Abstract
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.
Rienk de Jong*, Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands. rienk.dejong@naturalis.nl
John G. Coutsis, 4 Glykonos Street, Athens 10675, Greece. kouts@otenet.gr
Introduction
The skipper genus Gegenes Hübner, 1819, as conceived by Evans (1937, 1949) consists of four relatively small, rather drab-colored and swift-flying species, which are easily overlooked in the field and often neglected by collectors because of their dull colors. The genus is, with two or three species, distributed in scattered colonies in the Mediterranean area and eastward through the eremic regions as far as the Punjab (North India) and one species even to Bengal (Evans 1949). It occurs with a single species in Oman (Musandam Peninsula, Dhofar); three species can be found in present-day Yemen, while two species can be found further north in the Arabian Peninsula (Larsen 1983). Further, it occurs with at least three species throughout sub-Saharan Africa from Senegal to Eritrea and south to South Africa (Evans 1937), not counting the northern G. nostrodamus, that has reached Sudan (Evans 1949, Larsen 1983).
While the African species are not difficult to distinguish as males, the Mediterranean species are easily confused. A summary of relevant literature and distinguishing characters for the two species in the eastern Mediterranean was given by Coutsis & Olivier (1993), based on males only, leaving some uncertainty about females. Recently, the second author studied the female genitalia of the Mediterranean species (Coutsis 2012). Therefore a re-appraisal of the characters of the entire genus is timely. Synonymy is kept to a minimum; interested readers should consult the relevant literature, particularly Evans (1937, 1949) and Larsen (1982a).
All genitalia drawings (many more than presented here) were made by the second author, the other illustrations by the first author. Illustrated material is in the collection of the Naturalis Biodiversity Center, Leiden, unless indicated otherwise.
The traditional genus Gegenes
The genus belongs to the tribe Baorini of the Hesperiinae (Warren et al. 2009), a strictly Old World grouping of 14 mostly dark to grey brown genera (Fan et al. 2016). The justification for the erection of the genus Gegenes, first described by Hübner (1819), has never been challenged.
The genus is easily distinguished from all other Baorini by the very short antennae, only one third the length of the costa of the forewing, the short club consisting of 11 segments and the minute apiculus consisting of one or two (at most three) segments. The strongly reduced apiculus distinguishes the genus from all other Old-World Hesperiidae except Thymelicus Hübner, 1819, a genus of about ten Palaearctic Hesperiidae (Hesperiinae: Hesperiini), all with an ochreous and unspotted or vaguely spotted upperside (Evans 1949), and Taractrocera Butler, 1870, a genus of 16 Indo-Australian Hesperiidae (Hesperiinae: Taractrocerini), mostly with large ochreous spots in a brown ground color (De Jong 2004). The mid-tibiae are heavily spined in Gegenes, as is the case in some other genera of Baorini, such as Pelopidas Walker, 1870, and Baoris Moore, 1881, but according to Fan et al. (2016), spines on the mid-tibiae are not a useful taxonomic character.
Subdivision of Gegenes
History
Watson (1893: 104) was the first author to place the species dealt with in this paper together in the genus Gegenes. However, his subdivision and synonymy were rather confusing. On the basis of the extensive collections in the British Museum (Natural History), London, he recognized three species, with synonyms, as follows (with our present-day interpretation in square brackets): G. nostrodamus Fabricius, 1793, with synonyms pygmaeus Hübner [recte Cyrilli], 1887 (nec Fabricius), pumilio Hoffmansegg, 1804, and lefebvrii Rambur, 1842 [at present the last three names are considered to belong to a single species, G. pumilio, distinct from the first one, nostrodamus (Evans 1937, 1949, and others)], G. karsana Moore, 1874 [at present considered a junior synonym of nostrodamus (Evans 1937, 1949)], and G. hottentota Latreille, 1823, with letterstedti Wallengren, 1857, as junior synonym [at present considered a separate species]. The last synonymy by Watson is remarkable, since the two taxa are very different externally in the male sex. It is the more remarkable since Watson remarked: “No secondary sexual characters on the wings of the male except in a single specimen of an unidentified species in the British Museum from 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 conspicuous spot that so easily separates G. hottentota from letterstedti, indicating that Watson had misjudged the real character of hottentota.
Holland (1896) followed Watson in accepting the synonymy of Hesperia nostrodamus with Papilio pumilio, and of Hesperia hottentota with Hesperia letterstedti. In the case of the last two names he was of the opinion that the males with and without the large black patch on the upper side of the forewing were two morphs of the same species, since he had numerous specimens collected together, as well as specimens of both “morphs” caught in coitu with females which were identical.
Evans (1937) finally recognized the four species nostrodamus, pumilio, letterstedti and hottentota, a concept that was challenged for the first time by Larsen (1982a), as described below.
Two groups
The traditional genus Gegenes (sensu Evans 1937, 1949) easily falls apart into two groups that differ as much in external (color, design) and structural characters (mainly male and female genitalia) as any two genera of the Baorini, reason for us to separate the species into two genera, Gegenes Hübner, 1819, and Afrogegenes genus novum, the former widespread, from the Mediterranean to N India and throughout sub-Saharan Africa, the latter restricted to Africa (with some extension into the Arabian peninsula).
