Lycaenidae

Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina & Grishin, Nick V., 2023, Butterfly classification and species discovery using genomics, The Taxonomic Report of the International Lepidoptera Survey 11 (3), pp. 1-94 : 46-48

Family Lycaenidae [Leach], [1815] ( Lycaena [Fabricius], 1807)

Subfamily Curetinae Distant, 1884 ( Curetis Hübner, [1819] )

Subfamily Liphyrinae Doherty, 1889 ( Liphyra Westwood, [1864] ), stat. rest.

Subfamily Miletinae Reuter, 1896 ( Miletus Hübner, [1819] )

Tribe Lachnocnemini Clench, 1955 ( Lachnocnema Trimen, 1887 )

Tribe Miletini Reuter, 1896 ( Miletus Hübner, [1819] )

Subtribe Miletina Reuter, 1896 ( Miletus Hübner, [1819] )

Subtribe Megalopalpina Grishin, subtrib. n. ( Megalopalpus Röber, 1886 )

Tribe Spalgini Toxopeus, 1929 ( Spalgis F. Moore, 1879 )

Subtribe Spalgina Toxopeus, 1929 ( Spalgis F. Moore, 1879 )

Subtribe Tarakina Eliot, 1973 , stat. rev. ( Taraka Doherty, 1889 )

Tribe Liptenini Röber, 1892 ( Liptena Westwood, [1851] )

Subtribe Durbaniina Clench, 1955 ( Durbania Trimen, 1862 )

Subtribe Pentilina Aurivillius, 1914 ( Pentila Westwood, [1851] )

Subtribe Liptenina Röber, 1892 ( Liptena Westwood, [1851] ); includes Mimacraeina Stempffer, 1961 Subtribe Iridanina Clench, 1965 ( Iridana Aurivillius, 1920 )

Subtribe Epitolina Jackson, 1962 ( Epitola Westwood, [1851] )

Subtribe Cooksoniina Sáfián, Boyle & Pierce, 2023 ( Cooksonia H. H. Druce, 1905 )

Tribe Poritiini Doherty, 1886 ( Poritia F. Moore, [1866] )

Subfamily Aphnaeinae Distant, 1884 ( Aphnaeus Hübner, [1819] ); not monophyletic with Theclinae ! Tribe Axiocersini Grishin, trib. n. ( Axiocerses Hübner, [1819] ) Tribe Cigaritini Grishin, trib. n. ( Cigaritis Donzel, 1848 ) Subtribe Pseudaletidina Grishin, subtrib. n. ( Pseudaletis H. H. Druce, 1888 ) Subtribe Cigaritina Grishin ( Cigaritis Donzel, 1848 ) Tribe Aphnaeini Distant, 1884 ( Aphnaeus Hübner, [1819] ) Subtribe Aloeidina Grishin, subtrib. n. ( Aloeides Hübner, [1819] ) Subtribe Phasisina Grishin, subtrib. n. ( Phasis Hübner, [1819] ) Subtribe Aphnaeina Distant, 1884 ( Aphnaeus Hübner, [1819] )

Subfamily Lycaeninae [Leach], [1815] ( Lycaena [Fabricius], 1807)

Subfamily Theclinae Swainson, 1830 ( Thecla [Fabricius], 1807)

Tribe Theclini Swainson, 1830 ( Thecla [Fabricius], 1807)

Tribe Surendrini Koçak & Seven, 1997 , stat. nov. ( Surendra F. Moore, 1879 )

Tribe Arhopalini Bingham, 1907 ( Arhopala Boisduval, 1832 )

Tribe Drinini Grishin, trib. n. ( Drina Nicéville, 1890 )

Tribe Candalidini Eliot, 1973 ( Candalides Hübner, [1819] )

Tribe Hypochrysopini Grishin, trib. n. ( Hypochrysops C. Felder & R. Felder, 1860 )

Tribe Luciini Waterhouse & Lyell, 1914 ( Lucia W. Swainson, 1833 )

Tribe Ogyrini Waterhouse & Lyell, 1914 ( Ogyris Angas, 1847 )

Tribe Jalmenini Grishin, trib. n. ( Jalmenus Hübner, 1818 )

