Aphnaeinae Distant, 1884

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 : 34-46

Tribes in Aphnaeinae Distant, 1884

Inspection of the genomic tree reveals three strongly supported clades in the subfamily Aphnaeinae Distant, 1884 that we define as tribes ( Figs. 20, 21): nominotypical, “Apharitini” Chou et al., 1994, and the third one that does not have a name and is described below. The name “Apharitini” was published in Chou et al. (1994) in an informal way without description, definition, or references to them (fails ICZN Art. 13.) and is a nomen nudum. The name for this tribe that satisfies the requirements of the ICZN Code is proposed below. We define three tribes and not two because the phylogenetic affinities of the unnamed tribes with the nominotypical one are not particularly strong (86%), and combining their clades may not result in a monophyletic taxon if this topology is incorrect. The three clades are the same in the recently published large-scale tree (Kawahara et al. 2023), but the topology is different in a tree based on a more limited set of gene markers (but a larger set of taxa) (Boyle et al. 2015).

http://zoobank.org/ E44D9831-7137-4120-86CB-700DBCC8D752

Type genus. Cigaritis Donzel, 1848 .

Definition. The clade with Cigaritis (type species Cigaritis zohra Donzel, 1848 ) and related genera is genetically differentiated from other clades and is at the tree level of a tribe ( Figs. 20, 21). This new tribe is morphologically diverse and can be distinguished from other members of the subfamily Aphnaeinae Distant, 1884 by the shape of uncus, which is highly variable, but nevertheless bilobed or divided, with centrally concave distal margin (see Stempffer (1967) for illustrations and more detailed descriptions): in Lipaphnaeus Aurivillius, 1916 , subtriangular uncus is deeply divided with narrow, tooth-like pointed lobes; in Chloroselas Butler, 1886 , the lobes are small and subtriangular uncus is with a central notch separating finely serrated lobes; in Cigaritis Donzel, 1848 , uncus is broader than long, nearly rectangular or trapezoidal in dorsal view (sometimes nearly vestigial), lobes widely separated, uncus mostly concave (may be with irregularities) between the lobes; in Chrysoritis Butler, 1898 and Crudaria Wallengren, 1875 , uncus is similar, trapezoidal or heart-shaped, but less broad, with rounded lobes and convex margin between them; in Pseudaletis H. H. Druce, 1888 , uncus is divided for nearly its entire length with finger-like terminally rounded lobes. In other Aphnaeinae tribes, uncus in undivided and not strongly bilobed: typically, from almost square to broadly rectangular, with nearly flat (with some irregularities) distal margin, or dome-shaped with convex distal margin, which is only slightly concave in Trimenia Tite & Dickson, 1973 (type species Zeritis wallengrenii Trimen, 1887 ), thus resembling Chrysoritis , but uncus is narrower and without clearly defined lobes expanded laterally (as in Chrysoritis and Crudaria ). Furthermore, Trimenia has saccus, which is not developed in Chrysoritis . A combination of the following nuclear genomic base pairs is diagnostic: cce332.12.5:A61T, cce1232.20.1:A596G, cce1351.40.3:A349T, cce5072.9.1:A67C, cce2368.14.7:G95T.

Genera included. The type genus (i.e., Cigaritis Donzel, 1848 ), Chloroselas Butler, 1886 , Chrysoritis Butler, 1898 , Crudaria Wallengren, 1875 , Lipaphnaeus Aurivillius, 1916 , and Pseudaletis H. H. Druce, 1888 , including their subgenera and synonyms (e.g., Cesa Seven, 1997, Vansomerenia Heath, 1997 , and Apharitis Riley, 1925 ).

Parent Taxon. Subfamily Aphnaeinae Distant, 1884 .

Comment. This tribe corresponds to “Apharitini” of Chou et al. (1994), published without description, definition, or references to them: a nomen nudum (fails ICZN Art. 13.). Apharitis Riley, 1925 (type species Polyommatus epargyros Eversmann, 1855 ) is a junior subjective synonym of Cigaritis .

Pseudaletidina Grishin, new subtribe

http://zoobank.org/ D8A065B1-F0CF-4A7F-BADA-F06FDEECA919

Type genus. Pseudaletis H. H. Druce, 1888 .

Definition. Pseudaletis (type species Pseudaletis agrippina H. H. Druce, 1888 ) is in the lineage that is sister to all other Cigaritini Grishin, trib. n. and is genetically differentiated from them at the subtribal level ( Figs. 20, 21). Therefore, we propose to treat this lineage as a subtribe. This new subtribe is diagnosed by a combination of the following characters, as given for the Pseudaletis section by Eliot (1973): palpi short—much less than half of the head length—covered in appressed scales, proboscis very short (but functional), forewing appears disproportionately large comparatively to hindwing; uncus divided for nearly its entire length with finger-like terminally rounded lobes, falces rudimentary, pointed processes inflexibly fused to tegumen; female abdomen with a prominent tuft of specialized scales, which are spoon-shaped with long “handles”. A combination of the following nuclear genomic base pairs is diagnostic: cce1853.23.8:T1384A, cce133.4.1:A287G, cce4260.4.1:G34A, cce 1367.9.3:G1252A, cce17940.6.2:C71A.

Genera included. Only the type genus.

Parent Taxon. Tribe Cigaritini Grishin, trib. n.

http://zoobank.org/ 1BBFE9EC-AB8B-4709-A250-4DA082ED2B4E

Type genus. Axiocerses Hübner, [1819] .

