Cycad phylogeny predicts host plant use of Eumaeus butterflies.

Eumaeus butterflies are obligate herbivores of Zamia, the most diverse neotropical genus of cycads. Eumaeus-Zamia interactions have been characterized mainly for species distributed in North and Central America. However, larval host plant use by the southern Eumaeus clade remains largely unknown, precluding a comprehensive study of co-evolution between the genera. Here, we combine fieldwork with museum and literature surveys to expand herbivory records for Eumaeus from 21 to 38 Zamia species. We inferred a time-calibrated phylogeny of Eumaeus to test for distinct macroevolutionary scenarios of larval host plant conservatism and co-evolution. We found a remarkable coincidence between Eumaeus and Zamia diversification, with the butterfly stem group diverging at the same time as the most recent radiation of Zamia in the Miocene. Cophylogenetic reconciliation analyses show a strong cophylogenetic signal between cycads and their butterfly herbivores. Bipartite model-based approaches indicate that this is because closely related Zamia species are used by the same Eumaeus species, suggesting larval host plant resource tracking by the butterfly herbivores. Our results highlight a case of tight evolution between Eumaeus butterflies and cycads, pointing to the generality of correlated evolution and phylogenetic tracking in plant-herbivore interactions across seed plants.

Butterflies (Lepidoptera: Papilionoidea) of Iguaçu National Park and surrounding areas in southern Brazil: a long-term survey, with six new records for the Brazilian fauna / As Borboletas (Lepidoptera: Papilionoidea) do Parque Nacional do Iguaçu e arredores no Sul do Brasil: um inventário de longo prazo, com seis novos registros para a fauna brasileira

Abstract Iguaçu National Park is the second largest (1852.62 km²) protected area in the Atlantic Forest domain and harbors the largest area of semideciduous seasonal forest in Brazil. In this study, we present 795 subspecies and 787 species of butterflies that occur in this protected area and its surrounding areas, collected over 15 years and ten months using different non-standardized sampling methods. We also searched for additional records in the literature, entomological collections, and citizen science platforms on the internet. Among the sampled taxa, six are recorded for the first time in Brazil: Emesis orichalceus Stichel, 1916, Theope p. pakitza Hall & Harvey, 1998 (Riodinidae), Elbella v. viriditas (Skinner, 1920), Apaustus gracilis ssp. n. (Hesperiidae), Deltaya sp. n. (Nymphalidae), and Symbiopsis sp. n. (Lycaenidae). Another six are listed as endangered in lists of butterflies of conservation concern. The records for some species significantly increase previously documented distributions.

Resumo O Parque Nacional do Iguaçu é a segunda maior Unidade de Conservação (1.852,62 km²) no domínio Mata Atlântica, abrigando a maior área de Floresta Estacional Semidecídua no Brasil. Neste estudo apresentamos uma lista com 795 subespécies e 787 espécies de borboletas que ocorrem nesta Unidade de Conservação e seus arredores, coligida ao longo de 15 anos e dez meses através do uso de diferentes métodos de amostragem não padronizados. Nós também procuramos por registros adicionais na literatura, coleções entomológicas e plataformas de ciência cidadã na internet. Dentre os táxons amostrados, seis são registrados pela primeira vez para o Brasil: Emesis orichalceus Stichel, 1916, Theope p. pakitza Hall & Harvey, 1998 (Riodinidae), Elbella v. viriditas (Skinner, 1920), Apaustus gracilis ssp. n. (Hesperiidae), Deltaya sp. n. (Nymphalidae) e Symbiopsis sp. n. (Lycaenidae). Outras seis espécies são consideradas ameaçadas de extinção em listas de borboletas de interesse para a conservação. Os registros de algumas espécies aumentam significativamente as suas distribuições previamente documentadas.

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.

