Genus Metadelphax Wagner, 1963

[Back to Higher classification of Delphacidae]

Family Delphacidae Leach, 1815

Subfamily Delphacinae Leach, 1815

Tribe Delphacini Leach, 1815

Genus Metadelphax Wagner, 1963: 176.

Generic synonymy

= Metadelphax Wagner, 1963: 176; syn. with Toya by implication Fennah 1964: 142 (Type species moved to Toya); Nast 1972: 65, Linnavuori 1973: 107 (generic synonymy listed) (Metadelhax used as valid in Mitjaev 1971). Name restored by Ding 2006: 511; see also Gonzon & Bartlett 2008.
= Meadelphax Wagner, 1963: Missp. by Kuoh et al. 1983: 155.

Type species (in original combination): Delphax propinqua Fieber, 1866.

Recognized species

Five currently recognized species worldwide as follows:

Metadelphax argentinensis (Muir, 1929: 82)  – Argentina, Bolivia, Brazil, Chile, Columbia, Paraguay, Peru, Venezuela.
= Delphacodes argentinensis Muir, 1929: 82.
= Syndelphax argentinensis (Muir, 1929); comb. by Fennah, 1971: 323.
= Delphacodes variabilis De Remes, Lenikov & Teson, 1978; syn. by Teson & Remes Lenicov 1983: 322.
= Toya argentinensis (Muir, 1929); comb. by Teson & Remes Lenicov 1989: 101-102
= Metadelphax argentinensis (Muir, 1929); comb. by Gonzon & Bartlett 2008: 234-237.

Metadelphax dentata Gonzon and Bartlett, 2008 – Costa Rica, Ecuador, Nicaragua, Panama, Venezuela.

Metadelphax pero (Fennah 1971: 321) – Cayman Islands, Cuba
= Syndelphax pero Fennah, 1971: 321.
= Metadelphax pero (Fennah, 1971); comb. by Gonzon and Bartlett, 2008.

Metadelphax propinqua (Fieber, 1866: 525) – Cosmopolitan (found everywhere), especially in tropics and warm temperate regions; from Argentina to Canada, South Africa to Europe, and central Asia.
= Delphax propinqua Fieber, 1866b: 525.
= Delphax hamulata Kirschbaum, 1868: 38; syn. by Fieber 1872: 5.
= Liburnia propinqua (Fieber, 1866b); comb. by Fieber 1872: 5.
= Delphacodes propinqua (Fieber, 1866b); comb. by Muir 1917: 335.
= Liburnia terminalis Van Duzee, 1907: 49; syn. by Muir & Giffard 1924: 31.
= Liburnia tuckeri Van Duzee, 1912a: 506; syn. by Muir & Giffard 1924: 31.
= Delphacodes neopropinqua Muir, 1917: 335; syn. by Muir & Giffard 1924: 31.
= Delphacodes subfusca Muir, 1919b: 38; syn. by Muir & Giffard 1924: 31.
= Liburnia albicollis Haupt, 1935 (nec Motschulsky 1863): 144; syn. by Metcalf 1943: 498; see also Wagner 1954: 217-8 (syn. with Liburnia albicollis Motschulsky 1863, e.g., Lindberg 1936: 18; China 1938a: 54, Linnavuori 1952: 191, in error).
= Calligypona propinqua (Fieber, 1866b); comb. by Dlabola 1954: 14, 64 (comb. by implication in China 1954: 165).
= Delphacodes shirozui Ishihara, 1949: 53-54; syn. by Fennah 1956b: 122.
= Metadelphax propinqua (Fieber, 1866b); comb. by Wagner 1963: 170.
= Toya propinqua (Fieber, 1866b); comb. by Fennah 1964: 142 (from Delphacodes); see also Fennah 1965: 56 (syn. of Metadelphax by implication).
= Calligypona (Metadelphax) propinqua (Fieber, 1866b); comb. by Linnavuori 1964: 341-342.
= Delphacodes proquiqua (Fieber, 1866b); missp. by Wray 1967.
= M[etadelphax] propingua (Fieber, 1866b); missp. by Mitjaev 1971: 66.
= Toya (Calligypona) propinqua (Fieber, 1866b); comb. by Linnavuori 1973: 107.
= Delphax cataniae Matsumura, 1910: 35; syn. by Nast 1975: 4.
= Toya propinqua neopropinqua (Muir, 1917); status by Kuoh et al. 1983: 155.
= Liburnia marshalli Scott, 1873: 104; syn. by Webb & Wilson 1986: 286.
= Delphax graminicola Matsamura, 1910: 17; syn. by Yang 1989: 4.
= Toya porpinqua (Fieber, 1866b); missp. by Yang 1989: 219.
= Toya propincua (Fieber, 1866b); missp. by Rodriguez-León et al. 1994: 19.
= Metadelphax propinqua (Fieber, 1866b); restored comb. by Ding 2006: 511, 513-514; Gonzon & Bartlett, 2008: 242-246.

