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Type: Monograph
Published: 2008-06-16
Page range: 1–106
Abstract views: 85
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Identification guide to species in the scale insect tribe Iceryini (Coccoidea: Monophlebidae)

Department of Entomology, University of California, One Shields Ave., Davis, CA 95616
Department of Entomology, University of California, One Shields Ave., Davis, CA 95616
Hemiptera Crypticerya Echinicerya Gigantococcus Gueriniella Icerya

Abstract

We recently revised the scale insect tribe Iceryini (Coccoidea: Monophlebidae) based on a molecular phylogenetic hypothesis and supporting morphological evidence. As a result of this revision, we proposed new generic concepts that required significant reorganization of species in three of the five iceryine genera: Crypticerya Cockerell, Gigantococcus Pesson & Bielenin and Icerya Signoret. The other two genera, Echinicerya Morrison and Gueriniella Fernald, remain unchanged. Here we discuss taxonomic characters important for iceryine identification and provide a comprehensive review of all described iceryine species. We include keys to the genera and to the species of each genus of Iceryini based on features of the adult females. We present new diagnoses for all the species for which type material was available for examination and summarize original descriptions for species for which we were unable to examine material. We designate lectotypes for the following species names (valid name given in parentheses): Llaveia abrahami Newstead (C. abrahami), Icerya brasiliensis Hempel (C. brasiliensis), Icerya colimensis Cockerell (C. colimensis), Icerya genistae Hempel (C. genistae), Icerya (Crypticerya) littoralis Cockerell (C. littoralis), Icerya littoralis mimosae Cockerell (C. littoralis),Icerya littoralis tonilensis Cockerell (C. littoralis), Crypticerya rosae mexicana Cockerell & Parrott (C. mexicana), Icerya minima Morrison (C. minima), Icerya montserratensis Riley & Howard (C. montserratensis), Palaeococcus morrilli Cockerell (C. morrilli), Icerya palmeri Riley & Howard (C. palmeri), Icerya rileyi Cockerell (C. rileyi), Icerya rileyi larreae Cockerell (C. rileyi), Steatococcus tabernicolus Ferris (C. tabernicola), Icerya (Crypticerya) townsendi Cockerell (C. townsendi), Icerya (Crypticerya) townsendi plucheae Cockerell (C. townsendi), Icerya zeteki Cockerell (C. zeteki), Icerya seychellarum albolutea Cockerell (Gi. alboluteus), Crypticerya bicolor DeLotto (Gi. bicolor), Icerya brachystegiae Newstead (Gi. brachystegiae), Palaeococcus cajani Newstead (Gi. cajani), Palaeococcus caudata Newstead (Gi. caudatus), Icerya euphorbiae Brain (Gi. euphorbiae), Aspidoproctus gowdeyi Newstead (Gi. gowdeyi), Icerya longisetosa Newstead (Gi. longisetosus), Icerya nigroareolata Newstead (Gi. nigroareolatus), Icerya sulfurea pattersoni Newstead (Gi. pattersoni), Palaeococcus theobromae Newstead (Gi. theobromae), Crossotosoma aegyptiacum Douglas (I. aegyptiaca), Icerya rosae australis Maskell (I. australis), Icerya crocea Green (I. crocea), Icerya formicarum Newstead (I. formicarum), Icerya jacobsoni Green (I. jacobsoni), Icerya koebelei Maskell (I. koebelei), Icerya minor Green (I. minor), Icerya (Crypticerya) nuda Green (I. nuda), Icerya nudata Maskell (I. nudata), Icerya hyperici Froggatt (I. nudata), Palaeococcus dymocki Froggatt (I. nudata), Icerya pilosa Green (I. pilosa), Icerya purchasi citriperda Hempel (I. purchasi), Icerya purchasi crawii Cockerell (I. purchasi), Icerya purchasi maskelli Cockerell (I. purchasi), and Icerya schrottkyi Hempel (I. schrottkyi).

References

  1. Aisagbonhi, C. I., Nwana, I. E. & Agwu, S. I. (1985) Preliminary analysis of a field population of Aspidotus destructor Signoret (Homoptera: Diaspididae) and some soft scales on coconut palms. Nigerian Journal of Entomology, 6, 24–32.

