Previously unrecognized diversity of Afrotropical Melanitini butterflies (Nymphalidae, Satyrinae): doubling the number of species and genera

The Paleotropical tribe Melanitini (Nymphalidae, Satyrinae) has until now been considered to be represented in the Afrotropical region by two genera - Melanitis Fabricius and Gnophodes Westwood - and six species. Here, we demonstrate that the former genus Gnophodes comprises two lineages well-defined by morphological and genetic traits. According to the new arrangement Gnophodes sensu nov. comprises five species. Gnophodes betsimena (Boisduval) is split into three allopatric species including - G. betsimena, stat.rev. restricted to Madagascar, G. parmeno, stat.reinst. found in the main African rain forest block, and G. diversa, stat.reinst. occurring in East and Southern Africa. Two species - G. grogani Sharpe and G. heroni Pyrcz & Collins, sp.n. are montane specialists. The new genus Haydonia Pyrcz & Collins, gen.n. comprises four species previously placed in Gnophodes, including H. chelys (Fabricius), comb.n., two taxa, H. pythia (Fabricius), comb.n., stat.reinst. and H. harpa (Karsch), comb.n., stat.reinst., previously considered as synonyms of H. chelys, reinstated as valid species, and one new species - H. hassoni Pyrcz & Collins, sp.n. - from Katanga. At the same time, four other taxa are confirmed as synonyms of H. pythia, H. chelys or H. harpa. The two genera differ most immediately by the shape and position of male alar organs, elsewhere designated as androconia. They also differ notably in many aspects of female genitalia, however male genitalia vary little between the species and genera of Melanitini. A parapatric pattern of distribution is uncovered, with related species replacing each other along an elevational gradient. Montane species, H. chelys and G. heroni sp.n., present disjunct distributions being found in the mountains of western Uganda and eastern D.R. Congo, and again in the highlands of Cameroon. In addition, one species is removed from the genus Melanitis and placed in a new monobasic genus Ducarmeia Pyrcz gen.n. Ducarmeia ansorgei (Rothschild), comb.n. based on a salient synapomorphy of adults, a reflector patch situated on hindwing upperside anal fold. Further evidence is found in venation, female genitalia and molecular data. COI data were obtained for nine species, confirming the above findings, with the exception of the unresolved G. parmeno clade. Early stages are partially described for four species.