Libytheana Michener, 1943

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 : 14-17

Libytheana Michener, 1943 is a genus distinct from Prolibythea Scudder, 1889

Considered synonyms in some publications ( Kawahara 2009; Sohn et al. 2012), fossil Prolibythea Scudder, 1889 (type species Prolibythea vagabunda Scudder, 1889 ) and contemporary Libytheana Michener, 1943 (type species Libythea bachmanii Kirtland, 1851 , which is regarded as a subspecies of Papilio carinenta Cramer, 1777 ), both American, are separated by at least 30 million years according to the age estimate of the fossil as late Priabonian ( Sohn et al. 2012). The typical age of congeners is not larger than 20 million years. Furthermore, the divergence between Libytheana and the Old World genus Libythea [Fabricius], 1807 (type species Papilio celtis Laicharting, 1782 ) was dated to about 12 Mya (Kawahara et al. 2023), which is more recent than 30 Mya. Thus, it is most likely that Prolibythea lived before the divergence between Libytheana and Libythea . Therefore, if Libytheana and Libythea are treated as separate genera, then Prolibythea is not congeneric with Libytheana to avoid paraphyly.

The tribe Vagrantini Pinratana & Eliot, 1996 as currently defined is paraphyletic

Currently, the tribe Vagrantini Pinratana & Eliot, 1996 consists of ten genera: Vagrans Hemming, 1934 , Cupha Billberg, 1820 , Phalanta Horsfield, 1829 , Smerina Hewitson, 1874 , Terinos Boisduval, 1836 , Algia Herrich-Schäffer, 1864 ), Algiachroa Parsons, 1989 , Cirrochroa E. Doubleday, 1847 , Lachnoptera E. Doubleday, 1847 , and Vindula Hemming, 1934 ( Wahlberg 2019). However, our genomic tree reveals that Vagrantini , defined to include all ten genera, is paraphyletic with respect to Argynnini Swainson, 1833 with the highest support ( Fig. 6). The first five genera listed above form a clade sister to Argynnini. This clade includes Vagrans , which is the type genus of Vagrantini . Therefore, to restore monophyly, we restrict Vagrantini to include only these five genera: Vagrans , Cupha , Phalanta , Smerina , and Terinos . The remaining five genera previously included in Vagrantini form a clade sister to both Vagrantini and Argynnini ( Fig. 6) and, therefore, belong to other tribes. No published family-group names have been formed from any of these five genera; hence, these other tribes are new. They are described below.

Vindulini Grishin, new tribe

http://zoobank.org/ 151AC163-E036-49BE-A49C-48D7B9F0108F

Type genus. Vindula Hemming, 1934 .

Definition. Vindula (type species Papilio arsinoe Cramer, 1777 ) constitutes a lineage sister to four other genera that were previously included in Vagrantini Pinratana & Eliot, 1996 but did not belong to this tribe (see above). This lineage diverged from these other genera at about the same level as (if not earlier than) Vagrantini from Argynnini Swainson, 1833, and therefore corresponds to a tribe ( Fig. 6). This new tribe is distinguished from its relatives by sclerotized subpapillary glands in females and forked humeral vein ( Penz and Peggie 2003). A combination of the following nuclear genomic base pairs is diagnostic: hm2013347-RA.4:T162C, hm2013540-RA.5:G265A, hm2015146-RA.7:G80A, hm2015146-RA.7:G79A, hm2013347-RA.4:A220C.

Genera included. Only the type genus.

Parent Taxon. Subfamily Heliconiinae Swainson, 1822 .

http://zoobank.org/ AE49C6E7-8E51-4E4E-9947-151B37A467E8

Type genus. Algia Herrich-Schäffer, 1864 .

Definition. This tribe corresponds to the second major subclade in the clade that is sister to both Argynnini Swainson, 1833 and Vagrantini Pinratana & Eliot, 1996 . This subclade is sister to Vindulini trib. n., diverging from it at about the same level as (if not earlier than) Argynnini from Vagrantini ( Fig. 6). Due to this prominent genetic differentiation, it is defined as a tribe. This new tribe is distinguished from its relatives by unsclerotized subpapillary glands in females, forked humeral vein, and/or smooth eyes and undifferentiated androconial scales, or by an oval patch of androconial scales in males in the apical area of dorsal hindwing covering 1/7–1/6 of its surface ( Penz and Peggie 2003). A combination of the following nuclear genomic base pairs is diagnostic: hm2000037-RA.1:C698G, hm2008858-RA.12: T1021C, hm2008858-RA.12:C1022T, hm2016824-RA.4:C154A, hm2017493-RA.1:A2480G.

Genera included. The type genus (i.e., Algia Herrich-Schäffer, 1864 ), Algiachroa Parsons, 1989 , Cirrochroa E. Doubleday, 1847 , and Lachnoptera E. Doubleday, 1847 .

Parent Taxon. Subfamily Heliconiinae Swainson, 1822 .

Lachnopterina Grishin, new subtribe

http://zoobank.org/ 8BBABF1F-145B-4A67-94F5-F498130FB857

Type genus. Lachnoptera E. Doubleday, 1847 .

