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The classification of the largest subfamily of leafhoppers, Deltocephalinae, including 38 tribes, 923 genera, and 6683 valid species, is reviewed and revised. An updated phylogeny of the subfamily based on molecular (28S, Histone H3) and morphological data and an expanded taxon sample (37 taxa not included in previous analyses) is presented. Based on the results of these analyses and on the morphological examination of many representatives of the subfamily, the classification of the tribes and subtribes of Deltocephalinae is revised. Complete morphological descriptions, illustrations, lists of the included genera, and notes on their distribution, ecology, and important vector species are provided for the 38 recognized tribes and 18 subtribes. A dichotomous key to the tribes is provided. All names in the taxonomic treatments are hyperlinked to online resources for individual taxa which are supported by a comprehensive database for Deltocephalinae compiled using the taxonomic database software package 3I. The online functionality includes an interactive key to tribes and subtribes and advanced database searching options. Each taxon (subspecies through subfamily) has a unique taxon webpage providing nomenclatural information, lists of included taxa, an automated description (if available), images (if available), distributional information, bibliographic references and links to outside resources. Some observations and trends regarding the history of taxonomic descriptions in Deltocephalinae are reported. Four new tribes are described: Bahitini tribe nov. (25 genera), Bonsapeiini tribe nov. (21 genera), Phlepsiini tribe nov. (4 genera), and Vartini tribe nov. (7 genera). The circumscription and morphological characterization of Scaphoideini Oman, 1943 (61 genera) is substantially revised. Eleven new species are described: Acostemma stilleri sp. nov., Arrugada linnavuorii sp. nov., Drabescus zhangi sp. nov., Parabolopona webbi sp. nov., Goniagnathus emeljanovi sp. nov., Hecalus hamiltoni sp. nov., Scaphoideus omani sp. nov., Dwightla delongi sp. nov., Abimwa knighti sp. nov., Gannia viraktamathi sp. nov., and Doratulina dmitrievi sp. nov. Some family-group level taxonomic changes are made: Platymetopiini Haupt, 1929, Anoterostemmini Haupt, 1929, and Allygidiina Dmitriev, 2006 are synonymized with Athysanini Van Duzee, 1892, syn. nov.; Procepitini Dmitriev, 2002 is synonymized with Cicadulini Van Duzee, 1892, syn. nov.; Listrophorini Boulard, 1971 is synonymized with Chiasmini Distant, 1908, syn. nov.; Adamini Linnavuori & Al-Ne’amy, 1983, Dwightlini McKamey, 2003, and Ianeirini Linnavuori, 1978 are synonymized with Selenocephalini Fieber, 1872 syn.nov., and all three are now recognized as valid subtribes in their parent tribe. New placements of many genera to tribe and subtribe are made, and these are described in individual taxon treatments.
Five species of the subgenus Clessiniella of Cochlostoma (Cochlostomatidae) are recognized, viz. Cochlostoma (Clessiniella) villae (Strobel, 1851), Cochlostoma (Clessiniella) tergestinum (Westerlund, 1878), Cochlostoma (Clessiniella) waldemari (A.J. Wagner, 1897), Cochlostoma (Clessiniella) anomphale Boeckel, 1939 and Cochlostoma (Clessiniella) stelucarum sp. nov. The shells and the genitalia are described and the distributional data are summarized for all the species.
A new fossil diatom species, Tertiarius minutulus sp. nov., is described from a sediment sequence DEEP-5045-1 of Lake Ohrid. The species is characterized by small valves (3.0–8.0 μm) with a round shape, a marginal area with radially arranged costae and a central uneven area with scattered areolae. Externally, the alveoli are occluded by cribra perforated by irregularly arranged pores. Internally, the alveoli are simple and areolae are occluded with domed cribra. The marginal fultoportulae are situated on costae close to the valve margin, located on every 5th or 7th thick internal costa. One to three fultoportulae are present on the valve face, each surrounded by two to three satellite pores. One rimoportula is present, positioned on a costa at the valve face / mantle junction. The species is compared with morphologically similar taxa and a detailed differential diagnosis is provided. Tertiarius minutulus sp. nov. is known only as a fossil taxon from Lake Ohrid, with a first occurrence during the early stages of lake basin development.
The Iranian species of the genus Anomalon Panzer, 1804 (Hymenoptera: Ichneumonidae: Anomaloninae) are reviewed. Four species, Anomalon amseli (Hedwig, 1961), A. chinense (Kokujev, 1915), A. cruentatum (Geoffroy, 1785) and A. narinae Zardouei & Rakhshani sp. nov., are found to occur in Iran. The female of A. amseli is described for the first time. Anomalon chinense is a new record for Iran. A key to the known Anomalon species of Iran is provided.
