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 Afroracotis gen. n. Type species: Ophthalmodes squalida Butler, 1878 by present designation. Genetic analyses ( Fig. 340 , Text figs 1–3) Preliminary ML analyses of COI barcode data for 287 specimens of Racotis s.l. in BOLD (>500 bp, worldwide) showed a pattern of several well-separated clusters diverging at distances of 5–7% from each other. All Indomalayan species clustered into two clades, one which included the type species of the genus, Racotis boarmiaria and the other R. maculata Lucas and associated taxa, while all African taxa clustered separately. In order to obtain a clearer picture of the phylogenetic relationship between the Asian and African lineages of Racotis s.l. , as well as within the Afrotropical taxa, 179 morphologically identified specimens were analysed. The phylogenetic inferences obtained from BI and ML analyses recovered similar topologies, although the BI tree was not fully resolved with the Indomalayan boarmiaria and the Afrotropical apodosima , argillacea and Rwandaracotis subgen. n. lineages recovered as a polytomy. Due to its better resolution, the ML tree was chosen for illustration in this paper ( Fig. 340 , Text figs 1–3). In the ML analysis, the outgroup taxon Eulycia grisea ( Warren, 1897 ) + Asian Racotis clade was recovered as sister to all other African taxa. All African taxa were recovered as monophyletic, although with weak support (BS: 12; PP: -) but considering their genital morphology, the African Racotis are recognised here as belonging to a distinct genus: Afroracotis gen. n. Surprisingly, the analyses also suggested that Racotis ugandaria ( Swinhoe, 1904 ) —a species which has long been treated as the sole Afrotropical member of the genus Chorodna Walker ( Scoble & Hausmann 2007 ) and recently transferred to Racotis ( Murillo-Ramos et al. 2021 ) —does not belong to either the Indomalayan Racotis or the Afrotropical Afroracotis lineages but represents a distinct genus which is described in this paper: Chorocotis gen. n. A morphologically rather distinct, hitherto undescribed species from Rwanda was recovered as a basal sister branch to all other Afroracotis clades (BS: 12; PP: -), and it is recognised here as a representative of a monotypic subgenus Rwandaracotis subgen. n. Another surprising result was the morphologically very similar A. argillaceaapodosima group (BS: 9; PP: -) and the A. deportata cluster of undescribed species (BS: 16; PP: -) being recovered as paraphyletic. Based on their homologous habitus and genital morphology, both clades are recognised here as representatives of a distinct subgenus, Herbuloracotis subgen. n. (Text fig. 2). Due to the incongruencies between the genetic and morphological analyses, the species-group subdivision of this subgenus is based on morphological features corresponding with the ML topology in the A. apodosima and A. deportata species-groups, but the A. argillacea species-group remains paraphyletic. The A. squalida + zebrina lineage (BS: 55; PP: 97) is clearly divided into two well-supported monophyletic units with clear diagnostic morphological characters representing two subgenera: Afroracotis subgen. n. (Text fig. 1) and Zebracotis subgen. n. (Text fig. 3). Afroracotis is subdivided into two species-groups (BS: 96; PP: 100) clearly corresponding with morphology, whereas in Zebracotis both genetic analyses provided a rather heterogenous topology often conflicting with the constant diagnostic morphological traits of the species. The results of the genetic analyses are further discussed in detail under each subgenera. Besides the four subgenera delimited through integrative analyses, a fifth subgenus ( Sokokeracotis subgen. n. ) is also described in this paper based merely on its unique genital morphology, as the single accessed specimen could not be successfully sequenced. Diagnosis . Afroracotis gen. n. shares the general external habitus with the Indomalayan-Australasian Racotis Moore, [1887] ( type species: Hypochroma boarmiaria Guenée, [1858] described from eastern India , ( Figs 1 , 206 )), characterised by the dark greenish-brown, often mossy-green colouration usually darkened in the distal third of the wings, the typically speckled wing pattern with the diffuse, interrupted transverse lines often replaced by rows of variably sized patches only and the well-defined, relatively large ovoidal discal spot. The pattern of the underside of the wings