Coprophanaeus dardanus (MacLeay, 1819)

Fig. 22B

Phanaeus dardanus MacLeay, 1819: 126 (original description). Type locality: Brazil. Name-bearing type: Unknown typification status and whereabouts (Edmonds & Zídek 2010).

Phanaeus bitias Harold, 1863: 163 (original description). Type locality: Somewhere in the Amazon or the Atlantic Forest. Originally said to be Mexico: Oaxaca: Tuxtepec: “hot regions” (“ Mexique (Tustepec), régions chaudes”), but this is mistaken. The species is exclusively South American (Edmonds & Zídek 2010; Cupello & Vaz-de-Mello 2013a). The type locality is here corrected following Recommendation 76A.2 of the Code. Name-bearing type: a male holotype fixed by monotypy and originally from Chevrolat’s personal collection (Harold 1863), but whose current whereabouts is unknown (Edmonds & Zídek 2010). It was not found by Arnaud (1982) in the MNHN, where most of the Harold collection material has been housed since the 1950s (see Cupello 2020), nor by Olaf Jäger in the collection under his care in SMTD (pers. com. to MC, 3 March 2023), where the Chevrolat Scarabaeinae collection has been preserved since the 1910s via the Carl Felsche collection (Horn et al. 1990a; Cupello 2020). Bates (1887) informed that the holotype (cited as “the type-specimen”) was in Sallé’s collection. If correct, Sallé would have somehow acquired it from Chevrolat and the specimen would now most likely be in the BMNH, where most of Sallé’s collection of Mexican and Central American material has resided since the early 20 th century (Cupello 2021). Alas, we did not have the opportunity to go to London to confirm this. The synonymy between Phanaeus bitias and P. dardanus ‒ and, by extension, the incorrectness of the type locality of P. bitias ‒ are, nevertheless, confirmed by the illustrations of the P. bitias holotype provided by Bates (1887: pl. III fig. 19, 19a). In them, we can clearly see the diagnostic bidentate cephalic process of males of the species presently called Coprophanaeus dardanus (see Edmonds & Zídek 2010). This synonymy had been initially entertained by Bates (1887) himself and was later formalized by Nevinson (1892); we are the first to revisit it ever since. The degree of development of both the cephalic and the pronotal processes of the holotype, also judging from Bates’ illustrations, is that of a medium- to small-sized male.

Phanaeus jasion Felsche, 1901: 153 (original description). Type locality: French Guiana. Name-bearing type: holotype (SMTD), not examined.

Phanaeus arrowi d’Olsoufieff, 1924: 68 (original description). Type locality: Brazil: Bahia. Name-bearing type: lectotype (MNHN), designated by Arnaud (1982), not examined.

Phanaeus dardanus – Nevinson 1892: 3 (list). — Arnaud 2002b: 32 (cited).

Phanaeus bitias – Nevinson 1892: 3 (cited as synonym of Phanaeus dardanus). — Arnaud 2002b: 32 (cited as synonym of Coprophanaeus dardanus).

Phanaeus jasion – Gillet 1911b: 83 (catalogue). — Blackwelder 1944: 210 (checklist). — Arnaud 2002b: 32 (cited as synonym of Coprophanaeus dardanus).

Phanaeus (Coprophanaeus) dardanus – d’Olsoufieff 1924: 69 (distribution). — Pessôa 1934: 300 (comments).

Coprophanaeus dardanus – Edmonds 1972: 843 (recombination). — Forsyth & Gill 1993: 70 (list). — Larsen 2011: 99 (list for Suriname). — Feer 2000: 32 (list); 2013: 766 (list for French Guiana). — Feer & Pincebourde 2005: 30 (list, ecology). — Larsen et al. 2008: 1294 (list). — Gámez & Acconcia 2009: 394 (behaviour, table 1). — Brûle et al. 2011a: 193 (list); 2011b: 121 (list); 2014: 183 (list). — Brûle & Dalens 2012: 37 (list). — Brûle & Touroult 2013: 40 (list). — Feer & Boissier 2015: 169 (list). — Boilly et al. 2016: 89, 90 (key, comments catalogue). — Ferrer-Paris et al. 2013: 109 (list). — Silva et al. 2017: 491 (ecology).

Coprophanaeus (Coprophanaeus) dardanus – Edmonds 1972: 843 (recombination). — Martínez & Clavijo 1990: 7 (notes). — Forsyth & Gill 1993: 70 (list). — Larsen 2011: 99 (list for Suriname). — Feer 2000: 32 (list); 2013: 766 (list for French Guiana). — Vaz-de-Mello 2000: 192 (checklist for Brazil). — Arnaud 2002b: 32 (key). — Feer & Pincebourde 2005: 30 (list, ecology). — Gámez et al. 2006: 101 (ecology). — Larsen et al. 2008: 1294 (list). — Gámez & Acconcia 2009: 394 (behaviour, table 1). — Edmonds & Zídek 2010: 76, 80 (revision). — Brûle et al. 2011a: 193 (list); 2011b: 121 (list). — Brûle & Dalens 2012: 37 (list). — Brûle & Touroult 2013: 40 (list). — Cupello & Vaz-de-Mello 2013a: 361, 364–365, fig. 14 (key, distribution, comments). — Brûle et al. 2014: 183 (list). — Feer & Boissier 2015: 169 (list). — Pacheco & Vaz-de-Mello 2015: 3, 7 (checklist, key). — Boilly et al. 2016: 89, 90 (key, comments). — Ferrer-Paris et al. 2013: 109 (list). — Silva et al. 2017: 491 (ecology). — Hielkema & Hielkema 2019: 94 (catalogue for the Guianas).

Phanaeus arrowi – Arnaud 2002b: 32 (cited as synonym of Coprophanaeus dardanus).

Distribution

Colombia, Venezuela, Trinidad and Tobago, Guyana, Suriname, French Guiana, and Brazil.

Subregions of Venezuela

Maracaibo Depression, Peneplain of the Casiquiare River–Upper Orinoco, System of hills and low piedmont mountains of the Guiana Shield, System of low mountains and hills Imataca-Cuyuní of Northeast Guyana Shield, Central Coast Mountain Range, Oriental Coast Range, and Guiana Shield.

Literature records

Martínez & Clavijo 1990: 7 (Venezuela: Amazonas: Atures and Atabapo). — Arnaud 2002b: 32 (Venezuela). — Gámez et al. 2006: 101 (Venezuela: Amazonas). — Larsen et al. 2008: 1294 (Venezuela: Bolívar state). — Gámez & Acconcia 2009: 394 (Venezuela: Zulia: Colón). — Edmonds & Zídek 2010: 76, 80 (Venezuela: Amazonas, Aragua, Bolívar, Miranda and Monagas). — Ferrer-Paris et al. 2013: 109 (Venezuela: Aragua [Guárico]: Altagracia de Orituco and Bolívar: Isla de Anacoco).