Genus Aphelonema Uhler, 1876

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[Back to “North American Caliscelidae“]

Family Caliscelidae Amyot & Serville, 1843

Subfamily Caliscelinae Amyot & Serville, 1843

Tribe Peltonotellini Emeljanov, 2008

Genus Aphelonema Uhler, 1876

Type species: Aphelonema simplex Uhler, 1876.


 = Peltonotus Mulsant & Rey 1855 (preoccupied; nec. Burmeister 1847) (Type species Peltonotus raniformis Mulsant & Rey 1855).
Peltonotellus Puton 1886; replacement name for unavailable Peltonotus Mulsant & Rey 1855 (nec. Burmeister 1847); syn. by Melichar 1906: 36, 53; removed from syn. and redefined (subgenus) by Emeljanov 1996: 994; revised status (to genus) by Emeljanov (2008).
Mushya Kato, 1933; syn. by Chan & Yang 1994: 8.


Widespread in US, Canada, Central America and Palearctic.

Distribution of Aphelonema

Distribution of Aphelonema (including Protrocha, and unplaced taxa; excluding other subgenera) from FLOW (as of 6 June 2017); subgenus Aphelonema is New World.

Recognized species

By my count, 32 valid species, plus subspecies.  The following list may need to be updated (esp. viz. Emeljanov 2008)

Notes: Emeljanov (1996) revised the taxonomy of the subgenera; these subgenera have been inconsistently (and without explanation) used as full genera, but are retained as subgenera here (despite e.g., Emeljanov 2008). The North American taxa are – I think – inadequately diagnosed at this time to consider as full genera, and until that happens I will retain them as subgenera.

Distribution information for Palearctic species is incomplete (based largely on Metcalf 1958).

Subgenus Aphelonema Uhler, 1876

Aphelonema decorata (Van Duzee, 1908) – USA: MS, NJ, SC, FL
Aphelonema simplex Uhler, 1876 – USA: AL, CO, CT, DE, FL, IA, IL, KS, MD, MI, MO, MS, NC, NE, NJ, OH, SD, TX, VA; CAN: MB, ON, SK
Aphelonema rosa Metcalf, 1923: 188; syn. by Doering 1941: 190.
   Aphelonema simplex var. dorsata Ball, 1926 – USA: FL

Subgenus Nenema Emeljanov 1996 (Type species Peltonotellus bivittatus Ball 1902b)

Aphelonema bivittata (Ball, 1902) – USA: AZ, CA, CO, IA, KS, ND, NE, NM, OK, TX; CAN: AB, SK; Mexico (Chihuahua, Coahuila, Federal District)
Peltonotellus bivittatus Ball 1902: 263; also Maw et al. 2000: 82.
Aphelonema bivittata (Ball, 1902); comb. by Van Duzee 1916a: 80.
Aphelonema concinna Doering, 1941 – USA: NV
Aphelonema confragosa Doering, 1941 – USA: AZ, NM, NV, TX
Aphelonema convergens Bunn, 1930 – USA: AZ, CO, NM
   Aphelonema convergens var. canyonensis Bunn, 1930 – USA: AZ
Aphelonema histrionica (Stål, 1862) – USA: AZ, CA, CO, IA, IL, MA, MD, MI, MN, MO, MT, NH, NM, NV, NY, OR, PA, WI; CAN: AB, BC, MB, NB, NF, NS, ON, PE, QC, SK, YK
Peltonotus histrionicus Stål, 1862: 310.
= Peltonotellus histrionicus (Stål, 1862); comb. by Ball 1902b: 263; also Maw et al. 2000:82.
Aphelonema histrionica (Stål, 1862); comb. by Van Duzee 1916a: 80.
Aphelonema histrionica var. giffardi Van Duzee, 1917a – USA: CA, CO
Aphelonema rugosa (Ball, 1902) – USA: AZ, CA, CO, IA, ID, KS, MN, MT, NE. NM, OK, OR, SD, TX, UT, WA, WY; CAN: AB, SK (MD record possibly in error); Belize
Peltonotellus rugosus Ball, 1902: 263; Maw et al. 2000: 82.
Aphelonema rugosa (Ball, 1902); comb. by Kirkaldy 1913: 25-26.
10 Aphelonema virgata Doering, 1941 – USA: AZ, NM

Subgenus Protrocha Emeljanov 1996 (Type species Aphelonema orbiculata Ball, 1935).

