(Superfamily Membracoidea: Aetalionidae, Melizoderidae, and Membracidae)

photo of a treehopper Images

Selected Links

Selected References

The common name treehoppers broadly applies to three families--Aetalionidae, Melizoderidae, and Membracidae--which together include at least 3270 described species. Treehoppers are known for the remarkable diversity of their shapes and behaviors. Membracidae is the largest treehopper family (more than 400 genera and 3200 species), followed by Aetalionidae (6 genera, 42 species), and Melizoderidae (2 genera, 8 species).


Although treehoppers occur in all major zoogeographic regions, Europe has only a few species, and no indigenous species are known from New Zealand or Madagascar. Aetalionids are primarily Neotropical, but two species of Aetalion are recorded from the Nearctic region (Arizona and Florida), and one genus (Darthula, with two species) occurs in the Indomalayan region. Melizoderids are known only from Chile and Argentina. Membracidae, the largest and most widespread family, has 9 subfamilies, 8 of which are restricted to the New World (one Nearctic species of Smiliinae: Ceresini is introduced and widespread in the Palearctic). The remaining subfamily, Centrotinae, is the largest, with 6 New World tribes and 17 Old World tribes.


Treehoppers, which have 3 or fewer longitudinal rows of enlarged setae on the hind tibia, often have an oddly-shaped pronotum. In nearly all adults of the family Membracidae (2-24 mm in length) the pronotum is noticeably enlarged and extends posteriorly over the scutellum. Also, in membracid males, distinct lateral plates are differentiated from abdominal tergum IX, and, in nymphs (many of which have dorsal spines), abdominal segment IX forms an unbroken sheath around segment X. Melizoderid adults (4-6 mm in length) have an anterior pronotal horn, but lack posterior pronotal processes--in melizoderid nymphs abdominal tergum IX completely conceals the anal orifice dorsally. Aetalionids (3-30 mm) may have various modifications of the head (simple or with a pair of dorsal horns), pronotum (simple or with a middorsal process), or scutellum (simple or with an elevated keel). Also, in aetalionids the prothoracic trochanter and femur are fused, and females have abdominal tergum IX produced posteroventrally.

Ecology and Behavior

Treehoppers, like many Auchenorrhyncha, communicate by vibrational songs transmitted through stems and leaves of their host plants. Treehoppers are recorded from many herbaceous and woody hosts representing more than 100 plant families. Polyphagy is widespread among tropical treehoppers, but most Nearctic species are found on a single host genus or family. As a by-product of ingesting large quantities of plant sap (phloem), treehoppers secrete a sugary substance called "honeydew", which may serve as food for various opportunistic ants, bees, or wasps. Thus, treehoppers frequently benefit from interactions with ants that provide "house cleaning" services and sometimes protect the hoppers from predators in exchange for the honeydew secretions. Many treehoppers are gregarious or even subsocial, with some even guarding their eggs and immatures. Both parental care and ant-mutualism are common among aetalionids as well as membracids. Aetalionids and some membracids deposit egg masses on the surface of plants. Most membracids, however, insert the eggs into plant tissue--in so doing, a few species injure their hosts so seriously that they are considered plant pests.

Treehoppers prepared by L. L. Deitz, M. S. Wallace, C. H. Dietrich, S. H. McKamey, and M. J. Rothschild. 1 December 2008.


Explore the amazing diversity among treehoppers through GigaPan images from the NCSU Insect Museum (includes the collections of Z. P. Metcalf, D. A. Young, Wilhem Wager, and L. L. Deitz). All three treehopper families are represented:




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.

Treehoppers: Aetalionidae, Melizoderidae, and Membracidae (Hemiptera). [L.L. Deitz and M.S. Wallace].

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

Treehoppers [C.H. Dietrich].

Clowns of the Insect World: Treehoppers [Duane Flynn].

Australian Faunal Directory: references on Membracoidea [sensu lato].

Interactive Key to Tribes of New World Membracidae [C.H. Dietrich].

Anointing Behaviors and Malpighian Tubule Products in Membracidae [R.A. Rakitov].

Phylogeny and Systematics of Centrotinae [M.S. Wallace dissertation: differs slightly from published version].

Thorn Bug Umbonia crassicornis. [F.W. Mead].

Selected References on Treehoppers (Hemiptera: Membracoidea: Aetalionidae and Membracidae): 1956-2008

Prepared by L. L. Deitz and P. A. Alvarez. 1 December 2008.

Albertson, J. L.; Dietrich, C. H. 2005a. Systematics and phylogeny of the Neotropical treehopper subfamily Nicomiinae (Hemiptera, Membracidae). Rev. Brasileira Zool. 22(1): 231-283. [with Portuguese abstract]

Ananthasubramanian, K. S. 1996a. Fauna of India. Homoptera Membracidae. Zoological Survey of India, Calcutta, India. xviii + 534 pp.

