Genus Nilaparvata Distant, 1906

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Family Delphacidae Leach, 1815

Subfamily Delphacinae Leach, 1815

Tribe Delphacini Leach, 1815

Genus Nilaparvata Distant, 1906: 473.

Synonyms

= Kalpa Distant, 1906 (Type species Kalpa aculeata Distant 1906); syn. by Muir 1919: 7-8.

Distribution:

Pantropical, with most species in the Old World tropics.

Type species (in original combination): Nilaparavata greeni Distant, 1906: 473, a junior synonym of Delphax lugens Stal, 1854.

Recognized species

19 recognized species worldwide as follows:

New World

1 Nilaparvata caldwelli Metcalf, 1955 –  Belize, Puerto Rico, Belize, French Guiana, Hispaniola (D.R.), Mexico (Oaxaca); USA: Florida, North Carolina (Senegal, Sudan; reports from Africa need confirmation)
= Nilaparvata muiri Caldwell, 1951: 193 (in Caldwell & Martorell 1951) (nec China, 1925).
= Nilaparvata caldwelli Metcalf, 1955: 262; replacement name for unavailable N. muiri Caldwell, 1951.
= Nilaparvata bis Nast 1984: 396; replacement name for unavailable N. muiri Caldwell, 1951; syn. by Bartlett, 2007: 51.

2 Nilaparvata gerhardi (Metcalf, 1923: 210) – Canada (BC, Manitoba, Ontario, Quebec,  Saskatchewan); USA: Delaware, Illinois, Louisiana, Missouri,  New Hampshire, New Jersey, North Carolina, Texas, Utah, Virginia; ? Guatemala
= Liburnia gerhardi Metcalf, 1923: 210, pls. 62 & 70.
= Euidella gerhardi (Metcalf 1923); comb. by Metcalf 1949: 58.
= Euides gerhardi (Metcalf 1923); comb. by implication Metcalf 1952: 230-231.
= Nilaparvata gerhardi (Metcalf 1923); comb. by Bartlett 2007: 55.

3 Nilaparvata serrata Caldwell, 1951 – Bahamas, Belize, Costa Rica, Mexico (Distrito Federal, (Chiapas, Oaxaca, Veracruz), Nicaragua, Puerto Rico, Cayman Islands, ? Bolivia; USA: Florida

4 Nilaparvata wolcotti Muir and Giffard, 1924 – USA: Arizona, California, Colorado, Delaware, Florida, Georgia, Maryland, New Jersey, Oklahoma, Utah; Puerto Rico, Mexico (Guanajuato, Veracruz), Costa Rica

Old World (distribution records may be incomplete)
1 Nilaparvata albotristriata (Kirkaldy, 1907)  – Australia
= Delphax albotristriatus Kirkaldy 1907a: 154.
= Delphax thyestes Kirkaldy 1907a: 152 syn by Bellis & Donaldson 2016.
= Nilaparvata albotristriata (Kirkaldy); comb. apparently by Ding, 1981; (see also Metcalf 1943: 322).
= Chloriona thyestes (Kirkaldy); comb. by Metcalf 1943: 328.

2 Nilaparvata angolensis Synave, 1959  – Angola

3 Nilaparvata bakeri (Muir, 1917) –  Australia (Northern Territory, Queensland); China (Guangdong), Japan (Honshu, Kyushu, Shikoku); Nansei-shoto (Ryukyu Islands), Philippines (Luzon), Sri Lanka, Taiwan
= Delphacodes bakeri Muir, 1917, original combination
= Nilaparvata bakeri (Muir, 1917), comb. by Muir (1923)

