Pareuptychia, 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 : 18

Pareuptychia clade

The ‘ Pareuptychia clade’ was proposed first by Murray and Prowell (2005) and corroborated by increasingly comprehensive subsequent studies ( Espeland et al., 2019a; Marín et al., 2017; Peña et al., 2006, 2010). The clade contains 11 described genera, Neonympha (2 species), Pareuptychia (9 species), Erichthodes (monotypic), Megeuptychia (3 species), Satyrotaygetis (2 species), Euptychoides (4 species), Taydebis (3 species), Nhambikuara (11 species), Optimandes (2 species), Amiga (2 species), Xenovena gen. n. (monotypic) and an undescribed genus. The ‘ Pareuptychia clade’ is embedded within a larger clade also containing the ‘ Taygetis clade’, ‘ Splendeuptychia clade’ and ‘ Archeuptychia clade’, and we also include the unplaced genus Chloreuptychia in the discussions here ( Figures 1, S1 – S 3 ). In both FULL trees, Chloreuptychia is sister to the ‘ Pareuptychia clade’, while in both 4GENES trees the genus is sister to the ‘ Archeuptychia clade’. In the backbone tree ( Figure S1 ) the ‘ Pareuptychia clade’ is sister to the Taygetis clade, and here Chloreuptychia is found to be sister to a clade including the ‘ Taygetis clade’, ‘ Splendeuptychia clade’ and ‘ Archeuptychia clade’. These relationships are not well supported and need to be further investigated.

The small genus Amiga is strongly supported as sister group to the remainder of the clade in all datasets. Most generic relationships are well-supported ( Figures 9 , S 2 and S 3 ). The clade is divided into two well-supported subclades, the Optimandes subclade ( Optimandes , Nhambikuara ) and the Pareuptychia subclade ( Xenovena gen.n., Pareuptychia , Erichthodes , Neonympha , Euptychoides , Satyrotaygetis , and an undescribed generic-level clade containing some species currently in Erichthodes ), consistent with previous molecular studies ( Murray & Prowell, 2005; Peña et al., 2010). The placement of Taydebis needs to be further investigated; although it is found as sister to Optimandes + Nhambikuara in all trees, the support for this branch is always low.

Since the checklist Lamas (2004), a number of papers have helped revise the generic classification of the ‘ Pareuptychia clade’, with the description of new genera such as Taydebis ( Freitas, 2003) , Nhambikuara (Freitas, Barbosa, et al., 2018), Amiga (Nakahara, Lamas, et al., 2019) and Optimandes ( Willmott et al., 2019) , the sorting of species in previously described genera ( Andrade-Correa et al., 2019) and descriptions of 11 new species ( Benmesbah, 2015; Brévignon, 2005; Freitas et al., 2011; Freitas, Barbosa, et al., 2018; Huertas, 2011; Huertas et al., 2009; Nakahara, Marin, & Neild, 2016; Willmott et al., 2018, 2019).

We found strong support for a clade containing the type species of Nhambikuara , N. cerradensis , along with nine other species formerly placed in Splendeuptychia ( Lamas, 2004) , a relationship partly indicated in Freitas, Barbosa, et al. (2018). Morphological characters also support this as a relatively compact group of species ( Huertas, 2014) and we thus transfer the nine former Splendeuptychia species into Nhambikuara (comb.n.). We found strong support for the recently described genus Macrocissia ( Andrade-Correa et al., 2019) not being monophyletic, and the two species previously included in this genus are transferred to Satyrotaygetis ( Satyrotaygetis iris [C. Felder & R. Felder, 1867]) and Euptychoides ( Euptychoides inani [Staudinger, 1886]) in concordance with the phylogeny. The combination of morphological characters used by Andrade-Correa et al. (2019) to support Macrocissia , ‘vestigial recurrent vein into the discal cell’ in the forewing, and ‘valvae with a hook-like distal process’ and a ‘finely serrate cucullus in the male genitalia’, is a combination of characters present in previously described genera such as Satyrotaygetis and Euptychoides , and we thus synonymize Macrocissia syn.n. with Satyrotaygetis .

The ‘ Pareuptychia clade’ contains 44 described species, with the genera Pareuptychia and Nhambikuara , with nine and 11 species respectively, being the most diverse. Taxonomic revisions for these two genera are in progress, in addition to work on Taydebis , Euptychoides and Erichthodes . Members of the ‘ Pareuptychia clade’ have a wide distribution, ranging from southern Canada to northern Argentina, and from sea level to almost 2600 m, with the highest diversity in the Amazon basin and the Andes (Marín et al. in preparation). According to Marín et al. (2017), possible synapomorphies (not unique) and/or distinctive characters for the clade include the following: (a) obvious sclerotized cornuti in aedeagus; (b) in the valva, posterior ending of the cucullus acute. Hostplants for members of this clade are mostly bamboos or grasses in the family Poaceae , with records of Cyperaceae in Pareuptychia ( Janzen & Hallwachs, 2022) and Neonympha ( Hamm et al., 2013) , and Marantaceae in Megeuptychia ( Janzen & Hallwachs, 2022) . Immature stages have been described or illustrated for Amiga , Taydebis , Pareuptychia , Optimandes , Satyrotaygetis , Megeuptychia and Nhambikuara (Corahua-Espinoza, Nakahara, Kabir, et al., 2022; Corahua-Espinoza, Nakahara, Shellman, et al., 2022; Freitas, 2003; Freitas, Barbosa, & Marín, 2016; Janzen & Hallwachs, 2022; Murray, 2001b; Willmott et al., 2019). These are morphologically simple and similar to those of several other species of Neotropical Euptychiina larvae, lacking body scoli, and having short- to medium-sized head horns and caudal filaments, and the pupae are short and smooth.

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 9 Phylogeny of the ’Pareuptychia 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.