Basal subtribes of the Nymphidiini (Lepidoptera: Riodinidae): phylogeny and myrmecophily

Abstract

Three cladistic analyses, based predominantly on adult morphology, are presented for myrmecophilous riodinid butterflies in the tribe Nymphidiini. The first is a species-level analysis of all 22 species recognized here in Aricoris (= Audre auctt.) using 27 characters. The second is a species-level analysis of all 24 species recognized here in Synargis using 53 characters. The third is a generic-level analysis of all six genera ( Aricoris, Ariconias n. gen., Lemonias, Thisbe, Juditha, and Synargis) recognized here as belonging in the basal clades of the Nymphidiini (=Lemoniadina auctt.) using 17 characters, with members of the Theopeina and Nymphidiina included with the ingroup to assess the monophyly of subtribes. Almost all characters are illustrated. The first analysis indicates that Aricoris consists of five monophyletic species groups, with the relationship (( constantius gr. + colchis gr.) + ( chilensis gr. + ( aurinia gr. + epulus gr.))), and contains the type species of Eiseleia and Audre, which are synonymized with Aricoris ( n. syns.). The second analysis indicates that Synargis consists of four monophyletic species groups, with the relationship ( phliasus gr. + ( regulus gr. + ( pittheus gr. + abaris gr.))), and contains the type species of Ematurgina and Thysanota, which are synonymized with Synargis ( n. syns.). The third analysis indicates that the Lemoniadina, or basal clades of the Nymphidiini, are paraphyletic with respect to the Theopeina and Nymphidiina, with the generic relationship (( Aricoris + Ariconias) + ((( Lemonias + Thisbe) + ( Juditha + Synargis)) + (Theopeina + Nymphidiina))). We therefore restrict the Lemoniadina to include Lemonias, Thisbe, Juditha, and Synargis and provide the name Aricorina n. subtribe for Aricoris and Ariconias. All nymphidiine subtribes are characterized and a synonymic checklist for the Aricorina and Lemoniadina is presented. A “supertree” composed of the species-level phylogenies derived here for Aricoris and Synargis, and those derived elsewhere for Ariconias, Lemonias, Thisbe, and Juditha, is used to map the distribution of larval host plant and attending ant taxa and the occurrence of aphytophagy. Observed patterns are discussed.