Definition. Lachnoptera (type species Papilio iole Fabricius, 1781 ) forms a lineage that splits from all other Algiini trib. n. at the tree level of subtribes ( Fig. 6), therefore representing a subtribe. This new subtribe is distinguished from its relatives by unsclerotized subpapillary glands in females ( Penz and Peggie 2003) and an oval patch of androconial scales in males in the apical area of dorsal hindwing covering 1/7–1/6 of its surface. A combination of the following nuclear genomic base pairs is diagnostic: hm2008958-RA.5: T118 G, hm2008958-RA.5:G119 T, hm2006642-RA.2:A2275C, hm2006706-RA.1:A709 T, hm2006706-RA.1:A748 T, hm2015589-RA.1:A1852A (not C), hm2014529-RA.4:A139A (not G), hm2012118- RA.6:A79A (not G), hm2016492-RA.5:A1813A (not G), hm2016492-RA.5:C1831C (not A) .

Genera included. Only the type genus.

Parent Taxon. Tribe Algiini Grishin, trib. n.

Terinosina Grishin, new subtribe

http://zoobank.org/ 31DFAAA5-EA63-431B-AC9D-BF686549FEC8

Type genus. Terinos Boisduval, 1836 .

Definition. Terinos (type species Terinos clarissa Boisduval, 1836 ) forms a deep-diverging lineage in the tribe Vagrantini Pinratana & Eliot, 1996 at about the tree level corresponding to subtribes ( Fig. 6) and, therefore, represents a subtribe. This new subtribe is distinguished from its relatives by a combination of the following characters: larval head with scoli, hindwing cell closed, a fold across the forewing between R and M veins, vein R 2 arising from discal cell, vein R 4 arising at about the end of R 2 , humeral vein simple and straight, gnathos arms not ventrally fused, valva with a long (about 2/3 of valva length) projection off its base inside ( Penz and Peggie 2003). A combination of the following base pairs in the nuclear genome is diagnostic: hm2007153-RA.3:A305G, hm2014625-RA.4: T887 A, hm2012596-RA.1: C853G, hm2011273-RA.1: T484 G, hm2011273-RA.1:G486A .

Genera included. Only the type genus.

Parent Taxon. Tribe Vagrantini Pinratana & Eliot, 1996 .

http://zoobank.org/ 06DF6620-9B17-4522-9E4B-BD5342AC8DF5

Type genus. Smerina Hewitson, 1874 .

Definition. Smerina (type species Smerina vindonissa Hewitson, 1874 ) forms a deep-diverging lineage in the tribe Vagrantini Pinratana & Eliot, 1996 at about the tree level corresponding to subtribes ( Fig. 6) and therefore represents a subtribe. This new subtribe is distinguished from its relatives by a combination of the following characters: papilla anales moderately retracted (not deeply) inside the body, aedeagus not broadened at the tip in ventral view, costa of valva with one spiny process ( Penz and Peggie 2003), forewing apex more produced and hindwing margin evenly curved (not wavy, no tails). A combination of the following base pairs in the nuclear genome is diagnostic: hm2015462-RA.1: T360 C, hm2015689-RA. 5: T57 C, hm2009280-RA.3: T94 G, hm2010526-RA.2:A67G, hm2017391-RA.1:A139C .

Genera included. Only the type genus.

Parent Taxon. Tribe Vagrantini Pinratana & Eliot, 1996 .

Lebadeini Grishin, new tribe

http://zoobank.org/ F26F931E-9141-4329-92B6-6E86A9F369E9

Type genus. Lebadea C. Felder, 1861 .

Definition. Lebadea (type species Limenitis ismene E. Doubleday, 1848 , which is a junior subjective synonym of Papilio martha Fabricius, 1787 ), currently in Neptini Newman, 1870 ( Dhungel and Wahlberg 2018; Wahlberg 2019) is not monophyletic with it and instead is placed as sister to the clade of Chalingini Hemming, 1960 and Limenitidini Behr, 1864 but with weak support; therefore, it is distinct from them ( Fig. 6). Thus, this lineage, currently consisting only of Lebadea , represents a tribe that does not have a name. This new tribe is diagnosed by a combination of the following characters: tegumen and uncus are smaller than typical for Limenitidini , uncus is more gracile, thus more similar to Neptis [Fabricius], 1807, and differs from Neptis in having a well-defined and projecting anteriad lobe (not just a hump) on the dorsal side of the segment A 2 in the pupa (Willmott 2003). A combination of the following nuclear genomic base pairs is diagnostic: hm2005025-RA.3:C373A, hm2006832-RA.2:C97A, hm2004700-RA.1: C590A, hm2004700-RA.1:G351T, hm2005025-RA.3:G319A.

Genera included. Only the type genus.

Parent Taxon. Subfamily Limenitidinae Behr, 1864 .

Comment. Using morphological analysis, Willmott (2003) has placed Lebadea in the “ Limenitis group” of genera away from Neptis , which seems to agree more with our genomic results.