Definition. Axiocerses (type species Papilio perion Stoll, 1782 , which is a junior subjective synonym of Papilio harpax Fabricius, 1775 ) and Zeritis Boisduval, 1836 (type species Zeritis neriene Boisduval, 1836 ) are confident sisters in the genomic tree ( Figs. 20, 21), consistently with similarities in their morphology reported previously ( Stempffer 1967). The two genera form a clade that originates early in the radiation of the subfamily Aphnaeinae Distant, 1884 , and, therefore, it corresponds to a tribe. This new tribe is distinguished from the relatives by a combination of the following characters: uncus very broad and short, distal margin straight or convex, lobes nearly triangular or rounded, falces thick at the base, then strongly angled with free branch long and slender, ventral side with apophysis, tegumen with uncus hood-shaped, tegumen with convex anterior margin; palpi short, not extending or slightly extending beyond the fronts, 2 nd segment of palpi with long scales and hairs; tarsus unsegmented, with spines below; forewing with only 10 veins ( Stempffer 1967; Henning and Henning 1996). The similarity in uncus, falces, and tegumen unifies Axiocerses and Zeritis Boisduval, 1836 ( Stempffer 1967). A combination of the following nuclear genomic base pairs is diagnostic: cce243.9.9:A1417C, cce15587.11.3:G103A, cce127.4.3:A149T, cce980.22.7:C902T, cce 2423.1.2:T2315C.

Genera included. The type genus (i.e., Axiocerses Hübner, [1819] ) and Zeritis Boisduval, 1836 .

Parent Taxon. Subfamily Aphnaeinae Distant, 1884 .

Aloeidina Grishin, new subtribe

http://zoobank.org/ A32E1EE2-A06D-4100-AFD9-3214F38A2C93

Type genus. Aloeides Hübner, [1819] .

Definition. Within Aphnaeini Distant, 1884 , the clade of several genera that includes Aloeides (type species Papilio pierus Cramer, 1779 ) corresponds to the subtribal level in the tree ( Figs. 20, 21). This new subtribe is distinguished from its relatives by a combination of these characters, as discussed and illustrated by Stempffer (1967), Tite & Dickson (1973), and Eliot (1973): foreleg and midleg tibiae with apical spurs, palpi smooth, with equal length scales (in several species with some scattered long ribbonshaped blunt scales), or with uniquely long and bristly white scales (in Erikssonia Trimen, 1891 ), forewing vein R 4+5 originates at or beyond the junction of the discocellular vein and vein M 1, vein R 2 originates next to the origin of vein R 4+5. A combination of the following nuclear genomic base pairs is diagnostic: cce5760.10.2:C484T, cce2790.13.2:T63C, cce2790.13.2:A183G, cce 1354.3.7:A61T, cce 1354.3.7: T1903A.

Genera included. The type genus (i.e., Aloeides Hübner, [1819] ), Argyraspodes Tite & Dickson, 1973 , Erikssonia Trimen, 1891 , and Trimenia Tite & Dickson, 1973 .

Parent Taxon. Tribe Aphnaeini Distant, 1884 .

Phasisina Grishin, new subtribe

http://zoobank.org/ B6D8239D-DAF0-4865-9F3E-18677ABDEA6F

Type genus. Phasis Hübner, [1819] .

Definition. Within Aphnaeini Distant, 1884 , the clade of several genera that includes Phasis (type species Papilio salmoneus Stoll, 1781 , which is a junior subjective synonym of Papilio thero Linnaeus, 1764 ) corresponds to the subtribal level in the tree ( Figs. 20, 21). This new subtribe is distinguished from its relatives by a combination of the following characters, as discussed and illustrated by Tite & Dickson (1973): tibia of all legs without apical spurs, forewing vein M 2 originates much closer to the origin of vein longer than 16 mm. A combination of the following nuclear genomic base pairs is diagnostic: cce288.3.2: A79C, cce33.3.3:A234G, cce 2859.6.1:T729C, cce511.6.1:A54G, cce14215.5.1:T850C, cce33280.1.7:T944T (not A), cce3467.4.4:A374A (not G), cce7428.5.1:G156G (not A), cce4435.12.1:C616C (not A), cce 1239.1. 3:T235T (not G).

Genera included. The type genus (i.e., Phasis Hübner, [1819] ) and Tylopaedia Tite & Dickson, 1973 .

Parent Taxon. Tribe Aphnaeini Distant, 1884 .

Surendrini Koçak & Seven, 1997 is a tribe

Currently placed in the tribe Arhopalini Bingham, 1907 , Surendrina Koçak & Seven, 1997 is not monophyletic with it and instead is sister to Theclini Swainson, 1830 ( Fig. 20). It is conceivable to place Surendrina in Theclini , but the two groups are more distinct from each other than subtribes and are at the tree level of tribes. Therefore, we propose to treat the former taxon as a tribe Surendrini Koçak & Seven, 1997 , stat. nov.

Drinini Grishin, new tribe

http://zoobank.org/ 2001E28A-4C8C-45D9-B228-E34D99B26E85

Type genus. Drina Nicéville, 1890 .