    Ben-Dov, Y. (2005) A Systematic Catalogue of the Scale Insect Family Margarodidae (Hemiptera: Coccoidea) of the World. Intercept Ltd., Wimborne, U.K., 400 pp.

    Berlese, A. (1909) Gli insetti: loro organizzazione, sviluppo, abitudini e rapporti coll'uomo. Societá Editrice Libraria, Milan, 1009 pp.

    Bhatti, S. (1990) A new monophlebine genus (Homoptera: Coccoidea: Margarodidae: Monophlebinae) on Melaleuca L. in Australia. Invertebrate Taxonomy, 3, 495–517.

    Bhatti, S. & Gullan, P. J. (1990) New margarodid species (Homoptera: Coccoidea: Margarodidae: Monophlebinae) from New Guinea. Invertebrate Taxonomy, 3, 877–911.

    Borchsenius, N. S. (1949) A new genus and new species of hard and soft scales (Homoptera, Coccoidea) of USSR fauna. Entomologicheskoe Obozrenye, 30, 334–353.

    Boyer de Fonscolombe, E. L. J. H. (1834) Description des Kermes qu'on trouve aux environs d'Aix. Annales de la Société Entomologique de France, 3, 201–218.

    Brain, C. K. (1915) The Coccidae of South Africa. Transactions of the Royal Society of South Africa, 5, 65–194.

    Castel-Branco, A. J. F. (1952) Descrição de trés espécies novas de cochonilhas da Africa Oriental Portuguesa. Crypticeria rodriguesi n. sp., Cerococcus fradei n. sp., Pulvinaria vayssierei n. sp. Anais Estudos de Zoologia. Ministerio do Ultramar, Junta de Investigacoes do Ultramar. Lisboa, 7, 22–39.

    Cockerell, T. D. A. (1895) VIII. Some new insects. Pysche, 7, 15–16.

    Cockerell, T. D. A. (1896a) A check list of the Coccidae. Bulletin of the Illinois State Laboratory of Natural History, 4, 318–339.

    Cockerell, T. D. A. (1896b) New species of insects taken on a trip from the Mesilla Valley to the Sacramento Mts., New Mexico. Journal of the New York Entomological Society, 4, 201–207.

    Cockerell, T. D. A. (1897) Two forms of fluted scale. Psyche, 8, 94.

    Cockerell, T. D. A. (1898a) New Coccidae from Mexico. Annals and Magazine of Natural History (ser. 7), 1, 426–440.

    Cockerell, T. D. A. (1898b) Two new Coccidae from Lagos, W. Africa. The Entomologist, 31, 259.

    Cockerell, T. D. A. (1899a) Article VII. - First supplement to the check-list of the Coccidae. Bulletin of the Illinois State Laboratory of Natural History, 5, 389–398.

    Cockerell, T. D. A. (1899b) Insecta. Rhynchota, Hemiptera-Homoptera. Volume II, Part 2, Biologia Centrali-Americana. Published for the Editors by R.H. Porter, London, 37 pp.

    Cockerell, T. D. A. (1899c) Tables for the determination of the genera of Coccidae. The Canadian Entomologist, 31, 273–279, 330–333.

    Cockerell, T. D. A. (1902a) A contribution to the classification of the Coccidae. Entomologist, 35, 232–233; 257–260.

    Cockerell, T. D. A. (1902b) Tables for the determination of Mexican Coccidae. I. Subfamily Monophlebinae. Memorias y Revista de la Sociedad Científica 'Antonio Alzate', 18, 79–82.

    Cockerell, T. D. A. (1905) Some Coccidae from the Philippine Islands. Proceedings of the Davenport Academy of Sciences. Davenport, Iowa, 10, 127–136.

    Cockerell, T. D. A. (1914a) A new Coccid from Arizona (Hemip.). Entomological News, 25, 110.

    Cockerell, T. D. A. (1914b) A new cotton scale from Panama. Journal of Economic Entomology, 7, 148.

    Cockerell, T. D. A. & Parrott, P. J. (1899) Contributions to the knowledge of the Coccidae. The Industrialist, 25, 276–284.

    De Lotto, G. (1959) Two new Margarodidae (Homopt: Coccoidea) from Kenya. Journal of the Entomological Society of Southern Africa, 22, 385–389.

