Platyja Hübner, [1823]. In 1816–[1825], Verzeichniss bekannter Schmettlinge [sic] (17): 268. Type species: Phalaena Noctua umminia Cramer, [1780], by subsequent designation of Moore ([1885] in 1884–1887).

Synonymy

See Zilli et al. (2021), to which the following generic synonyms must be added:

=Facidia Walker, 1865 syn. n.

=Megacephalomana Strand, 1943 syn. n.

The former is however retained at subgeneric level by virtue of ICZN (1999: art. 61.3.1)—see below.

Diagnosis

See Holloway (2005), Zilli & De Vos (2021), and Zilli et al. (2021).

Facidia Walker, 1865. List of the Specimens of lepidopterous insects in the collection of the British Museum XXXIII (Supplement 3): 951. Type species: Facidia nigrofusca Walker, 1865, by monotypy. Currently considered as a junior subjective synonym of Tavia? vacillans Walker, 1858.

Synonymy

=Megacephalomana Strand, 1943 syn. n. Folia zoologica et hydrobiologica 12 (1): 211. An objective replacement name for Megacephalon Saalmüller, 1880, Bericht über die Senckenbergische naturforschende Gesellschaft in Frankfurt am Main 1879–1880: 286. A junior primary homonym of Megacephalon ‘Temminck, 1844’ (recte Gray 1846 in 1844–1849: [489]) [Aves]. Type species: Megacephalon rivulosum Saalmüller, 1880, by monotypy.

Diagnosis

Species of Platyja of the subgenus Facidia are best characterised by features of the male genitalia, notably the vinculum arms cojoining ventrally and bending inward, forming a long, narrow saccus, the portion of the diaphragma closing the tegumen with paired, thin, arched rods facing the periphallic membrane laterally at its base (such rods possibly being reduced, modified transtillae that lost their junction with the valvae), the small juxta consisting of two symmetrical, subtriangular plates separated by a narrow membrane, and the fairly simple, compact, saccate vesica with main trigonal-trilobate corpus sometimes with sclerotised plates or scobination but no nail-like cornuti. In the female abdomen, the last pre-genital segment (A7) has sternum not modified into a special lodix plate and the bursa copulatrix is consistently ovoid across all known taxa.

Distribution

All Sub-Saharan Africa from Western to Southern and Eastern Africa, eastwards to the Madagascan subregion.

Remarks

Examination of the type species of Facidia and Megacephalomana and of all African species currently attributed to these genera has shown that there is a core group of species whose separation at the generic level is unsubstantiated and which are fully compatible with the broader concept of Platyja, as a homogeneous subset within this genus. For this reason, Facidia and Megacephalomana are subsumed under Platyja at subgeneric level under the name taking priority. However, there are two outlier species that, despite sharing the groundplan features with the revised concept of Facidia proposed here, essentially stand out because of some remarkable autapomorphies. They will be discussed at the end of the taxonomic part.

As noted above, in the past, some taxa of this group have also been attributed to Facidina Hampson, 1926 (type species: Ischyja polystigma Lower, 1903) (e.g., Gaede 1939), but the concept of this genus is nowadays centred around a group of Australo-Papuan species corresponding in overall appearance to the type species (Edwards 1996; Holloway 2005). With the exception of F. spilophracta (Turner, 1933), originally described by Turner (1933: 165) as Erceia [sic] spilophracta and still to be properly morphologically analysed but possibly belonging to Platyja, typical Facidina are deprived of the characteristic Platyja pattern—as defined in Zilli et al. 2021—and sport a normally shaped postmedial line of the forewing, filiform male antennae and, in the male genitalia, a basally very long valva, articulated with a similarly long vinculum, that terminates with long processes, a strongly modified, very long juxta, and a very long and thin phallus sigmoid in shape.

Tavia? vacillans Walker, 1858. List of the Specimens of lepidopterous insects in the collection of the British Museum XV: 1820. Type material: Holotypus ♀, by monotypy, in NHMUK [examined]. Type locality: Africa [verbatim holotype label: W - Africa].

=Facidia nigrofusca Walker, 1865. List of the Specimens of lepidopterous insects in the collection of the British Museum XXXIII (Supplement 3): 952. Type material: Holotypus ♂, by monotypy, in NHMUK [examined]. Type locality: Natal [verbatim holotype label: Pt Natal].

=Megacephalon fenestratum Möschler, 1887. Abhandlungen herausgegebenvonderSenckenbergischenNaturforschenden Gesellschaft 15 (1): 84. Type material: Holotypus ♀, by monotypy, in MFNB [picture examined]. Type locality: Accra.

=Facidia horrida Holland, 1894. Psyche, a Journal of Entomology 7 (217): pl. 2, fig. 6; (218/221): 143. Type material: Syntypi ♂♂ (number not given), in CMNH, NHMUK [1 syntypus (NHMUK) examined]. Type locality: West Africa [verbatim syntypus label: Ogove River].

