Distributional patterns and possible origin of leafhoppers (Homoptera, Cicadellidae)

Nielson, Mervin W.; Knight, William J.

doi:10.1590/S0101-81752000000100010

Abstract

The zoogeographical distribution of 42 cicadellid subfamilies and their assigned tribes and genera is compiled with distributional maps and proposed dispersal pathways of genera that are shared interzoogeographically. Possible origin of the subfamilies and tribes is proposed in an ancestral context from which the more modern extant groups evolved whereas origin of genera is in a more modern context. Notwithstanding their complex biogeography, the distributional data of the higher groups indicate that all of the cosmopolitan and near cosmopolitan subfamilies arose during early Cretaceous or possibly the late Jurassic period (140-116 m.y.a.) when continental drift was in its early stages. Nearly all of the New World and some Old World subfamilies are considered of more recent origin (late Cretaceous-Tertiary). Ninety percent of the known genera (2,126) are endemic to their respective zoogeographical region and subregion, thus indicating relatively high host specificity and low rate of dispersal. The majority (76%) of known extant genera are pantropical in origin, suggesting early or possible Gondwanaland origin of their ancestors. Dispersal pathways of genera shared by more than one zoogeographical region were generally south to north (Neotropical/Nearctic, Oriental/Palaearctic) or west to east (Palaearctic/Nearctic, Oriental/Australian), from regions of high diversity to regions of low diversity and from warmer climates to cooler climates. The most diverse and richest leafhopper fauna are present in the Neotropical and Ethiopian regions although taxal affinities between them are poorest. The most depauperate fauna are in the Nearctic region and in Australia, reflecting the impact of isolating and ecological factors on distribution and radiation. Ecological barriers were more evident between the Ethiopian and Oriental fauna than between any other zoogeographical combination. Taxal affinities appeared to be correlated with close continental proximities. Vicariance (physical) was the principal event that appealed to explain the distribution of many subfamilies and tribes whereas dispersal accounted for distribution of the majority of interzoogeographical genera.

Leafhoppers; Cicadellidae; distribution; dispersal; vicariance; origin; classificaton; habitats

Distributional patterns and possible origin of leafhoppers (Homoptera, Cicadellidae)

Mervin W. Nielson^I; William J. Knight^II

^IMonte L. Bean Museum, Brigham Young University, Provo, UT 8460, USA

^IIThe Natural History Museum, London, UK, SW7 5BD

ABSTRACT

The zoogeographical distribution of 42 cicadellid subfamilies and their assigned tribes and genera is compiled with distributional maps and proposed dispersal pathways of genera that are shared interzoogeographically. Possible origin of the subfamilies and tribes is proposed in an ancestral context from which the more modern extant groups evolved whereas origin of genera is in a more modern context. Notwithstanding their complex biogeography, the distributional data of the higher groups indicate that all of the cosmopolitan and near cosmopolitan subfamilies arose during early Cretaceous or possibly the late Jurassic period (140-116 m.y.a.) when continental drift was in its early stages. Nearly all of the New World and some Old World subfamilies are considered of more recent origin (late Cretaceous-Tertiary).

Ninety percent of the known genera (2,126) are endemic to their respective zoogeographical region and subregion, thus indicating relatively high host specificity and low rate of dispersal. The majority (76%) of known extant genera are pantropical in origin, suggesting early or possible Gondwanaland origin of their ancestors. Dispersal pathways of genera shared by more than one zoogeographical region were generally south to north (Neotropical/Nearctic, Oriental/Palaearctic) or west to east (Palaearctic/Nearctic, Oriental/Australian), from regions of high diversity to regions of low diversity and from warmer climates to cooler climates.

The most diverse and richest leafhopper fauna are present in the Neotropical and Ethiopian regions although taxal affinities between them are poorest. The most depauperate fauna are in the Nearctic region and in Australia, reflecting the impact of isolating and ecological factors on distribution and radiation. Ecological barriers were more evident between the Ethiopian and Oriental fauna than between any other zoogeographical combination. Taxal affinities appeared to be correlated with close continental proximities. Vicariance (physical) was the principal event that appealed to explain the distribution of many subfamilies and tribes whereas dispersal accounted for distribution of the majority of interzoogeographical genera.

