The sunny butterflies: new species of high montane pierids of the Catasticta poujadei group from the Peruvian Andes (Pieridae, Pierinae, Aporiina).

The Catasticta poujadei group, within the subgenus Hesperochoia Reissinger, is revised. Two new species, C. copernicus Pyrcz & Fhraeus sp. nov., and C. buszkoi Boyer & Pyrcz sp. nov. occurring near the timberline in Junn and Apurmac are described. Catasticta eximia Rber is reinstated as a species separate from C. poujadei, and a new subspecies, C. eximia tapuna ssp. nov., is described. The affinities of the species of the C. poujadei group are evaluated based on COI barcodes. Their distribution and habitats are described.

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 high-altitude species of Penaincisalia Johnson, 1990 (Lepidoptera, Lycaenidae) from the Peruvian Andes.

A new Penelfin species, Penaincisalia jadwigae sp. nov. is described from very high elevations, 45104620 m, in the Department of Huancavelica, Peru. Superficially it is somewhat similar to P. perezi Blint, 2001, but can easily be recognized by wing colour and pattern. A key for the identification of species in the Penaincisalia s. str. group with orange dorsal wing surface colouration is provided. A list of the local butterfly fauna is given, and spatial distribution, thermal regulation and discoloration are briefly discussed. The taxon Shapiroana matusikorum Johnson, 1992 is reinstated from synonymy with Penaincisalia aurulenta Johnson, 1990, as Rhamma matusikorum (Johnson, 1992), stat. rev., comb. n. 17 figures are provided.

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.

Alpha-taxonomy and phylogeny of African Junoniini butterflies based on morphological data, with an emphasis on genitalia, and COI barcode (Lepidoptera: Nymphalidae).

The tribe Junoniini is a predominantly Paleotropical group of the cosmopolitan butterfly subfamily Nymphalinae (Nymphalidae), with highest diversity in the Afrotropical region. Its systematics and relationships are not entirely resolved. Question marks remain concerning the validity of some genera; and the apparently close relationship between the Indo-Australian genus Yoma and the Afrotropical Protogoniomorpha, as evidenced by molecular phylogenies, remains a puzzle. Here, we present a cladistic analysis, based on 42 characters of the male and female genitalia of 41 species of Junoniini belonging to six genera, nearly all of them continental Afrotropical, and 3 species of two Indo-Australian genera Yoma and Rhinopalpa. A ML COI-based tree is produced for 36 species of Afrotropical Junoniini and Yoma. The molecular data are consistent with previous studies. However, morphological analysis does not confirm a close relationship between Protogoniomorpha and Yoma. Despite the evolution of a number of modifications, the male genitalia within all genera and species of the Junoniini share a cohesive build plan, in particular a transformed sacculus, from which Yoma is highly divergent. The position of the genus Kamilla, previously synonymized with Junonia, is discussed. Three East African coast taxa, Junonia elgiva stat. reinst., Protogoniomorpha nebulosa stat. reinst. and Salamis amaniensis stat. reinst., and one from central Africa, Precis silvicola stat. reinst. are raised to species level, based on comparative analysis of their male genitalia.

Two new species of Manerebia Staudinger from paramo habitat in the Colombian Eastern Cordillera of the Andes (Lepidoptera: Nymphalidae, Satyrinae, Pronophilina).

Two new species of the genus Manerebia Staudinger, 1897 (Nymphalidae, Satyrinae) are described from paramo habitat on the eastern slopes of the Colombian Eastern Cordillera in the area of Pisba and La Colorada: Manerebia bernito n. sp. and Manerebia clarita n. sp. Both are morphologically most similar to Manerebia levana and Manerebia pervaga, and the possible phylogenetic relationship between them is hypothesized. Their habitat is described and a proposal of their conservation status is made. With the addition of the two new species described in this study, a total of 15 species of Manerebia are reported from Colombia. However, a molecular analysis should be carried out to validate the taxonomic status of several species of Manerebia, in particular having in mind a potentially high cryptic diversity of this genus.

Contributions to the knowledge of Neotropical Lycaenidae: taxonomy of Johnsonita/ Salazar amp; Constantino, 1995 with descriptions of seven new species (Theclinae: Eumaeini).

