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One of the key objectives in biological research is understanding how evolutionary processes have produced Earth’s diversity. A critical step toward revealing these processes is an investigation of evolutionary tradeoffs—that is, the opposing pressures of multiple selective forces. For millennia, nocturnal moths have had to balance successful flight, as they search for mates or host plants, with evading bat predators. However, the potential for evolutionary trade-offs between wing shape and body size are poorly understood. In this study, we used phylogenomics and geometric morphometrics to examine the evolution of wing shape in the wild silkmoth subfamily Arsenurinae (Saturniidae) and evaluate potential evolutionary relationships between body size and wing shape. The phylogeny was inferred based on 782 loci from target capture data of 42 arsenurine species representing all 10 recognized genera. After detecting in our data one of the most vexing problems in phylogenetic inference—a region of a tree that possesses short branches and no “support” for relationships (i.e., a polytomy), we looked for hidden phylogenomic signal (i.e., inspecting differing phylogenetic inferences, alternative support values, quartets, and phylogenetic networks) to better illuminate the most probable generic relationships within the subfamily. We found there are putative evolutionary trade-offs between wing shape, body size, and the interaction of fore- and hindwing (HW) shape. Namely, body size tends to decrease with increasing HW length but increases as forewing (FW) shape becomes more complex. Additionally, the type of HW (i.e., tail or no tail) a lineage possesses has a significant effect on the complexity of FW shape. We outline possible selective forces driving the complex HW shapes that make Arsenurinae, and silkmoths as a whole, so charismatic. [Anchored hybrid enrichment; Arsenurinae; geometric morphometrics; Lepidoptera; phylogenomics; Saturniidae.]

Butterflies and moths (Lepidoptera) are one of the most studied, diverse, and widespread animal groups, making them an ideal model for climate change research. They are a particularly informative model for studying the effects of climate change on species ecology because they are ectotherms that thermoregulate with a suite of physiological, behavioural, and phenotypic traits. While some species have been negatively impacted by climatic disturbances, others have prospered, largely in accordance with their diversity in life‐history traits. Here we take advantage of a large repertoire of studies on butterflies and moths to provide a review of the many ways in which climate change is impacting insects, animals, and ecosystems. By studying these climate‐based impacts on ecological processes of Lepidoptera, we propose appropriate strategies for species conservation and habitat management broadly across animals.

Amateur butterfly and moth collectors in the United States have procured more Lepidoptera specimens than professional scientists. The advent of large government-supported database efforts has made a quantitative examination of the impact of amateur collecting of these insects possible. We reviewed trends in Lepidoptera collecting since 1800, using more than 1 million United States lepidopteran specimens that have been collected into public databases. Our findings show a steep rise in the collection of specimens after World War II, followed by a short plateau and sharp decline in the late 1990s. In contrast, the rate of observations submitted to database groups dramatically increased around 2005. Declining acquisition of Lepidoptera specimens may compromise critically important testing of contemporary and future ecological, conservation, and evolutionary hypotheses on a grand scale, particularly given documented declines in insect populations. We suggest that increasing collaboration between professional and community-based scientists could alleviate the decline in amateur-collected specimens.

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 complex forces that shape butterfly wings have long been a subject of experimental and comparative research. Butterflies use their wings for flight, camouflage, mate recognition, warning, and mimicry. However, general patterns and correlations among wing shape and size evolution are still poorly understood. We collected geometric morphometric measurements from over 1400 digitized museum specimens of Papilio swallowtails and combined them with phylogenetic data to test two hypotheses: 1) forewing shape and size evolve independently of hindwing shape and size and 2) wing size evolves more quickly than wing shape. We also determined the major axes of wing shape variation and discovered that most shape variability occurs in hindwing tails and adjacent areas. We conclude that forewing shape and size are functionally and biomechanically constrained, whereas hindwings are more labile, perhaps in response to disruptive selective pressure for Batesian mimicry or against predation. The development of a significant, re-usable, digitized data resource will enable further investigation on tradeoffs between flight performance and ecological selective pressures, along with the degree to which intraspecific, local-scale selection may explain macroevolutionary patterns. [Batesian mimicry; Lepidoptera; geometric morphometrics; museum specimens.]

Background The subfamily Geometrinae (Lepidoptera: Geometridae), commonly known as emerald moths, is an ecologically diverse group of moths with over 2,500 described species. Many taxonomic and phylogenetic studies of Geometrinae have been undertaken in the past decade, resulting in hundreds of new taxonomic changes since online publication of the most recent checklist in December 2007. New information This review synthesises the last 12 years of alpha-taxonomic research in Geometrinae. A comprehensive list of Geometrinae genus- and species-group descriptions, synonymies, combinations and other taxonomic changes, made since 2007, is provided. Since 2007, the known species richness of Geometrinae has increased from 2,529 to 2,642 species; an updated list of all these species is presented in a supplementary spreadsheet.

The global increase in species richness toward the tropics across continents and taxonomic groups, referred to as the latitudinal diversity gradient, stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluate several of these hypotheses to explain spatial diversity patterns in a butterfly family, the Nymphalidae, by assessing the contributions of speciation, extinction, and dispersal, and also the extent to which these processes differ among regions at the same latitude. We generate a time-calibrated phylogeny containing 2,866 nymphalid species (~45% of extant diversity). Neither speciation nor extinction rate variations consistently explain the latitudinal diversity gradient among regions because temporal diversification dynamics differ greatly across longitude. The Neotropical diversity results from low extinction rates, not high speciation rates, and biotic interchanges with other regions are rare. Southeast Asia is also characterized by a low speciation rate but, unlike the Neotropics, is the main source of dispersal events through time. Our results suggest that global climate change throughout the Cenozoic, combined with tropical niche conservatism, played a major role in generating the modern latitudinal diversity gradient of nymphalid butterflies., F.L.C is supported by an “Investissements d’Avenir” grant managed by the Agence Nationale de la Recherche (CEBA, ref. ANR-10-LABX-25-01). D.J.L. was supported by grants DEB-1541557 from NSF and WW-227R-17 from the National Geographic Society. R.I.V.W. was supported by Leverhulme Trust emeritus programme. M.E. was supported by an ATIP grant, a grant from the Human Frontier Science Program (RGP0014/2016) and a grant from the French National Research Agency (ANR CLEARWING ANR-16-CE02-0012). E.O.A. was supported by Sigma-Xi (G20100315153261), Center for Systematic Entomology and the Council of the Linnean Society and the Systematics Association for the Systematics Research Fund. S.N. was supported by the Swedish Research Council (2015-04218 and 2019-03441). P.M.M. was supported by a Czech Science Foundation grant (Junior GAČR, 20-18566Y). R.V. was supported by projects PID2019-107078GB-I00 / AEI / 10.13039/501100011033 (Agencia Estatal de Investigación) and 2017-SGR-991 (Generalitat de Catalunya). N.W. acknowledges support by the Swedish Research Council (2015-04441) and a start-up grant from the Department of Biology, Lund University.

Phylogenomics is progressing rapidly, allowing large strides forward into our understanding of the tree of life. In this study, we generated transcriptomes from ethanol-preserved specimens of 13 tiger beetle species (Coleoptera: Cicindelinae) and one Scaritinae outgroup. From these 14 transcriptomes and seven publicly available transcriptomes, we recovered an average of 2538 loci for phylogenetic analysis. We constructed an evolutionary tree of tiger beetles to examine deep-level relationships and examined the extent to which the composition of the dataset, missing data, gene tree inconsistency and codon position saturation impacted phylogenetic accuracy. Ethanol-preserved specimens yielded similar numbers of loci to specimens originally preserved in costly reagents, showcasing more flexibility in transcriptomics than anticipated. The number of loci and gene tree inconsistency had less impact on downstream results than third codon position saturation and missing data. Our results recovered tiger beetles as sister to Carabidae with strong support, confirming their taxonomic status as an independent family within Adephaga. Within tiger beetles, phylogenetic relationships were robust across all nodes. This new phylogenomic backbone represents a useful framework for future endeavours in tiger beetle systematics and serves as a starting point for the development of less costly target capture toolkits to expand the taxonomic breadth of the future tiger beetle tree of life.

