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This work was supported through funding from the Australian Government’s Australian Biological Resources Study (ABRS) Bush Blitz Strategic Taxonomy Grants Scheme for a project entitled ‘Taxonomic revision of Candalides and allied genera in the tribe Candalidini (Lepidoptera: Lycaenidae)’ to MFB, National Geographic grant WW-227R-17 to DJL and National Science Foundation grants DEB-1541500 to AYK, DEB-1541557 to DJL and DEB-1541560 to NEP

The origin of taxa presenting a disjunct distribution between Africa and Asia has puzzled biogeographers for more than a century. This biogeographic pattern has been hypothesized to be the result of transoceanic long-distance dispersal, Oligocene dispersal through forested corridors, Miocene dispersal through the Arabian Peninsula or passive dispersal on the rifting Indian plate. However, it has often been difficult to pinpoint the mechanisms at play. We investigate biotic exchange between the Afrotropics and the Oriental region during the Cenozoic, a period in which geological changes altered landmass connectivity. We use Baorini skippers (Lepidoptera, Hesperiidae) as a model, a widespread clade of butterflies in the Old World tropics with a disjunct distribution between the Afrotropics and the Oriental region. We use anchored phylogenomics to infer a robust evolutionary tree for Baorini skippers and estimate divergence times and ancestral ranges to test biogeographic hypotheses. Our phylogenomic tree recovers strongly supported relationships for Baorini skippers and clarifies the systematics of the tribe. Dating analyses suggest that these butterflies originated in the Oriental region, Greater Sunda Islands, and the Philippines in the early Miocene c. 23 Ma. Baorini skippers dispersed from the Oriental region towards Africa at least five times in the past 20 Ma. These butterflies colonized the Afrotropics primarily through trans-Arabian geodispersal after the closure of the Tethyan seaway in the mid-Miocene. Range expansion from the Oriental region towards the African continent probably occurred via the Gomphotherium land bridge through the Arabian Peninsula. Alternative scenarios invoking long-distance dispersal and vicariance are not supported. The Miocene climate change and biome shift from forested areas to grasslands possibly facilitated geodispersal in this clade of butterflies., Directorate for Biological Sciences. Grant Numbers: DEB‐1541500, DEB‐1541560.

The Gelechioidea (>18 000 species), one of the largest superfamilies of Lepidoptera, are a major element of terrestrial ecosystems and include important pests and biological model species. Despite much recent progress, our understanding of the classification, phylogeny and evolution of Gelechioidea remains limited. Building on recent molecular studies of this superfamily and a recently revised family/subfamily classification, we provide an independent estimate of among-family relationships, with little overlap in gene sample. We analysed up to five nuclear genes, totalling 6633 bp, for each of 77 gelechioids, plus up to 14 additional genes, for a total of 14 826 bp, in 45 of those taxa and all 19 outgroup taxa. Our maximum-likelihood (ML) analyses, like those of previous authors, strongly support monophyly for most multiply-sampled families and subfamilies, but very weakly support most relationships above the family level. Our tree looks superficially divergent from that of the most recent molecular study of gelechioids, but when the previous tree is re-rooted to accord maximally with ours, the two phylogenies agree entirely on the deepest-level divergences in Gelechioidea, and strongly though incompletely on among-family relationships within the major groups. This concordance between independent studies is evidence that the groupings (or at least the unrooted branching order) are probably accurate, despite the low bootstrap values. After re-rooting, both trees divide the families into three monophyletic groups: a ‘Gelechiid Assemblage,’ consisting of Gelechiidae and Cosmopterigidae; a ‘Scythridid Assemblage,’ consisting of Stathmopodidae, Scythrididae, Blastobasidae, Elachistidae, Momphidae, Coleophoridae and Batrachedridae; and a ‘Depressariid Assemblage,’ consisting of Autostichidae, Xyloryctidae, Lecithoceridae, Oecophoridae, Depressariidae and Lypusidae. Within the largest family, Gelechiidae, our results strongly support the pairing of Anomologinae with Gelechiinae, in accordance with a recent study of this family. Relationships among the other subfamilies, however, conflict moderately to strongly between studies, leaving the intrafamily phylogeny unsettled. Within the ‘Scythridid Assemblage,’ both trees support an ‘SSB clade’ consisting of Blastobasidae + (Scythrididae + Stathmopodidae), strongly resolved only in our results. Coleophoridae + Batrachedridae is supported, albeit weakly, in both trees, and only Momphidae differ in position between studies. Within the ‘Depressariid Assemblage,’ both trees support an ‘AXLO’ clade consisting of Autostichidae, Xyloryctidae, Lecithoceridae and Oecophoridae. The monophyly of this clade and relationships therein are supported weakly in previous results but strongly in ours. The recently re-defined family Depressariidae is paraphyletic in our tree, but the evidence against depressariid monophyly is very weak. There is moderate support for a core group of Depressariidae consisting, among the seven subfamilies we sampled, of Depressariinae, Aeolanthinae and Hypertrophinae. We show that gelechioids have a higher total number and percentage of species that are saprophagous as larvae than any other apoditrysian superfamily, that saprophagy is concentrated primarily in the ‘AXLO clade,’ and that the ancestral gelechioid condition was probably feeding on live plants. Among the living-plant feeders, concealed external feeding was probably the ancestral state. The multiple origins of internal feeding of various kinds, including leaf mining (otherwise almost unknown in Apoditrysia), are restricted mostly to the Scythridid and Gelechiid Assemblages. The traits that predispose or permit lineages to adopt these unusual life histories are worthy of study.

