Keywords
apomorphy
gnathos
venation
genitalia
How to Cite
Abstract
Metarbelidae are an intriguing group of Cossoid moths that have recently received considerable attention. Although several new genera have been described from the Oriental region in recent years, the morphological diversity of the family remains poorly investigated. In addition, the unusual genus Psychidarbela and the rarely encountered diurnal family Ratardidae share several features overlapping with Metarbelidae. Consequently, Oriental Metarbelidae have remained difficult to delimit, and apomorphic characters are challenging to identify. In this study, we address this gap through an examination of the morphological characters of all Oriental genera. Using a maximum parsimony cladistic analysis based on 39 characters, we find Ratardidae nested within Metarbelidae. Accordingly, we synonymize the two families and recognize Ratardidae, stat. nov., following the ICZN principles of priority. Furthermore, we establish the family-group name Psychidarbelidae Roepke, 1938, stat. nov., for the genus Psychidarbela. Detailed morphological features supporting this arrangement are discussed. This proposal aims to improve diagnostic clarity and stabilize classification; however, it is tentative, and a comprehensive molecular analysis will be required to finalize broader phylogenetic relationships.
References
Chen Q, Deng M, Dai X, Wang W, Wang X, Chen L-S, Huang G-H (2025) Phylogenomic data exploration with increased sampling provides new insights into the higher-level relationships of butterflies and moths (Lepidoptera). Journal of Systematics and Evolution 63: 1133–1153. https://doi.org/10.1111/jse.13180
Choochuen T, Foit J (2025) Cossid moths (Lepidoptera: Cossidae) as pests of woody plants – A review. Agricultural and Forest Entomology 27: 512–531. https://doi.org/10.1111/afe.12689
Edwards ED, Gentili P, Horak M, Kristensen NP, Nielsen ES (1998) The cossoid/sesioid assemblage. In: Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Systematics, and Biogeography. Handbuch der Zoologie/Handbook of Zoology. Volume IV Arthropoda: Insecta Part 35. https://doi.org/10.1515/9783110804744.181
Heikkilä M, Mutanen M, Wahlberg N, Sihvonen P, Kaila L (2015) Elusive ditrysian phylogeny: an account of combining systematized morphology with molecular data (Lepidoptera). BMC Evolutionary Biology 15: 260. https://doi.org/10.1186/s12862-015-0520-0
Heppner JB, Matusik DD (2017) A new species of Ratarda from lowland Cambodia (Ratardidae). Lepidoptera Novae 10(1–2): 21–24.
Holloway JD (1986) The Moths of Borneo: Key to Families: Families Cossidae, Metarbelidae, Ratardidae, Dudgeonidae, Epipyropidae and Limacodidae. Malayan Nature Journal 40: 1–166.
Kawahara AY, Plotkin D, Espeland M, Meusemann K, Toussaint EFA, Donath A, Gimnich F, Frandsen PB, Zwick A, dos Reis M, Barber JR, Peters RS, Liu S, Zhou X, Mayer C, Podsiadlowski L, Storer C, Yack JE, Misof B, Breinholt JW (2019) Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences 116(45): 22657–22663. https://doi.org/10.1073/pnas.1907847116
Kobes LWR, Ronkay L (1990) The Ratardidae of Sumatra. Heterocera Sumatrana 6: 79–100. https://doi.org/10.1086/293219
Lehmann I (2008) Ten new species of Metarbelidae (Lepidoptera: Cossoidea) from the coastal forests and the Eastern Arc Mountains of Kenya and Tanzania, including one species from two upland forests. Journal of East African Natural History 97(1): 43–82. https://doi.org/10.2982/0012-8317
Lehmann I (2009) Six new species of Metarbelidae (Lepidoptera: Cossoidea) from the Eastern Arc Mountains of Tanzania, including one new species from Marenji Forest in southeast coastal Kenya. Journal of East African Natural History 97(2): 187–206. https://doi.org/10.2982/0012-8317-97.2.187
Lehmann I (2010a) A new genus of Metarbelidae (Lepidoptera: Cossoidea) from the Afrotropical Region with the description of seven new species. Esperiana Memoir 5: 294–321.