BackgroundThe silkmoths and their relatives constitute the ecologically and taxonomically diverse superfamily Bombycoidea, which includes some of the most charismatic species of Lepidoptera. Despite displaying some of the most spectacular forms and ecological traits among insects, relatively little attention has been given to understanding their evolution and the drivers of their diversity.ResultsTo begin to address this problem, we created a new Bombycoidea-specific Anchored Hybrid Enrichment (AHE) probe set and sampled up to 571 loci for 117 taxa across all major lineages of the Bombycoidea, producing a well-supported phylogeny. The tree was overall consistent with prior morphological and molecular studies, although some taxa (e.g.,ArotrosSchaus) were misplaced in the Bombycidae and here formally transferred to Apatelodidae. We identified important evolutionary patterns (e.g., morphology, biogeography, and differences in speciation and extinction), and our analysis of diversification rates highlights the stark increases that exist within the Sphingidae (hawkmoths) and Saturniidae (wild silkmoths).ConclusionsWe postulate that these rate shifts are due to the well-documented bat-moth “arms race” and differences in selective pressures from insectivorous bats. The study establishes a backbone for future evolutionary, comparative, and taxonomic studies, and presents a modified DNA extraction protocol that allows Lepidoptera specimens to be readily sequenced from pinned natural history collections, succeeding in samples up to 30 years old. Our research highlights the flexibility of AHE to generate genomic data from a wide range of museum specimens, both age and preservation method, and will allow researchers to tap into the wealth of biological data residing in natural history collections around the globe.

Relationships within satyrine butterflies have been notoriously difficult to resolve using both morphology and Sanger sequencing methods, and this is particularly true for the mainly Neotropical subtribe Euptychiina, which contains about 400 described species. Known larvae of Euptychiina feed on grasses and sedges, with the exception of the genus Euptychia, which feed on mosses and lycopsids, and the butterflies occur widely in rainforest, cloudforest and grassland habitats, where they are often abundant. Several previous molecular and morphological studies have made significant progress in tackling the systematics of the group, but many relationships remain unresolved, with long-branch-attraction artifacts being a major problem. Additionally, the monophyly of the clade remains uncertain, with Euptychia possibly not being closely related to the remainder of the clade. Here we present a backbone phylogeny of the subtribe based on 106 taxa, 368 nuclear loci, and over 180,000 bps obtained through hybrid enrichment. Using both concatenation and species tree approaches (IQ-TREE, EXABAYES, ASTRAL), we can for the first time strongly confirm the monophyly of Euptychiina with Euptychia being the sister group to the remainder of the clade. The Euptychiina is divided into nine well supported clades, but the placement of a few genera such as Hermeuptychia, Pindis and the Chloreuptychia catharina group still remain uncertain. As partially indicated in previous studies, the genera Cissia, Chloreuptychia, Magneuptychia, Megisto, Splendeuptychia and Euptychoides, among others, were found to be highly polyphyletic and revisions are in preparation. The phylogeny will provide a strong backbone for the analysis of datasets in development that are much more taxonomically comprehensive but have orders of magnitude fewer loci. This study therefore represents a critical step towards resolving the higher classification and studying the evolution of this highly diverse lineage.

Background The availability of thousands of genomes has enabled new advancements in biology. However, many genomes have not been investigated for their quality. Here we examine quality trends in a taxonomically diverse and well-known group, butterflies (Papilionoidea), and provide draft, de novo assemblies for all available butterfly genomes. Owing to massive genome sequencing investment and taxonomic curation, this is an excellent group to explore genome quality. Findings We provide de novo assemblies for all 822 available butterfly genomes and interpret their quality in terms of completeness and continuity. We identify the 50 highest quality genomes across butterflies and conclude that the ringlet, Aphantopus hyperantus, has the highest quality genome. Our post-processing of draft genome assemblies identified 118 butterfly genomes that should not be reused owing to contamination or extremely low quality. However, many draft genomes are of high utility, especially because permissibility of low-quality genomes is dependent on the objective of the study. Our assemblies will serve as a key resource for papilionid genomics, especially for researchers without computational resources. Conclusions Quality metrics and assemblies are typically presented with annotated genome accessions but rarely with de novo genomes. We recommend that studies presenting genome sequences provide the assembly and some metrics of quality because quality will significantly affect downstream results. Transparency in quality metrics is needed to improve the field of genome science and encourage data reuse.

Alternative life-history strategies (ALHS) are genetic polymorphisms generating phenotypes differing in life histories that generally arise due to metabolic resource allocation tradeoffs. Althouigh ALHS are often be limited to a single sex or populations of a species, they can, in rare cases, be found among several species across a genus. In the butterfly genus Colias, at least a third of the species have a female limited ALHS called Alba. While many females develop brightly pigmented wings, Alba females reallocate nitrogen resources used in pigment synthesis to reproductive development, producing white-winged, more fecund females. Whether this ALHS evolved once or many times, and whether it has moved among species via introgression or been maintained via long-term balancing selection, has not been established. Answering these questions presents an opportunity to investigate the genetic basis and evolutionary forces acting upon ALHS, which have rarely been studied at a genus level. Here we identify the genetic locus of Alba in a second Colias species, allowing us to compare this with previous results in a larger phylogenetic context. Our findings suggest Alba has a singular origin and has been maintained in Colias through a combination of balancing selection and introgression for nearly one million years and at least as many generations. Finally, using CRISPR/Cas9 deletions in the cis-regulatory region of the Alba allele, we demonstrate that the Alba allele is a modular enhancer for the BarH1 gene and is necessary for the induction of the ALHS, which potentially facilitates its long-term persistence in the genus.

Male butterflies in the hyperdiverse tribe Eumaeini possess an unusually complex and diverse repertoire of secondary sexual characteristics involved in pheromone production and dissemination. Maintaining multiple sexually selected traits is likely to be metabolically costly, potentially resulting in trade-offs in the evolution of male signals. However, a phylogenetic framework to test hypotheses regarding the evolution and maintenance of male sexual traits in Eumaeini has been lacking. Here, we infer a comprehensive, time-calibrated phylogeny from 379 loci for 187 species representing 91% of the 87 described genera. Eumaeini is a monophyletic group that originated in the late Oligocene and underwent rapid radiation in the Neotropics. We examined specimens of 818 of the 1096 described species (75%) and found that secondary sexual traits are present in males of 91% of the surveyed species. Scent pads and scent patches on the wings and brush organs associated with the genitalia were probably present in the common ancestor of Eumaeini and are widespread throughout the tribe. Brush organs and scent pads are negatively correlated across the phylogeny, exhibiting a trade-off in which lineages with brush organs are unlikely to regain scent pads and vice versa. In contrast, scent patches seem to facilitate the evolution of scent pads, although they are readily lost once scent pads have evolved. Our results illustrate the complex interplay between natural and sexual selection in the origin and maintenance of multiple male secondary sexual characteristics and highlight the potential role of sexual selection spurring diversification in this lineage., Proceedings of the Royal Society B: Biological Sciences, 288 (1950), ISSN:0080-4649, ISSN:0950-1193, ISSN:1471-2954, ISSN:0962-8452

Opsins, combined with a chromophore, are the primary light-sensing molecules in animals and are crucial for color vision. Throughout animal evolution, duplications and losses of opsin proteins are common, but it is unclear what is driving these gains and losses. Light availability is implicated, and dim environments are often associated with low opsin diversity and loss. Correlations between high opsin diversity and bright environments, however, are tenuous. To test if increased light availability is associated with opsin diversification, we examined diel niche and identified opsins using transcriptomes and genomes of 175 butterflies and moths (Lepidoptera). We found 14 independent opsin duplications associated with bright environments. Estimating their rates of evolution revealed that opsins from diurnal taxa evolve faster—at least 13 amino acids were identified with higher dN/dS rates, with a subset close enough to the chromophore to tune the opsin. These results demonstrate that high light availability increases opsin diversity and evolution rate in Lepidoptera., Yash Sondhi et al. annotate the genomes and transcriptomes of 175 Lepidoptera species and identify 14 independent opsin duplications associated with bright environments. From estimating their rates of evolution, the authors reveal that opsins from diurnal taxa evolve faster, and that high light availability increases opsin diversity and evolution rate in this order.