Key words: Leafhoppers, Cicadellidae, distribution, dispersal, vicariance, origin, classificaton, habitats

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ACKNOWLEDGMENTS. We thank the following for their constructive and useful comments which immeasurably improved the content of the paper: Christopher H. Dietrich (Illinois Natural History Survey, Urbana, USA), Gabriel Mejdalani (Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil) and two anonymous reviewers. We express also our appreciation! to Randal Baker, Graphics Designer (Monte L. Bean Museum, Brigham Young University) for his computer-generated faunal distribution and taxal pathway maps. This study was facilitated by the Monte L. Bean Life Science Museum, Brigham Young University.

REFERENCES

All leafhopper references cited before 1956 are found in:

Metcalf, Z.P. 1964. General Catalogue of the Homoptera. Fascicle VI. Cicadelloidea. Bibliography of the Cicadelloidea (Homoptera: Auchenorrhyncha). U.S. Department of Agriculture, Agricultural Research Service, 349p.

Those cited between 1956-1985 are found in:

Oman, P.W.; W.J. Knight & M.W. Nielson. 1990. Leafhoppers (Cicadellidae): A bibliography, generic check-list and index to the world literature 1956-1985. Wallingford, Oxon, CAB. International Institute of Entomology, 368p.

All other references including non-leafhopper papers are cited below.

Abdul-Nour, H. 1988. Deux nouveaux genres et especes de Cicadellidae due Liban (Homoptera). Nouv. Rev. Entomol. 5(1): 35-41.

Arzone, A.; E. Vidano & A. Alma. 1987. Auchenorrhyncha introduced into Europe from the Nearctic region: Taxonomic and phytopathological problems, p.3-17. In: M.R. Wilson & L.R. Nault. (Eds). Proceedings of the 2^nd International Workshop on Leafhoppers & Planthoppers of Economic Importance. Provo, UT, C.A.B. International Institute of Entomology.

Bekker-Migdisova, E.E. 1949. Mesozoic Homoptera of middle Asia. Proc. Paleontol. Inst. USSR Acad. Sci. 22: 1-68.

Blocker, H.D.; Q. Fang & W.C. Black. 1995. Review of Nearctic deltocephaline-like leafhoppers. Ann. Entomol. Soc. Amer. 88 (3): 294-315.

Blocker, H.D. & K.J. Larsen. 1991. A new leaihopper genus, Cocrassana (Homoptera:Cicadellidae) from Mexican Tripsacinae and a synopsis of related genera. Jour. Kans. Entomol. Soc. 64 (2): 123-126.

Blocker, H.D. & M.D. Webb. 1990. The leafhopper genus Bythonia (Homoptera: Cicadellidae). Entomol. News 101 (5): 297-300.

Briggs, J.C. 1989. The historic biogeography of India: Isolation or Contact? Syst. Zool. 38 (4): 322-332.

Brown, J.H. & A.C. Gibson. 1983. Biogeography. St. Louis, The C.V. Mosby Company, 643 pp.

Brundin, L. 1967. Insects and the problem of austral disjunctive distribution. Ann. Rev. Entomol. 12: 149-168.

Crisci, J.V.; M.M. Cigliano; J.J. Morrone & S. Roig-Junent. 1991. Historical biogeography of southern South America. Syst. Zool. 40 (2): 152-171.

Darlington, P.J. 1957. Zoogeography: the geographical distribution of animals. New York, John Wiley & Sons Inc.

De Boer, A.J. & J.P. Duffels. 1996a. Historical biogeography of the cicadas of Wallacea, New Guinea and the West Pacific: a geotectonic explanation. Palaeo 124: 153-177.

______. 1996b. Biogeography of Indo-Pacific cicadas east of Wallace's line, p. 297-330. In: A. Keast & S.E. Miller (Eds). The origin and evolution of Pacific Island biotas, New Guinea to Eastern Polynesia: patterns and processes. Amsterdam, SPB Academic Publishing.

Dietrich, C.H. 1993. Phylogenetic analysis of leafhoppers (Homoptera: Cicadellidae) family-groups: Problems and prospects, p.27-28. In: S. Drosopoulos; P.V. Petrakis; M.F. Claridge & P.W. de Frijer (Eds). Proc. 8^th Auchenorrhyncha Congress, Delphi, Greece.

Dietrich, C.H. 1994. Systematics of the leafhopper genus Draeculacephala Ball (Homoptera: Cicadellidae). Trans. Amer. Entomol. Soc. 120 (2): 87-112.

