June 2008 METAMORPHOSIS, VOL. 19, No. 2 41

METAMORPHOSIS ISSN 1018-6490 CONTENTS

Eighteen new species, five new subspecies, and interesting data on other African butterflies - Fourth ABRI research paper By Steve Collins1 & Torben B. Larsen2 ...... 42

Front cover: Liptena bia ♂: Sz. Sáfián Back cover: Euphaedra herberti ♂: A. Coetzer 42 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Eighteen new species, five new subspecies, and interesting data on other African butterflies - Fourth ABRI Research Paper

Steve Collins1 & Torben B. Larsen2

1 African Butterfly Research Institute, P. O. Box 14308, 0800 Nairobi, Kenya ([email protected]) 2 Torben B. Larsen, Jacobys alle 2, 1806 Frederiksberg, Denmark ([email protected])

Abstract: The primary objective of this paper is to describe 18 species and five subspecies mainly based on recent collecting under the auspices of the African Butterfly Research Institute, Nairobi (ABRI). The 23 new taxa are: Graphium almansor dufranei Collins & Larsen, ssp. nov.; Cooksonia abri Collins & Larsen, sp. nov.; Cooksonia ginettae Collins & Larsen, sp. nov.; Liptena lloydi Collins & Larsen, sp. nov.; Liptena minziro Collins & Larsen, sp. nov.; Liptena liberti Collins, Larsen & Libert, sp. nov.; Liptena bia Larsen & Warren-Gash, sp. nov.; Iridana agneshorvathae Collins, Larsen & Sáfián, sp. nov.; Hewitsonia kuehnei Collins & Larsen, sp. nov.; Aphnaeus marci Collins & Larsen, sp. nov.; Iolaus (Philiolaus) ofere Collins & Larsen, sp. nov.; Iolaus (Epamera) adorabilis Collins & Larsen, sp. nov.; Bicyclus sealeae Collins & Larsen, sp. nov.; Bicyclus pareensis sp. nov. Collins & Kielland; Bebearia staudingeri okomu Collins & Larsen, ssp. nov.; B. staudingeri carensis Collins & Larsen, ssp. nov.; Pyrrhochalcia iphis dejongi Collins & Larsen, ssp. nov.; Abantis ja usheri Collins & Larsen, ssp. nov.; Gorgyra warreni Collins & Larsen, sp. nov.; Meza gardineri Collins & Larsen, sp. nov.; Acleros bobiri Collins & Larsen, sp. nov.; Platylesches heathi Collins & Larsen, sp. nov.; and P. hassani Collins & Larsen, sp. nov. A dozen nearly unknown and/or misunderstood species are also discussed, and a number of these have their types figured in colour for the first time: Colias marnoana, Eresina cornucopiae, Falcuna melandeta, Iolaus likpe, Thermoniphas bibundana, T. leucocyanea, Bebearia denticula, Pseudathyma legeri, Acraea eugenia ochreata, Platylesches langa, and P. batangae. Pteroteinon reali Berger, 1962, hitherto treated as a subspecies of P. pruna Evans, 1937, is elevated to specific rank (stat. nov.) and is a senior subjective synonym of Caenides na Lindsey & Miller, 1965 (syn. nov.). The species Euriphene larseni is formally downgraded to a junior subjective synonym of E. saphirina itanii. We also hope that the paper will provide some impression of the research processes at ABRI. The status of our present knowledge of African butterflies has improved considerably during the past 15 years, but much remains to be done.

Keywords: Afrotropical Region, Papilionoidea, Hesperioidea, new taxa, new synonymy June 2008 METAMORPHOSIS, VOL. 19, No. 2 43

Introduction: Since the last of three previous contributions to this series of papers from the African Butterfly Research Institute (ABRI Research Papers) (Collins & Larsen, 1998, 2000; Collins, Larsen & Warren-Gash, 2004) went to press, a large number of new species and subspecies were described by our colleague Michel Libert and published in his revisions of the African Deudorigini, and the genera Ornipholidotos, Torbenia, Telipna, Pseudaletis, Oxylides, Syrmoptera, Deudorix s.l., and Cupidopsis (Libert, 2004a, b, c, 2005a, 2007a). This prodigious labour has greatly increased our understanding of the African Lycaenidae. Though Libert visited most major museums with African holdings, much of the work, the bulk of the types, and most of the photographs were based on the ABRI collection: thus, no less than 70 percent of the 1,670 Pseudaletis examined were from ABRI, including all or most of the type material of new species. This author is currently working on Anthene Doubleday and related genera (Libert, in prep.). The present paper aims at describing additional new taxa that were not in genera scheduled for in-depth revisions, as well as discussion of poorly known species on which new data have recently been procured, such as undescribed females, significant range extensions, and other points of interest. In all 18 new species and five new subspecies are described. We hope the paper will also give some impression of the work of ABRI and the process of discovering new butterflies in Africa. In order to speed up this process, we find it important that new species should be described as early as possible and most of the new taxa in this paper were first discovered during the past ten years. A dozen or so almost unknown or misunderstood species are also covered, as are descriptions of hitherto unknown females. We are particularly pleased to have been able to pin down two elusive skippers: Caenides na, was known only from the Liberia holotype (in terrible condition) was finally collected again in Guinea and proved to be a junior subjective synonym of Pteroteinon reali; the latter in turn is raised to specific status, having been considered a subspecies P. pruna in the past. Examination of the holotype of Platylesches batangae in the Carnegie Museum led to the discovery of a male and to the conclusion that all treatment of this taxon since the catalogue of African skippers by Evans (1937) refers to other, undescribed species. It is a sobering thought that the background research, description, photography, and genitalia drawings for each of the new taxa has averaged more than a full working day – though much was learned during this process. This is despite the fact that ABRI must presently be the best place to conduct this kind of research: the collection is well organized and has more African species than any other institution. The key literature is on the premises, as is a large picture gallery of types and interesting material from collections. We have excellent cooperation with other institutions and the internet is increasingly useful. Without the ABRI collection this paper would have taken even more time and many of the new taxa would still have been in doubt. 44 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Technical notes: For each of the species we list references to the original description, junior synonyms, as well as references to taxonomic papers that have special bearing on the taxa being discussed.

ABRI leg. implies that material was obtained by ABRI collectors in the field who cannot individually be identified.

Measurements are given as the length of the forewing from the base of the wing to the farthest point of the wing apex.

The following taxa, of which fresh material was available, will be sent for DNA barcoding when the paper has been accepted for publication: Cooksonia abri, Cooksonia ginettae, Liptena lloydi, Liptena minziro, Iridana agneshorvathae, Hewitsonia kuehnei, Aphnaeus marci, Iolaus ofere, Iolaus likpe, Iolaus adorabilis, Bicyclus sealeae, Bicyclus pareensis, Abantis ja usheri, Gorgyra warreni, and Acleros bobiri.

Genitalia references (e.g. SCC 565) refer to ABRI material dissected by Larsen.

Systematic Part: The listing of species follows the sequence in Larsen (2005): Papilionidae, Pieridae, Lycaenidae (Lipteninae, Theclinae, Polyommatinae), Nymphalidae (Satyrinae, Limenitidinae, Heliconiinae), and Hesperiidae (Coeliadinae, Pyrginae, and Hesperiinae).

Papilionidae

Graphium almansor dufranei Collins & Larsen, ssp. nov. Papilio almansor Honrath, 1884. Berliner Entomologische Zeitschrift 28: 210 (203- 212) (TL: Guinea, Ashanti [Patria falsa]). Papilio carchedonius Karsch, 1895. Entomologische Nachrichten. Berlin 21: 285 (275-286). [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum (Entomology Series) 70 (2): 633 (503-719))] (TL: Togo: Bismarckburg, Misahöhe). Papilio escherichi Gaede, 1915. Internationale Entomologische Zeitschrift 9: 71 (38-40, 71-74). (TL: Cameroon: “Neu-Kamerun, Carnot).

When reviewing the status of Graphium almansor Honrath, 1884 in West Africa and its distinctness from the commoner and more widespread G. adamastor Boisduval, 1836, Larsen (1996) concluded that G. almansor west of the Dahomey Gap was limited to the mountains of the Ghana/Togo border (the Volta Region refuge) in the endemic ssp. carchedonius Karsch, 1895 and that records from further west were erroneous. The faunistic distinctness of the Volta Region is discussed by Larsen (2006). June 2008 METAMORPHOSIS, VOL. 19, No. 2 45

G. almansor was described from 'Ashanti, Guinea' and was accepted from Guinea by Villiers (1957), despite the fact that Berger (1950) had already pointed out that the holotype of G. almansor was similar to Angolan material, and that the term 'Guinea' was often employed in a very wide sense. Since Pogge, the collector, did most of his collecting in Angola ('lower Guinea'), this interpretation is surely correct. 'Ashanti' is probably a misinterpretation of some local name in Angola. No material resembling the holotype was ever refound in West Africa. Condamin & Roy (1963) felt quite certain that G. almansor does not occur in the Nimba area. However, while the manuscript was in press we came across a photograph on a Flora & Fauna International website that illustrated the species. The photograph was taken October, 2003 by Jeremy Holden in the Nimba Mountains 20 km on the Guinea side of the border with Liberia. Graphium carchedonius was described from Togo, Bismarckburg as a species distinct from G. almansor, though closely related. Most subsequent authors have considered it the West African subspecies of G. almansor. It is consistently, though not strongly, differentiated from G. almansor escherichi (Gaede, 1915) from the mountains of the Nigeria/Cameroun border and in hilly country in central Cameroun. In 1996 Larsen had been unable to find any other records or material of G. almansor from west of the Volta River except for the descriptions of Papilio (Cosmodesmus) charcedonius [sic!] ab. guineensis Dufrane, 1946 (Guinea: Vallée de Mamou) and Papilio (Cosmodesmus) charcedonius [sic!] ab. houzeaui (Dufrane, 1946) (Guinea: kil. 30 au N. de Mamou, route vers Labé). A quick perusal at the time convinced Larsen that they were forms of G. adamastor – which they probably are. We have not traced the types, but the two names are clearly infrasubspecific and thus not available. However, a series of undoubted G. almansor was recently obtained by ABRI from the Fouta Djalon in Guinea. The specimens differ consistently from those of ssp. carchedonius in the Volta Region and are described as a subspecies below – formalizing the suggestion made by Larsen (2005).

Holotype: ♂, Guinea, Fouta Djalon, Labé (11o32'N 12o28'W), ii.2001 (ABRI leg. et coll. )

Paratypes: 12 ♂, 5 ♀, same data, various dates (one female is from Dubreka)

Diagnosis: Male forewing 40 mm. In the male the white cell-spot and that in space 3 are much larger than in carchedonius and fused with the small spot in 4 to form an almost solid patch: these three spots are well separated in the latter. The round spot in space 2 is usually prominent, normally being absent or small in carchedonius: sometimes it is nearly as wide as the spot in 3 and almost fuses with it. The new subspecies shares with carchedonius the long and somewhat irregular light subapical streaks, rather than the short, well-defined streaks of the equatorial and Angolan subspecies. Both subspecies are figured in Larsen (2005) and will not be illustrated here. When 46 METAMORPHOSIS, VOL. 19, No. 2 June 2008

viewed in series, both sexes of the Guinea subspecies seem to have a creamy tinge that is absent in carchedonius. The female is virtually identical with the male.

Discussion: Ssp. carchedonius is very locally and seasonally numerous in the Ghana/Togo Mountains, inhabiting the forest/savannah transition zone as far north as the Kyabobo National Park. None is known from Ghana west of the Volta, nor from Côte d'Ivoire, Liberia, or Sierra Leone. The Guinea population thus appears to be geographically isolated from the Ghana/Togo one, and also seems to be strongly localized. Records of ssp. escherichi are rather sporadic also in Nigeria and Cameroun (map in Libert, 2007b). Everywhere the species seems to be found mainly in hilly country (600-1,200 m) in woodland areas of the forest/savannah transition zone. A number of other subspecies are found further east, and then to the south, neatly avoiding the forest zone proper. It may occur also in the westernmost parts of Sierra Leone, though it may not be sufficiently hilly. Males are avid mud- puddlers and sometimes assemble by the dozen. The single specimen on the photograph from the Nimba Mountains cannot be confidently attributed to any of the subspecies, but appears closest to ssp. dufranei.

Etymology: We dedicate this subspecies to Abel Dufrane, who between 1929 and 1954 described a large number of species and aberrations of African butterflies. It is not impossible that one of his names for aberrations of G. charchedonius [sic] from Guinea actually does refer to this taxon. In his series of papers on the butterflies of Kivu, based on material sent to him by his son Alberic, who was a mining engineer, numerous new taxa were described. Eleven of his names remain valid, though many more have been relegated to synonymy. When the wheat is sorted from the chaff – not an easy task – Dufrane did make a significant contribution to the fauna of Kivu, still badly under-researched. The Liptena minziro that we describe in this paper was described by him in meticulous detail as an aberration, obsoleta, of L. eukrines. We dedicate this new subspecies to him: the only other tributes to this apparently rather eccentric man seem to be Papilio demodocus f. dufranei (Berger, 1950) and Spindasis dufranei (Bouyer, 1991).

Pieridae

Colias marnoana Rogenhofer, stat. rev. Colias marnoana Rogenhofer, 1884. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 33: 22 (21-25). (TL: Sudan).

Illustration of a little known species and confirmation of its specific rank. C. marnoana was described as a valid species and compared with C. hyale (Linné, 1758), but has subsequently usually been considered a subspecies of C. erate (Esper, 1803). The species is very rare in the Afrotropical parts of Yemen and Saudi Arabia and little material is available from elsewhere in its range, which covers June 2008 METAMORPHOSIS, VOL. 19, No. 2 47

parts of Sudan, Eritrea, and possibly Ethiopia. The species is specifically adapted to these dry areas, though irregular local seasonal migration is a possibility. If the species were truly migratory it would surely have made its way also to Egypt. In the recent monograph on the genus Colias, Verhulst (2000) retains the name marnoana as a subspecies of C. erate but d oes not illustrate the taxon. Many of the relatively few records are from the area immediately surrounding Khartoum in Sudan, from where ABRI recently received a long series of freshly hatched males and females collected by P.R.P . Ward: he was advised by Collins to try his luck in the alfalfa fields along the Nile, and on 27.vii.2007 he caught eight males and four yellow females, only just lighter than the male – white females are also known. This population lives under desert oasis conditions – the Nile is ecologically one long oasis – with temperatures often exceeding 40oC degrees and with little or no rain. It is more than 1,500 km from the nearest population of C. erate, and that is the Turkish population which has no yellow spots in the black forewing margin. All three points speak in favour of reinstating C. marnoana to its original species rank. Having collected and observed C. erate nilagiriensis Felder & Felder (sensu Verhulst) in the Longwood Shola [type locality] in the Nilgiri Mountains, as well as in other locations around Kotagiri and Kodanad in South India during the 1950s and in 1986, Larsen strongly believes that this taxon should also be considered specifically distinct. Colias marnoana: Male upperside (left) and female (right); male underside (below)

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Lycaenidae Lipteninae

Pentila condamini Stempffer, [1963] Pentila condamini Stempffer, 1964 [1963]. Bulletin de l'Institut Français d'Afrique Noire (A) 25: 954 (954-964). (TL: Senegal: “Basse Casamance: forêt classée de Santiaba Mandjak”)

Confirmation of specific status and range extension of a nearly unknown species: This species was described from the Basse Casamance in Senegal in 1963 and a single specimen was illustrated with identical genitalia from Harbel in Liberia by Clench in Fox et al. (1965). Larsen (2005) figured a Guinea male in colour and suspected that some specimens in the Belcastro collection were also P. condamini. Larsen was skeptical about a male from Gagnoa in Côte d'Ivoire, mentioned in a paper by Stempffer. Recently Szabolcs Sáfián collected a single male in Bia National Park in Ghana that seemed to match the species – Larsen is sure he never saw it there. Larsen collected a series at Bumbuna in north-central Sierra Leone in 2006. Reviewing the data, it now seems that P. condamini occurs all the way from Senegal to western Ghana (Senegal, Guinea, Sierra Leone, Liberia, Côte d'Ivoire, and western Ghana). Careful examination of the genitalia confirmed their stability and that the species cannot be considered a western subspecies of P. nigeriana Stempffer & Bennett, 1961.

Eresina cornucopiae (Holland), comb. nov. D'Urbania cornucopiae Holland, 1892, Annals and Magazine of Natural History (6) 10: 285 (TL: Ogowe, Gabon)

The holotype and genus of a much misunderstood species: This little butterfly has been mentioned in various lists and catalogues since its publication, but always as an uncertain entity and placed by various authors in a number of different genera, beginning with Durbania. Aurivillius in Seitz (1908-1924) says that “if the description was really taken from a male, the species is probably most closely allied to Pseuderesia russulus”. Ackery et al. (1995) placed it in Eresiomera, while Williams (2005) continued to follow Aurivillius. No-one claims to have refound it since its description. However, the female type in the Carnegie Museum is clearly a female Eresina in frightful condition. It strongly resembles the female of E. likouala Stempffer, 1962, itself an almost unknown entity from Kellé in Congo (Brazzaville), in the same biogeographical subregion as Gabon. It would seem rash to synonymize the two without more material since there do seem to be minor differences between Holland's type and the allotype of E. likouala in the Natural History Museum, London: their significance cannot be gauged in the absence of examination of male genitalia. Were the two species synonymized, Holland's cornucopiae would still take precedence: the name likouala is not yet sufficiently well established to be conserved under the criteria laid down in Section 23.9.2 of the current Code of Zoological Nomenclature (ICZN, 1999). June 2008 METAMORPHOSIS, VOL. 19, No. 2 49

The female upperside (left) and underside (right) of the type of Eresiomera cornucopiae in the Carnegie Museum, Pittsburgh (1.5 x natural size)

The genus Cooksonia H. H. Druce Transactions of the Entomological Society of London 1905: 256 (251-262) (Type species: Cooksonia trimeni Druce, 1905).

