TL;DR: The goal of this paper is to test previous hypotheses of nonditrysian phylogeny against a data set consisting of 61 nond itrysian species plus 20 representative Ditrysia and eight outgroups (Trichoptera), nearly all sequenced for 19 nuclear genes (up to 14 700 bp total).
Abstract: Within the insect order Lepidoptera (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 Ditrysia and eight outgroups (Trichoptera), nearly all sequenced for 19 nuclear genes (up to 14700bp 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 Lepidoptera minus Micropterigidae and Agathiphagidae, here termed Angiospermivora, but no definitive resolution of the position of Agathiphagidae, although support is strongest for alliance with Micropterigidae, consistent with another recent molecular study. (ii) There is very strong support for Glossata, which excludes Heterobathmiidae, but weak support for relationships among major homoneurous clades. Eriocraniidae diverge first, corroborating the morphological clade Coelolepida, but the morphological clades Myoglossata and Neolepidoptera are never monophyletic in the molecular trees; both are contradicted by strong support for Lophocoronoidea+Hepialoidea, the latter here including Mnesarchaeoidea syn.n. (iii) The surprising grouping of Acanthopteroctetidae+Neopseustidae, 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 Nepticuloidea+Eulepidoptera. Relationships within Nepticuloidea 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 Tischeriidae+Palaephatidae+Ditrysia, here termed Euheteroneura, and a moderately supported clade uniting Andesianidae with Adeloidea. (vi) Relationships within Adeloidea are strongly resolved and Tridentaformidae fam.n. is described for the heretofore problematic genus Tridentaforma Davis, which is strongly supported in an isolated position within the clade. (vii) Within Euheteroneura, the molecular evidence is conflicting with respect to the sister group to Ditrysia, but strongly supports paraphyly of Palaephatidae. 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 Neopseustidae, which needs confirmation, suggests that larvae of this family may be parasitoids. (Less)
TL;DR: The first discovery since the 1970s of a new extant family of homoneurous moths, based on the small Aenigmatinea glatzella sp.n.
Abstract: We report the first discovery since the 1970s of a new extant family (Aenigmatineidae fam.n.) of homoneurous moths, based on the small Aenigmatinea glatzella sp.n. from Kangaroo Island off southern Australia. It exhibits a combination of extraordinary anatomical characters, and, unlike most homoneurous moths, its larva is a conifer-feeder (stem mining in Callitris, Cupressaceae). While the adult's mouthparts are strongly regressed, evidence from other morphological characters and from a Bayesian analysis of 25 genetic loci convincingly places the taxon among Glossata ('tongue moths'). An unexpected tongue moth clade including Acanthopteroctetidae and Neopseustidae, suggested with low support in recent molecular analyses, remarkably becomes strongly supported when Aenigmatinea is included in the molecular analysis; the new taxon becomes subordinated in that clade (as sister group to Neopseustidae) and the clade itself appears as the sister group of all Heteroneura, representing the vast majority of all Lepidoptera. Including Aenigmatinea into the analysis thereby strengthens the surprising indication of non-monophyly of Myoglossata, and the new phylogeny requires an additional number of ad hoc assumptions of convergence/character reversals in early Lepidoptera evolution. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:44393B52-1889-431A-AB08-6BBCF8F946B8. (Less)
TL;DR: A comparative investigation of the galeal anatomy in representatives of all major taxa revealed that the intrinsic g Galeal muscles are derived from the basal g Galeae musculature, a groundplan autapomorphy of the Ditrysia.
Abstract: The suctorial proboscis of adult Lepidoptera represents a key morphological innovation that enabled these insects to gain access to new food sources. In the ancestral condition of the lepidopteran proboscis only extrinsic galeal muscles are present in the basal joint region. The presence of additional muscles (i.e., the intrinsic galeal muscles) is regarded as a morphological novelty of the Myoglossata that evolved after the galeae were modified to form suctorial mouthparts. The present comparative investigation of the galeal anatomy in representatives of all major taxa revealed that the intrinsic galeal muscles are derived from the basal galeal musculature. In the examined Neopseustoidea, Exoporia, Nepticuloidea, Incurvarioidea, and Tischerioidea all galeal muscles have their origin in the stipes-galea joint and/or in the proximal region of the galea. Two muscle units form the basal galeal musculature of the joint region and one to three longitudinal muscles extend into the galea lumen. Multiple intrinsic galeal muscles, of which both the origin and attachment sites are markedly distal from the basal joint region are regarded as a groundplan autapomorphy of the Ditrysia. Some slightly oblique muscles may occur along the lateral wall; these were lost in species with extremely slender galeae. In most investigated Obtectomera two series of intrinsic galeal muscles occur; these are the (1) oblique lateral intrinsic galeal muscles, which are arranged one upon the other along the lateral proboscis wall and (2) the median intrinsic galeal muscles, which run more or less longitudinally along the ventral wall. Oblique muscle arrangement probably evolved in concert with the functional demands of a long lepidopteran proboscis. A likely evolutionary pathway to account for the serial arrangement of galeal muscles is propo sed.
TL;DR: In this paper, the authors compare the morphological evidence for and against each grouping and conclude that there is very strong support for Lepidoptera minus Micropterigi-dae and Agathiphagidae, here termed Angiospermivora, but no definitive resolution of the position of Agathaphagidae.
Abstract: Within the insect order Lepidoptera (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 frame- work 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 nondit- rysian species plus 20 representative Ditrysia and eight outgroups (Trichoptera), 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 Lepidoptera minus Micropterigi- dae and Agathiphagidae, here termed Angiospermivora, but no definitive resolution of the position of Agathiphagidae, although support is strongest for alliance with Micropterigidae, consistent with another recent molecular study. (ii) There is very strong support for Glossata, which excludes Heterobathmiidae, but weak support for relationships among major homoneurous clades. Eriocraniidae diverge first, corrobo- rating the morphological clade Coelolepida, but the morphological clades Myoglossata and Neolepidoptera are never monophyletic in the molecular trees; both are contra- dicted by strong support for Lophocoronoidea + Hepialoidea, the latter here including