Butterfly Classification and Species Discovery Using Genomics

Authors

Jing Zhang, University of Texas Southwestern Medical Center
Qian Cong, University of Texas Southwestern Medical Center
Jinhui Shen, University of Texas Southwestern Medical Center
Leina Song, University of Texas Southwestern Medical Center
Nick V. Grishin, University of Texas Southwestern Medical CenterFollow

Date of this Version

10-2-2023

Document Type

Article

Citation

The Taxonomic Report (2023) 11(3): 1-94

Comments

Copyright 2023, the International Lepidoptera Survey. Open access material

License: Creative Commons Attribution-ShareAlike-NonCommercial 4.0 International (CC BY-SA-NC 4.0 International)

Abstract

Genomic sequencing of worldwide butterfly fauna followed by phylogenetic analysis of protein-coding genes informs butterfly classification throughout the taxonomic hierarchy, from families to species. As a rule, we attribute the same taxonomic rank to more prominent clades of comparable divergence (that is, at the same level in the tree). For species delimitation, we use criteria based on relative genetic differentiation and the extent of gene exchange between populations. We analyze the current taxonomic classification of butterflies in the light of genomic phylogenies and encounter clades that correspond to yet unnamed taxa. As a result, 11 tribes, 33 subtribes, 2 genera, 11 subgenera, and 12 species are proposed as new. New tribes are: in family Nymphalidae Rafinesque, 1815: in subfamily Heliconiinae Swainson, 1822: Vindulini Grishin, trib. n. (type genus Vindula Hemming, 1934) and Algiini Grishin, trib. n. (type genus Algia Herrich-Schäffer, 1864), Lebadeini Grishin, trib. n. (type genus Lebadea C. Felder, 1861, in subfamily Limenitidinae Behr, 1864), and Amnosiini Grishin, trib. n. (type genus Amnosia E. Doubleday, 1849, in subfamily Pseudergolinae Jordan, 1898) and in family Lycaenidae [Leach], [1815]: in subfamily Aphnaeinae Distant, 1884: Cigaritini Grishin, trib. n. (type genus Cigaritis Donzel, 1848) and Axiocersini Grishin, trib. n. (type genus Axiocerses Hübner, [1819]) and in subfamily Theclinae: Drinini Grishin, trib. n. (type genus Drina Nicéville, 1890), Hypochrysopini Grishin, trib. n. (type genus Hypochrysops C. Felder & R. Felder, 1860), Jalmenini Grishin, trib. n. (type genus Jalmenus Hübner, 1818), Pseudalmenini Grishin, trib. n. (type genus Pseudalmenus H. H. Druce, 1902), and Rapalini Grishin, trib. n. (type genus Rapala F. Moore, 1881). New subtribes are: in Papilionidae Latreille, [1802], Meandrusina Grishin, subtrib. n. (type genus Meandrusa F. Moore, 1888, in Papilionini Latreille, [1802]); in Pieridae Swainson, 1820: Gandacina Grishin, subtrib. n. (type genus Gandaca F. Moore, 1906, in Coliadini Swainson, 1821), Hebomoiina Grishin, subtrib. n. (type genus Hebomoia Hübner, [1819], in Anthocharidini Scudder, 1889), and Pseudopierina Grishin, subtrib. n. (type genus Pseudopieris Godman & Salvin, 1890, in Dismorphiini Schatz, 1886); in Nymphalidae: Lachnopterina Grishin, subtrib. n. (type genus Lachnoptera E. Doubleday, 1847, in Algiini Grishin, trib. n.), in