Definition. Currently in Loxurini Swinhoe, 1910 , Drina (type species Myrina donina Hewitson, 1865 ) is not monophyletic with it and is not closely related to that tribe, instead forming a separate lineage originating early in the diversification of Theclinae Swainson, 1831 ( Fig. 20). Therefore, we propose that this lineage corresponds to a tribe. This new tribe is diagnosed by a combination of the following characters, as given for the Drina section by Eliot (1973): male genitalia with significantly reduced (nearly vestigial) tegumen (not wider than valva in lateral view), uncus, and falces, and long and narrow vinculum; forewing with 11 veins, the three M veins are nearly at the same distance from each other, hindwing with a single tail at vein CuA 2. A combination of the following nuclear genomic base pairs is diagnostic: cce305.14.6:G130A, cce305.14.6:T131C, cce 1105.7.5:G93A, cce144.8.2:G223A, cce204.17.10: C49A, cce3081.9.2:G97G (not T), cce 2598.5.13:G484G (not C), cce 2598.5.13:C485C (not A), cce948.2.2: T73T (not A), cce948.2.2: A139A (not G).

Genera included. Only the type genus.

Parent Taxon. Subfamily Theclinae Swainson, 1831 .

Hypochrysopini Grishin, new tribe

http://zoobank.org/ 71317481-C0FE-4839-97E2-341444327DD7

Type genus. Hypochrysops C. Felder & R. Felder, 1860 .

Definition. Currently placed in Luciini Waterhouse & Lyell, 1914 , Hypochrysops (type species Papilio polycletus Linnaeus, 1758 ) and related genera are not monophyletic with it and instead form a distinct clade within Theclinae Swainson, 1831 not confidently associated with any other subtribe ( Fig. 20). Therefore, this clade represents a subtribe. This new tribe is diagnosed by a combination of the following characters, as given for the Hypochrysops section by Eliot (1973): uncus not produced, rounded, falces prominent, strongly curved, juxta absent or vestigial, aedeagus very thick, just slightly narrower than valva, valva rhomboidal, saccus not developed; ventral wing surface typically with obsolete or distorted patterns with metallic silver or green spots and bands and red blotches. A combination of the following nuclear genomic base pairs is diagnostic: cce3313.6.1:G1784C, cce3313.6.1:G2131C, cce67043.1.6:C55G, cce67043.1.6:A56T, cce1184.15.22:G142C.

and Titea Eliot, 1973 .

Parent Taxon. Subfamily Theclinae Swainson, 1831 .

Jalmenini Grishin, new tribe

http://zoobank.org/ D58D0D0A-382D-4E6D-B879-75D3DE7A3670

Type genus. Jalmenus Hübner, 1818 .

Definition. Currently in Zesiusinae Swinhoe, 1912, Jalmenus (type species Jalmenus evagoras Hübner, 1818 , which is a junior homonym of Papilio evagoras Donovan, 1805 ) is far removed from the genus Zesius Hübner, [1819] (type species Zesius chrysomallus Hübner, 1823 ) in the genomic tree, and instead forms a distinct lineage within Theclinae Swainson, 1831 not confidently associated with other genera ( Fig. 20). Therefore, this lineage represents a tribe. This new tribe is diagnosed by a combination of the following characters, as given for the Jalmenus section (excluding Pseudalmenus H. H. Druce, 1902 ) by Eliot (1973): uncus and tegumen narrower in lateral view, longer than in relatives, ventral side not expanded, falces smaller, tegumen not expanded anteriad, valva without costal process, simple and rounded; palpi with 3 rd segment very long, 2 nd segment with bristle-like scales. A combination of the following nuclear genomic base pairs is diagnostic: cce9330.11.2:G69A, cce7486.3.3:T58G, cce9657.10. 14:T3166C, cce10680.1.1:T28C, cce4529.2.2:A67G.

Genera included. Only the type genus.

Parent Taxon. Subfamily Theclinae Swainson, 1831 .

Pseudalmenini Grishin, new tribe

http://zoobank.org/ BB6B6C59-EAD7-42B4-B7E6-CA95E94A419D

Type genus. Pseudalmenus H. H. Druce, 1902 .

Definition. In the genomic tree, Pseudalmenus (type species Ialmenus myrsilus Westwood, 1851 , which is a subspecies of Thecla chlorinda Blanchard, 1848 ) is a weakly supported (54%) sister of Jalmenus Hübner, 1818 , and therefore may not be monophyletic with it ( Fig. 20). Hence, we propose that this lineage represents a tribe. This new tribe was included in the Jalmenus section by Eliot (1973) and is diagnosed by a combination of the following characters: 3 rd segment of palpi shorter than in Jalmenini trib. n., 2 nd segment hairy; uncus with ventral portion expanded, protruding posteriad of the dorsal margin, falces long and strongly curved, tegumen well-developed, expanded anteriad, valva with the curved costal process giving it a crab claw-like appearance. A combination of the following nuclear genomic base pairs is diagnostic: cce10780.3.1:G247A, cce 1122.11.2:C1079G, cce11670.2.2:C750T, cce24738.4.8:A1793G, cce10780.2.2:A2954G.

Genera included. Only the type genus.

Parent Taxon. Subfamily Theclinae Swainson, 1831 .

