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 leuce . Apanthesis leuce Scudder, 1889 Incertae sedis. USA , Colorado , Teller County , Florissant; late Priabonian, late Eocene. Depository: MCZH (holotype, 16354). Published figures: Emmel et al . (1992 : Fig. 1 /10); Scudder (1889: Pl. LII Figs. 12, 13 ) . One forewing only, with venation and design well preserved. Radial formula 1, 2, 3+(4+5), udc absent, M1, R3+(4+5) and mdc almost originating from the same point; ldc weak. According to Scudder, the radial branching is typical for the “tribe Vanessidi” ( Nymphalidae ), but I do not know what is so typical about it, except for the weak ldc. Comstock (1961: 181) observed a closer similarity to Limenitis , with which also the markings correspond with a uniformously brown hue with a submarginal band parallel to termen and consisting of light-coloured, slightly kidney-shaped spots, and vague spots between the band and termen in intervenal areas). Because Limenitis and Anaea share characters in the male genitalia, Comstock (1961) considered the discovery of Apanthesis leuce an indication of a northern origin of Anaea . This is, however, an unwarranted speculation. The venational arrangement could as well be plesiomorphic and is, for instance, also similar to the arrangement in the pierid genus Eronia . Euronia was considered a primitive member of the pierid tribe Pierini by Klots (1931) based on morphological grounds. Braby et al . (2006) concluded from a molecular analysis of the Pieridae , that Euronia was grouped in the Colotis group of genera, the first offshoot of the Pierinae . The weakness of ldc may be an argument in favor of Nymphalidae , but it remains to be seen whether in a fossil of some 35 Ma of age the weak ldc in, e.g., Limenitis , can be distinguished from the rather weak ldc in Eronia . The only reason for considering the fossil a member of the Nymphalidae is, therefore, the similarity in wing markings to some recent species, a rather weak argument. Emmel et al . (1992) followed Scudder's decision; they did not mention Comstock’s remarks. I think it is wise to consider the fossil as without clear affinities, as long as elements of the wing design remain unstudied in a phylogenetic context.