Search

Butterflies are a diverse and charismatic insect group that are thought to have evolved with plants and dispersed throughout the world in response to key geological events. However, these hypotheses have not been extensively tested because a comprehensive phylogenetic framework and datasets for butterfly larval hosts and global distributions are lacking. We sequenced 391 genes from nearly 2,300 butterfly species, sampled from 90 countries and 28 specimen collections, to reconstruct a new phylogenomic tree of butterflies representing 92% of all genera. Our phylogeny has strong support for nearly all nodes and demonstrates that at least 36 butterfly tribes require reclassification. Divergence time analyses imply an origin ~100 million years ago for butterflies and indicate that all but one family were present before the K/Pg extinction event. We aggregated larval host datasets and global distribution records and found that butterflies are likely to have first fed on Fabaceae and originated in what is now the Americas. Soon after the Cretaceous Thermal Maximum, butterflies crossed Beringia and diversified in the Palaeotropics. Our results also reveal that most butterfly species are specialists that feed on only one larval host plant family. However, generalist butterflies that consume two or more plant families usually feed on closely related plants., Funding came from the US National Science Foundation (NSF) GoLife ‘ButterflyNet’ collaborative grant (DEB-1541500, 1541557, 1541560) to A.Y.K., R.P.G., D.J.L. and N.E.P. Specimen collection and preservation was funded by NSF DBI-1349345, 1601369, DEB-1557007 and IOS-1920895 (A.Y.K.), NSF DEB-1120380 (D.J.L.), grants 9285-13 and WW-227R-17 from the National Geographic Society (D.J.L.), NSF DBI-1256742 (A.Y.K. and K.R.W.), NSF DEB-0639861 (K.R.W.) and NSF SES-0750480, DEB-0447244 and DEB-9615760 (N.E.P.). M.E. was supported by the Research Council of Norway (no. 204308) and the Hintelmann Scientific Award for Zoological Systematics. F.L.C. was supported by the European Research Council under the European Union’s Horizon 2020 research and innovation programme (project GAIA, no. 851188). M.P.B. was supported by the Swedish Research Council (IPG no. 2020‐06422). R.V. was supported by the Spanish Ministry of Science and Innovation grant PID2019-107078GB-I00/AEI/10.13039/501100011033. G.T. was supported by the Spanish Ministry of Science and Innovation (grants PID2020-117739GA-I00/AEI/10.13039/501100011033 and RYC2018-025335-I). V.L. was supported by the Russian Science Foundation (grant 19-14-00202) and by the Ministry of Science and Higher Education of the Russian Federation (grant 075-15-2021-1069). M.Y. was supported by MEXT KAKENHI no. 19916010 and JSPS KAKENHI grants 13010131, 23570111, 26440207, 17K07528 and 21H02215. A.B.B.M., H.P.R. and N.O.M. were supported by CNPQ grants proc 563332/2010-7 and 304273/2014-7., Main Results and discussion Methods Data availability Code availability References Acknowledgements Author information Ethics declarations Peer review Additional information Supplementary information Rights and permissions About this article

Temperature is thought to be a key variable explaining global patterns of species richness. However, to investigate this relationship carefully, it is necessary to study clades with broad geographic ranges that are comprised of species inhabiting diverse biomes with well- characterized species ranges. In the present study, we investigate the link between temperature and diversification in the butterfly family Pieridae (sulfurs and whites) by combining Next Generation sequences and published molecular data with fine-grained distribution information. After building the most comprehensive phylogeny of the group, with almost 600 species and all higher taxa (subfamilies, tribes and subtribes), we found strong support for the following relationships within the family: Dismorphiinae + (Coliadinae + (Pseudopontiinae + Pierinae)). With a curated distribution dataset of over 800,000 occurrences, we conducted multiple comparative phylogenetic analyses that provided strong evidence that species in environments with more stable daily temperatures or with cooler maximum temperatures in the warm seasons have higher diversification rates. We also found a positive correlation between speciation and extinction with paleotemperature: as global temperature decreased through geological time, so did diversification rates. Although many studies demonstrate higher diversity in the tropics, we have been able to identify specific climate variables associated with changes in diversification, while also inferring the most robust and well sampled phylogenomic framework for Pieridae to date.