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

Dlabola, J. 1987. Neue Zikadenarten von Saudi Arabien (Homoptera: Auchenorrhyncha). Annot. Zoolog. et Bot. 177: 1-11.

Dworakowska, I. 1972. Revision of the genus Aguriahana Dist. (Auchenorrhyncha, Cicadellidae, Typhlocybinae). Polskie Pismo Entomologiczne 42 (2): 273-312.

Eskov, K.Y. & S.I. Golovatch. 1986. On the origin of trans-Pacific disjunctions. Zool. Jahrb. Syst. 113: 265-285.

Fang, Q.; H.D. Blocker & W. C. Black. 1995. Cladistical analyses of Nearctic deltocephaline-like leafhoppers (Homoptera: Cicadellidae) using morphological and molecular data. Ann. Entomol. Soc. Amer. 88 (3): 316-323.

Felix, M. & G. Mejdalani. 1998. A new species of the Neotropical genus Bythonia (Hemiptera: Cicadellidae) and the female of B. consensa. Entomol. News 109: 307-314.

Freytag, P.H. 1987. Two new genera, Brevicisana and Minimana and four new species of Gyponinae (Homoptera: Cicadellidae). Ohio Jour. Sci. 87 (1): 33-35.

Freytag, P.H. 1989. Three new genera and six new species of South American gyponine leafhoppers (Homoptera: Cicadellidae). Trans. Kentucky Acad. Sci. 50 (1-2): 1-8.

______. 1990. Two new genera, Mediocerus and Mexicanocerus, and five new species of Idiocerinae (Homoptera: Cicadellidae). Trans. Kentucky Acad. Sci. 51 (1-2): 32-35.

Godoy, C. & M.D. Webb. 1994. Recognition of a new subfamily of Cicadellidae from Costa Rica based on phenetic analysis with similar taxa (Hemiptera: Homoptera: Auchenonhyncha). Trop. Zool. 7: 131-144.

Good, R. 1974. The geography of flowering plants. New York, John Wiley & Sons Inc., 4ª ed., 557p.

Hamilton, K.G.A. 1985. Review of Draeculacephala Ball (Homoptera, Auchenorrhyncha, Cicadellidae). Entomol. Abhand. Mus. Tierk. Dresden 49 (5): 83-103.

______. 1990. Insects from the Santana formation, lower Cretaceous, of Brazil. Homoptera. Bul. Amer. Mus. Nat. Hist. 195: 82-122.

Hayashi, M. & K. Aral 1990. Five new species of Pagaronia Ball (Homoptera, Cicadellidae - Cicadellinae) from Central Honshu. Esakia 1: 5-13.

Hayashi, M. & K. Yoshida. 1995. New species of the genus Pagaronia (Homoptera, Cicadellidae, Cicadellinae) from Japan. Japan Jour. Syst. Entomol. 1 (1): 73-93.

Hennig, W. 1965. Phylogenetic systematics. Ann. Rev. Entomol. 10: 97-116.

Kellogg, J.N. & V. Vega. 1995. Tectonic development of Panama, Costa Rica, and the Colombian Andes: Constraints from global positioning system geodetic studies and gravity, p.75-87, In: P. Mann (Ed.). Geological and tectonic development of the Caribbean plate boundary in southern Central America. Boulder, CO. Geological Society of America Special Paper 295.

Knight, W.J. & M.W. Nielson. 1986. The higher classification of the Cicadellidae. Tymbal 8: 10-14.

Knight, W.J. & M.D. Webb. 1988. Evidence for an Indo-Pacific origin of Hawaiian endemics in Balclutha and related genera (Cicadellidae: Homoptera: Macrostellini). Great Basin Nat. Mem. 12: 86-102.

______. 1993. The phylogenetic relationships between virus vector and other genera of macrosteline leafhoppers, including descriptions of new taxa (Homoptera: Cicadellidae: Deltocephalinae). Syst. Entomol. 18: 11-55.

Metcalf, Z.P. 1949. Zoogeography of the Homoptera. XIII Intern. Congr. Zool. 1948, p. 539-544.

Nielson, M.W. 1996. A new species of Myerslopia from Chile (Homoptera: Cicadellidae). Entomol. News 107 (5): 322-326.

