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.

Genomics-based higher classification of the species-rich Hairstreaks (Lepidoptera: Lycaenidae: Eumaeini).

We propose a higher classification of the lycaenid hairstreak tribe Eumaeini - one of the youngest and most species-rich butterfly tribes - based on autosome, Lepidopteran Z sex chromosome, and mitochondrial protein-coding genes. The subtribe Neolycaenina Korb is a synonym of Callophryidina Tutt, and subtribe Tmolusina Bálint is a synonym of Strephonotina K. Johnson, Austin, Le Crom, & Salazar. Proposed names are Rhammina Prieto & Busby, new subtribe; Timaetina Busby & Prieto, new subtribe; Atlidina Martins & Duarte, new subtribe; Evenina Faynel & Grishin, new subtribe; Jantheclina Robbins & Faynel, new subtribe; Paiwarriina Lamas & Robbins, new subtribe; Cupatheclina Lamas & Grishin, new subtribe; Parrhasiina Busby & Robbins, new subtribe; Ipideclina Martins & Grishin, new subtribe; and Trichonidina Duarte & Faynel, new subtribe. Phylogenetic results from the autosome and Z sex chromosome analyses are similar. Future analyses of datasets with hundreds of terminal taxa may be more practical time-wise by focussing on the smaller number of sex chromosome sequences (2.6% of nuclear protein-coding sequences). The phylogenetic classification and biological summaries for each subtribe suggest that a variety of factors affected Eumaeini diversification. About a dozen kinds of male secondary sexual organs with frequent evolutionary gains and losses occur in Atlidina, Evenina, and Jantheclina (141 species combined). Females have been shown to use these organs to discriminate between conspecific and non-conspecific males, facilitating sympatry among close relatives. Eumaeina, Rhammina, and Timaetina (140 species combined) are overwhelmingly montane with some evidence for a higher incidence of sympatric diversification. Seven Neotropical lineages in five subtribes invaded the temperate parts of the Nearctic Region with a diversification increase in the Callophryidina (262 species). North American Satyrium and Callophrys then invaded the Palearctic at least once each, with a major species-richness increase in Satyrium. The evolution of litter feeding detritivores within Calycopidina (172 species) resulted in an increase in diversification rate compared with its flower-feeding sister lineage. Atlidina, Strephonotina, Parrhasiina, and Strymonina (562 species combined) each contain a mixture of genera that specialize on one or two caterpillar food plant families and genera that are polyphagous. These would be appropriate subtribes to assess how the breadth of caterpillar food plants and the frequency of host shifts affected diversification.

Evolutionary trade-offs between male secondary sexual traits revealed by a phylogeny of the hyperdiverse tribe Eumaeini (Lepidoptera: Lycaenidae).

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.

Four hundred shades of brown: Higher level phylogeny of the problematic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae) based on hybrid enrichment data.

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.

Into the Andes: multiple independent colonizations drive montane diversity in the Neotropical clearwing butterflies Godyridina.

Understanding why species richness peaks along the Andes is a fundamental question in the study of Neotropical biodiversity. Several biogeographic and diversification scenarios have been proposed in the literature, but there is confusion about the processes underlying each scenario, and assessing their relative contribution is not straightforward. Here, we propose to refine these scenarios into a framework which evaluates four evolutionary mechanisms: higher speciation rate in the Andes, lower extinction rates in the Andes, older colonization times and higher colonization rates of the Andes from adjacent areas. We apply this framework to a species-rich subtribe of Neotropical butterflies whose diversity peaks in the Andes, the Godyridina (Nymphalidae: Ithomiini). We generated a time-calibrated phylogeny of the Godyridina and fitted time-dependent diversification models. Using trait-dependent diversification models and ancestral state reconstruction methods we then compared different biogeographic scenarios. We found strong evidence that the rates of colonization into the Andes were higher than the other way round. Those colonizations and the subsequent local diversification at equal rates in the Andes and in non-Andean regions mechanically increased the species richness of Andean regions compared to that of non-Andean regions ('species-attractor' hypothesis). We also found support for increasing speciation rates associated with Andean lineages. Our work highlights the importance of the Andean slopes in repeatedly attracting non-Andean lineages, most likely as a result of the diversity of habitats and/or host plants. Applying this analytical framework to other clades will bring important insights into the evolutionary mechanisms underlying the most species-rich biodiversity hotspot on the planet.

