A Re-Appraisal of Gegenes Hübner, 1819 (Lepidoptera, Hesperiidae
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Tijdschrift voor Entomologie 160 (2017) 41-60 A re-appraisal of Gegenes Hübner, 1819 (Lepidoptera, Hesperiidae) based on male and female genitalia, with the description of a new genus, Afrogegenes Rienk de Jong & John G. Coutsis Examination of male and female genitalia as well as external characters of all species of the current genus Gegenes Hübner, 1819 (Hesperiidae, Lepidoptera) led to the observation that two well-marked groups can be distinguished, differing in external characters as well as in male and female genitalia. For one of the two groups a new genus is erected, Afrogegenes. Conventionally, the group for which the new genus is erected contains two species, Gegenes letterstedti (Wallengren, 1857) and G. hottentota (Latreille, 1823), but G. hottentota ocra Evans, 1937, has been given full species rank here, raising the number of species in Afrogegenes to three. G. letterstedti has long been known in the literature as G. niso (Linnaeus, 1764), but this is based on an incorrect lectotype designation. Male and female genitalia are described and figured for all taxa, in some cases for the first time. Keywords: Lepidoptera; Hesperiidae; genitalia; Gegenes; Afrogegenes Rienk de Jong*, Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands. [email protected] John G. Coutsis, 4 Glykonos Street, Athens 10675, Greece. [email protected] Introduction The skipper genus Gegenes Hübner, 1819, as con- (Evans 1937), not counting the northern G. nostro- ceived by Evans (1937, 1949) consists of four rela- damus, that has reached Sudan (Evans 1949, Larsen tively small, rather drab-colored and swift-flying 1983). species, which are easily overlooked in the field and While the African species are not difficult to dis- often neglected by collectors because of their dull tinguish as males, the Mediterranean species are eas- colors. The genus is, with two or three species, dis- ily confused. A summary of relevant literature and tributed in scattered colonies in the Mediterranean distinguishing characters for the two species in the area and eastward through the eremic regions as far eastern Mediterranean was given by Coutsis & Ol- as the Punjab (North India) and one species even to ivier (1993), based on males only, leaving some un- Bengal (Evans 1949). It occurs with a single species certainty about females. Recently, the second author in Oman (Musandam Peninsula, Dhofar); three spe- studied the female genitalia of the Mediterranean cies can be found in present-day Yemen, while two species (Coutsis 2012). Therefore a re-appraisal of species can be found further north in the Arabian the characters of the entire genus is timely. Synony- Peninsula (Larsen 1983). Further, it occurs with at my is kept to a minimum; interested readers should least three species throughout sub-Saharan Africa consult the relevant literature, particularly Evans from Senegal to Eritrea and south to South Africa (1937, 1949) and Larsen (1982a). Tijdschrift voor Entomologie 160: 41–60, Table 1, Figs 1–67. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 21 July 2017. DOI 10.1163/22119434-16001003 *Corresponding author Downloaded from Brill.com09/27/2021 04:06:06PM via free access <UN> 42 Tijdschrift voor Entomologie, volume 160, 2017 All genitalia drawings (many more than presented in the male sex. It is the more remarkable since Wat- here) were made by the second author, the other il- son remarked: “No secondary sexual characters on lustrations by the first author. Illustrated material is the wings of the male except in a single specimen of in the collection of the Naturalis Biodiversity Center, an unidentified species in the British Museum from Leiden, unless indicated otherwise. Victoria Nyanza, in which there is a large ill-defined rounded spot of appressed scales on the upper side of the forewing below end of cell.” It is exactly this The traditional genus Gegenes conspicuous spot that so easily separates G. hotten- The genus belongs to the tribe Baorini of the Hes- tota from letterstedti, indicating that Watson had periinae (Warren et al. 2009), a strictly Old World misjudged the real character of hottentota. grouping of 14 mostly dark to grey brown genera Holland (1896) followed Watson in accepting (Fan et al. 2016). The justification for the erection the synonymy of Hesperia nostrodamus with Papilio of the genus Gegenes, first described by Hübner pumilio, and of Hesperia hottentota with Hesperia let- (1819), has never been challenged. terstedti. In the case of the last two names he was The genus is easily distinguished from all other of the opinion that the males with and without the Baorini by the very short antennae, only one third