Taxonomic study of the genus Unkanodes (Hemiptera, Fulgoroidea, Delphacidae) from Pakistan, with description of a new species
expand article infoKamran Sohail, Hassan Naveed§, Daozheng Qin, Yalin Zhang
‡ Northwest A&F University, Yangling, China
§ Leshan Normal University, Leshan, China
Open Access


Unkanodes (Kwonianella) malamjabbensis sp. nov. (Hemiptera, Delphacidae) is described and illustrated and U. latespinosa (Dlabola, 1957) is newly recorded from Malamjabbah, Swat, Pakistan. These two species represent the first records of the genus Unkanodes Fennah, 1956 from Pakistan. A key to the world’s species of the genus Unkanodes is provided.


Distribution, Fulgoromorpha, key, morphology, taxonomy


The planthopper family Delphacidae currently consists of 2217 species in 427 genera (Bourgoin 2020). Delphacids are small insects that can be easily distinguished by the presence of a large, movable spur (the “calcar”) at the apex of the hind tibiae (Bartlett 2014). Most delphacids are grass-feeders, although some feed on other monocots such as sedges and rushes, and some feed on dicots (Bartlett 2019). A number of species feed on economically important crops, such as sugarcane, maize and rice (Wilson and O’Brien 1987; Wilson et al. 1994). The plant order Poales accounts for more than 70% of their hosts, while all other plant orders provide only 3%. In the Poales group, the family Poaceae has the highest (52%) percentage of hosts, followed by Cyperaceae (16.5%), while the remaining families account for only 3% of host records (Bourgoin 2020).

The delphacid fauna of Pakistan has been poorly studied, with only ten species previously recorded from this country (Bourgoin 2020). This figure, consisting of about 0.4% of the world’s described species, likely largely underrepresents the actual diversity of delphicids in this country. The genus Unkanodes was established by Fennah (1956) with the type species Unkanodes sapporona (Matsumura, 1935) from Che-Kiang (Zhejiang Province of China). Currently, this genus comprises nine species (Bartlett 2019) occurring in Afghanistan, Alaska, Austria, China, Denmark, Estonia, Finland, Germany, Greece, Iran, Japan, Lithuania, Mongolia, Poland, Russia, South Korea, Sweden, Taiwan, Turkey, Ukraine, Yugoslavia, and the U.S.A. (Bourgoin 2020). The genus Unkanodes is economically important and its members are vectors of many diseases in rice and cereals and also the causative agents of hopper burn diseases. Unkanodes albifascia is responsible for transmission of NCMV (Northern cereal mosaic virus), RBSDV (Northern cereal mosaic virus) and stripe disease. Unkanodes sapporona is involved in the transmission of NCMV, RSV (Rice stripe tenuivirus), and RBSDV. Unkanodes tanasijevici is reported to be a vector of IMMV (Iranian Maize Mosaic Nucleorhabdovirus), MIMV (Maize Iranian Mosaic Virus), MRDV (Maize Rough Dwarf Fiji Virus), and RBSDV (Bartlett 2019).

In this study, U. (Unkanodes) latespinosa (Dlabola, 1957) is recorded for the first time from Pakistan and a new species U. (Kwonianella) malamjabbensis sp. nov., is described.

Materials and methods

Specimens were collected from Pakistan and deposited at the Entomological Museum of Northwest A&F University (NWAFU) Yangling, Shaanxi, China. Morphological terminology follows Asche (1985), Wilson (2005) and Bartlett et al. (2014). The method for genitalia preparation and clearing follows Wilson and McPherson (1980) and Wilson (2005). Morphological characters were observed using the stereomicroscope Olympus SZX10. Measurements of characters are given in millimeters (mm). Photographs of the adults were taken using a Zeiss AxioCam ICc 5. Adobe Photoshop was used for labeling and plate composition of the obtained images.


Family Delphacidae Leach, 1815

Subfamily Leach, 1815

Tribe Delphacini Leach, 1815

Unkanodes Fennah, 1956

Unkanodes Fennah, 1956.

Unkanodes sapporona (Matsumura, 1935), comb. by Fennah 1956: 474.

Type species

Unkana sapporona Matsumura, 1935: 131, by original designation.