Insects constitute the vast majority of known animal species and are ubiquitous across terrestrial ecosystems, playing key ecological roles. As prey, they are critical to the survival of countless other species, including the majority of bats, birds, and freshwater fishes (1). As herbivores, predators, and parasites, they are major determinants of the distribution and abundance of innumerable plants and animals. The majority of flowering plants, the dominant component of most terrestrial ecosystems, depend on insects for pollination and hence reproduction. As consumers of waste products, insects are essential to the recycling of nutrients. Humans and their agriculture rely heavily on such “ecosystem services” provided by insects (Fig. 1 A – J ), which together have at least an annual value of ∼$70 billion (2020 valuation) in the United States (2). Insects also provide humans with honey, silk, wax, dyes, and, in many cultures, food. Insects have become essential subjects in medical and basic biological research. Furthermore, insects are one of the most easily accessible forms of wildlife, with a diversity of morphology, life history, and behavior that seems ready-made for inspiring appreciation of nature and its conservation (Fig. 1 K – T ). Fig. 1. Examples of insects that are beneficial to humans ( A – J ), and examples of amazing things that insects do ( K – T ). A , Pollinator: Honey bee ( Apis mellifera ). B , Bumble bee ( Bombus sp.). C , Decomposer: Dung beetle (Scarabaeidae: Scarabaeinae). D , Biocontrol: Ladybird beetle ( Harmonia sp.). E , Ecosystem service: Ants (Formicidae). F , Silk production: Silk moth ( Bombyx mori ). G , Research: Fruit fly ( Drosophila melanogaster ). H , Vaccine development, including coronavirus: Alfalfa looper moth ( Autographa californica ). I , Dye production: Cochineal scale insect ( Dactylopius coccus ). J , Environmental assessment: Stonefly (Plecoptera). K , Visual systems: Dragonflies (e.g., Neurothemis sp.) have near 360° vision. L , Visual defense: … [↵][1]1To whom correspondence may be addressed. Email: kawahara{at}flmnh.ufl.edu. [1]: #xref-corresp-1-1

International audience; Phylogenomics seeks to use next‐generation data to robustly infer an organism's evolutionary history. Yet, the practical caveats of phylogenomics motivate investigation of improved efficiency, particularly when quality of phylogenies are questionable. To achieve improvements, one goal is to maintain or enhance the quality of phylogenetic inference while severely reducing dataset size. We approach this by assessing which kinds of loci in phylogenomic alignments provide the majority of support for a phylogenetic inference of cockroaches in Blaberoidea. We examine locus substitution rate, saturation, evolutionary divergence, rate heterogeneity, stabilizing selection, and a priori information content as traits that may determine optimality. Our controlled experimental design is based on 265 loci for 102 blaberoidean taxa and 22 outgroup species. Loci with high substitution rate, low saturation, low sequence distance, low rate heterogeneity, and strong stabilizing selection derive more support for phylogenetic relationships. We found that some phylogenetic information content estimators may not be meaningful for assessing information content a priori. We use these findings to design concatenated datasets with an optimized subsample of 100 loci. The tree inferred from the optimized subsample alignment was largely identical to that inferred from all 265 loci but with less evidence of long branch attraction, improved statistical support, and potential 4‐6x improvements to computation time. Supported by phylogenetic and morphological evidence, we erect three newly named clades (Anallactinae Evangelista & Wipfler subfam. nov., Orkrasomeria tax. nov. Evangelista, Wipfler, & Béthoux and Hemithyrsocerini Evangelista tribe nov.) and propose other taxonomic modifications. The diagnosis of Pseudophyllodromiidae Grandcolas, 1996 is modified to accommodate Anallactinae and Pseudophyllodromiinae Vickery & Kevan, 1983. The diagnosis of Ectobiidae Brunner von Wattenwyl, 1865 is modified to add novel morphological characters.

Predators can have strong influences on ecological processes through impacts on individuals at lower trophic levels, and changes in predator-prey dynamics can alter ecosystem functioning. However, much of what we currently know about interactions across trophic levels comes from mesic or relatively fertile systems, with fewer studies examining trophic interactions and resulting ecosystem processes in arid or infertile systems. To address this knowledge gap, we excluded avian predators from shrubs during the growing season using netting in a sagebrush steppe environment. We compared arthropod abundance, shrub herbivory damage, physiology (gas exchange and chlorophyll fluorescence), litter chemistry (C/N ratios and concentrations of phenolic compounds) and decomposition between netted and un-netted (control) sagebrush (Artemisia tridentata ssp. wyomingensis) shrubs across a growing season. While there were clear seasonal patterns in measurements, we observed no statistically significant differences between netting treatments in any of these measurements, though abundances of arthropods in the sap-feeding trophic guilds were appreciably greater (although not significantly) on netted compared to un-netted shrubs. Our results suggest that in the short-term, either top-down effects in this sage-steppe ecosystem are minimal, and/or inter-trophic interactions (vertebrate predators-arthropods-plants) are relatively weak and more dependent on bottom-up processes that are linked with abiotic variables.

DNA analysis of predator faeces using high-throughput amplicon sequencing (HTS) enhances our understanding of predator-prey interactions. However, conclusions drawn from this technique are constrained by biases that occur in multiple steps of the HTS workflow. To better characterize insectivorous animal diets, we used DNA from a diverse set of arthropods to assess PCR biases of commonly used and novel primer pairs for the mitochondrial gene, cytochrome oxidase C subunit 1 (COI). We compared diversity recovered from HTS of bat guano samples using a commonly used primer pair "ZBJ" to results using the novel primer pair "ANML." To parameterize our bioinformatics pipeline, we created an arthropod mock community consisting of single-copy (cloned) COI sequences. To examine biases associated with both PCR and HTS, mock community members were combined in equimolar amounts both pre- and post-PCR. We validated our system using guano from bats fed known diets and using composite samples of morphologically identified insects collected in pitfall traps. In PCR tests, the ANML primer pair amplified 58 of 59 arthropod taxa (98%), whereas ZBJ amplified 24-40 of 59 taxa (41%-68%). Furthermore, in an HTS comparison of field-collected samples, the ANML primers detected nearly fourfold more arthropod taxa than the ZBJ primers. The additional arthropods detected include medically and economically relevant insect groups such as mosquitoes. Results revealed biases at both the PCR and sequencing levels, demonstrating the pitfalls associated with using HTS read numbers as proxies for abundance. The use of an arthropod mock community allowed for improved bioinformatics pipeline parameterization.

