Digitonthophagus catta (Fabricius, 1787)

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 : 24-28

Digitonthophagus catta (Fabricius, 1787)

( Figs. 10 , 29–30 , 58 , 74 , 93–94, 122, 144–146; Map 5)

Scarabaeus Catta Fabricius, 1787a: 12 (original description) Scarabaeus Catta — Olivier 1789: 125 (redescription)

Scarabaeus Catta — Gmelin 1790: 1534 (diagnosis)

Scarabaeus Catta — Fabricius 1792: 44 (diagnosis)

Scarabaeus Catta — Illiger 1800: 232 (monograph)

Copris Catta — Fabricius 1801: 35 (diagnosis)

Copris Catta — Illiger 1804: 149 (comment taxonomy)

Copris Catta — Schönherr 1806: 49 (mentioned as synonym) Onthophagus Catta — Dejean 1821: 53 (mentioned as synonym) Copris Catta — LePeletier & Audinet-Serville 1828: 354 (comment taxonomy) Copris Catta — Klug 1833: 163 (mentioned as synonym)

[ Onthophagus ] Catta — Dejean 1836: 157 (mentioned as synonym) Onthophagus Catta — Hope 1837: 32 (catalog)

Ontophagus Catta — Reiche 1840: 243 (comment taxonomy) [Ontophagus] Catta — Laporte 1840: 85 (mentioned as synonym) Onthophagus Catta — Sturm 1843: 107 (mentioned as synonym) Onthophagus catta — Motschulsky 1860: 154 (comment taxonomy) Onthophagus catta — Harold 1869: 1029 (mentioned as synonym) Onthophagus catta — Gerstaecker 1871: 50 (distribution)

Onthophagus catta — Gerstaecker 1873: 130 (monograph)

Onthophagus catta — Quedenfeldt 1884: 272 (distribution)

Onthophagus catta — Kolbe 1887: 246 (distribution)

Onthophagus catta —Kolbe 1897: 150 (mentioned as synonym) Onthophagus catta — Péringuey 1901: 181 (mentioned as synonym) Onthophagus catta — Gillet & Boucomont 1927: 138 (mentioned as synonym) Onthophagus catta — Arrow 1931: 230 (monograph)

Onthophagus catta — Boucomont 1934: 26 (mentioned as valid species) Onthophagus catta — Boucomont 1935: 284 (distribution)

Onthophagus catta — Paulian 1937: 12 (distribution)

Onthophagus catta — Paulian 1948: 151 (distribution)

Onthophagus catta — Janssens 1953: 1549 (distribution)

Onthophagus catta — Frey 1958: 73 (mentioned as synonym) [ Scarabaeus ] catta — Balthasar 1963: 365 (mentioned as synonym) Onthophagus catta — Ferreira 1967: 1169 (mentioned as synonym) Onthophagus catta — Ferreira 1968: 593 (mentioned as synonym)

Onthophagus catta — Ferreira 1972: 780 (mentioned as synonym)

Onthophagus catta — Paulian 1986: 107 (catalog)

Onthophagus gazella — Veenakumari & Veeresh 1996: 252 (natural history) Onthophagus catta — Gaikwad & Bhawane 2015: 1538 (natural history) Digitonthophagus catta — Génier & Davis 2017: 497 (comment taxonomy, distribution)

Diagnosis. Species distributed from Pakistan eastward; left lateral apophysis of FLP sclerite moderately developed ( Fig. 144 ).