Cunoniaceae, nuevo registro de familia de plantas hospederas para los géneros altoandinos Penaincisalia y Micandra (Lycaenidae: Theclinae: Eumaeini) / Cunoniaceae, new family food plant record for the high altitude Andean genera Penaincisalia and Micandra (Lycaenidae: Theclinae: Eumaeini)

Resumen Orugas de dos especies altoandinas de Eumaeini (Lycaenidae: Theclinae) fueron halladas consumiendo hojas de Weinmannia crassifolia Ruiz & Pav., 1802 (Cunoniaceae). La colecta se realizó usando la técnica de colecta por golpeo. Esta familia de plantas es registrada por primera vez como planta alimenticia de los Eumaeini. Aunque no pudimos completar la crianza, las especies de Lycaenidae fueron identificadas mediante sus códigos de barra de ADN. Las orugas de Penaincisalia purpurea (K. Johnson, 1992) y Micandra dignota (Draudt, 1919) se describen e ilustran por primera vez. También se presentan datos sobre su ecología y relaciones con especies similares. Una tercera especie, registrada en estadio pupal en la misma planta de W. crassifolia fue un Nymphalidae, Perisama diotima (Hewitson, 1852).

Abstract Caterpillars of two Andean species of Eumaeini (Lycaenidae: Theclinae) were found eating leaves of Weinmannia crassifolia Ruiz & Pav., 1802 (Cunoniaceae). They were collected by "beating" the leaves. This plant family is reported for the first time as a food plant for Eumaeini. Although it was not possible to rear the caterpillars to the adult stage, we identified them using DNA barcodes. The caterpillars of Penaincisalia purpurea (K. Johnson, 1992) and Micandra dignota (Draudt, 1919) are described and illustrated for the first time. Data on their ecology and relationships with similar species are also presented. A third species collected on the same plant of W. crassifolia in the pupal stage was a Nymphalidae, Perisama diotima (Hewitson, 1852).

The butterflies of Cristalino Lodge, in the Brazilian southern Amazonia: An updated species list with a significant contribution from citizen science / As borboletas do Cristalino Lodge, no sul da Amazônia: Uma lista atualizada de espécies com contribuição significativa da ciência cidadã

Abstract The richest butterfly communities in the world are found in the Amazon rainforest. Despite of this, and the importance of species inventories for the knowledge of diversity patterns, there are few comprehensive lists of butterflies for localities in the Brazilian Amazon. Here, we present an updated list of the butterflies of Cristalino Lodge (Alta Floresta, Mato Grosso, Brazil), in southern Amazonia, based on specimens collected by researchers and photographic records taken by ecotourists, butterfly watchers, and tour guides. With 1010 species recorded, this is currently the largest list of butterflies published for a single locality in Brazil and the first to reach (and surpass) 1000 species, with more than one third of the records coming from citizen science. The region has about 29% of the butterfly species in Brazil and one of the greatest richnesses known in the country, inferior only to areas in the western Amazon. Its fauna is mainly composed of species widely distributed in lowland Amazonia, with the addition of some species typical of the Cerrado. It has a relatively low number of species of the tribe Ithomiini (Nymphalidae: Danainae), generally considered a good indicator of the total butterfly diversity in neotropical forests, which points to the need for caution when using a single taxonomic group as a surrogate of richness of entire communities. The present work highlights the importance of citizen science and ecotourism centers for inventories and data on species distribution in diverse tropical forests.

Resumo As comunidades de borboletas mais ricas do mundo são encontradas na Amazônia. Apesar disso, e da importância dos inventários de espécies para o conhecimento dos padrões de diversidade, existem poucas listas abrangentes de borboletas para localidades da Amazônia brasileira. Aqui, apresentamos uma lista atualizada das borboletas do Cristalino Lodge (Alta Floresta, Mato Grosso, Brasil), no sul da Amazônia, baseada em espécimes coletados por pesquisadores e em registros fotográficos feitos por ecoturistas, observadores de borboletas e guias turísticos. Com 1010 espécies registradas, essa é atualmente a maior lista de borboletas publicada para uma localidade no Brasil e a primeira a atingir 1000 espécies, sendo mais de um terço dos registros provenientes da ciência cidadã. A região apresenta cerca de 29% das espécies de borboletas do Brasil e uma das maiores riquezas conhecidas no país, inferior apenas a áreas no oeste da Amazônia. Sua fauna é composta principalmente por espécies amplamente distribuídas na planície amazônica, com adição de algumas típicas do Cerrado. Possui um número relativamente baixo de espécies da tribo Ithomiini (Nymphalidae: Danainae), que é geralmente considerada uma boa indicadora da riqueza total de borboletas em florestas neotropicais, o que aponta para a necessidade de cautela ao se usar um grupo taxonômico como previsor da riqueza de comunidades inteiras. O presente trabalho destaca a importância da ciência cidadã e dos centros de ecoturismo para inventários e dados sobre distribuição de espécies em florestas tropicais diversas.

