Digitonthophagus bonasus ( Fabricius, 1775 )

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 : 15-19

Digitonthophagus bonasus ( Fabricius, 1775)

( Figs. 7 , 23–24 , 55 , 71 , 87–88, 119, 135–137, 183; Map 2)

Scarabaeus Bonasus Fabricius, 1775: 23 (original description) Scarabaeus Bonasus — Goeze 1777: 69 (diagnosis)

Scarabaeus Bonasus — Fabricius 1781: 26 (redescription) Scarabaeus bonasus — Pallas 1781: 5 (redescription)

Scarabaeus Bonasus — Herbst 1786: 153 (diagnosis, distribution) Scarabaeus Bonasus — Fabricius 1787a: 14 (diagnosis)

Scarabaeus Bonasus — Olivier 1789: 121 (redescription)

Scarabaeus Bonasus — Herbst 1789 2: 180 (monograph)

Scarabaeus Bonasus — Gmelin 1790: 1541 (diagnosis)

Scarabaeus Bonasus — Fabricius 1792: 50 (diagnosis)

Scarabaeus Bonasus — Illiger 1800: 228 (monograph)

Copris Bonasus — Fabricius 1801: 40 (diagnosis)

Copris Bonasus — Illiger 1804: 149 (comment taxonomy)

Copris Bonasus — Schönherr 1806: 42 (catalog)

Ontophagus Bonasus — Sturm 1826: 177 (catalog)

Onthophagus bonasus — Latreille 1827: 282 (misidentification)

Onthophagus bonasus — LePeletier & Audinet-Serville 1828: 353 (identification key, comment) Onthophagus Bonasus — Hope 1837: 32 (catalog)

Ontophagus bonasus — Laporte 1840: 85 (diagnosis, distribution) Onthophagus Bonasus — Sturm 1843: 107 (catalog)

Onthophagus bonasus — Motschulsky 1860: 154 (comment taxonomy) Onthophagus bonasus — Motschulsky 1863: 457 (checklist)

Onthophagus bonasus — Harold 1869: 1026 (catalog)

Onthophagus (O.) bonasus — d’Orbigny 1900a: 218 (monograph) Onthophagus bonasus — Boucomont 1914: 241 (distribution)

Onthophagus bonasus — Boucomont 1921: 29 (identification key, distribution) Onthophagus bonasus — Gillet & Boucomont 1927: 133 (catalog) Onthophagus bonasus — Arrow 1931: 231 (monograph)

Onthophagus (O.) bonasus — Paulian 1945: 101 (monograph)

Onthophagus bonasus — Petrovitz 1955: 277 (distribution)

Onthophagus bonasus — Balthasar 1956: 435 (distribution)

Onthophagus (Digitonthophagus) bonasus — Balthasar 1959: 464 (genotype designation) Onthophagus (Digitonthophagus) bonasus — Balthasar 1963: 296 (monograph) Digitonthophagus bonasus — Zunino 1981: 413 (new combination) Onthophagus (Matashia) bonasus — Masumoto 1988: 140 (distribution) Digitonthophagus bonasus — Zunino & Halffter 1988: 20 (comment taxonomy) Digitonthophagus bonasus —Barbero & Palestrini 2003: 333 (phylogeny) Digitonthophagus bonasus —Cabral-de-Mello et al. 2008: 1248 (chromosomal formula) Digitonthophagus bonasus — Tarasov & Solodovnikov 2011: 4 (phylogeny) Digitonthophagus bonasus — Génier 2013: 2 (comment taxonomy) Digitonthophagus bonasus — Roggero et al. 2016a: [2] (phylogeny) Digitonthophagus bonasus — Roggero et al. 2016b: 2 (phylogeny) Digitonthophagus bonasus — Breeschoten et al. 2016: 90 (phylogeny)

Type locality. Tranquebariae [= Tharangambadi, Tamil Nadu State, India].

Diagnosis. Male and female pronotal anteromedian tubercle similar, well developed, forming an angular projection in lateral view (Fig. 88); pronotal anterior median longitudinal sulcus wide and deep ( Figs. 55 , 87).

