The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem Author Cupello, Mario Author Ribeiro-Costa, Cibele S. Author Vaz-De-Mello, Fernando Z. text Zoological Journal of the Linnean Society 2022 2021-07-12 194 973 1034 journal article 10.1093/zoolinnean/zlab015 cc82accd-7fe1-4f9f-a60d-c7c121456114 0024-4082 42E7A370-FCD8-492A-8604-C42863BDE6E2 BOLBITES HAROLD, 1868 Bolbites Harold, 1868a: 81 ; Harold, 1869a: 1016 ; Harold, 1869b: 62 ; Burmeister, 1874: 129 ; Jaeger, 1874: 225 ; Van Lansberge, 1875: 12 ; Waterhouse, 1891: 59 ; Fabre, 1899: 74 , 95; Kolbe, 1905: 531 ; Lagôa, 1905: 28 ; Kolbe, 1907: 31 ; Enderlein, 1908: 335 ; Bruch, 1911: 189 ; Gillet, 1911: 80 ; Bruch, 1915: 540 ; Fabre, 1919: 242 ; Lucas, 1920: 141 ; d’Olsoufieff, 1924: 9 , 12, 17, 19, 57–58, 133, 138; Balthasar, 1941: 350 ; Pessôa & Lane, 1941: 470 ; Blackwelder, 1944: 208 ; Balthasar, 1951: 335 ; Halffter, 1952: 83 ; Martínez, 1959: 94 ; Halffter & Matthews, 1966: 17 , 30, 123, 131, 196, 210, 257; Edmonds, 1972: 816 , 818, 854; Halffter 1974: 253 ; Halffter & Edmonds, 1982: 137 ; Zunino, 1985a: 22–23 , fig. 1; Zunino, 1985b: 107 , 109, fig. 4; Martínez, 1987: 50 ; Hanski & Cambefort, 1991: 466 ; Cabrera Walsh & Gandolfo, 1996 : fig. 7; Monteresino et al. , 1996: 109 ; Philips et al. , 2004: 43, 45, 50–54, 58, figs 6–8; Hamel-Leigue et al. , 2006: 3; Krajcik, 2006: 16 ; Hamel-Leigue et al. , 2009: 56 ; Price, 2009: 148 ; Scholtz, 2009b: 243 , 246–248, fig. 12.4; Scholtz et al. , 2009: 537 ; Vaz-de-Mello & Grossi, 2010: 220; Vaz-de-Mello et al. , 2011: 4 , 10, 17, 25, 32, 40, 44; Krajcik, 2012: 54 ; Cupello & Vaz-de-Mello, 2013: 439 , 441–443, 460; Cupello & Vazde-Mello, 2015: 16, 18, 21–23; Tissiani et al. , 2017: 411 , 413; Cupello et al. , 2020: 2 ; Gillett & Toussaint, 2020: 662 , 666, 671–672, 676, fig. 3. Bolites [incorrect subsequent spelling]: Fabre, 1919: 268 . Type species: Bolbites onitoides Harold, 1868 , by original monotypy. Etymology: According to Harold (1869a) , the genusgroup name Bolbites is derived from a Greek word supposedly spelled as βῐλβιτον and which he translated into Latin as ‘ stercus ’ (‘dung’). We did not find any word spelled like that in Greek dictionaries, but similar forms such as βόβλιτον ( bóbliton , neuter in gender), βόλβιθος ( bólbithos , masculine), βόλβιτος ( bólbitos , masculine), and βόλιτον ( bóliton , neuter) are listed and indeed translated into English as ‘(of) cow dung’ or ‘cowpat’ (e.g. Liddell & Scott, 1897 ; Wiktionary, 2020 ). Therefore, Harold’s translation seems to be correct, even though his exact transliteration of the word into the Latin script (i.e. the spelling Bolbites ) does not correspond to the way that any of the variants we found would be transliterated (see, for example, transliteration guidelines by: Grensted, 1958 ; Papavero, 1994 ; Vlachos, 2015 ). It seems that instead of transliterating one of those words directly, Harold used the stem of the word βόλβιτος (namely, βόλβι-, ‘ bolbi -’) in combination with the Greek masculine suffix -ῑ́της (- ites ), expressing the idea of relationship ( Brown, 1956 ; Papavero, 1994 ), to render the noun βόλβῑ́της, i.e. Bolbites . An English translation of the genus name is thus ‘the one from the cowpat’. Based on the gender agreement that Harold (1869a) proposed for the adjectival specific name of Gromphas dichroa when he transferred the species to Bolbites (viz., as dichrous ), it is clear that he regarded the grammatical gender of Bolbites as masculine. In reason of the gender of the suffix -ῑ́τηςbeing masculine, a modern interpretation of the gender of Bolbites under Article 30.1.2 of the Code agrees with Harold’s and the name must indeed be treated as masculine. It should be noted that a feminine version of the noun, βόλβῑ́τις(‘ bolbitis ’), is listed in Liddell & Scott (1897) and Brown (1956) as referring to ‘a small kind of cuttle-fish with a strong smell’ ( Liddell & Scott, 1897 ). This ‘strong smell’ was likely the reason for the reference to cow dung in the name of this cephalopod. In any case, there is no evidence that Harold (1868b) derived the name of his new genus from this word; if he had, he would have most probably mentioned it in the etymology given in 1869. Species composition: Authors have been almost unanimous in regarding B. onitoides as the sole extant species of Bolbites . This, as argued below, is indeed correct. The only two exceptions include Harold (1869a) , who, in his world Scarabaeidae catalogue, listed Gromphas dichroa Blanchard, 1846 as a second species of Bolbites with no explicit justification. Soon thereafter, Harold (1869b) examined Blanchard’s type material and returned the species to Gromphas , where it indeed belongs ( Cupello & Vaz-de-Mello, 2013 , 2016). Additionally, Van Lansberge (1875) , in his monograph of the onitids, mentioned that one of the two species described by Perty (1830) in Onitis – O. aeruginosus Perty, 1830 or O. chalcomelas Perty, 1830 – should, in fact, belong to Bolbites , but without stating which one, or giving any further explanation. Nevertheless, he was most probably referring to O. aeruginosus , which had already been correctly and independently transferred to Gromphas by Sturm (1843) and Harold (1859) ( Cupello & Vaz-de-Mello 2013 , 2014a , 2015 ); O. chalcomelas , in turn, is a Phanaeus MacLeay, 1819 ( Edmonds, 1994 , and elsewhere).