taxonID	type	description	language	source
7B3FB814D321473F93BFFE10FDA5FCC7.taxon	type_taxon	Type species. Onthophagus mopsus (Fabricius, 1792), by original designation.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D321473F93BFFE10FDA5FCC7.taxon	diagnosis	Diagnosis. Onthophagus species of small or medium size (length 3.0 to 10.0 mm), colour blackish brown, sometimes with green or blue metallic lustre, elytra sometimes with dark red basal and apical spots, in one case elytra yellow ochre with symmetrical v-shaped black spots. Antennae yellow, more or less dark. Setation pale-yellow. Head short, clypeus broadly round, barely or not at all emarginate anteriorly and not sinuate at sides; antenna with 9 antennomeres; clypeofrontal carina distinct, sometimes barely discernible, bent backward, in major males usually halfway between the base of the horn and the anterior clypeal margin, in one case nearly adjacent to the base of the horn; major males with the occipital carina extended in a long horn arising sometimes between eyes or slightly forward, sometimes in front of them, flat and more or less broad at base, directed backward, following the curvature of pronotum, often extending past the middle of the body; occipital carina reduced to a short tubercle in minor males or to a more or less straight carina, placed between eyes, or nearly so, in females (Fig. 1 a); clypeal surface with transversely rugose, setigerous punctures, frontal surface sparsely punctate, punctures of two sizes. Pronotum convex, declivous anteriorly, major males with a more or less appreciable anteromedian groove, with or without a tubercle on each side; minor males and females with an anteromedian transverse gibbosity, sometimes hardly appreciable, sometimes divided in two lobes; both sexes with a posterolateral area, more or less wide and depressed, dull with stronger microreticulation, on either side near pronotal posterior angles (Fig. 2), sometimes anteriorly with a tuft of few longer setae; anterior angles distinctly produced, sides not sinuate behind them; dorsal surface setigerously punctate, punctures more or less slightly impressed, usually subregular in distribution, bearing pale-yellow setae. Elytral striae usually shiny, distinctly impressed, with punctures slightly larger than strial width and barely crenulating interstrial sides; interstriae flat to convex, all rather regularly densely granulate, granules with a more or less short thin setae. Pygidium basally ridged, with rather regularly distributed, setigerous punctures; setae pale-yellow, thin, usually longer that those of elytra. Apical margin of male protibiae either oblique or perpendicular to the inner protibial margin, in the latter case with a tuft of short pale-yellow bristles, always with a small denticle strongly curved downward at the inner angle of protibial apex; metatibiae distinctly widened apically. Male genital armature. Parameres short, apices usually bent ventrally and diverging apically, without basolateral plate denticle, round at apex; endophallus with the presence of accessory endophallites, one (in O. ensifer), usually two evident raspulae and a U-shaped lamella copulatrix, with arms more or less slender, sometimes with spinelike processes or apically bifurcate. Epipharynx (Fig. 1 b). Rounded laterally, front edge not sinuate; acropariae long, rather densely arranged; epitorma very slender, almost indistinguishable, with some bristles along its length, the distal ones longer and stouter; corypha without spinules; chaetopariae stout; tomae rounded at the end.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D321473F93BFFE10FDA5FCC7.taxon	distribution	Distribution. East Palaearctic, east Afrotropical and west Oriental ecozone (Schoolmeesters 2023). Historical review. The first author that somehow grouped most of the species treated in this paper was Boucomont (1914). He drawn up a key to some Onthophagus species with the following characteristics: “ tête ♂ avec une ited by G. Fiumi. corne plus ou moins longue, non lamiforme, thorax ♂ sans saillies ni tubercules latéraux ou discaux, tout au plus avec deux petites dents au sommet; intervalles des élytres à ponctuation râpeuse; insectes concolores sans taches accusées ” [head ♂ with a more or less long, not lamiform horn, pronotum without projections or lateral tubercles, at most with two small teeth anteriorly; interstriae with raspish punctation; insects unicoloured, without definite spots]. Five species were placed in this “ group ”: Onthophagus spinifex (Fabricius, 1781), O. ensifer Boucomont, 1914, O. turbatus Walker, 1858, O. gracilicornis (Germar, 1813) [now deemed as junior synonym of O. mopsus, see below] and O. hastifer van Lansberge, 1885. Arrow (1931) proposed a “ mopsus group ” (Group 25) for O. mopsus, O. turbatus, O. hastifer, O. ensifer and for O. nitidulus Klug, 1845, together with other seventeen species. It’s worth noting that Arrow (1931) placed the male of Onthophagus spinifex in the Group 13 (marginalis group), whereas the female was placed in the Group 6 (rudis group). Biswas and Chatterjee (1985), probably referring to Arrow (1931), mentioned a “ mopsus group ” without specifying any morphological characteristic. Again, also Kabakov and Napolov (1999) placed O. hastifer in a vaguely defined “ mopsus group ”. Later on, Kabakov (2006) described the subgenus Indonthophagus for all the species quoted by Boucomont (1914) and adding Onthophagus nitidulus. The subgenus was then accepted and used by Ziani and Bezděk (2016), Král and Batelka (2017), Montreuil (2017), Gupta et al. (2018), Ziani et al. (2019), Ghosh et al. (2020), Kharel et al. (2020), Gupta et al. (2022) and Schoolmeesters (2023), even though none of them dealt with Indonthophagus more in depth. Phonetic remarks. As explained some years ago (Ziani 2020 a), Indonthophagus, having Onthophagus as suffix, is a proparoxytone word, that is, it has a heavy stress on the third-to-last syllable. It has to be pronounced Indontòfagus.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D321473F93BFFE10FDA5FCC7.taxon	biology_ecology	Biology. All the species cited in this paper should be strictly coprophagous. To confirm that, Mittal (1986) reported O. mopsus and O. spinifex attracted by human dung and Mittal (1999) by cattle and horse dung. Thomas et al. (2006) recorded O. ensifer in a trap baited with Indian elephant dung. Again, O. ensifer with O. turbatus have been recorded in gaur dung from Southern Western Ghats (India) by Thomas et al. (2007). According to Asha et al. (2022), this last two species (O. ensifer and O. turbatus) are specialists of non-ruminant dung, such as elephant and wild boar. And always regarding O. ensifer and O. turbatus, Sathiandran et al. (2021) specified that, in southern India, the two species attend dung of deer, elephant, gaur and especially boar, whereas only O. turbatus has been collected in macaque dung. For what I know, other eating preferences have never been reported. Taiwanese specimens of O. hastifer were collected in buffalo dung deposited in open pastures (D. Král, personal communication). In Western Ghats (India), O. turbatus seems to be abundant both in open and in shaded areas, with both day and night activity, whereas O. spinifex is a nocturnal species present in open areas (Asha et al. 2021). Three species at least (O. turbatus, O. mopsus and O. spinifex) present positive phototaxis, and were collected at light in Pakistan (unpublished data).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D323473193BFFC42FB06FB4F.taxon	description	(Fig. 3)	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D323473193BFFC42FB06FB4F.taxon	materials_examined	Type locality. “ Erythrée ” [Eritrea]. Type material. Holotype, a minor ♂, fixed by monotypy, examined by photos, in d’Orbigny collection (MNHN). Diagnostic features. Length 4.0 to 6.5 mm. Colour dark brown, moderately shiny, with distinct isodiametric microreticulation, elytra sometimes with dark red irregular humeral spots, clypeal sides dark red. Antennae yellow. Dorsal pubescence pale-yellow. Head short, clearly wider than long, with clypeus broadly round, only very feebly sinuate anteriorly, sides not sinuate; clypeofrontal carina distinct and bent backward, slightly closer to the base of the horn than to the anterior margin of clypeus; occipital carina extended in a long horn arising between the eyes, where it is broad, broader than clypeofrontal carina, and flat, directed backward, tapering gradually to the extremity and following the curvature of pronotum, with moderately long yellow setae mainly at sides, in major males, reduced to a narrow straight lamina in minor males (holotype); clypeal surface with very transversely rugose, setigerous punctures, frontal surface doubly sparsely punctate. Pronotum convex, declivous anteriorly, with a distinct anteromedian depressed area in major males (in minor male (holotype) without depression) and a small hardly distinguishable posterolateral dull area on either side near pronotal posterior angles; anterior angles distinctly produced, sides not sinuate behind them in dorsal view; dorsal surface setigerously punctate, punctures sub-regular in distribution, separated by 1 - 2 diameters on disc, smaller and more spaced on the anteromedian furrow; each puncture bears a small granule at its anterior margin, and a long pale-yellow seta, slightly longer that those on clypeus. Elytral striae shiny, distinctly impressed, with punctures slightly larger than strial width and barely crenulating interstrial sides; interstriae flat to barely convex, all rather regularly granulate; granules slightly smaller than strial punctures; posterior margin of each granule with a small, indistinct, setigerous puncture; setae pale-yellow, thin, as long as pronotal ones. Pygidium with rather regularly distributed, setigerous punctures; setae yellow, long, thin. Males with protibial spur clearly bent downward apically, and with a small denticle strongly curved downward on the inner angle of protibial apex. Female not examined. According to d’Orbigny (1913), females have the occipital carina extended in a backward bent lamina, placed between the eyes, in the middle of them, and a pronotal anteromedian transverse gibbosity, sometimes very slightly divided in two lobes. Male genital armature. Parameres short, apices bent ventrally, obviously diverging apically, without basolateral plate denticle, round at apex (Fig. 3 b – c); endophallus with the presence of accessory endophallites, two evident raspulae and a bracket-shaped lamella copulatrix, very thin (Fig. 3 d).