Cyllopsis, R. Felder, 1869

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 : 8

Cyllopsis clade

The ‘ Cyllopsis clade’ was recognized in Espeland et al. (2019a), where it was placed as a strongly supported sister group to a much larger clade including major, species-rich clades such as the ‘ Splendeuptychia clade’, ‘ Taygetis clade’, ‘ Megisto clade’, ‘ Pareuptychia clade’. The clade was similarly recovered in previous molecular phylogenetic studies ( Murray & Prowell, 2005; Nakahara, Janzen, et al., 2015; Peña et al., 2010), and in the recent study of Rosa et al. (2021). Currently, four genera are recognized in the ‘ Cyllopsis clade’, namely Cyllopsis (30 species), Paramacera (5 species), Inbio (monotypic) and Atlanteuptychia (monotypic), and our molecular data suggest further taxonomic changes at the generic level are not needed ( Figures 3 , S 2 and S 3 ). The relationships between all genera are stable and well-supported in all analyses, with Atlanteuptychia being sister to the remainder of the clade, Inbio being sister to Paramacera , and these two again being sister to Cyllopsis . Thus, the generic classification is perhaps the most stable in comparison with other clades in the subtribe that contain multiple genera. Based on our unpublished data, the only known undescribed species within this clade are in Cyllopsis . Only two Cyllopsis species have been described more recently, including the unique Cyllopsis emilia , which appears to be one of the few euptychiine species that may be involved in mimicry with Pieridae butterflies ( Chacon & Nishida,2002; Warren et al., 2018). Conversely, there are likely several currently recognized Cyllopsis species that may represent seasonal forms, so some revision to the species taxonomy is likely needed (Warren, unpublished data). Members of the ‘ Cyllopsis clade’ are relatively small butterflies, with the forewing length of most species not exceeding 25 mm. Possible synapomorphies (not unique) and distinctive characters for the clade include the following: (a) brachia short (about half of uncus in length or shorter),described as ‘not freely articulated’ in Miller (1974); (b) smooth vesica lacking obvious sclerotized cornuti. Unlike other clades in the subtribe, members of the ‘ Cyllopsis clade’ are most diverse in Central America, especially in southern Mexico and Guatemala. In fact, the monotypic genus Atlanteuptychia is the only taxon in this clade found east of the Andes, namely from the Atlantic Forest of Brazil ( Freitas et al., 2013), resulting in a disjunct distribution for the clade. Habitat preferences of species in this clade range from lowland rainforest to cloud forest at over 2000 m, with Cyllopsis being one of only three Euptychiina genera to have diversified extensively in montane regions (the others are Forsterinaria and Moneuptychia ). The immature stages have been described for Inbio hilara feeding on Rhipidocladum ( Poaceae : Bambusoideae: Bambuseae) (Nakahara,Janzen, et al., 2015), and various Cyllopsis species have been reported feeding on bamboo and other grass genera ( Beccaloni et al., 2008), but otherwise there are few comprehensive, detailed descriptions or hostplant records for most species in the clade.

Figures

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.