Four new species of the genus Coecobrya, C. gejianbangi sp. nov., C. annulata sp. nov., C. ciliata sp. nov., and C. oculata sp. nov., are described from Guangxi caves as the representative of the genus in China. Coecobrya oculata sp. nov. of the boneti-group has 1+1 eyes and a serrate outer edge of the unguiculus. The other three species, devoid of eyes and with a tiny outer tooth on the unguiculus, are assigned here to the tenebricosa-group, assuming that the large tooth on the unguiculus is transformed into a tiny one in cave-obligate species. Clypeal chaetae in Entomobryoidea are systematically surveyed for the first time, and are found to be well diversified at species level. They have a potential taxonomical value in discriminating taxa of morphologically conserved groups.
A century and a half since the time of Hewitson, we are experiencing a renaissance in species discovery fueled by whole genome sequencing. A large-scale genomic analysis of Hesperiidae Latreille, 1809 (Lepidoptera), including primary type specimens, reveals a deluge of species new to science. One hundred of them (one in a new genus) are described here from the New World (type localities are given in parenthesis): Drephalys (Drephalys) diovalis Grishin, new species (Ecuador: Napo), Euriphellus panador Grishin, new species (Ecuador: Esmeraldas), Euriphellus panamicus Grishin, new species (Panama: Panama), Cecropterus (Thorybes) viridissimus Grishin, new species (Ecuador: Zamora-Chinchipe), Cecropterus (Murgaria) dariensis Grishin, new species (Panama: Darien), Urbanus (Urbanus) mericuti Grishin, new species (Ecuador: Napo), Telegonus (Telegonus) pastus Grishin, new species (Panama: Panama), Autochton (Autochton) dora Grishin, new species (Ecuador: Pastaza), Astraptes centralis Grishin, new species (Panama: Colón), Aguna claxonica Grishin, new species (Ecuador: Napo), Aguna esmeralda Grishin, new species (Ecuador: Esmeraldas), Aguna lata Grishin, new species (Guyana), Ridens angulinea Grishin, new species (Peru: Cuzco), Pythonides lera Grishin, new species (Peru: Cuzco), Pythonides latemarginatus Grishin, new species (Panama: Panama), Gindanes variegatus Grishin, new species (Brazil: Mato Grosso), Milanion (Milanion) virga Grishin, new species (Brazil: Rondônia), Milanion (Milanion) furvus Grishin, new species (Panama: Panama), Milanion (Milanion) laricus Grishin, new species (Ecuador: Napo), Charidia ronda Grishin, new species (Brazil: Rondônia), Pseudodrephalys tinas Grishin, new species (Peru: Loreto), Pseudodrephalys argus Grishin, new species (Suriname: Para), Achlyodes calvus Grishin, new species (Brazil: Santa Catarina), Spioniades artemis Grishin, new species (Panama: Panama), Spioniades artemidoides Grishin, new species (Brazil: Santa Catarina), Myrinia orieca Grishin, new species (Ecuador: Orellana), Myrinia aragua Grishin, new species (Venezuela: Aragua), Myrinia maculosa Grishin, new species (Guatemala), Myrinia manchada Grishin, new species (Guyana), Polyctor (Fenops) lamperus Grishin, new species (Panama: Darien), Nisoniades (Nisoniades) lutum Grishin, new species (Mexico: Guerrero. ), Bolla (Stolla) vena Grishin, new species (Venezuela: Aragua), Staphylus (Vulga) vula Grishin, new species (Mexico: Veracruz), Staphylus (Vulga) vulga Grishin, new species (Panama: Darien), Staphylus (Staphylus) rotundalus Grishin, new species (Ecuador: Napo), Staphylus (Staphylus) yucatanus Grishin, new species (Mexico: Quintana Roo/Yucatan), Heliopetes (Heliopetes) lana Grishin, new species (Guatemala), Canesia ella Grishin, new species (Venezuela: Barinas), Paches (Paches) loxeca Grishin, new species (Ecuador: Morona-Santiago), Clito congruens Grishin, new species (Panama: Colón), Cycloglypha corax Grishin, new species (Brazil: Rio de Janeiro), Festivia peruvia Grishin, new species (Peru: Huánuco), Decinea notata Grishin, new species (Ecuador: Napo), Pompeius fuscus Grishin, new species (Brazil: Minas Gerais), Vernia clara Grishin, new species (Panama: Chiriquí), Oligoria (Oligoria) obtena Grishin, new species (Ecuador: Napo), Thespieus mandal Grishin, new species (Brazil: Rio de Janeiro), Psoralis (Saniba) magnamacus Grishin, new species (Panama: Darien), Alychna ayonis Grishin, new species (Ecuador: Napo), Wahydra banios Grishin, new species (Ecuador: Tungurahua), Wahydra cuzcona Grishin, new species (Peru: Cuzco), Cynea (Cynea) aureofimbra Grishin, new species (Ecuador), Cynea (Nycea) quada Grishin, new species (Ecuador: Napo), Cynea (Quinta) achirae Grishin, new species (Mexico: Tamaulipas), Eutus