Myrinini Toxopeus, 1929 is a tribe

Currently in Amblypodiini Doherty, 1886 , close relatives Myrina [Fabricius], 1807 (type species Papilio alcides Cramer, 1776 ) and Iraota F. Moore, 1881 (type species Hesperia maecenas Fabricius, 1793 ) are distantly related to Amblypodia Horsfield, 1829 (type species Thecla narada Horsfield, 1828 ). The clade of these three genera is not strongly supported (42%) in our tree ( Fig. 20), and their union is not monophyletic in a global phylogeny of butterflies (Kawahara et al. 2023). Therefore, we propose a status of a tribe for Myrinini Toxopeus, 1929 , stat. nov., which consists of two genera ( Myrina and Iraota ).

Horagina Swinhoe, 1910 and Loxurina Swinhoe, 1910 are subtribes of Cheritrini Swinhoe, 1910

Currently regarded as distinct tribes, Cheritrini Swinhoe, 1910 , Horagini Swinhoe, 1910 , and Loxurini Swinhoe, 1910 ( Eliot 1973) are closely related to each other and are at the tree level of subtribes ( Fig. 20). Being combined, all three constitute one tribe. The priority of these names could not be determined because they were proposed (as subfamilies) on the same page of the same work (i.e., issued on the same date). As the first revisers, we give priority to Cheritrini Swinhoe, 1910 , because this group includes more genera and species than the other two. As a result, we propose that Horagina Swinhoe, 1910 , stat. nov. and Loxurina Swinhoe, 1910 , stat. nov. are subtribes of Cheritrini Swinhoe, 1910 .

Rapalini Grishin, new tribe

http://zoobank.org/ 6B76C64C-1C57-4CC8-BBA2-925D4AFBCF7D

Type genus. Rapala F. Moore, 1881 .

Definition. Currently, in Deudorigini Doherty, 1886 , several genera, including Rapala (type species Thecla varuna Horsfield, 1829 ), form a clade that is not confidently monophyletic with it: only 26% partitions in the genomic tree support their grouping with Deudorigini . Low support values indicate a more distant relationship and a possibility of incomplete lineage sorting or gene exchange around the time of origin of these clades. The clade consisting of Rapala (mostly Oriental) and three Afrotropical genera closely related to Pilodeudorix H. H. Druce, 1891 (type species Pilodeudorix barbatus H. H. Druce, 1891 ) is most confidently supported (100%) and corresponds to the level of a tribe in the tree ( Fig. 20). This new tribe differs from relatives by male genitalia having conjoined valvae that evenly taper to narrow rounded or pointed apices, uncus and tegumen broad, two lobes of uncus with a concave margin between them, hood-shaped; secondary sexual characters in males: an oval brand of small androconia near the base of cell Sc+R 1 -RS (at least in non-African species), and typically a hair tuft on ventral forewing near inner margin to complement the brand, sometimes with other brands, including those on abdomen; hindwing tailed, forewing veins R 4+5 and M 1 originate separately, although may be very narrowly separated at their origins ( Eliot 1973). Most confidently distinguished by DNA and a combination of the following base pairs in the nuclear genome is diagnostic: cce303273.1.1:G197A, cce 1093.2.1:A4672C, cce3516.7.1:A173T, cce12299.7.3:G134A, cce4160.2.2:A207G.

Genera included. The type genus (i.e., Rapala F. Moore, 1881 ), Pilodeudorix H. H. Druce, 1891 , Hypomyrina H. H. Druce, 1891 , and Paradeudorix Libert, 2004 .

Parent Taxon. Subfamily Theclinae Swainson, 1831 .

Pilodeudorigina Grishin, new subtribe

http://zoobank.org/ 5826F9A8-6834-487C-B207-DC883CE19307

Type genus. Pilodeudorix H. H. Druce, 1891 .

Definition. Afrotropical clade of Rapalini trib. n. that includes Pilodeudorix (type species Pilodeudorix barbatus H. H. Druce, 1891 , which is a junior subjective synonym of Sithon camerona Plötz, 1880 ) is genetically differentiated from non-Arican (mostly Oriental) species at the tree level of a subtribe ( Fig. 20). This new subtribe is diagnosed by a combination of the following characters: abdomen frequently with scent brands and hindwings with hair tufts; aedeagus shorter and less gracile that in the nominotypical tribe, frequently expanded terminally, vesica with many small cornuti and usually with large cuneus; basally fused valvae with a rectangular gap between them in many species, in others, each valva with a process, hook-like in ventral view, uncus lobes are frequently less separated from each other than in the nominotypical subtribe ( Stempffer 1967). Best distinguished by DNA and a combination of the following nuclear genomic base pairs is diagnostic: cce303351.9.14:A88G, cce303351.9.14:A89G, cce14215.8.1:C143T, cce881.1.6:A79G, cce462.24.2:C89G.

Genera included. The type genus (i.e., Pilodeudorix H. H. Druce, 1891 ), Hypomyrina H. H. Druce, 1891 , and Paradeudorix Libert, 2004 .

Parent Taxon. Tribe Rapalini Grishin, trib. n.

