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.

The evolution and genetics of sexually dimorphic 'dual' mimicry in the butterfly Elymnias hypermnestra.

Sexual dimorphism is a major component of morphological variation across the tree of life, but the mechanisms underlying phenotypic differences between sexes of a single species are poorly understood. We examined the population genomics and biogeography of the common palmfly Elymnias hypermnestra, a dual mimic in which female wing colour patterns are either dark brown (melanic) or bright orange, mimicking toxic Euploea and Danaus species, respectively. As males always have a melanic wing colour pattern, this makes E. hypermnestra a fascinating model organism in which populations vary in sexual dimorphism. Population structure analysis revealed that there were three genetically distinct E. hypermnestra populations, which we further validated by creating a phylogenomic species tree and inferring historical barriers to gene flow. This species tree demonstrated that multiple lineages with orange females do not form a monophyletic group, and the same is true of clades with melanic females. We identified two single nucleotide polymorphisms (SNPs) near the colour patterning gene WntA that were significantly associated with the female colour pattern polymorphism, suggesting that this gene affects sexual dimorphism. Given WntA's role in colour patterning across Nymphalidae, E. hypermnestra females demonstrate the repeatability of the evolution of sexual dimorphism.

Use of species delimitation approaches to tackle the cryptic diversity of an assemblage of high Andean butterflies (Lepidoptera: Papilionoidea).

Cryptic biological diversity has generated ambiguity in taxonomic and evolutionary studies. Single-locus methods and other approaches for species delimitation are useful for addressing this challenge, enabling the practical processing of large numbers of samples for identification and inventory purposes. This study analyzed an assemblage of high Andean butterflies using DNA barcoding and compared the identifications based on the current morphological taxonomy with three methods of species delimitation (automatic barcode gap discovery, generalized mixed Yule coalescent model, and Poisson tree processes). Sixteen potential cryptic species were recognized using these three methods, representing a net richness increase of 11.3% in the assemblage. A well-studied taxon of the genus Vanessa, which has a wide geographical distribution, appeared with the potential cryptic species that had a higher genetic differentiation at the local level than at the continental level. The analyses were useful for identifying the potential cryptic species in Pedaliodes and Forsterinaria complexes, which also show differentiation along altitudinal and latitudinal gradients. This genetic assessment of an entire assemblage of high Andean butterflies (Papilionoidea) provides baseline information for future research in a region characterized by high rates of endemism and population isolation.

DNA barcodes and citizen science provoke a diversity reappraisal for the "ring" butterflies of Peninsular Malaysia (Ypthima: Satyrinae: Nymphalidae: Lepidoptera).

The "rings" belonging to the genus Ypthima are amongst the most common butterflies in Peninsular Malaysia. However, the species can be difficult to tell apart, with keys relying on minor and often non-discrete ring characters found on the hindwing. Seven species have been reported from Peninsular Malaysia, but this is thought to be an underestimate of diversity. DNA barcodes of 165 individuals, and wing and genital morphology, were examined to reappraise species diversity of this genus in Peninsular Malaysia. DNA barcodes collected during citizen science projects-School Butterfly Project and Peninsular Malaysia Butterfly Count-recently conducted in Peninsular Malaysia were included. The new DNA barcodes formed six groups with different Barcode Index Numbers (BINs) representing four species reported in Peninsular Malaysia. When combined with public DNA barcodes from the Barcode Of Life Datasystems, several taxonomic issues arose. We consider the taxon Y. newboldi, formerly treated as a subspecies of Y. baldus, as a distinct species. DNA barcodes also supported an earlier suggestion that Y. nebulosa is a synonym under Y. horsfieldii humei. Two BINs of the genus Ypthima comprising DNA barcodes collected during citizen science projects did not correspond to any species previously reported in Peninsular Malaysia.

Diversification of the cold-adapted butterfly genus Oeneis related to Holarctic biogeography and climatic niche shifts.

Both geographical and ecological speciation interact during the evolution of a clade, but the relative contribution of these processes is rarely assessed for cold-dwelling biota. Here, we investigate the role of biogeography and the evolution of ecological traits on the diversification of the Holarctic arcto-alpine butterfly genus Oeneis (Lepidoptera: Satyrinae). We reconstructed the molecular phylogeny of the genus based on one mitochondrial (COI) and three nuclear (GAPDH, RpS5, wingless) genes. We inferred the biogeographical scenario and the ancestral state reconstructions of climatic and habitat requirements. Within the genus, we detected five main species groups corresponding to the taxonomic division and further paraphyletic position of Neominois (syn. n.). Next, we transferred O. aktashi from the hora to the polixenes species group on the bases of molecular relationships. We found that the genus originated in the dry grasslands of the mountains of Central Asia and dispersed over the Beringian Land Bridges to North America several times independently. Holarctic mountains, in particular the Asian Altai Mts. and Sayan Mts., host the oldest lineages and most of the species diversity. Arctic species are more recent, with Pliocene or Pleistocene origin. We detected a strong phylogenetic signal for the climatic niche, where one lineage diversified towards colder conditions. Altogether, our results indicate that both dispersal across geographical areas and occupation of distinct climatic niches promoted the diversification of the Oeneis genus.

Cruising the rain forest floor: butterfly wing shape evolution and gliding in ground effect.

