Shinichi Nakahara

Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails

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.

Description of a new genus for Euptychia hilara (C. Felder & R. Felder, 1867) (Lepidoptera: Nymphalidae: Satyrinae)

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 species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae: Satyrini) from Mount Roraima, Guyana

A new nymphalid species in the subtribe Euptychiina, Euptychia roraima Nakahara, Fratello & Harvey n. sp., is described from Mount Roraima, Guyana. Both internal and external morphology of E. roraima are compared against several Euptychia species and the relationship between E. roraima and congeners is briefly discussed. A strong case is put forth for further and extensive exploration of the Pantepui region concerning its poorly known butterfly fauna.

A Potentially Endangered New Species of Euptychia Hubner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the Atlantic Coastal Forest of Brazil

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.

Remarkable sexual dimorphism, rarity and cryptic species: a revision of the ' aegrota species group' of the Neotropical butterfly genus Caeruleuptychia Forster, 1964 with the description of three new species (Lepidoptera, Nymphalidae, Satyrinae)

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 Brevignon, 2010, syn. nov. , and Magneuptychia keltoumae Brevignon & 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.

Graphita gen. nov., a New Genus for Neonympha griphe C. Felder & R. Felder, 1867 (Lepidoptera, Nymphalidae, Satyrinae)

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.

A new species of Cyllopsis R. Felder, 1869 from the highlands of Chiapas, Mexico (Lepidoptera: Nymphalidae: Satyrinae)

A new species of Cyllopsis R. Felder, 1869, is described and illustrated from the highlands of Chiapas, Mexico. Cyllopsis tomemmeli A. Warren Nakahara, sp. nov., is currently known from 13 specimens (9 males and 4 females) collected on March 26-28, 1959, southeast of San Cristóbal de Las Casas. Despite extensive studies on the butterfly fauna of this region, this species has not since been encountered. We discuss possible relationships between this new species and other species of Cyllopsis.

Description of a new species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the western Andes.

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.

Nomenclatural status of Euptychia mollina Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae).

The purpose of this note is to clarify the nomenclatural status of Euptychia mollina Hübner, 1818, the type species of Euptychia Hübner, 1818, as there seems to be confusion regarding its year of publication. Due to an unfortunate oversight, Lamas (2004) listed the name as Euptychia mollina (Hübner, [1813]), and this mistake has been repeated in the subsequent literature (e.g. Brévignon 2005; Warren et al. 2014; Neild et al. 2014).

Two new species of Euptychia Hübner, 1818 from the upper Amazon basin (Lepidoptera, Nymphalidae, Satyrinae).

Abstract 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.