This is a small genus with three long described species, which until recently were rarely collected. While we were preparing this paper a fourth species was described as C. nozolinoi by Mendes & Bivar de Sousa (2007) from central Angola, about which more later. The species are very large by the standards of the Lycaenidae. The largest females surpass all other African lycaenids in size. The colour patterns make them excellent mimics of day-flying moths and various Acraea species. They are members of the Lipteninae, as evidenced also by their early stages, but their placement within the subfamily is uncertain. Williams (2005) includes them in the tribe Mimacraeini, but the genitalia do not support such a view: Stempffer (1967) thought that the genitalia were “commonplace” for the Lipteninae, while Mimeresia and Mimacraea share a unique genital pattern. We agree with Stempffer on this but can take the issue no further at present. The genitalia of C. neavei and C. aliciae are well figured by Stempffer (1967). They differ mainly in the width of the uncus, the valves being quite similar. The first to be described was C. trimeni Druce, 1905, the type species of the genus. It is known from the Shaba Province of the DRC. A subspecies was described as terpsichore Talbot, 1935, also from Shaba, but this must be a junior synonym: we have not seen the type. Next came Sheffieldia neavei Druce, 1912, in a new genus described by Druce in the same paper, from the Uehe District in then German East Africa; it is now known from several localities in Tanzania. It clearly belongs in Cooksonia, as does its subspecies C. n. rhodesiae Pinhey, 1962 from Zimbabwe and Zambia. It was followed by C. aliciae Talbot, 1935 from Malawi, still known from nowhere else. ABRI now has long series of two quite different additional species that are described below.

Cooksonia abri Collins & Larsen, sp. nov.

The first of these new species was collected in northern Cameroun, a most unlikely locality since it is about 2,200 km from the recent Angola record and almost as far further north, while it is 1,900 km west of where the other new species, C. ginettae, flies. The species is less sexually dimorphic than the other members of the genus. It 50 METAMORPHOSIS, VOL. 19, No. 2 June 2008

was first found in 2004 but it took repeated visits over the next three years for ABRI collectors to obtain an adequate series.

Holotype: ♂, Northern Cameroun, Waak [Wak] (07 o 70'N 13o 55'E), v.2007 (ABRI leg. et coll.)

Paratypes: 25 ♂♂, 23 ♀♀, same data as holotype, but with different dates (mostly 2007) Cooksonia abri: column:Left Male (holotype) (above) and underside (below) Right column: Female upperside (above) and underside (below)

Diagnosis: Male forewing 25 mm. The ground-colour is orange-brown above, with a black costa, subapical patch, and margin. The apex has white spots in spaces 5-7. The cell has a black basal streak, a black spot in its centre, and another at the end of the cell. All veins are blackened and there are two small black spots in space 1b towards the base. The hindwing is orange-brown with a broad black margin (3 mm at its widest). There is a black spot in the cell and another at its end, as well as two basal spots. The underside has a more complex pattern. The forewing has a marginal band of creamy rectangular spots finely bordered with black on the margin, the veins, and at the inner edges; the spots gradually increase in width towards the apex. Spaces 5 to 7 each has a white, wedge-shaped spot that is bordered on all sides by black and increasing in size towards the apex. The black inner edge to these spots is broad and regular. There are a variety of black spots in the forewing cell and beyond, as well as at the base of 1b. The hindwing has the same marginal band as on the forewing, bordered by white June 2008 METAMORPHOSIS, VOL. 19, No. 2 51

triangular spots that are edged by strong black chevrons. There is a full and regular line of rather delicate postdiscal spots and about ten larger round spots in the discal and basal area, two of which in the cell and one at its end. The female (forewing 29 mm) is as usual in the genus much larger than the male, but otherwise quite similar. Sexual dimorphism is less than in any other member of the genus. On the upperside the forewing veins are not as blackened and there is a small orange area beyond the cell, cut into three by finely black veins. On the underside the postdiscal spots are better marked.

Male genitalia: The male genitalia were viewed and are very similar to those of C. aliciae figured by Stempffer (1967).

Discussion: The presence of a Cooksonia in dry northern Cameroun came as a real surprise, so far away from any other member of its genus. None is present in northern Tanzania, Kenya, or Uganda. The ancestor of C. abri probably established itself during one of several dry periods in the Miocene or Pleistocene when the forest left a savannah gap on Kalahari sands in the equatorial zone in central DRC, resulting in strong disjunctions of the current rainforest belt (see also discussion under Platylesches langa later in this paper).

Etymology: The species is named for the African Butterfly Research Institute and in appreciation of its collectors in Cameroun who have provided a large amount of valuable material and many other new species.

Cooksonia ginettae Collins & Larsen, sp. nov.

The second of the new species is morphologically the one that differs most from the three already known species. Its occurrence on the Equator in the Kivu Province of the DRC represents a major northwards extension in the range of the genus in the east. The female of this species is probably the largest of all the African Lycaenidae: the largest lycaenid in the world is probably Liphyra brassolis Westwood, [1864] in the Oriental and Australian regions.

Holotype: ♂, DRC, East Kivu, Mitumba Mountains, Kasuo, 1,800 m (00o 14'S 29o 01'E), 14.iv.2007 (R. Ducarme leg., coll. Musée Royal de l'Afrique Centrale (MRAC), Tervuren).

Paratypes: 6 ♂, 16 ♀, Kithikolo-Lubero area, i-iii.2007 (coll. ABRI) same data; ♀ DRC, Kivu, Lume (same area) (R. Ducarme leg. et coll.); 2 ♀♀, DRC, Kivu, Kasungo (R. Ducarme leg., coll. ABRI). Another 3 ♂♂ and 18 ♀♀ paratypes are in coll. Ducarme 52 METAMORPHOSIS, VOL. 19, No. 2 June 2008

from various locations in the same general area (all contained within the square composed of coordinates 00 o 20'N 00 o 15'S; and 29 o 01'E 29 o 34'E)

Cooksonia ginettae: Left column: M ale (above) and underside (below) Right column: Female upperside (above) and underside (below)

Diagnosis: Male forewing 25 mm. The male forewing margin is slightly convex between veins 1 and 4, giving it a different aspect from the other species. The ground-colour is a more intense reddish-orange than in the other species. The upperside is very like that of an Acraea. The costa is broadly black, as is most of the apical half of the wing and a broad margin that terminates at the tornus with a width of 2.5 mm. There is a black basal streak in the cell, as well as mid- and end-cell spots that are fused with the costa. The veins are blackened, leaving isolated streaks of orange beyond the cell in spaces 2, 3, and 4. The hindwing has two basal black spots and an end-cell spot, as well as a moderate black margin, at its widest in spaces 1b and 2 (a single male has 5 or 6 additional spots, three in the cell and three in the neighbouring spaces). The underside is a lighter orange, more intense on the forewing discal area. It is very lightly marked compared with the other species. The distal ends of the veins are blackened on both wings, the length of the blackening increasing towards the forewing apex: the inner part of the apical area is overlaid with blueish-white scaling. There is a moderate black end-cell spot. The hindwing has two small basal spots, a tiny mid-cell spot and another below the cell, and a small end-cell spot. All veins are bordered by lighter scaling except where they are blackened at the margin, the true orange ground-colour only showing through as narrow wedges that form an almost imperceptible band of orange marginal lunules. The female (forewing 33 mm) is a much lighter shade of orange. The forewing has a smallish end-cell spot, a wide black apical area that contains three well-developed white subapical streaks, well separated by the black veins. The black area is continued as a moderate margin June 2008 METAMORPHOSIS, VOL. 19, No. 2 53

to the tornus. The hindwing is unmarked except for a narrow black margin, in which there are two tiny white tornal spots, as well as two larger spots in spaces 1b and 2. The female underside is like that of the male except that the orange ground-colour is more apparent in between the lighter scaling bordering the veins. The females vary in tone, some having a redder ground-colour. Such individuals have deeper colours and more contrast on the underside than the one figured. We take this to be some form of seasonal variation or response to local ecological conditions. The new Angolan species, C. nozolinoi, is known from just a single female, figured in the description. The forewing upperside is like that of C. ginettae except that there are no white apical spots. The hindwing is also similar, but the margin is much wider. The underside is quite different from the nearly unmarked C. ginettae, having the same submarginal pattern as C. abri: the forewing has a single black end-cell spot and on the hindwing there is no submarginal line of spots and the few discal and basal spots are small. We find it impossible to venture a guess as to how its male might look.

Male genitalia: The male genitalia were viewed and are very similar to those of C. aliciae figured by Stempffer (1967).

Discussion: A few specimens were found over a period of years in eastern Kivu by R. Ducarme, on which he deserves our congratulations. In 2007 an initiative to get more resulted in the fine series being collected, some of which were sent to ABRI for description. They were from a number of locations within a small geographical area, where the butterflies fly during the dry season in a single annual brood.

Etymology: We take pleasure in naming this fine species after Ginette Ducarme, the wife of Robert Ducarme, who first found the species and has done so much to elucidate our knowledge of Kivu butterflies.

The Liptena eukrines species complex

For many years, small numbers of orange Liptena apparently related to L. eukrines Druce, 1905 remained unidentified in the ABRI collection. Recently more comparative material was obtained, especially with the incorporation of the Kielland collection, and we decided to look more closely at this complex. L. eukrines has always been considered a somewhat unusual species since its distribution was centered on the drier southern forest zone in northwestern Zambia (especially Ikelenge) and of the southern DRC (especially Shaba and Lualaba), and as a species without close relatives. The new material was of smaller size than typical L. eukrines and with much less consistent dark banding on the grey hindwing underside. Material from the Minziro Forest in the Bukoba area in Tanzania was identified as L. eukrines by Collins & Congdon (1998), but specimens were smaller in size and with less consistent banding on the underside. Additional material from Cameroun and the Central African Republic seemed to differ from the other two. The genitalia showed that all three populations were closely related. Examination of the genitalia 54 METAMORPHOSIS, VOL. 19, No. 2 June 2008

showed that the Minziro population was very similar to that of Zambian L. eukrines, while the Cameroun/CAR material had more distinctive genitalia. Accordingly two new species are described as L. lloydi and L. minziro respectively.

Liptena lloydi Collins & Larsen, sp. nov. Liptena eukrines Druce, 1905. Transactions of the Entomological Society of London 1905: 253 (251-262) (TL: [Democratic Republic of Congo], not Zambia (see Cookson, 1954): “North-east Rhodesia”.

Holotype: ♂, Central African Republic, Bookoko [near Bangui] (04o 30'N 17o 45'E), xii.1996 (ABRI leg. et coll. )

Paratypes: 1 ♀, as holotype, but in October; 2 ♂♂, Central African Republic, Yakoli, iii.2000; 1 ♂, Bonkoui, ix.1995; 4 ♀♀, Cameroun, Ebogo, ix.2001, ix.2002, ix.2005, x.2006; 1 ♂, Cameroun, Man, vi.2000

Diagnosis: Male forewing 14 mm, somewhat smaller than the typical L. eukrines (15-16 mm), though smaller specimens do occur. The ground-colour is a light orange, the tone of which is deeper than in L. eukrines. The black markings are similar, but slightly more extensive. The costa is black to beyond the end of the cell, where the black widens and extends to vein 4 as a black bar. There is then a gap where orange penetrates to a fine black line at the costa, before the apical patch begins. The inner edge of the apical patch is more regular than in L. eukrines and runs from the costa to vein 4, from where a narrow black marginal line runs to the tornus. The hindwing margin is bordered by a narrow black line that is broken by orange at the veins. On the underside the forewing markings are essentially as on the upperside, except that the costa and the apex are overlaid with ochreous scaling. The hindwing underside is light ochreous with irregular chocolate-brown spotting and banding. There are five weak, irregular spots in the basal part of the wing, followed by a discal band that clearly consists of an end-cell spot and a large costal bar that stretches past the end cell spot without touching it: it almost meets another smaller brown bar that extends from vein 1b to 2. There is a weak and broken postdiscal band, obsolete in some specimens, while a submarginal band is more substantial, running unbroken from the tornus to the costa. There is also a fine brown marginal line. However, all markings are subject to variation. The ochreous underside is in contrast to the greyer underside of the L. eukrines hindwing with its much neater and regular banding. The females can be told apart from the males only by the shape of the abdomen and the genital opening.

Male genitalia: The genitalia (SCC 560 (holotype) and 561) are considerably larger than in L. eukrines but their basic structure is the same, differing from other members of the genus by the astoundingly large subunci. The uncus consists of two lobes that are separated by a deep V-shaped gap; the outer edge of each uncus-lobe is gently rounded at the tip, but the inner edges of the V are straight: the uncus is June 2008 METAMORPHOSIS, VOL. 19, No. 2 55

proportionally longer than in L. eukrines, as is the tegumen, so that the whole uncus/tegumen structure is narrower and more elongated. The subunci are long, ending in a huge, flat triangular structure. The shape of this is very like the axes used for beheadings in medieval Europe, with the blade curving gently throughout. In L. eukrines this structure is of a different shape, being abruptly curved inwards just before the middle and ending in a much narrower and pointed manner. The valve ends in two projections, one a flap of the normal texture and the other a chitinized thorn. They are difficult to mount consistently, but it is clear that both are much longer than in L. eukrines. The slender penis has no special features. Finally the saccus is huge (longer than the valve), broad, and squared off at the end without the large lobes of the two other species. In situ the saccus of all three species projects forwards.

Discussion: We were initially uncertain to what extent the species was related to L. eukrines, but the genitalia clearly show the relationship to be very close. The morphological characters make the species closer to the next species, but the genital characters do not.

Etymology: The species is named after the late Patrick ('Paddy') Lloyd, a friend of Collins' during his youth in Kitale, Kenya. Paddy was an avid pupil of his neighbour, T.H.E. ('Pinkie') Jackson, one of the greatest lepidopterists that Africa has produced. One of Paddy's finest expeditions was to the eastern DRC, where his managed to scoop the master by finding several rare species that he had never seen – even though Jackson thought he “owned” the butterflies in that part of Africa.

Left: Liptena lloydi male holotype upperside (above), underside (below). Centre: Liptena minziro male holotype upperside (above), underside (below). Right: Liptena eukrines male upperside (above), underside (below) 56 METAMORPHOSIS, VOL. 19, No. 2 June 2008

The male genitalia of Liptena lloydi (left), L. minziro (centre), and L. eukrines (right). Probably these species are never sympatric.

Liptena minziro Collins & Larsen, sp. nov. Liptena eukrines Druce, 1905. Transactions of the Entomological Society of London 1905: 253 (251-262) (TL: [Democratic Republic of Congo], not Zambia (see Cookson, 1954): “North-east Rhodesia”. Liptena eukrines f. obsoleta Dufrane, 1953. Bulletin et Annales de la Société Royale Entomologique de Belgique 89: 49 (41-57) (TL: Democratic Republic of Congo: “Kamituga”)

When the Minziro population was examined it proved impossible to find decisive morphological characters that set it apart from L. lloydi described above, but on the other hand the male genitalia were distinctly smaller and much closer to those of L. eukrines in both size and structure. On the basis of genitalia alone it would just be possible to consider the Minziro population a subspecies of L. eukrines. The pattern of the hindwing makes that nearly impossible, since it relates more closely to that of L. lloydi. The most logical approach is therefore to describe it as yet another member in a complex of three closely related species (we are uncertain as to the affinities of L. eukrinaria Bethune-Baker, 1926 since we have not been able to dissect a male).

Holotype: ♂, NW Tanzania, Bukoba, Minziro Forest (01o20'S 31o49'E), 23.xi.1994 (Colin Congdon, Martin Hassan leg., coll. ABRI) June 2008 METAMORPHOSIS, VOL. 19, No. 2 57

Paratypes: 11 ♂♂, 10 ♀♀, same locality as the holotype, mainly on Kere [Kele] Hill, i, ii, iii, iv, vi, vii, ix, xi, xii.1993-1997 (Jan Kielland, Colin Congdon, Ivan Bampton, Martin Hassan leg., coll. ABRI); 1 ♂ Kenya, Kakamega, v.1995 (S.C. Collins leg., coll. ABRI)

Diagnosis: Male forewing 14 mm. The upperside is effectively identical to that of L. lloydi described above, but the orange ground-colour is marginally lighter in tone. The underside has a more washed out appearance than in L. lloydi and the darker markings are lighter. There is more diffuse dark spotting in the basal half of the hindwing, while the well-developed submarginal band is almost missing. The hindwing colour is ochreous without the grey tone of L. eukrines. When checking the undersides in the ABRI series, one specimen was picked out as being atypical – it turned out to be a misplaced L. lloydi from Cameroun! It should usually be possible to tell the two apart, but they probably never occur sympatrically. Confusion with L. eukrines should not be possible.

Male genitalia: The male genitalia (SCC 563) are about the same size as in the larger L. eukrines and distinctly smaller than in L. lloydi, which is about the same size. The structure is the same as in L. eukrines but differs slightly: the uncus/tegumen is more slender, and the two uncus lobes are divided more deeply. The huge subunci are slightly larger and their outer edge bends inwards to form a point more than half way from the base; in L. eukrines this bend is before the half-way point (and in L. lloydi it does not bend at all, but is evenly rounded), There are no evident differences in the other structures.

Discussion: We are confident that three distinct species are involved in what may be termed the L. eukrines-complex. One possibility for the evolution of such a complex is that their joint ancestor was widely distributed in the equatorial forests during mesic conditions, but then became isolated during one of several dry periods, when savannah woodland on Kalahari sands widely separated the Cameroun/Gabon/Congo refuge from that in the Ituri/Kivu refuge. The population in Ituri/Kivu was then able to move south under later more mesic periods, there to develop the special underside pattern of L. eukrines with little modification of the genital features. ABRI only has material from Minziro in NW Tanzania and a single Kenyan male. Judging by the description of L. eukrines f. obsoleta Dufrane, 1953 (TL: DRC, “Kamituga, southern Kivu) it must be the same as L. minziro. On present evidence L. eukrines is adapted to somewhat drier habitats than the other two. Etymology: We are pleased to give this butterfly the name of the Minziro Forest in Tanzania. Following the publication of his book on the butterflies of Tanzania, Jan Kielland realized that a revision was necessary in order to include the butterflies of the Bukoba area on the border with Uganda, on the western side of Lake Victoria. This is the part of Tanzania most influenced by the main equatorial forests and in the 58 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Minziro Forest he found the ideal location. Species new to Tanzania and to science were found well beyond any expectations: some 800 species were caught in northwestern Tanzania during this time, including 250 not included in the book on Tanzanian butterflies (Kielland, 1990): many species new to science were among them. He compiled exhaustive notes on his findings before his untimely death in 1995. Ivan Bampton, Colin Congdon, Martin Hassan, and Peter Walwanda continued his researches and the joint results were published in 1998 (Congdon & Collins, 1998). The Minziro list of species is one of the longest from any forest in Africa: if butterfly biodiversity is so high, that will be the case for other organisms as well. We hope that Liptena minziro will assist in the conservation of this priceless habitat.