Description. Alloreferent ♂ ( Fig. 7 ). Measurements. Length 14.5 mm, width 9.0 mm. Head ( Figs. 23–24 ). Anterior clypeal edge slightly sinuous on median seventh in dorsal view; clypeofrontal carina broadly arcuate and interrupted at gena; vertex with an acute median conical process higher than wide basally, surface with dense irregular granules. Horns long, obliquely divergent in frontal view, gradually tapering from base to apex, posterointernal edge unmodified basally, internal surface coarsely and densely granulated; genal edge slightly upturned and arcuate on anterior third, forming a broad angle with clypeal edge. Pronotum ( Fig. 55 ). Surface with granules restricted to median third, lacking distinct umbilicate punctures, punctures weaker but distinct posterolaterally with distinct, minute punctures throughout. Anteromedian tubercle well developed, forming an angular projection in lateral view and forming two short blunt longitudinal carinae in dorsal view; median longitudinal sulcus wide and deep; surface behind the eyes with a small round depression; surface of anterior angles sloping laterally; anterior half of lateral edge strongly sinuous in dorsal and lateral view; posterior angles produced into a obliquely-oriented tooth in dorsal view. Anterior hypomeral ridge broadly arcuate anteriorly with anterior most portion crenulate, anterior hypomeral depression surface slightly darker in color. Elytra ( Fig. 7 ). Intervals 2 and 4 with few fine granules on apical fourth. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus ( Fig. 119 ). Parameres with dorsal and ventral edges diverging toward apex in lateral view. Internal sac sclerites ( Figs. 135–137 ). Axial sclerite strongly sclerotized, crescent shaped. Subaxial sclerite extending straight approximately in line with apex right lateral fold apical portion, with fine villi.

Frontolateral peripheral sclerite basoventral apophysis moderately developed; two medioventral carinae present; right lateral fold produced into a rather large everted and open apically conical process with extremely irregular apical edge; left lateral lobe absent; subapicodorsal lobe membranous, narrow, not reaching anterior edge, apex set medially in dorsal view; apical lobe round and directed obliquely on left side, apical villi regular in shape; subapicoventral lobe short not reaching apical edge of apical lobe, oriented obliquely.

Variation. Measurements (111 ♂♂, 103 ♀♀). Length: male 11.0– 16.5 mm (13.9 ± 1.3 mm), female 10.0–16.0 mm (13.4 ± 1.2 mm). Female lectotype. Cephalic outline in dorsal view similar to Fig. 71 ; vertex with horns wide basally and regularly tapering to apex, posterointernal edge with an acute tooth basally; anterior pronotal configuration (Figs. 87–88) similar to males. Protibia short, with external teeth more robust. A number of females from various localities in Asia east of India present much shorter cephalic horns. The horns in these females ( Fig. 183 ) are wide and taper abruptly just before the apex, which is internally diverging in frontal view. Males collected with these females are identical to typical male D. bonasus and we consider this variation as intraspecific.

Primary type data. Lectotype ♀ (ZMUK) present designation: [ZMUC / 00021685]; WORLD / SCARAB. / DATABASE / WSD00032218] barcode label; [LECTOTYPE ♀ / Scarabaeus / bonasus / Fabricius, 1775 / dés. F. Génier, 2016] red card.

MAP 2. Distribution of Digitonthophagus bonasus .