Relatively slender, head slightly narrower than pronotum. Vertex longer than broad, its width at base not exceeding width of an eye, shallowly rounded at apical margin; carinae of vertex and frons distinct. Frons parallel-sided, about 2.0–2.5 times as long as wide, lateral margins parallel, narrowing upwards in apical 1/3; median carina of frons bifurcates near fastigium. Lateral carinae of pronotum diverging, vanishing before reaching posterior margin. Calcar with 10–20 well-developed teeth; apical tooth separate from the remaining teeth. Posterior margin of pygofer with a cut on the sides. Segment X (anal tube) with a pair of teeth or teeth absent. Styli flattened, diverging or more or less parallel beyond middle, with complex apices, zigzag-shaped bent and wide or narrowed and slanting outwards. Armature of diaphragm (bridge of pygofer) bearing a pair of teeth directed upwards or a projection with 2 apices. Aedeagus more or less straight, or bent ventrad, elbow-shaped, slightly asymmetrical due to location of gonopore and arrangement of teeth on aedeagal shaft (after Fennah 1956 and Anufriev and Emeljanov 1988).

Checklist of species of the genus Unkanodes Fennah

Subgenus Unkanodes (Chilodelphax Vilbaste, 1968)

Unkanodes (Chilodelphax) silvaticus Vilbaste, 1968

Unkanodes silvaticus Vilbaste, 1968: 24.

Chilodelphax silvaticus (Vilbaste, 1968); comb. by Kwon 1982: 4.

Unkanodes (Chilodelphax) silvaticus Vilbaste, 1968; comb. by Anufriev and Emeljanov 1988: 409.

Subgenus Unkanodes (Unkanodes Fennah, 1956)

Unkanodes (Unkanodes) excisa (Melichar, 1898)

Liburnia excisa Melichar, 1898: 67.

Delphax excisa (Melichar, 1898); comb. by Puton 1899: 108.

Liburnia elymi Jensen-Haarup, 1917: 3; syn. by Jensen-Haarup 1920: 53.

Delphacodes excisa (Melichar, 1898); comb. by Metcalf 1943: 436.

Elymodelphax excisa (Melichar, 1898); comb. by Wagner 1963: 167.

Unkanodes excisa (Melichar, 1898); comb. by implication Dlabola 1965: 86.

Unkanodes (Unkanodes) latespinosa (Dlabola, 1957)

Calligypona latespinosa Dlabola, 1957.

Unkanodes latespinosa (Dlabola, 1957), comb. apparently by Dlabola 1964: 240 (see also Dlabola 1967: 53; Emeljanov 1977: 113).

Unkanodes (Unkanodes) paramarginata (Dlabola, 1961: 275)

Unkanodes (Unkanodes) sapporona (Matsumura, 1935)

Unkana sapporona Matsumura, 1935.

Unkanodes sapporona (Matsumura, 1935), comb. by Fennah 1956: 474.

Unkanodes (Unkanodes) tanasijevici (Dlabola, 1965)

Elymodelphax tanasijevici Dlabola, 1965.

Calligypona zeravshanica Dubovsky, 1967; syn. by Emeljanov 1982: 98.

Ribautodelphax notabilis Logvinenko, 1970

Subgenus Unkanodes (Kwonianella Anufriev, 1988)

Unkanodes (Kwonianella) albifascia (Matsumura, 1900: 268)

Liburnia albifascia Matsumura, 1900: 268.

Delphax albifascia (Matsumura, 1900); comb. by Oshanin, 1907: 330.

Delphacodes albifascia (Matsumura, 1900); comb. by Metcalf 1943: 400.

Unkanodes (Chilodelphax) albifascia (Matsumura, 1900); comb. by Vilbaste 1968: 26.

Chilodelphax albifascia (Matsumura, 1900); status by Kwon 1982: 4.

Unkanodes (Kwonianella) albifascia (Matsumura, 1900); comb. by Anufriev and Emeljanov 1988: 409.

Unkanodes (Kwonianella) insularis Anufriev, 1988

Unkanodes (Kwonianella) insularis Anufriev & Emeljanov, 1988: 409.

Unkanodes (Kwonianella) sympaticus Anufriev, 1988

Unkanodes (Kwonianella) sympaticus Anufriev & Emeljanov, 1988: 409.

Key to subgenera and species of Unkanodes of the world

This key is modified from Anufriev and Emeljanov (1988). Bartlett and contributors (2017) treated Ribautodelphax notabilis Logvinenko, 1970 as a synonym of Unkanodes tanasijevici (Dlabola, 1965) based on Nast (1987). In the present key, characters mentioned for U. tanasijevici are from the description of Ding (2006). Unkanodes (Unkanodes) paramarginata is not included in the key due to limited literature.