Checklist of Yunnan Papilionidae (Lepidoptera: Papilionoidea) with nomenclatural notes and descriptions of new subspecies.

A checklist of the Papilionidae of Yunnan is presented, with nomenclatural and taxonomic changes made. In the nomenclatural section, the junior homonym Papilio bootes nigricans Rothschild, 1895 is replaced by Papilio bootes nigricauda Lamas & Cotton nom. nov., Chilasa (Cadugoides) epycides muhabbet Koak, 2005 is synonymised with Papilio epycides camilla Rousseau-Decelle, 1947 syn. nov., Graphium cloanthus nyghmat Koak & Kemal, 2000 is placed as a junior objective synonym syn. nov. of Graphium cloanthus clymenus (Leech, 1893), and Papilio astorion Westwood, 1842 is shown to have priority over Papilio varuna White, 1842, thus the valid species name is Atrophaneura astorion (Westwood, 1842) comb. nov. In the main checklist, five new subspecies are described: Parnassius cephalus haba Hu & Cotton ssp. nov., Lamproptera curius hsinningae Hu, Zhang & Cotton ssp. nov., Lamproptera curius yangtzeanus Hu & Cotton ssp. nov., Graphium macareus vadimi Cotton & Hu ssp. nov., and Papilio krishna benyongi Hu & Cotton ssp. nov. The First Reviser Principle under the ICZN Code is invoked to solve four taxonomic problems, and 18 names are synonymised with explanations, notably Papilio machaon venchuanus Moonen, 1984 syn. nov., which is synonymised with Papilio machaon schantungensis Eller, 1936. Byasa genestieri (Oberthr, 1918) stat. nov. is separated from Byasa latreillei (Donovan, 1826), and Papilio everesti Riley, 1927 stat. nov. and P. verityi Fruhstorfer, 1907 stat. nov. are separated from Papilio machaon Linnaeus, 1758 as species. Taxa that require further confirmation of their presence in Yunnan and those that do not occur in Yunnan are enumerated.

Lost and Found: Taxonomic Notes on Some Butterflies (Lepidoptera: Papilionoidea) from J.F. Zikán found on F. Dissmann's Collection.

J. F. Zikán described several new taxa of butterflies from the Neotropical region. The majority of the butterfly types of J. F. Zikán was deposited at the Instituto Oswaldo Cruz, Rio de Janeiro, Brazil (IOC). However, some type series were found to be incomplete. The present paper discusses and illustrate some type specimens of Papilionidae and Nymphalidae from J. F. Zikán found in Dissmann's collection, that now is deposited at Museu de Diversidade Biológica from University of Campinas - UNICAMP, Campinas, São Paulo, Brazil (ZUEC).

A global phylogeny of butterflies reveals their evolutionary history, ancestral hosts and biogeographic origins.

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.

Neotype designation for Papilio fulgerator Walch, 1775 (Hesperiidae: Eudaminae).

The discovery that a skipper butterfly Telegonus fulgerator (Walch, 1775), previously placed in the genus Astraptes Hübner, [1819], is a complex of many similar-looking species-level taxa with different COI barcodes, caterpillar foodplants and body patterns, and subtle differences in adult phenotypes raised a question about which species is the original T. fulgerator. To answer this question, being unable to locate its holotype, we designate the neotype of Papilio fulgerator Walch, 1775, a female specimen from Suriname in the Zoological State Collection, Munich, Germany. This neotype will form the foundation for a comprehensive revision of the T. fulgerator complex based on genomic sequencing and analysis augmented with phenotypic considerations.

A peculiar new species of Dione (Agraulis) Boisduval & Le Conte (Lepidoptera, Nymphalidae, Heliconiinae) associated with Malesherbia Ruiz & Pavón (Passifloraceae) in xeric western slopes of the Andes.

Butterflies associated with xerophytic environments of the Andes have been little studied, and they exhibit high levels of endemism. Herein Dione (Agraulis) dodona Lamas & Farfán, sp. nov. (Nymphalidae; Heliconiinae) is described, distributed on the western slopes of the Andes of Peru and northern Chile, between 800 and 3,000 m elevation. Adults of both sexes, and the immature stages, are described and illustrated based on light and scanning electron microscopy. The immature stages are associated with MalesherbiatenuifoliaD. Don (Passifloraceae) found in xeric environments, representing a new record of this genus as a host plant for the subfamily Heliconiinae. Conspicuous morphological differences are presented for all stages at the generic level. Based on a phylogenetic analysis of the COI barcode mitochondrial gene fragment, D. (A.) dodona Lamas & Farfán, sp. nov. is distinguished as an independent lineage within the Agraulis clade of Dione, with ca. 5% difference to congeneric species.

