Herbuloracotis, László & Hausmann & Karisch, 2023

László, Gyula M., Hausmann, Axel & Karisch, Timm, 2023, Integrative taxonomic revision of the African taxa of the Racotis Moore, 1887 generic complex (Lepidoptera, Geometridae, Ennominae, Boarmiini), Zootaxa 5308 (1), pp. 1-109 : 31-33

Herbuloracotis subgen. n.

Type species: Racotis apodosima Prout, 1931

Genetic analyses (Text fig. 2). The results of the genetic and morphological analyses of this lineage do not show a fully congruent picture. The morphologically well-characterised subgenus was revealed as paraphyletic in the ML analysis where one cluster comprised the A. apodosima species-group (mossy-green species) and true A. argillacea ( Warren, 1900) (ochreous-brown species) (BS: 9; PP: -), while the other lineage includes the monobasic A. deportata species-group (mossy-green species) and the A. longicornuta sp. n. and A. aristophanousi sp. n. species-pair (ochreous-brown species) (BS: 16; PP: -). To complicate the situation further, based on both genital and external morphology, A. argillacea (recovered in cluster A) and the A. longicornuta — aristophanousi species-pair (recovered in cluster B) form a clear morphological unit without any trait which would support the monophylies recovered in the ML and the polytomy in the BI analyses.As the paraphyly of the subgenus could not be resolved by morphological investigations, the infrasubgeneric classification of this subgenus is based merely on morphological characters disregarding the results of the genetic analyses. It is worth noting, however, that the morphology-based delineations of the A. apodosima (BS: 48; PP: 93) and A. deportata (BS: 23; PP: 66) species-groups are well-supported by the ML topology.

Diagnosis. The species of the subgenus are conspicuously bright mossy-green or ochreous-yellow, rendering them easily distinguishable from the much darker, rather greyish-green or brownish-grey species of other Afroracotis subgenera. The subgenus is characterized by the combination of the following traits: moderately long, distally tapered simple uncus; tubular, rather thick, inflated, somewhat expandable vesica bearing a single, robust distal cornutus erected at the base of vesica ejaculatorius; the cornutus in the A. apodosima species-group is highly regressed. The configuration of the cornutus is an important specific character. The female genitalia of the species of the subgenus are highly diverse interspecifically, with shared characters such as the medium long ovipositor, the short and narrow, moderately sclerotised antrum, the relatively long ductus bursae and the usually strongly sclerotised cervix bursae fully fused with the corpus bursae.

Description

External features of body and wings ( Figs 41–75, 79–80 , 151–172 ). Forewing length 18–25 mm. Male antenna bipectinate-fasciculate, rami short or moderately long, distal section filiform, length of which varying interspecifically between 2/5 and 1/2. Female antenna filiform or bipectinate-fasciculate (in A. argillacea , A. longicornuta sp. n. and A. aristophanousi sp. n.). Head moderately large, proboscis well-developed, labial palp short, length 1.5–2.0 times the diameter of eye, porrect, first and second segment dilated, third segment rather short and thin. Compound eye moderately large. Frons, vertex, collar (patagium), tegula, thorax and abdomen concolorous with wing upperside. Legs rather long, colouration as of wing underside, index of spurs 0-2-4. Forewing broad, triangular, costa straight, basal and apical section gently curved, termen slightly arched, moderately scalloped, ventral margin barely arched. Ground colour varying interspecifically from bright mossy to lemon, speckled with groups of brownish scales. Basal and medial area often paler than terminal one, postmedial area with dark blotches of variable size between M3 and CuA1. Transverse lines diffuse, interrupted. Basal line deleted, antemedial line detectable, more or less straight, medial line shadow-like, moderately S-curved. Postmedial line gently arched, undulate, subterminal line represented by a series of patches of variable size, or deleted. Terminal line interrupted, consisting of small black dots, short dashes or arches between veins. Fringe (cilia) short, colour as of forewing, occasionally chequered. Discal spot obliterated, patch-like, round or ovoid. Hindwing apex rounded, termen evenly arcuate, moderately scalloped, anal margin straight. Ground colour as of forewing, densely speckled with groups of brownish scales. Basal line deleted, antemedial line diffuse, often rather broad, evenly arched, wavy. Medial line well-defined, evenly arched, undulate. Postmedial line diffuse, interrupted, subterminal line poorly visible or deleted. 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 suffused with greyish scales, distally with broad, dark postmedial band protruding to termen at upper third. Transverse lines present, discal spot well-defined.

