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Myrmecophilous butterflies can establish complex symbiotic relationships with ants. A caterpillar wandering among the brood of the aggressive ponerine ant Neoponera villosa was found inside the core of a nest built in the myrmecophytic bromeliad Aechmea bracteata. This is the first caterpillar found living inside a ponerine ant nest. Its DNA barcode was sequenced, and an integrative approach was used to identify it as Pseudonymphidia agave, a poorly known member of the subtribe Pachythonina in the riodinid tribe Nymphidiini. The cuticle of the tank-like caterpillar lacks projections or tubercles and is covered dorsally by specialized flat setae that form an armor of small plates. Ant-organs potentially related to caterpillar-ant signaling, such as perforated cupola organs and tentacle nectary organs, are present. These morphological traits, together with evidence of social integration (direct contact with host brood, protective morphology, slow movement, no host aggressiveness), suggest that P. agave is a symbiotic, social parasite of N. villosa, preying on its host brood. However, several knowledge gaps remain, including oviposition site, dependence on bromeliad association, steps to colony integration, and larval diet through development. Carnivory has been reported in all known members of the subtribe Pachythonina (caterpillars prey on honeydew-producing hemipterans) suggesting a shift to myrmecophagy inside the ant nests as a possible evolutionary transition.

The high resemblance between closely related mimetic species poses a challenge to taxonomists who describe and define these species. An emblematic example is found within Stalachtis Hubner, 1818 (Lepidoptera: Riodinidae), a small Neotropical genus of brightly colored butterflies engaged in mimicry rings. Its systematic position within Riodinidae is difficult to define based on morphology alone, and while its taxonomy is inflated due to intraspecific color polymorphisms, phenotypic variation might be concealing cryptic species. Thus, we examined the subspecific diversity within S. phlegia (Cramer, 1779) through an integrative approach based on both molecular and morphological data, aiming to investigate taxon limits. Both morphology and molecular data support the status of S. phlegia and Stalachtis susanna stat. rev. as distinct species. S. susanna stat. rev. is a medium sized butterfly with an orange-black wing color pattern with a few white dots. Furthermore, S. susanna stat. rev. has no subspecies, and is endemic to the Atlantic Forest, while the four subspecies within S. phlegia are widely distributed throughout the Amazon, the Cerrado savanna and in parts of the Atlantic Forest. We consider S. phlegia a promising model for studies of the effect of deforestation on mimetic color patterns in the Amazon–Cerrado transition.

Myrmecophilous butterflies can establish complex symbiotic relationships with ants. An elusive symbiotic caterpillar wandering among the brood of the aggressive ponerine ant, Neoponera villosa, was found inside the core of a nest built in the myrmecophytic bromeliad Aechmea bracteata. This is the first caterpillar found in symbiosis with a species of the subfamily Ponerinae. Tissue sample was barcoded, and an integrative approach was used to identify it as Pseudonymphidia agave, a poorly known Pachythonina member in the riodinid myrmecophilous tribe Nymphidiini. The caterpillar has a general tank-like body morphology, without projections or tubercles, and is covered dorsally by specialized flat setae that form an armor of small plates. Ant-organs potentially related to the caterpillar-ant signaling, such as perforated cupola organs and tentacle nectary organs, are present. These functional traits, together with the stable social integration, suggest that P. agave is a specialized social parasite of N. villosa. However, several knowledge gaps remain, notably the oviposition site, dependence on bromeliad association, steps to colony integration, and the larval diet through ontogeny. Carnivory has been reported in all known members of the Pachythonina subtribe (caterpillars prey on honeydew-producing hemipterans), and a shift to myrmecophagy inside the ant nests is a possible evolutionary step.