Material examined (169 ♂♂, 164 ♀♀, 377 sex not recorded), distribution (Map 2): BANGLADESH : CHITTAGONG, Chittagong Hill Tracts (22°20'N, 92°13'E), 1883, H.M. Parish— 1 ♀, 1 ♂ ( BMNH) ; DHAKA , Dacca [= Dhaka] (23°42'N, 90°22'30''E), 5.v.1945, D. Leston— 1 ♀ ( BMNH) ; CAMBODIA: KOH KONG, 20 km SE Koh Kong, Tatai river , 50–300 m (11°34'N, 103°7'E), 17.x–15.xi.2011, Z. Linek—2 ♀♀, 2 ♂♂ (FGIC); PHNOM PENH, Kompong Luong (11°50'N, 104°48'E), [no date], [anonymous]— 1 ♀ ( MNHN) ; SIEM REAP, Kompong Chikreng (13°5'N, 104°18'E), 1912, G. Thomas— 1 ♂ ( MNHN) ; INDIA: Côte de Malabar, 1900, T. Deschamps— 4 ♀♀, 6 ♂♂ ( MNHN) ; HIMACHAL PRADESH, Near Kullu, Pirpandzhal range, 1500 m (31°57'30''N, 77°6'30''E), 21.vii.2003 , A. Gorodinski— 1 ♂ (CMD); Yol, Kangra Valley , 1300 m (32°10'N, 76°12'E), viii–ix.1943, C. Lomi—1 ♀, 1 ♂ (CMNC); KARNATAKA, Kanara [=Coastal Karnataka], [no date], [anonymous]— 2 ♂♂ ( BMNH) ; Belgaum District, Belgaum (15°51'N, 74°33'0''E), [no date], [anonymous]— 1 ♀, 3 ♂♂ ( BMNH) ; Kodagu District, Mercara [= Madikeri ] (12°25'14''N, 75°44'23''E), v.1973, T.R.S. Nathan— 1 ♂ ( RMNH) ; Mysore District, Mysore (12°18'N, 76°39'E), [no date], [anonymous]— 1 ♀, 1 ♂ ( BMNH) ; Shimoga District, Bhadravati (13°50'24''N, 75°42'7.2''E), vii.1938, [anonymous]— 1 ♀ ( FETC) ; Shimoga District, Bhadravati (13°50'24''N, 75°42'7.2''E), 13.v.1935, [anonymous]— 1 ♀ ( CMNC) ; Shimoga (13°55'N, 75°34'E), vi.1897, [anonymous]— 1 ♀, 1 ♂ ( MNHN) ; Shimoga (13°55'N, 75°34'E), [no date], [anonymous]— 1 ♂ ( NMPC); Thirthahalli , 700 m (13°42'N, 75°13'48''E), 14.xii.1989, A. Kumar— 2 ♀♀ ( OUMNH) ; KERALA, Kumily (9°37'6''N, 77°9'29''E), v.1986, T.R.S. Nathan— 1 ♀, 1 ♂ ( CMNC) ; Kumily (9°37'6''N, 77°9'29''E), vi.1986, T.R.S.

