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Comparison of best ML trees across analyses. The best ML trees from (A) the concatenated dataset, (B) combined nuclear dataset and (C) degenerated dataset (degen_1). The values at nodes in (A) and (B) represent support from 1,000 bootstrap trees analyzed with TIGER partitions/gene partitions. In (C), the values at the nodes are the output from the analyses using TIGER partitions only and the nodes without any value have bootstrap support <20. The other degenerated datasets (nt_12, noLRall1 + nt2) have similar or more fluctuating rearrangements for the nodes with BS < 20.
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Despite multiple attempts to infer the higher-level phylogenetic relationships of skipper butterflies (Family Hesperiidae), uncertainties in the deep clade relationships persist. The most recent phylogenetic analysis included fewer than 30% of known genera and data from three gene markers. Here we reconstruct the higher-level relationships with a r...
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... The subfamily Pyrginae in Europe, Asia and Australia was further divided into three 'genus groups', namely Celaenorrhinus Group, Tagiades Group and Pyrgus Group, which are more or less equivalent to current tribes. A number of studies based on molecular data in recent years have revised Evans' classification system, using from several (Warren et al., 2008(Warren et al., , 2009Yuan et al., 2015) to dozens (Sahoo et al., 2016(Sahoo et al., , 2017 or even as many as hundreds of genes. Some members of the Celaenorrhinus Group or the tribe Celarnorrhinini were transferred from Pyrginae to other subfamilies: Euschemon to Euschemoninae (Atkins, 2005;Warren et al., 2009) and Lobocla to Eudaminae (Warren et al., 2009). ...
... Nevertheless, a robust phylogenetic framework remains crucial for understanding butterfly diversity, adaptations, and ecological roles. Numerous phylogenetic studies have focused on the superfamily Papilionoidea, encompassing butterflies as a whole, and the individual families (Warren et al., 2008(Warren et al., , 2009Wahlberg et al., 2009;Wahlberg et al., 2014;Espeland et al., 2018;Kawahara et al., 2019;Li et al., 2019;Sahoo et al., 2016Sahoo et al., , 2017Toussaint et al., 2018;Yuan et al., 2015b;Zhang et al., 2019). More specifically, considerable progress has been made in the resolution of taxonomic issues within the family Hesperiidae using molecular data (Warren et al., 2008;Sahoo et al., 2016Sahoo et al., , 2017Toussaint et al., 2018Toussaint et al., , 2021aLi et al., 2019;Zhang et al., 2019Zhang et al., , 2022a. ...
... Numerous phylogenetic studies have focused on the superfamily Papilionoidea, encompassing butterflies as a whole, and the individual families (Warren et al., 2008(Warren et al., , 2009Wahlberg et al., 2009;Wahlberg et al., 2014;Espeland et al., 2018;Kawahara et al., 2019;Li et al., 2019;Sahoo et al., 2016Sahoo et al., , 2017Toussaint et al., 2018;Yuan et al., 2015b;Zhang et al., 2019). More specifically, considerable progress has been made in the resolution of taxonomic issues within the family Hesperiidae using molecular data (Warren et al., 2008;Sahoo et al., 2016Sahoo et al., , 2017Toussaint et al., 2018Toussaint et al., , 2021aLi et al., 2019;Zhang et al., 2019Zhang et al., , 2022a. However, the systematic positions of many taxa, especially those from Asian and African lineages, remain uncertain. ...
... Based on both molecular and morphological evidence, Warren et al. (2008Warren et al. ( , 2009) revised the classical classification framework of the family Hesperiidae, which was adopted after Watson (1893) and Evans (1937Evans ( , 1949Evans ( , 1951Evans ( , 1952Evans ( , 1953Evans ( , 1955. Since then, several molecular works have focused on exploring deeper-level relationships among skipper butterflies (Sahoo et al., 2016(Sahoo et al., , 2017Toussaint et al., 2018;Cong et al., 2019;Li et al., 2019;Zhang et al., 2019). These studies have proposed numerous taxonomic changes and sparked lively debates surrounding the validity of generic names, significantly advancing our understanding of the family Hesperiidae Cong et al., 2019;Li et al., 2019;Zhang et al., 2019Zhang et al., , 2022a. ...
Despite considerable research efforts in recent years, the deeper phylogenetic relationships among skipperbutterflies (Hesperiidae) remain unresolved. This is primarily because of limited sampling, especially withinAsian and African lineages. In this study, we consolidated previous data and extensively sampled Asian andAfrican taxa to elucidate the phylogenetic relationships within Hesperiidae. The molecular dataset comprisedsequences from two mitochondrial and two nuclear gene regions from 563 species that represented 353 genera.Our analyses revealed seven subfamilies within Hesperiidae: Coeliadinae, Euschemoninae, Eudaminae, Pyrginae,Heteropterinae, Trapezitinae, and Hesperiinae. The systematics of most tribes and genera aligned with those ofprior studies. However, notable differences were observed in several tribes and genera. Overall, the position oftaxa assigned to incertae sedis in Hesperiinae is largely clarified in this study. Our results strongly support themonophyly of the tribe Tagiadini (Pyrginae), and the systematics of some genera are clarified with compre-hensive discussion. We recognize 15 tribes within the subfamily Hesperiinae. Of these, nine tribes are discussedin detail: Aeromachini, Astictopterini, Erionotini, Unkanini (new status), Ancistroidini, Ismini (confirmed status),Plastingini (new status), Gretnini (confirmed status), and Eetionini (confirmed status). We propose four subtribeswithin Astictopterini: Hypoleucina subtrib.n., Aclerosina, Cupithina, and Astictopterina. Furthermore, wedescribe a new genus (Hyarotoides gen.n.) and reinstate two genera (Zea reinst.stat. and Sepa reinst.stat.) asvalid. Additionally, we propose several new combinations: Zea mytheca comb.n., Sepa bononia comb.n. &reinst.stat., and Sepa umbrosa comb.n. This study, with extensive sampling of Asian and African taxa, greatlyenhances the understanding of the knowledge of the skipper tree of life.
... The correct delineation and identification of species is not always possible based on characters of external morphology alone, especially in such taxonomically challenging groups as Colias [23][24][25][26]. Over the last decades, the rapid development and implementation of new molecular techniques and approaches validated mitochondrial (e.g., DNA barcodes) and nuclear DNA markers as a useful and efficient tool for species identification, detecting and analyzing cryptic diversity, revealing interspecific and deep-level relationships and phylogenetic structure in taxonomically challenging groups of insects, solving longstanding taxonomic problems [23,[27][28][29][30][31][32][33][34][35][36][37][38][39][40][41]. Despite the broad and constantly growing usage of DNA-based techniques in butterfly taxonomy and molecular systematics, surprisingly few studies focused, so far, on the genus Colias [6,[42][43][44][45][46]. ...
