New butterfly taxa and findings from genomic analyses

Continuing our genomics-driven exploration of butterfly taxonomy, we integrate phylogenetic trees from all protein-coding genes with existing taxonomic and phenotypic knowledge and uncover further insights into butterfly systematics. As a result, one subgenus, 44 species, and 11 subspecies are proposed as new (type species in original combinations or type localities are listed in parentheses): <i>Porphaurea</i> Grishin, <b>subgen. n</b>. (<i>Phoenicops porphyropis</i> Meyrick & Lower, 1902) of <i>Chaetocneme</i> C. Felder, 1860 in Hesperiidae Latreille, 1809; <i>Dione</i> (<i>Agraulis</i>) <i>lamasi</i> Grishin, <b>sp. n</b>. (Peru: Cuzco) in Nymphalidae Rafinesque, 1815; <i>Lasaia</i> (<i>Lasaia</i>) <i>chiapis</i> Grishin, <b>sp. n</b>. (Mexico: Chiapas), <i>Lasaia</i> (<i>Lasaia</i>) <i>occalla</i> Grishin, <b>sp. n</b>. (Mexico: Oaxaca), <i>Lasaia</i> (<i>Lochris</i>) <i>oilenor</i> Grishin, <b>sp. n</b>. (Belize), <i>Lasaia</i> (<i>Lochris</i>) <i>oilepanor</i> Grishin, <b>sp. n</b>. (Panama: Colón), <i>Lasaia</i> (<i>Lochris</i>) <i>oilemarca</i> Grishin, <b>sp. n</b>. (Peru: Cajamarca), and <i>Emesis</i> (<i>Tenedia</i>) <i>peripore</i> Grishin, <b>sp. n</b>. (Trinidad) in Riodinidae Grote, 1895 (1827); and <i>Burara danata</i> Grishin, <b>sp. n</b>. (China: central Fujian), <i>Burara lawana</i> Grishin, <b>sp. n</b>. (Philippines: Palawan), <i>Bungalotis quadra</i> Grishin, <b>sp. n</b>. (Panama: Chiriquí), <i>Bungalotis barbalotis</i> Grishin, <b>sp. n</b>. (Peru: Loreto), <i>Cecropterus</i> (<i>Murgaria</i>) <i>eryssus</i> Grishin, <b>sp. n</b>. (Ecuador: Sucumbíos), <i>Cephise panuspe</i> Grishin, <b>sp. n</b>. (Panama: Panamá), <i>Noctuana haematoesta</i> Grishin, <b>sp. n</b>. (Colombia: Valle del Cauca), <i>Noctuana statonama</i> Grishin, <b>sp. n</b>. (Panama: Chiriquí), <i>Diaeus piura</i> Grishin, <b>sp. n</b>. (Peru: Piura), <i>Zopyrion</i> (<i>Zopyrion</i>) <i>cruise</i> Grishin, <b>sp. n</b>. (Mexico: Veracruz), <i>Chaetocneme triuna</i> Grishin, <b>sp. n</b>. (Papua New Guinea: Oro), <i>Chaetocneme brazza</i> Grishin, <b>sp. n</b>. (Indonesia: West Papua), <i>Ge pina</i> Grishin, <b>sp. n</b>. (Philippines: Mindanao), <i>Parnara guda</i> Grishin, <b>sp. n</b>. (India: West Bengal), <i>Parnara sulawesa</i> Grishin, <b>sp. n</b>. (Indonesia: South Sulawesi), <i>Tisias panamyna</i> Grishin, <b>sp. n</b>. (Panama: Panamá), <i>Xeniades</i> (<i>Tixe</i>) <i>lora</i> Grishin, <b>sp. n</b>. (Peru: Loreto), <i>Oligoria</i> (<i>Oligoria</i>) <i>costaria</i> Grishin, <b>sp. n</b>. (Costa Rica: Alajuela), <i>Oligoria</i> (<i>Oligoria</i>) <i>bahia</i> Grishin, <b>sp. n</b>. (Brazil: Bahia), <i>Alychna ventanilla</i> Grishin, <b>sp. n</b>. (Ecuador: Napo), <i>Thoon cuadius</i> Grishin, <b>sp. n</b>. (Ecuador: Napo), <i>Viridina viridella</i> Grishin, <b>sp. n</b>. (Peru: Cuzco), <i>Tricrista lingulata</i> Grishin, <b>sp. n</b>. (Brazil: Rondônia), <i>Tricrista taxolla</i> Grishin, <b>sp. n</b>. (Peru: Madre de Dios), <i>Vettius fuscapicalis</i> Grishin, <b>sp. n</b>. (Ecuador: Napo), <i>Mnasitheus oaxaceus</i> Grishin, <b>sp. n</b>. (Mexico: Oaxaca), <i>Tarmia bolivia</i> Grishin, <b>sp. n</b>. (Bolivia: La Paz), <i>Artines tines</i> Grishin, <b>sp. n</b>. (French Guiana), <i>Calpodes salma</i> Grishin, <b>sp. n</b>. (Panama: