Leafhoppers

(Superfamily Membracoidea: Cicadellidae and Myerslopiidae)

Images Selected Links Selected References

Leafhoppers are common and abundant insects worldwide. They are currently placed among two families: Myerslopiidae, with only two genera, and the enormous family Cicadellidae, which, with more than 22,000 described species and 2600 genera, ranks among the 10 largest insect families. Phylogenetic analyses suggest that leafhoppers may have given rise to treehoppers, and it seems likely that additional families of leafhoppers will be recognized once the evolutionary relationships are better understood. Indeed, a number of previously recognized leafhopper "families" (including Eurymelidae, Hylicidae, Ledridae, and Ulopidae) are currently placed within Cicadellidae. As noted below, numerous leafhopper species are considered serious plant pests, and those that vector plant diseases inflict untold economic loses in agricultural crops.

Distribution

Leafhoppers occur in virtually all places inhabited by vascular plants. While the primitive family Myerslopiidae is known only from Chile and New Zealand, the much larger family Cicadellidae is widespread and abundant in all major zoogeographic regions. Within the Cicadellidae, some subfamilies and many tribes seem to be restricted to a single continent.

Morphology

Members of the Cicadellidae range in length from 2 to 32 mm and characteristically have four rows of enlarged, spine-like setae on the hind tibia. In this regard, they resemble treehoppers, which have up to three such rows of setae on the hind tibia, but most treehoppers have a posterior pronotal process that is absent in leafhoppers. Among all insects, only cicadellids produce minute granules called brochosomes. These intricately shaped structures form in a specialized region of the Malpighian tubules. The brochosomes are secreted and applied (using the legs) to the body as a hydrophobic coating. This curious anointing behavior is usually observed after each molt. Members of the small leafhopper family Myerslopiidae are peculiar flightless insects that have leathery forewings and vestigial ocelli.

Ecology and Behavior

A prominent group of insect herbivores, leafhoppers feed on an astonishing variety of vascular plants including conifers, grasses, sedges, and many families of broad-leafed woody and herbaceous plants. This feeding may mechanically injure plant tissues or may promote the spread of plant pathogens, including bacteria, viruses, and mycoplasma-like organisms. With more than 150 species reported as plant disease vectors, the family Cicadellidae ranks very high among insect groups that transmit pathogens among agricultural crops. Eggs are deposited in the host plant tissue, which may also injure the host. Some female "sharpshooters" (subfamily Cicadellinae) cover their eggs with specialized brochosomes. Leafhoppers seem to have 4 to 6 (usually 5) nymphal instars and 1 to 5 generations per year. Most leafhoppers feed on plant phloem. Cicadellinae, however, feed on xylem, and Typhlocybinae feed on parenchyma. Migratory behavior is documented for some leafhoppers. Because of their widespread abundance, leafhoppers are a significant source of food for predatory insects, spiders, birds, lizards, and other insectivores. Ant-attendance has been reported in several leafhopper lineages, including eurymelines, which usually occur in groups, and in some idiocerines that have been observed guarding their eggs.

Leafhoppers prepared by L. L. Deitz, C. H. Dietrich, S. H. McKamey, R. A. Rakitov, P. S. Southern, G. R. Balme, and P. A. Alvarez. 1 December 2008.

Images

Explore the extraordinary diversity among leafhoppers through GigaPan images from the NCSU Insect Museum, which holds nearly 200 drawers of Cicadellidae including the collections of Z. P. Metcalf, D. A. Young, Wilhem Wager, others.

Cicadellidae

Selected Links

Compiled by P. A. Alvarez, L. L. Deitz, and D. R. Nimocks. 1 December 2008. Potentially useful sites may have been omitted because reviewers [Acknowledgments] noted significant misidentifications or other concerns.

Leafhoppers [C.H. Dietrich].

Brochosomes [R.A. Rakitov].

Checklist of Leafhopper Species: 1758-1955 [S.H. McKamey].

Sharpshooter Leafhoppers of the World

Illustrated Key to the Economically Important Leafhoppers of Australia (Hemiptera: Cicadellidae) [M.J. Fletcher].

Interactive Key to Tribes of Leafhoppers (Cicadellidae) [D. Dmitriev].

Cicadellinae Database Search [D. Takiya and D. Dmitriev].

Les Cicadelles du Qu�bec [Claude Pilon: The Leafhoppers of Quebec]. In French.

Sharpshooters, Leafhoppers (Cicadellidae) [M. R.Wilson, J. A., Turner & S. H. McKamey].

Xylella fastidiosa: Leafhopper Vector and Transmission.

Australian Faunal Directory: Membracoidea

Membracoidea of the World Database. [Stuart H. McKamey].