Nielson, M.W. & C. Godoy. 1992 (1995). New species of Colladonus (Homoptera: Cicadellidae) from Costa Rica, with key to species. Brenesia 38: 37-44.

Oman, P.W.; W.J. Knight & M.W. Nielson. 1990. Leafhoppers (Cicadellidae): A bibliography, generic check-list and index to the world literature, 1956-1985. C.A.B. Internat. Inst. Entomol., 368p.

Shcherbakov, D.E. 1991. Diagnostics of the families of Auchenorrhyncha (Homoptera) as based on wings. I. Fore wing. Entomol. Oboz. 60: 828-843.

______. 1992. The earliest leafhoppers (Hemiptera: Karajassidae, n. fam.) from Jurassic of Karatau. Neue Jahrb. Geol. Paleontol. Monatsh. (1): 39-51.

______. 1993. Geological history of Auchenorrhyncha, p.3-4. In: S. Drosopoulos; P.V. Petrakis; M.F. Claridge & P.W. de Vrijer (Eds). Proc. 8^th Auchenorrhyncha Congr. Delphi, Greece, 112 P.

Smiley, C.J. 1979. Pre-tertiary phytogeographical and continental drift - some apparent discrepancies, p.311-319. In: J. Gray & A.J. Boucot (Eds). Historical biogeography, plate tectonics and the changing environment. Corvallis, Oregon State University Press.

Thapa, V.K. 1989. Some higher Himalayan typhlocybine leafhoppers (Homoptera: Cicadellidae) of Nepal. Insecta Matsumurana 42: 93-110.

Theron, J.G. 1986. New genera and species of southern African Coelidiinae (Homoptera: Ciadellidae), with description of the new tribe Equeefini. Phytophyl. 18: 153-163.

Thorne, R.F. 1973. Floristic relationships between tropical Africa and tropical America, p.27-47. In: B.J. Meggers; E.S. Ayensu & W.D. Duckworth (Eds.). Tropical forest ecosystems in Africa and South America: a comparative review. Washington, D.C., Smithsonian Institute Press.

Thulborn, R.A. 1986. Early Triassic tetrapod faunas of southeastern Gondwana. Alcheringa 10:297-313.

Viraktamath, C.A. & C.W. Wesley. 1988. Revision of the Nirvaninae (Homoptera: Cicadellidae) of the Indian subcontinent. Great Basin Nat. Mem. 12: 182-223.

Wallace, A.R. 1876. The geographical distribution of animals. London, Macmillan, Vol. 2.

Webb, M.D. 1987. The endemic Macrostelini of the island of St. Helen (Homoptera, Cicadellidae). Rev. Zool. Afric. 100: 453-464.

Wegner, A. 1924. Die Entstehung der Kontinente und Ozeane. Braunschweig, Vieweg.

Weijermars, R. 1989. Global tectonics since the breakup of Pangea 180 million years ago: Evolution maps and lithospheric budget. Earth-Science Rev. 26: 113-162.

Whitcomb, R.F. & A.L. Hicks. 1988. Genus Flexamia: New species, phylogeny, and ecology. Great Basin Nat. Mem. 12: 224-318.

Young, D.A. 1986. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 3. Old World Cicadellini. North Carolina State Univ. Tech. Bui. 281: 1-639p.

______. 1993. New genus and five new species of mileewine leafhoppers from New Guinea (Homoptera: Cicadellidae). Proc. Entomol. Soc. Wash. 95 (2): 228-240.

Zhang, Y. 1990. A taxonomic study of Chinese Cicadellidae (Homoptera). Tianze Eldonejo, Yangling, Shaanxi, China, 218p.

______. 1994. A taxonomic study of Chinese Coelidiinae (Homoptera, Cicadellidae). Henan Sci. & Techn. Press, 151p.

Zhang, Y. & M.D. Webb. 1993. A generic revision of the Austro-Oriental Selenocephalinae (Homoptera: Cicadellidae): Work in progress., p.35. In: S. Drosopoulos; P.V. Petrakis; M.F. Claridge & P.W.F. de Vrijer (Eds). Proc. 8^th Internat. Auchenorrhyncha Congr., Delphi, Greece, 112p.

Recebido em 23.VII.1998; aceito em 03.II.2000.