Tribe Pseudalmenini Grishin, trib. n. ( Pseudalmenus H. H. Druce, 1902 )

Tribe Amblypodiini Doherty, 1886 ( Amblypodia Horsfield, 1829 )

Tribe Myrinini Toxopeus, 1929 , stat. nov. ( Myrina [Fabricius], 1807)

Tribe Cheritrini Swinhoe, 1910 ( Cheritra F. Moore, 1881 )

Subtribe Cheritrina Swinhoe, 1910 ( Cheritra F. Moore, 1881 )

Subtribe Horagina Swinhoe, 1910 , stat. nov. ( Horaga F. Moore, 1881 )

Subtribe Loxurina Swinhoe, 1910 , stat. nov. ( Loxura Horsfield, [1829] )

Tribe Iolaini Riley, 1958 ( Iolaus Hübner, [1819] )

Tribe Catapaecilmatini Eliot, 1973 ( Catapaecilma Butler, 1879 )

Tribe Zesiusini Swinhoe, 1912 ( Zesius Hübner, [1819] )

Tribe Tomarini Eliot, 1973 ( Tomares Rambur, 1840 )

Tribe Rapalini Grishin, trib. n. ( Rapala F. Moore, 1881 )

Subtribe Rapalina Grishin ( Rapala F. Moore, 1881 )

Subtribe Pilodeudorigina Grishin, subtrib. n. ( Pilodeudorix H. H. Druce, 1891 )

Tribe Deudorigini Doherty, 1886 ( Deudorix Hewitson, 1863 )

Tribe Eumaeini Doubleday, 1847 ( Eumaeus Hübner, [1819] )

Subtribe Eumaeina Doubleday, 1847 ( Eumaeus Hübner, [1819] )

Subtribe Rhammina Prieto & Busby, 2022 ( Rhamma K. Johnson, 1992 )

Subtribe Timaetina Busby & Prieto, 2022 ( Timaeta K. Johnson, Kruse & Kroenlein, 1997 )

Subtribe Atlidina Martins & Duarte, 2022 ( Atlides Hübner, [1819] )

Subtribe Evenina Faynel & Grishin, 2022 ( Evenus Hübner, [1819] )

Subtribe Jantheclina Robbins & Faynel, 2022 ( Janthecla Robbins & Venables, 1991 )

Subtribe Paiwarriina Lamas & Robbins,2022 ( Paiwarria Kaye, 1904 )

Subtribe Cupatheclina Lamas & Grishin, 2022 ( Cupathecla Bálint, 2005 )

Subtribe Parrhasiina Busby & Robbins, 2022 ( Parrhasius Hübner, [1819] )

Subtribe Ipideclina Martins & Grishin, 2022 ( Ipidecla Dyar, 1916 )

Subtribe Calycopidina Duarte & Robbins, 2010 ( Calycopis Scudder, 1876 )

Subtribe Strymonina Tutt, 1907 ( Strymon Hübner, 1818 )

Subtribe Strephonotina K. Johnson, Austin, Le Crom & Salazar, 1997 ( Strephonota K. Johnson et al., 1997 ) Subtribe Trichonidina Duarte & Faynel, 2022 ( Trichonis Hewitson, 1865 )