Taxonomic discoveries enabled by genomic analysis of butterflies.

The comparative genomics of butterflies yields additional insights into their phylogeny and classification that are compiled here. As a result, 3 genera, 5 subgenera, 5 species, and 3 subspecies are proposed as new, i.e., in Hesperiidae: Antina Grishin, gen. n. (type species Antigonus minor O. Mielke, 1980), Pompe Grishin and Lamas, gen. n. (type species Lerema postpuncta Draudt, 1923), and Curva Grishin, gen. n. (type species Moeris hyagnis Godman, 1900); in Lycaenidae: Fussia Grishin, subgen. n. (type species Polyommatus standfussi Grum-Grshimailo, 1891) and Pava Grishin, subgen. n. (type species Thecla panava Westwood, 1852); in Hesperiidae: Monoca Grishin, subgen. n. (type species Tagiades monophthalma Plötz, 1884), Putuma Grishin, subgen. n. (type species Tisias putumayo Constantino and Salazar, 2013), and Rayia Grishin, subgen. n. (type species Mastor perigenes Godman, 1900); Cissia wahala Grishin, sp. n. (Nymphalidae; type locality in Mexico: Oaxaca); in Hesperiidae: Hedone mira Grishin and Lamas, sp. n. (type locality in Peru: Apurímac), Vidius pompeoides Grishin, sp. n. (type locality in Brazil: Amazonas), Parphorus hermieri Grishin, sp. n. (Hesperiidae; type locality in Brazil: Rondônia), and Zenis par Grishin, sp. n. (Hesperiidae; type locality in Peru: Cuzco); in Pieridae: Glutophrissa drusilla noroesta Grishin, ssp. n. (type locality in USA: Texas, Cameron Co.) and Pieris marginalis siblanca Grishin, ssp. n. (type locality in USA: New Mexico, Lincoln Co.), and Argynnis cybele neomexicana Grishin, ssp. n. (Nymphalidae; type locality in USA: New Mexico, Sandoval Co.). Acidalia leto valesinoides-alba Reuss, [1926] and Acidalia nokomis valesinoides-alba Reuss, [1926] are unavailable names. Neotypes are designated for Mylothris margarita Hübner, [1825] (type locality in Brazil) and Papilio coras Cramer, 1775 (type locality becomes USA: Pennsylvania, Montgomery Co., Flourtown). Mylothris margarita Hübner, [1825] becomes a junior objective synonym of Pieris ilaire Godart, 1819, currently a junior subjective synonym of Glutophrissa drusilla (Cramer, 1777). Lectotypes are designated for Hesperia ceramica Plötz, 1886 (type locality in Indonesia: Seram Island), Pamphila trebius Mabille, 1891 (type locality Colombia: Bogota), Methionopsis modestus Godman, 1901 and Papias microsema Godman, 1900 (type locality in Mexico: Tabasco), Hesperia fusca Grote & Robinson, 1867 (type locality in USA: Georgia), Goniloba corusca Herrich-Schäffer, 1869, and Goniloba devanes Herrich-Schäffer, 1869; the type localities of the last two species, together with Pamphila stigma Skinner, 1896 and Carystus (Argon) lota (Hewitson, 1877), are deduced to be in South America. Type locality of Junonia pacoma Grishin, 2020 is in Sinaloa, not Sonora (Mexico). Abdomen is excluded from the holotype of Staphylus ascalon (Staudinger, 1876). Furthermore, a number of taxonomic changes are proposed. Alciphronia Koçak, 1992 is treated as a subgenus, not a synonym of Heodes Dalman, 1816. The following genera are treated as subgenera: Lafron Grishin, 2020 of Lycaena [Fabricius], 1807, Aremfoxia Real, 1971 of Epityches D'Almeida, 1938, Placidina D'Almeida, 1928 of Pagyris Boisduval, 1870, and Methionopsis Godman, 1901 of Mnasinous Godman, 1900. Polites (Polites) coras (Cramer, 1775) is not a nomen