Spittlebugs

(Superfamily Cercopoidea)

photo of a spittlebug Images

Selected Links

Selected References

Spittlebugs, or froghoppers, constitute the superfamily Cercopoidea, which currently includes up to 5 families: Aphrophoridae (treated as a subfamily of Cercopidae by some), Cercopidae, Clastopteridae, Epipygidae, and Machaerotidae (treated as a subfamily of Clastopteridae by some). This superfamily embraces approximately 2500 described species and 330 genera. The common name "spittlebug" arises from the lifestyle of many species, whose nymphs form spittle masses in which they develop, concealed from predators and protected from desiccation.

Distribution

Two spittlebug families, Cercopidae and Aphrophoridae, occur worldwide, with many species in the tropics, but relatively few in the Holarctic region. In contrast, Machaerotidae occur only in the Afrotropical, Australian, Indomalayan, and Palearctic regions. With the exception of a single Philippine genus, Clastopteridae are limited to the New World. Epipygidae occur only in the Neotropics.

Morphology

Spittlebugs range from 3 to 27 mm in length (including the wing tips). Most species resemble leafhoppers, but the hind tibiae have at most 1 or 2 strong lateral spines rather than longitudinal rows of enlarged setae. Also, the hind tibiae are shorter than those of leafhoppers, and the tibial apex has a crown of enlarged radiating spines. In froghoppers, the frontotclypeus is large and not only covers most of the face but also extends dorsally. Although many froghoppers have cryptic coloration, many species of the family Cercopidae are brightly colored, with some combination of red, red-orange, yellow, and black, perhaps representing warning coloration associated with reflex bleeding. Some adult Machaerotidae have a posteriorly directed spinelike scutellar process, superficially resembling the pronotal process of treehoppers. As an apparent adaptation for life within spittle masses, froghopper nymphs have the abdominal tergites extended downward as flaps, forming a completely or partially closed "breathing tube" that functions in respiration and the production of the spittle mass.

Ecology and Behavior

Spittlebugs, like cicadas and one subfamily of leafhoppers, feed on plant xylem sap. The specialized Malpighian tubules of immature froghoppers synthesize products that make the filtered sap viscous. Nymphs use the tip of the abdominal breathing tube to introduce bubbles of air into these excreta to form spittle. Many nymphal cercopids feed on roots, forming spittle masses at or below the soil surface. Clastopterids and most aphrophorids develop in spittle masses on aerial plant parts. Finally, machaerotid nymphs produce calcareous tubes or cases in which they live immersed in fluid secretions. When molting to the adult stage (and in some cases at other molts), machaerotid nymphs have been observed to exit their tubes and produce spittle. Many froghoppers are preferentially associated with nitrogen-fixing angiosperms: cercopids with nitrogen-fixing grasses, clastopterids with actinorhizal plants, and aphrophorids with legumes. In many parts of the world, grass-feeding spittlebugs are serious pests of pastures, sugarcane, and maize. Certain Costa Rican cercopids are facultatively aquatic. Their nymphs may either live submerged in the water-filled bracts of Heliconia flowers or may develop in spittle masses just like other spittlebugs. These are the only well-documented aquatic auchenorrhynchans.

Spittlebugs prepared by L. L. Deitz, V. Thompson, R. A. Rakitov, C. H. Dietrich, J. R. Cryan, and P. A. Alvarez. 1 December 2008.

Images

Explore the diversity among spittlebugs through GigaPan images from the NCSU Insect Museum (includes Z. P. Metcalf's collection). Four of the five families are represented (as subfamilies) of Cercopidae

Cercopidae

(drawers 1-8 = Cercopidae (as Cercopidae: Cercopinae); drawers 9-10 = Clasptopteridae (as Cercopidae: Clasptopterinae); drawers 11-17 = Aphrophoridae (as Cercopidae: Aphrophorinae); drawer 18, columns 3-4 = Machaerotidae (as Cercopidae: Machaerotinae).

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.

Cercopoidea Organized On Line (COOL).

Australian Faunal Directory: references on Cercopoidea.

Saratoga Spittlebug: Aphrophora saratogensis [L.F. Wilson].

Saratoga Spittlebug: Aphrophora saratogensis.

Selected References on Spittlebugs (Hemiptera: Cercopoidea): 1956-2014

Prepared by L. L. Deitz, V. Thompson, P. A. Alvarez, M. K. Whitley, and N. D. Caldwell. 1 December 2008 (updated 24 June 2014).

Bell, A. J.; Svenson, G. J.; Cryan, J.R. 2014a. The phylogeny and revised classification of Machaerotidae, the tube-making spittlebugs (Hemiptera: Auchenorrhyncha: Cercopoidea). Syst. Entomol. 39(3): 474-485.

Carvalho, C. S.; Webb, M. D. 2006a [dated 2005]. Cercopid Spittle Bugs of the New World (Hemiptera, Auchenorrhyncha, Cercopidae). Pensoft, Sofia, [Bulgaria]. 271 pp.

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

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

Drosopoulos, S. 2003a. New data on the nature and origin of colour polymorphism in the spittleug genus Philaneus (Hemiptera: Aaphrophoridae). Ann. Soc. Entomol. France (n.s.) 39(1): 31-42.

Drosopoulos, S.; Remane, R. 2000a. Biogeographic studies on the spittlebug Philaenus signatus Melichar, 1896 species group (Hemiptera: Aphrophoridae) with the description of two new allopatric species. Ann. Soc. Entomol. France (n..s.) 36(3): 269-277.

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.

