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

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. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 89, 627–673.

[1]  T. Simonsen Glands, muscles and genitalia. Morphological and phylogenetic implications of histological characters in the male genitalia of Fritillary butterflies (Lepidoptera: Nymphalidae: Argynnini) , 2006 .

[2]  T. Simonsen The male genitalia segments in fritillary butterflies: Comparative morphology with special reference to the "rectal plate" in Issoria (Lepidoptera: Nymphalidae) , 2006 .

[3]  T. Simonsen Boloria phylogeny (Lepidoptera: Nymphalidae): tentatively reconstructed on the basis of male and female genitalic morphology , 2005 .

[4]  K. Brown,et al.  Phylogeny of the Nymphalidae (Lepidoptera). , 2004, Systematic biology.

[5]  G. Lamas,et al.  Yramea lynx sp. nov. (Lepidoptera: Nymphalidae, Heliconiini) from the Andes of central Perú , 2004 .

[6]  C. Penz,et al.  Phylogenetic relationships among Heliconiinae genera based on morphology (Lepidoptera : Nymphalidae) , 2003 .

[7]  G. Robinson,et al.  Hostplants of the moth and butterfly caterpillars of the Oriental region , 2001 .

[8]  A. Brower Phylogenetic relationships among the Nymphalidae (Lepidoptera) inferred from partial sequences of the wingless gene , 2000, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[9]  C. Penz Higher level phylogeny for the passion‐vine butterflies (Nymphalidae, Heliconiinae) based on early stage and adult morphology , 1999 .

[10]  T. Larsen The Butterflies of Kenya : And Their Natural History , 1996 .

[11]  R. Vane-Wright,et al.  Carcasson's African Butterflies , 1995 .

[12]  K. Bremer,et al.  BRANCH SUPPORT AND TREE STABILITY , 1994 .

[13]  C. Häuser Die inneren weiblichen Genitalorgane der Tagfalter (Rhopalocera): Vergleichende Morphologie und phylogenetische Interpretation (Insecta, Lepidoptera) , 1993 .

[14]  K. Bremer THE LIMITS OF AMINO ACID SEQUENCE DATA IN ANGIOSPERM PHYLOGENETIC RECONSTRUCTION , 1988, Evolution; international journal of organic evolution.

[15]  P. Ackery Hostplants and classification: a review of nymphalid butterflies , 1988 .

[16]  P. Opler,et al.  The Butterflies of Costa Rica and Their Natural History: Papilionidae, Pieridae, Nymphalidae. , 1989 .

[17]  R. Vane‐Wright,et al.  Milkweed Butterflies: Their Cladistics and Biology , 1984 .

[18]  N. P. Kristensen The male genitalia of Agathiphaga (Lepidoptera: Agathiphagidae) and the lepidopteran ground plan , 1984 .

[19]  E. Nielsen,et al.  South American micropterigid moths: two new genera of the Sabatinca-group (Lepidoptera: Micropterigidae) , 1982 .

[20]  R. A. Arnold,et al.  Operational Mechanisms of Copulation and Oviposition in Speyeria (Lepidoptera: Nymphalidae) , 1977 .

[21]  L. G. Higgins,et al.  The classification of European butterflies , 1975 .

[22]  T. Shirozu A generic classification of the genus Argynnis and its allied genera(Lepido-ptera:Nymphalidae) , 1973 .

[23]  T. Shirozu The systematic position of Argynnis argyrospilata Kotzsch(Lepidoptera Nymphalidae) , 1973 .

[24]  Robert E. Miller Welcoming Address for Rocky Mountain Section, Geological Society of America , 1968 .

[25]  L. D. Miller The higher classificationc phylogeny and zoogeography of the Satyridae lLepidopterar , 1968 .

[26]  F. Hemming The generic names of the butterflies and their type-species lLepidopterac Rhopalocerar , 1967 .

[27]  T. Reuss Systematischer Überblick der Dryadinae T. Rss. mit einigen Neubeschreibungen (Lep. Rhopal) , 2022 .

[28]  W. Howe The Butterflies of North America , 1874, The American Naturalist.