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D323473193BFFC42FB06FB4F.taxon	distribution	Distribution. Eritrea (d’Orbigny 1902). Sudan, Ethiopia, Kenya (d’Orbigny 1913). Ivory Coast (Balthasar 1969, doubtful record). Yemen (Ziani and Keith 2023).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D323473193BFFC42FB06FB4F.taxon	materials_examined	Material examined. Eritrea: “ Erythrée ” [Eritrea], examined by photos (holotype ♂ of O. aeneopiceus, MNHN). Yemen: Ibb gov., W of Jabal Hadish, 30. iii. 1976, R. Naviaux leg. 1 ♂ (DKCC); Ibb gov., 5 km SW Najd al Jumai - Wadi Maytam 1600 m, 13 ° 52 ’ 28 ″ N 44 ° 17 ’ 45 ” E, 26 - 27. x. 2005, D. Král leg. 1 ♂ (NMPC). Historical review. Despite O. aeneopiceus being described more than a century ago, its nomenclatorial and systematic history is relatively poor. According to d’Orbigny (1902), the holotype, a male, the sole available specimen for the description, was collected in Eritrea. Eleven years later, d’Orbigny (1913), evidently after examining further material, specified the presence of a long curved backward cephalic horn in major males, described the female and added other localities from three more countries. Balthasar (1963), whilst specifying that O. aeneopiceus was an Afrotropical species, hypothesized its presence also in the Palaearctic Africa. Balthasar (1969) reported the species for Ivory Coast, but this record, completely off the known distribution area of the species, was rightly regarded dubiously by Schoolmeesters (2023). Lastly, Ziani and Keith (2023) published the first record for the Palaearctic ecozone (sensu Löbl and Löbl 2016) after two males collected in Yemen.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D323473193BFFC42FB06FB4F.taxon	discussion	Remarks. Ziani and Keith (2023), although preferring not to insert O. aeneopiceus in any known subgenus of Onthophagus, pointed out the similarity between the lamella copulatrix and the overall structures inside the endophallus of O. aeneopiceus with those of O. mopsus. It is noteworthy that O. aeneopiceus is, so far, the sole species belonging to the group in which the male protibial spur is sinuate, clearly bent downward apically. This characteristic is common in those Onthophagus species somehow associated with burrows or nests of small mammals (Ziani and Gudenzi 2009). Only further investigation will be able to clarify whether the biology of this species is connected to pholeophily.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32D473293BFFACAFC34FC6B.taxon	description	(Fig. 4)	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32D473293BFFACAFC34FC6B.taxon	materials_examined	Type locality. “ Nilghiris: Coonour ”; “ Madura distr.: Shembaganur ”; “ Madras ” [South India, Tamil Nadu state]. Type material. Number of type specimens not specified in the original description, but more than one. One specimen, a major ♂, labelled as “ type ” and with a label handwritten by Boucomont himself, here considered a syntype, examined by photos (MNHN). Diagnostic features. Length 5.0 to 7.5 mm. Colour brownish black, elytra moderately shiny, with distinct isodiametric microreticulation, head and pronotum shinier, with cupreous or greenish metallic lustre, almost lacquered, not microreticulate. Antennal scape, pedicel and funicle dark red, antennal club brown. Dorsal pubescence whitish yellow. Head short, clearly wider than long, with clypeus broadly round, slightly or not at all sinuate anteriorly, sides not sinuate; clypeofrontal carina distinct, sometimes barely discernible, bent backward, clearly closer to the base of the horn than to clypeal anterior margin in major males, extended in a slightly sinuate lamina, placed between the eyes, in the middle of them, in females; occipital carina extended in a long horn arising in correspondence of the anterior margin of eyes, broad and flat at base, directed backward, tapering gradually to the extremity and following the curvature of pronotum, often extending past the middle of the body, sometimes bifurcate apically, in major males, reduced to a narrow straight lamina in minor males and females; clypeal surface with transversely rugose, setigerous punctures, frontal surface doubly sparsely punctate. Pronotum convex, declivous anteriorly, with a distinct anteromedian depressed smooth and unpunctured area and an anterolateral small tubercle on either side, a little behind the front margin, in major males, without depression and with an anteromedian transverse row, usually bilobate, besides the anterolateral tubercle, sometimes hardly appreciable, on either side, in minor males and females; both sexes with a small, hardly distinguishable, in some specimens almost undiscernible, posterolateral groove on either side near pronotal posterior angles; anterior angles distinctly produced, sides not sinuate behind them in dorsal view; dorsal surface setigerously punctate, punctures broad, sub-regular in distribution, separated by 1 / 2 to 1 diameter on disc, absent in the median depression, bearing pale-yellow setae, shorter on disc, barely longer at sides. Elytral striae barely shiny, distinctly impressed, with punctures slightly larger than strial width and barely crenulating interstrial sides; interstriae convex, particularly the external ones, all rather regularly granulate; granules smaller than strial punctures; posterior margin of each granule with a small, indistinct, setigerous puncture; setae short, pale-yellow, thin. Pygidium with rather regularly distributed, setigerous punctures; setae pale-yellow, thin, clearly longer that those of elytra. Males with protibial spur directed outward, and with a small denticle strongly curved downward at the inner angle of protibial apex. Male genital armature. Parameres short, apices bent ventrally, slightly diverging apically, without basolateral plate denticle, round at apex (Fig. 4 b – c); endophallus with the presence of accessory endophallites, a single raspula and a U-shaped lamella copulatrix, frayed at one end, enlarged and bifurcate at the other (Fig. 4 d).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32D473293BFFACAFC34FC6B.taxon	distribution	Distribution. South India (Boucomont 1914). Pakistan (Petrovitz 1961). North India (Sewak 1986). Nepal (Shrestha 2005).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32D473293BFFACAFC34FC6B.taxon	materials_examined	Material examined. India: Tamil Nadu state, “ Nilghiris / M. Maindron ”, “ Coonour / 15 - 30 Juill. 1901 / 1500 - 2000 m. alt. ”, “ Boucomont det. 1914 / Onthophagus / ensifer n. sp. ♂ ”, “ Typus ”, Muséum Paris / 1936 / Coll. A. Boucomont ”, examined by photos (syntype ♂ of O. ensifer, MNHN); Tamil Nadu state, “ Nilgiri Hills, 1922, H. L. Andrewes Bequest, compared with type. G. J. A [rrow] ”, 2 ♂♂ and 1 ♀ (MCSN); Tamil Nadu state, “ Shembaganur ” [Senbahanoor], 3 ♂♂ and 4 ♀♀ (Balthasar collection, NMPC). Historical review. Boucomont (1914) described O. ensifer on an unspecified number of specimens coming from three different localities, “ Coonour ”, “ Shembaganur ” and “ Madras ” whose present names are respectively Coonor, Senbahanoor and Chennai, all in the state of Tamil Nadu, South India. According the Article 73.2.3 of the Code (ICZN 1999), the type locality of the taxon encompasses all these places. Boucomont (1914) also stated that the new species could be a junior synonym of Onthophagus pardalis (Fabricius, 1798), described, as Copris, from “ India ”. Unfortunately, always according to Boucomont (1914), Fabricius description was not sufficient for establishing the synonymy, and what’s more, the type of O. pardalis was lost. Instead, Arrow (1931), citing Fabricius’ species, added an extensive description of both male and female and gave new records from central and southern India. He confirmed that pardalis ’ type had disappeared, but its original description appeared to him on the whole enough to apply to the concerned taxon. Therefore (Arrow 1931) considered it bona species, not conspecific with O. ensifer. Balthasar (1963), placed O. pardalis in the subgenus Colobonthophagus despite expressing the same considerations exposed by Arrow (1931), concerning its relationship with O. ensifer and O. mopsus. Shortly after, Zimsen (1964) added a little twist to the story, reporting the existence of a specimen of O. pardalis type series, preserved in Kiel Museum (Germany). The existence of this specimen gave Scheuern (1996) the opportunity to provide another systematic interpretation of the issue. According to this author, the placement of O. pardalis by Balthasar (1963) was incorrect, as the species that Balthasar – and before him Arrow (1931) – called O. pardalis was not conspecific with Fabricius’ species nor with O. ensifer or O. mopsus. Scheuern (1996), after having designated the lectotype of O. pardalis, claimed not to be able to insert the species in any known subgenera of Onthophagus. Despite this, Thomas et al. (2011), Karimbumkara and Rayan (2013), Mittal and Jain (2015), Karel et al. (2020) and Gupta et al. (2022) considered O. pardalis a Colobonthophagus, probably referring to the O. pardalis sensu Arrow and sensu Balthasar, that Scheuern (1996) named Onthophagus (Colobonthophagus) neocolobus Scheuern, 1996. I could examine pictures of the lectotype of Onthophagus pardalis (Fabricius, 1798), and can confirm its exclusion to the group treated in this paper, as already stated by Scheuern (1996).