Subfamily Polyommatinae Swainson, 1827 ( Polyommatus Latreille, 1804 ) Tribe Oxylidini Eliot, 1973 ( Oxylides Hübner, [1819] ) Subtribe Oxylidina Eliot, 1973 ( Oxylides Hübner, [1819] ) Subtribe Hemiolaina Grishin, subtrib. n. ( Hemiolaus Aurivillius, 1922 ) Tribe Remelanini Eliot, 1973 ( Remelana F. Moore, 1884 ) Tribe Hypolycaenini Swinhoe, 1910 ( Hypolycaena C. Felder & R. Felder, 1862 ) Tribe Lycaenesthini Toxopeus, 1929 ( Lycaenesthes F. Moore, 1866 ) Tribe Hypotheclini Eliot, 1973 ( Hypothecla G. Semper, 1890 ) Subtribe Hypotheclina Eliot, 1973 ( Hypothecla G. Semper, 1890 ) Subtribe Cupidopsina Grishin, subtrib. n. ( Cupidopsis Karsch, 1895 ) Tribe Polyommatini Swainson, 1827 ( Polyommatus Latreille, 1804 ) Subtribe Niphandina Sibatani & Ito, 1942 , stat. conf. ( Niphanda F. Moore, 1875 ) Subtribe Theclinesthina Grishin, subtrib. n. ( Theclinesthes Röber, 1891 ) Subtribe Azanina Grishin, subtrib. n. ( Azanus F. Moore, 1881 ) Subtribe Unina Grishin, subtrib. n. ( Una Nicéville, 1890 ) Subtribe Danina Koçak & Seven, 1997 ( Danis [Fabricius], 1807) Subtribe Ionolycina Grishin, subtrib. n. ( Ionolyce Toxopeus, 1929 ) Subtribe Pithecopina Grishin, subtrib. n. ( Pithecops Horsfield, 1828 ) Subtribe Zizulina Grishin, subtrib. n. ( Zizula Chapman, 1910 ) Subtribe Brephidiina Stempffer, 1957 ( Brephidium Scudder, 1876 ) Subtribe Celastrinina Tutt, 1907 ( Celastrina Tutt, 1906 ) Subtribe Jamidina Grishin, subtrib. n. ( Jamides Hübner, [1819] ) Subtribe Zizeeriina Chapman, 1910 ( Zizeeria Chapman, 1910 ) Subtribe Fameganina Grishin, subtrib. n. ( Famegana Eliot, 1973 ) Subtribe Catochrysopina Toxopeus, 1929 ( Catochrysops Boisduval, 1832 ) Subtribe Oboroniina Grishin, subtrib. n. ( Oboronia Karsch, 1893 ) Subtribe Castaliina Distant, 1884 ( Castalius Hübner, [1819] ) Subtribe Scolitantidina Tutt, 1907 ( Scolitantides Hübner, [1819] ) Subtribe Uranothaumatina Grishin, subtrib. n. ( Uranothauma Butler, 1895 ) Subtribe Actizerina Koçak, 1996 ( Actizera Chapman, 1910 ) Subtribe Leptotina Wagener, 1995 ( Leptotes Scudder, 1876 ) Subtribe Lampidina Tutt, 1907 ( Lampides Hübner, [1819] ) Subtribe Everina Tutt, 1907 ( Everes Hübner, [1819] ) Subtribe Polyommatina Swainson, 1827 ( Polyommatus Latreille, 1804 )

Our classification of Polyommatini Swainson, 1827 compared to that by Stradomsky (2016)

In his pioneering work based on limited DNA sequence data and careful genitalic comparison, Stradomsky (2016) proposed to partition the tribe Polyommatini into 22 subtribes. Stradomsky’s classification fares well compared to our genomic results (see above), and his study is impressive in its insight. Out of our 23 subtribes, 20 match Stradomsky’s in names, and one, Celastrinina Tutt, 1907 , was referred to by its junior subjective synonym, Lycaenopsina Swinhoe, 1910. The remaining two subtribes, Ionolycina subtrib. n. and Unina subtrib. n., were placed by Stradomsky into his Danina Koçak & Seven, 1997 and “ Azanina ,” respectively. Una Nicéville, 1890 cannot belong to “ Azanina ,” because it is in the clade with Danis [Fabricius], 1807 and not with Azanus F. Moore, 1881 (see Fig. 22 and Kawahara et al. (2023)). While it is indeed possible to unify these four groups into a single subtribe, Danina , genetic differentiation within this subtribe would be larger than in other subtribes in Polyommatini ( Fig. 22). Therefore, dividing the group into four subtribes brings the classification in line with the genetic differentiation within other Polyommatini subtribes. Finally, Stradomsky’s “Cacyreina” is merged into our Lampidina Tutt, 1907 due to genetic similarities, and Stradomsky’s phylogeny based on a small number of gene markers is poorly supported around these taxa. We put names of subtribes used by Stradomsky in quotes because they were proposed somewhat informally: authorship of previously published names was not indicated, and for the names we could not find in previous publications, Stradomsky did not explicitly indicate that they were intentionally new (fails ICZN Art. 16.1.) and did not specify what their type genera were (fails ICZN Art. 16.2.). Moreover, it remains unclear from the text of intended to propose new ICZN-compliant names in his work.