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.

A switch to feeding on cycads generates parallel accelerated evolution of toxin tolerance in two clades of Eumaeus caterpillars (Lepidoptera: Lycaenidae).

We assembled a complete reference genome of Eumaeus atala, an aposematic cycad-eating hairstreak butterfly that suffered near extinction in the United States in the last century. Based on an analysis of genomic sequences of Eumaeus and 19 representative genera, the closest relatives of Eumaeus are Theorema and Mithras We report natural history information for Eumaeus, Theorema, and Mithras Using genomic sequences for each species of Eumaeus, Theorema, and Mithras (and three outgroups), we trace the evolution of cycad feeding, coloration, gregarious behavior, and other traits. The switch to feeding on cycads and to conspicuous coloration was accompanied by little genomic change. Soon after its origin, Eumaeus split into two fast evolving lineages, instead of forming a clump of close relatives in the phylogenetic tree. Significant overlap of the fast evolving proteins in both clades indicates parallel evolution. The functions of the fast evolving proteins suggest that the caterpillars developed tolerance to cycad toxins with a range of mechanisms including autophagy of damaged cells, removal of cell debris by macrophages, and more active cell proliferation.

Phylogenetic Classification of the Atlides Section of the Eumaeini (Lepidoptera, Lycaenidae).

This paper provides the first phylogenetic classification of the Atlides Section (Lycaenidae: Theclinae, Eumaeini).  It is based on a recently published morphological phylogenetic study, in which the Atlides Section proved to be monophyletic in all analyses. In particular, that study identified a hindwing cleft anal lobe in all members of the Atlides Section that is lacking in all other Eumaeini (except for some species of Panthiades Hübner). The included genera are Theritas Hübner; Arcas Swainson; Lucilda d'Abrera Bálint; Pseudolycaena Wallengren; Brangas Hübner; Atlides Hübner; and Denivia K. Johnson. Each is characterized by synapomorphies. Margaritheclus Bálint and Dabreras Bálint are new generic synonyms of Lucilda and Brangas, respectively. We recognize 71 species. Distribution and habitat are provided for each, and nomenclature is updated. Atlides centralis Salazar Henao is a new synonym of Atlides browni Constantino, Salazar K. Johnson; Brangas contrastus Bálint is a new synonym of Brangas felderi (Goodson); Denivia grava Bálint, K. Johnson Kroenlein is a new synonym of Denivia deniva (Hewitson); Denivia ponsanota Bálint, K. Johnson Kroenlein is a new synonym of Atlides zava (Hewitson); Denivia striata Bálint, K. Johnson Kroenlein is a new synonym of Atlides zava (Hewitson); Margaritheclus boliboyeri Bálint Wojtusiak is a new synonym of Margaritheclus boliboyerus Bálint Wojtusiak; and Theritas gozmanyi Bálint Wojtusiak is a new synonym of Theritas paupera (C. Felder R. Felder). New combinations are Lucilda margaritacea (Draudt), Lucilda danaus (C. Felder R. Felder), Lucilda dabrerus (Bálint), Lucilda boliboyerus (Bálint Wojtusiak), Denivia curitabaensis (K. Johnson), Denivia augustinula (Goodson), Denivia theocritus (Fabricius), Denivia augustula (W.F. Kirby), Denivia arene (Goodson), Denivia monica (Hewitson), Denivia hemon (Cramer), Denivia phegeus (Hewitson), Denivia acontius (Goodson), Denivia chaluma (Schaus), Denivia viresco (H.H. Druce), Denivia silma (Martins, Faynel Robbins), and Denivia lisus (Stoll).

