Abstract
The Afrotropical taxa of the Racotis s.l. generic complex are revised utilising integrative taxonomical methods. Based on the evaluation of genital morphology and analyses of DNA barcodes, a new genus, Afroracotis gen. n. is established to include the Afrotropical “Racotis” species. The new genus is subdivided into 5 subgenera: Afroracotis subgen. n., Herbuloracotis subgen. n., Rwandaracotis subgen. n., Zebracotis subgen. n. and Sokokeracotis subgen. n.. A new monotypic genus is described to include Boarmia ugandaria Swinhoe, 1904 which was combined earlier with the genus Chorodna and recently with Racotis: Chorocotis gen. n.. Two species formerly assigned to Cleora are moved to Afroracotis: A. albitrigonis (Prout, 1927) comb. n., A. atriclava (Prout, 1926) comb. n.. Two species are transferred from Racotis to Colocleora: C. breijeri (Prout, 1922) comb. n., C. incauta (Prout, 1916) comb. n.. Seventeen new Afroracotis species (A. aliena, A. stadiei, A. violetteae, A. fiebigi, A. turlini, A. dargei, A. longicornuta, A. aristophanousi, A. muscivirens, A. chaineyi, A. lydiae, A. smithi, A. ochsei, A. milesi, A. helicalis, A. takanoi and A. staudei spp. n.) and 5 new subspecies (A. squalida thomensis, A. argillacea morettoi, A. longicornuta congolana, A. longicornuta ugandana and A. lydiae orientalis sspp. n.) are described, totalling 27 species and 8 subspecies contained in the genus Afroracotis. Adults and genitalia of all taxa are illustrated in 210 colour and 129 black and white figures demonstrating the intraspecific variability. The distribution of all taxa is illustrated in 6 dot maps. The results of the genetic analyses are figured in four phylograms.
References
- Aurivillius, C. (1925) s.n. Wissenschaftliche Ergebnisse der zweiten Deutschen Zentralafrika-Expedition 1910–1911 unter Führung Adolf Friedrichs, Herzog zu Mecklenburg, Band 1 Zoologie, Lieferung 18, Lepidoptera IV., 1 (18), 1243–1359, pls. 49–50.
- Booth, A.H. (1958) The Niger, the Volta and the Dahomey Gap as Geographic Barriers. Evolution, 12 (1), 48–62. DOI: https://doi.org/10.1111/j.1558-5646.1958.tb02927.x
- Butler, A.G. (1878) Descriptions of some new genera and species of Lepidoptera from Old Calabar and Madagascar. Annals and Magazine of Natural History, Series 5, 2 (12), 455–465. DOI: https://doi.org/10.1080/00222937808682457
- Chainey, J. & Karisch, T. (2017) A new species of Racotis Moore, [1887] (Lepidoptera: Geometridae: Ennominae) from Madagascar. Metamorphosis, 28, 49–53.
- Curtis, J. (1825) British Entomology: being illustrations and descriptions of the genera and insects found in Great Britain and Ireland: containing coloured figures from nature of the most rare and beautiful species, and in many instances of the plants upon which they are found. Vol. VI. printed for the author, London, 170 pp.
- Guenée, A.M. (1858) Uranides et Phalénites Tome I. In: Boisduval J.A.B & Guenée, A. Histoire naturelle des insectes. Spécies general des Lépidoptères, 9, pp. 1–xxxvii + 1–514.
- Guenée, A.M. (1862) Lépidoptères. In: Maillard, L., Notes sur l’île de la Réunion (Bourbon), Annexe G, pp. 1–72, pls. 12–13.
- Hacker, H.H. (2019) Biogeography; Boletobiinae (Lepidoptera, Noctuoidea, Erebidae). Moths of Africa. Systematic and illustrated Catalogue of the Heterocera (Lepidoptera) of Africa, 1, 1–816.
- Hebert, P.D.N., Braukmann, T.W.A, Prosser, S.W.J., Ratnasingham, S., deWaard, J.R., Ivanova, N.V., Janzen, D.H., Hallwachs, W., Naik, S., Sones, J.E. & Zakharov, E.V. (2018) A sequel to Sanger: amplicon sequencing that scales. BMC Genomics, 19, 219. https://doi.org/10.1186/s12864-018-4611-3 DOI: https://doi.org/10.1186/s12864-018-4611-3
- Herbulot, C. (1970) Lepidoptera Geometridae du Tsaratanana (Madagascar Nord). Mémoires O.R.S.T.O.M., 37, 157–166.
- Herbulot, C. (1973) Nouveaux Ennominae du Cameroun (Lepidoptera, Geometridae). Annales de la Faculté des Sciences du Cameroun, 1973 (14), 141–148.