    Deitz, L. L. & Tocker, M. F. (1980) W.M. Maskell's Homoptera: Species-group names and type-material. New Zealand Department of Scientific and Industrial Research, Information Series, 146, 1–76.

    Douglas, J. W. (1888) Notes on some British and exotic Coccidae (No. 10). Entomologist's Monthly Magazine, 25, 86–89.

    Douglas, J. W. (1889) Notes on some British and exotic Coccidae (No. 13). Entomologist's Monthly Magazine, 25, 232–235.

    Douglas, J. W. (1890) Notes on some British and exotic Coccidae (No. 15). Entomologist's Monthly Magazine, 26, 79–81.

    Fabricius, J. C. (1775) Systema Entomologiae, sistens Insectorum Classes, Ordines, Genera, Species, Adiectis Synonymis, Locis, Descriptionibus, Observationibus. VII. Ryngota. Flensburgi et Lipsiae, Korte, 832 pp.

    Fernald (1903) A catalogue of the Coccidae of the world. Bulletin of the Hatch Experiment Station of the Massachusetts Agricultural College, 88, 1–360.

    Ferris, G. F. (1921) Report upon a collection of Coccidae from Lower California. Stanford University Publications, Biological Sciences. Palo Alto, 1, 61–132.

    Foldi, I. & Ben-Dov, Y. (2001) A nomenclatural note on "Steatococcus anonae Newstead" (Hem., Coccoidea, Margarodidae). Bulletin de la Societe Entomologique de France, 106, 461.

    Franks, D. J. (1965) Data Contributory to Analysis of Mitotic Mechanisms in Steatococcus tuberculatus Morrison (Coccoidea-Homoptera). Thesis. Department of Zoology, Duke University, Durham, North Carolina.

    Froggatt, W. W. (1906) Mealy Bugs. No. I - Monophlebinae. Agricultural Gazette of New South Wales, 17, 770–779.

    Froggatt, W. W. (1919) Insects and St. John's Wort. Agricultural Gazette of New South Wales, 30, 470–472.

    Froggatt, W. W. (1921) A descriptive catalogue of the scale insects ('Coccidae') of Australia. Part III. Department of Agriculture, New South Wales Science Bulletin, 19, 1–43.

    Froggatt, W. W. (1923) Forest Insects of Australia. Forestry Commisioners: Sydney, N.S.W. 171 pp.

    Green, E. E. (1896) Catalogue of Coccidae collected in Ceylon. Indian Museum Notes, 4, 2–10.

    Green, E. E. (1908) Remarks on Indian scale insects (Coccidae), with descriptions of new species. Indian Museum Notes, 5, 93–103.

    Green, E. E. (1913) Remarks on Coccidae collected by Mr. Eward Jacobson, of Samarang, Java, with descriptions of two new species. Tijdschrift voor Entomologie, 55, 311–318.

    Green, E. E. (1922) The Coccidae of Ceylon, Part V. Dulau & Co., London, 457 pp.

    Green, E. E. (1930) Fauna sumatrensis (Bijdrag nr. 65) Coccidae. Tijdschrift voor Entomologie, 73, 279–297.

    Green, E. E. (1932) A note on Icerya aegyptiaca Dougl. (Coccidae) and some nearly related species. Stylops: A Journal of Taxonomic Entomology, 1, 31–33.

    Green, E. E. (1937) An annotated list of the Coccidae of Ceylon, with emendations and additions to date. Ceylon Journal of Science Section B. Zoology and Geology, 20, 277–341.

    Guérin-Méneville, F. E. (1852) Note sur une cochenille indigène qui vit sur la fève de Marais, et qui semble propre à donner une matière colorante abondante et susceptible d'etre employée dans l'industrie. Compte Rendu de l'Academie des Sciences. Paris, 34, 334–335.

    Guérin-Méneville, F. E. (1867) [Comment on] M. ed. Morin adresse une lettre sur les ravages produits a l'Ile de la Réunion par des insectes qui attaquent la canne à sucre. Revue et Magasin de Zoologie, 19, 450–451.

    Gullan, P. J. (1984) A revision of the gall-forming coccoid genus Apiomorpha Rübsaamen (Homoptera : Eriococcidae : Apiomorphinae). Australian Journal Zoology Supplementary Series, 97, 1–203.

