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Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L., Willmott, Keith R. (2023): Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae). Zoological Research 2023: 1-73, DOI: 10.1111/syen.12590

We here establish a new genus in the nymphalid butterfly subtribe Euptychiina, Cisandina Nakahara & Espeland, n. gen. to harbor five species hitherto placed within two polyphyletic genera, namely Magneuptychia Forster, 1964 and Euptychoides Forster, 1964. We compiled data from over 350 specimens in 17 public and private collections, as well as DNA sequence data for all relevant species, to revise the species-level classification of this new genus. According to our multi-locus molecular phylogeny estimated with the maximum likelihood approach, Cisandina lean. comb., Cisandina philippan. comb. & reinst. stat., Cisandina fidan. comb., Cisandina sanmarcosn. comb., and Cisandina trinitensisn. comb. are proposed as new taxonomic combinations, since these species are distantly related to the type species of Magneuptychia and Euptychoides and cannot reasonably be accommodated in any other genus. Lectotypes are designated for Papilio lea Cramer, 1777, Papilio junia Cramer, 1780, Euptychia philippa Butler, 1867, and Eupytchia fida Weymer, 1911. Two new species of Cisandinan. gen. are named and described herein, C. esmeralda Nakahara & Barbosa, n. sp. and C. castanya Lamas & Nakahara, n. sp., increasing the described species diversity of the genus to seven. The immature stages of C. castanyan. sp. and C. philippan. comb. & reinst. stat. are documented along with their natural hostplants, representing the first two species of the genus with known life history information. We describe a new subspecies, Cisandina fida directa Nakahara & Willmott, n. ssp., based on a limited number of specimens from southern Ecuador and central Peru. We were unable to obtain genetic data for the nominate race of C. fidan. comb., and thus, this taxonomic hypothesis is currently based solely on phenotypic characters.

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.

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.

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.

A new species of the butterfly satyrine subtribe Euptychiina is described: Carminda surpresa sp. nov. This species is apparently restricted to high altitude wet grasslands in southeastern Brazil. The description is based on morphological features, mainly from the wings, and male and female genitalia, and molecular data were also obtained and are used to validate the new species. Information about the geographic distribution, habitat and immature stages morphology is also provided. The systematic position of C. surpresa sp. nov. is discussed based on a molecular analysis which includes all described species of Carminda in addition to other species of Satyrinae.

A systematic revision of Paryphthimoides, a satyrine genus widely distributed in Central and South America, is presented based on an integrative approach. Molecular phylogeny inferred using multiple methods with nuclear (GAPDH, RpS5) and mitochondrial (COI) genes indicates that Paryphthimoides, as currently circumscribed, is polyphyletic. A new taxonomic arrangement is proposed reflecting the monophyly based on molecular and morphological evidence. Paryphthimoides now comprises 14 taxa, namely: P. poltys poltys (Prittwitz, 1865), P. poltys numilia (C. Felder & R. Felder, 1867) stat. nov., P. vestigiata (Butler, 1867), P. fridae Zacca, Casagrande, Huertas & Nakahara sp. nov., P. jorupe Zacca, Casagrande, Checa & Willmott sp. nov., P. terrestris terrestris (Butler, 1867) comb. nov., P. terrestris muyrakytan Zacca, Casagrande & Mielke ssp. nov., P. terrestris araguaianus Zacca, Casagrande & Mielke ssp. nov., P. terrestris grevei Zacca, Casagrande & Mielke ssp. nov., P. touloulou (Benmesbah, 2015) comb. nov., P. brixius brixius (Godart, [1824]) comb. nov., P. brixius madre Zacca, Mielke, Casagrande & Lamas ssp. nov., P. brixius rouranensis Brévignon ssp. nov. and P. flavofascia (Zacca & Siewert, 2014) comb. nov. Euptychia zeredatha Butler, 1869 and P. bahneri Anken, 1994 are new synonyms of P. p. poltys, and Euptychia binalinea Butler, 1867 and E. poltys bahiana Bryk, 1953 are new synonyms of P. poltys numilia. Lectotypes are designated for Euptychia zeredatha, Neonympha numilia, Euptychia vestigiata, Euptychia terrestris and Euptychia brixiola, Butler, 1867 and a neotype is designated for Satyrus brixius. In addition, female morphology is described for the first time for all available species, and biological data and distributional maps are provided for all species. Last instar and pupae of Paryphthimoides p. poltys are also described.

