A new species of Pseudaletis Druce, 1888 from northern Angola (Lepidoptera: Lycaenidae: Aphnaeinae).

ZOOBANK: https://zoobank.org/urn:lsid:zoobank.org:pub:C48E2654-0C83-4F19-818B-832423474EF5 ANNALES MUSEI HISTORICO-NATURALIS HUNGARICI Volume 117 Budapest, 2025 pp. 205–216 DOI: https://doi.org/10.53019/AnnlsMusHistNatHung.2025.117.205 HU-ISSN 0521-4726 (print) ISSN 2786-1368 (online) published: 2025. 12. 10. arrived: 2025. 11. 28. A new species of Pseudaletis Druce, 1888 from northern Angola (Lepidoptera: Lycaenidae: Aphnaeinae) Szabolcs Sáfián Hungarian National Museum Public Collection Centre, Budapest – Hungarian Natural History Museum, Department of Zoology, H-1088 Budapest, Baross utca 13, Hungary. Hungarian Natural Heritage Trust, H-9945 Kercaszomor, Fő út 57, Hungary. E-mail: [email protected]; https://orcid.org/0000-0002-0614-4203 Abstract – A new aphnaeine species Pseudaletis (Lepidoptera: Lycaenidae: Aphnaeinae) found in northern Angola is described. The type specimens were collected in a unique mixture of miombo and secondary forest outlier south of the Congo Basin situated on a plateau north of the Kwanza River valley, isolated from other rainforest areas. The new species belongs to the P. clymenus species-group and may be endemic to Angola as recent studies provide evidence of the uniqueness of the forests of northern Angola. With 17 figures. Key words – Afrotropics, ant-association, canopy, miombo, Nobel laureate, Pseudaletis krasznahorkaii sp. nov., species-groups INTRODUCTION The aphnaeine genus Pseudaletis Druce, 1888 is purely Afrotropical, distributed across the Guineo-Congolian Forest Zone. Adults are usually very difficult to come by as both sexes tend to stay in the high canopy or on the subcanopy of dense rainforest, with the exception of a few species, where males were observed displaying near Crematogaster-infested ant-trees (Larsen 2005). They were known to be ant-associated, and single evidence was documented in central Cameroon where pupae were found inside a Crematogaster ant nest and a single male Pseudaletis ‘zebra’ successfully hatched from the nest kept in vivarium (Bouyer 2013). A comprehensive revision of the genus was published by Libert (2007) and since this work only a few further descriptions or revisional notes have seen the light (Bouyer 2013, 2014a, 2014b), not always in agreement with the revision. Sz. Sáfián 206 Annls Mus. hist.-nat. hung. 117, 2025 As part of the country-wide butterfly survey the author visited a peculiar location in northern Malanje Province in November 2024, March 2025 and October 2025, where rare and endemic forest birds are known to occur (Sinclair et al. 2007). The area is a mixed miombo (Brachystegia-Julbernardia) woodland and patches of closed canopy forest with no particularly high trees, probably as a result of continuous human disturbance, and with very thick shrub layer and a dense network of creepers. In the site Crematogaster ants were swarming on the creepers of the forest interior and quite a few species of ant-associated liptenine lycaenids (Lycaenidae, Poritiinae: Liptenini) were also present. Among many interesting forest and miombo butterflies and a few certainly undescribed taxa collected in a dense secondary forest, there were one male and three females of an undescribed Pseudaletis, clearly belonging to the P. clymenus species-group (sensu Libert 2007). The aim of the present paper is to describe and to name this new species, and to place it amongst the relatives. The specimens were captured using conventional butterfly net with expendable handler (Bioquip Tropics net). Forewing length was measured in a straight line on costa between the base and tip of the apex. Etymology for wing pattern and venation follows Libert (2007). Genitalia were dissected in CEPUJ using KOH solution to dissolve soft abdominal tissue. For examination and photography, Nikon SMZ25 stereomicroscope was used with Nikon DS-Fi1 digital camera adapter and NIS Elements imaging software. Digital images of adult butterflies in vivo and in vitro were taken using Canon 7D Mark II DSLR camera and Canon EF 100 IS macro photo lens. Colour plates and the distribution map were edited in various versions of Adobe Photoshop and Adobe InDesign. Abbreviations – ABRI = African Butterfly Research Institute (Nairobi, Kenya); AJG = Alan John Gardiner’s research collection (Hoedspruit, South Africa); CAR = Central African Republic; CEPUJ = Nature Education Centre of the Jagiellonian University (Kraków, Poland); DRC = Democratic Republic of the Congo; HNHM = Hungarian National Museum Public Collection Centre – Hungarian Natural History Museum (Budapest, Hungary). TAXONOMY Ordo LEPIDOPTERA Linnaeus, 1758 Superfamily PAPILIONOIDEA Latreille, 1802 Family LYCAENIDAE Leach, 1815 Subfamily APHNAEINAE Distant, 1884 Tribe Cigaritini Grishin, 2023 A new Pseudaletis from Angola 207 Annls Mus. hist.