Fennah, R. G. 1968a. Revisionary notes on the New World genera of cercopid froghoppers (Homoptera: Cercopidae). Bull. Entomol. Res. 58(1): 165-190.

Halkka, O.; Halkka, L. 1990a. Population genetics of the polymorphic spittlebug, Philaenus spumarius (L.). Evol. Biol. 24: 149-191.

Hamilton, K. G. A. 1982b. The spittlebugs of Canada (Homoptera: Cercopidae). Insects and Arachnids of Canada Part 10. Biosystematics Research Institute, Ottawa, Ontario. 102 pp.

Hamilton, K. G. A. 2001a. A new family of froghoppers from the American tropics (Hempitera: Cercopoidea: Epipygidae). Biodiversity 2(3): 15-21.

Hamilton, K. G. A.; Morales, C. F. 1992a. Cercopidae (Insecta: Homoptera). Fauna New Zealand 25: 1-37. [available online]

Komatsu, T. 1997a-c. A revision of the froghopper genus Aphrophora Germar (Homoptera, Cercopoidea, Aphrophoridae) from Japan, [parts 1-3]. Japanese J. Entomol. 65(1): 81-96 [1997a, part 1], 65(2): 369-383 [1997b, part 2], 65(3): 502-514 [1997c, part 3].

Liang, A. P. 1998a. Cladistic biogeography of Cercopoidea and Fulgoroidea (Insecta: Homoptera) in China and adjacent regions. Acta Zootaxonom. Sinica 23 (Suppl.): 132-166.

Liang, A. P. 2003a. A new tibial gland in male spittlebugs with descriptions of two new species of Augustohahnia Schmidt (Hemiptera: Cercopidae) from New Guinea. J. Zool. 261: 173-180.

Liang, A. P.; Fletcher, M. J. 2003a. Review of the Australian aphrophorid spittlebugs (Hemiptera: Aphrophoridae). Australian J. Entomol. 42(1): 84-93.

Maa, T. C. 1963a. A review of the Machaerotidae (Hemiptera: Cercopoidea). Pacific Insects Monogr. 5: 1-166.

Martin, R. M.; Cox, J. R.; Alston, D. G.; Ibarra F., F. 1995a. Spittlebug (Homoptera: Cercopidae) life cycle in buffelgrass in Northwestern Mexico. Ann. Entomol. Soc. Amer. 88(4): 471-478.

Metcalf, Z. P. 1960a. A Bibliography of the Cercopoidea (Homoptera: Auchenorhyncha). North Carolina State College [now University], Raleigh. iv + 262 pp.

Metcalf, Z. P. 1960b. Fascicle VII, Cercopoidea. Part 1. Machaerotidae. General Catalogue of the Homoptera (North Carolina State College [now University], Raleigh) 7(1): [i]-vi, 1-49. [with introduction by Metcalf, Z. P., and Young, D. A.]

Metcalf, Z. P. 1961a. Fascicle VII, Cercopoidea. Part 2. Cercopidae. General Catalogue of the Homoptera (North Carolina State College [now University], Raleigh) 7(2): [i]-vii, 1-607. [with introduction by Metcalf, Z. P., and Young, D. A.]

Metcalf, Z. P. 1962a. Fascicle VII, Cercopoidea. Part 3. Aphrophoridae. General Catalogue of the Homoptera (North Carolina State College [now University], Raleigh) 7(3): [i]-vii, 1-600. [with introduction by Metcalf, Z. P., and Young, D. A.]

Metcalf, Z. P. 1962b. Fascicle VII, Cercopoidea. Part 4. Clastopteridae. General Catalogue of the Homoptera (North Carolina State College [now University], Raleigh) 7(4): [i]-vi, 1-59. [with introduction by Metcalf, Z. P., and Young, D. A.]

Peck, D. C. 1999a. Seasonal fluctuations and phenology of Prosapia spittlebugs (Homoptera: Cercopidae) in upland pastures of Costa Rica. Environ. Entomol. 28(3): 372-386.

Peck, D. C. 2000a. Reflex bleeding in froghoppers (Homoptera: Cercopidae): variation in behavior and taxonomic distribution. Ann. Entomol. Soc. Amer. 93(5): 1186-1194.

Rakitov, R. A. 2002b. Structure and function of the Malphighian tubules, and related behaviors in juvenile cicadas: evidence of homology with spittlebugs (Hemiptera: Cicadoidea & Cercopoidea). Zool. Anz. 241: 117-130.

Stewart, A. J. A.; Lees, D. R. 1996a. The colour/pattern polymorphism of Philaenus spumarius (L.) (Homoptera: Cercopidae) in England and Wales. Phil. Trans. R. Soc. London, B, 351: 69-89.

Thompson, V. 1994a. Spittlebug indicators of nitrogen-fixing plants. Ecol. Entomol. 19: 391-398.

Thompson, V. 1997a. Spittlebug nymphs (Homoptera: Cercopidae) in Heliconia flowers (Zingiberales: Heliconiaceae): preadaptation and evolution of the first aquatic Homoptera. Rev. Biol. Trop. 45(2): 905-912.

Tishechkin, D. Y. 2003a. Vibrational communication in Cercopoidea and Fulgoroidea (Homoptera: Cicadina) with notes on classification of higher taxa. Russian Entomol. J. 12(2): 129-181. [with Russian abstract]

Wade, V. 1963a. Fascicle VII, Cercopoidea. Species index. General Catalogue of the Homoptera (North Carolina State College [now University], Raleigh) 7 (species index): [i-ii], 1-31. [NCSU Libraries: QL521.G4]

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