Comprehensive phylogeny of Pieridae butterflies reveals strong correlation between diversification and temperature.

Temperature is thought to be a key factor influencing global species richness patterns. We investigate the link between temperature and diversification in the butterfly family Pieridae by combining next generation DNA sequences and published molecular data with fine-grained distribution data. We sampled nearly 600 pierid butterfly species to infer the most comprehensive molecular phylogeny of the family and curated a distribution dataset of more than 800,000 occurrences. We found strong evidence that species in environments with more stable daily temperatures or cooler maximum temperatures in the warm seasons have higher speciation rates. Furthermore, speciation and extinction rates decreased in tandem with global temperatures through geological time, resulting in a constant net diversification.

Patterns of speciation in a parapatric pair of Saturnia moths as revealed by target capture.

The focus of this study has been to understand the evolutionary relationships and taxonomy of a widely distributed parapatric species pair of wild silk moths in Europe: Saturnia pavonia and Saturnia pavoniella (Lepidoptera: Saturniidae). To address species delimitation in these parapatric taxa, target enrichment and mtDNA sequencing was employed alongside phylogenetic, admixture, introgression, and species delimitation analyses. The dataset included individuals from both species close to and farther away from the contact zone as well as two hybrids generated in the lab. Nuclear markers strongly supported both S. pavonia and S. pavoniella as two distinct species, with hybrids forming a sister group to S. pavoniella. However, the Maximum Likelihood (ML) tree generated from mtDNA sequencing data presented a different picture, showing both taxa to be phylogenetically intermixed. This inconsistency is likely attributable to mitonuclear discordance, which can arise from biological factors (e.g., introgressive hybridization and/or incomplete lineage sorting). Our analyses indicate that past introgressions have taken place, but that there is no evidence to suggest an ongoing admixture between the two species, demonstrating that the taxa have reached full postzygotic reproductive isolation and hence represent two distinct biological species. Finally, we discuss our results from an evolutionary point of view taking into consideration the past climatic oscillations that have likely shaped the present dynamics between the two species. Overall, our study demonstrates the effectiveness of the target enrichment approach in resolving shallow phylogenetic relationships under complex evolutionary circumstances and that this approach is useful in establishing robust and well-informed taxonomic delimitations involving parapatric taxa.

A new species of the genus Eterusia Hope from western China (Lepidoptera, Zygaenidae, Chalcosiinae).

A new species Eterusia sinoraja Huang & Horie sp. n. is described from western Sichuan Province, western China. The comparison is made with the reminiscent E. raja Moore, 1859 from northeastern India. The adult and genitalia of both sexes of the aforementioned taxa are illustrated.

A review of the genus Agalope Walker (Lepidoptera, Zygaenidae, Chalcosiinae) with taxonomic notes and descriptions of three new genera and three new species.

