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
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 descriptions 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
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 deeply indented medially entire, slightly indented pre- and post-vaginal plates not overlapping overlapping ductus bursae heavily sclerotized membranous corpus bursae rounded oblong
description did not match with Hübner’s own fig- cucullus sharply curving up distally, apex partly or ures, which were of Papilio pygmaeus Cyrilli, 1787. entirely overlapping distal costal process, upper edge Butler (1870) fixed the latter as genotype, but since of cucullus without proximal extension. Aedeagus: Cyrilli’s name was a junior homonym of Fabricius’ vesica with two pronounced cornuti, dorsal aperture name, it had to be replaced by Papilio pumilio Hoff- of ductus seminalis near proximal end of pre-zonal mansegg, 1804. part, post-zonal part long. Female genitalia. Pre-vaginal plate deeply indented medially, smooth, not overlapping post-vaginal Description/Diagnosis plate. Post-vaginal plate oblong with rounded ends. Antennal shaft. Underside plain creamy white or Ductus bursae heavily sclerotized. Corpus bursae lightly chequered. rounded. Club. Eleven segments, one of which is the tiny The two species traditionally recognized, G. pum- apiculus. ilio and G. nostrodamus, are not easily distinguished Wings. Upper side brown to very dark brown, may externally, particularly in the female sex (see Coutsis be paler outwardly; males unspotted, females with & Olivier 1993 for differences). For this reason, the variable number of pale ochreous spots in, at most, second author studied the previously undocumented spaces r3–5, m1–2, m3, cua1, cua2. Underside female genitalia (Coutsis 2012). This is the more im- forewing with spots as on upper side; underside portant since the two species overlap geographically, hindwing brown with more or less dense greyish although they are rarely found in the same locality, superscaling (overlaying scales), discal spots more or and part of the older literature records may be based less indicated. on confusion between the two. Lorkovic´ (1971) de- Mid tibia. Usually 6 long spines, but number may scribed an easily visible character to distinguish the vary from five to eight. two species in spread specimens (see the descriptions Male genitalia. In dorsal and ventral aspect tegumen of the species) but his short note was largely over- + uncus (the two are difficult to separate) constricted looked. Gegenes pumilio and G. nostrodamus have dif- just before middle, strongly expanding in distal half. ferent ecological preferences, the first being mainly Uncus deeply indented, the widely separated lateral found in open Mediterranean garrigue (a type of arms with shallow dorsal flap and irregularly formed low, soft-leaved scrubland) and dried out river beds, apex. Gnathos consisting of two large, smooth, often near the coast, while G. nostrodamus is more more or less reniform structures, medially widely shade-loving, frequenting areas with scattered trees, separated, in ventral view superimposed on uncus and may occur far inland, as in Spain (García-Barros arms, ventrally broadly and imperceptibly passing et al. 2013). into uncus sides. Saccus relatively long, more than half the length of tegumen + uncus. Valva elongate, Gegenes pumilio (Hoffmansegg, 1804) proximal edge about vertical, costa almost as long Figs 1–9, 18–25, 28 as ventral edge and broadly parallel to it, more or less straight, slightly curving up distally, ventral edge External characters. Antennal shaft chequered on un- nearly straight to slightly concave in proximal half, derside: apical part of segments darker than basal Downloaded from Brill.com09/27/2021 04:06:06PM via free access
Fig. 1. Gegenes pumilio, type species of Gegenes Hübner, 1819. SW Turkey, Bodrum, 18.iv.2016, R. de Jong.