The genus Colias Fabricius, 1807 includes numerous taxa and forms with uncertain status and taxonomic position. Among such taxa are Colias mongola Alphéraky, 1897 and Colias tamerlana Staudinger, 1897, interpreted in the literature either as conspecific forms, as subspecies of different but morphologically somewhat similar Colias species or as distinct species-level taxa. Based on mitochondrial and nuclear DNA markers, we reconstructed a phylogeographic pattern of the taxa in question. We recover and include in our analysis DNA barcodes of the century-old type specimens, the lectotype of C. tamerlana deposited in the Natural History Museum (Museum für Naturkunde), Berlin, Germany (ZMHU) and the paralectotype of C. tamerlana and the lectotype of C. mongola deposited in the Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia (ZISP). Our analysis grouped all specimens within four (HP_I–HP_IV) deeply divergent but geographically poorly structured clades which did not support nonconspecifity of C. mongola–C. tamerlana. We also show that all studied females of the widely distributed haplogroup HP_II were infected with a single Wolbachia strain belonging to the supergroup B, while the males of this haplogroup, as well as all other investigated specimens of both sexes, were not infected. Our data highlight the relevance of large-scale sampling dataset analysis and the need for testing for Wolbachia infection to avoid erroneous phylogenetic reconstructions and species misidentification.
... The seven subfamilies include Coeliadinae, Euschemoninae, Eudaminae, Pyrginae, Heteropterinae, Trapezitinae, and Hesperiinae. Nonetheless, the higher-level relationships among skipper butterflies have not yet been adequately resolved and there is still ambiguity regarding the relationships among the major clades within the subfamily Pyrginae (Sahoo et al. 2016). Furthermore, support for relationships among the monophyletic subfamilies Heteropterinae, Trapezitinae, and Hesperiinae was weak to moderate (Sahoo et al. 2016). ...
... Nonetheless, the higher-level relationships among skipper butterflies have not yet been adequately resolved and there is still ambiguity regarding the relationships among the major clades within the subfamily Pyrginae (Sahoo et al. 2016). Furthermore, support for relationships among the monophyletic subfamilies Heteropterinae, Trapezitinae, and Hesperiinae was weak to moderate (Sahoo et al. 2016). In the last two decades, there has been an enormous increase in the use of molecular data (DNA sequences) to assess genetic variation among various taxa. ...
... While in the same cluster, O. angulatum of Pyrginae with U. folus of Hesperiinae is resided with weak support (Fig. 2). This result indicated that Pyrginae is paraphyletic, which is consistent with the findings obtained by Sahoo et al. (2016). They worked with 270 hesperiid genera, and their studies showed that the Pyrginae subfamily displayed inconsistent topologies because it was paraphyletic. ...
In the present study, the COI genes of thirteen species were sequenced, and NCBI GenBank homology searches of these species revealed significant similarity across diverse countries of origin. The sequences of thirteen species of hesperiid butterflies were then submitted to NCBI's GenBank, and assigned accession number. The average nucleotide composition of 585 bp of these genes was analyzed, and the results showed the mean base compositions of the COI sequences were 40.01% T, 16.10% C, 30.21% A, and 13.66% G. The present investigation revealed a significant preference for Adenine+Thymine (A+T), with a prevalence of 70.22%. Afterward, the genetic divergence among thirteen species of hesperiid was analyzed using the Kimura 2-parameter of the MEGA10 program, and the results indicated that the interspecific nucleotide divergence among thirteen species of hesperiid ranged from 0.0532 to 0.1790%. Moreover, a phylogenetic tree was constructed using the Neighbor-Joining (NJ) method, and the result revealed three clusters within the Pyrginae, Hesperiinae, and Coeliadinae subfamilies under the family Hesperiidae. The present investigation revealed Pyrginae as paraphyletic, while Coeliadinae was shown to be the sister group to Hesperiinae. As a final point, the analyses of the data support the following relationships: [(P. dan+T. japetus) + (P. guttata+P. bada+P. agna+B. cinnara+I. semamora+I. salsala+T. colon+U. folus + (O. angulatum)) + (B. oedipodea+S. gremius)]. The findings may be applicable to future research in the disciplines of molecular taxonomy to identify hesperiid species and may reveal a higher level of phylogenetic relationship among the subfamilies of Hesperiidae. Biodivers. Conserv. Bioresour. Manag. 2023, 9(2): 21-28
... 2200 species (Peña and Wahlberg 2008) and grass skippers (Hesperiidae: Hesperiinae) with ca. 2000 species (Sahoo et al. 2016). These lineages have independently evolved the ability to feed on grasses, and phylogenetic comparative analyses have indicated that grass feeding was a key innovation that increased diversification rates in these groups (Sahoo et al. 2017). ...
Polyphagous butterflies, i.e., those that feed on multiple host plants, need to evolve adaptations against the defenses of many plants. Studies have focused on chemical defenses of host plants and counter-adaptations of their herbivores. Physical defenses, despite being the first line of defense of many plants, have received little attention. Grass feeding butterflies are among the most speciose adaptive radiations among insects. Grasses rely primarily on physical rather than chemical defenses. Therefore, it is interesting to understand how butterflies have adapted to grass feeding. Mandibles are the principal biting and chewing organs in insect larvae, and mandible morphology should affect the ability of butterflies to effectively feed on their host plants. We here ask whether grass feeding butterflies have plasticity in mandibular morphology to cope with grass physical defenses, using the widespread grass feeding butterfly Mycalesis mineus as a model. We first show that physical defenses of grasses, such as leaf toughness, presence of silica deposits and trichomes, negatively affect body size of the butterfly. Our results further indicate that larvae of M. mineus show mandible plasticity, with increased mandibular length and relative mass investment, when feeding on tougher leaves and leaves with silica deposits. We suggest that the evolution of mandible plasticity may have been a key component of the coevolutionary arms race between grasses and herbivorous insects. This plasticity may have also allowed butterflies such as M. mineus to colonize a broad range of habitats and geographic regions.
... Over the last decades, the rapid development and implementation of new molecular techniques and approaches validated mitochondrial (e.g. DNA barcodes) and nuclear DNA markers as a useful and efficient tool for species identification, detecting and analyzing cryptic diversity, revealing interspecific and deep-level relationships and phylogenetic structure in taxonomically challenging groups of insects, solving long-standing taxonomic problems [23,[27][28][29][30][31][32][33][34][35][36][37][38][39]. Despite broad and constantly growing usage of DNA-based techniques in butterfly taxonomy and molecular systematics, surprisingly few studies focused so far on the genus Colias [6, [42][43][44][45][46]. ...
The genus Colias Fabricius, 1807 includes numerous taxa and forms with uncertain status and taxonomic position. Among such taxa are Colias mongola Alphéraky, 1897 and Colias tamerlana Staudinger, 1897, interpreted in the literature either as conspecific forms, as subspecies of different but morphologically somewhat similar Colias species, or as distinct species-level taxa. Based on mitochondrial and nuclear DNA markers, we reconstruct a phylogeographic pattern of the taxa in question. We recover and include in our analysis DNA barcodes of the century-old type specimens, the lectotype of C. tamerlana deposited in the Natural History Museum (Museum für Naturkunde), Berlin, Germany (ZMHU), the paralectotype of C. tamerlana and the lectoptype of C. mongola deposited in the Zoological Institute, Russian Academy of Sciences, Saint-Petersburg, Russia (ZISP). Our analysis grouped all specimens within four (HP_I-HP-IV) deeply divergent but geographically poorly structured clades which did not support nonconspecifity of C. mongola–C. tamerlana. We also show that all studied females of the widely distributed haplogroup HP_II were infected with a single Wolbachia strain belonging to the supergroup B, while the males of this haplogroup, as well as all other investigated specimens, were not infected. Our data underline the relevance of large-scale sampling dataset analysis and the need of testing for Wolbachia infection to avoid erroneous phylogenetic reconstructions and species misidentification.
