Montane diversification as a mechanism of speciation in neotropical butterflies.

The mountains in the Atlantic Forest domain are environments that harbor a high biodiversity, including species adapted to colder climates that were probably influenced by the climatic variations of the Pleistocene. To understand the phylogeographic pattern and assess the taxonomic boundaries between two sister montane species, a genomic study of the butterflies Actinote mantiqueira and A. alalia (Nymphalidae: Acraeini) was conducted. Analyses based on partial sequences of the mitochondrial gene COI (barcode region) failed to recover any phylogenetic or genetic structure discriminating the two species or sampling localities. However, single nucleotide polymorphisms gathered using Genotyping-by-Sequencing provided a strong isolation pattern in all analyses (genetic distance, phylogenetic hypothesis, clustering analyses, and F ST statistics) which is consistent with morphology, separating all individuals of A. alalia from all populations of A. mantiqueira. The three sampled mountain ranges where A. mantiqueira populations occur-Serra do Mar, Serra da Mantiqueira, and Poços de Caldas Plateau-were identified as three isolated clusters. Paleoclimate simulations indicate that both species' distributions changed according to climatic oscillations in the Pleistocene period, with the two species potentially occurring in areas of lower altitude during glacial periods when compared to the interglacial periods (as the present). Besides, a potential path between their distribution through the Serra do Mar Mountain range was inferred. Therefore, the Pleistocene climatic fluctuation had a significant impact on the speciation process between A. alalia and A. mantiqueira, which was brought on by isolation at different mountain summits during interglacial periods, as shown by the modeled historical distribution and the observed genetic structure.

Hybrid speciation driven by multilocus introgression of ecological traits.

Hybridization allows adaptations to be shared among lineages and may trigger the evolution of new species1,2. However, convincing examples of homoploid hybrid speciation remain rare because it is challenging to demonstrate that hybridization was crucial in generating reproductive isolation3. Here we combine population genomic analysis with quantitative trait locus mapping of species-specific traits to examine a case of hybrid speciation in Heliconius butterflies. We show that Heliconius elevatus is a hybrid species that is sympatric with both parents and has persisted as an independently evolving lineage for at least 180,000 years. This is despite pervasive and ongoing gene flow with one parent, Heliconius pardalinus, which homogenizes 99% of their genomes. The remaining 1% introgressed from the other parent, Heliconius melpomene, and is scattered widely across the H. elevatus genome in islands of divergence from H. pardalinus. These islands contain multiple traits that are under disruptive selection, including colour pattern, wing shape, host plant preference, sex pheromones and mate choice. Collectively, these traits place H. elevatus on its own adaptive peak and permit coexistence with both parents. Our results show that speciation was driven by introgression of ecological traits, and that speciation with gene flow is possible with a multilocus genetic architecture.

Immature Stages, Natural History and Population Biology of Opoptera syme (Hübner, 1821), (Lepidoptera: Nymphalidae: Satyrinae) from Southeastern Brazil.

The present study describes the immature stages of Opoptera syme (Hübner, [1821]) using SEM and Micro-CT to generate 2D and 3D models to study the morphology and chaetotaxy, and present information on the natural history, behavior, and population biology. In laboratory, eggs were laid singly, and the isolated larvae passed through six to seven instars. Host plant in the study site is unknown, but larvae are known to feed on bamboos; in laboratory, larvae accepted the ornamental bamboo Bambusa textilis McClure, 1940. Adults are diurnal and univoltine, flying from December to April (the austral summer). The present study adds information to the biology and natural history of Brassolini, an iconic and still poorly known tribe of Neotropical butterflies.

Systematic Position, Immature Stages, and Geographic Distribution of Glennia pylotis (Godart, 1819) (Lepidoptera: Papilionoidea: Pieridae).

The biology, immature stages, geographic distribution, and systematic position of Glennia pylotis (Godart, 1819) are presented. This species is mostly restricted to the southeastern coastal Atlantic Forest, from the States of São Paulo to Bahia, with sparse records in the interior of Brazil, Argentina, and Bolivia (this locality is most probably spurious-see below). Data on immature stages are based on textual descriptions; pupal skins were illustrated and compared with those of other members of the subtribe Pierina. Based on molecular data, G. pylotis was recovered as a member of the "Leptophobia clade," as the sister group of all other genera in this clade except for Leptophobia. The immature stages are similar to, and host plants are the same, of several other related genera within Pierina, especially the "Leptophobia clade." By compiling all available data, searching for unpublished information in museums (including the finding of empty pupal cases) and adding molecular evidence for G. pylotis, not only its systematic position was elucidated, but its true conservation status could also be assessed.