Butterflies are a diverse and charismatic insect group that are thought to have diversified via coevolution with plants and in response to dispersals following key geological events. These hypotheses have been poorly tested at the macroevolutionary scale because a comprehensive phylogenetic framework and datasets on global distributions and larval hosts of butterflies are lacking. We sequenced 391 genes from nearly 2,000 butterfly species to construct a new, phylogenomic tree of butterflies representing 92% of all genera and aggregated global distribution records and larval host datasets. We found that butterflies likely originated in what is now the Americas, ∼100 Ma, shortly before the Cretaceous Thermal Maximum, then crossed Beringia and diversified in the Paleotropics. The ancestor of modern butterflies likely fed on Fabaceae, and most extant families were present before the K/Pg extinction. The majority of butterfly dispersals occurred from the tropics (especially the Neotropics) to temperate zones, largely supporting a “cradle” pattern of diversification. Surprisingly, host breadth changes and shifts to novel host plants had only modest impacts.

A new species, Daplasa medoga sp. nov., is described from Tibet, China. The new species can be distinguished morphologically from all other known Daplasa species by its male genitalia with a sagittate uncus and distally broadly crotched valvae. The validity of new species is well supported by the molecular phylogenetic analyses of one mitochondrial gene (COI) and two nuclear genes (EF-1α & RPS5), with a total length of 2,324 bp.

Recent advances in obtaining reduced representation libraries for next-generation sequencing permit phylogenomic analysis of species-rich, recently diverged taxa. In this study, we performed sequence capture with homemade PCR-generated probes to study diversification among closely related species in a large insect genus to examine the utility of this method. We reconstructed the phylogeny of Neptis Fabricius, a large and poorly studied nymphalid butterfly genus distributed throughout the Old World. We inferred relationships among 108 Neptis samples using 89 loci totaling up to 84 519 bp per specimen. Our taxon sample focused on Palearctic, Oriental and Australasian species, but included 8 African species and outgroups from 5 related genera. Maximum likelihood and Bayesian analyses yielded identical trees with full support for almost all nodes. We confirmed that Neptis is not monophyletic because Lasippa heliodore (Fabricius) and Phaedyma amphion (Linnaeus) are nested within the genus, and we redefine species groups for Neptis found outside of Africa. The statistical support of our results demonstrates that the probe set we employed is useful for inferring phylogenetic relationships among Neptis species and likely has great value for intrageneric phylogenetic reconstruction of Lepidoptera. Based on our results, we revise the following two taxa: Neptis heliodore comb. rev. and Neptis amphion comb. rev. (Less)

Heracula discivitta Moore is an uncommon moth species currently recorded from India, Nepal and China. Although this species has traditionally been placed in Lymantriinae, its systematic position in Macroheterocera has been enigmatic due to its unique morphological features. Here we used molecular and morphological data to explore the systematic position of H. discivitta. Our molecular phylogenetic analyses indicate that this species is sister to Pseudobiston pinratanai Inoue, a member of a recently established monotypic family Pseudobistonidae. The examinations of morphological features further show that H. discivitta shares synapomorphies with Pseudobistonidae. Based on the analysis results, we propose a new subfamily of Pseudobistonidae (Heraculinae subfam.n.) to accommodate H. discivitta. The resemblance of the habitus to that of the brahmaeid genus Calliprogonos Mell & Hering is discussed. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:urn:lsid:zoobank.org:pub:63D17850-6D51-4E03-A5D6-F9EF6E7AF402. (Less)

The tussock moth genus Daplasa Moore, 1879 is reviewed from China. Hitherto four species are recognized, where one new species, D. nivisala sp. n., is described and one new combination D. melanoma (Collenette, 1938) comb. n. is proposed. Adults and their genital structures are provided, together with a key to the species of Daplasa with the occurrence in China.

A new species of the genus Panolis Hübner, [1821], P. xundian sp. nov., is described and illustrated from southwestern China. It is well-defined morphologically by the male genitalia, with a well-developed pollex at the distal terminal of the sacculus and a broad, ventrally concave cucullus, the female corpus bursae with four long signum-stripes. Based on a 658 bp segment of the mitochondrial cytochrome c oxidase subunit I gene, we report the pairwise genetic distance of 2.5% from its allied species P. exquisita Draudt, 1950. Molecular phylogenetic analyses using three genes (2189 bp in total length) indicate that the new species belongs to the P. exquisita species group.