The most obvious differences between the two genera are given in Table 1. For more details, see the descriptions below.
Early stages are known for the two species of Gegenes and for at least one species of Afrogegenes. A summary for the African species is given by Cock & Congdon (2012). All species live on various grasses. A. letterstedti has even been observed on introduced grasses: Zea (maize, corn) and Oryza (rice). So far, characters of the caterpillars do not support the division of the species into two genera. This also holds for the pupae, which moreover are similar to the pupae of other Baorini genera as Pelopidas Walker, 1870, Borbo Evans, 1949 and Zenonia Evans, 1935, in being green, elongate and with a frontal spike.
Gegenes Hübner, 1819
The sole species included by Hübner (1819) in his new genus was Papilio pygmaeus Fabricius 1775 (now in the genus Aeromachus). However, Fabricius’ description did not match with Hübner’s own figures, which were of Papilio pygmaeus Cyrilli, 1787. Butler (1870) fixed the latter as genotype, but since Cyrilli’s name was a junior homonym of Fabricius’ name, it had to be replaced by Papilio pumilio Hoffmansegg, 1804.
Description/Diagnosis
Antennal shaft. Underside plain creamy white or lightly chequered.
Club. Eleven segments, one of which is the tiny apiculus.
Wings. Upper side brown to very dark brown, may be paler outwardly; males unspotted, females with variable number of pale ochreous spots in, at most, spaces r3–5, m1–2, m3, cua1, cua2. Underside forewing with spots as on upper side; underside hindwing brown with more or less dense greyish superscaling (overlaying scales), discal spots more or less indicated.
Mid tibia. Usually 6 long spines, but number may vary from five to eight.
Male genitalia. In dorsal and ventral aspect tegumen + uncus (the two are difficult to separate) constricted just before middle, strongly expanding in distal half. Uncus deeply indented, the widely separated lateral arms with shallow dorsal flap and irregularly formed apex. Gnathos consisting of two large, smooth, more or less reniform structures, medially widely separated, in ventral view superimposed on uncus arms, ventrally broadly and imperceptibly passing into uncus sides. Saccus relatively long, more than half the length of tegumen + uncus. Valva elongate, proximal edge about vertical, costa almost as long as ventral edge and broadly parallel to it, more or less straight, slightly curving up distally, ventral edge nearly straight to slightly concave in proximal half, cucullus sharply curving up distally, apex partly or entirely overlapping distal costal process, upper edge of cucullus without proximal extension. Aedeagus: vesica with two pronounced cornuti, dorsal aperture of ductus seminalis near proximal end of pre-zonal part, post-zonal part long.
Female genitalia. Pre-vaginal plate deeply indented medially, smooth, not overlapping post-vaginal plate. Post-vaginal plate oblong with rounded ends. Ductus bursae heavily sclerotized. Corpus bursae rounded.
The two species traditionally recognized, G. pumilio and G. nostrodamus, are not easily distinguished externally, particularly in the female sex (see Coutsis & Olivier 1993 for differences). For this reason, the second author studied the previously undocumented female genitalia (Coutsis 2012). This is the more important since the two species overlap geographically, although they are rarely found in the same locality, and part of the older literature records may be based on confusion between the two. Lorkovic´ (1971) described an easily visible character to distinguish the two species in spread specimens (see the descriptions of the species) but his short note was largely overlooked. Gegenes pumilio and G. nostrodamus have different ecological preferences, the first being mainly found in open Mediterranean garrigue (a type of low, soft-leaved scrubland) and dried out river beds, often near the coast, while G. nostrodamus is more shade-loving, frequenting areas with scattered trees, and may occur far inland, as in Spain (García-Barros et al. 2013).
Gegenes pumilio (Hoffmansegg, 1804)
External characters. Antennal shaft chequered on underside: apical part of segments darker than basal part. Length of forewing male 13.1−16.9 mm (n = 25, mean = 13.8 mm, s = 0.6) , female 13.8−16.6 mm (n = 25, mean = 14.0 mm, s = 0.7). Upper side dark blackish brown, forewing unmarked in male, with obscure spots in spaces 1b, 2 and 3, and at apex in female. Underside pale grey–brown with, in female, indistinct spots on forewing as on upper side, hindwing also with some indistinct spots. Forewing underside with densely packed dark brown scales in basal part of discoidal cell clearly visible in spread specimens (Lorkovic´ 1971) (in butterflies and skippers this patch of specialized scales is part of a wing coupling mechanism, see Kuijten 1974), not overlapped by the fringe of fine short hairs along the basal part of the front edge of the hindwing underside (compare G. nostrodamus). This character occurs in both sexes. It is visible in Figs 28 and 29. In honour of the discoverer we call this character the “Lorkovic´ character”.
Male genitalia. Saccus: length about 3/4 of vertical length of vinculum. Valva: cucullus relatively short, scarcely overlapping costa; upper margin smooth in proximal quarter or third, distal two-thirds or three-quarters unevenly dentate. Aedeagus: vesica with two cornuti of even size, left one dentate in side view, right one plain.