Leafhoppers (Cicadellidae) of the Pacific. [William J. Knight].

Deltocephalinae Database Search [J. Zahniser].

Megophthalminae Database Search [A. C. Gon�alves].

Selected References on Leafhoppers (Hemiptera: Membracoidea: Cicadellidae and Myerslopiidae): 1956-2013

Prepared by L. L. Deitz, P. A. Alvarez, C. H. Dietrich, S. H. McKamey, E. L. Blinka, and R. E. Baumler. 1 December 2008. (Updated 24 June 2014).

Burnside, V. W. 1971a. General Catalogue of The Homoptera. Fasc. VI, Cicadelloidea. Index to genera and species with addenda and corrigenda to parts 1-17. United States Department of Agriculture, Agricultural Research Service. 269 pp. [Burnside, V. W., formerly Wade, V.] [available online]

Cryan, J. R. 2005a. Molecular phylogeny of Cicadomorpha (Insecta: Hemiptera: Cicadoidea, Cercopoidea and Membracoidea): adding evidence to the controversy. Syst. Entomol. 30(4): 563-574.

Day, M. F.; Fletcher, M. J. 1994a. An annotated catalogue of the Australian Cicadelloidea (Hemiptera: Auchenorrhyncha). Invertebr. Taxon. 8(5): 1117-1288.

Dietrich, C. H. 2004a. Phylogeny of the leafhopper subfamily Evacanthinae with a review of Neotropical species and notes on related groups (Hemiptera: Membracoidea: Cicadellidae). Syst. Entomol. 29(4): 455-487.

Dietrich, C. H. 2005a. Keys to the families of Cicadomorpha and subfamilies and tribes of Cicadellidae (Hemiptera: Auchenorrhyncha). Florida Entomol. 88(4): 502-517.

Dietrich, C. H. 2006a. Annotated Bibliography of Literature Useful for Identification of Leafhoppers and Related Cicadomorpha. [organized by morphological studies, catalogues and bibliographies, regional works, and major taxonomic works] Online: http://www.inhs.uiuc.edu/~dietrich/LHbibliog.html.

Dietrich, C. H.; Rakitov, R. A.; Holmes, J. L. ; Black, W. C. IV. 2001a. Phylogeny of the major lineages of Membracoidea (Insecta: Hemiptera: Cicadomorpha) based on 28S rDNA sequences. Mol. Phylogenet. & Evol. 18(2): 293-305.

Dietrich, C. H.; Vega, F. E. 1995a. Leafhoppers (Homoptera: Cicadellidae) from Dominican amber. Ann. Entomol. Soc. Amer. 88(3): 263-270.

Emel'yanov, A. F.; Kirillova, V. I. 1989a,1990a. [Trends and modes of karyotype evolution in the Cicadina (Homoptera): I. Karyotypic features and evolutionary changes in the karyotypes of leafhoppers of superfamily Cicadelloidea.] Entomol. Obozr. 68(3): 587-603 (1989a). [in Russian, with English summary; English translation: Entomol. Rev. (Washington, D.C) 69(3): 62-80 (1990a); first author sometimes cited as Emeljanov]

Evans, J. W. 1966a. The leafhoppers and froghoppers of Australia and New Zealand (Homoptera: Cicadelloidea and Cercopoidea). Australian Mus. Mem. 12: 1-347.

Evans, J. W. 1977a. The leafhoppers and froghoppers of Australia and New Zealand (Homoptera: Cicadelloidea and Cercopoidea). Part 2. Rec. Australian Mus. 31(3): 83-129.

Hamilton, K. G. A. 1983a. Classification, morphology and phylogeny of the family Cicadellidae (Rhynchota: Homoptera). pp. 15-37. In Knight, W. J.; Pant, N. C.; Robertson, T. S.; Wilson, M. R. (eds.). Proceedings of the 1st International Workshop on Biotaxonomy, Classification and Biology of Leafhoppers and Planthoppers (Auchenorrhyncha) of Economic Importance, London, 4-7 October 1982. [Cover title: 1st International Workshop on Leafhoppers and Planthoppers of Economic Importance.]. Commonwealth Institute of Entomology, London. [ii] + 500 pp.

Hamilton, K. G. A. 1983b. Introduced and native leafhoppers common to the Old and New Worlds (Rhynchota: Homoptera: Cicadellidae). Canad. Entomol. 115 (5): 473-511.

Hamilton, K. G. A. 1985d,1985e. Leafhoppers of ornamental and fruit trees in Canada. Agric. Canada Publ. 1779/E: [1]-71 (1985d). [also available in French: Cicadelles des arbres ornementaux et fruitiers du Canada. Agric. Canada Publ. 1779/F: [1]-72 (1985e).]