Male genitalia ( Figs 232–252 ). Uncus short, simple, distally tapered, apically rounded. Tegumen short and broad, gnathos well-developed, with rounded medial plate. Valva short, broad at base, distally tapered, costal margin sclerotised, often setose, with short sclerotised apical arm expanding over the membranous ventral part of valva. Ventral margin of valva gently arched, medially often slightly concave, sparsely setose; sacculus short and broad, weakly sclerotised, without process. Juxta short, weakly sclerotised, shield-like without arms. Vinculum short or moderately long and broad, U- or V-shaped, in A. atriclava narrow, digitiform, apically rounded. Aedeagus medium thick, short, slightly curved, without carina. Vesica thick, relatively long in most species, very long and expandable in A. muscivirens sp. n., armed with a single robust cornutus; vesica with vestigial cornutus in A. apodosima species-group.

Female genitalia ( Figs 306–318 ). Ovipositor medium long, papilla analis narrow triangular, apically rounded, sparsely setose, apophysis posterioris moderately long. Eighth tergite medium long, trapezoidal, distal margin more or less straight, proximal margin gently undulate, anterior apophysis shorter than posterior apophysis. Ostium bursae broad, membranous, antrum moderately sclerotised, short, cup- or goblet-shaped. Ductus bursae membranous, short and narrow. Cervix bursae well-developed, heavily sclerotised, often elongate tubular, being direct continuation of ductus bursae (in several species of A. argillacea species-group) or forming a rounded bulge of ductus bursae (in A. apodosima and A. deportata species-groups) or may be fully merged with corpus bursae forming a sclerotised section of bursa copulatrix. Corpus bursae varied in size interspecifically, may be highly reduced and fused with cervix bursae or well-developed and separated from cervix; may bear a strongly sclerotised disc-shaped plate supposedly serving as receptive surface of cornutus vesicae during copulation; signum absent.

Key to the species-groups of the subgenus Herbuloracotis based on male genitalia characters

1. Vesica with a weakly sclerotised, small cornutus or without it................................... A. apodosima sp.-gr.

-. Vesica with a large, acute cornutus........................................................................ 2

2. Uncus short, broad...................................................................... A. deportata sp.-gr.

-. Uncus elongate, narrow................................................................. A. argillacea sp.-gr.

Figures

FIGURES 1–20. Adults, dorsal view. 1. Racotis boarmiaria, LT, ♁, India (NHMUK); 2. Afroracotis squalida squalida, HT of Ophthalmodes squalida, ♀, Nigeria (NHMUK); 3. Ibidem, HT of Boarmia uhligi, ♀, Tanzania (MfN); 4. Ibidem, ♁, Cameroon (ANHRT); 5. Ibidem, ♁, Zambia (ANHRT); 6. Ibidem, ♁, Togo (ANHRT); 7. Ibidem, ♁, Gabon (ANHRT); 8. Ibidem, ♁, Ethiopia (RCDS); 9. Ibidem, ♀, Ethiopia (RCDS); 10. Ibidem, ♀, Cameroon (ANHRT); 11. A. s. voeltzkowii, LT of Boarmia voeltzkowii, ♁, Madagascar (MfN); 12. Ibidem, ♁, Comoros (ZSM); 13. Ibidem, ♁, Madagascar (NHMUK); 14. Ibidem, ♀, Madagascar (NHMUK); 15. A. s. thomensis ssp. n., HT, ♁, São Tomé (ANHRT); 16. Ibidem, PT, ♀, São Tomé (ANHRT); 17. Ibidem, ♀, PT, São Tomé (NHMUK); 18. A. incompletaria incompletaria, ♀, Réunion (ZSM); 19. A. i. atrilunaria, HT of Boarmia atrilunaria, ♀, Mauritius (NHMUK); 20. Ibidem, ♀, Mauritius (ANHRT).

