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

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

Wing pattern diversity in Eunica butterflies (Nymphalidae: Biblidinae): phylogenetic analysis implies decoupled adaptive trends in dorsal sexual dimorphism and ventral eyespot evolution.

Butterfly eyespots are wing patterns reminiscent of vertebrate eyes, formed by concentric rings of contrastingly coloured scales. Eyespots are usually located close to the wing margin and often regarded as the single most conspicuous pattern element of butterfly wing colour displays. Recent efforts to understand the processes involved in the formation of eyespots have been driven mainly by evo-devo approaches focused on model species. However, patterns of change implied by phylogenetic relationships can also inform hypotheses about the underlying developmental mechanisms associated with the formation or disappearance of eyespots, and the limits of phenotypic diversity occurring in nature. Here we present a combined evidence phylogenetic hypothesis for the genus Eunica, a prominent member of diverse Neotropical butterfly communities, that features notable variation among species in eyespot patterns on the ventral hind wing surface. The data matrix consists of one mitochondrial gene region (COI), four nuclear gene regions (GAPDH, RPS5, EF1a and Wingless) and 68 morphological characters. A combined cladistic analysis with all the characters concatenated produced a single most parsimonious tree that, although fully resolved, includes many nodes with modest branch support. The phylogenetic hypothesis presented corroborates a previously proposed morphological trend leading to the loss of eyespots, together with an increase in the size of the conserved eyespots, relative to outgroup taxa. Furthermore, wing colour pattern dimorphism and the presence of androconia suggest that the most remarkable instances of sexual dimorphism are present in the species of Eunica with the most derived eyespot patterns, and are in most cases accompanied by autapomorphic combinations of scent scales and "hair pencils". We discuss natural and sexual selection as potential adaptive explanations for dorsal and ventral wing patterns.

Reassessment of the type locality of Euptychiastigmatica Godman, 1905, with the description of two new sibling species from Amazonia (Lepidoptera, Nymphalidae, Satyrinae, Satyrini).

A brief historical review regarding the type locality of Euptychiastigmatica Godman, 1905 was conducted, which suggests that its type locality is actually Rio de Janeiro, Brazil, rather than northeastern Argentina, as previously purported. Consequently, E.stigmatica and its senior synonym E.cyanites Butler, 1871, are regarded to be two species-group names representing a taxon in the euptychiine genus Caeruleuptychia Forster, 1964 known from the Brazilian Atlantic Forest. A lectotype is designated for E.cyanites. Additionally, two closely related species are named and described using an integrative approach with morphological and molecular evidence. Caeruleuptychiaharrisi Nakahara & Freitas, sp. nov. and C.aemulatio Nakahara & Willmott, sp. nov. both occur in Amazonia and COI barcode data recovered these taxa as part of the caerulea clade of Caeruleuptychia.

Mutualistic interactions shape global spatial congruence and climatic niche evolution in Neotropical mimetic butterflies.

Understanding the mechanisms underlying species distributions and coexistence is both a priority and a challenge for biodiversity hotspots such as the Neotropics. Here, we highlight that Müllerian mimicry, where defended prey species display similar warning signals, is key to the maintenance of biodiversity in the c. 400 species of the Neotropical butterfly tribe Ithomiini (Nymphalidae: Danainae). We show that mimicry drives large-scale spatial association among phenotypically similar species, providing new empirical evidence for the validity of Müller's model at a macroecological scale. Additionally, we show that mimetic interactions drive the evolutionary convergence of species climatic niche, thereby strengthening the co-occurrence of co-mimetic species. This study provides new insights into the importance of mutualistic interactions in shaping both niche evolution and species assemblages at large spatial scales. Critically, in the context of climate change, our results highlight the vulnerability to extinction cascades of such adaptively assembled communities tied by positive interactions.

The sunny butterflies: new species of high montane pierids of the Catasticta poujadei group from the Peruvian Andes (Pieridae, Pierinae, Aporiina).

