Splendeuptychia Forster 1964
Creators: name: Espeland, Marianne
name: Nakahara, Shinichi
name: Zacca, Thamara
name: Barbosa, Eduardo P.
name: Huertas, Blanca
name: Marín, Mario A.
name: Lamas, Gerardo
name: Benmesbah, Mohamed
name: Brévignon, Christian
name: Casagrande, Mirna M.
name: Fåhraeus, Christer
name: Grishin, Nick
name: Kawahara, Akito Y.
name: Mielke, Olaf H. H.
name: Miller, Jacqueline Y.
name: Nakamura, Ichiro
name: Navas, Vanessa
name: Patrusky, Brooke
name: Pyrcz, Tomasz W.
name: Richards, Lindsay
name: Tan, Denise
name: Tyler, Stephanie
name: Viloria, Angel
name: Warren, Andrew D.
name: Xiao, Lei
name: Freitas, André V. L.
name: Willmott, Keith R.
Publisher: name: Zenodo
Year: 2023
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Lepidoptera
Nymphalidae
Splendeuptychia

description: <i>Splendeuptychia</i> clade The ‘ <i>Splendeuptychia</i> clade’ was proposed by Peña et al. (2010), and was partially recovered by Espeland et al. (2019a) with the addition of <i>Magneuptychia</i> and <i>Cepheuptychia</i>; the clade also contains those species included within the ‘ <i>Cissia</i> clade’ of Murray and Prowell (2005). In our present study, this clade comprises the genera <i>Splendeuptychia</i> (9 described species), <i>Huberonympha</i> (monotypic), <i>Stevenaria</i> (3 species), <i>Cepheuptychia</i> (3 species), <i>Omacha</i> (monotypic), <i>Vareuptychia</i> (2 species), <i>Magneuptychia</i> (2 species), <i>Capronnieria</i> (2 species), <i>Caeruleuptychia</i> (26 species), <i>Emeryus</i> (3 species), <i>Colombeia</i> (3 species), <i>Malaveria</i> (17 species), <i>Modestia</i> (8 species), <i>Scriptor</i> (monotypic), <i>Paryphthimoides</i> (12 species), <i>Trico</i> <b>gen. n.</b> (monotypic), <i>Occulta</i> <b>gen.n.</b> (monotypic), <i>Deltaya</i> <b>gen.n.</b> (6 species), and <i>Modica</i> <b>gen.n.</b> (5 species). The clade itself, and all its genera, are well-supported in all trees (Figures 10, S 2 and S 3), but the relationships among genera are typically only weakly supported, with a few exceptions. There is relatively good support in both the FULL and 4GENES datasets for a clade including <i>Huberonympha</i>, <i>Stevenaria</i> and <i>Cepheuptychia</i> being the sister group to the remainder of the clade, followed by <i>Trico</i> <b>gen.n</b>. and <i>Occulta</i>, which either form a clade or a grade. There is also good support for a clade consisting of <i>Malaveria</i>, <i>Colombeia</i>, <i>Emeryus</i>, <b> <i>Deltayagen</i>.n.</b> , <i>Scriptor</i>, and <i>Parypthimoides</i> (all trees <b>≥</b> 100/95), which, with the exception of <i>Emeryus</i> and <i>Scriptor</i>, which were not included there, were also found in the backbone tree in this study and in Espeland et al. (2019a). The placement of genera within this clade is, however, rather uncertain, and the positions of <i>Deltaya</i> <b>gen.n.</b> and <i>Emeryus</i> seem especially unstable. In the best FULL tree <i>Deltaya</i> <b>gen.n.</b> is sister to <i>Scriptor</i> (Figure 10), in the second best FULL tree it is sister to <i>Capronnieria</i>, and in both 4GENES trees it is sister to <i>Scriptor</i> + <i>Colombeia</i>, but no placement is well-supported. <i>Emeryus</i> similarly moves around, and in the best FULL tree it is sister to <i>Malaveria</i> + <i>Colombeia</i> (on a very short branch) and in the second best FULL tree it is sister to <i>Modica</i> <b>gen.n</b>. Both 4GENES trees again give different placements of this genus; in the best tree it is sister to <i>Parypthimoides</i> and in the second best tree it is instead sister to a clade consisting of <i>Scriptor</i>, <b> <i>Deltayagen</i>.n.</b> and <i>Modica</i> <b>gen.n.</b> (again on a very short branch). Three genes or more were available for at least some samples of both <i>Deltaya</i> <b>gen. n.