=Platyja phaeophoenica Hampson, 1905. Annals of the South African Museum 3 (9): 434. Type material: Holotypus ♀, by original designation, in NHMUK [examined]. Type locality: Natal, Durban.

=Facidia sassana Strand, 1918 syn. rev. Zeitschrift des Österreichischen Entomologen-Vereines 3: 100. Type material: Holotypus ♂, by monotypy, in MRAC [picture examined]. Type locality: [Cameroon] Sassagebiet [= Région de Sassa].

Diagnosis

A very variable species whose males differ in the different degree of darkening of the ground colour, from chocolate-brown to nearly black, and show partially clinal variation in the expression of the Platyja pattern, generally indistinct in western populations and more evident in southeastern ones. The females have broader wings than the males and more regularly show the typical Platyja pattern. The African species most closely resembling P. (F.) vacillans are P. (F.) saalmuelleri comb. n. and P. (F.) ennomoides sp. n.

Besides allopatry, vacillans and saalmuelleri can be separated by the less pronounced sexual dimorphism in the wing shape of the latter, whose males have broader wings, especially the hind one, which is also less convex at the termen and more angular at the tornus, and the generally better expressed Platyja pattern. This last feature shows geographic congruence, whereby vacillans populations with the most emphasized pattern are those in closer geographic proximity to those of saalmuelleri. As regards the leg tufting, this is more strongly emphasised in males of the Madagascan species, especially in the easily examinable hindlegs. Females of the two species are virtually indistinguishable based on external morphology, though a clue may be found again in the more shallowly convex hindwing of the insular taxon. In the male genitalia, the differences between vacillans and saalmuelleri are weak, which together with the similarity in habitus provides clear evidence of their close relationship. Male vacillans have (1) valva more elongate with a slenderer apical expansion at the anal angle that confers a ‘foot-shape’ to the cucullus, contrary to the broader valva with shorter and subtriangular expansion in the Madagascan taxon, (2) sacculus triangular with distinct saccular corner, not subrectangular with the anterior edge parallel to the ventral margin of valva, (3) distal opening of phallus with a continuous row of very small, adpressed spines along the right side (vs. fewer, erect, much stouter apical spines in saalmuelleri), (4) vesica with shorter, forward-projecting diverticulum flanked internally by a rather large, conical diverticulum, (5) big, backward-oriented diverticulum has a weaker, less expanded scobination at base internally, and (6) laterally projected diverticulum with smaller sclerotised plate (Figs. 46–48, 58–60). In the female genitalia, the main differences occur in the ductus bursae and bursa copulatrix. Compared to the short, lyra-shaped ductus bursae of saalmuelleri (Fig. 70), that of vacillans is longer and with much less sinuous sides (Fig. 69), whereas the bursa of the latter is distinctly smaller and, with the exception of an irregular plate-like posterior signum, almost entirely membranous, contrary to the more broadly developed and more heavily sclerotised cervix bursae of the Madagascan taxon.

With respect to ennomoides sp. n., whose female is still unknown, male vacillans (and saalmuelleri) can easily be recognised by the less heavily pectinated antennae, brown- instead of white-scaled dorsally on the flagellum shaft, and the evenly curved outer margin of the forewing, while in the male genitalia the dorsal margin of the valva is smoothly concave, without a short, digitiform costal process.

Distribution

Widespread in Sub-Saharan Africa; Gambia, Guinea, Liberia, Sierra Leone, Ivory Coast, Ghana, Benin, Nigeria, São Tomé, Príncipe, Cameroon, Gabon, Congo (E), Kenya, Tanzania, Malawi, Zambia, Zimbabwe and South Africa.

Remarks

Facidia sassana, correctly treated by Gaede (1939) as a taxon subordinate to vacillans (misspelt as vaccillans), was misplaced by POOLE (1989) as a valid species in Falcimala Hampson, 1895, a genus of the subfamily Herminiinae.

Platyja (Facidia) vacillans and P. (F.) saalmuelleri comb. n. constitute a pair of allopatric sister species, respectively distributed in continental Africa (and islands in the Gulf of Guinea) and in the Madagascan subregion.

During the course of this study, it became clear that in W Africa there is a male morph partially sympatric with the commonest African one, which differs from the latter by features of the wings' underside. This morph, illustrated in Figs. 16 and 17, lacks the black suffusion that in typical vacillans extends over at least the discal area of the forewing underside (Fig. 18) and is here patterned as a female; it is possibly also more scalloped at the wing termens. Overall, five specimens of this morph have been examined, from Gambia, Guinea (Ditinn: Chute de Ditinn; Dalaba: Forêt de Tinka), and Sierra Leone (Northern Province: nr Kamabai, Kalainkay), the last two countries at least also hosting normally patterned males. Feeble differences in the genitalia of the specimen from Gambia (Figs. 45, 57) prompted dissections of other males of the female-like form and sympatric females to assess whether it might belong to a W African cryptic species close to vacillans, but this hypothesis could not be in any way morphologically substantiated. Accordingly, the presence of these morphs in W Africa is here provisionally inferred to be a case of male-limited dimorphism, in which the commoner and more widespread African morph with a black underside is exclusive to the male sex.

