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Type: Article
Published: 2018-06-26
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A non-destructive virtual dissection by micro-CT reveals diagnostic characters in the type specimen of Caloptilia stigmatella (Lepidoptera: Gracillariidae)

Hunterian Museum, University of Glasgow, G3 8AW, United Kingdom.
Electronic and Electrical Engineering, University of Strathclyde, Glasgow, G1 1RD, United Kingdom.
Museo Entomológico UNAB, Universidad Nacional de Colombia, Bogotá, Colombia.
Royal Belgian Institute of Natural Sciences, Brussels, Belgium.
Electronic and Electrical Engineering, University of Strathclyde, Glasgow, G1 1RD, United Kingdom.
Lepidoptera ampelipennella computed tomography genus-level diagnosis holotype nomen nudum non-destructive dissection visual method


Nearly a century ago, wing venation was introduced in gracillariid taxonomy as a means to diagnose closely related genera and species groups. Recent advances in non-destructive virtual micro-dissections suggest promising approaches with which to revisit the relevance of wing venation characters on historic primary type specimens. Many unique type specimens in Gracillariidae and other microlepidoptera groups preserved in museum collections are in poor condition, and over the course of history have suffered loss or damage to their abdomens. Consequently, genitalia morphology is not available for diagnoses and comparisons. In this paper we emphasize the need to include the type species and type specimens into the broader context of taxonomic studies on micro-moths in general and the family Gracillariidae in particular. The genus Caloptilia has a world-wide distribution and has been the subject of research for more than 200 years, yet the generic boundaries and groupings within the genus are still unresolved due to the lack of a reliable set of taxonomic characters obtained from the primary types. We describe a method of virtual descaling of the fore- and hindwings using the unset micro-moth type specimen of Caloptilia stigmatella Fabricius, 1781, in order to demonstrate that the study of historic and fragile type specimens and diagnoses of their internal morphological characters becomes possible by applying new and non-destructive technology.



  1. Albrecht, A. & Kaila, L. (1997) Variation of wing venation in Elachistidae (Lepidoptera: Gelechioidea): methodology and implications to systematics. Systematic Entomology, 22, 185–198.

    Bang-Haas, A. (1875) Fortegnelse over de I Danmark levende Lepidoptera (Sluttet). Naturhistorisk Tidsskrift, 3 (10), 1 (10), 1–56.

    Biesenbaum, W. (2016) Ein Beitrag zur Kleinschmetterlingsfauna des Niederbergischen Landen (Nordrhein-Westfalen) (Microlepidoptera). Melanargia, Nachrichten der Arbeitsgemeinschaft rheinisch-westfälischer Lepidopterologen, 28 (2), 41–49.

    BOLDSYSTEMS (2014) International Barcode of Life. Available from: (accessed 23 February 2018)

    Brower, A.E. (1984) A List of the Lepidoptera of Maine, Part 2: The Microlepidoptera, Section 2; Cosmopterigidae through Hepialidae. Technical Bulletin of the Maine Agricultural Experiment Station, 114, i–x + 1–70.

    Busck, A. (1904) Tineid moths from British Columbia, with descriptions of new species. Proceedings of the United States National Museum, 27, 1375, 745–778.

    Büttner, F.O. (1880) Die pommerschen, insbesondere die Stettiner Mikrolepidoptern [sic]. Stettiner entomologische Zeitung, 41, 383–473.

    Caradja, A. (1899) Zusammenstellung der bisher in Rumänien beobachteten Microlepidopteren. Deutsche Entomologische Zeitschrift "Iris", Dresden, 12, 171–218.

    Chambers, V.T. (1872) Micro-Lepidoptera. Canadian Entomologist, 4 (1–5), 7–12, 25–29, 41–44, 65–69, 88–92.

    Chambers, V.T. (1879) Micro-Lepidoptera. Canadian Entomologist, 11, 72–75, 89–93, 118–119, 125–127.

    Costa, O.G. (1836) Fauna del Regno di Napoli ossia enumerazione di tutti gli animali che abitano le diverse regioni di questo regno e le acque che le bagnano contenente la descrizione de nuovi o poco esattamente conosciuti. Lepidotteri. Da' Torchi del Tramater, Napoli, 24 + 8 + 4 pp., 14 pls.

    de Graaf, H.W. (1853) Nederlandsche schubvleugelige insekten [Lepidoptera]. In: Herklots, J.A., Bouwstoffen voor eene fauna van Nederland. Eerste deel. E. J. Brill, Leiden, 57 pp.