Beirne, B. P. 1961a [not dated]. The Cicadas (Homoptera: Cicadidae) and Treehoppers (Homoptera: Membracidae) of Canada. Scientific Information Section, Research Branch, Canada Department of Agriculture, [Ottawa]. ii + 54 pp. + 9 pls. [year assumed to be 1961]

Broomfield, P. S. 1971a. A catalogue of the membracid types (Homoptera: Membracidae) in the British Museum (Natural History). Bull. Brit. Mus. (Natur. Hist.), Entomol. 25(8): 325-386.

Capener, A. L. 1962a. The taxonomy of the African Membracidae. Part 1. The Oxyrhachinae. Repub. South Africa, Dep. Agric. Tech. Serv., Entomol. Mem. 6: [i], [1]-164.

Capener, A. L. 1968a. The taxonomy of the African Membracidae. Part 2. The Centrotinae. Repub. South Africa, Dep. Agric. Tech. Serv., Entomol. Mem. 17: [i-iii], [1-124].

Creão-Duarte, A. J.; Sakakibara, A. M. 1998a. Anáise cladítica dos gêneros de Membracinae (Hemiptera, Auchenorrhyncha, Membracidae). [Cladistic analysis of the Membracinae genera (Hemiptera, Auchenorrhyncha, Membracidae).] Rev. Brasileira Zool. 15(4): 823-846. [in Portuguese, with English abstract]

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

Cryan, J. R.; Deitz, L. L. 1999a-2000a. Review of the New World treehopper tribe Stegaspidini (Hemiptera: Membracidae: Stegaspidinae): [parts I-III]. Proc. Entomol. Soc. Washington 101(3): 469-489 ([1999a: part I); 101(4): 760-778 (1999b: part II); 102(1): 82-98 (2000a: part III).

Cryan, J. R.; Wiegmann, B. M.; Deitz, L. L.; Dietrich, C. H.; Whiting, M. F. 2004a. Treehopper trees: phylogeny of Membracidae (Hemiptera: Cicadomorpha: Membracoidea) based on molecules and morphology. Syst. Entomol. 29: 441-454.

Cryan, J. R.; Robertson, J. A.; Deitz, L. L. 2004a [dated 2003]. The New World Treehopper Tribe Microcentrini (Hemiptera: Membracidae: Stegaspidinae): Monographic Revision and Phylogenetic Position. Thomas Say Monographs, Entomological Society of America, Lanham, Maryland. [iii] + 103 pp.

Day, M. F. 1999a. The genera of Australian Membracidae (Hemiptera: Auchenorrhyncha). Invertebr. Taxon. 13(4): 629-747.

Deitz, L. L. 1975a. Classification of the higher categories of the New World treehoppers (Homoptera: Membracidae). North Carolina Agric. Exp. Sta. Tech. Bull. 225: [i-ii], 1-177.

Deitz, L. L. 1989a. Bibliography of the Membracoidea (Homoptera: Aetalionidae, Biturritiidae, Membracidae, and Nicomiidae) 1981-1987. North Carolina Agric. Res. Serv. Tech. Bull. 290: [1]-31.

Deitz, L. L.; Bartlett, C. R. 2004a. Publications of Thomas Kenneth Wood (1942-2002). Proc. Entomol. Soc. Washington 106(3): 586-591. [lists his 50 works on treehoppers]

Deitz, L. L.; Dietrich, C. H. 1993a. Superfamily Membracoidea (Homoptera: Auchenorrhyncha). I. Introduction and revised classification with new family-group taxa. Syst. Entomol. 18(4): 287-296.

Deitz, L. L.; Kopp, D. D. 1987a. Bibliography of the Membracoidea (Homoptera: Aetalionidae, Biturritiidae, Membracidae, and Nicomiidae) 1956-1980. North Carolina Agric. Res. Serv. Tech. Bull. 284: [i-iii], 1-39.

Deitz, L. L.; Wallace, M. S. 2012a. Richness of the Nearctic treehopper fauna (Hemiptera: Aetalionidae and Membracidae. Zootaxa 3423: 1-26. [available as online publication]

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.; Deitz, L L. 1991a. Revision of the Neotropical treehopper tribe Aconophorini (Homoptera: Membracidae). North Carolina Agric. Res. Serv. Tech. Bull. 293: [1]-134 pp. [with Spanish abstract]

Dietrich, C. H.; Deitz, L L. 1993a. Superfamily Membracoidea (Homoptera: Auchenorrhyncha). II. Cladistic analysis and conclusions. Syst. Entomol. 18(4): 297-311.

Dietrich, C. H.; McKamey, S. H. 1995a. Two new Neotropical treehopper genera and investigation of the phylogeny of the subfamily Membracinae (Homoptera: Membracidae). Proc. Entomol. Soc. Washington 97(1): 1-16.

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.; McKamey, S. H.; Deitz, L. L. 2001a. Morphology-based phylogeny of the treehopper family Membracidae (Hemiptera: Cicadomorpha: Membracoidea). Syst. Entomol. 26(3): 213-239.