A backbone molecular phylogeny of Mimallonidae, based on 47 species and 515 loci, was recently published. That study resolved some of the major relationships in the family, but taxon sampling was limited and a classification of the family was not formally presented for all species. Here morphological phylogenetic analyses in parsimony and maximum likelihood (ML) frameworks were conducted that included 192 species and 55 morphological characters. A phylogenetic analysis was also conducted on the morphological dataset with a topological constraint based on the 515 locus tree from the previous study. Results show that nearly all species can be confidently placed in a genus using morphological phylogenetics. The presence of a frenulum, a character that was historically used to distinguish major groups of Mimallonidae, varies within and among genera. Based on our phylogenetic results, the classification of Mimallonidae is revised, which now includes 291 species in 41 genera. Descriptions of three new genera are included:Fatellallagen. n.,Citrallagen. n., andLepismallagen. n.The following taxonomic changes were made in the present article: 43 new/revived combinations (inAceclostriaVuillot,ArcinnusHerbin,BedosiaSchaus,BedosialloSt Laurent & Kawahara,CicinnusBlanchard,Citralla,DruenticaStrand,Fatellalla,LacosomaGrote,Lepismalla,MimalloHübner,ProcinnusHerbin,PsychocampaGrote,RoelmanaSchaus, andThaeliaHerbin), two new species-level synonyms (meliniBryk is synonymized withviemandaSchaus,jarugaJones is synonymized withhamataWalker), one revived synonymy (roscidaDognin is resynonymized withexternaMoore), seven new statuses (inDruentica,MacessogaSchaus, andTrogopteraHerrich-Schäffer), six revived statuses (inAceclostria,Cicinnus,Druentica,Psychocampa, andZaphantaDyar), and one new designation ofnomen nudum. In order to alleviate nomenclatural problems, twelve lectotypes are designated (forTolypidaamaryllis(Schaus),TrogopteraalthoraSchaus,AdalgisacroesaSchaus,Alheitapulloides(Dognin),LacosomabriasiaSchaus,LacosomadiedericaSchaus,LacosomaraydelaSchaus,Psychocampalacuna(Schaus),CicinnuscorallinaDognin,CicinnuslatrisSchaus,CicinnussolvensSchaus,CicinnustuisanaSchaus) as well as a neotype forMimallodespectaWalker (=Cicinnusdespecta). This paper also provides apomorphies for each genus and a morphological key to genera. Annotations are given to aid researchers in understanding all changes made herein, and images of male and female and their genitalia are present for nearly all type species.

A new species of cicinnine Mimallonidae, Cicinnus chambersisp. nov., is described from the Sky Islands Region of southern Arizona, USA. The new species is closely related to C. mexicana (Druce), type locality Veracruz, Mexico, based on morphology and genetics. The other Cicinnus species known from the United States, the common C. melsheimeri (type locality Pennsylvania, USA) is morphologically and genetically distinct from both C. chambersi and C. mexicana. The new species is compared to C. mexicana and C. melsheimeri, as well as other Mexican Cicinnus. The life history of C. chambersi is unknown, but its description should facilitate future studies on this rarely reported North American mimallonid, a species which may have only recently become established in the United States. Cicinnus chambersi is the fifth known Mimallonidae species from the United States, and the first described from the country in nearly half a century.

The latitudinal diversity gradient (LDG) is arguably one of the most striking patterns in nature. The global increase in species richness toward the tropics across continents and taxonomic groups stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluated several of these hypotheses to explain spatial diversity patterns in the butterfly family, Nymphalidae, by assessing the contributions of speciation, extinction, and dispersal to the LDG, and also the extent to which these processes differ among regions at the same latitude. We generated a new, time-calibrated phylogeny of Nymphalidae based on 10 gene fragments and containing ca. 2,800 species (∼45% of extant diversity). Neither speciation nor extinction rate variations consistently explain the LDG among regions because temporal diversification dynamics differ greatly across longitude. For example, we found that Neotropical nymphalid diversity results from low extinction rates, not high speciation rates, and that biotic interchanges with other regions were rare. Southeast Asia was also characterized by a low speciation rate but, unlike the Neotropics, was the main source of dispersal events through time. Our results suggest that global climate change throughout the Cenozoic, particularly during the Eocene-Oligocene transition, combined with the conserved ancestral tropical niches, played a major role in generating the modern LDG of butterflies.

Opsins are the primary light-sensing molecules in animals. Opsins have peak sensitivities to specific wavelengths which allows for color discrimination. The opsin protein family has undergone duplications and losses, dynamically expanding and contracting the number of opsins, throughout invertebrate evolution, but it is unclear what drives this diversity. Light availability, however, appears to play a significant role. Dim environments are associated with low opsin diversity in deep-sea fishes and cave-dwelling animals. Correlations between high opsin diversity and bright environments, however, are tenuous. Insects are a good system to test whether opsin expansion is associated with greater light availability because they are enormously diverse and consequently display large variation in diel activity. To test this, we used 200 insect transcriptomes and examined the patterns of opsin diversity associated with diel-niche. We focused on the butterflies and moths (Lepidoptera) because this group has significant variation in diel-niche, substantial opsin recovery (n=100), and particularly well-curated transcriptomes. We identified opsin duplications using ancestral state reconstruction and examined rates of opsin evolution, and compared them across diel-niches. We find Lepidoptera species active in high light environments have independently expanded their opsins at least 10 times. Opsins from diurnal taxa also evolve faster; 13 amino acids were identified across different opsins that were under diversifying selection. Structural models reveal that four of these amino acids overlap with opsin color-tuning regions. By parsing nocturnal and diurnal switches, we show that light environment can influence gene diversity, selection, and protein structure of opsins in Lepidoptera.

Lepidoptera (butterflies and moths) are one of the most taxonomically diverse insect orders with nearly 160,000 described species. They have been studied extensively for centuries and are found on nearly all continents and in many environments. It is often assumed that adult butterflies are strictly diurnal and adult moths are strictly nocturnal, but there are many exceptions. Despite the broad interest in butterflies and moths, a comprehensive review of diel (day-night) activity has not been conducted. Here, we synthesize existing data on diel activity in Lepidoptera, trace its evolutionary history on a phylogeny, and show where gaps lie in our knowledge. Diurnality was likely the ancestral condition in Lepidoptera, the ancestral heteroneuran was likely nocturnal, and more than 40 transitions to diurnality subsequently occurred. Using species diversity estimates across the order, we predict that roughly 75-85% of Lepidoptera are nocturnal. We also define the three frequently used terms for activity in animals (diurnal, nocturnal, crepuscular), and show that literature on the activity of micro-moths is significantly lacking. Ecological factors leading to nocturnality/diurnality is a compelling area of research and should be the focus of future studies.

The larva of Lacosoma arizonicum Dyar is figured for the first time. This species was previously known to feed on oaks (Quercus), but no specific Quercus species has been reported. We studied the life history of L. arizonicum and confirm four larval host plants from field-collected larvae: Quercus arizonica, Q. emoryi, Q. gambelii, and Q. hypoleucoides. Larvae were successfully reared to pupation in captivity on Q. virginiana (in Florida). A tachinid fly in the genus Lespesia, was reared from a single L. arizonicum larva.