... Based on the significantly different male genitalia, Shirôzu and Saigusa, 1962 separated several members from the genus Coladenia and described a new genus Pseudocoladenia with Coladenia dan fabia Evans, 1949 as its type species, with this classification being widely accepted by subsequent researchers (Chou, 1994;Huang and Xue, 2004;Hsu et al. 2007;Warren et al., 2008Warren et al., , 2009Chiba et al., 2009;Yuan et al., 2015;Sahoo et al., 2016;Wu and Hsu, 2017;Li et al., 2019;Zhang et al., 2019;Cong et al., 2019;Huang, 2021). A study by Li et al. (2019) claimed that Pseudocoladenia is a genus in the tribe Celaenorrhinini of the subfamily Tagiadinae. ...
... Whenever possible, we included samples from the type localities, and we used samples from different localities to identify the species/subspecies with a wide distribution. In total, 69 specimens were used as ingroup taxa, along with six species from the tribes Tagiadini and Celaenorrhinini as outgroup taxa, based on recent phylogenetic findings (Warren et al., 2009;Sahoo et al., 2016;Cong et al. 2019;Zhang et al., 2022). We sequenced most specimens used in this study. ...
The range of the butterfly genus Pseudocoladenia includes several biodiversity hotspots, such as the Himalayas, mountains of southwestern China, and Sundaland. However, the taxonomic status of some of its species/subspecies remain controversial, and no previous phylogenetic or biogeographic analyses have been conducted. Herein, we determined the systematic relationships and biogeographic history of this genus by reconstructing its phylogeny based on six genes and 69 specimens as representatives of all known species/subspecies. Two species delimitation methods (Bayes Poisson Tree Processes and Bayesian Phylogenetics and Phylogeography) were also employed to assess the status of each taxon. Based on these results and morphological evidence, we identified 12 species and three subspecies in the genus and subsequently classified these into three species groups: P. fatih, P. dea, and P. dan. Five taxa, P. sadakoe (Sonan and Mitono, 1936) stat. nov., P. celebica (Fruhstorfer, 1909) stat. nov., P. fulvescens (Elwes and Edwarda, 1897) stat. nov., P. eacus (Latreille, 1823) stat. nov., and P. fabia (Evans, 1949) stat. nov. were all recognized as independent species. Additionally, two taxa, P. eacus sumatrana (Fruhstorfer, 1909) comb. nov. and P. eacus dhyana (Fruhstorfer, 1909) comb. nov., were placed under P. eacus (Latreille, 1823) stat. nov. as subspecies. Another new species distributed in N. Yunnan, Pseudocoladenia yunnana Fan, Cao & Hou sp. nov., was discovered and described. Divergence time and ancestral range estimation indicated that the most recent common ancestor of Pseudocoladenia was distributed in the Himalayas-Hengduan Mountain region and Indochina and diverged approximately 14.00 Ma. Continuous and episodic dispersal, vicariance, and extinction were used to determine the current geographic distribution of the genus. The P. fatih group had a prominently disjunct distribution between the Himalaya-Hengduan Mountain and Taiwan. Meanwhile, the P. dan group was first derived in Indochina and subsequently dispersed into the southeastern Asian archipelagoes. This study provides a reference for the evolutionary route of transoceanic distributed species in Asia and elaborates on the causes of biodiversity.
... Within this family, seven major clades correspond to seven subfamilies: Coeliadinae, Euschemoninae, Eudaminae, Pyrginae, Heteropterinae, Trapezitinae, and Hesperiinae. Similarly to previous studies 4,7,9,16,17,29 , (1) Coeliadinae was branched out at the base of the family; (2) Euschemoninae was sister to the other Hesperiidae excluding Coeliadinae; and (3) Pyrginae sensu lato was a monophyletic group. Within the subfamily Pyrginae, tribes Tagiadini and Celaenorrhinini were sister groups. ...
Here, we present new molecular and morphological evidence that contributes towards clarifying the phylogenetic relations within the family Hesperiidae, and overcomes taxonomic problems regarding this family. First, nine new complete mitogenomes, comprising seven newly sequenced species and two samples of previously sequenced species collected from different localities, were obtained and assembled to analyze characteristics. The length of the mitogenomes ranges from 15,284 to 15,853 bp and encodes 13 protein-coding genes, two ribosomal RNA (rRNA) genes, 22 transfer RNA (tRNA) genes, and a control region. Two model-based methods (maximum likelihood and Bayesian inference) were used to infer the phylogenetic relationships. Based on the mitogenomic phylogenetic analyses and morphological evidence, we claim that the lineage that comprises two Asian genera, Apostictopterus Leech and Barca de Nicéville, should be a tribe Barcini stat. nov. of the subfamily Trapezitinae, Pseudocoladenia dea (Leech, 1894), P. festa (Evans, 1949), and Abraximorpha esta Evans, 1949 are considered distinct species. Finally, we suggest that Lotongus saralus chinensis Evans, 1932 should belong to the genus Acerbas de Nicéville, 1895, namely Acerbas saralus chinensis (Evans, 1932) comb. nov..
... A seminal paper about phylogeny of the family by Warren et al. (2008;2009) triggered intensive discussions about the subfamily and tribal boundaries. Later on, Sahoo et al. (2016) and Toussaint et al. (2018) carried out the anchored hybrid enrichment sequencing and obtained a robust support for most subfamilial and tribal level relationships . One year later, published a large-scale genomic study of skipper butterflies. ...
The butterfly tribe Tagiadini Mabille, 1878 is a large group of skippers. Although there are a few species which are limited in distribution to some countries in the Oriental or Afrotropical region, the main radiation of the tribe is in China. In the present study, we constructed the first detailed molecular phylogeny of the Chinese species in the tribe based on mitochondrial and nuclear genes, and investigated its history of larval hostplants association. Maximum likelihood phylogeny strongly supports for monophyly of Tagiadini from China and confirms the most recent accepted species in the tribe. The controversial position of some species are presented and discussed. The systematic relationship of larval hostplants and Tagiadini species match well in phylogeny tree. They show conservatism in evolution of larval hostplant use. Moreover, the divergence time estimates and ancestral-area reconstructions provide a detailed description about the historical biogeography of the Tagiadini from China. The Chinese species of the tribe very likely originated from a combined ancestral area between Himalaya-Hengduan Mountains and Southern China region in the late Ecocene and expanded to the other regions. A dispersal-vicariance analysis suggests that dispersal events have played essential roles in the promotion of diversification in this tribe and distribution of extant species, and geological and climatic changes have been important factors driving current distribution patterns.
... The pioneering work of Warren et al. (2008Warren et al. ( , 2009) put the higher classification of Hesperiidae on solid footing with revolutionary methodology to combine DNA sequences of several genes with traditionally scored morphological characters. A number of subsequent studies used many more genes in their analyses, ranging from dozens to hundreds (Sahoo et al. 2016(Sahoo et al. , 2017Toussaint et al. 2018). Then, genome-scale revisionary and phylogenetic work has followed Li et al. 2019;Zhang et al. 2019b,c;Toussaint et al. 2021a,b). ...