    De Prins, J. & De Prins, W. (2018) Global Taxonomic Database of Gracillariidae (Lepidoptera). Available from: (accessed 1 May 2018)

    De Prins, J., Gumovsky, A. & De Coninck, E. (2015) Discovery of a new species of Caloptilia (Lepidoptera: Gracillariidae) from east and central Africa with its suggested associated host (Gentianales: Rubiaceae) and natural enemies (Hymenoptera: Eulophidae). Zootaxa, 3957 (4), 383–407.

    De Prins, W. (1985) On a Gracillaria species from Morocco with some supplementary notes on the genus Gracillaria Haworth (Lepidoptera: Gracillariidae). Nota lepidopterologica, 8 (4), 351–362.

    De Prins, W. & De Prins, J. (2005) World Catalogue of Insects. Vol. 6. Gracillariidae. Apollo Books, Stenstrup, 502 pp.

    De Sélys-Longchamps, E. (1844) Enumération des insectes Lépidoptères de la Belgique. Mémoires de la Société royale des Sciences de Liége, 2, 1–32.

    Duponchel, P.A.J. (1838–1840) Godart, J.-B., Histoire naturelle des Lépidoptères ou Papillons de France. Tome 11. Nocturnes. Tome huitième. Méquignon-Marvis, Libraire-Éditeur, Paris, 720 pp., pls. 287–314.

    Duponchel, P.A.J. (1842–1845) In: Godart, J.-B., Histoire naturelle des Lépidoptères ou Papillons de France. Nocturnes. Supplément aux tome quatrième et suivants. Méquignon-Marvis, Libraire-Éditeur, Paris, 554 pp., pls. 51–90.

    Dyar, H.G. (1903) A list of the North American Lepidoptera and key to the literature of this order of Insects. Bulletin of the United States National Museum, 52 (1902), i–xix + 1–723 pp.

    Ely, C.R. (1918) A revision of the North-American Gracilariidae [sic] from the standpoint of venation. Proceedings of the Entomological Society of Washington, 19B (1917), 29–77.

    Ermolaev, V.P. (1977) A review of the fauna and ecology of miner-moths (Lepidoptera, Gracillariidae) of the Primorye Territory. In: The Fauna of Insects of the Far East. Trudy Zoologicheskogo Instituta, Akademija Nauk SSSR, 70, 98–116.

    Fabricius, J.C. (1781–1782) Species insectorum exhibentes eorum differentias specificas, synonyma autorum, loca natalia, metamorphosin adiectis observationibus, descriptionibus. Tom II. Impensis Carol. Ernest, Bohnii, Hamburgi et Kilonii, 494 pp. (1781) [Appendix: pp. 495–514 1782, Index: pp. 515–517 (1782)]

    Faulwetter, S., Dailianis, T., Vasileiadou, K., Kouratoras, M. & Arvanitidis, C. (2014) Can micro-CT become an essential tool for the 21st century taxonomist? An evaluation using marine polychaetes. Microscopy and Analysis, Tomography Supplement, 2014, 9–12.

    Fletcher, T.B. (1929) A list of the generic names used for Microlepidoptera. Memoirs of the Department of Agriculture in India, Entomological Series, 11 (i–ix), 1–244.

    Fletcher, T.B. (1933) Life histories of Indian Microlepidoptera. Second Series. Cosmopterygidae to Neopseustidae. Scientific Monograph. Vol. 4. Imperial Council of Agricultural Research, India, 85 pp.

    Forbes, W.T.M. (1923) The Lepidoptera of New York and neighbouring states. Part I. Primitive forms, Microlepidoptera, Pyraloids, Bombyces. Memoirs of Cornell University Agricultural Experiment Station, 68, 1–729.

    Fourcroy, A.F. (1785) Entomologia Parisiensis; sive, Catalogus insectorum quae in Agro Parisiensi reperiuntur. Part 1 & 2. sur Privilegio Academiae, Parisiis, 231 & 544 pp.

    Frey, H. (1856) Die Tineen und Pterophoren der Schweiz. Meyer und Zeller, Zürich, xii + 430 pp.

    Frey, H. & Boll, J. (1876) Einige Tineen aus Texas. Entomologische Zeitung, herausgegeben von dem entomologischen Vereine zu Stettin, 37, 209–228.

    Gershenzon, Z.S. & Kholtshenkov, V.A. (1974) In: Vasiljev, V.P. (Ed), [Pests of agricultural crops and forestry plantations. Vol. 2. Pests (continuation), Vertebrata]. Urozhai, Kiev, pp. 236–244.