The advent of next-generation sequencing technology has allowed for thecollection of large portions of the genome for phylogenetic analysis. Hybrid enrichment and transcriptomics are two techniques that leverage next-generation sequencing and have shown much promise. However, methods for processing hybrid enrichment data are still limited. We developed a pipeline for anchored hybrid enrichment (AHE) read assembly, orthology determination, contamination screening, and data processing for sequences flanking the target "probe" region. We apply this approach to study the phylogeny of butterflies and moths (Lepidoptera), a megadiverse group of more than 157,000 described species with poorly understood deep-level phylogenetic relationships. We introduce a new, 855 locus AHE kit for Lepidoptera phylogenetics and compare resulting trees to those from transcriptomes. The enrichment kit was designed from existing genomes, transcriptomes, and expressed sequence tags and was used to capture sequence data from 54 species from 23 lepidopteran families. Phylogenies estimated from AHE data were largely congruent with trees generated from transcriptomes, with strong support for relationships at all but the deepest taxonomic levels. We combine AHE and transcriptomic data to generate a new Lepidoptera phylogeny, representing 76 exemplar species in 42 families. The tree provides robust support for many relationships, including those among the seven butterfly families. The addition of AHE data to an existing transcriptomic dataset lowers node support along the Lepidoptera backbone, but firmly places taxa with AHE data on the phylogeny. Combining taxa sequenced for AHE with existing transcriptomes and genomes resulted in a tree with strong support for (Calliduloidea $+$ Gelechioidea $+$ Thyridoidea) $+$ (Papilionoidea $+$ Pyraloidea $+$ Macroheterocera). To examine the efficacy of AHE at a shallow taxonomic level, phylogenetic analyses were also conducted on a sister group representing a more recent divergence, the Saturniidae and Sphingidae. These analyses utilized sequences from the probe region and data flanking it, nearly doubled the size of the dataset; resulting trees supported new phylogenetics relationships, especially within the Saturniidae and Sphingidae (e.g., Hemarina derived in the latter). We hope that our data processing pipeline, hybrid enrichment gene set, and approach of combining AHE data with transcriptomes will be useful for the broader systematics community.

Anthropogenic noise is a widespread and growing form of sensory pollution associated with the expansion of human infrastructure. One specific source of constant and intense noise is that produced by compressors used for the extraction and transportation of natural gas. Terrestrial arthropods play a central role in many ecosystems, and given that numerous species rely upon airborne sounds and substrate‐borne vibrations in their life histories, we predicted that increased background sound levels or the presence of compressor noise would influence their distributions. In the second largest natural gas field in the United States (San Juan Basin, New Mexico, USA), we assessed differences in the abundances of terrestrial arthropod families and community structure as a function of compressor noise and background sound level. Using pitfall traps, we simultaneously sampled five sites adjacent to well pads that possessed operating compressors, and five alternate, quieter well pad sites that lacked compressors, but were otherwise similar. We found a negative association between sites with compressor noise or higher levels of background sound and the abundance of five arthropod families and one genus, a positive relationship between loud sites and the abundance of one family, and no relationship between noise level or compressor presence and abundance for six families and two genera. Despite these changes, we found no evidence of community turnover as a function of background sound level or site type (compressor and noncompressor). Our results indicate that anthropogenic noise differentially affects the abundances of some arthropod families. These preliminary findings point to a need to determine the direct and indirect mechanisms driving these observed responses. Given the diverse and important ecological functions provided by arthropods, changes in abundances could have ecological implications. Therefore, we recommend the consideration of arthropods in the environmental assessment of noise‐producing infrastructure.

Ambulycini are a cosmopolitan tribe of the moth family Sphingidae, comprised of 10 genera, 3 of which are found in tropical Asia, 4 in the Neotropics, 1 in Africa, 1 in the Middle East, and 1 restricted to the islands of New Caledonia. Recent phylogenetic analyses of the tribe have yielded conflicting results, and some have suggested a close relationship of the monobasic New Caledonian genus CompsulyxHolloway, 1979 to the Neotropical ones, despite being found on opposite sides of the Pacific Ocean. Here, we investigate relationships within the tribe using full mitochondrial genomes, mainly derived from dry-pinned museum collections material. Mitogenomic data were obtained for 19 species representing nine of the 10 Ambulycini genera. Phylogenetic trees are in agreement with a tropical Asian origin for the tribe. Furthermore, results indicate that the Neotropical genus Adhemarius Oiticica Filho, 1939 is paraphyletic and support the notion that OrectaRothschild & Jordan 1903 and TrogolegnumRothschild & Jordan, 1903 may need to be synonymized. Finally, in our analysis the Neotropical genera do not collectively form a monophyletic group, due to a clade comprising the New Caledonian genus Compsulyx and the African genus BatocnemaRothschild & Jordan, 1903 being placed as sister to the Neotropical genus ProtambulyxRothschild & Jordan, 1903. This finding implies a complex biogeographic history and suggests the evolution of the tribe involved at least two long-distance dispersal events.

Over 300 million arthropod specimens are housed in North American natural history collections. These collections represent a “vast hidden treasure trove” of biodiversity −95% of the specimen label data have yet to be transcribed for research, and less than 2% of the specimens have been imaged. Specimen labels contain crucial information to determine species distributions over time and are essential for understanding patterns of ecology and evolution, which will help assess the growing biodiversity crisis driven by global change impacts. Specimen images offer indispensable insight and data for analyses of traits, and ecological and phylogenetic patterns of biodiversity. Here, we review North American arthropod collections using two key metrics, specimen holdings and digitization efforts, to assess the potential for collections to provide needed biodiversity data. We include data from 223 arthropod collections in North America, with an emphasis on the United States. Our specific findings are as follows: (1) The majority of North American natural history collections (88%) and specimens (89%) are located in the United States. Canada has comparable holdings to the United States relative to its estimated biodiversity. Mexico has made the furthest progress in terms of digitization, but its specimen holdings should be increased to reflect the estimated higher Mexican arthropod diversity. The proportion of North American collections that has been digitized, and the number of digital records available per species, are both much lower for arthropods when compared to chordates and plants. (2) The National Science Foundation’s decade-long ADBC program (Advancing Digitization of Biological Collections) has been transformational in promoting arthropod digitization. However, even if this program became permanent, at current rates, by the year 2050 only 38% of the existing arthropod specimens would be digitized, and less than 1% would have associated digital images. (3) The number of specimens in collections has increased by approximately 1% per year over the past 30 years. We propose that this rate of increase is insufficient to provide enough data to address biodiversity research needs, and that arthropod collections should aim to triple their rate of new specimen acquisition. (4) The collections we surveyed in the United States vary broadly in a number of indicators. Collectively, there is depth and breadth, with smaller collections providing regional depth and larger collections providing greater global coverage. (5) Increased coordination across museums is needed for digitization efforts to target taxa for research and conservation goals and address long-term data needs. Two key recommendations emerge: collections should significantly increase both their specimen holdings and their digitization efforts to empower continental and global biodiversity data pipelines, and stimulate downstream research.

Significance Lepidoptera play key roles in many biological systems. Butterflies are hypothesized to have evolved contemporaneously with flowering plants, and moths are thought to have gained anti-bat defenses in response to echolocating predatory bats, but these hypotheses have largely gone untested. Using a transcriptomic, dated evolutionary tree of Lepidoptera, we demonstrate that the most recent common ancestor of Lepidoptera is considerably older than previously hypothesized. The oldest moths in crown Lepidoptera were present in the Carboniferous, some 300 million years ago, and began to diversify largely in synchrony with angiosperms. We show that multiple lineages of moths independently evolved hearing organs well before the origin of bats, rejecting the hypothesis that lepidopteran hearing organs arose in response to these predators., Butterflies and moths (Lepidoptera) are one of the major superradiations of insects, comprising nearly 160,000 described extant species. As herbivores, pollinators, and prey, Lepidoptera play a fundamental role in almost every terrestrial ecosystem. Lepidoptera are also indicators of environmental change and serve as models for research on mimicry and genetics. They have been central to the development of coevolutionary hypotheses, such as butterflies with flowering plants and moths’ evolutionary arms race with echolocating bats. However, these hypotheses have not been rigorously tested, because a robust lepidopteran phylogeny and timing of evolutionary novelties are lacking. To address these issues, we inferred a comprehensive phylogeny of Lepidoptera, using the largest dataset assembled for the order (2,098 orthologous protein-coding genes from transcriptomes of 186 species, representing nearly all superfamilies), and dated it with carefully evaluated synapomorphy-based fossils. The oldest members of the Lepidoptera crown group appeared in the Late Carboniferous (∼300 Ma) and fed on nonvascular land plants. Lepidoptera evolved the tube-like proboscis in the Middle Triassic (∼241 Ma), which allowed them to acquire nectar from flowering plants. This morphological innovation, along with other traits, likely promoted the extraordinary diversification of superfamily-level lepidopteran crown groups. The ancestor of butterflies was likely nocturnal, and our results indicate that butterflies became day-flying in the Late Cretaceous (∼98 Ma). Moth hearing organs arose multiple times before the evolutionary arms race between moths and bats, perhaps initially detecting a wide range of sound frequencies before being co-opted to specifically detect bat sonar. Our study provides an essential framework for future comparative studies on butterfly and moth evolution.