-nat. hung. 117, 2025 Genus Pseudaletis Druce, 1888 (type species: Pseudaletis agrippina Druce, 1888) In his monograph Libert (2007) revised the genus and established the P. clymenus species-group with two subgroups: P. clymenus and P. busoga. The latter one is monotypic, and known only from females, but the former one has similar looking black and white patterned males with a series of black costal bars (“barres costales” sensu Libert 2007). Since the revision, a further species in the group was described and P. rileyi was synonymised with P. lusambo (Bouyer 2014a). P. subangulata was also elevated to species rank from a subspecies of P. zebra (Bouyer 2013). The updated checklist of the P. clymenus species-groups with known country records (“Distribution”) is presented here; species are listed in alphabetical order. Pseudaletis clymenus subgroup Pseudaletis clymenus (Druce, 1885): CAR, Cameroon, DRC, Nigeria Pseudaletis ducarmei Libert, 2007: DRC Pseudaletis ginettae Bouyer, 2014: DRC Pseudaletis krasznahorkaii Sáfián, sp. nov.: Angola (Figs 1–6) Pseudaletis lusambo Stempffer, 1961 (= P. rileyi Libert, 2007): CAR, DRC Pseudaletis subangulata Talbot, 1935: Ghana, Guinea, Ivory Coast, Liberia, Sierra Leone Pseudaletis taeniata Libert, 2007: Cameroon, DRC, Nigeria Pseudaletis zebra Holland, 1891: Cameroon, CAR, DRC, Gabon, Nigeria Note. The taxonomic status of Pseudaletis cf. lusambo sensu Libert, 2007 is unclear. Pseudaletis busoga subgroup Pseudaletis busoga van Someren, 1939: Cameroon, Uganda, Tanzania Pseudaletis krasznahorkaii sp. nov. (Figs 1–15) Type material – Holotype: male, set dorsally, forewing costa length: 19 mm, in good condition, ANGOLA: Malanje Province, Calandula Municipality, Forest-Miombo Mosaic along the road to Kinjila village. 07–08.10.2025. Leg.: Szabolcs Sáfián; deposited in HNHM (Figs 1–2). Paratypes (n = 3), all female, with holotype data, deposited in HNHM (paratype no. 1; Figs 3–4), ABRI (paratype no. 2; Figs 5–6), AJG (paratype no. 3). Sz. Sáfián 208 Annls Mus. hist.-nat. hung. 117, 2025 Figures 1–6. Type material of Pseudaletis krasznahorkaii sp. n. in dorsal and ventral views: 1–2 = HNHM holotype; 3–4 = HNHM paratype female no. 1; 5–6 = ABRI paratype female no. 2; scale bar: 10 mm. Diagnosis – Males in the P. clymenus species-group are very similar and identification can be difficult. In this case, based on a single male, species diagnosis may not be appropriate, but the straight inner edge of the broad black margin between inner margin and vein 2 (meeting the inner margin in a right angle), seems to be unique to the holotype of P. krasznarhorkaii sp. n. as all other species’ males in the P. clymenus species-group have their inner edge of the black margin either angled or curved at the tornus or appears with a slight protrusion or diffusion of black into the whitish central area of the wing. Female differs from any other species in the P. clymenus subgroup (P. clymenus species-group) (sensu Libert 2007) by the very narrow black outer margin on the forewing upperside, which is broad to very broad in species P. ginettae, P. lusambo, P. subangulata, and P. zebra, while the black colour covers approximately half the forewing surface in P. clymenus and P. taeniata and much over half of it in P. ducarmei. Description – Male. Wings: Forewing length: 19 mm (n = 1); upperside ground colour creamy white, on forewing with a broad black costa and margin and three parallel bars (“barres costales” = BC 1–3 sensu Libert 2007) transversally that intruding the cell, the basal being the shortest and the postdiscal one with curved edge, almost fused with the broad black apical area that encompassing bars BC 4 and 5, inner edge of very broad black margin straight in spaces 1a and 1b, not turning inwards along inner margin, reaching the inner margin at a right 1 2 3 4 5 6 A new Pseudaletis from Angola 209 Annls Mus. hist.