Figs 2–17. Upper (even numbers) and under sides (odd numbers) of Gegenes species. — 2, 3, G. pumilio pumilio, male, Italy, Lazio, Sperlonga, 4.x.1975, leg. T. Racheli; 4, 5, G. pumilio pumilio, female, Italy, Formia, Monti Au- runci, 16–30.x.1910; 6, 7, G. pumilio gambica, male, Madagascar, Ft. Dauphin, 14.iv.1952, leg. E.W. Diehl; 8, 9, G. pumilio pumilio, female, Pakistan, Rawalpindi, 19–20.vi.1981, leg. W.L. Blom; 10, 11, G. nostrodamus, male, Greece, Ahaía, Río (Pátra), 18.vii.1983, leg. A. Riemis; 12, 13, G. nostrodamus, female, Spain, Granada, Salobrena del Mar, 9.vi.1973, leg. J.C. Weiss; 14, 15, G. nostrodamus, male, Iran, Isfahan, Yazdabad, 12–13.v.1977, leg. W.L. Blom; 16, 17, G. nostrodamus, male, Sudan, W Medani, 1.xii.1981, leg. J.H. Lourens. All figures natural size. part. Length of forewing male 13.1−16.9 mm (n = female, indistinct spots on forewing as on upper side, 25, mean = 13.8 mm, s = 0.6) , female 13.8−16.6 hindwing also with some indistinct spots. Forewing mm (n = 25, mean = 14.0 mm, s = 0.7). Upper side underside with densely packed dark brown scales in dark blackish brown, forewing unmarked in male, basal part of discoidal cell clearly visible in spread with obscure spots in spaces 1b, 2 and 3, and at specimens (Lorkovic´ 1971) (in butterflies and skip- apex in female. Underside pale grey–brown with, in pers this patch of specialized scales is part of a wing Downloaded from Brill.com09/27/2021 04:06:06PM via free access
Figs 18–19. Male genitalia of Gegenes pumilio gambica. Left to right, upper part: inside and outside of right valva saccus, underside and upper side of tegumen + uncus; lower part: lateral view and dorsal view of aedeagus, right lateral view of tegumen + uncus. — 18, Tchad, Moundou, Bebedjia, 12.ix.1970, leg. J.H. Lourens, prep. no. 5349; 19, Madagascar, Ft Dauphin, 14.iv.1952, leg. E.W. Diehl, prep. no. 5346. Downloaded from Brill.com09/27/2021 04:06:06PM via free access
Figs 20–23. Gegenes pumilio pumilio, male genitalia, variation in dentation of valva. — 20, Tunisia, Tabarka, 17.iv.1990, prep. no. 3001; 21, France, Cannes, La Boca, 30.viii.1945, prep. no. 2999; 22, Italy, Messina, Isole Li- pari, 12.viii.1991, prep. no. 3000; 23, Spain, Mallorca, Sta Marico, 15.vii.1982, prep. no. 2998. All in coll. Coutsis.
Figs 24–27. Female genitalia of Gegenes. — 24, 25, G. pumilio: 24, Greece, Sími Island, 24.iv.2010; 25, Greece, Spétses Island, 24.ix.1979; 26, 27, G. nostrodamus: 26, Greece, Spétses Island, 10.ix.2001; 27, Greece, Attikí, Ekáli, 25.viii.1963. All in coll. Coutsis. coupling mechanism, see Kuijten 1974), not over- proximal quarter or third, distal two-thirds or three- lapped by the fringe of fine short hairs along the bas- quarters unevenly dentate. Aedeagus: vesica with two al part of the front edge of the hindwing underside cornuti of even size, left one dentate in side view, (compare G. nostrodamus). This character occurs in right one plain. both sexes. It is visible in Figs 28 and 29. In honour In order to find possible differences between G. of the discoverer we call this character the “Lorkovic´ pumilio and G. gambica we examined the genitalia character”. of specimens from France (Cannes), Italy (Lipari Male genitalia. Saccus: length about 3/4 of vertical island, north of Sicily) and Spain (Mallorca) in Eu- length of vinculum. Valva: cucullus relatively short, rope, Tunisia (Tabarka) in NW Africa, and Chad scarcely overlapping costa; upper margin smooth in (Bebedjia), Kenya (Arabuko Forest) and Madagascar Downloaded from Brill.com09/27/2021 04:06:06PM via free access
Figs 28–29. Lorkovic´ character. — 28, G. pumilio (detail of Fig 5); 29, G. nostrodamus (detail of Fig 13).
(Taolagnaro) in sub-Saharan Africa, in addition to some 800 km between SE Iran and the Pakistan/NW specimens from numerous Greek localities. They India localities. In addition, the species is found in S were all similar and although there was some slight and SE Arabian Peninsula (Yemen, Dhofar, Oman; individual variation, there was no indication of geo- Larsen 1980, 1982b, 1983, 1984, Larsen & Larsen graphical variation. 