Oxylidini Eliot, 1973 , Remelanini Eliot, 1973 , and Hypolycaenini Swinhoe, 1910 belong to Polyommatinae Swainson, 1827 , not to Theclinae Swainson, 1831

Traditionally placed among hairstreaks ( Theclinae Swainson, 1831 ) due to their appearance (frequently with very long hindwing tails), species from the tribes Oxylidini Eliot, 1973 , Remelanini Eliot, 1973 , and Hypolycaenini Swinhoe, 1910 are confidently placed as a sister clade to “Blues” and are not monophyletic with the subfamily Theclinae ( Figs. 20, 22). Therefore, they do not belong to Theclinae . Their genetic differentiation from the clade that consists of traditional Polyommatinae Swainson, 1827 smaller than that for other subfamilies of Lycaenidae . Insufficient differentiation argues against treating Oxylidini , Remelanini, and Hypolycaenini as a subfamily of their own. Therefore, we place them in the subfamily Polyommatinae . We see that the three subfamilies, Lycaeninae , Theclinae , and Polyommatinae , are closely related to each other ( Figs. 20, 22), as expected and evidenced by morphology ( Eliot 1973). The genetic distinction of Lycaeninae from the two other subfamilies is more pronounced, and the tree branches separating them are more prominent ( Fig. 20). However, Theclinae and Polyommatinae are less distinct from each other, and “blues” originate from within “hairstreaks”, not next to them, like butterflies originate from within moths, birds from within reptiles, and tetrapods from within fishes. Furthermore, due to their close relationship, it is conceivable to unify Theclinae and Polyommatinae into one subfamily ( Polyommatinae ). We are not taking this step here and continue with the traditional tri-subfamily arrangement for now, simply rearranging tribes among Theclinae and Polyommatinae to restore monophyly. See our proposed subtribal classification of Lycaenidae below.

Hemiolaina Grishin, new subtribe

http://zoobank.org/ 45FFC7EB-6CFA-49A6-A688-EC98E721BA62

Type genus. Hemiolaus Aurivillius, 1922 .

Definition. Hemiolaus (type species Jolaus caeculus Hopffer, 1855 ) forms a lineage sister to other Oxylidini Eliot, 1973 , but is genetically differentiated from them at the subtribal level, and therefore represents a subtribe ( Fig. 22). This new subtribe is diagnosed by a combination of the following characters, as given for the Hemiolaus section by Eliot (1973): juxta enlarged, longer than half of valva, shaped as the end of a nail puller with a deep cleft; ventral hindwing of males with a scent patch beneath a hair brush; antennal club short, nudum confined to the club, about 32 segments, palpi with 3 rd segment slightly less than half of the 2 nd, tarsus of male foreleg terminally tapered and downturned. A combination of the following nuclear genomic base pairs is diagnostic: cce54422.3.2:C373T, cce993.15.2:A922T, cce303329.2.7:A2399C, cce2502.15.1:A320C, cce5483.2.11:T370A.

Genera included. Only the type genus.

Parent Taxon. Tribe Oxylidini Eliot, 1973 .

Cupidopsina Grishin, new subtribe

http://zoobank.org/ 8AFD9453-3743-4ECB-BA87-4DAC0A3297FB

Type genus. Cupidopsis Karsch, 1895 .

Definition. Confidently placed in the tribe Hypotheclini Eliot, 1973 , the lineage with Cupidopsis (type species Lycaena jobates Hopffer, 1855 ) is genetically differentiated from the Hypothecla Semper, 1890 lineage at the tree level of subtribes ( Fig. 22) and therefore represents a subtribe. This new subtribe is diagnosed by a combination of the following characters as given for the Cupidopsis section by Eliot (1973): only 10 veins on the forewing, secondary sexual characters absent, aedeagus with developed coecum and ductus entrance on dorsal side, saccus smaller than in relatives, nearly vestigial, tegumen with uncus comparatively massive, the same length as valva in lateral view. A combination of the following nuclear genomic base pairs is diagnostic: cce9377.2.3:A136T, cce4053.12.2:A3235C, cce332.21. 2:G117T, cce199. 20.2:A244C, cce3203.9.2:A172C.

Genera included. Only the type genus.

Parent Taxon. Tribe Hypotheclini Eliot, 1973 .

Comments. A family-group name formed from the same genus was proposed as a nomen nudum (fails ICZN Art. 13.) by Koçak (1996). The genomic tree demonstrates that this subtribe (as other Hypotheclini ) belongs to Polyommatinae Swainson, 1827 ( Figs. 20, 22), contrary to the hypothesis of Stradomsky

Niphandina Sibatani & Ito, 1942 is a subtribe

In agreement with Stradomsky (2016), we find that Niphandini Sibatani & Ito, 1942 clusters closely with Polyommatini Swainson, 1827 , being at the tree level with subtribes ( Fig. 22). Therefore, we confirm its treatment as a subtribe Niphandina Sibatani & Ito, 1942 , stat. conf.

Theclinesthina Grishin, new subtribe

http://zoobank.org/ 32E65131-DDFB-4EBC-BE37-4F42E8F5F6D3

Type genus. Theclinesthes Röber, 1891 .

Definition. The clade with Theclinesthes (type species Plebeius (Theclinesthes) eremicola Röber, 1891 , which is a junior subjective synonym of Nacaduba miskini gaura Doherty, 1891 ) originates near the base of Polyommatini Swainson, 1827 ( Fig. 22) and therefore represents a subtribe. This new subtribe is diagnosed by a combination of the following characters, as given for the Theclinesthes section by Eliot (1973) and Stradomsky (2016): uncus lobes and (vestigial) falces directed ventrad, vinculum in lateral view much broader than in relatives, as broad as valva, aedeagus basally bulbous and apically tapered, ductus enters at anterior end, valva constricted in the middle. A combination of the following nuclear genomic base pairs is diagnostic: cce2737.15.2:A259G, cce3516.7.8:A98T, cce10374.3.2:A67C, cce 2234.8. 11:G1960A, cce2399.18.8:A86G.