The family Epicopeiidae is a small group of day-flying moths, known for mimicking many different groups of butterflies and moths. So far, there still lacks a reliable phylogenetic framework of Epicopeiidae that is necessary to our understanding of the evolutionary process of their mimicry. In this study, we sequenced 94 nuclear protein-coding markers for 56 epicopeiid samples and 11 outgroups, covering all ten genera of Epicopeiidae. We used homemade PCR-generated baits to capture target sequences, which allowed us to utilize old and dried specimens that were difficult to handle by conventional PCR + Sanger sequencing. Maximum likelihood and Bayesian analyses of the newly obtained dataset (86,388 bp) at both DNA and protein levels produced identical phylogenies with strong support. The non-mimicry genus Deuveia is the sister group of other epicopeiid genera. Epicopeia and Nossa are not monophyletic, and these two genera nest together to form a clade. We also estimated divergence times of Epicopeiidae and found that their initial diversification happened in Eocene about 41 million years ago. The ancestral state reconstruction of mimicry type for this family suggested that the last common ancestor of epicopeiid moths is non-mimetic, and the Riodinidae-mimicry type evolved first. In summary, our work provides a comprehensive and robust time-calibrated phylogeny of Epicopeiidae that provides a sound framework for revising their classification and interpreting character evolution.

A new species of the genus Synersaga Gozmany, Synersaga atriptera Xu & Wang sp. nov. (Lepidoptera: Lecithoceridae) is described from China. Adults, wing venation and male genitalia are illustrated. The new species is similar to S. nigriptera, but is distinguished by the wing venation and the male genitalia. The ground color of wings of S. atriptera sp. nov. is darker than that of the wings of S. nigriptera, especially the hind wings; the forewing of S. atriptera sp. nov. has a broad and blackish transverse outer line, which it is not present in S. nigriptera; and the fringe of S. nigriptera has a paler basal line, which it is not obvious in S. atriptera sp. nov. Juxta of S. atriptera sp. nov. has a pair of claviform lateral lobes, which are separated at the base. Juxta of S. nigriptera also has a pair of claviform lateral lobes, but they are connected at the base by a heavily sclerotized band. Type specimens were deposited in the Insect Collection of the Department of Entomology, South China Agricultural University, Guangzhou, China.

Two new species of the genus Longipenis Wu (Lepidoptera: Lecithoceridae), L. paradeltidius M. Wang and Xiong, sp. nov. and L. dentivalvus H. Wang and M. Wang, sp. nov. are described from South China. Longipenis paradeltidius is very similar to L. deltidius Wu in external morphology, but can be easily distinguished from the latter by M3 free from CuA1 in the hindwing and cornutus as long as 2/3 length of aedeagus. Longipenis dentivalvus differs from the 2 preceding species by uncus not apically forked. Adults, male genitalia, and wing venation are illustrated. A key and a distribution map of the species are given. The type specimens are deposited in the Department of Entomology, South China Agricultural University, Guangzhou, China.

A new species, Lymantria (Nyctria) furvinis H. Wang, Kishida and M. Wang sp. nov., is described from Guangdong Nanling National Nature Reserve, South China. The new species can be distinguished from all previously described species by the basal area on the forewing yellow with black spots, valve with a transverse sclerite connecting a long digitate process medially. Illustrations of the adults, wing venation and genitalia are provided.

To understand the evolutionary history of Lymantriinae and test the present higher-level classification, we performed the first broad-scale molecular phylogenetic analysis of the subfamily, based on 154 exemplars representing all recognized tribes and drawn from all major biogeographical regions. We used two mitochondrial genes (cytochrome c oxidase subunit I and 16S ribosomal RNA) and six nuclear genes (elongation factor-1 alpha, carbamoylphosphate synthase domain protein, ribosomal protein S5, cytosolic malate dehydrogenase, glyceraldehyde-3-phosphate dehydrogenase and wingless). Data matrices (in total 5424 bp) were analysed by parsimony and model-based evolutionary methods (maximum likelihood and Bayesian inference). Based on the results of the analyses, we present a new phylogenetic classification for Lymantriinae composed of seven well-supported tribes, two of which are proposed here as new: Arctornithini, Leucomini, Lymantriini, Orgyiini, Nygmiini, Daplasini trib. nov. and Locharnini trib. nov. We discuss the internal structure of each of these tribes and address some of the more complex problems with the genus-level classification, particularly within Orgyiini and Nygmiini.