Metcalf, Z.P. 1964. General Catalogue of the Homoptera. Fascicle VI. Cicadelloidea. Bibliography of the Cicadelloidea (Homoptera: Auchenorrhyncha) U.S. Department of Agriculture, Agricultural Research Service, 349p.
Oman, P.W.; W.J. Knight & M.W. Nielson. 1990. Leafhoppers (Cicadellidae): A bibliography, generic check-list and index to the world literature 1956-1985. Wallingford, Oxon, CAB. International Institute of Entomology, 368p.
Abdul-Nour, H. 1988. Deux nouveaux genres et especes de Cicadellidae due Liban (Homoptera). Nouv. Rev. Entomol. 5(1): 35-41.
Arzone, A.; E. Vidano & A. Alma. 1987. Auchenorrhyncha introduced into Europe from the Nearctic region: Taxonomic and phytopathological problems, p.3-17. In: M.R. Wilson & L.R. Nault. (Eds). Proceedings of the 2^nd International Workshop on Leafhoppers & Planthoppers of Economic Importance Provo, UT, C.A.B. International Institute of Entomology.
Bekker-Migdisova, E.E. 1949. Mesozoic Homoptera of middle Asia. Proc. Paleontol. Inst. USSR Acad. Sci. 22: 1-68.
Blocker, H.D.; Q. Fang & W.C. Black. 1995. Review of Nearctic deltocephaline-like leafhoppers. Ann. Entomol. Soc. Amer. 88 (3): 294-315.
Blocker, H.D. & K.J. Larsen. 1991. A new leaihopper genus, Cocrassana (Homoptera:Cicadellidae) from Mexican Tripsacinae and a synopsis of related genera. Jour. Kans. Entomol. Soc. 64 (2): 123-126.
Blocker, H.D. & M.D. Webb. 1990. The leafhopper genus Bythonia (Homoptera: Cicadellidae). Entomol. News 101 (5): 297-300.
Briggs, J.C. 1989. The historic biogeography of India: Isolation or Contact? Syst. Zool. 38 (4): 322-332.
Brown, J.H. & A.C. Gibson. 1983. Biogeography St. Louis, The C.V. Mosby Company, 643 pp.
Brundin, L. 1967. Insects and the problem of austral disjunctive distribution. Ann. Rev. Entomol. 12: 149-168.
Crisci, J.V.; M.M. Cigliano; J.J. Morrone & S. Roig-Junent. 1991. Historical biogeography of southern South America. Syst. Zool. 40 (2): 152-171.
Darlington, P.J. 1957. Zoogeography: the geographical distribution of animals. New York, John Wiley & Sons Inc.
De Boer, A.J. & J.P. Duffels. 1996a. Historical biogeography of the cicadas of Wallacea, New Guinea and the West Pacific: a geotectonic explanation. Palaeo 124: 153-177.
______. 1996b. Biogeography of Indo-Pacific cicadas east of Wallace's line, p. 297-330. In: A. Keast & S.E. Miller (Eds). The origin and evolution of Pacific Island biotas, New Guinea to Eastern Polynesia: patterns and processes. Amsterdam, SPB Academic Publishing.
Dietrich, C.H. 1993. Phylogenetic analysis of leafhoppers (Homoptera: Cicadellidae) family-groups: Problems and prospects, p.27-28. In: S. Drosopoulos; P.V. Petrakis; M.F. Claridge & P.W. de Frijer (Eds). Proc. 8^th Auchenorrhyncha Congress, Delphi, Greece.
Dietrich, C.H. 1994. Systematics of the leafhopper genus Draeculacephala Ball (Homoptera: Cicadellidae). Trans. Amer. Entomol. Soc. 120 (2): 87-112.
Dietrich, C.H. & F.E. Vega. 1995. Leafhoppers (Homoptera: Cicadellidae) from Dominican amber. Ann. Entomol. Soc. Amer. 88 (3): 263-270.
Dlabola, J. 1987. Neue Zikadenarten von Saudi Arabien (Homoptera: Auchenorrhyncha). Annot. Zoolog. et Bot. 177: 1-11.
Dworakowska, I. 1972. Revision of the genus Aguriahana Dist. (Auchenorrhyncha, Cicadellidae, Typhlocybinae). Polskie Pismo Entomologiczne 42 (2): 273-312.
Eskov, K.Y. & S.I. Golovatch. 1986. On the origin of trans-Pacific disjunctions. Zool. Jahrb. Syst. 113: 265-285.
