Revisional Notes on the Cloud Forest Butterfly Genus Oxeoschistus Butler in Central America (Lepidoptera: Nymphalidae: Satyrinae).

So far, six species of Oxeoschistus Butler, including its junior synonym Dioriste Thieme, were listed from Central America, with five of them from Costa Rica alone, which appears to represent the highest regional diversity of this Neotropical montane butterfly genus. Our research based on field work, morphological studies and barcode analysis proved that one record is a misunderstanding perpetuated in scientific literature for over a century: Oxeoschistus cothonides Grose-Smith is identified here as an individual form of the female of O. cothon Salvin. The presence of Oxeoschistus tauropolis (Westwood) in Costa Rica, subject to some controversy, is confirmed, and a new local subspecies is described from Costa Rica, O. tauropolis mitsuko Pyrcz & Nakamura n. ssp. Specific status of O. euriphyle Butler is reinstated based on morphological and molecular data. A new subspecies O. hilara lempira Pyrcz n. ssp. is described from Honduras. O. puerta submaculatus Butler is reported for the first time from the Darién region on the Panama-Colombia border. Species relationships are preliminarily evaluated based on COI data concluding, among others, that O. hilara and O. euriphyle are less closely related than previously assumed. Altitudinal and distributional data are revised, and ecological and behavioural information of all the species of Central American Oxeoschistus is provided.

Considerations on the Taxonomy of the Genus Arhuaco Adams and Bernard 1977, and its Relationships with the Genus Pronophila Doubleday [1849] (Nymphalidae, Satyrinae).

Arhuaco Adams & Bernard (1977) is one of the least known genera of Neotropical Satyrinae. It comprises two species and presents an unusual disjunct distribution, with A. ica Adams & Bernard (1977), endemic to the isolated Colombian Sierra Nevada de Santa Marta, and A. dryadina (Schaus 1913) found in the mountains of Costa Rica and Panama. Here, the female of A. dryadina is described, and a new generic diagnosis is presented. Affinities with other genera of the subtribe Pronophilina, in particular the potential closest relatives, such as Pronophila Doubleday (1849), are investigated based on morphological, molecular, ecological, and behavioral data. Results from molecular and morphological sources are incongruent. Molecular data indicate that Arhuaco is paraphyletic, with A. dryadina segregating within the Pronophila clade. Morphological data, by contrast, indicate a closer affinity between the two species currently placed in Arhuaco, favoring the monophyly of the genus, and show no consistent synapomorphies for Arhuaco + Pronophila. A vicariance biogeographical scenario is evaluated.

Uncovered Diversity of a Predominantly Andean Butterfly Clade in the Brazilian Atlantic Forest: a Revision of the Genus Praepedaliodes Forster (Lepidoptera: Nymphalidae, Satyrinae, Satyrini).

The genus Praepedaliodes Forster, 1964, the only representative of the mega-diverse mostly Andean Pedaliodes complex lineage in the Brazilian Atlantic Forest, is revised. Prior to this study, four species were known, P. phanias (Hewitson, 1862), P. granulata (Butler, 1868), P. amussis (Thieme, 1905) and P. exul (Thieme, 1905). Here, a further six are described, all from SE Brazil, expanding to 10 the number of species in this genus. Lectotypes are designated for P. phanias, P. granulata and P. amussis. The genus is most diverse in the Serra da Mantiqueira (São Paulo, Rio de Janeiro, Minas Gerais) and in the Serra Geral (Paraná, Santa Catarina) with seven species occurring in both ranges. Praepedaliodes phanias is the most widespread species and the only one found in the western part of the Atlantic Forest; only this species and P. duartei Dias, Dolibaina & Pyrcz n. sp. occurring to near sea level. Other species, P. zaccae Dolibaina, Dias & Pyrcz n. sp., P. francinii Freitas & Pyrcz n. sp., P. sequeirae Pyrcz, Dias & Dolbaina n. sp., P. landryi Pyrcz & Freitas n. sp. and P. pawlaki Pyrcz & Boyer n. sp. are strictly montane and the highest species richness is reached at 1400-1800 m. One species, P. sequeirae n. sp., is a narrow endemic found only at timberline in the Agulhas Negras massif above 2300 m. Immature stages are described for two species, P. phanias and P. landryi n. sp. Molecular data (barcode region of cytochrome oxidase, subunit I) and adult morphology, including male and female genitalia, support the genus as monophyletic, belonging to a predominantly Andean clade of the Pedaliodes Butler, 1867 complex. Morphological evidences, in particular female genitalia comparative analysis, indicate the genera Physcopedaliodes Forster, 1964 and Panyapedaliodes Forster, 1964 as possibly the closest relatives to Praepedaliodes. Molecular data are inconclusive in this respect.

