Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera: Eupterotidae: Janinae)
Abstract
Takano, Hitoshi (2025): Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera: Eupterotidae: Janinae). European Journal of Taxonomy 1022: 134-175, DOI: 10.5852/ejt.2025.1022.3085, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3085/13751
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134 European Journal of Taxonomy 1022: 134–175 https://doi.org/10.5852/ejt.2025.1022.3085 europeanjournaloftaxonomy.eu ISSN 2118-9773 2025 · Takano H. This work is licensed under a Creative Commons Attribution License (CC BY 4.0) Received: 21 February 2025 • Accepted: 15 July 2025 • Published: 9 October 2025 Topic editor: Tony Robillard • Section editor: Jurate De Prins • Desk editor: Eva-Maria Levermann Research article urn:lsid:zoobank.org:pub:F9263916-BFA7-4E2F-ABD1-E4DDFFC280C7 Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera: Eupterotidae: Janinae) Hitoshi TAKANO African Natural History Research Trust, Street Court, Leominster, HR6 9QA, UK. Email: [email protected]g.uk Abstract. The distinctive monkey moth genus Camerunia Aurivillius, 1893 is revised, resulting in the implementation of several taxonomic and nomenclatural changes. Two genera historically considered to be allied to Camerunia, Hemijana Aurivillius, 1901 syn. nov. and Catajana Strand, 1910 syn. nov. are synonymised with it and as a result, the genus now contains six species: C. orphne (Schaus, 1893), C.flava Aurivillius, 1904, C. albida Aurivillius, 1901, C. subrosea (Aurivillius, 1893) comb. nov., C. bimaculata (Dewitz, 1879) comb. rev. and a new species, C. smithi sp. nov. described from Tanzania. To preserve the stability of nomenclature, lectotypes are designated for Camerunia insignis Aurivillius, 1893, Cameruniaflava Aurivillius, 1904, Jana subrosea Aurivillius, 1893 and Hemijana ruberrima Rothschild, 1917. Examination of the unlabelled type of Hemijana griseola Rothschild, 1917 syn. nov. has confirmed that it is conspecific with Poloma nigromaculata (Aurivillius, 1893) and it is synonymised accordingly. Keywords. Afrotropics, monkey moths, barcodes, systematics, Hemijana, Catajana. Takano H. 2025. Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera: Eupterotidae: Janinae). European Journal of Taxonomy 1022: 134–175. https://doi.org/10.5852/ejt.2025.1022.3085 Introduction The Afrotropical Eupterotidae Swinhoe, 1892 have remained poorly studied despite their large size and colourful appearance. With the exception of checklists (e.g., Vári et al. 2002; Nässig & Oberprieler 2008; Kitching et al. 2018), descriptions of distinct taxa (e.g., Bouyer 2014) or notes on life histories (e.g., Raviglione et al. 2022), there have been very few taxonomic overviews of the Afrotropical genera or species, with many in need of holistic revision (Takano & László 2022). The subfamily Janinae Aurivillius, 1892 currently comprises 22 genera and 149 species, all of which are restricted to the Afrotropical realm (Kitching et al. 2018). Species of three of these genera, Camerunia Aurivillius, 1893, Hemijana Aurivillius, 1901 and Catajana Strand, 1910, have historically been treated in the literature as being closely allied to each other (e.g., Forbes 1955), and in this current paper, these affiliations are investigated further. The genus Camerunia was initially erected by Aurivillius (1893) for the new species C. insignis Aurivillius, 1893 from Central Africa. In his review of the African Eupterotidae (or Striphnopterygidae
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 135 as it was referred to then), Aurivillius (1901b) transferred Homochroa orphne Schaus, 1893, described from Sierra Leone, to Camerunia and synonymised C. insignis with it, whilst also describing a second species in the genus, C. albida Aurivillius, 1901 from Tanzania. He also tentatively placed Dreata bimaculata Dewitz, 1879 from Angola in Camerunia, although this species was later deemed different enough by Strand (1910) – notably the orientation of the palps, the presence of protibial epiphyses and differing wing venation – to merit a new genus, Catajana. A third species of Camerunia,C.flava Aurivillius, 1904 was later described from East Africa with some uncertainty as to its generic placement due to slight differences in wing venation. Aurivillius (1901b) established the genus Hemijana for Jana subrosea Aurivillius, 1893 based once again on differences in the forewing venation, to which were later added three further taxa, H. ruberrima Rothschild, 1917, H. griseola Rothschild, 1917 and H. variegata Rothschild, 1917. Since Gaede (1927), H. ruberrima has been synonymised with the type species of the genus (Vári et al. 2002) and H. variegata transferred to the genus Poloma Walker, 1855 (Takano & László 2022), leaving two valid species in Hemijana. Forbes (1955) divided the Eupterotidae into several subfamilies and stated that Camerunia and Hemijana were closely related, although rather than comparing with the type species of the latter genus, he treated Catajana bimaculata as belonging to Hemijana. In the most recent catalogues including the Eupterotidae (Nässig & Oberprieler 2008; Kitching et al. 2018), Camerunia comprised three species, Catajana remained monobasic, whilst Hemijana contains two species after the removal of H. variegata as mentioned above. The taxon semijana Aurivillius, 1893, assigned to Hemijana by Kitching et al. (2018), could not be traced in the literature despite determination labels attached to specimens of H. subrosea in NHMUK, and it is treated here as a manuscript name. Based on the rich holdings of Eupterotidae held at the African Natural History Research Trust and in conjunction with specimens in institutional and private collections, the aforementioned genera are herein comprehensively reviewed, resulting in several taxonomic changes. Both Hemijana and Catajana are considered to be junior subjective synonyms of Camerunia based on similar genital morphology and DNA barcodes of their respective type species, and the following combinations are proposed: Camerunia subrosea (Aurivillius, 1893) comb. nov. and Camerunia bimaculata (Dewitz, 1879) comb. rev. Furthermore, the remaining species of Hemijana, H. griseola syn. nov., is deemed to be conspecific with Poloma nigromaculata (Aurivillius, 1893) and is synonymised accordingly. Aside from the two aforementioned Camerunia taxa, the genus sensu novo includes six species, C. orphne, C.flava, C. albida and a new species from the Eastern Arc Mountains of Tanzania, C. smithi sp. nov. Material and methods The preparation of genitalia followed Lafontaine & Mikkola (1987). The phalluses were stained with Eosin red, and the dissected genitalia were mounted on microscope slides or stored in glycerol. Genitalia were photographed using a Canon EOS 700D camera mounted on a Leitz Diaplan compound microscope. Adults were photographed using a Canon EOS 80D with a Canon 100 mm Macro lens. All images were edited in Adobe Photoshop. Primary label data have been transcribed verbatim with “//” denoting a different label and “/” denoting a line break followed by a comment for each label in square brackets (if no indication is given, the label is printed on plain card). Genital terminology follows Forbes (1955) and Oberprieler et al. (2003). Aside from the data extracted from examined specimens, photographic records that could be unequivocally identified from iNaturalist (https://www.inaturalist.org) have been included in the distribution maps. To test the monophyly of Camerunia, DNA barcodes were obtained by removing tarsal segments from 14 adult specimens, belonging to five putative taxa in the three genera, and then submitted to the Canadian Centre for DNA Barcoding (CCDB, Biodiversity Institute of Ontario, University of Guelph). The Janinae genus Hibrildes Druce, 1888 was selected as the outgroup taxon. Sequences were obtained using Single Molecule Real-Time sequencing through the Sequel II (PacBio) pipeline at CCDB (Hebert et al. 2018)