Vagrantini Pinratana & Eliot, 1996: Terinosina Grishin, subtrib. n. (type genus Terinos Boisduval, 1836) and Smerinina Grishin, subtrib. n. (type genus Smerina Hewitson, 1874), in Adoliadini Doubleday, 1845: Evenaina Grishin, subtrib. n. (type genus Evena Westwood, 1850) and Pseudathymina Grishin, subtrib. n. (type genus Pseudathyma Staudinger, 1891), and Kumothalina Grishin, subtrib. n. (type genus Kumothales Overlaet, 1940, in Cymothoini Dhungel & Wahlberg, 2018); in Riodinidae Grote, 1895 (1827): Teratophthalmina Grishin, subtrib. n. (type genus Teratophthalma Stichel, 1909, in Mesosemiini Bates, 1859), Argyrogrammanina Grishin, subtrib. n. (type genus Argyrogrammana Strand, 1932, in Symmachiini Reuter, 1896), in the tribe Calydnini Seraphim, Freitas & Kaminski, 2018: Echenaidina Grishin, subtrib. n. (type genus Echenais Hübner, [1819]) and Echydnina Grishin, subtrib. n. (type genus Echydna J. Hall, 2002), and Cariina Grishin, subtrib. n. (type genus Caria Hübner, 1823, in Riodinini Grote, 1895 (1827); in Lycaenidae: Megalopalpina Grishin, subtrib. n. (type genus Megalopalpus Röber, 1886, in Miletini Reuter, 1896), Pseudaletidina Grishin, subtrib. n. (type genus Pseudaletis H. H. Druce, 1888, in Cigaritini Grishin, trib. n.), in Aphnaeini Distant, 1884: Aloeidina Grishin, subtrib. n. (type genus Aloeides Hübner, [1819]) and Phasisina Grishin, subtrib. n. (type genus Phasis Hübner, [1819]), Pilodeudorigina Grishin, subtrib. n. (type genus Pilodeudorix H. H. Druce, 1891, in Rapalini Grishin, trib. n.), Hemiolaina Grishin, subtrib. n. (type genus Hemiolaus Aurivillius, 1922, in Oxylidini Eliot, 1973), Cupidopsina Grishin, subtrib. n. (type genus Cupidopsis Karsch, 1895, in Hypotheclini Eliot, 1973), and in tribe Polyommatini Swainson, 1827: Theclinesthina Grishin, subtrib. n. (type genus Theclinesthes Röber, 1891), Azanina Grishin, subtrib. n. (type genus Azanus F. Moore, 1881), Unina Grishin, subtrib. n. (type genus Una Nicéville, 1890), Ionolycina Grishin, subtrib. n. (type genus Ionolyce Toxopeus, 1929), Pithecopina Grishin, subtrib. n. (type genus Pithecops Horsfield, 1828), Zizulina Grishin, subtrib. n. (type genus Zizula Chapman, 1910), Jamidina Grishin, subtrib. n. (type genus Jamides Hübner, [1819]), Fameganina Grishin, subtrib. n. (type genus Famegana Eliot, 1973), Oboroniina Grishin, subtrib. n. (type genus Oboronia Karsch, 1893), and Uranothaumatina Grishin, subtrib. n. (type genus Uranothauma Butler, 1895); and in Hesperiidae Latreille, 1809, Cupithina Grishin, subtrib. n. (type genus Cupitha F. Moore, 1884, in Astictopterini Swinhoe, 1912). The new genera are Balenga Grishin, gen. n. (type 2 species Proteides balenge Holland, 1891, in Gretnini Grishin, 2019) and Tulsia Grishin, gen. n. (type species Parnara tulsi Nicéville, 1884, in Baorini Doherty, 1886). New subgenera are: in Pieridae, Lirinia Grishin, subgen. n. (type species Terias lirina H. Bates, 1861, in genus Pyrisitia A. Butler, 1870); in Nymphalidae: Hyperanartia Grishin, subgen. n. (type species Vanessa dione Latreille, [1813], in genus Hypanartia Hübner, [1821]) and Paranartia