A new genus, Nhambikuara Freitas, Barbosa & Zacca gen. nov., and species, Nhambikuara cerradensis Freitas, Barbosa & Zacca sp. nov., of the highly diverse Neotropical butterfly subtribe Euptychiina are described. Nhambikuara cerradensis sp. nov. is the type species for the genus, and Euptychia mima Butler, 1867 is also transferred to the new genus, as Nhambikuara mima (Butler, 1867) comb. nov., from the genus Zischkaia Forster, 1964. The taxonomy, phylogenetic relationships, geographic distribution and natural history of species of the genus are also discussed.http://zoobank.org/urn:lsid:zoobank.org:pub:41AD7568-3490-4F63-A019-32D5592A8C44 Keywords: Cerrado, Paryphthimoides, Savanna, Splendeuptychia, Zischkaia

The genus Praepedaliodes Forster, 1964, the only representative of the mega-diverse mostly Andean Pedaliodes complex lineage in the Brazilian Atlantic Forest, is revised. Prior to this study, four species were known, P. phanias (Hewitson, 1862), P. granulata (Butler, 1868), P. amussis (Thieme, 1905) and P. exul (Thieme, 1905). Here, a further six are described, all from SE Brazil, expanding to 10 the number of species in this genus. Lectotypes are designated for P. phanias, P. granulata and P. amussis. The genus is most diverse in the Serra da Mantiqueira (São Paulo, Rio de Janeiro, Minas Gerais) and in the Serra Geral (Paraná, Santa Catarina) with seven species occurring in both ranges. Praepedaliodes phanias is the most widespread species and the only one found in the western part of the Atlantic Forest; only this species and P. duartei Dias, Dolibaina & Pyrcz n. sp. occurring to near sea level. Other species, P. zaccae Dolibaina, Dias & Pyrcz n. sp., P. francinii Freitas & Pyrcz n. sp., P. sequeirae Pyrcz, Dias & Dolbaina n. sp., P. landryi Pyrcz & Freitas n. sp. and P. pawlaki Pyrcz & Boyer n. sp. are strictly montane and the highest species richness is reached at 1400–1800 m. One species, P. sequeirae n. sp., is a narrow endemic found only at timberline in the Agulhas Negras massif above 2300 m. Immature stages are described for two species, P. phanias and P. landryi n. sp. Molecular data (barcode region of cytochrome oxidase, subunit I) and adult morphology, including male and female genitalia, support the genus as monophyletic, belonging to a predominantly Andean clade of the Pedaliodes Butler, 1867 complex. Morphological evidences, in particular female genitalia comparative analysis, indicate the genera Physcopedaliodes Forster, 1964 and Panyapedaliodes Forster, 1964 as possibly the closest relatives to Praepedaliodes. Molecular data are inconclusive in this respect.

The present paper describes the immature stages of the common Neotropical satyrine butterfly Pareuptychia ocirrhoe interjecta (R.F. d'Almeida, 1952). The solitary eggs are white and round, turning black 4 to 6 hours after oviposition. The four solitary larval instars are predominantly green and feed on grasses (Poaceae), including Setaria in nature and several other native and introduced species in captivity. The pupa is short and smooth and entirely green. Except for the black eggs, the immature stages are similar to those of other forest species of Euptychiina. The most remarkable and unique characteristic of P. ocirrhoe interjecta is the shiny black eggs, a possible synapomorphy for Pareuptychia not known in any other Euptychiina.

The male genitalic characters of Hexapoda are well known for their great taxonomic and systematic value. Despite insect male genitalia displaying large diversity, variation, and modification across orders, some structures are consistently present, and such characters can serve as the basis for discussion regarding homology. In the order Lepidoptera, a male genitalic structure widely known as the ‘juxta’ is present in many taxa and absence or modification of this character can be phylogenetically informative at the generic or higher level. We here focus on the systematics of the so-called ‘Taygetis clade’ within the nymphalid subtribe Euptychiina, and report an unusual case of ‘juxta loss’ in a single species, Taygetina accacioi Nakahara & Freitas, n. sp., a new species from Brazil named and described herein. Additionally, we describe another west Amazonian Taygetina Forster, 1964 species, namely Taygetina brocki Lamas & Nakahara, n. sp., in order to better document the species diversity of Taygetina. Our most up-to-date comprehensive molecular phylogeny regarding ‘Taygetis clade’ recovered these two species as members of a monophyletic Taygetina, reinforcing the absence of juxta being a character state change occurring in a single lineage, resulting in an apomorphic condition, which we report here as a rare case in butterflies (Papilionoidea).