- Herbulot, C. (1991) Un nouveau Racotis africain (Lepidoptera Geometridae). Bulletin de la Société entomologique de Mulhouse, 21 (1), 16.
- Herbulot, C. (1998) Lépidoptères récoltés par le Dr. J.-G. Canu dans l’île de Bioko (ex Fernando Poo) (Lepidoptera). Lambillionea, 98 (2), 287–292.
- Holloway, J.D. (1994) The moths of Borneo: family Geometridae, subfamily Ennominae, Part 11. Malayan Nature Journal, 47, 1–309, pls. 1–19, 593 figs.
- Ivanova, N.V., Dewaard, J.R. & Hebert, P.D.N. (2006) An inexpensive, automation-friendly protocol for recovering high-quality DNA. Molecular Ecology Notes, 6, 998–1002. https://doi.org/10.1111/j.1471-8286.2006.01428.x DOI: https://doi.org/10.1111/j.1471-8286.2006.01428.x
- Jiang, N., Li, X., Hausmann, A., Cheng, R., Xue, D. & Han, H. (2017) A molecular phylogeny of the Palaearctic and Oriental members of the tribe Boarmiini (Lepidoptera : Geometridae : Ennominae). Invertebrate Systematics, 31, 427–441. https://doi.org/10.1071/IS17005 DOI: https://doi.org/10.1071/IS17005
- Kimura, M. (1980) A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16, 111–120. DOI: https://doi.org/10.1007/BF01731581
- Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Molecular Biology and Evolution, 35, 1547–1549. DOI: https://doi.org/10.1093/molbev/msy096
- Lafontaine, J.D. & Mikkola, K. (1987) Lock-and-key system in the inner genitalia of Noctuidae (Lepidoptera) as taxonomic character. Entomologiske Meddelelser, 55, 161–167.
- László, G.M. & Hausmann, A. (2022) Taxonomic review of the genus Morabia Hausmann & Tujuba, 2020 with descriptions of two new species and introducing five new generic combinations (Lepidoptera, Geometridae, Ennominae). Zootaxa, 5134 (2), 215–237. https://doi.org/10.11646/zootaxa.5134.2.3 DOI: https://doi.org/10.11646/zootaxa.5134.2.3
- Mabille, P. (1893) Descriptions de Lépidoptères nouveaux. Annales de la Société entomologique de Belgique, 37, 50–65.
- Moore, F. (1887) s.n. In: The Lepidoptera of Ceylon, London, 3, pp. 1–578, pls. 144–215.
- Murillo-Ramos, L., Chazot, N., Sihvonen, P., Õunap, E., Jiang, N., Han, H., Clarke, J.T., Davis, R.B., Tammaru, T. & Wahlberg, N. (2021) Molecular phylogeny, classification, biogeography and diversification patterns of a diverse group of moths (Geometridae, Boarmiini). Molecular Phylogenetics and Evolution, 162 (2021), 107198, 1–10. DOI: https://doi.org/10.1016/j.ympev.2021.107198
- Pagenstecher, A. (1907) Lepidoptera—Heterocera (Uraniidae, Geometridae, Noctuidae, Pyralidae, Thyrididae, Tortricidae, Tineidae, Pterophoridae) von Madagaskar, den Comoren und Ostafrika. In: Voeltzkow, A., Reise in Ostafrika in den Jahren 1903–05. Wissenschaftliche Ergebnisse 2. Systematische Arbeiten, 2, pp. 93–146.
- Prout, L.B. (1916) New South African Geometridae. Annals of the Transvaal Museum, 5 (3), 151–179.
- Prout, L.B. (1922) Geometridae. In: Prout, L.B. & Prout, A.E., New South African Heterocera. Annals of the Transvaal Museum, 8 (3), pp. 149–186.
- Prout, L.B. (1926) New Geometridae in the Tring Museum. Novitates Zoologicae, 33, 179–188. DOI: https://doi.org/10.5962/bhl.part.21147
- Prout, L.B. (1927) A list of the Geometridae (Lep. Het.) known to occur in the Island of Saõ Thomé, with descriptions of some new species collected by Mr. T.A. Barns. Transactions of the entomological Society of London, 75 (1), 187–200. DOI: https://doi.org/10.1111/j.1365-2311.1927.tb00071.x
- Prout, L.B. (1931) Geometrid descriptions and notes. Novitates Zoologicae, 36 (2), 151–180. DOI: https://doi.org/10.5962/bhl.part.10110
- Prout, L.B. (1929–1938) Die Spannerartigen Nachtfalter (Geometridae). Die Afrikanischen Spanner. Geometridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde. Eine Systematische Bearbeitung der bis jetzt bekannten Gross–Schmetterlinge, 16, pp. 1–160.