Metadelphax wetmorei (Muir and Giffard, 1924: 30) – USA: Florida, Georgia, Louisiana, North Carolina.
= Megamelus terminalis Crawford, 1914: 623 [nec Van Duzee 1907].
= Delphacodes wetmorei Muir & Giffard, 1924: 30 (Replacement name for unavailable Megamelus terminalis Crawford, 1914).
= Metadelphax wetmorei (Muir & Giffard, 1924); comb. by Gonzon & Bartlett 2008: 246-248.

Plant associations

Metadelphax propinqua is reported on about 15 grass species, including sugarcane (Saccharum officinarum L), barley (Hordeum vulgare L.), Bermuda grass (Cynodon dactylon (L.) Pers.), maize (Zea mays L.), and rice (Oryza sativa L).  Metadelphax propinqua reaches high population levels on Bermuda grass. Rizwan et al. (2020) reported this species in Pakistan in the highest numbers on  Leptochloa chinensis L., but also on Cynodon dactylon L., Helianthus allus L., Medicago polymorpha L., Trifolium alexandrinum L., Sorghum bicolor L., and Zea mays L.

Metadelphax argentinensis is reported from Oats (Avena sativa L.), barley (Hordeum vulgare L.), sorghum (Sorghum bicolor (L.) Moench), rye (Secale cereale L.), beet (Beta vulgaris var. cicla (L.) Koch, tomato (Solanum lycopersicum L.), pepper (Capiscum L.), and maize (Zea mays L.).

See Wilson et al. 1994; also see FLOW for hosts I may have missed.

Parasites

All for Metadelphax propinqua

Anagrus flaveolus Waterhouse, 1913 (Insecta, Hymenoptera, Mymaridae)(Virla & Rafael 1996: 34).
Elenchus tenuicornis (Kirby, 1815) (Insecta, Strepsiptera, Elenchidae);  Remes Lenicov & Teson 1990, Virla & Rafael 1996: 34)
Pseudogonatopus chilensis Olmi, 1989 (Insecta, Hymenoptera, Dryinidae); Virla (1995): 124

The other three species have no reported hosts.

Economic importance

Metadelphax propinqua is known to transmit Cynodon chlorotic streak nucleorhabdovirus (CCSV) in corn and bermudagrass, Maize rough dwarf virus (also here) (MRDV), Barley yellow striate mosaic cytorhabdovirus (also here) (BYSMV).

Metadelphax argentinensis has been taken on corn and other crops in Argentina, but its economic impact is unclear.

Recognition

Metadelphax is very similar to Toya in most respects.  They are both medium-sized and usually straw-colored.  Most species have darker markings alongside the carinae of the face.  The genera and species are best separated by features of the male genitalia.  Females (without males) usually can not be identified to species.

Metadelphax argentinensis

Metadelphax argentinensis is a relatively common species of temperate South America.  Additional images from Gonzon and Bartlett (2008) are here:

Metadelphax dentata

Metadelphax dentata is a recently described South and Central American species that had been confused with M. wetmorei.   Additional images from Gonzon and Bartlett (2008) are here:

Metadelphax pero (these photos are of the holotype)

This is a rarely encountered species.  Other than the type specimen from the Caymen islands, it has been reported from Cuba.  Additional images of the type specimen are here.

Metadelphax propinqua – a very common species

Metadelphax wetmorei

Metadelphax wetmorei is an uncommon species of the southeastern United States (possibly Mexico).  Additional images from Gonzon and Bartlett (2008) are here.

Online resources

American Insects
EOL
FLOW
Discover Life
Bugguide

Web Images

Flickr (search = Metadelphax)

Molecular resources

Molecular resources for Metadelphax are limited. BOLD lists data for M. propinqua.  A Genbank search in the taxonomy browser provides several gene sequences for M. propinqua.