Description. Alloreferent ♂ ( Fig. 10 ): Measurements. Length 11.5 mm, width 6.5 mm. Head ( Figs. 29–30 ). 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 moderately long, arcuate in frontal view, parallel sided on basal four-fifths; distinctly tapering externally before apex and narrower on apical sixth; 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. 58 ). Surface with granulate punctures restricted to anterior half medially, with smaller, irregularlyscattered, simple punctures on posterior half of disc; punctures lacking 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 slightly concave, surface of anterior angles concave, anterior angles unmodified; anterior half of lateral edge arcuate in dorsal and slightly sinuous anteriorly in lateral view; posterior angles simply arcuate in dorsal view. Anterior hypomeral ridge broadly arcuate, anterior hypomeral depression surface slightly darker in color medially. Elytra ( Fig. 10 ). Interval 2 with few fine granules on apical declivity, interval 4 lacking granules from base to apex. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus ( Fig. 122 ). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites ( Figs. 144–146 ). Axial sclerite moderately sclerotized, shorter than subaxial sclerite, almost straight. Subaxial sclerite, larger basally and gradually tapering, extending straight and interrupted at apex of frontolateral peripheral sclerite apical portion, with villi on apical sixth. Frontolateral peripheral sclerite basoventral apophysis moderately developed, angular in lateral view; with a single acute medioventral carinae; right lateral fold produced into a uneven bifurcate process, lateral edge with few indentations; left lateral apophysis present, moderately developed; left lateral lobe absent; subapicodorsal lobe well sclerotized, reaching anterior edge, partly fused to apical lobe, with villi similar to apical lobe, right edge round before apex; apical lobe round apically, longitudinally folded and partly fused to subapicodorsal lobe, apical villi regular in shape; subapicoventral lobe atrophied, oriented ventrally.

Variation. Measurements (107 ♂♂, 106 ♀♀). Length: male 7.0–13.0 mm (10.6 ± 1.0 mm), female 7.5–12.5 mm (10.2 ± 1.0 mm). Female specimen. Cephalic outline in dorsal view similar to Fig. 74 ; 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 approximately parallel in dorsal view (Fig. 93), anterolateral surface obliquely concave, anterosuperior edge slightly arcuate in dorsal view, lateral portion of anterosuperior edge slightly upturned (Fig. 94). Protibia short, with external teeth more robust.

Primary type data. Lectotype ♀ (ZMUK) present designation: [WORLD / SCARAB. / DATABASE / WSD00032221] barcode label; [LECTOTYPE ♀ / Scarabaeus / catta / Fabricius, 1787 / des. F. Génier, 2016] red card.