</b> and <i>Emeryus</i>, but none of these genera were included in the backbone, so long-branch-attraction might be one of the reasons why these genera move around. More data is needed to possibly resolve these relationships, but the very short branches in the backbonemight make this task difficult. Several recent papers have provided generic revisions and descriptions of new taxa from this clade (Zacca et al., 2014; Zacca, Casagrande, et al., 2017; Zacca, Casagrande, Mielke, Huertas, Barbosa, Freitas, et al., 2020 [<i>Paryphthimoides</i>], Zacca, Casagrande, Mielke, Huertas, Espeland, Freitas, et al., 2020 [<i>Vareuptychia</i>], Zacca, Casagrande, Mielke, Huertas, Barbosa, Freitas, &amp; Willmott, 2020 [<i>Emeryus</i>]; Costa et al., 2016 [<i>Huberonympha</i>, <i>Stevenaria</i>, <i>Magneuptychia</i>]; Benmesbah et al., 2018, 2021 [<i>Malaveria</i>, <i>Modestia</i>]; Nakahara, Zacca, et al., 2018 [<i>Caeruleuptychia</i>], Nakahara, Lamas, et al., 2020 [<i>Scriptor</i>], Nakahara et al., 2023 [<i>Caeruleuptychia</i>]; Andrade-Correa et al., 2019 [<i>Omacha</i>]; Viloria, 2022 [<i>Malaveria</i>]). Andrade-Correa et al. (2019) described <i>Colombeia</i> as a monotypic genus with <i>Euptychia mycalesis</i> as type species, but our molecular data show that species to be a member of a strongly supported clade containing two other species, which we here also transfer to <i>Colombeia</i> (<i>Colombeia nossis</i> <b>comb.n.</b> and <i>Colombeia hotchkissi</i> <b>comb.n.</b>). The genus <i>Malaveria</i> was recently described with <i>Euptychia nebulosa</i> as type species (Benmesbah et al., 2021), with three other described species moved into the genus and a further four species described. These last species descriptions highlighted hitherto unrecognized important morphological and molecular variation, but all of the described species are allopatric with respect to their closest relatives, or with only limited evidence for sympatry, and further research is needed to confirm their status. Barbosa et al. (2022) showed that five additional species also form a clade with <i>Malaveria</i> species and are best placed in that genus, and a number of other new species require description. We also provisionally transfer <i>Euptychia argyrospila</i> Butler, 1867 from <i>Yphthimoides</i> to <i>Malaveria</i> (<b>comb.n.</b>) since partial DNA barcode data support this placement, while the male genitalia illustrated by Forster (1964) also appear to be similar to those of other <i>Malaveria</i>, but this classification requires confirmation. Most recently, Viloria (2022) described the new genus <i>Rudyphthimoides</i> with a new species, <i>R</i>. <i>iseai</i>, as type species, and also included several species placed by Barbosa et al. (2022) in <i>Malaveria</i>. While it seems likely that <i>Rudyphthimoides</i> forms a clade (containing <i>M</i>. <i>maepius</i>, <i>M</i>. <i>affinis</i> and <i>M</i>. <i>mythra</i> in Figure 10), retaining this generic name would restrict the name <i>Malaveria</i> to a very closely related, and weakly supported, clade, and require description of a new monotypic genus for <i>M</i>. <i>grimon</i>. Alternatively, recognizing <i>Malaveria</i> as including <i>M</i>. <i>grimon</i>, <i>Rudyphthimoides</i>, and remaining <i>Malaveria</i> results in a strongly supported clade, separated from other genera by a relatively long branch (Figure 10), which is likely to remain stable. We therefore prefer this latter solution and treat <i>Rudyphthimoides</i> (<b>syn.n.