A preliminary review of the genus Agalope Walker, 1854 is presented. Two new genera are established for four species-groups: Rotundagalope S.-Y. Huang & Horie, gen. n. (type species Agalope immaculata Leech, 1898, for the immaculata species-group), Paragalope S.-Y. Huang & Horie, gen. n. (type species Chelura pica Wileman, 1910, for the pica, glacialis and dejeani species-groups). An additional new genus, Agacysma S.-Y. Huang & Horie, gen. n., related to Agalope and Elcysma, is erected for the new species Agacysma sinica S.-Y. Huang & Horie sp. n. (mainland China: Chongqing, Hubei & Shaanxi). Two new species of the genus Agalope are described: A. geoffi S.-Y. Huang & Horie sp. n. (mainland China: SE. Xizang) and A. liuzihaoi S.-Y. Huang & Horie sp. n. (mainland China: SE. Xizang), forming a species-group of their own which is clearly different from congeners in their male genitalia. The taxonomic issues between Paragalope haoi (S.-Y. Huang, 2022) comb. n. and P. bieti (Oberthür, 1886) comb. n. are discussed. Moreover, following the erection of Paragalope, the homonymy of Agalope glacialis Butler, 1881 and A. glacialis (Moore, 1872) ends; hence the replacement name of the former, A. butleri Owada & Horie, 2000 syn. n., is unnecessary and hence synonymized. The following new combinations are given: Rotundagalope immaculata (Leech, 1898) comb. nov., Paragalope ardjuna (Roepke, 1936) comb. nov., P. aurelia (Oberthür, 1923) comb. nov., P. basiflava (Moore, 1879b) comb. nov., P. bieti (Oberthür, 1886) comb. nov., P. chayuensis (S.-Y. Huang & Pan, 2022) comb. nov., P. haoi (S.-Y. Huang, 2022) comb. nov., P. owadai (S.-Y. Huang, 2022) comb. nov., P. pica (Wileman, 1910) comb. nov., P. trimacula (Matsumura, 1927) comb. nov., P. wangi (Owada, 1992) comb. nov., P. dejeani (Oberthür, 1893) comb. nov., P. lucia (Oberthür, 1923) comb. nov., P. glacialis glacialis (Moore, 1872) comb. nov., P. glacialis parthenie (Jordan, 1907b) comb. nov., P. glacialis angustifasciata (Hering, 1922) comb. nov., P. glacialis postfasciata (Holloway, 2011) comb. nov. A checklist of the species and genera mentioned in the present study is given. Adults and genitalia of the newly described taxa and related ones are illustrated.

Rapid radiation of ant parasitic butterflies during the Miocene aridification of Africa.

Africa has undergone a progressive aridification during the last 20 My that presumably impacted organisms and fostered the evolution of life history adaptations. We test the hypothesis that shift to living in ant nests and feeding on ant brood by larvae of phyto-predaceous Lepidochrysops butterflies was an adaptive response to the aridification of Africa that facilitated the subsequent radiation of butterflies in this genus. Using anchored hybrid enrichment we constructed a time-calibrated phylogeny for Lepidochrysops and its closest, non-parasitic relatives in the Euchrysops section (Poloyommatini). We estimated ancestral areas across the phylogeny with process-based biogeographical models and diversification rates relying on time-variable and clade-heterogeneous birth-death models. The Euchrysops section originated with the emerging Miombo woodlands about 22 million years ago (Mya) and spread to drier biomes as they became available in the late Miocene. The diversification of the non-parasitic lineages decreased as aridification intensified around 10 Mya, culminating in diversity decline. In contrast, the diversification of the phyto-predaceous Lepidochrysops lineage proceeded rapidly from about 6.5 Mya when this unusual life history likely first evolved. The Miombo woodlands were the cradle for diversification of the Euchrysops section, and our findings are consistent with the hypothesis that aridification during the Miocene selected for a phyto-predaceous life history in species of Lepidochrysops, with ant nests likely providing caterpillars a safe refuge from fire and a source of food when vegetation was scarce.

Metabarcoding dietary analysis in the insectivorous bat Nyctalusleisleri and implications for conservation.

In this study, we aim to uncover diet preferences for the insectivorous bat Nyctalusleisleri (Leisler's bat, the lesser noctule) and to provide recommendations for conservation of the species, based on the analysis of prey source habitats. Using a novel guano trap, we sampled bat faeces at selected roosts in a forest in Germany and tested two mitochondrial markers (COI and 16S) and three primer pairs for the metabarcoding of bat faecal pellets. We found a total of 17 arthropod prey orders comprising 358 species in N.leisleri guano. The most diverse orders were Lepidoptera (126 species), Diptera (86 species) and Coleoptera (48 species), followed by Hemiptera (28 species), Trichoptera (16 species), Neuroptera (15 species) and Ephemeroptera (10 species), with Lepidoptera species dominating in spring and Diptera in summer. Based on the ecological requirements of the most abundant arthropod species found in the bat guano, we propose some recommendations for the conservation of N.leisleri that are relevant for other insectivorous bat species.

DNA barcoding unveils a high diversity of caddisflies (Trichoptera) in the Mount Halimun Salak National Park (West Java; Indonesia).

