Integrative taxonomic revision of the African taxa of the Racotis Moore, 1887 generic complex (Lepidoptera, Geometridae, Ennominae, Boarmiini) Author László, Gyula M. 0000-0001-9862-8290 gyulalaszlo@anhrt.org.uk Author Hausmann, Axel 0000-0002-0358-9928 hausmann.a@snsb.de Author Karisch, Timm 0000-0001-9862-8290 gyulalaszlo@anhrt.org.uk text Zootaxa 2023 2023-06-26 5308 1 1 109 http://dx.doi.org/10.11646/zootaxa.5308.1.1 journal article 10.11646/zootaxa.5308.1.1 1175-5326 8080929 CCA5F817-6B5F-4BE5-BEFB-EDE98C07A0EE Zebracotis subgen. n. Type species: Racotis zebrina Warren, 1899 Genetic analyses (Text fig. 3). The subgenus was recovered as a well-supported sister clade of the subgenus Afroracotis with high support in both analyses (BS: 55; PP: 97). The DNA barcode-based infrageneric classification of the subgenus, however, is rather inconsistent often conflicting with the clearly distinguished morphotaxa. Based on certain constant morphological characters discussed below, the subgenus contains ten taxa (nine species and one subspecies), while only seven BINs in the complex were assigned in BOLD. The topologies of the BI and ML trees are nearly identical and both are rather incongruent with the morphology-based taxon delineations, where a single morphological taxon is often split into two separate genetic clusters without any diagnostic characters. Such an example is A. milesi sp. n. , which was recovered as paraphyletic: in a cluster of exclusively Gabonese specimens BIN-sharing with A. zebrina , A. chaineyi sp. n. and A. lydiae orientalis ssp. n. and in a distinct cluster with its own BIN URI comprising A. milesi samples from Zambia , Cameroon and Gabon . Another example of such incongruency between genetics and morphology is A. lydiae sp. n. which is morphologically subdivided into a West (nominotypical taxon) and East African subspecies ( A. lydiae orientalis ssp. n. ), however, the West African subspecies is recovered as a genetically largely separate cluster with its own BIN URI, whereas the East African subspecies is BIN-sharing with the predominantly West African A. zebrina , the mainly central African A. milesi sp. n. and the more widespread A. smithi sp. n. . The only taxa showing genetic features being congruent with morphology are A. madagascariensis ( Chainey & Karisch, 2017 ) which is recovered as a sister clade to all other Zebracotis taxa in both analyses (BS: 67%, PP: 100%), A. ochsei sp. n. , A. takanoi sp. n. and A. helicalis sp. n. , although the last one clusters as a sister species to A. lydiae lydiae despite their largely dissimilar genital morphology. It is difficult to reliably establish the reason of such an extent of conflicts between genetics and morphology in the Zebracotis subgenus. Seemingly, DNA barcodes have limited taxonomic relevance in this group, possibly due to repeated genetic introgression. Alternatively, the members of the subgenus may be evolutionary very young taxa “in statu nascendi”, where the speciation has been manifested in an early divergence in genital morphology before a clear differentiation in the external morphology and COI DNA sequences could have taken place. It is also worth noting that the single COI-5P gene region used in the genetic analyses is relatively short and an in-depth genetic analysis based on multiple gene regions (including nDNA) would presumably result in a better resolution of the phylogeny. Overall, as the tree topology of the Zebracotis subgenus is not entirely congruent with the morphological species concept, the taxa in this subgenus have been delimited based primarily on the remarkable and constant differences observed in the genital morphology. TEXT FIGURE 3 . ML tree of the subgenus Zebracotis based on analysis of COI barcodes. Bootstrap values are indicated above, posterior probability values below the branches. Samples which have not been confirmed by dissection are labelled as ‘indet’. Diagnosis. The species of the subgenus are characterized by the broad, triangular forewing, the dark greyish-green colouration of densely speckled scales on the upperside, the well-defined, ovoidal discal spots, and the variably contrasting row of round black patches representing the subterminal line. As the wing pattern varies largely both infra- and interspecifically, the facies of most of the species do not provide reliable distinctive features for species-level identification. In the males of all members of the subgenus, the third abdominal sternite bears a small tuft of fine setae, a character which is also found in A. squalida and in the monotypic subgenus Sokokeracotis subgen. n. within Afroracotis , however, it is also present in the sister genus Chorocotis gen. n. . The diagnostic male genitalia characters of the subgenus are as follows: Zebracotis has a clasping apparatus similar to that of Herbuloracotis , but it has a longer uncus, a medially inverse V-shaped gnathos without a shield-like medial plate, a ventro-medially more concave valva, and a considerably more elongate juxta, medially tapered into a narrow distal projection. The vesica of the Zebracotis species bears highly distinctive characters allowing reliable species-level identification. It is characterized by the medium long and moderately inflated main vesica chamber, in most species with a well-developed diverticulum with an apical cornutus of specific shape and