-nat. hung. 117, 2025 angle. Hindwing creamy white with only a black marginal line and tornal spot, and with two black tails (tornal one longer). Underside ground colour creamy white, on forewing, broad black apical area and margin zebra-striped with three fine creamy lines, the three bars (BC 1–3) being completely separate, connected only with fine black costal line. On hindwing underside slight asymmetry appearing: one of the irregular tornal patches appearing golden/ochreous, other one mixed black and golden ochreous. Black lines typical for genus present, all starting from tornal patch, that along vein 1 (“trait abdominal” sensu Libert 2007) prominent tornally ochre (in one third length), and basally black and prominent (in two thirds length), the other three lines being vestigial. Transverse line between tornus and costa (“bande médiane” sensu Libert 2007) black, continuous between vein 2 and costa. Submarginal line (“ligne marginale” sensu Libert 2007) black, diffuse, those along inner margin fine, black. At the base of these lines, in subtornus, silver scaling present. Two silver-edged black dots at tornus present (Figs 1–2). Body. Head black, antennae black dorsally, with tiny white dots ventrally, slightly shorter than half of the length of costa. Thorax black dorsally, sparsely covered with golden/ochreous hairs. Legs black with some ochreous mottle. Abdomen black dorsally, with greyish-white rings on each segment. Ventrally golden/ochreous, with the rings on each segment also visible. Tip of abdomen thickly covered with short golden/brown hairs (Fig. 7). Genitalia. Uncus deeply bi-furcate, spine-like, fully fused with shorter, laterally prickle-shaped subuncus, latter with a small, triangular sting-like tip. Gnathos simple, almost rectangular. Saccus back-curving, showel-shaped. Shape of valva rather triangular, with angled dorsal edge. Basal adjacent longer, with an almost right angle and shorter opposite. Ventral edge (hypotenuse) incurving, top tapering down to a narrow, but not particularly acute incurving tip. Fultura inferior open dorsally, broadly and deeply incised with evenly curving bottom of incision. Aedeagus slightly shorter than 2 mm. Basally broader, membranous, in lateral view both edges upcurving, ventral edge only slightly, curving down terminally, ending in laterally thumb-like, in dorsal view more acute tip. Dorsal edge more deeply curved basally, with a slight serration of three teeth before turning down at a flat angle towards tip (Figs 9–12). Female. Wings. Forewing length: 23–24.5 mm (n = 2). Upperside ground colour creamy yellow, with fine black costa and margin, slightly broader black apical area. Black bars barely visible, sub-basal one appearing as black shading, cell-closing one appearing as an elongate black spot, while post-median one appearing as two faint blackish lines between costa and vein 2. Base with dark, orangish-yellow patch. Hindwing clear, creamy yellow, with only a faint black transverse line (shade from underside), and a fine black marginal line. Tornus outer half black, inner half dark, orangish-yellow. Tails black (Figs 3–6). Body. Head black, antennae black, slightly longer than one-third of the length of costa. Thorax black dorsally, covered with golden/ochreous hairs. Legs black with some ochreous mottle. Abdomen black dorsally, with creamy white rings Sz. Sáfián 210 Annls Mus. hist.-nat. hung. 117, 2025 on each segment. Ventrally golden/ochreous. Spongy looking tip of abdomen thickly covered with short golden/brown hairs (Fig. 8). Genitalia. Length of papillae anales dorsoventrally slightly longer than 1 mm. Posterior edge gently curved, dorsally broader, sparely scattered with fine hairs. Apophyses posterior fine, unevenly curved, short (<1 mm). Lamella antevaginalis posteriorly bilobed, with equally sized, evenly rounded lobes. Inconspicuous apophyses anteriores straight, shorter than posterior ones. Ductus bursae almost entirely membranous, longitudinally finely grooved, gently sclerotised only near lamella antevaginalis. Broadening significantly at entrance of bursa copulatrix. Bursa oval, only approximately three times broader than ductus. Very finely grooved longitudinally, signum gently sclerotised, appearing as a dark line, slightly shorter than 1 mm. Anal segment very broad, over five times broader than dorsoventral length of papillae anales (Figs 13–15). Etymology. – Noun in apposition, in honour to novelist László Krasznahorkai Nobel laureate in literature in 2025, who in his newest book “A magyar nemzet biztonsága. Vadászat pillangóra” (Security of the Hungarian Nation. Hunting Butterflies) highlights a lepidopterist researcher. He, with the other protagonist, carries the author’s name, search for the origin of life. Krasznahorkai’s life has multiple parallel lines with those of the author. They did not only escape from the same hometown, Gyula in Eastern Hungary, but they also participated a journey, a pathway of endless learning and wish to return their vision/knowledge to the human world they never felt to be fit in. Figures 7–8. Pseudaletis krasznahorkaii sp. n. abdominal tips in lateral view: 7 = holotype male; 8 = ABRI paratype female nr. 3; scale bar: 1 mm. 7 8 A new Pseudaletis from Angola 211 Annls Mus. hist.