amazonicus Grishin, new species (Peru: Madre de Dios), Eutus incus Grishin, new species (Peru: Cuzco), Eutus septemaculatus Grishin, new species (Brazil: Mato Grosso), Godmia viridicapita Grishin, new species (Ecuador: Napo), Rhomba pulla Grishin, new species (Peru: Cuzco), Niconiades victoria Grishin, new species (Mexico: Tamaulipas), Lancephallus purpurus Grishin, new genus and new species (Guyana), Mnasicles (Remella) ecua Grishin, new species (Ecuador: Pichincha), Amblyscirtes (Amblyscirtes) aeratus Grishin, new species (Mexico: Oaxaca), Amblyscirtes (Mastor) chrysoplea Grishin, new species (Mexico: Oaxaca), Amblyscirtes (Mastor) chrysomisa Grishin, new species (Mexico: Chiapas), Amblyscirtes (Flor) meridus Grishin, new species (Mexico: Veracruz), Rectava chiriquensis Grishin, new species (Panama: Chiriquí), Cobalopsis adictys Grishin, new species (Panama: Veraguas), Cymaenes melaporphyrus Grishin, new species (Mexico: San Luis Potosí), Lerema (Morys) ecuadorica Grishin, new species (Ecuador: Pichincha), Saturnus obscurior Grishin, new species (Panama: Darien), Cantha zoirodicta Grishin, new species (Peru: Madre de Dios), Cantha meiodicta Grishin, new species (Peru: Madre de Dios), Phlebodes duplex Grishin, new species (Guatemala: Cayuga), Lychnuchus (Enosis) valle Grishin, new species (Colombia: Valle), Eutychide ochoides Grishin, new species (Peru: Cuzco), Dion bora Grishin, new species (Panama: Darien), Dion occida Grishin, new species (Peru: Madre de Dios), Eprius (Eprius) veledinus Grishin, new species (Ecuador: Pichincha), Radiatus panamensis Grishin, new species (Panama: Panama), Pheraeus pulcher Grishin, new species (Peru: Madre de Dios), Callimormus rades Grishin, new species (Panama: Panama), Gubrus lubens Grishin, new species (Ecuador: Loja), Ludens labens Grishin, new species (Panama: Darien), Rigga isa Grishin, new species (Ecuador: Napo), Flaccilla lactea Grishin, new species (Peru: Cuzco), Falga athena Grishin, new species (Panama: Darien), Panoquina jay Grishin, new species (Peru: Loreto), Calpodes salianus Grishin, new species (Peru: Madre de Dios), Calpodes stingo Grishin, new species (Ecuador: Sucumbíos), Aides nobra Grishin, new species (Panama: Colón), Thracides pavo Grishin, new species (Mexico: Tabasco), Talides eluta Grishin, new species (Peru: Cuzco), Talides laeta Grishin, new species (Peru: Cuzco), Neoxeniades angustior Grishin, new species (Brazil: Rio de Janeiro), Damas zea Grishin, new species (Guyana), Tromba xantha Grishin, new species (Mexico: Veracruz), Perichares fura Grishin, new species (Ecuador: Pichincha), Carystoides (Balma) goliath Grishin, new species (Colombia: Valle), and Agathymus galeana Grishin, new species (Mexico: Nuevo Leon). Additionally, we present evidence to support 22 taxa as species (not subspecies or synonyms) and synonymize one genus and four species. Namely, the following taxa are species: Milanion pilta Evans, 1953 (not Milanion pilumnus Mabille and Boullet, 1917), Milanion latior Mabille and Boullet, 1917 (not a synonym of Milanion marciana Godman and Salvin, 1895), Charidia pilea Evans, 1953, and Charidia pocus Evans, 1953 (not Charidia lucaria (Hewitson, 1868)), Paches (Paches) gloriosus Röber, 1925 and Paches (Paches) loxana Evans, 1953 (not Paches (Paches) loxus (Westwood, 1852)), Spioniades anta Evans, 1953 (not Spioniades abbreviata (Mabille, 1888)), Decinea onasima (Hewitson, 1877) and Decinea formosus (Hayward, 1940) (not Decinea dama (Herrich-Schäffer, 1869)), Thespieus guerreronis (Dyar, 1913) (not Thespieus dalman (Latreille, [1824])), Cynea (Nycea) erebina (Möschler, 1879) and Cynea (Nycea) cleochares (Mabille, 1891) (not Cynea (Cynea) diluta (Herrich-Schäffer, 1869)), Amblyscirtes (Mastor) repta Evans, 1955 (not Amblyscirtes (Flor) florus (Godman, 1900)), Saturnus tiberius (Möschler, 1883), Saturnus conspicuus (E. Bell, 1941), Saturnus meton (Mabille, 1891), and Saturnus obscurus (E. Bell, 1941) (not Saturnus reticulata (Plötz, 1883)), Phlebodes sifax Evans, 1955 (not Phlebodes campo (E. Bell, 1947)), Eutychide ochus Godman, 1900 and Eutychide rogersi (Kaye, 1914) (not a subspecies and a synonym, respectively, of Eutychide subcordata (Herrich-Schäffer, 1869)), Falga mirabilis Evans, 1955, Falga jacta Evans, 1955, and Falga ombra Evans, 1955 (not Falga jeconia (A. Butler, 1870)); and the following taxa are junior subjective synonyms: Libra Evans, 1955 (of Phemiades Hübner, [1819]), Papilio clito Fabricius, 1787 of Milanion hemes hemes (Cramer, 1777), Pamphila hycsos Mabille, 1891 of Cynea (Nycea) erebina (Möschler, 1879), Hesperia olympia Plötz, 1882 of Eutychide subcordata (Herrich-Schäffer, 1869), and Hesperia ocrinus Plötz, 1882 of Aides aegita (Hewitson, 1866). Furthermore, we propose new combinations for genus-species: Lychnuchus (Enosis) ponka (Evans, 1955) (not Thoon Godman, 1900), and species-subspecies: Charidia pocus mayo Evans, 1953 (not Charidia lucaria (Hewitson, 1868)), Decinea onasima boliviensis (E. Bell, 1930) (not Decinea dama (Herrich-Schäffer, 1869)), Cynea (Nycea) erebina somba Evans, 1955 (not Pamphila hycsos Mabille, 1891), Saturnus tiberius suffuscus (Hayward, 1940) (not Saturnus reticulata (Plötz, 1883)), and Falga mirabilis odol Evans, 1955 (not Falga jeconia (A. Butler, 1870)). Then, Milanion pilumnus var. hemestinus Mabille and Boullet, 1917 is a junior subjective synonym of Milanion pilumnus pilumnus Mabille and Boullet, 1917, not of Milanion leucaspis (Mabille, 1878). Lectotypes are designated for nine taxa (names in original combinations below): Pellicia bromias Godman and Salvin, 1894 (Mexico: Veracruz, Atoyac), Nisoniades perforata Möschler, 1879 (Colombia), Helias ascalaphus Staudinger, 1876 (central Panama), Pamphila hycsos Mabille, 1891 (Colombia), Amblyscirtes fluonia Godman, 1900 (Mexico: Guerrero, Xocomanatlan), Mastor anubis Godman, 1900 (Mexico: Guerrero, Omiltemi), Eutychide ochus Godman, 1900 (Mexico: Veracruz, Atoyac), Cobalus subcordata Herrich-Schäffer, 1869 (Southeast Brazil), and Thracides xanthura Godman, 1901 (Panama: Chiriquí Province, Bugaba). A neotype is designated for Eudamus briccius Plötz, 1881 (Guyana: Iwokrama Forest).
ZooBank registration. urn:lsid:zoobank.org:pub:ACDF923B-906D-460E-9707-259E0ECDBCA8
Analyses of whole genomic shotgun datasets, COI barcodes, morphology, and historical literature suggest that the following 13 butterfly species from the family Hesperiidae (Lepidoptera: Papilionoidea) in Texas, USA are distinct from their closest named relatives and therefore are described as new (type localities are given in parenthesis): Spicauda atelis Grishin, new species (Hidalgo Co., Mission), Urbanus (Urbanus) rickardi Grishin, new species (Hidalgo Co., nr. Madero), Urbanus (Urbanus) oplerorum Grishin, new species (Hidalgo Co., Mission/Madero), Telegonus tsongae Grishin, new species (Starr Co., Roma), Autochton caballo Grishin, new species (Hidalgo Co., 6 mi W of Hidalgo), Epargyreus fractigutta Grishin, new species (Hidalgo Co., McAllen), Aguna mcguirei Grishin, new species (Cameron Co., Brownsville), Polygonus pardus Grishin, new species (Hidalgo Co., McAllen), Arteurotia artistella Grishin, new species (Hidalgo Co., Mission), Heliopetes elonmuski Grishin, new species (Cameron Co., Boca Chica), Hesperia balcones Grishin, new species (Travis Co., Volente), Troyus fabulosus Grishin, new species (Hidalgo Co., Peñitas), and Lerema ochrius Grishin, new species (Hidalgo Co., nr. Relampago). Most of these species are known in the US almost exclusively from the Lower Rio Grande Valley in Texas. Nine of the holotypes were collected in 1971-1975, a banner period for butterfly species newly recorded from the Rio Grande Valley of Texas; five of them collected by William W. McGuire, and one by Nadine M. McGuire. At the time, these new species have been recorded under the names of their close relatives. A Neotype is designated for Papilio fulminator Sepp, [1841] (Suriname). Lectotypes are designated for Goniurus teleus Hübner, 1821 (unknown, likely in South America), Goniloba azul Reakirt, [1867] (Mexico: Veracruz) and Eudamus misitra Plötz, 1881 (Mexico). Several taxonomic changes are proposed. The following taxa are species (not subspecies): Spicauda zalanthus (Plötz, 1880), reinstated status (not Spicauda teleus (Hübner, 1821)), Telegonus fulminator (Sepp, [1841]), reinstated status (not Telegonus fulgerator (Walch, 1775), Telegonus misitra (Plötz, 1881), reinstated status (not Telegonus azul (Reakirt, [1867])), Autochton reducta (Mabille and Boullet, 1919), new status (not Autochton potrillo (Lucas, 1857)), Epargyreus gaumeri Godman and Salvin, 1893, reinstated status (not Epargyreus clavicornis (Herrich-Schäffer, 1869)), and Polygonus punctus E. Bell and W. Comstock, 1948, new status (not Polygonus savigny (Latreille, [1824])). Urbanus ehakernae Burns, 2014 and Epargyreus socus chota Evans, 1952 are junior subjective synonyms of Urbanus alva Evans, 1952 and Epargyreus clavicornis (Herrich-Schäffer, 1869), respectively, and Epargyreus gaumeri tenda Evans, 1955, new combination is not a subspecies of E. clavicornis.