Material examined (188 ♂♂, 199 ♀♀, 855 sex no recorded), distribution (Map 5): AFGHANISTAN: KHYBER PAKHTUNKHWA, Hazara, Abbottabad (39°10'N, 73°13'E), vi.2002, [anonymous] —2 ♀♀, 2 ♂♂ (CEMT); BANGLADESH: RANGPUR, Dinajpur (25°45'N, 88°43'E), iv.1970, Barbe— 1 ♂ ( CMNC) ; INDIA: Coromandel Coast, [no date], Zschuck— 4 ♀♀ (lectotype, 3 paralectotypes) ( ZMUC) ; ANDAMAN AND NICOBAR ISLANDS, [unspecified locality] (12°30'N, 92°45'E), [no date], Roepstorff— 1 ♂ ( ZMUC) ; GOA, Calangute (15°32'30''N, 73°45'45''E), 15.xii.1976, [anonymous]— 4 ♀♀, 3 ♂♂ ( OMOC) ; [unspecified locality] (15°22'N, 74°4'E), 22–23.x.1986, J.L. & A. Nicolas— 1 ♀, 1 ♂ (CMD) ; JHARKHAND, Ranchi (23°20'N, 85°19'E), 1970, Dr. Soběslavský— 1 ♀ ( CVM) ; Ranchi (23°20'N, 85°19'E), [no date], R.P. Bretaudeau— 1 ♀, 1 ♂ ( MNHN) ; KARNATAKA, 15 km W Gadag , 670 m (15°25'48''N, 75°28'39''E), 13.viii.1984, B. Gill— 1 ♀ ( BDGC) ; 20 km SW Hubli , 650 m (15°12'3''N, 74°59'34''E), 11.vii.1984, B. Gill— 3 ♀♀, 1 ♂ ( BDGC) ; Bangalore , 970 m (12°58'N, 77°34'E), 17–19.vii.1984, B. Gill— 7 ♀♀, 2 ♂♂ ( BDGC) ; Chikkangalur , près de Bangalore [=Chickmagaluru] (13°18'53''N, 74°46'29''E), vii–ix.1899, R.P. Tabourel— 1 ♀ ( MNHN) ; Hampi (15°20'6''N, 76°27'36''E), 18.x.1986, J.L. Nicolas— 2 ♀♀, 2 ♂♂ (CMD); Harihar , 750 m (14°30'30''N, 75°48'30''E), 15.vii.1984, B. Gill— 2 ♀♀, 5 ♂♂ ( BDGC) ; Hubli , 670 m (15°21'42''N, 75°5'6''E), 7–14.vii.1984, B. Gill— 5 ♀♀, 8 ♂♂ ( BDGC) ; Sindhanur (15°46'N, 76°45'E), 21.xii.1989, C. Mahalingappa— 1 ♀, 2 ♂♂ ( OUMNH) ; Mysore District, Mysore (12°18'N, 76°39'E), [no date], [anonymous]— 1 ♀, 1 ♂ ( MNHN) ; Shimoga District, Bhadravati (13°50'24''N, 75°42'7.2''E), 13.v.1935, [anonymous]— 1 ♀, 1 ♂ ( CMNC) ; Shimoga (13°55'N, 75°34'E), v.1897, [anonymous]— 1 ♀, 1 ♂ ( MNHN) ; Shimoga (13°55'N, 75°34'E), vi.1897, [anonymous]— 1 ♀, 1 ♂ ( MNHN) ; Shimoga (13°55'N, 75°34'E), iv.1936, [anonymous]— 1 ♂ ( PMOC) ; KERALA, Chembra Peak , 1067 m (11°33'12''N, 76°5'14''E), iv.1970, T.R.S. Nathan— 1 ♂ ( CMNC) ; Thenmala (8°58'N, 77°4'E), v.1988, T.R.S. Nathan— 1 ♀, 1 ♂ ( CMNC) ; Thiruvananthapuram District, Poonmudi Range , 914 m (8°45'27''N, 77°6'54''E), v.1989, T.R.S. Nathan— 1 ♀, 1 ♂ ( CMNC) ; KERALA / TAMIL NADU, Anaimalai Hills , 1067 m (10°22'N, 77°7.5'E), v.1963, P.S. Nathan— 1 ♀ ( CNC) ; Anaimalai Hills , 1067 m (10°22'N, 77°7.5'E), v.1977, [anonymous]— 4 ♀♀, 4 ♂♂ ( CNC) ; Anaimalai Hills , 1067 m (10°22'N, 77°7.5'E), iv.1963, P.S. Nathan— 2 ♂♂ ( CNC) ; Anaimalai Hills , 1067 m (10°22'N, 77°7.5'E), x.1977, T.R.S. Nathan— 1 ♂ ( RMNH) ; Cinchona, Anaimalai Hills , 1067 m, iv.1964, P.S. Nathan— 1 ♂ ( CNC) ; MADHYA PRADESH, Jabalpur (23°10'N, 79°56'E), x.1957, Nathan— 1 ♀, 1 ♂ ( CMNC) ; Maihar (24°16'10''N, 80°45'24''E), x.1972, [anonymous]— 1 ♀ ( ZMUC) ; Pachmarhi, Satputa Hills , 1067 m (22°28'.12''N, 78°25'59.88''E), x.1970, T. Nathan— 1 ♀, 1 ♂ ( CNC) ; Ratlam (23°20'N, 75°2'E), [no date], [anonymous]— 1 ♂ ( MNHN) ; Satna (24°36'N, 80°50'E), [no date], Stloukal— 1 ♂ ( CMNC) ; MAHARASHTRA, Pune (18°31'N, 73°51'E), viii.1984, S. Pokorný— 2 ♀♀, 2 ♂♂ ( CVM) ; ODISHA, Brahmapur (19°19'N, 84°47'E), xi.1972, [anonymous]— 1 ♂ ( ZMUC) ; ORISSA, Berhampur (19°18'30''N, 84°47'26''E), [no date], Atkinson— 1 ♂ ( OUMNH) ; Balasore District, Balasore (21°30'N, 86°56'E), [no date], R.P. Gengler— 1 ♀, 1 ♂ ( MNHN); PONDICHERRY , Karaikal District, Karaikal (10°56'N, 79°50'E), viii.1991, T.R.S. Nathan— 1 ♀ ( CMNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), x.1991, T.R.S. Nathan— 1 ♂ ( CMNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), iv.1962, P.S. Nathan— 1 ♀, 1 ♂ ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), x.1963, P.S. Nathan— 1 ♀, 1 ♂ ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), ix.1952, P.S. Nathan— 2 ♀♀ ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), vi.1962, P.S. Nathan— 1 ♂ ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), vii.1963, P.S. Nathan— 1 ♂ ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), i.1962, P.S. Nathan— 11 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), ii.1962, P.S. Nathan— 36 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), iii.1962, P.S. Nathan— 34 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), vii.1962, P.S. Nathan— 40 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), viii.1962, P.S. Nathan— 74 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), ix.1962, P.S. Nathan— 146 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), x.1962, P.S. Nathan— 83 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), xi.1962, P.S. Nathan— 39 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), xii.1962, P.S. Nathan— 9 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), i.1963, P.S. Nathan— 16 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), ii.1963, P.S. Nathan— 12 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), iii.1963, P.S. Nathan— 34 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), iv.1963, P.S. Nathan— 12 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), vi.1963, P.S. Nathan— 4 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), x.1966, P.S. Nathan— 13 specimens ( CNC) ; Karaikal District, Karaikal (10°56'N, 79°50'E), viii.1963, T.R.S. Nathan— 2 ♀♀, 2 ♂♂ ( MMUE) ; Karaikal District, Karaikal (10°56'N, 79°50'E), i–ii.1964, T.R.S. Nathan— 4 ♀♀, 4 ♂♂ ( MMUE) ; Karaikal District, Karaikal (10°56'N, 79°50'E), vii.1970, P. Susai Nathan— 1 ♂ ( RMNH) ; Nedungadu (10°58'N, 79°46'E), [no date], [anonymous]— 1 ♂ ( NMPC) ; RAJASTHAN, Barmer (25°45'N, 71°23'E), ix.1973, T.R.S. Nathan— 1 ♀ ( RMNH) ; Kota , 250 m (25°10'28''N, 75°50'14''E), x.1981, T.R.S. Nathan— 2 ♀♀, 2 ♂♂ ( OUMNH) ; Thar desert around Jaisalmer (26°55'N, 70°56'E), 2.viii.2007, I. Nádai— 4 ♀♀, 7 ♂♂ ( PMOC) ; TAMIL NADU, 20 km SE Sagar , 600 m (14°6.37'N, 75°8.93'E), 12.v.2005, M. Halada— 1 ♀ ( OUMNH) ; 20 km SW Karamadai , 450 m (11°12.6'N, 76°44.9'E), 27.iv.2005, M. Halada— 1 ♂ ( OUMNH) ; Cap Comorin [=Kanyakumari] (8°5'N, 77°32'E), 1888, R.P. Castets— 16 specimens ( MNHN) ; Coimbatore , 427 m (11°0'N, 76°57'E), xi.1969, P.S. Nathan— 1 ♀, 3 ♂♂ ( CMNC, CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), iv.1959, P.S. Nathan— 1 ♀ ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), iv.1962, P.S. Nathan— 25 specimens ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), v.1962, P.S. Nathan— 19 specimens ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), vii.1962, P.S. Nathan— 18 specimens ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), ix.1962, P.S. Nathan— 14 specimens ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), x.1962, P.S. Nathan— 3 specimens ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), xi.1962, P.S.

