On the taxonomy of Meganola cretacea (Hampson, 1914) with description of two new species (Lepidoptera, Nolidae, Nolinae)

The taxonomic position of Meganola cretacea (Hampson, 1914) is clarified and the species is transferred to the hitherto monotypic Oriental genus Ezishnola László, Ronkay & Witt, 2010: Ezishnola cretacea (Hampson, 1914) comb. nov. The male and female genitalia of E. cretacea are illustrated and described for the first time. Two new species from Zambia ( E. inopinata sp. nov. ) and Uganda ( E. carcassoni sp. nov. ) are described. 14 colour and 10 black and white diagnostic figures and a distribution map are provided.


Introduction
The peculiar Noline species Roeselia cretacea Hampson, 1914 has long been known from its two female syntype specimens housed in the collections of NHMUK. One of the two syntypes is missing its abdomen and is therefore inadequate for morpho-taxonomic studies. In his revision of the Afrotropical Nolini, Hacker illustrated a male specimen from Ivory Coast, collected at Danane as the third known specimen of the taxon (Hacker et al. 2012). The external appearance of the male specimen indeed matches well with that of the female syntypes and though Hacker illustrated the male genitalia of the species, he did not provide information on the female genitalia. As the main characters of the male genitalia of the illustrated specimen considered to be M. cretacea by Hacker (Hacker et al. 2012) fit well with the general ground plan of Meganola, he transferred the species to this genus. During the course of studying Heterocera materials recently collected in West Africa, numerous specimens of M. cretacea have been found in the collections of ANHRT and HNHM. Despite the fact that the external appearance and wing pattern of all these examined specimens are nearly identical with that of the type specimen of M. cretacea, the male genitalia of all of the 6 dissected West African specimens have turned out to be fundamentally different from the one illustrated in Hacker et al. (2012). The reason for this discrepancy could be either an editorial error in Esperiana Vol. 17 or a misplaced abdomen or confused genitalia preparation of the specimen examined by Hacker. This assumption is supported by the fact, that the otherwise rather characteristic genitalia referred by Hacker et al. (2012) as M. cretacea seems to be identical with that of M. genitaliana Hacker, 2012 illustrated 17 pages later (cf p. 496 and 513 of the same publication). Either way an error has occurred and it is certain that the real genitalia morphology of M. cretacea has remained unknown until now.
A number of specimens of both sexes of true M. cretacea have been located during the examination of the Nolini material of the ANHRT and HNHM, and the genitalia morphology of the species is clarified for the first time in this present paper. Surprisingly, based on both male and female genital characters, this externally distinct species turned out to belong not to Meganola, but to Ezishnola László, Ronkay & Witt, 2010, a hitherto monotypic genus that has been known from the Oriental region only. The placement of M. cretacea in the genus Ezishnola is supported by the similarly broad, rather quadrangular forewing with similar arrangement of transverse lines, the large, medially dilated, rounded, tongue-shaped valva and the long, spike-like harpe situated in the medial plate of the valva in the male. In addition, the female genitalia display very similar configurations of the ovipositor and eighth segment, the short sclerotized and long membranous sections of the ductus bursae and the ovoid corpus bursae bearing a pair of rounded plates of the signum bursae.
Further specimens reminiscent of E. cretacea with similar habitus but with a considerably darker greyish forewing ground colour were found in the collection of ANHRT collected recently in Zambia. Despite the rather different coloration, the Zambian specimens show almost identical configuration in male genitalia with those of the West African E. cretacea. Interestingly however, the female genitalia of the Zambian specimens differ considerably from the West African ones, and taking into consideration their strikingly different external habitus, the E. cretacea-like specimens from Zambia are described here as new to science.
In the course of sorting the mixed Heterocera accessions of the NHMUK, a male and female of an E. cretacea-like insect from Uganda were found. The examination of their genitalia revealed that despite the fact that their external habitus is similar to E. cretacea from West Africa, the configuration of the female genitalia of the Ugandan specimen shows a closer relationship to the new Zambian species. The male genitalia of the Ugandan specimen is very similar to the other two taxa but the female genitalia differs markedly. Thus, the specimens from Uganda represent a third African Ezishnola taxon and is described here as new to science. As all three Ezishnola species bear distinctive characters in their female genitalia, female specimens have been designated as the holotypes of the two new species.

