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Fritillary phylogeny, classification, and larval host plants: reconstructed mainly on the basis of male and female genitalic morphology (Lepidoptera: Nymphalidae: Argynnini)

Biological journal of the Linnean Society 2006 v.89 no.4 pp. 627-673
Dryas, Nymphalidae, Passiflora, Salix, Vaccinium, Viola, butterflies, eggs, food plants, genitalia, larvae, phylogeny
Species-level phylogeny of the Butterfly tribe Argynnini is established based on 141 characters derived from wing and genitalia morphology of both sexes. The Argynnini can be divided into three subtribes; Yrameina comprising Yramea and Boloria s.l., Argynnina comprising Prokuekenthaliella, Issoria, Brenthis, and all the 'large fritillary' species joined in the genus Argynnis s.l. and a new subtribe Euptoietina comprising only the genus Euptoieta. The classical genus Issoria s.l. is polyphyletic regarding Yramea and possibly paraphyletic regarding the two Afrotropic species baumanni and hanningtoni; these two species are tentatively transferred to the old genus/subgenus Prokuekenthaliella. Surprisingly, one Afrotropic species, Issoria smaragdifera is closely related to the East Palaearctic Issoria species. A revised classification of Argynnini is proposed based on the obtained phylogeny. Studies of larval host plants based on the obtained phylogeny suggest that the ancestral Argynnini used Passiflora and Violaceae, but already the ancestor of Yrameina + Argynnina was probably specialized on Violaceae. Whereas the Boloria species have turned to other food plants such as Dryas, Vaccinium and Salix on several occasions, only Brenthis among the Argynnina use other host plants than Viola (mainly Rosaceae). The habit of laying eggs away from the food plant has probably evolved twice within Argynnina.