DNA Barcoding of an Assembly of Montane Andean Butterflies (Satyrinae): Geographical Scale and Identification Performance.

DNA barcoding is a technique used primarily for the documentation and identification of biological diversity based on mitochondrial DNA sequences. Butterflies have received particular attention in DNA barcoding studies, although varied performance may be obtained due to different scales of geographic sampling and speciation processes in various groups. The montane Andean Satyrinae constitutes a challenging study group for taxonomy. The group displays high richness, with more of 550 species, and remarkable morphological similarity among taxa, which renders their identification difficult. In the present study, we evaluated the effectiveness of DNA barcodes in the identification of montane Andean satyrines and the effect of increased geographical scale of sampling on identification performance. Mitochondrial sequences were obtained from 104 specimens of 39 species and 16 genera, collected in a forest remnant in the northwest Andes. DNA barcoding has proved to be a useful tool for the identification of the specimens, with a well-defined gap and producing clusters with unambiguous identifications for all the morphospecies in the study area. The expansion of the geographical scale with published data increased genetic distances within species and reduced those among species, but did not generally reduce the success of specimen identification. Only in Forsterinaria rustica (Butler, 1868), a taxon with high intraspecific variation, the barcode gap was lost and low support for monophyly was obtained. Likewise, expanded sampling resulted in a substantial increase in the intraspecific distance in Morpho sulkowskyi (Kollar, 1850); Panyapedaliodes drymaea (Hewitson, 1858); Lymanopoda obsoleta (Westwood, 1851); and Lymanopoda labda Hewitson, 1861; but for these species, the barcode gap was maintained. These divergent lineages are nonetheless worth a detailed study of external and genitalic morphology variation, as well as ecological features, in order to determine the potential existence of cryptic species. Even including these cases, DNA barcoding performance in specimen identification was 100% successful based on monophyly, an unexpected result in such a taxonomically complicated group.

Description of a New Species of the Andean Butterfly Genus Forsterinaria Gray (Lepidoptera: Nymphalidae) with Considerations on an Apparently New Structure in Male Genitalia.

The butterfly genus Forsterinaria Gray is the only strictly montane representative of the diverse Neotropical subtribe Euptychiina (Nymphalidae, Satyrinae), with 24 described species. Recent research in some of the most isolated and highly diverse Andean regions, such as central Peru, show that its total species richness is still underestimated. An example is the new species described here, Forsterinaria emo n. sp., which is particularly interesting because of an unusual structure discovered in its male genitalia which consists of a bunch of bristle-like processes, composing a fringe-like formation on the dorsum of the tegumen. No similar, homologous structure was found in any congener, nor indeed, in any species of diurnal Lepidoptera. Scanning electron microscope studies revealed that the microstructure of the processes resembles a membrane lining the tegumen. Its function is unknown but two hypotheses are discussed based on a comparative study with other genital structures of butterflies. We argue that it may help stabilizing the partners in the process of mating or it may serve as a 'mating plug', preventing the female from multiple copulations.

The presence-absence situation and its impact on the assemblage structure and interspecific relations of Pronophilina butterflies in the Venezuelan Andes (Lepidoptera: Nymphalidae).

Assemblage structure and altitudinal patterns of Pronophilina, a species-rich group of Andean butterflies, are compared in El Baho and Monte Zerpa, two closely situated and ecologically similar Andean localities. Their faunas differ only by the absence of Pedaliodes ornata Grose-Smith in El Baho. There are, however, important structural differences between the two Pronophilina assemblages. Whereas there are five co-dominant species in Monte Zerpa, including P. ornata, Pedaliodes minabilis Pyrcz is the only dominant with more than half of all the individuals in the sample in El Baho. The absence of P. ornata in El Baho is investigated from historical, geographic, and ecological perspectives exploring the factors responsible for its possible extinction including climate change, mass dying out of host plants, and competitive exclusion. Although competitive exclusion between P. ornata and P. minabilis is a plausible mechanism, considered that their ecological niches overlap, which suggests a limiting influence on each other's populations, the object of competition was not identified, and the reason of the absence of P. ornata in El Baho could not be established. The role of spatial interference related to imperfect sexual behavioral isolation is evaluated in maintaining the parapatric altitudinal distributions of three pairs of phenotypically similar and related species of Pedaliodes, Corades, and Lymanopoda.