Cases of di- or polymorphic males with monomorphic females seem to be very rare in Lepidoptera. In a review of polymorphism in butterflies, Vane-Wright (1975) suggested only one likely case, that of the Neotropical satyrine nymphalid Manataria maculata (Hopffer, 1874) (Lepidoptera: Nymphalidae), currently ranked as a subspecies of Manataria hercyna (Hübner, [1821]), but called for further insights into this possibly doubtful example. The same author (R. I. Vane-Wright, pers. comm.) drew our attention to the oriental limenitidine nymphalid Euthalia monina (Fabricius, 1787) (Lepidoptera: Nymphalidae), whose males occur in at least three morphs, “although intergrades are not rare” (Corbet et al. 2020), and whose females do not vary. Last but not least, a well-known case is that of polymorphic male populations of Hepialus humuli (Linnaeus, 1758) (Lepidoptera: Hepialidae) in the Shetland and Faroe Islands, also in this case with some transitional phenotypes (Kaaber et al. 2009). Admittedly, Gerould (1923) generally mentioned cases of male dimorphism in Lycaenidae and Arctiidae (now Erebidae: Arctiinae), but his statement is difficult to put in proper context as he did not specify whether only the male sex was dimorphic, and there are endless examples of Lepidoptera species whose males are as polymorphic as the females.

At a first glance, characters shown only by one sex seem ascribable to sex-linked inheritance, that is to the presence on the sexual chromosomes (viz. allosomes) of genes responsible for such traits. However, the fact that Lepidoptera have a ZW sex-determination system and that females are heterogametic does not support this scenario in vacillans, because if the gene responsible was on the Z chromosome the females should show that character at the same phenotypic frequency in the male, given that sperm will be produced at the same ratio and females bear only one Z chromosome (obviously, the gene cannot be on the W chromosome, as otherwise the trait would occur only in females and be unknown in males). However, females putatively bearing this gene would not develop the dark underside if there was another gene suppressing its expression in that sex. It is thus the suppressor gene that could be sex-linked if hosted by the W chromosome, regardless of the position of the gene for the dark phenotype on the Z allosome or any somatic chromosomes (autosomes). Nevertheless, the suppressor might also be located on autosomes and act as a switch on some regulatory mechanisms that prevent the expression of the dark phenotype whenever it happens to operate in a female genetic environment.

Remarks

A male singleton originally present in the collection of Charles Oberthür (now NHMUK), but unfortunately without locality data, matches with Platyja (Facidia) vacillans in many respects, but differs by its larger size, slightly longer antennal pectinations, valva less slender at middle, thinner phallus, and less prominent backward-projecting lobe of vesica (Figs. 49, 61). Although there are no sufficient elements to comfortably proceed with the description of a new taxon, attention is simply drawn to this outlier specimen until new material clarifying its relation to vacillans is found.

Megacephalon saalmülleri Viette, 1965. Lambillionea 64 (9/10): 41, pl. 1, fig. 7. Type material: Holotypus ♂, by original designation, in MNHN [picture examined]. Type locality: Madagascar Est, environs de Perinet, forêt d'Analamazaotra, 910 m [verbatim holotype label: forêt d'Analamazoatra].

Diagnosis

Details for distinguishing this species from its close relative Platyja (Facidia) vacillans comb. n. are given above under that species. Male genitalia as in Figs. 50 and 62, female genitalia as in Fig. 70.

Distribution

Endemic to Madagascar and surrounding islands (Nosy Boraha).

Remarks

As noted above, this species and Platyja (Facidia) vacillans comb. n. are closely related species distributed, respectively, in the Madagascan subregion and mainland Africa (also São Tomé and Príncipe islands). The two taxa are indeed very close, and it would be somewhat surprising that saalmuelleri was described as a distinct species if not for the fact that Viette (1965) did not take vacillans into account at all and described the Madagascan taxon only comparing it against the very different, sympatric Platyja (Facidia) rivulosum comb. n. With careful analysis, however, saalmuelleri and vacillans can be distinguished (see Remarks under vacillans), and that the taxonomic rank of saalmuelleri as a valid species is retained.

Alamis semifimbria Walker, [1858]. List of the Specimens of lepidopterous insects in the collection of the British Museum XIII: 1048. Type material: Holotypus ♀, by monotypy, in NHMUK [examined]. Type locality: Sierra Leone.