Hamilton, K. G. A. 1999a. The ground-dwelling leafhoppers Myerslopiidae, new family, and Sagmatiini, new tribe (Homoptera: Membracoidea). Invertebr. Taxon. 13(2): 207-235.

Jones, J. R., and L. L. Deitz. 2009a. Phylogeny and systematics of the leafhopper subfamily Ledrinae (Hemiptera: Cicadellidae). Zootaxa 2186: [1]-120. [available online]

Kirillova, V. I. 1987a, 1988a. Chromosome numbers of the leafhoppers (Homoptera, Auchenorrhyncha) of the world fauna. II. Karyotypic peculiarities of leafhoppers of the superfamily Cicadelloidea. Entomol. Obozr. 66(2): 321-337 (1987a). [in Russian; English translation: Kirillova 1988a. Entomol. Rev. (Washington, D.C.) 67(1): 80-108 (1988a).]

Knight, W. J. 2010. Leafhoppers (Cicadellidae) of the Pacific. An annotated systematic checklist of the leafhoppers recorded in the Pacific region during the period 1758-2000. http://www.tymbal.org/KnightCatalogue.pdf

Le Quesne, W. J. 1965a. Cicadomorpha (excluding Deltocephalinae and Typhlocybinae). Handbooks for the Identification of British Insects (Royal Entomological Society of London) 2(2a): 1-64.

Le Quesne, W. J. 1969a. Hemiptera, Cicadomorpha, Deltocephalinae. Handbooks for the Identification of British Insects (Royal Entomological Society of London) 2(2b): 65-148.

Le Quesne, W. J.; Payne, K. R. 1981a. Cicadellidae (Typhlocybinae), with a check list of the British Auchenorrhyncha. Handbooks for the Identification of British Insects (Royal Entomological Society of London) 2(2): [1]-195.

Maramorosch, K.; Harris, K. F. (eds.). 1979a. Leafhopper Vectors and Plant Disease Agents. Academic Press, New York. xvi + 654 pp.

McKamey, S. H. 2000a [not dated, with periodic revisions]. Checklist of Leafhopper Species 1758-1955 (Hemiptera: Membracoidae: Cicadellidae and Myserslopiidae) with Synonymy and Distribution [Catalogue of the Homoptera, Fascicle 6, Abridged]. Online: http://www.bio-nica.info/biblioteca/McKameyCicadellidae.pdf [pdf version]; http://www.ars.usda.gov/News/docs.htm?docid=12015 [html version].

Mejdalani, G. L. F. 1998a. Morfologia externa dos Cicadellinae (Homoptera, Cicadellidae): comparação entre Versigonalia ruficauda (Walker) (Cicadellini) e Tretogonia cribrata Melichar (Proconiini), com notas sobre outras espécies e anáise da terminologia. [External morphology of the Cicadellinae (Homoptera, Cicadellidae): comparison between Versigonalia ruficauda (Walker) (Cicadellini) and Tretogonia cribrata Melichar (Proconiini), with notes on other species and an analysis of the structural terminology.] Rev. Brasileira Zool. 15(2): 451-544. [in Portuguese, with English abstract]

Metcalf, Z. P. 1962d. Cicadelloidea, Part 2, Hylicidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(2): [i-ii], 1-18. [with introduction by Young, D. A.; listed as Metcalf 1962a by Oman et al. 1990a] [available online:]

Metcalf, Z. P. 1962e. Cicadelloidea, Part 3, Gyponidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(3): [i-ii], 1-229. [with introduction by Young, D. A.; listed as Metcalf 1962b by Oman et al. 1990a] [available online:]

Metcalf, Z. P. 1962f. Cicadelloidea, Part 4, Ledridae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(4): [i]-vii, 1-147. [with introduction by Young, D. A.; listed as Metcalf 1962c by Oman et al. 1990a][available online:]

Metcalf, Z. P. 1962g. Cicadelloidea, Part 5, Ulopidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(5): [i]-vii, 1-101. [with introduction by Young, D. A.; listed as Metcalf 1962d by Oman et al. 1990a] [available online:]

Metcalf, Z. P. 1963c. Cicadelloidea, Part 6, Evacanthidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(6): [i-vi, 1-63. [with introduction by Young, D. A.; listed as Metcalf 1963a by Oman et al. 1990a] [available online:]

Metcalf, Z. P. 1963d. Cicadelloidea, Part 7, Nirvanidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(7): [i]-vi, 1-35. [with introduction by Young, D. A.; listed as Metcalf 1963b by Oman et al. 1990a] [available online:]