Our expanded efforts in genomic sequencing to cover additional skipper butterfly (Lepidoptera: Hesperiidae) species and populations, including primary type specimens, call for taxonomic changes to restore monophyly and correct misidentifications by moving taxa between genera and proposing new names. Reconciliation between phenotypic characters and genomic trees suggests three new tribes, two new subtribes, 23 new genera, 17 new subgenera and 10 new species that are proposed here: Psolosini Grishin, new tribe (type genus Psolos Staudinger, 1889), Ismini Grishin, new tribe (type genus Isma Distant, 1886), Eetionini Grishin, new tribe (type genus Eetion de Nicéville, 1895), Orphina Grishin, new subtribe (type genus Orphe Godman, 1901), Carystoidina Grishin, new subtribe (type genus Carystoides Godman, 1901), Fulvatis Grishin, new genus (type species Telegonus fulvius Plötz, 1882), Adina Grishin, new genus (type species Nascus adrastor Mabille and Boullet, 1912), Ornilius Grishin, new genus (type species Ornilius rotundus Grishin, new species), Tolius Grishin, new genus (type species Antigonus tolimus Plötz, 1884), Lennia Grishin, new genus (type species Leona lena Evans, 1937), Trida Grishin, new genus (type species Cyclopides barberae Trimen, 1873), Noxys Grishin, new genus (type species Oxynthes viricuculla Hayward, 1951), Gracilata Grishin, new genus (type species Enosis quadrinotata Mabille, 1889), Hermio Grishin, new genus (type species Falga ? hermione Schaus, 1913), Eutus Grishin, new genus (type species Cobalus rastaca Schaus, 1902), Gufa Grishin, new genus (type species Phlebodes gulala Schaus, 1902), Godmia Grishin, new genus (type species Euroto chlorocephala Godman, 1900), Rhomba Grishin, new genus (type species Eutychide gertschi Bell, 1937), Rectava Grishin, new genus (type species Megistias ignarus Bell, 1932), Contrastia Grishin, new genus (type species Hesperia distigma Plötz, 1882), Mit Grishin, new genus (type species Mnasitheus badius Bell, 1930), Picova Grishin, new genus (type species Vorates steinbachi Bell, 1930), Lattus Grishin, new genus (type species Eutocus arabupuana Bell, 1932), Gubrus Grishin, new genus (type species Vehilius lugubris Lindsey, 1925), Koria Grishin, new genus (type species Hesperia kora Hewitson, 1877), Corta Grishin, new genus (type species Eutychide lycortas Godman, 1900), Calvetta Grishin, new genus (type species Hesperia calvina Hewitson, 1866), Oz Grishin, new genus (type species Astictopterus ozias Hewitson, 1878), Praxa Grishin, new subgenus (type species Nascus prax Evans, 1952), Bron Grishin, new subgenus (type species Papilio broteas Cramer, 1780), Turis Grishin, new subgenus (type species Pyrgus (Scelothrix) veturius Plötz, 1884), Tiges Grishin, new subgenus (type species Antigonus liborius Plötz, 1884), Ocrypta Grishin, new subgenus (type species Notocrypta caerulea Evans, 1928), Tixe Grishin, new subgenus (type species Cobalus quadrata Herrich-Schäffer, 1869), Nycea Grishin, new subgenus (type species Pamphila hycsos Mabille, 1891), Nausia Grishin, new subgenus (type species Oenus [sic] nausiphanes Schaus, 1913), Flor Grishin, new subgenus (type species Stomyles florus Godman, 1900), Geia Grishin, new subgenus (type species Pamphila geisa Möschler, 1879), Rotundia Grishin, new subgenus (type species Enosis schausi Mielke and Casagrande, 2002), Volus Grishin, new subgenus (type species Eutocus volasus Godman, 1901), Pseudopapias Grishin, new subgenus (type species Papias tristissimus Schaus, 1902), Septia Grishin, new subgenus (type species Justinia septa Evans, 1955), Brasta Grishin, new subgenus (type species Lychnuchus brasta Evans, 1955), Bina Grishin, new subgenus (type species Cobalus gabina Godman, 1900), Balma Grishin, new subgenus (type species Carystoides balza Evans, 1955), Ornilius rotundus Grishin, new species (type locality in Brazil: Santa Catarina), Salantoia metallica Grishin, new species (type locality in Guyana: Acarai Mts.), Dyscophellus australis Grishin, new species (type locality in Paraguay: Sapucay), Dyscophellus basialbus Grishin, new species (type locality in Brazil: Rondônia), Telegonus subflavus Grishin, new species (type locality in Ecuador: Riobamba), Decinea colombiana Grishin, new species (type locality in Colombia: Bogota), Lerema lucius Grishin, new species (type locality in Panama: Colón), Cynea rope Grishin, new species (type locality in Nicaragua: Chontales), Lerodea sonex Grishin, new species (type locality in Peru: Cuzco), and Metiscus goth Grishin, new species (type locality in Costa Rica). Lectotypes are designated for the following 17 taxa: Telegonus gildo Mabille, 1888, Netrocoryne damias Plötz, 1882, Telegonus erythras Mabille, 1888, Telegonus galesus Mabille, 1888, Eudamus cretellus Herrich-Schäffer, 1869, Leucochitonea chaeremon Mabille, 1891, Antigonus aura Plötz, 1884, Pamphila voranus Mabille, 1891, Hesperia pupillus Plötz, 1882, Cobalus lumina Herrich-Schäffer, 1869, Cobalus stigmula Mabille, 1891, Megistias isus Godman, 1900, Cobalopsis latonia Schaus, 1913, Pamphila nubila Mabille, 1891, Metiscus atheas Godman, 1900, Mnasalcas amatala Schaus, 1902, and Hesperia ina Plötz, 1882. The lectotype of Hesperia infuscata Plötz, 1882 is invalid because it does not agree with the original description and illustration by Plötz, is not from the locality listed in the original description, and therefore is not a syntype. Neotypes are designated for the following five taxa: Telegonus corentinus Plötz, 1882, Hesperia dido Plötz, 1882, Hesperia distigma Plötz, 1882, Hesperia infuscata Plötz, 1882, and Hesperia pruinosa Plötz, 1882. As a result, the following five taxa are junior objective synonyms: Telegonus diophorus Möschler, 1883 of Telegonus corentinus Plötz, 1882, Pamphila puxillius Mabille, 1891 of Hesperia pupillus Plötz, 1882, Cobalus