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32D473293BFFACAFC34FC6B.taxon	discussion	Remarks. I was able to study specimens coming from Tamil Nadu state (southern India) only. The records listed in literature from northern India, Pakistan and Nepal are reasonable but anyway should be confirmed. The morphology of the aedeagus, particularly of paramera (Fig. 4 b – c), short and not elongate as in the other species treated here, casts some doubts on the placement of O. ensifer in the subgenus Indonthophagus. Nevertheless, for now I prefer to keep the species in this group, based on the shape of the lamella copulatrix that follows the pattern of the other species assigned to the subgenus.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32E473793BFFC17FAEFF92B.taxon	description	(Fig. 5, 12 a, c)	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32E473793BFFC17FAEFF92B.taxon	materials_examined	Type locality. “ Minhla, Birmania ” [Minhla, Bago Region, Myanmar]. Type material.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32E473793BFFC17FAEFF92B.taxon	description	Head short, slightly wider than long, with clypeus broadly round, slightly or not at all sinuate anteriorly, sides not sinuate; clypeofrontal carina distinct, bent backward, placed nearly halfway between base of the horn and anterior clypeal margin in major males; occipital carina with an extremely long and slender thread-like horn, slightly enlarged at base (Fig. 12 a), arising between the eyes and curving backward, sometimes extending beyond the hind margin of pronotum, in major males, reduced to a short transverse tubercle between the eyes in minor males, and into a straight or slightly sinuate transverse carina (Fig. 12 c), subequal in length to the clypeofrontal carina, in females; clypeal surface with transversely rugose, setigerous punctures, frontal surface doubly sparsely punctate. Pronotum convex, declivous anteriorly, with a shallow anteromedian narrow groove in major males, with an anteromedian transverse gibbosity, sometimes hardly appreciable, in females; both sexes with a slightly depressed posterolateral area, duller than pronotal disc, on either side near pronotal posterior angles, anteriorly with a tuft of longer setae; anterior angles distinctly produced, sides not sinuate behind them in dorsal view; dorsal surface setigerously punctate, punctures slightly impressed, sub-regular in distribution, separated by 1 to 2 diameters on disc, subequal in size to the punctures of elytral striae, bearing pale-yellow setae, shorter on disc, barely longer at sides. Elytral striae shiny, distinctly impressed, with punctures slightly larger than strial width and barely crenulating interstrial sides; interstriae flat to barely convex, all rather regularly densely granulate; granules smaller than strial punctures; posterior margin of each granule with a small, indistinct, setigerous puncture; setae pale-yellow, thin. Pygidium with rather regularly distributed, setigerous punctures; setae hook-like at the extremity, pale-yellow, thin, clearly longer that those of elytra. Males with protibial spurs directed outward. Distal margin of male protibiae at right angles to the inner margin, with a tuft of short pale-yellow bristles, and with a small denticle strongly curved downward at the inner angle of protibial apex. Male genital armature. Parameres short, apices bent ventrally, diverging apically, without basolateral plate denticle, round at apex (Fig. 5 b – c); endophallus with the presence of accessory endophallites, two evident raspulae of different sizes and a U-shaped lamella copulatrix, bifurcate at one end, clearly divided in two arms at the other (Fig. 5 d).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32E473793BFFC17FAEFF92B.taxon	distribution	Distribution. Myanmar (van Lansberge 1885). South-Eastern China, “ Indochina ” (Boucomont 1929). Taiwan (Arrow 1931). Central China (Balthasar 1963). Thailand (Hanboonsong et al. 1999; Kabakov and Napolov 1999). Vietnam (Kabakov and Napolov 1999). All the western records (Nepal (Shrestha 2005); India (Mittal and Jain 2015); Sri Lanka (Schoolmeesters 2023 )) have to be considered incorrect until proven otherwise.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32E473793BFFC17FAEFF92B.taxon	materials_examined	Material examined. Myanmar: “ Minhla / Birmania / D [onavit]. Comotto 18 __ ”, “ Typus ”, “ hastifer / Lansb. / ♂ ”, “ Onthophagus / Hastifer / Lansberge [Lansberge’s handwriting] ”, “ MUSEO CIVICO / GENOVA ”, “ HOLOTYPE / Onthophagus / hastifer / Lansberge ”, (holotype ♂ of O. hastifer, MCSN); “ Rangoon [Yangon] ”, L. Fea leg., A. Boucomont det. ”, 2 ♀♀, examined by photos (MCSN); “ Bhamò [Bhamo], vi. 1886, L. Fea leg., A. Boucomont det. ”, 1 minor ♂ (MCSN); “ Da Yonang-Young / a Mandalay, v. 1886, L. Fea leg., A. Boucomont det. ” 1 ♂ (MCSN). Thailand: Pak Chong, i-iii 1989, S. Steinke leg. 1 ♂ and 1 ♀, J. Schönfeld det. (MCSN); Chonburi prov., Pattaya, 5 km E Elefantcamp, 30. i. 1995, A. Weigel leg. 1 ♂ (JSCS). Laos: Champassak prov., 25 km SE Pakxe, 200 m, bank of Banglieng river, 14 ° 58 ’ N 105 ° 55 ’ E, 30. iii. 1998, O. Merkl and G. Csorba leg. 2 ♂♂ (JSCS); Viang Chan prov., Ban Pa Kho resort, 50 km NE Vientiane, 90 m, 9 - 14. vi. 2007, M. Štrba leg. 2 ♂♂ (JSCS). Vietnam: “ Hué ”, southern Vietnam, 2 ♂♂ (JSCS). China: Hong Kong, 2 ♂♂ (NHMUK); Fujian prov., Guangze, 24. ix. 1937, J. Klapperich leg. 1 ♂ (JSCS). Taiwan: “ Tainan / Formosa / H. Sauter, VII. 11 [date handwritten]. ” (2 syntypes ♀♀ of O. turmalis, NHMUK); “ Chokakurai / Da Ibu, Formosa, 28 - vi- 1936, Coll. Y. Yano ”, examined by photos (holotype ♂ of O. agilis, EIHU); “ Chokakurai / Da Ibu, Formosa, 28 - vi- 1936, Coll. Y. Yano ”, examined by photos (holotype ♀ of O. putealis, EIHU); Pingtung Co., Hengchun / Shuiwaku, 14. ix. 2021, B. - H. Ho leg. 1 minor ♂ and 1 ♀ (SZCM); Pingtung, N of Shuiwaku, 80 m, 21 ° 56 ′ 29 ″ N 120 ° 50 ′ 23 ″ E, 20. vi. 2023, D. Král and J. Růžička leg. 1 major ♂ and 4 ♀♀ (SZCM). Historical review. Onthophagus hastifer was described by van Lansberge (1885) most probably on a major male only, from Minhla, Myanmar. According to R. Poggi (MCSN, personal communication), there is a sole specimen, a major male, in the collection of the Genoa Museum, with a label written by van Lansberge. This specimen is to be considered the holotype. The female was subsequently described by Boucomont (1914) on a specimen also from Myanmar and deposited in MCSN. Boucomont and Gillet (1921) ran into a misprint, and with a misplaced asterisk, excluded O. hastifer from the fauna of French Indochina, in favour of O. turbatus, described from Sri Lanka and most probably not distributed east of Bhutan (see below). Such mistake was later partially amended (Boucomont and Gillet 1927) with the citation of O. turbatus Boucomont, 1921, not Walker, junior synonym of O. hastifer. Onthophagus turmalis Gillet, 1924 was described from Taiwan on at least a male and a female, and was synonymized with O. hastifer by Arrow (1931) after 7 years from its description. A misspelling involved this taxon, wrongly cited as “ O. turmaris ” by Miwa (1930), Matsumura (1938) and Masumoto (1976 b). Krajcik (2006) listed twice Onthophagus turmalis both as valid species (as Onthophagus turmalis Gillet, 1930, wrong year) and, doubtfully, a junior synonym of O. hastifer. Later, the same author (Krajcik 2012, 2013) confirmed the validity of the species (always dated 1930). Matsumura (1938) described 13 new species of Onthophagini from “ Formosa ” [Taiwan]. Of which only five are presently considered bonae species. Among those that were synonymized, there are two species treated in this paper: Onthophagus agilis and Onthophagus putealis, described from the same locality (Chokakurai), collected on the same day (28. vi. 1936) and by the same entomologist (Y. Yano). O. agilis, described on a single male, was treated as a doubtful species by Balthasar (1963) and Masumoto (1976 a; 1976 b). This last author (Masumoto 1976 b) affirmed “ I still don’t know what it is ” (verbatim, from the Japanese). But it was from Masumoto et al. (2006) that O. agilis was treated as a junior synonym of O. hastifer. Onthophagus putealis, instead, has a little different nomenclatural history. Balthasar (1963), quoting Nakane (1956), considered O. putealis Matsumura, 1938, described on a sole female, as a junior synonym of Onthophagus viduus von Harold, 1874. Actually, Nakane (1956) excluded O. putealis from the twenty species consider as junior synonym of O. viduus. The synonymy by Balthasar (1963) is to be considered incorrect. So, as stated for the first time by Masumoto (1976 a), the correct synonymy is Onthophagus putealis Matsumura, 1938 = Onthophagus hastifer van Lansberge, 1885. It should be noted that O. putealis has been listed twice in Löbl et al. (2006) and in Ziani and Bezděk (2016), both as a junior synonym of O. viduus and, as O. puetalis, misprint, as junior synonym of O. hastifer. As mentioned above, the first reference is wrong and should be deleted. There is another species described by Matsumura (1938), Onthophagus komabellus, supposed to be a junior synonym of O. hastifer by Balthasar (1942). Later though, Nakane (1956) proposed O. komabellus as junior synonym of O. viduus, and all the subsequent authors, from Balthasar (1963), followed this synonymy.