    Hall, W. J. (1940) Observations on the Coccidae of Southern Rhodesia. - IX. Transactions of the Royal Entomological Society of London, 90, 487–496.

    Hempel, A. (1900) As coccidas Brazileiras. Revista do Museu Paulista. São Paulo, 4, 365–537.

    Hempel, A. (1912) Catalogos da Fauna Brasileira Editados pello Museu Paulista S. Paulo - Brasil. Diario Official, São Paulo, 77 pp.

    Hempel, A. (1918) Descripção de sete novas espécies de coccidas. Revista do Museu Paulista. São Paulo, 10, 193–208.

    Hempel, A. (1920) Descripçoes de coccidas novas e pouco conhecidas. Revista do Museu Paulista. São Paulo, 12, 329–377.

    Hempel, A. (1923) Hemiptera (Homoptera) familia Coccidae. Sub-familia Monophlebinae. Revista do Museu Paulista. São Paulo, 13, 510–512.

    Hempel, A. (1932) Descripção de vinte a duas espécies novas de coccideos (Hemiptera - Homoptera). Revista de Entomologia, 2, 310–339.

    Hodges, G. (2006) Icerya genistae (Hemiptera: Margarodidae) A new exotic scale insect for Florida. Available from: http://www.doacs.state.fl.us/pi/enpp/ento/i.genistae.html (acessed 8 March 2007).

    Hughes-Schrader, S. (1925) Cytology of hermaphroditism in Icerya purchasi (Coccidae). Zeitschrift für Zellforschung und Mikroskopische Anatomie, 2, 264–292.

    Hughes-Schrader, S. (1946) A new type of spermiogenesis in iceryine coccids, with linear alignment of chromosome in the sperm. Journal of Morphology, 78, 43–84.

    Hughes-Schrader, S. (1963) Hermaphroditism in an African coccid, with notes on other margarodids (Coccoidea—Homoptera). Journal of Morphology, 113, 173–184.

    Hughes-Schrader, S. & Monahan, D. F. (1966) Hermaphroditism in Icerya zeteki Cockerell, and the mechanism of gonial reduction in iceryine coccids (Coccoidea: Margarodidae Morrison). Chromosoma, 20, 15–31.

    Hughes-Schrader, S. & Ris, H. (1941) The diffuse spindle attachment of coccids, verified by the mitotic behavior of induced chromosome fragments. Journal of Experimental Zoology, 87, 429–456.

    Hughes-Schrader, S. & Tremblay, E. (1966) Gueriniella and the cytotaxonomy of iceryine coccids (Coccoidea: Margarodidae). Chromosoma, 19, 1–13.

    International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature (4th ed.). International Trust for Zoological Nomenclature, London, 306 pp.

    Jashenko, R. V. & Danzig, E. M. (1992) A new genus and three new species of the subfamily Monophlebinae Maskell (Homoptera, Margarodidae) from Middle Asia, Afghanistan and Vietnam. Entomologicheskoe Obozrenye, 71, 84–90.

    Kieffer, J. J. (1909) Description de galles et d'insectes gallicoles d'Asie. Marcellia (Rivista Internazionale di Cecidologia), 7, 149–167.

    Koteja, J. (1974) Comparative studies on the labium in the Coccinea (Homoptera): Zeszyty Naukowe Akademii Rolniczej w Warszawie, Rozprawy Naukowe, 89, 1 –162.

    Kuwana, S. I. (1907) Coccidae of Japan, I. A synoptical list of Coccidae of Japan with descriptions of thirteen new species. Bulletin of the Imperial Central Agricultural Experiment Station, Japan, 1, 177–212.

    Leonardi, G. (1907) Notizie sopra alcune cocciniglie dell'isola di giava raccolte dal Prof. O. Penzig. Annali della R. Scuola Superiore di Agricoltura. Portici, 7, 1–22.

    Leonardi, G. (1911) Contributo alla conoscenza delle coccinglie della Repúbblica Argentina. Bollettino del R. Laboratorio di Entomologia Agraria di Portici, 5, 237–284.

    Lindinger, L. (1913) Afrikanische Schildläuse. V. Die Schildläuse Deutsche-ostafrikas. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 30, 59–95.

    Lindinger, L. (1932) Beiträge zur Kenntnis der Schildläuse. Konowia, 11, 177–205.