A highly resolved food web for insect seed predators in a species-rich tropical forest.

The top-down and indirect effects of insects on plant communities depend on patterns of host use, which are often poorly documented, particularly in species-rich tropical forests. At Barro Colorado Island, Panama, we compiled the first food web quantifying trophic interactions between the majority of co-occurring woody plant species and their internally feeding insect seed predators. Our study is based on more than 200 000 fruits representing 478 plant species, associated with 369 insect species. Insect host-specificity was remarkably high: only 20% of seed predator species were associated with more than one plant species, while each tree species experienced seed predation from a median of two insect species. Phylogeny, but not plant traits, explained patterns of seed predator attack. These data suggest that seed predators are unlikely to mediate indirect interactions such as apparent competition between plant species, but are consistent with their proposed contribution to maintaining plant diversity via the Janzen-Connell mechanism.

Hairstreak butterflies (Lepidoptera, Lycaenidae) and evolution of their male secondary sexual organs.

Hairstreak butterflies in the Atlides Section of the Eumaeini are biologically notable for a diverse array of male secondary sexual organs. A "species recognition" hypothesis postulates that females use these organs to choose between conspecific and non-conspecific males, thereby promoting reproductive isolation. Alternately, a "sexual selection" hypothesis posits that females use these organs to choose among conspecific males. These hypotheses need not be mutually exclusive but make different predictions about the evolutionary gain and loss of male secondary sexual organs. We analysed most of the Atlides Section (Theclinae, Eumaeini) phylogenetically. Sister lineages were sympatric at 22 of 37 nodes. Nine evolutionary gains occurred in lineages that were sympatric with their phylogenetic sister, and one occurred in a lineage that was allopatric/parapatric with its sister. By contrast, seven of ten evolutionary losses occurred in lineages that were allopatric/parapatric with their sisters. These results are significantly different from those predicted by a sexual selection hypothesis. We conclude that male secondary sexual organs in the Atlides Section function primarily for species recognition and thereby promote sympatric diversification.

Butterflies (Lepidoptera: Papilionoidea) of an urban park in northeastern Brazil / As borboletas (Lepidoptera: Papilionoidea) de um parque urbano do nordeste do Brasil

Abstract Despite being one of the groups most well studied in Brazil, the butterfly fauna of northeastern Brazil, especially north of the mouth of São Francisco River, is poorly known. The aim of this study was to inventory the butterfly fauna in a remnant of Atlantic Forest in the state of Pernambuco as a contribution to understanding the distribution of these insects. The study was carried out in a 384.7 hectares fragment of Parque Estadual Dois Irmãos, in the metropolitan region of Recife. The butterflies were sampled monthly with insect nets from August 2011 to July 2012 and from January to August 2016. Additional records were obtained from two entomological collections and from two previous visits to the area in July 2003 and August 2006. A total of 273 species was sampled in 464 hours, with an additional 15 species recorded from the Entomological Collections, totaling 288 butterfly species recorded. The richest family was Hesperiidae (108 species), followed by Nymphalidae (80), Lycaenidae (43) Riodinidae (37), Pieridae (16) and Papilionidae (4). The fauna was dominated by generalist species with a broad geographical distribution, many of them commonly found on open areas, forest edges and secondary vegetation. Roeberella lencates (Hewitson, 1875) and Pheles atricolor atricolor (Butler, 1871) (Riodinidae) represent new records for northeastern Brazil.