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32E473793BFFC17FAEFF92B.taxon	discussion	Remarks. Present literature (Ziani and Bezděk 2016 and Schoolmeesters 2023) considers O. turmalis, O. agilis and O. putealis as junior synonyms of O. hastifer. After studying directly two syntypes of the first species and examining high resolution images of holotypes of the other two species, I can confirm such synonymies.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32B472B93BFF950FA05F99E.taxon	description	(Fig. 1, 6 – 7)	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32B472B93BFF950FA05F99E.taxon	materials_examined	Type locality. “ … in India orientali ” [East Indies]. Type material. Scarabaeus mopsus Fabricius, 1792: lectotype, a minor ♂, designated by Arrow (1931), examined by photos (OUMNH); 4 paralectotypes, 2 ♂♂ and 2 ♀♀, all examined by photos (ZMUK). Copris gracilicornis Germar, 1813: a syntype ♂, examined by photos (ZMB). Diagnostic features. Length 6.0 to 8.0 mm. Colour blackish brown or bronzed, sometimes with slight metallic lustre, moderately shiny, with distinct isodiametric microreticulation. Elytra sometimes with dark red irregular humeral and apical spots. Antennae reddish yellow more or less pale. Dorsal pubescence whitish yellow. Head short, slightly wider than long, with clypeus broadly round, slightly or not at all sinuate anteriorly, sides not sinuate; clypeofrontal carina distinct, bent backward; occipital carina of major males extended in an extremely long and slender thread-like horn, its base a little enlarged and very close, nearly adjacent to the clypeofrontal carina, horn arising in front of the eyes and curving backward, sometimes extending beyond the hind margin of pronotum, reduced to a very short, erected conical tubercle, placed between the eyes, in minor males, to a straight carina between the eyes, sometimes slightly downwards at the ends, shorter than or subequal to clypeofrontal carina, in females (Fig. 1 a); clypeal surface transversely rugose, setigerous punctate, frontal surface doubly sparsely punctate. Pronotum convex, declivous anteriorly, with a distinct anteromedian smooth and unpunctured area delimited by a strong tubercle on either side, a little behind the front margin in major males, with a smooth vertical anterior margin, delimited by a short straight carina in the middle of its upper edge, sometimes divided in two, in minor males and females; both sexes with a small – in some specimens, particularly females, almost undiscernible – posterolateral depressed area on either side near pronotal posterior angles, sometimes distinguishable only because it is slightly duller than the surrounding surface; anterior angles distinctly produced, sides not sinuate behind them in dorsal view; dorsal surface setigerously punctate, punctures sub-regular in distribution, separated by 2 to 3 diameter on disc, bearing pale-yellow setae. Elytral striae moderately shiny, distinctly impressed, with punctures slightly larger than strial width and barely crenulating interstrial sides; interstriae flat to barely convex, all rather regularly granulate; granules smaller than strial punctures; posterior margin of each granule with a small, indistinct, setigerous puncture; setae short, pale-yellow, thin. Pygidium with rather regularly distributed, setigerous punctures; setae pale-yellow, thin, a little longer than those of elytra. Males with protibial spur outward, and with a small denticle strongly curved downward at the inner angle of protibial apex. Male genital armature. Parameres short, apices bent ventrally, little diverging apically, without basolateral plate denticle, latero-apical angle round or almost so (Fig. 6 b – c); endophallus with the presence of accessory endophallites, two evident raspulae and a U-shaped very thin lamella copulatrix, sharp at one end, broad and slightly fringed or bifurcate at the other (Fig. 6 d).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32B472B93BFF950FA05F99E.taxon	distribution	Distribution. West Bengal [India] (von Harold 1869). Nepal (Boucomont 1914, as O. gracilicornis). Pakistan, North India, Kashmir (Arrow 1931). Afghanistan (Balthasar 1956). South India (Rajan 2006).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32B472B93BFF950FA05F99E.taxon	materials_examined	Material examined. Pakistan. Islamabad Capital Territory, Islamabad, 6. vi. 1991, S. Prepsl leg. 1 ex. (SZCM); Islamabad Capital Territory, Marghalla hills, 1000 m, 15. vii. 2010, G. Sabatinelli leg. 3 exx. (SZCM); Islamabad Capital Territory, Islamabad, 15. ix. 2010, G. Sabatinelli leg. 5 exx. (SZCM); Islamabad Capital Territory, Marghalla hills, 23. iv. 2011, S. Ziani leg. 10 exx. (SZCM); Islamabad Capital Territory, Islamabad, sect. E 7, 600 m, 20. vi. 2011, G. Sabatinelli leg. 1 ex. (SZCM); Islamabad Capital Territory, Marghalla hills, 1060 m, 22. vi. 2011, G. Sabatinelli leg. 1 ex. (SZCM); Islamabad Capital Territory, Marghalla hills, 1060 m, 16. vii. 2012, G. Sabatinelli leg. 3 exx. (SZCM); Balochistan, Avaran (Khuzdar), 4. iv. 1993, Bečvář leg. 3 exx. (SZCM); Balochistan, Bela, 21. iv. 1993, Bečvář leg. 1 ex. (SZCM); Punjab, Kallar Kahar, 14. iii. 2007, Z. Ahmed leg. 1 ex. (SZCM); Punjab, Bhan Kotly Sathia, 25. iv. 2011, S. Ziani leg. 6 exx. (SZCM); Punjab, Melod, 27. iv. 2011, S. Ziani leg. 5 exx. (SZCM); Punjab, Chakri, 26. vi. 2011, Z. Ahmed leg. 2 exx. (SZCM); Punjab, Ransial (Kallar Kahar), 800 m, 11. iv. 2012, S. Ziani leg. 2 exx. (SZCM); Gilgit - Baltistan, Goner Farm / 25 km E Chilas, 1100 m, 2. vi. 2007, G. Carpaneto leg. 2 exx. (SZCM); Khyber Pakhtunkhwa, Mansehra, vi. 2009, G. Sabatinelli leg. 3 exx. (SZCM); Khyber Pakhtunkhwa, Mansehra – Jaba Dara, 28. iv. 2011, S. Ziani leg. 5 exx. (SZCM); Khyber Pakhtunkhwa, Mansehra, 10. vii. 2011, G. Sabatinelli leg. 1 ex. (SZCM); Khyber Pakhtunkhwa, Mansehra, viii. 2011, G. Sabatinelli leg. 3 exx. (SZCM); Khyber Pakhtunkhwa, 21 km N Mingora, 1024 m, 7. iv. 2012, S. Ziani leg. 3 exx. (SZCM); Azad Kashmir, Sawa Hill, 3566 m, 26. iv. 2011, G. Sabatinelli leg. 1 ex. (SZCM); Azad Kashmir, Rawala Kot – Banjosa lake 1980 m, 26. iv. 2011, G. Sabatinelli leg. 2 exx. (SZCM). India: “ Mopsus / [illegible] ”, “ in India or. / Monson mopsus Fabr. ”, “ Onthophagus / mopsus / type ”, “ Type / Onthophagus / mopsus, F. / G. J. Arrow det. / Faun. Brit. Ind. / Lamellicornia. / iii, Coprinae. / Dec. 1931 ”, “ Type Col: 458 / Onthophagus / mopsus Fabr. / Hope Dept. Oxford ”, examined by photos (lectotype of O. mopsus, OUMNH); “ Mopsus ”, “ Type i Oxford ”, examined by photos (4 paralectotypes, 2 ♂♂ and 2 ♀♀, of O. mopsus, ZMUC); Himachal Pradesh state, W Himalaya - Pirpandzhal range near Kullu, 1500 m, 21. vii. 2003, A. Gorodinski leg. 3 exx. (SZCM). Nepal: Bagmati Prov., Chitwan National Park – Sauraha 213 m, 4. vi. 1983, M. J. D. Brendell leg. 19 exx. (NHMUK); Bagmati Prov., Chitwan National Park – Sauraha, 11. iv. 2009, L. Nádai leg. 1 ex. (SZCM); Karnali prov., Humla distr., between Simikot – 29 ° 56 ′ 33 ″ N 81 ° 51 ′ 27 ″ E – and 29 ° 54 ′ 23 ″ N 81 ° 55 ′ 7 ″ E, 2975 m- 2990 m, 16. vi. 2022, D. Telnov leg. 4 exx. (NHMUK). Without locality: “ Germari / typ. ”, “ gracilicornis / Gr. ”, “ Holotype ♂ ”, “ Ex-Musaeo / E. Harold ”, examined by photos (syntype ♂ of O. gracilicornis, ZMB). Historical review. Fabricius (1792) described Onthophagus mopsus, as Scarabaeus mopsus, from “ East Indies ”, that presently encompasses, in its broadest context, India, the mainland Southeast Asia, the Malay Archipelago including the Philippines, and the Republic of Indonesia (Encyclopaedia Britannica 2018). It is not possible to know exactly where the specimen or the specimens described by Fabricius came from, but according to the present distribution of the species, it is reasonable to think that the type locality was in the Western portion of this area. “ Onthophagus brunneus Meg [erle] ” was given as a possible synonym of O. mopsus by Dejean (1821, 1833, 1836). Later von Harold (1869) pointed out that Onthophagus brunneus Megerle von Mühlfeld was a “ nomen nudum ” and, as such, not available in the light of the current nomenclatural rules (Article 12 of the Code, ICZN 1999). In the taxonomic history of taxa involved with O. mopsus, there are another two possible cases of “ nomina nuda ”. The first one concerns Onthophagus rupicapra Illiger, cited by Dejean (1833, 1836) as “ var. ” of O. mopsus. As far as I know, Illiger never described a taxon named as such. Therefore, the same nomenclatural rule applies for Onthophagus rupicapra Illiger and consequently the name is not available. Beware, though, because the epithet ruficapra is currently in use for a valid Australian species, Onthophagus rupicapra Waterhouse, 1894 (Schoolmeesters 2023). A second case is represented by “ Onthophagus longicornis Westermann ”, deemed as a synonym of O. mopsus by Dejean (1833, 1836). Bernt Wilhelm Westermann (1781 – 1868) was a Danish merchant, amateur entomologist and tropical