In order to find possible differences between G. pumilio and G. gambica we examined the genitalia of specimens from France (Cannes), Italy (Lipari island, north of Sicily) and Spain (Mallorca) in Europe, Tunisia (Tabarka) in NW Africa, and Chad (Bebedjia), Kenya (Arabuko Forest) and Madagascar (Taolagnaro) in sub-Saharan Africa, in addition to specimens from numerous Greek localities. They were all similar and although there was some slight individual variation, there was no indication of geographical variation.
Female genitalia. Lamella antevaginalis deeply divided into a bifid structure with two narrow arms, lamella postvaginalis slightly more than twice as broad as high, heavily sclerotized ductus bursae of even width throughout, and corpus bursae less than twice as long as sterigma.
Distribution. G. pumilio is widely distributed in scattered populations in NW Africa (Algeria and Tunisia; Tennent 1996). Europe: Spain, Mallorca (García-Barros et al. 2013; according to these authors not known from the Spanish mainland, although mentioned by Evans (1937, 1949), SE France (Lafranchis 2000) and eastward through Italy and Sicily (Verity 1940) to the Balkans and Greece (Rebel & Zerny 1931, Thurner 1964, Schaider & Jaksic´ 1989, Pamperis 2009, Verovnik & Popovic´ 2013). Further east it occurs in Turkey (Hesselbarth et al. 1995), Cyprus (Makris 2001), Syria (Evans 1949), Lebanon (Larsen 1974), Israel (Benyamini 1997), Iraq (Wiltshire 1957, Tshikolovets et al. 2014) and Iran (Evans 1949, Nazari 2003, Tshikolovets et al. 2014, mainly along the Persian Gulf) to Pakistan (not mentioned by Roberts 2001, but records from Chitral, Waziristan and Khyber in Evans 1949); latter locality could as well be Afghanistan, but not mentioned by Sakai (1981) from that country; there is a female from Rawalpindi in the collection of the Naturalis Biodiversity Center (Figs 8, 9), and NW India (Punjab, NW Himalayas; Evans 1949). Not recorded from Turkmenistan, Uzbekistan, Tajikistan or Kyrgyzstan (Tshikolovets 1998, 2000, 2003, 2005), where the very similar G. nostrodamus is widespread, and (most of) Afghanistan, leaving a gap of some 800 km between SE Iran and the Pakistan/NW India localities. In addition, the species is found in S and SE Arabian Peninsula (Yemen, Dhofar, Oman; Larsen 1980, 1982b, 1983, 1984, Larsen & Larsen 1980). Detailed distribution data also provided by Kudrna et al. (2011) and Tshikolovets (2011). It occurs throughout sub-Saharan Africa (Evans 1937, Kielland 1990, Larsen 1991, 2005), including the island of Socotra east of the Horn of Africa, politically belonging to Yemen (Rebel 1907, Evans 1937), in suitable habitats, but absent from most of Cape Province, South Africa, (Dickson & Kroon 1978, Pringle et al. 1994, Henning et al. 1997). Not mentioned from Gabon (Vande weghe 2010) and Cameroon (personal data). It has not been recorded from Madagascar so far, but there is a single specimen in the collection of the Naturalis Biodiversity Center, Leiden, from Ft Dauphin ( = Taolagnaro) (Figs 6–7, 19).
Geographic variation. Gegenes pumilio has been divided into three subspecies (Evans 1937, 1949), a northern one, north of the Sahara and east to NW India (the nominal subspecies), a southern one throughout Africa (ssp. gambica Mabille) and the third (ssp. monochroa Rebel) restricted to the island of Socotra. Ssp. gambica differs from the nominate form in having more prominent markings on the underside of the hindwing and reduced spots in the female forewing. Ssp. monochroa is smaller and darker with barely visible spots.
It has been found, however, that the chromosome numbers are different in the western and eastern Mediterranean. De Lesse (1960, 1967) reported a haploid chromosome number of n = 24 for specimens from Nice and Algiers, and n = 41 for a specimen from Gülek near Adana, Turkey. Larsen (1982b) confirmed the latter number for specimens from Beirut and Yemen. In view of the huge distribution area of the species, the importance of such scattered findings should not be overestimated, but as pointed out by Larsen (1982b), if seen in combination with the large distribution gaps between Tunisia and Israel (more than 2000 km) and between Adana and Rhodos (700 km, according to Larsen; actually about 600 km), it is suggestive of the existence of two species with different chromosome number, one in the western and central Mediterranean, and the other in the eastern Mediterranean, leaving the position of the sub-Saharan African populations uncertain. Meanwhile, the large gap in Turkey has been bridged by findings of the species around Alanya, near Antalya, and even as far west as the Turkish west coast around Kuc´adası (Hesselbarth et al. 1995), close to Aegean islands like Chios, from where it has been reported by Cuvelier (2014). Equally, the first author found the species not uncommon around Bodrum in the southwest of Turkey in April 2016 (Fig. 1).