-nat. hung. 117, 2025 Figures 9–12. Pseudaletis krasznahorkaii sp. n. holotype male genitalia: 9 = lateral view, 10 = aedeagus, 11 = posterior view, 12 = dorsal view; scale bar: 1 mm. Figures 13–15. Pseudaletis krasznahorkaii sp. n. female genitalia: 13 = lateral view with last segment of abdomen, 14 = lamella antevaginalis and bursa copulatrix (lateral view), 15 = lamella antevaginalis and bursa copulatrix (dorsal view); scale bar: 1 mm. 9 10 11 12 13 14 15 Sz. Sáfián 212 Annls Mus. hist.-nat. hung. 117, 2025 DISCUSSION Ant-association The immediate surrounding of the location of the capture is a thick secondary forest with the trees and understorey thickly tangled with creepers. Most of the larger trees hosted Crematogaster ants and several ant-associated liptenine lycaenids (Lycaenidae, Poritiinae) (as extensively discussed in Larsen 2005) were also captured (e.g. Epitola urania, Eresina sp., Liptena undularis, Micropentila sp., Stempfferia sp.) in the same small area. Figure 16. Female Pseudaletis krasznahorkaii sp. n. in resting position. After finding another specimen in the same situation, it became apparent that the temperature in the top of the forest canopy was too hot, and the butterflies were seeking shelter in the semi-shaded lower canopy of the secondary forest. This phenomenon is well recorded for forest-dwelling Iolaini in the subfamily Theclinae (e.g. Larsen 2005). A new Pseudaletis from Angola 213 Annls Mus. hist.-nat. hung. 117, 2025 Behaviour One of the female paratypes of the new species dropped from the high canopy onto a tree of approximately 6 metres height after midday in hot and sunny weather; it was clearly freshly hatched and was possibly disturbed by birds or insects before it could fly properly. The male holotype was captured on the same day by 2 pm. It perched high but still within reach (6–8 m) on the foliage of a tree along the dirt track some 50 metres away from the collecting spot of the female in hot and sunny weather. Further two female specimens were collected on the following day, in hot and sunny weather after midday, where they behaved that they were hiding from the heat of the direct sun that hit the top of the forest canopy (Fig. 16). This behaviour resembled to that of the Palaearctic thecline species Favonius quercus (Linnaeus, 1758) and Thecla betulae (Linnaeus, 1758). In the summer heat it is not rare to see hundreds, occasionally thousands of specimens resting in the shade of the subcanopy or in the shrub layer during the hottest hours (pers. obs.; Ebert & Rennwald 1991: 172–173; Bálint 2022: 30). In Afrotropical rainforests, multiple species of essentially canopy-dwelling Iolaini (Lycaenidae, Theclinae) tend to rest under leaves of trees and bushes in the shade along the forest edges or road verges during the hottest hours in dry season (pers. obs.; Larsen 2005). Habitat The general habitat is a forest-miombo mosaic landscape and riverine forest along the Nzanda rivulet near Kinjila village in Malanje Province (Fig. 17), which is known to host rare and near endemic bird species, including the Red-crested Turaco (Tauraco erythrolophus) and the White-headed Robin-chat (Cossypha heinrichi). While the former one is more widely distributed in the Northern Escarpment forests in Cuanza Norte Province, the latter one has a very restricted distribution in Northern Angola and the southern Democratic Republic of the Congo. The area is listed as a Key Biodiversity Area (KBA) and an Important Bird Area (BirdLife International 2025, Key Biodiversity Areas Partnership 2025). The butterfly studies carried out by the author and his team, support the recognition of the area as a KBA, since the fauna consists of a rich mixture of Congolian rainforest species and miombo specialists, similarly to that in northwestern Zambia (Heath et al. 2002). Several other butterfly species from this area appear to differ from all known taxa and could also prove undescribed.