stigmula Mabille, 1891 of Hesperia distigma Plötz, 1882, Mnasalcas amatala Schaus, 1902 of Hesperia infuscata Plötz, 1882, and Hesperia pruinosa Plötz, 1882 of Hesperia uza Hewitson, 1877. Morys valerius valda Evans, 1955 is fixed as the type species of Morys Godman, 1900, and Pamphila compta Butler, 1877 is reaffirmed as the type species of Euroto Godman, 1900. Furthermore, the following taxonomic changes are suggested. Prosopalpus Holland, 1896, Lepella Evans, 1937, and Creteus de Nicéville, 1895 are placed in Aeromachini Tutt, 1906. Triskelionia Larsen and Congdon, 2011 is transferred from Celaenorrhinini Swinhoe, 1912 to Tagiadini Mabille, 1878. Kobelana Larsen and Collins, 2013 is transferred from Tagiadini Mabille, 1878 to Celaenorrhinini Swinhoe, 1912. The following nine genus-group names are resurrected from synonymy and treated as valid genera: Abaratha Moore, 1881 (not in Caprona Wallengren, 1857), Bibla Mabille, 1904 (not in Taractrocera Butler, 1870), Kerana Distant, 1886 and Tamela Swinhoe, 1913 (not in Ancistroides Butler, 1874), Metrocles Godman, 1900 (not in Metron Godman, 1900), Alerema Hayward, 1942 (not in Tigasis Godman, 1900), Metiscus Godman, 1900 (not in Enosis Mabille, 1889), Vistigma Hayward, 1939 (not in Phlebodes Hübner, [1819]), and Mnasalcas Godman, 1900 (not in Mnasitheus Godman, 1900). The genus-group names Daimio Murray, 1875 and Pterygospidea Wallengren, 1857 are resurrected from synonymy and treated as valid subgenera of Tagiades Hübner, [1819]. We confirm Apallaga Strand, 1911 as a valid genus. The following 24 genera are placed as subgenera, new status: Pseudonascus Austin, 2008 of Nascus Watson, 1893; Albiphasma Huang, Chiba, Wang and Fan, 2016 of Pintara Evans, 1932; Ctenoptilum de Nicéville, 1890 of Tapena Moore, [1881]; Odontoptilum de Nicéville, 1890 of Abaratha Moore, 1881; Caprona Wallengren, 1857 of Abantis Hopffer, 1855; Timochreon Godman and Salvin, 1896 of Zopyrion Godman and Salvin, 1896; Pulchroptera Hou, Fan and Chiba, 2021 of Heteropterus Duméril, 1806; Stimula de Nicéville, 1898 of Koruthaialos Watson, 1893; Udaspes Moore, [1881] and Notocrypta de Nicéville, 1889 of Ancistroides Butler, 1874; Cravera de Jong, 1983 of Xeniades Godman, 1900; Cobaloides Hayward, 1939 of Oligoria Scudder, 1872; Saniba O. Mielke and Casagrande, 2003 of Psoralis Mabille, 1904; Quinta Evans, 1955 of Cynea Evans, 1955; Styriodes Schaus, 1913 and Remella Hemming, 1939 of Mnasicles Godman, 1901; Repens Evans, 1955 of Eprius Godman, 1901; Morys Godman, 1900 of Lerema Scudder, 1872; Enosis Mabille, 1889 of Lychnuchus Hübner, [1831]; Penicula Evans, 1955 of Vistigma Hayward, 1939; Mnasinous Godman, 1900 of Methionopsis Godman, 1901; and Moeros Evans, 1955, Argon Evans, 1955, and Synale Mabille, 1904 of Carystus Hübner, [1819]. The following 20 genera are treated as junior subjective synonyms: Leucochitonea Wallengren, 1857 of Abantis Hopffer, 1855; Sapaea Plötz, 1879 and Netrobalane Mabille, 1903 of Caprona Wallengren, 1857; Parasovia Devyatkin, 1996 of Sebastonyma Watson, 1893; Pemara Eliot, 1978 of Oerane Elwes and Edwards, 1897; Ankola Evans, 1937 of Pardaleodes Butler, 1870; Arotis Mabille, 1904 of Mnaseas Godman, 1901; Chalcone Evans, 1955, Hansa Evans, 1955, and Propertius Evans, 1955 of Metrocles Godman, 1900; Jongiana O. Mielke and Casagrande, 2002 of Cobaloides Hayward, 1939; Pamba Evans, 1955 of Psoralis Mabille, 1904; Brownus Grishin, 2019 of Styriodes Schaus, 1913; Mnasilus Godman, 1900 of Papias Godman, 1900; Sucova Evans, 1955 of Mnasitheus Godman, 1900; Pyrrhocalles Mabille, 1904 and Asbolis Mabille, 1904 of Choranthus Scudder, 1872; Miltomiges Mabille, 1903 of Methionopsis Godman, 1901; Sacrator Evans, 1955 of Thracides Hübner, [1819]; and Lychnuchoides Godman, 1901 of Perichares Scudder, 1872. Arunena Swinhoe, 1919 is a junior subjective synonym of Stimula de Nicéville, 1898 (not of Koruthaialos Watson, 1893). The following 27 names are species-level taxa (some in new combinations) reinstated from synonymy: Salantoia gildo (Mabille, 1888) (not Salatis cebrenus (Cramer, 1777)), Bungalotis corentinus (Plötz, 1882) (not Bungalotis midas (Cramer, 1775)), Telegonus cretellus (Herrich-Schäffer, 1869) (not Telegonus cassander (Fabricius, 1793)), Santa palica (Mabille, 1888) (not Chiothion asychis (Stoll, 1780)), Camptopleura cincta Mabille and Boullet, 1917 (not Camptopleura auxo (Möschler, 1879)), Camptopleura orsus (Mabille, 1889) (not Nisoniades mimas (Cramer, 1775)), Metron voranus (Mabille, 1891) and Metron fasciata (Möschler, 1877) (not Metron zimra (Hewitson, 1877)), Limochores catahorma (Dyar, 1916) (not Limochores pupillus (Plötz, 1882)), Pares viridiceps (Mabille, 1889) (not Thoon modius (Mabille, 1889)), Tigasis wellingi (Freeman, 1969) (not Tigasis arita (Schaus, 1902)), Rectava sobrinus (Schaus, 1902) (not Papias phainis Godman, 1900), Nastra subsordida (Mabille, 1891) (not Adlerodea asema (Mabille, 1891), previously in Eutychide Godman, 1900), Lerema pattenii Scudder, 1872 (not Lerema accius (J. E. Smith, 1797)), Lerema (Morys) ancus (Möschler, 1879) (not Cymaenes tripunctus theogenis (Capronnier, 1874)), Cobalopsis zetus (Bell, 1942) (not Cobalopsis nero (Herrich-Schäffer, 1869)), Lerema (Geia) etelka (Schaus, 1902) (not Lerema (Geia) geisa (Möschler, 1879), previously in Morys Godman, 1900), Cymaenes isus (Godman, 1900) (not Cymaenes trebius (Mabille, 1891)), Vehilius labdacus (Godman, 1900) (not Vehilius inca (Scudder, 1872)), Papias amyrna (Mabille, 1891) (not Papias allubita (Butler, 1877), previously in Mnasilus Godman, 1900), Papias integra (Mabille, 1891) (not Papias subcostulata (Herrich-Schäffer, 1870)), Metiscus atheas Godman, 1900 (not Hesperia achelous Plötz, 1882), Dion agassus (Mabille, 1891) (not Dion uza (Hewitson, 1877), previously in Enosis Mabille, 1889), Picova incompta (Hayward, 1942) (not Lerema (Morys) micythus (Godman, 1900), previously in Morys Godman, 1900), Lucida melitaea (Draudt, 1923) (not Lucida lucia (Capronnier, 1874)), Methionopsis modestus Godman, 1901 (not Methionopsis ina (Plötz, 1882)), and Thargella (Volus) volasus (Godman, 1901) (not Eutocus facilis (Plötz, 1884)). The following 57 taxa are elevated from subspecies to species, new status (some in new combinations): Dyscophellus doriscus (Hewitson, 1867) (not Dyscophellus porcius (C. Felder and R. Felder, 1862), Phocides vida (A. Butler, 1872) (not Phocides urania (Westwood, 1852)), Tagiades (Daimio) ceylonica Evans, 1932 (not Tagiades litigiosa Möschler, 1878), Tagiades (Daimio) tubulus Fruhstorfer, 1910 (not Tagiades sambavana Elwes and Edwards, 1897), Tagiades (Daimio) kina Evans, 1934, Tagiades (Daimio) sheba Evans, 1934, Tagiades (Daimio) martinus Plötz, 1884, Tagiades (Daimio) sem Mabille, 1883, and Tagiades (Daimio) neira Plötz, 1885 (not Tagiades trebellius (Hopffer, 1874)), Tagiades (Daimio) korela Mabille, 1891 and Tagiades (Daimio) presbyter Butler, 1882 (not Tagiades nestus (C. Felder, 1860)), Tagiades obscurus Mabille, 1876, Tagiades ravi (Moore, [1866]), Tagiades atticus (Fabricius, 1793), Tagiades titus Plötz, 1884, Tagiades janetta Butler, 1870, Tagiades inconspicua Rothschild, 1915, and Tagiades hovia Swinhoe, 1904 (not Tagiades japetus (Stoll, [1781])), Tagiades silvia Evans, 1934 and Tagiades elegans Mabille, 1877 (not Tagiades gana (Moore, [1866])), Tapena bornea Evans, 1941 and Tapena minuscula Elwes and Edwards, 1897 (not Tapena thwaitesi Moore, [1881]), Darpa dealbata (Distant, 1886) (not Darpa pteria (Hewitson, 1868)), Perus manx (Evans, 1953) (not Perus minor (Schaus, 1902)), Canesia pallida (Röber, 1925) (not Carrhenes canescens (R. Felder, 1869)), Carrhenes conia Evans, 1953 (not Carrhenes fuscescens (Mabille, 1891)), Anisochoria extincta Hayward, 1933 and Anisochoria polysticta Mabille, 1876 (not Anisochoria pedaliodina (Butler, 1870)), Anisochoria verda Evans, 1953 (not Anisochoria minorella Mabille, 1898), Bralus alco (Evans, 1953) (not Bralus albida (Mabille, 1888)), Ephyriades jamaicensis (Möschler, 1879) (not Ephyriades brunnea (Herrich-Schäffer, 1865)), Koruthaialos (Stimula) frena Evans, 1949 (not Koruthaialos focula (Plötz, 1882)), Euphyes kiowah (Reakirt, 1866) (not Euphyes vestris (Boisduval, 1852)), Mnaseas inca Bell, 1930 (not Mnaseas bicolor (Mabille, 1889)), Metron hypochlora (Draudt, 1923) (not Metrocles schrottkyi (Giacomelli, 1911), previously in Metron Godman, 1900), Decinea huasteca (H. Freeman, 1969), Decinea denta Evans, 1955, and Decinea antus (Mabille, 1895) (not Decinea decinea (Hewitson, 1876)), Xeniades pteras Godman, 1900 (not Xeniades chalestra (Hewitson, 1866)), Xeniades difficilis Draudt, 1923 (not Xeniades orchamus (Cramer, 1777)), Xeniades hermoda (Hewitson, 1870) (not Tisias quadrata (Herrich- Schäffer, 1869)), Hermio vina (Evans, 1955) (not Hermio hermione (Schaus, 1913), previously in Lento Evans, 1955), Cymaenes loxa Evans, 1955, (not Cymaenes laureolus (Schaus, 1913)), Niconiades peri (Evans, 1955) (not Rhinthon bajula (Schaus, 1902), previously in Neoxeniades Hayward, 1938), Gallio danius (Bell, 1941) (not Vehilius seriatus (Mabille, 1891)), Gallio massarus (E. Bell, 1940) (not Gallio garima (Schaus, 1902) previously in Tigasis Godman, 1900), Cymaenes edata (Plötz, 1882), Cymaenes miqua (Dyar, 1913) and Cymaenes aequatoria (Hayward, 1940) (not Cymaenes odilia (Burmeister, 1878)), Lychnuchus (Enosis) demon (Evans, 1955) (not Lychnuchus (Enosis) immaculata (Hewitson, 1868), previously in Enosis Mabille, 1889), Naevolus naevus Evans, 1955 (not Naevolus orius (Mabille, 1883)), Lucida scopas (Mabille, 1891), Lucida oebasus (Godman, 1900), and Lucida leopardus (Weeks, 1901) (not Lucida lucia (Capronnier, 1874)), Corticea schwarzi (E. Bell, 1941) and Corticea sylva (Hayward, 1942) (not Corticea mendica (Mabille, 1898)), and Choranthus orientis (Skinner, 1920) (not Choranthus antiqua (Herrich-Schäffer, 1863), previously in Pyrrhocalles Mabille, 1904). Borbo impar bipunctata (Elwes and J. Edwards, 1897) is a valid subspecies, not a synonym of Borbo impar tetragraphus (Mabille, 1891), here placed in synonymy with Lotongus calathus (Hewitson, 1876), new synonym. We confirm the species status of Telegonus cassius (Evans, 1952) and Lerema (Morys) valda Evans, 1955. Euphyes chamuli Freeman, 1969 is placed as a subspecies of Euphyes kiowah (Reakirt, 1866), new status. The following 41 taxa are junior subjective synonyms, either newly proposed or transferred from synonymy with other species or subspecies: Telegonus mutius Plötz, 1882 of Euriphellus phraxanor (Hewitson, 1876), Telegonus erythras Mabille, 1888 of Dyscophellus damias (Plötz, 1882), Aethilla jaira Butler, 1870 of Telegonus cretellus (Herrich-Schäffer, 1869), Paches era Evans, 1953 of Santa palica (Mabille, 1888), Antigonus alburnea Plötz, 1884 of Tolius tolimus robigus (Plötz, 1884) (not of Echelatus sempiternus simplicior (Möschler, 1877)), Echelatus depenicillus Strand, 1921 of E. sempiternus simplicior (not of T. tolimus robigus), Antigonus aura Plötz, 1884 of Theagenes dichrous (Mabille, 1878) (not of Helias phalaenoides palpalis (Latreille, [1824])), Achlyodes impressus Mabille, 1889 of Camptopleura orsus (Mabille, 1889), Augiades tania Schaus, 1902 of Metron voranus (Mabille, 1891), Pamphila verdanta Weeks, 1906 of Metron fasciata (Möschler, 1877), Niconiades viridis vista Evans, 1955 of Niconiades derisor (Mabille, 1891), Pamphila binaria Mabille, 1891 of Conga chydaea (A. Butler, 1877) (not of Cynea cynea (Hewitson, 1876)), Psoralis concolor Nicolay, 1980 of Ralis immaculatus (Hayward, 1940), Hesperia dido Plötz, 1882 of Cynea (Quinta) cannae (Herrich-Schäffer, 1869) (not of Lerema lochius (Plötz, 1882)), Proteides