Liptena liberti Collins, Larsen & Rawlins, sp. nov.

In 1993 Larsen located a member of the genus Liptena during a visit to the Carnegie Museum and pointed out to John Rawlins, the curator, that it ought to be described. It was an unusual, and unusually pretty, species characterized by the presence on the hindwing underside of a broad orange band, contrasting strongly with the five other bands that had the usual tan colour of its sub-group within the genus. However, the visit was for the purpose of studying West African material during the first stages of researching the West Africa book (Larsen, 2005); all memory of it was erased under pressure of the primary objective. During a visit to Carnegie Museum in 2007 the specimen again came to light, Rawlins actually remembering that he had been shown that very butterfly 14 years ago. Enquiries showed that none was at ABRI, but that Michel Libert had also seen this specimen and had two additional specimens from Cameroun that were identical. Libert plans eventually to revise the genus Liptena and it was resolved that we should name the species after him in anticipation of such a revision.

Holotype: ♂, Cameroun, Bikoman (03 o26'N 11 o52'E), 16.iv.1989 (M. Libert leg., coll. Muséum National d'Histoire Naturelle, Paris)

Paratypes: 1 ♀, as holotype but in coll. Libert; 1 ♀, Cameroun, [Central Province, Nyong-et- Soo Division], Métet (03°26'30"N 11°45'30"E), xi.1921 (E. Lippert leg., coll. Carnegie Museum, Pittsburgh).

Liptena liberti: ♀ paratype (Carnegie Museum), upperside left, underside right June 2008 METAMORPHOSIS, VOL. 19, No. 2 59

Diagnosis: Male forewing 14 mm; female 15 mm. The upperside is orange, somewhat deeper in tone than in other species with a similar type of hindwing underside banding (L. rochei Stempffer, 1951, L. eketi Bethune-Baker, 1926, and L. seyboui Larsen & Warren-Gash, 2004). The forewing costa is rather broadly black extending well into the cell and almost obscuring two black teeth in and at the end of the cell. The costa is slightly invaded by orange just where a large black subapical patch begins. The inner margin of the patch runs straight, if somewhat irregularly, to join a narrow margin that tapers somewhat towards the tornus. The final millimetre of vein 1 is marked by a wedge of black. The hindwing has a black margin and a well-marked black submarginal band that extends to the margin along the veins, creating a series of orange submarginal lunules. The species could be recognized by the upperside alone. The forewing underside is orange. The costa at the end of the cell is more strongly invaded by orange, while the inner margin of the subapical black area is smaller in extent than on the upperside. A line of whitish submarginal, slightly lunular, spots stretches from space 3 to the costa, the one nearest the apex in space 7 being large and wedge-shaped. Three tiny spots in spaces 8-10 complete the series. An inner line of narrow, slightly lunular spots that are divided by the veins precede a narrow black margin. The hindwing has seven well-developed black bands on a light tan background and a narrow black margin; in total almost half the underside area is black. The four inner tan bands thus created are straight, the inner two more or less crossing the wing straight to the costa. The outer two are stopped by black at the edge of the cell. The fifth, postdiscal, band is broader and of an intense orange colour, giving the species a unique appearance. A particularly large orange spot is created just beyond the cell in space 4 and 5: here the band is interrupted by black, recessed inwards, and continued to the costa. The next band consists of rather large, sagittate, tan lunules, separated by dark veins. A submarginal band of less developed lunules abuts a narrow black margin. Even if the postdiscal band were not orange, this underside would be most distinctive.

Male genitalia: The species is so unique that description of the genitalia will await a revision of the genus.

Discussion: Being recognizable at a glance it seems amazing that just three specimens of this butterfly appear to be known. The two localities are separated by only 18 km. However, many of the Lipteninae, even those with quite wide distributions, are very rarely captured. Presumably their micro-habitat is limited by a number of factors, one of which is the associated ants; another may be the need for display areas. Some species may also spend most of their life-cycles only just below the main canopy. Structured field observation on liptenine behaviour would really be most interesting.

Provenance of the Carnegie specimen: Many specimens in the Carnegie Museum are simply labelled “Metet Cameroon”. A search under this name reveals a number of possible localities. Study of Museum archives revealed that all material with this label was obtained from Mrs. Lippert, the 60 METAMORPHOSIS, VOL. 19, No. 2 June 2008

wife of Dr. A.B. Lippert. The Lipperts worked as missionaries in Cameroun between 1909 and 1926 and were instrumental in setting up a hospital in Métet (Central Province, Nyong-et-Soo Division] (3°26' 30"N 11°45'30"E). Métet is on the paved road between Mbalmayo and Sangmélima [Sangmelina is an alternative spelling (Jeff Boyd pers. comm.)] and the Bikoman type locality is just 18.2 km NW of there.

Etymology: We take pleasure in naming this species after Michel Libert. His prodigious recent work on revisions of African Lycaenidae has already been covered in the introduction to the paper. He hopes one day to tackle the genus Liptena, which in some respects will be an even greater challenge than some of those he has already met.

Liptena bia Larsen & Warren-Gash, sp. nov. Liptena rochei Stempffer, 1951. Bulletin de la Société Entomologique de France 56: 66 (66-71) (TL: Nigeria, Lagos)

When surveying the butterflies of Bia and Ankasa National Parks in Ghana, Larsen (1981) found a butterfly that differed from his experience with L. rochei in Nigeria, and it was presumed to be a new species: “Liptena bia (Larsen & Warren-Gash, manuscript) was a most surprising capture during the present mission. Nothing close has been recorded from West Africa. In the field it was identified as being close to L. eukrinaria Bethune-Baker, 1926, but comparison shows that even if it is related to that species, it is clearly distinct”. The description of Liptena bia fell through owing to lack of comparative material and the fact that the projected co-authors were living in Manila and Abidjan, respectively. ABRI now has a large material of this group both from the Volta Region and Ghana proper, as well as a few from Côte d'Ivoire. It is now clear that this Bia specimen was a female of the variable western population of L. rochei, with limited b rown markings on the hindwing that were more broken up than usual. Stempffer (1967) had stated that “specimens from Côte d'Ivoire are more orange than those from Nigeria, while those from material from Ghana are intermediate”: he almost certainly only had material from the Volta Region collected by Father Maessen. Closer examination indicates that the western population actually constitutes a distinct species, which is described below. Holotype : ♂, Ghana, Central Region, Sagamase (00o 17'N 34o 45'E), xii.1997 (ABRI leg. et coll.)

Paratypes: 5 ♂♂, 4 ♀♀ from localities in central and western Ghana (3 ♂♂, 3 ♀♀ locality as holotype but various years in i, vii, ix, x, xii (1997-2005); 1 ♀, Ghana, Western Region, Bia National Park xii.2003; 2 ♂♂, Côte d'Ivoire, Abengourou, xi.1991 (all leg. et coll. ABRI). June 2008 METAMORPHOSIS, VOL. 19, No. 2 61

Liptena bia: male holotype to the left: upperside (above), underside (below). To the right the same for Nigerian L. rochei.

Diagnosis: Male forewing 15 mm, probably on average slightly smaller than L. rochei. The ground-colour is distinctly deeper orange than the light ochreous of L. rochei and the black markings on the forewing costa and along the hindwing margin are usually better developed, but there is considerable variation in both species. The undersides are even better characterized by their deeper orange colour. The pattern of parallel bands tends to be rather more broken up, in extreme cases being almost as broken up as they are in L. eukrinaria. Females are usually quite similar to the males but the dark markings are less developed.

To the left are the genitalia of Liptena rochei (from Stempffer, 1967). To the right the sub- uncus (above) and valve (below) of L. bia 62 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Male genitalia: The male genitalia (SCC 565) are quite difficult to mount on slides in a consistent manner and the differences and illustration are partly from completely dissecting the genitalia of two males. Compared with Stempffer's original genitalia illustration (shown above and identical with an old mount made by Larsen from Lagos) the new species has the same basic ground-plan, but differs in two main features. The subunci are distinctly more massive and differing in shape, the outer of the two processes curving up, ending almost parallel to the main process. The valves are much broader, the two distal processes longer as well as more widely separated. We tried all different modes of observing the valves as Stempffer would have mounted them (he taught Larsen this skill forty years ago): with four specimens it was not possible to mount them so that they had the same appearance as Stempffer's.

Discussion: This is one of those cases where the choice of specific or subspecific status is a delicate one. Were it not for the fact that the genitalia supported the colour differences, and that a distinct species west of the Dahomey Gap is biogeographically plausible, we would have opted for subspecific status. The population from the Volta Region is generally a lighter orange than that from west of the Volta Lake/River, but the genitalia are definitely closer to those of L. bia. An intriguing thought is that L. rochei evolved in the Volta Region and then spread both east and west, evolving in different directions on either side. L. rochei has a narrow distribution, being limited to western Nigeria. L. bia is known from Sierra Leone to Ghana. We shall revisit the Volta Region population, which has been excluded from the type series and may deserve a subspecies of its own. The closest relative west of the Dahomey Gap is L. seyboui Larsen & Warren-Gash, 2004 but confusion between these two is impossible: L. flavicans Grose-Smith & Kirby, 1891 is also very different.

Etymology: The species is given the name of Bia National Park in western Ghana, a world heritage site, and one of the most important conservation sites in Ghana. There are probably 650 species of butterfly in Bia and it is proposed to include the forest in a long-term monitoring project that will culminate in an in-depth survey around 2100.

Falcuna melandeta (Holland) Larinopoda melandeta Holland, 1893. Entomological News 4: 25 (22-28). (TL: Gabon: “Talaguga, Upper Valley of the Ogové”)

Illustration of a unique holotype: This species was described by Holland from a single Gabon specimen, almost certainly male. Stempffer & Bennett (1963) never saw it, but concluded on the basis of the description that it belonged in their new genus Falcuna and included it as such. The holotype in the Carnegie Museum has June 2008 METAMORPHOSIS, VOL. 19, No. 2 63

never been figured and we take the opportunity of doing so. There can be little doubt that it is indeed a very distinctive Falcuna, with a minimum of hindwing markings. Most members of the genus have a white tooth at the end of the forewing cell that breaks the black costa almost in two. F. melandeta thus falls into the small F. campimus-group. The abdomen is pinned in a very fragile envelope below the specimen and was not examined, but it does seem to be a male – sexual dimorphism in the genus hardly exists. The holotype remains unique and quite different from any other Falcuna: there is no reason to suspect it to be an aberration.

The holotype of Falcuna melandeta (upperside left, underside right)

Iridana agneshorvathae Collins, Larsen & Sáfián, sp. nov. Iridana obscura Stempffer, 1964. Bulletin de l'Institut Français d'Afrique Noire (A) 26: 1259 (1226-1287). (TL: Uganda: “Bwamba, Hakitengya”) Iridana ghanana Stempffer, 1964. Bulletin de l'Institut Français d'Afrique Noire (A) 26: 1249 (1226-1287). (TL: Ghana: “Juase [Ashanti]”)

In late September, 2006 Szabolcs Sáfián collected a single fine male specimen of an Iridana that to his great surprise came at night to light on his moth screen together with myriads of moths and nine other butterflies in Ghana's Bia National Park: there are actually quite a number of records of Iridana coming to light, including the first record of I. exquisita Grose-Smith, 1898 from Ghana (Larsen, 2005) and four specimens of I. incredibilis Staudinger, 1891collected by Sáfián. The species has a very dark underside, which led him to believe it might be I. obscura, an almost unknown taxon from Uganda. The holotype of this is illustrated by D'Abrera (1980) and we have found no further records. This is a smaller butterfly and on the forewing underside there are additional white discal spots. Of the Iridana known from West Africa none is as extensively blackened on the underside. ABRI has by far the largest collection of Iridana anywhere, but none that resembles this specimen, and nothing indicates that it might be a melanic aberration. It is therefore described as a new species in comparison with I. ghanana, a species not illustrated by Larsen (2005) and the status of which was considered uncertain. However, a fine male and a matching dark brown female were found in the Maessen collection in the McGuire Center in October 2007and the male is illustrated here for the first time in colour. 64 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Top row: The male holotype of Iridana agneshorvathae, upperside (left) and underside (right). Bottom row: The male of I. ghanana, upperside (left) and underside (right).

Holotype: ♂, Ghana, Western Region, Bia National Park (05 o 46'N 08o 67'W), 1.x.2006 (Sz. Sáfián & K. Aduse-Poku leg., coll. ABRI).

Diagnosis: Forewing 16 mm. The male forewing is more squat than in other Ghana members of the genus. The male upperside is a brilliant, shimmering cobalt blue that covers the lower one-third of the cell and most of the discal areas, much deeper than in I. ghanana. The costa and two-thirds of the cell are black, extending to form a large subapical patch that continues as a broad margin tapering towards the tornus. In West Africa the upperside comes closest to that of I. ghanana, but the black apical patch is larger (only just under half of space 3 is blue, almost three-quarters in the latter). The hindwing margin of the new species is also wider. In both, the black of the abdominal fold of the hindwing reaches the lower cell vein, but turns towards the margin so that the blue intrudes slightly into space 1c. The underside has the usual pattern of silvery-green bands and spots on both wings: these are almost identical in the West African species, except for the small I. hypocala Eltringham, 1929. The chief character of the new species is its very dark underside: in particular the postdiscal half of the wing beyond the green discal band is so overlaid with dark scales that it is effectively black (except for the costal area). The forewing is also dark: there is a large brownish-white double spot just beyond the cell and a darker, diffuse white patch in space 1b. In the small I. obscura these spots are lighter, almost white, there are additional discal spots, and spaces 1a and 1b are very light. The hindwing is angled at vein 3.

Male genitalia: The male genitalia were not considered useful by Stempffer (1964); we suspect there may be subtle differences but do not wish to experiment with the only known specimen till this has been researched further. June 2008 METAMORPHOSIS, VOL. 19, No. 2 65

Discussion: The single male is so distinctive that we have no hesitation in describing it as a new species. The more squat forewing, the reduced amount of blue on the forewing, the wider hindwing margin, and the darker ground-colour preclude it from being a melanic aberration of I. ghanana. Larsen (2005) expressed doubts about the validity of the latter, but the specimen in the Maessen collection fully validates it: he placed the underside ground-colour in between grey and the cinnamon or brick-red of I. exquisita and I. nigeriana Stempffer, 1964. However, the distal half of the hindwing does have a strong dusting of reddish scales, but it is still more grey than cinnamon.

Etymology: The species is named after Ágnes Horváth, the partner of Szabolcs Sáfián. By naming this most beautiful Iridana after her, he wishes to express his gratitude for her understanding and acceptance of his lifetime commitment to the study of butterflies – including his extended trips to Africa.

Hewitsonia kuehnei Collins & Larsen, sp. nov. Hewitsonia inexpectata Bouyer, 1997. Lambillionea 97 (1) (Tome II): 89 (82-98). (TL: Democratic Republic of Congo: “Zaïre, Sankuru Katako-Kombe, 14-III-1953 (Dr M. Fontaine).” In MRAC, Tervuren.

When Bouyer (1997) revised the genus Hewitsonia he had at hand an ABRI male and a female of his new species H. inexpectata Bouyer, 1997 from Kenya, commenting that “these two specimens were slightly different from the population in eastern Zaïre [DRC] but that it is too early to say whether it concerns a distinct subspecies [translated from the French]”. ABRI now has a large material from Kakamega, which actually justifies the description of a distinct species.

o o Holotype: ♂, Kenya, Kakamega Forest (00 17'N 34 45'E), vi.2001 (S.C. Collins leg., coll. ABRI)

Paratypes: 33 ♂♂ 23 ♀♀, same locality, various dates, years, and different collectors (coll. ABRI).

We are only including Kakamega material as formal paratypes, but one male from Mongiro, Bwamba in Uganda and three males and a female from Minziro Forest, NW Tanzania also seem to match. Two of the Tanzanian males have black marginal triangles on the hindwing but these are seen sporadically in populations of other Hewitsonia species, though not in the Kakamega series. 66 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Hewitsonia kuehnei: Male upperside (left), female upperside (right). Hewitsonia inexpectata from Ghana figured below these to show the difference in wing shape.

Diagnosis: Male forewing 22 mm, making it slightly smaller than H. inexpectata from Ghana, with which it is compared (24 mm). The wing shape is very different, the forewing being narrower (the ratio between the costa and the distance between tornus and apex is 9% higher in H. inexpectata) and the hindwing smaller and more angular: the margin is straight from the tornus to vein 3, whereas evenly rounded in H. inexpectata. This is a consistent feature in the long series. H. amieti Bouyer, 1997 has the same type of forewing (also consistently), but the hindwing is more rounded. The ground-colour is slightly more silvery than in H. inexpectata and more restricted in extent, especially the large blue postdiscal patch in space 2 is more of a narrow streak and there is hardly any blue scaling in space 1a. The hindwing costa is white, though somewhat darker in a few paratypes. The female also differs in the shape of the wings. The size is again smaller than in H. inexpectata (24 versus 26 mm). The spots of the subapical band of the forewing are distinctly smaller and the outer spot in space 4 is completely detached, usually separated by the width of the spot itself: H. inexpectata has a much wider true band, and if the spot in 4 is detached, it is only by a sliver of black. The khaki-grey ground-colour of spaces 1a and 1b of the forewing and all of the hindwing is lighter in tone than in H. inexpectata.

