Genus Toya Distant, 1906

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Family Delphacidae Leach, 1815

Subfamily Delphacinae Leach, 1815

Tribe Delphacini Leach, 1815

Genus Toya Distant, 1906: 472.

Type species (in original combination): Toya attenuata Distant, 1906.

Generic Synonyms

=  Himeunka Matsumura and Ishihara 1945 (Type species Unkana kushiana Matsumura, 1945 in Matsumura and Ishihara 1945: 70): Synonymy made by Anufriev 1977: 865. Removed from synonymy by Kuoh et al. 1981: 190.

=  Metadelphax Wagner, 1963 (type species Delphax propinqua Fieber, 1866): Synonymy made by Fennah 1964: 142 (by implication; type species moved to Toya); Nast 1972: 65, Linnavuori 1973: 107 (generic synonymy listed). Removed from synonymy by Ding 2006: 511.  See also Gonzon and Bartlett 2008)

Distribution

Widespread, but most species in Old World tropics.

 

Recognized species

About 44 currently recognized species worldwide, and one nomen dubium.  Here are included several names that are doubtfully valid but for which I have no better information at present.  Any errors will be corrected as I find them.  This list below is not exactly the same as that presented in Gonzon and Bartlett 2008 (dated 2007).

New World

1 Toya boxi (Muir and Giffard, 1924) – USA: FL; Guyana, Paraguay, Puerto Rico, St. Lucia, Trinidad and Tobago
= Delphacodes boxi Muir, 1926; pp. 32-33, figs. 120-122.
= Toya boxi (Muir, 1926); new combination by Fennah, 1965a: 96.

2 Toya dietrichi Gonzon and Bartlett, 2008 – Mexico (Colima)

3 Toya goliai Gonzon and Bartlett, 2008 – USA: Florida

4 Toya idonea (Beamer, 1947) – USA: AL, FL, LA, MS, NC; Bahamas (South Bimini, Andros, Great Inagua), Belize, Brazil, Bolivia, Cayman Islands, Colombia, Guatemala, Honduras, Panama, Paraguay, Surinam, Uruguay, Venezuela
= Delphacodes idonea Beamer, 1947; p. 66, figs. 10a-10c.
= Toya idonea (Beamer, 1947) ; comb. by Gonzon and Bartlett 2008: 222.

5 Toya nigra (Crawford, 1914) – USA: FL; Argentina, Bahamas (Cat, Exuma Cays, Long); Belize, Brazil, Bolivia, Cayman Islands, Columbia, Costa Rica, Cuba, Dominica, Guyana, Guadeloupe, Jamaica, Mexico (Guerrero, Veracruz), Panama, Puerto Rico (inc. Mona and Vieques Is.), St. Lucia, Trinidad and Tobago, Venezuela
= Megamelus erectus niger Crawford, 1914: 608, 624.
= Delphacodes nigra (Crawford, 1914), new combination and emendation by Muir and Giffard, 1924: 32.
= Delphacodes axonopi Fennah, 1945: 434-435, fig. 107-110, synonymy by Gonzon and Bartlett 2008: 224.

6 Toya recurva Gonzon and Bartlett, 2008 – Mexico (Veracruz)

7 Toya venilia (Fennah, 1959) – Bahama Islands (Andros, Abaco Cay, Great Inagua); British Virgin Islands (Guana, Necker, Anegada, Beef, Great Camanoe, Great Dog, Great Thatch, Jost van Dyke, Tortola, Virgin Gorda); US Virgin Islands (St. John); Montserrat, Puerto Rico (incl. Muertos Is., Vieques Is.), Turks and Caicos Islands
= Delphacodes venilia Fennah, 1959: 261-262, figs. 7a-7c.
= Toya venilia (Fennah, 1959); combination by Fennah, 1965a: 96.

nomen dubium
Toya iaxartes (Fennah, 1959) – St. Lucia (see Gonzon and Bartlett 2008)

Old World

(Synonyms are incomplete, as well as some referring authorities; status of some names unclear)

1 Toya actaeon (Fennah, 1958) – Africa [Sudan?]
= Delphacodes actaeon Fennah, 1958
= Toya (Delphacodes) actaeon (Fennah, 1958); comb. by Linnavouri 1973: 107. (note: Linnavouri ‘s parenthetical notation of  Delphacodes should be understood to mean that this species came from Delphacodes, not that Delphacodes was a subgenus; my intent is just to report the notation as Linnavouri  provided it.)