Background: Trichoptera are one of the most diverse groups of freshwater insects worldwide and one of the main bioindicators for freshwater quality. However, in many areas, caddisflies remain understudied due to lack of taxonomic expertise. Meanwhile, globally increasing anthropogenic stress on freshwater streams also threatens Trichoptera diversity. Methods: To assess the Trichoptera diversity of the area within and around the Mount Halimun Salak National Park (MHSNP or Taman Nasional Gunung Halimun Salak) in West Java (Indonesia), we conducted a molecular-morphological study on Trichoptera diversity using larvae from a benthic survey and adults from hand-netting. In addition to morphological identification, we applied four different molecular taxon delimitation approaches (Generalized Mixed Yule Coalescent, Bayesian Poisson Tree Processes, Automatic Barcode Gap Discovery and Assemble Species by Automatic Partitioning) based on DNA barcoding of Cytochrome-C-Oxidase I (COI). Results: The molecular delimitation detected 72 to 81 Operational Taxonomic Units (OTU). Only five OTUs could be identified to species level by comparing sequences against the BOLD database using BLAST, and four more to the genus level. Adults and larvae could be successfully associated in 18 cases across six families. The high diversity of Trichoptera in this area highlights their potential as bioindicators for water quality assessment. Conclusions: This study provides an example of how molecular approaches can benefit the exploration of hidden diversity in unexplored areas and can be a valuable tool to link life stages. However, our study also highlights the need to improve DNA barcode reference libraries of Trichoptera for the Oriental region.

How butterfly wings got their pattern.

Gene regulatory elements play a crucial role in the pattern formation of butterfly wings.

Contribution to the knowledge of the genus Agalope Walker from mainland China, with descriptions of four new species (Lepidoptera, Zygaenidae, Chalcosiinae).

New information of the genus Agalope Walker, 1854 from mainland China is presented. Three new species of the Agalope pica species-group are described: A. haoi S.-Y. Huang sp. n. from Weixi County, Yunnan, A. chayuensis S.-Y. Huang Pan sp. n. from Chayu County, Southeastern Xizang and A. owadai S.-Y. Huang sp. n. from Bomi and Jiali Counties, Southeastern and Eastern Xizang. The little-known Agalope aurelia Oberthr, 1923 and A. lucia Oberthr, 1923 were rediscovered, with the male of the latter reported for the first time. Based on the newly discovered male, Agalope lucia is found to be a close relative of A. dejeani, hence it is excluded from the A. pica species-group and transferred to the A. bieti species-group. A new species of the Agalope hyalina species-group is also described: A. jianqingi S.-Y. Huang sp. n. from Pianma, Western Yunnan. Adults and genitalia of the aforementioned and related taxa are illustrated. An updated checklist of the genus is also provided.

Resolving two centuries of mistaken identity: Reinterpretation of Papilio marcus Fabricius, 1787 (Lepidoptera: Nymphalidae, Hesperiidae).

Here, we untangle an oversight surrounding the application of the name Papilio marcus Fabricius, 1787 (Hesperiidae: Hesperiinae), currently in Troyus Warren & Turland, 2012, which has eluded taxonomists for nearly two centuries. First, we note that P. marcus is a junior primary homonym of Papilio marcus Schaller, 1785, a species currently in Morpho Fabricius, 1807 (Nymphalidae: Satyrinae). Second, we designate a lectotype for P. marcus Fabricius, 1787, and recognize it as conspecific with Papilio phyllus Cramer, 1777, a species currently in Vettius Godman, 1901. Therefore, P. marcus is also a junior subjective synonym of V. phyllus (new synonym). Third, aided by genomic sequencing of the lectotype of Vettius phyllides Röber, 1925, we find that this species is not conspecific with V. phyllus and represents instead a valid species of Troyus Warren & Turland, 2012, so that the relative epithet, currently considered as a junior subjective synonym of P. marcus, has to be reappreciated as Troyus phyllides (reinstated status, new combination). Moreover, T. phyllides is apparently the species that has been misidentified as P. marcus since 1832. As a result of this study, the name P. marcus Fabricius nec Cramer falls in synonymy with P. phyllus (currently in Vettius), and the species currently known as T. marcus becomes Troyus phyllides.

