Egeona, Zhang & Cong & Shen & Song & Grishin, 2025

Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina & Grishin, Nick V., 2025, Advancing butterfly systematics through genomic analysis, The Taxonomic Report of the International Lepidoptera Survey 12 (5), pp. 1-201 : 11-13

Egeona Grishin , new subgenus

http://zoobank.org/ BFF752DF-DA63-4043-91F6-34E579F956F8

Type species. Taxila egeon Westwood, 1851 .

Definition. This is the second new subgenus of Dodona Hewitson, 1861 (type species Melitaea durga Kollar, 1844 ) (see above for discussions) ( Fig. 5 green) that is sister to the nominal subgenus ( Fig. 5 magenta). COI barcodes between these sister taxa differ by 7.3% (48 bp). This new subgenus differs from its relatives by the following combination of characters: the phallus is usually longer and stronger curved, the phallobase is straighter and the connection between the phallus and phallobase is less bent; males have brown wings with orange spots and stripes above (four stripes on the forewing: the apical stripe—which is sometimes vestigial—not merged with the submarginal stripe) but without white areas and stripes characteristics of Balonca and with wings and orange spots less round and spots less uniform than in the subgenus Dodona , and differs from several similar-looking species of Balonca either by more extensive orange coloration, especially of the ventral side, or by not having brown framing on the basal side of pale hindwing streaks. For genitalia illustrations of some representative species in the new subgenus, see Wu et al. (2018). In DNA, a combination of the following characters is diagnostic in the nuclear genome: cne3991.3.1:T363C, cne3991.3.1:T384C, cne9860.2.4:C37T, cne 1134.1.1:A336G, cne3461.1.15:C2524A; and in COI barcode: T127A or T463C, A202C or T206C, T479T, T484T, T571C or T574A, T533T.

Etymology. The name is formed from the name of the type species and is a feminine noun in the nominative singular.

Species included. The type species (i.e., Taxila egeon Westwood, 1851 ), Dodona adonira Hewitson, 1866 , Dodona chrysapha Fruhstorfer, 1910 , Dodona eugenes H. Bates, 1867 , Dodona formosana Matsumura, 1919 , Dodona hoenei Forster, 1951 , Dodona maculosa Leech, 1890 , Dodona phuongi Monastyrskii & Devyatkin, 2000 , Dodona speciosa Monastyrskii & Devyatkin, 2000 , and Dodona windu Fruhstorfer, 1894 , including their subspecies and synonyms.

Parent taxon. Genus Dodona Hewitson, 1861 .

http://zoobank.org/ 83817CD8-BC22-4AA1-BF5C-7F07CA4DF9D7 ( Figs. 6 part, 7, 8b)

Definition and diagnosis. Nuclear genome analysis of Lasaia H. Bates, 1868 (type species Papilio meris Stoll, 1781 ) reveals a clade ( Fig. 6a red) that is sister to both Lasaia sula Staudinger, 1888 (type locality in Honduras) ( Fig. 6 blue) and Lasaia peninsularis Clench, 1972 (type locality in Mexico: Veracruz) ( Fig. 6a purple), thus representing a new species. The three species (the new one, L. sula , and L. peninsularis ) are genetically differentiated from each other to a similar degree in the nuclear genome ( Fig. 6a) (Fst 0.37 and 0.44 between the new one and L. sula and L. peninsularis , respectively) but do not strongly differ in the mitochondrial genome ( Fig. 6b) and, consequently, also in the COI barcode. The new species differs from its relatives by males having better developed dark spots and dashes, the hindwing with stronger developed dark dashes, similarly to the forewing (weaker than on the forewing or absent dashes in both L. sula and L. peninsularis ), and some specimens having less blue above, greyer, thus somewhat resembling Lasaia maria maria Clench, 1972 (type locality in Mexico: Jalisco) and Lasaia sessilis Schaus, 1890 (type locality in Mexico: Veracruz), but are paler and patterned more like L. sula . In male genitalia ( Fig. 8), the transtilla ( McAlpine 1971) is pointed in the middle as in L. peninsularis and not flattened as in L. sula ( Fig. 8 green arrow 1); lateral lobes of the transtilla are narrower than in L. sula and are more similar to L. peninsularis ( Fig. 8 green arrow 2), if not smaller; and the scobinate bulla (Clench 1972) ( Fig. 8 green arrow 3) is more robust than in the other two species. Due to the cryptic nature of this species and