The need for robust estimates of times of divergence is essential for downstream analyses, yet assessing this robustness is still rare. We generated a time-calibrated genus-level phylogeny of butterflies (Papilionoidea), including 994 taxa, up to 10 gene fragments and an unprecedented set of 12 fossils and 10 host-plant node calibration points. We compared marginal priors and posterior distributions to assess the relative importance of the former on the latter. This approach revealed a strong influence of the set of priors on the root age but for most calibrated nodes posterior distributions shifted from the marginal prior, indicating significant information in the molecular data set. Using a very conservative approach we estimated an origin of butterflies at 107.6 Ma, approximately equivalent to the latest Early Cretaceous, with a credibility interval ranging from 89.5 Ma (mid Late Cretaceous) to 129.5 Ma (mid Early Cretaceous). In addition, we tested the effects of changing fossil calibration priors, tree prior, different sets of calibrations and different sampling fractions but our estimate remained robust to these alternative assumptions. With 994 genera, this tree provides a comprehensive source of secondary calibrations for studies on butterflies.

We evaluated how the taxonomic, phylogenetic and functional diversities of butterflies and their community-weighted traits are affected by urbanization in the southeastern Brazilian Atlantic Forest. For this purpose, a dataset of Nymphalidae species distributed across 15 urban, semiurban, and rural fragments was analyzed. Urbanization was defined by a set of environmental variables. Furthermore, the total area of each fragment was also considered in the analyses but did not influence the results, in which disturbance level and patch connectivity drove the environmental variation across the urban matrix. Species diversity increased towards the more connected fragments, while phylogenetic and functional diversity did not vary in relation to urbanization. A high forewing:hindwing ratio and the frequency of tiger-like wings were positively related to the urban fragments, while a low forewing:hindwing ratio and iridescent wings were related to the semiurban and rural fragments. The suitability of highly interconnected rural habitats for the maintenance of butterfly diversity was corroborated as expected. Nonetheless, our results also showed that semiurban fragments preserved the ecologically relevant traits of butterflies related to forested habitats, expressed in butterfly groups possibly linked with dispersal capability to avoid predation. Careful management of semiurban fragments and urban landscaping, including highly structured and native vegetation outside urban parks, may increase the functional and taxonomic diversities or at least maintain the current levels of functionality in the urban matrix. Thus, it is possible to preserve the biological diversity of native fauna and flora and recover relevant ecosystem services, ensuring the conservation of Neotropical urban centers.

Species losses are increasing and may have an impact on our understanding of patterns of evolutionary pathways and phylogenetic relationships among the groups being lost. The knowledge of such patterns can contribute to preventing future losses by identifying which lineages have higher or lower diversification rates, thus informing conservation strategies. Recent years have seen a significant growth in studies of butterfly systematics, allowing a better understanding of evolutionary relationships among most groups and revealing significant taxonomic chaos in several groups. One of the latter groups is the nymphalid subtribe Euptychiina (Satyrinae), which has been shown to include a number of non-monophyletic genera based on recent molecular phylogenetic analyses. Among others, these genera include Yphthimoides, which is widespread throughout the Neotropical region but particularly diverse in the southeastern Neotropics, and a pair of related genera, Pharneuptychia Forster, 1964 and Moneuptychia Forster, 1964. Using molecular data, this study scope and aims were to provide a phylogenetic hypothesis that corroborates Yphthimoides as presently conceived being non-monophyletic, a result reinforced by a comparative study of the male genitalic morphology. Our results also show that Pharneuptychia and Moneuptychia, plus a species misplaced elsewhere in the Euptychiina, Euptychoides castrensis (Schaus, 1902), form a well supported clade, and that the latter 'species' is a complex of cryptic species. We therefore propose a number of taxonomic rearrangements in the present work to resolve these issues: Yphthimoides eriphule (A. Butler, 1867) will be moved to a new genus; Y. affinis (A. Butler, 1867), Y. maepius (Godart, [1824]), Y. mimula (Hayward, 1954), Y. neomaenas (Hayward, 1967) and Y. mythra (Weymer, 1911) are being transferred to Malaveria ViloriaBenmesbah, 2021; Pharneuptychia innocentia (Godart, [1824]) will be moved to another genus to be described; and Euptychoides castrensis, Pharneuptychia romanina (Bryk, 1953) and Yphthimoides viviana (Romieux, 1927) are being moved to Moneuptychia. The dating of divergences points to a split between the ancestral lineage of Yphthimoides and its sister group, Carminda Ebert and Dias, inDias 1998, during the last half of the Miocene, around 11.86 Mya, and to the diversification of the Pharneuptychia during the same time 11.35 (±3.52) Mya. Biogeographic analysis showed that the most recent common ancestor of Yphthimoides started to diversify either in the the Brazilian Cerrado savannas or in a combined area of Cerrado and South Atlantic Forest, with a possible change in the ancestral habitat of Carminda. Furthermore, ancestral character mapping favors a savanna origin hypothesis over a forest origin hypothesis.

