Abstract
Members of the Smiliini, the nominotypical tribe of the large New World subfamily Smiliinae, are predominately Nearctic in distribution. This tribe included 169 mostly tree-feeding species in 23 genera. A parsimony-based phylogenetic analysis of an original dataset comprising 89 traditional and newly discovered morphological characters for 69 species, including representatives of 22 of the 23 described genera of Smiliini and five other previously recognized tribes of the subfamily, resulted in a single most parsimonious tree with three major clades. The broad recent concept of Smiliini (including Telamonini as a junior synonym) was not recovered as monophyletic by the analysis. Instead, the analysis supported narrower definitions of both Telamonini, here reinstated from synonymy, and Smiliini. A key and diagnoses are given to define these tribes, along with discussions of their phylogeny, biogeography, and host plant associations. The genera Antianthe Fowler, Hemicardiacus Plummer, Smilirhexia McKamey, and Tropidarnis Fowler are placed as Smiliinae, incertae sedis. Based on the phylogeny, several genera from both tribes including Atymna Stål, Cyrtolobus Goding, Heliria Stål, and Telamona Fitch are not monophyletic. Diagnostic characters emphasizing the morphological differences between the Smiliini and Telamonini include the dorsal margin of the head, the shape of the pronotum, the size of the pronotal humeral angles, the presence or absence of pronotal longitudinal rugae, the size of forewing cells, variations in the fusion of veins R and M apically in both the foreand hind wing, and the shape of the apex of the female second valvulae. Mapping geographic distribution onto the phylogeny suggests that the common ancestor of the ingroup (all three clades) occurred in Central America and Mexico, with multiple dispersals to temperate North America. Many Smiliini and Telamonini feed on various species of oak (Quercus) and the close evolutionary association between these insects and their hosts is discussed.References
Andrade, G.S. de. (2004) As espécies do gênero Ceresa Amyot & Serville (Hemiptera, Auchenorrhyncha, Membracidae). Revista Brasileira de Zoologia, 21(4), 671–738.
Ball, E.D. (1931) A monographic revision of the treehoppers of the tribe Telamonini of North America. Entomologica Americana, 12, 1−69.
Ballou, C.H. (1935) Insect notes from Costa Rica in 1934. The Insect Pest Survey Bulletin, 15, 163−212.
Bartlett, C.R., Deitz, L.L., Rothschild, M.J., & Wallace, M.S. (2008) Treehopper diversity (Hemiptera: Membracidae) of Little Orleans, Allegany Co., Maryland. Proceedings of the Entomological Society of Washington, 110, 130−143.
Beirne, B.P. (1961) The cicadas (Homoptera: Cicadidae) and treehoppers (Homoptera: Membracidae) of Canada. Scientific Information Section, Research Branch, Canada Department of Agriculture, Ottawa, 54 pp.
Cast, M.S. (2002 [unpublished]) A Molecular Phylogenetic Analysis of the Treehopper Subfamily Smiliinae (Hemiptera: Membracidae). M.S. thesis, Department of Entomology, University of Delaware, Newark, 51 pp.
Cocroft, R.B., & Rodríguez, R.L. (2005) The behavioral ecology of insect vibrational communication. Bioscience, 55(4), 323–334.
Cryan, J.R., Wiegmann, B.M., Deitz, L.L., Dietrich, C.H., & Whiting, M.F. (2004)Treehopper trees: phylogeny of Membracidae (Hemiptera: Cicadomorpha: Membracoidea) based on molecules and morphology. Systematic Entomology, 29, 441−454.
Dallwitz, M.J., Paine, T.A., & Zurcher, E.J. (1993) User’s guide to the DELTA System: a general system for processing taxonomic descriptions. 4thedition. Available from http://biodiversity.uno.edu/delta/ (accessed 23 December 2010)
Dallwitz, M.J., Paine, T.A., & Zurcher, E.J. (1999) User’s guide to the DELTA Editor. Available from http://biodiversity.uno.edu/delta/ (accessed 23 December 2010)
Deitz, L.L. (1974) Mild mannered mini‑monsters. Smithsonian, 5, 64−66, 70−71.
Deitz, L.L. (1975) Classification of the higher categories of the New World treehoppers (Homoptera: Membracidae). North Carolina Agricultural Experiment Station Technical Bulletin, 225, 1–177.
Deitz, L.L., & Wallace, M.S. (2010 and updates) Treehoppers: Aetalionidae, Melizoderidae, and Membracidae (Hemiptera). Available from http://treehoppers.insectmuseum.org (accessed 29 January 2011)
Dennis, C.J. (1952) Genitalia of the Membracidae of Wisconsin. The Canadian Entomologist, 84(6), 157−173.
Dennis, C.J. (1963) Membracidae in a southern Oklahoma woods. Proceedings of the Oklahoma Academy of Science, 43, 95−98.
Dennis, C.J. (1964) Observations on treehopper behavior (Homoptera, Membracidae). American Midland Naturalist, 71(2), 452−459.