dubium but a valid species. The following are species-level taxa (not subspecies or synonyms of taxa given in parenthesis): Lycaena pseudophlaeas (Lucas, 1866) and Lycaena hypophlaeas (Boisduval, 1852) (not Lycaena phlaeas (Linnaeus, 1761), Satyrium dryope (W. H. Edwards, 1870) (not Satyrium sylvinus (Boisduval, 1852)), Apodemia cleis (W. H. Edwards, 1882) (not Apodemia zela (Butler, 1870)), Epityches thyridiana (Haensch, 1909), comb. nov. (not Epityches ferra Haensch, 1909, comb. nov.), Argynnis bischoffii W. H. Edwards, 1870 (not Argynnis mormonia Boisduval, 1869), Argynnis leto Behr, 1862 (not Argynnis cybele (Fabricius, 1775)), Boloria myrina (Cramer, 1777) (not Boloria selene ([Denis & Schiffermüller], 1775)), Phyciodes jalapeno J. Scott, 1998 (not Phyciodes phaon (W. H. Edwards, 1864)), Phyciodes incognitus Gatrelle, 2004 and Phyciodes diminutor J. Scott, 1998 (not Phyciodes cocyta (Cramer, 1777)), Phyciodes orantain J. Scott, 1998 (not Phyciodes tharos (Drury, 1773)), Phyciodes anasazi J. Scott, 1994 (not Phyciodes batesii (Reakirt, [1866])), Cercyonis silvestris (W. H. Edwards, 1861) (not Cercyonis sthenele (Boisduval, 1852)), Paramacera allyni L. Miller, 1972 and Paramacera rubrosuffusa L. Miller, 1972 (not Paramacera xicaque (Reakirt, [1867])), Cissia cheneyorum (R. Chermock, 1949), Cissia pseudocleophes (L. Miller, 1976), and Cissia anabelae (L. Miller, 1976) (not Cissia rubricata (W. H. Edwards, 1871)), Tarsoctenus gaudialis (Hewitson, 1876) (not Tarsoctenus corytus (Cramer, 1777)), Nisoniades inca (Lindsey, 1925) (not Nisoniades mimas (Cramer, 1775), Xenophanes ruatanensis Godman & Salvin, 1895 (not Xenophanes tryxus (Stoll, 1780)), Lotongus shigeoi Treadaway & Nuyda, 1994, Lotongus balta Evans, 1949, Lotongus zalates (Mabille, 1893), and Lotongus taprobanus (Plötz, 1885) (not Lotongus calathus (Hewitson, 1876)), Oxynthes martius (Mabille, 1889) (not Oxynthes corusca (Herrich-Schäffer, 1869)), Notamblyscirtes durango J. Scott, 2017 (not Notamblyscirtes simius W. H. Edwards, 1881), Hedone praeceps Scudder, 1872, Hedone catilina (Plötz, 1886), and Hedone calla (Evans, 1955) (not Hedone vibex (Geyer, 1832)), Atalopedes huron (W. H. Edwards, 1863) (not Atalopedes campestris (Boisduval, 1852)), Papias microsema Godman, 1900 (not Mnasinous phaeomelas (Hübner, [1829]), comb. nov.), Papias unicolor (Hayward, 1938) and Papias monus Bell, 1942 (not Papias phainis Godman, 1900), Nastra leuconoides (Lindsey, 1925) (not Nastra leucone (Godman, 1900)), Nastra fusca (Grote & Robinson, 1867) (not Nastra lherminier (Latreille, [1824])), Zenis hemizona (Dyar, 1918) and Zenis janka Evans, 1955 (not Zenis jebus (Plötz, 1882)), Carystus (Argon) argus Möschler, 1879 (not Carystus (Argon) lota Hewitson, 1877), and Lycas devanes (Herrich-Schäffer, 1869) (not Lycas argentea (Hewitson, 1866)). Borbo impar ceramica (Plötz, 1886), comb. nov. is not a synonym of Pelopidas agna larika (Pagenstecher, 1884) but a valid subspecies. Parnassius smintheus behrii W. H. Edwards, 1870 and Cercyonis silvestris incognita J. Emmel, T. Emmel & Mattoon, 2012 are subspecies, not species. The following are junior subjective synonyms: Shijimiaeoides Beuret, 1958 of Glaucopsyche Scudder, 1872, Micropsyche Mattoni, 1981 of Turanana Bethune-Baker, 1916, Cyclyrius Butler, 1897 of Leptotes Scudder, 1876, Mesenopsis Godman & Salvin, 1886 of Xynias Hewitson, 1874, Carystus