNC) ; Zone inférieure, Kilimandjaro, i–iv.1904, C. Alluaud— 1 ♀, 3 ♂♂ (4 paratypes) ( MNHN) ; MANYARA, 100 km NNE Singida, 1650 m (4°31'S, 35°21'E), 20.xii.2006, F. Kantner— 5 ♀♀, 2 ♂♂ (7 paratypes) ( SMNS) ; Giting, Mount Hanang , 1964 m (4°24'14''S, 35°24'56''E), 22–26.xi.2011, R. Smith & H. Takano— 8 ♀♀, 2 ♂♂ (10 paratypes) ( BMNH) ; MARA, Shirati (1°9'S, 34°2'E), 1909, [anonymous]— 5 ♀♀, 1 ♂ (paratypes) ( SMF) ; MOROGORO, Llonga (9°2'S, 36°51'E), 21.x.1966, Robertson— 1 ♀, 2 ♂♂ (3 paratypes) ( FETC) ; Majawanga , 1400 m (6°5'S, 36°50'E), 14–18.iii.2004, S. Tind Nielsen— 1 ♀, 3 ♂♂ (4 paratypes) ( ZMUC) ; RUKWA, Tumba (7°7'S, 31°10'E), 15.xii.1950, H.-J. Brédo— 1 ♀, 2 ♂♂ (3 paratypes) ( FGIC, IRSNB) ; SHINYANGA , Lake Ndutu area, Serengeti National Park (2°59'S, 35°1'E), 15.xi.1990, R. Foster— 3 ♀♀, 2 ♂♂ (allotype, 4 paratypes) ( OUMNH) ; Lake Ndutu area, Serengeti National Park (2°59'S, 35°1'E), 17.xii.1990, R. Foster— 3 ♀♀, 1 ♂ (paratypes) ( OUMNH) ; Lake Ndutu area, Serengeti National Park (2°59'S, 35°1'E), 18.xii.1990, R. Foster— 1 ♂ (paratype) ( OUMNH) ; Lake Ndutu area, Serengeti National Park (2°59'S, 35°1'E), 22.xii.1990, R. Foster— 1 ♂ (paratype) ( OUMNH) ; Lake Ndutu area, Serengeti National Park (2°59'S, 35°1'E), 26.xii.1990, R. Foster— 1 ♂ (paratype) ( OUMNH) ; Lake Ndutu area, Serengeti National Park (2°59'S, 35°1'E), 18.v.1991, R. Foster— 2 ♂♂ (holotype, 1 paratype) ( OUMNH) ; Old Shinyanga (3°34'S, 33°25'E), 1.xii.1935, E. Burtt— 1 ♂ (paratype) ( BMNH) ; Old Shinyanga (3°34'S, 33°25'E), 26.i.1936, E. Burtt— 1 ♂ (paratype) ( BMNH) ; Old Shinyanga (3°34'S, 33°25'E), 10.iv.1936, E. Burtt— 1 ♀ (paratype) ( BMNH) ; TABORA, Tabore (5°2'S, 32°49'E), 1886, Révoil— 1 ♂ (paratype) ( MNHN) ; UGANDA: CENTRAL, Entebbe (0°4'N, 32°28'E), ii–vii.1972, H. Falke— 10 ♀♀, 9 ♂♂ (19 paratypes) ( CNC, FGIC) ; Entebbe (0°4'N, 32°28'E), ii.1972, H. Falke— 2 ♂♂ (2 paratypes) ( CNC) ; Entebbe (0°4'N, 32°28'E), 22.vi.1912, C . A. Wiggins— 1 ♀ (paratype) ( OUMNH); Entebbe (0°4'N, 32°28'E), 15.viii.1912, C . A. Wiggins— 1 ♂ (paratype) ( OUMNH) ; Entebbe (0°4'N, 32°28'E), 9.ii.1914, [anonymous]— 1 ♂ (paratype) ( BMNH) ; Entebbe (0°4'N, 32°28'E), 11.ii.1914, [anonymous]— 1 ♀ (paratype) ( BMNH) ; Entebbe (0°4'N, 32°28'E), 20–28.v.1914, C.C. Gowdey— 2 ♂♂ (2 paratypes) ( BMNH) ; Kampala (0°21'N, 32°35'E), 1–10.i.1918, C.C. Gowdey— 1 ♀, 5 ♂♂ (6 paratypes) ( BMNH) ; Kampala (0°21'N, 32°35'E), 11.ii.1918, [anonymous]— 1 ♀, 1 ♂ (paratypes) ( BMNH) ; Région du Bouganda (0°27'S, 31°55'E), 1906, [anonymous]— 1 ♂ (paratype) ( IRSNB) ; [unspecified locality] (0°15'N, 32°15'E), i–ii.1909, C. Alluaud— 1 ♀, 3 ♂♂ (4 paratypes) ( MNHN); EASTERN , Jinja, 1151 m (0°26'N, 33°12'E), 14.ii.1906, C . A. Wiggins— 1 ♂ (paratype) ( OUMNH) ; WESTERN, Kafu River near Hoima , Kampala Road (1°18'N, 31°27'E), 29–31.xii.1911, S . A. Neave— 1 ♂ (paratype) ( BMNH) ; Province d’Unyoro, Région de l’Albert-Nyanza (1°17'N, 31°10'E), ii.1909, C. Alluaud— 1 ♂ (paratype) ( MNHN) ; ZAMBIA: NORTHERN PROVINCE, Mbala (8°50'S, 31°28'E), 15.xii.1943, H.-J. Brédo— 2 ♂♂ (2 paratypes) ( IRSNB) .

MAP 8. Distribution of Digitonthophagus biflagellatus .

Etymology. Biflagellatus is a Latin adjective referring to the configuration of the axial and subaxial sclerites of the internal sac.

Natural history. The single specimen with habitat data was collected in woodland in Eastern Province, Uganda. The few specimens with collecting data indicate light and dung traps.

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 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).

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.