1980). Detailed distribution data also provided by Female genitalia. Lamella antevaginalis deeply di- Kudrna et al. (2011) and Tshikolovets (2011). It oc- vided into a bifid structure with two narrow arms, curs throughout sub-Saharan Africa (Evans 1937, lamella postvaginalis slightly more than twice as Kielland 1990, Larsen 1991, 2005), including the is- broad as high, heavily sclerotized ductus bursae of land of Socotra east of the Horn of Africa, politically even width throughout, and corpus bursae less than belonging to Yemen (Rebel 1907, Evans 1937), in twice as long as sterigma. suitable habitats, but absent from most of Cape Prov- Distribution. G. pumilio is widely distributed in ince, South Africa, (Dickson & Kroon 1978, Pringle scattered populations in NW Africa (Algeria and et al. 1994, Henning et al. 1997). Not mentioned Tunisia; Tennent 1996). Europe: Spain, Mallorca from Gabon (Vande weghe 2010) and Cameroon (García-Barros et al. 2013; according to these authors (personal data). It has not been recorded from Mada- not known from the Spanish mainland, although gascar so far, but there is a single specimen in the col- mentioned by Evans (1937, 1949), SE France lection of the Naturalis Biodiversity Center, Leiden, (Lafranchis 2000) and eastward through Italy and from Ft Dauphin ( = Taolagnaro) (Figs 6–7, 19). Sicily (Verity 1940) to the Balkans and Greece (Reb- Geographic variation. Gegenes pumilio has been di- el & Zerny 1931, Thurner 1964, Schaider & Jaksic´ vided into three subspecies (Evans 1937, 1949), a 1989, Pamperis 2009, Verovnik & Popovic´ 2013). northern one, north of the Sahara and east to NW Further east it occurs in Turkey (Hesselbarth et al. India (the nominal subspecies), a southern one 1995), Cyprus (Makris 2001), Syria (Evans 1949), throughout Africa (ssp. gambica Mabille) and the Lebanon (Larsen 1974), Israel (Benyamini 1997), third (ssp. monochroa Rebel) restricted to the island Iraq (Wiltshire 1957, Tshikolovets et al. 2014) and of Socotra. Ssp. gambica differs from the nominate Iran (Evans 1949, Nazari 2003, Tshikolovets et al. form in having more prominent markings on the un- 2014, mainly along the Persian Gulf) to Pakistan derside of the hindwing and reduced spots in the fe- (not mentioned by Roberts 2001, but records from male forewing. Ssp. monochroa is smaller and darker Chitral, Waziristan and Khyber in Evans 1949); with barely visible spots. latter locality could as well be Afghanistan, but not It has been found, however, that the chromo- mentioned by Sakai (1981) from that country; there some numbers are different in the western and is a female from Rawalpindi in the collection of the eastern Mediterranean. De Lesse (1960, 1967) re- Naturalis Biodiversity Center (Figs 8, 9), and NW ported a haploid chromosome number of n = 24 for India (Punjab, NW Himalayas; Evans 1949). Not specimens from Nice and Algiers, and n = 41 for a recorded from Turkmenistan, Uzbekistan, Tajiki- specimen from Gülek near Adana, Turkey. Larsen stan or Kyrgyzstan (Tshikolovets 1998, 2000, 2003, (1982b) confirmed the latter number for specimens 2005), where the very similar G. nostrodamus is wide- from Beirut and Yemen. In view of the huge distri- spread, and (most of) Afghanistan, leaving a gap of bution area of the species, the importance of such Downloaded from Brill.com09/27/2021 04:06:06PM via free access
Fig. 30. Gegenes nostrodamus, male genitalia; Greece, Kríti, nr Haniá, 16.iv.1990, leg. et coll. A. Olivier, prep. no. 2016. Inside and outside of right valva, upper side tegumen + uncus, left lateral view of tegumen + uncus, ventral view of top of aedeagus, lateral view of complete aedeagus.
basal part of hindwing underside (Lorkovic´ 1971) over corpus bursae, which is about three times as (the Lorkovic´ character, see under G. pumilio). Ob- long as sterigma. viously, this character is only helpful in specimens Distribution. Gegenes nostrodamus from the western in which the hairs or scales at the frontal edge of the Mediterranean to India, but contrary to G. pumilio, hindwing have not been rubbed off by age or rough has not been found in sub-Saharan Africa, except for handling. It is found in both sexes. See Figs 28 and 29. a few Sudanian localities. It is widespread in NW Af- Male genitalia. Saccus as long as vertical length of rica (Tennent 1996). In Europe, it has been found in vinculum. Valva: cucullus longer than in G. pumilio, Spain (García-Barros et al. 2013) and the southern more broadly overlapping costa and distally extend- half of Portugal (Aguiar et al. 2003), Italy (Verity ing beyond it, upper margin evenly dentate, also in 1940) and eastward through the Balkans and Greece proximal part. Aedeagus: vesica with two cornuti of (Rebel & Zerny 1931, Thurner 1964, Schaider & uneven size, left one small and dentate, right one Jakšic´ 1989, Pamperis 2009, Verovnik & Popovic´ much larger and without teeth. 