The butterfly family Nymphalidae contains some of the most important non-drosophilid insect model systems for evolutionary and ecological studies, yet the evolutionary history of the group has remained shrouded in mystery. We have inferred a robust phylogenetic hypothesis based on sequences of 10 genes and 235 morphological characters for exemplars of 400 of the 540 valid nymphalid genera representing all major lineages of the family. By dating the branching events, we infer that Nymphalidae originated in the Cretaceous at 90 Ma, but that the ancestors of 10 -12 lineages survived the end-Cretaceous catastrophe in the Neotropical and Oriental regions. Patterns of diversification suggest extinction of lineages at the Cretaceous/Tertiary boundary (65 Ma) and subsequent elevated speciation rates in the Tertiary.

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A generic-level phylogeny for the butterfly family Nymphalidae was produced by cladistic analysis of 234 characters from all life stages. The 95 species in the matrix (selected from the 213 studied) represent all important recognized lineages within this family. The analysis showed the taxa grouping into six main lineages. The basal branch is the Libytheinae, with the Danainae and Ithomiinae on the next branch. The remaining lineages are grouped into two main branches: the Heliconiinae-Nymphalinae, primarily flower-visitors (but including the fruit-attracted Coeini); and the Limenitidinae (sensu strictu), Biblidinae, and the satyroid lineage (Apaturinae, Charaxinae, Biinae, Calinaginae, Morphinae, Brassolinae, and Satyrinae), primarily fruit-attracted. Data partitions showed that the two data sets (immatures and adults) are very different, and a partitioned Bremer support analysis showed that the adult characters are the main source of conflict in the nodes of the combined analysis tree. This phylogeny includes the widest taxon coverage of any morphological study on Nymphalid butterflies to date, and supports the monophyly and relationships of most presently recognized subgroups, providing strong evidence for the presently accepted phylogenetic scheme.

Global biodiversity peaks in the tropical forests of the Andes, a striking geological feature that has likely been instrumental in generating biodiversity by providing opportunities for both vicariant and ecological speciation. However, the role of these mountains in the diversification of insects, which dominate biodiversity, has been poorly explored using phylogenetic methods. Here we study the role of the Andes in the evolution of a diverse Neotropical insect group, the clearwing butterflies. We used dated species-level phylogenies to investigate the time course of speciation and to infer ancestral elevation ranges for two diverse genera. We show that both genera likely originated at middle elevations in the Andes in the Middle Miocene, contrasting with most published results in vertebrates that point to a lowland origin. Although we detected a signature of vicariance caused by the uplift of the Andes at the Miocene-Pliocene boundary, most sister species were parapatric without any obvious vicariant barrier. Combined with an overall decelerating speciation rate, these results suggest an important role for ecological speciation and adaptive radiation, rather than simple vicariance.

The Atlantic Forest region (wide sense) includes very complex tropical environments, increasingly threatened by extensive anthropogenic conversion (>90%). Ecologically specialized, short-generation insects (butterflies) are evaluated here as indicators for monitoring community richness, landscape integrity, and sustainable resource use in the region. The > 2100 butterfly species in the Atlantic Forest region have been censused in many sites over 35 years, giving comparable daily, weekly, monthly, and long-term site lists. The 21 most thoroughly studied sites include 218-914 species, of which half can be censused in a week or less. The butterfly communities are divided into six relatively distinct faunal regions, centered in the northeast, the central coastal tablelands, the southeast coastal plain, the mountains plus interior of the southeastern states, thc central plateau, and the southern states. Species richness shows the highest values in coastal mountains from 15 to 23"s. Local butterfly communities show a high turnover, with 20 to 40 percent of the species, especially small Lycaenidae and Hesperiidae, recorded only as unstable populations or "tourists." Easily sampled species in the family Nymphalidae, and especially its bait-attracted subfamilies, are best correlated with the entire butterfly fauna and can bc used as surrogates for species diversity. In most butterfly groups, species richness is well predicted by landscape connectivity alone, or by composite indices of environmental heterogeneity, natural disturbance, and (negatively) anthropogenic disturbance. Principal components and redundancy analyses showed that the richness and proportions of different butterfly groups in the local fauna are variably explained by disturbance, seasonality, temperature, vegetation, soils, and landscape connectivity. Various groups thus can be used as rapid indicators of different types of change in the community, its environment, and the landscape. Threatened and rare species also can be used as indicators of the most unique Atlantic Forest communities (paleoenvironments), which need special attention. RESUMOA regizo da Mata Atlintica latu sensu inclui ambientes tropicais muito complexos, cada vez mais ameagados por extensa convers%o antr6pica (>90%). Insetos pequenos, especializados, e de ciclo ripido (borboletas) sPo avaliados neste trabalho como indicadores para o monitoramento da riqueza de comunidades, integridade de paisagens, e us0 sustentivel de recursos na regido. As >2100 espCcies de borboletas na regido da Mata Atlhntica t&m sido recenseadas em muitos sitios durante 0s Liltimos 35 anos, dando listas compariveis diirias, semanais, mensais e totais para cada sitio. 0 s 21 sitios mais intensivamente estudados incluem 218-914 espCcies, das quais metade pode ser amostrada em uma semana ou menos. As comunidades de borboletas sdo divididas em seis subregibes faunisticas relativamente distintas, centradas no nordeste, nos tabuleiros baianos, no litoral do sudeste, nas regi6es montanhosas no interior dos estados do s...

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.

The Brazilian cerrado savanna covers nearly 2 million km2 and has a high incidence on foliage of various liquid food sources such as extrafloral nectar and insect exudates. These liquid rewards generate intense ant activity on cerrado foliage, making ant-plant-herbivore interactions especially prevalent in this biome. We present data on the distribution and abundance of extrafloral nectaries in the woody flora of cerrado communities and in the flora of other habitats worldwide, and stress the relevance of liquid food sources (including hemipteran honeydew) for the ant fauna. Consumption by ants of plant and insect exudates significantly affects the activity of the associated herbivores of cerrado plant species, with varying impacts on the reproductive output of the plants. Experiments with an ant-plant-butterfly system unequivocally demonstrate that the behavior of both immature and adult lepidopterans is closely related to the use of a risky host plant, where intensive visitation by ants can have a severe impact on caterpillar survival. We discuss recent evidence suggesting that the occurrence of liquid rewards on leaves plays a key role in mediating the foraging ecology of foliage-dwelling ants, and that facultative ant-plant mutualisms are important in structuring the community of canopy arthropods. Ant-mediated effects on cerrado herbivore communities can be revealed by experiments performed on wide spatial scales, including many environmental factors such as soil fertility and vegetation structure. We also present some research questions that could be rewarding to investigate in this major neotropical savanna.

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