A new euptychiine butterfly species from south Brazil and taxonomic rearrangements for Taydebis Freitas, 2013 and Hermeuptychia Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae).

A new species of Taydebis Freitas, 2003 from south Brazil is described using comparative morphology and species distributions. Also, based on morphology, we transfer Neonympha melobosis Capronnier, 1874 (formerly placed in Paryphthimoides Forster, 1964) to Taydebis, and recognize Euptychia peculiaris Butler, 1874 as its junior synonym (syn. nov.). Furthermore, the monotypic Prenda Freitas Mielke, 2011 is herein treated as junior synonym of Taydebis based on morphology, molecular and ecological evidence. Species of Taydebis are endemic and restricted to south Brazil, and now comprises three species: Taydebis guria Zacca, Casagrande Mielke sp. nov., T. melobosis comb. nov. and T. clarissa Freitas Mielke comb. nov. To continue clarifying Euptychiina taxonomy, Euptychia undulata Butler, 1867 (also formerly placed in Paryphthimoides) is transferred to Hermeuptychia Forster, 1964, and we provide additional information on its taxonomy, morphology, and distribution. Diagnoses, illustrations, and distributional maps are provided for all taxa except T. clarissa comb. nov.

Is it time to describe new species without diagnoses?A comment on Sharkey et al. (2021).

New methods in taxonomy and systematics can influence the overall practice of formally naming and describing biodiversity. DNA barcoding has been controversial since its emergence, but now, large scale species descriptions exclusively based on barcodes have created what can be called a 'new quality of performance. Its limitations are discussed from different perspectives: nomenclature, general pragmatism, and problems of DNA-based species delimitation in the light of the central aim of achieving a robust and stable nomenclature of organisms, essential for all applications of biodiversity research. This issue needs to be addressed to prevent restraining the progress of taxonomy and its ability to contribute to modern science.

The Genome Assembly and Annotation of the Apollo Butterfly Parnassius apollo, a Flagship Species for Conservation Biology.

Conservation genomics has made dramatic improvements over the past decade, leveraging the power of genomes to infer diverse parameters central to conservation management questions. However, much of this effort has focused upon vertebrate species, despite insects providing similar flagship status with the added benefit of smaller genomes, shorter generation times and extensive historical collections in museums. Here we present the genome of the Apollo butterfly (Parnassius apollo, Papilionidae), an iconic endangered butterfly, which like many species in this genus, needs conservation genomic attention yet lacks a genome. Using 68.7 Gb of long-read data (N50 = 15.2 kb) we assembled a 1.4 Gb genome for the Apollo butterfly, making this the largest sequenced Lepidopteran genome to date. The assembly was highly contiguous (N50 = 7.1 Mb) and complete (97% of Lepidopteran BUSCOs were single-copy and complete) and consisted of 1,707 contigs. Using RNAseq data and Arthropoda proteins, we annotated 28.3K genes. Alignment with the closest-related chromosome-level assembly, Papilio bianor, reveals a highly conserved chromosomal organization, albeit genome size is highly expanded in the Apollo butterfly, due primarily to a dramatic increase in repetitive element content. Using this alignment for superscaffolding places the P. apollo genome in to 31 chromosomal scaffolds, and together with our functional annotation, provides an essential resource for advancing conservation genomics in a flagship species for insect conservation.

A New Species of Butterfly of the Genus Protopedaliodes from the Isolated Tramen Tepui in the Guyana Shield (Lepidoptera, Nymphalidae, Satyrinae).