The present paper describes the immature stages of the Neotropical satyrine butterflyEuptychiamollina(Hübner, [1813]) from the Brazilian Amazon Forest. Eggs were laid singly on the under surface of the fronds of its host plant,Selaginellasp. (Bryophyta: Selaginellaceae). Larvae are solitary in all instars, presenting a color pattern and shape that make them cryptic on its host plant. The pupa is short, smooth and varies from rusty brown to green. Despite the lack of a close phylogenetic relationship, larvae ofEuptychiaare very similar to those of the paleotropical satyrinesRagadiaandAcrophtalmia, suggesting that camouflage is likely to be one of the factors explaining the similarities among them.

A revision of Godartiana Forster, 1964 (Satyrinae: Euptychiina) is provided, including the description of a new species, G. amadoi Paluch, Zacca & Freitas sp. nov. from Bahia and Pernambuco, northeastern Brazil. Two genera, Cercyeuptychia L. Miller & Emmel, 1971 syn. nov. and Praefaunula Forster, 1964 syn. nov., are herein synonymised with Godartiana, based on morphological and molecular data from mitochondrial gene Cytochrome C Oxidase I and the nuclear genes GAPDH and RpS5. A new taxonomic arrangement is proposed for Godartiana, increasing to five the number of species known in this genus and also synonymising Euptychia liturata Butler, 1867 syn. nov. and E. vesper Butler, 1867 syn. nov. with G. armilla (Butler, 1867) comb. nov. The female genitalia of all species of Godartiana are described and illustrated, information is given on host plants and biology, and the previously unknown immature stages of G. muscosa are described and illustrated.

This paper describes two new species of Neotropical butterflies: Yphthimoides blanquita Barbosa, Marin and Freitas sp. nov., from the dry forest of northwestern Colombia, and Yphthimoides nareta Barbosa & Freitas sp. nov. from northeastern Brazil, based on morphological and molecular data. Adult morphology, including wing shape and pattern as well as male genitalia, is described in detail. Furthermore, analysis of the mitochondrial CoxI ‘barcode’ showed that both new species are quite distinct from all similar Yphthimoides species and additionally, the ‘renata clade’ is defined based on the presence of cornuti in the aedeagus.http://zoobank.org/urn:lsid:zoobank.org:pub:22CA85FF-9682-4016-A300-AB11C71CEDA2

The present paper describes a new species of Actinote (Nymphalidae, Heliconiinae, Acraeini) from southeastern Brazil, and describes the morphology of the adults and immature stages of this species. Actinote mantiqueira sp. nov. occurs in the Serra da Mantiqueira and Serra do Mar in the Atlantic Forest. Adults from this population are very similar to other species of the “orangish red mimicry complex”, including Actinote alalia (C. Felder & R. Felder, 1860), its sister species, restricted to the mountains of southern Brazil. Actinote mantiqueira sp. nov. and A. alalia are distinguishable by wing pattern, male genitalia and larval morphology, and have strongly allopatric distributions. A redescription of Actinote alalia is also provided. Keywords: Atlantic Forest, Araucaria Forest, Actinote alalia, Species redescription, Immature stages

Euptychia saltuensis Hayward, 1962, new synonym, currently regarded as a nomen dubium and possibly a junior subjective synonym of Yphthimoides manasses (C. Felder & R. Felder, 1867), is here treated as a junior subjective synonym of Yphthimoides patricia (Hayward, 1957), based on morphological characters of the male genitalia and the DNA barcode. The taxonomic status of Y. patricia is re-examined, and a detailed redescription of the adult morphology, including the male genitalia, is presented. Information on the distribution, habitat and immature stages of Y. patricia is also provided. Yphthimoides patricia is clearly a distinct species from Y. manasses based on the analysis of DNA barcode sequences and the morphology of the male genitalia.

Three new species belonging to the butterflies satyrine subtribe Euptychiina are described: (1) Yphthimoides gabriela n. sp. occurring in low to medium altitudes in the coastal forests between the states of Rio de Janeiro and Bahia, (2) Yphthimoides bella n. sp., from Brazilian Cerrado savannas, known from only two localities in the State of Goias, and (3) Yphthimoides iserhardi n. sp., occurring in high altitudes in the rocky outcrops of Chapada Diamantina, in the interior of the Bahia state. Descriptions are based on wing shape, wing pattern, and morphology of male genitalia. Furthermore, molecular data from the “DNA barcode” (Cytochrome C Oxidase I, ca. 658 bp) was obtained and used to validate the three new species. Additionally, information about geographic distribution and habitat for the three new species is provided, and the systematic position of each of the three species is discussed based on a molecular analysis using another 11 additional species of Yphthimoides.