Lehmann I (2010b) A revision of the genus Arbelodes Karsch (Lepidoptera: Cossoidea: Metarbelidae) from southeast-central and southern Africa with the description of thirteen new species. Published by the author, Hamburg & Wismar, 3–81+ 8 b/w pls., 5 colour pls. https://doi.org/10.5962/bhl.title.79419
Lehmann I (2011) The description of a new genus and twenty-three new species of Metarbelidae (Lepidoptera: Cossoidea) from the lowland tropical rain forests of the Guineo-Congolian Region with notes on habitats and biogeography. Published by the author, Hamburg, 1–67 + 10 b/w pls., 6 colour pls., 1 coloured map. https://doi.org/10.5962/bhl.title.79417
Lehmann I (2012) Description of a new genus and species of Metarbelidae (Lepidoptera, Cossoidea) from the Albertine Rift region of Tanzania, East Africa. Norwegian Journal of Entomology 59: 234–240.
Lehmann I (2013) Description of two new genera and ten new species of Metarbelidae (Lepidoptera: Cossoidea) from western, north-central and eastern Africa with notes on habitats and biogeography. Published by the author, S&K NEUE Hamburger Digitaldruck + Medien GmbH, Hamburg, 82 pp. https://doi.org/10.5962/bhl.title.79421
Lehmann I (2014) Description of two new genera and two new species of Metarbelidae (Lepidoptera, Cossoidea) from the Northeastern Congolian Lowland Forests Ecoregion (Central Africa). Zootaxa 3895(4): 570–580. http://doi.org/10.11646/zootaxa.3895.4.6
Lehmann I (2019a) First revision of the family Metarbelidae Strand, 1909 (Lepidoptera, Cossoidea Leach, 1815) and a phylogeny based on adult morphology of 60 genera from the Afrotropical and Oriental Region. Bonn, 397 pp.
Lehmann I (2019b) Description of two new genera and two new species of Metarbelidae (Lepidoptera, Cossoidea) from Nepal and Sumatra (Indonesia), Oriental Region. Heterocera Sumatrana 13(2): 47–72.
Lehmann I (2025) The new Genera Kayamuhakaia, Finsterwaldeia, Lichterfeldia (Lepidoptera: Cossoidea: Metarbelidae Strand, 1909) with thirty-six new species from lowland, submontane and montane areas of eastern, central and south-central Africa (Afrotropical Region). Published by the author, Berlin, 347 pp.
Lehmann I, Dalsgaard Th (2023) Revision of Saalmulleria Mabille, 1891 (Lepidoptera, Metarbelidae) from Madagascar with the description of three new genera and fifteen new species. Evolutionary Systematics 7: 133–182. http://doi.org/10.3897/evolsyst.7.85204
Lehmann I, Rajaei H (2013) Description of a new genus and three new species of Metarbelidae (Lepidoptera: Cossoidea) from East and central Africa, with notes on biogeography. Bonn zoological Bulletin 62(1): 100–110. https://doi.org/10.5962/bhl.title.79421
Lehmann I, Roberts M, Mathiu JM, Butynski TM (2018) Undescribed species and genera of Metarbelidae moths (Lepidoptera, Cossoidea) from Lolldaiga Hills Ranch, central Kenya. Newsletter Lolldaiga Hills Research Programme, January–February 2018: 3–4.
Lehmann I, Zahiri R, Husemann M (2023) Revision of the Metarbelodes Strand, 1909 genus-group (Lepidoptera: Cossoidea: Metarbelidae) with descriptions of two new genera and 33 new species from high elevations of eastern and southern Africa. Zootaxa 5267(1): 1–106. http://doi.org/10.11646/zootaxa.5267.1.1
Mayer C, Dietz L, Call E, Kukowka S, Martin S, Espeland M (2021) Adding leaves to the Lepidoptera tree: capturing hundreds of nuclear genes from old museum specimens. Systematic Entomology 46: 649–671. https://doi.org/10.1111/syen.12481
Mey W (2005) Metarbela naumanni sp. nov. from southern Africa (Lepidoptera, Cossidae: Metarbelidae). Entomologische Zeitschrift 115(1): 10–12.
Mey W (2012) Arbelodes goellnerae sp. nov. aus dem südlichen Namibia (Lepidoptera: Cossoidea, Metarbelidae). Entomologische Zeitschrift 122(3): 103–105.