ZooBank registration. https://zoobank.org/D5462F9E-E08D-46C6-898D-76EE7466DD19
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 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 (HerrichSchäffer, 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 genusspecies 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 (HerrichSchäffer, 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.
Genomic analysis of Pyrginae Burmeister, 1878 (Lepidoptera: Hesperiidae Latreille, 1809) with an emphasis on the tribes Achlyodini Burmeister, 1878 and Carcharodini Verity, 1940 reveals many inconsistencies between the resulting phylogeny and the current classification. These problems are corrected by proposing new taxa, changing the ranks of others, or synonymizing them, and transferring species between genera. As a result, five subtribes, one genus, 20 subgenera, and one species are proposed as new: Cyclosemiina Grishin, new subtribe (type genus Cyclosemia Mabille, 1878), Ilianina Grishin, new subtribe (type genus Iliana E. Bell, 1937), Nisoniadina Grishin, new subtribe (type genus Nisoniades Hübner, [1819]), Burcina Grishin, new subtribe (type genus Burca E. Bell and W. Comstock, 1948), and Pholisorina Grishin, new subtribe (type genus Pholisora Scudder, 1872), all in Carcharodini; Lirra Grishin, new genus (type species Leucochitonea limaea Hewitson, 1868) in Pythonidina Grishin, 2019; Trifa Grishin, new subgenus (type species Tagiades jacobus Plötz, 1884), Tuberna Grishin, new subgenus (type species Pythonides contubernalis Mabille, 1883), Ebona Grishin, new subgenus (type species Quadrus eboneus E. Bell, 1947), Noctis Grishin, new subgenus (type species Achlyodes accedens Mabille, 1895), and Cyrna Grishin, new subgenus (type species Achlyodes cyrna Mabille, 1895) of Quadrus Lindsey, 1925; Liddia Grishin, new subgenus (type species Helias pallida R. Felder, 1869), Minna Grishin, new subgenus (type species Achlyodes minna Evans, 1953), and Thilla Grishin, new subgenus (type species Eurypterus later Mabille, 1891) of Eantis Boisduval, 1836; Torgus Grishin, new subgenus (type species Ouleus gorgus E. Bell, 1937) of Iliana E. Bell, 1937; Fenops Grishin, new subgenus (type species Cabares enops Godman and Salvin, 1894) of Polyctor Evans, 1953; Bezus Grishin, new subgenus (type species Pellicia bessus Möschler, 1877) and Macarius Grishin, new subgenus (type species Pellicia macarius Herrich-Schäffer, 1870) of Nisoniades Hübner, [1819]; Quadralis Grishin, new subgenus (type species Pterygospidea extensa Mabille, 1891) of Gorgopas Godman and Salvin, 1894; Menuda Grishin, new subgenus (type species Nisoniades menuda Weeks, 1902) and Narycus Grishin, new subgenus (type species Pythonides narycus Mabille, 1889) of Perus Grishin, 2019; Bovaria Grishin, new subgenus (type species Achlyodes cyclops Mabille, 1876), Sebia Grishin, new subgenus (type species Nisoniades eusebius Plötz, 1884), and Stolla Grishin, new subgenus (type species Pholisora balsa E. Bell, 1937) of Bolla Mabille, 1903; Vulga Grishin, new subgenus (type species Achlyodes vulgata Möschler, 1879) and Capilla Grishin, new subgenus (type species Helias aurocapilla Staudinger, 1876, currently a junior subjective synonym of Hesperia musculus Burmeister, 1875) of Staphylus Godman and Salvin, 1896; and Quadrus (Zera) vivax Grishin, new species (type locality in Brazil: Rio de Janeiro). The following 10 are subgenera, not genera or synonyms: Ouleus Lindsey, 1925 and Zera Evans, 1953 of Quadrus Lindsey, 1925; Atarnes Godman and Salvin, 1897 and Eburuncus Grishin, 2012 of Milanion Godman and Salvin, 1895; Pachyneuria Mabille, 1888 and Austinus O. Mielke and Casagrande, 2016 of Sophista Plötz, 1879; Hemipteris Mabille, 1889 and Mictris Evans, 1955 of Pellicia