11 Aphelonema brevata Caldwell, 1945 – Mexico (Puebla, Veracruz)
12 Aphelonema impercepta Doering, 1941 – USA: AZ
13 Aphelonema minuta Bunn, 1930 – USA: AZ
14 Aphelonema nesolitaria Caldwell, 1945 – Mexico (Federal District)
15 Aphelonema nigriviridia Ball, 1926 – USA: FL
16 Aphelonema obscura Van Duzee, 1912 – USA: FL, GA, IA, KS, MS
17 Aphelonema orbiculata Ball, 1935 – USA: NM, TX; Mexico (Federal District, Mexico, Puebla, Veracruz)
18 Aphelonema robusta Caldwell, 1945 – Mexico (Michoacan)
19 Aphelonema rubranota Caldwell, 1945 – Mexico (Puebla)
20 Aphelonema solitaria Ball, 1935a – USA: AZ
21 Aphelonema viridis Dozier, 1928 – USA: MS

Subgenus (or genus) Ceragra Emeljanov 1996 (Type species Aphelonema oxyura Kusnezov, 1930).
22 Aphelonema brunneolutea Dlabola, 1993 – Iran
23 Aphelonema deserticola Emeljanov, 1964 – Kazakhstan
24 Aphelonema oxyura Kusnezov, 1930 – Uzbekistan.

Subgenus (or genus) Acromega Emeljanov 1996 (Type species Peltonotus scurrilis Stål, 1862)
25 Aphelonema scurrilis (Stål, 1862) – Mongolia, Russia (Siberia)

Subgenus (or genus) Peltonotellus Puton, 1886 (Type species Peltonotellus raniformis Mulsant & Rey, 1855)
26 Aphelonema ellisi Dlabola, 1974 – Corsica, France
27 Aphelonema eoa Kusnezov, 1930 – Kyrgyzstan
Aphelonema zonata Dlabola, 1961, syn. by Dubovsky 1966, but see Emeljanov 2008. [Uzbekistan].
28 Aphelonema fasciata Chan & Yang, 1994 – Taiwan, Japan
Mushya quadrivittata Kato, 1933; syn. Chan & Yang, 1994
29 Aphelonema melichari Horváth, 1897 – Austria, Italy, Yugoslavia
30 Aphelonema punctifrons (Horváth, 1895) – Crimea , Hungary, Russia (South European Russia), Serbia, Yugoslavia, Urkraine
31 Aphelonema quadrivittata (Fieber, 1876) – Austria, Hungary, Italy, Yugoslavia, Germany
32 Aphelonema raniformis (Mulsant & Rey, 1855) – France, Italy.
33 Peltonotellus turgidus Emeljanov, 2008 – Kazakhstan
34 Peltonotellus niger Meng, Gnezdilov & Wang, 2015 – China North-Central (Gansu)
35 Peltonotellus brevis Meng, Gnezdilov & Wang, 2015 – China [Inner Mongolia (Ningxia)]
36 Peltonotellus labrosus Emeljanov, 2008 – China (Qinghai)
37 Peltonotellus pelidnus Emeljanov, 2008 – Kazakhstan
38 Peltonotellus mitjaevi Emeljanov, 2008 – Kyrgyzstan
39 Peltonotellus diffusus Emeljanov, 2008 – Kyrgyzstan, Tadzhikistan

Distribution of Peltonotellus from Flow

Distribution of Peltonotellus from FLOW (as of June 6 2017)

Peltonotellus is most frequently treated as a separate genus; Meng et al. 2015 provides an emended diagnosis.