The Transandean-Andean genus Johnsonita Salazar Constantino, 1995 is revised on the basis of wing and genitalia morphology. Apart from eight species formerly placed in Johnsonita (Thecla assula Draudt, 1919; Thecla auda Hewitson, 1867; Thecla catadupa Hewitson, 1869; Thecla chaluma Schaus, 1902, Thecla chlamydem Druce, 1907; Thecla pardoa d'Abrera, 1995; Johnsonita johnsoni Salazar Constantino, 1995 and Johnsonita johnbanksi Bálint, 2003), seven new species are described: Johnsonita carpia Bálint, Boyer Pyrcz, sp. n. (Ecuador), Johnsonita iacinta Bálint, Boyer Pyrcz, sp. n. (Peru), Johnsonita ianusca Bálint, Lorenc-Brudecka Pyrcz, sp. n. (Ecuador), Johnsonita oxalida Bálint, Boyer Pyrcz, sp. n. (Peru), Johnsonita subcunicula Bálint, Cerdeña Pyrcz, sp. n. (Peru), Johnsonita turquisca Bálint, Boyer Pyrcz, sp. n. (Ecuador) and Johnsonita zubkova Bálint, Boyer Lorenc-Brudecka, sp. n. (Peru). One species, erroneously recorded from Bolivia under the name Thecla chaluma Schaus, 1902, is not formally described due to insufficient material. An identification key, type material revision, descriptions or redescriptions, and diagnosis for all the taxa recognized are given. Lectotypes for the nominal species Thecla assula Draudt, 1919 and Thecla chlamydem Druce, 1907 are designated. Male secondary wing characters are tabulated. A brief discussion on classification, life-history and male secondary characters of Johnsonita is presented.

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.

Two remarkable new species of Argyrophorus Blanchard from the Peruvian high Andes (Lepidoptera, Nymphalidae, Satyrinae).

Two new species of butterflies of the genus Argyrophorus Blanchard (Nymphalidae, Satyrinae)-A. idealis n. sp. and A. rubrostriata n. sp.-are described from the Cordillera Negra in north-central Peru. Both, as indicated by colour patterns, male and female genitalia, are related to Argyrophorus lamna (Thieme) which occurs further east and southwards. The new species occur in open puna grassland at approximately 4000 m. One of them is exceptional because both fore and hindwings are uniform shining silvery, in this respect similar only to the Chilean A. argenteus Blanchard.

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.

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.

Taxonomy and distribution pattern of the African rain forest butterfly genus Euphaedra Hübner sensu stricto with the description of three new subspecies of Euphaedra cyparissa (Cramer) and one of E. sarcoptera (Butler) (Lepidoptera, Nymphalidae, Limenitidinae, Adoliadini).

Updated data on the distribution, ecology and taxonomy of Euphaedra cyparissa (Cramer) and Euphaedra sarcoptera (Butler) are presented. Three new subspecies of Euphaedra cyparissa and one of Euphaedra sarcoptera are described and their geographic distribution is presented. The monophyly of the genus Euphaedra sensu Hecq is assessed based on morphological, in particular male and female genitalia, and behavioural traits. Possible evolutionary reasons for the convergence of colour pattern between the sympatric subspecies of Euphaedra cyparissa and Euphaedra sarcoptera are discussed.

Stegosatyrus, a new genus of Euptychiina from the grasslands of neotropical realm (Lepidoptera: Nymphalidae: Satyrinae).

Three species of Pampasatyrus Hayward, 1953 (Satyrinae, Pronophilina) are transferred to Stegosatyrus n. gen. (Euptychiina) based on morphological evidence: S. imbrialis (Weeks, 1901) n. comb. from Bolivia (Cochabamba) and northern Argentina; S. ocelloides (Schaus, 1902) n. comb. from Paraguay (Hernandarias and Caaguazú) and Brazil (Midwest, Southeast and South regions); and S. periphas (Godart, [1824]) n. comb. from southern Brazil, northeastern Argentina (Buenos Aires) and Uruguay. A new species is described, Stegosatyrus hemiclara Pyrcz, Boyer & Zacca, n. sp. from the Andes of southern Peru. The neotype of Satyrus periphas Godart, [1824] and the lectotypes of Epinephele imbrialis Weeks, 1901 and Euptychia ocelloides Schaus, 1902 are designated. Redescriptions and illustrations are presented, including information on geographical and temporal distribution, and habitats. The oviposition process, eggs and first instars of S. periphas are published for the first time.

New species of high elevation cloud forest butterflies of the genus Pedaliodes Butler from the northern Colombian Andes (Lepidoptera, Nymphalidae, Satyrinae).

Four new species of Pedaliodes Butler (i.e., P. adrianae, n. sp., P. haydoni, n. sp., P. philinae, n. sp. and P. rodriguezi, n. sp.) are described from the high elevation cloud forests in the Frontino massif in the Colombian Western Cordillera and the northern part of the Central and Eastern Cordilleras. The affinities of these new species with other congeners are discussed. The total number of described species of Colombian Pronophilina is increased to 208, with a particularly high total number of species (88) and proportion of endemic species (26%) in the Western Cordillera.