Colón), <i>Calpodes anabara</i> Grishin, <b>sp. n</b>. (Brazil: Rio de Janeiro), <i>Talides megamaca</i> Grishin, <b>sp. n</b>. (Mexico: Veracruz), <i>Perichares guatine</i> Grishin, <b>sp. n</b>. (Guatemala), <i>Perichares solamancha</i> Grishin, <b>sp. n</b>. (Peru: Cuzco), <i>Perichares indivisa</i> Grishin, <b>sp. n</b>. (Bolivia: La Paz), <i>Perichares amaletes</i> Grishin, <b>sp. n</b>. (Peru: Madre de Dios), <i>Perichares aura</i> Grishin, <b>sp. n</b>. (Peru: Cuzco), <i>Lycas gabriel</i> Grishin, <b>sp. n</b>. (Peru: Madre de Dios). <i>Burara danata himavata</i> Grishin, <b>ssp. n</b>. (Nepal), <i>Burara gomata burmana</i> Grishin, <b>ssp. n</b>. (Myanmar: Sagaing Region), <i>Burara gomata namata</i> Grishin, <b>ssp. n</b>. (China: Hong Kong), <i>Burara lorquini vichitra</i> Grishin, <b>ssp. n</b>. (unknown, likely in or around the Philippines), <i>Cecropterus</i> (<i>Murgaria</i>) <i>chales estales</i> Grishin, <b>ssp. n</b>. (USA: Texas, Starr Co.), <i>Pholisora mejicanus yuesanus</i> Grishin, <b>ssp. n</b>. (USA: Colorado, El Paso Co.), <i>Celotes sabinus verdinus</i> Grishin, <b>ssp. n</b>. (USA: Arizona, Yavapai Co.), <i>Ephyriades brunnea sansalva</i> Grishin, <b>ssp. n</b>. (Bahamas: San Salvador Island), <i>Ephyriades brunnea turcaica</i> Grishin, <b>ssp. n</b>. (Turks & Caicos Islands), <i>Ephyriades arcas norleewa</i> Grishin, <b>ssp. n</b>. (Anguilla), and <i>Calpodes hewitsoni supernalis</i> Grishin, <b>ssp. n</b>. (Mexico: Chiapas) in Hesperiidae. <i>Metron fas</i> Grishin, <b>nom. nov</b>. (Costa Rica: Heredia) is proposed as a new substitute name that replaces <i>Metron fascia</i> Grishin, 2025 preoccupied by <i>Metron fascia</i> Draudt, 1923. The following are valid species or subspecies (not subspecies or synonyms of taxa listed in parentheses): <i>Burara lara</i> (Leech, 1893), <b>stat. nov</b>., <i>Burara vajra</i> (Fruhstorfer, 1911), <b>stat. nov</b>., <i>Burara lorquini</i> (Mabille, 1876), <b>stat. rest</b>., and <i>Burara radiosa</i> (Plötz, 1885), <b>stat. rest</b>. (not <i>Burara gomata</i> (Moore, [1866])), <i>Cecropterus</i> (<i>Murgaria</i>) <i>chales</i> (Godman & Salvin, 1893), <b>stat. rest</b>. (not <i>Cecropterus</i> (<i>Murgaria</i>) <i>doryssus</i> (Swainson, 1831)), <i>Noctuana bipuncta</i> (Plötz, 1884), <b>stat. rest</b>. (not <i>Noctuana lactifera</i> (Butler & H. Druce, 1872)), <i>Zopyrion</i> (<i>Timochreon</i>) <i>forta</i> (Evans, 1953), <b>stat. nov</b>. and <i>Zopyrion</i> (<i>Timochreon</i>) <i>tampa</i> (Evans, 1953), <b>stat. nov</b>. (not <i>Zopyrion</i> (<i>Timochreon</i>) <i>satyrus</i> (C. Felder & R. Felder, 1867)), <i>Parnara mangala</i> (F. Moore, 1866), <b>stat. rev</b>. (not <i>Parnara guttatus</i> (Bremer & Grey, [1852])), <i>Parnara philotas</i> (de Nicéville, 1895), <b>stat. rest</b>., <i>Parnara daendeli</i> (Plötz, 1885), <b>stat. rest</b>., and <i>Parnara bada nondoa</i> (Plötz, 1886), <b>stat. nov</b>. (not <i>Parnara bada bada</i> (F. Moore, 1878)), <i>Xeniades corna</i> Evans, 1955, <b>stat. nov</b>. (not <i>Xeniades</i> (<i>Xeniades</i>) <i>chalestra</i> (Hewitson, 1866)), <i>Oligoria</i> (<i>Oligoria</i>) <i>pindar</i> (Schaus, 1913), <b>stat. rest</b>. (not <i>Oligoria</i> (<i>Oligoria</i>) <i>lucifer</i> (Hübner, [1829]), <i>Perichares crotona</i> (Hewitson, 1866), <b>stat. rest</b>. (not <i>Perichares