Resumo Apesar de ser um dos grupos mais bem estudados no Brasil, as borboletas da região Nordeste, especialmente ao norte da foz do Rio São Francisco, são ainda pouco conhecidas. O objetivo deste estudo foi realizar um inventário da fauna de borboletas em um fragmento de Floresta Atlântica no estado de Pernambuco, como uma contribuição para compreensão sobre a distribuição desses insetos. O estudo foi realizado em um fragmento de Floresta Atlântica de 384.7 hectares do Parque Estadual Dois Irmãos, localizado na região metropolitana de Recife. As borboletas foram amostradas mensalmente, de agosto de 2011 a julho de 2012 e de janeiro a agosto de 2016, com auxílio de redes entomológicas. Registros adicionais foram obtidos de duas Coleções Entomológicas e de duas coletas prévias em julho de 2003 e agosto de 2006. Um total de 273 espécies foi registrado em 464 horas e 15 espécies adicionais registradas a partir de Coleções Entomológicas, totalizando 288 espécies de borboletas registradas. A família mais rica foi Hesperiidae (108 espécies), seguida por Nymphalidae (80), Lycaenidae (43), Riodinidae (37), Pieridae (16) e Papilionidae (4). A fauna de borboletas foi dominada por espécies generalistas com uma ampla distribuição geográfica, muitas delas encontradas comumente em áreas abertas, bordas de mata e em vegetação secundária. Roeberella lencates (Hewitson, 1875) e Pheles atricolor atricolor (Butler, 1871) (Riodinidae) representam um novo registro para o Nordeste do Brasil.

When COI barcodes deceive: complete genomes reveal introgression in hairstreaks.

Two species of hairstreak butterflies from the genus Calycopis are known in the United States: C. cecrops and C. isobeon Analysis of mitochondrial COI barcodes of Calycopis revealed cecrops-like specimens from the eastern US with atypical barcodes that were 2.6% different from either USA species, but similar to Central American Calycopis species. To address the possibility that the specimens with atypical barcodes represent an undescribed cryptic species, we sequenced complete genomes of 27 Calycopis specimens of four species: C. cecrops, C. isobeon, C. quintana and C. bactra Some of these specimens were collected up to 60 years ago and preserved dry in museum collections, but nonetheless produced genomes as complete as fresh samples. Phylogenetic trees reconstructed using the whole mitochondrial and nuclear genomes were incongruent. While USA Calycopis with atypical barcodes grouped with Central American species C. quintana by mitochondria, nuclear genome trees placed them within typical USA C. cecrops in agreement with morphology, suggesting mitochondrial introgression. Nuclear genomes also show introgression, especially between C. cecrops and C. isobeon About 2.3% of each C. cecrops genome has probably (p-value < 0.01, FDR < 0.1) introgressed from C. isobeon and about 3.4% of each C. isobeon genome may have come from C. cecrops. The introgressed regions are enriched in genes encoding transmembrane proteins, mitochondria-targeting proteins and components of the larval cuticle. This study provides the first example of mitochondrial introgression in Lepidoptera supported by complete genome sequencing. Our results caution about relying solely on COI barcodes and mitochondrial DNA for species identification or discovery.

Natural history of the mistletoe-feeding Thereus lomalarga (Lepidoptera, Lycaenidae, Eumaeini) in Colombia.

The natural history and morphology of the immature stages of the butterfly Thereus lomalarga Robbins, Heredia & Busby are described and illustrated. The food plant is Oryctanthus alveolatus (H.B.K.) Kuijt (Loranthaceae). Chaetotaxy of the first instar is described and compared with that of three locally studied Thereus species. Larvae have four instars, and the dorsal nectary organ becomes functional in the third instar. They are facultatively tended by ants belonging to seven genera that are attracted to O. alveolatus by floral disc nectaries, honeydew producing Hemiptera, and secretory wounds produced by Hemiptera on the fleshy inflorescence rachis. The average period from egg to eclosion under lab conditions was 35.68 days. Females emerged before males. Adults of both sexes feed on nectar from the flowers of the food plant and on hemipteran secretions; adults were not observed feeding on other flowers. Campopleginae and Chalcidinae were the most important parasitoids.

Complete genomes of Hairstreak butterflies, their speciation, and nucleo-mitochondrial incongruence.