European Journal of Taxonomy 1022: 134–175 (2025) 136 and, together with 12 publicly available sequences from the BOLD (https://boldsystems.org) database, were aligned using MUSCLE in MEGA ver. X (Kumar et al. 2018). Genetic distances were calculated using Kimura’s two-parameter model (Kimura 1980). The final dataset comprised 17 individuals of Camerunia (three taxa), four of Hemijana (one taxon) and five of Catajana (one taxon). Phylogenetic tree searches were performed using Bayesian Inference (BI) and Maximum Likelihood (ML). BI analyses were performed using MrBayes ver. 3.2.7a (Ronquist et al. 2012). Metropolis-coupled Markov chain Monte Carlo (MCMC) analyses were run with four chains (one cold and three heated) for 10 000 000 generations sampling every 100 generations, discarding the first 25% as burn-in. The two runs converged with the standard deviation of split frequencies 0.002. ML analyses were performed using raxmlGUI ver. 2.0 (Edler et al. 2021) using a general time reversible model (GTR-GAMMA). Support for clades was evaluated for BI using posterior probabilities and ML using rapid bootstrapping with 1000 replicates. Trees were visualised and annotated in FigTree ver. 1.4.4 and Adobe Photoshop. Institutional abbreviations AMNH = American Museum of Natural History, New York, USA ANHRT = African Natural History Research Trust, Leominster, U.K. MCSN = Museo Civico di Storia Naturale Giacomo Doria, Genoa, Italy MfN = Museum für Naturkunde, Berlin, Germany MZUF = Museo di Storia Naturale “La Specola”, Florence, Italy NHRS = Naturhistoriska riksmuseet, Stockholm, Sweden NHMUK = Natural History Museum, London, U.K. NHMW = Naturhistorisches Museum, Vienna, Austria OUMNH = Oxford University Museum of Natural History, Oxford, U.K. RCPB = Research collection of Patrick Basquin, Yvetot-Bocage, France RMCA = Royal Museum for Central Africa, Tervuren, Belgium USNM = National Museum of Natural History, Washington, D.C., USA Results Synthesis of molecular and morphological data Phylogenetic inferences based on BI and ML recovered identical topologies (ML tree figured in Fig. 1) wherein Camerunia was recovered as paraphyletic with regard Hemijana and Catajana. Camerunia orphne was recovered as sister to a moderately well-supported (Hemijana + Catajana) + (Camerunia smithi sp. nov. + Cameruniaflava) clade (PP: 85; BS: 59). The recovery of the Hemijana + Catajana (PP: 100; BS: 92) clade was unsurprising considering their respective type species share similar wing patterns as well as genital morphology. Hemijana was originally separated from all other Janinae based on a different length ratio of the lower and middle discocellular veins (Aurivillius 1901b), but the general appearance and genital configuration of its type species is very much that of the Camerunia-form. Aurivillius relied heavily upon venation to arrange and classify the Bombycoidea Latreille, 1802 and Lasiocampoidea Harris, 1841, but its suitability in Eupterotidae was considered doubtful by Mell (1930) and more recently has been shown to be uninformative in Lasiocampidae Harris, 1841 (Takano & László 2024). Similarly, for Catajana wing venation and the presence of an epiphysis on the foretibia were the traits outlined to differentiate it from Camerunia. Forbes (1955) considered Hemijana and Catajana to be congeneric and, referring to the type species of Catajana, Forbes (1955: 95) noted that if it were not for the venation, “the genus would hardly be considered distinct from Camerunia”. Moreover, Forbes (1955) correctly indicated that C. bimaculata lacked an epiphysis on the foretibia and suggested that Strand (1910) had confused it with the epiphyses of the mesotibia. Perhaps the most unexpected result from the phylogenetic analyses was the recovery of C.flava with C. smithi in a well-supported clade (PP: 99; BS: 67), and not C. orphne, which shares with C.flava tufts of ciliate scales on the abdomen and the flattened lateral processes of the tegumen in the male genitalia. Seemingly, C. smithi is a peculiar species
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 137 with wing patterns closest to C. orphne whereas the genitalia are most similar to the Hemijana / Catajana species. In summary, both Hemijana syn. nov. and Catajana syn. nov. are here treated as junior subjective synonyms of Camerunia based on external and genital morphology as well as the phylogenetic results. Taxonomy Class Insecta Linnaeus, 1758 Order Lepidoptera Linnaeus, 1758 Superfamily Bombycoidea Latreille, 1802 Family Eupterotidae Swinhoe, 1892 Subfamily Janinae Aurivillius, 1892 Genus Camerunia Aurivillius, 1893 Camerunia Aurivillius, 1893: 211. Type species: Camerunia insignis Aurivillius, 1893, by monotypy. Hemijana Aurivillius 1901b: 21. Syn. nov. Type species: Jana subrosea Aurivillius, 1893, by monotypy. Fig. 1. Maximum likelihood tree of the Camerunia Aurivillius, 1893 species, with Hibrildes Druce, 1888 as outgroup, based on DNA barcodes. Numbers above branches indicate posterior probabilities (PP) and bootstrap (BS) below. Scale bar indicates substitution rates per site.
European Journal of Taxonomy 1022: 134–175 (2025) 138 Catajana Strand 1910: 98. Syn. nov. Type species: Dreata bimaculata Dewitz, 1879, by original designation. Camerunia – Aurivillius 1901b: 27; 1904: 698. — Gaede 1927: 306. — Forbes 1955: 94, 129. — Fletcher & Nye 1982: 29. — Vári et al. 2002: 150. — Nässig & Oberprieler 2008: 56. — Kitching et al. 2018: suppl. material 1. Hemijana – Gaede 1927: 302. — Forbes 1955: 94, 129. — Fletcher & Nye 1982: 75. — Vári et al. 2002: 150. — Nässig & Oberprieler 2008: 56. — Kitching et al. 2018: suppl. material 1. Catajana – Gaede 1927: 306. — Forbes 1955: 129 (as synonym of Hemijana). — Fletcher & Nye 1982: 32. — Vári et al. 2002: 150. — Nässig & Oberprieler 2008: 56. — Kitching et al. 2018: suppl. material 1. Diagnosis Camerunia species are mediumto large-sized, brightly-coloured, highly distinctive monkey moths with broad, papery wings and, compared to other Janinae, have a proportionally small body. There is a superficial resemblance to the peculiar genus Hibrildes, which, unlike Camerunia, displays a high degree of sexual dimorphism (Poulton 1929). Male Hibrildes have almost translucent wings with markings mostly limited to densely packed dark brown scales along well-defined veins. The females, in contrast, are well-marked and polymorphic, displaying aposematic colouration that varies from pale yellow through to orange, with a dark discal bar-like marking present on both pairs of wings. The male genital morphology of the two genera is, however, very different; those of Hibrildes are highly modified with lateral processes on the uncus and tripartite valves (Minet 1994; Oberprieler et al. 2003). Redescription of the adult Antenna bipectinate in both sexes, rami shorter in females of some species. Ground colour of body and forewing varying from white, through yellow to brown; hindwing white, yellow or pink. Forewing triangular, rounded at apex and outer margin gently arcuate; hindwing ovoid, outer margin arcuate. Both wings often with transverse fasciae; forewing may display basal and discal markings. Vein CuP of the forewing present and highlighted with brown scaling in some species. Male genitalia. Uncus reduced, fused with tegumen. Tegumen broad with a pair of long apically rounded lateral projections. Gnathos with spinose or clavate process. Valve trapezoidal or triangular with well-defined costa and sacculus, cleft in some species. Juxta trapezoid, emarginate distally. Vinculum V-shaped. Saccus cylindrical, rounded at apex. Phallus as long as valve, slightly curved medially; carina scobinate in some species; coecum rounded. Vesica with scobination. Molecular characterisation The barcode region of Camerunia species was variable with intraspecific pairwise distances (PWD) for the five sampled taxa ranging from 0.0–5.8%, while interspecific PWDs were 6.0–13.1%. They diverged from the outgroup taxon Hibrildes by 13.6–16.5%. Aside from C. subrosea (Aurivillius, 1893) comb. nov. and C. smithi sp. nov., each of the remaining species was assigned numerous Barcode Index Numbers (BIN) in BOLD as a result of the large intraspecific divergences. Although the BINs in general showed geographical concordance, no differences in habitus or genital morphology were observed in the individuals assigned to each BIN, and thus any further taxonomic splitting (which would ultimately be based solely on barcodes) could not be justified.