4 Nilaparvata camilla Fennah, 1969 – Sudan

5 Nilaparvata castanea Huang and Ding, 1979 – China

6 Nilaparvata chaeremon Fennah, 1975 – Sri Lanka

7 Nilaparvata diophantus Fennah, 1958a – Portuguese Guinea

8 Nilaparvata lineolae Huang and Tian, 1979 – China

9 Nilaparvata lugens (Stal, 1854: 246) – Widespread in tropical and temperate Asia and Pacific Islands (Bangladesh, Brunei, Cambodia, Caroline Islands, China (Hainan, Hong Kong, Hubei, Zhejiang; Jiangsu); Fiji, India, Indonesia, Japan (Hokkaido, Honshu, Kyushu, Shikoku); Korea, Laos, Malaysia Mariana islands, Myanmar (ex Burma), Nansei-shoto (Ryukyu Islands), Nepal, Pakistan (palearctic), Palau, Papua New Guinea, Philippines, Singapore, Solomon Islands, Sri Lanka, Thailand, Vietnam )
= Delphax lugens Stål, 1854: 246, original combination.
= Delphax sordescens De Motschulsky, 1863: 109; synonym by Melichar, 1903: 102, 225; Muir & Giffard (1924)
= Nilaparvata greeni Distant, 1906: 473; synonym according to Muir (1919: 7, 1922: 350); Muir & Giffard (1924)
= Kalpa aculeata Distant, 1906: 474; synonym according to Muir (1919: 8, 1922: 350); Muir & Giffard (1924)
= Delphax parysatis Kirkaldy, 1907: 153; synonym according to Muir & Giffard (1924).
= Delphax ordovix Kirkaldy, 1907: 152; synonym according to Muir & Giffard (1924).
= Liburnia sordescens (De Motschulsky, 1863); comb. by Melichar, 1903: 102, 225.
= Dicranotropis anderita Kirkaldy, 1907: 133; syn. by Muir 1919: 7.
= Delphacodes anderita (Kirkaldy, 1907); comb. by Muir, 1917: 311.
= Delphacodes parysatis (Kirkaldy, 1907); comb. by Muir, 1917: 333.
= Delphacodes ordovix (Kirkaldy, 1907), comb. by Muir 1917: 333.
= Delphacodes sordescens (De Motschulsky, 1863); comb. by Muir, 1919: 8 (transfer implicit).
= Nilaparvata sordescens (De Motschulsky, 1863); comb. by Muir, 1922: 350.

10 Nilaparvata maeander Fennah, 1958 – Guinea, Sudan, Mauritius

11 Nilaparvata muiri China, 1925 – South Korea, China (Hubei, Yunnan, Zhejiang); Japan (Hokkaido, Honshu, Kyushu, Shikoku), Taiwan, Vietnam .

12 Nilaparvata myersi Muir, 1923 – New Zealand (North), Australia (Tasmania)

13 Nilaparvata nigritarsis Muir, 1926 – South Africa, Sudan
Nilaparvata oryzae (Matsumura, 1907) [Japan, Korea, Formosa, China, Manchuria [I think this is a missed synonym of N. lugens)]
= Delphax oryzae Matsumura, 1907 original combination
= Nilaparvata oryzae (Matsumura, 1907); comb. by Kaburaki and Imamura, 1932

14 Nilaparvata seminula Melichar, 1914: 110 – Philippine Islands (Java)

15 Nilaparvata terracefrons Guo and Liang, 2005 – China (Guangdong)

Note: Some species of Nilaparvata, in particular N. lugens, have long-distance migration.

Known hosts:

Nilaparvata lugens, N. bakeri, N. maeander and N. muiri – Rice (Oryza sativa L.)

Nilaparvata lugens – Oryza sativa L., Saccharum officinarum L. Zizania sp. (Poales, Poaceae) (Hasegawa 1955: 132, Sigsgaard 2007);
Nilaparvata bakeri, N. muiri – Cutgrasses (Leersia spp.); Cui et al. (2013) reports these species on Leersia hexandra Sw. and Leersia sayanuka Ohwi in laboratory experiments (i.e., not on rice)

Nilaparvata bakeri – Leersia oryzoides (L.) Sw. (Poaceae) by Hasegawa (1955: 132)

Nilaparvata gerhardi has been reported from Schoenoplectus americanus (Pers.) Volkart ex Schinz & R. Keller  (reported as Scirpus americana) (bulrush, a sedge – Cyperaceae)

Nilaparvata muiriLeersia japonica (Makino ex Honda) Honda (Poaceae)

Nilaparvata wolcotti – I have collected this species on Schoenoplectus, but I am not sure which species it was.  They were found by D-vacing the base of the plant.

Hosts from Wilson et al. (1994) and Cui et al. (2013); see also FLOW.  Host names from USDA PLANTS or Tropicos.

Predators and parasites
Nilaparvata Lugens:
Agamermis changshaensis Bao, Luo and Luo, 1992 (Nematoda, Mermithida, Mermithidae) (Bao et al. 1992)
Agamermis unka Kaburaki et Imamura 1932 (Nematoda, Mermithida, Mermithidae) (Choo et al. 1989 , Choo & Kaya 1990: 513)
Anagrus sp. (Hymenoptera, Mymaridae) (Swaminathan & Ananthakrishnan 1984: 7).

Economic importance:

Of the four Nilaparvata species in the New World, none are known to be pests on economic crops, although Nilaparvata wolcotti has been recorded from sugarcane. Nilaparvata planthoppers endemic to the New World are often found at lights.