    MacGillivray, A. D. (1921) The Coccidae. Tables for the Identification of the Subfamilies and Some of the More Important Genera and Species, together with Discussions of their Anatomy and Life History. Scarab,Urbana, Illinois, 502 pp.

    Mallis, A. (1971) American Entomologists. Rutgers University Press, New Brunswick, New Jersey.

    Mamet, R. (1951) Notes on the Coccoidea of Madagascar. - II. Mémoires de l'Institut Scientifique de Madagascar (Ser. A), 5, 213–254.

    Maskell, W. M. (1879) On some Coccidae in New Zealand. Transactions and Proceedings of the New Zealand Institute, 11, 187–228.

    Maskell, W.M. (1892) A new Icerya, and some other new coccids from Australia. Entomologist's Monthly Magazine, 28, 183–184.

    Maskell, W. M. (1893) Further coccid notes: with descriptions of new species from Australia, India, Sandwich Islands, Demerara, and South Pacific. Transactions and Proceedings of the New Zealand Institute, 24, 1–64.

    Maskell, W. M. (1894) Further coccid notes with descriptions of several new species, and discussion of various points of interest. Transactions and Proceedings of the New Zealand Institute, 26, 65–105.

    Maskell, W. M. (1896) Further coccid notes: with descriptions of new species, and discussion of questions of interest. Transactions and Proceedings of the New Zealand Institute, 28, 380–411.

    Morales, C. F. (1991) Margarodidae (Insecta: Hemiptera). Fauna of New Zealand / Ko te Aitanga Pepeke o Aotearoa. In: C. T. Duval (Ed). DSIR Plant Protection, Aukland, New Zealand, 123 pp.

    Morrison, H. (1919) A report on a collection of Coccidae from Argentina with descriptions of apparently new species (Hom.). Proceedings of the Entomological Society of Washington, 21, 63–91.

    Morrison, H. (1921) Some nondiaspine Coccidae from the Malay peninsula, with descriptions of apparently new species. The Philippine Journal of Science, 18, 637–677.

    Morrison, H. (1927) Descriptions of the genera and species belonging to the coccid family Margarodidae. Proceedings of the Biological Society of Washington, 40, 99–109.

    Morrison, H. (1928) A classification of the higher groups and genera of the coccid family Margarodidae. United States Department of Agriculture Technical Bulletin, 52, 1–239.

    Morrison, H. (1930) An interesting new genus of iceryine coccid. Proceedings of the Biological Society of Washington, 43, 17–20.

    Morrison, H. (1941) A new Steatococcus from Mexico (Hemiptera: Coccoidea). Proceedings of the Entomological Society of Washington, 43, 140–141.

    Morrison, H. & Morrison, E. R. (1923) The scale insects of the subfamilies Monophlebinae and Margarodinae treated by Maskell. Proceedings of the United States National Museum, 62, 1–47.

    Morrison, H. & Morrison, E.R. (1966) An annotated list of generic names of the scale insects (Homoptera: Coccoidea). Miscellaneous Publication United States Department of Agriculture, 1015, 1–206.

    Moses, M. J. (1966a) Chromosome reorganization and migration during spermateleosis in a coccid (Steatococcus tuberculatus). Journal of Cell Biology, 31, A78.

    Moses, M. J. (1966b) Microtubules in aflagellate sperm of a cocid (Steatococcus tuberculatus). Anatomical Record, 154, 391.

    Moses, M. J. & Wilson, M. H. (1970) Spermiogenesis in an iceryine coccid, Steatococcus tuberculatus Morrison: an electron microscopic study. Chromosoma, 30, 373–429.

    Newstead, R. (1894) Scale insects in Madras. Indian Museum Notes, 3, 21–32.

    Newstead, R. (1896) Observations on Coccidae (No. 15). Entomologist's Monthly Magazine, 32, 132–134.

    Newstead, R. (1897) Observations on Coccidae (No. 16). Entomologist's Monthly Magazine, 33, 165–171.

    Newstead, R. (1908a) On the gum-lac insect of Madagascar, and other coccids affecting the citrus and tobacco in that island. Quarterly Journal. Institute of Commercial Research in the Tropics. University of Liverpool, 3, 3–13.