The genus Panolis is a small group of noctuid moths with six recognized species distributed from Europe to East Asia, and best known for containing the widespread Palearctic pest species P. flammea, the pine beauty moth. However, a reliable classification and robust phylogenetic framework for this group of potentially economic importance are currently lacking. Here, we use morphological and molecular data (mitochondrial genes cytochrome c oxidase subunit I and 16S ribosomal RNA, nuclear gene elongation factor-1 alpha) to reconstruct the phylogeny of this genus, with a comprehensive systematic revision of all recognized species and a new one, P. ningshan sp. nov. The analysis results of maximum parsimony, maximum likelihood and Bayesian inferring methods for the combined morphological and molecular data sets are highly congruent, resulting in a robust phylogeny and identification of two clear species groups, i.e., the P. flammea species group and the P. exquisita species group. We also estimate the divergence times of Panolis moths using two conventional mutation rates for the arthropod mitochondrial COI gene with a comparison of two molecular clock models, as well as reconstruct their ancestral areas. Our results suggest that 1) Panolis is a young clade, originating from the Oriental region in China in the Late Miocene (6–10Mya), with an ancestral species in the P. flammea group extending northward to the Palearctic region some 3–6 Mya; 2) there is a clear possibility for a representative of the Palearctic clade to become established as an invasive species in the Nearctic taiga.

Thirteen species belonging to seven genera of the tribe Nygmiini are recognized from Nanling National Nature Reserve, Guangdong province, China. A new species, Arna bicostata sp. nov., is described, and two species are transferred as new combinations: Artaxa angulata comb. nov. and Toxoproctis croceola comb. nov.. Adults and their genitalia are illustrated.

Hainan Island, with an area of about 33920 km2 located at the northern margin of tropical Asia, (Zhang 2004), has exceptional high species richness and is considered a hotspot region for biodiversity research (Myers et al. 2000). On 11 Apr 2009, we conducted a survey of Lepidoptera in Jianfengling National Nature Reserve, south west of Hainan Island, as a part of Lepidoptera inventory initiated several years ago (Wang & Huang 2005). Two male lymantrid specimens of Nygmiini collected by light traping were con firmed as Euproctis wilemani based on the male genitalia compared with the literature. The spe cies was first described by Collenette in 1929 and previously recorded only from regions of the Ma lay Archipelago including The Philippines, Borneo, and Sumatra (Holloway 1999). We report here the occurrence of E. wilemani from Hainan Island as well as China for the first time and de scribe morphological characters. Male adult (Figs. 1 and 2): Wingspan 22 mm. Head covered with yellow scales. Antenna bipec tinate, pale yellow. Labial palpus short, upturned, thickened with somewhat rough yellow scales. Thorax and tegula yellow. Forewing evenly yel low, fringe white; venation with R2 branching off more distally than R5, R5 stalked with R2+3+4, M1 arising from upper angle of distal cell, M3 from under angle of distal cell, M2, M3 and CuAx iso lated. Hindwing yellowish white, fringe white; ve nation with Rs and Ml stalked at basal 1/3, M2 and M3+CuA1 arising from under angle of distal cell, M3 stalked with CuAj at 1/5 basally. Abdomen dark yellow. Male genitalia (Fig. 3): Uncus bifid at base. Valves square, cucullus slightly convex medially. Saccus strong, V-shaped. Juxta deeply convex on anterior margin at middle. Aedeagus wide, longer than valves in length; a single, heavily sclerotized cornutus present in the vesica, broad basally, acute apically. The present discovery greatly extends the dis tribution range of E. wilemani from the Malay Ar chipelago, which is mainly a tropical rainforest cli mate, to Hainan Island, which has a tropical mon soon climate. However, Hainan Island has close as sociation and shares a common origin with Malay Archipelago in some insect species (Zhang 2004; Huang 2002). Euproctis wilemani from Hainan has no distinct difference from those of Borneo and Sumatra, except the Hainan specimens are Figs. 1 and 2. Adult and venation of Euproctis wile mani Collenette, 1929, 1. Male adult, upperside. 2. Ve nation, male.