seneca</i> (Latreille, [1824])), <i>Perichares fulvimargo</i> (Butler, 1873), <b>stat. rest</b>. and <i>Perichares luscinia</i> (Plötz, 1882), <b>stat. rest</b>. (not <i>Perichares deceptus</i> (Butler & H. Druce, 1872), <i>Perichares philetes trinitad</i> (Lucas, 1857), <b>stat. rev</b>. (not <i>Perichares philetes philetes</i> (Gmelin, [1790])), and <i>Lycas boisduvalii</i> (Ehrmann, 1909), <b>stat. rest</b>. (not <i>Lycas godart</i> (Latreille, [1824])). The following are new species-subspecies combination: <i>Burara lorquini minda</i> Chiba & Tsukiyama, 2009 (not <i>Burara gomata</i> (Moore, [1866])) and <i>Parnara mangala ormuzd</i> (Grum-Grshimailo, 1888) (not <i>Parnara guttatus</i> (Bremer & Grey, [1852])). The following are new synonyms: <i>Parnara guttatus batta</i> Evans, 1949, <b>syn. nov</b>. of <i>Parnara mangala</i> (F. Moore, 1866), <b>stat. rev</b>. and <i>Parnara kuwanoi</i> Seok, 1937, <b>syn. nov</b>. of <i>Pelopidas sinensis</i> (Mabille, 1877); and a new synonym placement, <i>Eudamus electra</i> Lintner, 1881, under <i>Ephyriades brunnea floridensis</i> E. Bell & W. P. Comstock, 1948, not <i>Ephyriades brunnea brunnea</i> (Herrich-Schäffer, 1865). Furthermore, we discuss and illustrate the following particulars: wing pattern differences between <i>Cercyonis silvestris</i> (W. H. Edwards, 1861) and <i>Cercyonis sthenele</i> (Boisduval, 1852) in the Sierra-Cascade ranges of California, USA; <i>Dione</i> (<i>Agraulis</i>) <i>galapagensis</i> (W. Holland, 1890) as a species-level taxon confirmed by genomic sequencing; female paratypes of recently described Hesperiidae species with strong sexual dimorphism not shown in the original descriptions; the third male of <i>Cecropterus</i> (<i>Thorybes</i>) <i>viridissimus</i> Grishin, 2023; all sequenced specimens of <i>Urbanus</i> (<i>Urbanus</i>) <i>viterboana</i> (Ehrmann, 1907) establishing this species as sister to <i>Urbanus</i> (<i>Urbanus</i>) <i>dubius</i> Steinhauser, 1981; <i>Windia sonoria</i> Grishin, 2025, and not <i>Windia windi</i> Freeman, 1969, recorded from the USA (Arizona, Cochise Co.); <i>Zopyrion</i> (<i>Zopyrion</i>) <i>xerxes</i> Grishin, 2025 as a species-level taxon confirmed by sequencing of additional specimens extending its range from Honduras to include El Salvador, Nicaragua, and Costa Rica; a second specimen of <i>Gorgythion cerrada</i> Grishin, 2025; a possible paralectotype of <i>Antigonus ruptifasciata</i> Plötz, 1884, also from Jamaica as deduced by genomic comparison; the holotype of <i>Pamphila philino</i> Möschler, 1879; a female of <i>Oligoria</i> (<i>Cobaloides</i>) <i>unica</i> (de Jong, 1983) from French Guiana; and a male of <i>Lychnuchus</i> (<i>Enosis</i>) <i>valle</i> Grishin, 2023 from Ecuador. We provide an updated list of the subtribe Euremina Grote, 1898 that comprises five genera (rather than one) and five additional subgenera. Finally, lectotypes are designated for eight taxa: <i>Helias haematospila</i> C. Felder & R. Felder, 1867 (Venezuela), <i>Tagiades editus</i> Plötz, 1885 (Indonesia: Maluku Province, Dobo), <i>Hesperia kolantus</i> Plötz, 1885 (India), <i>Hesperia daendeli</i> Plötz, 1885 (Indonesia: West Java, Jakarta), <i>Celaenorhinus</i> [sic] <i>lucifer</i> Hübner, [1829] (Suriname), <i>Cobalus pindar</i> Schaus, 1913 (Costa Rica: Limón), <i>Proteides hyas</i> Mabille, 1891 (Colom