Tomasz W. Pyrcz

Diversity and distribution patterns of Pronophilina butterflies (Lepidoptera: Nymphalidae: Satyrinae) along an altitudinal transect in north-western Ecuador.

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 brachypterous butterfly

Butterflies of the genus Redonda Adams & Bernard (Lepidoptera: Nymphalidae: Satyrinae) are endemic to the Andes of Venezuela. They comprise a monophyletic group of five allopatric taxa, females of which show various degrees of wing reduction and ability to fly. The female of Redonda bordoni Viloria & Pyrcz sp. nov. appears to be brachypterous and incapable of sustained flight, a phenomenon previously unknown within the Rhopalocera.

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.

Latitudinal gradient and spatial covariance in species richness of tropical Lepidoptera in the Andes

We investigated whether Lepidoptera species richness in the tropical Andes is distributed evenly across the latitudinal gradient, explored the spatial correlations between different taxonomic groups, and investigated historical and ecological factors responsible for shaping the observed patterns. We interpolated species richness from collecting records of nine ecologically diverse taxa of montane Lepidoptera, eight of butterflies (Ithomiini, Acraeini, Callicorini, Pronophilina, Aporiina, Leptophobia, Penaincisalia, and Forsterinaria) and one of moths (Geometridae: Erateina) along elevational transects from 1000 to 3500 m every two degrees in latitude between 11°N and 27°S. We used multiple regression to assess the role of climatic and historical variables in shaping their species richness patterns, and Spearmans correlation coefficient to examine the correlation of species richness gradients between pairs of taxa. We found that Lepidoptera species richness in the tropical Andes is distributed unevenly across the latitudinal gradient, differing considerably between the least and the most diverse transects, with a peak at 3–11°S, and is strongly correlated among all nine taxa. Geological age of the Andes, area and climatic seasonality explained the most variation in species richness, with age dominating in the north and seasonality in the south. The latitudinal gradient in east Andean Lepidoptera species richness is likely a result of historical and contemporary factors, in particular temperature seasonality and the greater age of the Andes south of the Equator. Local aspects of species richness patterns are due to small‐scale geological or climatic phenomena, for example, the peak at 3–5°S which results from local mixing of east‐west Andean faunas in the Huancabamba Deflection area.

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.

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.

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.

Differences in butterfly (Nymphalidae) diversity between hillsides and hilltop forest patches in the northern Andes

Andean montane butterflies have unique diversity patterns in each mountain system, and the knowledge on how their community diversity changes at local scale has potential important implications on designing regional conservation strategies. The multiplicative partition of the diversity and the effective number of species provides a useful tool in studying the patterns of diversity in heterogeneous Andean cloud forest habitats. Here, we evaluated diversity in three nearby sites in the Andean Central Cordillera of Colombia, two hillsides – on western (Cauca canyon) and eastern (Aburrá valley) slopes – and one hilltop. We collected a total of 1039 individuals belonging to 55 species of butterflies. Alpha diversity differs between hillsides and the hilltop. Beta diversity in the three areas show differences in richness (q0) and in diversity of the order of magnitude of q1 and q2, with the major differences between hillsides and hilltop. This study shows important differences in butterfly (Nymphalidae) assemblages on a local scale, and suggests that any efforts directed towards comprehensive protection of a given area in a heterogeneous mountain landscape must be focused on the complete forested area, not being limited to a hilltop or to a single hillside.

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.

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.