Megisto, Hübner, 1819

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 : 11

Megisto clade

The ‘Megisto clade’ currently comprises 10 genera, Megisto (with 2 species), Pharneuptychia (5 species, in addition to ‘ Pharneuptychia ’ innocentia in the ‘ Amphidecta clade’), Moneuptychia (11 species), Graphita (1 species), Stegosatyrus (4 species), Koutalina (1 species), Cissia (10 species), Carminda (4 species), Vanima (3 species) and Yphthimoides (21 species). The relationships between genera in our analyses ( Figures 4 , S 2 and S 3 ) match those of previous molecular studies ( Murray & Prowell, 2005; Peña et al., 2010) and are stable, with the exception of the placement of the genus Koutalina . In the best FULL tree Koutalina is found as sister to the remainder of the clade, except Megisto, but is placed deeper within the clade as sister to Graphita in the second best FULL tree, and as sister to Moneuptychia + Pharneuptychia in the best 4GENES tree. This genus was not included in the backbone tree, and only COI sequences are available for the included samples, which likely explains the instability of this taxon. The recently described Yphthimoides kinyoni (Nakahara, Barbosa, Nakamura, et al., 2021) is found within the genus Yphthimoide s, as expected, in the best FULL tree, but in the second best FULL tree and in the best two 4GENES trees it is placed as sister to Stegosatyrus within Cissia . This instability is likely also caused by its lack of very close relatives within the genus and only short COI sequences being available, but a placement within Yphthimoides is also justified morphologically (Nakahara, Barbosa, Nakamura, et al., 2021).

It is estimated that the diversity of this clade will increase with the description of new species for Moneuptychia and Pharneuptychia ( Barbosa et al., 2022; Freitas et al., in preparation) in forthcoming papers. Some recent papers provided generic taxonomic revisions and description of new taxa in this clade ( Zacca et al., 2013 [ Stegosatyrus ], Zacca, Casagrande, et al., 2018 [ Cissia ], Zacca, Casagrande, Mielke, Huertas, Espeland, Freitas, et al., 2020 [ Vanima ]; Barbosa et al., 2015 [ Yphthimoides ], Barbosa et al., 2016 [ Yphthimoides ], Barbosa et al., 2020 [ Yphthimoides ]; Freitas et al., 2015 [ Moneuptychia ]; Nakahara, Barbosa, et al., 2016 [ Graphita ], Nakahara, Barbosa, Nakamura, et al., 2021 [ Yphthimoides ]; Henao-Bañol & Meneses, 2018 [ Pharneuptychia ]; Zhang et al., 2020 [ Cissia ], Zhang et al., 2022 [ Cissia ]; Benmesbah et al., 2021 [ Koutalina ]). Barbosa et al. (2022) provided a molecular phylogeny of Yphthimoides and relatives, and investigated the biogeography of the group. Possible synapomorphies and relatively distinctive characters for the clade include the following: (a) bifid distal region of the aedeagus, (b) saccus relatively short compared with other euptychiines; (c) lamella antevaginalis well developed (but somewhat variable,may be less developed in some species); (d) spiracle on the lateral plate of the 8th abdominal segment in females. Most of the species of the clade occur in Central and South America, except the Nearctic Megisto cymela and Oriental Megisto opalina . Several species of the clade (e.g. Moneuptychia , Stegosatyrus and Yphthimoides ) are typical of montane natural grasslands in southern Brazil with remarkable examples including Stegosatyrus hemiclara in high elevation grasslands in the Bolivian and Peruvian Andes ( Zacca et al., 2013) and Graphita griphe (Nakahara, Barbosa, et al., 2016) in the Andean cloud forests in Venezuela, Colombia, Peru and Ecuador. Adults vary from small to medium size and most species are characterized by their dull brown coloration on the dorsal and ventral wings, except for the remarkable marbled ventral pattern of some species of Carminda . Adults of both sexes are commonly attracted to traps baited with rotting fruit.Some species of Moneuptychia exhibit territorial behaviour and two or more males may make a clicking noise when they fly together, produced by the callus on the forewing subcostal vein ( Freitas et al., 2015; Freitas, Rosa, et al., 2018; Murillo-Hiller, 2006). The larvae feed on various genera of Poaceae and Cyperaceae ( Ackery, 1988; Beccaloni et al., 2008; DeVries, 1987; Freitas et al., 2015; Freitas, Barbosa, & Carreira, 2021; Freitas, Rosa, et al., 2018; Freitas, Rosa, & Kaminski, 2021; Janzen & Hallwachs,2022; Piovesan et al., 2022; Singer et al., 1983), and the last instars of Moneuptychia , Pharneuptychia , Yphthimoides , Carminda , and Cissia show a common body pattern with thin longitudinal zigzag stripes ( Barbosa et al., 2018; Freitas et al., 2015; Freitas, 2007; Freitas, Barbosa, & Carreira, 2021; Freitas, Kaminski, et al., 2012; Freitas,Rosa,et al., 2018; Freitas pers.obs.).

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.

FIGURE 4 Phylogeny of the Megisto 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.