FIGURES 41–60. Adults, dorsal view. 41. Afroracotis apodosima, HT of Racotis apodosima, ♁, Mozambique (NHMUK); 42. Ibidem, ♁, Tanzania (ZSM); 43. Ibidem, ♁, Zambia (ANHRT); 44. Ibidem, ♀, Mozambique (ANHRT); 45. A. dargei sp. n., HT, ♁, Cameroon (ZSM); 46. Ibidem, PT, ♁, Cameroon (ZSM); 47. Ibidem, PT, ♀, Cameroon (ZSM); 48. A. deportata, ♁, Madagascar (RCHS); 49. Ibidem, HT of Racotis deportata, ♁, Madagascar (MNHN); 50. Ibidem, ♀, Comoros (NHMUK); 51. A. argillacea argillacea, HT of Alcis argillacea, ♁, DRC (NHMUK); 52. Ibidem, ♁, DRC (NHMUK); 53. Ibidem, ♁, DRC (ZSM); 54. Ibidem, ♁, Cameroon (RCMO); 55. A. argillacea cedrici, HT of Racotis cedrici, ♀, Bioko (ZSM); 56. Ibidem, PT, ♀, Bioko (ZSM); 57. Ibidem, ♀, Bioko (RCTK); 58. A. argillacea morettoi ssp. n., HT, ♁, Ivory Coast (ANHRT); 59. Ibidem, PT, ♀, Ivory Coast (ANHRT); 60. Ibidem, PT, ♀, Ivory Coast (ZSM).

FIGURES 61–80. Adults, dorsal view. 61. Afroracotis longicornuta sp. n., HT, ♁, Liberia (ANHRT); 62. Ibidem, PT, ♁, Guinea (ANHRT); 63. A. l. congolana ssp. n., HT, ♁, Cameroon (RCMO); 64. Ibidem, PT, ♁, Cameroon (ZSM); 65. Ibidem, PT, ♁, DRC (NHMUK); 66. Ibidem, PT, ♀, Cameroon (ZSM); 67. A. l. ugandana ssp. n., HT, ♁, Uganda (RCHS); 68. Ibidem, PT, ♀, Uganda (RCHS); 69. A. aristophanousi sp. n., HT, ♁, Liberia (ANHRT); 70. Ibidem, PT, ♁, Cameroon (ZSM); 71. Ibidem, PT, ♁, Uganda (NHMUK); 72. Ibidem, PT, ♀, Cameroon (ZSM); 73. A. muscivirens sp. n., HT, ♁, Liberia (ZSM); 74. Ibidem, PT, ♁, Ivory Coast (ZSM); 75. Ibidem, PT, ♀, Liberia (ANHRT); 76. A. turlini sp. n., HT, ♁, Rwanda (ZSM); 77. Ibidem, PT, ♁, Rwanda (ZSM); 78. Ibidem, PT, ♀, Rwanda (ZSM); 79. A. atriclava, ST of Cleora atriclava, ♁, Madagascar (NHMUK); 80. Ibidem, ♀, Madagascar (ZSM).