Diagnosis

Easily recognisable by the strongly variegated pattern with conspicuously undulated crosslines, dots and other elements highlighted with silvery grey and by the lack of secondary sexual dimorphism in the pattern of the moths. The only other African species with which it may be confused is Platyja (Facidia) rivulosum comb. n., whose males have more oblong-elongated wings and a more shallowly convex outer margin of the hindwing, besides being restricted to the Madagascan subregion and hence allopatric. Females of semifimbria and rivulosum are very similar, though those of the Madagascan taxon show more oblong wings and a straighter postmedial line of the hindwing. In any case, the best diagnostic hint for distinguishing the two species in both sexes is the spatial relationships between the outer lobe of the median field and the submarginal line of the forewing. In rivulosum the lobe is more outwardly produced and the submarginal line starts from the costa before the apex, so the postmedial line bounding the lobe and the submarginal line do actually overlap, the latter often being visible as venular dots between the lobes of the undulated postmedial. In contrast, in semifimbria the lobe is less prominent and the submarginal line more distal, starting from the wing apex; as a result, the two lines (postmedial and submarginal) do not coincide and there is some space between them.

In the male genitalia, the valva of semifimbria has a broader sacculus compared to rivulosum, with a conspicuous hump at the saccular corner; it also is narrowest at the middle and has a bigger distal superior process, whereas in the vesica the relative development of the three main lobes is totally different (Figs. 51, 63). In the female genitalia, the most evident diagnostic feature is that of the antrum-ductus bursae system, shorter and wider in semifimbria (Fig. 71).

Distribution

Widespread in Sub-Saharan Africa; Sierra Leone, Liberia, Ivory Coast, Ghana, Kenya, Tanzania, Mozambique, Zambia, Zimbabwe, and South Africa.

Remarks

This species was treated by Gaede (1939) and Poole (1989) as belonging in Facidina. Holloway (2005) correctly recognised its similarities with Platyja and moved it to this genus, though he did not treat the taxonomic position of the other close African relatives of semifimbria.

Megacephalon rivulosum Saalmüller, 1880. Bericht über die Senckenbergische naturforschende Gesellschaft in Frankfurt am Main 1879–1880: 287. Type material: Syntypi ♂♂(by implication from original description but number not given), 1♀; traced 2♂♂ and 1♀ in SMF [examined]. It should be noted that specimens in SMF labelled as “Typus” (1♂, Loucoubè, Stumpff 82/508; also illustrated by DE PRINS & De Prins 2011–2021) and “Paratypoid” (1♂, sine data/508a; 1♀, Nossi-Bé Stumpff e[nde] 89/508b; 2♂♂, Loucoubè, STUMPFF 82/508c & 508d) [all examined] are all posterior to the original description and are therefore not part of the type series. In contrast, of another series of three specimens, labelled respectively “Loucoubé, Stumpff80” [hand corrected into 82] (1♂), “Loucoubé, Ebenau 80” (1♂), and “Loucoubé Stumpff80” (1♀) [all examined], the last two are fully compatible with information given by Saalmüller (1880) and are therefore regarded as syntypes. The first male is also considered to be a syntype as there is evidence that the correction of the year from [18]80 to [18]82 is subsequent to the original labelling of the specimen and hence spurious. In fact, until 1880 the labels recording the collecting locality by Stumpff were printed with the acute accent (Loucoubé), while later the grave accent was used (Loucoubè) [see above], and such specimen clearly bears the first kind of labels. The apparent discrepancy in the original locality, given by Saalmüller (1880: 258, 1891: 449) as Nossi-Bé, while the specimens are labelled as from Loucoubé [= Lokobe], is simply ascribable to the fact that the latter is a precise site in the island of Nossi-Bé. Type locality: Madagascar, Nossi-Bé [= Nosy Be] [verbatim syntypes labels given here above].

Diagnosis

Hints for distinguishing this species from the externally similar Platyja (Facidia) semifimbria have been given under that species. In the male genitalia, the Madagascan taxon can be distinguished from its close relative by the shape of valva, of uniform width along its middle section, with slenderer sacculus and smaller distal superior process (Fig. 52), while the size and configuration of the vesical lobes are different (cf. Fig. 64). In the female genitalia, the antrum-ductus bursae system of rivulosum is appreciably longer and thinner (Fig. 72).

Distribution

Madagascar and surrounding islands (Grande Comore, Nosy Be).

Remarks

This species replaces Platyja (Facidia) semifimbria in the Madagascan subregion. Both species closely resemble in pattern some Indo-Australian members of the subgenus Mocrendes Nye, 1975 of Platyja, namely those of the lemur-lemurella-carpentariella lineage (see Zilli et al. 2021), but analysis of structural features shows that they are not directly related. Another similar-looking Australian species is Facidina spilophracta (Turner, 1933), which still needs to be properly morphologically analysed.

Type material

Holotypus: ♂, Tanzania, Mal[ariological] Inst[itute] Amani, III.[19]64; NHMUK014165500, in NHMUK.

Paratypi (2♂♂): 1♂, Tanzania, Amani, Usambara, IV.1969, Wathlege [leg.?]; NHMUK014165496. – 1♂, [Zimbabwe] South Rhodesia, Vumba Mts, Umtali, III–IV.1963, [ex] Nat. Mus. S[outhern] R[hodesia]; NHMUK0141654951. Both in NHMUK.