insect collector (Dohrn 1868). He published only one paper on entomology (Westermann 1821), where Onthophagus is not cited. Therefore, in agreement with von Harold (1869), it can be concluded that O. longicornis Westermann is a “ nomen nudum ”, never published. The name was probably assigned by Westermann himself to one or more specimens present in his collection. Germar (1813) mentioned Copris gracilicornis, attributing its authorship to Megerle von Mühlfeld, who, indeed, described the species and made the name available in his paper dated 1803 (Megerle von Mühlfeld 1803). Nevertheless, this last work, together with ten more authored by Megerle von Mühlfeld between 1801 and 1805, the so-called “ J. C. Megerle’s (1801 – 1805) auction catalogues of insects ” (ICZN 1993), have been suppressed for nomenclatural purposes by the International Commission on Zoological Nomenclature (ICZN 1993). Almost all the names featured in those publications were ruled not to be available, including Copris gracilicornis. Because of it, the name Copris gracilicornis became available from Germar (1813), who redescribed it on one male and one female at least. Furthermore, it must also be said that “ America ”, the specimen (s) locality indicated by Germar (1813), is clearly incorrect, as evidenced by the collecting place (“ Beng. ” [Bengal]) presented in Megerle von Mühlfeld (1803) paper. Arrow (1931) distinguished two “ gracilicornis ”: the first, as Copris gracilicornis, attributed to Germar (1813), the second, as Onthophagus gracilicornis, attributed to Boucomont (1914). Both taxa are deemed junior synonyms of O. mopsus. Why this distinction, since Boucomont never described an Onthophagus gracilicornis? Was it a simple mistake? It would be unusual for Arrow. Maybe he referred to the claim of Boucomont (1914) that Onthophagus ensifer, just described in that paper, was present as O. gracilicornis in most entomological collections at that time. But this explanation is not convincing since Arrow (1931) did list also O. ensifer Boucomont, 1914 as valid species. I was unable to find a nomenclatorial or systematic meaning for this distinction and for the citation of Onthophagus gracilicornis Boucomont, 1914, done by Arrow (1931). Unfortunately, the same distinction, maybe a copy and paste operation, is present also in Rajan (2006). Therefore, I am forced to leave the problem unsolved. The name gracilicornis has a nomenclatorial convoluted history. Howden and Cartwright (1963) mentioned O. gracilicornis Sturm, 1843 as junior homonym of O. gracilicornis Germar. But in this case such authorship is clearly a mistake: Howden and Cartwright (1963) missed the fact that Sturm (1843) never described the species and indeed, he correctly attributed the name to Megerle von Mühlfeld. Kaur and Yadav (2014, 2018) considered gracilicornis Germar as a subspecies of O. mopsus. No other authors have subsequently shared this taxonomic concept. The name Onthophagus gracilicornis Germar, 1813 gave rise to two homonyms, at least: Onthophagus gracilicornis Fåhraeus, 1857 and Onthophagus gracilicornis Raffray, 1877, both Afrotropical. However, the nomenclatorial problems connected to these taxa are beyond the scopes of this paper.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D32B472B93BFF950FA05F99E.taxon	discussion	Remarks. It is not known how many specimens Fabricius (1792) examined for describing Scarabaeus mopsus. According to Zimsen (1964), the type series currently available is composed by six specimens, one housed in OUMNH, five in ZMUK. Among these six specimens, Arrow (1931) selected the specimen in OUMNH, naming it “ Type ” both in the book “ The Fauna of the British India … ” (Arrow 1931) and in the label that he added to the specimen in OUMNH (Fig. 7). According to the Article 74.5 of the Code (ICZN 1999), this has to be considered a valid designation of a lectotype. This designation, not taken into account in subsequent literature, turned out to be very important to stabilise the nomenclature, especially because the five specimens in ZMUK are not conspecific. After Arrow’s nomenclatorial act, such specimens became automatically paralectotypes, all but one, to which I added the label “ not conspecific with the lectotype and the paralectotypes of Scarabaeus mopsus Fabricius, 1792 ”.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D336472F93BFFF0EFD57FCFB.taxon	description	(Fig. 8)	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D336472F93BFFF0EFD57FCFB.taxon	materials_examined	Type locality. “ Benisuef ” [Beni Suef, Egypt]; “ Syenam ” [Aswan, Egypt]; “ Ambukohl ” [Ambigol, Sudan]. Type material.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D336472F93BFFF0EFD57FCFB.taxon	description	Head short, slightly wider than long, with clypeus broadly round, slightly sinuate anteriorly, sides not sinuate; clypeofrontal carina distinct, bent backward, placed slightly closer to the base of the horn than to the anterior margin of clypeus in major males; occipital carina with a long slender thread-like horn, thickened and flat at base, arising in correspondence of the anterior margin of eyes, directed backward and tapering gradually to the extremity, in major males, reduced to a short, erected conical tubercle, in minor males, to a sinuate carina, longer than the clypeofrontal one, placed between the eyes or little further back in females; clypeal surface with transversely rugose, setigerous punctures, frontal surface double sparsely punctate. Pronotum convex, declivous anteriorly, with an almost undistinguishable anteromedian smooth and unpunctured area, in major males, with an anteromedian slightly bilobate prominence, in minor males and females; both sexes with a small unpunctured area, duller than the surrounding surface, on either side near pronotal posterior angles; anterior angles distinctly produced, sides not sinuate behind them in dorsal view; dorsal surface setigerously punctate, punctures broad, sub-regular in distribution, separated by 2 to 3 diameter on disc, bearing pale-yellow setae. Elytral striae little or not at all shiny, barely impressed, with punctures slightly larger than strial width and crenulating interstrial sides; interstriae barely convex, all rather regularly granulate; granules spread, smaller than strial punctures, each granule with a small, indistinct, setigerous puncture; setae short, pale-yellow, thin. Pygidium with rather regularly distributed, setigerous punctures; setae pale-yellow, thin, longer than those of elytra. Males with protibial spur outward and with a strongly curved downward denticle on the inner angle of protibial apex. Male genital armature. Parameres short, apices slightly bent ventrally, slightly diverging apically, without latero-proximal denticle, round at apex (Fig. 8 b – c); endophallus with the presence of accessory endophallites, two evident raspulae and a U-shaped lamella copulatrix, with an arm straight, the other broad and clearly divided in two unequal branches (Fig. 8 d).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D336472F93BFFF0EFD57FCFB.taxon	distribution	Distribution. Egypt, Sudan (Klug 1845). Eritrea, “ Arabia ” (Gestro 1889). Iraq, Djibouti, Somalia, Saudi Arabia, Yemen, Pakistan (d’Orbigny 1898). Afghanistan (Balthasar 1956). Rajasthan, India (Frey 1956). Ethiopia (Balthasar 1963). Iran (Baraud 1968). United Arab Emirates, Oman (Gillett 1995).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D336472F93BFFF0EFD57FCFB.taxon	materials_examined	Material examined. Egypt: “ Aegypt Ehrenb. / […], “ nitidulus / Klug ”, examined by photos (4 syntypes of O. nitidulus, 1 ♂ and 3 ♀♀, ZMB). Saudi Arabia: Madinah prov., 10 km N Wadi Reem, 200 m. 17 ° 55 ′ 28 ″ N; 42 ° 15 ′ 20 ″ E, 30. iii. 2017, G. Magnani leg. 1 ex. (SZCM). Yemen: Abyan gov., W Lawdar, env. 1150 m, 13 ° 52 ’ 36 ” N 45 ° 48 ’ 01 ” E, 22. x. 2005, D. Král leg. 1 ex. (SZCM); Al Hudaydah gov., Al Munirah, 21 m, 15 ° 20 ′ 10 ″ N 42 ° 50 ′ 12 ″ E, 31. x. 2005, D. Král leg. 1 ex. (SZCM). Oman: Dhofar gov., Road 47 before Al Mughssyl, 61 m, 16 ° 54 ’ 11 ” N 53 ° 49 ’ 20 ” E, 11. ix. 2000, M. Dellacasa leg. 3 exx. (SZCM); Dhofar gov., Khor Rori, 37 m, 17 ° 02 ’ 27 ” N 54 ° 26 ’ 16 ” E, 12. ix. 2000, M. Dellacasa leg. 3 exx. (SZCM); Dhofar gov., Salalah - Wadi Darbat, 41 m, 17 ° 04 ’ 23 ” N 54 ° 25 ’ 56 ” E, 10. vii. 2018, I. Zappi leg. 1 ex. (SZCM); Dhofar gov., Rakhyut, 27. x. 2022, L. Melloni leg. 4 exx. (SZCM); Dhofar gov., 15 km W Salalah, 20 m, 16 ° 56 ’ 52 ” N 53 ° 57 ’ 37 ” E, 4. iv. 2023, S. Ziani leg. 8 exx. (SZCM); Dhofar gov., Wadi Ayun 840 m, 17 ° 15 ’ 14 ” N 53 ° 53 ’ 33 ” E, 5. iv. 2023, S. Ziani leg. 4 exx. (SZCM). Iran: Lorestan prov., Kumas, 2. iv. 2015 (SZCM); Hormozgan prov., Ahmadi, 10. iii. 2017, 2 exx. (SZCM); Hormozgan prov., Chah Faleh, 19 m, 30. iii. 2018, G. Sabatinelli leg. 2 exx. (SZCM). Pakistan: “ Kurrachee ” [Karachi], viii-ix, Maindron leg. 1 ♂, examined by photos (syntype ♂ of O. nitidulus var. mediofasciatus, d’Orbigny collection, MNHN); Punjab prov., Chakri, motorway for Islamabad, 26. vi. 2011 (SZCM). Historical review. The specimens (syntypes) examined and used by Klug (1845) for describing Onthophagus nitidulus came from three localities, “ Benisuef ” [Beni Suef], “ Syenam ” [Aswan] and “ Ambukohl ” [Ambigol], respectively in northern Egypt, in southern Egypt, and in northern Sudan. The type locality of the taxon encompasses all these three localities (Article 73.2.3 of the Code, ICZN 1999). Kolenati (1846) and von Harold (1867, 1869) mentioned Onthophagus sexstriatus Waltl [“ 6 - striatus Waltl. ”] and