Herrich-Schäffer, 1870; and Hesperopsis Dyar, 1905 and Scantilla Godman and Salvin, 1896 of Staphylus Godman and Salvin, 1896. The following 7 are species, not subspecies: Quadrus (Ebona) cristatus (Steinhauser, 1989) (not Quadrus (Ebona) negrus (Nicolay, 1980)), Quadrus (Quadrus) ophia (A. Butler, 1870) (not Quadrus (Quadrus) lugubris (R. Felder, 1869)), Quadrus (Zera) gellius (Mabille, 1903) and Quadrus (Zera) servius (Plötz, 1884) (not Quadrus (Zera) hyacinthinus (Mabille, 1877)), Mimia pazana Evans,1953 (not Mimia phidyle (Godman and Salvin, 1894)), Polyctor (Polyctor) dagua Evans, 1953 (not Polyctor (Polyctor) polyctor (Prittwitz, 1868)), and Staphylus (Vulga) satrap Evans, 1953 (not Staphylus (Vulga) saxos Evans, 1953); and these 8 are species, not synonyms: Quadrus (Zera) menedemus (Godman and Salvin, 1894) (not Quadrus (Zera) tetrastigma (Sepp, [1847])), Pellicia (Pellicia) bilinea Mabille, 1889 (not Pellicia (Pellicia) dimidiata Herrich-Schäffer, 1870), Pellicia (Hemipteris) nema Williams and Bell, 1939 (not Pellicia (Pellicia) theon Plötz, 1882), Bolla (Bovaria) sodalis Schaus, 1913 (not Bolla (Bolla) imbras (Godman and Salvin, 1896)), Bolla (Bovaria) aplica (E. Bell, 1937) (not Bolla (Sebia) eusebius (Plötz, 1884)), Bolla (Sebia) chilpancingo (E. Bell, 1937) (not Bolla (Bolla) subapicatus (Schaus, 1902)), and Bolla (Stolla) madrea (R. Williams and E. Bell, 1940) and Bolla (Stolla) hazelae (Hayward, 1940) (not Bolla (Stolla) zorilla (Plötz, 1886)). The following 2 are junior subjective synonyms: Achlyodes erisichthon Plötz, 1884 of Quadrus (Zera) servius (Plötz, 1884) (not a subspecies of Quadrus (Zera) tetrastigma (Sepp, [1847]) and Staphylus subapicatus Schaus, 1902 of Bolla (Bolla) imbras (Godman and Salvin, 1896). Furthermore, we propose the following additional new genus-species combination: Gindanes homer (Evans, 1953), Gindanes nides (O. Mielke and Casagrande, 2002), Gindanes maraca (O. Mielke and Casagrande, 1992), Gindanes jenmorrisae (Shuey and Ramírez. 2022), Gindanes tullia (Evans, 1953), Gindanes herennius (Geyer, [1838]), Gindanes proxenus (Godman and Salvin, 1895), Gindanes parallelus (Mabille, 1898), Gindanes braga (Evans, 1953), Gindanes hampa (Evans, 1953), Gindanes rosa (Steinhauser, 1989), Gindanes neivai (Hayward, 1940), Gindanes mundo (H. Freeman, 1979), Gindanes eminus (E. Bell, 1934), Quadrus (Trifa) francesius Freeman, 1969, Quadrus (Trifa) ineptus (Draudt, 1922), Quadrus (Trifa) jacobus (Plötz, 1884), Quadrus (Tuberna) lancea (Hewitson, 1868), Quadrus (Ebona) pescada (E. Bell, 1956), Lirra pteras (Godman and Salvin, 1895), and Lirra limaea (Hewitson, 1868) (not Pythonides Hübner, 1819); Quadrus (Cyrna) zora (Evans, 1953) (not Bolla Mabille, 1903); Eantis later (Mabille, 1891) and Eantis haber (Mabille, 1891) (not Aethilla Hewitson, 1868); Iliana (Torgus) gorgus (E. Bell, 1937) and Iliana (Torgus) taurus (Evans, 1953) (not Eantis Boisduval, 1836); Bolla (Stolla) evemerus (Godman and Salvin, 1896), Bolla (Stolla) chlora (Evans, 1953), Bolla (Stolla) astra (R. Williams and E. Bell, 1940), Bolla (Stolla) balsa (E. Bell, 1937), Bolla (Stolla) tridentis (Steinhauser, 1989), Bolla (Stolla) esmeraldus (L. Miller, 1966), Bolla (Stolla) chlorocephala (Latreille, [1824]), and Bolla (Stolla) incanus (E. Bell, 1932) (not Staphylus Godman and Salvin, 1896). Finally, lectotypes are designated for Achlyodes servius Plötz, 1884 (type locality in Brazil: Rio de Janeiro), Pellicia theon Plötz, 1882 (type locality in South America), and Nisoniades eusebius Plötz, 1884 (type locality in Central America). Unless stated otherwise, all subgenera, species, subspecies, and synonyms of mentioned genera and species are transferred with their parent taxa, and others remain as previously classified.
