Mario A. Marín

From the Phylogeny of the Satyrinae Butterflies to the Systematics of Euptychiina (Lepidoptera: Nymphalidae): History, Progress and Prospects

We review the various proposals of evolutionary and classification schemes for Satyrinae and particularly Euptychiina butterflies, assessing progress and prospects of research for the group. Among the highlights is the proposal to include Morphini, Brassolini and Amathusiini as part of Satyrinae. Although it is clear that this hypothesis requires further investigation, phylogenetic studies recently conducted recover this clade as part of Satyrinae with high support. The phylogenetic analyses for Euptychiina carried out to date recover the monophyly of the group and have identified a variety of genera as non-monophyletic. Further work is necessary to resolve the position of the subtribe and the evolutionary relationships of several genera.

Morphological and molecular marker contributions to disentangling the cryptic Hermeuptychia hermes species complex (Nymphalidae: Satyrinae: Euptychiina)

The genus Hermeuptychia is common and widespread through the Americas, from Argentina to the southern United States of America. All eight recognized species within Hermeuptychia are small and brown, with very similar interspecific external morphologies and intraspecifically variable ocelli patterns that render taxonomic identification based on morphology difficult. In our study, we surveyed variability within Hermeuptychia, and evaluated species boundaries based on molecular data (sequences of the ‘barcode’ mitochondrial DNA COI gene) and morphology (mainly male genitalia), using a phylogenetic approach. We found eight DNA‐based and 12 morphological groups in our sampling. Species names were assigned based mainly on comparisons with male genitalia morphology descriptions corresponding to name‐bearing type specimens. Morphological and DNA variability were highly congruent, with the exception of group H, the Hermeuptychia cucullina complex. Also, the barcode region showed a clear threshold for intra‐ and interspecific mean distances around 2%. Based on these results, we circumscribe the species boundaries in the genus Hermeuptychia and discuss conflicts between mitochondrial genes and classic morphological approaches for identifying and delimiting species. Our study revealed cryptic diversity within an ubiquitous genus of Neotropical butterflies.

Euptychia boulleti (Le Cerf) n. comb. (Lepidoptera: Nymphalidae: Satyrinae), a Rare and Endangered Butterfly from Southeastern Brazil

This paper discusses the systematic position of the rare and endangered satyrine butterfly Caenoptychia boulleti Le Cerf, the only included species in Caenoptychia (type species), based on adult morphology and molecular data. The results showed that Caenoptychia Le Cerf belongs to the Euptychia Hübner clade, and the genus is synonymized with Euptychia, new synonymy. Euptychia boulleti (Le Cerf) is a new combination. The male genitalia of E. boulleti showed at least one important synapomorphy with the other species of Euptychia, which is the presence of a posterior projection of the tegumen above the uncus. Molecular data reinforces the position of Caenoptychia within the genus Euptychia.

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.

Interspecific variation in mitochondrial serine transfer RNA (UCN) in Euptychiina butterflies (Lepidoptera: Satyrinae): structure and alignment.

The nucleotide variation and structural patterns of mitochondrial RNA molecule have been proposed as useful tools in molecular systematics; however, their usefulness is always subject to a proper assessment of homology in the sequence alignment. The present study describes the secondary structure of mitochondrial tRNA for the amino acid serine (UCN) on 13 Euptychiina species and the evaluation of its potential use for evolutionary studies in this group of butterflies. The secondary structure of tRNAs showed variation among the included species except between Hermeuptychia sp1 and sp2. Variation was concentrated in the ribotimidina-pseudouridine-cystosine (TψC), dihydrouridine (DHU) and variable loops and in the DHU and TψC arms. These results suggest this region as a potential marker useful for taxonomic differentiation of species in this group and also confirm the importance of including information from the secondary structure of tRNA to optimize the alignments.

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.

Morphology agrees with molecular data: phylogenetic affinities of Euptychiina butterflies (Nymphalidae: Satyrinae)

Euptychiina is the most species‐rich subtribe of Neotropical Satyrinae, with over 450 known species in 47 genera (14 monotypic). Here, we use morphological characters to examine the phylogenetic relationships within Euptychiina. Taxonomic sampling included 105 species representing the majority of the genera, as well as five outgroups. A total of 103 characters were obtained: 45 from wing pattern, 48 from genitalia and 10 from wing venation. The data matrix was analysed using maximum parsimony under both equal and extended implied weights. Euptychiina was recovered as monophyletic with ten monophyletic genera, contrasting previous DNA sequence‐based phylogenies that did not recover the monophyly of the group. In agreement with sequence‐based hypotheses, however, three main clades were recognized: the ‘Megisto clade’ with six monophyletic and three polyphyletic genera, the ‘Taygetis clade’ with nine genera of which three were monophyletic, and the ‘Pareuptyhia clade’ with four monophyletic and two polyphyletic genera. This is the first morphology‐based phylogenetic hypothesis for Euptychiina and the results will be used to complement molecular data in a combined analysis and to provide critical synapomorphies for clades and genera in this taxonomically confused group.

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.

Description of two new species of the Neotropical genus Yphthimoides Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae) from the ‘renata clade’

This paper describes two new species of Neotropical butterflies: Yphthimoides blanquita Barbosa, Marín and Freitas sp. nov., from the dry forest of northwestern Colombia, and Yphthimoides nareta Barbosa & Freitas sp. nov. from northeastern Brazil, based on morphological and molecular data. Adult morphology, including wing shape and pattern as well as male genitalia, is described in detail. Furthermore, analysis of the mitochondrial CoxI ‘barcode’ showed that both new species are quite distinct from all similar Yphthimoides species and additionally, the ‘renata clade’ is defined based on the presence of cornuti in the aedeagus. http://zoobank.org/urn:lsid:zoobank.org:pub:22CA85FF-9682-4016-A300-AB11C71CEDA2