Interrelationships and diversification of Argynnis Fabricius and Speyeria Scudder butterflies

Abstract

Diverse radiations of insects are often associated with adaptations to host plants, and well-resolved phylogenetic relationships are required to fully understand them. Palearctic Argynnis and related subgenera, together with North American Speyeria butterflies make up a radiation whose species hypotheses are confounded by shared wing colour patterns between sympatric populations of closely related recognized species. Previous studies of this group indicate that Speyeria is a lineage within Argynnis, but sampling in these studies has either involved too few Speyeria species or incomplete sampling of Argynnis species. Thus, no comprehensive phylogenetic analysis exists for all members that answers the question of monophyly of Speyeria, or other subgeneric taxa, and their relationship to Argynnis species. We completed a phylogenetic analysis of all North American Speyeria species and all but one species within Argynnis, using one mitochondrial (cytochrome c oxidase I, COI) and four nuclear genes [elongation factor 1 alpha (EF1α), wingless (WG), glyceraldehyde-3-phosphate dehydrogenase (GAPDH), and ribosomal protein S5 (RPS5)]. The results indicate three major lineages within Argynnis s.l.: two Palearctic and one containing both Palearctic and Nearctic species. In summary, the phylogenetic analyses suggest the need for reorganization into three natural groups: Argynnis, Fabriciana and Speyeria. Within each of these genera the phylogenetic hypothesis indicates an evolutionary history marked by rapid diversification and potential extinction, followed by ongoing lineage sorting. The position of North American Speyeria is nested within the Palearctic lineages, which indicates that the radiation began in Asia and was fuelled by existing Viola diversity in North America. Dating analyses of Viola and Speyeria corroborate this hypothesis. The current North American Speyeria species are mixed on the tree, indicating a recent and ongoing radiation. These results provide needed clarity on the evolution of this group, which contains species of conservation concern