2 Toya attenuata Distant, 1906: 472 – Sri Lanka, Taiwan, China
= Sogatella hedai Kuoh, 1977, original combination
= Toya hedai (Kuoh, 1977); comb. by Asche & Wilson (1990: 7)
= Toya hedai (Kuoh, 1977); synonym by Ding (2006: 504)

3 Toya beninu Fennah, 1975 – Sri Lanka

4 Toya bridwelli (Muir, 1920) –  AFR: Nigeria, Senegal, Sudan, Ivory Coast, Reunion; ORI: Taiwan, China, Thailand, Nansei-shoto (Ryukyu Islands); AUS: Australia (New South Wales, Northern Territory, Queensland, Western Australia).
= Delphacodes bridwelli Muir, 1920.
= Toya (Delphacodes) bridwelli (Muir, 1920); comb. by Linnavuori (1973: 107).
= Metadelphax bridwelli (Muir, 1920); comb. by Ding 2006.
= Toya bridwelli (Muir, 1920); combination restored by Gonzon & Bartlett (2007: 261)

Toya camena Fennah, 1969: 47 (Sudan), to Partoya camena (Fennah, 1969) by sche (1988: 200).

5 Toya canidia Fennah, 1969 – Sudan

6 Toya ceresensis (Muir, 1929) – South Africa, Cape Verde Is., Sudan, Nigeria, Ivory Coast
= Delphacodes ceresensis Muir, 1929, original combination.
= Toya (Delphacodes) ceresensis (Muir, 1929); comb. by Linnavuori (1973: 107).

7 Toya complexa (Muir, 1929) – South Africa, Natal, Uganda, Ethiopia
= Delphacodes complexa Muir, 1929, original combination.
= Toya complexa (Muir, 1929); comb. by Fennah (1969: 46).

8 Toya cularo Fennah, 1975 – Sri Lanka

9 Toya demophoon Fennah, 1964 – Cameroon, Ethiopia (as Abyssinia), Nigeria , Reunion

10 Toya dryope (Kirkaldy, 1907) – AUST: Australia, New Zealand, Samoa New Caledonia, Loyalty Is.
= Delphax dryope Kirkaldy, 1907, original combination
= Delphacodes dryope (Kirkaldy, 1907); comb. by Muir (1917).
= Toya dryope (Kirkaldy, 1907); comb. by Fennah (1965: 56).

Delphacodes epimelas Fennah, 1958 (original combination) to Toya (Delphacodes) epimelas (Fennah, 1958) by Linnavuori (1973: 107); to Rhombotoya epimelas (Fennah, 1958) by Asche (1988: 204).

11 Toya fulva (Yang, 1989) – Taiwan [species status uncertain]
= Sogatella fulva Yang, 1989, original combination
= Toya fulva (Yang, 1989) , comb. by Asche & Wilson (1990: 7)

12 Toya fusca Melichar, 1914 – Indonesia (Java)

Delphacodes hargreavesi Muir, 1929 (original combination), to Toya (Delphacodes) hargreavesi (Muir, 1929) by Linnavuori (1973: 107), to Spinidelphacella hargreavesi (Muir, 1929) by Asche (1988: 212).

13 Toya hessei (Muir, 1929) – South Africa, French Guinea
= Delphacodes hessei Muir, 1929, original combination
= Toya hessei (Muir, 1929), comb. by Fennah (1964: 142)

14 Toya hispidula (Lindberg, 1954) – Canary Island, Egypt, Madeira Archipelago, Morocco, Sudan
= Calligypona hispidula Lindberg, 1954, original combination.
= Toya (Calligypona) hispidula (Lindberg, 1954); comb. by Linnavuori (1973: 107).
= Toya hispidula (Lindberg, 1954); comb. by Nast (1975: 6).

