Higher - level phylogeny of the Ithomiinae ( Lepidoptera : Nymphali - 4 dae ) : classification , patterns of larval hostplant colonization and 5 diversification 6

13 We present a higher-level phylogenetic hypothesis for the diverse neotropical butterfly subfamily Ithomiinae, inferred from one of 14 the largest non-molecular Lepidoptera data sets to date, including 106 species (105 ingroup) and 353 characters (306 informative) 15 from adult and immature stage morphology and ecology. Initial analyses resulted in 1716 most parsimonious trees, which were 16 reduced to a single tree after successive approximations character weighting. The inferred phylogeny was broadly consistent with 17 other past and current work. Although some deeper relationships are uncertain, tribal-level clades were generally strongly 18 supported, with two changes required to existing classification. The tribe Melinaeini is polyphyletic and Athesis + Patricia require a 19 new tribe. Methona should be removed from Mechanitini into the restored tribe Methonini. Dircennini was paraphyletic in analyses 20 of all data but monophyletic based on adult morphology alone, and its status remains to be confirmed. Hypothyris, Episcada, 21 Godyris, Hypoleria and Greta are paraphyletic. A simulation analysis showed that relatively basal branches tended to have higher 22 partitioned Bremer support for immature stage characters. Larval hostplant records were optimized on to a reduced, generic-level 23 phylogeny and indicate that ithomiines moved from Apocynaceae to Solanaceae twice, or that Tithoreini re-colonized Apocynaceae 24 after a basal shift to Solanaceae. Ithomiine clades have specialized on particular plant clades suggesting repeated colonization of 25 novel hostplant niches consistent with adaptive radiation. The shift to Solanum, comprising 70% of neotropical Solanaceae, occurs 26 at the base of a clade containing 89% of all ithomiines, and is interpreted as the major event in the evolution of ithomiine larval 27 hostplant relationships. 28 The Willi Hennig Society 2006. 29