Nathan—20 specimens (CNC); Coimbatore , 427 m (11°0'N, 76°57'E), xii.1962, P.S. Nathan— 5 specimens ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), i.1963, P.S. Nathan— 6 specimens ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), iv.1963, P.S. Nathan— 50 specimens ( CNC) ; Coimbatore , 427 m (11°0'N, 76°57'E), xi.1966, P.S. Nathan— 20 specimens, 2 ♂♂ ( CNC, PMOC) ; Coimbatore , 427 m (11°0'N, 76°57'E), viii.1963, T.R.S. Nathan— 6 ♀♀, 6 ♂♂ ( MMUE) ; Coimbatore , 427 m (11°0'N, 76°57'E), ix.1964, T.R.S. Nathan— 1 ♀, 5 ♂♂ ( MMUE) ; Coimbatore , 427 m (11°0'N, 76°57'E), x–xi.1964, T.R.S. Nathan— 4 ♂♂ ( MMUE) ; Coimbatore , 427 m (11°0'N, 76°57'E), xi.1967, P.S. Nathan— 1 ♀ ( PMOC) ; Coimbatore , 427 m (11°0'N, 76°57'E), ix.1972, T.R.S. Nathan— 2 ♀♀, 1 ♂ ( PMOC, RMNH) ; Coimbatore , 427 m (11°0'N, 76°57'E), x.1972, T.R.S. Nathan— 2 ♀♀, 2 ♂♂ ( PMOC) ; Coimbatore , 427 m (11°0'N, 76°57'E), xii.1976, T.R.S. Nathan— 1 ♂ ( RMNH) ; Gingee (12°15'N, 79°25'E), [no date], M. André— 1 ♂ ( PMOC) ; Madurai (9°55'N, 78°7'E), x.1975, M. Coe— 1 ♀, 4 ♂♂ ( OUMNH) ; Maravakkadu (10°38'N, 79°27'E), v.1962, T.R.S. Nathan— 2 ♀♀ ( CNC) ; NE of Madurai , 200 m (10°8.7'N, 77°46.5'E), 6.v.2005, M. Halada— 1 ♀ ( OUMNH) ; Palni Hills (10°12'N, 77°30'E), 1898, R.P. Castets— 1 ♀, 1 ♂ ( MNHN) ; Point Calimere (10°17'35''N, 79°52'6''E), 1972, T.R.S. Nathan— 1 ♀ ( RMNH) ; Pudukkottai (10°23'N, 78°49'E), x.1984, T.R.S. Nathan— 1 ♀, 1 ♂ ( CMNC) ; Singara, Nilgiri Hills (11°22'N, 76°46'E), v.1963, T.R.S. Nathan— 2 ♀♀, 2 ♂♂ ( MMUE) ; Thambikkottai (10°23'N, 79°27'E), vi.1962, P.S. Nathan— 3 ♀♀ ( CNC) ; Tirunelveli (8°44'N, 77°42'E), x.1986, [anonymous]— 1 ♀, 1 ♂ ( OMOC) ; Tranquebar [= Tharangambadi] (11°2'N, 79°5'E), [no date], [anonymous]— 1 ♂ ( ZMUC) ; Vellore (12°55'N, 79°8'E), [no date], Löventhal— 2 ♂♂ ( ZMUC) ; Chennai District, Madras Airport [= Chennai Airport ] (12°59'38''N, 80°9'28''E), v.1962, P.S. Nathan— 64 ♀♀, 42 ♂♂ ( CNC) ; Chennai District, Madras Airport [= Chennai Airport ] (12°59'38''N, 80°9'28''E), viii.1962, P.S. Nathan— 19 specimens ( CNC) ; Chennai District, Madras Airport [= Chennai Airport ] (12°59'38''N, 80°9'28''E), ix.1962, P.S. Nathan— 9 specimens ( CNC) ; Chennai District, Madras Airport [= Chennai Airport ] (12°59'38''N, 80°9'28''E), x.1962, P.S. Nathan— 14 specimens ( CNC) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), x.1958, P.S. Nathan— 1 ♀ ( CNC) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), vi.1962, P.S. Nathan— 1 ♀ ( CNC) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), x.1976, [anonymous]— 1 ♀, 1 ♂ ( CNC) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), xi.1976, [anonymous]— 1 ♂ ( CNC) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), iv.1976, [anonymous]— 1 ♂ ( CNC) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), xi.1957, P.S. Nathan— 3 specimens ( CNC) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), xi.1962, P.S. Nathan— 12 specimens ( CNC) ; Coimbatore District, Coimbatore (11°0'N, 76°57'E), xi.1965, P.S. Nathan— 10 specimens ( CNC) ; Marudamalai Hills (11°2'46''N, 76°51'7''E), xi.1969, P.S. Nathan— 29 specimens, 1 ♀, 