2013). Detailed distribution data also provided by Female genitalia. Lamella antevaginalis with inden- Tshikolovets (2011). It extends through Turkey tion just over middle leaving two broad parts, hardly (Hesselbarth et al. 1995), Syria (Evans 1949), higher than wide, together covering most of sterig- Lebanon (Larsen 1974), Israel (Benyamini 1997), ma, lamella postvaginalis over three times as broad Jordania (Larsen 1974), Iraq (Wiltshire 1957, Tshi- as high, heavily sclerotized ductus bursae expanding kolovets et al. 2014) and Iran (Nazari 2003, north, Downloaded from Brill.com09/27/2021 04:06:06PM via free access
Figs 31–34. Gegenes nostrodamus, male genitalia, variation in dentation of valva. — 31, Spain, Nerja Malaga, 13- 23.v.1990, prep. No. 2989; 32, Israel, Ein Gedi, 12.iv.1996, prep. No. 2996; 33, Italy, Sicily, Monti Le Madonie, Palermo, 16.vii.1991, prep. No. 2990; 34, Morocco, Taroudannt, 4.iv.1989, prep. No. 2992. All in coll. Coutsis. central and east to Bojnurd, Tshikolovets et al. 2014) Description/Diagnosis to Turkmenistan (Tshikolovets 1998), Uzbekistan Antennal shaft. Plain pale ochreous on underside. (Evans 1949, Tshikolovets 2000), Tajikistan (Tshi- Club. Eleven segments, of which two (rarely three) kolovets 2003), Kyrgyzstan (Tshikolovets 2005), form the apiculus. Afghanistan (Sakai 1981), Pakistan (Evans 1949, Wings. Length of forewing male 11.5−14(−15.1) Roberts 2001) and NW India (Evans 1949; he mm, female 12−15.1 mm, upper side greyish or dark also mentioned “Bengal”, but since this is split be- brown; male with at most very obscure ochreous tween India and Pakistan, it is not clear from where spots in r3–5, m1–2 and m3 on forewing or spot- exactly this material originated. Since Larsen 2004 less, more or less ochreous superscaling in basal part; does not mention it from Bangladesh, Evans’ listing female pale ochreous spots in spaces r3–5, m1–2, may concern the Indian part of Bengal). It is rather m3, cua1 and cua2 on forewing, series of suffused widespread in the Arabian Peninsula (Larsen 1980, and partly obscure ochreous spots in (at most) spaces 1982b, 1983, 1984). In Egypt it is widespread in r4–cua2. Underside forewing male, greyish brown, the Delta and the Nile Valley (Larsen 1990) and it with ochreous superscaling when fresh, suffused and has been found in the Sinai Peninsula (Benyamini obscure spots in spaces r3–5, m3 and cua1; female 1984). The southernmost localities known are in as upper side, but rather strong ochreous superscal- Sudan: Khartoum (which may have been reached ing in apical third. Underside hindwing male and following the Nile valley; Larsen 1983), Kassala (Ev- female, ochreous with small dark markings and ans 1949) over 400 km east of Khartoum, and Wad patches indicating the edges of further disappeared Medani, ca 200 km southeast of Khartoum along the discal spots. Blue Nile (collection Museum Leiden), by far the Mid tibia. Usually 5, sometimes 6 or, as an exception, southernmost locality for the species overall. 7 long spines. Geographic variation. The populations from Af- Male genitalia. In dorsal and ventral aspect tegu- ghanistan, Pakistan and NW India have been men + uncus (the two are difficult to separate) with separated as subspecies karsana Moore, 1874 (type parallel sides, but narrower in distal quarter. Uncus locality Rawalpindi) by Evans (1937), leaving the squarish apically, slightly indented, the two apical rest of the distribution area to the nominate subspe- angles slightly protruding, shallow dorsal flaps before cies (type locality “Barbaria”, i.e. NW Africa), but in constricting to squarish apex. Gnathos narrow, elon- 1949 he decided that the differences observed (subsp. gate, slightly curved structures narrowly connected karsana being “smaller, paler, no pinkish tinge”) were to uncus sides, medially separated but interconnect- not meaningful or constant enough to justify a sub- ed apically, forming two short diverging protrusions species recognition, which conclusion we follow here. that, in dorsal view, peep out from under the uncus. Afrogegenes gen. n. Saccus much shorter than half length of tegumen + Type species. Hesperia hottentota Latreille, 1823. uncus. Valva more rounded than in Gegenes , proximal Downloaded from Brill.com09/27/2021 04:06:06PM via free access
