Phylogenetic systematics of Yphthimoides Forster, 1964 and related taxa, with notes on the biogeographical history of Yphthimoides species.

Species losses are increasing and may have an impact on our understanding of patterns of evolutionary pathways and phylogenetic relationships among the groups being lost. The knowledge of such patterns can contribute to preventing future losses by identifying which lineages have higher or lower diversification rates, thus informing conservation strategies. Recent years have seen a significant growth in studies of butterfly systematics, allowing a better understanding of evolutionary relationships among most groups and revealing significant taxonomic chaos in several groups. One of the latter groups is the nymphalid subtribe Euptychiina (Satyrinae), which has been shown to include a number of non-monophyletic genera based on recent molecular phylogenetic analyses. Among others, these genera include Yphthimoides, which is widespread throughout the Neotropical region but particularly diverse in the southeastern Neotropics, and a pair of related genera, Pharneuptychia Forster, 1964 and Moneuptychia Forster, 1964. Using molecular data, this study scope and aims were to provide a phylogenetic hypothesis that corroborates Yphthimoides as presently conceived being non-monophyletic, a result reinforced by a comparative study of the male genitalic morphology. Our results also show that Pharneuptychia and Moneuptychia, plus a species misplaced elsewhere in the Euptychiina, Euptychoides castrensis (Schaus, 1902), form a well supported clade, and that the latter 'species' is a complex of cryptic species. We therefore propose a number of taxonomic rearrangements in the present work to resolve these issues: Yphthimoides eriphule (A. Butler, 1867) will be moved to a new genus; Y. affinis (A. Butler, 1867), Y. maepius (Godart, [1824]), Y. mimula (Hayward, 1954), Y. neomaenas (Hayward, 1967) and Y. mythra (Weymer, 1911) are being transferred to Malaveria Viloria & Benmesbah, 2021; Pharneuptychia innocentia (Godart, [1824]) will be moved to another genus to be described; and Euptychoides castrensis, Pharneuptychia romanina (Bryk, 1953) and Yphthimoides viviana (Romieux, 1927) are being moved to Moneuptychia. The dating of divergences points to a split between the ancestral lineage of Yphthimoides and its sister group, Carminda Ebert and Dias, inDias 1998, during the last half of the Miocene, around 11.86 Mya, and to the diversification of the Pharneuptychia during the same time 11.35 (±3.52) Mya. Biogeographic analysis showed that the most recent common ancestor of Yphthimoides started to diversify either in the the Brazilian Cerrado savannas or in a combined area of Cerrado and South Atlantic Forest, with a possible change in the ancestral habitat of Carminda. Furthermore, ancestral character mapping favors a savanna origin hypothesis over a forest origin hypothesis.

Four hundred shades of brown: Higher level phylogeny of the problematic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae) based on hybrid enrichment data.

Relationships within satyrine butterflies have been notoriously difficult to resolve using both morphology and Sanger sequencing methods, and this is particularly true for the mainly Neotropical subtribe Euptychiina, which contains about 400 described species. Known larvae of Euptychiina feed on grasses and sedges, with the exception of the genus Euptychia, which feed on mosses and lycopsids, and the butterflies occur widely in rainforest, cloudforest and grassland habitats, where they are often abundant. Several previous molecular and morphological studies have made significant progress in tackling the systematics of the group, but many relationships remain unresolved, with long-branch-attraction artifacts being a major problem. Additionally, the monophyly of the clade remains uncertain, with Euptychia possibly not being closely related to the remainder of the clade. Here we present a backbone phylogeny of the subtribe based on 106 taxa, 368 nuclear loci, and over 180,000 bps obtained through hybrid enrichment. Using both concatenation and species tree approaches (IQ-TREE, EXABAYES, ASTRAL), we can for the first time strongly confirm the monophyly of Euptychiina with Euptychia being the sister group to the remainder of the clade. The Euptychiina is divided into nine well supported clades, but the placement of a few genera such as Hermeuptychia, Pindis and the Chloreuptychia catharina group still remain uncertain. As partially indicated in previous studies, the genera Cissia, Chloreuptychia, Magneuptychia, Megisto, Splendeuptychia and Euptychoides, among others, were found to be highly polyphyletic and revisions are in preparation. The phylogeny will provide a strong backbone for the analysis of datasets in development that are much more taxonomically comprehensive but have orders of magnitude fewer loci. This study therefore represents a critical step towards resolving the higher classification and studying the evolution of this highly diverse lineage.

Four new species of Moneuptychia (Lepidoptera: Satyrinae: Euptychiina) from Brazil.

This paper describes four new species of Moneuptychia as follows: M. montana Freitas, M. vitellina Freitas & Barbosa, M. pervagata Freitas, Siewert & Mielke and M. wahlbergi Freitas, Barbosa, Siewert & Mielke from south and southeastern Brazil. Details are presented on the morphology of adults of all species, and immature stages for two species, and we discuss the taxonomy and identification of the genus Moneuptychia. The mitochondrial CoxI "barcode" region was used for exploring the utility of this DNA marker to identify these species, giving strong support for all new species.