1 ♂ ( CMNC, CNC) ; Marudamalai Hills (11°2'46''N, 76°51'7''E), x.1969, P.S. Nathan— 2 ♀♀ ( CNC) ; Kancheepuram District, Manapakkam (13°0'N, 80°10'E), 20.ix.2002, S. Saluk— 1 ♂ ( PMOC) ; UTTAR PRADESH, Babina , 300 m (25°15'N, 78°28'E), viii.1984, T.R.S. Nathan— 1 ♀, 2 ♂♂ ( CMNC) ; Babina , 300 m (25°15'N, 78°28'E), viii.1987, T.R.S. Nathan— 3 ♀♀, 3 ♂♂ ( CMNC, FETC) ; UTTARAKHAND, Rishikesh (30°5'N, 78°16'E), 7.x.1981, F. Tagliaferri— 1 ♀ ( FETC) ; WEST BENGAL, Burasanti, Singur (22°49'N, 88°13'E), 14.vi.1971, Oppenheimer— 1 ♂ ( CMNC) ; Burasanti, Singur (22°49'N, 88°13'E), 27.vi.1971, Oppenheimer— 2 ♀♀ ( CMNC) ; Kalkata District, Calcutta [= Kolkata ] (22°34'N, 88°22'E), [no date], [anonymous]— 1 ♀ ( NMPC) ; NEPAL: CENTRAL, Adhabar, near Simra , 183 m (27°13'49''N, 84°59'14''E), 23.viii.1967, Canadian Nepal Expedition— 1 ♂ ( CNC) ; Adhabar, near Simra , 183 m (27°13'49''N, 84°59'14''E), 25.viii.1967, Canadian Nepal Expedition— 1 ♀ ( CNC) ; Adhabar, near Simra , 183 m (27°13'49''N, 84°59'14''E), 27.viii.1967, Canadian Nepal Expedition— 1 ♀ ( CNC) ; Adhabar, near Simra , 183 m (27°13'49''N, 84°59'14''E), 28.viii.1967, Canadian Nepal Expedition— 1 ♂ ( CNC) ; Chitwan District, Sauraha, Chitwan National Park , 300 m (27°34'29''N, 84°29'37''E), 1–4.vi.1999, M. Pejcha— 1 ♀ ( FETC) ; EASTERN, Dharan (26°49'N, 87°16'E), 4.iv.1959, K. Becker Larsen— 1 ♀ ( ZMUC) ; Dharan (26°49'N, 87°16'E), 8.vii.1959, K. Becker Larsen— 1 ♀ ( ZMUC) ; PAKISTAN: Thar , 18.ix.2007, Zubair & Atique— 2 ♂♂ ( BDGC) ; KHYBER PAKHTUNKHWA, Changla Gali, Murree, Islamabad area , 1981 m (33°59'42''N, 73°23'2''E), 5.vii.2006, Z. Ahmed— 1 ♀ ( BDGC) ; PENJAB, Chakwal (32°33'N, 72°30'36''E), 19.v.1990, Naeem— 1 ♀ ( CNC) ; Kallar Kahar (32°47'32''N, 72°43'34''E), 26.vi.2008, A. Zubair— 2 ♂♂ ( PMOC) ; Murree, Islamabad area , 1981 m (33°54'27''N, 73°23'29''E), vii.2006, Z. Ahmed— 2 ♀♀, 2 ♂♂ ( BDGC) ; SINDH, Karachi (24°57'N, 67°5'E), viii– ix, Maindron— 1 ♀, 1 ♂ ( MNHN) ; Karachi (24°57'N, 67°5'E), 26.i.2007, Hareem & Atige— 2 ♀♀, 2 ♂♂ ( CEMT) ; SRI LANKA: Wilpattu National Park (8°25'N, 80°1'E), 18.vii.1979, G. de Rougemont— 1 ♀ ( OUMNH) ; EASTERN, Pottuvil (6°52'N, 81°50'E), 1–12.vii.1983, O. Mehl— 1 ♀ ( ZMUC) ; NORTH CENTRAL, Anuradhapura (8°19'N, 80°24'E), 9.vii.1977, O. Mehl— 1 ♀ ( ZMUC) ; Girithale (8°0'N, 80°56'E), 16.viii.1979, G. de Rougemont— 1 ♀ ( OUMNH) ; NORTH WESTERN, Kalpitiya (8°14'N, 79°45.8'E), 12.xi.2009, Kuduvidanage et al.— 2 ♂♂ ( OUMNH) ; SOUTHERN, Galle (6°2'N, 80°13'E), 30.vii.1982, P. Moretto— 2 ♀♀, 2 ♂♂ ( PMOC) ; Hambantota District, Hambantota (6°7'2''N, 81°5'9''E), 10.xii.1979, V. Mahler— 1 ♀ ( ZMUC) ; Tissamaharama (6°17'N, 81°17'E), 12.xii.1979, [anonymous]— 1 ♂ ( ZMUC) ; Tissamaharama (6°17'N, 81°17'E), 1.viii.1982, P. Moretto— 2 ♀♀, 1 ♂ ( PMOC) ; W of Tissamaharama (6°16'N, 81°14'E), 20.ii.1986, R.W. Lekkerkerk— 1 ♂ ( RMNH) ; WESTERN, Negombo (7°13'N, 79°50'E), 17–20.vi.1985, O. Mehl— 1 ♀ ( ZMUC) ; Negombo (7°13'N, 79°50'E), 21.viii.1986, C. Lestrade— 1 ♀ ( PMOC) ; Negombo (7°13'N, 79°50'E), 23.viii.1986, C. Lestrade— 2 ♂♂ ( PMOC) ; Wadduwa (6°40'N, 79°56'E), [no date], v. de Poll— 1 ♂ ( PMOC) .

