Euptychia, Hubner, 1818

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 : 7-54

Euptychia

Euptychia is strongly supported as a sister to the remainder of the subtribe ( Figures 2, S2 and S 3 ) as also found in Espeland et al. (2019a). The systematic placement of Euptychia has been unstable in previous phylogenetic studies ( Murray & Prowell, 2005; Peña et al., 2010, 2011). Currently, 37 valid species are recognized ( Fratello et al., 2015; Freitas, Wahlberg, et al., 2012; Nakahara et al., 2014, 2017; Nakahara, Hall, et al., 2015; Nakahara, Janzen, et al., 2015; Nakahara, Llorente-Bousquets, et al., 2015; Nakahara, Vega, & Willmott, 2016; Neild et al., 2014, 2015). Only a subset of species was included here, and several taxonomic changes, in addition to the description of over a dozen species, will be made in a forthcoming revision of the genus, and thus the genus is estimated to include more than 50 species (Nakahara et al., in preparation). Members of Euptychia are relatively small for the subtribe, with the forewing length often less than 20 mm. Except for a few species, they have sexually monomorphic wing patterns often marked prominently with ventral bands and submarginal eyespots and many species exhibit translucence. Possible synapomorphies and/or distinctive characters for Euptychia include the following: (a) Third segment of labial palpi shorter than one-fourth of second segment in length; (b) absence of tibial spurs in midleg and hindleg; (c) absence of lateral spines on tibia; (d) presence of the forewing recurrent vein in the discal cell; (e) absence of basal swelling of the forewing cubital vein; (f) humeral vein relatively reduced in comparison with other Euptychiina ; (g) developed male eighth tergite and sternite; (h) presence of the sclerotized region of the eighth abdominal segment in the female, located at the very basal side of the eighth abdominal segment; (i) absence of the lateral sclerotization of the 8th abdominal segment of the female; (j) origin of the ductus seminalis at the posterior end of the ductus bursae In fact, many of these characters are unique to Euptychia (Nakahara, unpublished data), highlighting the distinctiveness of this lineage within the subtribe. Members of Euptychia range from north-western Mexico to south-eastern Brazil, and somewhat unusually for the subtribe, the genus appears to be more diverse in lower montane regions and adjacent lowlands than in lowland forest far from mountains. Adult males of some species exhibit perching behaviour, often with several individuals in close proximity, mostly in light gaps or sunflecks and often on hilltops. Although most known host plants for Euptychiina are either bamboo or other grasses in the family Poaceae , larvae of Euptychia notably feed instead on some species of Selaginella (Lycopsida) , as well as a single record utilizing Neckeropsis (Neckeraceae) ( Brévignon, 2008; Freitas, Mota, Barbosa, & Carreira, 2019; Hamm & Fordyce, 2016; Singer et al., 1971; Singer & Mallet, 1986).

Euptychia Hübner, 1818

= Caenoptychia Le Cerf, 1919

= Ristia Gagarin, 1936

alacristata Neild, Nakahara & Fratello, 2014 [ Neild et al. (2014, Tropical Lepidoptera Research, 24(1): 4-9)]

aquila Fratello, Nakahara & Brévignon, 2015 [ Fratello et al. (2015, Journal of the Lepidopterists' Society, 69(4): 293-306)]

atlantica Nakahara & Freitas, 2017 [ Nakahara et al. (2017, Neotropical Entomology, 46: 302-309)]

attenboroughi Neild, Nakahara, Fratello & Le Crom, 2015 [ Neild et al. (2015, ZooKeys, 541: 87-108)]

audacia Brévignon, Fratello & Nakahara, 2015 [ Fratello et al. (2015, Journal of the Lepidopterists' Society, 69(4): 293-306)]

boulleti (Le Cerf, 1919) [ Freitas et al. (2012, Neotropical Entomology, 41(6): 461-467)]

= virgata Joicey & Talbot, 1924 [ Freitas et al. (2012, Neotropical Entomology, 41(6): 461-467)]

= tigrina (Gagarin, 1936) [ Freitas et al. (2012, Neotropical Entomology, 41(6): 461-467)]

cesarense Pulido, Andrade, Peña & Lamas, 2011 [Pulido et al. (2011, Zootaxa, 2906: 43-51)]

- obtusa Nakahara, 2015 [ Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)]

- viloriai Andrade, Pulido, Peña & Lamas, 2011 [Pulido et al. (2011, Zootaxa, 2906: 43-51)]

efraini Ríos, 2019 [Ríos (2019, Anartia, 29: 49-53)]

= similis Henao, 2019 , preocc. (not Butler, 1867) [Ríos (2019, Anartia, 29: 49-53)]

enyita Nakahara, Lamas & Willmott, 2015 [ Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)]

enyo Butler, 1867

favonius Nakahara, Vega & Willmott, 2016 [ Nakahara et al. (2016, Zootaxa, 4184: 358-366)]

fernandae Nakahara & Willmott, 2015 [ Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)]

fetna Butler, 1870

granatina Nakahara, Le Crom & Hall, 2015 [ Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)]

hannemanni Forster, 1964

insolata Butler & Druce, 1872

= macrophthalma Staudinger, 1876

jesia Butler, 1869

juanjoi Le Crom, Nakahara & Lamas, 2015 [ Nakahara et al. (2015, Journal of Research on the Lepidoptera, 48: 59-63)]

lacandona Warren & Nakahara, 2015 [ Nakahara et al. (2015, Journal of Research on the Lepidoptera, 48: 51-57)]

marceli Brévignon, 2005 [ Brévignon (2005, Lambillionea, 105(3)(1): 393-404)]

- divisa Benmesbah, Costa, Attal & Viloria, 2021 [Costa et al. (2021, Antenor, 8(1): 2-28)]

meta Weymer, 1911

mollina (Hübner, [1818])

= mollina (Hübner, 1808) , nom. nud.

- suzannae Brévignon, 2005 [ Brévignon (2005, Lambillionea, 105(3)(1): 393-404)]

mollis Staudinger, 1876

neblina Warren & Nakahara, 2015 [ Nakahara et al. (2015, Journal of Research on the Lepidoptera, 48: 51-57)]

neildi Brévignon, 2005 [ Brévignon (2005, Lambillionea, 105(3)(1): 393-404)]

padroni Nakahara, Lamas & Willmott, 2015 [ Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)]

pegasus Nakahara & J. Hall, 2015 [ Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)]

picea Butler, 1867

pillaca Nakahara & Willmott, 2015 [ Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)]

roraima Nakahara, Fratello & Harvey, 2014 [ Nakahara et al. (2014, Zootaxa, 3881(3): 291-300)]

rubrofasciata L.D. Miller & J.Y. Miller, 1988

rufocincta Weymer, 1911

sarah Benmesbah & Viloria, 2019 [Costa et al. (2019, Anartia, 29: 20-48)]

sophiae Zacca, Nakahara, Dolibaina & Dias, 2015 [ Neild et al. (2015, ZooKeys, 541: 87-108)]

truncata Nakahara & J. Hall, 2015 [ Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)]

westwoodi Butler, 1867

- muli Brévignon, 2005 [ Brévignon (2005, Lambillionea, 105(3)(1): 393-404)]

woroina Viloria & Benmesbah, 2020 [Costa et al. (2020, Antenor, 7(1): 19-41)]

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.