Archeuptychia, Forster, 1964

Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), Zoological Research 2023, pp. 1-73 : 17

Archeuptychia clade

The ‘ Archeuptychia clade’ is a relatively small clade recognized in Espeland et al. (2019a), and that work showed its placement as a weakly supported sister group to the much larger ‘ Splendeuptychia clade’, which is also found in the extended backbone tree in this study ( Figure S1 ). In the FULL dataset it is, however, weakly supported as sister to a clade including Chloreuptychia and the Pareuptychia clade ( Figure S2 ), while in the 4GENES dataset ( Figure S3 ) it is sister to Chloreuptychia , and these two again are sister to the ‘ Splendeuptychia clade’. Previous molecular phylogenetic studies only sampled a few representatives and thus did not result in recognition of this clade ( Marín et al., 2017; Murray & Prowell, 2005) or subsumed it within the ‘ Pareuptychia clade’ ( Peña et al., 2010), but close relationships among Archeuptychia , species placed here in Pseudeuptychia , Chloreuptychia and Cisandina were variously implied in these studies. The clade ( Figure 8 ) currently includes 16 described species and at least three undescribed species. Currently, four genera, Pseudeuptychia (seven species), Archeuptychia (monotypic), Cisandina (seven species) and Stephenympha (three species) are recognized in the clade. The two sampled Stephenympha species are here found to be sister groups, with some support in the best trees (FULL 90/91, 4GENES 90.2/88), and are well-supported as members of the ‘ Archeuptychia clade’, but their exact placement within the clade needs to be investigated, since support of relationships within the group are rather low ( Figure 8 ). Here, we provisionally move four species to Pseudeuptychia (comb.n.) that were placed in Chloreuptychia prior to this study, since these species form a strongly supported clade that contains Euptychia languida Butler, 1871 , the type of Pseudeuptychia , but that excludes Papilio chloris Cramer, 1780 , the type species of Chloreuptychia ( Marín et al., 2017; Nakahara et al., in preparation). We do not know of any taxonomic issues surrounding Archeuptychia , and thus this monotypic genus does not require study at present. Members of the ‘ Archeuptychia clade’ are small to medium-sized butterflies, all rather typical of euptychiines in terms of size. However, unlike most other euptychiine groups, the clade harbours some of the more ‘showy’ euptychiines, with many species bearing iridescent blue or green dorsal and/or ventral coloration. Possible synapomorphies and distinctive characters for the clade include the following: (a) lamella antevaginalis at least somewhat sclerotized (the degree of sclerotization is apparently variable, ranging from a rather weakly sclerotized plate in species such as Pseudeuptychia herseis comb.n., to a more heavily sclerotized plate in species such as Archeuptychia cluena ). The male genitalia of species in Cisandina show some interesting features, such as a sclerotized portion of the manica extending from the phallobase, a character not observed in other euptychiines. Stephenympha eriphule has a uniformly sclerotized eighth tergite, which is an unusual feature for euptychiines otherwise known only for species in the genus Euptychia . However, a similar form of the eighth tergite is observed in Cristalinaia and species in Amphidecta , but the eighth tergite of these species appears to have a membranous portion in the middle, thus not being uniformly sclerotized. Members of the ‘ Archeuptychia clade’ are known exclusively from east of the Andes, except for Pseudeuptychia hemileuca , which is still known only from a single syntype, that was reportedly collected in western Colombia. Some species are confined to cloud forest along the slopes of the eastern Andes, while the majority of species occur in lowland rainforest from the Amazon basin and/or Guianas, with a few species found in the Atlantic coastal forest of Brazil. Information on the early stage biology for species in the ‘ Archeuptychia clade’ is limited, like many other major euptychiine clades, but the immature stages of Pseudeuptychia marica comb.n. and two species of Cisandina were recently described, with caterpillars feeding on the herbaceous bamboo genera Pariana , Olyra and Taquara ( Poaceae : Bambusoideae: Olyreae) (Nakahara, Rodríguez-Melgarejo, et al., 2022; Tejeira et al., 2021), and additional data will soon be published (Corahua-Espinoza et al., in preparation).

Figures

FIGURE 1 Summary tree based on the FULL dataset showing the major clades supported in this study. Colours are the same as in Espeland et al. (2019a). Support for major clades is shown. Interpretation of support values can be found in the legend.

FIGURE 2 Phylogeny of Euptychia based on the FULL dataset. Interpretation of support values can be found in the legend. The scale bars under the butterfly images equal 1 cm.

FIGURE 3 Phylogeny of the Cyllopsis clade based on the FULL dataset. Interpretation of support values can be found in the legend. The scale bars under the butterfly images equal 1 cm.

FIGURE 8 Phylogeny of the ’Archeuptychia clade’ + Chloreuptychia based on the FULL dataset. Interpretation of support values can be found in the legend. The scale bars under the butterfy images equal 1 cm.