Hybrid speciation driven by multilocus introgression of ecological traits.

Hybridization allows adaptations to be shared among lineages and may trigger the evolution of new species1,2. However, convincing examples of homoploid hybrid speciation remain rare because it is challenging to demonstrate that hybridization was crucial in generating reproductive isolation3. Here we combine population genomic analysis with quantitative trait locus mapping of species-specific traits to examine a case of hybrid speciation in Heliconius butterflies. We show that Heliconius elevatus is a hybrid species that is sympatric with both parents and has persisted as an independently evolving lineage for at least 180,000 years. This is despite pervasive and ongoing gene flow with one parent, Heliconius pardalinus, which homogenizes 99% of their genomes. The remaining 1% introgressed from the other parent, Heliconius melpomene, and is scattered widely across the H. elevatus genome in islands of divergence from H. pardalinus. These islands contain multiple traits that are under disruptive selection, including colour pattern, wing shape, host plant preference, sex pheromones and mate choice. Collectively, these traits place H. elevatus on its own adaptive peak and permit coexistence with both parents. Our results show that speciation was driven by introgression of ecological traits, and that speciation with gene flow is possible with a multilocus genetic architecture.

Convergent morphology and divergent phenology promote the coexistence of Morpho butterfly species.

The coexistence of closely-related species in sympatry is puzzling because ecological niche proximity imposes strong competition and reproductive interference. A striking example is the widespread wing pattern convergence of several blue-banded Morpho butterfly species with overlapping ranges of distribution. Here we perform a series of field experiments using flying Morpho dummies placed in a natural habitat. We show that similarity in wing colour pattern indeed leads to interspecific territoriality and courtship among sympatric species. In spite of such behavioural interference, demographic inference from genomic data shows that sympatric closely-related Morpho species are genetically isolated. Mark-recapture experiments in the two most closely-related species unravel a strong temporal segregation in patrolling activity of males. Such divergence in phenology reduces the costs of reproductive interference while simultaneously preserving the benefits of convergence in non-reproductive traits in response to common ecological pressures. Henceforth, the evolution of multiple traits may favour species diversification in sympatry by partitioning niche in different dimensions.

Genomics Reveal Admixture and Unexpected Patterns of Diversity in a Parapatric Pair of Butterflies.

We studied the evolutionary relationship of two widely distributed parapatric butterfly species, Melitaea athalia and Melitaea celadussa, using the ddRAD sequencing approach, as well as genital morphology and mtDNA data. M. athalia was retrieved as paraphyletic with respect to M. celadussa. Several cases of mito-nuclear discordance and morpho-genetic mismatch were found in the contact zone. A strongly diverged and marginally sympatric clade of M. athalia from the Balkans was revealed. An in-depth analysis of genomic structure detected high levels of admixture between M. athalia and M. celadussa at the contact zone, though not reaching the Balkan clade. The demographic modelling of populations supported the intermediate genetic make-up of European M. athalia populations with regards to M. celadussa and the Balkan clade. However, the dissimilarity matrix of genotype data (PCoA) suggested the Balkan lineage having a genetic component that is unrelated to the athalia-celadussa group. Although narrowly sympatric, almost no signs of gene flow were found between the main M. athalia group and the Balkan clade. We propose two possible scenarios on the historical evolution of our model taxa and the role of the last glacial maximum in shaping their current distribution. Finally, we discuss the complexities regarding the taxonomic delimitation of parapatric taxa.

A Contribution Towards Resolving the Systematics of the High-Altitude Tropical Andean Satyrine Genus Altopedaliodes Forster, 1964 (Lepidoptera, Nymphalinae: Satyrinae).

The first phylogenetic hypotheses for the high Andean satyrine butterfly genus Altopedaliodes is proposed based on sequence data from mitochondrial (COI and COII) and nuclear (EF-1α) genes. Four species previously included in the genus were found not to be closely related to the clade containing the type species for Altopedaliodes, and these species are therefore removed and placed in the appropriate genus: Pedaliodes cocytia (C. Felder and R. Felder, 1867) n. comb., Pedaliodes nebris Thieme 1905rev. comb., Neopedaliodes tamaensis (Pyrcz and Viloria 2007) n. comb., and Altopedaliodes similis Henao, Páez and Rodríguez-M., 2017; the last name is here synonymized with Pedaliodes nebris (Thieme 1905) n. syn. A taxonomic revision of the twelve species in Altopedaliodes as newly circumscribed is presented and all the currently recognized taxa are revised. We describe two new species, A. llanganati Padrón, Pyrcz and Willmott n. sp. (Ecuador, Napo) and A. pilimbala Pyrcz and Boyer n. sp. (Colombia, Cauca), and three new subspecies, A. reissi papallacta Padrón, Pyrcz and Willmott n. ssp. (Ecuador, Napo), A. reissi dominica Pyrcz and Padrón n. ssp. (Colombia, Valle del Cauca) and A. halli cagnoni Boyer, Pyrcz and Padrón n. ssp. (Ecuador, Morona-Santiago). A new status is proposed for A. nucea Pyrcz and Viloria 1999, n. stat., we combine Altopedaliodes scydmaena (Hayward, 1968) n. comb. and Altopedaliodes belmira (Pyrcz and Rodríguez, 2004), n. comb., n. stat. into the genus and treat the latter taxon as a species, and we newly treat Altopedaliodes scydmaena kruegeri Pyrcz 1999n. stat. as a subspecies.