Atlantic Forest White Morpho butterflies, currently classified as Morpho epistrophus and M. iphitus, are endemic to the Atlantic Forest, where they are widely distributed throughout heterogeneous environmental conditions. Studies with endemic butterflies allow to elucidate questions on both patterns of diversity distribution and current and past processes acting on insect groups in this biodiversity hotspot. In the present study, we characterized one mtDNA marker (COI sequences) and developed 11 polymorphic loci of microsatellite for 22 sampling locations distributed throughout the entire Atlantic Forest domain. We investigated both the taxonomic limits of taxa classified as White Morpho and the structure and distribution of the genetic diversity throughout their populations. Genetic markers and distribution data failed to identify species diversification, population structure, or isolation among subpopulations attributed to different taxa proposed for the White Morpho, suggesting that the current distinction between two species is unreasonable. The Bayesian coalescent tree based on COI sequences also failed to recover monophyletic clades for the putative species, and pointed instead to a north-south oriented pattern of genetic structure, with the northern clade coalescing later than the southern clade. Northern samples also showed more intragroup structure than southern samples based on mtDNA data. Clustering tests based on microsatellites indicated the existence of three genetic clusters, with turnover between the states of Parana and Sao Paulo. The north-south pattern found for the White Morpho populations is showed for the first time to a endemic AF insect and coincides with the two different bioclimatic domains previously described for vertebrates and plants. Population structure observed for these butterflies is related to climate- and landscape-associated variables, mainly precipitation and elevation.

In the first paragraph of page 627, under section ‘Diversity measures and functional composition’, the sentence "Positive or negative SES.MPD/MFD values indicate phylogenetic/functional clustering or overdispersion of species in the fragment, respectively (Webb et al. 2002).” was published incorrectly. The correct sentence should read "Positive or negative SES.MPD/MFD values indicate phylogenetic/functional overdispersion or clustering of species in the fragment, respectively (Webb et al. 2002).”

Parides ascanius (Lepidoptera: Papilionidae) is a butterfly endemic to the sand forests (“restingas”) of one of the most populated areas of Brazil (from Rio de Janeiro state to South Espirito Santo state), and was the first invertebrate officially recognized as being threatened in Brazil. Here we present a panel of eight polymorphic microsatellite loci and partial sequences of mitochondrial gene COI aiming to characterize this butterfly’s genetic diversity and understand its distribution among the extant populations. We estimate FST metrics, migration rates, cluster assignment, and spatial structure of genetic diversity. FST and statistics indicate low genetic structure and no evidence for endogamy, with all populations connected by high migration rates. Seven populations have low permanence rates (68–75 %) with increased migration probabilities for all populations. One population displays higher permanence rate (87.7 %), as the metropolitan matrix isolates it. Spatial analysis shows a global structure around the city of Rio de Janeiro and the Guanabara Bay; assignment analysis recovers six clusters evenly spread among sampled populations. These findings are consistent with a natural scenario of metapopulation dynamics for P. ascanius, with low genetic diversity and no endogamy, but progressively isolated by the metropolitan matrix. Conservation efforts should focus in connecting the isolated population, broaden the searches for new populations, and preserve all extant habitat patches where P. ascanius still occurs.