1 Armature of diaphragm with a pair of teeth slanting upwards or directed back; genital style with relatively wide apices; segment X (anal tube) with large widely-spaced processes; sub genus Unkanodes (Unkanodes) 2
Armature of diaphragm without teeth or with a tooth bifurcate at apex; genital style with narrow apices; segment X (anal tube) with or without such processes 5
2 Process of anal tube spaced more widely, weakly or strongly diverging; genital style with strong subapical lobe, apex wider 3
Processes of anal tube spaced less widely, more or less parallel; genital style with weak subapical lobe, apex narrower (Anufriev and Emeljanov 1988: fig. 310: 1–12) U. (Unkanodes) excisus
3 Process of segment X widely spaced weakly diverging; apex of genital style relatively wider (Anufriev and Emeljanov 1988: fig. 310: 13–17) U. (Unkanodes) sapporona
Process of segment X widely-spaced strongly diverging; apex of genital style comparatively less wider 4
4 Aedeagus elbow-shaped; process of segment X posteroventrally curved (Figs 6–11) U. (Unkanodes) latespinosa
Aedeagus straight with a strong tooth on dorsal aspect, ventrally with a weak lobe below the tooth; segment X not curved (Ding 2006: fig. 338A–L) U. (Unkanodes) tanasijevici
5 Dorsal and posterior margin of pygofer forming an obtuse angle in lateral view; segment X (anal tube) with large widely-spaced teeth; apical half of aedeagus straight; subgenus Chilodelphax (Anufriev and Emeljanov 1988: fig. 311: 1–15) U. (Chilodelphax) silvaticus
Dorsal and posterior margin of pygofer forming an acute angle in lateral view; segment X (anal tube) without or with narrowly spaced teeth; apical half of aedeagus directed dorsad (Anufriev and Emeljanov 1988); subgenus Kwonianella 6
6 Process of pygofer bridge very short, directed downwards, sometimes bifurcate at apex; genital style comparatively short with wide subapical lobe 7
Process of pygofer bridge slightly long, bifurcated and directed backwards; genital style longer with narrow subapical lobe or short with wide subapical lobe 8
7 Aedeagus near bent with a pair of long teeth perpendicular to shaft, the length of aedeagus matches with thickness of shaft (Anufriev and Emeljanov 1988: fig. 312: 17–19) U. (Kwonianella) sympatricus
Aedeagus near bent without long teeth, the length of which matches with thickness of the shaft (Anufriev and Emeljanov 1988: fig. 312: 1–11) U. (Kwonianella) albifascia
8 Aedeagus narrowing abruptly in apical 1/3, with a lobe-like process on ventral aspect in lateral view (Fig. 21) U. (Kwonianella) malamjabbensis sp. nov.
Aedeagus not narrowing abruptly in apical 1/3, without a lobe on ventral aspect in lateral view (Anufriev and Emeljanov 1988: fig. 312: 12–16) U. (Kwonianella) insularis

Unkanodes (Unkanodes) latespinosa (Dlabola, 1957)

Figs 1–11


Dlabola (1957) described this species based on specimens from Afghanistan and provides a detailed description. It can be distinguished from other species of Unkanodes by the large, widely-spaced processes of segment X (anal tube) and elbow-shaped aedeagus.

Material examined

3♂♂ (brachypterous), 7♂♂ (macropterous) Malamjabbah, Swat-Khyber Pakhtunkhwa, Pakistan, 35°13'21.76"N, 72°25'32.93"E, 2993.39 m, 5 vii 2018, sweeping grasses, coll. Kamran Sohail. The area has a very diverse habitat for fruits and vegetables, and this species was collected in grasses near vegetable fields. This species is newly recorded for the fauna of Pakistan.


Previously recorded from Afghanistan, Iran, Mongolia, Turkey and Yugoslavia. In this study it is recorded from Swat, Khyber Pakhtunkhwa-Pakistan.

Figures 1–11. 

Unkanodes (Chilodelphax) latespinosa (from Pakistan) 1 adult, dorsal view (macropterous) 2 adult, same species (brachypterous) 3 adult (macropterous), lateral view 4 vertex, pronotum and mesonotum, dorsal view 5 frons, ventral view 6, 7 male genitalia, caudal and lateral views 8–10 anal segment and aedeagus, lateral, dorsal and ventral views 11 genital style, lateral view.

Unkanodes (Kwonianella) malamjabbensis sp. nov.


Length of male (n=2) 1.4–1.6 mm.