This paper describes the immature stages of Pagyris cymothoe cymothoe (Hewitson, 1855) based on samples collected in Venezuela and Colombia in 1991 and 2013, respectively. Eggs were collected on Brugmansiacandida Pers. and Brugmansiasuaveolens (Willd.), and reared on B.suaveolens under laboratory conditions at 25 ± 2°C. Larvae are gregarious, passing through five instars; the duration of life cycle (from egg to adult) lasted about 28 days. Detailed morphology of the egg and first instar chaetotaxy is presented. This new life history is compared with the life histories of Pagyris ulla (Hewitson, 1857) and of the sister genus Placidina d'Almeida, 1928.

The recently described butterfly Magneu­ptychia flavofascia was formerly known from only two localities in the Brazilian Cerrado savanna. Here, we report new distribution records, DNA barcode sequences, and information on the habitat, adult behavior and systematic position of this species., LAK was supported by CNPq (163119/2013-9) and CAPES (3200-14-0), and RDE acknowledges CNPq (563134/2010-0). EPB acknowledges FAPESP (2012/03750-8) for a graduate fellowship. AVLF was supported by CNPq (fellowship 302585/2011-7), and the National Science Foundation (DEB-1256742). 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 BIOTA-FAPESP Program (2011/50225-3). Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (Brasil) Fundação de Amparo à Pesquisa do Estado de São Paulo National Science Foundation (US)

A new satyrine species in the subtribe Euptychiina, Euptychia atlantica Nakahara & Freitas sp. nov., is described from the Atlantic coastal forest of Brazil. Based on the existing museum specimens, E. atlantica sp. nov. is known from the coastal montane forests of Rio de Janeiro to south Bahia, a unique biogeographical region which is undergoing rapid degradation. Illustrations of adults and their genitalia, as well as a distribution map, are provided herein, in addition to a discussion of the relationships and conservation status of the new species.

Sepona Freitas and Barbosa, gen. nov. is proposed for the Neotropical satyrine butterfly species Euptychia punctata Weymer, 1911 and its junior subjective synonyms Euptychia griseola Weymer, 1911 and Taygetis indecisa Ribeiro, 1931. The new genus has a distinctive wing pattern and shape of the valvae in the male genitalia, the latter being a unique autapomorphy within the subtribe Euptychiina. Based on molecular data, this genus is not sister to any other single euptychiine genus, instead appearing as the sister to all remaining genera in the Taygetis clade. The present paper illustrates the complexity of the taxonomy of Euptychiina, and the importance of using different sources of evidence in taxonomic studies. Keywords: Harjesia, Integrative taxonomy, Pseudodebis, Satyrini, Yphthimoides

A new genus is described for Neonympha griphe C. Felder & R. Felder, 1867, to contribute toward a revision of the polyphyletic genus Euptychoides Forster, 1964. Based on DNA sequence data, Graphita Nakahara, Marin & Barbosa, gen. nov. is strongly supported as a member of a clade of predominantly southeastern Brazilian taxa, in which it is weakly supported as sister to a well-supported clade containing Pharneuptychia Forster, 1964, Moneuptychia Forster, 1964 and the E. castrensis (Schaus, 1902) species complex. The data show that Graphita griphe comb. nov. is not related to other Euptychoides and not very closely related to any other sampled euptychiines, and thus support the description of this new genus. In addition, we provide morphological illustrations and a distribution map for this taxon based on museum specimens.

The immature stages of the Neotropical mistletoe butterfly Cunizza hirlanda planasia Fruhstorfer (Pieridae: Anthocharidini) are described and illustrated for the first time from cerrado areas of Central Brazil Plateau. Eggs are laid in small loose clusters (2–7 eggs) on new red leaves of the mistletoe Passovia ovata (Pohl ex DC.) Kuijt (Loranthaceae). Larvae are semi-gregarious and underwent five instars. Host plant use, morphology, and behavior of immature stages are similar to those of related genera in the Hesperocharis group of Anthocharidini.