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)

Many insects possess conspicuous external circular ring markings that resemble the eye of a vertebrate. These ‘eyespots’ typically function to startle or otherwise deter predators, but few studies have examined how eyespots have evolved. We study the evolution of the posterior larval eyespot in the charismatic New World hawkmoth genus Eumorpha. While Eumorpha has a range of posterior larval eyespot shapes and sizes, little is known of how this trait has evolved because phylogenetic relationships of Eumorpha remain largely unknown. In this study, we included 62 individuals from 23 of 26 described Eumorpha species, and sequenced four genes (CAD, EF-1α, Wingless and COI), totaling 3773 base pairs. Maximum likelihood and Bayesian phylogenetic methods produced largely congruent trees with well-supported relationships. Our analyses reveal that Eumorpha probably had an ancestor with a posterior larval eyespot and that the eyespot was subsequently lost in at least three lineages. Eumorpha appears to have originated in Central and South America and expanded its distribution to North America.

The Afrotropical butterfly subfamily Pseudopontiinae (Pieridae) was traditionally thought to comprise one species, with two subspecies (Pseudopontia paradoxa paradoxa Felder & Felder and Pseudopontia paradoxa australis Dixey) differing in a single detail of a hindwing vein. The two subspecies also differ in their known geographic distributions (mainly north of versus south of the equator). Unlike most butterflies, Pseudopontia is white with no visible wing or body markings. We now report that males of P. paradoxa australis have an area of ultraviolet-reflecting scales along the anal vein of the forewing, whereas males of P. paradoxa paradoxa and all females do not. A total of 21 individuals of the northern subspecies, which were collected in three localities south of the equator, were found in the collection of the Royal Museum for Central Africa, indicating sympatry of the two traditional subspecies in the Congo River basin. To determine if additional cryptic species might be present, we sequenced three nuclear genes (CAD, DDC and wingless) as well as cytochrome oxidase I (COI), examined amplified fragment-length polymorphisms, and re-examined wing and genitalic morphology, using recently collected specimens from several regions of Africa. Phylogenetic analyses of the COI sequences and amplified fragment-length polymorphism data concur, and indicate the existence of at least five monophyletic, non-interbreeding populations, with a particularly deep divergence between three populations of P. paradoxa paradoxa and two of P. paradoxa australis. Despite the slow rate of evolution of the nuclear genes studied, individual gene trees and a concatenated three-gene tree demonstrate, with high bootstrap support, clear divergence among the five populations of Pseudopontia. In addition, consistent variations in details of wing vein stalks were found among four of the genetically distinct populations, which supports the hypothesis of multiple species. Division of Pseudopontia into five phylogenetic species is proposed, including the elevation of ssp. australis to species rank and the description of Pseudopontia mabira, Pseudopontia gola and Pseudopontia zambezi