On the other hand, Nilaparvata lugens, the brown planthopper (BPH) is an important and widespread pest of rice (see ricehoppers.net).  Nilaparvata lugens can reach high densities on rice, and vectors Rice Grassy Stunt Virus (RGSV), and Rice Ragged Stunt Virus (RRSV).  A good recent reference on rice diseases transmitted by Nilaparvata lugens is Cabauatan et a;. (2009) (citation below).

Nilaparvata lugens has been intercepted at US ports, but so far has not become established.

Nilaparvata bakeri, N. maeander and N. muiri are also known from rice.  None of these four species are known from the New World.

Other Nilaparvata on rice are of comparatively minor (or at least local) importance.  Nilaparvata muiri is also a vector of Rice Grassy Stunt Virus (RGSV), Nilaparvata bakeri has also been recorded as a vector of Rice Grassy Stunt Virus (RGSV), and Rice Ragged Stunt Virus (RRSV).  Photos and additional information on these species can be found here: http://naturalhistory.museumwales.ac.uk/vectors/Home.php

A good general source of information on rice planthoppers is the ricehoppers blog: http://ricehoppers.net/. (Which seems to be down now.)

Recognition:

The genus Nilaparvata can be recognized by the presence of a series of teeth on the first tarsal segment of the hind leg.  New World Nilaparvata were reviewed by Bartlett (2007).

Key to New World species of Nilaparvata (modified from Bartlett, 2007)

1. Aedeagus appearing bifid from base to apex, without subapical serrate flange, parameres relatively narrow, apex acute and directed dorsolaterally … Nilaparvata caldwelli
1.’ Aedeagus not appearing bifid, with serrate flange originating on right side; parameres relatively broad, apex blunt and laterally directed… 2

2. Segment X with long processes, approximated at base, originated subdorsally; parameres in widest view broadly laterally projecting, lateral margin nearly truncate …Nilaparvata serrata

2’. Segment X with short processes or a pair of teeth, originating dorsally; parameres in widest view with dorsal margin broad and truncate, lateral margin acutely pointed, directed sublaterally … 3

3. Segment X processes very short, often reduced to teeth; originating somewhat medially from dorsolateral margin; parameres in widest view with dorsal margin gradually sloped upwards to a dorsolaterally directed apex … Nilaparvata wolcotti
3’. Segment X processes longer, curved; originating from dorsolateral margin of segment X; parameres with dorsal margin truncate, lateral apex directed sublaterally … Nilaparvata gerhardi

 

New World Nilaparvata are externally quite similar and are best identified by their male genitalia, in particular the shape of the parameres and aedeagus.

Nilaparvata caldwelli Metcalf, 1955 (Dorsal view and face) (Photos by Kimberley Shropshire, University of Delaware)

Nilaparvata caldwelli Nilaparvata caldwelli

Nilaparvata caldwelli is the least common of the New World Nilaparvata.  Additional views of this species, including the genitalia are here.

Nilaparvata gerhardi (Metcalf, 1923)

Nilaparvata gerhardi

Nilaparvata gerhardi appears to be distributed in temperate eastern North America, and only doubtfully in the tropics.  Additional views of this species, including genitalia are here.

 

Nilaparvata lugens (Stal, 1854) (not found in the New World)

Nilaparvata lugensNilaparvata lugens

Nilaparvata lugens – the brown planthopper – is not known from the New World, but has been intercepted at ports.  It is a very important rice pest throughout the Old world.  Externally it is very similar to the endemic New World species, but the parameres and shape of the aedeagus are quite different (see figure here).

An extension fact sheet (from the Solomon Islands) is here.

 

Nilaparvata serrata Caldwell, 1951

Nilaparvata serrataNilaparvata serrata

Nilaparvata serrata is found mostly in the New World tropics, but is known from Florida.  The shape of the male paramere (the image on the right is a left paramere) and aedeagus are distinctive in this species and can be see in the figure here.

Nilaparvata wolcotti Muir and Giffard, 1924

Nilaparvata wolcottiNilaparvata wolcotti

Nilaparvata wolcotti is widespread in North America, and is also known from Mexico and Puerto Rico. Like all the new World Nilaparvata, it is best recognized by the male genitalia, which can be seen here.

I do not have photographs of Nilaparvata muiri or N. bakeri, but here are line drawings of the genitalia from Wilson & Claridge, 1991 (from here).

Websites: Nilaparvata on…

EOL
FLOW
Discover Life
Bugguide (this genus not on Bugguide, oddly)
Insects of Australia (4 species)
Ricehoppers
Leafhopper, Planthopper & Psyllid Vectors of Plant Disease (search=Nilaparvata, if that works)
BOLD.

ICAR-NBAIR (India).