unexplored individual variation, most reliable identification is achieved by DNA, and a combination of the following base pairs is diagnostic in the nuclear genome: cne 2812.5.8:A69 T, cne28857.1.4:G42A, cne28857.1.4:A65G, cne8137.3.6:C54 T, cne8137.3.6:C66 T. The COI barcode does not differ for L. sula . Barcode sequence of the holotype. Sample NVG-23103F05, GenBank PV 549979, 658 base pairs: AACTTTATATTTTATTTTTGGAATTTGAGCAGGAATAGTAGGTACATCATTAAGTTTATTAATTCGTATAGAATTAGGTATACCTGGATCATTAATTGGAGATGATCAAATTTATAATACT ATTGTTACAGCTCATGCTTTTATTATAATTTTTTTTATAGTTATACCCATTATAATTGGAGGATTTGGTAATTGATTAGTACCTTTAATATTAGGAGCTCCTGATATAGCATTTCCACGAA TAAATAATATAAGATTTTGACTTTTACCCCCATCTTTATTTCTATTAATTTCAAGAAGTATTGTAGAAAATGGAGCAGGAACTGGTTGAACAGTTTATCCCCCTTTATCTTCTAATATTGC CCACGGAGGATCCTCAGTAGATTTAGCTATTTTCTCTCTTCATTTAGCAGGAATTTCTTCAATTTTAGGAGCCATTAATTTTATTACAACTATTATTAATATACGAATTAATAATTTATCT TTTGATCAAATACCATTATTCGTATGATCCGTTGGTATTACTGCTTTATTATTATTATTATCATTACCTGTTTTAGCAGGAGCTATTACTATATTATTAACTGATCGTAATTTAAATACAT CTTTTTTTGATCCAGCAGGAGGAGGTGATCCAATTTTATATCAACATTTATTT

Type material. Holotype: ♂ deposited in the Carnegie Museum of Natural History, Pittsburgh, PA, USA ( CMNH), illustrated in Fig. 7, bears the following four rectangular labels (1 st handwritten, others printed), three white: [ MEXICO: Colima | Comala 2100 ft. | 1.X.1967 | R.G.Wind], [ R.G. Wind, leg. | Gift of F.M.Brown | C.M. Acc. 23123], [DNA sample ID: | NVG-23111F10 | c/o Nick V. Grishin ], and one red [HOLOTYPE ♂ | Lasaia cola | Grishin ]. Paratypes: 2♂♂ and 1♀ from Mexico, Colima: 1♂ NVG-24079H06 (leg DNA extraction, sequenced), NVG-25014D03 (abdomen DNA extraction and dissection) La Salada, 1000 ft, 4-Jan-1968, Robert G. Wind leg., genitalia NVG250517-02 ( Fig. 8b) [ MGCL] and (no locality details) [ SMF]: 1♂ NVG-23103F 05 May-1918 and 1♀ NVG-23103F 06 Oct-1926.

Type locality. Mexico: Colima, Comala , elevation 2100 ft.

Etymology. The name is formed from the type locality in Col [im] a and is a feminine noun in apposition.

Distribution. Currently known only from Colima in Mexico.

Comment. In Lasaia , valvae are partly (and weakly) sclerotized and are flexible, semi-transparent side flaps (with sparse setae) on the sides of the scobinate bulla (Clench 1972), as seen in Fig. 8a, ventral view.