Select references

(still incomplete)

Attié M., T. Bourgoin, J. Veslot and A. Soulier-Perkins. 2008. Patterns of trophic relationships between planthoppers (Hemiptera: Fulgoromorpha) and their host plants on the Mascarene Islands. Journal of Natural History 42(23-24): 1591-1638.

Ballou, C. H. 1936. Insect notes from Costa Rica in 1935. The Insect Pest Survey Bulletin 16: 437-497.

Bartlett, C.R. and G. Kunz. 2015. A new genus and species of delphacid planthopper (Hemiptera: Fulgoroidea: Delphacidae) from Central America with a preliminary regional species list. Zootaxa 3946(4): 510–518.(Erratum.Zootaxa 3963(4): 598-600).

Bartlett, C. R., L. B. O’Brien and S. W. Wilson. 2014. A review of the planthoppers (Hemiptera: Fulgoroidea) of the United States. Memoirs of the American Entomological Society 50: 1-287.

Bonfils, J., S. Quilici and B. Reynaud. 1994. Les Hémiptères Auchénorrhynques de l’Ile de la Réunion. Bulletin de la Société Entomologique de France 99: 227-240.

Borges P. A. V., A. Costa , R. Cunha, R. Gabriel, V. Gonçalves, A. F. Martins, I. Melo, M. Parente, P. Raposeiro, P. Rodrigues, R. S. Santos, L. Silva, P. Vieira and V. Vieira. 2010. Listagem dos organismos terrestres e marinhos dos Açores (A list of the terrestrial and marine biota from the Azores). Principia, Cascais (Portugal). 432 pp.

Buntin, G. D. 1988. Species composition and phenology of leafhoppers (Homoptera: Cicadellidae) and planthoppers (Homoptera: Delphacidae) in bermudagrass forage. Journal of Economic Entomology 81(1): 201-207.

China, W. E. 1938a. Die Arthropodenfauna von Madeira nach den Ergebnissen der Reise von Prof. Dr. O. Lundblad Juli – August 1935. III. Terrestrial Hemiptera (Hemiptera and Homoptera Auchenorrhyncha). Arkiv för Zoologi 30A (2): 1-68.

China, W. E. 1954.Delphacodes Fieber, 1866, versus Calligypona J. Sahlberg, 1871 (Homoptera: Delphacidae). Entomologist’s Monthly Magazine 90: 165.

Crawford, D. L. 1914a. A contribution toward a monograph of the homopterous insects of the family Delphacidae of North and South America. Proceedings of the United States National Museum 46:557-640, plus 6 plates.

Ding, J. H. 2006. Fauna Sinica Insecta Vol. 45 Homoptera Delphacidae. Science Press, Beijing, China. 776 pp + 20 plates.

Dlabola, J. 1954. Fauna CSR, Svazek l. Krisi -Homoptera [Fauna CSR, Volume l. Krisi – Homoptera]. Nakladatelstvi Ceskoslovenske Akademie Ved, Sekce Biologicka, Praha (Prague, Czech Republic). 1: 1-339 [In Slovakian with Russian and German summaries].

Endrestøl, A. 2013. Contribution to the Icelandic fauna of Auchenorrhyncha (Hemiptera, Fulgoromorpha & Cicadomorpha). Norwegian Journal of Entomology 60(1): 95-107.

Fennah, R. G. 1956b. Insects of Micronesia, Homoptera: Fulgoroidea. Bernice P. Bishop Museum Insects of Micronesia 6: 39-211.

Fennah, R. G. 1959. Delphacidae from the Lesser Antilles (Homoptera: Fulgoroidea). Bulletin of the British Museum (Natural History). Entomology 8: 245-365.

Fennah, R. G. 1964. Delphacidae from Madagascar and the Mascarene Islands (Homoptera: Fulgoroidea). Transactions of the Royal Entomological Society, London 116(7): 131-150.

Fennah, R. G. 1965. Delphacidae from Australia and New Zealand (Homoptera: Fulgoroidea). Bulletin of the British Museum (Natural History) Entomology 17(1): 1-59.

Fennah, R. G. 1971. Fulgoroidea from the Cayman Islands and adjacent Areas. Journal of Natural History 5: 299-342.