Material and methods
The genital apparatuses were dissected, stained with Eosin red and mounted in Euparal on microscope slides applying standard methods of preparation (Lafontaine & Mikkola 1987). Photos of adults were taken using either a Nikon D700 SLR camera equipped with Nikkor AF-S Micro 105 mm lens or a Nikon D90 SLR camera equipped with Nikkor AF Micro 60 mm lens. Genitalia were photographed using either a Canon EOS 700D camera mounted on a Leitz Diaplan compound microscope or a Canon EOS 5D SLR camera with a Canon MP-E 65 mm lens.

Description of the female genitalia
Ezishnola inopinata sp. nov. (Figs 9-12, 16, 21, 22) Holotype. Female. "Zambia, 1147m, Lukwakwa, West Lunga NP (Cryptosepalum forest/Dambo), S12°39'40", E24°26'13", 9-15 Diagnosis. E. inopinata is an allopatric sister species of E. cretacea distinguished by the following characters: E. inopinata has a considerably darker forewing ground colour with more sharply defined, dark greyish transverse lines and with much wider blackish irregular patches in the postmedial area compared to those of E. cretacea; in addition, the postmedial line of the new species is continuous, whereas that of the sister species consists of a row of dots. In the male genitalia, the differences between the two taxa are subtle, the new species has a somewhat shorter uncus, a narrower, medially less dilated valvae, and slightly shorter and narrower harpe compared to E. cretacea. The differences between the two species are well expressed in the configuration of the female genitalia: E. inopinata has slightly shorter papillae anales, much shorter and thinner apophyses, somewhat wider and shorter goblet-shaped ostium bursae, a somewhat thicker ductus bursae with considerably shorter medial sclerotization, a conspicuously shorter and thicker tubular distal part and much more elongated proximal part of corpus bursae compared to those of E. cretacea; however, the most conspicuous difference between the two species is expressed by the complete lack of signum bursae in E. inopinata, while its West African sister species has a pair of heavily sclerotized semi-circular signa bursae.
Description (Figs 9-12). Forewing length 9.5-11 mm in males, 11-12 mm in females. Antenna filiform in both sexes with somewhat longer and denser ciliation in males compared to that of females.
Head relatively large, labial palps medium long, curved dorsad, inner surface greyish white, lateral and ventral surface of first and second segments dark blackish grey, third segment pale greyish white; frons and vertex pale greyish white; compound eyes moderately large, globular. Thorax whitish grey, basal twothird of collar darker greyish, abdomen creamy whitebrownish white striped. Intraspecific variability limited, all known specimens are rather similar in coloration without noticeable alterations in wing pattern. Sexual dimorphism is moderately expressed in size (females slightly larger) and in coloration, namely females have slightly darker forewing and considerably darker hindwing ground colour compared to those of males.
Forewing relatively long and broad, rather quadrangular, apically rounded. Forewing ground colour pale graphite grey, with blackish grey irregular patches in the postmedian area. Sub-basal and basal lines diffuse, shadow-like, represented by groups of dark grey hair scales; antemedial line fine, dark grey, interrupted, almost straight, running angularly towards costa where it joins the medial line, forming a small but conspicuous dark grey quadrangular costal patch; medial line rather thick, slightly diffuse and shadowlike, medially angled, with lower section reaching ventral margin in right angle. Orbicular stigma rather large, rounded, consisting of raised pale whitish grey scales. Postmedial line relatively thick, very sharply defined by blackish scales, costal section broadly arcuate, ventral section straight, running parallel with medial line. Subterminal line diffuse, interrupted, shadow-like, consisted of dark graphite grey patches of different size; terminal line very fine, undulating, nearly continuous, blackish; terminal area with dark grey suffusion; cilia pale brownish grey chequered with dark grey.   (Hampson, 1914), LGNA 835, Guinea, Nimba Mts (ANHRT); 20, E. cretacea (Hampson, 1914) Hindwing pale whitish grey in basal half, gradually darkened towards termen in males, darker grey in females; cilia whitish grey in both sexes. Underside of forewing uniformly dark graphite grey, that of hindwing pale graphite grey, darkened along the margin in males, similar but darker in females, without traces of pattern. (Figs 21, 22). Uncus relatively short, rather narrow, distally slightly dilated, apically with a short, fine, pointed spine; subscaphium rather long and narrow poorly sclerotized; tegumen short and broad, rather triangular; valvae weakly sclerotized, medium long, medially moderately dilated, apically broadly rounded, tongue-shaped; harpe spike-like, rather long, relatively thick, slightly arcuate, apically pointed, situated in the central plate of valva; transtillae well-developed, semi-circular, medially divided; sacculus long and broad, basal third more strongly sclerotized than apical one; fultura inferior (juxta) very small, with a pair of ear-like rounded lobes; vinculum rather broad and long, V-shaped, saccus short, apically acutely pointed. Aedeagus short and narrow, caecum penis rather long, evenly rounded; carina without sclerotization; vesica membranous in full length, without cornuti.