15 Toya ibiturca Asche, 1980 – Turkey, France, Spain, Greece

16 Toya larymna Fennah, 1975 – Sri Lanka

17 Toya lazulis (Kirkaldy, 1907) – Australia (Queensland), Fiji
= Delphax lazulis Kirkaldy, 1907, original combination
= Delphacodes lazulis (Kirkaldy, 1907), combination by Fennah (1965: 58)

18 Toya lima (Yang, 1989)[genus status unclear] [Taiwan etc.)]
= Sogatella lima Yang, 1989, original combination
= Toya lima (Yang, 1989), comb. by Asche & Wilson (1990: 7)
= Herbalima lima (Yang, 1989), comb. by Emeljanov 1977: 117; see also  Anufriev and Emeljanov 1988

19 Toya lyraeformis (Matsumura, 1900) – Caroline Islands; Japan (Honshu, Kyushu, Shikoku), Mariana islands
= Liburnia lyraeformis Matsumura, 1900, original combination.
= Delphacodes lyraeformis (Matsumura, 1900), comb. by Metcalf (1943: 467).
= Falcotoya lyraeformis (Matsumura, 1900); comb. by Fennah (1969: 21).
= Toya lyraeformis (Matsumura, 1900), comb. by Fennah (1971: 581), Kuoh et al 1983: 154.

20 Toya mahensis (Distant, 1917) – Seychelles (Indian Ocean), Mauritius, Ghana, Mauritius
= Nilaparvata mahensis Distant, 1917
= Toya mahensis (Distant, 1917), comb. by Fennah (1964: 141)
= Delphacodes mahensis Muir, 1919; synonym of Nilaparvata mahensis Distant, 1917 by Metcalf 1943: 297.

21 Toya mamurra Fennah, 1969 – Ethiopia (as Abyssinia)

22 Toya mandonius Fennah, 1969 – Sudan

23 Toya mastanabal Fennah, 1969 – Ethiopia (as Abyssinia), Sudan, Reunion

24 Toya menedemus Fennah, 1969 – Sudan

25 Toya minutula (Melichar, 1903) – Sri Lanka
= Liburnia minutula Melichar, 1903, original combination.
= Delphacodes minutula (Melichar, 1903); comb. by Metcalf 1943: 469.
= Toya minutula (Melichar, 1903), comb. by Fennah (1975: 117)

26 Toya narcissus Fennah, 1969 – Sudan

Toya nigeriensis (Muir, 1920) – See Sogatella (Asche and Wilson 1990: 13).

27 Toya obtusangula (Linnavuori, 1957) – Corsica, Cyprus, Greece, Israel, Italy, Egypt, France, Spain
= Calligypona obtusangula Linnavuori, 1957, Original combination
= Toya obtusangula (Linnavuori, 1957), comb. by Dlabola (1974: 290)

28 Toya peruda Fennah, 1975 – Sri Lanka

29 Toya salambo Fennah, 1964 – Madagascar

30 Toya siaka Fennah, 1975 – Sri Lanka

31 Toya simulans (Dlabola, 1958) – Russia (Georgia), Kazakhstan, Tadzhikistan, Ukraine
= Calligypona simulans Dlabola, 1958.
= Metadelphax simulans (Dlabola 1958); comb. in Mitjaev, 1971: 66.
= Toya simulans (Dlabola, 1958); comb. by Nast, 1972: 65.

Delphacodes sporoboli Lindberg, 1958 (original combination); to Toya (Delphacodes) sporoboli (Lindberg, 1958) by Linnavuori (1973: 107); to Falcotoya sporoboli (Lindberg, 1358), comb. by Asche (1988: 174).

32 Toya suezensis (Matsumura, 1910) Azores, Canary Is., Crimea , Cyprus, Egypt, Iran, Iraq, Israel, Jordan, Lebanon, Madeira Archipelago, Morocco, Ukraine, Yugoslavia, Turkey
= Delphax suezensis Matsumura, 1910, original combination
= Liburnia suezensis (Matsumura, 1910); comb. by Debski (1924: 17).
= Sogatella suezensis (Matsumura, 1910; nec. Linnavuori); comb by. Linnavuori, 1964: 341, misidentification, see Asche and Wilson 1990: 22.
= Toya suezensis (Matsumura, 1910), comb. by Nast (1975: 5)