ZooBank registration. http://zoobank.org/B9AFA1A9-8664-4F31-B4D9-ACF7780C7CC6
New species of genus Atractides Koch, 1837 (Acari: Hydrachnidiae, Hygrobatidae) from Qinghai, China
(2022)
The paper deals with five new species of the genus Atractides Koch, 1837 (Acari: Hydrachnidiae, Hygrobatidae) collected from Qinghai Province, P.R. China, Atractides (Atractides) biprojectus Zhang, Li & Guo sp. nov., Atractides (Atractides) smiti Zhang, Li & Guo sp. nov., Atractides (Atractides) menyuanensis Zhang, Li & Guo sp. nov., Atractides (Atractides) longiprojectus Zhang, Li & Guo sp. nov. and Atractides (Atractides) xianmiensis Zhang, Li & Guo sp. nov. All the new species are described and illustrated in detail, and all the type specimens are deposited in the Institute of Entomology, Guizhou University, Guiyang, China (GUGC).
A new mesoserphid wasp from the Middle Jurassic of northeastern China (Hymenoptera, Proctotrupoidea)
(2017)
A new genus and species of Mesoserphidae (Hymenoptera), Juraserphus modicus gen. et sp. nov., is described based on a well-preserved fossil specimen from the Middle Jurassic Jiulongshan Formation of northeastern China. It is characterized by the following forewing features: the forking of Rs+M located approximately one-third of the distance between 1m-cu and 2r-rs, both 1cu-a and 2cu-a antefurcal; 1-M more than twice as long as 1m-cu and hind wing with cells r and rm closed. In addition, it has a short ovipositor, only extending slightly beyond the metasomal apex. Its new morphological characters broaden the diversity of Mesoserphidae in the Mesozoic and provide new insights into the evolution and relationships of Mesoserphidae.
New fossil records of Xyelidae (Hymenoptera) from the Middle Jurassic of Inner Mongolia, China
(2021)
A new genus and two new species of Xyelidae Newman, 1834 (Hymenoptera Linnaeus, 1758), Platyxyela tenuis sp. nov. and Scleroxyela daohugouensis gen. et sp. nov. are described and illustrated from the Middle Jurassic Daohugou Beds of Ningcheng, Inner Mongolia, China. Platyxyela tenuis sp. nov. (Xyelinae Newman, 1834, Liadoxyelini Rasnitsyn, 1966) can be distinguished from its congeners by short forewing length and ovipositor sheath strongly narrowed toward acute apex. Scleroxyela daohugouensis gen. et sp. nov. is placed within Macroxyelinae Ashmead, 1898, Xyeleciini Benson, 1945 based on pterositigma completely sclerotised and costal space sclerotised before ptrostigma, 1-Rs half as long as 1-M, and 1m-cu short. It is the first recorded species of Xyeleciini in Daohugou Beds. Furthermore, an updated key of identification of subfamilies and tribes of Xyelidae is provided. Our findings of new morphological data provide insights into the early evolution of Hymenoptera, as well as effectively enrich our understanding of the diversity of Xyelidae in the Mesozoic.
Two new species, Russula pseudopunicea C.L.Hou, G.Q.Cheng & H.Zhou sp. nov. and R. wulingshanensis C.L.Hou, G.Q.Cheng & H.Zhou sp. nov., from Yanshan mountains in North China are described herein based on morphological and phylogenetic analyses of nrITS, and nrLSU-rpb2-mtSSU gene regions. Morphologically, R. pseudopunicea sp. nov. is characterised by a reddish brown, light brown to brownish orange pileus with a greyish yellow margin, subglobose to broadly ellipsoid basidiospores with warts forming a partial reticulum and pleurocystidia turning grey to purplish red in sulfovanillin. Russula wulingshanensis sp. nov. is characterised by a purple pinkish pileus with a grey-white to grey-purple margin, subglobose to broadly ellipsoid basidiospores with isolated warts, and pileocystidia turning black in sulfovanillin. Phylogenetic and morphological analyses resolved the two species in Russula subg. Heterophyllidia. Russula pseudopunicea sp. nov. and R. wulingshanensis sp. nov. were placed in the lineages of subsect. Virescentinae and subsect. Griseinae, respectively.
