Considerations on the Systematics of Neotropical Pierina, with the Description of Two New Species of Phulia Herrich-Schäffer from the Peruvian Andes (Lepidoptera: Pieridae, Pierinae, Pierini).

A comparative analysis of high-Andean Pierina was carried out, including a total of 25 species. Based on morphological evidence, with an emphasis on venation and genitalia and molecular data, using three genetic markers, we confirm the recent subjective synonymy of the generic names Tatochila Butler, 1870, Piercolias, Staudinger, 1894, Hypsochila Ureta, 1955, Infraphulia Field, 1958, Pierphulia Field, 1958, and Theochila Field, 1958 with Phulia Herrich-Schäffer, 1867. Two new species are described, namely Phulia stoddardi Pyrcz & Cerdeña n. sp., from the Andes of Central Peru, which occurs at an unusually high altitude of close to 5000 m a.s.l. in dry puna habitat, and Phulia phantasma Lamas, Willmott & Boyer n. sp., from dry montane forests in northern Peru and southern Ecuador. An overview of high-elevation butterflies is presented, with some discussion on adaptations to this environment.

A new species of the enigmatic genus Oressinoma Doubleday from the Andes of central Peru (Lepidoptera: Nymphalidae, Satyrinae).

A new, and only the third known species of the Neotropical montane genus Oressinoma Doubleday is describedO. sorina n. sp., from the Andes of central Peru. It is distinguishable immediately from the other two congeners by the shape of the hindwing underside submarginal orange band, and by the male genitalia. The systematics of Oressinoma are reviewed. A preliminary analysis is carried out based on COI barcode confirming the separate specific status of O. sorina n. sp. in relation to other two congeners. Both barcode and genital morphology data suggest that the widespread O. typhla Doubleday may be a complex of allopatric or, locally parapatric species. The genus Oressinoma is the only neotropical member of the predominantly Australian subtribe Coenonymphina, represented in the entire Holarctic by one genus onlyCoenonympha Hbner, considered as the putative sister-genus of Oressinoma. Their origins and relationships are briefly discussed.

Conserved ancestral tropical niche but different continental histories explain the latitudinal diversity gradient in brush-footed butterflies.

The global increase in species richness toward the tropics across continents and taxonomic groups, referred to as the latitudinal diversity gradient, stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluate several of these hypotheses to explain spatial diversity patterns in a butterfly family, the Nymphalidae, by assessing the contributions of speciation, extinction, and dispersal, and also the extent to which these processes differ among regions at the same latitude. We generate a time-calibrated phylogeny containing 2,866 nymphalid species (~45% of extant diversity). Neither speciation nor extinction rate variations consistently explain the latitudinal diversity gradient among regions because temporal diversification dynamics differ greatly across longitude. The Neotropical diversity results from low extinction rates, not high speciation rates, and biotic interchanges with other regions are rare. Southeast Asia is also characterized by a low speciation rate but, unlike the Neotropics, is the main source of dispersal events through time. Our results suggest that global climate change throughout the Cenozoic, combined with tropical niche conservatism, played a major role in generating the modern latitudinal diversity gradient of nymphalid butterflies.

A New Species of Butterfly of the Genus Protopedaliodes from the Isolated Tramen Tepui in the Guyana Shield (Lepidoptera, Nymphalidae, Satyrinae).

A new species of Protopedaliodes Viloria & Pyrcz, a satyrine butterfly genus endemic to the highest part of the Guyana Shield, P. arekuna Pyrcz & Stachowicz n. sp., is described from the summit area, ca. 2400 m, of Tramen Tepui, an isolated mountain situated on the Venezuela-Guyana border. It is a remarkable finding as it is probably a narrow endemic, and only the fourth known member of the genus. Morphologically it most closely resembles P. kukenani Viloria & Pyrcz from the Roraima-Kukenán twin peaks. COI barcode analysis shows, however, high genetic distances between these two species, 9-10%. Protopedaliodes taxonomy is briefly revised, from the perspective of the monophyly of the genus based on preliminary molecular and morphological comparative data, including the female genitalia described for the first time for P. kukenani and P. ridouti Viloria & Pyrcz.

Solving the cryptic diversity of the genus Manerebia Staudinger in northern Peru: description of new species and considerations on the biogeographical role of the Huancabamba Deflection (Nymphalidae: Satyrinae: Pronophilina).

The Huancabamba Deflection in the Andes of northern Peru and southern Ecuador is a pivotal area for Neotropical biogeography, where exceptional biodiversity coincides with high rates of endemism. These characteristics are well illustrated within the montane butterfly genus Manerebia Staudinger (Nymphalidae, Satyrinae). Here, six new, apparently endemic species, and two new subspecies, are described from this region: M. inducta Pyrcz Willmott n. sp., M. ronda Pyrcz Boyer, n. sp., M. ronda amplia Pyrcz Boyer, n. ssp., M. punku Pyrcz Farfn n. sp., M. huamanii Cerdea Pyrcz n. sp., M. placida Pyrcz Willmott n. sp., M. granatus Willmott, Radford Pyrcz n. sp. and M. prattorum udima Pyrcz Boyer, n. ssp. A total of 22 species of Manerebia is reported from the study region, as much as half the total number of species in this genus distributed throughout the tropical Andes. The alpha-taxonomy of Manerebia is particularly demanding, as colour patterns have apparently converged among different species making identification virtually impossible in some cases without comparison of their genitalia, which prove taxonomically and phylogenetically highly valuable. On the other hand, several species differ markedly in wing colour patterns and occur at different elevations but have identical genitalia. Furthermore, our data show that DNA barcoding is only partly viable as an alpha-taxonomic tool, as some markedly different species of Manerebia, in terms of external morphology and ecological preferences, are genetically not separable using only COI markers. On the other hand, several species barely differing morphologically are identified based on barcode. Barcodes of 26 species, including 18 from the northern Andes, are studied here, showing strong differences within some taxa previously considered conspecific based on morphological characters, suggesting that their taxonomic status needs to be re-evaluated. In particular, M. trimaculata and the widely distributed polytypic M. inderena may be highly variable species or complexes of several species. A total of 16 species are found north of the Chamaya valley in southern Ecuador and extreme northern Peru, compared to 14 species south of it in northern Peru. The faunal (Jaccard) similarity coefficient of the two adjacent regions is low (0.3), reflecting the role of the Huancabamba Deflection as a biogeographical barrier.