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 139 Distribution Members of this genus are widely distributed from north-eastern South Africa as far north as Uganda and westwards to Sierra Leone and Guinea. With the exception of C. orphne and C. smithi sp. nov., which inhabit forested environments, all other species are found in open woodland and forest-savanna mosaics. Descriptions of the species Due to incomplete taxon sampling in the phylogeny, the species are presented below based on the configurations of the male genitalia. The genus naturally divides into two groups, one with flattened lateral processes of the tegumen (C. orphne, C.flava and C. albida) and another with cylindrical processes (C. smithi sp. nov., C. bimaculata (Dewitz, 1879) comb. rev. and C. subrosea (Aurivillius, 1893) comb. nov.). Camerunia orphne (Schaus, 1893) Figs 1–8, 37–38, 48, 54 Homochroa orphne Schaus, 1893: 30, pl. 1 fig. 7. Type locality: “Sierra Leone”. Camerunia insignis Aurivillius, 1893: 211. Type locality: “Camerun interior; Gabun”. Camerunia orphne – Aurivillius 1901b: 27. — Gaede 1927: 306, pl. 46 fig. c. — Kitching et al. 2018: suppl. material 1. Camerunia insignis – Aurivillius 1901b: 27 (syn.). Diagnosis The largest species of the genus, C. orphne cannot be easily confused with any other species of Camerunia, with perhaps the exception of C. smithi sp. nov., which is smaller and distributed allopatrically. In both these species, a dark brown basal patch is present on the forewing, but the discal markings are comprised of dots in C. smithi, whereas in C. orphne, it is a bar as wide as the cell, along the medial vein. The configuration of the male genitalia is most similar to C.flava and C. albida, with the flattened, apically rounded lateral processes of the tegumen, but the long, clavate, denticulate, apical process of the gnathos is unique to C. orphne. Type material Holotype of Homochroa Orphne SIERRA LEONE • ♂ (Fig. 2); “No. 19248 / W. Schaus / Collection. [black border; partially handwritten] // TYPE / No. / A.M.N.H. [red card] // Homochroa / orphne / Type. Schs, [handwritten in Schaus’s hand]”; AMNH. Lectotype of Camerunia insignis (by present designation) CAMEROON • ♂ (Fig. 3); “Kamer. / int. / Pr. // 272. [handwritten] // Camerunia / insignis / Aur. ♂ typ. [handwritten] // 16,1. [handwritten]”; MfN. Paralectotype of Camerunia insignis GABON • ♀ (Fig. 8); “Gabun / 1890. Mocquerys [black border] // Camerunia / insignis / Aur. ♀ typ. [handwritten] // 16,1. [handwritten]”; MfN. Other material examined (24 ♂♂, 10 ♀♀) CAMEROON • 1 ♂; [unspecified locality]; D.G. Rutherford leg.; OUMNH. – North • 1 ♀; Bascho [= Bashéo]; 28 Aug. 1906; A. Schultze leg.; MfN. – South • 1 ♂, 1 ♀; Bitje; 610 m a.s.l.; G. Bates leg.;
European Journal of Taxonomy 1022: 134–175 (2025) 140 NHMUK. – Southwest • 1 ♀; Bibundi; 15–30 Oct. 1904; G. Tessmann leg.; MfN • 1 ♂, 1 ♀; Victoria; P. Preuss leg.; MfN • 1 ♂; same locality as for preceding; 1892; G. Zenker leg.; MfN. CENTRAL AFRICAN REPUBLIC – Nana-Grébizi • 1 ♀; Fort Crampel; NHMUK. DEMOCRATIC REPUBLIC OF CONGO – Lualaba • 1 ♂; Kapanga; Mar. 1934; F.G. Overlaet leg.; RMCA. GABON – Estuaire • 1 ♂; Monts de Cristal; 8 Dec. 1990; 700 m a.s.l.; P. Basquin leg.; RCPB. GHANA – Ashanti • 2 ♂♂; [unspecified locality]; Mar. 1907; G.C. Dudgeon leg.; NHMUK • 1 ♀; Friapere Forest; 1913; NHMUK. GUINEA – Mamou • 2 ♂♂; Chute de Ditinn; 10°49′08″ N, 12°11′30″ W; 771 m a.s.l.; 18–25 Sep. 2019; M. Geiser, M. Leno, S. Koivagui, W. Miles, L. Mulvaney and S. Sáfián leg.; ANHRT. IVORY COAST – Montagnes • 1 ♂; Mont Tonkoui; 7°27′15″ N, 7°38′12.6″ W; 1200 m a.s.l.; Jun. 2016; ANHRT. – Woroba • 1 ♀; Dolla Ranch; 7°58′07.7″ N, 07°34′35.7″ W; 481 m a.s.l.; 1–5 Jun. 2018; M. Aristophanous, W. Miles, P. Moretto and Y. Outtara leg.; ANHRT. LIBERIA – Maryland • 1 ♂; Cape Palmas; Seaton leg.; NHMUK. – Nimba • 1 ♀; Bahn; 7°02′ N, 8°47′ W; 457 m a.s.l.; 24 Sep. 1953; W. Peters leg.; NHMUK • 1 ♂; Nimba Mountains, Mount Gangra summit; 7°32′45.82″ N, 08°38′09.36″ W; 974 m a.s.l.; 17–25 Mar. 2017; S. Sáfián and G. Simonics leg.; ANHRT. NIGERIA – Enugu • 1 ♀; Ogruga [= Ogrugru]; NHMUK. REPUBLIC OF CONGO – Cuvette-Ouest • 1 ♂; Odzala-Kokoua National Park, Bangassou Forest near Lobo; 0°32′50″ N, 14°51′47″ E; 400 m a.s.l.; 21–26 Sep. 2024; M. Bashford, G. László, M. Talani and A. Volynkin leg.; ANHRT. SIERRA LEONE • 2 ♂♂, 1 ♀; [unspecified locality, but most likely Freetown]; P. Bainbridge leg.; NHMUK. – North West • 1 ♂; Batkanu; F.G. FitzGerald leg.; NHMUK. – Northern • 3 ♂♂; Kalainkay, near Kamabai; 9°10′52″ N, 11°56′44″ W; 80 m a.s.l.; 3–6 Nov. 2015; R. Goff leg.; ANHRT. – Western Area • 1 ♂; Freetown; 19 Apr. 1917; P.A. Buxton leg.; OUMNH • 1 ♂; Wilberforce; 9 Nov. 1911; C.A. Foster leg.; OUMNH. TOGO – Plateaux • 2 ♂♂; Forêt de Missahohe, Kpalime; 6°56′41″ N, 0°34′30″ E; 610 m a.s.l.; 31 Jul. 2013; P. Moretto leg.; ANHRT. Redescription Male (Figs 2–6, 48) Forewing length. 28–40 mm. Upperside. Ground colour of head, thorax and abdomen brown; vertex creamy-white, patagium mahogany, tegula with paler beige scales turning darker mahogany posteriorly. Antenna bipectinate, brown. Abdomen covered with greyish scales and tufts of bronze, longer ciliate scales medially forming crest which runs along abdomen. Each abdominal segment with bronze scales along posterior margin, giving abdomen banded appearance. Eighth sternite truncate posteriorly, weakly sclerotised, its surface punctate. Wing veins and termen well-defined with dark brown scaling. Forewing triangular, rounded at apex, outer margin gently arcuate, almost straight; ground colour beige to brown. Basal marking
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 141 dark chocolate turning black towards anal margin, under half length of cell; its outer margin arcuate, slightly sinuate, with white fascia on inner (and sometimes outer) edge of border. Antemedial fascia dark brown, indistinct, following margin of basal marking. Discal marking black with greyish-white scaling internally, in form of bar along outer edge of medial vein. Postmedial fasciae dark brown, arcuate, evenly crenulate, outer of two more indistinct; fasciae arising almost perpendicularly from dark brown spot on costa, sharply curving inward at vein M2 and terminating almost perpendicularly along anal margin. Space CuA1 and dorsal portion of CuA2 with well-defined darker brown marking filling space between postmedial fasciae. Submarginal fasciae dark brown, crenulate, broadly running in parallel with postmedial fasciae; inner of two arising from small dark brown marking on costa, outer arising from dark brown rectangular sub-apical marking perpendicular to costa. Subterminal area irrorated with darker scales and dark brown triangular patch, its inner margin arising ventrad of apex, widening proximad as far as submarginal fascia in space M2 and terminating along termen around vein CuA1. Fringe brown with pale greyish-white scales forming dots at termination points of veins. hindwing. Outer margin arcuate; ground colour pale yellow, sometimes with ferruginous tint basally. Postmedial fascia dark brown, arcuate and angled at vein M2, weakly crenulate, terminating at dark brown spot along anal margin. Submarginal fascia dark brown, composed of dark brown lunules between veins running parallel with postmedial fascia. Subterminal fascia dark brown, strongly crenulate. Fringe as on forewing. Underside. Ground colour of body golden yellow; wings yellow, veins and termen well-defined with dark brown scaling. Forewing costa highlighted with gold scaling to apex. Postmedial and submarginal fasciae often absent but when present, similar to upperside but finer and not bilineate. Dark brown patch in subterminal area often present but less well-defined. Hindwing postmedial, submarginal and subterminal fasciae as on upperside but better defined. Both postmedial and submarginal fasciae often bilineate, outer of both less well-defined. Male genitalia (Figs 37–38). Uncus reduced, fused with tegumen. Tegumen broad with pair of long flattened, apically rounded, truncate lateral projections, its inner surface setose. Gnathos with long clavate process, serrate basally becoming smooth towards its rounded apex. Valve trapezoidal, apically bifurcate. Costa with short, blunt, distal process at apex; costal margin slightly concave. Sacculus welldefined, third width of valve at base, tapering towards apex before broadening into short, pointed distal process. Juxta trapezoid, proximally V-shaped, distally bifurcate, with broad u-shaped emargination; each process heavily serrated. Vinculum V-shaped. Saccus cylindrical, rounded at apex. Phallus slightly longer than valve, gently curved medially; carina scobinate; coecum rounded. Vesica with scobination. Female (Figs 7–8) Forewing length. 36–48 mm. Similar to male but larger, and rami of antenna half the length. Variation Specimens from southern Cameroon, Gabon and D.R. Congo are in general larger (especially the females) and the males often display a strong pinkish hue at the base of the hindwing (Figs 5, 8). In the male genitalia, the extent of the serrations at the base of the clavate process of the gnathos varies greatly among individuals across its range.