Discussion: H. inexpectata is known from Côte d'Ivoire, Ghana, Togo and then again from Zaïre [DRC] (TL Eala). Bouyer states that there are no significant distinctions between the populations of West Africa and those of Zaïre, despite its apparent absence from Cameroun, Gabon, Congo, and the Central African Republic. It is therefore curious that the populations east of the Rift Valley are significantly, if not dramatically different, but we have not had access to DRC material. However, such differences between the two sides of the Rift are biogeographically plausible. June 2008 METAMORPHOSIS, VOL. 19, No. 2 67

Etymology: The species is named after Lars Kühne who has devoted much effort to the conservation of Kakamega Forest. He collaborated with ABRI in drawing up an annotated list of the Kakamega butterflies, one of the most detailed from any African rainforest (Kühne et al., 2004). He is currently working on a book on Kakamega Forest which will have the latest updates on its moths and butterflies.

Theclinae

Aphnaeus marci Collins & Larsen, sp. nov.

This is another interesting new species that was collected at Waak in northern Cameroun, from where also Cooksonia abri is described in this paper, as was Iolaus (Philiolaus) christofferi Collins & Larsen, 2004. Many other rare butterflies have turned up here. Similar specimens of Aphnaeus from West Africa (assigned to A. jefferyi Hawker-Smith, 1928 in Larsen (2005)) all have the two tails, as does A. jefferyi from western Kenya (TL Kitale).

Holotype: ♂, Northern Cameroun, Waak (07 o 70'N 13o 55'E), v.2007 (ABRI leg. et coll.)

Paratypes: 12 ♂♂, s ame data.

The holotype of Aphnaeus marci : upperside left, underside centre, and underside of a specimen of A. jefferyi from a Cameroun forest to the right (note the two tails)

Diagnosis: Male forewing 15 mm. The hindwing has a single, long tail (two tails in the Aphnaeus jefferyi-complex), which it resembles in the greenish colouration). The upperside ground-colour is black with the following markings in a rather dark, dully- shining green, the green scaling being interspersed with black scales at on average a half-to-half basis: the base of the forewing cell is green with a green bar at mid-cell and another such bar at the end of the cell. There are few green scales in the basal half of space 3, more in the basal half of 2, while more than half of 1a and 1b are green. The hindwing is black with green scaling in the cell and much of spaces 2, 3, and 4. The underside differs from all other species in the lack of silver spots in the outer half 68 METAMORPHOSIS, VOL. 19, No. 2 June 2008

of both wings. The ground-colour is dark reddish-brown with silver spots as follows: a basal, middle, and end-cell bar in cell; two or three small subapical spots; a small silver spot in space 2; a similar spot in 1b, beneath which is a silver streak. The main character of the new species is the two very large silver spots on the costa of the hindwing (fused to a single long spot in about half the specimens). The spot in the cell is also large, almost quadrangular. The female was not found.

Male genitalia: The male genitalia are generally not diagnostic in the genus and have not been studied.

Discussion: The limited material of the A. jefferyi-complex available to us from across most of Africa indicates that the species probably ‘hides’ at least two, possibly more, different species. All differ from the present species in having two well-developed tails. Known material of the A. jefferyi-complex is from the forest zone, and the fact that Waak is a dry savannah area must also be taken into account.

Etymology: The species is named after Collins' son Marc and is pronounced with a hard c.

Iolaus (Philiolaus) likpe Collins & Larsen Iolaus (Iolaphilus) likpe Collins & Larsen, 2004. Metamorphosis 14: 90 (63-110).

Description of the unknown female: While collecting with Larsen in the Bobiri Butterfly Sanctuary in Ghana on 16.xi.2006, Szabolcs Sáfián collected a female Iolaus that differed from any previously seen by either. It was identified in the field as probably being the unknown female of I. likpe, a species so far known only from the male holotype from Likpe-Mate in Ghana's Volta Region – and the only record outside of the Volta biogeographical subregion. It turned out that ABRI had also recently obtained three females from Amedzofe in the Volta Region. Linking males and females in the genus without having material collected in copula is often difficult. The underside of the species is unique for the region, so we are almost certain that the true female is the one described below. Forewing 19 mm. The female is a duller blue than the male. There are no lighter areas. The black markings of the forewing are slightly different from the male holotype in that the base of space 3 is not black and the margin at the tornus is wider. The hindwing has the same tone of blue as the forewing. The costa is slightly lighter except at the base, which is powdered with blackish scales. The apical area of the hindwing is rather widely black, which continues as a 2.5 mm wide margin to the tornus. The tornus has two prominent red spots between the three tails, the one between the two upper tails being the larger. A dark postdiscal spot touches the marginal band just above the lower red tornal spot and two free spots are present in spaces 2 and 3. The underside is identical with that of the male as illustrated in the original description and in Larsen (2005); often females have darker edges and better developed submarginal and/or postdiscal lines on the forewing. Both male and female have a slight but clear red postdiscal line from the red spot in space 2 to the costa. The combination of the red tornal spots on the hindwing upperside, and the red postdiscal line on the hindwing underside are sufficient to tell the species from any other West African Iolaus. June 2008 METAMORPHOSIS, VOL. 19, No. 2 69

Iolaus (Philiolaus) likpe , upperside (left), underside (right)

Iolaus (Philiolaus) ofere Collins & Larsen, sp. nov.

A single large male Iolaus in frightful condition was collected on the Obudu Plateau during Collins' expedition in April 2007. Despite its condition, it clearly could not be the undescribed male of Argiolaus manasei Libert, 1993 from Mt Tabenken in the Cameroun Highlands since ABRI actually has both sexes of this, the male from the type locality (its placement in Argiolaus was since changed to Philiolaus by Collins, Larsen and Warren-Gash (2004)). We believed that Argiolaus should be defined also on the narrow genitalic grounds outlined by Stempffer & Bennett (1958) and not on host plant choice alone as advocated by Heath (1985).

The previously undescribed male of Iolaus manasei, upperside (left), underside (right)

I. manasei was described on the basis of a single female in good condition, well illustrated in the original description: we figure the male above. It looks almost exactly as might be expected from the female. The forewing upperside has a large black apical area and a broad margin. The hindwing has a prominent orange tornal spot and an additional red spot in space 2. The orange lines on the underside are well developed. The genitalia have not yet been examined. The Obudu male is described as a distinct species below since its genitalia are also distinct from all described species. 70 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Holotype: ♂, Nigeria, Cross River State, Obudu Plateau, 1,300 m (06 o26'N 09o19''E), 7/11.iv.2007(Ofere Agbor leg., coll. ABRI)

The holotype of Iolaus ofere , upperside ( left), underside (right)

Diagnosis: Male forewing 20 mm, making it quite a large species. The single specimen is unfortunately in such poor condition that an adequate description is impossible. However, microscopic examination indicates that the specimen is very dark. The costa is broadly black: no blue scales are present except very near the base. At the end of the cell the black apical area widens; there are just a few scales in space 3 and space 2 is blue only for a third of its length. The blue in space 1b leaves an unusually broad black margin (2 mm at least), which is in-turned along the inner margin to become wider still. The hindwing has a broad black margin (almost 2 mm) that appears to be of even width from tornus to apex. No large orange tornal spot can be seen. The grey androconial patch is situated in a large, black androconial area, devoid of scales, larger than in I. manasei. In contrast, the underside is lightly marked, effectively unmarked on the forewing, where the androconial brush is orange. On the hindwing the only discernible markings are two small red tornal spots, one smaller black such spot, a few rather faint submarginal black lunules, two tornal and one crowning the outer red spot, and an indistinct submarginal line in spaces 2-4.

The male genitalia of Iolaus (Philiolaus) ofere (holotype) June 2008 METAMORPHOSIS, VOL. 19, No. 2 71

Male genitalia: The genitalia (SCC 554) are not of the specialized types pertaining to the subgenera Argiolaus or Iolaphilus and thus place the species in Philiolaus. The uncus consists of two large, well-chitinized triangles, which are only narrowly separated from each other at their base (we are not aware that we have seen this configuration in any other species). There are two large, slightly curved subunci that swivel (ruling out any association with Argiolaus). The vinculum has two large lateral triangular flaps that are well chitinized. The valve has a simple tapering shape from an ovoid base, with a tiny serrated ridge on the ventral edge in the distal half. The fultura ends in a hoop where the two branches are fully fused. The penis is large, thickest at the distal end, which carries a ventral curved tooth that is strongly chitinized: on the dorsal side a movable cuneus projects. A slender cuneus is inside the penis (displaced during the cleaning process). The hoops formed by the fultura, the valves, and the penis are rather similar to those of I. aequatorialis Stempffer & Bennett, 1958, but the triangular flaps on the vinculum and the uncus are very different. We have not seen a similar uncus in any other members of the genus. The triangular flaps on the vinculum are larger even than in Argiolaus. Such flaps are present also in Philiolaus likpe and P. shaba Collins & Larsen, 1997, but are here much smaller and in the case of I. likpe placed closer to the tegumen. The genitalia fit no other described species.

Discussion: Many of the large Iolaus-species are procured only rarely during random collecting, but once a tree with the Loranthus host-plants is found, long series of a “rare” species can be obtained over time. On balance the likelihood is that this is a genuine submontane element, though several lowland Iolaus have been found in the submontane zone. It was fortunate the male genitalia were so different from any known species: without them it could not have been described. The Obudu Plateau is a unique habitat in Nigeria, the conservation of which must be of the highest priority. It is part of the Okwangwo Division of the Oban Hills National Park and as such has some protection. Recently the area has been opened up for tourism that, if carefully managed, should provide additional protection to the habitat, apart from the old Cattle Ranch area where the tourist facilities are based.

Etymology: We are pleased to dedicate this species to Ofere Agbor, a guide in the Obudu National Park. He usually takes people to catch a glimpse of the gorillas but he took so well to butterflies during Collins' brief visit that he came up with a new species.

Iolaus (Epamera) adorabilis Collins & Larsen, sp. nov.

During a visit to the Obudu Plateau on the Nigeria/Cameroun border in April 2007, Collins collected a series of a most distinctive species of Iolaus that could immediately be described solely on the basis of morphological characters. They have the smallest black apical area of any known Iolaus and the hindwing androconial patch is very small. On the underside there are dark bars closing the cells of both wings, which in West Africa is most unusual outside of the subgenus Iolaus. It is thus a singular butterfly that in both sexes can be recognized at a glance. 72 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Iolaus (Epamera) adorabilis, male upperside (left), male underside (centre), female upperside (right)

Holotype: ♂, Nigeria, Cross River State, Obudu Plateau, 1,300 m (06o 26'N 09o 19'E), 7/11.iv.2007 (S.C. Collins leg., coll. ABRI)

Paratypes: 3 ♂♂ 6 ♀♀, data as for holotype; 1♂, Cameroun, Bamenda Highlands, iv.1956 (F. Davey leg., coll. ABRI)

Diagnosis: Male forewing 15 mm. The frons is red. The forewing margin is unusually convex: it has a small lobe on the inner margin that matches the small androconial patch on the hindwing. The hindwing has only two well-developed tails, which is the norm in subgenus Epamera. The ground-colour is a beautiful brilliant, dark sky blue. The costa is narrowly black until after the cell, from where it widens to produce a black apical patch, the inner edge of which is sharply defined and curves regularly. It is widest along vein 5 and continues narrowly to the tornus. The extent of blue on the forewing is larger than in any other species known to us, except perhaps for I. carina Hewitson, 1873. The hindwing is also blue. The unusually small greyish-brown androconial patch is centered with black. There is a fine black marginal line, barely widening towards the apex, as well as a tiny tornal spot in space 1b. The underside is white. The bands of the underside are orange-yellow and not bordered by black, again unusual. The submarginal line of the forewing is black, the orange postdiscal line stops abruptly at vein 2. A well-developed orange bar closes the cell. On the hindwing the submarginal line is orange and much better developed. The bar closing the cell is narrower. The postdiscal line stretches from the costa almost reaching the eye-spots at the tornus, at which point the line turns black and continues some way up the abdominal fold. The margin is broadly orange in spaces 1a to 1c, with a black spot in the tornal lobe, a violet patch in 1b, and another black spot in 1c. There are two tails. The androconial brushes are blackish-grey. The female (forewing 16 mm) is slightly larger than the male. The blue ground-colour is darker and duller, but still quite bright. The black forewing margins are considerably wider than in the male, being about 2 mm at the tornus, though still less extensive than in most female Iolaus. The black hindwing margin widens to form a small black apical patch and the two black tornal spots are quite well developed. The underside is similar to that of the male. June 2008 METAMORPHOSIS, VOL. 19, No. 2 73

Male genitalia: The relatively simple genitalia (SCC 550) display a small, strongly chitinized uncus attached to a moderate tegumen. Where the uncus, which has no central depression, ends, there is a large non-chitinized gap where the solid subunci are loosely embedded in membranes: they seem ready to detach at any time. The subunci are strongly chitinized, with a bulbous base, tapering and finally expanding again at the tip, one side being rounded, the other ending in a point. The valves are simple and unadorned, being like a narrow ellipse with an in-turned small tip. There is a huge fultura, like a squat letter Y, with two large triangular extensions. The penis is relatively thick and without cornuti or other embellishments (not captured on the figure). The entire structure is very similar to that of the very rare I. flavilinea Riley, 1928.

The male genitalia of a paratype of Iolaus (Epamera) adorabilis

Discussion: The close relationship with I. flavilinea (TL: Bitje, Djah River) is apparent from the underside pattern as well as from the genitalia. Species of the subgenus Epamera are generally more readily met with than the larger species since they tend to descend low more frequently and to be more numerous where they occur in the forest zone. As the species can be recognized at a glance – and given that the single Cameroun male was also taken at altitude – it is presumable a genuinely submontane element, as are many of the other distinctive butterflies of the Obudu Plateau. Collins saw many specimens at tree-crown level where a forested scarp allowed access to the canopy, including females flying around Loranthus.

Etymology: The name stems from the fact that it is simply such an adorable little butterfly, with its intense blue colour and delicately marked underside. 74 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Polyommatinae

Thermoniphas leucocyanea Clench Cupido bibundana Grünberg, 1910. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1910: 478 (469-480). (TL: Cameroon, “Bibundi”) Thermoniphas leucocyanea Clench, 1961. Annals of the Carnegie Museum 36: 56 (49-62). (TL: Cameroon: “Lolodorf”)

Redescription and illustration of a nearly unknown species: In 1996 Larsen collected a small series of the nearly unknown Thermoniphas bibundana at Toko, 1,000 m, in the hills adjoining Korup National Park in Cameroun. They closely matched the original description and were figured in monochrome and re- described in Larsen (2005). Michel Libert (pers. comm.) kindly informed us that the types in the Museum für Naturkunde, Humboldt University, Berlin matched this description well. The outstanding characteristic is the total lack of discal and postdiscal spots on the underside. A long series is now available in ABRI from Bioko. Larsen (2005) speculated that T. leucocyanea might be a junior synonym of T. bibundana. The unique types (a single pair) of this taxon were photographed during a recent trip to the Carnegie Museum and are very similar to T. bibundana. The male lacks the white shading on the forewing discal area, and the white costa does not impinge on the discal area of the hindwing; however, scattered white scales are actually present in those areas. However, on the underside the postdiscal and submarginal bands are displaced even further towards the margin and the postdiscal band is very short in both sexes (resembling the single male from the Natural History Museum, London figured by D'Abrera (1980)). Bibundi (on the western slopes of Mt Cameroun (Mt. Buea)) and Korup are both inside the biogeographical subregion bordered by the Cross River in the west and the Sanaga River to the east, which has affinities also with Bioko. The types of T. leucocyanea are from Lolodorf, well to the southeast and in another biogeographical subregion (while the manuscript was in press, G. van de Weghe kindly informed us that he had caught the species also in Gabon). Examination of the male genitalia shows them to be virtually identical with the ones figured by Clench in the description of T. leucocyanea, the only difference seems to be one fewer of the long setae on the valve. Larsen (2005) illustrates the valve, which in his drawing is missing two of the fragile setae. We tentatively maintain T. leucocyanea as a distinct species, pending more material.

June 2008 METAMORPHOSIS, VOL. 19, No. 2 75

Top row: Thermoniphas bibundana: Male upperside and underside followed by female upper- and undersides (slightly enlarged). Bottom row: The same for male (holotype) and female of Thermoniphas leucocyanea

Thermoniphas bibundana

Thermoniphas leucocyanea

Nymphalidae Satyrinae

Bicyclus sealeae Collins & Larsen, sp. nov.

It is interesting that a brief visit to Bioko should have uncovered a new species of Bicyclus. Since Condamin (1973) provided his comprehensive review of the genus, recognizing 77 species, only nine additional species were described. Five are from Tanzania (uzungwensis Kielland, 1990, mahale Congdon, Kielland & Collins, 1998, kiellandi Condamin, 1986, tanzanicus Condamin, 1986, pareensis this paper), two from Cameroun (ewondo Libert, 1996 and amieti Libert, 1996), one from Gabon (ivindo van de Weghe, 2007), and one from Bioko (sealeae this paper). Seven of 76 METAMORPHOSIS, VOL. 19, No. 2 June 2008

these are more or less submontane, only B. amieti and B. ivindo being lowland. The new species was collected at 1,700 m and is probably wholly submontane. Doubtless many other species still await discovery in the mountains of Bioko.

Holotype: ♂, Equatorial Guinea, Bioko, nr Moka, 1,700 m (03o 33'N 08o 67'E), 8-13.iii.2007 (S.C. Collins leg., coll. ABRI).

