Systematics of Hypanartia (Lepidoptera: Nymphalidae: Nymphalinae), with a test for geographical speciation mechanisms in the Andes

A taxonomic review of the Neotropical nymphaline butterfly genus Hypanartia Hübner is presented, including notes on the taxonomy, biology and distribution of its component species, illustrations of all taxa and the male genitalia of all species, and the description of four new species and two new subspecies: Hypanartia celestia sp.n., H. cinderella sp.n., H. dione disjuncta ssp.n., H. fassli sp.n., H. trimaculata sp.n. and H. trimaculata autumna ssp.n. Hypanartia arcaei (Salvin) is placed as a subspecies of H. dione (Latreille) (stat.n.) and lectotypes are designated for eight nominal taxa. Fourteen species are recognized, with the centre of diversity being in high Andean cloud forest habitats. A cladistic analysis was conducted, based on fifty-three illustrated characters of male genitalic and abdominal morphology, and external facies, to investigate phylogenetic relationships. The resulting phylogenetic hypothesis was used to test four different geographical mechanisms of speciation in the Andes: colonization from temperate latitudes, speciation across elevational gradients, radiation within the Andes and allopatric speciation between the Andes and other montane regions. There is evidence that speciation across an elevational gradient occurred twice, both times into elevations largely unoccupied by the genus, and in both cases followed by subsequent, elevationally sympatric, in situ radiation. Differentiation in allopatry between montane regions has apparently been of recent influence only, causing infraspecific variation in two species. These results parallel several recent studies of Andean bird speciation.