The Catasticta poujadei group, within the subgenus Hesperochoia Reissinger, is revised. Two new species, C. copernicus Pyrcz & Fhraeus sp. nov., and C. buszkoi Boyer & Pyrcz sp. nov. occurring near the timberline in Junn and Apurmac are described. Catasticta eximia Rber is reinstated as a species separate from C. poujadei, and a new subspecies, C. eximia tapuna ssp. nov., is described. The affinities of the species of the C. poujadei group are evaluated based on COI barcodes. Their distribution and habitats are described.

The benefits of contributing to the citizen science platform iNaturalist as an identifier.

As the number of observations submitted to the citizen science platform iNaturalist continues to grow, it is increasingly important that these observations can be identified to the finest taxonomic level, maximizing their value for biodiversity research. Here, we explore the benefits of acting as an identifier on iNaturalist.

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.

Revision of the "celia clade" of Pseudodebis Forster, 1964, with Two New Species and Notes on Papilio phorcys Fabricius, 1793 (Lepidoptera: Nymphalidae: Satyrinae).

The species-level classification of the "celia clade" of the nymphalid butterfly genus Pseudodebis Forster, 1964, is revised as part of ongoing revisionary work on this genus. The "celia clade" contains three species, of which two, Pseudodebis darrenthroopi Nakahara & Willmott, n. sp. and P. tigrillo Nakahara & Willmott, n. sp., are described and named herein based on morphology and molecular data. Consequently, we increase the described species diversity of Pseudodebis to 13, with a remarkable six species occurring in the trans-Andean region. We discuss five specific epithets associated with the clade and designate a neotype for Papilio celia Cramer, 1780, and lectotype for Taygetis keneza Butler, 1869, based on the same specimen, thus retaining the status of the former name as a senior objective synonym. We also provide a brief historical review for Papilio phorcys Fabricius, 1793, an enigmatic name currently synonymized under Papilio celia. Nevertheless, we were unable to locate the syntype(s) for this name and the identity of Papilio phorcys remains uncertain, so we retain the current synonymy as a parsimonious approach.

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.

Fifty years without a name: a new species of Splendeuptychia Forster (Lepidoptera: Nymphalidae: Satyrinae).

A new satyrine butterfly species, Splendeuptychia tupinamba Freitas, Huertas Rosa, sp. nov. (Nymphalidae: Satyrinae), is described. This species is found throughout a large geographical range in Brazil, Argentina and Paraguay, predominantly in the Cerrado domain, with some records in neighboring Amazonia and Atlantic Forest. Morphology and molecular data indicate that this species is part of a clade that includes Splendeuptychia ashna (the type species of the genus Splendeuptychia), and several species placed in the recently described genus Nubila Viloria, Andrade Henao, 2019.

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

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

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.

Hard to catch: experimental evidence supports evasive mimicry.

Most research on aposematism has focused on chemically defended prey, but the signalling difficulty of capture remains poorly explored. Similar to classical Batesian and Müllerian mimicry related to distastefulness, such 'evasive aposematism' may also lead to convergence in warning colours, known as evasive mimicry. A prime candidate group for evasive mimicry are Adelpha butterflies, which are agile insects and show remarkable colour pattern convergence. We tested the ability of naive blue tits to learn to avoid and generalize Adelpha wing patterns associated with the difficulty of capture and compared their response to that of birds that learned to associate the same wing patterns with distastefulness. Birds learned to avoid all wing patterns tested and generalized their aversion to other prey to some extent, but learning was faster with evasive prey compared to distasteful prey. Our results on generalization agree with longstanding observations of striking convergence in wing colour patterns among Adelpha species, since, in our experiments, perfect mimics of evasive and distasteful models were always protected during generalization and suffered the lowest attack rate. Moreover, generalization on evasive prey was broader compared to that on distasteful prey. Our results suggest that being hard to catch may deter predators at least as effectively as distastefulness. This study provides empirical evidence for evasive mimicry, a potentially widespread but poorly understood form of morphological convergence driven by predator selection.