Reassessment of the type locality of Euptychiastigmatica Godman, 1905, with the description of two new sibling species from Amazonia (Lepidoptera, Nymphalidae, Satyrinae, Satyrini).

A brief historical review regarding the type locality of Euptychiastigmatica Godman, 1905 was conducted, which suggests that its type locality is actually Rio de Janeiro, Brazil, rather than northeastern Argentina, as previously purported. Consequently, E.stigmatica and its senior synonym E.cyanites Butler, 1871, are regarded to be two species-group names representing a taxon in the euptychiine genus Caeruleuptychia Forster, 1964 known from the Brazilian Atlantic Forest. A lectotype is designated for E.cyanites. Additionally, two closely related species are named and described using an integrative approach with morphological and molecular evidence. Caeruleuptychiaharrisi Nakahara & Freitas, sp. nov. and C.aemulatio Nakahara & Willmott, sp. nov. both occur in Amazonia and COI barcode data recovered these taxa as part of the caerulea clade of Caeruleuptychia.

Morphology and behavior of the early stages of the skipper, Urbanus esmeraldus, on Urera baccifera, an ant-visited host plant.

The Neotropical genus Urbanus (Hübner) (Lepidoptera: Hesperiidae) contains around 34 described species, and is widely distributed from the extreme southern United States to Argentina. Here, we describe the larval morphology and behavior of Urbanus esmeraldus (Hübner)(Hesperiidae) in Urera baccifera (Urticaceae), a plant producing food rewards and fleshy fruits that attract ants (including predacious species) in a Brazilian forest. Larvae pass through five instars and construct two kinds of leaf shelters. Experiments with ejected fecal pellets showed that these can serve as cues to ground-dwelling ants that climb onto host plants and potentially attack the larvae. Manipulation with pellets placed at different distances suggests that ejection behavior decreases larval vulnerability to ant predation. Larval preference for mature leaves may be related with increased predation risk at ant-visited young leaves. The study shows that a combination of natural history and experimental data can help understand the life history of a butterfly using a plant with high predation risk.

Two new species of Hermeuptychia from North America and three neotype designations (Nymphalidae: Satyrinae).

Two new species of Hermeuptychia Forster, 1964 are described. Hermeuptychia sinuosa Grishin, sp. n. (type locality Guatemala: El Progreso, Morazán) is an isolated member of the genus that does not readily fit into known species groups, as suggested by its distinct male and female genitalia and COI DNA barcode sequences. It is distinguished from its congeners by prominently wavy submarginal lines, rounder wings and distinctive genitalia, and can typically be identified by a white dot, instead of an eyespot, near the ventral hindwing apex. Hermeuptychia occidentalis Grishin, sp. n. (type locality Mexico: Guerrero, Acapulco) belongs to the Hermeuptychia sosybius group as indicated by the presence of androconia on the dorsal surface of the wings, genitalia and COI DNA barcodes, and in addition to DNA characters, differs from its relatives in the shape of the uncus and female genitalia. Neotypes of Oreas strigata canthe Hübner, [1811] (type locality Suriname: Gelderland, Suriname River), Megisto acmenis Hübner, 1823 (type locality Argentina: Buenos Aires), and Satyrus cantheus Godart, [1824] (type locality USA: Florida, Pinellas Co., St. Petersburg) and lectotype of Euptychia celmis var. bonaërensis [sic] Burmeister, 1878 (type locality Argentina: Buenos Aires) are designated. These designations establish Hermeuptychia canthe as a valid species widely distributed in South America from Colombia to Bolivia and Southeast Brazil, Euptychia celmis var. bonaërensis [sic] Burmeister, 1878 as a junior objective synonym of Yphthimoides acmenis, and S. cantheus as a junior subjective synonym of Hermeuptychia sosybius (Fabricius, 1793). Papilio camerta Cramer, 1780 is treated as nomen dubium requiring further studies to determine an identity that is consistent with the original description, as it may be conspecific with Paryphthimoides poltys (Prittwitz, 1865) instead of being a Hermeuptychia species as currently assumed.