Mey W (2018) Dukearbela translucens gen. nov., spec. nov. – a remarkable taxon from South Africa (Lepidoptera: Metarbelidae). Metamorphosis 28: 11–13. https://doi.org/10.4314/met.v29i1.5
Mey W (2024) New records of Metarbelidae from Namibia and description of a new species from the Great Escarpment (Lepidoptera, Cossoidea). Metamorphosis 35: 25–30. https://doi.org/10.4314/met.v35i1.6
Minet J (1991) Tentative reconstruction of the ditrysian phylogeny (Lepidoptera: Glossata). Entomologica scandinavica 22: 69–95. https://doi.org/10.1163/187631291X00327
Mutanen M, Wahlberg N, Kaila L (2010) Comprehensive gene and taxon coverage elucidates radiation patterns in moths and butterflies. Proceedings of the Royal Society B: Biological Sciences 277(1695): 2839–2848. https://doi.org/10.1098/rspb.2010.0392
Nonaka T (2021) Discovery of a new genus and species of Limacodidae (Lepidoptera) from Laos. Tinea 26(1): 60−62.
Owada M (1993) The Systematic Position of Shisa excellens (Lepidoptera and Ratardidae). Japanese Journal of Entomology 61(2): 251–260.
Pavlova PD, Müller GC, Prozorov AM, Yakovlev RV (2025) Axelhausmannia – a new genus of Metarbelidae (Lepidoptera: Cossoidea) from the equatorial Africa. Ecologica Montenegrina 93: 36–42. https://doi.org/10.37828/em.2025.93.3
Pavlova PD, Müller GC, Prozorov AM, Yakovlev RV (2026a) A new Genus of the Family Metarbelidae (Lepidoptera: Cossoidea) from Central and Western Africa. Far Eastern Entomologist 542: 1–10. https://doi.org/10.25221/fee.542.1
Pavlova PD, László GM, Müller GC, Tujuba TF, Prozorov AM, Yakovlev RV (2026b) On the taxonomy of Salagena tesselata Distant, 1897 with description of a new genus and three new species (Lepidoptera, Cossoidea, Metarbelidae). Acta Biologica Sibirica 12: 85–98. https://doi.org/10.5281/zenodo.18337242
Roepke W (1938) Psychidarbela kalshoveni n. g., n. sp. (Lep., Cossidae). Entomologische mededeelingen van Nederlandsch-Indië 4(1): 22−26.
Roepke W (1957) The cossids of the Malay Region (Lepidoptera: Heterocera). Verhandelingen der Koninklijke Nederlandse Akademie van Weterschappen, Afd. Natuurkunde (Tweede Reeks) Deel LII (1): 1−60.
Rota J, Twort V, Chiocchio A, Peña C, Wheat CW, Kaila L, Wahlberg N (2022) The unresolved phylogenomic tree of butterflies and moths (Lepidoptera): assessing the potential causes and consequences. Systematic Entomology 47: 531–550. https://doi.org/10.1111/syen.12545
Schoorl JW (1990) A phylogenetic study on Cossidae (Lepidoptera: Ditrysia) based on external adult morphology. Zoologische Verhandelingen 263: 1−295.
Sugi S (1995) Ratardidae and Metarbelidae. In Haruta T (Ed.) Moths of Nepal. Part 4. Tinea 14(Suppl. 2): 117–118.
van Nieukerken EJ, Kaila L, Kitching IJ, Kristensen NP, Lees DC, Minet J, Mitter C, Mutanen M, Regier JC, Simonsen T J, Wahlberg N, Yen S-H, Zahiri R, Adamski D, Baixeras J, Bartsch D, Bengtsson BÅ, Brown JW, Bucheli SR, Davis DR, De Prins J, De Prins W, Epstein ME, Gentili-Poole P, Gielis C, Hättenschwiler P, Hausmann A, Holloway JD, Kallies A, Karsholt O, Kawahara AY, Koster S (JC), Kozlov MV, Lafontaine JD, Lamas G, Landry J-F, Lee S, Nuss M, Park K-T, Penz C, Rota J, Schintlmeister A, Schmidt BC, Sohn J-C, Solis MA, Tarmann GM, Warren AD, Weller S, Yakovlev RV, Zolotuhin VV, Zwick A (2011) Order Lepidoptera Linnaeus, 1758. In: Zhang Z-Q (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 212–221. https://doi.org/10.11646/zootaxa.3148.1.41
Wahlberg N, Wheat CW, Peña C (2013) Timing and Patterns in the Taxonomic Diversification of Lepidoptera (Butterflies and Moths). PLoS ONE 8(11): e80875. https://doi.org/10.1371/journal.pone.0080875
Wright CJ, Shirey VM, Condamine FL, Wahlberg N, Kawahara AY (2025) Evolution, genomics and conservation of butterflies and moths. Nature Reviews Biodiversity 2: 186–203. https://doi.org/10.1038/s44358-025-00128-8