Onthophagus niloticus Reiche, presumably as junior synonyms of O. nitidulus. As specified by von Harold (1867), in the first half of the nineteenth century, specimens of this last species were often found in entomological collections under the names “ niloticus ” and “ 6 - striatus ”. It does not appear that Joseph Waltl and Louis Jérome Reiche published papers where, respectively, Onthophagus sexstriatus and Onthophagus niloticus have been described. Therefore, as specified by von Harold (1869), it can be reasonably presumed that these names are “ nomina nuda ”, hence not available. It should be noted that, later on, Montrouzier (1855) and von Harold (1879) described, respectively, Onthophagus sexstriatus from “ Woodlark ” [Papua New Guinea] and Onthophagus niloticus from “ Kordofan, Nilus coeruleus ” [Sudan], valid names of taxa that, obviously, have nothing to do with Onthophagus sexstriatus Waltl, nomen nudum, with Onthophagus niloticus Reiche, nomen nudum, and with Onthophagus nitidulus Klug. Ten years after having described Onthophagus nitidulus, Klug (1855) described “ another ” Onthophagus nitidulus – clearly different from Onthophagus nitidulus Klug, 1845 – from Sena, Mozambique, description confirmed a few years later (Klug 1862) with, in addition, the statement “ nova spec. ” after the name of the species, as if the 1855 paper had never been published. Whatever happened, Klug most likely forgot his 1845 paper and his “ first ” Onthophagus nitidulus. Aware of this clear primary homonymy (Onthophagus nitidulus Klug, 1845 and Onthophagus nitidulus Klug, 1855), von Harold (1867) replaced the junior homonym with the new name virescens. Onthophagus tetraspilus Fairmaire, 1887 was described from the then “ Côte française des Somalis ”, presently Djibouti, and considered close to the female of O. nitidulus. Later d’Orbigny (1898) deemed O. tetraspilus as a simple chromatic variation of O. nitidulus, with elytral spot only on the 3 rd and 7 th interstriae or even just on 7 th. In 1898 d’Orbigny described the variety mediofasciatus of O. nitidulus, from Cairo, Egypt, and from Karachi, Pakistan, with elytral spots joint to form a transversal V-shaped band, sometimes interrupted on the 6 th interstria. Since d’Orbigny’s taxon was published before 1961, it must be considered of subspecific rank according the Article 45.6.4 of the Code (ICZN 1999). From Winkler (1929), mediofasciatus was deemed to be an aberration of O. nitidulus, hence its junior synonym. Janíková (1998) and Bunalski (2014) inserted O. nitidulus in the subgenus Palaeonthophagus Zunino, 1979 without any explanation. No subsequent authors followed this systematic concept.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D336472F93BFFF0EFD57FCFB.taxon	discussion	Remarks. The current systematic literature (Balthasar 1963; Krajcik 2006; Löbl et al. 2006; Ziani and Bezděk 2016; Schoolmeesters 2023) considers O. tetraspilus and O. nitidulus ssp. mediofasciatus junior synonyms of O. nitidulus. This paper follows such direction. According to Zunino (1981), O. nitidulus has phylogenetic affinities with Onthophagus sellatus Klug, 1845 (tentatively inserted in the subgenus Furconthophagus Zunino, 1979 by Ziani 2020 b), and Onthophagus margaritifer d’Orbigny, 1898 (inserted in Onthophagus sensu lato, without a subgeneric placement, by Ziani and Bezděk 2016). After studying the external morphology and the structures of the endophallus of the last two species, I prefer not to insert them in Indonthophagus.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D333472393BFFC80FC7EFB6E.taxon	materials_examined	Type localities. “ Ceylon ” [Sri Lanka]. Type material. 2 syntypes, a medium ♂ and a ♀, studied (NHMUK). Diagnostic features. Length 7.0 to 8.0 mm. Colour reddish brown or bronzed, with slight metallic lustre, moderately shiny, with distinct isodiametric microreticulation. Elytra dark red, sometimes with lighter red irregular basal and apical spots. Antennal scape, pedicel and funicle reddish yellow, antennal club yellow. Dorsal pubescence whitish yellow. Head short, distinctly wider than long, with clypeus broadly round, slightly or not at all sinuate anteriorly, sides not sinuate; clypeofrontal carina distinct, bent backward, placed nearly halfway between base of the horn and anterior clypeal margin; occipital carina with a long and slender thread-like horn, flat and clearly moderately enlarged at base (Fig. 12 b), curving backward, sometimes reaching and also exceeding the pronotal hind margin, in major males, reduced to a short transverse tubercle between the eyes in minor males, and into a straight, sometimes sinuate transverse carina in females (Fig. 12 d), clearly broader than clypeofrontal one, almost reaching inner margin of eyes; clypeal surface with transversely rugose, setigerous punctures, frontal surface doubly sparsely punctate. Pronotum convex, declivous anteriorly, with a slight anteromedian depression in major males, front pronotal margin vertical in the middle, the upper edge of the declivity forming a more or less evident anteromedian prominence in minor males and in females, in the latter the prominence being slightly bilobate; medium males, including the syntype, with horn shortened sometime by half, and a deep wide pronotal anteromedian hollow, its surface smooth and scarcely simply punctate; both sexes with a slightly depressed posterolateral area (Fig. 2), duller than pronotal disc, on either side near pronotal posterior angles, anteriorly with a tuft of longer setae; anterior angles distinctly produced, sides not sinuate behind them in dorsal view; dorsal surface setigerously punctate, punctures slightly impressed, sub-regular in distribution, separated by 1 to 3 diameters on disc, smaller than the punctures of elytral striae, bearing pale-yellow setae, shorter on disc, barely longer at sides. Elytral striae shiny, distinctly impressed, with punctures slightly larger than strial width and barely crenulating interstrial sides; interstriae flat to barely convex, all rather regularly granulate; granules smaller than strial punctures, each granule with a short, pale-yellow, thin seta. Pygidium with regularly or sub-regularly distributed, setigerous punctures; setae pale-yellow, thin, clearly longer that those of elytra. Males with protibial spur outward. Terminal margin of male protibiae at right angle to the inner margin, with a tuft of short pale-yellow bristles, and with a small denticle curved downward on the inner angle of protibial apex. Male genital armature. Parameres short, apices bent ventrally and rhombus-shape, diverging apically, without basolateral plate denticle (Fig. 10 b – c); endophallus with the presence of accessory endophallites, two evident and large raspulae of different sizes and a U-shaped lamella copulatrix, with one arm bifurcate apically, the other with a spinelike process, more or less long but never reaching the apex that appears clearly lobate (Fig. 10 d).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D333472393BFFC80FC7EFB6E.taxon	distribution	Distribution. Sri Lanka (Walker 1858). India (Boucomont 1914). First records for Pakistan, Nepal and Bhutan.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D333472393BFFC80FC7EFB6E.taxon	materials_examined	Material examined. Pakistan: Islamabad Capital Territory, Marghalla hills, 1000 m, 15. viii. 2010, G. Sabatinelli leg. 3 ♂♂ and 2 ♀♀ (SZCM); Islamabad Capital Territory, Islamabad, sect. E 7, 600 m, 1. ix. 2012, G. Sabatinelli leg. 2 ♂♂ at light (SZCM); Khyber Pakhtunkhwa, Jaba Dara (Manshera), 28. iv. 2011, S. Ziani leg. 3 ♂♂ and 1 ♀ (SZCM). India: Kerala state, Cardamon H., Periyar, env. Aranya Nivas, 950 m, 4. xi. 1972, C. Besuchet, I. Löbl, R. Mussard leg. 1 ♀ (MHNG); Karnataka state, Bangalore University Campus, Bangalore, 6. vii. 2004, R. Ramalingam leg. 3 major ♂♂ and 1 medium ♂, examined by photos (AIMB); West Bengal state, Siliguri distr., Sevoke, 200 m. 7. vi. 2008, V. Patrikeev leg. 1 minor ♂ (STCH); Arunachal Pradesh state, West Kameng, Bhalukpong, 150 m, 27 ° 00 ′ 48 ″ N 92 ° 39 ′ 08 ″ E, 1 / 8. v. 2012. L. Dembický leg. 1 major ♂ (SZCM); Kerala state, Nelliyampathy / Palakkad, 97 m, 23. iii. 2022, K. A. Sobhana leg. 1 medium ♂, examined by photos (ZSIK); Kerala state, Meppadi / Wayanad, 987 m, 29. xi. 2022, J. Subha Babu leg. 2 medium ♂♂, examined by photos (ZSIK). Sri Lanka: “ Typus ”, “ Ceylon ”, “ Onth. turbatus / Walker / (type) ”, “ turbatus ” (syntype ♂ of O. turbatus, NHMUK); “ Ceylon ”, “ Onth. turbatus / Walker / (type) ” (syntype ♀ of O. turbatus, NHMUK); Central Province, Kandy, 1892, E. Simon leg. 1 ♂ and 1 ♀ (Balthasar collection, NMPC); Uva prov., Inginiyagala, 12. ii. 1970, Mussard, Besuchet, Löbl leg. 1 ♂ (NMPC); Sabaragamuwa prov., Pinnawala – Rambukkana, 2. ii. 2024, M. Rivalta and P. Subini leg. 1 ♀ (SZCM). Nepal: Central region, Annapurna - Himalaya / near Tatopani, 1300 m, 3. vi. 1996, L. Nádai leg. 1 ♂ (SZCM). Bhutan: Punakha, Sobsokha, 1200 m, 13. vii. 2016, S. Ziani leg. 2 ♀♀ (SZCM). Historical review. Onthophagus turbatus was described on one male and one female at least, since Walker (1858) has featured both sexes. The “ types ” present in NHMUK, with Walker’s handwritten labels, are therefore to be considered syntypes. The taxon did not appear to be born under a lucky star: only four years after its description, O. turbatus was synonymized by von Harold (1862). He affirmed that Walker described the species on a female of O. spinifex (Fabricius, 1781), and later confirmed such synonymy (von Harold 1869). On the contrary, Arrow (1907), claiming