a group of cornuti erected distally at the base of the vesica ejaculatorius. The shared characters of the Zebracotis species in the female genitalia are the sclerotised, broad cup-shaped antrum, the absence of the membranous ductus bursae replaced by the variably elongate heavily sclerotised cervix bursae which is fully merged into the corpus bursae forming a sclerotised posterior section. The lack of true ductus bursae clearly separates the Zebracotis subgenus from all other lineages of Afroracotis . Description External features of body and wings ( Figs 81–119 , 175–204 ). Forewing length 17–24 mm . Male antenna bipectinate-fasciculate, rami moderately long, distal section filiform, varying from 1/2 to 2/5 of length of antenna. Female antenna filiform. Head moderately large, proboscis well-developed, labial palp short, length 1.2–1.5 times the diameter of eye, porrect, first and second segment dilated, third segment rather short and thin. Compound eyes moderately large. Frons, vertex, collar (patagium), tegula, thorax and abdomen colouration as of wing upperside. Legs rather long, colouration as of wing underside, index of spurs 0-2-4. Forewing broad, triangular, costa straight, apical section slightly arched, termen straight to slightly arched, ventral margin almost straight. Ground colour varying inter- and intraspecifically from dark rusty-brown to greenish-grey usually with mossy tinge, densely speckled with groups of brownish-grey scales, basal and medial area often lighter than terminal one, postmedial area with series of large dark blotches. Transverse lines diffuse, interrupted. Basal, subbasal and antemedial line poorly detectable, interrupted, often partly deleted, more or less straight, or gently arched. Medial line often interrupted, shadow-like, slightly S-curved, postmedial line diffuse, slightly wavy, subterminal line represented by a row of patches of variable size, terminal line narrow, often interrupted, consisting of black dashes or arches between veins. Fringe (cilia) short, concolourous with forewing. Discal spot well-developed, round or ovoid with lighter inner area. Hindwing apex rounded, termen evenly arcuate, slightly scalloped, anal margin straight. Ground colour as of forewing, densely speckled, transverse lines diffuse, shadow-like, basal line evenly arched, wavy or sharply angled medially, medial line undulate, postmedial line interrupted, terminal line sometimes interrupted, consisting of black arches between veins. Discal spot as of forewing. Fringe (cilia) short, concolourous with hindwing. Wing underside with pale brownish-grey basal and medial area speckled densely with greyish-brown scales, terminal area with broad, dark transverse band. Traces of transverse lines present, diffuse, shadow-like, discal spot and postmedial patches well-defined. Male genitalia ( Figs 254–283 ). Uncus simple, moderately long, distally tapered, apex narrowly rounded. Tegumen short and broad, gnathos well-developed, apically rounded. Valva short, broad at base, distally tapered, costal margin slightly concave, sclerotised, sparsely setose, without apical projection.Ventral margin of valva convex basally and subapically, smoothly concave medially; sacculus short and broad, weakly sclerotised, without process. Juxta weakly sclerotised, distally tapered, elongate shield-like without arms. Vinculum moderately long and broad, V-shaped, apically rounded.Aedeagus rather thick, short to medium long, straight or slightly curved, without carina. Vesica thick, relatively long, in some species with protractile diverticulum armed with a robust apical cornutus, and, in all but one species ( A. zebrina ) with a small group of cornuti situated near the base of vesica ejaculatorius. Female genitalia ( Figs 320–338 ). Ovipositor medium long, papilla analis narrow triangular, apically rounded, sparsely setose, apophysis posterioris moderately long. Eighth tergite medium long, trapezoidal, distal margin slightly arcuate, proximal margin undulate with a shallow medial depression, anterior apophysis shorter than posterior apophysis. Ostium bursae broad, membranous, antrum moderately sclerotised, short, funnel-, cup-, goblet-shaped or inverse trapezoidal. Distal section of ductus bursae membranous, very short and narrow, may be fully reduced; proximal part tubular or funnel-shaped, varying in length interspecifically, directly continued in sclerotised distal part of corpus bursae. Cervix bursae well-developed, heavily sclerotised, fully integrated into corpus bursae forming a sclerotised, longitudinally striate distal section of bursa copulatrix. Membranous anterior part of corpus bursae varied in size interspecifically, may be almost fully reduced; corpus bursae with a strongly sclerotised, smooth-surfaced disc-like or quadrangular plate supposedly serving as receptive area of cornutus vesicae during copulation; signum absent. Species content Afroracotis (Zebracotis) zebrina ( Warren, 1899 ) comb. nov. Afroracotis (Zebracotis) chaineyi sp. n. Afroracotis (Zebracotis) lydiae sp. n. Afroracotis (Zebracotis) lydiae orientalis ssp. n. Afroracotis (Zebracotis) madagascariensis ( Chainey & Karisch, 2017 ) comb. nov. Afroracotis (Zebracotis) smithi sp. n. Afroracotis (Zebracotis) milesi sp. n. Afroracotis (Zebracotis) ochsei sp. n. Afroracotis (Zebracotis) helicalis sp. n. Afroracotis (Zebracotis) takanoi sp. n.