32 Toya superba (Emeljanov, 1964) – Kazakhstan, Mongolia
= Chloriona superba Emeljanov, 1964, original combination
= Toya superba (Emeljanov, 1964), comb. by Emeljanov (1977: 117)

Toya tateyamaella (Matsumura, 1935) – Japan (Honshu, Kyushu) (placement in Toya depends on whether syn of Himeunka with Toya is accepted)
= Unkana tateyamaella Matsumura, 1935, original combination
= Toya tateyamaella (Matsumura, 1935) according to Anufriev (1975: 864).
= Himeunka tateyamaella (Matsumura, 1935); comb. in Matsumura & Ishihara 1945: 71.

34 Toya terryi (Muir, 1917) –  China; Japan (Honshu, Kyushu), Nansei-shoto (Ryukyu Islands), Taiwan; see Hayashi & Fujinuma (2016: 341)
= Delphacodes terryi Muir, 1917, Original combination
= Toya terryi (Muir, 1917) synonym of Toya tuberculosa (Distant, 1916) by Fennah (1975: 117).
= Toya terryi (Muir, 1917) status revivisco according to Ding (2006: 506).

Note: Toya terryi (Muir) was synonymized with T. tuberculosa (Distant) by Fennah 1975: 117, however; Ding (2006: 506) reinstates T. terryi as a senior synonym of Sogatella fulva Yang 1989, S. lima Yang 1989, and “Toya tuberculosa Fennah (nec Distant 1916)”)

35 Toya thomasseti (Muir, 1925) – Mauritius (Rodriguez Islands, Indian Ocean); Reunion, Saint Helena, Sudan
= Delphacodes thomasseti Muir, 1925, original combination
= Toya thomasseti (Muir, 1925) comb. by Fennah (1964: 141)
= Toya (Delphacodes) thomasseti (Muir, 1925;: comb. by Linnavuori (1973: 107).
= Opiconsiva modesta Distant, 1917; synonym of Toya thomasseti (Muir, 1925) in Fennah (1964: 141) (not noted as new syn.)
= Delphacodes distanti Metcalf, 1943; synonym of Toya thomasseti (Muir, 1925) in Fennah (1964: 141) (not noted as new syn.)

36 Toya tricolorata (Dlabola, 1961) – Kazakhstan, Tadzhikistan
= Calligypona tricolorata Dlabola, 1961, original combination.
= Toya tricolorata (Dlabola, 1961), comb. in Nast 1972: 65.

37 Toya trophonius Fennah, 1967 – Samoa (not listed in Gonzon and Bartlett 2008)

38 Toya tuberculosa (Distant, 1916) – Greece, Italy (inc. Sicily), Spain, Turkey; China (Guangdong), Taiwan, Indonesia (Java), Sri Lanka, Japan;  South Africa, Sudan; Doubtful occurances: Reunion ? (Attié et al. 2008: 1621; refuted by Fennah (1975: 117); Saint Helena (Fennah 1976: 270; refuted by Fennah 1975: 117); Vietnam Fennah 1978: 221; refuted by Fennah 1975: 117)
= Liburnia varia Hesse, 1925, original combination
= Delphacodes varia (Hesse, 1925); comb. by Muir (1929: 220).
= Toya (Delphacodes) varia (Hesse, 1925); by Linnavuori (1973: 107).
= Toya varia (Hesse, 1925); comb. by Fennah (1964: 142); synonym of Toya tuberculosa (Distant) by Fennah (1976: 269)
= Toya terryi (Muir, 1917); synonym of Toya tuberculosa (Distant) by Fennah (1975: 117) (refuted in Ding 2006: 506)
= Liburnia tuberculosa Distant, 1916, original combination
= Toya tuberculosa (Distant, 1916); comb. by Fennah (1976: 269)
= Toya hispijimena Asche, 1980; synonym of Toya tuberculosa (Distant) by Asche (1988: 224)
= Toya terryi (Muir, 1917) status revivisco by Ding (2006: 506)
= Delphacodes terryi Muir, 1917, original combination

 Toya yanoi (Ishihara, 1952) – Japan (Shikoku) – see Opiconsiva albicollis (Fujinuma 2013)

“ Toya sp.” Jayanthi and Baskaran 1989, 1990; Described, but not named, by Jayanthi and Baskaran 1989, 1990 on Urochloa mutica (Forssk.) T.Q. Nguyen (as Brachiaria mutica) (Poaceae). Not clearly attributable to a named species.