Solving the cryptic diversity of the genus Manerebia Staudinger in northern Peru: description of new species and considerations on the biogeographical role of the Huancabamba Deflection (Nymphalidae: Satyrinae: Pronophilina).

The Huancabamba Deflection in the Andes of northern Peru and southern Ecuador is a pivotal area for Neotropical biogeography, where exceptional biodiversity coincides with high rates of endemism. These characteristics are well illustrated within the montane butterfly genus Manerebia Staudinger (Nymphalidae, Satyrinae). Here, six new, apparently endemic species, and two new subspecies, are described from this region: M. inducta Pyrcz Willmott n. sp., M. ronda Pyrcz Boyer, n. sp., M. ronda amplia Pyrcz Boyer, n. ssp., M. punku Pyrcz Farfn n. sp., M. huamanii Cerdea Pyrcz n. sp., M. placida Pyrcz Willmott n. sp., M. granatus Willmott, Radford Pyrcz n. sp. and M. prattorum udima Pyrcz Boyer, n. ssp. A total of 22 species of Manerebia is reported from the study region, as much as half the total number of species in this genus distributed throughout the tropical Andes. The alpha-taxonomy of Manerebia is particularly demanding, as colour patterns have apparently converged among different species making identification virtually impossible in some cases without comparison of their genitalia, which prove taxonomically and phylogenetically highly valuable. On the other hand, several species differ markedly in wing colour patterns and occur at different elevations but have identical genitalia. Furthermore, our data show that DNA barcoding is only partly viable as an alpha-taxonomic tool, as some markedly different species of Manerebia, in terms of external morphology and ecological preferences, are genetically not separable using only COI markers. On the other hand, several species barely differing morphologically are identified based on barcode. Barcodes of 26 species, including 18 from the northern Andes, are studied here, showing strong differences within some taxa previously considered conspecific based on morphological characters, suggesting that their taxonomic status needs to be re-evaluated. In particular, M. trimaculata and the widely distributed polytypic M. inderena may be highly variable species or complexes of several species. A total of 16 species are found north of the Chamaya valley in southern Ecuador and extreme northern Peru, compared to 14 species south of it in northern Peru. The faunal (Jaccard) similarity coefficient of the two adjacent regions is low (0.3), reflecting the role of the Huancabamba Deflection as a biogeographical barrier.

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

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

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

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

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

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

A revision of the new genus Amiga Nakahara, Willmott & Espeland, gen. n., described for Papilioarnaca Fabricius, 1776 (Lepidoptera, Nymphalidae, Satyrinae).

We here propose a new, monotypic genus, Amiga Nakahara, Willmott & Espeland, gen. n., to harbor a common Neotropical butterfly, described as Papilioarnaca Fabricius, 1776, and hitherto placed in the genus Chloreuptychia Forster, 1964. Recent and ongoing molecular phylogenetic research has shown Chloreuptychia to be polyphyletic, with C.arnaca proving to be unrelated to remaining species and not readily placed in any other described genus. Amigaarnaca gen. n. et comb. n. as treated here is a widely distributed and very common species ranging from southern Mexico to southern Brazil. A neotype is designated for the names Papilioarnaca and its junior synonym, Papilioebusa Cramer, 1780, resulting in the treatment of the latter name as a junior objective synonym of the former. A lectotype is designated for Euptychiasericeella Bates, 1865, which is treated as a subspecies, Amigaarnacasericeella (Bates, 1865), comb. n. et stat. n., based on molecular and morphological evidence. We also describe two new taxa, Amigaarnacaadela Nakahara & Espeland, ssp. n. and Amigaarnacaindianacristoi Nakahara & Marín, ssp. n., new subspecies from the western Andes and eastern Central America, and northern Venezuela, respectively.

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.