Figs 35–50. Upper (odd numbers) and under sides (even numbers) of Afrogegenes species. — 35, 36, A. letterstedti, male, Kenya, Maralal, 31.x–6.xi.1975, leg. R. de Jong; 37, 38, A. letterstedti, female, Ethiopia, Jimma, 16–24. xi.1969, leg. R.H. Cobben; 39, 40, A. letterstedti, male, Swaziland, Matsapha, 30.v.1991, leg. J.A.W. Lucas; 41, 42, A. letterstedti, female, Swaziland, Matsapha, 30.v.1991, leg. J.A.W. Lucas; 43, 44, A. hottentota, male, Cameroun, Kribi, 11.vii.1974, leg. J.A.W. Lucas; 45, 46, A. hottentota, female, Cameroun, Kribi, 11.vii.1974, leg. J.A.W. Lucas; 47, 48, A. ocra, male, S Africa, Port Elizabeth, 12.xi.1938, leg. Clarck; 49, 50, A. ocra, male, S Africa, Cape Town, 13.x.1966, leg. B. Stuart. All figures natural size. edge oblique, making an angle of at least 30° with Afrogegenes letterstedti (Wallengren, 1857) ventral edge, costa straight to slightly curved in distal comb. n. half, ventral edge slightly concave in proximal quar- Figs 35–42, 51–64 ter, then curving to a more or less straight cucullus that projects well beyond costal end, proximal part Nomenclature. The nomenclatural history of the of dorsal edge of cucullus with strongly toothed ex- name is a bit tortuous. The species was known under tension pointing proximo-distally. Aedeagus: vesica the name Gegenes letterstedti (originally described in without cornuti, dorsal aperture of ductus seminalis the genus Hesperia by Wallengren 1857: 49), until roughly medially in pre-zonal part, post-zonal part 1946 when Evans decided for reasons not mentioned short. that it should be replaced by Papilio niso Linnaeus, Female genitalia. Pre-vaginal plate entire, slightly 1764, although in 1937 (p. 191) he had remarked indented at apex, densely spined, overlapping post- “Neither Linnaeus’ description nor Clerck’s figure vaginal plate. Post-vaginal plate oblong with pointed [see below] agree at all with letterstedti and the name ends. Ductus bursae entirely membranous and trans- cannot be applied with certainty to any African Hes- parent. Corpus bursae oblong. periid.” Sixty-four years before, Aurivillius (1882) Traditionally, two species are recognized, A. letter- had strongly advocated this synonymy, referring to stedti (generally known as niso) and A. hottentota. For (and presenting a copy of) an unpublished figure by reasons explained below we decided to raise Gegenes Clerck. Holland (1896) disagreed with this synon- hottentota ocra Evans, 1937, considered a subspe- ymy, maintaining that the description by Linnaeus cies of either one of the two species, to species rank, (1764) is “wholly inadequate”. In this he followed R. raising the total number of species for this genus to Trimen, expert in South African butterflies, who had three. expressed the same opinion in a letter to Aurivillius. Downloaded from Brill.com09/27/2021 04:06:06PM via free access
Figs 51–52. Male genitalia of Afrogegenes letterstedti. Left to right, upper part: inside and outside of right valve, sac- cus, underside and upper side of tegumen + uncus; lower part: lateral view and dorsal view of aedeagus, right lateral view of tegumen + uncus. — 51, Kenya, Thika, Fourteen Falls, 12.v.1974, leg. J.H. Lourens, prep. no. 5342; 52, Swaziland, Simunye Country Club, 1.vi.1991, leg. J.A.W. Lucas, prep. no. 5348.
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Figs 53–64. Ventral view of female genitalia of Afrogegenes letterstedti or hottentota. — 53–58, type A: 53, Swazi- land, Manzini, 7.v.1991, leg. J.A.W. Lucas, prep. no. 5350; 54, Kenya, Taita Distr., Wundanyi, 16–17.x.1974, leg. R. de Jong, prep. no. 5355; 55, Tanzania, Mufindi, Lugoda, 14.viii.1982, leg. T.C.E. Congdon, prep. no. 5354; 56, Swaziland, Manzini, 29.v.1991, leg. J.A.W. Lucas, prep. no. 5361 (lamella antevaginalis only); 57, Kenya, Mt Kenya, Naro Moru Route, 26.i.1971, leg. J.H. Lourens, prep. no. 5363 (lamella antevaginalis only); 58, S Africa, Umfolozi, 22.ii.1977, leg. A.T. Gay, prep. no. 5366 (lamella antevaginalis only). 59–64, type B: 59, Kenya, Kaka- mega Forest, 26–28.x.1974, leg. R. de Jong, prep. no. 5353; 60, Kenya, Maralal, 31.x–6.xi.1978, leg. R. de Jong, prep. no. 5351; 61, S Africa, Natal, Monaux Sources, 22.v.1991, leg. J.A.W. Lucas, prep. no. 5352; 62, Kenya, Naro Moru, 27–28.xi.1978, leg. R. de Jong, prep. no. 5365 (lamella antevaginalis only); 63, Yemen, El Udayn, 2.x.1988, prep. no. 5364 (lamella antevaginalis only); 64, Swaziland, Manzini, 20.v.1991, leg. J.A.W. Lucas, prep. no. 5362 (lamella antevaginalis only). For an explanation of type A and type B, see text.