Grishin, subgen. n. (type species Hypanartia hippomene Hübner, [1823], in genus Vanessa [Fabricius], 1807); in Lycaenidae, Auricirrus Grishin, subgen. n. (type species Papilio thysbe Linnaeus, 1764, in genus Chrysoritis Butler, 1898); and in Hesperiidae: Isocleros Grishin, subgen. n. (type species Pamphila (?) mackenii Trimen, 1868, in genus Acleros Mabille, 1885), Mesna Grishin, subgen. n. (type species Parnara leucophaea Holland, 1894, in genus Fresna Evans, 1937), Lippina Grishin, subgen. n. (type species Carystus telesinus Mabille, 1878, in genus Xanthoneura Eliot, 1978), Ganda Grishin, subgen. n. (type species Zophopetes ganda Evans, 1937, in genus Leona Evans, 1937), Zarida Grishin, subgen. n. (type species Hesperia lacida Hewitson, 1876, in genus Gretna Evans, 1937), and in genus Gegenes Hübner, 1819: Flanga Grishin, subgen. n. (type species Parnara perobscura H. H. Druce, 1912), and Havea Grishin, subgen. n. (type species Hesperia havei Boisduval, 1833). New species are: in Nymphalidae: Microtia elvira Grishin, sp. n. (type locality in the USA: Arizona, Pima/Santa Cruz Counties) and Cyllopsis brocki sp. n. (type locality in Mexico: Sonora, Yécora), in Riodinidae, Argyrogrammana astuta Grishin, sp. n. (type locality in Peru: Madre de Dios), in Hesperiidae: Cecropterus (Thorybes) rockiensis Grishin, sp. n. (type locality in USA: Colorado, Jefferson County), Cecropterus (Thorybes) floridianus Grishin, sp. n. (type locality in USA: Florida, Volusia County), Cecropterus (Thorybes) oaxacensis Grishin, sp. n. (type locality in Mexico: Oaxaca), Nascus (Bron) lux Grishin, sp. n. (type locality in Brazil: Amapá), Cogia chiagua Grishin, sp. n. (type locality in Guatemala), Celotes sabinus Grishin, sp. n. (type locality in USA: Arizona, Pima County), Acleros togo Grishin, sp. n. (type locality in Togo), Ceratrichia notata Grishin, sp. n. (type locality in Central African Republic), and Semalea malawi Grishin, sp. n. (type locality in Malawi). Furthermore, we elevate a tribe to subfamily, resurrect 3 tribes, 3 subtribes, 5 genera (and confirm 1), 3 subgenera, change the rank of 5 currently recognized tribes to subtribes (and confirm 1), 10 genera to subgenera, synonymize 4 genera, and present evidence to support 21 taxa as species instead of subspecies and 3 taxa as subspecies instead of synonyms. Namely, we reinstate Liphyrinae Doherty, 1889 as a subfamily (was a tribe of Miletinae Reuter, 1896), treat the following as tribes: Leptidiini Grote,1897 (in Dismorphiinae Schatz, 1886) and in Theclinae Swainson, 1830: Surendrini Koçak & Seven, 1997 (not a subtribe of Arhopalini Bingham, 1907) and Myrinini Toxopeus, 1929 (not a synonym of Amblypodiini Doherty, 1886), and subtribes: Callidryina Kirby, 1896 and Gonepterygina Verity, 1920 (in Coliadini Swainson, 1821), Abrotina Hemming, 1960 and Bebeariina Hemming, 1960 (in Adoliadini Doubleday, 1845), and Sarotina Bridges, 1988 (in Helicopini Stichel, 1928). We confirm Libytheana Michener, 1943 as a valid genus, not a junior subjective synonym of Prolibythea Scudder, 1889 and resurrect from synonymy the following genera: Pseudanaphaeis