Se describe dos nuevas especies de Hermeuptychia Forster, 1964. Hermeuptychia sinuosa Grishin, sp. n. (localidad tipo Guatemala: El Progreso, Morazán), es un componente aislado del género que no encaja fácilmente en los grupos de especies conocidas, como lo indica su distintiva genitalia masculina y femenina y las secuencias de ADN del código de barras COI. Se distingue de sus congéneres por tener líneas submarginales prominentemente onduladas, alas más redondas y genitales diferentes, y se puede identificar típicamente por un punto blanco, en lugar de una mancha ocular, cerca del ápice ventral del ala anterior. Hermeuptychia occidentalis Grishin, sp. n. (localidad tipo México: Guerrero, Acapulco) pertenece al grupo de Hermeuptychia sosybius como lo indica la presencia de androconia en las alas anteriores, la estructura genital y secuencias de ADN de la región del código de barras COI, y además de caracteres del ADN, se diferencia de sus parientes en la forma del uncus y la genitalia femenina. Se designa neotipos para Oreas strigata canthe Hübner, [1811] (localidad tipo Surinam: Gelderland, Río Surinam), Megisto acmenis Hübner, 1823 (localidad tipo Argentina: Buenos Aires), y Satyrus cantheus Godart, [1824] (localidad tipo Estados Unidos: Florida, Pinellas Co., St. Petersburg), y el lectotipo de Euptychia celmis var. bonaërensis [sic] Burmeister, 1878 (localidad tipo Argentina: Buenos Aires). Estas designaciones establecen a Hermeuptychia canthe como una especie válida ampliamente distribuida en América del Sur desde Colombia hasta Bolivia y el sureste de Brasil, a Euptychia celmis var. bonaërensis [sic] Burmeister, 1878 como sinónimo objetivo más reciente de Yphthimoides acmenis, y a S. cantheus como sinónimo subjetivo más reciente de Hermeuptychia sosybius (Fabricius, 1793). Papilio camerta Cramer, 1780 es tratado como un nomen dubium requiriendo más estudios para determinar una identidad que sea consistente con la descripción original, ya que puede ser coespecífica con Paryphthimoides poltys (Prittwitz, 1865) en lugar de ser una especie de Hermeuptychia como se asume actualmente.

Fifty years without a name: a new species of Splendeuptychia Forster (Lepidoptera: Nymphalidae: Satyrinae).

A new satyrine butterfly species, Splendeuptychia tupinamba Freitas, Huertas Rosa, sp. nov. (Nymphalidae: Satyrinae), is described. This species is found throughout a large geographical range in Brazil, Argentina and Paraguay, predominantly in the Cerrado domain, with some records in neighboring Amazonia and Atlantic Forest. Morphology and molecular data indicate that this species is part of a clade that includes Splendeuptychia ashna (the type species of the genus Splendeuptychia), and several species placed in the recently described genus Nubila Viloria, Andrade Henao, 2019.

A new species of butterfly (Lepidoptera: Nymphalidae) from the highlands of Southeastern Brazil.

A new species of the butterfly satyrine subtribe Euptychiina is described: Carminda surpresa sp. nov. This species is apparently restricted to high altitude wet grasslands in southeastern Brazil. The description is based on morphological features, mainly from the wings, and male and female genitalia, and molecular data were also obtained and are used to validate the new species. Information about the geographic distribution, habitat and immature stages morphology is also provided. The systematic position of C. surpresa sp. nov. is discussed based on a molecular analysis which includes all described species of Carminda in addition to other species of Satyrinae.

Redescription of Yphthimoides patricia (Hayward, 1957), with taxonomic notes on the names Euptychia saltuensis Hayward, 1962 and Yphthimoides manasses (C. Felder R. Felder, 1867) (Nymphalidae: Satyrinae).

Euptychia saltuensis Hayward, 1962, new synonym, currently regarded as a nomen dubium and possibly a junior subjective synonym of Yphthimoides manasses (C. Felder R. Felder, 1867), is here treated as a junior subjective synonym of Yphthimoides patricia (Hayward, 1957), based on morphological characters of the male genitalia and the DNA barcode. The taxonomic status of Y. patricia is re-examined, and a detailed redescription of the adult morphology, including the male genitalia, is presented. Information on the distribution, habitat and immature stages of Y. patricia is also provided. Yphthimoides patricia is clearly a distinct species from Y. manasses based on the analysis of DNA barcode sequences and the morphology of the male genitalia.

Description of a new and highly distinctive genus and species of Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) from the Brazilian southern Amazon

ABSTRACT A new genus and species of Euptychiina (Satyrinae), Cristalinaia vitoria Mota, Zacca & Freitas gen. et sp. nov., is described based on three specimens collected in the region of the Cristalino River, Alta Floresta, Mato Grosso, Brazil. This rare species is known only from this region, where it flies inside the dense bamboo patches typical of that area. The last instar larva and the pupa are described; the larva was observed feeding on mature leaves of the common bamboo Guadua aff. paniculata Munro.urn:lsid:zoobank.org:pub:D61EDE8D-CAE9-41C6-B24D-BB789873566E

Before it is too late: description of a new genus and species of butterfly from a highly threatened Brazilian biome

Abstract A new genus, Nhambikuara Freitas, Barbosa & Zacca gen. nov., and species, Nhambikuara cerradensis Freitas, Barbosa & Zacca sp. nov., of the highly diverse Neotropical butterfly subtribe Euptychiina are described. Nhambikuara cerradensis sp. nov. is the type species for the genus, and Euptychia mima Butler, 1867 is also transferred to the new genus, as Nhambikuara mima (Butler, 1867) comb. nov., from the genus Zischkaia Forster, 1964. The taxonomy, phylogenetic relationships, geographic distribution and natural history of species of the genus are also discussed.http://zoobank.org/urn:lsid:zoobank.org:pub:41AD7568-3490-4F63-A019-32D5592A8C44