It remains obscure why Aurivillius referred to subreversis fuscis: primoribus supra punctis quat- Clerck’s unpublished figure since Linnaeus did not. tuor albis sparsis.” (“with wings with smooth edges, It is unknown where the specimen figured by Clerck brown on the underside: on forewings with four came from. As Holland (1896: 58) put it: “The cop- white scattered dots.”). We can only agree with ies of Clerk’s figures given by Prof. Aurivillius do Holland in that it, indeed, could refer to a number not carry conviction with them. They may apply to of African Hesperiidae. However, it cannot refer several other obscure African forms as well as to the to A. letterstedti, since in the males of this species species named by Latreille, and the description by the spots on the upper side are absent or, at most, Linnaeus is wholly inadequate. We shall forever be obscure and ochreous, and the female has spots of in the dark as to the species intended by Linnaeus.” a suffused ochreous tinge, and certainly not white. We think it possible that Clerk’s figures represent a Linnaeus (1764) added that Papilio niso was very Gegenes (in the old sense), but without knowing its similar to “P. Tages” [a Palaearctic species described provenance and being a female, it could be either by Linnaeus (1758), at present considered to be- letterstedti or hottentota. However, since Linnaeus long to the genus Erynnis], a species that only did not refer to it, Clerck’s figure is irrelevant for agrees with Hesperia letterstedti in being brown, be the correct identification of the species described by it another tinge of brown, and for the rest is very Linnaeus. different in wing shape and otherwise. Even when Linnaeus’ (1764: 339) description reads: “157. totally worn, it is difficult to confuseErynnis tages PAPILIO P[lebeji]. U[rbicolae]. alis integerrimis with any Gegenes. Downloaded from Brill.com09/27/2021 04:06:06PM via free access
Finally, in the 12th edition of Systema Naturae, Male — Upper side forewing dark grey–brown, Linnaeus (1767: 796) repeated the description and usually more ochreous near base, suffused small added: “Antennae apice acuto, uti P. Comma.” (“an- ochreous spots in at most spaces r3–5, m1–2, m3, tennae with sharp apex, like P. Comma”). Once more but spots may be practically missing; upper side it shows that Papilio niso and Hesperia letterstedti hind wing unspotted dark grey–brown; underside cannot be the same species. Although the apiculus of forewing more ochreous with suffused but well vis- Hesperia comma is smalI, it is definitely longer (con- ible pale ochreous spots in spaces r3–5, m1–2, m3 sisting of four segments) and seen as a sharp point and cua1; underside hindwing ochreous with some to the antennal club, very different from the hardly darker lines and patches, leaving a band of median visible apiculus of Hesperia letterstedti. spots in spaces r5–cua1 and (less evident) basal spots Honey & Scoble (2001) studied the types of the in space cua1, cell and space r4. butterflies described by Linnaeus in detail. They Female — Upper side forewing dark brown with found a female Gegenes (in the old sense) in the col- pale ochreous spots in spaces 1b–8; upper side hind- lection of the Linnean Society of London with a wing female dark brown with obscure pale ochreous label “Niso” in Linnaeus’ handwriting and made it spots in spaces 2–6; underside forewing dark brown, the lectotype of Papilio niso. This was an unfortunate suffused ochreous in apical third, spots as on upper decision, since the specimen does not match the de- side; underside hindwing as in male. scription at all (in other words, it seems unlikely that Male genitalia. Valva 3.0 to 3.2 times as long as high. Linnaeus had this specimen in front of him when de- Ventral edge of cucullus slightly, but evenly convex. scribing niso) and its provenance is unknown. The as- Apex of cucullus upturned. Proximal extension of sumption that it came from the Cape region is based cucullus relatively short (less than one third of dorsal on the other assumption that Linnaeus described this length of cucullus) and tapering. Aedeagus: proximal particular specimen when giving Cape of Good Hope end without prominent olecranon. as origin, which is unlikely since the description does Female genitalia. See below, under A. ocra. not match. The attached label is no guarantee either, Distribution. Widely distributed throughout sub- since Honey & Scoble (2001: 285) themselves re- Saharan Africa in open areas in forests, savannah, mark that during subsequent curation the position of grassland, park and gardens, in central highlands of Linnean labels was probably changed twice. Since the Kenya going as high as at least 3000 m; extending specimen is female and females of A. letterstedt and eastward to the Arabian Peninsula: Asir (Saudi Ara- A. hottentota cannot yet be separated with certainty, bia), Yemen and Aden (politically part of Yemen) it could as well belong to A. hottentota (which is not (Larsen 1983) and southward reaching the Cape known from the Cape of Good Hope Region), if it Peninsula of South Africa (Evans 1937, Kielland originated from a more northern locality. 