European Journal of Taxonomy 1022: 134–175 (2025) 142 Figs 2–8. Camerunia orphne (Schaus, 1893), adults. 2. Holotype of Homochroa orphne Schaus, 1893, ♂ (AMNH). 3. Lectotype of C. insignis Aurivillius 1893, ♂ (MfN). 4. ♂, Sierra Leone, Kalainkay (ANHRT). 5. ♂, Cameroon, Bitje (NHMUK). 6. ♂, Guinea, Chute de Ditinn (ANHRT). 7. ♀, Ivory Coast, Dolla Ranch (ANHRT). 8. C. insignis, paralectotype, ♀ (MfN).
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 149 Figs 9–16. Cameruniaflava Aurivillius, 1904, adults. 9. Lectotype, ♂ (NHMUK). 10. Holotype of C.lactiflora Wichgraf, 1914, ♂ (NHMUK). 11. ♂, Zambia, Chilambwe Falls (ANHRT). 12. ♂, Cameroon, Wack (ANHRT). 13. ♂, Malawi, Zomba (NHMUK). 14. ♂, Sudan, Tambura (NHMUK). 15. ♀, Malawi, Zomba (NHMUK). 16. ♀, Zambia, Chilambwe Falls (ANHRT).
European Journal of Taxonomy 1022: 134–175 (2025) 150 Molecular characterisation This species has been assigned three BINs: BOLD:AFG4533 (Cameroon), BOLD:AAU2961 (Tanzania), BOLD:ABX5434 (D.R. Congo) and BOLD:AFG4806 (Zambia). Intraspecific PWDs were 0.2–5.8% (n = 9) diverging from its nearest neighbour, C. smithi sp. nov. by 8.8–9.4% (n=2). Distribution (Fig. 55) This species is widely distributed throughout woodland habitats and forest-savanna transitional zones in central and eastern Africa, with an isolated population in Cameroon. It is absent from the rainforests of the Congo Basin and is found no further west than the Cameroon highlands, and no further south than the Zambezi River. This species is known from Cameroon, D.R. Congo, South Sudan, Uganda, Rwanda, Tanzania, Malawi, Zambia and Angola. It is likely present in Burundi, Kenya and Mozambique, and may be found in the Central African Republic. Remarks The two male and two female syntypes of C.flava are all held in NHMUK. To preserve the stability of nomenclature by fixing the published name to a single specimen, the best-preserved male specimen from the Stevenson Road, collected by Reverend Alexander Dewar, is designated as the lectotype (Fig. 9). The Stevenson Road was a thoroughfare that linked the northern end of Lake Malawi with the southern end of Lake Tanganyika, running broadly parallel to the Tanzania-Zambia border. Dewar served with the Free Church of Scotland at Livingstonia, Malawi, and founded the mission station at Mwenzo, where he was stationed between 1894–1896 (Jack 1901); it is most probable that the two male specimens were captured there. George Hampson noted that the two female specimens in the syntype series from Zomba, Malawi, and ‘Tanganyika’ differed slightly in appearance (Aurivillius 1904: 698). The Malawian specimen was collected by Brigadier-General Sir William Manning, deputy commissioner of British Central Africa at the time, and the Tanzanian specimen was collected by Alexander Carson, a London Missionary Society agent, likely originating from the southern half of Lake Tanganyika based on his published observations (Carson 1892) and other specimens he collected (Butler 1895). In the original description of C. lactiflora, Wichgraf (1914) provided the type locality as Lake Tanganyika, but the label attached to the specimen suggests the specimen was taken at Lake Victoria (Fig. 10). It is presumed, however, that the former is the true type locality, as Wichgraf (1914) stated “Vom Bahnbau in der Nähe des Tanganjikasees [From the railway construction near Lake Tanganyika]” for another species described in the same paper. The Tanganyika Railway (Tanganjikabahn) linking Dar es Salaam with Kigoma was completed in early 1914, with the line to Kalenge, 30 km from the lake, opening to the public by October 1913 (Baltzer 1916). Although a railway line towards Lake Victoria (Ruandabahn) had been planned, it was only approved in 1914 (Baltzer 1916), and thus the locality data on the holotype specimen is likely a typographical error by Wichgraf. Gaede’s (1927) reference to Ethiopia for var. lactiflora is difficult to interpret and although possible that he had seen material from this region, it is unlikely. Camerunia albida Aurivillius, 1901 Figs 17–19, 41, 50, 56 Camerunia albida Aurivillius, 1901b: 27, fig. 17. Type locality: [Tanzania] “Nyangao”. Camerunia albida – Gaede 1927: 306, pl. 46 fig. c. — Kitching et al. 2018: suppl. material 1.