Not placed / not sure
39 Aphelonema vespertina Kirkaldy, 1913: 25 – Mexico (Federal District)
40 Aphelonema registanica (Dlabola, 1961) – Iran

Economic Importance


Plant associations

Aphelonema decorata – sedgesSpartina alterniflora Loisel. (cordgrass, Poaceae)
Aphelonema scurrilis – Carex duriuscula C.A. Mey. (needleleaf sedge, Cyperaceae)
Aphelonema simplex – Spartina patens (Aiton) Muhl. (cordgrass, Poaceae)
Aphelonema decorata  – Spartina patens (Aiton) Muhl. (cordgrass, Poaceae), Deschampsia flexuosa (L.) Trin. (Poaceae), Panicum repens L. (torpedo grass) (all from specimen label data)
Aphelonema rugosa – Agropyron cristatum (L.) Gaertn. (crested wheatgrass, Poaceae), Bromus L. (Brome grass, Poaceae) (both specimen label data)
Hosts from Wilson et al. 1994 or specimen label data; plant names from USDA PLANTS or Tropicos.


Head not weevil-like, legs not modified (i.e., not expanded as in Fitchiella), vertex broader than long (vs. Papagona)

Key to subgenera (rearranged from Emeljanov 1996, comments in brackets; the key was not illustrated or otherwise explained so the meaning of some features remains obscure to me).

1- lateral parts of pronotum with only one foveola [i.e., sensory pit, see illustration below from Doering 1941] … 4
1’-  lateral parts of pronotum with no less than 2, usually 3 or more foveolae … 2

Aphelonema lateral view of head

Aphelonema lateral view of head from Doering 1941

2-  intermediate carinae of metopa [~frons] gently convex outside, inner parts forming longitudinal elongate dolioform [barrel-like] figure … 3
2’- Intermediate carinae of metopa strongly convex outside, inner parts forming a rather irregular, slightly transverse oval … s.g. Acromega Emeljanov 1996 [not found in New World]

3- Metopa with lower foveola of inner row strongly shifted mediad … s.g. Ceragra Emeljanov, 1996 [not found in New World]
3’- Metopa with lower foveola of inner row not shifted, lying along same line as others … s.g. Nenema Emeljanov 1996

4- Abdomen with lateral group of only 2 foveolae in oblique row, and median foveola group of only 1 row; Lateral lobes of metopa in upper half with 2 parallel rows of foveolae … s.g. Aphelonema
4’- Abdomen with lateral group of 3 (forming triangle) or more foveolae. Median group having additional row or individual foveolae coming out of row; Lateral lobes of metopa with additional 3rd row of foveolae … 5

5- Abdomen with lateral group of more than 3 foveolae. Metopal intermediate carinae separately running into clypeal suture … s.g. Peltonotellus Puton
5’- Abdomen with lateral group of only 3 foveolae. Metopal intermediate carinae more or less merging before clypeal region … s.g. Protrocha Emeljanov, 1996

Key to species of Aphelonema north of Mexico transcribed (and slightly modified) from Doering (1941) (there may be OCR errors I missed)

1. Vertex extremely broad, width at least five or six times median length; frons greatly exposed above, roundingly protruded as viewed from the side. … 2
1’. Vertex much longer, width between eyes two to three times median length; frons through middle not exposed from above, as viewed from the side either at right angles to body or distinctly receded … 4

2. (1) Small species, length of female not over 2.55 mm, male, 1.76 mm; central tablet of frons heart-shaped approximately one-fourth longer than greatest width …Aphemonema obscura Van Duzee.
2’. Larger species, length of female, 3.00 to 3.75 mm; males, 2.5 to 2.7 mm; central tablet of frons subequal in width and length … 3

3. (2) Head and thorax orange-tan, rest of dorsum blackish brown; central frontal tablet pointed below … Aphemonema decorata (Van Duzee).
3’. Uniformly pale ochreous color; central frontal tablet almost circular… Aphemonema simplex Uhler.

4. (1) As viewed from the side head distinctly conically produced in front of eyes; frons conspicuously receded; vertex somewhat triangular, anterior margin narrowed … 5
4’. Head not conically produced in front of eyes, sometimes shortened instead; frons more or less vertical; vertex with three anterior margins subequal … 11

5. (4) Vertex triangular, median anterior margin less than length of lateral margin against eye; median frontal tablet longer than wide … 6
5.’ Vertex not so narrowed anteriorly, median anterior margin wider than lateral margin; median frontal tablet wider than long, except Aphemonema histrionica … 8