Diagnosis

The new species may easily be mistaken for some homogeneously coloured forms of the sympatric Platyja (Facidia) vacillans comb. n., less so with the allopatric sister species of the latter, P. (F.) saalmuelleri comb. n., but its male can be distinguished from these by the more heavily pectinated antennae with dorsal white scaling on the flagellum, the outer margin of the forewing (termen) not smoothly convex but feebly angular at middle, and the paler hindwing, not as dark as the forewing. In the male genitalia, the presence of a thumb-like costal process on the valva, the superiorly narrowly elongated juxta, and the longer phallus with different vesical configuration (Figs. 53, 65) allow an easy separation of the new species from the other two.

Description

Male. Wingspan 44.0–47.5 mm apex to apex but forewings widest at middle of termen, up to 49 mm (n = 3). Head very large, dark chocolate-brown-coloured, frons narrow and bulged, vertex thickly and roughly scaled, eye globular, antenna conspicuously pectinated except at very tip, with very long lateral rami and long, reclinate ventral lamellae, with dorsal scaling of flagellum almost pure white, labial palpus compactly scaled with stout, suberect 2nd joint and rod-like 3rd joint, straw-coloured at very tip; haustellum present.

Thorax uniformly dark chocolate-brown, with patagium, tegula, and notum thickly and roughly-scaled; forewing broad, with slightly prominent apex and convex termen almost angular at middle, ground colour uniformly dark chocolate-brown, with faint or indistinct Platyja pattern, antemedial line faded, with a few minute silvery dots in correspondence with veins, postmedial line slightly better expressed, feebly darker than ground colour and also with some silvery dotting across veins, orbicular a tiny dark brown dot, reniform crescent-shaped, outer section of median field very narrow, with very small, dull grey-filled pedunculate lobe; adterminal area and fringe concolorous with ground colour; hindwing rounded, chocolate-brown, paler at base; legs conspicuously hairy except tarsi; tarsomeres appearing pale beige ringed at distal ends. Underside of wings brown with minute pinkish-white irroration, postmedial line of forewing almost indistinct, that of hindwing feebly expressed as a darker brown line followed by tiny whitish dots in correspondence with veins.

Abdomen uniformly brown. Male genitalia: tegumen voluminous, dome-shaped, vinculum long and robust, longer than tegumen, valva long and sinuous, with a comparatively small, narrowly elongated subtriangular sacculus, broad, long, slightly excurved apical expansion and small, thumb-like costal process; juxta sub-rhomboid basally, very narrow and elongated apically; uncus comparatively short and thick, with prominent recurved apical hook; tuba analis large, with distinctly sclerotised scaphium; phallus long and slender, slightly curved at middle, vesica as in Fig. 65.

Female. Unknown.

Etymology

The name of the new species is based on the external resemblance of the male with numerous geometrid moths of the subfamily Ennominae. The name is meant as a feminine adjective.

Distribution

So far known only from south-eastern Africa (Tanzania and Zimbabwe).

Remarks

The particularly smooth, homogeneous colouring of the forewings and the shared presence of a costal process on the valvae are suggestive of a systematic relatedness between Platyja (Facidia) ennomoides sp. n. and P. (F.) stygium (Saalmüller, 1881) comb. n.

Megacephalon stygium Saalmüller, 1881. Entomologische Zeitung (Stettin) 42 (4/6): 217. Type material: Holotypus ♂, by (likely) monotypy, in SMF [examined]. Type locality. Madagascar... Nossi-Bé [verbatim holotype label: Loucoubé. See explanation above under P. (F.) rivulosum].

Diagnosis

An unmistakable species, easily distinguishable from all relatives, besides other characters, by the white-coloured basal area of the hindwing in both sexes. Notably, the hindwing has a nearly straight termen in the male. Genitalia as in Figs. 54, 66, and 73.

Distribution

Endemic to Madagascar.

Remarks

Together with Platyja (Facidia) ennomoides sp. n. and P. (“F.”) remaudi comb. n., this is another African species of the group with a costal process on the valva.

Taxonomic treatment of outlier species

The following two species agree in most respects with those in the subgenus Facidia treated above, but show some outstanding features that might even be used to justify their placement each in a separate subgenus. On one hand, if the outlier was only one, the erection of a monobasic subgenus would be untenable in a cladistically inspired classification, because this would leave Facidia as paraphyletic, and this consideration would apply to either of the two outliers in relation to the main group of species. Nevertheless, as in this case the outliers are two, and because they are hardly reconcilable with one another, they cannot be comfortably associated to Facidia together. Cladistic orthodoxy would require assessing which of the two outliers is actually the sister group of the core Facidia, combine them under the same subgeneric concept, and separate the second one within a monobasic subgenus. However, this task is hampered by the fact that the autapomorphies of the two outliers are remarkable, pertain to different character systems, and have no counterparts in the core Facidia, so that they cannot easily be placed along a phylogenetic sequence. Further to this, their autapomorphies also involve some sort of developmental modulations, which adds further difficulty to ascertaining any sister-group relationships with other Facidia. For this reason, also awaiting possibly resolutive molecular information, the following two species will provisionally be ascribed to “Facidia” in a broad sense.