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 151 Type material Holotype TANZANIA • ♂ (Fig. 17); “Typus [red card with black border] // Deutsch. Ost- / Afrika // Camerunia / albida Aur. typ [handwritten in Aurivillius’ hand] // NHRS-SRAH / 000001416”; NHRS. Diagnosis This is the smallest species in the genus, and its creamy-white habitus is highly distinctive and perhaps most reminiscent of male Hibrildes, although in the latter, the wings are more translucent and lack markings except for the well-defined veins. The discal markings are similar to C. bimaculata (Dewitz, 1879) comb. rev. and C. subrosea, (Aurivillius, 1893) comb. nov. but in these two species, the markings are composed of two or three spots, rather than the four in C. albida. The apical patch and subterminal markings are not dissimilar to C. orphne and C. smithi sp. nov., but the ground colour of both these species is straw-yellow and the forewings display a distinctive dark basal patch. The eighth sternite of C. albida shares similarities with C. smithi in the heavily setose posterior margin and emarginate posterior margin, but in the latter, the sternite tapers posteriorly and the margin is deeply emarginate. The ground plan of the male genital capsule is most similar to C.flavabut in the latter, the lateral projections of the tegumen are narrower, lacking the two long spines, and instead an apically arcuate pad with a heavily spinulose outer margin is present near the base of the inner surface. Furthermore, in C.flava, the gnathos is smaller and the spines are considerably shorter and more densely packed. Redescription Male (Figs 17, 50) Forewing length. 27 mm. Upperside. Ground colour of head, thorax and wings creamy-white; abdomen pale brown. Vertex and patagium irrorated with brown scaling. Antenna bipectinate, dark brown. Abdomen irrorated with creamy-white scaling, abdominal segments with pale yellow scaling posteriorly giving banded appearance; anal tuft pale yellow. Eighth sternite gently concave anteriorly and weakly emarginate posteriorly, with some sclerotisation along midline; its surface covered throughout in fine punctures, heavily setose along posterior margin with longer setae medially. Forewing triangular, rounded at apex, outer margin gently arcuate; costa with brown scaling basally. Antemedial fascia dark brown, sinuate, indistinct. Cell bisected with longitudinal brown fascia running from base to medial vein. Dark brown scaling along vein CuP from base to outer margin. Discal marking black, quadruple, pair on either side of medial vein; outer pair larger with curved margins distad, inner pair smaller, one placed costad indistinct. Postmedial fascia dark brown, arcuate, gently crenulate; fascia arising perpendicularly just over halfway along costa and gently curved inwards at vein M1, terminating almost perpendicularly to anal margin. Large, broadly diamond-shaped, dark brown marking present at intersection of postmedial fascia and vein CuA1, about 5 mm wide and almost reaching veins on either side. Submarginal fascia dark brown, sinuate, running in line with postmedial fascia. Triangular apical patch, dark brown, its inner and outer edges placed equally just distad of submarginal fascia and proximad of subterminal area, respectively. Subterminal area dark brown, demarked proximally by sinuate margin arising near apex, broadening to be at its widest at vein M2, tapering towards vein CuA1 and petering out near tornus. Two indistinct dark brown spots placed proximad of the subterminal area in spaces CuA1 and CuA2. Fringe creamy-white (although unclear based on the limited remnants of the holotype). h indwing . Outer margin arcuate. Postmedial fasciae dark brown, arcuate, gently crenulate and generally indistinct, although better defined towards anal margin. Submarginal fasciae composed of dark brown lunules between veins, widest in space CuA1. Subterminal area irrorated with dark brown scaling.
European Journal of Taxonomy 1022: 134–175 (2025) 152 Underside. Ground colour as on upperside, but body entirely creamy white except for brown legs and some pale yellow scaling. Postmedial fascia as on upperside but indistinct. Submarginal fascia as on upperside but slightly displaced distad and running closely in line with inner edge of apical patch. Subterminal margin as on upperside but indistinct. Hindwing postmedial and submarginal fasciae as on upperside, but finer. Male genitalia (Fig. 41). Uncus reduced, fused with tegumen. Tegumen broad with pair of long, flattened, apically rounded, truncate lateral projections, angled medially; its inner surface heavily setose with two spines, one long spine (about half the length of lateral projection) three-quarters of way along dorsal margin and smaller spine half the size, ventrally near base. Gnathos emarginate proximally, with pair of long curved spines dorsally and three pairs of long spines, each shorter proximally, either side of midline. Valve triangular and cleft (about two-thirds of way along valve). Costa blunt at apex. Sacculus well-defined, tapering apically into gently curved distal process. Juxta trapezoid, proximally V-shaped, lateral margins tapering dorsad, distally weakly V-shaped. Vinculum V-shaped. Saccus triangular, rounded at apex. Phallus slightly longer than valve, gently curved medially, tapering towards carina; coecum rounded. Vesica with sparse scobination. Female Unknown. Variation Unknown. Figs 17–21. Species of Camerunia Aurivillius, 1893, adults. 17–19. C. albida Aurivillius, 1901, holotype, ♂ (NHRS). 17. Upperside view (left), underside view (right). 18. Figure from Aurivillius (1901b). 19. Figure from Gaede (1927). 20. C. smithi sp. nov., holotype, ♂ (ANHRT), upperside view (left), underside view (right). 21. C. smithi, paratype ♂ (ANHRT).
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 153 Larval foodplant Unknown. Molecular characterisation Unknown. Distribution (Fig. 56) This species is known only from the holotype male from coastal south-east Tanzania, a region that harbours numerous endemic species, including butterflies such as Triskelionia compacta (Evans, 1951) and other bombycoids such as the hawkmoth, Rufoclanis maccleeryi Carcasson, 1968. It has been reported that much of the original forest in this general area has been degraded (Larsen & Congdon 2011). Remarks Aurivillius (1901b) gave the type locality as Nyangao in south-eastern Tanzania, despite there being no indication of this on the labels attached to the type specimen (Fig. 17). Nyangao was a German mission station set up in 1896 (Hassing 1970) and a search of the literature revealed that at the turn of the 20th century, most entomological specimens from this locality were in the collection of Johann Ertl, who had in turn obtained them from the Missionary Benedictines of St Ottilien (Suffert 1904). Aurivillius was well-acquainted with Ertl and his collection, having described numerous Tanzanian species of Coleoptera Linnaeus, 1758 (mainly Cerambycidae Latreille, 1802) over a ten-year period (e.g., Aurivillius 1903). It is also known that Aurivillius (1901a) had specimens of Lepidoptera from Ertl in front of him at the time of the description of C. albida, and although only a single specimen is known of this taxon, its provenance is currently not in question. The species has been figured twice in the literature, the first in the original description (reproduced here in Fig. 18) and once more in Gaede (1927) (reproduced here in Fig. 19). Despite the differences in the depictions, the former being the most accurate and the latter with some artistic stylisation, it is believed they refer to the same unique holotype specimen. Camerunia smithi sp. nov. urn:lsid:zoobank.org:act:319163B9-7853-44DD-AF06-C2B338E45C96 Figs 1, 20–21, 42–43, 51, 56 Diagnosis This distinctive species can only be confused with C. orphne, sharing with it the dark brown basal marking on the forewing but the latter is a larger insect, and the forewing discal marking forms a bar along the medial vein. Moreover, these two species are found in allopatry on either side of Rift Valley and thus there is unlikely to be any confusion in specimens with well-documented provenance. The male genitalia of C. smithi sp. nov. are of the Camerunia bimaculata / subrosea type, but those of C. bimaculata / subrosea differ from that of the new species in the gently curved lateral processes of the tegumen (angled in C. smithi), the smaller but more spinulose gnathos, the apices of the valves hooked ventrad (straight in C. smithi) and the longer, narrower saccus. Etymology The new species is dedicated with great pleasure to Richard Smith, Chairman of the Board of Trustees, ANHRT, for his outstanding and continued support of and contribution to African entomology through the institution he founded.