General body colour dark brown to black. Vertex pale, compartments with three distinct yellow spots. Carina on frons pale, intercarinal region dark brown, gena concolourous with intercarinal region, compound eyes greyish. Antenna yellowish slightly darker at junction of scape and pedicel. Pronotum and mesonotum medially with a white stripe; darker at adjoining areas, extreme lateral margins and median carina white, lateral carina concolourous with adjoining regions. Forewings dark brown to black, apical and anal margins pale. Legs yellowish, spines with black apices. Abdominal tergites darker, segments IX and X lighter, pygofer brown.


Head narrower than pronotum, eyes extending beyond posterior margin of vertex (Figs 12, 14). Vertex ca. 2X longer than wide; stem of Y-shaped carina of vertex obsolete, lateral and posterior margins distinct, arms of submedian carina meeting at fastigium (Fig. 14). Frons parallel-sided; widest near basal 1/4 of eyes, narrower in apical 1/3, median carina bifurcate near fastigium (Fig. 15). Antennal scape about as long as wide, ca 1/2 x length of pedicel, pedicel bearing many sensory pits arranged in longitudinal rows dorsally from base to apex (Figs 13, 15). Frontoclypeal suture distinct, slightly arched; median carina on postclypeus visible, rostrum elongate, reaching hind coxae (Fig. 15). Pronotum much wider than long at midlength; lateral carinae strongly diverging, vanishing before reaching posterior margin and not in line with mesonotal lateral carinae, anterior margin straight at vertex, posterior margin slightly concave medially (Fig. 14). Mesonotum tricarinate, subequal to length of pronotum; median carina not extending to apex of scutellum, lateral carinae slightly diverging reaching hind margin, tegula inconspicuous (Fig. 14). Forewing covering only half of abdomen; veins granulate (Figs 12, 16). Metatibiae with two lateral spines on shaft, first near tibiofemoral articulation, second after middle. Metatibial spur tectiform, distally narrowed bearing row of 18 black-tipped teeth on outer margin, inner margin straight (Fig. 17). Spinal formula of hind leg 5/7/4.

Figures 12–17. 

Unkanodes (Kwonianella) malamjabbensis sp. nov. 12, 13 adult, dorsal and lateral views 14 head and thorax, dorsal view 15 frons, ventral view 16 forewing 17 metatibial spur.

Male genitalia

In caudal view, pygofer wider than long widest at mid length, dorsolaterally nearly straight (Figs 18, 20); diaphragm armature well-developed, V-shaped, pair of lobes located near the parameres directed upwards, pygofer bridge bearings two distinct tooth-like processes widely diverging, directed backwards forming blunt apex (Fig. 20). In lateral view, anterior margin nearly straight gradually arched apically, dorsal and posterior margins acutely rounded (Fig. 19). Segment X (anal tube) bearing large, widely spaced posteroventrally curved acute processes (Figs 18, 19). Parameres longer than wide, apically narrow, subapical lobe wider, posterior margin straight (Fig. 24). Aedeagus elongate and narrow, basal 1/3 straight, bent gradually forming an obtuse angle, apical 1/3 gradually curved ventrad (Fig. 21); in lateral view, with lobe or hump-like process on ventral aspect, with a large tooth just above the lobe on the dorsal aspect (Fig. 21). Suspensorium angling circled laterally, apically wider (Figs 22, 23).

Figures 18–25. 

Unkanodes (Kwonianella) malamjabbensis sp. nov. 18, 19 male genitalia, caudal and lateral view 20 pygofer, caudal view 21 aedeagus, lateral view 22, 23 same, dorsal and ventral views 24 genital style, lateral view 25 medioventral process of armature of diaphragm.

Type materials

Holotype: ♂ Malamjabbah, Swat-Khyber Pakhtunkhwa, Pakistan, 35°13'21.76"N, 72°25'32.93"E, 2993.39 m, 5 vii 2018, sweeping grasses, coll. Kamran Sohail. Paratype: 1♂, same data as holotype.


This new species was collected in a grass habitat near ponds. The Type locality is an understudied habitat for fulgoroids and the region reflects a true diversity of planthoppers for future prospects.




The new species is named after the type locality ‘Malamjabba’.


