Onthophagus (Indonthophagus) mopsus ( Fabricius, 1792 )

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), pp. 1-42 : 13-17

Onthophagus (Indonthophagus) mopsus ( Fabricius, 1792)

( Fig. 1 , 6–7 )

Scarabaeus mopsus Fabricius 1792: 58 .

Copris mopsus, Fabricius 1801: 49 ; Schönherr 1806: 50.

Onthophagus mopsus, Dejean 1833: 142 (with a question mark); von Harold 1869: 1033; von Harold 1872: 206; Boucomont and Gillet 1927: 141; Arrow 1931: 328; Balthasar 1956: 435; Petrovitz 1961: 102; Balthasar 1963: 441; Petrovitz 1967: 479; Balthasar 1968: 153; Endrödi 1974: 3; Mittal 1981: 80; Gupta and Mittal 1987: 52; Gupta 1989: 25; Hashmi and Tashfeen 1992: 148; Mittal 1999: 34; Biswas and Ghosh 2000: 590; Mittal 2000: 264; Chandra 2005: 149; Mittal 2005: 46; Krajcik 2006: 119; Rajan 2006: 128; Chandra and Gupta 2012: 100; Krajcik 2012: 182; Karimbumkara and Rajan 2013: 176; Krajcik 2013: 174; Kaur and Yadav 2014: 4; Mittal and Jain 2015: 399; Singh et al. 2017: 18; Kaur and Yadav 2018: 405; Singh 2020: 522.

Onthophagus (incertae sedis) mopsus, Löbl et al. 2006: 175 ; Siddiqui and Kamaluddin 2011: 349.

Onthophagus (Furconthophagus) mopsus, Siddiqui et al. 2014: 304 .

Onthophagus (Indonthophagus) mopsus, Kabakov 2006: 154 (footnote); Ziani and Bezděk 2016: 188; Gupta et al. 2018: 484; Ghosh et al. 2020: 243; Kharel et al. 2020: 368; Ghosh et al. 2022a: 11; Ghosh et al. 2022b: 42; Gupta et al. 2022: 425; Schoolmeesters 2023.

Copris gracilicornis Germar 1813: 114 [type locality: “America”, patria errata]; Arrow 1931: 328 (as junior synonym of O. mopsus ).

Onthophagus gracilicornis, Dejean 1821: 53 (as “ gracilicornis Meg. ”); Sturm 1826: 178 (as “ gracilicornis Meg. ”); Dejean 1833: 141 (as “ O. gracilicornis Dej. ”); Dejean 1836: 157 (as “ O. gracilicornis Dej. ”); Sturm 1843: 107 (as “ gracilicornis Megerl. ”); von Harold 1869: 1030; van Lansberge 1885: 381; Boucomont 1914: 220; Boucomont and Gillet 1927: 138; Arrow 1931: 328 (as O. gracilicornis Boucomont, 1914 , junior synonym of O. mopsus ); Balthasar 1963: 441 (as junior synonym of O. mopsus ); Krajcik 2006: 119 (as junior synonym of O. mopsus ); Löbl et al. 2006: 175 (as junior synonym of O. mopsus ); Rajan 2006: 128 (as O. gracilicornis Boucomont, 1914 , junior synonym of O. mopsus ); Ziani and Bezděk 2016: 188 (as junior synonym of O. mopsus ).

Onthophagus mopsus ssp. gracilicornis, Kaur and Yadav 2014: 4 ; Kaur and Yadav 2018: 405.

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. 1a ); 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. 6b–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. 6d ).

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).

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.

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 ”.

Figures

Figure 1. Onthophagus (Indonthophagus) mopsus (Fabricius, Figure 2. Onthophagus (Indonthophagus) turbatus Walker, 1792) [Pakistan, Islamabad Capital Territory]. 1a) ♀ head, 1858 [Sri Lanka, Uva prov.]. Detail of pronotal posterolateral from above. 1b) Epipharynx. Photos by A. Degiovanni, ed- area. Photos by A. Degiovanni, edited by G. Fiumi.

Figure 6. Onthophagus (Indonthophagus) mopsus (Fabricius, 1792) [major ♂: Pakistan, Islamabad Capital Territory]. 6a) Dorsal habitus; 6b) Parameres, lateral view. 6c) Parameres, dorsal view. 6d) Endophallus. AcS: accessory endophallites. LC: lamella copulatrix. Rs: raspulae. Photos by A. Degiovanni, edited by G. Fiumi.

Figure 7. Onthophagus (Indonthophagus) mopsus (Fabricius, 1792), lectotype labels. Photo by Katerine Child (OUMNH).