Web Images

Nilaparvata myersi Muir, 1923 (holotype) on Landcare Research – Manaaki Whenua (New Zealand)

(more to add)

Molecular resources

There are a wide variety of molecular resources available for Nilaparvata lugens including a variety of genes, an EST library, and an ongoing genome project.  From a molecular standpoint, Nilaparvata lugens is probably the best known delphacid.  A link to Nilaparvata lugens molecular resources on Genbank is (should be) here.  As of this writing, BOLD reports 138 233 specimen barcodes available here.

Resources for the remaining species of Nilaparvata are sparse, although Urban et al. (2010) sequenced four genes for Nilaparvata wolcotti (18s, 28, CO1, WG).

Selected References (always more to add and links to BHL always in progress):

There is a voluminous literature on Nilaparvata lugens; search for Nilaparvata here.

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.

Bao, Xue-Chun, Qi-Gui Luo and Da-Ming Luo. 1992. A new species of the genus Agamermis from China (Nematoda: Mermithidae). Acta Zootaxonomica Sinica 17(1): 20-24.

Bartlett, C. R.  2007. A review of the planthopper genus Nilaparvata (Hemiptera: Delphacidae) in the New World. Entomological News 118(1): 49-66.

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.

Bellis, G. A. and J. F. Donaldson. 2016. Reassessment of some of Kirkaldy’s Australian species of Delphacini (Hemiptera: Delphacidae: Delphacinae). Austral Entomology 55: 247–260. DOI: 10.1111/aen.12180.

Bellis, G. A., J. F. Donaldson, V. Quintao, A. Rice, D. Tenakanai and L. Tran-Nguyen. 2013. New records of Delphacini (Hemiptera: Delphacidae: Delphacinae) from Australia, Timor Leste and Papua New Guinea, and an updated checklist of Delphacini from Australia. Australian Journal of Entomology 53(2): 167-174. doi:10.1111/aen.12069.

Caldwell, J. S. and L. F. Martorell. 1951 [dated1950]. Review of the Auchenorynchous [sic] Homoptera of Puerto Rico. Part II. The Fulgoroidea except Kinnaridae. Journal of Agriculture of the University of Puerto Rico 34(2): 133-269.

Cabauatan, P. Q., R. C. Cabunagan and I. R. Choi. 2009. Rice viruses transmitted by the brown planthopper Nilaparvata lugens Stål. Pp 357-368. In: K.L. Heong and B. Hardy, (eds.) . 2009. Planthoppers: new threats to the sustainability of intensive rice production systems in Asia. Los Baños (Philippines): International Rice Research Institute.

Catindig, J. L. A., G. S. Arida, S. E. Baehaki, J. S. Bentur, L. Q. Cuong, M. Norowi, W. Rattanakarn, W. Sriratanasak, J. Xia and Z. Lu. 2009.  Situation of planthoppers in Asia.  Pp. 191-220. In: K. L. Heong and B. Hardy (eds.). Planthoppers: new threats to the sustainability of intensive rice production systems in Asia, International Rice Research Institute, Los Banos (Philippines).

China, W. E. 1925. The Hemiptera collected by Prof. J. W. Gregory’s expedition to Yunnan, with synonymic notes on allied species. Annals and Magazine of Natural History. London. (Ser. 9) 16: 449-485.

Choo H. Y., H. K. Kaya and J. B. Kim. 1989. Agamermis unka (Mermithidae) parasitism of Nilaparvata lugens in Rice Fields in Korea. Journal of Nematology 21(2): 254-259.

Choo H. Y. and H. K. Kaya 1990. Parasitism of Brown Planthopper and White backed Planthopper by Agamermis unka in Korea. Journal of Nematology 22(4): 513-517.

Cui, Y. L., J. C. He, J. Luo, F. X. Lai and Q. Fu. 2013. Host plants of Nilaparvata muiri China and N. bakeri (Muir), two sibling species of N. lugens (Stål). Chinese Journal of Rice Science 27(1): 105-110.

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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.

Hasegawa, H. 1955. Some notes on the Japanese species of the genus Nilaparvata. Bulletin of the National Institute of Agricultural Sciences (Japan) Ser.C, 5: 117-138.

Hayashi, M. and S. Fujinuma. 2016. Part Fulgoromorpha. Pp. 323-355. 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.

Jairin, J. P. Leelagud, A. Pongmee and K. Srivilai. 2017. Chromosomal location of a recessive red-eye mutant gene in the brown planthopper Nilaparvata lugens (Stål) (Insecta: Hemiptera). Advances in Entomology 5: 33-39. doi: 10.4236/ae.2017.51003. http://www.scirp.org/journal/PaperInformation.aspx?paperID=73605.

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