Fennah, R. G. 1975. Homoptera: Delphacidae from Ceylon. Entomologica Scandinavica (Supplement) 4: 79-136.

Fieber, F. X. 1866b. Grundzüge zur generischen Theilung der Delphacini. Verhandlungen der Kaiserlich-Königlichen Zoologish-botanischen Gesellschaft in Wien 16: 517-534.

Fieber, F. X. 1872. Katalog der europäischen Cicadinen, nach Originalien mit Benützung der neuesten Literatur. Carl Gerold’s Sohn, Wein [Vienna, Austria]. Pp. i-iv, 1-19.

Gonzon, A. T. and C. R. Bartlett. 2008[dated 2007]. Systematics of Hadropygos n. g., Metadelphax Wagner and New World Toya Distant (Hemiptera: Delphacidae). Transactions of the American Entomological Society 133(3/4): 205-277.

Haupt, H. 1935. Unterordnung: Gleichflügler, Homóptera. In Die Tierwelt Mitteleuropas. Ein Handbuch zu ihrer Bestimmung als Grundlage für faunistisch-zoogeographische Arbeiten. Unter Mitwirkung hervorragender Fachleute herausgegeben von P. Brohmer, P. Ehrmann und G. Ulmer. 4(3): 115-221.

Hayashi, M. and S. Fujinuma. 2016. Part Fulgoromorpha. In: Entomological Society of Japan 2016 – Catalogue of the insects of Japan. Volume 4 Paraneoptera (Psocodea, Thysanoptera, Hemiptera), 4. Editorial Committee of Catalogue of the Insects of Japan. p. 323-355.

Horváth, G. 1887. Izlet u Podsused u Zagrebackoj okolici mjeseca novembra. Glasnik Hrvatskoga Naravoslovnoga Druztva. Societas Historico-Naturalis Croatica 2: 267-274.

Ishihara, T. 1949. Revision of the Araeopidae of Japan, Ryuku Islands and Formosa (Hemiptera). Scientific Reports of the Matsuyama Agricultural College 2: 1-102 + 17 plates.

Kirschbaum, C. L. 1868. Die Cicadinen der gegend von Wiesbaden und Frankfurt A. M. nebst einer anzahl neuer oder Schwer zu unterscheidender Arten aus anderen Gegenden Europa’s Tabellarisch Beschrieben. Jahrbücher des Vereins für Naturkunde im Herzogthum Nassau. 21-22: 1-202.

Kuoh, C. L., J.-H. Ding, L. X. Tian and C. L. Hwang. 1983. Economic Insect Fauna of China. Fasc. 27. Homoptera, Delphacidae. Science Press, Beijing.

Leach, W. E. 1815. Entomology. The Edinburgh Encyclopedia 9: 57-172 [p. 125].

Lindberg, H. 1936. Die Cicadinen der Kanarischen Inseln. Commentationes Biologicae. Societas Scientiarum Fennica 6 (9): 1-19.

Linnavuori, R. 1952. Contributions to the Hemipterous fauna of Palestine. Annales Entomologici Fennici 18(4): 188-195

Linnavuori, R. 1964. Hemiptera of Egypt, with remarks on some species of the adjacent Eremian region. Annales Zoologici Fennici 1: 306-356.

Linnavuori, R. 1973. Hemiptera of the Sudan, with remarks on some species of the adjacent countries 2. Homoptera Auchenorrhyncha: Cicadidae; Cercopidae, Machaerotidae, Membracidae and Fulgoridae (Zoological contribution to the Sudan no. 33). Notulae Entomologicae 53(3): 65-137.

Llano C. A., C. R. Bartlett and G. Guevara. 2016. First record of the subfamily Asiracinae and Copicerus irroratus (Hemiptera: Auchenorrhyncha: Delphacidae) in Colombia. Florida Entomologist 99(1): 120-122.

Matsumura, S. 1910. Neue Cicadinen aus Europa und dem Mittelmeergebiet (Schluss). Journal of the College of Science, Imperial University of Tokyo 27 (18): 1-38.

Metcalf, Z. P. 1943. General catalogue of the Hemiptera. Fasc. IV. Fulgoroidea, Part 3. Araeopidae (Delphacidae), Smith College, Northampton, Massachusetts. 552 pp.