Understanding the evolutionary mechanisms governing the uneven distribution of species richness across the tree of life is a great challenge in biology. Scientists have long argued that sexual conflict is a key driver of speciation. This hypothesis, however, has been highly debated in light of empirical evidence. Recent advances in the study of macroevolution make it possible to test this hypothesis with more data and increased accuracy. In the present study, we use phylogenomics combined with four different diversification rate analytical approaches to test whether sexual conflict is a driver of speciation in brush-footed butterflies of the tribe Acraeini. The presence of a sphragis, an external mating plug found in most species among Acraeini, was used as a proxy for sexual conflict. Diversification analyses statistically rejected the hypothesis that sexual conflict is associated with shifts in diversification rates in Acraeini. This result contrasts with earlier studies and suggests that the underlying mechanisms driving diversification are more complex than previously considered. In the case of butterflies, natural history traits acting in concert with abiotic factors possibly play a stronger role in triggering speciation than does sexual conflict. [Acraeini butterflies; arms race; exon capture phylogenomics; Lepidoptera macroevolution; sexual selection; sphragis.]

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.

Insects are possibly the most taxonomically and ecologically diverse class of multicellular organisms on Earth. Consequently, they provide nearly unlimited opportunities to develop and test ecological and evolutionary hypotheses. Currently, however, large-scale studies of insect ecology, behavior, and trait evolution are impeded by the difficulty in obtaining and analyzing data derived from natural history observations of insects. These data are typically highly heterogeneous and widely scattered among many sources, which makes developing robust information systems to aggregate and disseminate them a significant challenge. As a step towards this goal, we report initial results of a new effort to develop a standardized vocabulary and ontology for insect natural history data. In particular, we describe a new database of representative insect natural history data derived from multiple sources (but focused on data from specimens in biological collections), an analysis of the abstract conceptual areas required for a comprehensive ontology of insect natural history data, and a database of use cases and competency questions to guide the development of data systems for insect natural history data. We also discuss data modeling and technology-related challenges that must be overcome to implement robust integration of insect natural history data.

Butterflies and moths exhibit a spectacular diver sity of wing shape and size. The extent of wing variation is particularly evident in wild silk moths (Saturniidae), which have large wing shape and size variation. Some species have jagged wing margins, rounded forewing apical lobes, or narrow hind wings with long tails, while others lack these traits entirely. Surprisingly, very little work has been done to formally quantify wing variation within the fa mily. We analyzed the hind wing shape and size of 76 saturniid species representing 52 genera across five subfamilies using geometric morphometrics. We identified fifteen landmarks that we predict can be applied to families across Lepidoptera. PCA analyses grouped saturniid hind wings into six distinct morphological clusters. These groups did not appear to follow species relatedness—some phylogenetically and genetically distantly related taxa clustered in the same morphological group. We discuss ecological factors that might have led to the extraordinary wing variation within Saturniidae.

Predators can induce extreme stress and profound physiological responses in prey. Insects are the most dominant animal group on Earth and serve as prey for many different predators. Although insects have an extraordinary diversity of anti-predator behavioral and physiological responses, predator-induced stress has not been studied extensively in insects, especially at the molecular level. Here, we review the existing literature on physiological predator-induced stress responses in insects and compare what is known about insect stress to vertebrate stress systems. We conclude that many unrelated insects share a baseline pathway of predator-induced stress responses that we refer to as the octopamine-adipokinetic hormone (OAH) axis. We also present best practices for studying predator-induced stress responses in prey insects. We encourage investigators to compare neurophysiological responses to predator-related stress at the organismal, neurohormonal, tissue, and cellular levels within and across taxonomic groups. Studying stress-response variation between ecological contexts and across taxonomic levels will enable the field to build a holistic understanding of, and distinction between, taxon- and stimulus-specific responses relative to universal stress responses.

Primary biodiversity data records that are open access and available in a standardised format are essential for conservation planning and research on policy-relevant time-scales. We created a dataset to document all known occurrence data for the Federally Endangered Poweshiek skipperling butterfly [Oarismapoweshiek(Parker, 1870; Lepidoptera: Hesperiidae)]. The Poweshiek skipperling was a historically common species in prairie systems across the upper Midwest, United States and Manitoba, Canada. Rapid declines have reduced the number of verified extant sites to six. Aggregating and curating Poweshiek skipperling occurrence records documents and preserves all known distributional data, which can be used to address questions related to Poweshiek skipperling conservation, ecology and biogeography. Over 3500 occurrence records were aggregated over a temporal coverage from 1872 to present. Occurrence records were obtained from 37 data providers in the conservation and natural history collection community using both “HumanObservation” and “PreservedSpecimen” as an acceptable basisOfRecord. Data were obtained in different formats and with differing degrees of quality control. During the data aggregation and cleaning process, we transcribed specimen label data, georeferenced occurrences, adopted a controlled vocabulary, removed duplicates and standardised formatting. We examined the dataset for inconsistencies with known Poweshiek skipperling biogeography and phenology and we verified or removed inconsistencies by working with the original data providers. In total, 12 occurrence records were removed because we identified them to be the western congenerOarismagarita(Reakirt, 1866). This resulting dataset enhances the permanency of Poweshiek skipperling occurrence data in a standardised format.This is a validated and comprehensive dataset of occurrence records for the Poweshiek skipperling (Oarismapoweshiek) utilising both observation and specimen-based records. Occurrence data are preserved and available for continued research and conservation projects using standardised Darwin Core formatting where possible. Prior to this project, much of these occurrence records were not mobilised and were being stored in individual institutional databases, researcher datasets and personal records. This dataset aggregates presence data from state conservation agencies, natural heritage programmes, natural history collections, citizen scientists, researchers and the U.S. Fish & Wildlife Service. The data include opportunistic observations and collections, research vouchers, observations collected for population monitoring and observations collected using standardised research methodologies. The aggregated occurrence records underwent cleaning efforts that improved data interoperablitity, removed transcription errors and verified or removed uncertain data. This dataset enhances available information on the spatiotemporal distribution of this Federally Endangered species. As part of this aggregation process, we discovered and verified Poweshiek skipperling occurrence records from two previously unknown states, Nebraska and Ohio.

This paper provides new taxonomic and biological data on a complex of gracillariid moths in the endemic genus Philodoria Walsingham, 1907 that are associated with Myrsine (Primulaceae) in the Hawaiian Islands, United States. Two new species, Philodoriakauaulaensis Kobayashi, Johns & Kawahara, sp. n. (host: Myrsinelanaiensis, M.lessertiana, and M.sandwicensis) and P.kolea Kobayashi, Johns & Kawahara, sp. n. (host: M.lessertiana) are described. Biological data are provided for two previously described species that also feed on Myrsine: P.auromagnifica Walsingham, 1907 and P.succedanea Walsingham, 1907. For the first time we detail and illustrate genital structures, immature stages, biology, and host plants of P.auromagnifica and P.succedanea. Philodoriakolea, P.auromagnifica, and P.succedanea occur in sympatry on the island of Hawaii (Big Island), but each species differs in behavioral characters: P.kolea utilizes leaves of seedlings and forms a serpentine mine, whereas the latter two utilize leaves of larger plants, and form linear or serpentine to blotch mines. More broadly, leaf mine forms and diagnostic characteristics of the Myrsine-feeding species complex of Philodoria (as currently known) are reviewed and illustrated.