Facidia luteilinea Hampson, 1926. Descriptions of new genera and species of Lepidoptera Phalaenae of the subfamily Noctuinae (Noctuidae) in the British Museum (Natural History): 94. Type material: Holotypus ♂, by original designation, in NHMUK [examined]. Type locality: S. Nigeria, Old Calabar.

Diagnosis

An unmistakable, large-sized species with broad wings. In the male, the forewing is particularly high, with costa strongly bowed before apex and paired, elliptical, loosely scaled areas, especially on the underside, associated to a warped and modified venation (detailed hereunder). Genitalia as in Figs. 55, 67, and 74.

Distribution

So far known Western-Central Africa: Ivory Coast, Nigeria, and Gabon.

Remarks

Obtaining adequate knowledge about the features of this taxon was long hampered by the existence of only one known specimen, the male holotype, which was never illustrated. Examination of supplementary material from West Africa led to the circumscription of a further series of males closely matching the holotype in their structural features. These males show different degrees of expression of the pattern elements, but when considered overall they represent a consistent set of continuously varying phenotypes whose pattern elements may be differentially enhanced, especially the crosslines and distal banding of the wings, which can be more or less emphasized with dull yellow, and this either along their entirety or only in definite sections. One male from Gabon (in CPB) shows a fully expressed pattern that entirely matches that of some West African females that had long remained taxonomically unidentified. The bridging of the gaps in how the various pattern elements are expressed, both within and between the sexes, together with their morphoanatomical and geographical congruence, provides sufficient evidence that these specimens are conspecific and allows to spare the historical type from dissection. The previously unknown female of luteilinea deserves formal description, as detailed hereafter.

Description of the female

Wingspan 48–55 mm (n = 4). Head very large, dark chocolate-brown, frons bulged, vertex thickly and roughly scaled, eye large, globular, antenna filiform, labial palpus semi-erect, its first two joints compactly scaled, the third long, rod-like, white at very tip, haustellum present.

Thorax robust, uniformly dark chocolate-brown, patternless; forewing large and elongated, with crenulate termen; ground colour dark chocolate-brown, Platyja pattern (defined in Zilli et al. 2021) variably expressed as in male (in one worn specimen, not illustrated, it is equivalent to the male in Fig. 20), if with well evident white highlight, subbasal present, antemedial line subvertical, slightly oblique from costa to anal margin and feebly angular at veins (especially on 1A+2A), postmedial line concave (either smoothly arched or feebly waved) from costa and outwardly-directed towards submarginal line, where an acute tooth is produced, followed below by three shallow lobes, ‘drop’ of the typical Platyja pedunculate lobe well distinct and large, in shape of transverse ellipse that is joined by a median streak to tooth of inferior section of postmedial line; all projections of superior section of postmedial line (tooth, lobes, drop) closely adpressed to submarginal line, which runs obliquely albeit a little irregularly from apex to tornus; terminal line darker brown than ground colour, with white or golden dots in correspondence with veins, fringe brown with some sparse golden irroration at base and facing veins; hindwing broad and rounded, with crenulate termen, ground colour dark chocolate-brown, postmedial and submarginal lines incomplete, white, the former from middle of disc to anal margin, strongly waved, the latter straight and diffuse, only in lower third of wing; terminal area as in forewing. Legs slender, tarsomeres off-white ringed terminally. Underside of wings chocolate-brown with some white sprinkles and thin white strokes on veins in correspondence with postmedial lines, especially on hindwing.

Abdomen brown, shortly scaled. Genitalia: segment A8 wide, weakly sclerotised, apophyses anteriores short and flimsy; ostium bursae large, leading to broad, strongly sclerotised, trapezoidal antrum the inner wall of which extends posteriorly to ostium slit, antrum separated by narrow membranous section from short and wide, sub-quadrangular sclerotised section of ductus bursae, corpus bursae spheroidal, with narrow, elongated, conical appendix bursae from left posterior end of corpus, oblique; intersegmental membrane A8–A9 very short; papillae anales massive, increasingly setose towards posterior edge, apophyses posteriores rod-like, slightly longer than anteriores (Fig. 74).

