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      Symmetry systems on the wings of Dichromodes Guenée (Lepidoptera: Geometridae) are unconstrained by venation

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      PeerJ
      PeerJ Inc.
      Color pattern, Evolution, Macroheterocera, Scale, Venation

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          Abstract

          The nymphalid groundplan, an idealized schematic illustrating the essential elements of butterfly wing patterns, predicts a consistent relationship between color pattern and wing venation. Moths in the family Geometridae have wing shapes and patterns that often resemble those of butterflies, and until recently, this family was believed to be among butterflies’ closest relatives. However, an examination of the geometrid genus Dichromodes Guenée, 1858 shows no consistent relationship between the central symmetry system and wing venation. Whereas the distal edge of the central symmetry system is predicted to reach the costal margin proximal to the Subcostal vein in butterflies and acronictine moths, it has no consistent relationship with the Subcostal, Radius, or Radial Sector 1 veins in Dichromodes. This finding highlights developmental diversity that was previously overlooked due to the overwhelming preference for butterflies in studies of lepidopteran wing patterns.

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          The industrial melanism mutation in British peppered moths is a transposable element.

          Discovering the mutational events that fuel adaptation to environmental change remains an important challenge for evolutionary biology. The classroom example of a visible evolutionary response is industrial melanism in the peppered moth (Biston betularia): the replacement, during the Industrial Revolution, of the common pale typica form by a previously unknown black (carbonaria) form, driven by the interaction between bird predation and coal pollution. The carbonaria locus has been coarsely localized to a 200-kilobase region, but the specific identity and nature of the sequence difference controlling the carbonaria-typica polymorphism, and the gene it influences, are unknown. Here we show that the mutation event giving rise to industrial melanism in Britain was the insertion of a large, tandemly repeated, transposable element into the first intron of the gene cortex. Statistical inference based on the distribution of recombined carbonaria haplotypes indicates that this transposition event occurred around 1819, consistent with the historical record. We have begun to dissect the mode of action of the carbonaria transposable element by showing that it increases the abundance of a cortex transcript, the protein product of which plays an important role in cell-cycle regulation, during early wing disc development. Our findings fill a substantial knowledge gap in the iconic example of microevolutionary change, adding a further layer of insight into the mechanism of adaptation in response to natural selection. The discovery that the mutation itself is a transposable element will stimulate further debate about the importance of 'jumping genes' as a source of major phenotypic novelty.
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            A Large-Scale, Higher-Level, Molecular Phylogenetic Study of the Insect Order Lepidoptera (Moths and Butterflies)

            Background Higher-level relationships within the Lepidoptera, and particularly within the species-rich subclade Ditrysia, are generally not well understood, although recent studies have yielded progress. We present the most comprehensive molecular analysis of lepidopteran phylogeny to date, focusing on relationships among superfamilies. Methodology / Principal Findings 483 taxa spanning 115 of 124 families were sampled for 19 protein-coding nuclear genes, from which maximum likelihood tree estimates and bootstrap percentages were obtained using GARLI. Assessment of heuristic search effectiveness showed that better trees and higher bootstrap percentages probably remain to be discovered even after 1000 or more search replicates, but further search proved impractical even with grid computing. Other analyses explored the effects of sampling nonsynonymous change only versus partitioned and unpartitioned total nucleotide change; deletion of rogue taxa; and compositional heterogeneity. Relationships among the non-ditrysian lineages previously inferred from morphology were largely confirmed, plus some new ones, with strong support. Robust support was also found for divergences among non-apoditrysian lineages of Ditrysia, but only rarely so within Apoditrysia. Paraphyly for Tineoidea is strongly supported by analysis of nonsynonymous-only signal; conflicting, strong support for tineoid monophyly when synonymous signal was added back is shown to result from compositional heterogeneity. Conclusions / Significance Support for among-superfamily relationships outside the Apoditrysia is now generally strong. Comparable support is mostly lacking within Apoditrysia, but dramatically increased bootstrap percentages for some nodes after rogue taxon removal, and concordance with other evidence, strongly suggest that our picture of apoditrysian phylogeny is approximately correct. This study highlights the challenge of finding optimal topologies when analyzing hundreds of taxa. It also shows that some nodes get strong support only when analysis is restricted to nonsynonymous change, while total change is necessary for strong support of others. Thus, multiple types of analyses will be necessary to fully resolve lepidopteran phylogeny.
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              Comprehensive gene and taxon coverage elucidates radiation patterns in moths and butterflies.

              Lepidoptera (butterflies and moths) represent one of the most diverse animals groups. Yet, the phylogeny of advanced ditrysian Lepidoptera, accounting for about 99 per cent of lepidopteran species, has remained largely unresolved. We report a rigorous and comprehensive analysis of lepidopteran affinities. We performed phylogenetic analyses of 350 taxa representing nearly 90 per cent of lepidopteran families. We found Ditrysia to be a monophyletic taxon with the clade Tischerioidea + Palaephatoidea being the sister group of it. No support for the monophyly of the proposed major internested ditrysian clades, Apoditrysia, Obtectomera and Macrolepidoptera, was found as currently defined, but each of these is supported with some modification. The monophyly or near-monophyly of most previously identified lepidopteran superfamilies is reinforced, but several species-rich superfamilies were found to be para- or polyphyletic. Butterflies were found to be more closely related to 'microlepidopteran' groups of moths rather than the clade Macrolepidoptera, where they have traditionally been placed. There is support for the monophyly of Macrolepidoptera when butterflies and Calliduloidea are excluded. The data suggest that the generally short diverging nodes between major groupings in basal non-tineoid Ditrysia are owing to their rapid radiation, presumably in correlation with the radiation of flowering plants.
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                Author and article information

                Contributors
                Journal
                PeerJ
                PeerJ
                peerj
                peerj
                PeerJ
                PeerJ Inc. (San Diego, USA )
                2167-8359
                2 January 2020
                2020
                : 8
                : e8263
                Affiliations
                [-1] Department of Geological Sciences, Stanford University , Stanford, CA, United States of America
                Article
                8263
                10.7717/peerj.8263
                6942684
                86d72399-23cc-4bee-a44f-4254aaf2410a
                ©2020 Schachat

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, reproduction and adaptation in any medium and for any purpose provided that it is properly attributed. For attribution, the original author(s), title, publication source (PeerJ) and either DOI or URL of the article must be cited.

                History
                : 29 August 2019
                : 21 November 2019
                Funding
                Funded by: National Science Foundation Graduate Research Fellowship
                Award ID: DGE-1125191
                Funded by: Graduate Research Opportunities Worldwide, a joint initiative of Universities Australia and the National Science Foundation
                Funded by: Chair’s Fellowship in the Department of Geological Sciences, Stanford University
                Funded by: Coleman F. Fung Interdisciplinary Graduate Fellowship, Vice Provost of Graduate Education, Stanford University
                Funded by: Bio-X, Stanford University
                Funding was provided by National Science Foundation Graduate Research Fellowship grant no. DGE-1125191; by Graduate Research Opportunities Worldwide, a joint initiative of Universities Australia and the National Science Foundation; by the Chair’s Fellowship in the Department of Geological Sciences, Stanford University; by the Coleman F. Fung Interdisciplinary Graduate Fellowship, Vice Provost of Graduate Education, Stanford University; and by Bio-X, Stanford University. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Biodiversity
                Entomology
                Evolutionary Studies
                Zoology

                color pattern,evolution,macroheterocera,scale,venation
                color pattern, evolution, macroheterocera, scale, venation

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