Taygetis, Hubner, 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 : 14

Taygetis clade

The ‘ Taygetis clade’ is one of the most stable generic groups that has been recognized within the Euptychiina , identified first by Murray and Prowell (2005) and corroborated by increasingly comprehensive subsequent molecular and morphological studies ( Espeland et al., 2019a; Marín et al., 2017; Matos-Maraví et al., 2013; Peña et al., 2006, 2010, 2011). The clade is fully supported in all datasets ( Figures 7 , S 2 and S 3 ) and contains 10 genera, Forsterinaria (27 species), Harjesia (3 species), Orotaygetis (monotypic), Parataygetis (2 species), Posttaygetis (monotypic), Pseudodebis (13 species), Taguaiba gen.n. (5 species), Sepona (monotypic), Taygetina (8 species) and Taygetis (25 species). It is embedded within a larger clade also containing the ‘ Pareuptychia -clade’, ‘ Splendeuptychia -clade’ and ‘ Archeuptychia -clade’. Multiple recent papers have helped revise the generic classification and describe new species ( Matos-Maraví et al., 2013; Siewert et al., 2013 [covering species included here in Taguaiba gen.n.]; Freitas, Barbosa, Willmott, et al., 2016 [ Sepona ]; Nakahara, Willmott, et al., 2018 [ Orotaygetis ], Nakahara, Matos-Maraví, et al., 2019 [ Taygetina ], Nakahara, Matos-Maraví, Willmott, et al., 2021; Nakahara, Matos-Maraví, Schwartz, et al., 2021; Nakahara, Janzen, et al., 2022 [ Pseudodebis ]). The relationships found here are similar to previous studies, with Sepona being the sister group to the remainder of the clade in all trees, and the remaining genera falling into two well-supported subclades; the Forsterinaria subclade including Forsterinaria, Paratygetis , Posttaygetis and Harjesia , and the Taygetis subclade containing the remaining genera ( Matos-Maraví et al., 2013; Nakahara, Willmott, et al., 2018). All genera are monophyletic and well-supported in all analyses, except for in the best 4GENES tree, where Forsterinaria quantius is sister to Parataygetis instead of the remainder of Forsterinaria , which is the case in all other trees. It has long been known that a new genus is needed for the former ‘ Taygetis ’ ypthima and relatives in the Taygetis subclade ( Matos-Maraví et al., 2013; Nakahara, Matos-Maraví, et al., 2019; Nakahara, Willmott, et al., 2018; Peña et al., 2010; Siewert et al., 2013), and below we describe the new genus Taguaiba gen.n. for this species group. The ‘ Taygetis clade’ contains 86 described species and a number of currently known undescribed species, with the latter concentrated in particular in Forsterinaria , Pseudodebis and Taygetis . Taxonomic revisions are in progress for all three of the above-mentioned genera, in addition to Taygetina , while most of the remaining genera are monotypic or bitypic and are of less taxonomic concern. Members of the ‘ Taygetis -clade’ are relatively large, and indeed Taygetis contains the largest species in the subtribe. Perhaps as a result of their size and presumably greater conspicuousness to predators, adults of some species show remarkable intra-specific variation in their ventral wing patterns, which resemble dead leaves (e.g. Sepona , Taguaiba gen.n., Taygetis mermeria , Taygetis larua ). Males of many species have darker areas on the dorsal forewing and/or hindwing with variably distributed brush-like androconial scales that are widespread among euptychiines, and most species have a ventral surface with somewhat to very undulate dark discal, postdiscal and/or submarginal lines. Possible synapomorphies (not all unique) and distinctive characters for the clade include the following: (a) vesica lacking obvious sclerotized cornuti; (b) corpus bursae extending to second abdominal segment (ductus bursae and corpus bursae occupying almost entire abdomen), except in a few taxa, such as Posttaygetis penelea ; (c) lamella antevaginalis sclerotized; (d) spatula-like primary head capsule setae in the first instar larva. Furthermore, the female abdomen of members of this clade shows some unusual characters, such as notably weakly sclerotized tergites and sternites (barely more sclerotized than the intersegmental membrane) in genera such as Taygetis and Pseudodebis , and two weakly sclerotized patches on the dorsal surface of the inter-segmental membrane of the seventh and eighth abdominal segments that are filled with scales in Forsterinaria quantius . Members of the clade range from Mexico to Paraguay, Argentina and south-eastern Brazil, with a slight centre of diversity in the south-western Amazon (southern Peru and adjacent areas of Brazil) driven largely by Taygetis . Otherwise, the clade is notable for two genera that have diversified in areas of lower euptychiine diversity, namely Taguaiba gen.n. in the Brazilian Atlantic Forest domain ( Siewert et al., 2013), and Forsterinaria in montane regions, particularly the tropical Andes ( Peña & Lamas, 2005). Adults of some species appear to be crepuscular ( DeVries, 1987; Murray, 2001a, 2001b; Young, 1972, 1984) and both sexes are attracted to rotting fruit (e.g. Freitas, Carreira, Santos, & Barbosa, 2016; Uehara-Prado & Freitas, 2019; Young, 1972), with males of Forsterinaria in particular also strongly attracted to carrion (Willmott & Pyrcz, pers. obs.). The immature stages feed mostly on various genera of Poaceae , with infrequent records on Marantaceae and Cyperaceae ( Baine et al., 2019; Corahua-Espinoza et al., 2023; Freitas, 2017; Freitas, Carreira, Santos, & Barbosa, 2016; Hurtado et al., 2021; Janzen & Hallwachs, 2022; Murray, 2001a, 2001b). The close spatial association of adults of many species with stands of Guadua (lowlands) and Chusquea bamboos ( Poaceae : Bambusoideae: Bambuseae) suggests that these plants are also widely used as hostplants throughout the clade. Indeed, some evidence suggests that species in Posttaygetis , Pseudodebis and Taygetina have a narrow diet breadth, feeding exclusively on Guadua or Rhipidocladum ( Corahua-Espinoza et al., 2023; Janzen & Hallwachs, 2022; Murray, 2001a, 2001b).

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 7 Phylogeny of the ’Taygetis clade’ based on the FULL dataset. Interpretation of the support values can be found in the legend. The scale bars under the butterfly images equal 1 cm.