Table 1 (revised from Gonzon and Bartlett 2008).  Summary of world Toya species. Names used as valid since Metcalf 1953; species epithets in bold currently placed in Toya (I have corrected some – but not all – errors that I have found).

Specific epithet and author (original genus when not Toya)	Synoptic Distribution[1]	Comments
actaeon (Fennah, 1958) (Delphacodes)		Moved to Toya by Linnavouri 1973: 107.
albicollis (Motschulsky, 1863) (Delphax)		See Vilbaste (1968: 37). Possibly error for albicollis Haupt, 1935 (junior syn of M. propinqua); Current combination Opiconsiva albicollis (Motschulsky)  by Fennah 1975: 112
argentinensis (Muir, 1929a) (Delphacodes)	NEO: Argentina (Teson et al. 1989)	Moved to Metadelphax by Gonzon and Bartlett 2008
attenuata  Distant, 1906	ORI: Sri Lanka, Taiwan (Metcalf 1943, Fennah 1975), China (Ding 2006)	Type of genus
beninu Fennah, 1975	ORI: Sri Lanka (Fennah 1975)
boxi (Muir and Giffard, 1924) (Delphacodes)	NEO: British Guiana (Fennah 1965a)	Placed in Toya by Fennah, 1965a: 96
bridwelli (Muir, 1920) (Delphacodes)	AFR: Nigeria, Senegal, Sudan, Ivory Coast; ORI: Taiwan, China (Metcalf 1943, Fennah 1958, Linnavuori 1973, Medler 1980, Asche 1988, Yang 1989, Ding 2006)	Reinstated into Toya from Metadelphax by Gonzon and Bartlett 2008
camena Fennah, 1969a		Placed in Partoya Asche by Asche (1988: 200)
canidia Fennah, 1969a	AFR: Sudan (Fennah 1969a, Linnavouri 1973, Nast 1975)
ceresensis (Muir, 1929b) (Delphacodes)	AFR: South Africa, Cape Verde Is., Sudan, Nigeria, Ivory Coast (Muir 1929b, Lindberg 1958, Linnavuori 1973, Medler 1980, Asche 1988)	Placed in Toya from Delphacodes by Linnavuori 1973: 107
complexa (Muir, 1929b) (Delphacodes)	AFR: South Africa, Natal (Metcalf 1943, Uganda, Ethiopia (as Abyssinia) (Fennah 1969a)	Placed in Toya by Fennah  (1969a: 46)
cularo Fennah, 1975	IND: Sri Lanka (Fennah 1975)
demophoon Fennah, 1963	IND: Cameroon, Ethiopia (as Abyssinia), Nigeria (Fennah 1963, 1969a; Linnavouri 1973, Medler 1980)
dryope (Kirkaldy, 1907) (Delphax)	AUST: Australia, New Zealand, Samoa New Caledonia, Loyalty Is. (Metcalf 1943, Fennah 1965b, 1969a)	Placed in Toya by Fennah (1965b: 56)
epimelas (Fennah 1958) (Delphacodes)		Placed in Toya by Linnavouri (1973: 107), moved to Rhombotoya by Asche 1988: 204.
euonymus Fennah, 1965b		Listed in Syndelphax by Fennah, 1975: 109-110.
fulva (Yang, 1989) (Sogatella)	IND: Taiwan (Yang 1989; Asche and Wilson 1990)	Moved from Sogatella to Toya by Asche and Wilson (1990: 7)
fusca Melichar, 1914	IND: Java (Metcalf 1943)
hargreavesi (Muir 1929b) (Delphacodes)		Placed in Toya by Linnavouri 1973: 107; current combination, Spinidelphacella hargreavesi (Muir) by Asche 1988: 212-213
hedai (Kuoh, 1977) (Sogatella)		Moved from Sogatella to Toya by Asche and Wilson (1990: 7); Junior synonym of T. attenutata Distant (Ding 2006: 504)