Paratypes: 10 ♂♂, 2 ♀♀, same data as holotype

Bicyclus sealeae: male holotype (left), female upperside (right), and male underside (below)

Diagnosis: Male upperside 23 mm. The ground-colour is a dark blackish-brown. The forewing has a diffuse white band stretching from the upper part of space 3 but not reaching the costa. The subapical spot is fully developed and impinges somewhat on the band. The large eye-spot in space 2 is obscured and the pupil hardly visible. The white band stops above this eye-spot. The hindwing is the same colour as the forewing, the margin of the wing being slightly lighter. The following androconial features are present: on the forewing vein 1 is slightly bent and above the vein is a small black hair-tuft directed outwards. The hindwing has a black costal hair-tuft that is placed in a large grey androconial area. A smaller such hair-tuft is in space 1b and a yet smaller one at the base of space 2. There is also a small tuft inside the cell next to the one in 2. The male underside is brown with the orange tinge also found in B. trilophus, B. sciathis, and B. procora. The forewing apical eye-spot is well developed and the June 2008 METAMORPHOSIS, VOL. 19, No. 2 77

eye-spot in 2 is large, impinging strongly on spaces 1b and 3. There are faint traces of a brown sub-basal line on both wings, almost invisible to the naked eye. On the forewing the white apical area is well delineated and purer white than on the upperside: it fills out the entire area between the discal line, the two eye spots, and the inner of the two submarginal lines, but not quite reaching the costa. The discal line is relatively straight, though bending inwards at the level of the cell. The submarginal eye-spots of the hindwing are fully developed, that in space 2 being the largest and that in 6 being somewhat larger than the rest. All eye-spots are firmly ringed with yellow. The female is larger than the male (forewing 25 mm). The ground-colour of both wings is lighter. On the forewing upperside the subapical band is larger, purer white, better defined, and extending closer to the costa. She is otherwise similar to the male. The male is more or less unmistakable, since no other male with a similar underside has a white subapical band in the Cameroun/Bioko area. The female is quite similar to that of B. sciathis: the best distinction lies in the fact that the white subapical area curls round much of the apical eye-spot, while it does not even reach the eye-spot in B. sciathis.

Male genitalia: Were it not for the fact that we had already noted that the arrangement of androconial brushes on both wings was similar to that in B. trilophus, we would have been very surprised to see the genitalia (SCC 553), which closely resemble those of that species, but differ strongly from those of all other Bicyclus species. The main difference lies in the much shorter length of the tegumen. The valve is slightly broader. The impossibly long subunci seem a little less dramatic. We noted a tuft of hair very firmly attached to the small area where the tegumen fuses with the uncus: it could not readily be brushed off and appeared to be an integral part of the genital structure. The vinculum is so small that the valves might as well have been attached directly to the tegumen itself in both species – it is a feature that differs from all other Bicyclus.

The male genitalia of Bicyclus sealeae (left) and B. trilophus (right) (after Condamin, 1973) 78 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Discussion: Before comparing the genitalia we thought it might just be possible to consider B. sealeae a subspecies of B. trilophus, but apart from the prominent white band in both sexes and the genitalia, there are others differences: inter alia the upperside ground- colour is darker and more blackish; the large forewing upperside eye-spot in space 2 is less clearly ringed with yellow; the light discal band on the underside is of a different tone (blueish-white rather than light tan); and the sub-basal lines of both wings are less prominent. Interestingly, the species was flying together with B. sciathis, the female of which is almost identical. The species is almost certainly an endemic of the submontane zone of Bioko, though it might just hide somewhere in the highest mountains of Cameroun. B. feae Aurivillius, 1910 is another Bicyclus species endemic to Bioko.

Etymology: We are pleased to name this butterfly after Jennifer Seale who has contributed much to conservation efforts on Bioko and who helped facilitate Collins' visit to the interesting submontane area of the type locality.

Bicyclus pareensis Collins & Kielland… , sp. nov.

This species was sent for description in a Dutch journal by Jan Kielland shortly before his death, but he included it as a manuscript name with a colour illustration in his book on Tanzanian butterflies (Kielland 1990). It was again mentioned as pending publication by Congdon & Collins (1998) in their update of Kielland's book. D'Abrera (1997) included and figured the species as a valid taxon on the authority of Kielland. Williams (2005) cautions that it is a manuscript name. It was thus never formally described. The species was extensively discussed with Collins by Kielland at the time, and he also commented on the draft paper. It proved impossible to track down copies of the manuscript to secure its posthumous publication after Jan's untimely death in 1995. It is here formally described; nothing is included that would not have been in Kielland's own paper, except that special efforts from ABRI have procured a much larger type material. We consider it appropriate to publish the name with Kielland as a posthumous co-author as a late testimony to his prodigious work with the butterflies of Tanzania and beyond.

Holotype: ♂, Tanzania, South Pare Mts. (03o 45'S 37o 52'E), 14.iv.1978 (Jan Kielland leg., coll. Kielland in ABRI)

Paratypes: 26 ♂♂ 12 ♀♀ from the South Pare Mountains, by various collectors at various times are formal paratypes (coll. ABRI).

Kielland distributed pre-publication paratypes to many institutions and individuals: these ought to be paratypes since all are from the South Pare Mountains, but cannot formally be designated as such. June 2008 METAMORPHOSIS, VOL. 19, No. 2 79

Bicyclus pareensis: Top row male (holotype) upperside (left) and underside (right). Bottom row female upperside (left) and underside (right)

Diagnosis: Male forewing 23 mm. The species is close to B. danckelmani Rogenhofer, 1891 which flies in large parts of east-central Tanzania. The two are not sympatric, though the latter flies in the neighbouring Usambara Hills. This is a larger species with a more blackish ground-colour. The ground-colour of B. pareensis is dark brown. There is a small subapical eye-spot on the forewing and a larger such spot in space 2: the spots are black with tiny pupils and any yellow ring is poorly developed. Between spaces 1b and 3 the narrow white band of the underside shows through as a yellowish band, with varying degrees of development. On the hindwing the outer half is clearly lighter than the inner. The male has an inner costal hair-tuft of greyish-white hairs and a smaller such tuft in darker grey at mid-costa. The underside is characterized by a narrow white postdiscal band that divides the wings into a dark basal two-thirds and a lighter distal one-third. The eye-spots are not divided from this band by one of the light lines that usually run along the row of eye-spots in other species. The inner two-thirds of the wings are very dark, almost blackish, and the sub-basal line running through the cells is hardly visible. The white postdiscal line is abruptly angled at vein 4 and turns slightly outwards to just under the large eye-spot in space 2. On the hindwing the discal line is almost straight, but slightly curved outwards in the area of veins 3 to 5. The outer third of the wings is lighter and the area of the eye-spots is overlaid with violet. The forewing has a small subapical eye-spot and a moderate spot in space 2. The hindwing has a complete row of rather small eye-spots, that at the apex and the three most tornal ones being the largest. The base of space 1b on the forewing forms a white triangle surmounting a small white androconial patch on vein 1, evidently 80 METAMORPHOSIS, VOL. 19, No. 2 June 2008

linked to the inner androconial tuft, and a grey patch at the middle of the vein, linked to the outer tuft. Vein 1 is twisted in its basal half. The female is larger (25 mm) and much lighter brown. The forewing eye-spots are larger and the yellowish postdiscal line stretches as far as on the underside. The underside is also lighter allowing the sub-basal lines to be fully visible. There is only modest seasonal variation in either sex. The species is quite unmistakable.

The male genitalia of Bicyclus pareensis

Male genitalia: The genitalia (SCC 555) are close to those of B. danckelmani: the valve is proportionately wider in the basal half, the uncus is shorter, the subunci are shorter without being upturned at the tip, and the penis is strongly bent. The presence of the raised and serrated hump in the middle of the dorsal edge of the valve is unique to the three species of the B. danckelmani-group, which in addition to B. pareensis includes B. simulacris Kielland, 1990, from submontane areas in western Tanzania, south to Malawi and Zambia.

Discussion: The South Pare Mountains are not far from the main Usambara Mountains (of which the North Pare Mountains are biogeographically a part). That such a distinctive vicariant of the common B. danckelmani should have evolved there is rather surprising, but the species is well isolated since it rarely flies below 1,700-2,000m. The southwestern vicariant, B. simulacris, is much less differentiated from B. danckelmani.

Etymology: The species is named after the South Pare Mountains, where it is widespread and locally common. June 2008 METAMORPHOSIS, VOL. 19, No. 2 81

Limenitidini

Euriphene larseni Hecq, 1994, syn. nov. Diestogyna saphirina Karsch, 1894. Entomologische Nachrichten 20: 220 (209- 240). (TL: DRC, Ituri-Fähre (W. Albert Njansa)). Diestogyna itanii Carcasson, 1964. Journal of the East Africa Natural History Society & Coryndon Museum 24 (4): 64 (62-67) (TL: Tanzania: “Makuyu, Kigoma, Tanganyika) Euriphene larseni Hecq, 1994. Revue d'Entomologique Générale 8: 38 (1-62) (TL: Ghana, Aburi) [Patria falsa]

Species relegated to junior subjective synonym: The species was described from a poor photograph, taken by Larsen in late 1993, of a pair in the Father Theodor Maessen collection in the Allyn Museum, Sarasota (now in the McGuire Center, Gainesville) and sent to J. Hecq, who described it without further consultation in his revision of the genus Euriphene (Hecq 1994), then in proof stage. The labels indicated that they were from Aburi, Ghana, a well-known butterfly locality near Accra. This is now a popular tourist and picnic destination, which Maessen visited only intermittently, probably mainly on social occasions. Larsen (2005) wrote that it “seems increasingly likely that the Ghana specimens were mislabelled and that the name E. larseni is a junior synonym of E. saphirina or one of its subspecies”. We have since found that during the 1970s various species of Euriphene were exchanged between Maessen in Ghana and Jan Kielland in Tanzania, some of which were seen in the Maessen collection at the McGuire Center earlier this year. We are now wholly convinced that some simple mistake is involved – probably just mis-labelling. We consider E. larseni to be a junior subjective synonym of E. saphirina itanii (Carcasson, 1964) (syn. nov.), specimens of which were very likely sent from Kielland to Maessen: other Kigoma material of his is in the Maessen collection. No subspecies of E. saphirina, or of similar species, are known much west of the Rift Valley in the DRC.

Euriphene (Doricleana) paralysandra D'Abrera Euriphene paralysandra D'Abrera, 2004. Butterflies of the Afrotropical Region. Part I1 [revised]. Hill House Publishers, Melbourne & London: 366 (TL: Nigeria, Cross River Loop, Ikom)

Validation and range extension of a recently described species: This species was recently described from a single male and two females collected at Ikom in the Cross River Loop of Nigeria. Larsen (2005) wrote that “it certainly seems to be a valid species”. While curating the Euriphene at ABRI during 1995/96, we had actually already isolated a series of E. paralysandra from Nigeria (Okwangwo, Oban Hills, Makurdi) along the Cameroun coast as far southeast as Kribi as probably being a new species, but never formalized it (9 ♂♂ and 9 ♀♀). Thus, D'Abrera's description is fully justified, but the range extends much beyond Ikom to southeastern Cameroun and probably even Gabon. The species is well figured by both D'Abrera (2004) and Larsen (2005). While on average both sexes of E. paralysandra are larger than those of E. lysandra, there is some overlap. 82 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Bebearia staudingeri (Aurivillius) Euryphene staudingeri Aurivillius, 1893. Entomologisk Tidskrift 14: 199. (TL: Cameroun: “Camerun: N'Dian)

This fine species has always been considered among the most stable and unproblematic of the Bebearia, which was aided by the fact that it is generally rare. Large numbers of males from different areas in Africa are not available in one place. The range is generally given as Cameroon, Gabon and the DRC (Equateur, Tshuapa, Sankuru) and that is indeed where most material comes from. Larsen (2005) records two old specimens also from the Cross River Loop and another from Benin City in Nigeria. Robert Warren has since collected a series in the Okomu Nature Sanctuary in western Nigeria, near Benin City. These Nigeria specimens are smaller than those from Cameroun, the type locality, and the Cross River Loop, also having a distinctly narrower subapical band of the forewing. It is so far known only from around Benin and appears to be isolated from the Cross River Loop population. It is described as a new subspecies. The nominate subspecies is known from the Cross River Loop, as well as from many localities in Cameroun and Equatorial Guinea. To our surprise the population in the Central African Republic also differs quite strongly from the nominate subspecies in Cameroun and is also described as a new subspecies.

Bebearia staudingeri male upperside: Top row: the nominate ssp. staudingeri. Bottom row: ssp. carensis (left) and ssp. okomu (right) (0.9 x natural size) June 2008 METAMORPHOSIS, VOL. 19, No. 2 83

Bebearia staudingeri okomu Collins & Larsen, ssp. nov.

Holotype: ♂, Nigeria, Edo State, Okomu National Park (06o 26'N 05o 10'E), xii.2005 (R. Warren leg., coll. ABRI)

Paratypes: 2 ♂♂ 3 ♀♀, same data as holotype (various dates); 3 ♂♂ 1 ♀, same locality as holotype (various dates) (leg. et coll. R. Warren)

Diagnosis: Male forewing 34 mm, slightly less than in the nominate subspecies (37 mm). The forewing is more squat than in the other two subspecies. The hindwing is more rounded than in the nominate. The subapical band is much narrower and slightly deeper orange. The tone of the brown colour is a warmer chestnut. The white subapical band of the female is also narrower. There are no obvious underside differences.

Discussion: Larsen (2005) knew of only three old Nigerian records. Two were from the Cross River Loop, which is biogeographically the same as western Cameroun. One was from Benin City, quite near Okomu, where Robert Warren did much of his collecting, managing not only to get a series of this butterfly but even to photograph the species in the wild (photo in Larsen 2005). Okomu forms part of the Niger Delta refuge, which on one or more occasions has evolved its own flora and fauna. The flagship for this fauna is an endemic monkey, Cercopithecus erythrogaster. There are already three well-defined butterfly subspecies: Cymothoe hesiodutus nigeriensis Overlaet, 1952 [possibly even a distinct, endemic species] which recurs only east of the Sanaga River in Cameroun, C. hypatha okomu Hecq & Larsen, 1997, and Bebearia flaminia leventisi Hecq & Larsen, 1997, all of which are isolated from the main populations of their species. Robert Warren has only seen the species near streambeds and speculates that host-plants may be linked to water. Males are not usually seen flying more than a metre above the ground, but females may fly higher up.

Etymology: We are pleased to name this new subspecies after Okomu National Park, one of the most important conservation areas in Nigeria, since it protects a forest system that on occasion has been an isolated refuge.

Bebearia staudingeri carensis Collins & Larsen, ssp. nov.

Holotype: ♂, Central African Republic, Moloundou [near Bangui], (04o 30'N 17o 45'E), xii.1997) (ABRI leg. et coll.). 84 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Paratypes: 23 ♂♂ 12 ♀♀, Central African Republic, various localities and years (ABRI leg. et coll.).

Diagnosis: Male forewing 37 mm, as in the nominate subspecies. The hindwing is intermediate between the nominate subspecies and that of the rounded shape of ssp. okomu. The subapical band is narrower than in the nominate and is a of more insipid, lighter ochreous. The brown areas of both wings are a warmer and lighter brown. The three almost parallel bands of light spots on the forewing (discal, postdiscal, and submarginal) are brighter and more visible. The differences are especially apparent when two series of these subspecies are seen side-by-side. The female subapical band is also narrower than in the nominate subspecies. There are no obvious underside differences.

Discussion: The presence of a distinct subspecies in Central African Republic is not necessarily that surprising, since the assumption must be that it also occurs in parts of Congo and northwestern DRC.

Etymology: The name is based on the English abbreviation of the Central African Republic (CAR).

ABRI does not have sufficient material from the eastern DRC except to say that material from both Kivu and Equateur is a darker brown without the lighter overlay of the other subspecies, and that the females in these two areas differ in having almost half the hindwing black with a blue overlay, instead of just the apical area as in the subspecies discussed above.

Bebearia denticula Hecq Bebearia denticula Hecq, 2000. Nymphalidae IV, Bebearia. Butterflies of the world, Goecke & Evers, Keltern.

Validation of an almost unknown species and assignment of type locality: This species was described from a single female of uncertain origin (? Nigeria). Larsen (2005) did not accept it as a valid species for Nigeria. ABRI has a female from the Central African Republic (Bangui 1995). The type is almost certainly a specimen that was sent to Tervuren some 8-10 years ago from ABRI with other material collected at the same time near Bangui, which is herewith designated as the type locality. The second specimen is effectively identical with the holotype and is figured below: the falcate forewing is very unusual for a female. The light, matte greyish-brown tone is different from that of other species. Our guess is that the male is quite similar … but it might just be very different! June 2008 METAMORPHOSIS, VOL. 19, No. 2 85

Bebearia denticula ♀: Upperside left, underside right (0.9 x natural size)

Pseudathyma legeri Larsen & Boorman Pseudathyma legeri Larsen and Boorman, 1995. Lambillionea 95 (4) (Tome II): 611 (611-613). (TL: Nigeria: “Obudu Plateau, Cattle Ranch, 1500 m, IV.1959 (F. Davey)”

Description of a dimorphic female: This species has always been considered very rare, but since it was first discovered it been collected on a few occasions in the vicinity of the Cattle Ranch at Obudu, where Larsen last observed it in 1996. Still, less than a dozen were present in collections. Collins was fortunate to come across a mass hatching of this butterfly during a visit to Obudu in April 2007 and to his surprise found that the females were dimorphic, one being white-banded as the described female, the other being almost as yellow as the male. They appeared in a roughly 50:50 ratio. The dimorphism is genuine since no intermediates were caught or observed. This is the only Pseudathyma known to be dimorphic. Cases are found within other groups of the Adoliadini, notably in Aterica galene (Brown, 1776) and Bebearia mardania (Fabricius, 1793), but it is not at all frequent amongst the 425+ members of this forest tribe. The species is still endemic to the Obudu Plateau and has not been found in any of the Cameroun Highlands. It is a pleasure to report on such a vigorous population of this very special butterfly, the closest relative of which seems to be P. uluguru Kielland, 1985 from eastern Tanzania.

The typical white morph of Pseudathyma legeri and its yellow morph 86 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Acraea eugenia ochreata Grünberg Acraea eugenia var. ochreata Grünberg, 1910. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1910: 470 (469-480). Equatorial Guinea: “Spanisch-Guinea, Makomo, Ntumegebiet”.