Genera included. The type genus (i.e., Theclinesthes Röber, 1891 ), Neolucia G. Waterhouse & Turner, 1905 , and Sahulana Hirowatari, 1992 .

Parent Taxon. Tribe Polyommatini Swainson, 1827 .

Comment. The same name was published in a pioneering study by Stradomsky (2016) without explicitly indicating that the name was intentionally new (fails ICZN Art. 16.1.) and not stating what the type genus of this taxon was (fails ICZN Art. 16.2.). This name for the subtribe already discovered by Stradomsky and confirmed by our genomic analysis is simply formalized here to comply with the ICZN Code ( ICZN [International Commission on Zoological Nomenclature] 1999) .

Azanina Grishin, new subtribe

http://zoobank.org/ DF55B888-23ED-4B2E-94F9-B1B6685CBDA3

Type genus. Azanus F. Moore, 1881 .

Definition. Azanus (type species Papilio ubaldus Stoll, 1782 ) is confidently placed in a clade of Polyommatini Swainson, 1827 containing several subtribes, not confidently grouping with any of them ( Fig. 22). Therefore, the lineage with Azanus represents a subtribe. This new subtribe is diagnosed by a combination of the following characters, as given for the Azanus section by Eliot (1973): male genitalia elongated and flattened, uncus narrowly divided into separate lobes, forewings veins SC and R 1 come together and then diverge, androconia of two unusual types: nearly rectangular scales with concave bases and long padded scales, eyes hairy, ventral forewing with a dark streak below SC vein. A combination of the following nuclear genomic base pairs is diagnostic: cce9549.2.2:A815G, cce302383.7.1:T1153A, cce302383.7.1:C1154G, cce 2510.1.2:A686G, cce178.15.6:A356G.

Genera included. Only the type genus.

Parent Taxon. Tribe Polyommatini Swainson, 1827 .

Comment. The same name was proposed as a nomen nudum (fails ICZN Art. 13.) by Koçak (1996) and then published in a pioneering study by Stradomsky (2016) without explicitly indicating that the name was intentionally new (fails ICZN Art. 16.1.) and not stating what the type genus of this taxon was (fails by our genomic analysis is simply formalized here to comply with the ICZN Code (ICZN [International Commission on Zoological Nomenclature] 1999).

Unina Grishin, new subtribe

http://zoobank.org/ 2FD670E6-5ECD-404E-A1F7-842AEF97335C

Type genus. Una Nicéville, 1890 .

Definition. Una (type species Zizera? usta Distant, 1886 ) is confidently placed in the clade of Polyommatini Swainson, 1827 with several subtribes, not confidently grouping with any of them ( Fig. 22). Therefore, the lineage with Ionolyce represents a subtribe. This new subtribe is a union of the Una and Petrelaea sections of Eliot (1973), who listed characters for them, and is diagnosed as follows: male genitalia elongated and appear flattened, with gracile and long aedeagus and prominent saccus (absent in close relatives); forewings with 11 veins, veins SC and R 1 fuse at least for some distance. A combination of the following nuclear genomic base pairs is diagnostic: cce10730.5.6:A179T, cce7658.4.3:A184G, cce4822.3.3:A98T, cce5018.4.1:A78G, cce870.7.1:G452A.

Genera included. The type genus (i.e., Una Nicéville, 1890 ), Orthomiella Nicéville, 1890 , Petrelaea Toxopeus, 1929 , and Pseudonacaduba Stempffer, 1942 .

Parent Taxon. Tribe Polyommatini Swainson, 1827 .

Comment. The same name was proposed as a nomen nudum (fails ICZN Art. 13.) by Koçak & Seven (1997).

Ionolycina Grishin, new subtribe

http://zoobank.org/ 77081BED-FF23-424A-98B3-541E04BA53E5

Type genus. Ionolyce Toxopeus, 1929 .

Definition. Ionolyce (type species Ionolyce helicon javanica Toxopeus, 1929 ) is confidently placed in the clade of Polyommatini Swainson, 1827 with several subtribes, not confidently grouping with any of them ( Fig. 22). Therefore, the lineage with Ionolyce represents a subtribe. This new subtribe is diagnosed by a combination of the following characters, as given for Ionolyce by Tite (1963): cornuti in aedeagus are large and spine-like, ribs in androconial scales are ribbon-like with nodular irregularities mainly in the posterior third of the scale; fused part of veins SC and R 1 is typically longer than in relatives, and the free end of vein SC is faint. A combination of the following nuclear genomic base pairs is diagnostic: cce437.15.1:A182G, cce303334.4.2:C106G, cce3111.1.6:A86G, cce3368.2.2:T241A, cce29649.12.1:A151G.

Genera included. The type genus (i.e., Ionolyce Toxopeus, 1929 ) and Paraduba Bethune-Baker, 1906 .

Parent Taxon. Tribe Polyommatini Swainson, 1827 .

Pithecopina Grishin, new subtribe

http://zoobank.org/ 3E5E4A9A-64A9-4AFC-85DD-81B13BE53FBF

Type genus. Pithecops Horsfield, 1828 .

