Three new subfamilies of skipper butterflies (Lepidoptera, Hesperiidae) Author Zhang, Jing Author Cong, Qian Author Shen, Jinhui Author Brockmann, Ernst Author Grishin, Nick V. text ZooKeys 2019 861 91 105 http://dx.doi.org/10.3897/zookeys.861.34686 journal article http://dx.doi.org/10.3897/zookeys.861.34686 1313-2970-861-91 802AE175A8104FFCB3FDE289D0CC4191 Katreinae Grishin subfam. n. Type genus. Katreus Watson, 1893. Diagnosis . In appearance, most similar to Celaenorrhinus Huebner , [1819] and its relatives ( Evans 1937 ), and was placed in Celaenorrhinini Swinhoe, 1912 by Warren et al. (2008 , 2009 ) but differs by longer apiculus of antennae and hindwing produced at vein 1A+2A. Morphologically, distinguished from all Hesperiidae by the combination of the following characters. Abdomen short, shorter than inner margin of hindwing. Antennal club arcuate, bent in the middle, apiculus long, pointed. Second segment of palpi protrudes partly forward and partly upward (at an angle between the axis of the body and the axis perpendicular to it, =sub-erect). Males with hair pencil on hind tibiae, without stigmas or brands on wings. Forewing discal cell long, about 2/3 of the costa; vein M2 originates about midway between or closer to M1 than to M3 and vein CuA2 originates closer to the base of wing than to the end of discal cell. Hindwing produced at vein 1A+2A, vein 3A much shorter than vein CuA2. Male genitalia with a well-developed gnathos, which is not smaller than uncus, uncus bulging dorsad in lateral view, with small or tiny arms distant from each other, tegumen robust, extends caudad for the length of uncus, harpe longer than sacculus. See Larsen (2005 : 469-471) for illustrations of all representative species in this subfamily. In DNA, a combination of the following base pairs in the nuclear genome is diagnostic: aly528.10.2:G940C, aly925.27.5:A3610T, aly84.77.5:T1651G, aly595.14.2:G184C, aly2284.22.2:G967C, and in COI barcode region: C235T, A335T, C347T, and T349A. Genera included. Katreus with its invalid synonym Choristoneura Mabille, 1889 (junior homonym of Choristoneura Lederer 1859 in Lepidoptera : Tortricidae ) and subjective synonyms Loxolexis Karsch, 1895 and Daratus Lindsey, 1925 (replacement name for Choristoneura ) ( Fig. 1e ); and Ortholexis Karsch, 1895 with its subjective synonym Acallopistes Holland, 1896 ( Fig. 1a-d ). Comments. Taxonomy of these skippers has been confusing until it was resolved by Cock and Congdon (2011) . For the most part, they were all placed in the genus Katreus , until Larsen emphasized the differences in genitalia of those species placed in Ortholexis from true Katreus ( Larsen 2005 ). Indeed, the two genera are quite distinct in our genomic analysis. A recent study based on several genes placed this group (only Ortholexis holocausta (Mabille, 1891) was included in that study) as a sister of Pyrrhopygini Mabille, 1877 ( Sahoo et al. 2017 ), probably due to an insufficient number of genes included. In their study, Euschemon Doubleday, 1846 grouped with Eudaminae instead of being sister to all other Hesperiidae with exclusion of Coeliadinae Evans, 1937 ( Warren et al. 2009 ; Zhang et al. 2017 ; Toussaint et al. 2018 ); such problems are expected from smaller datasets. We find ( Fig. 2 ) that the Katreinae subfam. n. is an ancient and unique Afrotropical lineage that diverged from other Hesperiidae at the time when the family was diversifying into subfamilies.