6. (5) Elongate, length of female, 3.4 mm; length of male, 2.7 mm; median frontal tablet smooth, oval, no angles indicated at either end, two pustules between tablet and postclypeus, an extra median row of 5 pustules in each lateral compartment … Aphemonema orbiculata Ball.
6’. Extremely small species, length of female under 2.5 mm; of male under 1.9 mm; median frontal tablet truncate at ends, only one pustule between tablet and postclypeus, the usual two rows only of pustules in each lateral compartment … 7

7. (6) Body slender, sides of abdomen subparallel; base of median frontal tablet roundingly narrowed, carinae continuous with no basal margin indicated….Aphelonema impercepta
7’. Body stouter, abdomen extended at sides of tegmina; base of frontal tablet straight, about one-half the width through middle… Aphelonema minuta Bunn.

8. (5) Median frontal tablet elongate, apical pustules in lateral compartment of frons, and apices of lateral carinae widely separated; tegmina and body with conspicuous yellow and black stripes (Northern and Eastern) … Aphelonema histrionica
8’. Median frontal tablet wider than long; lateral frontal carinae joined or closely approaching each other; pale species with occasional dark markings … 9

9. (8) Vertex longer than median length of pronotum; frontal tablet twice as wide as long, lateral carinae meeting well before apex, lateral pustules evenly spaced, forming a complete circlet across face (Eastern) … Aphelonema nigriviridia
9’. Vertex subequal to median length of pronotum, not extended; frontal tablet not over one-half wider than length, lateral pustules not evenly spaced … 10

10. (9) Vertex about twice wider than median length; pale, robust species with a conspicuous round black spot at apex of abdomen (Eastern) … Aphelonema viridis
10.’ Vertex less than twice wider than long; longer species; a median dark stripe and lesser dark spot at apex of abdomen (Southwestern) … Aphelonema solitaria

11. (4) Pale green and yellow, no dark markings present … Aphelonema concinna
11’. Body and tegmina stramineous, spotted or striped with dark markings … 12

12. (11) Tegmina reticulate, veins conspicuously elevated; basal portion of postclypeus overhanging rest as a blunt tubercle (less pronounced in male). …Aphelonema rugosa.
12’. Tegmina not reticulate, only 1 or 2 veins visible; no tubercle on postclypeus … 13

13. (12) A conspicuous elevated vein on clavus; costal area only of tegmen dark … 14
13’. No claval vein indicated; tegmina and abdomen with a broad conspicuous black stripe on each side of median yellow stripe … 15

14. (13) Tegmina obliquely depressed through middle; black areas of abdominal segments inflated; claval vein light brown … Aphelonema confragosa
14’. Tegmina not depressed; abdominal segments smooth; claval vein outlined conspicuously in black … Aphelonema convergens

15. (13) Five anterior sides of vertex equal; mesal black stripes of abdomen not much wider than those of tegmina … Aphelonema bivittata
15’. Median anterior margin of vertex longer than either the angulate or eye margin, mesal abdominal black bands at one-third wider than those of tegmina. …Aphelonema virgata

Aphelonema decorata and histronica

Aphelonema (Aphelonema) decorata and Aphelonema (Nenema) histronica

Aphelonema (Nenema) rugosa

Aphelonema (Nenema) rugosa

Aphelonema histrionica

Aphelonema (Nenema) histrionica (Salvador Vitanza; Arivaca, Pima County, Arizona, February 16, 2020, 3,655 ft, sweeping roadside vegetation; used by permission)

Aphelonema histrionica

Aphelonema (Nenema) histrionica (Salvador Vitanza; Arivaca, Pima County, Arizona, February 16, 2020, 3,655 ft, sweeping roadside vegetation; used by permission)

Aphelonema histrionica

Aphelonema (Nenema) histrionica (Salvador Vitanza; Arivaca, Pima County, Arizona, February 16, 2020, 3,655 ft, sweeping roadside vegetation; used by permission)

Aphelonema (Aphelonema) simplex

Aphelonema (Aphelonema) simplex

Aphelonema species dorsal view from Doering 1941

Aphelonema species dorsal view from Doering 1941


Pages from Doering (1941)

Aphelonema species head from Doering 1941


Most often taken sweeping grasses. Aphelonema simplex and A. decorata are found in coastal marshes and can be locally abundant.