Dennis, C.J. (1965) Oklahoma treehoppers (Homoptera, Membracidae). Proceedings of the Oklahoma Academy of Science, 45, 50−64.
Dennis, C.J. (1969) The treehoppers of Wisconsin in relation to the tension zone (Homoptera, Membracidae). American Midland Naturalist, 81(1), 236−242.
Dennis, C.J. (1970) A new treehopper of the genus Cyrtolobus from Wisconsin (Homoptera, Membracidae). Entomological News, 81(8), 191–194.
Dietrich, C.H., Rothschild, M.J, & Deitz, L.L. (1999) Checklist and host plants of the treehoppers (Hemiptera: Membracidae) of North Carolina. Proceedings of the Entomological Society of Washington, 101, 242−262.
Dietrich, C.H., McKamey, S.H., & Deitz, L.L. (2001a) Morphology-based phylogeny of the treehopper family Membracidae (Hemiptera: Cicadomorpha: Membracoidea). Systematic Entomology, 26, 213−239.
Dietrich, C.H., Rakitov, R.A, Holmes, J.L., & Black IV, W.C. (2001b) Phylogeny of the major lineages of Membracoidea (Insecta: Hemiptera: Cicadomorpha) based on 28s rDNA sequences. Molecular Phylogenetics and Evolution, 18(2), 293−305.
Evangelista, O., & Sakakibara, A.M. (2007) Aurimastris, new genus of the treehopper tribe Amastrini (Hemiptera, Membracidae, Smiliinae). Zootaxa, 1652, 63–68.
Froeschner, R.C. (1968) Telamona archboldi, a new treehopper from Florida (Homoptera: Membracidae). Proceedings of the Entomological Society of Washington, 70(2), 154−155.
Frost, S.W. (1957) More about Membracidae at lights. Entomological News, 68, 77−78.
Funkhouser, W.D. (1917) Biology of the Membracidae of the Cayuga Lake Basin. Cornell Agricultural Experimental Station Memoirs, 2, 177−445.
Funkhouser, W.D. (1923) Family Membracidae. In: Britton's Guide to the insects of Connecticut. Part IV. The Hemiptera or sucking insects of Connecticut. Bulletin. State of Connecticut. State Geological and Natural History Survey, 34, pp. 163−206.
Funkhouser, W.D. (1951) Homoptera Fam. Membracidae. Genera Insectorum, 208: 1–383.
Goding, F.W. (1892) A synopsis of the subfamilies and genera of the Membracidae of North America. Transactions of the American Entomological Society, 19, 253–260.
Goding, F.W. (1893) Food plants of some N.A. Membracidae. Insect Life, 5, 92−93.
Grimaldi, D.A., & Engel, M.S. (2005) Evolution of the Insects. Cambridge University Press, Cambridge, 755 pp.
Johnson, M.P., & Freytag, P.H. (1997) Treehoppers (Homoptera: Membracidae) on pin oak in Kentucky. Journal of the Kansas Entomological Society, 70, 21−30.
Kopp, D.D., & Yonke, T.R. (1973a) The treehoppers of Missouri: Part 1. Subfamilies Centrotinae, Hoplophorioninae, and Membracinae (Homoptera: Membracidae). Journal of the Kansas Entomological Society, 46, 42−64.
Kopp, D.D., & Yonke, T.R. (1973b) The treehoppers of Missouri: Part 2. Subfamily Smiliinae; tribes Acutalini, Ceresini, and Polyglyptini (Homoptera: Membracidae). Journal of the Kansas Entomological Society, 46(2), 233−276.
Kopp, D.D., & Yonke, T.R. (1973c) The treehoppers of Missouri: Part 3. Subfamily Smiliinae; tribe Smiliini. Journal of the Kansas Entomological Society, 46, 375−421.
Kopp, D.D., & Yonke, T.R. (1974) The treehoppers of Missouri: Part 4. Subfamily Smiliinae; tribe Telamonini (Homoptera: Membracidae). Journal of the Kansas Entomological Society, 47, 80−130.
Lawson, P.B. (1922) The Membracidae of Kansas. University of Kansas Scientific Bulletin, 14, 27−110.
Maddison, D.R., & Maddison, W.P. (2003) ‘MacClade 4: Analysis of Phylogeny and Character Evolution.’ Version 4.06c. Sinauer Associates, Sunderland, MA.
Manos, P.S., & Stanford, A.M (2001) The historical biogeography of Fagaceae: tracking the Tertiary history of temperate and subtropical forests of the Northern hemisphere. International Journal of Plant Sciences, 162(6), 77−93.
McKamey, S.H. (1998) Taxonomic Catalogue of the Membracoidea (exclusive of leafhoppers): Second Supplement to Fascicle 1 — Membracidae of the General Catalogue of the Hemiptera. Memoirs of the American Entomological Institute 60, [1]−377.
McKamey, S.H. (2008) A new genus, Smilirhexia, of Smiliini (Hemiptera, Membracidae) from Costa Rica. ZooKeys, 3, 51−55.
