Müllerian mimicry among Neotropical Heliconiini butterflies is an excellent example of natural selection, associated with the diversification of a large continental-scale radiation. Some of the processes driving the evolution of mimicry rings are likely to generate incongruent phylogenetic signals across the assemblage, and thus pose a challenge for systematics. We use a data set of 22 mitochondrial and nuclear markers from 92% of species in the tribe, obtained by Sanger sequencing and de novo assembly of short read data, to re-examine the phylogeny of Heliconiini with both supermatrix and multispecies coalescent approaches, characterize the patterns of conflicting signal, and compare the performance of various methodological approaches to reflect the heterogeneity across the data. Despite the large extent of reticulate signal and strong conflict between markers, nearly identical topologies are consistently recovered by most of the analyses, although the supermatrix approach failed to reflect the underlying variation in the history of individual loci. However, the supermatrix represents a useful approximation where multiple rare species represented by short sequences can be incorporated easily. The first comprehensive, time-calibrated phylogeny of this group is used to test the hypotheses of a diversification rate increase driven by the dramatic environmental changes in the Neotropics over the past 23 myr, or changes caused by diversity-dependent effects on the rate of diversification. We find that the rate of diversification has increased on the branch leading to the presently most species-rich genus Heliconius, but the change occurred gradually and cannot be unequivocally attributed to a specific environmental driver. Our study provides comprehensive comparison of philosophically distinct species tree reconstruction methods and provides insights into the diversification of an important insect radiation in the most biodiverse region of the planet.

Abstract.-Müllerian mimicry among Neotropical Heliconiini butterflies is an excellent example of natural selection, associated with the diversification of a large continental-scale radiation. Some of the processes driving the evolution of mimicry rings are likely to generate incongruent phylogenetic signals across the assemblage, and thus pose a challenge for systematics. We use a data set of 22 mitochondrial and nuclear markers from 92% of species in the tribe, obtained by Sanger sequencing and de novo assembly of short read data, to re-examine the phylogeny of Heliconiini with both supermatrix and multispecies coalescent approaches, characterize the patterns of conflicting signal, and compare the performance of various methodological approaches to reflect the heterogeneity across the data. Despite the large extent of reticulate signal and strong conflict between markers, nearly identical topologies are consistently recovered by most of the analyses, although the supermatrix approach failed to reflect the underlying variation in the history of individual loci. However, the supermatrix represents a useful approximation where multiple rare species represented by short sequences can be incorporated easily. The first comprehensive, time-calibrated phylogeny of this group is used to test the hypotheses of a diversification rate increase driven by the dramatic environmental changes in the Neotropics over the past 23 myr, or changes caused by diversity-dependent effects on the rate of diversification. We find that the rate of diversification has increased on the branch leading to the presently most species-rich genus Heliconius, but the change occurred gradually and cannot be unequivocally attributed to a specific environmental driver. Our study provides comprehensive comparison of philosophically distinct species tree reconstruction methods and provides insights into the diversification of an important insect radiation in the most biodiverse region of the planet.

The ‘aegrota species group’ of the Neotropical nymphalid genus Caeruleuptychia Forster, 1964, in addition to three other superficially similar, enigmatic species in the genus, are revised. A lectotype is designated for Euptychia aegrota Butler, 1867, E. aetherialis Butler, 1877 stat. rev., E. helios Weymer, 1911 and E. pilata Butler, 1867, and C. aetherialis is resurrected from its synonymy with C. aegrota. Caeruleuptychia helios caelestissima Brévignon, 2010, syn. nov., and Magneuptychia keltoumae Brévignon & Benmesbah, 2012, syn. nov. are both regarded as junior subjective synonyms of C. helios (Weymer, 1911), as a result of the discovery and first illustration of the female of this taxon. The female of C. aegrota is also described and illustrated for the first time, and three new species, C. trembathi Willmott, Nakahara, Hall & Neild, sp. nov., C. scripta Nakahara, Zacca & Huertas, sp. nov., and C. maryzenderae Lamas & Nakahara, sp. nov. are described and named. We analyze morphological and molecular data separately, in addition to combining morphological data with molecular data, to provide the first phylogenetic hypothesis for the taxa treated in this revision.

A new species of Magneuptychia Forster, 1964 (Lepidoptera: Nymphalidae, Satyrinae), Magneuptychia andrei Zacca, Casagrande & Mielke sp. n., is described and illustrated from Venezuela, French Guiana, Trinidad and Tobago and northern Brazil. A comparative diagnosis between the new species and Magneuptychia ocypete (Fabricius, 1776), M. fugitiva Lamas, [1997] and Cissia terrestris (Butler, 1867) is also provided due the similarities in wing pattern.

Two new species of Euptychia Hübner, 1818 are described from the upper Amazon basin: Euptychia attenboroughi Neild, Nakahara, Fratello & Le Crom, sp. n. (type locality: Amazonas, Venezuela), and Euptychia sophiae Zacca, Nakahara, Dolibaina & Dias, sp. n. (type locality: Acre, Brazil). Their unusual facies prompted molecular and phylogenetic analyses of one of the species resulting in support for their classification in monophyletic Euptychia. Diagnostic characters for the two species are presented based on wing morphology, wing pattern, presence of androconial patches on the hindwing, and genitalia. Our results indicate that the projection of the tegumen above the uncus, previously considered a synapomorphy for Euptychia, is not shared by all species in the genus. The adults and their genitalia are documented, and distribution data and a map are provided.

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Se proveen datos y anotaciones sobre los representantes de los géneros Ocaria Clench, 1970 y Strymon Hübner, 1818 encontrados en el Pantepui y áreas adyacentes en el sur de Venezuela con particular enfoque en el endemismo.

Eryphanis zolvizora (Hewitson, 1877) is a rare Andean endemic butterfly, described from Bolivia, which has been historically classified either as a unique species, or as part of a group of three allopatric species from Bolivia, Ecuador and Colombia. In this paper, the group is revised using more than 200 specimens housed in 34 European, and North and South American public and private collections. For the first time, the presence of the group in Western Ecuador and Venezuela is confirmed, and important data on Peruvian populations are provided. In some populations, individual variations of genitalia are observed. Nevertheless, male genitalia allow the distinction of four geographical groups. Considering also habitus characters, eight taxa are distinguished and considered to be subspecies, of which five are new: Eryphanis zolvizora inca ssp. nov., Eryphanis zolvizora chachapoya ssp. nov., Eryphanis zolvizora casagrande ssp. nov.., Eryphanis zolvizora reyi ssp. nov., and Eryphanis zolvizora isabelae ssp. nov.  In the present state of knowledge, these taxa are allopatric, except for a possible geographic overlap in central Peru, where data are insufficient to prove sympatry. The “several subspecies vs. several species” dilemma is discussed, considering its impact for conservation action and policies.