Comparison of complete genomes of closely related species enables research on speciation and how phenotype is determined by genotype. Lepidoptera, an insect order of 150,000 species with diverse phenotypes, is well-suited for such comparative genomics studies if new genomes, which cover additional Lepidoptera families are acquired. We report a 729 Mbp genome assembly of the Calycopis cecrops, the first genome from the family Lycaenidae and the largest available Lepidoptera genome. As detritivore, Calycopis shows expansion in detoxification and digestion enzymes. We further obtained complete genomes of 8 Calycopis specimens: 3 C. cecrops and 5 C. isobeon, including a dry specimen stored in the museum for 30 years. The two species differ subtly in phenotype and cannot be differentiated by mitochondrial DNA. However, nuclear genomes revealed a deep split between them. Genes that can clearly separate the two species (speciation hotspots) mostly pertain to circadian clock, mating behavior, transcription regulation, development and cytoskeleton. The speciation hotspots and their function significantly overlap with those we previously found in Pterourus, suggesting common speciation mechanisms in these butterflies.

Phylogenetic position of Electrostrymon picoloro, a new high elevation hairstreak from western Colombia (Lepidoptera: Lycaenidae: Eumaeini).

The phylogenetic position of a new high-elevation butterfly species, which we describe as Electrostrymon picoloro Prieto & Robbins, new species, is assessed. The ventral wing pattern and genitalic structures of this species are unique among Electrostrymon. This species is also unique among Electrostrymon in that it occurs, so far as is known, only in habitats above 2800 m elevation. A phylogenetic analysis places this species in Electrostrymon as the sister of a multi-species clade. The ancestors of E. picoloro appear to have colonized its high elevation habitat relatively early in the phylogenetic history of Electrostrymon, but the species then did not recolonize lower elevation habitats nor did it diversify into other extant species.

Male secondary sexual structures and the systematics of the Thereus oppia species group (Lepidoptera, Lycaenidae, Eumaeini).

The Thereus oppia species group includes species with and without a scent pad, which is a histologically and morphologically characterized male secondary sexual structure on the dorsal surface of the forewing. To assess the hypothesis that these structures are lost evolutionarily, but not regained (Dollo's Law), the taxonomy of this species group is revised. Thereus lomalarga sp. n., and Thereus brocki sp. n., are described. Diagnostic traits, especially male secondary structures, within the Thereus oppia species group are illustrated. Distributional and biological information is summarized for each species. Three species have been reared, and the caterpillars eat Loranthaceae. An inferred phylogeny is consistent with the hypothesis that scent pads in the Thereus oppia species group have been lost evolutionarily twice (in allopatry), and not re-gained.

A butterfly with olive green eyes discovered in the United States and the Neotropics (Lepidoptera, Lycaenidae, Eumaeini).

We describe Ministrymon janevicroy Glassberg, sp. n., from the United States (Texas). Its wing pattern closely resembles that of the widespread and well-known lycaenid, Ministrymon azia (Hewitson). The new species is distinguished by the structure of its male and female genitalia, by the patterning of the ground color on the basal half of the ventral hindwing surface, and by the color of its eyes. Adults of Ministrymon janevicroy in nature have olive green eyes in contrast to the dark brown/black eyes of Ministrymon azia. Ministrymon janevicroy occurs in dry deciduous forest and scrub from the United States (Texas) to Costa Rica (Guanacaste) with disjunct populations on Curaçao and Isla Margarita (Venezuela). In contrast, Ministrymon azia occurs from the United States to southern Brazil and Chile in both dry and wet lowland habitats. Nomenclaturally, we remove the name Electrostrymon grumus K. Johnson & Kroenlein, 1993, from the synonymy of Ministrymon azia (where it had been listed as a synonym of Ministrymon hernandezi Schwartz & K. Johnson, 1992). We accord priority to Angulopis hernandezi K. Johnson & Kroenlein, 1993 over Electrostrymon grumus K. Johnson & Kroenlein, 1993, syn. n., which currently is placed in Ziegleria K. Johnson, 1993. The English name Vicroy's Ministreak is proposed for Ministrymon janevicroy. We update biological records of dispersal and caterpillar food plants, previously attributed to Ministrymon azia, in light of the new taxonomy.