tetragraphus Mabille, 1891 of Lotongus calathus parthenope (Plötz, 1886), Parnara bipunctata Elwes & J. Edwards, 1897 of Borbo impar ceramica (Plötz, 1886), Hesperia peckius W. Kirby, 1837 of Polites (Polites) coras (Cramer, 1775), and Lerodea neamathla Skinner & R. Williams, 1923 of Nastra fusca (Grote & Robinson, 1867). The following transfers are proposed: of species between genera (i.e., revised genus-species combinations): Nervia niveostriga (Trimen, 1864) (not Kedestes Watson, 1893), Leona lota Evans, 1937 (not Lennia Grishin, 2022), Leona pruna (Evans, 1937) and Leona reali (Berger, 1962) (not Pteroteinon Watson, 1893), Mnasinous phaeomelas (Hübner, [1829]) (not Papias Godman, 1900), Saturnus jaguar (Steinhauser, 2008) (not Parphorus Godman, 1900), Parphorus harpe (Steinhauser, 2008) (not Saturnus Evans, 1955), Parphorus kadeni (Evans, 1955) (not Lento Evans, 1955), and Calpodes chocoensis (Salazar & Constantino, 2013) (not Megaleas Godman, 1901); of subspecies between species (i.e., revised species-subspecies combinations): Melitaea sterope W. H. Edwards, 1870 of Chlosyne palla (Boisduval, 1852) (not Chlosyne acastus (W. H. Edwards, 1874)) and Panoquina ocola distipuncta Johnson & Matusik, 1988 of Panoquina lucas (Fabricius, 1793); and junior subjective synonym transferred between species: Rhinthon zaba Strand, 1921 of Conga chydaea (A. Butler, 1877), not Cynea cynea (Hewitson, 1876), Pamphila stigma Skinner, 1896 of Hedone catilina (Plötz, 1886), not Hedone praeceps Scudder, 1872, and Pamphila ortygia Möschler, 1883 of Panoquina hecebolus (Scudder, 1872), not Panoquina ocola (W. H. Edwards, 1863). Proposed taxonomic changes result in additional revised species-subspecies combinations: Lycaena pseudophlaeas abbottii (Holland, 1892), Satyrium dryope putnami (Hy. Edwards, 1877), Satyrium dryope megapallidum Austin, 1998, Satyrium dryope itys (W. H. Edwards, 1882), Satyrium dryope desertorum (F. Grinnell, 1917), Argynnis bischoffi opis W. H. Edwards, 1874, Argynnis bischoffi washingtonia W. Barnes & McDunnough, 1913, Argynnis bischoffi erinna W. H. Edwards, 1883, Argynnis bischoffi kimimela Marrone, Spomer & J. Scott, 2008, Argynnis bischoffi eurynome W. H. Edwards, 1872, Argynnis bischoffi artonis W. H. Edwards, 1881, Argynnis bischoffi luski W. Barnes & McDunnough, 1913, Argynnis leto letona (dos Passos & Grey, 1945), Argynnis leto pugetensis (F. Chermock & Frechin, 1947), Argynnis leto eileenae (J. Emmel, T. Emmel & Mattoon, 1998), Boloria myrina nebraskensis (W. Holland, 1928), Boloria myrina sabulocollis Kohler, 1977, Boloria myrina tollandensis (W. Barnes & Benjamin, 1925), Boloria myrina albequina (W. Holland, 1928), Boloria myrina atrocostalis (Huard, 1927), Boloria myrina terraenovae (W. Holland, 1928), Phyciodes anasazi apsaalooke J. Scott, 1994, Polites coras surllano J. Scott, 2006, and Curva darienensis (Gaviria, Siewert, Mielke & Casagrande, 2018). Specimen curated as the holotype of Acidalia leto valesinoides-alba Reuss, [1926] is Argynnis leto letona (dos Passos & Grey, 1945) (not A. leto leto Behr, 1862) from USA: Utah, Provo. A synonymic list of available genus-group names for Lycaeninae [Leach],[1815] is given. Unless stated otherwise, all subgenera, species, subspecies and synonyms of mentioned genera and species are transferred with their parent taxa, and others remain as previously classified.