Yakovlev RV (2018) World Catalogue of the Family Ratardidae (Lepidoptera, Cossoidea) with Description of a New Species from Thailand and Establishment of New Synonymy and a New Status for Ratarda javanica Roepke, 1937. Entomological Review 98(5): 644– 651. https://doi.org/10.1134/S001387381805007X
Yakovlev RV, Zolotuhin VV (2020) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. I. Introduction and genera Encaumaptera Hampson, 1893, Orgyarbela gen. nov., and Hollowarbela gen. nov. Ecologica Montenegrina 38: 84–101. https://doi.org/10.37828/em.2020.38.11
Yakovlev RV, Zolotuhin VV (2021a) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. II. Two monotypic genera – Ghatarbela gen. nov. and Micrarbela gen. nov. – from the Western Ghats and Sri Lanka biodiversity hotspot. Ecologica Montenegrina 42: 103–108. https://doi.org/10.37828/em.2021.42.6
Yakovlev RV, Zolotuhin VV (2021b) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. III. Genus Stueningeria Lehmann, 2019. Ecologica Montenegrina 43: 16–29. https://doi.org/10.37828/em.2021.43.2
Yakovlev RV, Zolotuhin VV (2021c) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. IV. Genus Tagoria Yakovlev & Zolotuhin, gen. nov. Ecologica Montenegrina 43: 38–43. https://doi.org/10.37828/em.2021.43.5
Yakovlev RV, Zolotuhin VV (2021d) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. V. Genus Marcopoloia Yakovlev & Zolotuhin, gen. nov. from Taiwan Island and Indo-Burma biodiversity hotspot. Ecologica Montenegrina 44: 103–108. https://doi.org/10.37828/em.2021.44.6
Yakovlev RV, Zolotuhin VV (2021e) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. VI. Genus Tearbela Yakovlev & Zolotuhin, gen. nov. from Bangladesh and Marcopoloia dea (Swinhoe, 1890), comb. nov. Ecologica Montenegrina 45: 43–47. https://doi.org/10.37828/em.2021.45.8
Yakovlev RV, Zolotuhin VV (2022) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. VII. Genus Aukorbela Yakovlev & Zolotuhin, gen. nov. from Central Vietnam. Ecologica Montenegrina 50: 67–70. https://doi.org/10.37828/em.2022.50.12
Yakovlev RV, de Vos, R, Hulsbosch R, Zolotuhin VV (2022a) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. VIII. Genus Lutzkobesia Lehmann, 2019. Ecologica Montenegrina 52: 49–52. https://doi.org/10.37828/em.2022.52.7
Yakovlev RV, Korshunov AV, Zolotuhin VV (2022b) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. IX. Description of Marcopoloia svetlanae sp. nov. from Central Thailand. Ecologica Montenegrina 53: 35–37. https://doi.org/10.37828/em.2022.53.4
Yakovlev RV, Hulsbosch R, Zolotuhin VV (2022c) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. X. Description of Stueningeria bretschneideri sp. nov. from the Andaman Islands (India). Ecologica Montenegrina 56: 10–13. https://doi.org/10.37828/em.2022.56.2
Yakovlev RV, de Vos R, Hulsbosch R (2022d) New genus of the family Ratardidae (Lepidoptera: Cossoidea) from New Guinea. Far Eastern Entomologists 452: 1–6. https://doi.org/10.25221/fee.452.1
Yakovlev RV, Hulsbosch R, Zolotuhin VV (2023) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. XI. Genus Psychidarbela Roepke, 1938. Ecologica Montenegrina 65: 51–59. https://doi.org/10.37828/em.2023.65.7
Yakovlev RV, Hulsbosch R (2024) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. XII. New species of Psychidarbela Roepke, 1938 from Vietnam. Ecologica Montenegrina 77: 110–112. https://doi.org/10.37828/em.2024.77.10
Yakovlev RV, Hulsbosch R, Ahmad J, Singh N (2025) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. XIII. Genus Indarbela Fletcher, 1922. Zootaxa 5661(4): 553–560. https://doi.org/10.11646/zootaxa.5661.4.5
Yakovlev RV, Müller GC, Prozorov AM, Pavlova PD (2026) Revision of the family Metarbelidae (Lepidoptera) of the Oriental Region. XIV. Genus Siamella gen. nov. from Northern Thailand. Ecologica Montenegrina 96: 30–34. https://doi.org/10.37828/em.2026.96.3
This work is licensed under a Creative Commons Attribution 4.0 International License.