- Pyrcz, T.W., Collins, S., Lachowska-Cierlik, D., Lees, D.C., Sáfián, Sz. & Florczyk, K. (2020) A Malagasy element in Continental Africa: a new subspecies of the rare Amauris nossima (Nymphalidae, Danainae) from the Kenyan coast. African Zoology, 2020, 1–13. https://doi.org/10.1080/15627020.2020.1824588 DOI: https://doi.org/10.1080/15627020.2020.1824588
- Rajaei, H., Hausmann, A., Scoble, M., Wanke, D., Plotkin, D., Brehm, G., Murillo-Ramos, L. & Sihvonen, P. (2022) An online taxonomic facility of Geometridae (Lepidoptera), with an overview of global species richness and systematics. Integrative Systematics, 5 (2), 145–192. https://doi.org/10.18476/2022.577933 DOI: https://doi.org/10.18476/2022.577933
- Ronquist, F., Teslenko, M., Van Der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61 (3), 539–542. https://doi.org/10.1093/sysbio/sys029 DOI: https://doi.org/10.1093/sysbio/sys029
- Rögl, F. (1999) Mediterranean and Paratethys. Facts and hypotheses of an Oligocene to Miocene paleogeography (short overview). Geologica Carpathica, 50 (4), 339–349.
- Sato, R. (1995) The genus Racotis Moore (Lepidoptera: Geometridae) from the Philippines, with descriptions of two new species. Tinea, 14, 89–92.
- Sato, R. (2004) Taxonomic notes on Racotis boarmiaria (Guenée) (Geometridae, Ennominae) and its allies from the Indo-Malayan region, with descriptions of four new species. Tinea, 18 (2), 130–139.
- Scoble, M.J. (1999) Geometrid Moths of the World: A Catalogue. CSIRO Publishing, Collingwood, Stenstrup, 1016 pp. https://doi.org/10.1071/9780643101050 DOI: https://doi.org/10.1071/9780643101050
- Scoble, M.J. & Hausmann, A. (2007) Online list of valid and available names of the Geometridae of the world. Available from: http://www.lepbarcoding.org/geometridae/species_checklists.php (accessed 31 May 2023)
- Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology, 57 (5), 758–771. https://doi.org/10.1080/10635150802429642 DOI: https://doi.org/10.1080/10635150802429642
- Strand, E. (1909) Lepidoptera aus Deutsch–Ost–Afrika, gesammelt von Herrn Dr. C. Uhlig. Internationale entomologische Zeitschrift, Guben, 3 (23), 128–130.
- Swinhoe, C. (1904) On the Geometridae of tropical Africa in the National Collection. Transactions of the entomological Society of London, 1904 (3), 497–590. DOI: https://doi.org/10.1111/j.1365-2311.1904.tb02755.x
- Toussaint, E.F.A., Vila, R., Yago, M., Chiba, H., Warren, A.D., Aduse-Poku, K., Storer, C., Dexter, K.M., Maruyama, K., Lohman, D.J. & Kawahara, A.Y. (2019) Out of the Orient: Post-Tethyan transoceanic and trans-Arabian routes fostered the spread of Baorini skippers in the Afrotropics. Systematic Entomology, 44, 926–938. https://doi.org/10.1111/syen.12365 DOI: https://doi.org/10.1111/syen.12365
- Tujuba, T.F., Hausmann, A. & Sciarretta, A. (2020) Revision of the Orbamia Herbulot, 1966 group of genera with description of two new genera, ten new species, and two new subspecies (Lepidoptera, Geometridae, Ennominae, Cassymini). ZooKeys, 929, 53–77. https://doi.org/10.3897/zookeys.929.50391 DOI: https://doi.org/10.3897/zookeys.929.50391
- Warren, W. (1894) New genera and species of Geometridae. Novitates Zoologicae, 1, 366–466. DOI: https://doi.org/10.5962/bhl.part.24566
- Warren, W. (1897) New genera and species of moths from the Old–World regions in the Tring Museum. Novitates Zoologicae, 4, 12–130. DOI: https://doi.org/10.5962/bhl.part.21182
- Warren, W. (1899) New species and genera of the families Drepanulidae, Thyrididae, Uraniidae, Epiplemidae, and Geometridae from the Old-World regions. Novitates Zoologicae, 6 (1), 1–66.
- Warren, W. (1900) New genera and species of Thyrididae and Geometridae from Africa. Novitates Zoologicae, 7, 90–98.