One Hundred Years of Solitude: rediscovery of Catasticta lycurgus (Godman amp; Salvin, 1880), a yellow butterfly from the Sierra Nevada de Santa Marta, Colombia (Lepidoptera: Pieridae).

Catasticta lycurgus is a striking endemic butterfly, restricted to high elevation habitats in the Sierra Nevada de Santa Marta, an isolated mountain range separate from the Andes in the north of Colombia. The type, which for almost a hundred years was the only known specimen, was collected in 1878 by Frederick Simons in the vicinity of Atánquez and was sent to the UK to be described by renowned naturalists Godman and Salvin in 1880. In 1972, explorers Adams and Bernard collected a second specimen of C. lycurgus in the locality of San Pedro at 2,900m of elevation in the Sierra Nevada de Santa Marta. These two specimens were the only known ones for many decades until recently, when Colombian entomologists found the species again in San Pedro de la Sierra and later, when a female was discovered in 2013. Here, we report the rediscovery of this rare and charismatic species, with new specimens collected near the type locality, which have not been reported previously. The female of C. lycurgus is described and illustrated for the first time as well the male genitalia of this species. We combine all information available to provide some insights on the systematic relationships of this species within the genus Catasticta Butler, discuss its distribution and provide a preliminary conservation assessment. Despite the newly collected specimens, the species remains very rare in the field and in collections.

Taxonomy of the Neolycaena (Rhymnaria) davidi (Oberthür, 1881) species group, with the description of two new taxa from Russia and China (Lepidoptera, Lycaenidae).

Poorly known taxa of the Neolycaena (Rhymnaria) davidi (Oberthür, 1881) species group are reviewed in the light of modern findings. The taxon kozlovi Zhdanko, 1996 previously considered a subspecies of N. (R.) davidi is recognised here as a separate species, N. (R.) kozlovi Zhdanko, 1996, stat. rev., including the subspecies N. (R.) kozlovi namkhaidorji Churkin, 2004, comb. nov., based on morphological differences, the geographic isolation and ecological specialisation. A new species, N. (R.) markhasiovi sp. n., is described from Xinjiang Uyghur Autonomous Region in China (Eastern Tian Shan, the Karlik Mountains) on the basis of distinct genitalia morphology and features of the external morphology, including colouration of the female abdomen. Populations of the subgenus Rhymnaria from southern Tuva in Russia (the Tannu-Ola Mountains) are described as a separate subspecies, N. (R.) davidi tannuola ssp. n. A checklist of the taxa of the N. (R.) davidi species group including data on their distribution and host plants is presented, as well as a key to the species of the group.

Micropylar and Perimicropylar Regions of the Egg Exochorion in Six Genera of Dismorphiinae (Lepidoptera: Pieridae).

Using scanning electron microscopy (SEM), we examine and compare the micropylar and perimicropylar regions of the egg exochorion in six genera of the subfamily Dismorphiinae (Pieridae): Leptidea Billberg, Enantia Hübner, Pseudopieris Godman Salvin, Lieinix Gray, Dismorphia Hübner, and Moschoneura Butler. We provide the first descriptions of the morphological structures of the chorion in these genera. Leptidea is the most primitive genus, and Dismorphia represents the most diverse. We examined the jethys complex of Enantia to study diversity of characters within the genus. We conclude that the exochorion of Dismorphiinae is the most plesiomorphic in relation to Coliadinae and Pierinae, owing to its simple morphology without a transition zone surrounding the wreath (perimicropylar region). Leptidea (Leptideini) shows the least derived characters of the subfamily, followed by Enantia (Dismorphiini). The latter genus exhibits several typical characters of the tribe that are distinctive from other more specialized genera. Exochorionic similarities among the four species of the jethys complex support their close relationship; E. mazai exhibits the most derived features among the species of the complex. Pseudopieris and Moschoneura exhibit several plesiomorphies, such as a large number of openings (micro-grid), but they also show some derived features, such as three semi-rings in the wreath. Dismorphia and Lieinix exhibit characters in their most derived states, such as a striated micro-grid (most conspicuous in Lieinix) and basal aeropyles (most conspicuous in Dismorphia), a character shared with Moschoneura. The purpose of this work is exploratory and descriptive and would be improved by a larger sample size. This contribution is the third in a series of studies on the chorion of Dismorphiinae eggs.