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

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

Atlantic flower-invertebrate interactions: A data set of occurrence and frequency of floral visits.

Encounters between flowers and invertebrates are key events for the functioning of tropical forests. Assessing the structure of networks composed of the interactions between those partners leads to a better understanding of ecosystem functioning and the effects of environmental factors on ecological processes. Gathering such data is, however, costly and time-consuming, especially in the highly diverse tropics. We aimed to provide a comprehensive repository of available flower-invertebrate interaction information for the Atlantic Forest, a South American tropical forest domain. Data were obtained from published works and "gray literature," such as theses and dissertations, as well as self-reports by co-authors. The data set has ~18,000 interaction records forming 482 networks, each containing between one and 1061 interaction links. Each network was sampled for about 200 h or less, with few exceptions. A total of 641 plant genera within 136 different families and 39 orders were reported, with the most abundant and rich families being Asteraceae, Fabaceae, and Rubiaceae. Invertebrates interacting with these plants were all arthropods from 10 orders, 129 families, and 581 genera, comprising 2419 morphotypes (including 988 named species). Hymenoptera was the most abundant and diverse order, with at least six times more records than the second-ranked order (Lepidoptera). The complete data set shows Hymenoptera interacting with all plant orders and also shows Diptera, Lepidoptera, Coleoptera, and Hemiptera to be important nodes. Among plants, Asterales and Fabales had the highest number of interactions. The best sampled environment was forest (~8000 records), followed by pastures and crops. Savanna, grasslands, and urban environments (among others) were also reported, indicating a wide range of approaches dedicated to collecting flower-invertebrate interaction data in the Atlantic Forest domain. Nevertheless, most reported data were from forest understory or lower strata, indicating a knowledge gap about flower-invertebrate interactions at the canopy. Also, access to remote regions remains a limitation, generating sampling bias across the geographical range of the Atlantic Forest. Future studies in these continuous and hard-to-access forested areas will yield important new information regarding the interactions between flowers and invertebrates in the Atlantic Forest. There are no copyright restrictions on the data set. Please cite this data paper if the data are used in publications and teaching events.

ATLANTIC ANTS: a data set of ants in Atlantic Forests of South America.

Ants, an ecologically successful and numerically dominant group of animals, play key ecological roles as soil engineers, predators, nutrient recyclers, and regulators of plant growth and reproduction in most terrestrial ecosystems. Further, ants are widely used as bioindicators of the ecological impact of land use. We gathered information of ant species in the Atlantic Forest of South America. The ATLANTIC ANTS data set, which is part of the ATLANTIC SERIES data papers, is a compilation of ant records from collections (18,713 records), unpublished data (29,651 records), and published sources (106,910 records; 1,059 references), including papers, theses, dissertations, and book chapters published from 1886 to 2020. In total, the data set contains 153,818 ant records from 7,636 study locations in the Atlantic Forest, representing 10 subfamilies, 99 genera, 1,114 ant species identified with updated taxonomic certainty, and 2,235 morphospecies codes. Our data set reflects the heterogeneity in ant records, which include ants sampled at the beginning of the taxonomic history of myrmecology (the 19th and 20th centuries) and more recent ant surveys designed to address specific questions in ecology and biology. The data set can be used by researchers to develop strategies to deal with different macroecological and region-wide questions, focusing on assemblages, species occurrences, and distribution patterns. Furthermore, the data can be used to assess the consequences of changes in land use in the Atlantic Forest on different ecological processes. No copyright restrictions apply to the use of this data set, but we request that authors cite this data paper when using these data in publications or teaching events.

A New Species of Dynamine Hübner, [1919] (Lepidoptera: Nymphalidae: Biblidinae) from the Atlantic Forest in Southeastern Brazil.