Flight is a key innovation in the evolutionary success of insects and essential to dispersal, territoriality, courtship and oviposition. Wing shape influences flight performance and selection likely acts to maximize performance for conducting essential behaviours that in turn results in the evolution of wing shape. As wing shape also contributes to fitness, optimal shapes for particular flight behaviours can be assessed with aerodynamic predictions and placed in an ecomorphological context. Butterflies in the tribe Haeterini (Nymphalidae) are conspicuous members of understorey faunas in lowland Neotropical forests. Field observations indicate that the five genera in this clade differ in flight height and behaviour: four use gliding flight at the forest floor level, and one utilizes flapping flight above the forest floor. Nonetheless, the association of ground level gliding flight behaviour and wing shape has never been investigated in this or any other butterfly group. We used landmark-based geometric morphometrics to test whether wing shapes in Haeterini and their close relatives reflected observed flight behaviours. Four genera of Haeterini and some distantly related Satyrinae showed significant correspondence between wing shape and theoretical expectations in performance trade-offs that we attribute to selection for gliding in ground effect. Forewing shape differed between sexes for all taxa, and male wing shapes were aerodynamically more efficient for gliding flight than corresponding females. This suggests selection acts differentially on male and female wing shapes, reinforcing the idea that sex-specific flight behaviours contribute to the evolution of sexual dimorphism. Our study indicates that wing shapes in Haeterini butterflies evolved in response to habitat-specific flight behaviours, namely gliding in ground effect along the forest floor, resulting in ecomorphological partitions of taxa in morphospace. The convergent flight behaviour and wing morphology between tribes of Satyrinae suggest that the flight environment may offset phylogenetic constraints. Overall, this study provides a basis for exploring similar patterns of wing shape evolution in other taxa that glide in ground effect.

Complete mitochondrial genome of the Minois paupera Alphéraky, 1888 (nymphalidae: satyrinae) and its phylogenetic analysis.

The present study firstly reported a complete mitochondrial genome of Minois paupera (Alphéraky, 1888), a Satyrinae species endemic to China. This mitogenome is circular, 15,213 bp in length, and consists of 37 typical mitochondrial genes, including 13 protein-coding genes (PCGs), 22 tRNAs, and two rRNAs. The phylogenetic position was inferred using 31 previously published complete mitogenomes, and the results reveal that M. paupera is the most closely related to Minois dryas. The complete mitogenome of M. paupera provides useful genetic information for further research on the phylogeography and phylogeny of the genus Minois.

Chromosome-Level Assembly and Annotation of the Pearly Heath Coenonympha arcania Butterfly Genome.

We present the first chromosome-level genome assembly and annotation of the pearly heath Coenonympha arcania, generated with a PacBio HiFi sequencing approach and complemented with Hi-C data. We additionally compare synteny, gene, and repeat content between C. arcania and other Lepidopteran genomes. This reference genome will enable future population genomics studies with Coenonympha butterflies, a species-rich genus that encompasses some of the most highly endangered butterfly taxa in Europe.

Characterization of the complete mitochondrial genome of an endemic species in China, Aulocera merlina (Lepidoptera: Nymphalidae: Satyrinae) and phylogenetic analysis within Satyrinae.

The mitochondrial genome (mitogenome) has been extensively used as molecular markers in determining the insect phylogenetic relationships. In order to resolve the relationships among tribes and subtribes of Satyrinae at the mitochondrial genomic level, we obtained the complete mitogenome of Aulocera merlina (Oberthür, 1890) (Lepidoptera: Nymphalidae: Satyrinae) with a size of 15,259 bp. The mitogenome consisted of 37 typical genes, including 13 protein-coding genes (PCGs), 2 ribosomal RNA genes (rRNAs), 22 transfer RNA genes (tRNAs), and an A + T-rich region. The gene organization and arrangement were similar to those of all other known Satyrinae mitogenomes. All PCGs were initiated with the canonical codon pattern ATN, except for the cox1 gene, which used an atypical CGA codon. Nine PCGs used the complete stop codon TAA, while the remaining PCGs (cox1, cox2, nad4, and nad5) were terminated with a single T nucleotide. The canonical cloverleaf secondary structures were found in all tRNAs, except for trnS1 which lacked a dihydrouridine arm. The 448 bp A + T-rich region was located between rrnS and trnM, and it included the motif ATAGA followed by a 19-bp poly-T stretch and a microsatellite-like (TA)6 element preceded by the ATTTA motif. The phylogenetic tree, inferred using Bayesian inference and maximum likelihood methods, generated similar tree topologies, revealing well-supported monophyletic groups at the tribe level and recovering the relationship ((Satyrini + Melanitini) + ((Amathusiini + Elymniini) + Zetherini)). The close relationship between Satyrina and Melanargiina within the Satyrini was widely accepted. Additionally, Lethina, Parargina, and Mycalesina were closely related and collectively formed a sister group to Coenonymphina. Moreover, A. merlina was closely related to Oeneis buddha within the Satyrina. These findings will provide valuable information for future studies aiming to elucidate the phylogenetic relationships of Satyrinae.

Immature Stages, Natural History and Population Biology of Opoptera syme (Hübner, 1821), (Lepidoptera: Nymphalidae: Satyrinae) from Southeastern Brazil.

The present study describes the immature stages of Opoptera syme (Hübner, [1821]) using SEM and Micro-CT to generate 2D and 3D models to study the morphology and chaetotaxy, and present information on the natural history, behavior, and population biology. In laboratory, eggs were laid singly, and the isolated larvae passed through six to seven instars. Host plant in the study site is unknown, but larvae are known to feed on bamboos; in laboratory, larvae accepted the ornamental bamboo Bambusa textilis McClure, 1940. Adults are diurnal and univoltine, flying from December to April (the austral summer). The present study adds information to the biology and natural history of Brassolini, an iconic and still poorly known tribe of Neotropical butterflies.