Metcalf, Z. P. 1963e. Cicadelloidea, Part 8, Aphrodidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(8): [i-ii], 1-268. [with introduction by Young, D. A.; listed as Metcalf 1963c by Oman et al. 1990a] [available online:]

Metcalf, Z. P. 1963f. Cicadelloidea, Part 9, Hecalidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(9): [i-ii], 1-123. [with introduction by Young, D. A.; listed as Metcalf 1963d by Oman et al. 1990a] [available online:]

Metcalf, Z. P. 1964a. Bibliography of the Cicadelloidea (Homoptera: Auchenorhyncha). General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(0): 1-349. [part number not indicated]

Metcalf, Z. P. 1964b. Cicadelloidea, Part 11, Coelidiidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(11): [i-ii], 1-182. [with introduction by Young, D. A.] [available online:]

Metcalf, Z. P. 1965a. Cicadelloidea, Part 1, Tettigellidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(1): [i-ii], 1-730. [with introduction by Young, D. A.] [available online:]

Metcalf, Z. P. 1965b. Cicadelloidea, Part 12, Eurymelidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(12): [i-ii], 1-43. [with introduction by Young, D. A.] [available online:]

Metcalf, Z. P. 1966a. Cicadelloidea, Part 13, Macropsidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(13): [i-ii], 1-261. [with introduction by Young, D. A.] [available online:]

Metcalf, Z. P. 1966b. Cicadelloidea, Part 14, Agalliidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(14): [i]-vi, 1-173. [with introduction by Young, D. A.] [available online:]

Metcalf, Z. P. 1966c. Cicadelloidea, Part 15, Iassidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(15): [i-ii], 1-229. [with introduction by Young, D. A.] [available online:]

Metcalf, Z. P. 1966d. Cicadelloidea, Part 16, Idioceridae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(16): [i-ii], 1-237. [with introduction by Young, D. A.] [available online:]

Metcalf, Z. P. 1968a. Cicadelloidea, Part 17, Cicadellidae. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(17): [i]-vii, 1-1513. [with introduction by Young, D. A.] [available online:]

Metcalf, Z. P. 1968b-d (dated 1967). Cicadelloidea, Part 10, Euscelidae. [In three sections]. General Catalogue of the Homoptera. United States Department of Agriculture, Agricultural Research Service, Washington, D.C. 6(10): [i-ii], i-v, 1-1077 (1968b); [i-iii], 1078-2074 (1968c); [i-ii], 2075-2695 (1968d). [with introduction by Young, D. A.; listed as Metcalf 1968a-c by Oman et al. 1990] [available online:] section 1, section 2, section 3

Nault, L. R.; Rodriguez, J. G.(eds.). 1985a. The Leafhoppers and Planthoppers. Wiley, New York. [xviii] + 500 pp.

Nickel, H. 2003a. The Leafhoppers and Planthoppers of Germany (Hemiptera: Auchenorrhyncha): Patterns and Strategies in a Highly Diverse Group of Phytophagous Insects. Pensoft, Sofia, [Bulgaria], and Goecke & Evers, Keltern, [Germany]. x + 460 pp.

Nielson, M. W. 1968b. The leafhopper vectors of phytopathogenic viruses (Homoptera: Cicadellidae): taxonomy biology, and virus transmission.. U. S. Dep. Agric. Tech. Bull. 1382: [i-ii], 1- 368.

Nielson, M. W. 1975a. A revision of the subfamily Coelidiinae (Homoptera: Cicadellidae). Tribes Tinobregmini, Sandersellini and Tharrini. Bull. Brit. Mus. (Natur. Hist.), Entomol., Suppl. 24: [1]-197.

Nielson, M. W. 1977a. A revision of the subfamily Coelidiinae (Homoptera: Cicadellidae). II. Tribe Thagriini. Pacific Insects Monogr. 34: [1]-218.

Nielson, M. W. 1979b. A revision of the subfamily Coelidiinae (Homoptera: Cicadellidae). III. Tribe Teruliini. Pacific Insects Monogr. 35: [1]-329.

Nielson, M. W. 1982e. A revision of the subfamily Coelidiinae (Homoptera: Cicadellidae). IV. Tribe Coelidiini. Pacific Insects Monogr. 38: [i-ii, 1]-318.

Nielson, M. W. 1983e. A revision of the subfamily Coelidiinae (Homoptera: Cicadellidae). V. New tribes Hikangiini, Youngolidiini, and Gabritini. Pacific Insects Monogr. 40: [i-ii,1]-78.