Review of the species level taxonomy of the neotropical butterfly genus Oenomaus (Lycaenidae, Theclinae, Eumaeini).

Seven new species of the Neotropical hairstreak genus Oenomaus are described: Oenomaus mancha Busby & Faynel, sp. n. (type locality Ecuador); Oenomaus gwenish Robbins & Faynel, sp. n. (type locality Panama); Oenomaus lea Faynel & Robbins, sp. n. (type locality Ecuador); Oenomaus myrteana Busby, Robbins & Faynel, sp. n. (type locality Ecuador); Oenomaus mentirosa Faynel & Robbins, sp. n. (type locality Peru); Oenomaus andi Busby & Faynel, sp. n. (type locality Ecuador) and Oenomaus moseri Robbins & Faynel, sp. n. (type locality Brazil, Santa Catarina). For each new Oenomaus species, we present diagnostic characters and notes on its habitat and biology. We illustrate adults, genitalia, and distribution. New distributional and biological data are presented for 21 previously described Oenomaus species. Oenomaus melleus guyanensis Faynel, 2008 is treated as a new synonym of Oenomaus melleus melleus (Druce, 1907). Females are described and associated with males for ten species using a variety of factors, including mitochondrial COI DNA "barcode" sequences. We summarize the reasons why the number of recognized Oenomaus species has grown in the past decade from one species to 28 species. Finally, we overview the habitats that Oenomaus species occupy and note that the agricultural pest on Annonaceae, Oenomaus ortygnus, is the only Oenomaus species that regularly occurs in greatly disturbed habitats.

Description and phylogenetic analysis of the Calycopidina (Lepidoptera, Lycaenidae, Theclinae, Eumaeini): a subtribe of detritivores / Descrição e análise filogenética de Calycopidina (Lepidoptera, Lycaenidae, Theclinae, Eumaeini): uma subtribo de detritívoros

Description and phylogenetic analysis of the Calycopidina (Lepidoptera, Lycaenidae, Theclinae, Eumaeini): a subtribe of detritivores. The purpose of this paper is to establish a phylogenetic basis for a new Eumaeini subtribe that includes those lycaenid genera in which detritivory has been recorded. Morphological characters were coded for 82 species of the previously proposed "Lamprospilus Section" of the Eumaeini (19 of these had coding identical to another species), and a phylogenetic analysis was performed using the 63 distinct ingroup terminal taxa and six outgroups belonging to four genera. Taxonomic results include the description in the Eumaeini of Calycopidina Duarte & Robbins new subtribe (type genus Calycopis Scudder, 1876), which contains Lamprospilus Geyer, Badecla Duarte & Robbins new genus (type species Thecla badaca Hewitson), Arzecla Duarte & Robbins new genus (type species Thecla arza Hewitson), Arumecla Robbins & Duarte, Camissecla Robbins & Duarte, Electrostrymon Clench, Rubroserrata K. Johnson & Kroenlein revalidated status, Ziegleria K. Johnson, Kisutam K. Johnson & Kroenlein revalidated status, and Calycopis. Previous "infratribe" names Angulopina K. Johnson & Kroenlein, 1993, and Calycopina K. Johnson & Kroenlein, 1993, are nomenclaturally unavailable and polyphyletic as proposed. New combinations include Badecla badaca (Hewitson), Badecla picentia (Hewitson), Badecla quadramacula (Austin & K. Johnson), Badecla lanckena (Schaus), Badecla argentinensis (K. Johnson & Kroenlein), Badecla clarissa (Draudt), Arzecla arza (Hewitson), Arzecla tarpa (Godman & Salvin), Arzecla canacha (Hewitson), Arzecla calatia (Hewitson), Arzecla tucumanensis (K. Johnson & Kroenlein), Arzecla sethon (Godman & Salvin), Arzecla nubilum (H. H. Druce), Arzecla paralus (Godman & Salvin), Arzecla taminella (Schaus), Arzecla albolineata (Lathy), Electrostrymon denarius (Butler & H.Druce), Electrostrymon guzanta (Schaus), Electrostrymon perisus...