is also similar in both genera characterised by the usually sharply divided dark greyish postmedial third and the considerably paler, variably speckled basal and antemedial areas. The new Afrotropical genus differs from Racotis by a conspicuous autapomorphic character of the male genitalia, the absence of the saccular process, which is well-developed in species of true Racotis ( Fig. 211 ). Additionally, the configuration of the gnathos seems to display constant differences between the species of Afroracotis and Racotis , being apically rounded in the former and pointed in the latter. The gound plan of the female genitalia in Afroracotis and Racotis are rather homologous (cf. Sato 2004 ), especially the configuration of the fused cervix and corpus bursae suggesting a close affinity between the genera. However, the Racotis females have a uniformly elongate, sack-like bursa copulatrix unlike the remarkably diverse corpus bursae configuration in Afroracotis . Description External features of body and wings ( Figs 1–120 , 131–205 ). Forewing length 16–30 mm . Male antenna bipectinate-fasciculate, rami short or moderately long, distal section filiform, ratio of which varying interspecifically between 1/6 and 1/2 length of entire antenna. Female antenna filiform or bipectinate-fasciculate with short rami in several taxa of A. argillacea species-group. Head moderately large, proboscis well-developed, labial palp short, length 1.2–2.5 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 concolourous with 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 arcuate, may be moderately scalloped, ventral margin straight. Venation of typical Ennominae ground plan, forewing with four branches of R veins, cell relatively short, wide distally, Sc and R1 running parallely, R1+R2, R3, R4, R5 stalked, M1, M2, M3, CuA1, CuA2 and A well-developed, a longitudinal fold between CuA2 and A present. Hindwing with Sc+R1, Rs, M1, M3, CuA1, CuA2, A2, A3 fully developed, M2 reduced to a fold. Wing venation of all subgenera identical. Ground colour varying interspecifically from greyish dark green usually with a mossy tinge to pale ochreous-yellow, characteristically speckled in most taxa, basal and medial area often paler than terminal one, postmedial area with large dark blotches between M3 and CuA 1 in several species. Transverse lines diffuse, interrupted. Basal and antemedial lines detectable, more or less straight, medial line shadow-like, moderately S-curved, postmedial line slightly wavy, subterminal line represented by a row of patches or deleted, terminal line interrupted, consisting of small black dots, short dashes or arches between veins. Fringe (cilia) short, concolourous with forewing. Discal spot well-developed, dot-like, round or ovoidal. Hindwing apex rounded, termen evenly arcuate, or medially angled and moderately scalloped in certain taxa, anal margin straight. Ground colour as of forewing, densely speckled, basal line often diffuse, evenly arched, wavy or sharply angled medially, medial line undulate, postmedial line diffuse, interrupted, terminal line interrupted, consisting of small black dots, short dashes or arches between veins. Discal spot as on forewing. Fringe (cilia) short, colour as of hindwing. Wing underside with broad, dark terminal area with exception for the A. argillacea species-group with limited darker area. Basal and medial areas considerably paler than terminal one, densely suffused with greyish scales. Traces of transverse lines present, often poorly visible, discal spot and postmedial patches well-defined. Male genitalia ( Figs 212–284 ). Uncus short, bifid with short apical projections or simple, moderately long, distally tapered. Tegumen short and broad, gnathos well-developed, apically rounded. Valva short, broad at base, distally tapered; costal margin sclerotised, often setose, may have a sclerotised apical projection outreaching the membranous ventro-distal section of valva. Ventral margin of valva moderately arched, medially often concave, slightly varying intra- and interspecifically; sacculus short and broad, weakly sclerotised, without process. Juxta weakly sclerotised, shield-like without arms. Vinculum short or moderately long and broad, V-shaped or medially tapered, apically rounded. Aedeagus rather thick, short to medium long, straight or slightly curved, without carina. Vesica thick, moderately long, in most species armed with robust cornutus, may bear additional