Thus, the supposed gap between the central and eastern Mediterranean is, actually, non-existent. There is, however, a gap in the western Mediterranean, since the species is not known from Morocco and the Iberian Peninsula, although it occurs on the island of Mallorca (García-Barros et al. 2013). Evans (1937) mentioned three males and two females from “S. Spain”, but the thorough study by García-Barros et al. (2013) has not revealed continental Spanish specimens. It must be added that the western and eastern Mediterranean populations cannot be separated morphologically.
Finally, there appears to be a wide gap on both sides of the Red Sea, since there are no records along the more than 2000 km between Israel in the north and Yemen and Ethiopia in the south. Possibly there is a more or less loose chain of populations from Yemen via the southern and eastern part of the Arabian Peninsula to Iraq and Iran. As described above, a further gap (of some 800 km) appears to exist between SE Iran and the Pakistan/NW India populations. Whether the chromosome number remains the same over the huge area from S Africa to NW India is unknown.
A solution to the question of the exact distribution of the two chromosomal numbers also has nomenclatural consequences. The type locality of G. pumilio is Napoli. It is tempting to suppose that here the chromosome number will be the same as in Nice, i.e. n = 24. This is, however, unknown. It would also be helpful, if the chromosome number of Greek populations were known. If it proves the same as in Nice, then it would be highly unlikely that the number in Napoli would be different. In other words, the chromosome number in the type locality of the species would be n = 24. Although there are some junior synonyms given to sub-Saharan populations (see Larsen 1982a), there are no names given to populations from the eastern Mediterranean and further east. So if these populations, with n = 41, are considered a separate species, then either they need a new name or, if the chromosome number of sub-Saharan populations proves to be n = 41 as well, they should have the oldest available name for such populations, viz. Pamphila gambica Mabille, 1878. Anticipating on this, Larsen (2005: 546) wrote “Ssp. gambica is found throughout the drier savannah zones of Africa, except for the Cape, as well as in Arabia and the Middle East to India”.
At the present state of knowledge we think it premature to take such far-reaching decisions. Since the sub-Saharan populations are, on the whole, slightly different externally from the northern populations, from NW Africa to India (the reason why Evans 1937 recognized them as ssp. gambica), we hesitate to take any taxonomic or nomenclatural action here before the chromosome number has been established in relevant localities, particularly in The Gambia, type-locality of ssp. gambica. Alternatively, DNA analysis across the entire distribution area would be helpful. Biogeographically it would not be surprising to find a close link between the West African and NW African/western Mediterranean populations (De Jong 1976). The possibility that a haploid chromosome number of 24 is found in West Africa, while in more eastern populations in Africa it is 41, cannot be ruled out without further research. It should also be kept in mind that the distance between the type-locality of ssp. gambica and Yemen is about 7000 km and much can happen in between. An example of the biogeographic link between West Africa and the west-Mediterranean biota that comes to mind is Colotis evagore Klug, 1829, a pierid that is widely distributed in dry parts of sub-Saharan Africa, NW Africa and Spain as far north as Catalonia (Fric 2005); eastward it extends to the western and southern part of the Arabian Peninsula, and to Hadhramaut (Larsen 1983) and Socotra (Butler 1881, as Teracolus niveus).
Gegenes nostrodamus (Fabricius, 1793)
External characters. Antennal shaft creamy white on underside. Length of forewing male 13.5−17.1 mm (n = 25, mean = 14.5 mm, s = 0.7), female 14.5−19.2 mm (n = 25, mean = 15.8 mm, s = 0.9). Upper side pale to brown, forewing unmarked in male, with pale indistinct spots in spaces 1b, 2 and 3, and at apex in female. Underside pale sandy brown, unmarked in male, female forewing with spots as on upper side, hindwing same vague discal spots. Forewing underside with densely packed dark brown scales in basal part of discoidal cell, in spread specimens hidden by fringe of long hairs or narrow scales at frontal edge of basal part of hindwing underside (Lorkovic´ 1971) (the Lorkovic´ character, see under G. pumilio). Obviously, this character is only helpful in specimens in which the hairs or scales at the frontal edge of the hindwing have not been rubbed off by age or rough handling. It is found in both sexes. See Figs 28 and 29.
Male genitalia. Saccus as long as vertical length of vinculum. Valva: cucullus longer than in G. pumilio, more broadly overlapping costa and distally extending beyond it, upper margin evenly dentate, also in proximal part. Aedeagus: vesica with two cornuti of uneven size, left one small and dentate, right one much larger and without teeth.
Female genitalia. Lamella antevaginalis with indention just over middle leaving two broad parts, hardly higher than wide, together covering most of sterigma, lamella postvaginalis over three times as broad as high, heavily sclerotized ductus bursae expanding over corpus bursae, which is about three times as long as sterigma.