FIGURES 151–170. Adults, ventral view. 151. Afroracotis apodosima, ♁, Zambia (ANHRT); 152. Ibidem, ♀, Mozambique (ANHRT); 153. A. dargei sp. n., PT, ♁, Cameroon (ZSM); 154. Ibidem, ♀, Cameroon (ZSM); 155. A. deportata, ♁, Madagascar (RCHS); 156. Ibidem, ♀, Comoros (NHMUK); 157. A. argillacea argillacea, ♁, Cameroon (ZSM); 158. Ibidem, ♁, Cameroon (ZSM); 159. A. argillacea cedrici, HT of Racotis cedrici, ♀, Bioko (ZSM); 160. A. argillacea morettoi ssp. n., HT, ♁, Ivory Coast (ANHRT); 161. Ibidem, PT, ♀, Ivory Coast (ANHRT); 162. A. longicornuta sp. n., HT, ♁, Liberia (ANHRT); 163. Ibidem, PT, ♁, Guinea (ZSM); 164. A. l. congolana ssp. n., PT, ♁, Cameroon (ZSM); 165. Ibidem, PT, ♀, Cameroon (ZSM); 166. A. l. ugandana ssp. n., PT, ♀, Uganda (RCHS); 167. A. aristophanousi sp. n., HT, ♁, Liberia (ANHRT); 168. Ibidem, PT, ♀, Cameroon (ZSM); 169. A. muscivirens sp. n., HT, ♁, Liberia (ZSM); 170. Ibidem, PT, ♀, Liberia (ANHRT).

FIGURES 171–190. Adults, ventral view. 171. Afroracotis atriclava, ST of Cleora atriclava, ♁, Madagascar (ZSM); 172. Ibidem, ST, ♀, Madagascar (ZSM); 173. A. turlini sp. n., PT, ♁, Rwanda (ZSM); 174. Ibidem, PT, ♀, Rwanda (ZSM); 175. A. zebrina, LT of Racotis zebrina, ♁, Nigeria (NHMUK); 176. A. zebrina, ♁, Guinea (ANHRT); 177. Ibidem, ♁, Liberia (ANHRT); 178. Ibidem, ♀, Ivory Coast (ANHRT); 179. A. chaineyi sp. n., HT, ♁, Gabon (ANHRT); 180. Ibidem, PT, ♁, Cameroon (ZSM); 181. Ibidem, PT, ♀, Bioko (ZSM); 182. A. lydiae lydiae sp. n., PT, ♁, Liberia (ANHRT); 183. Ibidem, PT, ♀, Guinea (ANHRT); 184. A. lydiae orientalis ssp. n., HT, ♁, Tanzania (ANHRT); 185. Ibidem, PT, ♀, Tanzania (ANHRT); 186. A. madagascariensis, ♁, Madagascar (NHMUK); 187. Ibidem, ♀, Madagascar (ANHRT); 188. A. smithi sp. n., HT, ♁, Zambia (ANHRT); 189. Ibidem, PT, ♀, Zambia (ANHRT); 190. Ibidem, PT, ♀, Cameroon (ZSM).

FIGURES 225–232. Male genitalia, a: clasping apparatus, b: aedeagus. 225. Afroracotis stadiei sp. n., PT, Rwanda, slide No. LG 4789 (ZSM); 226. A. violetteae sp. n., HT, Guinea, slide No. LG 5433 (ANHRT); 227. Ibidem, PT, Liberia, slide No. LG 4999 (ANHRT); 228. A. fiebigi sp. n., HT, Liberia, slide No. LG 5000 (ANHRT); 229. Ibidem, PT, Cameroon, slide No. LG 5070 (ZSM); 230. Ibidem, PT, Uganda, slide No. LG 4106 (NHMUK); 231. A. albitrigonis, São Tomé, slide No. LG 4800 (ANHRT); 232. A. apodosima, Mozambique, slide No. LG 4827 (ANHRT).