1990, Larsen 1991, 2005, Pennington in Pringle Further, although the nominate subspecies of et al. 1994, Henning et al. 1997, Vande weghe A. hottentota is not known from the Cape Region, 2010). Evans (1937) recognized a subspecies ocra that does Geographic variation. Wallengren (1857) described occur at the Cape of Good Hope (and northward to Hesperia letterstedti in a paper on the butterflies col- the Delagoa Bay). As explained below, later authors lected by J.A. Wahlberg during his travels in South (Pennington in Dickson & Kroon 1978, Henning Africa (“Caffraria”) in the years 1838–1845. Kaf- et al. 1997) moved the taxon to letterstedti (called fraria was a huge area stretching along the southeast niso by them), but we maintain that ocra is a separate coast of South Africa and situated in the eastern part species, the female of which cannot yet be distin- of what at present is known as the Eastern Cape Prov- guished from either the female of letterstedti or of ince. There are no indications of when and where hottentota. exactly the type material was collected. Evans (1937) In summary, the specimen selected as lectotype of recognized two subspecies, the nominate subspecies Papilio niso Linnaeus does not match with Linnaeus’ and brevicornis Plötz. The latter was described by description and cannot be identified to a particular Plötz (1884) as a new species from Angola (in the species, while labelling and provenance are not be- genus Thymelicus). Although the description is clear yond doubt. Our conclusion can only be that the enough to recognize an A. letterstedti or A. hottentota, lectotype designation is invalid, Papilio niso is a no- it is of a female and it is impossible to pinpoint to men dubium as long as no acceptable type material which of the two the name refers. In other words, has been found, and the name Hesperia letterstedti it is a nomen dubium, and the only way to preserve Wallengren, 1857, should be re-instated. the name is to designate an undoubted A. letterstedti External characters. Length of forewing, male 11.6– from Angola as neotype. However, we are not sure it 15.1 mm (n = 25, mean = 12.9 mm, s = 0.5; see also is worth the attempt. Apart from the nomenclatural under Geographic variation), female 12.5−15.1 aspect, the only difference between the subspecies mm (n = 25, mean = 13.0 mm, s = 1.04). (according to Evans, length of forewing in letterstedti Downloaded from Brill.com09/27/2021 04:06:06PM via free access
14 mm, in brevicornis 13 mm) shows much overlap. Female — Not known for sure. The forewing of the smallest and largest specimens Male genitalia. Valva 2.75 times as long as high. among 30 males from S Africa and Swaziland (letter- Ventral edge of cucullus slightly concave or at most stedti) varied from 12.5 to 13.7 mm with outliers of straight. Apex of cucullus straight. Proximal exten- 11.6 and 14.7 mm, while in over 50 specimens from sion of cucullus at least one third of dorsal length Kenya the length varied from 12.6 to 13.7 mm, with of cucullus and slenderer than in A. letterstedti, with an outlier of 11.7 mm (all specimens in the collec- practically parallel sides. Aedeagus: proximal end tion of Naturalis Biodiversity Center, Leiden). As a with prominent olecranon (a flat, rounded extension consequence, we do not see any value in recognizing at the coecum of the aedeagus). brevicornis as a separate subspecies. Female genitalia. See below. Distribution. A.ocra has a restricted range, from Cape Afrogegenes hottentota (Latreille, 1823) comb. n. of Good Hope, South Africa, to southern Mozam- External characters. Length of forewing, male bique. Evans (1937) described it as a subspecies of 11.5−13.8 mm (n = 25, mean = 12.6 mm, s = 0.56); A. hottentota, even though it overlapped with the lat- since females cannot be separated from females ter according to the original description (both taxa of A. letterstedti with certainty we give here the recorded from Natal and Delagoa Bay). same measurements as for the latter: 12.5−15.1 mm Justification for species status. Overlap without inter- (n = 25, mean = 13.0 mm, s = 1.04). breeding would indicate separate species, unless the Male — Upper side forewing dark yellow in basal difference was due to an individual variation, found two-thirds, dark brown in outer third and at apex, in only part of the population. This seems unlikely no traces of apical spots, large black subtornal patch for an androconial patch, since it must deeply in- of specialized scales, presumably androconia; up- fluence display behaviour and thus the presence or per side hindwing as forewing, unspotted, without absence of androconia may act as an isolating mecha- the black patch; underside forewing ochreous, dark nism. We checked whether A. ocra possibly had an- brown in basal third, large pale ochreous patch in droconia with the same color as the rest of the wing, space cua2 that is suffused outwardly, ochreous spots but this was not the case. in spaces r3–5, m1–2, m3 and cua1. The problem seemed solved, when Pennington, Female — As female A. letterstedti. in Dickson & Kroon (1978), decided that ocra was a Male genitalia. Valva 2.75 times as long as high. form of G. niso (i.e. A. letterstedti). This concept was Ventral edge of cucullus slightly concave or at followed by Henning et al. (1997), who saw ocra as most straight. Apex of cucullus upturned. Proximal a slightly yellower form that becomes more prevalent extension of cucullus at least one third of dorsal in the more southern populations. length of cucullus and slenderer than in A. lettersted- Earlier, Kroon (1973), referring to Evans (1937), ti, with practically parallel sides. Aedeagus: proximal had taken up the challenge to find differences in the end without olecranon. male genitalia between A. niso (i.e. A. letterstedti) and Female genitalia. See below, under