Distribution. Gegenes nostrodamus from the western Mediterranean to India, but contrary to G. pumilio, has not been found in sub-Saharan Africa, except for a few Sudanian localities. It is widespread in NW Africa (Tennent 1996). In Europe, it has been found in Spain (García-Barros et al. 2013) and the southern half of Portugal (Aguiar et al. 2003), Italy (Verity 1940) and eastward through the Balkans and Greece (Rebel & Zerny 1931, Thurner 1964, Schaider & Jakšic´ 1989, Pamperis 2009, Verovnik & Popovic´ 2013). Detailed distribution data also provided by Tshikolovets (2011). It extends through Turkey (Hesselbarth et al. 1995), Syria (Evans 1949), Lebanon (Larsen 1974), Israel (Benyamini 1997), Jordania (Larsen 1974), Iraq (Wiltshire 1957, Tshikolovets et al. 2014) and Iran (Nazari 2003, north, central and east to Bojnurd, Tshikolovets et al. 2014) to Turkmenistan (Tshikolovets 1998), Uzbekistan (Evans 1949, Tshikolovets 2000), Tajikistan (Tshikolovets 2003), Kyrgyzstan (Tshikolovets 2005), Afghanistan (Sakai 1981), Pakistan (Evans 1949, Roberts 2001) and NW India (Evans 1949; he also mentioned “Bengal”, but since this is split between India and Pakistan, it is not clear from where exactly this material originated. Since Larsen 2004 does not mention it from Bangladesh, Evans’ listing may concern the Indian part of Bengal). It is rather widespread in the Arabian Peninsula (Larsen 1980, 1982b, 1983, 1984). In Egypt it is widespread in the Delta and the Nile Valley (Larsen 1990) and it has been found in the Sinai Peninsula (Benyamini 1984). The southernmost localities known are in Sudan: Khartoum (which may have been reached following the Nile valley; Larsen 1983), Kassala (Evans 1949) over 400 km east of Khartoum, and Wad Medani, ca 200 km southeast of Khartoum along the Blue Nile (collection Museum Leiden), by far the southernmost locality for the species overall.
Geographic variation. The populations from Afghanistan, Pakistan and NW India have been separated as subspecies karsana Moore, 1874 (type locality Rawalpindi) by Evans (1937), leaving the rest of the distribution area to the nominate subspecies (type locality “Barbaria”, i.e. NW Africa), but in 1949 he decided that the differences observed (subsp. karsana being “smaller, paler, no pinkish tinge”) were not meaningful or constant enough to justify a subspecies recognition, which conclusion we follow here.
Afrogegenes gen. n.
Type species. Hesperia hottentota Latreille, 1823.
Description/Diagnosis
Antennal shaft. Plain pale ochreous on underside.
Club. Eleven segments, of which two (rarely three) form the apiculus.
Wings. Length of forewing male 11.5−14(−15.1) mm, female 12−15.1 mm, upper side greyish or dark brown; male with at most very obscure ochreous spots in r3–5, m1–2 and m3 on forewing or spotless, more or less ochreous superscaling in basal part; female pale ochreous spots in spaces r3–5, m1–2, m3, cua1 and cua2 on forewing, series of suffused and partly obscure ochreous spots in (at most) spaces r4–cua2. Underside forewing male, greyish brown, with ochreous superscaling when fresh, suffused and obscure spots in spaces r3–5, m3 and cua1; female as upper side, but rather strong ochreous superscaling in apical third. Underside hindwing male and female, ochreous with small dark markings and patches indicating the edges of further disappeared discal spots.
Mid tibia. Usually 5, sometimes 6 or, as an exception, 7 long spines.
Male genitalia. In dorsal and ventral aspect tegumen + uncus (the two are difficult to separate) with parallel sides, but narrower in distal quarter. Uncus squarish apically, slightly indented, the two apical angles slightly protruding, shallow dorsal flaps before constricting to squarish apex. Gnathos narrow, elongate, slightly curved structures narrowly connected to uncus sides, medially separated but interconnected apically, forming two short diverging protrusions that, in dorsal view, peep out from under the uncus. Saccus much shorter than half length of tegumen + uncus. Valva more rounded than in Gegenes, proximal edge oblique, making an angle of at least 30° with ventral edge, costa straight to slightly curved in distal half, ventral edge slightly concave in proximal quarter, then curving to a more or less straight cucullus that projects well beyond costal end, proximal part of dorsal edge of cucullus with strongly toothed extension pointing proximo-distally. Aedeagus: vesica without cornuti, dorsal aperture of ductus seminalis roughly medially in pre-zonal part, post-zonal part short.
Female genitalia. Pre-vaginal plate entire, slightly indented at apex, densely spined, overlapping post-vaginal plate. Post-vaginal plate oblong with pointed ends. Ductus bursae entirely membranous and transparent. Corpus bursae oblong.
Traditionally, two species are recognized, A. letterstedti (generally known as niso) and A. hottentota. For reasons explained below we decided to raise Gegenes hottentota ocra Evans, 1937, considered a subspecies of either one of the two species, to species rank, raising the total number of species for this genus to three.
Afrogegenes letterstedti (Wallengren, 1857) comb. n.
