Onthophagus (sensu lato) spinifex ( Fabricius, 1781 )

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 : 25-30

[ Onthophagus (sensu lato) spinifex ( Fabricius, 1781) ]

( Fig. 11 )

Scarabaeus spinifex Fabricius, 1781: 29 ; Fabricius 1787: 15; Jablonsky and Herbst 1789: 240; Olivier 1789: 148 (as S. spinifer , misprint); Gmelin 1790: 1543; Fabricius 1792: 58; Illiger 1800: 263 (as S. spinifer , misprint); Sturm 1802: 92 (as S. spinifer , misprint); Zimsen 1964: 28.

Copris spinifex, Olivier 1790: 170 ; Fabricius 1801: 49; Illiger 1802: 316; Schönherr 1806: 50.

Onthophagus spinifex, Hope 1837: 33 ; Hope 1838: 315 (as spinifer , misprinting); Westwood 1842: 2; Motschulsky 1859: 154; von Harold 1869: 1036; Arrow 1907: 430 (as spinifer , misprint); Boucomont 1914: 222; Boucomont and Gillet 1927: 147; Arrow 1931: 200; Balthasar 1963: 535; Gupta and Mittal 1987: 50; Gupta 1989: 24; Mittal 1999: 35; Biswas and Mulay 2001: 137; Krajcik 2006: 133; Chandra and Gupta 2011: 254; Chandra and Gupta 2012: 105; Chandra et al. 2012: 53; Jadhav and Sharma 2012: 491; Krajcik 2012: 186; Chandra and Gupta 2013a: 4665; Chandra and Gupta 2013b: 345; Karimbumkara and Rajan 2013: 176; Krajcik 2013: 257; Gupta et al. 2014: 233; Mittal and Jain 2015: 401; Ziani 2015: 19 ; Gajendra and Prasad 2016: 711; Kharel et al. 2020: 380; Ghosh et al. 2021: 387; Ghosh et al. 2022a: 11.

Onthophagus ( Indonthophagus ?) spinifex, Kabakov 2006: 154 (footnote).

Onthophagus (Onthophagus) spinifex, Sobhana et al. 2013: 95 ; Kalawate et al. 2021: 17583.

Onthophagus (incertae sedis) spinifex, Löbl et al. 2006: 176 ; Ziani and Bezděk 2016: 203; Gupta et al. 2018: 484; Ghosh et al. 2020: 246; Ghosh et al. 2022b: 120; Gupta et al. 2022: 426.

Scarabaeus aeneus Fabricius 1781: 34 [type locality: “ Coromandel ” (south-eastern India)]; Fabricius 1787: 18; Jablonsky and Herbst 1789: 328; Olivier 1789: 131; Gmelin 1790: 1557; Fabricius 1792: 59; Illiger 1800: 238; Sturm 1802: 80.

Copris aeneus, Olivier 1790: 163 .

Copris aenea, Fabricius 1801: 51 ; Schönherr 1806: 53.

Onthophagus aeneus, von Harold 1869: 1024 ; von Harold 1880: 154 (as junior synonym of O. spinifex );Shipp 1895: 179 (as junior synonym of O. spinifex ); Boucomont and Gillet 1927: 147 (as junior synonym of O. spinifex ); Arrow 1931: 200 (as junior synonym of O. spinifex ); Balthasar 1963: 535 (as junior synonym of O. spinifex ); Krajcik 2006: 133 (as junior synonym of O. spinifex ); Löbl et al. 2006: 176 (as junior synonym of O. spinifex ); Ziani and Bezděk 2016: 203 (as junior synonym of O. spinifex ).

Scarabaeus truncaticornis Herbst 1786: 154 ; [type locality: “Ostindien” (East India); Jablonsky and Herbst 1789: 209 (as junior synonym of S. truncaticornis Schaller, 1783 ); Fabricius 1792: 59 (as junior synonym of S. aeneus Fabricius, 1781 ); Illiger 1800: 239 (as junior synonym of S. aeneus ); Schönherr 1806: 53 (as junior synonym of Copris aenea ).

Onthophagus truncaticornis, von Harold 1869: 1024 (as junior synonym of O. aeneus ); von Harold 1880: 154 (as junior synonym of O. aeneus ); Shipp 1895: 179 (as junior synonym of O. aeneus and junior synonym of O. spinifex ); Boucomont and Gillet 1927: 147 (as junior synonym of O. spinifex ); Balthasar 1963: 535 (as junior synonym of O. spinifex ); Krajcik 2006: 133 (as O. truncaticornis Herbst in Jablonsky 1789, junior synonym of O. spinifex ); Löbl et al. 2006: 176 (as junior synonym of O. spinifex ); Ziani and Bezděk 2016: 203 (as junior synonym of O. spinifex ).

Onthophagus reflexicornis Redtenbacher 1868: 57 [type locality: “ Ceylon ” ( Sri Lanka)]; von Harold 1872: 206 (as junior synonym of O. spinifex ); Boucomont and Gillet 1927: 147 (as junior synonym of O. spinifex ); Arrow 1931: 200 (as junior synonym of O. spinifex ); Balthasar 1963: 535 (as junior synonym of O. spinifex ); Krajcik 2006: 133 (as junior synonym of O. spinifex ); Löbl et al. 2006: 176 (as “ O. reflexicornis L. Redtenbacher 1867 ”, junior synonym of O. spinifex ); Ziani and Bezděk 2016: 203 (as junior synonym of O. spinifex ); Schoolmeesters 2023 (as junior synonym of O. spinifex ).

Onthophagus bifossus d’Orbigny 1902: 145 [type locality: “ Sénégal ” ( Senegal)]; d’Orbigny 1908: 155 (as junior synonym of O. spinifex ); d’Orbigny 1913: 726 (as junior synonym of O. spinifex ); Boucomont and Gillet 1927: 144 (as junior synonym of O. spinifex ); Balthasar 1963: 535 (as junior synonym of O. spinifex ); Krajcik 2006: 133 (as junior synonym of O. spinifex ); Löbl et al. 2006: 176 (as junior synonym of O. spinifex ); Ziani and Bezděk 2016: 203 (as junior synonym of O. spinifex ); Schoolmeesters 2023 (as junior synonym of O. spinifex ).

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. 9b–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. 9d ).

Distribution. India ( Fabricius 1781). Sri Lanka ( Boucomont 1914). Nepal (Löbl et al. 2006). Pakistan ( Ziani 2015). Bangladesh ( Kharel et al. 2020).

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.

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.

Figures

Figure 9. Onthophagus (Indonthophagus) turbatus Walker, 1858 [syntype, medium ♂: Sri Lanka]. 9a) Male dorsal habitus [photo and editing by G. Fiumi]. 9b) Parameres, lateral view. 9c) Parameres, dorsal view. 9d) Endophallus. AcS: accessory endophallites. LC: lamella copulatrix. Rs: raspulae. Photos by A. Degiovanni, edited by G. Fiumi.

Figure 11. Onthophagus (incertae sedis) spinifex (Fabricius, 1781) [major ♂: Pakistan, Capital Territory].11a) Dorsal habitus. 11b) Parameres, lateral view.11c) Parameres, dorsal view.11d) Endophallus. AcS: accessory endophallites. LC: lamella copulatrix. Photos by A. Degiovanni, edited by G. Fiumi.