Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea) Author Jong, Rienk De text Zootaxa 2017 4270 1 1 63 journal article 32975 10.5281/zenodo.583183 6c479acc-8b18-4f0b-a6e5-85bcd6d7b6b7 1175-5326 583183 2D00AFF5-4FE2-4EC1-A328-C8670CFB8D6D pluto . Vanessa pluto Heer, 1849 Figs 11–13 . Pieridae : Coliadinae + Pierinae . Croatia, Radoboj, Croatia; Burdigalian, early Miocene. FIGURES 11–13. Ƒanessa pluto Heer, 1849 . Figures copied from (1875), Plate II, figures 1, 17 and 7, respectively. (A), copied by Scudder from original description by Heer; (B) and (C), figures sent to Scudder by Mr. Brunner de Wattenwyl, made after the same object. Note differences in wing shape and other details. Figs 14–16. Thaites ruminiana Scudder, 1875 . Figures copied from Scudder (1875), Plate III, figures 9, 1 and 3, respectively. (A), underside of fossil; (B), reconstruction of venation; (C), reconstruction of markings. In (A) two separate subcostal veins have been drawn in the forewing, an impossible configuration in Lepidoptera , and apparently a mistake, corrected in (B) and (C). Fig. 17. Cyllo sepulta Boisduval, 1841 , wing venation after Scudder (1875). Fig. 18. Butterfly fossil recorded by Wangrin (1939), but shown to be a fraud by Ansorge (2015). Photo kindly provided by J. Ansorge, Horst, Germany (see text for explanation). Depository: NHMW (holotype, no. 1940/0001/0011). Published figure: Murata (1998: Figs 27, 28) ; Pongrácz (1928: Fig. 30) ; Scudder (1875: Pl. II Figs 2 , 7, 17). This specimen consists of the greater part of both forewings and basal fragments of hindwings. Original description and figure copied in Scudder (1875: 44–50). Considered to belong to Nymphalidae by Heer (and followed by some other authors, see discussion by Scudder) on the basis of a peculiar and highly unlikely venational arrangement: Sc arising from R halfway between base and apex, together with the common stalk of R1 and R2 (which forks shortly thereafter) and the common stalk of R3, R4 and R5, while M1 originates from the radial stem before this trichotomy; the cell is open. Such an arrangement is not found in any extant species. Scudder’s interpretation is very different, with Sc from base as usual; the radial branching is not clearly visible, partly because the apex is broken, but it seems that M1 branches off an R-branch (formula R1, R2, (R3+M1)). This arrangement, with M1 branching off an R-branch, is found in a number of genera belonging to Coliadinae and Pierinae (see phylogeny by Wahlberg et al . 2014 ), and Scudder apparently was correct in considering the fossil attributable to the Pieridae . He placed it in his genus Mylothrites (followed by Kozlov 1988 ) and compared it to the recent genera Mylothris (Africa) and Hebomoia (Southeastern Asia) ( Pieridae : Pierinae ), both which have only three radial veins. Three radial veins are also found in the American genus Nathalis ( Pieridae : Coliadinae ), so we cannot identify the fossil more precisely than belonging to Coliadinae + Pierinae . Pongrácz (1928) re-examined the type and concluded that it was close to the satyrine genus Satyrus (Nymphalidae) . However, in view of the state of preservation of the fossil of which a photograph of the holotype has been published by Ponomarenko & Schultz 1988 : Pl. 7 Fig. 2 , Pongrácz’s reconstruction (his Fig. 30) must have been erroneous. Without providing reasons, Handlirsch (1908) mentioned the fossil under “ Papilionidae ”, Handlirsch (1925) under “ Papilioninae ”, and Zeuner (1942) under " Nymphalinae ".