    Newstead, R. (1908b) On the structural characters of three species of Coccidae affecting cocoa, rubber and other plants in Western Africa. Journal of Economic Biology, 2, 149–157.

    Newstead, R. (1911) On a collection of Coccidae and Aleurodidae, chiefly African, in the collection of the Berlin Zoological Museum. Mitteilungen aus dem Zoologischen Museum in Berlin, 5, 155–174.

    Newstead, R. 1913. Notes on scale-insects (Coccidae). Part I. Bulletin of Entomological Research, 4: 67--–-81.

    Newstead, R. (1914) Notes on scale-insects (Coccidae). Part II. Bulletin of Entomological Research, 4, 301–311.

    Newstead, R. (1917) Observations on scale-insects (Coccidae). - IV. Bulletin of Entomological Research, 8, 1–34.

    Newstead, R. (1920) Observations on scale-insects (Coccidae). - VI. Bulletin of Entomological Research, 10, 175–207.

    O'Brien, R. (1956) Deoxyribose nucleic acid in a haploid-diploid species (Steatococcus tuberculatus Morrison). Chromosoma, 8, 229–259.

    Pesson, P. & Bielenin, I. (1966) Gigantococcus nouveau genre de cochenille Margarodidae pour Icerya maxima Newstead 1914 = Icerya corticalis Vayssière 1926, redescription de l'espèce Africaine Gigantococcus maximus (Newstead). Annales de la Société Entomologique de France (N.S.), 2, 219–251.

    Rao, V. P. (1951a) Iceryine scale insects recorded from the Orient. Indian Journal of Entomology, 12, 39–66.

    Rao, V. P. (1951b) Iceryine scale insects recorded from the Orient II. Indian Journal of Entomology, 12, 127–158.

    Riley, C. V. & Howard, L. O. (1890a) An Icerya in Florida. Insect Life, 2, 333.

    Riley, C. V. & Howard, L. O. (1890b) Some new iceryas. Insect Life, 3, 92–106.

    Signoret, V. (1876) Essai sur les cochenilles ou gallinsectes (Homoptères - Coccides), 16e partie. Annales de la Société Entomologique de France, (serie 5), 5, 346–373.

    Silvestri, F. (1939) Compendio di Entomologia Applicata. Parte speciale. Tipografia Bellavista, Portici, 974 pp.

    Tang, F.-T. & Hao, J.-J. (1995) The Margarodidae and others of China. Chinese Agricultural Science Technology Press, Beijing, 738 pp.

    Targioni Tozzetti, A. (1868) Introduzione alla seconda memoria per gli studi sulle cocciniglie, e catalogo dei generi e delle specie della famiglia dei coccidi. Atti della Società italiana di scienze naturali, 11, 694–738.

    Townsend, C. H. T. (1892) Scale-insects in New Mexico. Bulletin of New Mexico Agricultural Experiment Station, 7, 1–23.

    UCEC (2003) CU Museum: Entomology Section - Species Type List. Available from: http://cumuseum.colorado.edu/Research/Entomology/Databases/Types/ (accessed 1 April 2007).

    Unruh, C. M. & Gullan, P. J. (2008) Molecular data reveal convergent reproductive strategies in iceryine scale insects (Hemiptera: Coccoidea: Monophlebidae), allowing re-interpretation of morphology and a revised generic classification. Systematic Entomology, 33, 8–50.

    Vayssière, P. (1926) Contribution à l'étude biologique et systématique des Coccidae. Annales des Epiphyties, 12, 187–382.

    Westwood, J. O. (1855) The Seychelles Dorthesia. Gardeners' Chronicle and Agricultural Gazette, 51, 836.

    Williams, D. J., Gullan, P. J., Englberger, K. & Moore, A. (2006) Report on the scale insect Icerya imperatae Rao (Hemiptera: Coccoidea: Margarodidae) seriously infesting grasses in the Republic of Palau. Micronesia, 38, 267–272.

    Williams, D. J. & Watson, G. W. (1990) The Scale Insects of the Tropical South Pacific Region. Part 3: The Soft Scales (Coccidae) and Other Families. C.A.B. International Institute of Entomology, Wallingford, Oxon, 267 pp.

    Zimsen, E. (1964) The Type Material of I.C. Fabricius. Munksgaard, Copenhagen, 656 pp.