Analysis of venation

By its external habitus, this species is extraordinary in many respects, notably the unusually shaped forewing of the male with long, elliptical, loosely scaled areas, one narrow, comprising the cell and the space beyond it, the other very broad below the cell, and the forewing venation of both sexes. The latter in particular shows features previously unrecorded—to the authors' best knowledge—in other Erebidae, and is out of the ordinary for Lepidoptera in general. It is dimorphic between the sexes, that of the female being less deviant from the commonest configuration found in Erebidae, which is why it will be discussed first, also to better ascertain homologies in the male. The forewing venation of the female is illustrated in Fig. 42, where veins of doubtful interpretation are marked X, Y and Z. By virtue of their position in the sequence with the other veins and the topology of branching from the discal cell, female M₁ and CuA₂ are unambiguously identifiable. Between them there are three veins, of which the anteriormost, free from the lower discocellular crossvein closing the cell, is certainly a medial (M_x), which is almost connate with a stalk situated below it and arising from the lower angle of cell, that of Y + Z. M_x could either be M₂ or M₃, or also the result of a complete fusion between the two (that should thence be noted as M₂-M₃). The interpretation of Y and Z depends on that of M_x, with four possible combinations, as follows (corresponding oddities between parentheses): M_x = M₂, Y = M₃, Z = CuA₁ (M₃ stalked with CuA₁); M_x = M₃, Y = CuA_1a, Z = CuA_1b (M₂ lost, CuA₁ split); M_x = M₂-M₃, Y = CuA_1a, Z = CuA_1b (M₂ and M₃ fused, CuA₁ split); M_x = M₂, Y = CuA_1a, Z = CuA_1b (M₃ lost, CuA₁ split). It may be noted that to account for the observed configuration, three of these options would imply the splitting of CuA₁ into two veins, while the remaining one would imply the stalking of M₃ with CuA₁. However, neither the splitting of the first anterior cubital nor the stalking of this with the last medial seem to have ever been recorded in Lepidoptera.

In the male (Fig. 43), the eccentricities are even more pronounced due to remarkable warping of the veins associated to the dilated, loosely scaled areas. There is a vein M_x that looks clearly homologous to the female one, and one long stalk proceeding from the cubitus beyond the lower angle of the cell that trifurcates into the veins noted here as Y, Z and W, with no other vein branching off from the cubitus below the cell. This trifurcation develops sequentially, first with a Y-(Z+W) split, then with a dichotomy of the last two, shortly stalked veins (i.e., Y is not connate with Z and W). It should be noted that what looks like a conspicuous vein longitudinally bisecting the large, loosely scaled area of the forewing is not a vein but a ridge/groove (on the upperside/underside, respectively), strongly lined with dark scales, that corresponds to the vannal fold, vein 1A+2A actually bordering the loosely scaled area below, next to the anal margin.

If all instances of losses, fusions and splits into supernumerary veins are considered, the possible combinations regarding the homologies of these veins in the male are very numerous, all implying some odd circumstances hitherto unrecorded in Lepidoptera. Nonetheless, if an ‘Occam’s razor’ approach is followed, there is one option that appears to be both more likely and more compatible for the two sexes, also in terms of visual similarity of the fork of Y and Z, i.e., M_x = M₂, Y = M₃, Z = CuA₁, and W in the male = CuA₂. In this case, the unusual occurrences would be those of M₃ (Y) stalked with CuA in both sexes, precisely CuA₁ in the female (Z), and male CuA₂ (W) distally shifted to branch off from CuA₁ (Z) following a short common stalk. As noted by Kristensen (2003), stalking of the forewing CuA fork is very unusual in Lepidoptera, but it is known in some Gelechioidea and thus there is support for considering the configuration of Z and W seen in male luteilinea as a developmental homoplasy that has some precedents in other moth groups.

The influence of developmental controls over the final configuration of venational patterns, best illustrated by the sexual dimorphism in this species for the position of CuA₂, sliding along the cubitus (if the above interpretation is correct), further calls for caution when using venational characters in classification, as already shown for another erebid species (Zilli 2021).

Megacephalomana remaudi Laporte, 1972. Bulletin du Muséum national d'Histoire naturelle (3) 59 (Zoologie 45): 664, fig. 6 (male genitalia). Type material: Holotypus ♂, by original designation, in MNHN [picture examined]. Type locality: Congo-Brazzaville, Loanga [verbatim holotype label: Luango].

Diagnosis

An unmistakable species, easily recognisable by the strongly modified albeit still fully distinct Platyja pattern of the forewing. In particular, the inferior sections of the antemedial and postmedial lines, both double, converge in the discal cell, where a big rusty-coloured patch is produced. This patch is proximal to the reniform stigma, i.e., in the shape of two acuminated marks slightly flexed with respect to one another and cojoined in the middle, and distal to a narrowly elongated orbicular stigma inside the cell. The rusty-brown ground colour and elongated forewing with apical area pale beige highlighted and notched apex are other outstanding external characteristics of this species. Genitalia as in Figs. 56, 68 and 75.

Distribution

Guinea, Liberia, Ivory Coast, Cameroon, Gabon, Congo (W, E).

Remarks

This is another greatly peculiar species of Platyja, especially for the strongly modified pattern and structure of the male antenna. The latter is not tripectinate but retains a midventral row of lamellae, which are short, blade-like and aligned along the flagellum to produce a midventral keel. Each flagellar antennomere bears, at the top of the keel and midway along the segment itself, a pair of transverse, oppositely oriented setae with bases not exactly juxtaposed but slightly shifted from one another. In the male genitalia, the thin arched rods of the diaphragma are poorly sclerotised and hardly recognisable.