Experimental bat-moth battles reveal that sonar sensing is a driving force in the repeated evolution of silk moth hindwings., Prey transmit sensory illusions to redirect predatory strikes, creating a discrepancy between what a predator perceives and reality. We use the acoustic arms race between bats and moths to investigate the evolution and function of a sensory illusion. The spinning hindwing tails of silk moths (Saturniidae) divert bat attack by reflecting sonar to create a misleading echoic target. We characterized geometric morphometrics of moth hindwings across silk moths, mapped these traits onto a new, robust phylogeny, and found that elaborated hindwing structures have converged on four adaptive shape peaks. To test the mechanism underlying these anti-bat traits, we pit bats against three species of silk moths with experimentally altered hindwings that created a representative gradient of ancestral and extant hindwing shapes. High-speed videography of battles reveals that moths with longer hindwings and tails more successfully divert bat attack. We postulate that sensory illusions are widespread and are underappreciated drivers of diversity across systems.

Background Butterflies (Papilionoidea) are perhaps the most charismatic insect lineage, yet phylogenetic relationships among them remain incompletely studied and controversial. This is especially true for skippers (Hesperiidae), one of the most species-rich and poorly studied butterfly families. Methods To infer a robust phylogenomic hypothesis for Hesperiidae, we sequenced nearly 400 loci using Anchored Hybrid Enrichment and sampled all tribes and more than 120 genera of skippers. Molecular datasets were analyzed using maximum-likelihood, parsimony and coalescent multi-species phylogenetic methods. Results All analyses converged on a novel, robust phylogenetic hypothesis for skippers. Different optimality criteria and methodologies recovered almost identical phylogenetic trees with strong nodal support at nearly all nodes and all taxonomic levels. Our results support Coeliadinae as the sister group to the remaining skippers, the monotypic Euschemoninae as the sister group to all other subfamilies but Coeliadinae, and the monophyly of Eudaminae plus Pyrginae. Within Pyrginae, Celaenorrhinini and Tagiadini are sister groups, the Neotropical firetips, Pyrrhopygini, are sister to all other tribes but Celaenorrhinini and Tagiadini. Achlyodini is recovered as the sister group to Carcharodini, and Erynnini as sister group to Pyrgini. Within the grass skippers (Hesperiinae), there is strong support for the monophyly of Aeromachini plus remaining Hesperiinae. The giant skippers (Agathymus and Megathymus) once classified as a subfamily, are recovered as monophyletic with strong support, but are deeply nested within Hesperiinae. Conclusions Anchored Hybrid Enrichment sequencing resulted in a large amount of data that built the foundation for a new, robust evolutionary tree of skippers. The newly inferred phylogenetic tree resolves long-standing systematic issues and changes our understanding of the skipper tree of life. These resultsenhance understanding of the evolution of one of the most species-rich butterfly families. Electronic supplementary material The online version of this article (10.1186/s12862-018-1216-z) contains supplementary material, which is available to authorized users.

The origins and evolution of Hawaiian biodiversity are a matter of controversy, and the mechanisms of lineage diversification for many organisms on this remote archipelago remain unclear. Here we focus on the poorly known endemic leaf-mining moth genusPhilodoria(Lepidoptera, Gracillariidae), whose species feed on a diversity of Hawaiian plant lineages, many of which are critically endangered. We use anchored hybrid enrichment to assemble the first phylogenomic dataset (507 loci) for any Hawaiian animal taxon. To uncover the timing and pattern of diversification of these moths, we apply two frequently used dating calibration strategies, biogeographic calibrations and secondary calibrations. Island calibrations on their own resulted in much younger and unrealistic dates compared to strategies that relied on secondary calibrations.Philodoriaprobably originated on the now partially sunken islands of Laysan or Lisianski, approximately 21 Ma, and were associated with host plants in the families Ebenaceae, Malvaceae or Primulaceae. Major feeding groups associated with specific host-plant families originated soon after the plants colonized the islands. Allopatric isolation and host shifts, in concert and independently, probably play major roles in the diversification ofPhilodoria. Our dating results indicate thatPhilodoriais among the oldest known Hawaiian arthropod lineages, and that island calibrations alone can lead to unrealistically young dates.

The subfamily Erebinae (Lepidoptera, Erebidae) includes approximately 10,000 species with many still undescribed. It is one of the most diverse clades within the moth superfamily Noctuoidea and encompasses a diversity of ecological habits. Erebine caterpillars feed on a broad range of host plants including several economically important crops. Adults possess a unique array of adaptations for predator defense, including some of the most sensitive hearing organs (tympana) across the Lepidoptera and striking wing coloration to startle visual predators. Despite the relevance of these moths to agriculture and ecological research, a robust phylogenetic framework is lacking. Here we used anchored hybrid enrichment, a relatively new approach in phylogenomics, to resolve relationships among the subfamily. Using the recently developed Lep1 anchored hybrid enrichment probe set, 658 gene fragments with an average length of 320 bp were captured from an exemplar set of 75 erebine species, representing 73 genera and 23 tribes. While the total number of erebine tribes is not firmly established, this represents at least 75% of known tribal level diversity. Anchored hybrid enrichment data were partitioned by locus and by codon position for maximum likelihood phylogenetic analysis and coalescent-based species-tree approaches. Results from our study provided strong nodal support (BP ≥ 95) for nearly all nodes in the partitioned ML tree, solidifying many relationships that were previously uncertain or moderately supported based on morphology or a smaller number of gene fragments. Likelihood analyses confidently resolved the placement of Acantholipini as a sister tribe to Sypnini and all other Erebinae. The remaining tribes were placed in a single, strongly supported clade split into two major subclades. Additionally, 25 tropical species that did not have previous tribal assignments are confidently placed on the phylogeny. Statistical comparisons with Shimodaira-Hasegawa (SH) tests found that our maximum likelihood trees were significantly more likely than alternative hypotheses. This study demonstrates the utility of anchored phylogenomics for resolving relationships within subfamilies of Lepidoptera.

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.

International audience; Background: Bombycoidea is an ecologically diverse and speciose superfamily of Lepidoptera. The superfamily includes many model organisms, but the taxonomy and classification of the superfamily has remained largely in disarray. Here we present a global checklist of Bombycoidea. Following Zwick (2008) and Zwick et al. (2011), ten families are recognized: Anthelidae, Apatelodidae, Bombycidae, Brahmaeidae, Carthaeidae, Endromidae, Eupterotidae, Phiditiidae, Saturniidae and Sphingidae. The former families Lemoniidae and Mirinidae are included within Brahmaeidae and Endromidae respectively. The former bombycid subfamilies Oberthueriinae and Prismostictinae are also treated as synonyms of Endromidae, and the former bombycine subfamilies Apatelodinae and Phitditiinae are treated as families.New information: This checklist represents the first effort to synthesize the current taxonomic treatment of the entire superfamily. It includes 12,159 names and references to their authors, and it accounts for the recent burst in species and subspecies descriptions within family Saturniidae (ca. 1,500 within the past 10 years) and to a lesser extent in Sphingidae (ca. 250 species over the same period). The changes to the higher classification of Saturniidae proposed by Nässig et al. (2015) are rejected as premature and unnecessary. The new tribes, subtribes and genera described by Cooper (2002) are here treated as junior synonyms. We also present a new higher classification of Sphingidae, based on Kawahara et al. (2009), Barber and Kawahara (2013) and a more recent phylogenomic study by Breinholt et al. (2017), as well as a reviewed genus and species level classification, as documented by Kitching (2018).