Etymology. The latin adjective inopinata means unexpected, referring to the surprising occurrence of a new Ezishnola species in NW Zambia. Ezishnola carcassoni sp. nov. (Figs 13, 14, 17, 23 Diagnosis. E. carcassoni sp. n. is an East African allopatric sister species of E. creatacea and E. inopinata known so far only from Uganda. The new species differ externally from both sister species by its conspicuously interrupted transverse lines consisting of a dense row of sharply defined blackish dots, whereas the crosslines are more continuous in the other species, although similar wing pattern is not unknown in E. cretacea. In the male genitalia, E. carcassoni differs from both sister species by its somewhat shorter uncus and valva, in addition more robust, almost straight harpe (those of the other two species are thinner and slightly arcuate) and considerably narrower, more elongate transtillae. In the female genitalia, the new species has somewhat shorter apophyses than those of E. cretacea, ovoid corpus bursae without tubular section and without sack-like cervix bursae (corpus bursae with conspicuously long tubular section and cervix bursae is sack-like in E. cretacea), in addition E. carcassoni lacks signum bursae. The differences between the female genitalia of the new species and E. inopinata are as follows: E. carcassoni has somewhat longer apophyses, narrower ductus bursae with longer sclerotized medial section compared to those of E. inopinata, in addition less elongate, more rounded corpus bursae with simple cervix bursae, while the corpus bursae of E. inopinata is more elongated with anteriorly dilated distal tubular section and its cervix bursae with a well-developed, membranous sack.
Description (Figs 13, 14). Forewing length 12 mm in male, 13 mm in female. Antenna filiform in both sexes with somewhat longer and denser ciliation in males compared to that of females.
Head relatively large, labial palps medium long, curved dorsad, inner surface brownish white, lateral and ventral surface of first and second segments brownish grey, third segment pale creamy white; frons and vertex pale brownish white; compound eyes moderately large, globular. Thorax brownish grey, basal twothird of collar darker greyish, abdomen brownish white. Intraspecific variability limited. Sexual dimorphism is moderate, expressed in size (female slightly larger) and in coloration, (female hindwing somewhat darker than that of male).
Forewing relatively long and broad, rather quadrangular, apically rounded. Forewing ground colour pale brownish grey, with some darker brown irregular patches in the postmedian area. Sub-basal and basal lines diffuse, shadow-like, represented by groups of brownish hair scales; antemedial line fine, brownish grey, interrupted, almost straight, running angularly towards costa; medial line rather thick, interrupted, diffuse, present only in the ventral third; Orbicular stigma rather large, rounded, consisting of raised pale whitish grey scales. Postmedial line relatively thick, interrupted, very sharply defined by dense row of dark brown scales, upper section broadly arched, lower section running parallel with medial line. Subterminal line rather undulating, interrupted, consisting of dark brown triangular patches of different size; terminal line thick, interrupted, consisting of a row of dark brown streaks; terminal area with some red-brown suffusion; cilia pale brownish grey chequered with darker brown.
Hindwing pale greyish white, with some darker brownish suffusion terminally in males, darker grey in females, turning to brownish grey in the terminal area; cilia whitish grey in both sexes. Underside of forewing uniformly dark brownish grey, that of hindwing paler brownish grey, darkened along the margin in males, similar but darker in females, without traces of pattern.