MAP 5. Distribution of Digitonthophagus catta .

Natural history. Grassy plot in West Bengal (India) and arid scrub forest in Sri Lanka. A few specimens collected in buffalo, cow, goat, northern plains gray langur ( Semnopithecus entellus ), and human dung. The nesting behavior and some aspects of larval natural history are described by Veenakumari & Veeresh (1996) and Gaikwad & Bhawane (2015).

Nomenclature and taxonomy. Scarabaeus catta Fabricius was described in the first volume of Mantissa Insectorum. This work, which contained the Coleoptera , was published in June 1787 ( Evenhuis 1997; Bousquet 2016). There is a misspelling of the type locality (Ceromandel) in the original description, but there is no doubt that it pertains to Coromandel Coast, which historically was covering the coastal area of the current states of Tamil Nadu and Andhra Pradesh. These specimens are most likely from the area of Tranquebar, a former Danish colony from 1620–1845. However, there is no biographical information on Zschuck the stated collector of the specimens (see Zimsen 1964). The four specimens forming the type series are unlabeled females from Kiel collection. Based on the pins used, these four specimens are unquestionably from Fabricius time. The three different types of pins used also suggest that these specimens are from more than a single collecting event. In order to stabilize nomenclature, we designate here the clean and unabraded specimen as the 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. The lectotype has a steel pin similar to those used to pin the syntypes of D. bonasus , which were collected from the same area in India. This female specimen, with the distinctly concave ventral surface of the median pronotal tubercle combined with the distinctly arcuate anterior hypomeral carina, matches the taxon present in India. One of the female syntypes is pinned with a Crataegus thorn and appears to have the same origin as the four male specimens from Kiel collection which have been prepared in the same way. This pinning method suggests that this fourth female specimen was from the same collecting event as the four males S. gazella from the Kiel collection. All of these specimens do also lack the typical small green label used to identify Fabrician types. This gives support to the hypothesis that Fabricius considered females as S. catta and males as S. gazella .

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 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 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.