A new species of Protopedaliodes Viloria & Pyrcz, a satyrine butterfly genus endemic to the highest part of the Guyana Shield, P. arekuna Pyrcz & Stachowicz n. sp., is described from the summit area, ca. 2400 m, of Tramen Tepui, an isolated mountain situated on the Venezuela-Guyana border. It is a remarkable finding as it is probably a narrow endemic, and only the fourth known member of the genus. Morphologically it most closely resembles P. kukenani Viloria & Pyrcz from the Roraima-Kukenán twin peaks. COI barcode analysis shows, however, high genetic distances between these two species, 9-10%. Protopedaliodes taxonomy is briefly revised, from the perspective of the monophyly of the genus based on preliminary molecular and morphological comparative data, including the female genitalia described for the first time for P. kukenani and P. ridouti Viloria & Pyrcz.

The roles of wing color pattern and geography in the evolution of Neotropical Preponini butterflies.

Diversification rates and evolutionary trajectories are known to be influenced by phenotypic traits and the geographic history of the landscapes that organisms inhabit. One of the most conspicuous traits in butterflies is their wing color pattern, which has been shown to be important in speciation. The evolution of many taxa in the Neotropics has also been influenced by major geological events. Using a dated, species-level molecular phylogenetic hypothesis for Preponini, a colorful Neotropical butterfly tribe, we evaluated whether diversification rates were constant or varied through time, and how they were influenced by color pattern evolution and biogeographical events. We found that Preponini originated approximately 28 million years ago and that diversification has increased through time consistent with major periods of Andean uplift. Even though some clades show evolutionarily rapid transitions in coloration, contrary to our expectations, these shifts were not correlated with shifts in diversification. Involvement in mimicry with other butterfly groups might explain the rapid changes in dorsal color patterns in this tribe, but such changes have not increased species diversification in this group. However, we found evidence for an influence of major Miocene and Pliocene geological events on the tribe's evolution. Preponini apparently originated within South America, and range evolution has since been dynamic, congruent with Andean geologic activity, closure of the Panama Isthmus, and Miocene climate variability.

Description of a new genus and species for a common and widespread Amazonian satyrine butterfly (Lepidoptera: Nymphalidae: Satyrinae: Satyrini).

We here propose a new monotypic butterfly genus Scriptor Nakahara & Espeland, n. gen. to accommodate a new species, S. sphenophorus Lamas & Nakahara, n. sp., described and named herein. Scriptor sphenophorus n. gen. and n. sp.is a relatively common and widespread butterfly species which is recovered as a member of the so-called "Splendeuptychia clade" in the nymphalid subtribe Euptychiina, based on our molecular phylogenetic analysis using a maximum likelihood approach. Nevertheless, its sister group is not confidently resolved in any analysis, supporting a relatively distant relationship to any described genus as well as our decision to establish a new monotypic genus. We further discuss the proposed taxonomy in the light of frequent criticism of the description of monotypic taxa, as well as emphasize the importance of incorporating multiple evidence when describing new genera, illustrated by reference to several recent generic descriptions in this subtribe.

Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman amp; Salvin, 1889 (Lepidoptera: Nymphalidae: Satyrinae).

Vareuptychia Forster, 1964 stat. rest. is revalidated and comprises two species, V. similis (Butler, 1867) comb. rest. and V. themis (Butler, 1867) comb. nov. Vanima Zacca, Casagrande Mielke gen. nov. is described to contain Euptychia labe Butler, 1870 (the type species), E. palladia Butler, 1867 and E. lesbia Staudinger, [1886]. The taxonomy of these two genera was initially revised based on morphological and distributional data, and subsequently tested and supported with a Maximum Likelihood analysis using four genes (COI, GAPDH, RpS5 and EF1-a). Lectotypes are designated for Euptychia similis Butler, 1867, E. themis Butler, 1867, E. undina Butler, 1870 and E. lesbia Staudinger, [1886]. No DNA sequences were obtained for Euptychia cleophes Godman Salvin, 1889 but its transfer to Megisto Hübner, [1819] is supported by morphological evidence. For all taxa treated in this study, a taxonomic catalog, diagnosis, (re)description and illustrations of adults, venation and genitalia are provided, as well as comments on intraspecific variation, sexual dichromatism, ecology and distribution maps.

Description of 47 new species of the New Caledonian endemic caddisfly genus Agmina Ward & Schefter (Trichoptera, Ecnomidae).