Nomenclature. The nomenclatural history of the name is a bit tortuous. The species was known under the name Gegenes letterstedti (originally described in the genus Hesperia by Wallengren 1857: 49), until 1946 when Evans decided for reasons not mentioned that it should be replaced by Papilio niso Linnaeus, 1764, although in 1937 (p. 191) he had remarked “Neither Linnaeus’ description nor Clerck’s figure [see below] agree at all with letterstedti and the name cannot be applied with certainty to any African Hesperiid.” Sixty-four years before, Aurivillius (1882) had strongly advocated this synonymy, referring to (and presenting a copy of) an unpublished figure by Clerck. Holland (1896) disagreed with this synonymy, maintaining that the description by Linnaeus (1764) is “wholly inadequate”. In this he followed R. Trimen, expert in South African butterflies, who had expressed the same opinion in a letter to Aurivillius.
It remains obscure why Aurivillius referred to Clerck’s unpublished figure since Linnaeus did not. It is unknown where the specimen figured by Clerck came from. As Holland (1896: 58) put it: “The copies of Clerk’s figures given by Prof. Aurivillius do not carry conviction with them. They may apply to several other obscure African forms as well as to the species named by Latreille, and the description by Linnaeus is wholly inadequate. We shall forever be in the dark as to the species intended by Linnaeus.” We think it possible that Clerk’s figures represent a Gegenes (in the old sense), but without knowing its provenance and being a female, it could be either letterstedti or hottentota. However, since Linnaeus did not refer to it, Clerck’s figure is irrelevant for the correct identification of the species described by Linnaeus.
Linnaeus’ (1764: 339) description reads: “157. PAPILIO P[lebeji]. U[rbicolae]. alis integerrimis subreversis fuscis: primoribus supra punctis quattuor albis sparsis.” (“with wings with smooth edges, brown on the underside: on forewings with four white scattered dots.”). We can only agree with Holland in that it, indeed, could refer to a number of African Hesperiidae. However, it cannot refer to A. letterstedti, since in the males of this species the spots on the upper side are absent or, at most, obscure and ochreous, and the female has spots of a suffused ochreous tinge, and certainly not white. Linnaeus (1764) added that Papilio niso was very similar to “P. Tages” [a Palaearctic species described by Linnaeus (1758), at present considered to belong to the genus Erynnis], a species that only agrees with Hesperia letterstedti in being brown, be it another tinge of brown, and for the rest is very different in wing shape and otherwise. Even when totally worn, it is difficult to confuse Erynnis tages with any Gegenes.
Finally, in the 12th edition of Systema Naturae, Linnaeus (1767: 796) repeated the description and added: “Antennae apice acuto, uti P. Comma.” (“antennae with sharp apex, like P. Comma”). Once more it shows that Papilio niso and Hesperia letterstedti cannot be the same species. Although the apiculus of Hesperia comma is smalI, it is definitely longer (consisting of four segments) and seen as a sharp point to the antennal club, very different from the hardly visible apiculus of Hesperia letterstedti.
Honey & Scoble (2001) studied the types of the butterflies described by Linnaeus in detail. They found a female Gegenes (in the old sense) in the collection of the Linnean Society of London with a label “Niso” in Linnaeus’ handwriting and made it the lectotype of Papilio niso. This was an unfortunate decision, since the specimen does not match the description at all (in other words, it seems unlikely that Linnaeus had this specimen in front of him when describing niso) and its provenance is unknown. The assumption that it came from the Cape region is based on the other assumption that Linnaeus described this particular specimen when giving Cape of Good Hope as origin, which is unlikely since the description does not match. The attached label is no guarantee either, since Honey & Scoble (2001: 285) themselves remark that during subsequent curation the position of Linnean labels was probably changed twice. Since the specimen is female and females of A. letterstedt and A. hottentota cannot yet be separated with certainty, it could as well belong to A. hottentota (which is not known from the Cape of Good Hope Region), if it originated from a more northern locality.
Further, although the nominate subspecies of A. hottentota is not known from the Cape Region, Evans (1937) recognized a subspecies ocra that does occur at the Cape of Good Hope (and northward to the Delagoa Bay). As explained below, later authors (Pennington in Dickson & Kroon 1978, Henning et al. 1997) moved the taxon to letterstedti (called niso by them), but we maintain that ocra is a separate species, the female of which cannot yet be distinguished from either the female of letterstedti or of hottentota.
In summary, the specimen selected as lectotype of Papilio niso Linnaeus does not match with Linnaeus’ description and cannot be identified to a particular species, while labelling and provenance are not beyond doubt. Our conclusion can only be that the lectotype designation is invalid, Papilio niso is a nomen dubium as long as no acceptable type material has been found, and the name Hesperia letterstedti Wallengren, 1857, should be re-instated.
External characters. Length of forewing, male 11.6–15.1 mm (n = 25, mean = 12.9 mm, s = 0.5; see also under Geographic variation), female 12.5−15.1 mm (n = 25, mean = 13.0 mm, s = 1.04).
Male — Upper side forewing dark grey–brown, usually more ochreous near base, suffused small ochreous spots in at most spaces r3–5, m1–2, m3, but spots may be practically missing; upper side hind wing unspotted dark grey–brown; underside forewing more ochreous with suffused but well visible pale ochreous spots in spaces r3–5, m1–2, m3 and cua1; underside hindwing ochreous with some darker lines and patches, leaving a band of median spots in spaces r5–cua1 and (less evident) basal spots in space cua1, cell and space r4.