The need for robust estimates of times of divergence is essential for downstream analyses, yet assessing this robustness is still rare. We generated a time-calibrated genus-level phylogeny of butterflies (Papilionoidea), including 994 taxa, up to 10 gene fragments and an unprecedented set of 12 fossils and 10 host-plant node calibration points. We compared marginal priors and posterior distributions to assess the relative importance of the former on the latter. This approach revealed a strong influence of the set of priors on the root age but for most calibrated nodes posterior distributions shifted from the marginal prior, indicating significant information in the molecular dataset. We also tested the effects of changing assumptions for fossil calibration priors and the tree prior. Using a very conservative approach we estimated an origin of butterflies at 107.6 Ma, approximately equivalent to the Early Cretaceous–Late Cretaceous boundary, with a credibility interval ranging from 89.5 Ma (mid Late Cretaceous) to 129.5 Ma (mid Early Cretaceous). This estimate was robust to alternative analyses changing core assumptions. With 994 genera, this tree provides a comprehensive source of secondary calibrations for studies on butterflies.

The riodinid Rhetus belphegor (Westwood) (n. comb., previously in the genus Nirodia) is a critically endangered butterfly confined to the “campos rupestres”; a high-altitude rocky outcrop vegetation from southeast Brazil. The aim of this study is to unveil its biology and evaluate its systematic position. Based on museum data and public contribution of data (in the context of citizen science), R. belphegor is restricted to the “Espinhaço Mountain Chain”, and occurs exclusively above 1000 m. Adults were found resting upside down on rock walls. Females searched for host plants during the hottest hours of the day, depositing 1–2 eggs on leaves of the herbaceous subshrub Microstachys serrulata (Euphorbiaceae). The non-myrmecophilous larvae developed through six instars and the developmental time from egg to adult was ~50 days. Larvae are covered with abundant setae. Morphology of immature stages and molecular phylogenetic analysis showed that Nirodia is part of Rhetus, justifying the generic change. Our data supports that Nirodia is the only species in its clade associated with high mountains, in contrast to its lowland congeners. The description of the immature biology and clarification on its systematic position are essential steps for the establishment of better and more effective conservation efforts for this magnificent Brazilian butterfly., LAK was supported by CNPq (163119/2013-9) and CAPES (3200-14-0). NS was supported by CNPq (141254/2013-0) and CAPES (3700/14-3). AVLF thanks the CNPq (fellowship 302585/2011-7), the National Science Foundation (DEB-1256742) and the BIOTA-FAPESP Program (11/50225-3). NW acknowledges funding from the Academy of Finland (265511). This publication is part of the RedeLep “Rede Nacional de Pesquisa e Conservação de Lepidópteros” SISBIOTA-Brasil/CNPq (563332/2010-7), of the project “Identificação Molecular de Biodiversidade de Invertebrados Terrestres” (Grant 564954/2010-1) included in the “Rede Nacional de Identificação Molecular da Biodiversidade—BR-BoL” (MCT/CNPq/FNDCT 50/2010), and of the collaborative grant ‘Dimensions US-BIOTA São Paulo: A multidisciplinary framework for biodiversity prediction in the Brazilian Atlantic forest hotspot’, US NSF, NASA and FAPESP (Grant 2013/50297-0).

Sertania Callaghan & Kaminski, gen. nov. is proposed for the Neotropical riodinid butterfly species “Emesis” guttata (Stichel, 1910). Both, morphological (wing shape pattern and characters in the male genitalia) and molecular data support this as a new genus distinct from all other described genera of Riodinidae. Based on morphology and molecular data, two described subspecies were raised to species status: Sertania jaibensis (Callaghan & Soares, 2001), comb. nov., stat. nov., and Sertania lambedor (P. Jauffret, J. Jauffret & Pessôa, 2008), comb. nov., stat. nov. In addition, taxonomic and ecological notes, distribution map and illustrations of adults are provided. Species in this genus are restricted to xeric environments along the South American ‘dry diagonal’ and can be a promising model for biogeography studies of Neotropical open vegetal physiognomies.

