No abstract

Numerous hypotheses on the evolution of Neotropical biodiversity have stimulated research to provide a better understanding of diversity dynamics and distribution patterns of the region. However, few studies integrate molecular and morphological data with complete sampling of a Neotropical group, and so there has been little synthesis of the multiple processes governing biodiversity through space and time. Here, a total-evidence phylogenetic approach is used to reconstruct the evolutionary history of the butterfly subgenus Heraclides. We used DNA sequences for two mitochondrial genes and one nuclear gene and coded 133 morphological characters of larvae and adults. A robust and well-resolved phylogeny was obtained using several analytical approaches, while molecular dating and biogeographical analyses indicated an early Miocene origin (22 Mya) in the Caribbean Islands. We inferred six independent dispersal events from the Caribbean to the mainland, and three from the mainland to the Caribbean, and we suggest that cooling climates with decreasing sea levels may have contributed to these events. The time-calibrated tree is best explained by a museum model of diversity in which both speciation and extinction rates remained constant through time. By assessing both continental and fine-scale biodiversity patterns, this study provides new findings, for instance that islands may act as source of diversity rather than as a sink, to explain spatio-temporal macroevolutionary processes within the Neotropical region.

We here propose a new, monotypic genus, Amiga Nakahara, Willmott & Espeland, gen. n., to harbor a common Neotropical butterfly, described as Papilioarnaca Fabricius, 1776, and hitherto placed in the genus Chloreuptychia Forster, 1964. Recent and ongoing molecular phylogenetic research has shown Chloreuptychia to be polyphyletic, with C.arnaca proving to be unrelated to remaining species and not readily placed in any other described genus. Amigaarnacagen. n. et comb. n. as treated here is a widely distributed and very common species ranging from southern Mexico to southern Brazil. A neotype is designated for the names Papilioarnaca and its junior synonym, Papilioebusa Cramer, 1780, resulting in the treatment of the latter name as a junior objective synonym of the former. A lectotype is designated for Euptychiasericeella Bates, 1865, which is treated as a subspecies, Amigaarnacasericeella (Bates, 1865), comb. n. et stat. n., based on molecular and morphological evidence. We also describe two new taxa, Amigaarnacaadela Nakahara & Espeland, ssp. n. and Amigaarnacaindianacristoi Nakahara & Marín, ssp. n., new subspecies from the western Andes and eastern Central America, and northern Venezuela, respectively.

Based on external morphology, food plant records for caterpillars, and molecular analysis, Euptychia hilara (C. Felder & R. Felder, 1867) is removed from Euptychia Hübner 1818. A new genus, Inbio Nakahara & Espeland gen. nov., is proposed for this taxon. Inbio hilara comb. nov. is a member of a monophyletic clade containing Cyllopsis Felder, 1869, Paramacera Butler, 1868, and Atlanteuptychia Freitas, Barbosa & Mielke, 2013, although it can be morphologically distinguished from these genera. Lectotypes for Neonympha hilara C. Felder & R. Felder, 1867 and Euptychia anacleta Butler, 1877 (a synonym of E. hilara) are designated herein.

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, Marín & 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 ‘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.

We describe a new nymphalid butterfly, Euptychia favonius sp. nov., from the western Andes. Adults and genitalia for this new taxon are illustrated and a distribution map is provided. The taxonomy of its sister species, E. meta Weymer, 1911, is discussed, and a lectotype is designated for E. meta.

We provide the first comprehensive taxonomic revision of the poorly known South American butterfly genus Zischkaia Forster, 1964, hitherto regarded as including three described species. A phylogenetic analysis based on DNA sequence data shows that Zischkaia is monophyletic and consists of two morphologically diagnosable clades. Morphological characters and DNA ‘barcodes’ support the recognition of twelve species in the genus, a significant increase even for the relatively poorly studied subtribe Euptychiina. Consequently, nine new species are described and named herein, including Z. arctoa Nakahara, sp. nov., Z. chullachaki Nakahara & Zacca, sp. nov., Z. baku Zacca, Dolibaina & Dias, sp. nov., Z. arenisca Nakahara, Willmott & Hall, sp. nov., Z. argyrosflecha Nakahara, L. Miller & Huertas, sp. nov., Z. abanico Nakahara & Petit, sp. nov., Z. josti Nakahara & Kleckner, sp. nov., Z. mielkeorum Dolibaina, Dias & Zacca, sp. nov. and Z. warreni Dias, Zacca & Dolibaina, sp. nov. In addition, a neotype is designated for Satyrus pacarus Godart, [1824], and lectotypes are designated for Euptychia amalda Weymer, 1911, Euptychia fumata Butler, 1867 and Euptychia saundersii Butler, 1867.