Metcalf, Z.P., & Wade, V. (1965) General Catalogue of the Homoptera. A Supplement to Fascicle I ‑ Membracidae of the General Catalogue of Hemiptera. Membracoidea. In Two Sections. North Carolina State University, Raleigh, NC, 1552 pp.
Osborn, H. (1940) The Membracidae of Ohio. Bulletin of the Ohio Biological Survey, 7, 51−101.
Plummer, C.C. (1936) New Membracidae (Homoptera) from Mexico, with notes on other species. Annals of the Entomological Society of America, 29, 682−693.
Plummer, C.C. (1938) The genus Atymna Stål and a new related genus (Homoptera: Membracidae).Journal of the New York Entomological Society, 46, 235−242.
Plummer, C.C. (1945) New Membracidae from Central America. Proceedings of the Entomological Society of Washington, 47, 39−44.
Quisenberry, S.S., Yonke, T.R., & Kopp, D.D. (1978) Key to the genera of certain immature treehoppers of Missouri with notes on their host plants (Homoptera: Membracidae). Journal of the Kansas Entomological Society, 51(1), 109−122.
Sakakibara, A.M. (1996) Taxonomic notes on some Polyglyptini: descriptions of new genus and new species (Homoptera, Membracidae, Smiliinae). Revista Brasileira de Zoologia, 13(2), 463–474.
Sakakibara, A.M. (1997) The tribe Acutalini Fowler (Homoptera, Membracidae, Smiliinae): new genera, new species and some nomenclatural changes. Revista Brasileira de Zoologia, 14(3), 659–674.
Sakakibara, A.M. (1999) A synopsis of the tribe Micrutalini Haupt (Homoptera, Membracidae, Smiliinae). Revista Brasileira de Zoologia, 16, (Supl. 1), 193–220.
Swofford, D.L. (2002) ‘PAUP*. Phylogenetic Analysis Using Parsimony (*and other methods). Version 4.’ Sinauer Associates, Sunderland, MA.
Stål, C. (1866) Hemiptera Homoptera Latr. Hemiptera Africana 4. Norstedt, Stockholm, Sweden, 276 pp.
USDA: NRCS. (2010) The PLANTS Database. National Plant Data Center, Baton Rouge, LA. Available from http://plants.usda.gov (accessed 23 December 2010).
Wallace, M.S. (2008) Occurrence of treehoppers (Hemiptera: Membracidae: Smiliinae) on oaks in Delaware Water Gap National Recreation Area, 2004−2006. Journal of Insect Science, 8, 59, available online: insectscience.org/8.59.
Wallace, M.S. (2010 and updates) Interactive keys to the genera and higher taxa of treehoppers (Hemiptera: Membracidae) of the United States. Parts 1–3. In: Deitz, L.L., and Wallace, M.S. (team leaders). Treehoppers: Aetalionidae, Melizoderidae, and Membracidae (Hemiptera). Available from http://treehoppers.insectmuseum.org/projects/74/public/site/treehoppers/home/primer (accessed 29 January 2011).
Wallace, M.S., & Deitz, L.L. (2004) Phylogeny and systematics of the treehopper subfamily Centrotinae (Hemiptera: Membracidae). Memoirs on Entomology International, 19, 1−377.
Wallace, M.S., & Deitz, L.L. (2006) Australian treehoppers (Hemiptera: Membracidae: Centrotinae: Terentiini): phylogeny and biogeography. Invertebrate Systematics, 20(2), 163–183.
Wallace, M.S., Deitz, L.L., & Rothschild, M.J. (2003) Checklist of the treehoppers (Hemiptera: Membracidae) of Great Smoky Mountains National Park. Proceedings of the Entomological Society of Washington, 105, 578−591.
Wallace, M.S., & Troyano, N.M. (2006) The oak-feeding smiliine treehoppers (Hemiptera: Membracidae) of Delaware Water Gap National Recreation Area. Journal of the Pennsylvania Academy of Science, 80, 28−34.
Wood, T.K. (1984) Life history patterns of tropical membracids (Homoptera: Membracidae). Sociobiology, 8, 3, 299–345.
Wood, T.K. (1993) Diversity in the New World Membracidae. Annual Review of Entomology, 38, 409−435.
Wood, T.K., & Olmstead, K.L. (1984) Latitudinal effects on treehopper species richness (Homoptera: Membracidae). Ecological Entomology, 9, 109−115.
Wood, T.K., & Pesek, J.D. (1992) Pronotal shape: a source of confusion or panacea in systematic studies of treehoppers (Homoptera: Membracidae)? In: Sorensen, J.T., & Foottit, R. (Eds.), Ordination in the Study of Morphology, Evolution and Systematics of Insects: Applications and Quantitative Genetic Rationals. Elsevier, Amsterdam, pp. 349−384.
Woodruff L.B. (1924) Critical observations in the membracid genus Cyrtolobus Goding. (Hemip.-Homop.). Journal of the New York Entomological Society, 32, 1−62.
Yothers, M.A., & Allen, P.B., Jr. (1941) Observations of the biology and control of the treehopper Heliria praealta (Fowler) in orchards of the Pacific Northwest. USDA Circular, 606, 1−12.