FIGURES 233–238. Male genitalia, a: clasping apparatus, b: aedeagus.; 233. Afroracotis apodosima, Zambia, slide No. LG 4930 (ANHRT); 234. Ibidem, Gabon, slide No. LG 4770 (ZSM); 235. A. dargei sp. n., HT, Cameroon, slide No. LG 4615 (ZSM); 236. Ibidem, PT, Cameroon, slide No. LG 5041 (ZSM); 237. A. deportata, Madagascar, slide No. LG 4067 (ANHRT); 238. A. argillacea argillacea, DRC, slide No. NHMUK 010315180 (NHMUK).

FIGURES 239–244. Male genitalia, a: clasping apparatus, b: aedeagus. 239. Afroracotis argillacea argillacea, Cameroon, slide No. LG 4405 (ZSM); 240. A. a. morettoi ssp. n., HT, Ivory Coast, slide No. LG 5416 (ANHRT); 241. A. longicornuta longicornuta sp. n., HT, Liberia, slide No. LG 5064 (ANHRT); 242. Ibidem, Guinea, slide No. LG 4424 (ZSM); 243. A. l. congolana ssp. n., HT Cameroon, slide No. LG 4088 (RCMO); 244. Ibidem, PT, Cameroon, slide No. LG 5043 (ZSM).

FIGURES 245–250. Male genitalia, a: clasping apparatus, b: aedeagus. 245. Afroracotis longicornuta congolana ssp. n., PT, DRC, slide No. NHMUK 010315179 (NHMUK); 246. A. l. ugandana ssp. n., HT, Uganda, slide No. LG 5529 (RCHS). 247. Ibidem, PT, Uganda, slide No. LG 5530 (RCHS); 248. A. aristophanousi sp. n., HT, Liberia, slide No. LG 5418 (ANHRT); 249. Ibidem, PT, Uganda, slide No. NHMUK 010315182 (NHMUK); 250. A. muscivirens sp. n., HT, Liberia, slide No. LG 4758 (ZSM).

FIGURES 251–255. Male genitalia, a: clasping apparatus, b: aedeagus. 251. Afroracotis muscivirens sp. n., PT, Ivory Coast, slide No. LG 4617 (ZSM); 252. A. atriclava, ST of Cleora atriclava, Madagascar, slide No. LG 5039 (ZSM); 253. A. turlini sp. n., HT, Rwanda, slide No. LG 4619 (ZSM). 254. Afroracotis zebrina, Ivory Coast, slide No. LG 4176 (ANHRT); 255. Ibidem, Liberia, slide No. LG 4965 (ANHRT).

FIGURES 306–313. Female genitalia. 306. Afroracotis apodosima, Tanzania, slide No.: LG 4767 (ZSM); 307. Ibidem, Mozambique, slide No.: LG 4826 (ANHRT); 308. A. dargei sp. n., PT, Cameroon, slide No.: LG 4771 (ZSM); 309. A. deportata, Madagascar, slide No.: LG 4082 (ANHRT); 310. A. argillacea cedrici, PT, Bioko, slide No.: HERB 7255 (ZSM); 311. A. a. morettoi ssp. n., PT, Ivory Coast, slide No.: LG 5417 (ANHRT); 312. A. longicornuta congolana ssp. n., PT, Cameroon, slide No.: LG 5044 (ZSM); 313. A. l. ugandana ssp. n., PT, Uganda, gen. slide No.: LG 5531 (RCHS).

FIGURES 314–319. Female genitalia. 314. Afroracotis aristophanousi sp. n., PT, Ghana, slide No.: NHMUK 010316165 (NHMUK); 315. Ibidem, PT, Cameroon, slide No.: LG 5045 (ZSM); 316. Ibidem, PT, Cameroon, slide No.: LG 5849 (RMCA); 317. A. atriclava, ST, Madagascar, slide No.: LG 5040 (ZSM); 318. A. muscivirens sp. n., PT, Liberia, slide No.: LG 5430 (ANHRT); 319. A. turlini sp. n., PT, Rwanda, slide No.: LG 4773 (ZSM).