Definition. The clade with Pithecops (type species Pithecops hylax corax Fruhstorfer, 1919 , which is a subspecies of Pithecops corvus Fruhstorfer, 1919 ) is confidently placed as sister to the “crown group” of Polyommatini Swainson, 1827 that undergoes extensive diversification ( Fig. 22), and we include it in the “crown group” despite its unusual wing patterns. The Pithecops clade does not have close relatives within Polyommatini and, therefore, represents a subtribe. This new subtribe is diagnosed by a combination of the following characters, as given for the Pithecops section by Eliot (1973): male genitalia elongated and and R 1 fuse at least for some distance, eyes not hairy, palpi hairy. A combination of the following nuclear genomic base pairs is diagnostic: cce59502.1.2:A118G, cce1246.19.3:A71G, cce9990.6.1:G740A, cce1317. 1.1:C772T, cce3516.4.2:G83T.

Genera included. Only the type genus.

Parent Taxon. Tribe Polyommatini Swainson, 1827 .

Comment. The same name was published in a pioneering study by Stradomsky (2016) without explicitly indicating that the name was intentionally new (fails ICZN Art. 16.1.) and not stating what the type genus of this taxon was (fails ICZN Art. 16.2.). This name for the taxon already discovered by Stradomsky and confirmed by our genomic analysis is simply formalized here to comply with the ICZN Code ( ICZN [International Commission on Zoological Nomenclature] 1999) .

Zizulina Grishin, new subtribe

http://zoobank.org/ A230CCD1-B083-4025-8B78-A31A89478932

Type genus. Zizula Chapman, 1910 .

Definition. The lineage with Zizula (type species Lycaena gaika Trimen, 1862 , which is a junior subjective synonym of Papilio hylax Fabricius, 1775 ) is a confident sister to Brephidiina Stempffer, 1957 , but is at the tree level that corresponds to subtribes in the “crown group” of Polyommatini Swainson, 1827 ( Fig. 22) and therefore represents a subtribe. This new subtribe is diagnosed by a combination of the following characters, as given for the Zizula section by Eliot (1973): male genitalia unusual, aedeagus stout and terminally divided into two processes of about half of its length, dorsal and ventral, together resembling a beak, valva with a rod-like process of about the same length as genitalia and long bristles twice of valval length, veins SC and R 1 fused towards costa, secondary sexual characters absent. A combination of the following nuclear genomic base pairs is diagnostic: cce993.15.2:A161G, cce993.15.2: T162C, cce811.10.3:A2207G, cce811.10.3:T2197C, cce1162.15.2:T151A.

Genera included. Only the type genus.

Parent Taxon. Tribe Polyommatini Swainson, 1827 .

Comment. The same name was proposed as a nomen nudum (fails ICZN Art. 13.) by Koçak (1996) and then published in a pioneering study by Stradomsky (2016) without explicitly indicating that the name was intentionally new (fails ICZN Art. 16.1.) and not stating what the type genus of this taxon was (fails ICZN Art. 16.2.). This name for the subtribe already discovered by Koçak and Stradomsky and confirmed by our genomic analysis is simply formalized here to comply with the ICZN Code (ICZN [International Commission on Zoological Nomenclature] 1999).

Jamidina Grishin, new subtribe

http://zoobank.org/ F3E26669-CDF0-4EFC-9539-F88A3D47B1F7

Type genus. Jamides Hübner, [1819] .

Definition. The lineage with Jamides (type species Papilio bochus Stoll, 1782 ) is a confident sister to Celastrinina Tutt, 1907 , but splits from the latter at the tree level of subtribes ( Fig. 22), thus representing a subtribe. This new subtribe is diagnosed by a combination of the following characters, as given for the Jamides section by Eliot (1973): male genitalia with vinculum lacking cephalad expansion, uncus directed ventrad, falces very long (nearly half of valva length), aedeagus longer than valva (usually by a third or more), valva deeply bilobed; forewing veins SC and R 1 are not fused for any distance, but connected with a short cross-vein. A combination of the following nuclear genomic base pairs is diagnostic: cce23510.3.2: T93C, cce23510.3.2:A432G, cce 1568.2.4:A5519G, cce 1568.2.4:A6856C, cce10823.2.3:T258C.

Parent Taxon. Tribe Polyommatini Swainson, 1827 .

Comment. The same name was proposed as a nomen nudum (fails ICZN Art. 13.) by Koçak (1996) and then published in a pioneering study by Stradomsky (2016) without explicitly indicating that the name was intentionally new (fails ICZN Art. 16.1.) and not stating what the type genus of this taxon was (fails ICZN Art. 16.2.). This name for the subtribe already discovered by Koçak and Stradomsky and confirmed by our genomic analysis is simply formalized here to comply with the ICZN Code (ICZN [International Commission on Zoological Nomenclature] 1999).

Fameganina Grishin, new subtribe

http://zoobank.org/ 07D24435-1EAD-4C80-9350-0A1FA6003972

Type genus. Famegana Eliot, 1973 .

Definition. The lineage with Famegana (type species Lycaena alsulus Herrich-Schäffer, 1869 , which is a junior subjective synonym of Lycaena nisa Wallace, 1866 ) is sister to Zizeeriina Chapman, 1910 with moderate confidence ( Fig. 22). Because the confidence of this grouping is not the highest and because this lineage originates at the tree level of subtribes, it represents a subtribe. This new subtribe is diagnosed by a combination of the following characters, as given for the Famegana section by Eliot (1973): in male genitalia, tegumen and uncus bulky, falces stout and nearly rigidly connected, uncus lobes terminally pointed and slightly downturned; veins SC and R 1 touch each other over a short distance, eyes not hairy, palpi with bristles. A combination of the following nuclear genomic base pairs is diagnostic: cce 1088.12. 3:A100T, cce 1088.12.3:G102A, cce18.45.4:G1256T, cce18.45.4:G1255A, cce10490.1.2:T1933A.