hessei (Muir, 1929b) (Delphacodes)	AFR: South Africa, French Guiana (Metcalf 1943, Fennah 1958, Asche 1988)	Placed in Toya by Fennah  (1964: 142)
hispidula (Lindberg, 1954) (Calligypona)	PAL: Canary Is., Egypt, Madeira Archipelago, Morocco, Sudan (Nast 1972, Linnavouri 1973, Nast 1975, Asche 1980)	Placed in Toya by Nast (1975: 6)
hispijimena Asche, 1980		Junior synonyn of Toya tuberculosa (Distant) by Asche (1988: 224-226)
iaxartes (Fennah, 1959) (Delphacodes)	NEO: St Lucia (Fennah 1959, 1965a)	Placed in Toya by Fennah, 1965a: 96; moved to nomen dubium by Gonzon and Bartlett 2008
ibiturca Asche, 1980	PAL: W. Turkey (Asche 1980), France (Giustina and  Remane 1999, 2001)
larymna Fennah, 1975	IND: Sri Lanka (Fennah 1975), China (Ding 2006)
lazulis (Kirkaldy, 1907) (Delphax)	AUST: Queensland (Metcalf 1943, Fennah 1965b)	Placed in Toya by Fennah (1965b: 58)
lima (Yang, 1989) (Sogatella)	IND: Taiwan (Yang 1989)	Moved from Sogatella to Toya by Asche and Wilson (1990: 7)
limbata Emeljanov, 1972		Placed in Herbalima Emeljanov, 1972 by Emeljanov 1977: 117 [see also Emeljanov 1988]
lyraeformis (Matsumura, 1900) (Liburnia)	IND: Korea (Lee 1979); China (Kuoh et al. 1983)	Placed in Toya by Fennah 1971b.  Placed in Falcotoya by Fennah, 1969b: 21.
mahensis (Distant, 1917) (Nilaparvata)	AFR: Seychelles (Indian Ocean) (Metcalf 1943), Mauritius, Ghana (Fennah 1964)	Placed in Toya by Fennah  (1964: 141)
mamurra Fennah, 1969a	AFR: Ethiopia (as Abyssinia) (Fennah 1969a)
mandonius Fennah, 1969a	AFR: Sudan (Fennah 1969a, Linnavouri 1973)
mastanabal Fennah, 1969a	AFR: Ethiopia (as Abyssinia) (Fennah 1969a)
menedemus Fennah, 1969a	AFR: Sudan (Fennah 1969a, Linnavouri 1973)
minutula (Melichar, 1903) (Liburnia)	IND: Sri Lanka (Metcalf 1943, Fennah 1975)	Placed in Toya by Fennah (1975: 117)
minuscula (Horváth, 1897) (Delphax)		Placed in Falcotoya by Emeljanov 1972: 79
narcissus Fennah, 1969a	AFR: Sudan (Fennah 1969a, Linnavouri 1973)
nigeriensis (Muir 1920) (Megamelus as M. furcifer (sic) nigeriensis)		Placed in Toya by Linnavouri 1973: 107; current combination, Sogatella nigeriensis (Muir) by Asche and Wilson (1990: 7, 13)
obtusangula (Linnavuori, 1957) (Calligypona)	PAL: Cyprus, Israel, Italy, Egypt (Nast 1972, Dlabola 1974, Nast 1975, Asche 1980)	Placed in Toya by Dlabola 1974: 290
peruda Fennah, 1975	IND: Sri Lanka (Fennah 1975)
propinqua (Fieber, 1866) (Delphax)	Widespread in Neotropical, Nearctic, Palearctic, Afrotropical, and Indomalayan regions (e.g., Metcalf 1947, Caldwell and Martorell 1951, Schiemenz 1970, Nast 1972, Lodos and Kalkandelen 1980, Yang 1989, Holzinger et al. 2003, Ding, 2006)	Reinstated to Metadelphax by Ding (2006: 511, 513)
salambo Fennah, 1964	AFR: Madagascar (Fennah 1964)
sapporonis Matsumura 1935		Current combination Kakuna velitchkovskyi (Melichar, 1913) by Anufriev, 1977: 861
siaka Fennah, 1975	IND: Sri Lanka (Fennah 1975)
simulans (Dlabola, 1958) (Calligypona)	PAL: USSR (Georgia, Kazakhstan, Tadzhikistan, Ukraine) (Nast 1972)