    Glerean, P. & Triberti, P. (2015) The Gracillariidae of the Usvelli's collection (Lepidoptera). Gortania Atti del Museo Friulano di Storia Naturale, 36 (2014), 61–68.

    Godfrey, G.L., Cashatt, E.D. & Glenn, M.O. (1987) Microlepidoptera from the Sandy Creek and Illinois River Region: An annotated checklist of the suborders Dacnonypha, Monotrysia, and Ditrysia (in part) (Insecta). Illinois Natural History Survey Special Publication, 7, 1–44.

    Goeze, J.A.E. (1783) Entomologische Beyträge zu des Ritter Linne zwölften Ausgabe des Natursystems. Band 4. Weidmanns Erben und Reich, Leipzig, xx + 178 pp.

    Grehan, J.R., Parker, B.L., Nielsen, G.R., Miller, D.H., Hedbor, J.D., Sabourin, M. & Griggs, M.S. (1995) Moths and butterflies of Vermont (Lepidoptera). A faunal checklist. Agricultural Experiment Station, University of Vermont. Vermont Miscellaeous Publication, 116, i–xi + 1–95.

    Hancock, E.G. (2015) The shaping role of Johan Christian Fabricius (1745–1808): William Hunter’s insect collection and entomology in eighteenth century London. In: Hancock, E.G., Pearce, N. & Campbell, M. (Eds.), William Hunter’s World: the art and science of eighteenth century collecting. Routledge, Ashgate, pp. 151–163.

    Handfield, L. (1997) Liste des Lépidoptères du Québec et du Labrador. Fabreries, 7 (Supplement), 1–155.

    Haworth, A.H. (1828) Lepidoptera Britannica sistens digestionem novam insectorum lepidopterorum quae in Magne Britannia reperiuntur, larvarum pabulo, temporeque pascendi, expansione alarum; mensibusque volandi; synonymis atque locis observationibusque variis. Pars IV cum indice finali. R. Taylor, London, 98 pp. [pp. 512–609]

    Heppner, J.B. (2007) Arthropods of Florida and neighbouring land areas. Vol. 17. Lepidoptera of Florida. Part 1. Introduction and catalogue. Florida Department of Agriculture & Consumer Services, Gainesville, 670 pp.

    Heydenreich, G.H. (1851) Lepidopterorum Europaeorum Catalogus Methodicus. Systematisches Verzeichniss der Europaeischen Schmetterlinge. Dritte Ausgabe. Julius Klinkhardt, Leipzig, 130 pp.

    Hübner, J. (1816–1826) Verzeichniss bekannter Schmettlinge [sic]. J. Hübner Verlag, Augsburg, 431 pp., 72 pls.

    Hübner, J. (1822) Systematisches-alphabetisches Verzeichniss aller bisher bey den Fürbildungen zur Sammlung europäischer Schmetterlinge angegebenen Gattungsbennungen; mit Vorbemerkung auch augsburgischer Gattungen. J. Hübner Verlag, Augsburg, vi + 81 pp.

    Hübner, J. (1796–1838) Sammlung europäischer Schmetterlinge. Achte Horde. Tineae Die Schaben; nach der Natur geordnet, beschrieben und vorgestellt. J. Hübner Verlag, Augsburg, 78 pp., 71 pls.

    ICZN (1999) International Code of Zoological Nomenclature. Fourth edition. The International Trust for Zoological Nomenclature, London, 306 pp. Avaiable from: (Accessed 25 Jun. 2018)

    Ivinskis, P. & Pakalniškis, S. (1984) Microlepidoptera of Lithuania (Gracillariidae). Lietuvos TSR Mokslu Akademijos darbai, 1 (85), 26–36.

    Kawahara, A.Y., Ohshima, I., Kawakita, A., Regier, J.C., Mitter, C., Cummings, M.P., Davis, D.R., Wagner, D.L., De Prins, J. & Lopez-Vaamonde, C. (2011) Increased gene sampling strengthens support for higher-level groups within leaf-mining moths and relatives (Lepidoptera: Gracillariidae). BMC Evolutionary Biology, 11, 182–196.