The new species is externally similar to U. (Kwonianella) albifascia which also has a white stripe on the thorax and median margins of the forewings. However, it can be separated by the distinctly separated process of the pygofer bridge, widely diverging in U. (Kwonianella) malamjabbensis sp. nov. but very short and bifurcate at the apex in U. (Kwonianella) albifascia (Anufriev and Emeljanov 1988, Figs 2, 6; p. 412); and apical half of aedeagus gradually curved ventrad bearing a lobe-like process on the ventral aspect in U. (Kwonianella) malamjabbensis sp. nov. versus the apical half of the aedeagus slanting dorsad without a lobe in U. (Kwonianella) albifascia (Anufriev and Emeljanov 1988, Fig. 4; pp. 409, 412). Unkanodes (Kwonianella) malamjabbensis sp. nov. is also close to U. (Kwonianella) insularis Anufriev and U. (Kwonianella) sympatricus Anufriev in external appearance but can be distinguished by the distinct shapes of the aedeagus and parameres.

Figure 26. 

Distribution map of species of Unkanodes in Pakistan.


We are sincerely thankful to Stephen W. Wilson (Professor Emeritus of Biology at University of Central Missouri Warrensburg, USA) for revising and providing comments on an early draft of this manuscript. We are also thankful to Prof. John Richard SCHROCK (Emporia State University, USA) for English revision of this manuscript and Dr. Tehseen Javed (College of Water Resources and Architecture Engineering, NWAFU) for providing the distributional map. We also extend our thanks to anonymous reviewers and the editor for their valuable suggestions and comments. This study was supported by the National Natural Science Foundation of China (31420103911) and The Ministry of Science and Technology of the People’s Republic of China (2005DKA21402).