O objetivo desse trabalho é estabelecer a base filogenética para uma nova subtribo de Eumaeini incluindo os gêneros com registros de detritivoria. Foram codificados caracteres morfológicos para as 82 espécies previamente arroladas na Seção Lamprospilus de Eumaeini (19 delas tiveram codificação idêntica a outras espécies); posteriormente foi conduzida uma análise filogenética incluindo 63 táxons terminais do grupo interno e seis grupos-externos pertencentes a quatro gêneros. Como resultados taxonômicos incluem-se a descrição de Calycopidina Duarte & Robbins subtribo nova (gênero-tipo Calycopis Scudder, 1876) e o reconhecimento de Lamprospilus Geyer, Badecla Duarte & Robbins gênero novo (espécie-tipo Thecla badaca Hewitson), Arzecla Duarte & Robbins gênero novo (espécie-tipo Thecla arza Hewitson), Arumecla Robbins & Duarte, Camissecla Robbins & Duarte, Elecrostrymon Clench, Rubroserrata K. Johnson & Kroenlein revalidado, Ziegleria K. Johnson, Kisutam K. Johnson & Kroenlein revalidado e Calycopis. Nomenclaturalmente, Angulopina K. Johnson & Kroenlein, 1993 e Calycopina K. Johnson & Kroenlein, 1993 são consideradas inválidas e polifiléticas tal como foram propostas. Novas combinações incluem Badecla badaca (Hewitson), Badecla picentia (Hewitson), Badecla quadramacula (Austin & K. Johnson), Badecla lanckena (Schaus), Badecla argentinensis (K. Johnson & Kroenlein), Badecla clarissa (Draudt), Arzecla arza (Hewitson), Arzecla tarpa (Godman & Salvin), Arzecla canacha (Hewitson), Arzecla calatia (Hewitson), Arzecla tucumanensis (K. Johnson & Kroenlein), Arzecla sethon (Godman & Salvin), Arzecla nubilum (H. H. Druce), Arzecla paralus (Godman & Salvin), Arzecla taminella (Schaus), Arzecla albolineata (Lathy), Electrostrymon denarius (Butler & H. Druce), Electrostrymon guzanta (Schaus), Electrostrymon perisus (H. H. Druce), Rubroserrata mathewi (Hewitson), Rubroserrata ecbatana (Hewitson), Kisutam micandriana (K. Johnson) e Kisutam syllis (Godman...

Integration of DNA barcoding into an ongoing inventory of complex tropical biodiversity.

Inventory of the caterpillars, their food plants and parasitoids began in 1978 for today's Area de Conservacion Guanacaste (ACG), in northwestern Costa Rica. This complex mosaic of 120 000 ha of conserved and regenerating dry, cloud and rain forest over 0-2000 m elevation contains at least 10 000 species of non-leaf-mining caterpillars used by more than 5000 species of parasitoids. Several hundred thousand specimens of ACG-reared adult Lepidoptera and parasitoids have been intensively and extensively studied morphologically by many taxonomists, including most of the co-authors. DNA barcoding - the use of a standardized short mitochondrial DNA sequence to identify specimens and flush out undisclosed species - was added to the taxonomic identification process in 2003. Barcoding has been found to be extremely accurate during the identification of about 100 000 specimens of about 3500 morphologically defined species of adult moths, butterflies, tachinid flies, and parasitoid wasps. Less than 1% of the species have such similar barcodes that a molecularly based taxonomic identification is impossible. No specimen with a full barcode was misidentified when its barcode was compared with the barcode library. Also as expected from early trials, barcoding a series from all morphologically defined species, and correlating the morphological, ecological and barcode traits, has revealed many hundreds of overlooked presumptive species. Many but not all of these cryptic species can now be distinguished by subtle morphological and/or ecological traits previously ascribed to 'variation' or thought to be insignificant for species-level recognition. Adding DNA barcoding to the inventory has substantially improved the quality and depth of the inventory, and greatly multiplied the number of situations requiring further taxonomic work for resolution.