Validation of an ignored subspecies: Acraea eugenia Karsch, 1893 was described from near Bismarckburg in Togo. It is sometimes numerous at the southern end of the Kyabobo National Park (just 30 km from the type locality) and at Wli Falls further south in Ghana's Volta Region. C. Belcastro collected one specimen from the Atewa Range in central Ghana: no-one else has recorded it from this heavily exploited locality: a single male is known from near Lagos in Nigeria, collected by Eva Larsen. These are probably strays from the Volta Region or Togo. There are very scattered records from Cameroun, Equatorial Guinea, Gabon, and Angola, as well as from (Shaba: Berger & ABRI) in DRC and Uganda. ABRI has a large series from Waak and Man i n Cameroun, one of which was figured by Larsen (2005). While the Lagos specimen is identical with those in Ghana, the Cameroun series is sufficiently distinct to deserve subspecific status. The colour of the hindwing is slightly darker ochreous and the spotting is more extensive, but there is much variation. A female from the dry area of Waak has virtually no hindwing margin while a male has the entire hindwing with the colour of the margin. The name var. ochreata is available, with a type locality that is now in mainland Equatorial Guinea (Makomo, Ntum area). The f orewing end-cell spot is better developed, the hindwing colour is deeper ochreous-brown, the black spotting larger, and the hindwing margin inwardly better defined. The differences are not that great but most apparent in series. We have only seen two specimens from Shaba: they may belong to a further subspecies. We did not see specimens from Uganda and Angola.

Acraea eugenia; A male of ssp. ochreata from Cameroun (Waak) and a male of the nominate subspecies from Ghana (Wli Falls)

Hesperiidae Coeliadinae

Pyrrhochalcia iphis dejongi Collins & Larsen, ssp. nov. Papilio phidias Cramer, [1779] and Papilio jupiter Fabricius, 1787 are the only synonyms. The type locality for both is Sierra Leone and they are thus typical P. iphis. June 2008 METAMORPHOSIS, VOL. 19, No. 2 87

During a large study of the phylogeny of the Coeliadinae based on morphological characters (preliminary results in de Jong, 2007), Rienk de Jong found a number of old males and females of this species in relatively poor condition, the wings of which had a blue metallic sheen. He was so intrigued by this that he checked the genitalia against West African material, finding no differences. He sent us pictures of these and we located three DRC males, which are here described and figured as a distinct subspecies. A specimen from ABRI is chosen as the holotype because of its near- perfect condition.

Holotype: ♂, DRC, Equateur, Kuluboku, 60 km from Mbandaka (00o 01'N 18o 20'E), 1996 (Ph. Oremans leg., coll. ABRI)

Paratypes: 2 ♂♂, same data as holotype; 1 ♂ DRC, Mayoumbé (coll. Musée Royal de l'Afrique Centrale (MRAC), Tervuren); 7♂♂, 22♀♀ Congo, Etoumbi, ex le Moult (coll. National Museum of Natural History, Leiden)

Pyrrhochalcia iphis dejongi: Male holotype (top) compared with a Ghana male of the nominate subspecies (bottom). 88 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Diagnosis: The male upperside of nominate P. iphis (Drury, [1773]) between Sierra Leone and most of Cameroun is a beautiful, profound, velvet black imperceptibly tinged with dark blue. Ssp. dejongi has a blue metallic sheen, except for the basal part of the hindwing, curiously reminiscent of the West African endemic Pyrrhiades lucagus (Cramer, [1777]). When viewed simultaneously the two are quite distinctive. Despite much experimentation we have found no way to produce a metallic sheen in specimens from Ghana, in the specimens themselves or in photographs. There are no obvious underside differences, nor do the females differ much, though de Jong tells us that there is also a slight blue sheen (we have not been able to view females).

Male genitalia: According to de Jong (pers. comm.) there are no genital differences between the two subspecies, so we did not re-examine the genitalia.

Discussion: The surprising suggestion by de Jong that a new subspecies of this characteristic and well-known butterfly needed description turned out to be correct. P. iphis is found from Sierra Leone through the West African rainforest zone, south to Gabon and the Congo Mayoumbé, and east to the Central African Republic and the westernmost parts of the DRC. A few specimens in the Natural History Museum, London from Bioko and one from southwestern Cameroun show signs of transition between the two subspecies. The new subspecies escaped notice until now due to a combination of factors: a) the species is so evident that it is never examined closely, b) material from the eastern end of its range is scarce in collections, c) few museums have significant holdings from Central Africa, and d) most material of ssp. dejongi is in poor condition, while there are hundreds of males of the nominate subspecies in perfect condition from the area between Sierra Leone and Cameroun. The existence of a distinct subspecies in the extreme east of the range of the species is biogeographically plausible: other species and subspecies cover the Congo, Gabon, Central African Republic subregion of the main equatorial forests. The absence of the species further east than recorded here is rather surprising.

Etymology: Shortly before his retirement from the National Museum of Natural History in Leiden, The Netherlands, Rienk de Jong drew our attention to the fact that anomalous specimens of the largest African skipper (P. iphis) were in their collections. We take pleasure in naming it after him, wishing him well in retirement, and hoping that his work on the Hesperiidae and biogeography will continue unabated: we very much look forward to his long awaited and much needed monograph on the genus Metisella.

Pyrginae

Celaenorrhinus sagamase Collins & Larsen Celaenorrhinus sagamase Collins & Larsen, 2005 in Larsen, 2005. Butterflies of West Africa: appendix 1: 556 (plate 125) (TL: Ghana, Atewa Range, Sagyamase Trail) June 2008 METAMORPHOSIS, VOL. 19, No. 2 89

Description of the unknown female. A female of this recently described species was located by Larsen amongst the Maessen material from the type locality at the McGuire Center, Gainesville in October 2007. This single female can simply be described as being very similar to the male, but with both pairs of wings slightly more rounded and with marginally larger hyaline spotting. An illustration would seem superfluous. An additional male from Begoro on the Kwahu Escarpment in Ghana was located in the collection of the National Museums of Scotland, Edinburgh, being the third known locality apart from the Atewa Range and Kakum National Park. As currently known the species is a Ghana endemic. Ugo Dall'Asta kindly confirmed that all Côte d'Ivoire material in the MRAC Collection pertains to C. rutilans (Mabille, 1877).

Abantis ja usheri Collins & Larsen, ssp. nov. Abantis ja Druce, 1909. Proceedings of the Zoological Society of London 1909: 408 (406-413). (TL: Cameroon: “Bitje, Ja River, Cameroons, 2000 feet”) Abantis tanobia Collins & Larsen, 2005. Butterflies of West Africa, appendix 1: 558- 559. (TL: Ghana, Tano Ofin)

Father Theodor Maessen collected a single female of Abantis ja on the Atewa Range in Ghana in 1980, which was discussed in detail by Usher (1984). He concluded that it probably pertained to a distinct West African subspecies. ABRI collectors obtained very small specimens of what at first seemed to be Abantis ja in 2003 and 2004, but they proved to be a valid species in its own right that was described as Abantis tanobia Collins & Larsen, 2005: the genitalia were clearly different from those of A. ja, and apart from other differences, the hyaline spot in space 2 is at the very base of space 2 in A. ja, while in A. tanobia it is displaced well outwards. This trait and the small size of the species have been confirmed by several additional specimens from the Atewa Range, but a male A. ja was also procured at Atewa. Usher's photo of the female A. ja was not all that clear, but Larsen photographed the specimen during a recent visit to the McGuire Center for Lepidoptera at the University of Florida, into which the Maessen collection has been incorporated. The four (two were found while the manuscript was being processed) Ghana specimens are clearly distinct from long series from Cameroun, Congo, and the Central African Republic at ABRI. We have no hesitation in following Usher's suggestion that it deserves subspecific status.

Holotype: ♂, Ghana, Central Region, Atewa Range, (06 o 16'N 00 o 34'W), ii.2007 (ABRI leg. et coll.)

Paratypes: 1 ♀, Ghana, same locality, 5.viii.1980 (T. Maessen leg., coll. McGuire Center); 2 ♀♀, Ghana, Western Region, Bibiani, i.2008 (R. Vorgas leg., coll. ABRI). 90 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Abantis ja usheri: Male holotype upperside (left) with the female to the right. Below a male of Abantis ja ja

Diagnosis: Male forewing 18 mm. This subspecies differs from the nominate A. ja ja in the following respects: the forewing is more pointed and the hindwing is more drawn out at the tornus (one nominate A. ja from Cameroun in ABRI also has this hindwing character, but none of about 50 has the forewing shape). The veins are broadly black as is a vein-like streak in the centre of space 1b. The interspaces are a dusky ochreous-grey, lighter on the hindwing. In all markings in both sexes the present subspecies is lighter than in nominate material. In the male of ssp. usheri the ochreous scaling diminishes rather abruptly distally, leaving a faintly darker margin to the wing that is not apparent in the nominate subspecies. The most vivid difference lies in the hyaline spots of the forewing: in the nominate subspecies all spots are well-defined and clear. In the new subspecies they are so narrow or small that they are almost invisible to the naked eye. There are four subapical spots that are narrow and elongate, compared with usually only three shorter, but more prominent, spots in the nominate subspecies. Both cell-spots are narrow and obscured streaks, the upper cell-spot being particular long, reaching the end of the cell (in the nominate subspecies the upper cell-spot is usually shorter than the lower, but both are much better defined). The spot in space 2 is both distinctly smaller and more obscure, that in 3 less so. The hindwings of both subspecies are lighter than the forewings, but those of ssp. usheri much lighter than in the nominate subspecies. Both have the cell occupied by two large hyaline spots, divided by a black longitudinal line and a further hyaline spot is in the space just above the cell. There is some black scaling distally of the hyaline spots, while the rest of the wing is scaled ochreous between the heavily blackened veins. The underside is largely ochreous, with some slightly darker areas, and the veins are finely black. The hyaline spots in the males remain less visible in ssp. usheri. The characters outlined will also set the new subspecies apart from the much smaller A. tanobia that differs significantly in the genitalia and in which the spot in space 3 is displaced outwards, far from the base of the cell. The female hardly differs from the male, except that the hindwing is more rounded and June 2008 METAMORPHOSIS, VOL. 19, No. 2 91

the faint forewing margin is not present. The two additional females collected in January 2008 do not differ from the one figured.

Male genitalia: We can see no significant difference between the genitalia (viewed) and those of A. ja ja (figured by Larsen (2005)).

Discussion: This is one of the cases where a decision on whether to choose subspecific or specific status is a difficult call. The external characters are an indication of specific status, but the genitalia are not, though other closely related species in the genus may have very similar genitalia. We describe it as a subspecies on the understanding that it could be upgraded to species rank once more material of both sexes is at hand. It is interesting that A. ja and A. tanobia should be sympatric on the Atewa Range in Ghana. There is a record of A. ja also from Danané in Côte d'Ivoire: this might be A. tanobia. The fact that only two West African specimens have been collected, at an interval of more than 25 years, demonstrates how difficult it can be to obtain material of this genus [just before going to press two additional females of A. ja usheri were collected near Bibiani in Ghana by ABRI collectors].

Etymology: We are pleased to dedicate this new subspecies to Michael B. Usher. He collected intensively in Ghana over a period of several years (1959-60, 1971-73 and 1975), recording many firsts for the country and writing a number of interesting and useful papers, in addition to the one on Abantis. His fine collection of Ghana butterflies was incorporated into the Nations Museums of Scotland, where it was reviewed by Larsen in May 2007. Hesperiinae

Gorgyra warreni Collins & Larsen, sp. nov.

In April 2007 Collins collected two males of an undescribed Gorgyra in the submontane zone of the Obudu Plateau in eastern Nigeria. These are the males are apparently conspecific with a single Obudu female in the MRAC collection, Tervuren (C.S. Lewis leg.), placed by L. Berger under the manuscript name Gorgyra dubia. Since the males belong without doubt to a distinct species we shall not use the manuscript name dubia, which would anyway misrepresent this interesting skipper. At first sight the species might seem to be either of two described species in the group of species with a well-developed androconial tuft at the tornus of the hindwing upperside, though with an extreme reduction in the size and number of spots: G. bina Evans, 1937 also has only two hindwing spots, but its abdomen is clearly tipped with white; G. kalinzu Evans, 1949 lacks the white abdominal tip, but always has three hyaline spots on the hindwing.

Holotype: ♂ Nigeria, Obudu Plateau (06 o 25'N 09o 19'E), iv.2007 (S.C. Collins leg., coll. ABRI) 92 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Paratypes: 1 ♂, same data as holotype; 1 ♀, Obudu Cattle Ranch (C.S. Lewis leg., coll. Musée Royal de l'Afrique Centrale (MRAC, Tervuren)

Top row: Gorgyra warreni (Holotype) upperside left and underside right. Bottom row: Gorgyra bina upperside and underside

Diagnosis: Forewing 15 mm, quite large for a Gorgyra of its group. This species is characterized by the extreme reduction in the number and size of the usual blueish-white hyaline spotting. The upperside has a tiny, defined upper cell-spot with no trace of a lower spot (in virtually all other Gorgyra the lower spot is distinctly larger than the upper, and if one spot is missing, it is the upper of the two). The spot in space 2 is narrow and slightly out-curved. The spot in space 3 is small and almost quadrate. There is a single lower subapical spot. The ground-colour is dark blackish-brown. The hindwing is the same colour with two small hyaline postdiscal spots, that in 2 smaller than the one in 3. The tornal hair-tuft is black. The cilia is blackish, with a greyer edge. The forewing underside is dark brown, with only modest lighter brown shading along the costa or in the subapical area. Spaces 1a and 1b are slightly lighter than the rest of the wing. The spotting is as on the upperside but with the addition of a smallish non-hyaline off-white patch in the centre of space 1b. The hindwing ground-colour is olivaceous-brown with a slight dusting of yellow scales. The hindwing has indistinct darker spots, including a diffuse submarginal row. It does not have a spot in the cell, where G. bina usually has a tiny white spot circled with black. The two white postdiscal spots are clearly marked and circled with black. As in several other species there is a diffuse yellowish radial streak along space 1b. The abdomen is not tipped with white. The forewing is more pointed than in G. bina. Ugo Dall'Asta kindly sent a photo of the G. dubia female, taken by Frans Desmet. It is almost certainly the female of G. warreni and is included in the type series. The hyaline spotting is better developed, with one large and two small subapical spots. The spot in 2 is almost quadrate and there is a clear spot in space 1b. The strongest indication that it is indeed the female of this taxon is the presence of a strongly developed upper cell-spot, without even traces of a lower spot. June 2008 METAMORPHOSIS, VOL. 19, No. 2 93

The genitalia of Gorgyra warreni (left) compared with those of G. bina (right) (penis missing)

Male genitalia: The male genitalia (SCC 556a) are close to those of G. bina. The uncus is somewhat stouter and the dorsal edge of the distal triangular area is less strongly spined. The overall aspect of the valve is somewhat more slender. We had expected stronger genital differences from other species, since in most of the recognized species these are quite clear.

Discussion: The fact that the species is known only from submontane zone of Obudu is interesting: most West African Gorgyra are species of lowland forest, only G. minima inhabiting Guinea savannah. The fact that the genitalia are so close to G. bina is surprising, though the large size and the morphological features make the species easily identifiable on morphological grounds. However, we also have a number of cases, especially within the G. diversata-group where several undescribed species show hardly any genitalic distinction. There is obviously no reason to expect that all Gorgyra have genitalia that are decisive in their determination, though fortunately most of them do. The two specimens at hand were caught 15 km from each other so the species is probably widespread at submontane level. So far the species is endemic to Obudu: Michel Libert informs us that he has not come across this species in Cameroun and there is none in the extensive material of Gorgyra in ABRI.

Etymology: We are pleased to dedicate this species to Robert Warren. His first major discovery in Nigeria was a large species of Bebearia, which he readily agreed should be named as B. omo Larsen & Warren (in Larsen, 2005) as a contribution to the conservation of a very important habitat, rather named after himself. He has made several other important contributions to the Nigerian butterfly fauna, not least the first Nigerian record of the hesperiid Celaenorrhinus bettoni Butler, 1902 and the continued presence and validity of Ceratrichia lewisi Collins & Larsen, 2000, both from the Obudu Plateau. He also facilitated Collins' successful visit to Obudu in April 2007, where three of the new species in this paper were collected. We have already dedicated a skipper endemic to Obudu to C.S. Lewis (Ceratrichia lewisi Collins & Larsen, 2000). 94 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Meza gardineri Collins & Larsen, sp. nov.

In June, 2007 Alan Gardiner collected two female skippers at Ikelenge in northwestern Zambia that he could not identify. Good photographs were immediately forwarded to us and they appeared to be of an undescribed species. Closer investigation showed differences from other known Meza of such a nature that we have no hesitation in describing it here, since two virtually identical specimens are available.

Meza gardineri female holotype: Upperside (left) and underside (right)

Holotype: ♀, Zambia, NW Province, Ikelenge (11 o 15'S 24 o 16'E), 13.vi.2007 (A. Gardiner leg., coll. Natural History Museum, London)

Paratype: 1♀, same data but in coll. A. Gardiner.

Diagnosis: Female forewing 17.1/17.5 mm. The upperside is dark brown. The forewing has the typical spotting of the genus. The two cell-spots are smaller than usual, while the spots in spaces 2 and 3 are of the usual size. There are three subapical spots. The usual non-hyaline white spot is present in space 1b. The hindwing has two smallish hyaline discal spots, as in most other species: there are traces of a non-hyaline white spot at the end of the cell. The costa is more prominently whitened (sepia) than usual in the genus. The forewing underside is as the upperside with only slight lighter shading along the costa and the subapical area. The basal and discal areas of the hindwing are a dirty beige that darkens into a broad brown margin. The costa is also dark brown, but its border with the lighter discal area runs sharply along vein 7. The two hyaline spots are clear. The end-cell has a small brown spot and a small circle, but neither has any traces hyaline.