to have examined the type material of both species, stated that O. turbatus and O. spinifex were absolutely not conspecific. Boucomont (1914), in a footnote at p. 222, asserted that “ Ce nom [O. turbatus] est probablement synonyme de mopsus F., (…), malheureusement le type de Fabricius est perdu et la description est insuffisante ” [This name [O. turbatus] is probably a synonym of mopsus F., (…), unfortunately Fabricius’ type has been lost and the description is insufficient]. Later, Boucomont, with Gillet, accepted the validity of the species, first in “ Faune entomologique de l’Indochine française ” (Boucomont and Gillet 1921), then in “ Coleopterorum Catalogus ” (Boucomont and Gillet 1927). It is worth noting that Boucomont and Gillet (1921) recorded this species from Vietnam (“ Tonkin: Quang Yen; Annam: Quang-tri), whereas Arrow (1931) specified that these records were wrong. Paulian (1945) hypothesized that O. turbatus sensu Boucomont [and Gillet] (1921), which is the species recorded from Vietnam by Boucomont and Gillet (1921) – and from Myanmar, China and Taiwan by himself – was not conspecific with O. turbatus Walker. This last taxon, always according to Paulian (1945), is distributed in Sri Lanka and south India, whereas in north-western India “ l’espèce est représentée par le vicariant O. mopsus Fabricius ” [the species is represented by the vicariant O. mopsus Fabricius]. O. turbatus sensu Boucomont [and Gillet] 1921, always according to Paulian (1945), is a junior synonym of O. hastifer. Lau (2019) quoted, in the “ Checklist of Insects of Hong Kong ”, “ Onthophagus turbatus Boucomont and Gillet (nec Walker, 1858) ”, but as bona species. Kharel et al. (2020) recorded the species, cited as O. turbatus Walker, also from Thailand. Regarding its subgeneric placement, the taxon was mentioned twice by Kalawate et al. (2021), either belonging to the subgenus Onthophagus and as “ species incertae sedis ”.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D333472393BFFC80FC7EFB6E.taxon	discussion	Remarks. After the study of type material of O. turbatus and O. hastifer, as well as other specimens belonging to both taxa, I fully agree with Paulian (1945) in considering O. turbatus sensu Boucomont [and Gillet] 1921 not conspecific with O. turbatus Walker, and a junior synonym of O. hastifer. Consequently, the distribution of O. turbatus Walker does not include localities eastward of Bhutan. Most probably O. turbatus and O. hastifer are sister species and, for what I know, their distributional areas are not overlapping. All the morphological descriptions of O. turbatus published in the literature (Boucomont 1914; Boucomont and Gillet 1921; Arrow 1931; Balthasar 1963), including the original description (Walker 1858), report as the main distinctive characters of males, the cephalic horn, short and far from reaching the pronotal hind margin, and the pronotal deep and wide antero-median depression. The authors who pointed out such features had clearly described the syntype male or had reported a previous description of the syntype male, without noticing, or stressing at least, that the specimen was not a major male, but a so-called “ medium ” male, with all the characteristics of moderately developed males, such as shorter cephalic horn and evident pronotal antero-median depression (Fig. 10 a). Actually, major males of O. turbatus have the cephalic horn reaching and sometimes exceeding the pronotal hind margin, and a very shallow anteromedian hollow, if any (Fig. 11). The morphology of lamella copulatrix leaves no doubts about the conspecificity of major and medium specimens. For these reasons, in the dichotomous key to the Indonthophagus species provided below, also other characters for discriminating O. turbatus from other species, particularly O. hastifer, the taxon in the same couplet, are used.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D33F472693BFFB14FC30FD40.taxon	description	(Fig. 11)	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D33F472693BFFB14FC30FD40.taxon	materials_examined	Type localities. “ Coromandel ” [south-eastern India]. Type material. Scarabaeus spinifex Fabricius, 1781: a syntype (holotype?), a major male, examined by photos (NHMUK). Scarabaeus aeneus Fabricius, 1781: a syntype (holotype?) female, examined by photos (NHMUK). Scarabaeus truncaticornis Herbst, 1786: holotype (?) not traced. Scarabaeus truncaticornis Schaller, 1783: type lost / destroyed (Hendrick Müller, Martin-Luther-Universität, Halle-Wittenberg, personal communication). Onthophagus reflexicornis Redtenbacher, 1868: holotype male, fixed by monotypy, not traced. Onthophagus bifossus d’Orbigny, 1902: holotype female, fixed by monotypy, examined by photos (MNHN). Diagnostic features. Length 7.5 to 10.0 mm. Colour dark metallic blue or dark metallic green, glossy, without distinct isodiametric microreticulation. Antennal scape, pedicel and funicle brownish reddish yellow, antennal club yellow. Dorsal pubescence yellow. Head very slightly wider than long, with clypeus broadly rounded, not sinuate anteriorly, sides not sinuate; clypeofrontal carina distinct, bent backward, very slightly closer to the occipital carina than to clypeal anterior margin in major males; occipital carina bearing a long slender horn, flattened and broad in its basal part, curving and tapering backward, sometimes extending beyond the middle of the body in major males, reduced to a short, erected conical tubercle, placed between the eyes, or to a transverse carina, in minor males, to a strongly elevated carina, curved backward, clearly longer than clypeofrontal carina and extended from eye to eye, in females; clypeal surface with transversely rugose, setigerous punctures, frontal surface sparsely punctate. Pronotum convex, declivous anteriorly, with a longitudinal smooth and almost unpunctured very slight median groove, in major males, with a short anteromedian steep declivity bearing a pair of small tubercles rather close to each other, in minor males, and with an anteromedian curved prominence, slightly elevated, a little depressed in the middle, with a smooth hollow on either side, in females; both sexes with a small posterolateral area on either side near pronotal posterior angles, slightly duller than the surrounding surface and simply sparsely punctate, anteriorly with a line of longer setae; anterior angles produced, sides not sinuate behind them in dorsal view; dorsal surface setigerously punctate, punctures small on disc, very sparse, simple, in major males, a slightly rough in minor males, clearly rough and closer in females, each puncture bearing a short pale-yellow seta, longer at sides. Elytral striae distinctly impressed, with punctures slightly larger than strial width and crenulating interstrial sides; interstriae barely convex, wrinkling granulate; granules spread, equal in size to strial punctures, each gran- Pygidium with rather regularly distributed, setigerous punctures; setae pale-yellow, thin, a clearly longer than those of elytra. Males with protibial spur very slightly bent downward, and with a small denticle curved downward on the inner angle of protibial apex. Male genital armature. Parameres very short, with two symmetrical denticles apically, and two spatulae, slightly diverging, ventrally (Fig. 9 b – c); endophallus with the presence of accessory endophallites and a sub-quadrangular lamella copulatrix, with three arms at least, one arms longer than the others, and a sinuate side (Fig. 9 d).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D33F472693BFFB14FC30FD40.taxon	distribution	Distribution. India (Fabricius 1781). Sri Lanka (Boucomont 1914). Nepal (Löbl et al. 2006). Pakistan (Ziani 2015). Bangladesh (Kharel et al. 2020).	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D33F472693BFFB14FC30FD40.taxon	materials_examined	Material examined. Pakistan: Azad Kashmir, Muzaffarabad, 800 m, 15. v. 2012, G. Sabatinelli leg. 17 exx. (SZCM); Azad Kashmir, Rawalakot – Tatapani, 7. vi. 2012, G. Sabatinelli leg. 1 ex. (SZCM); Khyber Pakhtunkhwa prov., Kalam, 1900 m, 28. ix. 2012, G. Sabatinelli leg. 1 ex. (SZCM); Islamabad Capital Territory, Islamabad, sect. E 7, 600 m, 15. vii. 2012, G. Sabatinelli leg. 1 ex. (SZCM); ibidem, 1. ix. 2012, G. Sabatinelli leg. 4 exx. (SZCM); ibidem, 15. ix. 2012, G. Sabatinelli leg. 1 ex. (SZCM); Islamabad Capital Territory, Marghalla hills, 1056 m, 16. vii. 2012, G. Sabatinelli leg. 2 exx. (SZCM). India: [Coromandel], “ Scarab. spinifex / Fabr. […] ”, “ Syntypus ”, examined by photos (syntype [holotype?] major ♂ of O. spinifex, NHMUK); [Coromandel], “ Scarab. aeneus. / Fabr. […] ”, examined by photos (syntype [holotype?] ♀ of O. aeneus, NHMUK). Sri Lanka: Western prov., Negombo, 1. ii. 1971, L. Lindgren leg. 1 ♂ (MZH). Northern Prov., Madhu, 8. ii. 1971, L. Lindgren leg. 1 ♀. (MZH); Uva prov., Kataragama env., 1 - 3. vii. 2003, O. Mehl leg. 1 ♀ (SZCM). Without locality: “ Senegal ” [patria errata] / Ex-Musaeo / Van Lansberge ”, “ H. d’Orbigny / Onth. Afr. 1902 ”, “ bifossus / n. sp. d’Orb. ”, “ Holotype ”, examined by photos (( syntype [holotype?] ♀ of O. bifossus, MNHN). Historical review. Fabricius (1781) described Onthophagus spinifex, as Scarabaeus spinifex, from “ Coromandel ”. Presently Coromandel is the name of a southeastern costal region of the Indian subcontinent, comprising the states of Tamil Nadu and Andhra Pradesh, but historically Coromandel stretch of coast was shorter, from the mouth of the river Krishna to, southward, the Cape of Calimere (“ Kodiakarai ”). Olivier (1789) wrongly reported the