    Kawahara, A.Y., Plotkin, D., Ohshima, I., Lopez-Vaamonde, C., Houlihan, P.R., Breinholt, J.W., Kawakita, A., Xiao, L., Regier, J.C., Davis, D.R., Kumata, T., Sohn, J.-C., De Prins, J. & Mitter, C. (2017) A molecular phylogeny and revised higher-level classification for the leaf-mining moth family Gracillariidae and its implications for larval host-use evolution. Systematic Entomology, 42 (1), 60–81.

    Simonsen, T.J. & Kitching, I.J. (2014) Virtual dissections through micro-CT scanning: a method for non-destructive genitalia ‘dissections’ of valuable Lepidoptera material. Systematic Entomology, 39, 606–618.

    Koçak, A.Ö. & Seven, S. (2001) Tentative Checklist of the Turkish Lepidoptera (except Papilionoidea and Hesperioidea). Available from: (accessed 25 February 2018)

    Kumata, T. (1982) A taxonomic revision of the Gracillaria group occurring in Japan (Lepidoptera: Gracillariidae). Insecta Matsumurana, New Series, 26, 1–186.

    Kuznetzov, V.I. (1981) Fam. Gracillariidae (Lithocolletiidae)—leaf blotch miners. In: Medvedev, G.S. (Ed.), A guide to the insects of the European part of the USSR. Lepidoptera. Vol. 4. Lepidoptera. Part 2. Nauka, Leningrad, pp. 149–311.

    Leubner, F. Wipfler, B. & Bradler, S. (2017) Comparative morphology of the orthopteran thorax with a discussion of phylogenetically relevant characters. Insect Systematics and Diversity, 1(1), 29–47.

    Lienig, F. (1846) Lepidopterische Fauna von Lievland und Curland. Isis, oder enzyklopädische Zeitung von Oken, 3–4 (1846), 175–302.

    Linnaeus, C. (1758) Systema Naturae per Regna Tria Naturae, secundum Classes, Ordines, Genera, Species, cum characteribus, differentiis, synonymis, Iocis. Tomus 1. Editio Decima, Reformata. Laurentii Salvii, Holmiae, iv + 824 pp.

    Martynova, E.F. (1952) Osobennosti fauny cheshuekrylych Juzhnogo Priuralja i ee zhachenie dlia stepnogo lesorazvedenija. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR, 11, 66–91.

    Massa, B., Rizzo, M.C. & Caleca, V. (2001) Natural alternative hosts of Eulophidae (Hymenoptera: Chalcidoidea) parasitoids of the citrus leafminer Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae) in the Mediterranean Basin. Journal of Hymenoptera Research, 10 (1), 91–100.

    Mendes D'Azevedo, C. (1913) Lepidopteros de S. Fiel. Supplemento. Broteria, Revista de Sciencias naturaes, Serie Zoologica, 11, 15–44.

    Meyrick, E. (1927) A revised handbook of British Lepidoptera. E. W. Classey ltd., London, 914 pp.

    Mutanen, M., Wahlberg, N. & Kaila, L. (2010) Comprehensive gene and taxon coverage elucidates radiation patterns in moths and butterflies. Proceedings of the Royal Society, series B Biological Sciences, 277 (1695), 2839–2848.

    Nielsen, M.C. (1998) Preliminary list of Michigan moths: the Microlepidoptera. Newsletter of the Michigan Entomological Society, 43 (4), 1, 4–14.

    Noreika, R. (1997) Gracillariidae. In: Ler, P.A. (Ed.), Key to the insects of Russian Far East. Vol. V. Trichoptera and Lepidoptera. Part 1. Dal'nauka, Vladivostok, pp. 373–429.

    Panzer, G.W.F. (1793–1844) Fauna Insectorum Germanicae initia oder Deutschlands Insecten. Heft 1–12 (1793), 13–24 (1794), 25–36 (1796), 37–48 (1797), 49–60 (1798), 61–72 (1799), 73–84 (1801), 85–96 (1805), 97–108 (1809), 109 (1813). In den Felseckerschen Buchhandlung, Nürnberg. [no pagination]

    Park, K.T. (1983) Microlepidoptera of Korea. Insecta Koreana, Series 3, 1–189, pls. 1–2.

    Pohl, G.R., Anweiler, G.G., Schmidt, B.C., Kondla, N.G. (2010) An annotated list of the Lepidoptera of Alberta, Canada. ZooKeys, 38, 1–549.

    Pohl, G.R., Landry, J.-F., Schmidt, B.C., Lafontaine, J.D., Troubridge, J.T., Macaulay, A.D., van Nieukerken, E.J., DeWaard, J.R., Dombroskie, J.J., Klymko, J., Nazari, V. & Stead, K. (2018) Annotated checklist of the moths and butterflies (Lepidoptera) of Canada and Alaska. Pensoft Publishers, 580 pp.