Molecular resources: 

At this writing, data for A. histrionica and 2 undet. Aphelonema appears on Genbank; there is data for Aphelonema simplex and A. decorata on Barcode of Life.

Selected references:

Amyot, C. J. B. and J. G. A. Serville. 1843a. Deuxieme partie. Homopteres. Homoptera Latr. Histoire Naturelle des Insectes. Hemipteres. lxxvi + 676 pp.  [plates are here] [from Google books]

Ball, E. D. 1902. New genera and species of North American Fulgoridae. Canadian Entomologist 34: 259-266.

Ball, E. D. 1935. Some new Issidae with notes on others (Homoptera: Fulgoridae). Bulletin of the Brooklyn Entomological Society 30: 37-41.

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.

Bunn, R. W. 1930a. Notes on the genus Aphelonema Uhler with descriptions of new species. Journal of the Kansas Entomological Society 3(3): 73-77.

Caldwell, J. S. 1945. Notes on Issidae from Mexico (Homoptera, Fulgoroidea). Annals of the Entomological Society of America 38: 89-120.

Chan, M. L. and Chung-Tu Yang. 1994. Issidae of Taiwan: (Homoptera: Fulgoroidea). National Chung Hsing University, Department of Entomology, Taichung, China (Taiwan).  168 pp.

Dlabola, J. 1961. Die Zikaden von Zentralasien, Dagestam und Trankav Kasien (Homoptera: Auchenorrhyncha). Acta Entomologica Musei Nationalis Pragae 34: 241-358.

Dlabola, J. 1974. Zur Taxonomia und Chorologie einiger Mediterraner Zikaden (Homoptera Auchenorrhyncha). Acta Zoologica Academiae Scientiarum Hungaricae 20(3-4): 289-308

Dlabola, J. 1994 [1993]. Erganzungen zur Iranischen, Israelischen und Benachbarten Zikadenfaunen mit Beschreibungen 30 neuer Taxone (Homoptera, Auchenorrhyncha). Sbornik Narodniho Muzea V Praze Rada B Prirodni Vedy 49(1-4): 41-110. [German with Czech summary]

Doering, K. C. 1939 [1940a]. A contribution to the taxonomy of the subfamily Issinae in America north of Mexico (Fulgoroidea: Homoptera. Part III. University of Kansas Science Bulletin. 26(2): 83-167.

Doering, K. C. 1941. A contribution to the taxonomy of the subfamily Issinae in America north of Mexico (Fulgoroidea: Homoptera. Part IV. University of Kansas Science Bulletin 27(10): 185-233.

Dozier, H. L. 1928a [dated 1922 or 1926]. The Fulgoridae or planthoppers of Mississippi, including those of possible occurrence. Technical Bulletin of the Mississippi Agricultural Experiment Station 14: 1-152.

Dubovskii, G. K. 1966. Tsikadovye (Anchenorrhyncha) Ferganskoi doliny. [Cicadinea (Auchenorrhyncha) of the Fergana Valley]. Uzbek “Fan” Press, Tashkent. 156 pp.

Emeljanov, A. F. 1964. New Cicadina from Kazakhstan (Homoptera, Auchenorrhyncha). [New species of insects of the fauna of Kazakhstan.] Trudy Zoologicheskogo Instituta AN SSSR 34: 3-51. [Russian]

Emeljanov, A. F. 1996. A new genus, Chirodisca gen. n., and new subgenera of the genus Aphelonema Uhl. (Homoptera, Fulgoroidea, Issidae). Entomological Review 76(8): 993-994.

Emeljanov, A. F. 1999. Notes on delimitation of families of the Issidae group with description of a new species of Caliscelidae belonging to a new genus and tribe (Homoptera, Fulgoroidea). Zoosystematica Rossica 8(1): 61-72.

Emeljanov, A. F. 2008. New species of the genus Peltonotellus Puton (Homoptera, Caliscelidae) from Kazakhstan, Middle and Central Asia. Tethys Entomological Research 16: 5-12.

Fitch, A. 1856. Third report on noxious and other insects of the State of New York. Transactions of the New-York State Agricultural Society 16: 315-490.

