Taxonomy, phylogeny and distribution of the genus Steromapedaliodes sensu novo in the Cordillera de Mérida, Venezuela (Lepidoptera: Nymphalidae: Satyrinae: Satyrini)

We revise the taxonomy and phylogeny of the genera Steromapedaliodes Forster, 1964 and Redonda Adams & Bernard, 1981, both subparamo and paramo endemics of the Cordillera de Mérida (Venezuela). Redonda is hereby synonymized with Steromapedaliodes, a decision supported by a comparative study evidencing several synapomorphies of both genera in external characters and genitalia, and the lack of salient synapomorphies specific to each genus in morphological, biogeographical and ecological traits. Also, Steromapedaliodes mavarezi sp.n. presents several intermediate characters in relation to the species traditionally assigned to Redonda and Steromapedaliodes. Furthermore, a molecular phylogenetic analysis with the use of one nuclear and three mitochondrial markers shows no support for reciprocal monophyly between Redonda and Steromapedaliodes but provides strong support for the monophyly of Steromapedaliodes sensu novo, with low levels of genetic divergence among species. Two new species of Steromapedaliodes are described, S. kahlua sp.n. and S. mavarezi sp.n. We describe five new subspecies of S. albonotata (Godman, 1905), two of S. albarregas (Adams & Bernard, 1981), and the females of S. empetrus empetrus (Thieme, 1905) comb.n. and S. lathraia Viloria & Camacho, 2015 comb.n. Several new status and synonymies are also established. Accordingly, Steromapedaliodes contains nine species, including four that are highly polytypic. Adult Seromapedaliodes share several striking morphological and genital similarities with another north Andean paramo genus, Dangond Adams & Bernard, 1979 from the Sierra de Perijá. However, our phylogenetic analysis shows that the two are not sister taxa and that they originated in separate clades of the speciose Pedaliodes complex. This shows that convergence in butterflies affects not only external morphology traits directly correlated with habitat selection but also genitalia. Steromapedaliodes has undergone an adaptive radiation in geographically isolated paramos resulting in unusual morphological and behavioural characters such as brachyptery and flightlessness of the females of S. bordoni Viloria & Pyrcz, 2003 comb.n. and S. empetrus (Thieme, 1905) comb.n., a unique adaptation among the world's butterflies