A new butterfly species, Dynamine sideria Rosa & Freitas sp. nov. (Nymphalidae: Biblidinae), is described from the Atlantic Forest in Southeastern Brazil. The new species is known from only seven adult specimens and three observation records (two from photographs) in five sites, including a large forest reserve, forest fragments on a disturbed landscape, and a backyard in a dense urban site. Illustrations of adults and male genitalia are provided, as well as a distribution map and limited natural history notes.

Knowledge gaps hamper understanding the relationship between fragmentation and biodiversity loss: the case of Atlantic Forest fruit-feeding butterflies.

BACKGROUND: A key challenge for conservation biology in the Neotropics is to understand how deforestation affects biodiversity at various levels of landscape fragmentation. Addressing this challenge requires expanding the coverage of known biodiversity data, which remain to date restricted to a few well-surveyed regions. Here, we assess the sampling coverage and biases in biodiversity data on fruit-feeding butterflies at the Brazilian Atlantic Forest, discussing their effect on our understanding of the relationship between forest fragmentation and biodiversity at a large-scale. We hypothesize that sampling effort is biased towards large and connected fragments, which occur jointly in space at the Atlantic forest. METHODS: We used a comprehensive dataset of Atlantic Forest fruit-feeding butterfly communities to test for sampling biases towards specific geographical areas, climate conditions and landscape configurations. RESULTS: We found a pattern of geographical aggregation of sampling sites, independently of scale, and a strong sampling bias towards large and connected forest fragments, located near cities and roads. Sampling gaps are particularly acute in small and disconnected forest fragments and rare climate conditions. In contrast, currently available data can provide a fair picture of fruit-feeding butterfly communities in large and connected Atlantic Forest remnants. DISCUSSION: Biased data hamper the inference of the functional relationship between deforestation and biodiversity at a large-scale, since they are geographically clustered and have sampling gaps in small and disconnected fragments. These data are useful to inform decision-makers regarding conservation efforts to curb biodiversity loss in the Atlantic Forest. Thus, we suggest to expand sampling effort to small and disconnected forest fragments, which would allow more accurate evaluations of the effects of landscape modification.

Seasonal Patterns of Host Plant Use in an Assemblage of Heliconiini Butterflies (Lepidoptera: Nymphalidae) in a Neotropical forest.

Insect-plant interactions involve physiological adaptations by insects to secondary metabolic compounds synthetized by host plants, which are considered essential for the determination of resources partitioning of these insects. Data on such phenomena are important to understand evolutionary and ecological processes. However, climatic factors also seem to play a key role in affecting these patterns. The present study aimed to investigate the influence of seasonal variation on patterns of host plant use (Passifloraceae) by Heliconiini butterflies (Nymphalidae: Heliconiinae) at a Neotropical site in Southeastern Brazil. A total of 12 species of Heliconiini were reported, with nine of them being resident and using five species of Passiflora (Passifloraceae) as larval host plants. Three host plant species accounted for 97% of the total use, and the use varied along the seasons highlighting the plasticity boundaries in Heliconiini and possible limiting factors.

Use of species delimitation approaches to tackle the cryptic diversity of an assemblage of high Andean butterflies (Lepidoptera: Papilionoidea).

Cryptic biological diversity has generated ambiguity in taxonomic and evolutionary studies. Single-locus methods and other approaches for species delimitation are useful for addressing this challenge, enabling the practical processing of large numbers of samples for identification and inventory purposes. This study analyzed an assemblage of high Andean butterflies using DNA barcoding and compared the identifications based on the current morphological taxonomy with three methods of species delimitation (automatic barcode gap discovery, generalized mixed Yule coalescent model, and Poisson tree processes). Sixteen potential cryptic species were recognized using these three methods, representing a net richness increase of 11.3% in the assemblage. A well-studied taxon of the genus Vanessa, which has a wide geographical distribution, appeared with the potential cryptic species that had a higher genetic differentiation at the local level than at the continental level. The analyses were useful for identifying the potential cryptic species in Pedaliodes and Forsterinaria complexes, which also show differentiation along altitudinal and latitudinal gradients. This genetic assessment of an entire assemblage of high Andean butterflies (Papilionoidea) provides baseline information for future research in a region characterized by high rates of endemism and population isolation.

