Alpha-taxonomy and phylogeny of African Junoniini butterflies based on morphological data, with an emphasis on genitalia, and COI barcode (Lepidoptera: Nymphalidae)

The tribe Junoniini is a predominantly Paleotropical group of the cosmopolitan butterfly subfamily Nymphalinae (Nymphalidae), with highest diversity in the Afrotropical region. Its systematics and relationships are not entirely resolved. Question marks remain concerning the validity of some genera; and the apparently close relationship between the IndoAustralian genus Yoma and the Afrotropical Protogoniomorpha, as evidenced by molecular phylogenies, remains a puzzle. Here, we present a cladistic analysis, based on 42 characters of the male and female genitalia of 41 species of Junoniini belonging to six genera, nearly all of them continental Afrotropical, and 3 species of two Indo-Australian genera Yoma and Rhinopalpa. A ML COI-based tree is produced for 36 species of Afrotropical Junoniini and Yoma. The molecular data are consistent with previous studies. However, morphological analysis does not confirm a close relationship between Protogoniomorpha and Yoma. Despite the evolution of a number of modifications, the male genitalia within all genera and species of the Junoniini share a cohesive build plan, in particular a transformed sacculus, from which Yoma is highly divergent. The position of the genus Kamilla, previously synonymized with Junonia, is discussed. Three East African coast taxa, Junonia elgiva stat. reinst., Protogoniomorpha nebulosa stat. reinst. and Salamis amaniensis stat. reinst., and one from central Africa, Precis silvicola stat. reinst. are raised to species level, based on comparative analysis of their male genitalia.