The pierid Charonias theano (Boisduval), an endangered butterfly species, has been rarely observed in nature, and has not been recorded in the state of São Paulo in the last 50 years despite numerous efforts to locate extant colonies. Based on museum specimens and personal information, C. theano was known from 26 sites in southeastern and southern Brazil. Recently, an apparently viable population was recorded in a new locality, at Serra do Japi, Jundiaí, São Paulo, with several individuals observed during two weeks in April, 2011. The existence of this population at Serra do Japi is an important finding, since this site represents one of the few large forested protected areas where the species could potentially persist not only in the state of São Paulo, but within its entire historical distribution.

This paper describes four new species of Moneuptychia as follows: M. montana Freitas, M. vitellina FreitasBarbosa, M. pervagata Freitas, SiewertMielke and M. wahlbergi Freitas, Barbosa, SiewertMielke from south and southeastern Brazil. Details are presented on the morphology of adults of all species, and immature stages for two species, and we discuss the taxonomy and identification of the genus Moneuptychia. The mitochondrial CoxI "barcode" region was used for exploring the utility of this DNA marker to identify these species, giving strong support for all new species.

[EN] Documenting how diversity patterns vary at fine- and broad scales may help answer many questions in theoretical and applied ecology. However, studies tend to compare diversity patterns at the same scale and within the same taxonomic group, which limits the applicability and generality of the results. Here, we have investigated whether vegetation-dwelling arthropods from different trophic ranks and with distinct life histories (i.e., ants, caterpillars, cockroaches, and spiders) have different beta-diversity patterns at multiple scales. Specifically, we compared their beta diversity across architecturally distinct plant species (fine-scale process) and a latitudinal gradient of sites (broad-scale process) along 2040 km of coastal restinga vegetation in the Neotropics. Over 50 percent of the compositional changes (β-diversity) in ants, caterpillars, and spiders and 41 percent of those in cockroaches were explained by plant identity within each site. Even groups that do not feed on plant tissues, such as omnivores and predators, were strongly affected by plant identity. Fine-scale variation was more important than large-scale processes for all studied groups. Performing a cross-scale comparison of diversity patterns of groups with distinct life histories helps elucidate how processes that act at regional scales, such as dispersal, interact with local processes to assemble arthropod communities., [PT] Entender como os padrões de diversidade variam em escalas finas e amplas pode responder muitas questões em ecologia aplicada e teórica. Porém, muitos estudos comparam padrões de diversidade na mesma escala e dentro do mesmo grupo taxonômico, o que limita a aplicabilidade e generalização dos resultados. Nós investigamos se múltiplos artrópodes que vivem na vegetação e que pertencem a diferentes ordens tróficas e com história de vida distinta (i.e., aranhas, baratas, formigas, e lagartas) apresentam padrões de diversidade beta diferentes em múltiplas escalas. Especificamente, comparamos a diversidade beta entre espécies de plantas com arquiteturas distintas (um processo de escala fina) e localidades diferentes ao longo do gradiente latitudinal (um processo de escala ampla) ao longo de 2.040 km em vegetação costeira de restinga na região Neotropical. Observamos que mais de 50% das mudanças na composição de espécies (diversidade beta) de formigas, lagartas e aranhas, e 41% de baratas foram explicados por diferenças na identidade das plantas em uma mesma localidade. Demonstramos que mesmo grupos que não se alimentam de tecidos vegetais, tais como onívoros e predadores, são fortemente afetados pela identidade da planta. A variação em escala fina foi mais importante que processos em escala ampla para todos os grupos estudados. Sugerimos que, ao desenvolver comparações dos padrões de diversidade entre diferentes escalas com grupos de ordens tróficas distintas, podemos compreender como os processos que atuam em escalas regionais, tais como dispersão, interagem com processos locais para organizar comunidades de artrópodes., This manuscript is part of TG.-S.' PhD thesis developed at the Graduate Program in Animal Biology (Universidade Estadual Paulista UNESP, São José do Rio Preto, Brazil). This study was supported by a doctoral fellowship from FAPESP to TG-S and by FAPESP and CNPq research grants to G.Q.R. L.A.K. was supported by CAPES (3200-14-0), and E.P.B. thanks FAPESP (12/03750-8) for a fellowship. AJS was financially supported by CNPq (proc. 308072/2012-0 and 475179/2012-9), FAPEMIG (PPM-00335-13) and Instituto Nacional de Ciência e Tecnologia dos Hymenoptera Parasitóides da Região Sudeste Brasileira (http:/www.hympar.ufscar.br/).