Nathan—1 ♂ (CMNC); KERALA / TAMIL NADU, Anaimalai Hills , 1067 m (10°22'N, 77°7.5'E), iv.1963, P.S. Nathan— 3 ♀♀, 3 ♂♂ ( CNC) ; ORISSA, Balasore District, Balasore (21°30'N, 86°56'E), [no date], R.P. Gengler— 1 ♀, 2 ♂♂ ( MNHN) ; PONDICHERRY, Karaikal District, Karaikal (10°56'N, 79°50'E), iii.1963, P.S. Nathan— 91 specimens, 1 ♀ ( BDGC, CNC) ; PONDICHERRY, Karaikal District, Karaikal (10°56'N, 79°50'E), iv.1963, P.S. Nathan— 20 specimens, 1 ♀ ( BDGC, CNC) ; Karaikal (10°56'N, 79°50'E), ix.1969, P.S. Nathan— 1 ♀ ( CMNC) ; Karaikal (10°56'N, 79°50'E), iii.1968, T.R.S. Nathan— 1 ♀ ( CMNC) ; Karaikal (10°56'N, 79°50'E), v.1988, T.R.S. Nathan— 1 ♀ ( CMNC) ; Karaikal (10°56'N, 79°50'E), viii.1991, T.R.S. Nathan— 1 ♀, 1 ♂ ( CMNC) ; Karaikal (10°56'N, 79°50'E), xii.1961, P.S. Nathan— 3 ♀♀, 2 ♂♂ ( CNC) ; Karaikal (10°56'N, 79°50'E), i.1962, P.S. Nathan— 5 ♀♀, 5 ♂♂ ( CNC) ; Karaikal (10°56'N, 79°50'E), ii.1962, P.S. Nathan— 5 ♀♀, 5 ♂♂ ( CNC) ; Karaikal (10°56'N, 79°50'E), iii.1962, P.S. Nathan— 5 ♀♀, 5 ♂♂ ( CNC) ; Karaikal (10°56'N, 79°50'E), vi.1962, P.S. Nathan— 1 ♀ ( CNC) ; Karaikal (10°56'N, 79°50'E), iv.1962, P.S. Nathan— 14 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), v.1962, P.S. Nathan— 2 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), vii.1962, P.S. Nathan— 45 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), viii.1962, P.S. Nathan— 21 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), ix.1962, P.S. Nathan— 63 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), x.1962, P.S. Nathan— 42 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), xii.1962, P.S. Nathan— 5 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), i.1963, P.S. Nathan— 36 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), ii.1963, P.S. Nathan— 18 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), vi.1963, P.S. Nathan— 8 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), x.1963, P.S. Nathan— 4 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), x.1966, P.S. Nathan— 4 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), iii.1967, P.S. Nathan— 4 specimens ( CNC) ; Karaikal (10°56'N, 79°50'E), [no date], [anonymous]— 1 ♀, 2 ♂♂ ( FETC) ; Karaikal (10°56'N, 79°50'E), i–ii.1964, T.R.S. Nathan— 4 ♀♀, 1 ♂ ( MMUE) ; Karaikal (10°56'N, 79°50'E), viii.1963, T.R.S. Nathan— 1 ♂ ( MMUE) ; Karaikal (10°56'N, 79°50'E), v.2005, Nathan— 1 ♀, 1 ♂ ( PMOC) ; Karaikal (10°56'N, 79°50'E), v.1973, T.R.S. Nathan— 1 ♂ ( RMNH) ; Kurumbagaram (10°59'N, 79°46'E), xii.1950, P.S. Nathan— 1 ♂ ( CMNC) ; Nedungadu (10°58'N, 79°46'E), v.1938, [anonymous]— 1 ♂ ( FETC) ; Nedungadu (10°58'N, 79°46'E), [no date], [anonymous]— 1 ♂ ( FETC) ; Nedungadu (10°58'N, 79°46'E), 1936, P.S. Nathan— 14 ♀♀, 10 ♂♂ ( MNHN) ; Nedungadu (10°58'N, 79°46'E), 1.vi.1931, P.S. Nathan— 1 ♂ ( BMNH) ; PUNJAB, Rormazara, Hoshiarpur (31°11'50.7''N, 76°12'58''E), 6–9.vii.2004, M. Uhler— 2 ♀♀ ( CVM) ; TAMIL NADU, Arcot (12°54'N, 79°19'E), ix.2012, [anonymous]— 1 ♀ (CMD); Coimbatore , 427 m (11°0'N, 76°57'E), iv.1962, P.S. Nathan— 10 ♀♀, 12 ♂♂ ( BDGC, CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), ix.1962, P.S. Nathan— 1 ♀ ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), x.1962, P.S. Nathan— 1 ♀, 4 ♂♂ ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), xi.1962, P.S. Nathan— 1 ♀, 2 ♂♂ ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), xii.1962, P.S. Nathan— 1 ♀, 3 ♂♂ ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), i.1963, P.S. Nathan— 1 ♀, 1 ♂ ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), iv.1963, P.S. Nathan— 10 ♀♀, 10 ♂♂ ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), viii.1963, T.R.S. Nathan— 1 ♀, 3 ♂♂ ( MMUE) ; Coimbatore , 427 m (11°0'N, 76°57'E), ix.1964, T.R.S. Nathan— 4 ♀♀, 2 ♂♂ ( MMUE) ; Coimbatore , 427 m (11°0'N, 76°57'E), x–xi.1964, T.R.S. Nathan— 1 ♀ ( MMUE) ; Pudukkottai (10°23'N, 78°49'E), x.1984, T.R.S. Nathan— 1 ♀ ( CMNC) ; Thambikkottai (10°23'N, 79°27'E), vi.1962, P.S. Nathan— 1 ♀ ( CNC) ; Tranquebar [= Tharangambadi] (11°2'N, 79°5'E), [no date], [anonymous]— 3 ♀♀, 1 ♂ (lectotype, paralectotypes, alloreferent) ( ZMUC, ZMUK) ; Chennai District, Madras [= Chennai ] (13°5'2''N, 80°16'12''E), xi.1907, Tomlin— 1 ♀ ( BMNH) ; Madras Airport [= Chennai Airport ] (12°59'38''N, 80°9'28''E), x.1962, P.S. Nathan— 3 ♀♀, 2 ♂♂ ( CNC) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), iii.1962, P.S. Nathan— 3 ♀♀ ( CNC) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), vii.1939, [anonymous]— 1 ♂ ( FETC) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), [no date], P.S. Nathan— 1 ♀, 2 ♂♂ ( MNHN) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), xii.1971, T.R.S. Nathan— 1 ♂ ( RMNH) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), 15.ix.1947, P.S. Nathan— 2 ♂♂ ( CMNC) ; Marudamalai Hills (11°2'46''N, 76°51'7''E), xi.2004, Nathan— 1 ♂ ( PMOC) ; Tiruchirappalli District, Trichinopoly [= Tiruchirappalli ] (10°48'18''N, 78°41'8''E), iii.1907, [anonymous]— 1 ♀ ( BMNH) ; Tiruchirappalli District, Trichinopoly [= Tiruchirappalli ] (10°48'18''N, 78°41'8''E), x.1919, C. Leigh— 1 ♂ ( BMNH) ; Tiruchirappalli District, Trichinopoly [= Tiruchirappalli ] (10°48'18''N, 78°41'8''E), [no date], R.P. Castets— 1 ♀, 1 ♂ ( MNHN) ; UTTAR PRADESH, Kheri Reserved Forest (25°21'11''N, 79°17'9''E), ii.1916, H.G. Champion— 1 ♂ ( BMNH) ; UTTARAKHAND, Haldwani District, Kumaon [= Haldwani ] (29°13'N, 79°31'E), [no date], H.G. Champion— 2 ♀♀ ( BMNH) ; WEST BENGAL, Howrah District, Sarda (22°20'N, 88°8'E), [no date], F.W. Champion— 2 ♂♂ ( BMNH) ; Kalkata District, Calcutta [= Kolkata ] (22°34'N, 88°22'E), [no date], [anonymous]— 1 ♂ ( BMNH) ; Kalkata District, Calcutta [= Kolkata ] (22°34'N, 88°22'E), [no date], Waagen— 1 ♀ ( SMF); LAOS : CHAMPASAK, Ban Sepian (15°9'N, 106°15'E), 1–7.iv.2003, R. N.— 1 ♂ ( CEMT) ; PHONGSALY, Gnot-Ou District, Mount Phu Nam Khé (22°5'N, 102°27'E), 15–20.vi.2006, Local collectors— 12 ♀♀, 7 ♂♂ ( FGIC, OUMNH) ; VIENTIANE, Phou Khao Khouay National Bio-Biversity Conservation Area, 700– 800 m (18°20.369'N, 102°48.523'E), 25–30.v.2008 , A. Solodovnikov & J. Pedersen— 6 ♀♀, 7 ♂♂ ( FGIC, ZMUC); MALAYSIA : MALACCA, Malacca (2°12'N, 102°15'E), [no date], [anonymous]— 1 ♂ ( SMF) ; MYANMAR: KAYIN, Hpapun (18°3'30''N, 97°26'30''E), 1932, R.P. Loizeau— 1 ♂ ( MNHN) ; YANGON, Hlegu- Goygon (17°5'43''N, 96°13'16''E), iii.1997, M. Klicha— 2 ♂♂ ( OUMNH) ; Htauk Kyant (17°2'40''N, 96°8'12''E), ii.1997, M. Klicha— 1 ♀, 1 ♂ ( OUMNH) ; Rangoon [= Yangon] (16°48'N, 96°9'E), v.1934, F.J. Meggitt— 3 ♀♀, 2 ♂♂ ( BMNH) ; Rangoon [= Yangon] (16°48'N, 96°9'E), 1933–1934, F.J. Meggitt— 5 ♀♀, 4 ♂♂ ( BMNH); NEPAL : CENTRAL, Chitwan District, Sauraha, Chitwan National Park , 213 m (27°31'57''N, 84°30'20''E), 4.vi.1983, M.J.D. Brendell— 1 ♂ ( BMNH) ; Chitwan District, Sauraha, Chitwan National Park , 213 m (27°31'57''N, 84°30'20''E), 1–4.vi.1999, M. Pejcha— 1 ♀ ( FETC) ; Chitwan District, Sauraha, Chitwan National Park , 213 m (27°31'57''N, 84°30'20''E), 4–6.vi.1999, A. Kudrna— 1 ♂ ( FETC) ; PAKISTAN: ISLAMABAD CAPITAL TERRITORY, Islamabad (33°43'N, 73°4'E), 6.vi.1991, S. Prepsl— 1 ♀ (CMD); PENJAB, Changa Manga Forest , 70 km S Lahore (31°5'N, 74°0'E), 19–21.viii.1998, L. Cerný— 3 ♀♀ ( CVM) ; Kallar Kahar (32°47'32''N, 72°43'34''E), 26.vi.2008, A. Zubair— 1 ♂ ( PMOC) ; SRI LANKA: EASTERN, Pottuvil (6°52'N, 81°50'E), 1– 12.vii.1983, O. Mehl— 1 ♂ ( ZMUC) ; NORTH CENTRAL, Girithale (8°0'N, 80°56'E), 16.viii.1979, G. de Rougemont— 1 ♀ ( OUMNH) ; Polonnaruwa (7°56'N, 81°0'E), 24.vii.1982, P. Moretto— 2 ♂♂ ( PMOC) ; NORTHERN, Jaffa (9°20.5'N, 80°24.3'E), 1.iii.2010, Kuduvidanage et al.— 1 ♂ ( OUMNH) ; SOUTHERN, Hambantota District, Tissamaharama (6°17'N, 81°17'E), 1.viii.1982, P. Moretto— 2 ♀♀, 2 ♂♂ ( PMOC) ; THAILAND: BANGKOK , Bangkok (13°45'N, 100°30'E), [no date], S.S. Flower— 1 ♀ ( BMNH) ; CHIANG MAI, Chiang Dao (19°25'N, 98°52'E), 17–24.v.1991, V. Kubáň— 1 ♀, 1 ♂ ( CVM) ; Chiang Mai (18°48'N, 99°0'E), v.1986, N. Koyama— 2 ♂♂ ( CEMT) ; Chiang Mai (18°48'N, 99°0'E), 28.vi.1990, R. Minetti— 1 ♀ ( FETC) ; Chom Thong (18°25'N, 98°40'E), 24–27.iv.1991, J. Horák— 1 ♀, 1 ♂ ( BDGC) ; Doi Saket District , [unspecified locality] (18°52'N, 99°8'E), 1.vii.1959, B. Degerbol— 1 ♂ ( ZMUC) ; MAE HONG SON, Ban Huai Suea Thao (19°15'35''N, 97°54'15''E), 11–12.v.1992, J. Strnad— 1 ♀ ( CVM) ; Mae Hong Son (19°18'N, 97°58'E), 6–9.v.1991, J. Horák— 1 ♀, 1 ♂ ( BDGC) ; Pai (19°21'32''N, 98°26'12''E), 1–14.v.2001, J. Hromádka— 1 ♀, 3 ♂♂ ( OUMNH) ; NAKHON SAWAN , Nakhon Sawan (15°42'N, 100°7'E), 25.v.1980 , A. Chaminade— 1 ♀ ( PMOC) ; Nakhon Sawan (15°42'N, 100°7'E), 22.v.1982 , A. Chaminade— 1 ♂ ( PMOC); VIETNAM : DONG NAI, Long Binh R . V.N., Bien Hoa Province (10°56'N, 106°53'E), 12.v.1970, W.H. Tyson— 1 ♂ ( BDGC) ; HO CHI MINH , Saigon [= Ho Chi Minh City] (10°49'N, 106°37'E), x.1982, S. Pokorný— 2 ♀♀, 4 ♂♂ ( CVM) .