Mitjaev, I. D. 1971. Leafhoppers of Kazakhstan (Homoptera – Cicadinea). Nauka, Alma-Ata (Kazakhstan).

Mochida, O. and T. Okada. 1971. A list of the Delphacidae (Homoptera) in Japan with special reference to host plants, transmission of plant diseases, and natural enemies. Bulletin of the Kyushu Agricultural Experiment Station 15: 737-843.

de Motschulsky, V. 1863. Essai d’un catalogues des insectes de l’île Ceylan. Bulletin de la Société Impériale des Naturalistes de Moscou 36: 1-153.

Mozaffarian, F. and M. R. Wilson. 2011. An annotated checklist of the planthoppers of Iran (Hemiptera, Auchenorrhyncha, Fulgoromorpha) with distribution data. ZooKeys 145: 1-57.

Muir, F.A.G. 1917. Homopterous notes. Proceedings of the Hawaiian Entomological Society 3(4): 311-338.

Muir, F.A.G. 1919. Some New American Delphacidae. Canadian Entomologist 51: 35-39

Muir, F.A.G. 1929. New and little-known South American Delphacidae (Homoptera, Fulgoroidea) in the collection of the British Museum. Annals and Magazine of Natural History (Ser. 10) 3: 75-85.

Muir, F.A.G. and W. M. Giffard. 1924. Studies in North American Delphacidae. Bulletin of the Hawaiian Sugar Planters’ Association Division of Entomology 15: 1-53.

Nast, J. 1972. Palaearctic Auchenorrhyncha (Homoptera). An annotated check list. Polish Scientific Publishers, Warsaw. 550 pp.

Nast, J. 1975. On some Mediterranean Delphacidae (Homoptera, Auchenorrhyncha) described by S. Matsumura. Annales Zoologici (Warsaw) 33: 1-14.

Oliveira, C. M. De, E. De Oliveira, I. R. P. De Souza, E. Alves, W. Dolezal, S. Paradell, A. M. M. de Remes-Lenicov and M. R. Frizzas. 2013. Abundance and species richness of leafhoppers and planthoppers (Hemiptera: Cicadellidae and Delphacidae) in Brazilian maize crops. Florida Entomologist 96(4): 1470-1481. https://doi.org/10.1653/024.096.0427 

Oromi, P., S. De la Cruz and M. Báez. 2010. Hemiptera. p. 234-253. In: Arechavaleta M., Rodriguez S., Zurita N. & Garcia A.  Lista de especies silvestres de Canarias. Hongos, plantas y animales terrestres. 2009, Gobierno de Canarias, Tenerife (Spain).

Quaglino, F., F. Sanna, A. Moussa, M. Faccincani, A. Passera, P. Casati, P. A. Bianco and N. Mori. 2019. Identification and ecology of alternative insect vectors of ‘Candidatus Phytoplasma solani’ to grapevine. Scientific Reports 9(article 19522, ePub): 11 pp. https://doi.org/10.1038/s41598-019-56076-9 [Metadelphax noted]

de Remes-Lenicov, A.M. M., E. G. Virla and G. Varela. 1997. Description of preimaginal stages of Toya propinqua (Fieber) and notes on its behavior under laboratory conditions. Neotropica (La Plata) 43(109-110): 85-91.

Raatikainen, M., T. Raatikainen and A. Vasarainen. 1994. Effect of feeding by Metadelphax propinqua (Fieber) (Homoptera, Delphacidae) on barley. Agricultural Science in Finland 3: 513-517.

Raatikainen, M. and A. Vasarainen. 1990. Biology of Metadelphax propinqua (Fieber) (Homoptera: Delphacidae). Entomologica Fennica 1(3): 145-149.

de Remes Lenicov, A. M. M. and A. Teson. 1990. Estrepsipteros en Argentina parasitoides de insectos homopteros fulgoroideos (Insecta, Strepsiptera). Anales de la Sociedad Cientifica Argentina 220: 1-6.

de Remes-Lenicov, A.M.M. and A. Tesón. 1978. Contribucion al estudio de los fulgoridos Argentinos I. (Homoptera, Fulgoroidea, Delphacidae). Revista Sociedad Entomologica Argentina 37(1-4): 17-22.

de Remes-Lenicov, A.M.M. and E. G. Virla. 1999. Delfácidos asociados al cultivo de maíz en la República Argentina (Insecta – Homoptera – Delphacidae). Revista de la Facultad de Agronomía (La Plata) 104(1): 1-15.

