Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae)

Abstract

The diverse, largely Neotropical subtribe Euptychiina is widely regarded as one of the most taxonomically challenging groups among all butterflies. Over the last two decades, morphological and molecular studies have revealed widespread paraphyly and polyphyly among genera, and a comprehensive, robust phylogenetic hypothesis is needed to build a firm generic classification to support ongoing taxonomic revisions at the species level. Here, we generated a dataset that includes sequences for up to nine nuclear genes and the mitochondrial COI ‘barcode’ for a total of 1280 specimens representing 449 described and undescribed species of Euptychiina and 39 out‐groups, resulting in the most complete phylogeny for the subtribe to date. In combination with a recently developed genomic backbone tree, this dataset resulted in a topology with strong support for most branches. We recognize eight major clades that each contain two or more genera, together containing all but seven Euptychiina genera. We provide a summary of the taxonomy, diversity and natural history of each clade, and discuss taxonomic changes implied by the phylogenetic results. We describe nine new genera to accommodate 38 described species: Lazulina Willmott, Nakahara & Espeland, gen.n., Saurona Huertas & Willmott, gen.n., Argentaria Huertas & Willmott, gen.n., Taguaiba Freitas, Zacca & Siewert, gen.n., Xenovena Marín & Nakahara, gen.n., Deltaya Willmott, Nakahara & Espeland, gen.n., Modica Zacca, Casagrande & Willmott, gen.n., Occulta Nakahara & Willmott, gen.n., and Trico Nakahara & Espeland, gen.n. We also synonymize Nubila Viloria, Andrade & Henao, 2019 (syn.n.) with Splendeuptychia Forster, 1964, Macrocissia Viloria, Le Crom & Andrade, 2019 (syn.n.) with Satyrotaygetis Forster, 1964, and Rudyphthimoides Viloria, 2022 (syn.n.) with Malaveria Viloria & Benmesbah, 2020. Overall, we revised the generic placement of 79 species (74 new generic combinations and five revised combinations), and as a result all but six described species of Euptychiina are accommodated within 70 named, monophyletic genera. For all newly described genera, we provide illustrations of representative species, drawings of wing venation and male and (where possible) female genitalia, and distribution maps, and summarize the natural history of the genus. For three new monotypic genera, Occulta gen.n., Trico gen.n. and Xenovena gen.n. we provide a taxonomic revision with a review of the taxonomy of each species and data from examined specimens. We provide a revised synonymic list for Euptychiina containing 460 valid described species, 53 subspecies and 255 synonyms, including several new synonyms and reinstated species.

Description of a new genus and species of Euptychiina (Lepidoptera: Nymphalidae) from the Brazilian Amazon rainforest

“…Although the precise phylogenetic relationships of Xikrin ueharapradoi with other lineages in the 'Archeuptychia clade' sensu Espeland et al (2019Espeland et al ( , 2023 cannot be assessed confidently based on the currently available data, the molecular phylogeny shows Xikrin as an isolated lineage with considerable genetic divergence from other members in the clade (see Table 3). Thus, the best decision at this time was to describe it as a monotypic genus, taking into account the low/moderate support for Xikrin as sister to any of the other lineages in the clade and the lack of strong morphological characters associating it with other genera in the clade, though the general features of the male genitalia are similar to its relatives.…”

“…The ML tree recovered Xikrin gen. nov. as part of the 'Archeuptychia clade' sensu Espeland et al (2019Espeland et al ( , 2023 along with the genera Archeuptychia Forster, 1964, Pseudeupytychia Forster, 1964, Cisandina and Stephenympha with strong support (SHaRLT = 99.7; aBayes = 1; UFBoot = 100) (Figure 1). In the clade, Xikrin gen. nov. appears as sister to Stephenympha + Cisandina, although with low/moderate support (SHaRLT = 66.8; aBayes = 0.827; UFBoot = 65), implying that its relationship with other genera in the clade could change in the future with the addition of more data.…”

“…The subtribe Euptychiina includes 460 valid species (Espeland et al 2023;Zacca et al 2018) distributed mainly in the Neotropical region, with few Nearctic species and a single species in Asia (Espeland et al 2019(Espeland et al , 2023Lamas 2004;Marín et al 2011Marín et al , 2017Peña et al 2010). This group used to be one of the least studied within Nymphalidae.…”

“…ufl.edu/museum-voices/euptychiina/ for a more detailed list of publications). However, although the phylogeny of Euptychina is now resolved to some extent (Espeland et al 2019(Espeland et al , 2023Marín et al 2017;Peña et al 2010), several groups are still largely unknown, without proper revision. Among these are some species-rich genera such as Taygetis Hübner, 1819 and Splendeuptychia Forster, 1964 (with 32 and 26 described species, respectivelybut see Espeland et al 2023).…”

“…However, although the phylogeny of Euptychina is now resolved to some extent (Espeland et al 2019(Espeland et al , 2023Marín et al 2017;Peña et al 2010), several groups are still largely unknown, without proper revision. Among these are some species-rich genera such as Taygetis Hübner, 1819 and Splendeuptychia Forster, 1964 (with 32 and 26 described species, respectivelybut see Espeland et al 2023). Moreover, the taxonomic diversity of Euptychiina is still growing: Many new genera and species are still being described, not only from lowland tropical forests (where the group is more diverse), but also from montane habitats and open vegetation (e.g., Barbosa et al 2020;Freitas et al 2018Freitas et al , 2019Nakahara et al 2019Nakahara et al , 2021Nakahara et al , 2022Rosa et al 2021).…”