osembo Möschler, 1883 of Cynea (Cynea) diluta (Herrich-Schäffer, 1869) (not of Cynea (Quinta) cannae (Herrich-Schäffer, 1869)), Cobalopsis brema E. Bell, 1959 of Eutus rastaca (Schaus, 1902), Psoralis panamensis Anderson and Nakamura, 2019 of Rhomba gertschi (Bell, 1937), Cobalus asella Herrich-Schäffer, 1869 of Amblyscirtes alternata (Grote and Robinson, 1867) (not of Amblyscirtes vialis (W. H. Edwards, 1862)), Papias trimacula Nicolay, 1973 of Nastra subsordida (Mabille, 1891), Pamphila bipunctata Mabille, 1889 and Sarega staurus Mabille, 1904 of Lerema pattenii Scudder, 1872 (not of Cymaenes lumina (Herrich-Schäffer, 1869), previously in Lerema Scudder, 1872), Hesperia aethra Plötz, 1886 of Lerema lineosa (Herrich-Schäffer, 1865) (not of Lerema (Morys) compta Butler, 1877), Megistias miaba Schaus, 1902 of Cobalopsis valerius (Möschler, 1879), Phanis sylvia Kaye, 1914 of Lerema etelka (Schaus, 1902) (not of Lerema (Geia) geisa (Möschler, 1879), previously in Morys Godman, 1900), Carystus odilia Burmeister, 1878, Pamphila trebius Mabille, 1891 and Megistias corescene Schaus, 1902 of Cymaenes lumina (Herrich-Schäffer, 1869), Hesperia phocylides Plötz, 1882 of Cymaenes edata (Plötz, 1882) (not of Lerema accius (J. E. Smith, 1797)), Pamphila xenos Mabille, 1898 of Vehilius inca (Scudder, 1872), Mnasilus guianae Lindsey, 1925 of Papias amyrna (Mabille, 1891), Pamphila nubila Mabille, 1891 of Papias integra (Mabille, 1891) (not of Cynea corisana (Plötz, 1882)), Enosis matheri H. Freeman, 1969 of Metiscus atheas Godman, 1900 (previously in Enosis Mabille, 1889), Hesperia infuscata Plötz, 1882 of Mnaseas derasa derasa (Herrich-Schäffer, 1870) (previously Arotis Mabille, 1904), (not of Papias subcostulata (Herrich-Schäffer, 1870)), Pamphila astur Mabille, 1891 of Metiscus angularis (Möschler, 1877) (not of Cymaenes tripunctus theogenis (Capronnier, 1874)), Anthoptus macalpinei H. Freeman, 1969 of Anthoptus inculta (Dyar, 1918), Methionopsis typhon Godman, 1901 of Methionopsis ina (Plötz, 1882), Methionopsis dolor Evans, 1955 of Thargella volasus (Godman, 1901), Hesperia cinica Plötz, 1882 of Dubiella dubius (Stoll, 1781), Cobalus disjuncta Herrich-Schäffer, 1869 of Dubiella dubius (Stoll, 1781) (not of Vettius lafrenaye (Latreille, [1824])), and Saliana vixen Evans, 1955 of Neoxeniades parna (Evans, 1955). The following are new and revised genus-species combinations: Euriphellus cebrenus (Cramer, 1777) (not Salatis Evans, 1952), Gorgopas extensa (Mabille, 1891) (not Polyctor Evans, 1953), Clytius shola (Evans, 1953) (not Staphylus Godman and Salvin, 1896), Perus narycus (Mabille, 1889) (not Ouleus Lindsey, 1925), Perus parvus (Steinhauser and Austin, 1993) (not Staphylus Godman and Salvin, 1896), Pholisora litus (Dyar, 1912) (not Bolla Mabille, 1903), Carrhenes decens (A. Butler, 1874) (not Antigonus Hübner, [1819]), Santa palica (Mabille, 1888) (not Chiothion Grishin, 2019), Bralus nadia (Nicolay, 1980) (not Anisochoria Mabille, 1876), Acerbas sarala (de Nicéville, 1889) (not Lotongus Distant, 1886), Caenides sophia (Evans, 1937) (not Hypoleucis Mabille, 1891), Hypoleucis dacena (Hewitson, 1876) (not Caenides Holland, 1896), Dotta tura (Evans, 1951) (not Astictopterus C. Felder and R. Felder, 1860), Nervia wallengrenii (Trimen, 1883) (not Kedestes Watson, 1893), Testia mammaea (Hewitson, 1876) (not Decinea Evans, 1955), Oxynthes trinka (Evans, 1955) (not Orthos Evans, 1955), Metrocles argentea (Weeks, 1901) (not Paratrytone Godman, 1900), Metrocles scitula (Hayward, 1951) (not Mucia Godman, 1900), Metrocles schrottkyi (Giacomelli, 1911) (not Metron Godman, 1900), Niconiades derisor (Mabille, 1891) (not Decinea Evans, 1955), Paratrytone samenta (Dyar, 1914) (not Ochlodes Scudder, 1872), Oligoria (Cobaloides) locutia (Hewitson, 1876) (not Quinta Evans, 1955), Psoralis (Saniba) laska (Evans, 1955) (not Vidius Evans, 1955), Psoralis (Saniba) arva (Evans, 1955) and Psoralis (Saniba) umbrata (Erschoff, 1876) (not Vettius Godman, 1901), Psoralis (Saniba) calcarea (Schaus, 1902) and Psoralis (Saniba) visendus (E. Bell, 1942) (not Molo Godman, 1900), Alychna gota (Evans, 1955) (not Psoralis Mabille, 1904), Adlerodea asema (Mabille, 1891) and Adlerodea subpunctata (Hayward, 1940) (not Eutychide Godman, 1900), Ralis immaculatus (Hayward, 1940) (not Mucia Godman, 1900), Rhinthon braesia (Hewitson, 1867) and Rhinthon bajula (Schaus, 1902) (not Neoxeniades Hayward, 1938), Cymaenes lochius Plötz, 1882 (not Lerema Scudder, 1872), Paracarystus ranka (Evans, 1955) (not Thoon Godman, 1900), Tricrista aethus (Hayward, 1951), Tricrista canta (Evans, 1955), Tricrista slopa (Evans, 1955), Tricrista circellata (Plötz, 1882), and Tricrista taxes (Godman, 1900) (not Thoon Godman, 1900), Gallio madius (E. Bell, 1941) and Gallio seriatus (Mabille, 1891) (not Vehilius Godman, 1900), Gallio garima (Schaus, 1902) (not Tigasis Godman, 1900), Tigasis corope (Herrich- Schäffer, 1869) (not Cynea Evans, 1955), Tigasis perloides (Plötz, 1882) (not Cymaenes Scudder, 1872), Amblyscirtes (Flor) florus (Godman, 1900) (not Repens Evans, 1955), Vidius fraus (Godman, 1900) (not Cymaenes Scudder, 1872), Nastra celeus (Mabille, 1891) (not Vehilius Godman, 1900), Nastra nappa (Evans, 1955) (not Vidius Evans, 1955), Vehilius warreni (Weeks, 1901) and Vehilius limae (Lindsey, 1925) (not Cymaenes Scudder, 1872), Cymaenes lumina (Herrich-Schäffer, 1869) (not Lerema Scudder, 1872), Cobalopsis valerius (Möschler, 1879) (not Cobalopsis Godman, 1900), Cobalopsis dictys (Godman, 1900) (not Papias Godman, 1900), Lerema (Morys) venias (Bell, 1942) (not Cobalopsis Godman, 1900), Papias latonia (Schaus, 1913) (not Cobalopsis Godman, 1900), Dion iccius (Evans, 1955) and Dion uza (Hewitson, 1877) (not Enosis Mabille, 1889), Vistigma (Vistigma) opus (Steinhauser, 2008) (not Thoon Godman, 1900), Saturnus