European Journal of Taxonomy 1022: 134–175 (2025) 154 Type material Holotype TANZANIA • ♂ (Fig. 20); “TANZANIE : Morogoro Re- / gion, Kaguru Mts., 1870m., / 06°22.367’S.,036°55.820’E., / 7-XI-2013 (Ph. Darge) // Ex. Coll. Ph. Darge / ANHRT:2019.20 // ANHRTUK / 00207258”; ANHRT. Paratypes TANZANIA • 6 ♂♂; same data as for holotype; ANHRT. Description Male (Figs 20–21, 51) Forewing length. 29–32 mm. Ground colour of body dark chocolate-brown. Vertex and thorax sparsely irrorated with white scales. Antenna bipectinate, dark brown. First abdominal segment with dense tuft of white scales anteriorly; first two segments darker than ground colour, irrorated with white scales. Remaining segments with tufts of longer ciliate scales medially forming crest which runs along abdomen. Eighth sternite with deep concavity posteriorly, and heavy sclerotisation latero-anteriorly; its surface evenly covered in fine punctures and heavily setose posteriorly. Anal tuft irrorated with white scales. Forewing triangular, rounded at apex, outer margin gently arcuate, almost straight; ground colour pale straw. Basal marking, dark chocolate, almost half length of cell; its outer margin arcuate, slightly sinuate, with white fascia on the inner edge of border. Costa with dark spot between basal marking and discal marking. Discal marking black, comprising three fine spots on outer edge of medial, one each at junctures of veins M1–M3, spot at M2 being smallest and longitudinally elongate. Postmedial fascia, dark brown, bilineate, arcuate, evenly crenulate, outer of two more indistinct; fasciae arising almost perpendicularly from dark brown spot on costa, sharply curving inward at vein M2 and terminating almost perpendicularly along anal margin. Space CuA1 and dorsal portion of CuA2 with well-defined darker brown marking filling space between postmedial fasciae. Submarginal fasciae dark brown, bilineate, crenulate, broadly running in parallel with postmedial fascia; inner of two arising from small dark brown marking on costa with two longitudinal dark brown markings proximad along veins M3 and CuA1; outer arising from three well-defined dark brown markings with rounded margins distad, sometimes conjoined proximad, forming subapical patch. Subterminal fascia dark brown, crenulate, following outer margin. Subterminal area with olive brown triangular patch, its inner margin arising at apex, widening proximad as far as subterminal fascia in space M2 and terminating along termen around vein CuA1. Fringe olive brown with pale straw-coloured scales forming dots at termination points of veins. hindwing. Outer margin arcuate; ground colour pale yellow. Postmedial fascia dark brown, bilineate, arcuate and angled at vein M2, weakly crenulate, outer of two indistinct, terminating at dark brown spot along anal margin. Submarginal fascia dark brown, bilineate, running in parallel with postmedial fascia, the inner crenulate and indistinct, the outer composed of dark brown lunules between veins, terminating at dark brown spot along anal margin. Subterminal fascia dark brown, crenulate. Termen highlighted with olive-brown scales. Fringe as on forewing but irrorated with yellow scales. Underside. Ground colour of body and wings yellow, legs brown. Forewing markings showing through from upperside, but only postmedial fascia and submarginal triangular patch weakly marked. Hindwing veins and termen well-defined with dark brown scaling. Postmedial, submarginal and subterminal fasciae as on upperside but better defined. M ale genitalia (Figs 42–43). Uncus reduced, fused with tegumen. Tegumen broad with pair of long, cylindrical, apically pointed lateral projections, medially hooked inward; its inner surface heavily setose.
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 155 Gnathos bilobed, with small, dense cluster of spines, either side of midline. Valve triangular and cleft (about half way along valve). Costa blunt at apex with tiny spine ventrally. Sacculus well-defined, tapering into curved distal process rounded at apex. Juxta trapezoid, proximally V-shaped, lateral margins gently tapering dorsad, distally V-shaped. Vinculum V-shaped. Saccus triangular, short (as long as gnathos), rounded at apex. Phallus slightly longer than valve, curved at coecum but otherwise straight; coecum rounded; carina weakly scobinate. Vesica with sparse scobination. Female Unknown. Variation Based on the short series of males, the extent of the discal markings varies slightly, the smallest longitudinally elongate spot missing in some individuals. Larval foodplant Unknown. Molecular characterisation The new species has been assigned the BIN BOLD:AFF5559. The intraspecific PWD was 0.0% (n = 2) diverging from its nearest neighbour, C.flava by 8.8–9.4% (n = 9). Distribution (Fig. 56) The new species is known only from the seven male specimens from the type locality in Mamiwa-Kisara Forest Reserve, Ukaguru Mountains, Tanzania, and it is almost certainly restricted to this forest block. The Ukagurus form part of the Eastern Arc Mountains, well-known for their exceptional biodiversity and single-locality endemics. The Mamiwa-Kisara Forest Reserve constitutes the largest tract of montane forest in the Ukagurus, which is home to numerous endemic species, although habitat degradation has recently been reported at this site (Lawson et al. 2023). Recent discoveries of narrow-ranged bombycoid species in this chain of mountains include Temnora smithi Takano, 2022, known only from a small patch of montane forest in the Mahenge Mountains (Takano 2022). Camerunia bimaculata (Dewitz, 1879) comb. rev. Figs 1, 22–27, 44–45, 52, 56 Dreata bimaculata Dewitz, 1879: 33, pl. 2 fig. 5. Type locality: [Angola] “dem 10° S.B. und zwischen dem 17–22° O.L. von Greenw.” Camerunia bimaculata – Aurivillius 1901b: 27 (comb.) Catajana bimaculata – Strand 1910: 98. — Gaede 1927: 306, pl. 46 fig. d. — Kiriakoff 1954: 11. — Vári et al. 2002: 150. — Kitching et al. 2018: suppl. material 1. — Krüger 2020: 138. Hemijana bimaculata – Forbes 1955: 94, 129. Diagnosis Camerunia bimaculata (Dewitz, 1879) comb. rev. is a highly distinctive insect taxon that cannot readily be confused with any of its congeners. It shares a yellow ground colour of the wings with C.flava, but in the latter, both sets of wings are of the same colour and the forewing lacks the discal markings as well as fasciae. The general pattern of the wings is somewhat reminiscent of that of C. subrosea (Aurivillius, 1893) comb. nov., but in the latter, the forewings generally have a slightly pinkish tinge, whilst the hindwings range from rosy-red to pink. The male genitalia of C. bimaculata are most similar to those