The subgenus Indonthophagus Kabakov, 2006 of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini) is reviewed. A morphological definition of it is provided. A diagnosis, based both on external and internal characters, of the seven species deemed to belong to the group according to literature, is supplied. As a result, Onthophagus spinifex (Fabricius, 1781), previously placed with some uncertainty in the subgenus, is definitively excluded. A key to Indonthophagus species is also provided, as well as images of male, aedeagus and endophallus for each species. New country records of Onthophagus turbatus Walker, 1858 for Pakistan, Nepal and Bhutan are given.
ZooBank registration. urn:lsid:zoobank.org:pub:9D0CE9F2-35CF-449D-8984-1B4C722762F7
Die Notwendigkeit des vorliegenden Beitrags zur Trichogramma-Fauna in Deutschland hat folgende Gründe: 1) Die Erfassung der Fauna dient der Kenntnis und der Erhaltung der genetischen und biologischen Vielfalt im Sinne der Konvention zur Biodiversität. 2) Die Gattung Trichogramma Westwood, 1833 (Hymenoptera: Trichogrammatidae) ist im biologischen Pflanzen- und Vorratsschutz weltweit von ökonomischem Nutzen. Das Spektrum möglicher Zielschädlinge umfasst in Deutschland mindestens 30 Schädlinge im Pflanzen- und Vorratsschutz (Zimmermann 2004). Als Eiparasitoide greifen sie bereits das Eistadium der Schädlinge an, was gegenüber allen anderen biologischen und chemischen Bekämpfungsmethoden ein entscheidender Vorteil ist. Trichogramma spp. werden in Mitteleuropa als einzige Nutzarthropoden großflächig im Freiland eingesetzt. Daher ist es von besonderer Bedeutung die natürlich auftretenden Arten und deren lokale Ökotypen zu charakterisieren, zu schützen und gegebenenfalls für die biologische Schädlingsbekämpfung zu nutzen. Nicht zuletzt im Hinblick auf eine zu erwartende Registrierungspflicht für Nützlinge in Deutschland ist es ein grundlegender Beitrag, die Wissenslücken hinsichtlich der einheimischen Trichogramma-Fauna aufzuzeigen.
The genus Raveniola Zonstein, 1987 is found to be represented in Western Asia by 16 species: ♂♀ R. adjarica sp. nov. (Georgia), ♂ R. anadolu sp. nov. (Turkey), ♂ R. arthuri Kunt & Yağmur, 2010 (Turkey), ♂ R. birecikensis sp. nov. (Turkey), ♂♀ R. dunini sp. nov. (Armenia, Azerbaijan, Iran), ♂♀ R. hyrcanica Dunin, 1988 (Azerbaijan), ♂ R. marusiki sp. nov. (Iran), ♂ R. mazandaranica Marusik, Zamani & Mirshamsi, 2014 (Iran), ♂♀ R. micropa (Ausserer, 1871) (Turkey), ♀ R. nana sp. nov. (Turkey), ♂♀ R. niedermeyeri (Brignoli, 1972) (Iran), ♂♀ R. pontica (Spassky, 1937) (Russia, Georgia), ♀ R. sinani sp. nov. (Turkey), ♂♀ R. turcica sp. nov. (Turkey), ♂♀ R. vonwicki Zonstein, 2000 (Iran) and ♂♀ R. zaitzevi (Charitonov, 1948) (Azerbaijan, Georgia) = ♀ Brachythele recki Mcheidze, 1983, syn. nov. Eight species are described as new; others are redescribed from types and/or conspecific material. Males of R. micropa and R. zaitzevi, hitherto unknown, are described for the first time. Data on the variability, relationships, distribution and ecology of all considered species are also provided.
The genus Trypheridium Brancucci, 1985 is endemic to the Hindu Kush Himalayan Region, and is currently known from a single species, T. nuristanicum (Wittmer, 1956). Here, the genus is reviewed, T. nuristanicum nom. emend. is re-described and T. kashmiricum sp. nov. is described from Kashmir Himalayan Region of India. Descriptions, diagnoses, high quality images, distribution maps and identification keys are presented. The morphology and distribution of Trypheridium are discussed and compared with those of the closely related genus Trypherus LeConte, 1851.
A new monotypic genus of flatid planthoppers (Hemiptera: Fulgoromorpha: Flatidae), Medleria gen. nov., is described for Medleria caudata gen. et sp. nov. (type species) from the island of Socotra (Yemen). Habitus, male and female external and internal genital structures of the new species are illustrated and compared with similar taxa. Medleria caudata gen. et sp. nov. is probably endemic to Socotra where it is known to date from a small area of the Dixam mountain plateau only.