The genus Hermeuptychia is common and widespread through the Americas, from Argentina to the southern United States of America. All eight recognized species within Hermeuptychia are small and brown, with very similar interspecific external morphologies and intraspecifically variable ocelli patterns that render taxonomic identification based on morphology difficult. In our study, we surveyed variability within Hermeuptychia, and evaluated species boundaries based on molecular data (sequences of the ‘barcode’ mitochondrial DNA COI gene) and morphology (mainly male genitalia), using a phylogenetic approach. We found eight DNA-based and 12 morphological groups in our sampling. Species names were assigned based mainly on comparisons with male genitalia morphology descriptions corresponding to name-bearing type specimens. Morphological and DNA variability were highly congruent, with the exception of group H, the Hermeuptychia cucullina complex. Also, the barcode region showed a clear threshold for intra- and interspecific mean distances around 2%. Based on these results, we circumscribe the species boundaries in the genus Hermeuptychia and discuss conflicts between mitochondrial genes and classic morphological approaches for identifying and delimiting species. Our study revealed cryptic diversity within an ubiquitous genus of Neotropical butterflies.

Submitted by Guilherme Lemeszenski (guilherme@nead.unesp.br) on 2013-08-22T19:05:05Z No. of bitstreams: 1 S1415-47572011000400028.pdf: 1141453 bytes, checksum: d1631d076ee46ec0e4fd65e336087e5e (MD5) Made available in DSpace on 2013-08-22T19:05:05Z (GMT). No. of bitstreams: 1 S1415-47572011000400028.pdf: 1141453 bytes, checksum: d1631d076ee46ec0e4fd65e336087e5e (MD5) Previous issue date: 2011-01-01 Made available in DSpace on 2013-09-30T20:08:48Z (GMT). No. of bitstreams: 2 S1415-47572011000400028.pdf: 1141453 bytes, checksum: d1631d076ee46ec0e4fd65e336087e5e (MD5) S1415-47572011000400028.pdf.txt: 27402 bytes, checksum: 491d55e9c41d523f7f74808b54ab9f19 (MD5) Previous issue date: 2011-01-01 Submitted by Vitor Silverio Rodrigues (vitorsrodrigues@reitoria.unesp.br) on 2014-05-20T15:17:32Z No. of bitstreams: 2 S1415-47572011000400028.pdf: 1141453 bytes, checksum: d1631d076ee46ec0e4fd65e336087e5e (MD5) S1415-47572011000400028.pdf.txt: 27402 bytes, checksum: 491d55e9c41d523f7f74808b54ab9f19 (MD5) Made available in DSpace on 2014-05-20T15:17:32Z (GMT). No. of bitstreams: 2 S1415-47572011000400028.pdf: 1141453 bytes, checksum: d1631d076ee46ec0e4fd65e336087e5e (MD5) S1415-47572011000400028.pdf.txt: 27402 bytes, checksum: 491d55e9c41d523f7f74808b54ab9f19 (MD5) Previous issue date: 2011-01-01 Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) The applicability of mitochondrial nad6 sequences to studies of DNA and population variability in Lepidoptera was tested in four species of economically important moths and one of wild butterflies. The genetic information so obtained was compared to that of cox1 sequences for two species of Lepidoptera. nad6 primers appropriately amplified all the tested DNA targets, the generated data proving to be as informative and suitable in recovering population structures as that of cox1. The proposal is that, to obtain more robust results, this mitochondrial region can be complementarily used with other molecular sequences in studies of low level phylogeny and population genetics in Lepidoptera. Universidade de São Paulo Escola Superior de Agricultura Luiz de Queiroz Departamento de Entomologia e Acarologia Universidade Estadual Paulista Júlio de Mesquita Filho Instituto de Biociências Departamento de Zoologia Universidade Estadual de Campinas Departamento de Biologia Animal Programa de Pós-Graduação em Ecologia Monsanto do Brazil Universidade Estadual Paulista Júlio de Mesquita Filho Instituto de Biociências Departamento de Zoologia CAPES: 0103/08-0 CNPq: 578509/2008-3 CNPq: 480619/2008-5