large black patch on the upper side of the forewing the length of the costa of the forewing, the short club were two morphs of the same species, since he had consisting of 11 segments and the minute apiculus numerous specimens collected together, as well as consisting of one or two (at most three) segments. specimens of both “morphs” caught in coitu with fe- The strongly reduced apiculus distinguishes the males which were identical. genus from all other Old-World Hesperiidae ex- Evans (1937) finally recognized the four species cept Thymelicus Hübner, 1819, a genus of about nostrodamus, pumilio, letterstedti and hottentota, a ten Palaearctic Hesperiidae (Hesperiinae: Hesperi- concept that was challenged for the first time by ini), all with an ochreous and unspotted or vaguely Larsen (1982a), as described below. spotted upperside (Evans 1949), and Taractrocera Butler, 1870, a genus of 16 Indo-Australian Hesperi- Two groups idae (Hesperiinae: Taractrocerini), mostly with large The traditional genus Gegenes (sensu Evans 1937, ochreous spots in a brown ground color (De Jong 1949) easily falls apart into two groups that differ as 2004). The mid-tibiae are heavily spined in Gegenes, much in external (color, design) and structural char- as is the case in some other genera of Baorini, such acters (mainly male and female genitalia) as any two as Pelopidas Walker, 1870, and Baoris Moore, 1881, genera of the Baorini, reason for us to separate the but according to Fan et al. (2016), spines on the species into two genera, Gegenes Hübner, 1819, and mid-tibiae are not a useful taxonomic character. Afrogegenes genus novum, the former widespread, from the Mediterranean to N India and throughout Subdivision of Gegenes sub-Saharan Africa, the latter restricted to Africa (with some extension into the Arabian peninsula). History The most obvious differences between the two Watson (1893: 104) was the first author to place the gen era are given in Table 1. For more details, see the species dealt with in this paper together in the genus descrip tions below. Gegenes. However, his subdivision and synonymy Early stages are known for the two species of Ge- were rather confusing. On the basis of the extensive genes and for at least one species of Afrogegenes. A collections in the British Museum (Natural History), summary for the African species is given by Cock & London, he recognized three species, with synonyms, Congdon (2012). All species live on various grasses. as follows (with our present-day interpretation in A. letterstedti has even been observed on introduced square brackets): G. nostrodamus Fabricius, 1793, grasses: Zea (maize, corn) and Oryza (rice). So far, with synonyms pygmaeus Hübner [recte Cyrilli], characters of the caterpillars do not support the divi- 1887 (nec Fabricius), pumilio Hoffmansegg, 1804, sion of the species into two genera. This also holds and lefebvrii Rambur, 1842 [at present the last three for the pupae, which moreover are similar to the names are considered to belong to a single species, pupae of other Baorini genera as Pelopidas Walker, G. pumilio, distinct from the first one, nostrodamus 1870, Borbo Evans, 1949 and Zenonia Evans, 1935, (Evans 1937, 1949, and others)], G. karsana Moore, in being green, elongate and with a frontal spike. 1874 [at present considered a junior synonym of nostrodamus (Evans 1937, 1949)], and G. hottentota Latreille, 1823, with letterstedti Wallengren, 1857, Gegenes Hübner, 1819 as junior synonym [at present considered a separate The sole species included by Hübner (1819) in his species]. The last synonymy by Watson is remark- new genus was Papilio pygmaeus Fabricius 1775 able, since the two taxa are very different externally (now in the genus Aeromachus). However, Fabricius’ Downloaded from Brill.com09/27/2021 04:06:06PM via free access <UN> De Jong & Coutsis: Re-appraisal of Gegenes and description of Afrogegenes 43 Table 1. Characters distinguishing Gegenes and Afrogegenes. Character Gegenes Afrogegenes apiculus 1 segment 2 segments hindwing underside greyish brown, hardly visible pale dots ochreous, with small dark markings and patches spines mid tibia usually 6, varying from 5 to 8 5, sometimes 6, rarely 7 tegumen + uncus constricted before middle, strongly expanding distally parallel-sided, narrowing distally uncus apically deeply indented with widely separated arms squarish, apically slightly indented, apical angles slightly protruding gnathos two large reniform structures two narrow, elongate, sligthtly curved structures gnathos–uncus broadly connected narrowly connected saccus > half length uncus + tegumen << half length uncus + tegumen valva, proximal edge about vertical oblique, angle with ventral edge about 300 aedeagus two pronounced cornuti without cornuti pre-vaginal plate