</b>) as a synonym of <i>Malaveria</i>. The genus <i>Modestia</i> was recently described with <i>Euptychia modesta</i> as type species (Benmesbah et al., 2021), with the inclusion of one other described species and one newly described species. Molecular data recovered a strongly supported clade containing the three original species of <i>Modestia</i> along with four species previously placed in <i>Magneuptychia</i> (Lamas, 2004), which are here transferred to <i>Modestia</i>. The genus <i>Nubila</i> was described by Andrade-Correa et al. (2019) with <i>Euptychia gera</i> Hewitson, 1850 as type species, with five additional closely related species transferred from <i>Magneuptychia</i> and <i>Splendeuptychia</i>. These species do indeed form a well-supported clade in our study (Figure 10), which is placed without strong support as sister to a clade containing the recently described <i>Splendeuptychia tupinamba</i> and an undescribed species. Both these clades are sister to a clade containing the type species of <i>Splendeuptychia</i>. <i>Splendeuptychia tupinamba</i> and the undescribed species are phenotypically intermediate between <i>Nubila</i> and the clade containing other described <i>Splendeuptychia</i>, and since morphological data also support a close relationship among these species (Huertas, 2014), we believe the best taxonomic solution is to regard them as a single genus, so we therefore synonymize <i>Nubila</i> <b>syn.n.</b> with <i>Splendeuptychia</i>. Considering the high diversity in the wing pattern, venation, and genitalia exhibited by the members of the ‘ <i>Splendeuptychia</i> clade’, it is difficult to identify possible morphological synapomorphies and distinctive characters for the clade. Most of the species of the clade occur in South America, except for members of <i>Vareuptychia</i> and some <i>Paryphthimoides</i> species that occur in Central America. Habitat preferences of species in this clade range from lowland rainforest and savanna to cloud forest up to 2200 m. Adults of <i>Magneuptychia</i>, <i>Emeryus</i> and <i>Paryphthimoides</i> are commonly attracted to traps baited with rotting fruit (LourenÇo et al., 2019; Melo et al., 2019; Santos et al., 2018) and some <i>Caeruleuptychia</i>, <i>Paryphthimoides</i> and <i>Splendeuptychia</i> species show hilltopping behaviour (Benmesbah et al., 2018; Nakahara, Zacca, et al., 2018). Known larvae of the species in this clade feed on various genera of Poaceae (e.g. Beccaloni et al., 2008; Brown, 1992; Corahua-Espinoza, Nakahara, Kabir, et al., 2022; Corahua-Espinoza, Nakahara, Shellman, et al., 2022; DeVries, 1987; Janzen &amp; Hallwachs, 2022; Núñez-Bustos, 2010), with additional records of Cyperaceae and Marantaceae for <i>Vareuptychia themis</i> (Janzen &amp; Hallwachs, 2022) and Heliconiaceae for <i>Modica confusa</i> <b>comb.n.</b> (Janzen &amp; Hallwachs, 2022). For most species of this clade, published information on the natural history, immatures and biology is still unknown, with a few exceptions (Corahua-Espinoza, Nakahara, Kabir, et al., 2022; Corahua-Espinoza, Nakahara, Shellman, et al., 2022; Freitas, 2022; Janzen &amp; Hallwachs, 2022; Kaminski &amp; Freitas, 2008; Nakahara, Hoffman, et al., 2020).
description: Published as part of <i>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. &amp; Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023</i> on pages 18-19, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395
description: {"references": ["Pena, C., Nylin, S., Freitas, A. V. L. &amp; Wahlberg, N. (2010) Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae). Zoologica Scripta, 39, 243 - 258.", "Espeland, M., Breinholt, J., Barbosa, E. P., Casagrande, M., Huertas, B., Lamas, G. et al. (2019 a) Four hundred shades of brown: higher level phylogeny of the problematic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae) based on hybrid enrichment data. Molecular Phylogenetics and Evolution, 131, 116 - 124.", "Murray, D. &amp; Prowell, D. P. (2005) Molecular phylogenetics and evolutionary history of the neotropical Satyrine subtribe Euptychiina (Nymphalidae: Satyrinae). Molecular Phylogenetics and Evolution, 34, 67 - 80.", "Zacca, T., Siewert, R. R., Mielke, O. H. H. &amp; Casagrande, M. M. (2014) A new species of Magneuptychia Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae) from Brazilian savanna. Zootaxa, 3795, 71 - 77.", "Costa, M., Viloria, A\u00b4. 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