Description of five new species of Epitola sensu lato (Lepidoptera: Lycaenidae: Poritiinae) from West and Central Africa.

Five new species of Epitola s.l. (Lycaenidae: Poritiinae) are described in the genera Stempfferia Jackson, 1962 and Cephetola Libert, 1999. Stempfferia salonga sp. n. was found in the 'Cuvette Centrale' of the Democratic Republic of Congo and a single specimen was identified also from southern Cameroon. The similar S. buea sp. n. is known only from the unique holotype collected at the foothills of Mount Cameroon, while S. michelliberti sp. n. is distributed in lowland rainforest between southeast Ivory Coast and Guinea. The new species of Cephetola were found only in Liberia. Cephetola praecox sp. n. is known from a single upland forest locality in the Putu Range, whereas C. wologizi sp. n. was collected also in upland forest in the Wologizi Mountains.

Toward standards in practices and techniques on ootaxonomy in the Pieridae (Lepidoptera: Papilionoidea).

We present a proposal on the standards used on ootaxonomy practices and techniques in the butterfly family Pieridae Duponchel (Lepidoptera: Papilionoidea) in five stages: 1) getting the specimens, 2) integration into a collection, 3) dissections to recover the exochorion, 4) elaboration of images of it, and 5) the preparation of its description with the necessary diagrams and tabulations. Also, we present the detailed techniques applied in observation and graphic representation, based on the methylene blue staining techniques and those required for the use with the scanning electron microscope (SEM). We compare the result of the standardized techniques with those from macro photography, drawings, and photographs with SEM-all of them found in books with descriptions and images of eggs of the Pieridae. We present a glossary and general aspects of the exochorion in the Pieridae as an Appendix to this article. Standardized techniques show more accurate and extensive character retrieval for systematics. For the scale in which they work, these techniques recovered more information than those present from oviposited eggs, where the exochorionic base is not seen. Also, the descriptions contain detailed data on more structures-which are comparable to each other-than are absent in the references mentioned. We present the recovered characters with the techniques found in the literature as three synthetically supplementary materials.

Alpha-taxonomy and phylogeny of African Junoniini butterflies based on morphological data, with an emphasis on genitalia, and COI barcode (Lepidoptera: Nymphalidae).

The tribe Junoniini is a predominantly Paleotropical group of the cosmopolitan butterfly subfamily Nymphalinae (Nymphalidae), with highest diversity in the Afrotropical region. Its systematics and relationships are not entirely resolved. Question marks remain concerning the validity of some genera; and the apparently close relationship between the Indo-Australian genus Yoma and the Afrotropical Protogoniomorpha, as evidenced by molecular phylogenies, remains a puzzle. Here, we present a cladistic analysis, based on 42 characters of the male and female genitalia of 41 species of Junoniini belonging to six genera, nearly all of them continental Afrotropical, and 3 species of two Indo-Australian genera Yoma and Rhinopalpa. A ML COI-based tree is produced for 36 species of Afrotropical Junoniini and Yoma. The molecular data are consistent with previous studies. However, morphological analysis does not confirm a close relationship between Protogoniomorpha and Yoma. Despite the evolution of a number of modifications, the male genitalia within all genera and species of the Junoniini share a cohesive build plan, in particular a transformed sacculus, from which Yoma is highly divergent. The position of the genus Kamilla, previously synonymized with Junonia, is discussed. Three East African coast taxa, Junonia elgiva stat. reinst., Protogoniomorpha nebulosa stat. reinst. and Salamis amaniensis stat. reinst., and one from central Africa, Precis silvicola stat. reinst. are raised to species level, based on comparative analysis of their male genitalia.

Two new species of Manerebia Staudinger from paramo habitat in the Colombian Eastern Cordillera of the Andes (Lepidoptera: Nymphalidae, Satyrinae, Pronophilina).