The Temporal Dynamics of Two Morpho Fabricius, 1807 Species (Lepidoptera: Nymphalidae) are Affected Differently by Fire in the Brazilian Savanna.

Fire is one of the main disturbance agents globally and one of the main threats on the Brazilian cerrado (Neotropical savanna), acting as environmental filter for species selection. Individuals of Morpho helenor achillides (C. Felder and R. Felder, 1867) and M. menelaus coeruleus (Perry, 1810) were captured weekly using entomological nets and fruit-baited traps over a 36-mo period (from May 2005 to April 2008). The present study analyzed the impact of a fire event in both the above species, showing that they present different responses to this disturbance. Morpho helenor was persistent as adults during the dry season, which could fly away from the flames to neighbor unburned areas and return after dry-season bushfires. Conversely, Morpho menelaus persists only as caterpillars during the dry season, which are unable to escape from the flames resulting in high impacts on local population (the species was not captured up to the end of this study). In addition, based on host plant species reported in the literature, we assigned a broader host plant range to M. helenor, suggesting that a generalist diet could help in the maintenance of individuals during the dry season, as they have more options to breed all year round. A better understanding of the temporal dynamics of adult and immature stages could help predict the amplitude of the impacts of dry season fire events on insects, especially when preventive fires are strategically used inside protected areas.

Priors and Posteriors in Bayesian Timing of Divergence Analyses: The Age of Butterflies Revisited.

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.

Atlantic butterflies: a data set of fruit-feeding butterfly communities from the Atlantic forests.

Butterflies are one of the best-known insect groups, and they have been the subject of numerous studies in ecology and evolution, especially in the tropics. Much attention has been given to the fruit-feeding butterfly guild in biodiversity conservation studies, due to the relative ease with which taxa may be identified and specimens sampled using bait traps. However, there remain many uncertainties about the macroecological and biogeographical patterns of butterflies in tropical ecosystems. In the present study, we gathered information about fruit-feeding butterfly species in local communities from the Atlantic Forests of South America. The ATLANTIC BUTTERFLIES data set, which is part of ATLANTIC SERIES data papers, results from a compilation of 145 unpublished inventories and 64 other references, including articles, theses, and book chapters published from 1949 to 2018. In total, the data set contains 7,062 records (presence) of 279 species of fruit-feeding butterflies identified with taxonomic certainty, from 122 study locations. The Satyrini is the tribe with highest number of species (45%) and records (30%), followed by Brassolini, with 13% of species and 12.5% of records. The 10 most common species correspond to 14.2% of all records. This data set represents a major effort to compile inventories of fruit-feeding butterfly communities, filling a knowledge gap about the diversity and distribution of these butterflies in the Atlantic Forest. We hope that the present data set can provide guidelines for future studies and planning of new inventories of fruit-feeding butterflies in this biome. The information presented here also has potential use in studies across a great variety of spatial scales, from local and landscape levels to macroecological research and biogeographical research. We expect that such studies be very important for the better implementation of conservation initiatives, and for understanding the multiple ecological processes that involve fruit-feeding butterflies as biological indicators. No copyright restrictions apply to the use of this data set. Please cite this Data paper when using the current data in publications or teaching events.

A new species of Actinote Hübner (Nymphalidae: Heliconiinae: Acraeini) from southeast Brazil

Abstract 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.

'Species' from two different butterfly genera combined into one: description of a new genus of Euptychiina (Nymphalidae: Satyrinae) with unusually variable wing pattern

ABSTRACT Sepona Freitas and Barbosa, gen. nov. is proposed for the Neotropical satyrine butterfly species Euptychia punctataWeymer, 1911 and its junior subjective synonyms Euptychia griseolaWeymer, 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.

The "Taygetis ypthima species group" (Lepidoptera, Nymphalidae, Satyrinae): taxonomy, variation and description of a new species.

A new species of Taygetis Hübner, [1819] (Lepidoptera, Nymphalidae, Satyrinae) from southeastern Brazil is described: Taygetis drogoni sp. n. In addition, T. servius Weymer, 1910 and T. fulginia d'Almeida, 1922 are resurrected from synonymy and a taxonomic discussion on the species T. ypthima Hübner, [1821] and T. rectifascia Weymer, 1907 is provided. A dichotomous key for the species is also provided.