First description of the early stage biology of the genus Mygona: the natural history of the satyrine butterfly, Mygona irmina in eastern Ecuador.

The immature stages and natural history of Mygona irmina Doubleday (Lepidoptera: Nymphalidae: Satyrinae: Pronophilina) from northeastern Ecuadorian cloud forests are described based on 17 rearings. The dwarf bamboo, Chusquea c.f. scandens Kunth (Poaceae, Bambusoidea) is the larval food plant. Eggs are laid singly on the bottom side of mature host plant leaves. Larvae take 102-109 days to mature from egg to adult. Adults are encountered most frequently on sunny days, flying rapidly over areas dominated by their food plant or feeding on the ground at mammal feces. Males are often encountered inside large forest gaps near patches of bamboo guarding perches in the mid-canopy.

Diversity and distribution patterns of Pronophilina butterflies (Lepidoptera: Nymphalidae: Satyrinae) along an altitudinal transect in north-western Ecuador / Diversidad y patrones de distribución de mariposas de la subtribu Pronophilina (Lepidoptera: Nymphalidae: Satyrinae) en un transecto altitudinal en el nor-oeste de Ecuador

Se realizó en Ecuador un muestreo de mariposas de la subtribu Pronophilina con el fin de evaluar los efectos de altitud sobre los patrones de distribución, diversidad y estructura de la comunidad en un transecto altitudinal. Se demostró una correlación significativa de todos los índices de diversidad y altitud. El máximo de diversidad expresado por la riqueza de especies, fue reportado a 2600 m. Se identificaron dos grupos de especies en la parte inferior y superior del transecto. La comparación de los coeficientes de similitud indicó valores menores en la franja de altitud intermedia. Se demostró que varios pares de especies relacionadas morfo y ecológicamente tienen distribuciones altitudinales mutuamente exclusivas. La comparación con estudios semejantes revelaron una congruencia muy notable en cuanto a los patrones de diversidad altitudinal de los Pronophilina en varias áreas de la cordillera Andina. En particular, el índice de Shannon llega a valores máximos entre 2600 m y 2850 m, aproximadamente 400 - 500 por debajo del limite superior del bosque nublado. El aumento de la diversidad de los Pronophilina con la altitud puede ser relacionado tan solo de manera marginal con la mayor disponibilidad de los recursos limitados. La menor presión por parte de los depredadores, parasitoides y ectoparásitos en mayores altitudes puede contribuir a la mayor abundancia pero no directamente a la diversidad. Esta última, está relacionada con las características intrínsicas del grupo, tales como el solapamiento de grupos faunísticos elevacionales y la tasa de especiación más rápida en el límite superior del bosque.

Samplings of Pronophilina, a species-rich group of neotropical montane butterflies, were carried out along an elevational transect in Ecuador to assess the effect of altitude on their distribution patterns, diversity and community structure. All diversity indices were significantly correlated with altitude. Maximum diversity expressed in species-richness, Shannon index and Fisher alpha was recorded at 2600 m. Two assemblages of species were identified in the lower (below 2100 m) and upper (above 2300 m) sections of the transect by means of correspondence (CA) and cluster analysis. A comparison of Sørensen similarity coefficients showed lower values, thus higher turnover in the intermediate elevational band. Several closely related morphologically and ecologically species were found to have mutually exclusive altitudinal distribution patterns. A comparison with similar studies in Venezuela, Colombia and Peru revealed far reaching congruency of the patterns of altitudinal diversity of Pronophilina in distant areas of the Andes. In particular, the Shannon index reaches its maximum values at 2600-2850 m, which invariably correspond to ca. 400-500 m below the upper limit of cloud forest. Increase of diversity of Pronophilina with altitude is marginally related to higher limited resource availability. The lower pressure of predators and parasites at higher elevation can contribute with higher abundance, but cannot be directly correlated with higher diversity. Higher diversity is related with intrisic characteristics of the group, such as aggregated diversity by overlapping of elevational faunal assemblages and higher speciation ratio towards high elevations, particularly near timberline.

A new species of Eretris Thieme (Lepidoptera: Nymphalidae, Satyrinae) from the Elbow of the Andes region in Bolivia.

A new species of cloud forest butterfly, Eretris julieta n. sp. is described from a region of south-central Bolivia known as the Elbow of the Andes. It is the southernmost known representative of the genus, hitherto known only from a restricted area of interandean valleys in the department of Santa Cruz. Its affinities with other congeners are evaluated.