Discussion: Only one similar Meza species is so far known from Zambia (Heath et al., 2002), the much smaller M. larea Neave, 1910, which has a uniformly blackish-brown hindwing underside. Other species lacking a hyaline cell-spot on the hindwing differ as follows: M. mabea Holland, 1894 also has a uniformly dark brown hindwing, June 2008 METAMORPHOSIS, VOL. 19, No. 2 95

though with a chocolate tone; M. leucophaea Holland, 1894 has a prominent white cilia and extensive white marginal shading on the greyish-black hindwing underside; M. elba Evans, 1937 comes slightly closer to the new species but the two hindwing hyaline spots are more prominent and almost twice as long as they are wide. These spots are especially prominent on the brownish underside, where they are faint in the new species: there is no trace of the broad brown margin and the tornal area is whitish. None of the species mentioned is known anywhere close to Zambia. A. Gardiner also has a specimen of M. cybeutes cybeutes Holland, 1892 [not the eastern ssp. pallida Holland, 1892] from Zambia, but this species has a very prominent hyaline spot in the cell of the hindwing and differs in many other respects. This is a most interesting range extension of the latter species that has never been recorded from Zambia. The type locality of M. gardineri is in one of two areas in Zambia with equatorial forest conditions, but like M. elba it might also be found in denser types of adjoining woodlands. The species was flying fast and low in woodland/degraded riparian forest in the company of M. larea: the light undersides were immediately recognizable.

Etymology: We take pleasure in naming this butterfly after Alan Gardiner who has already made several interesting butterfly discoveries and contributed to conservation activities in Zambia, Zimbabwe, and Botswana. He kindly donated the holotype to ABRI. Several species that he found as new to Burkina Faso were first published by Larsen (2005).

Acleros bobiri Collins & Larsen, sp. nov. =Acleros bala Berger. Manuscript name (TL: Ghana, Volta Region)

This new species was recognized by the late L. Berger among the Father Maessen material in the collections at the Musée Royal de l'Afrique Centrale (MRAC), Tervuren under the manuscript name bala, under which it was also cited and figured by Larsen (2005). We do not know the etymology of this name and prefer to give it the name of the Bobiri Butterfly Sanctuary in Ghana.

Holotype: ♂, Ghana, Hohoe, Volta Region (06o60'N 00o46'E), (T. Maessen leg., coll. McGuire Center, Gainesville). The holotype is illustrated as A. bala Berger, ms. in Larsen (2005).

Paratypes: 7 ♂♂, 2 ♀♀, Ghana, Volta Region (T. Maessen leg., coll. McGuire Center, Gainesville); 2 ♂♂, Ghana, Volta Region, Hohoe (T. Maessen leg., coll. Carnegie Museum), 1 ♂, Ghana, Kakum National Park (T.B. Larsen leg. et coll.); 1 ♂, Ghana, Bobiri Butterfly Sanctuary: 1 ♂ 1 ♀, Ghana, Volta Region, (T.B. Larsen leg. et coll.); 1 ♂, Ghana, Suhien (leg. et coll. ABRI). 96 METAMORPHOSIS, VOL. 19, No. 2 June 2008

Top row uppersides: Acleros bobiri male (left), A. nigrapex male (centre) and A. nigrapex female (right). Bottom row undersides: Acleros bobiri male (left), Acleros nigrapex (right). All are from Amedzofe in Ghana (slightly enlarged)

Diagnosis: Forewing 15 mm. The male is similar to A. nigrapex Strand, 1913 though usually slightly larger. The ground-colour is dark brown, with a chocolate tinge, lighter than in the more blackish A. nigrapex Strand, 1913 (and most other Acleros). The forewing usually has a distinct white spot just inside the cell at the base of space 2. An additional, smaller spot in the cell may be present. The extent of white on the hindwing is more extensive, with the white streak on the abdominal fold reaching almost halfway to the base and the main area of white reaching the level of end-cell in spaces 1c and 2. The forewing underside is brown: the two white spots are usually more accentuated. Space 1b is centered with a diffuse, sagittate white spot. There is a whitish marginal scaling, more widely but also more diffusely, than in A. nigrapex, where it is concentrated in spaces 2-4 close to the margin. The hindwing is whiter, and more uniformly so, but retains the dark apical patch that gave A. nigrapex its name, though less prominently. However, most of the brown markings are vestigial, especially the rather solid dark postdiscal patch in space 1b. Viewed from above at least half the abdomen is white, against one-third or less in A. nigrapex – this difference is more apparent than it would seem from this description. When males in good condition are compared side by side, the distinguishing characters are usually clear-cut; with specimens in poor condition it is less easy. The female is also characterized by being slightly larger on average, by having whiter marginal/tornal markings on the hindwing upperside, and by having the hindwing underside much less marked with brown. The forewing pattern of white varies, as it does in all Acleros. The amount of white on the abdomen is also more extensive than in A. nigrapex, though not as much as in the males. The females are less easily told apart than the males.

Male genitalia: As might be expected, the genitalia of A. bobiri are similar to those of A. nigrapex, but the overall structure is more robust. The tegumen is proportionately larger, the uncus is not somewhat upturned, and the small, toothed excrescence at the dorsal tip of the June 2008 METAMORPHOSIS, VOL. 19, No. 2 97

valve is much better developed. These characters have been checked in several specimens.

The male genitalia (left) of Acleros bobiri (Ghana, Bobiri) and of A. nigrapex (Ghana, Kakum) (right)

Discussion: Despite the examination of long series of Acleros in many collections, we have only found a limited number of A. bobiri from Ghana and its Volta Region, thus making it a Ghana endemic, as well as an endemic of the Ghana sub-region of Africa west of the Dahomey Gap (see Larsen (2005) for definition).

Etymology: We give this new skipper the name of the Bobiri Butterfly Sanctuary in Ghana, which has demonstrated the value of butterflies in ecotourism and where a long-term monitoring of butterflies is planned to culminate in the 2100 African Butterfly Survey.

Pteroteinon reali Berger, 1962 stat. nov. Pteroteinon pruna reali Berger, 1962. Bulletin de l'Institut Français d'Afrique Noire (A) 24: 458 (447-463). Ivory Coast: “Côte d'Ivoire: Adiopodoumé”. Erroneously synonymized with P. pruna pruna Evans, 1937 by Larsen, 2005a: 526 [Holotype not in MRAC, Tervuren: it may be in the IFAN (Dakar) but was most likely lost with much of the collection that was formed at the Adiopodoumé Research Station during late colonial times]. Pteroteinon pruna Evans, 1937. A catalogue of the African Hesperiidae indicating the classification and nomenclature adopted in the British Museum: 153 (212 pp.) (TL: Cameroon: “Cameroons (Bitje)”) Caenides na Lindsey and Miller, 1965 in Fox, et al., 1965. Memoirs of the American Entomological Society No. 19: 127 (438 pp.) (TL: Liberia (northwest), Yendamalahoun) (syn. nov.) Leona na (Lindsey and Miller, 1965); Larsen (2005) Butterflies of West Africa (text and colour plate (holotype))

Re-evaluation of Pteroteinon pruna reali as a distinct species and as a senior synonym of Caenides na: This species was described from Adiopodoumé near Abidjan in Côte d'Ivoire as a Côte d'Ivoire subspecies of C. pruna, in contrast to material from Cameroun and Central Africa. During, and following, a visit to ABRI in 1999, Larsen examined all the limited material available of Pteroteinon pruna and 98 METAMORPHOSIS, VOL. 19, No. 2 June 2008

made several genitalia preparations (about fifty specimens examined in ABRI; coll. Larsen; MRAC, Tervuren; and NHM, London). He concluded that no valid subspecies of P. pruna were present in West Africa, but that there was variation in spot size. P. pruna reali was relegated to a junior synonym of nominate P. pruna (Larsen, 2005) but none of the specimens examined was actually true ssp. reali. Caenides na Lindsey & Miller, 1965 was described after a single male from near Nzérékoré in the Guinea/Liberia mountains. The unique type was in such a poor condition that it could not be adequately described (the holotype in frightful condition is figured for the first time by Larsen (2005: plate 121)), but it was clear that hyaline spotting was effectively absent. The genitalia, however, were distinctive and appeared most closely related to species such as Caenides (Leona) leonora (Plötz, 1879); Larsen (2005) therefore listed it as a Leona. A single male was recently collected by Eric Vingerhoedt in the Diecké Forest in Guinea (v.2005), a species-rich lowland forest locality among the northern Guinea/Liberia mountains. The outer half of the forewing had a strong violet-blue sheen as in Pteroteinon pruna Evans, 1937 and we realized that their genitalia were actually very similar. When S. Sáfián reported collecting “Caenides na” also in Bia National Park in western Ghana, while this paper was at proof stage, we realized that P. pruna reali was actually a senior synonym of Caenides na (syn. nov.) as well as being a species distinct from nominate P. pruna (stat. nov.), but with overlapping ranges.

Pteroteinon reali :Male upperside (left) and underside (right), compared with P. pruna (below)

Redescription and distribution of Pteroteinon reali: Forewing 20 mm. The shape of the forewing is narrower than that of P. pruna with the apical part more pointed (standardized for size, the distance between the tornus and the apex of the forewing is 12% longer in P. pruna). The male upperside appears uniformly brown with a strong violet-blue sheen on the distal half of the wing. However, there are the faintest traces of a narrow hyaline spot in space 2 and a tiny spot in 3 (the visibility to the human eye is exaggerated in the photos). Overall the underside is darker with a chestnut tone June 2008 METAMORPHOSIS, VOL. 19, No. 2 99

that is absent from P. pruna. On the forewing underside the narrow spot in 2 and the tiny spot in 3 are more visible than on the upperside. In addition there are two tiny, almost conjoined spots in the cell. A single subapical spot in space 6 is barely visible even under the microscope. Spaces 1a and 1b are darker than in P. pruna and without trace of the large white spot in 1b. The hindwing underside is uniformly dark brown tinged with chestnut and much darker than in P. pruna. There are four faint postdiscal spots in spaces 2 to 6. The male genitalia (SCC 551) are similar, differing only in two respects: i) The distal edge of the valve curves back along the ventral edge, which at this point is moderately spined, the extent of the spines stretching to about the middle of the distal edge. In P. pruna the entire distal edge is more strongly spined: in fact the largest spines are where the distal edge meets the ventral part of the valve and is thence drawn out almost to a point. ii) The two lamellae that form the gnathos are distinctly larger in P. reali. However, the differences in genitalia structure are not great, and doubtless the valves may vary somewhat (those of our specimen are actually slightly asymmetrical). We believe the differences in wing-shape, the size of the spots, the deep chestnut-tinged underside, and the genital characters are sufficient to validate the two species, especially since their distributions overlap. A female from near Abidjan is said by Berger to have the light spotting much less reduced than in the males ['la diminution et la réduction des macules sont nettement moins accusées]. P. reali is currently known from five males and a female from the Guinea/Liberia Mountains (Yendamalahoun (TL of C. na) and Diecké Forest in Guinea), the Nimba area in Côte d'Ivoire (Gopoupleu near Danané (paratype) and Adiopodoumé near Abidjan) as well as the recent record from Bia NP in Ghana; there are several records of P. pruna also from this area. The range of P. pruna is wider, ranging from Freetown in Sierra Leone through West Africa to Nigeria, the Central African Republic, and much of west-central DRC (Tshuapa, Uele, and Mayoumbé): it is not a common butterfly. Both species seem limited to wetter forests in good condition.

The male genitalia of Pteroteinon reali (left) and P. pruna (right) 100 METAMORPHOSIS, VOL. 19, No. 2 June 2008

The genus Platylesches Holland Proceedings of the Zoological Society of London 1896: 72 (2-107). Type-species: Parnara (?) picanini Holland, by original designation

The genus Platylesches was last revised by Evans (1937) and the most recent compilation lists 20 species (Williams, 2005): of these, six were described by Evans in his revision, in addition to a few subspecies. He grouped the genus in eminently practical groups based on morphological characters, at least some of which probably also reflect a genuine phylogeny (a seventh species, P. villa. was subsequently downgraded to subspecies by Evans (1956). This remained the status for nearly 50 years until the description of P. rossii Belcastro, 1986 [mis-spelled rossi by Larsen (2005)], a species now known from Guinea, Sierra Leone (TL), Côte d'Ivoire, and Ghana. Kielland (1978) raised P. langa to species rank and described a new species as P. larseni (eastern Tanzania, Zambia, and Shaba in DRC) (Kielland, 1992). The most recent addition to the genus is P. dolomitica Henning & Henning, 1997 from South Africa. Platylesches robusta Neave, 1910, common in eastern and southern Africa, has two strongly disjunct subspecies in the shape of ssp. villa Evans, 1937 from Cameroun (TL) and the Central African Republic (ABRI), of which only two males are known, and ssp. fofi Larsen & Mei, 1998 known only from three or four specimens. Both probably need upgrading to species rank. During a visit to the Carnegie Museum, Pittsburgh in 2007, Larsen inspected the holotype of P. batangae Holland, 1894, which proved actually to be female, and found the first genuine male of this elusive species, which is re-described below. While browsing for additional material in the ABRI collection, and examining the genitalia of odd-looking males, two new species were found and are described below. However, this also showed that the genus is more complex than it seemed. Larsen has commenced an in-depth revision of the genus in collaboration with ABRI.

Platylesches langa Evans Pamphila ayresii Trimen & Bowker, 1889. South-African Butterflies: a monograph of the extra-tropical species 3 Papilionidae and Hesperiidae: 321 (438 pp.). London (TL: South Africa, Transvaal) Platylesches ayresii langa Evans, 1937. A catalogue of the African Hesperiidae indicating the classification and nomenclature adopted in the British Museum: 169 (212 pp.) (TL: Malawi. “Nyasaland, Mlanje”) Platylesches larseni Kielland, 1992. Metamorphosis 3: 148 (148-153). (TL: Tanzania: “Tanzania, Katuma River, Mpanda, 1600 m”)

Confirmation of a southern species in West Africa: Platylesches ayresii langa Evans, 1937 was first formally raised to species rank by Kielland (1978), with illustrations of the genitalia of the two. P. langa was illustrated in colour together with “P. ayresii” in the book on Tanzanian butterflies (Kielland, 1990). Larsen (1992) pointed out to Kielland that he had raised P. langa in comparison with a species that was not actually P. ayresii presumably an undescribed taxon. Kielland (1992) promptly described the latter as P. larseni Kielland, 1992, with the type from the eastern parts of Lake Tanganyika the late L. Berger at the Musée Royal de l'Afrique Centrale (MRAC), Tervuren had actually isolated the same species from Shaba (Katanga), in the DRC on the western side of the lake under the manuscript name straeleni, but never published it. June 2008 METAMORPHOSIS, VOL. 19, No. 2 101

In the Natural History Museum, London is a Nigerian male of this group, more than a thousand kilometres from the closest records from Shaba and Tanzania. Larsen (2005) included it as P. langa, stating that they [erroneously referring to two females] “looked authentic and are definitely [not] P. ayresii or P. larseni” and that males would be necessary to be sure. A single male is now available from even further away, from Likpe Mate in Ghana's Volta Region (illustrated here). Its white spots are more developed than usual, but the genitalia viewed are identical with material from Zimbabwe, southern DRC and eastern Tanzania, where it is sympatric with P. larseni. It may deserve subspecies status, but in this difficult genus we do not wish to act on a single specimen, the Nigerian male (Zungeru, xi.1910, J.W. McAfie leg.) being intermediate between the Ghana and Zimbabwe males. It is one of a small group of species with an interesting biogeographical pattern, occurring in southern Africa and then again in West Africa, being absent from northern Tanzania, Kenya, the northern DRC, and most of the equatorial forest zone (e.g. Caprona adelica Karsch, 1892 and Astictopterus abjectus (Snellen, 1872)).

Platylesches batangae (Holland) Parnara batangae (Holland, 1894). Entomological News 5: 92. (TL: German West Africa [Cameroun], Batanga).The unique type is figured in monochrome. Platylesches batangae Holland, 1896. Proceedings of the Zoological Society of London 1896: 72 (2-107) Platylesches batangae Holland, 1894 [in error]. Sierra Leone, Upper Kasai [DRC] (Evans, 1937) Platylesches batangae Holland, 1894 [in error]. Basse Casamance, Senegal (Berger, 1968) Platylesches batangae Holland, 1894 [in error]. Gambia (Gillies 1982 [1985]) Platylesches b atangae Holland, 1894 [in error]. Sierra Leone (Belcastro, 1986) Platylesches batangae Holland, 1894 [in error]. Zambia (Heath et al., 2002) Platylesches batangae Holland, 1894 [in error]. Zambia (Williams, 2005) Platylesches batangae Holland, 1894 [in error]. West Africa (Larsen, 2005)

This species was described and illustrated from a single 'male' in the Carnegie Museum, Pittsburgh collected at Batanga, Cameroun. It has been misunderstood ever since, mainly due to its treatment by Evans (1937). In October, 2007 the type was examined by Larsen and found actually to be female. However, a male from Efulen, Cameroun matched the female extremely well and is here used as the basis for a redescription of the species: its genitalia cannot be the same as those figured in the rough sketches that Evans made, nor do they match those glued under the specimen examined by him (which are difficult to examine since they are embedded in congealed glue that hides most of the features except the clearly different valves). No matching specimens were found in any other collection inspected and both sexes are quite different from, and much larger than, the series in the Natural History Museum, London, which Evans considered to be P. batangae. The two localities are in southwestern Cameroun, situated some 70 km apart (Efulen to the north). Description of the unknown male: The male forewing ( 15.5 mm) is longer than that of P. chamaeleon (14.0 mm in West Africa). The upperside ground-colour is a lighter brown than the latter, lacking its somewhat silky sheen: this is a qualitative 102 METAMORPHOSIS, VOL. 19, No. 2 June 2008

difference that cannot be due to fading. Both sexes have a single, well-defined subapical spot and the forewing markings are the same as in other members with a single such spot; however, the spot in 2 is wider than usual. The hindwing has a line of four discal spots, the two closest to the abdominal fold being strongly overlaid with ochreous-brown scales. The lighter brown colour is even more evident on the underside, which also lacks any trace of the usual purplish sheen of P. chamaeleon and P. affinissima. The hindwing has a clear white streak along the inner edge of the abdominal fold and two somewhat diffuse whitish spots are present on the hindwing, the two that are obscured by brown scaling missing. The palps and the underside of the thorax are densely clad in off-white hairs: they are usually distinctly tinged with sulphur-yellow in P. chamaeleon, but are apparently off-white also in P. affinissima. The female holotype is very similar to the male, but larger: there are two additional small, faint white spots in the cell of the hindwing underside. The fact that there is no trace of more than one subapical spot even in the female is a significant factor linking the two specimens. However, Holland (1920) is adamant that he saw a specimen of P. batangae from Basoko in the DRC, of which he says that “I have carefully compared [it] with the type and find it to be identical”. This may well be true but the specimen has not been viewed; it is from the Lang-Chapin Expedition to Congo and all the Lepidoptera were sent to Holland in 1903 to enable him to write the 1920 paper. The locality is Basoko (01o 20'N 23 o 35'E) well to the north of other DRC material of P. “batangae” and very far east of Cameroun, and on the other hand Holland writes that it is “dwarfed and also somewhat damaged”. Most likely it is actually P. chamaeleon (Mabille, 1891) that is known from this part of the DRC and was not in the Carnegie Museum at the time.