Subspecies-level systematics and affinities of Cheimas Thieme-an endemic genus of the subparamo of the Venezuelan Cordillera de Mérida (Lepidoptera: Nymphalidae, Satyrinae).

The validity of the monobasic neotropical butterfly genus Cheimas Thieme (Nymphalidae, Satyrinae, Satyrini, Pronophilina) is discussed, and confirmed based on morphological and molecular data. Cheimas opalinus (Staudinger), endemic to the Venezuelan Cordillera de Mérida, and considered prior to this study to be monotypic and restricted to the central part of the range, is demonstrated to be polytypic and more widely distributed. Five subspecies are recognised, differing mostly in their dorsal patterns, in particular the shape and colour of hindwing greenish-blue patch. Mitochondrial DNA sequences (COI) were obtained for three of them. The nominate subspecies is found in the central part of the range, in the Sierra Nevada and La Culata. The other subspecies are found as follows: C. opalinus dominici n. ssp.; in the Santo Domingo valley in the centre-north; C. opalinus cristalinus n. ssp. in the north; C. opalinus iosephi n. ssp. on the eastern slopes, and C. opalinus rosalinus n. ssp. in the southern Páramo El Batallón massif. A hybrid zone between the latter two subspecies was detected in the northern part of the Batallón massif based on unusual individual variation and intermediate phenotypes. All the populations of Cheimas opalinus occur in the forest-paramo ecotone at 2800-3400 m a.s.l., with the notable exception of C. opalinus cristalinus n. ssp. found also in mid-elevation forests down to 2300 m a.s.l.

Systematic revision of the Andean butterfly genus Orophila Staudinger, 1886 (Lepidoptera: Nymphalidae: Biblidinae).

A systematic review of the Neotropical butterfly genus Orophila Staudinger, 1886 based on morphological characters and geographical distribution is presented. Four species are recognized: O. campaspe (Hewitson, 1869) stat. rest., O. cardases (Hewitson, 1869), O. cecidas (Hewitson, 1869) stat. rest. and O. diotima (Hewitson, 1852). Other changes proposed include treating Orophila diotima footei (Lamas, 2003) syn. n. and O. cardases zurita (Fruhstorfer, 1916) syn. n. as synonyms of other taxa. Illustrations of adult males and male genitalia of all taxa are presented and the importance of morphological characters in both males and females is evaluated. Female specimens and genitalia are illustrated for the first time for O. c. cardases, O. cecidas and O. diotima. Lectotypes are designated for two nominal taxa.

Páramo de Belmira as an important centre of endemism in the northern Colombian Andes: new evidence from Pronophilina butterflies (Lepidoptera: Nymphalidae, Satyrinae, Satyrini).

Páramo de Belmira (Páramo de Santa Inés) is the highest part of the Andean Central Cordillera in the Colombian department of Antioquia. It harbours a pocket of highlands grassland vegetation isolated from the nearest southerly other large paramo extensions by some 150 km. Butterflies sampling was carried out for over three years in the cloud forest-paramo mosaic and open grassland at 2650-3350 m. As a result, several new taxa of various butterflies groups were identified, including three species and two subspecies belonging to the Satyrinae subtribe Pronophilina, described herein: Lymanopoda casneri n. sp., L. ionius lilliput n. ssp., Panyapedaliodes rojasi n. sp., Pedaliodes nutabe n. sp. and Apexacuta orsedice mariadelmarae n. ssp. L. casneri is particularly interesting from the biogeographical perspective as its nearest relative is L. hazelana Brown, found in south-central Ecuador, some 1000 km southwards. Overall, 48 species of Pronophilina were identified. Considering the occurrence of several endemic species and subspecies of butterflies in the extremely restricted paramo grasslands in the study area, and the anthropogenic pressure from surrounding locations, it is worth attributing the Páramo de Belmira a protection area status.

Hypandrium as a key character in resolving species-level taxonomy on the example of Perisama oppelii (Latreille) (Lepidoptera: Nymphalidae, Biblidinae).

A detailed comparative study of the subspecies of Perisama oppelii (Latreille, [1809]) is carried out. Ten out of eleven subspecies are illustrated, including the females of P. o. bleuzeni Attal & Crosson du Cormier, 1996 and P. o. cristal Attal & Crosson du Cormier, 1996 for the first time. Male genitalia of P. o. amalia Oberthür, 1916, P. o. bleuzeni, P. o. cristal, P. o. erebina Oberthür, 1916, P. o. oppelii (Latreille, [1809]), P. o. viridinota Butler, 1873, and P. o. xanthica (Hewitson, 1868), as well as female genitalia of P. o. bleuzeni, P. o. cristal and P. o. oppelii are also presented for the first time. Based on the characters of hypandrium, a separate specific status is proposed for P. bleuzeni stat. n. Taxonomical value of the hypandrium is evaluated. The co-evolution of male and female genitalia is discussed.