Two new species of the genus Manerebia Staudinger, 1897 (Nymphalidae, Satyrinae) are described from paramo habitat on the eastern slopes of the Colombian Eastern Cordillera in the area of Pisba and La Colorada: Manerebia bernito n. sp. and Manerebia clarita n. sp. Both are morphologically most similar to Manerebia levana and Manerebia pervaga, and the possible phylogenetic relationship between them is hypothesized. Their habitat is described and a proposal of their conservation status is made. With the addition of the two new species described in this study, a total of 15 species of Manerebia are reported from Colombia. However, a molecular analysis should be carried out to validate the taxonomic status of several species of Manerebia, in particular having in mind a potentially high cryptic diversity of this genus.

Jamides wananga, a new species from Papua New Guinea and Australia (Lepidoptera: Lycaenidae).

Jamides wananga sp. n. is described and illustrated from Madang Province of Papua New Guinea and Dauan Island in the Torres Strait (Queensland, Australia). The new species is similar in appearance to several other Jamides Hübner species in the bochus-group. DNA sequence data and morphology were used to distinguish the nominotypical subspecies found on the New Guinea mainland from J. w. roxina subsp. n. on Dauan Island. Notes on the habitat and behaviour are also provided.

A brief history of Lepidoptera section contributions to Zootaxa.

As a reviewer, John Brown received the first Lepidoptera manuscript submitted to Zootaxa in 2002. Within a year he was persuaded by a colleague to volunteer as its first Lepidoptera section editor. As submissions increased, he realized that he needed assistance, so in July 2005 he enlisted Robert Robbins (U.S. National Museum of National History), and the two editors split the submissions-Brown covered moths and Robbins butterflies. As submissions continued to grow, Robbins stepped down and Brown was again the sole editor. Owing to the ever-increasing manuscript load, in 2007 Brown submitted a proposal to several colleagues, inviting them to become Lepidoptera section editors, with the concept that the more editors there were, the fewer manuscripts each would have to handle, and their duties would include papers primarily in their area of expertise. The solicitation was successful, with four new subject editors coming on board in 2007: Lawrence Gall for macrolepidoptera families, Michael Toliver for butterflies, Jean-François Landry for microlepidoptera families, and Shen-Horn Yen for Pyraloidea and Zygaenoidea; the last two are still section editors today. Over the next 13 years, numerous editors came and went-turnover in editorship was always viewed as a positive way to involve new scientists and interject fresh ideas. From 2001 to 2020, a cumulative total of 21 scientists have served as Lepidoptera Section editors (Table 1), representing 14 different countries.

Geographical distribution of Emesis/ Fabricius (Lepidoptera: Riodinidae) in Mexico: Updated checklist and temporal patterns.

We present a synthesis of the existing information on the genus Emesis Fabricius in Mexico concerning biogeographical patterns and taxonomical aspects. Emesis is the most diverse genus of Emesidini with 57 species and subspecies, with Mexico as the northern limit of this Neotropical genus. We analyzed 5434 specimens of the Lepidoptera Collection of the MZFC, UNAM and compared them with specimens from collections of Mexico, Central and South America. Taxonomic determination and corroboration were made by analysis of wing patterns and genitalia. Geographic distribution and phenology were obtained from the database MARIPOSA. We present an updated list of Emesis of Mexico, with 17 species and subspecies. For each species, we provide information on phenology, geographic, altitudinal, and vegetation distributions. We discuss taxonomic and undersampling concerns for some species, as well as spatial and temporal patterns with special reference to vegetation types and biogeographic provinces in Mexico.

Notes on some Tanzanian butterfly specimens in the Suffert Collection: a case of patria/ falsa/.

A number of mislabelled butterfly specimens in the Suffert collection purporting to be from Lindi in southeast Tanzania belong to taxa restricted to the Usambara Mountains in northeast Tanzania thus suggesting patria falsa. The history of Suffert's collection and the possible reasons behind this mislabelling event are discussed in more detail. One of the mislabelled specimens, the holotype of Papilio illyris hamatus Joicey Talbot, 1918 described from Tanzania, is the senior name for Graphium illyris girardeaui Guilbot Plantrou, 1978, the latter name formally synonymised herein with the former. A brief biography of the little-known Ernst Suffert is presented.

Revised species definitions and nomenclature of the blue and purple/rose Cithaerias/ butterflies (Lepidoptera, Nymphalidae, Satyrinae).

This study reassesses the taxonomic status of Neotropical blue and purple/rose-colored Cithaerias butterflies, thus complementing a previous study of the rose-colored species. Based on comparative study of wing coloration and genitalia morphology, I revise species definitions and the taxonomic status of: Cithaerias andromeda, C. azurina STAT. REV., C. esmeralda STAT. REV., C. bandusia STAT. REV., C. pyropina, and C. songoana STAT. REV. Photographs of adults and illustrations of male and female genitalia are provided for all species. Of particular importance are the genitalia illustrations of male and female C. azurina, presented here for the first time, as well as finding a putative first female of Ecuadorean subspecies C. pyropina julia.