specific name “ spinifer ” for Scarabaeus spinifex. This misprint is to be considered an incorrect subsequent spelling, also because the same author, one year later (Olivier 1790), returned to use the original spelling spinifex, a clear evidence that his 1789 change was not an emendation – i. e. an intentional change – but simply a misprint. As such, the name “ spinifer ”, as incorrect subsequent spelling, is not available and does not enter in homonymy (Article 33.3 of the Code, ICZN 1999). Unfortunately the “ copy and paste ” operations were very fashionable also more than two centuries ago. And so, Illiger (1800) [who two year later (Illiger 1802) realized the mistake and remedied], Sturm (1802), Hope (1838) and even Arrow (1907) fell into Olivier’s misprint and wrongly used “ spinifer ”. Fabricius (1781) described also Scarabaeus aeneus from “ Coromandel ”, just in the same volume in which he described Scarabaeus spinifex. Almost a century later, von Harold (1880), claiming that Fabricius’s aeneus was nothing but the female of spinifex, acted as First Reviser (Article 24.2.1 of the Code, ICZN 1999) and gave the precedence to the name Onthophagus spinifex. The synonymy was followed by all the subsequent authors. According to some recent literature, Onthophagus aeneus, as Scarabaeus, was described twice, by two different authors. Let’s see how. At page 131 of “ Entomologie, ou Histoire Naturelle des Insectes ... ” Olivier (1789) quoted Scarabaeus aeneus Fabricius, 1781. It was simply a citation, not a description of a new species. Also, in “ Dictionnaire encyclopédique de l’histoire naturelle ”, the same author (Olivier 1790), even referring to the image of the beetle in Olivier (1789), clearly gave the authorship of Scarabaeus aeneus to Fabricius (1781). But despite this, some authors (among them Balthasar 1963, Löbl et al. 2006, Rossini et al. 2014, Ziani and Bezděk 2016) considered Scarabaeus aeneus as described twice, the first time in 1781 by Fabricius, then in 1789 by Olivier. In particular, according to Balthasar 1963, Löbl et al. 2006 and Ziani and Bezděk 2016, Scarabaeus aeneus Fabricius, 1781 is a junior synonym of Onthophagus spinifex, whereas Scarabaeus aeneus Olivier 1789 is a junior synonym of Onthophagus dama (Fabricius, 1789). Actually, as stated above, Olivier (1789) cited the Scarabaeus aeneus described by Fabricius (1781), with indications and bibliographical references, even with the same type locality (“ Coromandel ”, south-eastern India), of Fabricius’s species. Therefore, Scarabaeus aeneus Olivier, 1789 was never described and doesn’t enter in any kind of synonymy or homonymy. Blanchard (1847) described Onthophagus aeneus from Argentina. Since this name was a secondary homonym of Onthophagus aeneus (Fabricius, 1781), it was replaced by von Harold (1859) with Onthophagus brasiliensis nomen novum. This taxon is currently considered a junior synonym of Onthophagus hircus Billberg, 1815 (Schoolmeesters 2023). There is also a Scarabaeus aeneus described by Linnaeus (1764) from South Africa but, as already noticed by Jablonsky and Herbst (1789), it is a Cetoniinae (Scarabaeidae) and all the issues related to its homonymies or synonymies are not discussed here. Since it also concerns O. aeneus, it is now necessary to mention the very complicated and tangled story, under a systematic / nomenclatorial point of view, of Onthophagus truncaticornis. It was described for the first time, as Scarabaeus truncaticornis, by Schaller (1783), on an unknown number of specimens from Malabar (southwestern India). Three years later Herbst (1786) claimed the authorship of the name, based on reasons irrelevant under nomenclatorial conventions. He asserted that “ his ” S. truncaticornis differed from Schaller S. truncaticornis only by the presence of head carinae. Strangely enough, Jablonsky and Herbst (1789) gave the precedence to Schaller as author of Scarabaeus truncaticornis. Are the two Scarabaeus truncaticornis really different species? We cannot say it with certainty. According to von Harold (1880) and the current literature, they are so. Anyway, despite the confusion about the authorship, the systematic fate of Scarabaeus truncaticornis intertwined that of Scarabaeus aeneus Fabricius, 1781 – as stated before, junior synonym of O. spinifex – after Fabricius (1792), who affirmed that Herbst’s Scarabaeus truncaticornis was nothing but the female of Scarabaeus aeneus. Later, Sturm (1800) followed Fabricius (1792) synonymy, namely S. truncaticornis Herbst, 1786 = S. aeneus Fabricius, 1781 but added to the synonymy “ the other ” Scarabaeus truncaticornis by Schaller (1783). Gmelin (1790) gave the authorship of S. truncaticornis to Schaller (1781) only. Fabricius (1792) resumed the synonymy Scarabaeus aeneus versus Scarabaeus truncaticornis Schaller, confirming that S. truncaticornis Herbst was the female of S. aeneus. According to Illiger (1800), S. truncaticornis Schaller was a junior synonym of S. aeneus, whereas according to Schönherr (1802), was instead S. truncaticornis Herbst a junior synonym of S. aeneus (as Copris aenea). As it can be seen, nomenclatorial and taxonomic chaos were dominant. Only after Latreille (1802), when all these taxa had been moved into the genus Onthophagus by subsequent authors, it became less mystifying mainly thanks to von Harold (1869) and especially to von Harold (1880). In this last paper von Harold clearly suggested, after seeing the type of Onthophagus truncaticornis (Herbst) preserved in Berlin Museum, that the latter was absolutely conspecific with O. aeneus (Fabricius) – in turn conspecific with O. spinifex – but different from O. truncaticornis (Schaller). All the subsequent authors followed von Harold’s systematic statement, so the nomenclatorial chaos problem is seemingly solved. The choice by Krajcik (2006) to reference “ Herbst in Jablonsky, 1789 ” as the authorship and year of description of O. truncaticornis is not clear to me and even, also in view of Bousquet (2016), seems to be incorrect. And Schaller’s Onthophagus truncaticornis? It is presently accepted as a species recorded from India (Schoolmeesters 2023) and belonging to the O. pusillus group (Palestrini 1983). The treatment of this taxon is, however, out of the topic of this paper. Finally, to make nomenclatorial matter worse, Boheman (1860) described “ another ” Onthophagus truncaticornis from Botswana (Africa), a secondary homonym replaced by von Harold (1870) with the new name Onthophagus trucidatus. O. trucidatus is presently considered a junior synonym of O. quadrinodosus Fåhraeus, 1857 (Schoolmeesters 2023). Onthophagus reflexicornis, described by Redtenbacher (1868) on a single male from Ceylon, was soon synonymized with O. spinifex by von Harold (1872), after examining the type. This synonymy was followed by all subsequent authors. Onthophagus bifossus, described by d’Orbigny (1902) on a single female from Senegal, was synonymized with O. spinifex by d’Orbigny himself (d’Orbigny 1908), on the ground of a possible mislabelled specimen.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D33F472693BFFB14FC30FD40.taxon	discussion	Remarks. Some characteristics of O. spinifex external morphology, such as the depressed and dull area on either side near pronotal posterior angles and the major males’ long thin horn, curved backward, suggest the species could belong to Indonthophagus. And in that subgenus the species was inserted by Kabakov (2006) based on these characters. Actually, females of O. spinifex have a strongly elevated occipital carina, curved backward, clearly longer than the clypeofrontal carina and extended from eye to eye, whereas female members of Indonthophagus have a simple occipital carina, nearly straight, placed between the eyes and more or less extended but never from eye to eye. As to male genitalia, O. spinifex parameres and structures of the endophallus are definitively far from the ones of the other species here included in the subgenus Indonthophagus. Particularly, the lamella copulatrix is not U-shaped but sub-quadrangular. Additionally, the endophallus is lacking raspulae. For the abovementioned reasons, I decided to exclude definitively O. spinifex from the subgenus Indonthophagus, and to insert it in mare magnum of the Onthophagus species needing a subgeneric placement. There is a little mystery in Ghosh et al. (2021): in the geographical distribution of O. spinifex, the code “ CH ” is cited. Nevertheless, such code is not present in the section of that paper called “ abbreviations ” which specifies countries matching the initialisms. What does “ CH ” mean? From a geographical point of view, CH can mean “ China ”, as in Löbl and Smetana (2006) and Löbl and Löbl (2016), but this Country, although not in O. spinifex distribution, is already present in Ghosh et al. (2021) under the code “ CN ”. Switzerland, the other Country that is often abridged as “ CH ”, can be definitely ruled out. So, is CH a misprint for CN, which would be China? If this is the case, should China be added to the distribution of the species? For the moment and until verified records are available, I prudently exclude O. spinifex from the Chinese fauna.	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
7B3FB814D305471893BFFA01FA04FB6F.taxon	description	— Elytra monochromatic dark brown, dark red or bronzed, at most with basal or apical red spots ..... 3	en	Ziani, Stefano (2024): Historical and morphological review of the subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Insecta Mundi 2024 (38): 1-42, DOI: 10.5281/zenodo.11450108