Rizwan, M., B. Atta, A.M.M. de Remes Lenicov, R. Mariani, A. Makhdoom Sabir, M. Afzal. 2020. An Annotated List of Planthoppers with Alternate Hosts from Kallar Tract of Punjab, Pakistan. Punjab University Journal of Zoology. 35(2): 195-202. https://dx.doi.org/10.17582/journal.pujz/2020.35.2.195.202

Rodriguez-León, R., M. M. Hidalgo-Gatov and N. Novo. 1994. Fulgoroideos en la cana de azúcar y la vegetación sinantrópica [Fulgoroidea on sugar cane and border plant communities]. Ciencias Biologicas Academia de Ciencias de Cuba 27: 18-23.

Scott, J. 1873. Description of three new continental and one British species of Liburnia. Entomologist’s Monthly Magazine 10: 104-107.

Sokolov, I., X. Chen, R. M. Strecker and L. M. Hooper-Bùi. 2018. An Annotated List of Auchenorrhyncha and Heteroptera Collected in the Coastal Salt Marshes of the Mississippi Delta in Louisiana. Psyche 2018. 1-16. 10.1155/2018/1808370.

Tesón, A., and A.M.M. de Remes Lenicov. 1983. Contriuccion al estudio de los Fulgoridos Argentinos III. (Homoptera, Fulgoroidea, Delphacidae). Revista de la Sociedad Entomologica Argentina 42(1-4): 313-323.

Tesón, A., and A.M.M. de Remes Lenicov. 1989. Contribucion al estudio de los fulgoroideos argentinos. 4. (Homoptera, Fulgoroidea). Observaciones sobre tres especies halladas en cultivos de maiz (Zea mays L.). Revista Sociedad Entomologica Argentina 47(1-4): 101-107.

Van Duzee, E. P. 1907. Notes on Jamaican Hemiptera: A report on a collection of Hemiptera made on the Island of Jamaica in the spring of 1906. Bulletin of the Buffalo Society of Natural Sciences 8(5): 3-79.

Van Duzee, E. P. 1912. Hemipterological gleanings. Bulletin of the Buffalo Society of Natural Sciences. 10: 477-512.

Virla, E. G. and J. A. Rafael. 1996. Datos bionomicos preliminares y descripcion de la hembra de Cephalops penepauculus (Hardy) (Diptera: Pipunculidae) un parasitoide de Delphacidae (Homoptera: Auchenorryncha) en Argentina. Cirpon, Revista de Investigaciones 10(1-4): 33-36.

Virla, E. G. 1995. Biologia de Pseudogonatopus chilensis Olmi 1989 (Hymenoptera: Dryinidae). Acta Entomologica Chilena 19: 123-127.

Wagner, W. 1954. Die Fulgoroidea der Omer-Cooper-Expedition in die Lybische Wueste (Hemiptera-Homoptera). Bulletin de la Societe Fouad Ier d’Entomologie 38: 211-224.

Wagner, W. 1963 [dated 1962]. Dynamische Taxionomie, angewandt auf die Delphaciden Mitteleuropas. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 60: 111-189.

Webb, M. D. and M. R. Wilson. 1986. The Homoptera types described by John Scott in the family Delphacidae (Hemiptera). Entomologica Scandinavica 17(3): 279-289.

Wilson, B. H., S. Phillips and H. M. Harris. 1973. Species and seasonal occurrence of leafhoppers and planthoppers in a ‘Coastal’ bermudagrass pasture in the Macon Ridge area of Louisiana. Journal of Economic Entomology 66: 1346-1347.

Wilson, S. W., C. Mitter, R. F. Denno and M. R. Wilson. 1994. Evolutionary patterns of host plant use by delphacid planthoppers and their relatives. In: R.F. Denno and T.J. Perfect, (eds.). Planthoppers: Their Ecology and Management. Chapman and Hall, New York. Pp. 7-45 & Appendix [host references in appendix].

Wray, D. L. 1967. Insects of North Carolina, Third supplement. North Carolina Department of Agriculture, Raleigh, North Carolina.

Yang, C. T. 1989. Delphacidae of Taiwan II (Homoptera: Fulgoroidea). NSC Special Publications 6: 1-334