Guglielmino, A. and C. Bückle. 2007. Contribution to the knowledge on the Auchenorrhyncha fauna of the Tuscanian-Emilian Apennines. Bulletin of Insectology 61(1): 151-152. [record of Peltonotellus quadrivittatus]

Holzinger, W.E., I. Kammerlander and H. Nickel. 2003. Fulgoromorpha, Cicadomorpha excluding Cicadellidae. Volume 1. The Auchenorrhyncha of Central Europe. Brill Academic Publishing, Leiden, Netherlands.

Holzinger, W. E. 2007. Redescriptions of Ordalonema faciepilosa, Peltonotellus melichari and P. raniformis, with a key to Western Palaearctic Caliscelidae genera (Insecta: Hemiptera: Fulgoromorpha). European Journal of Entomology 104(2): 277-283.

Horváth, G. 1895a. Hémiptères nouveaux d’Europe et des pays limitrophes. Revue d’entomologie 14: 152-165.

Horváth, G. 1897a. Description d’Hémiptères nouveaux et notes diverses. Revue d’entomologie 16: 81-97.

Kato, M. 1933. Notes on Japanese Homoptera, with descriptions of one new genus and some new species. The Entomological World 1: 452-471.

Kirkaldy, G. W. 1913. On some new species of leafhoppers. Part 1. Bulletin. Hawaiian Sugar Planters’ Association Experiment Station. Division of Entomology. 12: 7-27.

Kusnezov, V. 1930. Ubersicht der asiatischen Caliscelaria-Arten (Homoptera-Issidae). Entomologische Zeitung. Herausgegeben von dem entomologischen Vereine zu Stettin 91: 267-278.

Maw, H.E.L., R. G. Foottit, K.G.A. Hamilton and G.G.E. Scudder. 2000. Checklist of the Hemiptera of Canada and Alaska. NRC Research Press, Ottawa.

Melichar, L. 1906. Monographie der Issiden. (Homoptera). Abhandlungen der Kaiserlich-Königliche Zoologisch-Botanischen Gesellschaft in Wien 3: 1-327.

Melichar, L. 1907. Bemerkungen zur Monographie der Issiden (Homoptera). Wiener Entomologische Zeitung 26: 323-324.

Meng Rui, Gnezdilov V. M. & Wang Ying-Lun 2015. Two new species of the genus Peltonotellus Puton (Hemiptera: Fulgoromorpha: Caliscelidae) from northwestern China with a world checklist. Zootaxa 4052(4): 465-477 [467].

Metcalf, Z. P. 1923. A Key to the Fulgoridae of Eastern North America with descriptions of new species. Journal of the Elisha Mitchell Scientific Society 38(3): 139-230. [see]

Metcalf, Z. P. 1958. General Catalogue of the Homoptera. Fascicle IV, Fulgoroidea, Part 15, Issidae. Waverly Press, Inc., Baltimore, Maryland.

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.

Mulsant M. E. and C. Rey. 1855. Description de quelques Hémiptères-Homoptères nouveaux ou peu connus. Annales de la Société linnéenne de Lyon. (Ser. 2) 2: 197-249 [see p. 207].

Puton, A. 1886. Homoptera Am. Serv. (Gulaerostria Zett. Fieb.) Sect. 1. Auchenorhyncha Dumér. Cicada Burm. p. 67-89. In: A. Puton.  Catalogue des Hémiptères de la faune paléarctique. Hétéroptères, Cicadines et Psyllides – 3e Edition, 1886.  [see p. 70] [BHL has 4th edition]

Stål, C. 1862. Novae vel minus cognitae Homopterorum formae et species. Berliner Entomologische Zeitschrift 6: 303-315.

Uhler, P. R. 1876. List of Hemiptera of the Region West of the Mississippi River, including those collected during the Hayden Exploration of 1873. Bulletin of the United States Geological and Geographical Survey of the Territories 1(5): 269-361.

Van Duzee, E. P. 1908. Studies in North American Fulgoridae. Proceedings of the Academy of Natural Sciences of Philadelphia 1907: 467-498.

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

Van Duzee, E. P. 1916a. Check list of Hemiptera (excepting the Aphididae, Aleurodidae and Coccidae) of America North of Mexico. New York Entomological Society, New York. 111 pp.

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.

last edit: July 2018

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