A. ocra. A. hottentota and he did find and illustrate them. Re- Distribution. Widely distributed throughout sub- markably, together with C.G.C. Dickson, Kroon ed- Saharan Africa in short grasslands and along forest ited the first edition of Pennington (1978), but did edges, wetlands and gullies, not as far into the dri- not refer to his earlier paper and changed the alloca- est zones as G. letterstedti (Evans 1937, Dickson & tion of ocra for unknown reasons, maybe as a tribute Kroon 1978, Kielland 1990, Larsen 1991, 2005, to Pennington who died before he could finish his Pringle et al. 1994, Henning et al. 1997, Vande book. This allocation was followed in the second edi- weghe 2010), extending to the Arabian Peninsula: tion of Pennington (Pringle et al. 1994) and by Hen- Asir (Saudi Arabia), Yemen and Aden (politically ning et al. (1997). When we checked the genitalia of part of Yemen) (Larsen 1983); not further south a male of ocra, we found them close to the type de- than the former South African provinces Transvaal scribed and illustrated by Kroon for A. hottentota and (Gauteng-Mpupalanga) and Natal (Kwazulu-Natal), not for A. niso (i.e. A. letterstedti). As a consequence and “homeland” Transkei (at present in Eastern we had little choice but to raise ocra to species level. Cape Province) (Henning et al. 1997). We have checked males from the following localities: Afrogegenes ocra (Evans, 1937) comb. n. A. letterstedti — Kenya (4), Swaziland (2), A. hot- Figs 47–50, 67 tentota — Kenya (2), Tanzania (2), Swaziland (2), A. ocra — South Africa (1), with results as given External characters. Male — As male of A. hottentota, above. but large black patch on upper side forewing miss- While the males of the three species are easily ing; about same size, 12.5−14.8 mm (n = 7, mean = separated by external characters, the females pose a 13.5 mm, s = 1.9), but the material available is too problem. According to Evans (1937) the discal band limited to get a fair idea of the variation. on the underside of the hindwing is constricted Downloaded from Brill.com09/27/2021 04:06:06PM via free access
Figs 65–66. Male genitalia of Afrogegenes hottentota. Left to right, upper part: inside and outside of right valve, saccus, underside and upper side of tegumen + uncus; lower part: lateral view and dorsal view of aedeagus, right lateral view of tegumen + uncus. — 65, Kenya, Kakamega Forest, 26–28.x.1974, leg. R. de Jong, prep. no. 5341; 66, Swaziland, Matsapha, Figueredo Dam, 30.v.1991, leg. J.A.W. Lucas, prep. no. 5344.
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Fig. 67. Male genitalia of Afrogegenes ocra. Left to right, upper part: inside and outside of right valve, saccus, un- derside and upper side of tegumen + uncus; lower part: lateral view and dorsal view of aedeagus, right lateral view of tegumen + uncus. S Africa, Port Elizabeth, Summerstand, 9.xi.1966, leg. B. Stuart, prep. no. 5367.
along vein 6 in A. letterstedti, and without constric- lamella appears more strongly bilobed. Both types tion in A. hottentota, but we can only agree with were found in Kenya, Swaziland and South Africa, Larsen (2005), that the two cannot be separated for the single specimen from Tanzania had a type A sure externally. Evans’ justification for the identity lamella, the only specimen from Yemen had a type of the females was the presence of two pairs taken B lamella. Since A. letterstedti and A. hottentota are in copula in the British Museum, one of each of both known from Yemen (Larsen 1983), the speci- the species. Obviously, this smallest possible sample men could belong to either species. So even if this size is not enough for evaluation of the constancy in variation is specific, we do not yet know which type such a minute character. Evans (1937) added that belongs to which species or if two or three species “The genitalia [Evans only studied male genitalia] were dissected. of hottentota and niso [i.e. A. letterstedti] seem alike”, which was not correct either, as was shown by Kroon (1973). Concluding remarks Evans (1937) justly realized that for a correct Since the genus has a distribution area covering three identification of females recourse must be found in continents, Europe, Asia and Africa, and relevant pairs taken in copula. Apart from the two pairs men- literature never dealt with the three areas together, tioned by Evans, we do not know of any other pairs. the marked differences between G. pumilio and G. So, as for the females, we are faced with three spe- nostrodamus on the one hand, and the purely African cies, which have very similar females. We dissected species on the other, in external characters as well as the genitalia of twelve females, from the following in male and female genitalia, has not received suf- localities: Kenya (5), Tanzania (1), Swaziland (3), ficient attention so far. The differences have led us to South Africa (2), and Yemen (1), hoping to find at distinguish two genera, Gegenes and Afrogegenes. Ex- least two different types. In fact, we found two types amination of the male genitalia showed A. ocra to be of pre-vaginal plate, type A (Figs 53−58) shallowly different from both A. hottentota and A. letterstedti. indented giving a weakly bilobed lamella, and type While the females of the Gegenes species can be sepa- B (Figs 59−64) more deeply indented so that the rated by their genitalia, discrimination of the females Downloaded from Brill.com09/27/2021 04:06:06PM via free access
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