European Journal of Taxonomy 1022: 134–175 (2025) 156 of C. subrosea, but in the latter, a digitate basal process is present along the ventral margin of the lateral process of the tegumen (absent in C. bimaculata), the spines of the gnathos are more densely packed together arising closer to the midline, the apex of the saccular process is triangular and pointed (rounded in C. bimaculata), and the phallic tube is shorter and more robust. Type material Holotype ANGOLA • ♂ (Fig. 22); “21768 // guinea / inter. / Pogge [green paper; handwritten] // Bimaculata / Dwtz. * [green paper; handwritten] // Bimaculata / Dwtz. Mittheill. / J. Münchener ent. / Vereins. 1879. p.33. / t.II. f.5 [green paper; handwritten]”; MfN. Other material examined (58 ♂♂, 7 ♀♀) ANGOLA – Bié • 1 ♀; Bihé [= Kuito]; F.S. Arnot leg.; NHMUK. – Malanje • 1 ♂; Fort Don Carlos; 20 Sep. 1903; W.J. Ansorge leg.; NHMUK. – Moxico • 1 ♂; Moxico District; 1219 m a.s.l.; Oct. 1928; T.A. Barns leg.; NHMUK. DEMOCRATIC REPUBLIC OF CONGO – Haut-Katanga • 1 ♂; Elisabethville [= Lubumbashi]; 14 Jan. 1948; C. Seydel leg.; RMCA • 1 ♂; same collecting data as for preceding; 26 Nov. 1955; RMCA • 1 ♂; Likasi; 1924; NHMUK • 1 ♂; Luena; 25 Sep. 1983; RMCA • 1 ♂; Mwera; 23 Feb. 1982; RMCA • 1 ♂; Parc National de l’Upemba, Kaswabilenga; 700 m a.s.l.; 15 Sep.–6 Nov. 1947; Mission G.F. de Witte leg.; RMCA. – Lualaba • 1 ♂; Kafakumba; Mar. 1929; NHMUK • 1 ♀; Kangagi [= Kangaji]; Jan. 1927; F.G. Overlaet leg.; RMCA • 1 ♂; Kolwezi; Jan. 1953; V. Allard leg.; RMCA. MALAWI – Southern • 1 ♂; between Fort Mangoche and Chikala Boma; 1219 m a.s.l.; 21 Mar. 1910; S.A. Neave leg.; NHMUK. TANZANIA – Rukwa • 1 ♂; Kalambo Forest Reserve; 8°21.596′ S, 31°15.221′ E; 1632 m a.s.l.; 22 Nov. 2009; P. Darge leg.; ANHRT. ZAMBIA – Central • 1 ♂; Kasanka River Pontoon, Kasanka National Park; 12°34′23″ S, 30°14′05″ E; 1191 m a.s.l.; 2–4 Dec. 2012; R. Smith and H. Takano leg.; ANHRT. – Copperbelt • 1 ♀; Mwengwa; 23 Feb. 1914; H.C. Dollman leg.; NHMUK. – Luapula • 1 ♂; Mumbuluma Falls; 1190 m; 12 Jan. 2018; P. Schmit leg.; RCPB. – Muchinga • 3 ♂♂; Benyanga village; 10°40′41″ S, 33°27′45″ E; 1250 m a.s.l.; 7–12 Dec. 2023; G. László, L. Morgan and A. Volynkin leg.; ANHRT • 1 ♂; Kalungu; 9°40′52″ S, 32°42′50″ E; 1280 m a.s.l.; 22–24 Nov. 2016; R. Smith, H. Takano and D. Oram leg.; ANHRT • 6 ♂♂; same locality as for preceding; 5–8 Mar. 2017; D. Oram, W. Miles and L. Smith leg.; ANHRT • 3 ♂♂; same data as for preceding; 11 Mar. 2017; ANHRT • 1 ♂; Mama Muwowo’s Lodge, Muyombe; 10°32′40″ S, 33°26′05″ E; 1230 m a.s.l.; 29–30 Nov. 2023; M. Bashford, A. Collins, G. László, L. Morgan and A. Volynkin leg.; ANHRT • 3 ♂♂; same locality as for preceding; 6–7 Dec. 2023; G. László, L. Morgan and A. Volynkin leg.; ANHRT • 1 ♀; Mpika; Nov. 1950; H.B.D. Kettlewell leg.; NHMUK. – Northwestern • 1 ♂, 1 ♀; Chitunta Plain; 11°29′12″ S, 24°24′18″ E; 1396 m a.s.l.; 29 Nov.–4 Dec. 2019; M. Bashford, W. Miles and L. Mulvaney leg.; ANHRT • 1 ♂; dambo near Kanyama; 11°25′36″ S, 24°40′00″ E; 1375 m a.s.l.; 4–7 Dec. 2019; M. Bashford, W. Miles and L. Mulvaney leg.; ANHRT • 2 ♂♂; Hillwood, Ikelenge; 11°16′02″ S, 24°18′59″ E; 1400 m a.s.l.; 23–30 Nov. 2019; M. Bashford, W. Miles, L. Mulvaney and R. Smith leg.; ANHRT • 1 ♂; Jiwundu Swamp; 11°51′54″ S, 25°33′20″ E; 1340 m a.s.l.; 25–30 Oct. 2017; M. Carter, A. Lloyd, W. Miles, D. Oram and R. Smith leg.; ANHRT • 1 ♂; same locality as for preceding; 29 Oct.–4 Nov. 2018; M. Aristophanous, V. Dérozier, G. László and D. Oram leg.; ANHRT • 1 ♂; Kambishi, Jiwundu; 11°54′39″ S, 25°29′05″ E; 1340 m a.s.l.; 18–19 Oct. 2014; R. Smith, H. Takano and D. Oram leg.; ANHRT • 1 ♂; Lukwakwa, West Lunga National Park; 12°39′40″ S, 24°26′13″ E; 1147 m a.s.l.; 4–8 Nov. 2013; R. Smith, H. Takano and D. Oram leg.; ANHRT
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 157 • 2 ♂♂; same locality as for preceding; 9–15 Nov. 2018; M. Aristophanous, V. Dérozier, G. László and D. Oram leg.; ANHRT • 1 ♂; Nkwaji, Mwinilunga; 11°36′22″ S, 24°33′17″ E; 1316 m a.s.l.; 29 Oct.–3 Nov. 2013; R. Smith and H. Takano leg.; ANHRT • 1 ♂; same locality as for preceding; 3–10 Nov. 2017; M. Carter, A. Lloyd, W. Miles, D. Oram and R. Smith leg.; ANHRT • 1 ♂; Solwezi; 2 Nov. 1917; H.C. Dollman leg.; NHMUK • 1 ♀; Zambezi Rapids; 11°07′30″ S, 24°11′06″ E; 1205 m a.s.l.; 4–9 Nov. 2018; M. Aristophanous, V. Dérozier, G. László and D. Oram leg.; ANHRT. – Southern • 7 ♂♂; Bruce-Miller Farm, Choma; 16°38′12″ S, 27°01′30″ E; 1179 m a.s.l.; 28 Feb.–8 Mar. 2019; Dérozier, M. Imakando, W. Miles and L. Mulvaney leg.; ANHRT • 1 ♂; Kalomo; C.W. Brames Hall leg.; NHMUK • 3 ♂♂; Musuku Lodge, Bruce-Miller Farm, Choma; 16°38′44.4″ S, 27°03′32.6″ E; 1227 m a.s.l.; 28 Feb.–8 Mar. 2019; V. Dérozier, M. Imakando, W. Miles and L. Mulvaney leg.; ANHRT. – Western • 1 ♂; 30 km S of Mongu; 15°36.78′ S, 23°19.13′ E; 4 Dec. 2017; R. Minetti leg.; ANHRT • 2 ♂♂; Ndanda; 15°04′44″ S, 23°45′59″ E; 1090 m a.s.l.; 10–11 Nov. 2013; R. Smith, H. Takano and D. Oram leg.; ANHRT. ZIMBABWE • 1 ♂; Mashonaland; H.B. Dobbie leg.; NHMUK. – Harare • 1 ♀; Salisbury [= Harare]; Dec. 1898; G.A.K. Marshall leg.; NHMUK. Redescription Male (Figs 22–25, 52) Forewing length. 30–35 mm. Upperside. Ground colour of head, thorax, forewings and abdomen beige; vertex dark greyish-brown, patagium slightly golden, hindwing yellow. Antenna bipectinate, dark brown. Eighth sternite truncate posteriorly, weakly sclerotised; its surface covered evenly in fine punctures with scattering of deeper punctures. Forewing triangular, rounded at apex, outer margin gently arcuate; costa with golden scaling. Antemedial fascia indicated by dark brown dots along veins, sinuate, poorly defined, arising at cubital and terminating along anal margin. Discal marking black, double, one on either side of medial vein, inner smaller than outer; outer sometimes split in half by vein M2. Postmedial fascia dark brown, arcuate, with darker dots along veins; fascia arising perpendicularly just over halfway along costa and gently curved inwards at vein M1, terminating perpendicularly to anal margin. Submarginal fascia indicated by dark brown dots along veins, running in line with the postmedial fascia. Fringe pale beige irrorated with golden scales. hindwing. Outer margin arcuate, slightly angled at vein M2. Postmedial and submarginal fasciae dark brown, arcuate, showing through from the underside. Fringe pale beige irrorated with golden scales. Underside. Legs and to lesser degree abdomen densely covered in long carmine ciliate scales, giving different appearance to upperside. Ground colour of wings golden with ferruginous tinge. Postmedial fascia dark brown, fine, placed slightly distad of and straight than that on upperside. Submarginal fascia dark brown, fine, placed slightly distad of and more arcuate than that on upperside. Hindwing veins well-defined with dark brown scaling. Postmedial and submarginal fasciae dark brown, arcuate, almost running parallel to each other, inner of two slightly broader, the outer gently crenulate. Subterminal fascia dark brown, sinuate, indistinct or absent in some individuals. M ale genitalia (Figs 44–45). Uncus reduced, fused with tegumen. Tegumen broad with pair of long, cylindrical, apically pointed lateral projections, gently curved inward; their inner surface weakly setose. Gnathos with sparse cluster of spines of varying length, either side of midline. Valve triangular and cleft (about one-third of way along valve). Costa curved ventrad at apex, tapering to blunt point. Sacculus well-defined with curved distal process, rounded at apex. Juxta trapezoid, proximally V-shaped, lateral margins tapering cranially, distally with deep U-shaped concavity. Vinculum V-shaped. Saccus
European Journal of Taxonomy 1022: 134–175 (2025) 158 cylindrical, rounded at apex. Phallus as long as valve, gently curved medially; coecum rounded. Vesica with scobination. Female (Figs 26–27) Forewing length. 34–37 mm. Similar to male with no difference in rami length of the antenna. Some individuals with a salmon hue to the hindwing upperside. Figs 22–27. Camerunia bimaculata (Dewitz, 1879) comb. rev., adults. 22. Holotype of Dreata bimaculata Dewitz, 1879, ♂ (MfN). 23–25. ♂♂, Zambia, Kalungu (ANHRT). 26. ♀, Zambia, Zambezi Rapids (ANHRT). 27. ♀, Zambia, Chitunta Plain (ANHRT).