The Burmese python, Python bivittatus Kuhl, is a well-established invasive species in the greater Everglades ecosystem of southern Florida, USA. Most research on its ecological impacts focuses on its role as a predator and its trophic interactions with native vertebrate species, particularly mammals. Beyond predation, there is little known about the ecological interactions between P. bivittatus and native faunal communities. It is likely that established populations of P. bivittatus in southern Florida serve as hosts for native mosquito communities. To test this concept, we used mitochondrial cytochrome c oxidase subunit I DNA barcoding to determine the hosts of blood fed mosquitoes collected at a research facility in northern Florida where captive P. bivittatus and Argentine black and white tegu, Salvator merianae (Duméril and Bibron), are maintained in outdoor enclosures, accessible to local mosquitoes. We recovered python DNA from the blood meals of three species of Culex mosquitoes: Culex erraticus (Dyar and Knab), Culex quinquefasciatus Say, and Culex pilosus (Dyar and Knab). Culex erraticus conclusively (P = 0.001; Fisher's Exact Test) took more blood meals from P. bivittatus than from any other available host. While the majority of mosquito blood meals in our sample were derived from P. bivittatus, only one was derived from S. merianae. These results demonstrate that local mosquitoes will feed on invasive P. bivittatus, a recently introduced host. If these interactions also occur in southern Florida, P. bivittatus may be involved in the transmission networks of mosquito-vectored pathogens. Our results also illustrate the potential of detecting the presence of P. bivittatus in the field through screening mosquito blood meals for their DNA.

Aim The aim was to determine processes driving the latitudinal diversity gradient (LDG) in New World swallowtail butterfly diversity. We tested three mechanisms commonly invoked to explain the LDG: ecological opportunity, evolutionary rates and biogeographical history. Location New World and Eurasia. Time period Oligocene–Present. Major taxa studied New World swallowtail butterfly clade (Papilio: Agehana, Alexanoria, Chilasa, Heraclides and Pterourus). Methods We integrated data from the most complete current phylogeny of this clade with geographical distributions of each species inferred from ecological niche models (ENMs). We tested for correlation between breadth of available abiotic ecological niche space, latitude and differential rates of diversification between tropical and non-tropical lineages. The clade's history of climatic and geographical occupancy was also reconstructed using both continuous ancestral character reconstructions and biogeographical history inferred under a dispersal–extinction–cladogenesis model. We considered both latitudinal and climatic definitions of tropicality in our reconstructions. Results There was no strong support for ecological opportunity or macroevolutionary processes as latitudinal diversity gradient drivers. Instead, we recovered discordant patterns in phylogenetic reconstructions of latitudinal geographical range and suitable abiotic climate conditions. Heraclides are likely to have originated and diversified in climatically and latitudinally tropical environments before some lineages dispersed to temperate habitats. The Alexanoria + Chilasa + Pterourus clade is likely to have originated in climatically and latitudinally temperate habitat before dispersing and diversifying; some lineages are likely to have dispersed into the latitudinal tropics via highland temperate-analogue environments. Main conclusions The LDG in New World swallowtails results from complex interactions between ecological niche evolution and biogeographical history; both out-of-the-tropics and into-the-tropics processes have contributed to the LDG. Our results present an example where temperate zones appear to be a source, instead of a sink, for biodiversity. Our results emphasize the need to consider biogeographical history not only from the perspective of shifts in geographical space, but also in terms of constraints enforced by ecological niche conservatism.

Summary Butterflies (Papilionoidea), with over 18,000 described species [1], have captivated naturalists and scientists for centuries. They play a central role in the study of speciation, community ecology, biogeography, climate change, and plant-insect interactions and include many model organisms and pest species [2, 3]. However, a robust higher-level phylogenetic framework is lacking. To fill this gap, we inferred a dated phylogeny by analyzing the first phylogenomic dataset, including 352 loci (> 150,000 bp) from 207 species representing 98% of tribes, a 35-fold increase in gene sampling and 3-fold increase in taxon sampling over previous studies [4]. Most data were generated with a new anchored hybrid enrichment (AHE) [5] gene kit (BUTTERFLY1.0) that includes both new and frequently used (e.g., [6]) informative loci, enabling direct comparison and future dataset merging with previous studies. Butterflies originated around 119 million years ago (mya) in the late Cretaceous, but most extant lineages diverged after the Cretaceous-Paleogene (K-Pg) mass-extinction 65 mya. Our analyses support swallowtails (Papilionidae) as sister to all other butterflies, followed by skippers (Hesperiidae) + the nocturnal butterflies (Hedylidae) as sister to the remainder, indicating a secondary reversal from diurnality to nocturnality. The whites (Pieridae) were strongly supported as sister to brush-footed butterflies (Nymphalidae) and blues + metalmarks (Lycaenidae and Riodinidae). Ant association independently evolved once in Lycaenidae and twice in Riodinidae. This study overturns prior notions of the taxon's evolutionary history, as many long-recognized subfamilies and tribes are para- or polyphyletic. It also provides a much-needed backbone for a revised classification of butterflies and for future comparative studies including genome evolution and ecology.

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.

International audience; Numerous hypotheses on the evolution of Neotropical biodiversity have stimulated research to provide a better understandingof diversity dynamics and distribution patterns of the region. However, few studies integrate molecular and morphological datawith complete sampling of a Neotropical group, and so there has been little synthesis of the multiple processes governing biodi-versity through space and time. Here, a total-evidence phylogenetic approach is used to reconstruct the evolutionary history ofthe butterfly subgenusHeraclides. We used DNA sequences for two mitochondrial genes and one nuclear gene and coded 133morphological characters of larvae and adults. A robust and well-resolved phylogeny was obtained using several analyticalapproaches, while molecular dating and biogeographical analyses indicated an early Miocene origin (22 Mya) in the CaribbeanIslands. We inferred six independent dispersal events from the Caribbean to the mainland, and three from the mainland to theCaribbean, and we suggest that cooling climates with decreasing sea levels may have contributed to these events. The time-cali-brated tree is best explained by a museum model of diversity in which both speciation and extinction rates remained constantthrough time. By assessing both continental and fine-scale biodiversity patterns, this study provides new findings, for instancethat islands may act as source of diversity rather than as a sink, to explain spatio-temporal macroevolutionary processes withinthe Neotropical region.

Recent advancements in molecular sequencing techniques have led to a surge in the number of phylogenetic studies that incorporate large amounts of genetic data. We test the assumption that analyzing large number of genes will lead to improvements in tree resolution and branch support using moths in the superfamily Bombycoidea, a group with some interfamilial relationships that have been difficult to resolve. Specifically, we use a next-gen data set that included 19 taxa and 938 genes (∼1.2M bp) to examine how codon position and saturation might influence resolution and node support among three key families. Maximum likelihood, parsimony, and species tree analysis using gene tree parsimony, on different nucleotide and amino acid data sets, resulted in largely congruent topologies with high bootstrap support compared with prior studies that included fewer loci. However, for a few shallow nodes, nucleotide and amino acid data provided high support for conflicting relationships. The third codon position was saturated and phylogenetic analysis of this position alone supported a completely different, potentially misleading sister group relationship. We used the program RADICAL to assess the number of genes needed to fix some of these difficult nodes. One such node originally needed a total of 850 genes but only required 250 when synonymous signal was removed. Our study shows that, in order to effectively use next-gen data to correctly resolve difficult phylogenetic relationships, it is necessary to assess the effects of synonymous substitutions and third codon positions.