    Rebel, H. (1904) Studien über die Lepidopterenfauna der Balkanländer. II. Teil. Bosnien und Herzegowina. Annalen des kaiserlich-königlichen Naturhistorischen Hofmuseums, Wien, 19 (2–3), 97–377, pls. iv–v.

    Regier, J.C., Mitter, C., Zwick, A., Bazinet, A.L., Cummings, A.L., Kawahara, A.Y., Sohn, J.-C., Zwickl, D.J., Cho, S., Davis, D.R., Baixeras, J., Brown, J., Parr, C., Weller, S., Lees, D.C. & Mitter, K.T. (2013) A large-scale, higher-level, molecular phylogenetic study of the insect order Lepidoptera (moths and butterflies). PLoS One, 8 (3), e 58568.

    Rungs, C.E.E. (1979) Catalogue raisonné des Lepidoptères du Maroc. Inventaire faunistique et obsevations écologiques. Tome 1. Travaux de l'Institut Scientifique, Série Zoologie, 39, 1–222.

    Saito, K., Nomura, S., Yamamoto, S., Niiyama, R. & Okabe, Y. (2017) Investigation of hindwing folding in ladybird beetles by artificial elytron transplantation and microcomputed tomography. Proceedings of the National Academy of Sciences of the United States of America, 114 (22), 5624–5628.

    Sherniyazova, R.M. (1975) K faune i ekologi nizskikh cheshuekrylykh ivivykh yuzhnogo sklona Gissarskogo khrebta. Entomologiya Tadzhikistana, 1, 187–190.

    Simonsen, T.J. & Kitching, I.J. (2014) Virtual dissections through micro-CT scanning: a method for non-destructive genitalia ‘dissections’ of valuable Lepidoptera material. Systematic Entomology, 39, 606–618.

    Skala, H. (1937) Minen aus Mittel- und Südeuropa (Fortsetzung und Schluss). Zeitschrift des Österreichischen Entomologen-Vereines, 22, 10–11 + 19–20.

    Skala, H. (1938) Einiges über Falter-Minen aus dem Mediterrangebiet. Zeitschrift des Österreichischen Entomologen-Vereines, 23, 8–10 + 30–31 + 43–46.

    Stainton, H.T. (1849) An attempt at a systematic catalogue of the British Tineidae & Pterophoridae. John van Voorst, London, iv + 32 pp.

    Stainton, H.T. (1854) Lepidoptera: Tineina. In: Insecta Britannica. Vol. 3. Lovell Reeve, London, 313 pp., 10 pls.

    Stainton, H.T. (1864) The Natural History of the Tineina. Vol. VIII. containing Gracilaria, Part I and Ornix. Part I. John van Voorst, London, vii + 315 pp., 8 pls.

    Strand, E. (1901) Beitrag zur Schmetterlingsfauna Norwegens. Nyt Magazin for Naturvidenskaberne, 39, 25–72.

    Szöcs, J. (1981) Angaben über die minierenden Motten aus Budapest und Umgebung. Folia Entomologica Hungarica / Rovartani Közlemények, 42 (2), 209–220.

    Tengström, J.M.J. (1848) Bidrag till Finlands Fjäril-Fauna. Notiser ur Sällskapets pro Fauna et Flora Fennica Förhandlingar, 1, 69–164.

    van Orden Covell, C. (1999) The butterflies and moths (Lepidoptera) of Kentucky: an annotated checklist. Kentucky State Nature Preserves Commission Scientific and Technical Series, 6, i–liv + 1–220.

    Vári, L. (1961) South African Lepidoptera. Vol. I. Lithocolletidae. Transvaal Museum Memoir, 12, 1–238, 112 pls.

    Wagner-Rollinger, C. (1974) Les lépidoptères du Grand-Duché de Luxembourg (et des régions limitrophes). Archives de l'Institut Grand-Ducal, Section des Sciences, 36, 285–358.

    Wallengren, H.D.J. (1881) Genera nova Tinearum. Entomologisk Tidskrift, 1 (2), 94–97.

    Zeller, P.C. (1847) Die Gracilarien. Linnaea Entomologica, 2, 303–383, 585–586, pl. II.

    Zimmerman, E.C. (1978) Insects of Hawaii. Vol. 8. Pyraloidea. University of Hawaii Press, Honolulu, 881pp.