  • Anufriev GA, Emeljanov AF (1988) Volume II: Homoptera and Heteroptera. In: P.A. Lehr (ed.). Keys to the Insects of the Far East of the USSR in Six Volumes, Transliteration of the Russian title: Opredelitel’ nasekomykh Dal’nego Vostoka SSSR v shesti tomakh (Vol. 2). Ravnokrylye i poluzhestkokrylye. In: Lehr PA (Ed.) Keys to the Insects of the Far East of the USSR in Six Volumes. Nauka, Leningrad.
  • Asche M (1985) Zur Phylogenie der Delphacidae Leach, 1815 (Homoptera: Cicadina: Fulgoromorpha). Marburger Entomologische Publikationen 2(1), Vols. 1–2: 1–398. [399–910.]
  • Bartlett CR (2014) New species of the planthopper genus Parkana (Hemiptera: Fulgoroidea: Delphacidae) from Mesoamerica. Transactions of the American Entomological Society 140: 185–208.
  • Bartlett CR, O’Brien LB, Wilson SW (2014) A review of the planthoppers (Hemiptera: Fulgoroidea) of the United States. Memoirs of the American Entomological Society 50: 1–287.
  • Bourgoin T (2020) FLOW (Fulgoromorpha Lists On the Web): a world knowledge base dedicated to Fulgoromorpha. Version 8, updated 16th January 2020. [accessed 21st January 2020]
  • Ding JH (2006) Fauna Sinica Insecta (Vol. 45). Homoptera Delphacidae. Science Press, Beijing.
  • Dlabola J (1957) Die Zikaden Afghanistans (Homoptera Auchenorrhyncha) nach der Ergebnisse der von Herrn J. Klapperich in den Jahren 1952–1953 nach Afghanistan untergenommenen Expedition. Mitteilungen der Münchner Entomologischen Gesellschaft 47: 265–303.
  • Dlabola J (1961) Die Zikaden von Zentralasien, Dagestan und Transkaukasien (Homopt. Auchenorrhyncha). Acta Faunistica Entomologica Musei Nationalis Pragae 34(587): 241–358.
  • Dlabola J (1964) Die Zikaden Afghanistans (Homoptera Auchenorrhyncha) II Teil. Ergebnisse der Sammelreisen von Dr. H. G. Amsel, G. Ebert, Dr. Erichson, J. Klapperich und Dr. K. Lindberg. Mitteilungen der Münchner Entomologischen Gesellschaf 54: 237–255.
  • Dlabola J (1965) Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei. 54 Homoptera-Auchenorrhyncha. Acta Faunistica Entomologica Musei Nationalis Pragae 11(100): 79–136.
  • Dlabola J (1967) Ergebnisse der 1. mongolisch-tschechoslowakischen entomologisch-botanischen Expedition in der Mongolei. Nr. 3: Homoptera, Auchenorrhyncha (Erganzung). Acta Faunistica Entomologica Musei Nationalis Pragae 12(118): 51–102.
  • Dubovsky GK (1967) Novye vidy cikadovykh (Auchenorrhyncha) iz Uzbekistana in Poleznye i vrednye bezpozvonochnye zhivotnye Uzbekistana, Tashkent, 56–59. [Calligypona zeravshanica Dubovsky, 1967]
  • Emeljanov AF (1977) Leaf-hoppers (Homoptera, Auchenorrhyncha) from the Mongolian People’s Republic based mainly on materials of the Soviet-Mongolian zoological expeditions (1967–1969). Nasekomye Mongolii [Insects of Mongolia] 5: 96–195. [In Russian]
  • Emeljanov AF (1982) Fulgoroidea (Homoptera) collected in the Mongolian People’s Republic by the entomofaunistical group of the Soviet Mongolian complex biological expedition in 1970–1975. Nasekomye Mongolii [Insects of Mongolia] 8: 69–122. [In Russian]
  • Fennah RG (1956) Fulgoroidea from southern China. Proceedings of the California Academy of Sciences. San Francisco 28(4): 441–527. [474]
  • Jensen-Haarup AC (1915) Danmarks Cikader. Flora och Fauna 1915: 137–144. [Liburnia elymi Jensen-Haarup, 1915]
  • Jensen-Haarup AC (1920) Cikader. In: Danmarks Fauna Illustrerede Haandbøger over den Danske dyreverden med Statsunderstøttelse udgivne af Dansk Naturhistorisk Forening. G. E. C. Gads, København, 189 pp. [Liburnia elymi Jensen-Haarup, 1915]
  • Kwon YJ (1982) New and little known planthoppers of the family Delphacidae (Homoptera: Auchenorrhyncha). Korean Journal of Entomology 12(1): 1–11.
  • Leach WE (1815) Entomology. The Edinburgh Encyclopedia 9: 57–172. [125]
  • Logvinenko VN (1970) New and little-known leafhoppers of the family Delphacidae (Homoptera, Auchenorrhyncha) from Southern regions of the USSR. Entomologicheskoye Oboszrenie 49(3): 624–633. [in Russian; Ribautodelphax notabilis Logvinenko, 1970]
  • Matsumura S (1900) Uebersicht der Fulgoriden Japans. Entomologische Nachrichten 26: 257–269.
  • Matsumura S (1935) Revision of Stenocranus Fieb. (Hom.) and its allied species in Japan Empire. Insecta Matsumurana 9: 125–140.
  • Metcalf ZP (1943) General Catalogue of the Hemiptera. Fascicle IV, Fulgoroidea, Part 3, Araeopidae (Delphacidae). Smith College, Northampton, Massachusetts.
  • Nast J (1987) The Auchenorrhyncha (Homoptera) of Europe. Annales Zoologici 40(15): 535–661. [Ribautodelphax notabilis Logvinenko, 1970]
  • Oshanin VT (1907) Verzeichnis der palaearktischen hemipteren, mit besonderer berücksichtigung ihrer verteilung im Russischen reiche. II. Band. Homoptera. I. Lieferung. Annuaire du Musée Zoologique de l’Académie Impériale des Sciences de St. Pétersbourg 12: 193–384.
  • Puton A (1899) Homoptera. Am. Serv. (Gulaerostria Zett. Fieb.) Sect. I. Auchenorrhyncha Dumér. (Cicadina Burm.). Catalogue des Hémiptères (Hétéroptères, Cicadines et Psyllides) de la faune Paléarctique. 4e Ed. Bureau de la Société Française d’Entomologie, Caen, 3–121.
  • Vilbaste J (1968) On the Cicadine fauna of the Primosk region. Valgus Publishing, Tallinn, 195 pp. [In Russian, see p. 26]
  • Wagner W (1963) Dynamische Taxionomie, angewandt auf die Delphaciden Mitteleuropas. Mitteilungen des Hamburger Zoologischen Museums und Instituts 60: 111–180.
  • Wilson SW, McPherson JM (1980) Keys to the planthoppers, or Fulgoroidea of Illinois (Homoptera). Transactions of the Illinois State Academy of Science 73(2): 1–61.
  • Wilson SW, Mitter C, Denno RF, Wilson MR (1994) Evolutionary patterns of host plant use by delphacid planthoppers and their relatives. In: Denno RF, Perfect TJ (Eds) Planthoppers: their ecology and management. Chapman Hall, New York, 7–45.
  • Wilson SW, O’Brien LB (1987) A survey of planthopper pests of economically important plants (Homoptera: Fulgoroidea). In: Wilson MR, Nault LR (Eds) Proceedings of the 2nd International Workshop on Leafhoppers and Planthoppers of Economic Importance. CAB International Institute of Entomology, London, 343–360.