Female — Upper side forewing dark brown with pale ochreous spots in spaces 1b–8; upper side hindwing female dark brown with obscure pale ochreous spots in spaces 2–6; underside forewing dark brown, suffused ochreous in apical third, spots as on upper side; underside hindwing as in male.
Male genitalia. Valva 3.0 to 3.2 times as long as high. Ventral edge of cucullus slightly, but evenly convex. Apex of cucullus upturned. Proximal extension of cucullus relatively short (less than one third of dorsal length of cucullus) and tapering. Aedeagus: proximal end without prominent olecranon.
Female genitalia. See below, under A. ocra.
Distribution. Widely distributed throughout sub-Saharan Africa in open areas in forests, savannah, grassland, park and gardens, in central highlands of Kenya going as high as at least 3000 m; extending eastward to the Arabian Peninsula: Asir (Saudi Arabia), Yemen and Aden (politically part of Yemen) (Larsen 1983) and southward reaching the Cape Peninsula of South Africa (Evans 1937, Kielland 1990, Larsen 1991, 2005, Pennington in Pringle et al. 1994, Henning et al. 1997, Vande weghe 2010).
Geographic variation. Wallengren (1857) described Hesperia letterstedti in a paper on the butterflies collected by J.A. Wahlberg during his travels in South Africa (“Caffraria”) in the years 1838–1845. Kaffraria was a huge area stretching along the southeast coast of South Africa and situated in the eastern part of what at present is known as the Eastern Cape Province. There are no indications of when and where exactly the type material was collected. Evans (1937) recognized two subspecies, the nominate subspecies and brevicornis Plötz. The latter was described by Plötz (1884) as a new species from Angola (in the genus Thymelicus). Although the description is clear enough to recognize an A. letterstedti or A. hottentota, it is of a female and it is impossible to pinpoint to which of the two the name refers. In other words, it is a nomen dubium, and the only way to preserve the name is to designate an undoubted A. letterstedti from Angola as neotype. However, we are not sure it is worth the attempt. Apart from the nomenclatural aspect, the only difference between the subspecies (according to Evans, length of forewing in letterstedti 14 mm, in brevicornis 13 mm) shows much overlap. The forewing of the smallest and largest specimens among 30 males from S Africa and Swaziland (letterstedti) varied from 12.5 to 13.7 mm with outliers of 11.6 and 14.7 mm, while in over 50 specimens from Kenya the length varied from 12.6 to 13.7 mm, with an outlier of 11.7 mm (all specimens in the collection of Naturalis Biodiversity Center, Leiden). As a consequence, we do not see any value in recognizing brevicornis as a separate subspecies.
Afrogegenes hottentota (Latreille, 1823) comb. n.
External characters. Length of forewing, male 11.5−13.8 mm (n = 25, mean = 12.6 mm, s = 0.56); since females cannot be separated from females of A. letterstedti with certainty we give here the same measurements as for the latter: 12.5−15.1 mm (n = 25, mean = 13.0 mm, s = 1.04).
Male — Upper side forewing dark yellow in basal two-thirds, dark brown in outer third and at apex, no traces of apical spots, large black subtornal patch of specialized scales, presumably androconia; upper side hindwing as forewing, unspotted, without the black patch; underside forewing ochreous, dark brown in basal third, large pale ochreous patch in space cua2 that is suffused outwardly, ochreous spots in spaces r3–5, m1–2, m3 and cua1.
Female — As female A. letterstedti.
Male genitalia. Valva 2.75 times as long as high. Ventral edge of cucullus slightly concave or at most straight. Apex of cucullus upturned. Proximal extension of cucullus at least one third of dorsal length of cucullus and slenderer than in A. letterstedti, with practically parallel sides. Aedeagus: proximal end without olecranon.
Female genitalia. See below, under A. ocra.
Distribution. Widely distributed throughout sub-Saharan Africa in short grasslands and along forest edges, wetlands and gullies, not as far into the driest zones as G. letterstedti (Evans 1937, Dickson & Kroon 1978, Kielland 1990, Larsen 1991, 2005, Pringle et al. 1994, Henning et al. 1997, Vande weghe 2010), extending to the Arabian Peninsula: Asir (Saudi Arabia), Yemen and Aden (politically part of Yemen) (Larsen 1983); not further south than the former South African provinces Transvaal (Gauteng-Mpupalanga) and Natal (Kwazulu-Natal), and “homeland” Transkei (at present in Eastern Cape Province) (Henning et al. 1997).
Afrogegenes ocra (Evans, 1937) comb. n.
External characters. Male — As male of A. hottentota, but large black patch on upper side forewing missing; about same size, 12.5−14.8 mm (n = 7, mean = 13.5 mm, s = 1.9), but the material available is too limited to get a fair idea of the variation.
Female — Not known for sure.
Male genitalia. Valva 2.75 times as long as high. Ventral edge of cucullus slightly concave or at most straight. Apex of cucullus straight. Proximal extension of cucullus at least one third of dorsal length of cucullus and slenderer than in A. letterstedti, with practically parallel sides. Aedeagus: proximal end with prominent olecranon (a flat, rounded extension at the coecum of the aedeagus).