A New Species of Aporia/ (Lepidoptera: Pieridae) from Northwest Yunnan, China with Taxonomic Notes on Its Similar Sympatric Taxa.

The northwest corner of Yunnan Province, China is the world-class diversity centre of the genus Aporia Hübner, 1819 (Lepidoptera: Pieridae). During our expeditions to this area in 2019 and 2020, a new species of Aporia was discovered from the upper Yangtze River valley in northwest Yunnan, China. The new taxon, A. chunhaoi sp. nov., is similar to A. lhamo (Oberthür, 1893) but can be easily identified by larger size, much paler (creamy) male hindwing, as well as different genitalic structures. Our field surveys and comparative studies also confirmed a new range of A. lhamo in the upper Yangtze River valley, and the morphological variation of this species is discussed herein.

Contributions to the knowledge of Neotropical Lycaenidae: taxonomy of Johnsonita/ Salazar amp; Constantino, 1995 with descriptions of seven new species (Theclinae: Eumaeini).

The Transandean-Andean genus Johnsonita Salazar Constantino, 1995 is revised on the basis of wing and genitalia morphology. Apart from eight species formerly placed in Johnsonita (Thecla assula Draudt, 1919; Thecla auda Hewitson, 1867; Thecla catadupa Hewitson, 1869; Thecla chaluma Schaus, 1902, Thecla chlamydem Druce, 1907; Thecla pardoa d'Abrera, 1995; Johnsonita johnsoni Salazar Constantino, 1995 and Johnsonita johnbanksi Bálint, 2003), seven new species are described: Johnsonita carpia Bálint, Boyer Pyrcz, sp. n. (Ecuador), Johnsonita iacinta Bálint, Boyer Pyrcz, sp. n. (Peru), Johnsonita ianusca Bálint, Lorenc-Brudecka Pyrcz, sp. n. (Ecuador), Johnsonita oxalida Bálint, Boyer Pyrcz, sp. n. (Peru), Johnsonita subcunicula Bálint, Cerdeña Pyrcz, sp. n. (Peru), Johnsonita turquisca Bálint, Boyer Pyrcz, sp. n. (Ecuador) and Johnsonita zubkova Bálint, Boyer Lorenc-Brudecka, sp. n. (Peru). One species, erroneously recorded from Bolivia under the name Thecla chaluma Schaus, 1902, is not formally described due to insufficient material. An identification key, type material revision, descriptions or redescriptions, and diagnosis for all the taxa recognized are given. Lectotypes for the nominal species Thecla assula Draudt, 1919 and Thecla chlamydem Druce, 1907 are designated. Male secondary wing characters are tabulated. A brief discussion on classification, life-history and male secondary characters of Johnsonita is presented.

A new species of the genus Teratozephyrus/ Sibatani, 1946 from China (Lepidoptera, Lycaenidae, Theclinae).

A new species, T. yaolihuoi Huang, Jiang Song sp. nov. of the genus Teratozephyrus Sibatani, 1946 is described from southeastern China. Among the other representatives of the genus, the new species resembles only T. hinomaru Fujioka, 1994 from southwestern China, from which the new species can be separated easily by examining details in both wing pattern and the male genitalia. Wing patterns and genitalia of the new species and T. hinomaru are illustrated and compared.

Chromosome Level Assembly of the Comma Butterfly (Polygonia c-album).

The comma butterfly (Polygonia c-album, Nymphalidae, Lepidoptera) is a model insect species, most notably in the study of phenotypic plasticity and plant-insect coevolutionary interactions. In order to facilitate the integration of genomic tools with a diverse body of ecological and evolutionary research, we assembled the genome of a Swedish comma using 10X sequencing, scaffolding with matepair data, genome polishing, and assignment to linkage groups using a high-density linkage map. The resulting genome is 373 Mb in size, with a scaffold N50 of 11.7 Mb and contig N50 of 11,2Mb. The genome contained 90.1% of single-copy Lepidopteran orthologs in a BUSCO analysis of 5,286 genes. A total of 21,004 gene-models were annotated on the genome using RNA-Seq data from larval and adult tissue in combination with proteins from the Arthropoda database, resulting in a high-quality annotation for which functional annotations were generated. We further documented the quality of the chromosomal assembly via synteny assessment with Melitaea cinxia. The resulting annotated, chromosome-level genome will provide an important resource for investigating coevolutionary dynamics and comparative analyses in Lepidoptera.