A molecular phylogeny for the oldest (nonditrysian) lineages of extant L epidoptera, with implications for classification, comparative morphology and life‐history evolution
Abstract Within the insect order L epidoptera (moths and butterflies), the so‐called nonditrysian superfamilies are mostly species‐poor but highly divergent, offering numerous synapomorphies and strong morphological evidence for deep divergences. Uncertainties remain, however, and tests of the widely accepted morphological framework using other evidence are desirable. The goal of this paper is to test previous hypotheses of nonditrysian phylogeny against a data set consisting of 61 nonditrysian species plus 20 representative D itrysia and eight outgroups ( T richoptera), nearly all sequenced for 19 nuclear genes (up to 14 700 bp total). We compare our results in detail with those from previous studies of nonditrysians, and review the morphological evidence for and against each grouping The major conclusions are as follows. (i) There is very strong support for L epidoptera minus M icropterigidae and A gathiphagidae, here termed A ngiospermivora, but no definitive resolution of the position of A gathiphagidae, although support is strongest for alliance with M icropterigidae, consistent with another recent molecular study. (ii) There is very strong support for G lossata, which excludes H eterobathmiidae, but weak support for relationships among major homoneurous clades. Eriocraniidae diverge first, corroborating the morphological clade C oelolepida, but the morphological clades M yoglossata and N eolepidoptera are never monophyletic in the molecular trees; both are contradicted by strong support for L ophocoronoidea + H epialoidea, the latter here including M nesarchaeoidea syn.n. (iii) The surprising grouping of A canthopteroctetidae + N eopseustidae, although weakly supported here, is consistent with another recent molecular study. (iv) Heteroneura is very strongly supported, as is a basal split of this clade into N epticuloidea + E ulepidoptera. Relationships within N epticuloidea accord closely with recent studies based on fewer genes but many more taxa. (v) Eulepidoptera are split into a very strongly supported clade consisting of T ischeriidae + P alaephatidae + D itrysia, here termed E uheteroneura, and a moderately supported clade uniting A ndesianidae with A deloidea. (vi) Relationships within A deloidea are strongly resolved and T ridentaformidae fam.n. is described for the heretofore problematic genus Tridentaforma D avis, which is strongly supported in an isolated position within the clade. (vii) Within E uheteroneura, the molecular evidence is conflicting with respect to the sister group to D itrysia, but strongly supports paraphyly of P alaephatidae. We decline to change the classification, however, because of strong morphological evidence supporting palaephatid monophyly. (viii) We review the life histories and larval feeding habits of all nonditrysian families and assess the implications of our results for hypotheses about early lepidopteran phytophagy. The first host record for N eopseustidae, which needs confirmation, suggests that larvae of this family may be parasitoids. This published work has been registered in ZooBank: http://zoobank.org/urn:lsid:zoobank.org:pub:C17BB79B‐EF8F‐4925‐AFA0‐2FEF8AC32876 .