Brassolines are among the largest Neotropical butterflies, with large eyespots resembling vertebrate eyes in their wings. Some subspecies of co-occurring brassolines show striking similarities in wing traits to the extent that mimicry within the tribe has been suggested. When adults exhibit phenotypic convergence, data on immature stages can be used to uncover similarities derived from common descent. Here, evidence for future phylogenies to support Selenophanes relatedness to either Opsiphanes or Catoblepia are provided by comparing novel molecular (barcode sequence) and morphological (immature) data. While barcoding showed Selenophanes sequences to be nested within Catoblepia, with intergeneric distances smaller than between Catoblepia species, the state of the art of immature morphological descriptions showed that Catoblepia and Opsiphanes were more similar than either were to Selenophanes. Despite their popularity, brassolines are difficult to find in nature, partly because they are active during dawn and dusk. S. cassiope is a rare species, so the hitherto unknown female of S. cassiope guaranyCasagrande, 1992 is described, as well as morphologically and ecologically relevant adult traits. Analysis of sensory organs revealed a possible reversal of brassoline adaptations to the crepuscular habit. The pigment class of wing cells from several regions with different colors was melanin, except for white (colorless) scales. Multitrophic interactions with an Arecaceae host plant, and egg (Anastatus sp., Galeopsomyia sp.) and pupal (Eurytoma sp.) parasitoids are also reported. This integrative approach benefits from a group with great potential to explore the interplay, from molecules to ecology, of adaptive evolution.

The Neotropical genus Urbanus (Hubner) (Lepidoptera: Hesperiidae) contains around 34 described species, and is widely distributed from the extreme southern United States to Argentina. Here, we describe the larval morphology and behavior of Urbanus esmeraldus (Hubner)(Hesperiidae) in Urera baccifera (Urticaceae), a plant producing food rewards and fleshy fruits that attract ants (including predacious species) in a Brazilian forest. Larvae pass through five instars and construct two kinds of leaf shelters. Experiments with ejected fecal pellets showed that these can serve as cues to ground—dwelling ants that climb onto host plants and potentially attack the larvae. Manipulation with pellets placed at different distances suggests that ejection behavior decreases larval vulnerability to ant predation. Larval preference for mature leaves may be related with increased predation risk at ant—visited young leaves. The study shows that a combination of natural history and experimental data can help understand the life history of a butterfly using a plant with high predation risk.

We provide the first information on the morphology of the immature stages (egg, larva, and pupa), oviposition and larval behavior, and host plant, for the Brazilian crescent butterfly Ortilia liriope (Cramer), based on material from Santarem Municipality, Para State, Northern Brazil. Females of O. liriope lay eggs in clusters. After hatching, larvae eat the exochorion and remain gregarious in all but the final instar. The host plant recorded in the study site is Justicia sp. (Acanthaceae). Despite the scarcity of data on the immature stages of Neotropical Melitaeini, we can already say that some morphological and behavioral traits observed in the immature stages of O. liriope are also present in all known genera in this tribe.

This paper describes four new species of Moneuptychia as follows: M. montana Freitas, M. vitellina Freitas & Barbosa, M. pervagata Freitas, Siewert & Mielke and M. wahlbergi Freitas, Barbosa, Siewert & Mielke from south and southeastern Brazil. Details are presented on the morphology of adults of all species, and immature stages for two species, and we discuss the taxonomy and identification of the genus Moneuptychia. The mitochondrial CoxI "barcode" region was used for exploring the utility of this DNA marker to identify these species, giving strong support for all new species.

This paper describes a new, abundant and widespread species of Yphthimoides Forster from the Atlantic forests of southern Brazil, Paraguay, and northern Argentina (Misiones) in open and secondary vegetation and forest edges. Adult and immature stage morphology is described, molecular data are provided, and the placement of the new species within the genus Yphthimoides is discussed.

A new genus and species of Euptychiina (Satyrinae), Cristalinaia vitoria Mota, Zacca & Freitas gen. et sp. nov., is described based on three specimens collected in the region of the Cristalino River, Alta Floresta, Mato Grosso, Brazil. This rare species is known only from this region, where it flies inside the dense bamboo patches typical of that area. The last instar larva and the pupa are described; the larva was observed feeding on mature leaves of the common bamboo Guadua aff. paniculata Munro.urn:lsid:zoobank.org:pub:D61EDE8D-CAE9-41C6-B24D-BB789873566E Keywords: Amazonia, Immature stages, Morphology, Molecular phylogeny, Taxonomy