Natural history. A single specimen collected in arid scrub forest in Sri Lanka and 13 specimens collected near strongly disturbed primary forest in Laos. A few specimens were collected in cow dung, pitfall traps baited with human dung, and at light.

Remarks. The redescription is based on a male alloreferent (ZMUK) from the type locality and matching the female lectotype of S. bonasus .

Nomenclature and taxonomy. Digitonthophagus bonasus was the first species described included in the genus Digitonthophagus . The type series consist of three females. The larger and less abraded specimen has been chosen as lectotype (present designation). A lectotype is designated in order to choose a single specimen as name bearing type in the event that the other specimens of the syntype series could belong to a different cryptic species in this genus. This single specimen also shows better diagnostic characters as it is relatively undamaged and the original cephalic outline is clearly delimited. It is important to mention that these specimens together with the other Fabrician types were borrowed from the ZMUC but will eventually be deposited in the ZMUK collection.

Figures

FIGURES 7 – 10. Male dorsal habitus. 7. Digitonthophagus bonasus, alloreferent (Tamil Nadu State, India); 8. D. uks, holotype; 9. D. sahelicus, holotype; 10. D. catta, alloreferent (Tamil Nadu State, India).

FIGURES 23 – 30. 23, 25, 27, 29. Male head, dorsal view; 24, 26, 28, 30. Male head, frontal view. 23 – 24. Digitonthophagus bonasus, alloreferent (Tamil Nadu State, India); 25 – 26. D. uks, holotype; 27 – 28. D. sahelicus, holotype; 29 – 30. D. catta, alloreferent (Tamil Nadu State, India).

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 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.

FIGURES 135 – 146. Internal sac sclerites (FLP, A + SA). 135, 138, 141, 144. Dorsal view; 136, 139, 142, 145. Ventral view. 137, 140, 143, 146. Right lateral view. 135 – 137. Digitonthophagus bonasus, alloreferent (Tamil Nadu State, India); 138 – 140. D. uks, holotype; 141 – 143. D. sahelicus, holotype; 144 – 146. D. catta, alloreferent.

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.