Considerations on the Systematics of Neotropical Pierina, with the Description of Two New Species of Phulia Herrich-Schäffer from the Peruvian Andes (Lepidoptera: Pieridae, Pierinae, Pierini).

A comparative analysis of high-Andean Pierina was carried out, including a total of 25 species. Based on morphological evidence, with an emphasis on venation and genitalia and molecular data, using three genetic markers, we confirm the recent subjective synonymy of the generic names Tatochila Butler, 1870, Piercolias, Staudinger, 1894, Hypsochila Ureta, 1955, Infraphulia Field, 1958, Pierphulia Field, 1958, and Theochila Field, 1958 with Phulia Herrich-Schäffer, 1867. Two new species are described, namely Phulia stoddardi Pyrcz & Cerdeña n. sp., from the Andes of Central Peru, which occurs at an unusually high altitude of close to 5000 m a.s.l. in dry puna habitat, and Phulia phantasma Lamas, Willmott & Boyer n. sp., from dry montane forests in northern Peru and southern Ecuador. An overview of high-elevation butterflies is presented, with some discussion on adaptations to this environment.

LepTraits 1.0 A globally comprehensive dataset of butterfly traits.

Here, we present the largest, global dataset of Lepidopteran traits, focusing initially on butterflies (ca. 12,500 species records). These traits are derived from field guides, taxonomic treatments, and other literature resources. We present traits on wing size, phenology,voltinism, diapause/overwintering stage, hostplant associations, and habitat affinities (canopy, edge, moisture, and disturbance). This dataset will facilitate comparative research on butterfly ecology and evolution and our goal is to inspire future research collaboration and the continued development of this dataset.

Two new species of Hermeuptychia from North America and three neotype designations (Nymphalidae: Satyrinae).

Two new species of Hermeuptychia Forster, 1964 are described. Hermeuptychia sinuosa Grishin, sp. n. (type locality Guatemala: El Progreso, Morazán) is an isolated member of the genus that does not readily fit into known species groups, as suggested by its distinct male and female genitalia and COI DNA barcode sequences. It is distinguished from its congeners by prominently wavy submarginal lines, rounder wings and distinctive genitalia, and can typically be identified by a white dot, instead of an eyespot, near the ventral hindwing apex. Hermeuptychia occidentalis Grishin, sp. n. (type locality Mexico: Guerrero, Acapulco) belongs to the Hermeuptychia sosybius group as indicated by the presence of androconia on the dorsal surface of the wings, genitalia and COI DNA barcodes, and in addition to DNA characters, differs from its relatives in the shape of the uncus and female genitalia. Neotypes of Oreas strigata canthe Hübner, [1811] (type locality Suriname: Gelderland, Suriname River), Megisto acmenis Hübner, 1823 (type locality Argentina: Buenos Aires), and Satyrus cantheus Godart, [1824] (type locality USA: Florida, Pinellas Co., St. Petersburg) and lectotype of Euptychia celmis var. bonaërensis [sic] Burmeister, 1878 (type locality Argentina: Buenos Aires) are designated. These designations establish Hermeuptychia canthe as a valid species widely distributed in South America from Colombia to Bolivia and Southeast Brazil, Euptychia celmis var. bonaërensis [sic] Burmeister, 1878 as a junior objective synonym of Yphthimoides acmenis, and S. cantheus as a junior subjective synonym of Hermeuptychia sosybius (Fabricius, 1793). Papilio camerta Cramer, 1780 is treated as nomen dubium requiring further studies to determine an identity that is consistent with the original description, as it may be conspecific with Paryphthimoides poltys (Prittwitz, 1865) instead of being a Hermeuptychia species as currently assumed.