Fang, Q.; H.D. Blocker & W. C. Black. 1995. Cladistical analyses of Nearctic deltocephaline-like leafhoppers (Homoptera: Cicadellidae) using morphological and molecular data. Ann. Entomol. Soc. Amer. 88 (3): 316-323.
Felix, M. & G. Mejdalani. 1998. A new species of the Neotropical genus Bythonia (Hemiptera: Cicadellidae) and the female of B. consensa. Entomol. News 109: 307-314.
Freytag, P.H. 1987. Two new genera, Brevicisana and Minimana and four new species of Gyponinae (Homoptera: Cicadellidae). Ohio Jour. Sci. 87 (1): 33-35.
Freytag, P.H. 1989. Three new genera and six new species of South American gyponine leafhoppers (Homoptera: Cicadellidae). Trans. Kentucky Acad. Sci. 50 (1-2): 1-8.
______. 1990. Two new genera, Mediocerus and Mexicanocerus, and five new species of Idiocerinae (Homoptera: Cicadellidae). Trans. Kentucky Acad. Sci. 51 (1-2): 32-35.
Godoy, C. & M.D. Webb. 1994. Recognition of a new subfamily of Cicadellidae from Costa Rica based on phenetic analysis with similar taxa (Hemiptera: Homoptera: Auchenonhyncha). Trop. Zool. 7: 131-144.
Good, R. 1974. The geography of flowering plants. New York, John Wiley & Sons Inc., 4ª ed., 557p.
Hamilton, K.G.A. 1985. Review of Draeculacephala Ball (Homoptera, Auchenorrhyncha, Cicadellidae). Entomol. Abhand. Mus. Tierk. Dresden 49 (5): 83-103.
______. 1990. Insects from the Santana formation, lower Cretaceous, of Brazil. Homoptera. Bul. Amer. Mus. Nat. Hist. 195: 82-122.
Hayashi, M. & K. Aral 1990. Five new species of Pagaronia Ball (Homoptera, Cicadellidae - Cicadellinae) from Central Honshu. Esakia 1: 5-13.
Hayashi, M. & K. Yoshida. 1995. New species of the genus Pagaronia (Homoptera, Cicadellidae, Cicadellinae) from Japan. Japan Jour. Syst. Entomol. 1 (1): 73-93.
Hennig, W. 1965. Phylogenetic systematics. Ann. Rev. Entomol. 10: 97-116.
Kellogg, J.N. & V. Vega. 1995. Tectonic development of Panama, Costa Rica, and the Colombian Andes: Constraints from global positioning system geodetic studies and gravity, p.75-87, In: P. Mann (Ed.). Geological and tectonic development of the Caribbean plate boundary in southern Central America. Boulder, CO. Geological Society of America Special Paper 295.
Knight, W.J. & M.W. Nielson. 1986. The higher classification of the Cicadellidae. Tymbal 8: 10-14.
Knight, W.J. & M.D. Webb. 1988. Evidence for an Indo-Pacific origin of Hawaiian endemics in Balclutha and related genera (Cicadellidae: Homoptera: Macrostellini). Great Basin Nat. Mem. 12: 86-102.
______. 1993. The phylogenetic relationships between virus vector and other genera of macrosteline leafhoppers, including descriptions of new taxa (Homoptera: Cicadellidae: Deltocephalinae). Syst. Entomol. 18: 11-55.
Metcalf, Z.P. 1949. Zoogeography of the Homoptera. XIII Intern. Congr. Zool. 1948, p. 539-544.
Nielson, M.W. 1996. A new species of Myerslopia from Chile (Homoptera: Cicadellidae). Entomol. News 107 (5): 322-326.
Nielson, M.W. & C. Godoy. 1992 (1995). New species of Colladonus (Homoptera: Cicadellidae) from Costa Rica, with key to species. Brenesia 38: 37-44.
Oman, P.W.; W.J. Knight & M.W. Nielson. 1990. Leafhoppers (Cicadellidae): A bibliography, generic check-list and index to the world literature, 1956-1985 C.A.B. Internat. Inst. Entomol., 368p.
Shcherbakov, D.E. 1991. Diagnostics of the families of Auchenorrhyncha (Homoptera) as based on wings. I. Fore wing. Entomol. Oboz. 60: 828-843.