Nielson, M. W.; Godoy, C. 1995b. The Agalliinae of Central America (Homoptera: Cicadellidae). Contrib. Entomol., Int.1(2): 103-181.

Nielson, M. W.; Knight, W. J. 2000a. Distributional patterns and possible origin of leafhoppers (Homoptera: Cicadellidae). Rev. Brasileira Zool. 17(1): 81-156.

Oman, P. W.; Knight, W. J.; Nielson, M. W. 1990a. Leafhoppers (Cicadellidae): a Bibliography, Generic Check-list and Index to the World Literature 1956-1985. CAB International Institute of Entomology, Wallingford, Oxon, United Kingdom. iii + 368 pp.

Rakitov, R. A. 1998a [dated 1997]. On differentiation of cicadellid leg chaetotaxy (Homoptera, Auchenorrhyncha, Membracoidea). Russian Entomol. J. 6(3-4): 7-27.

Szwedo, J. 2004a. Annotated checklist on Myerslopiidae with notes on the distribution and origin of the group (Hemiptera: Cicadomorpha). Zootaxa 425: 1-15.

Vilbaste, J. 1982a. Preliminary key for the identification of the nymphs of north European Homoptera Cicadinea. II. Cicadelloidea. Ann. Zool. Fennici 19(1):1-20.

Viraktamath, C. A. 1998a. A revision of the leafhopper tribe Paraboloponini (Hemiptera: Cicadellidae: Selenocephalinae) in the Indian subcontinent. Bull. Natur. Hist. Mus., Entomol. Ser. 67(2): 153-207.

Wilson, M. R.; Claridge, M. F. 1991a. Handbook for the Identification of Leafhoppers and Planthoppers of Rice. CAB International for International Institute of Entomology in Association with Natural Resources Institute Wallingford, Oxon, UK. x + 142 pp.

Wilson, M. R.; Nault, L. R. (eds.). 1987a. Proceedings of 2nd International Workshop on Leafhoppers and Planthoppers of Economic Importance. CAB Int. Inst. Entomol., London. xiii + 368 pp.

Wilson, M. R.; Turner, J. A. 2007a. Progress in the study of sharpshooter leafhoppers (Hemiptera: Auchenorrhyncha: Cicadellidae) over 150 years: monographs, museums and individuals. Tijdschr. Entomol. 150: 289-303.

Wilson, M. R; Turner, J. A.; McKamey, S. H. 2009a. Sharpshooter Leafhoppers (Hemiptera: Cicadellinae). An Illustrated Checklist. Part 1: Old World Cicadellini. Studies in Terrestrial and Freshwater Biodiversity and Systematics from the National Museum of Wales. BIOTIR Reports 4: [1]-229.

Young, D.A., Jr. 1957c. The leafhopper tribe Alebrini (Homoptera: Cicadellidae). Proc. U. S. Natl. Mus. 107:127 277.

Young, D.A., [Jr.]. 1965a. Western Hemisphere Mileewanini (Homoptera, Cicadellidae). Zool. Beitr. (N.F.) 11(1 2): [369] 380. [Mileewini]

Young, D. A., [Jr.] 1968a. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 1. Proconiini. U. S. Natl. Mus. Bull. 261: [i-vii], [1]-287.

Young, D. A., [Jr.] 1977a. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New World Cicadellini and the genus Cicadella. North Carolina Agric. Exp. Sta. Tech. Bull. 239: [i]-vi, 1-1135.

Young, D. A., [Jr.] 1986a. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 3. Old World Cicadellini. North Carolina Agric. Res. Serv. Tech. Bull. 281: [i]-v, [1]-639.

Zahniser, J. N.; Dietrich, C. H. 2008a. Phylogeny of the leafhopper subfamily Deltocephalinae (Insecta: Auchenorrhyncha: Cicadellidae) and related subfamilies based on morphology. Syst. & Biodiv. 6(1): 1-24.

Zahniser, J.N.; Dietrich, C. H.  2013a. A review of the tribes of Deltocephalinae (Hemiptera: Auchenorrhyncha: Cicadellidae). Eur. J. Taxon. 45: 1-211.  [available online]

Zhang, Y. 1990a. A Taxonomic Study of Chinese Cicadellidae (Homoptera). T`ien tse ch`u pan she, Shan hsi sheng hsin hua shu tien fa hsing, Shan hsi [Shanxi]. iv + 218 pp. [in Chinese, with English summary; author sometimes cited as Chang, Y.]

Zhang Y.; Webb, M. D. 1996a. A revised classification of the Asian and Pacific selenocephaline leafhoppers (Homoptera: Cicadellidae). Bull. Natur. Hist. Mus., Entomol. Ser. 65 (1): 1-103.