Se describe dos nuevas especies de Hermeuptychia Forster, 1964. Hermeuptychia sinuosa Grishin, sp. n. (localidad tipo Guatemala: El Progreso, Morazán), es un componente aislado del género que no encaja fácilmente en los grupos de especies conocidas, como lo indica su distintiva genitalia masculina y femenina y las secuencias de ADN del código de barras COI. Se distingue de sus congéneres por tener líneas submarginales prominentemente onduladas, alas más redondas y genitales diferentes, y se puede identificar típicamente por un punto blanco, en lugar de una mancha ocular, cerca del ápice ventral del ala anterior. Hermeuptychia occidentalis Grishin, sp. n. (localidad tipo México: Guerrero, Acapulco) pertenece al grupo de Hermeuptychia sosybius como lo indica la presencia de androconia en las alas anteriores, la estructura genital y secuencias de ADN de la región del código de barras COI, y además de caracteres del ADN, se diferencia de sus parientes en la forma del uncus y la genitalia femenina. Se designa neotipos para Oreas strigata canthe Hübner, [1811] (localidad tipo Surinam: Gelderland, Río Surinam), Megisto acmenis Hübner, 1823 (localidad tipo Argentina: Buenos Aires), y Satyrus cantheus Godart, [1824] (localidad tipo Estados Unidos: Florida, Pinellas Co., St. Petersburg), y el lectotipo de Euptychia celmis var. bonaërensis [sic] Burmeister, 1878 (localidad tipo Argentina: Buenos Aires). Estas designaciones establecen a Hermeuptychia canthe como una especie válida ampliamente distribuida en América del Sur desde Colombia hasta Bolivia y el sureste de Brasil, a Euptychia celmis var. bonaërensis [sic] Burmeister, 1878 como sinónimo objetivo más reciente de Yphthimoides acmenis, y a S. cantheus como sinónimo subjetivo más reciente de Hermeuptychia sosybius (Fabricius, 1793). Papilio camerta Cramer, 1780 es tratado como un nomen dubium requiriendo más estudios para determinar una identidad que sea consistente con la descripción original, ya que puede ser coespecífica con Paryphthimoides poltys (Prittwitz, 1865) en lugar de ser una especie de Hermeuptychia como se asume actualmente.

Conserved ancestral tropical niche but different continental histories explain the latitudinal diversity gradient in brush-footed butterflies.

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.

The anatomy of a 'suture zone' in Amazonian butterflies: a coalescent-based test for vicariant geographic divergence and speciation.

Attempts by biogeographers to understand biotic diversification in the Amazon have often employed contemporary species distribution patterns to support particular theories, such as Pleistocene rainforest refugia, rather than to test among alternative hypotheses. Suture zones, narrow regions where multiple contact zones and hybrid zones between taxa cluster, have been seen as evidence for past expansion of whole biotas that have undergone allopatric divergence in vicariant refuges. We used coalescent analysis of mutilocus sequence data to examine population split times in 22 pairs of geminate taxa in ithomiine and heliconiine butterflies. We test a hypothesis of simultaneous divergence across a suture zone in NE Peru. Our results reveal a scattered time course of diversification in this suture zone, rather than a tight cluster of split times. Additionally, we find rapid diversification within some lineages such as Melinaea contrasting with older divergence within lineages such as the Oleriina (Hyposcada and Oleria). These results strongly reject simple vicariance as a cause of the suture zone. At the same time, observed lineage effects are incompatible with a series of geographically coincident vicariant events which should affect all lineages similarly. Our results suggest that Pleistocene climatic forcing cannot readily explain this Peruvian suture zone. Lineage-specific biological traits, such as characteristic distances of gene flow or varying rates of parapatric divergence, may be of greater importance.

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.

A new species of Mathania Oberthür, 1890 from Peru (Lepidoptera, Pieridae).

A new species of the genus Mathania Oberthür, 1890, M. hughesi Lamas, Farfán Cerdeña, sp. n. is described from the southwestern slopes of the Andes of Peru, associated with xerophytic environments, between 2300 and 3500 m elevation. This new species is distinguishable from its congeners by the following external character: a black band on forewing dorsal surface at the end of the discal cell, extending from the costal margin to the base of cell M3-CuA1. Adults and male and female genitalia are illustrated and compared to other species of Mathania. In addition, we report Ligaria cuneifolia (R. et P.) Tiegh. (Loranthaceae) as host plant of M. hughesi.

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.

A multipronged approach to the study of peruvian ethnomedicinal plants: a legacy of the ICBG-Peru Project.

A multidisciplinary and international team of scientists was assembled in the early 1990s to conduct an ethnobotanical study of plants used by the Aguaruna people of the Peruvian Amazon forest. The initial ethnobotanical project, carried out under the auspices of an International Cooperative Biodiversity Grant (ICBG), led to the collection of approximately 4000 plant species. Some members of the original team of scientists have continued this collaboration by focusing on potential sources of new anticancer, anti-infective, and wound-healing agents. This effort has uncovered several secondary metabolites representing a wide variety of chemical diversity. In this short review we describe some bioactive compounds of interest as part of our continuing collaboration.