Bernardi, 1953 (not Belenois Hübner, [1819]), Charmion Nicéville, 1894 (not Celaenorrhinus Hübner, [1819]), Sape Mabille, 1891 (not Sarangesa F. Moore, [1881]), Milena Evans, 1912 (not Caltoris Swinhoe, 1893) and subgenera: Neofieldia Özdikmen, 2008 and Bassaris Hübner, [1821] of Vanessa [Fabricius], 1807 and Spindasis Wallengren, 1857 of Cigaritis Donzel, 1848. We change the status of the following taxa from tribes to subtribes: Nathalina Bálint, 2022 and Kricogonina Bálint, 2022 of Euremini Grote, 1898, Tarakina Eliot, 1973 of Spalgini, 1929, Horagina Swinhoe, 1910 and Loxurina Swinhoe, 1910 of Cheritrini Swinhoe, 1910, and Niphandina Sibatani & Ito, 1942 of Polyommatini; from genus to subgenus: Leucidia E. Doubleday, 1847 of Abaeis Hübner, [1819], Cesa Seven, 1997 of Crudaria Wallengren, 1875, Paralycaeides Nabokov, 1945 of Itylos Draudt, 1921, Eldoradina Balletto, 1993 of Nabokovia Hemming, 1960, Pyrrhochalcia Mabille, 1904 of Coeliades Hübner, 1818, Paracleros Berger, 1978 of Acleros Mabille, 1885, Mopala Evans, 1937 of Leona Evans, 1937, Afrogegenes Jong & Coutsis, 2017 and Torbenlarsenia Kemal & Koçak, 2020 of Gegenes Hübner, 1819, and Zenonoida Fan & Chiba, 2016 of Zenonia Evans, 1935; and from genus to junior subjective synonym: Vansomerenia Heath, 1997 of Chloroselas Butler, 1886, Ceratricula Larsen, 2013 of Paronymus Aurivillius, 1925, Xanthodisca Aurivillius, 1925 of Semalea Holland, 1896, Perrotia Oberthür, 1916 of Galerga Mabille, 1898. The following taxa are species, not subspecies or synonyms: Pyrisitia mayobanex (M. Bates, 1939), stat. nov. and Pyrisitia memulus (A. Butler, 1871), stat. rest. (not Pyrisitia dina (Poey, 1832)), Abaeis gratiosa (E. Doubleday, 1847), stat. rest. and Abaeis angulata (Wallengren, 1860), stat. rest. (not Abaeis arbela (Geyer, 1832)), Teriocolias doris (Röber, 1909), stat. rest. (not Teriocolias deva (E. Doubleday, 1847)), Vanessa madegassorum (Aurivillius, 1899), stat. nov. (not Vanessa hippomene (Hübner, 1823)), Eresia (Anthanassa) seminole Skinner, 1911, stat. rev. (not Eresia (Anthanassa) texana (W. H. Edwards, 1863)), Cecropterus (Thorybes) albosuffusa (H. Freeman, 1943), stat. nov. and Cecropterus (Thorybes) indistinctus (Austin & J. Emmel, 1998), stat. nov. (not Cecropterus (Thorybes) pylades (Scudder, 1870)), Cogia hiska Evans, 1953, stat. nov. (not Cogia hippalus (W. H. Edwards, 1882)), Cogia moschus (W. H. Edwards, 1882), stat. rest. (not Cogia caicus (Herrich-Schäffer, 1869)), Pholisora albicirrus Glassberg 2023, stat. nov. (not Pholisora catullus (Fabricius, 1793)), Acleros (Isocleros) instabilis Mabille, 1889, stat. rest. and Acleros (Isocleros) olaus (Plötz, 1884), stat. rest. (not Acleros (Isocleros) mackenii (Trimen, 1868)), Fresna (Mesna) bassa (Lindsey & L. Miller, 1965), stat. nov., comb. nov. (not Meza leucophaea (Holland, 1894)), Paronymus volta (L. Miller, 1971), stat. nov., comb. nov. (not Meza cybeutes (Holland, 1894)), Paronymus indeterminabilis (Strand, 1912), stat. rest., comb. nov. and Paronymus congdoni (Larsen, 2013), stat. nov., comb. nov. (not Ceratricula semilutea (Mabille, 1891)), Semalea