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 165 at some stage and is instead pinned to the holotype of Poloma variegata (Rothschild, 1917). This taxon is conspecific with Poloma nigromaculata (Aurivillius, 1893) as determined by Gaede and is herein synonymised with Aurivillius’s taxon; thus Hemijana griseola Rothschild, 1917 syn. nov. = Poloma nigromaculata (Aurivillius, 1893). See Takano & Morgan (2025) for further figures of this species. Genus composition Camerunia orphne (Schaus, 1893) = Camerunia insignis Aurivillius, 1893 Cameruniaflava Aurivillius, 1904 = Camerunialactiflora Wichgraf, 1914 Camerunia albida Aurivillius, 1901 Camerunia smithi sp. nov. Camerunia bimaculata (Dewitz, 1879) comb. rev. Camerunia subrosea (Aurivillius, 1893) comb. nov. = Hemijana ruberrima Rothschild, 1917 Identification key to the species of Camerunia Aurivillius, 1893 1. Forewing with dark basal marking ................................................................................................... 2 – Forewing without dark basal marking .............................................................................................. 3 2. Forewing discal marking as wide as cell. Dark spot along forewing postmedial fasciae in space CuA1 diffuse. Hindwing fringe brown with white markings at the termination of veins. Large species found in West and Central Africa .........................................................................C. orphne (Schaus, 1893) – Forewing discal marking indicated by two or three spots. Dark spot along forewing postmedial fasciae in space CuA1 well defined. Hindwing fringe uniformly pale yellow. Small species endemic to Tanzania .............................................................................................................. C. smithi sp. nov. 3. Ground colour of hindwings yellow ................................................................................................. 4 – Ground colour of hindwings not yellow ........................................................................................... 5 4. Forewing with large discal marking ................................. C. bimaculata (Dewitz, 1879) comb. rev. – Forewing without discal marking .............................................................. C. flava Aurivillius, 1904 5. Ground colour of hindwings white .......................................................... C. albida Aurivillius, 1901 – Ground colour of hindwings pink .................................. C. subrosea (Aurivillius, 1893) comb. nov. Discussion As a result of taxonomic analyses integrating morphological and genetic traits, the genus Camerunia is comprehensively revised and its generic boundary has been delimited. Two allied genera, Hemijana and Catajana, both described based mainly on differences in wing venation, are considered to be junior subjective synonyms of Camerunia. The genus now contains six species, the three species originally belonging to Camerunia, the type species of Hemijana and Catajana, and the new species described above. Based on the genital morphology, the six species fall into two groups, one with flattened lateral processes of the tegumen (C. orphne, C.flava and C. albida) and another with cylindrical processes (C. smithi sp. nov., C. bimaculata (Dewitz, 1879) comb. rev. and C. subrosea (Aurivillius, 1893) comb. nov.). However, the results of the barcode analyses were discordant with this observation and moreover, when mapped onto the phylogram, species grouped on other perceived character sets (e.g., wing pattern or
European Journal of Taxonomy 1022: 134–175 (2025) 166 Figs 37–41. Camerunia Aurivillius, 1893 species, male genitalia, clasping apparatus (above), phallus (below). 37–38. C. orphne (Schaus, 1893). 37. Sierra Leone (HT 025). 38. Gabon (HT 029). 39– 40. C.flavaAurivillius, 1904. 39. Zambia (HT 011). 40. Cameroon (HT 010). 41. C. albida Aurivillius, 1901, Tanzania, holotype.
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 167 Figs 42–47. Camerunia Aurivillius, 1893 species, male genitalia, clasping apparatus (above), phallus (below). 42. C. smithi sp. nov., Tanzania, holotype (HT 086). 43. C. smithi sp. nov., Tanzania, paratype (HT 015). 44–45. C. bimaculata (Dewitz, 1879 comb. rev., Zambia. 44. HT 087. 45. HT 027. 46– 47. C. subrosea (Aurivillius, 1893). 46. South Africa (HT 028). 47. Zambia (HT 030).
European Journal of Taxonomy 1022: 134–175 (2025) 168 ciliate scale tufts of the abdomen) were equally incongruent. It must be noted that the genetic analyses were based on a small fragment of the mitochondrial gene and taxon sampling was incomplete. Further analyses utilising other mitochondrial and nuclear markers, together with the inclusion of C. albida, an isolated species with unique characters in the male genitalia, are required to clarify relationships among the taxa. Although members of this genus are widely distributed, two of the six species, C. albida and C. smithi sp. nov., are narrow-range species known only from their type localities in East Africa, and the type species of the genus, C. orphne, is found mainly in the Guinean forests and their transition zones. The distribution of the remaining three species, C.flava, C. bimaculata (Dewitz, 1879 comb. rev. and C. subrosea (Aurivillius, 1893) comb. nov. are centred around the miombo woodlands of southern Central Africa. Although their modern-day ranges broadly overlap, it is interesting to note that in the south, C.flavadoes not appear to cross the Zambezi River, but C. bimaculata and C. subrosea reach South Africa, and in the north C. subrosea and C.flava cross the Rukwa Rift into Tanzania, whereas C. bimaculata does not. Over 400 specimens were examined as part of this revisionary work and, as reported in other groups of Janinae (Takano 2024b), female specimens were found to be uncommon in collections, constituting only one in eight specimens (and unknown in two species). Females of Camerunia are infrequently attracted to light but when they are, large numbers of males often congregate as observed on at least two occasions in northern Zambia by the present author. Furthermore, over three-quarters of the material examined was collected in the past 20 years, indicating the general scarcity of specimens in historical collections and Figs 48–53. Camerunia Aurivillius, 1893 species, male eighth abdominal sternite. 48. C. orphne (Schaus, 1893) (HT 029). 49. C.flavaAurivillius, 1904 (HT 011). 50. C. albida Aurivillius, 1901, holotype (ANHRT). 51. C. smithi sp. nov. holotype (HT 086). 52. C. bimaculata (Dewitz, 1879) comb. rev. (HT 087). 53. C. subrosea (Aurivillius, 1893) (HT 028).
TAKANO H., Taxonomic revision of Camerunia Aurivillius, 1893 and allied genera (Lepidoptera) 169 the need for careful and considerate faunistic assessments in an era when biodiversity conservation is paramount. The discovery of a highly distinct new species of Camerunia from a small forest fragment in the Eastern Arc Mountains of Tanzania highlights once more the high levels of endemism in this extremely biodiverse region and reinforces the importance in undertaking further biological inventories, especially for nocturnal insects, and protecting what remains of these extraordinary ecosystems (Takano 2022). Figs 54–55. Distribution maps of Camerunia Aurivillius, 1893 species. 54. C. orphne (Schaus, 1893). 55. C.flavaAurivillius, 1904.
European Journal of Taxonomy 1022: 134–175 (2025) 170 The synonymy of two genera with Camerunia leaves 20 genera in Janinae, some remaining in a state of disarray (e.g., Jana Herrich-Schäffer, 1854). But despite their large size and beautiful appearance, natural groupings are unclear due to the extreme plasticity of the genitalia (Forbes 1955) and paraphyletic assemblages, such as the “Ganisa-group” proposed by Oberprieler et al. (2003), have arisen as a result. Moreover, the DNA barcode region of Janinae is highly variable in some genera (e.g., Camerunia) Figs 56–57. Distribution maps of Camerunia Aurivillius, 1893 species. 56. C. bimaculata (Dewitz, 1879) comb. rev. (red circles), C. smithi sp. nov. (blue triangle) and C. albida Aurivillius, 1901 (yellow star). 57. C. subrosea (Aurivillius, 1893) (open circles represent records from iNaturalist).
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