______. 1992. The earliest leafhoppers (Hemiptera: Karajassidae, n. fam.) from Jurassic of Karatau. Neue Jahrb. Geol. Paleontol. Monatsh (1): 39-51.
______. 1993. Geological history of Auchenorrhyncha, p.3-4. In: S. Drosopoulos; P.V. Petrakis; M.F. Claridge & P.W. de Vrijer (Eds). Proc. 8^th Auchenorrhyncha Congr. Delphi, Greece, 112 P.
Smiley, C.J. 1979. Pre-tertiary phytogeographical and continental drift - some apparent discrepancies, p.311-319. In: J. Gray & A.J. Boucot (Eds). Historical biogeography, plate tectonics and the changing environment. Corvallis, Oregon State University Press.
Thapa, V.K. 1989. Some higher Himalayan typhlocybine leafhoppers (Homoptera: Cicadellidae) of Nepal. Insecta Matsumurana 42: 93-110.
Theron, J.G. 1986. New genera and species of southern African Coelidiinae (Homoptera: Ciadellidae), with description of the new tribe Equeefini. Phytophyl. 18: 153-163.
Thorne, R.F. 1973. Floristic relationships between tropical Africa and tropical America, p.27-47. In: B.J. Meggers; E.S. Ayensu & W.D. Duckworth (Eds.). Tropical forest ecosystems in Africa and South America: a comparative review Washington, D.C., Smithsonian Institute Press.
Thulborn, R.A. 1986. Early Triassic tetrapod faunas of southeastern Gondwana. Alcheringa 10:297-313.
Viraktamath, C.A. & C.W. Wesley. 1988. Revision of the Nirvaninae (Homoptera: Cicadellidae) of the Indian subcontinent. Great Basin Nat. Mem. 12: 182-223.
Wallace, A.R. 1876. The geographical distribution of animals London, Macmillan, Vol. 2.
Webb, M.D. 1987. The endemic Macrostelini of the island of St. Helen (Homoptera, Cicadellidae). Rev. Zool. Afric. 100: 453-464.
Wegner, A. 1924. Die Entstehung der Kontinente und Ozeane Braunschweig, Vieweg.
Weijermars, R. 1989. Global tectonics since the breakup of Pangea 180 million years ago: Evolution maps and lithospheric budget. Earth-Science Rev. 26: 113-162.
Whitcomb, R.F. & A.L. Hicks. 1988. Genus Flexamia: New species, phylogeny, and ecology. Great Basin Nat. Mem. 12: 224-318.
Young, D.A. 1986. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 3. Old World Cicadellini. North Carolina State Univ. Tech. Bui. 281: 1-639p.
______. 1993. New genus and five new species of mileewine leafhoppers from New Guinea (Homoptera: Cicadellidae). Proc. Entomol. Soc. Wash. 95 (2): 228-240.
Zhang, Y. 1990. A taxonomic study of Chinese Cicadellidae (Homoptera). Tianze Eldonejo, Yangling, Shaanxi, China, 218p.
______. 1994. A taxonomic study of Chinese Coelidiinae (Homoptera, Cicadellidae) Henan Sci. & Techn. Press, 151p.
Zhang, Y. & M.D. Webb. 1993. A generic revision of the Austro-Oriental Selenocephalinae (Homoptera: Cicadellidae): Work in progress., p.35. In: S. Drosopoulos; P.V. Petrakis; M.F. Claridge & P.W.F. de Vrijer (Eds). Proc. 8^th Internat. Auchenorrhyncha Congr., Delphi, Greece, 112p.

Publication Dates

Publication in this collection
22 May 2009
Date of issue
Mar 2000

History

Received
23 July 1998
Accepted
03 Feb 2000

This work is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License.

[1] Metcalf, Z.P. 1964. General Catalogue of the Homoptera. Fascicle VI. Cicadelloidea. Bibliography of the Cicadelloidea (Homoptera: Auchenorrhyncha) U.S. Department of Agriculture, Agricultural Research Service, 349p.

[2] Oman, P.W.; W.J. Knight & M.W. Nielson. 1990. Leafhoppers (Cicadellidae): A bibliography, generic check-list and index to the world literature 1956-1985. Wallingford, Oxon, CAB. International Institute of Entomology, 368p.

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Distributional patterns and possible origin of leafhoppers (Homoptera, Cicadellidae)

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