corvinus (Mabille, 1890), stat. rest. (not Semalea sextilis (Plötz, 1886)), Xanthoneura patmapana (Fruhstorfer, 1911), stat. nov. (not Xanthoneura corissa (Hewitson, 1876)), Gretna capra Evans, 1937, stat. nov. (not Gretna carmen Evans, 1937), and Lerodea dysaules Godman, 1900, stat. rest. (not a synonym of Lerodea arabus (W. H. Edwards, 1882)), while Vernia verna sequoyah (H. Freeman, 1942), stat. rest. (not a synonym of Vernia verna W. H. Edwards, 1862) and Euphyes vestris osceola (Lintner, 1878), stat. rev. (not a synonym of Euphyes vestris vestris (Boisduval, 1852)) are subspecies. In addition, we propose new genus-species combinations: Pyrisitia amelia (Poey, [1852]), Pyrisitia lirina (H. Bates, 1861), Abaeis paulina (H. Bates, 3 1861), Abaeis xantochlora (Kollar, 1850), Abaeis fabiola (C. Felder & R. Felder, 1861), Abaeis tupuntenem (Lichy, 1976), and Abaeis adamsi (Lathy, 1898) (not Eurema Hübner, [1819]), Shijimia potanini (Alphéraky, 1889) (not Tongeia Tutt, 1908), Fresna (Mesna) larea (Neave, 1910), Fresna (Mesna) leucophaea (Holland, 1894), Fresna (Mesna) mabea (Holland, 1894), Paronymus banda (Evans, 1937), Paronymus cybeutes (Holland, 1894), Paronymus elba (Evans, 1937), Paronymus gardineri (Collins & Larsen, 2008), Paronymus indusiate (Mabille, 1891), and Paronymus mabillei (Holland, 1893) (not Meza Hemming, 1939), Paronymus punctata (Holland, 1896) (not Ceratrichia Butler, 1870), Galerga ariel (Mabille, 1878) (not Xanthodisca Aurivillius, 1925), Isoteinon anomoeus (Plötz, 1879), Isoteinon bruno (Evans, 1937), Isoteinon inornatus (Trimen, 1864), and Isoteinon punctulata (A. Butler, 1895) (not Astictopterus C. Felder & R. Felder, 1860), Borbo gemella (Mabille, 1884) (not Torbenlarsenia Kemal & Koçak, 2020), and Gegenes (Torbenlarsenia) cottrelli (Larsen, 2013), Gegenes (Torbenlarsenia) fallax (Gaede, 1916), Gegenes (Torbenlarsenia) fanta (Evans, 1937), Gegenes (Torbenlarsenia) micans (Holland, 1896), and Gegenes (Torbenlarsenia) sirena (Evans, 1937) (not Borbo Evans, 1949); and a new species-subspecies combination Cogia hiska hester Evans, 1953 (not Cogia hippalus). Acleros nyassicola Strand, 1921 is a junior subjective synonym of Acleros (Isocleros) olaus (Plötz, 1884), not of Acleros (Isocleros) mackenii (Trimen, 1868). We transferred the tribes Oxylidini Eliot, 1973, Remelanini Eliot, 1973, and Hypolycaenini Swinhoe, 1910 from Theclinae Swainson, 1831 to Polyommatinae Swainson, 1827. We conclude that [No genus] osibius Draudt, 1924 is an unavailable name. The lectotype is designated for Eudamus caicus Herrich-Schäffer, 1869 (type locality likely in Mexico: Oaxaca, as deduced by sequence comparison), and the neotype is designated for Telemiades solon Plötz, 1882 (type locality becomes Brazil: Bahia). The type localities of Ceratrichia punctata Holland, 1896 and Osmodes staudingeri Holland, 1896 are Sierra Leone: Freetown and Cameroon: Efoulan, respectively, as determined from the locality labels of primary type specimens. Finally, we provide taxonomic lists for Euremini (to worldwide subgenera and American species) and Lycaenidae (to subtribes).