cornuti field and well-developed diverticulum. Configuration of vesica-cornuti-diverticulum complex is an important diagnostic species-level character. Female genitalia ( Figs 290–339 ). Ovipositor medium long, papilla analis narrow triangular, apically rounded, sparsely setose, apophysis posterioris moderately long. Eighth tergite medium long, quadrangular or trapezoidal, distal margin more or less straight, proximal margin often undulate, apophysis anterioris shorter than apophysis posterioris. Ostium bursae broad, membranous, antrum moderately sclerotised, short, funnel-, cup- or goblet-shaped. Ductus bursae membranous, short and narrow, may be fully reduced, substituted by tubular distal part of cervix-corpus bursae complex. Cervix bursae well-developed, heavily sclerotised, often present as a rounded bulge or may be fully merged with corpus bursae forming a sclerotised, distal section of bursa copulatrix. Corpus bursae varying in size interspecifically, may be highly reduced and fully fused with sclerotised, often rugose cervix bursae or well-developed and separated from cervix; may bear a strongly sclerotised disc-shaped plate supposedly serving as receptive surface of cornutus during copulation; signum absent. Taxonomic overview of the genus Afroracotis Afroracotis subgen. n. The Afroracotis squalida species-group Afroracotis (Afroracotis) squalida squalida ( Butler, 1878 ) comb. n. Afroracotis (Afroracotis) squalida voeltzkowii ( Pagenstecher, 1907 ) comb. n. Afroracotis (Afroracotis) squalida thomensis ssp. n. Afroracotis (Afroracotis) incompletaria incompletaria ( Guenée, 1862 ) comb. n. Afroracotis (Afroracotis) incompletaria atrilunaria ( Mabille, 1893 ) comb. n. , stat. n. Afroracotis (Afroracotis) aliena sp. n. The Afroracotis angulosa species-group Afroracotis (Afroracotis) angulosa ( Herbulot, 1973 ) comb. n. Afroracotis (Afroracotis) canui ( Herbulot, 1991 ) comb. n. Afroracotis (Afroracotis) stadiei sp. n. Afroracotis (Afroracotis) violetteae sp. n. Afroracotis (Afroracotis) fiebigi sp. n. Afroracotis (Afroracotis) albitrigonis ( Prout, 1927 ) comb. n. Rwandaracotis subgen. n. Afroracotis (Rwandaracotis) turlini sp. n. Herbuloracotis subgen. n. The Afroracotis apodosima species-group Afroracotis (Herbuloracotis) apodosima ( Prout, 1931 ) comb. n. Afroracotis (Herbuloracotis) dargei sp. n. The Afroracotis deportata species-group Afroracotis (Herbuloracotis) deportata ( Herbulot, 1970 ) comb. n. The Afroracotis argillacea species-group Afroracotis (Herbuloracotis) argillacea argillacea ( Warren, 1900 ) comb. n. Afroracotis (Herbuloracotis) argillacea morettoi ssp. n. Afroracotis (Herbuloracotis) argillacea cedrici ( Herbulot, 1998 ) stat. n. Afroracotis (Herbuloracotis) longicornuta longicornuta sp. n. Afroracotis (Herbuloracotis) longicornuta congolana ssp. n. Afroracotis (Herbuloracotis) longicornuta ugandana ssp. n. Afroracotis (Herbuloracotis) aristophanousi sp. n. Afroracotis (Herbuloracotis) muscivirens sp. n. Afroracotis (Herbuloracotis) atriclava ( Prout, 1926 ) comb. n. Zebracotis subgen. n. Afroracotis (Zebracotis) zebrina ( Warren, 1899 ) comb. n. Afroracotis (Zebracotis) chaineyi sp. n. Afroracotis (Zebracotis) madagascariensis ( Chainey & Karisch, 2017 ) comb. n. Afroracotis (Zebracotis) lydiae lydiae sp. n. Afroracotis (Zebracotis) lydiae orientalis ssp. n. 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. Sokokeracotis subgen. n. Afroracotis (Sokokeracotis) staudei sp. n. Key to the subgenera of the genus Afroracotis based on male genitalia characters 1. Uncus short, bifid.................................................................................... 2 -. Uncus moderately long, simple, pointed................................................................... 3 2. Valva tip straight, valva without medial crest....................................................... Afroracotis -. Valva tip curved medio-caudad, valva with medial crest............................................. Sokokeracotis 3. Vesica with a single, usually large (reduced in apodosima group), acute distal cornutus.............................. 4 -. Vesica with a diverticular cornutus, or a distal cluster of cornuti or both................................... Zebracotis 4. Vesica extremely long and thin................................................................ Rwandaracotis -. Vesica short and thick....................................................................... Herbuloracotis