Genera included. Only the type genus.

Parent Taxon. Tribe Polyommatini Swainson, 1827 .

Comment. The same name was proposed as a nomen nudum (fails ICZN Art. 13.) by Koçak & Seven (1997) and then published in a pioneering study by Stradomsky (2016) without explicitly indicating that the name was intentionally new (fails ICZN Art. 16.1.) and not stating what the type genus of this taxon was (fails ICZN Art. 16.2.). This name for the subtribe already discovered by Koçak& Seven and Stradomsky and confirmed by our genomic analysis is simply formalized here to comply with the ICZN Code (ICZN [International Commission on Zoological Nomenclature] 1999).

Oboroniina Grishin, new subtribe

http://zoobank.org/ FB96541D-672A-41A6-9C47-2EDAADEC0D60

Type genus. Oboronia Karsch, 1893 .

Definition. The clade with Oboronia (type species Oboronia staudingeri Hemming, 1960 , which is a junior subjective synonym of Plebeius punctatus Dewitz, 1879 ) is placed within the “crown group” of Polyommatini Swainson, 1827 without strongly supported phylogenetic affinity to any subtribe ( Fig. 22) and therefore represents a subtribe of its own. This new subtribe is diagnosed by a combination of the following characters, as given for the Euchrysops section by Eliot (1973): male genitalia with long falces, vinculum broadening in the middle in lateral view, long and narrow valva, massive rod-shaped aedeagus with anterior ductus entrance; veins SC and R 1 not fused. A combination of the following nuclear genomic base pairs is diagnostic: cce349.2.1:C166A, cce349.2.1:A167T, cce935.8.2:A66G, cce 2073.8.1: A230G, cce178.15.6:A190G.

Genera included. The type genus (i.e., Oboronia Karsch, 1893 ), Euchrysops Butler, 1900 , Lepidochrysops Hedicke, 1923 , Orachrysops Vári, 1986 , and Thermoniphas Karsch, 1895 .

Parent Taxon. Tribe Polyommatini Swainson, 1827 .

indicating that the name was intentionally new (fails ICZN Art. 16.1.) and not stating what the type genus of this taxon was (fails ICZN Art. 16.2.). This name for the subtribe already discovered by Stradomsky and confirmed by our genomic analysis is simply formalized here to comply with the ICZN Code ( ICZN [International Commission on Zoological Nomenclature] 1999) .

Uranothaumatina Grishin, new subtribe

http://zoobank.org/ 57D2B8F3-36FA-4457-88B2-FC99DADC2EF8

Type genus. Uranothauma Butler, 1895 .

Definition. The lineage with Uranothauma (type species Uranothauma crawshayi Butler, 1895 ) is confidently placed as sister to Scolitantidina Tutt, 1907 ( Fig. 22), but it was not traditionally included in the latter subtribe. Additionally, because it originates at the tree level corresponding to subtribes, it represents a subtribe. This new subtribe is a union of the Uranothauma and Phlyaria sections of Eliot (1973), who listed characters for them, and is diagnosed as follows (see also Stradomsky (2016) for genitalia illustrations): in male genitalia, saccus absent, falces developed, as long as tegumen with uncus, vinculum expanded in the middle in lateral view, aedeagus shorter than valva, rod-shaped with ductus entrance dorso-cephalad, valva elongated, undivided; veins SC and R 1 touch each other or fuse at least for some distance, eyes hairy, palpi hairy or bristly. A combination of the following nuclear genomic base pairs is diagnostic: cce993.29.2:A515G, cce103.22.12:A47G, cce462.35.1:G193T, cce912.1.1:C56A, cce 1162.12.1:G739A.

Genera included. The type genus (i.e., Uranothauma Butler, 1895 ) and Phlyaria Karsch, 1895 .

Parent Taxon. Tribe Polyommatini Swainson, 1827 .

Comment. The same name was published in a pioneering study by Stradomsky (2016) without explicitly indicating that the name was intentionally new (fails ICZN Art. 16.1.) and not stating what the type genus of this taxon was (fails ICZN Art. 16.2.). This name for the subtribe already discovered by Stradomsky and confirmed by our genomic analysis is simply formalized here to comply with the ICZN Code ( ICZN [International Commission on Zoological Nomenclature] 1999) .

Higher classification of Lycaenidae to the subtribal level

Based on our genome-scale phylogeny ( Figs. 20–22) complemented with other studies ( Talavera et al. 2012; Boyle et al. 2015; Robbins et al. 2022; Boyle et al. 2023; Kawahara et al. 2023), we propose the following provisional classification of Lycaenidae into subfamilies, tribes, and subtribes. We partition the family into eight subfamilies. If no tribes and subtribes are listed for a subfamily, we consider that subfamily to be monotypic. If no subtribes are listed for a tribe, we consider that tribe to be monotypic. The type genus name for each taxon is given in parentheses. New taxa and status changes are shown in red font. Synonymy is not provided.