In addition to these features, it is the pattern that provides the most surprising characteristic. Indeed, considering that the origin of the postmedial line of the forewing from costa is well evident, the more proximal segment from the costa to the rusty-coloured patch inside the cell may easily be recognised as the anterior section of the antemedial line, which is in fact double, like the posterior section reaching the anal margin. However, this would imply that the orbicular stigma is proximal to this line, which is something completely unusual for Noctuoidea (Fig. 44). Whatever the case, this arrangement shows the remarkable modifications occurring in remaudi and calls for a thorough assessment of the homologies of its pattern elements. The dotted anterior section of the postmedial line, the much reduced pedunculate lobe of the median field, and the apical pale beige mark on the notched apex further highlight the unusual appearance of this species, which finds a likely explanation in the disruptive appearance of the moth at rest. The posterior sections of the antemedial lines of the two wings closely align with the similarly coloured abdominal tergites 1–2 to produce a sort of transverse striga that terminates with a starred configuration at both ends.

Nomina dubia

The following nominal taxa originally or secondarily combined with Megacephalomana are of doubtful identity, and it is unlikely that this will ever be resolved due to the apparent loss of the types and to the original descriptions, hardly referable to any extant specimens or species.

Megacephalomana divisa (Walker, 1865) (Poole 1989)

Hypaetra divisa Walker, 1865. List of the Specimens of lepidopterous insects in the collection of the British Museum XXXIII (Supplement 3): 963. Type material: Syntypi ♂♂(number not given). Type locality: Ceylon.

Remarks

After its description, Hypaetra divisa was generally treated in Anereuthina Hübner, 1823 (type species: Anereuthina renosa Hübner, 1823), of which Hypaetra Guenée, 1852 is a junior objective synonym, e.g., by Moore ([1885] in 1884–1887) and Cotes & Swinhoe (1888). This taxon was transferred by Poole (1989) to Megacephalomana, but this action seems fully unsubstantiated by the fact that the type(s) went missing early on (Hampson 1893: 54, 1895: 104, 1913: 334), which compelled Hampson not to treat this unrecognizable species; ultimately, there are no clues about its identity. Strangely enough, the name of this taxon had already been mentioned by Motschoulsky (1863), attributed to Walker, in advance of its formal publication. As quoted by Motschoulsky, the name is nudum and thence unavailable. However, this circumstance shows that there were contacts between these two authors prior to the publication of Walker's (1865) volume.

Megacephalomana pilosum (Pagenstecher, 1888) (Poole 1989)

Megacephalon pilosum Pagenstecher, 1888. Jahrbücher des Nassauischen Vereins für Naturkunde 41: 156. Type material: Holotypus ♂, by monotypy, in MFNB (coll. Staudinger) [not traced]. Type locality. Amboina.

Remarks

This nominal taxon of doubtful identity has already been extensively discussed in Zilli et al. (2021).

Provisionally placed species

The following taxon is a validly recognised species, but all evidence indicates that it has always been misplaced in Megacephalomana. Due to the synonymisation of Megacephalomana with Facidia as a subgenus of Platyja, such species has to provisionally follow the new taxonomic arrangement for its original generic name in order not to be left floating in a taxonomic ‘limbo’. However, it is recommended that it is cited with the genus-group names between quotation marks, to stress that this is a temporary assignment pending further studies that will hopefully clarify its systematic relationships.

Megacephalomana laportei Berio, 1974. Annali del Museo civico di Storia naturale “Giacomo Doria” 80: 224, fig. 12 (male genitalia). Type material: Holotypus ♂, by original designation, in MRAC [picture examined]. Type locality: Congo, Sankuru, Katako-Kombe.

Remarks

This species from the Congo basin was attributed to Megacephalomana by Berio (1974), who also relied on the advice of Bernard Laporte, great connoisseur of African Noctuoidea and author of another species of Megacephalomana (namely remaudi). During the present study, no material of this taxon was available. However, analysis of the wing pattern and features of the male genitalia, all well represented by De Prins & De Prins (2011–2021), strongly support the view that it does not belong to the Platyja genus group. In fact, the habitus has no relation at all to the characteristic Platyja pattern—it rather shows similarities with the Saroba genus group (sensu Holloway 2005)—and the genitalia have a short vinculum not produced into a long, narrow, inwardly-bent saccus, the valva bears a robust clasper protruding towards the middle and has strong sparse setae along the distal edge, normal transtillae are present, and the uncus is long-necked with strongly setose, dorsally sulcate apical club: all features not seen in any other member of the subgenus Facidia (= Megacephalomana). The holotype, the only male known to the present authors, seems to have lost its antennae, and thus their structure cannot be checked.

Distribution

Known only from E Congo.