Male genitalia: The male genitalia were found to be slightly damaged, with parts missing from one valve; it appears as if a crude attempt has been made to view them in situ, though it could also be pest damage, which was observed in one other contemporaneous hesperiid from the Museum. Fortunately the two valves complement each other so that the complete configuration can be reconstructed. The uncus and tegumen are of the usual shape in the group. The uncus ends rather narrowly and is slightly bifid. The gnathos is rather small and upturned, much as in P. chamaeleon, but the two elements are fused (see illustrations of the latter in Larsen (2005) and Miller (1965)). The valve is long with practically parallel ventral and dorsal edges and ends by being rather bluntly rounded in an even manner. The dorsal half of the rounded edge is finely serrated. The dorsal edge of the valve has a small ridge with such serration before the middle of the valve. Even by the standards of the genus the slender penis is very long (1.77 times the length of the valve, proportionally longer than any other species studied). The valves are rather similar to those of P. picanini Holland, 1894 and P. iva Evans, 1937, but lack the vertical dorsal thorn characteristic of the valve of these species (see Larsen, 2005). The uncus is short, being only 0.39 of the length of the combined uncus and tegumen.

Misrepresentations of P. batangae: As already mentioned, Evans (1937) certainly misunderstood this species. In addition to the points already made, the basis for his statement that the wings were more rounded than in other species was that the draughtsman of the drawing in the June 2008 METAMORPHOSIS, VOL. 19, No. 2 103

description had not corrected for the fact that the wings were drooping significantly. Evans lists a male and female from Sierra Leone and four males from Upper Kasai in the DRC. Berger (1968) records P. batangae from the Basse Casamance in Senegal, which Larsen (2005) [wrongly] suggested might be P. rossii Belcastro, 1986. Gillies (1982) records a few from the Gambia, now on the Collection of M. Newport, who kindly sent photographs of the specimens in question. Belcastro (1986) recorded it from Sierra Leone, in which he was followed by Larsen (2005). Heath et al. (2002) recorded and figured it from Zambia. Examination of the material in the Musée Royal de l'Afrique Centrale (MRAC), Tervuren showed a long series from the southern parts of the DRC and a single male from Côte d'Ivoire idenfied by L. Berger as P. batangae. These all resemble each other in most characters but are quite distinct from true P. batangae. They seem to pertain to three distinct species: 1) the Zambian population, usually with two or three subapical spots, has very distinctive genitalia and is described as P. heathi in this paper; 2) the West African population, usually with two or three subapical spots, has genitalia that are closest to those of P. moritili Wallengren, 1857, but quite different from the DRC population (illustrated and inconclusively discussed by Larsen (2005)); 3) the population from the DRC consists of small specimens, usually with just a single subapical spot and with sulphur-yellow palpi, but with genitalia that are closest to those of P. moritili, though clearly different from the undescribed West African taxon. They all share a dark brown hindwing underside with some sheen and the presence of five or six small, irregularly placed, discal spots but lacking the diffuse white band of the common P. moritili. These two additional species will be described in Larsen's proposed revision of the genus that was prompted by the study of P. batangae.

Top row: Platylesches batangae male from Efulen, The female Holotype of Cameroun in the Carnegie Museum, upperside (left) Platylesches batangae. and underside (right). Bottom row: P. chamaeleon from Harbel, Liberia (its genitalia are figured in Fox et al., 1965). Upperside (left) and underside (right). 104 METAMORPHOSIS, VOL. 19, No. 2 June 2008

The male genitalia of the Carnegie Museum male of P. batangae

Platylesches heathi Collins & Larsen, sp. nov.

While researching the status of P. batangae we studied the photos in the book on Zambian butterflies (Heath et al., 2002), also shown on the Williams cd-rom (2005), purporting to be P. batangae. It has three subapical spots, thus precluding a correct identification. This specimen was included in the Heath collection donated to ABRI and formed part of a series (the specimen has been reversed in the reproduction process and has since lost an antenna). A specimen with matching data proved to have genitalia that were not only very different from those of P. batangae but also from all known Platylesches. It is here described as a new species with the male in the published photo designated as the holotype.

Holotype: ♂, Zambia, Mundwiji Plain, 40 km E of Mwinilunga (11o 45'S 24 o 26'E), 27.x.1979 (A. Heath leg., coll. ABRI)

Paratypes: 5 ♂♂ 1 ♀, data as holotype; 1 ♂ 1 ♀, Zambia, Ikelenge, Hillwood, 21.xi.1979 (A. Heath leg., coll. ABRI)

Platylesches heathi: Male holotype upperside (left) and underside (right) June 2008 METAMORPHOSIS, VOL. 19, No. 2 105

Diagnosis: Male forewing 15 mm. The ground-colour is dark brown, but not as dark and glossy as species of the P. chamaeleon/affinissima group with a single subapical spot. There are usually two or three subapical spots on the forewing upperside but the upper spots are often only faintly visible and in a few cases wholly missing (in such cases they can be seen on the underside). The usual complement of spots is present, including one in space 1b (cream-coloured). The upper cell-spot is usually very small. The hindwing has three or four clear cream postdiscal spots. The cilia is greyer than the ground-colour. The disk of the forewing underside is blackish. The spotting is as on the upperside except that the spot in 1b is surrounded by much white scaling (more than 2 mm). The costa, apex, and margin are chestnut-brown. The usual narrow basal streak along the costa is beige. The hindwing is mainly silky chestnut- brown but the colour is uneven. There is a white spot in 1c followed by an irregular row of five cream spots, those nearer the costa being diffuse. The central discal and the apical areas are slightly lighter than the rest of the wing and the tornal triangle darker. The cilia of the tornal angle is not as white as in P. moritili. The female is almost identical to the male.

The male genitalia of Platylesches heathi with its very large uncus twisted

Male genitalia: The male genitalia (SCC 557) are of the usual type for the genus but match no described species. The most prominent characteristic is the massive uncus/tegumen structure, larger than in any other species so far dissected by us, and as long as the narrow valve. The uncus is also very wide, but of the same basic shape as P. chamaeleon, ending in two small points that are widely separated by a small rounded depression: in P. chamaeleon the tegumen is very small. The main part of the uncus structure folds smoothly underneath the dorsal surfaces as in many other species in the genus. The gnathos has two slender branches that are not fused, but which support membranous material (drawn before the uncus was twisted). The valve is rather narrow. Towards its tip is a large dorsal thorn, after which a strongly serrated edge continues until the very tip, where it meets the ventral edge which is not 106 METAMORPHOSIS, VOL. 19, No. 2 June 2008

serrated. The penis is 1.5 times longer than the valve.

Discussion: The species has two or three subapical spots and cannot be confused with the species that have one spot, such as P. chamaeleon, P. affinissima, and P. panga, which otherwise look rather similar. While it is easy, especially for males, to lose subapical spots (even losing all three), we do not recollect having seen an undoubted male of the group with one spot having an additional spot. Similarly upper cell-spots may occur where there is usually only a lower spot, and the upper spot may disappear in two-spotted forms. The key in Evans (1937) would lead this species to be determined as P. moritili (Wallengren, 1857). It is actually rather close to that species, which rarely has any spotting as such on the hindwing underside – and if there is, then any spots are more or less obliterated by whitish scaling forming a diffuse greyish-white band. Confusion with P. shona Evans, 1937 is ruled out by the presence of a well- developed spot in space 1b of the forewing upperside.

Etymology: We are pleased to name this species after Alan Heath, who has contributed so much to the knowledge of African butterflies. His revision of the classification of the Aphnaeini, his work on myrmecophily, and his leading role in compiling the book on butterflies of Zambia are only highlights of a long research career. Given the confusion that has surrounded P. batangae, it is hardly surprising that he was wrong about what is now P. heathi!

Platylesches hassani Collins & Larsen, sp. nov.

Among many other Platylesches collected at Hillwood, Ikelenge in Zambia during an ABRI expedition (Colin Congdon, Ivan Bampton, Martin Hassan, and Peter Namakana) that was specially asked to obtain as many members of the genus as possible and to look for their early stages, were various unusual specimens. The team collected a number of the scarce P. tina Evans, 1937 of typical appearance, with limited forewing spotting, as well as five somewhat larger males with almost the usual complement of spots for the genus. These were examined and proved to belong to a new species that is described below.

Platylesches hassani : Male holotype upperside (left) and underside (right) (slightly enlarged) June 2008 METAMORPHOSIS, VOL. 19, No. 2 107

Holotype: ♂, Zambia, Ikelenge, Hillwood (11 o 15'S 24 o 16'E), iii/iv.1999 (ABRI team leg., coll. ABRI)

Paratypes: 4 ♂♂, same data as holotype

Diagnosis: Male forewing 12 mm, somewhat larger than in P. tina. The ground-colour is blackish without any sheen. The spotting is white and well marked, with two or three subapical spots (there are usually at least traces of the third spot on the underside when missing on the upperside). The two cell-spots are not large but very clear. Below them is a large spot in space 2, slightly wider than it is high. There are smaller but well-developed white spots in spaces 3 and 4. The usual spot in space 1b is absent. The hindwing has a row of diffuse light ochreous discal spots. The underside colour is chestnut. The forewing has the same spotting as above but in addition there is a somewhat diffuse white area in space 1b. The margin and apex have some violet scaling but not very pronounced. The hindwing has a rather prominent white spot in space 1b and the faintest of traces can be seen of the upperside postdiscal spots. The hindwing has hardly any purple overlay, which is prominent in P. tina. The female should be similar. The subapical spots, the one in 2, the hindwings spots, and the white area in 1b of the forewing underside are missing in P. tina, while the violet overlay is much stronger. The two cannot be confused.

The male genitalia of Platylesches hassani with the large ventral thorn on the valve

Male genitalia: The tegumen (paratype genitalia SCC 558) is considerably longer than the uncus and is also relatively tall. The uncus tapers to a single blunt point. The gnathos has two L- shaped branches that are lightly fused distally. The dorsal edge of the valve is straight for 3/4 of its length and then turns up at a 30o angle. At its apex, it turns at a right angle to form a lightly serrated distal end to the valve: where they meet, the ventral edge of the valve continues, forming a strong, lightly serrated tooth. The dorsal edge of the valve has two small depressions where the chitinized edge weakens. The ventral 108 METAMORPHOSIS, VOL. 19, No. 2 June 2008

edge of the valve is also nearly straight, running completely parallel with the dorsal till it starts turning up towards the distal end. The penis is only 1.2 times the length of the valve. The entire structure is similar to that of P. tina, but clearly different in details: the gnathos is less developed, the dorsal part of the valve has only one small depression, the ventral edge of the valve is concave in its basal half, and the distal half of the valve is more extended and ending in a much broader tooth.

Discussion: The type locality, Hillwood Farm, is extremely rich in Platylesches. In all, the following twelve species are known: galesa, langa, larseni, robustus, heathi sp. nov., moritili, tina, hassani sp. nov., picanini, affinissima, chamaeleon, and panga (two new to science in this paper, with larseni and panga being new country records for Zambia). The species is presumably rare since no others are present in the ABRI collection and none has been encountered elsewhere. It is, however, quite likely that other material is in collections, placed as “unusual” specimens of P. tina (in fact one was found in the MRAC collection, Tervuren from southeastern DRC under the manuscript name nina Berger while the manuscript was in press).

Etymology: We are pleased to dedicate this new species to Martin Hassan who has worked with ABRI on butterfly collecting and breeding for many years. The breeding of species of Platylesches will in the next few years help to illuminate additional complexities within the genus.

Acknowledgements: Ugo Dall'Asta and Frans Desmet of the Musée Royal de l'Afrique Centrale (MRAC), Tervuren kindly sent us a good photo of Gorgyra dubia (Berger ms) to assist in the description of one species. Keith Bland, of the National Museums of Scotland facilitated Larsen's 2007 visit to study the M. B. Usher collection from Ghana. Apart from his assistance at the Museum, he and his wife Val put up and put up with Larsen during his visit, resulting in many stimulating discussions outside of 'office hours'. We were joined by Cedric Holmes whose recent monograph on the Bebearia cocalia-complex is a fine example of scholarship. Jeff Boyd of the Central Africa Liaison, Presbyterian Church USA in Yaoundé took the trouble to send us the precise details on Mrs and Dr Lippert and the exact location of Métet, for which we are grateful. Robert Ducarme kindly allowed us to describe the magnificent Cooksonia ginettae, donating 3 females to ABRI since procured by local assistants, yet one of his amazing finds in the rich and ecologically diverse Kivu Province in the DRC. Alan Gardiner entrusted us with the description of a new species of Meza from Zambia: we are pleased to name it after him in honour of his general contributions to the fauna of the Zambezian area. We are grateful to Eddie John who screened a late version of the manuscript and did much to improve the style, consistency, and language of the paper. Michel Libert, as usual, kindly provided useful comments on a number of specific issues. Jacqueline and Lee Miller helped us with access to the Maessen Collection from Ghana, which has been transferred from Sarasota to June 2008 METAMORPHOSIS, VOL. 19, No. 2 109

Gainesville and remains one of the most important sources for West African butterflies. Sadly, Lee died while this manuscript was being processed. We are grateful to Claude Pierre-Baltus for sending a scan of the description of Acraea eugenia ochreata from a paper that is almost impossible to get hold of. We thank John Rawlins and his colleagues for access and permission to publish images and data from the important African holdings at the Carnegie Museum, Pittsburgh, from which further new taxa and interesting information remain to be mined at a later stage. The Holland material from Cameroun and Gabon and the Liberian holdings from the Fox collection remain two very important sources for African butterflies. Szabolcs Sáfián has collected butterflies in Ghana over the past five years and found many interesting species, including several new to the Ghana list. Together with Kwaku Aduse-Poku he took the initiative in starting Ghana Butterfly Conservation (www.ghanabutterflies.uw.hu). He kindly donated the unique holotype of Iridana agneshorvathae to ABRI. His hope is to undertake a doctorate on the African Lipteninae. Andrei Sourakov guided Larsen at the McGuire Center at Gainesville (University of Florida) and hosted him during his 2007 visit, also kindly following up on some issues. Eric Vingerhoedt kindly sent some of his smaller butterflies from the Guinea/Liberia border to ABRI and amongst these was the second specimen ever of what was named Caenides na, finally pinning down this elusive skipper after more than 40 years. P.R.P. Ward deserves thanks for procuring the excellent series of Colias marnoana, not least since he reported back that he had to give up collecting that day because of extreme heat exhaustion! Robert Warren kindly facilitated Collins' visit to the Obudu Plateau in 2007: he personally collected interesting material in that locality and we are pleased to thank him by naming a new Obudu species after him. His donations of material to the ABRI collection are also much appreciated. Mark Williams' constantly updated Catalogue of the Afrotropical Butterflies proved itself to be a most effective tool that saved many hours of work during the research process. Many people have collected for ABRI over the years, including Ivan Bampton, Colin Congdon, Martin Hassan, Malang Mane, Peter Namakana, Richard Vorgas, and Simon Yevu. Their prodigious efforts, often under difficult conditions in remote areas, have added much of the material now incorporated in the ABRI Collection. Larsen would like to thank the Carlsberg Foundation in Denmark for their continued financial support for his travel programme, which allowed the visit to ABRI as well as those to the McGuire Center and the Carnegie Museum in the USA. Without their generosity during the past thirty years the book on Butterflies of West Africa would have been much poorer than it is … possibly not even yet published. The cooperation, including permission to figure some almost unknown species, of the Carnegie Museum in Pittsburgh and the McGuire Center for Lepidoptera and Biodiversity in Gainesville contributed significantly to the paper. The Natural History Museum, London is part of the foundation for any serious work on butterfly taxonomy and the assistance and advice of Blanca Huertas, Jeff Martin, Campbell Smith, and the Library Staff are much appreciated. Finally, the editor of Metamorphosis and an anonymous referee improved the final version of the paper. 110 METAMORPHOSIS, VOL. 19, No. 2 June 2008

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Sponsor Members of LEPSOC

The following members, apart from their significant contributions to the Society as individuals, have also chosen to be Sponsor Members for 2008 and have through their generosity provided significant financial support which is much appreciated:

Dr. Jonathan Ball Dr. Bennie Coetzer Steve Collins Alf Curle Martin Curle Jeremy Dobson Dr. Dave Edge Owen Garvie Tim Gilbert Graham Henning Dr. Doug Kroon Dave McDermott Duncan McFadyen cc (E. Oppenheimer & Son) Andrew Morton Ian Richardson Harald Selb Hermann Staude Reinier Terblanche Prof. Mark Williams

Any member can volunteer to become a Sponsor Member on an annual basis and make a contribution of R600. As the Society does need all the financial support it can get it is hoped that more members will select to become Sponsor Members in the future. Donations to the Society will also be most welcome.

AFRICANABRI BUTTERFLY RESEARCH INSTITUTE This issue of Metamorphosis is proudly sponsored by ABRI June 2008 METAMORPHOSIS, VOL. 19, No. 2 115 116 METAMORPHOSIS, VOL. 19, No. 2 June 2008 June 2008 METAMORPHOSIS, VOL. 19, No. 2 117 118 METAMORPHOSIS, VOL. 19, No. 2 June 2008