New Caledonia has a rich Trichoptera fauna with over 200 known species, most of them endemic. The total diversity has been estimated as high as 300 to 600 species. The endemic genus Agmina Ward & Schefter (Ecnomidae, Trichoptera) includes 28 described species. Based on male genitalia morphology and previously published molecular data another 47 new species in the genus are described, namely Agmina tuberosa sp. nov., A. semiovale sp. nov., A. rocheta sp. nov., A. tenuisa sp. nov., A. multidentata sp. nov., A. cornuta sp. nov., A. sagittata sp. nov., A. circulata sp. nov., A. digitata sp. nov., A. longispina sp. nov., A. magnahamata sp. nov., A. longicordata sp. nov., A. campanula sp. nov., A. semicampanula sp. nov., A. cunicula sp. nov., A. cerritula sp. nov., A. monstrosa sp. nov., A. rectangulata sp. nov., A. chela sp. nov., A. piscaria sp. nov., A. amplexa sp. nov., A. caraffa, sp. nov., A. rostrata sp. nov., A. dathioensis sp. nov., A. rougensis sp. nov., A. viklundi sp. nov., A. lata sp. nov., A. falx sp. nov., A. guttula sp. nov., A. amieuensis sp. nov., A. spina sp. nov., A. complexa sp. nov., A. dognyensis sp. nov., A. mana sp. nov., A. anterohamata sp. nov., A. curvatacua sp. nov., A. recurvata sp. nov., A. taoensis sp. nov., A. triangulata sp. nov., A. bleuensis sp. nov., A. touhoensis sp. nov., A. wardi sp. nov., A. parallela sp. nov., A. christinae sp. nov., A. brevis sp. nov., A. ninguana sp. nov., and A. scopula sp. nov. Additionally, new records are provide for the species A. acula Ward, 2003, A. artarima Ward & Schefter, 2000, A. berada Ward & Schefter, 2000, A. bimaculata Ward & Schefter, 2000, A. cheirella Ward, 2003, A. comata Ward, 2003, A. diriwi Ward & Schefter, 2000, A. hamata Ward & Schefter, 2000, A. hastata Ward & Schefter, 2000, A. hirta Ward & Schefter, 2000, A. jepiva Ward & Schefter, 2000, A. joycei Ward & Schefter, 2000, A. kapiwa Ward & Schefter, 2000, A. kara Ward & Schefter, 2000, A. mariae Ward & Schefter, 2000, A. nodosa Ward, 2003, A. panda Ward & Schefter, 2000, A. padi Ward & Schefter, 2000, A. parie Ward & Schefter, 2000, A. rhara Ward & Schefter, 2000, A. urugi Ward & Schefter, 2000, and A. vuegi Ward & Schefter, 2000. With a total of 75 described species Agmina is one of the largest animal radiations in New Caledonia. Nothing is known about the early stages of any of the species in this genus.

Description of a new Peronia species (Gastropoda: Eupulmonata: Onchidiidae) from Iran, Persian Gulf.

Peronia J. Fleming, 1822 is an eupulmonate slug genus with a wide distribution in the Indo-Pacific Ocean. Currently, nine species are considered as valid. However, molecular data indicate cryptic speciation and more species involved. Here, we present results on a new species found in the Persian Gulf, a subtropical region with harsh conditions such as elevated salinity and high temperature compared to the Indian Ocean. Peronia persiae sp. nov. is described based on molecular, histological, anatomical, micro-computer tomography and scanning electron microscopy data. ABGD, GMYC and bPTP analyses based on 16S rDNA and cytochrome oxidase I (COI) sequences of Peronia confirm the delimitation of the new species. Moreover, our 14 specimens were carefully compared with available information of other described Peronia species. Peronia persiae sp. nov. is distinct in a combination of characters, including differences in the genital (ampulla, prostate, penial hooks, penial needle) and digestive systems (lack of pharyngeal wall teeth, tooth shape in radula, intestine of type II).

Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths.

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.