Godoy, C.; Miranda, X.; Nishida, K. 2006a. Membrácidos de la América Tropical. Treehoppers of Tropical America. 1 edit. Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica. 352 pp. [in Spanish and English].

Kopp, D. D.; Yonke, T. R. 1973a-1974a. The treehoppers of Missouri: [parts 1-4]. J. Kansas Entomol. Soc. 46(1): 42-64 [1973a (part 1)]; 46(2): 233-276 [1973b (part 2)]; 46(3): 375-421 [1973c (part 3)];; 47(1): 80-130 [1974a (part 4)].

Kopp, D. D.; Yonke, T. R. 1979a. A taxonomic review of the tribe Ceresini (Homoptera: Membracidae). Entomol. Soc. Amer. Misc. Publ. 11(2): 1-97.

Lin, C. P. [as C.-P.]; Danforth, B. N.; Wood, T. K. 2004a. Molecular phylogenetics and evolution of maternal care in membracine treehoppers. Syst. Biol. 53(3): 400-421.

McKamey, S. H. 1998a. Taxonomic Catalogue of the Membracoidea (exclusive of leafhoppers): Second Supplement to Fascicle 1 -- Membracidae of the General Catalogue of the Hemiptera. Mem. Amer. Entomol. Inst. 60: [1]-377.

McKamey, S. H.; Deitz, L. L. 1996a. Generic revision of the New World tribe Hoplophorionini (Hemiptera: Membracidae: Membracinae). Syst. Entomol. 21(4): 295-342.

Metcalf, Z. P.; Wade, V. 1963a. A Bibliography of the Membracoidea and Fossil Homoptera (Homoptera: Auchenorhyncha). North Carolina State [University], Raleigh. iv + 200 pp.

Metcalf, Z. P.; Wade, V. 1965a. 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. 1552 pp.

Quisenberry, S. S.; Yonke, T. R.; Kopp, D. D. 1978a. Key to the genera of certain immature treehoppers of Missouri with notes on their host plants (Homoptera: Membracidae). J. Kansas Entomol. Soc. 51(1): 109-122.

Roy, L. 2003a. Revue historique des Darnini (Hemiptera: Membracidae) au niveau generique. Notes Fauniques Gembloux 51: 33-47. [French, with English summary; includes updated list of genera and species]

Roy, L.; Guilbert, E.; Bourgoin, T. 2007a. Phylogenetic patterns of mimicry strategies in Darnini (Hemiptera: Membracidae). Ann. Soc. Entomol. France (n.s.) 43(3): 273-288. [with French abstract]

Sakakibara, A. M. 1999a. A synopsis of the tribe Micrutalini Haupt (Homoptera, Membracidae, Smiliinae). Rev. Brasileira Zool. 16 (supl. 1): 193-220.

Strümpel, H. 1972a. Beitrag zur Phylogenie der Membracidae Rafinesque. Zool. Jahrb., Abt. Syst., Ökol. & Geogr. Tiere 99(3): 313-407. [in German, with English summary]

Wade, V. 1966a. Species index of the Membracoidea and fossil Homoptera (Homoptera: Auchenorhyncha). A supplement to fascicle I - Membracidae of the General Catalogue of the Hemiptera. General Catalogue of the Homoptera. North Carolina State University, Raleigh. 40 pp.

Wallace, M. S. 2008a. Occurrence of treehoppers (Hemiptera: Membracidae: Smiliinae) on oaks in Delaware Water Gap National Recreation Area, 2004-2006. J. Insect Sci. 8: 59 [16 pp.]. [online publication; 42 color images representing 27 spp. of Smiliini]

Wallace, M. S. 2011a. Morphology-based phylogenetic analysis of the treehopper tribe Smiliini (Hemiptera: Membracidae: Smiliinae), with reinstatement of the tribe Telamonini. Zootaxa 3047: 1-42.) [available as online publication]

Wallace, M. S.; Deitz, L. L. 2004a. Phylogeny and systematics of the treehopper subfamily Centrotinae (Hemiptera: Membracidae). Mem. Entomol., Int. 19: [i]-iv, 1-377.

Wallace, M. S.; Deitz, L. L. 2006a. Australian treehoppers (Hemiptera: Membracidae: Centrotinae: Terentiini): phylogeny and biogeography. Invertebr. Syst. 20(2): 163-183.

Wood, T. K. 1984a. Life history patterns of tropical membracids (Homoptera: Membracidae). Sociobiology 8(3): 299-344.

Wood, T. K. 1993a. Diversity in the New World Membracidae. Annu. Rev. Entomol. 38: 409-435.

Yuan, F.; Chou. I. 2002a. Homoptera: Membracoidea: Aetalionidae [and] Membracidae. Fauna Sinica Insecta (Chinese Academy of Sciences, Beijing) 28: [i-v], [i]-xix, [1]-590 pp., Pls. I-IV. [in Chinese, with English summary]

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