Female genitalia. See below.
Distribution. A.ocra has a restricted range, from Cape of Good Hope, South Africa, to southern Mozambique. Evans (1937) described it as a subspecies of A. hottentota, even though it overlapped with the latter according to the original description (both taxa recorded from Natal and Delagoa Bay).
Justification for species status. Overlap without interbreeding would indicate separate species, unless the difference was due to an individual variation, found in only part of the population. This seems unlikely for an androconial patch, since it must deeply influence display behaviour and thus the presence or absence of androconia may act as an isolating mechanism. We checked whether A. ocra possibly had androconia with the same color as the rest of the wing, but this was not the case.
The problem seemed solved, when Pennington, in Dickson & Kroon (1978), decided that ocra was a form of G. niso (i.e. A. letterstedti). This concept was followed by Henning et al. (1997), who saw ocra as a slightly yellower form that becomes more prevalent in the more southern populations.
Earlier, Kroon (1973), referring to Evans (1937), had taken up the challenge to find differences in the male genitalia between A. niso (i.e. A. letterstedti) and A. hottentota and he did find and illustrate them. Remarkably, together with C.G.C. Dickson, Kroon edited the first edition of Pennington (1978), but did not refer to his earlier paper and changed the allocation of ocra for unknown reasons, maybe as a tribute to Pennington who died before he could finish his book. This allocation was followed in the second edition of Pennington (Pringle et al. 1994) and by Henning et al. (1997). When we checked the genitalia of a male of ocra, we found them close to the type described and illustrated by Kroon for A. hottentota and not for A. niso (i.e. A. letterstedti). As a consequence we had little choice but to raise ocra to species level. We have checked males from the following localities: A. letterstedti — Kenya (4), Swaziland (2), A. hottentota — Kenya (2), Tanzania (2), Swaziland (2), A. ocra — South Africa (1), with results as given above.
While the males of the three species are easily separated by external characters, the females pose a problem. According to Evans (1937) the discal band on the underside of the hindwing is constricted along vein 6 in A. letterstedti, and without constriction in A. hottentota, but we can only agree with Larsen (2005), that the two cannot be separated for sure externally. Evans’ justification for the identity of the females was the presence of two pairs taken in copula in the British Museum, one of each of the species. Obviously, this smallest possible sample size is not enough for evaluation of the constancy in such a minute character. Evans (1937) added that “The genitalia [Evans only studied male genitalia] of hottentota and niso [i.e. A. letterstedti] seem alike”, which was not correct either, as was shown by Kroon (1973).
Evans (1937) justly realized that for a correct identification of females recourse must be found in pairs taken in copula. Apart from the two pairs mentioned by Evans, we do not know of any other pairs. So, as for the females, we are faced with three species, which have very similar females. We dissected the genitalia of twelve females, from the following localities: Kenya (5), Tanzania (1), Swaziland (3), South Africa (2), and Yemen (1), hoping to find at least two different types. In fact, we found two types of pre-vaginal plate, type A (Figs 53−58) shallowly indented giving a weakly bilobed lamella, and type B (Figs 59−64) more deeply indented so that the lamella appears more strongly bilobed. Both types were found in Kenya, Swaziland and South Africa, the single specimen from Tanzania had a type A lamella, the only specimen from Yemen had a type B lamella. Since A. letterstedti and A. hottentota are both known from Yemen (Larsen 1983), the specimen could belong to either species. So even if this variation is specific, we do not yet know which type belongs to which species or if two or three species were dissected.
Concluding remarks
Since the genus has a distribution area covering three continents, Europe, Asia and Africa, and relevant literature never dealt with the three areas together, the marked differences between G. pumilio and G. nostrodamus on the one hand, and the purely African species on the other, in external characters as well as in male and female genitalia, has not received sufficient attention so far. The differences have led us to distinguish two genera, Gegenes and Afrogegenes. Examination of the male genitalia showed A. ocra to be different from both A. hottentota and A. letterstedti. While the females of the Gegenes species can be separated by their genitalia, discrimination of the females of the three species of Afrogegenes is yet uncertain; although slightly different types of female genitalia have been found they cannot be associated with species with certainty due to a lack of males and females found in copula, or mixed sexes reared from single females. It is even possible that all females examined belonged to a single species, and that we have not seen the females of two of the three species involved. The other important but still unanswered question is where the boundary lies between the populations of G. pumilio with haploid chromosome number 24 and those with 41. To answer this question fresh material or material freshly put in a preservative from crucial localities, like C. Italy, Greece, W. and E. Africa is needed. The answer has nomenclatural consequences. In short, as usual in taxonomic studies, more material is needed.
We are greatly indebted to Torben Larsen, who generously shared his extensive knowledge of African Hesperiidae with us. His premature death on 21 May 2015 came as a shock. The senior author is grateful to Hein Boersma (St Nicolaasga, Netherlands), Henk Geertsema (Stellenbosch, South Africa) and Jan Lourens (Manilla, Philippines) for providing material from various African localities and for showing him around in Ghana, Cameroon, Kenya and South Africa. Two reviewers made critical remarks that greatly improved the text in places.
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