fartuga (Schaus, 1902) (not Parphorus Godman, 1900), Phlebodes fuldai (E. Bell, 1930) (not Vettius Godman, 1901), Mnasitheus padus (Evans, 1955) (not Moeris Godman, 1900), Naevolus brunnescens (Hayward, 1939) (not Psoralis Mabille, 1904), Lamponia ploetzii (Capronnier, 1874) (not Vettius Godman, 1901), Mnestheus silvaticus Hayward, 1940 (not Ludens Evans, 1955), Rigga spangla (Evans, 1955) (not Sodalia Evans, 1955), Corticea vicinus (Plötz, 1884) (not Lento Evans, 1955), Mnasalcas thymoetes (Hayward, 1942) (not Mnasicles Godman, 1901), Mnasalcas boyaca (Nicolay, 1973) (not Pamba Evans, 1955), Vertica brasta (Evans, 1955) (not Lychnuchus Hübner, [1831]), Carystina discors Plötz, 1882 (not Cobalus Hübner, [1819]), Zetka irena (Evans, 1955) (not Neoxeniades Hayward, 1938), and Neoxeniades parna (Evans, 1955) (not Niconiades Hübner, [1821]). The following are new or revised species-subspecies combinations: Tagiades neira moti Evans, 1934, Tagiades neira canonicus Fruhstorfer, 1910, Tagiades sheba vella Evans, 1934, Tagiades sheba lola Evans, 1945, Tagiades korela biakana Evans, 1934, Tagiades korela mefora Evans, 1934, Tagiades korela suffusus Rothschild, 1915, Tagiades korela brunta Evans, 1949, Tagiades ravi ravina Fruhstorfer, 1910, Tagiades atticus carnica Evans, 1934, Tagiades atticus nankowra Evans, 1934, Tagiades atticus helferi C. Felder, 1862, Tagiades atticus balana Fruhstorfer, 1910, Tagiades inconspicua mathias Evans, 1934, Tagiades hovia kazana Evans, 1934, Tagiades elegans fuscata de Jong and Treadaway, 2007, Tagiades elegans semperi Fruhstorfer, 1910, Metron hypochlora tomba Evans, 1955, Decinea denta pruda Evans, 1955, and Choranthus orientis eleutherae (Bates, 1934) (previously in Pyrrhocalles Mabille, 1904). In addition to the abovementioned changes, the following new combinations involve newly proposed genus group names: Fulvatis fulvius (Plötz, 1882) and Fulvatis scyrus (E. Bell, 1934) (not Salatis Evans, 1952); Adina adrastor (Mabille and Boullet, 1912) (not Bungalotis Watson, 1893); Nascus (Praxa) prax Evans, 1952, Nascus (Bron) broteas (Cramer, 1780), and Nascus (Bron) solon (Plötz, 1882) (not Pseudonascus Austin, 2008); Chirgus (Turis) veturius (Plötz, 1884); Paches (Tiges) liborius (Plötz, 1884), and Paches (Tiges) mutilatus (Hopffer, 1874) (not Antigonus Hübner, [1819]); Paches (Tiges) exosa (A. Butler, 1877); Tolius tolimus (Plötz, 1884) and Tolius luctuosus (Godman & Salvin, 1894) (not Echelatus Godman and Salvin, 1894); Ancistroides (Ocrypta) caerulea (Evans, 1928), Ancistroides (Ocrypta) renardi (Oberthür, 1878), Ancistroides (Ocrypta) waigensis (Plötz, 1882), Ancistroides (Ocrypta) aluensis (Swinhoe, 1907), Ancistroides (Ocrypta) flavipes (Janson, 1886), and Ancistroides (Ocrypta) maria (Evans, 1949) (not Notocrypta de Nicéville, 1889); Lennia lena (Evans, 1937), Lennia binoevatus (Mabille, 1891), Lennia maracanda (Hewitson, 1876), and Lennia lota (Evans, 1937) (not Leona Evans, 1937); Trida barberae (Trimen, 1873) and Trida sarahae (Henning and Henning, 1998) (not Kedestes Watson, 1893); Noxys viricuculla (Hayward, 1951) (not Oxynthes Godman, 1900); Xeniades (Tixe) quadrata (Herrich-Schäffer, 1869), Xeniades (Tixe) rinda (Evans, 1955), Xeniades (Tixe) putumayo (Constantino and Salazar, 2013) (not Tisias Godman, 1901); Gracilata quadrinotata (Mabille, 1889) (not Styriodes Schaus, 1913); Hermio hermione (Schaus, 1913) (not Lento Evans, 1955); Cynea (Nycea) hycsos (Mabille, 1891), Cynea (Nycea) corisana (Plötz, 1882), Cynea (Nycea) popla Evans, 1955, Cynea (Nycea) iquita (E. Bell, 1941), Cynea (Nycea) robba Evans, 1955, Cynea (Nycea) melius (Geyer, 1832), and Cynea (Nycea) irma (Möschler, 1879); Eutus rastaca (Schaus, 1902) (not Eutychide Godman, 1900); Eutus yesta (Evans, 1955) (not Thoon Godman, 1900); Eutus mubevensis (E. Bell, 1932) (not Tigasis Godman, 1900); Gufa gulala (Schaus, 1902) (not Mucia Godman, 1900); Gufa fusca (Hayward, 1940) (not Tigasis Godman, 1900); Godmia chlorocephala (Godman, 1900) (not Onophas Godman, 1900); Rhomba gertschi (E. Bell, 1937) (not Justinia Evans, 1955); Mnasicles (Nausia) nausiphanes (Schaus, 1913) (not Tigasis Godman, 1900); Amblyscirtes (Flor) florus (Godman, 1900) (not Repens Evans, 1955); Rectava ignarus (E. Bell, 1932) (not Papias Godman, 1900); Rectava vorgia (Schaus, 1902) (not Cobalopsis Godman, 1900); Rectava nostra (Evans, 1955) (not not Vidius Evans, 1955); Lerema (Geia) geisa (Möschler, 1879) and Lerema (Geia) lyde (Godman, 1900) (not Morys Godman, 1900); Contrastia distigma (Plötz, 1882) (not Cymaenes Scudder, 1872); Mit (Mit) badius (E. Bell, 1930) (not Styriodes Schaus, 1913); Mit (Mit) gemignanii (Hayward, 1940), (not Mnasitheus Godman, 1900); Mit (Rotundia) schausi (Mielke and Casagrande, 2002), (not Enosis Mabille, 1889); Picova steinbachi (E. Bell, 1930) (not Saturnus Evans, 1955); Lattus arabupuana (E. Bell, 1932) (not Eutocus Godman, 1901); Gubrus lugubris (Lindsey, 1925) (not Vehilius Godman, 1900); Thargella (Pseudopapias) tristissimus (Schaus, 1902) (not Papias Godman, 1900); Koria kora (Hewitson, 1877) (not Justinia Evans, 1955); Justinia (Septia) septa Evans, 1955; Corta lycortas (Godman, 1900) (not Orthos Evans, 1955); Vertica (Brasta) brasta (Evans, 1955) (not Lychnuchus Hübner, [1831]); Calvetta calvina (Hewitson, 1866) (not Cobalus Hübner, [1819]); Neoxeniades (Bina) gabina (Godman, 1900) (not Orthos Evans, 1955); Oz ozias (Hewitson, 1878) and Oz sebastiani Salazar and Constantino, 2013 (not Lychnuchoides Godman, 1901); and Carystoides (Balma) balza Evans, 1955 and Carystoides (Balma) maroma (Möschler, 1877). Finally, unless stated otherwise, all subgenera, species, subspecies and synonyms of mentioned genera and species are transferred together with their parent taxa, and taxa not mentioned in this work remain as previously classified.