sp. Jayanthi and Baskaran		Described, but not named, by Jayanthi and Baskaran 1989, 1990 on Urochloa mutica (Forssk.) T.Q. Nguyen (as Brachiaria mutica).  (Poaceae) Not clearly attributable to a named species.
sporoboli (Lindberg, 1958) (Delphacodes)		Placed in Toya by Linnavouri (1973: 107), moved to Falcotoya by Asche 1988: 174
suezensis (Matsumura, 1910) (Delphax)	PAL: Azores, Canary Is., Cyprus, Egypt,  Iran, Iraq, Isreal, Jordan, Lebanon, Madeira Archipelago, Morocco, Ukraine, Yugoslavia (Nast 1972), Turkey (Lodos and Kalkandelen 1980)	Placed in Toya by Nast (1975: 5-6)
superba (Emeljanov, 1964) (Chloriona)	PAL: Kazakhstan (Nast 1972)	Placed in Toya by Emeljanov 1977: 117
tangira (Matsumura, 1910) (Delphax)	AFR: Morocco (Tangier) (Nast, 1975)	Placed in Toya by Nast (1975: 6-8); moved  to Corbulo Fennah, 1965b, by  Kwon 1982: 7
tateyamaella (Matsumura, 1935) (Unkana)	PAL: Japan (Honshu, Kyushu) (Nast 1972)	Placed in Toya by Anufriev 1975: 864; current combination Himeunka tateyamaella by Kuoh et al. 1981: 191
terryi (Muir, 1917) (Liburnia)	IND: China, Taiwan (Ding 2006)	Toya terryi (Muir) was synonymized with T. tuberculosa (Distant) by Fennah 1975: 117, however; Ding (2006: 506) reinstates T. terryi as a senior syn of Sogatella fulva Yang 1989, S. lima Yang 1989, and “Toya tuberculosa Fennah (nec Distant 1916)”
thomasseti (Muir, 1925) (Delphacodes)	AFR: Rodrigues Islands, (Mascarene Islands, Indian Ocean), Seychelles Is., Ghana, Sudan, Nigeria, Ivory Coast; St. Helena (Atlantic Ocean) (Metcalf 1943, Fennah 1964, 1969a, 1976; Linnavouri 1973, Medler 1980, Asche 1988)	Placed in Toya by Fennah  (1964: 141)
tricolorata (Dlabola, 1961) (Calligypona)	PAL: USSR (Kazakhstan, Tadzhikistan) (Nast 1972)	Placed in Toya by Nast 1972: 65
tuberculosa (Distant, 1916) (Liburnia)	Widely distributed Indomalayan, Aftrotropical, southern Mediterranean regions, plus St. Helena (Metcalf 1943, Fennah 1975, Medler 1980, Asche 1980, 1988; Yang 1989)	Placed in Toya by Fennah (1975: 117), see also Fennah (1976: 269); Fennah (1975: 117) placed Delphacodes terryi Muir 1917 as junior synonym of Toya tuberculosa (Distant), but see Ding (2006)
varia (Hesse, 1925) (Liburnia)		Placed in Toya by Fennah (1964: 142); Jr. syn of Toya tuberculosa (Distant) by Fennah (1976: 269)
venilia (Fennah, 1959) (Delphacodes)	NEO: Montserrat, ?Puerto Rico (Fennah 1959), Guana Is., (Bartlett 2000), British Virgin Islands (Denno et al. 2001)	Placed in Toya by Fennah, 1965a: 96
yanoi (Ishihara, 1952) (Sogata)	PAL: Japan (Shikoku) (Nast 1972)	Placed in Toya by Nast 1972: 65; see Opiconsiva albicollis (Fujinuma 2013)

[1] Biogeographic Region: AFR = Aftotropical, AUST = Australian, NEA = Nearctic, NEO = Neotropical, IND = Indomalayan, PAL = Palearctic.

Note: Some references such as Harder and Bakker (1973) and Abo and Sy (1997) list Delphacodes catilina Fennah, 1958 as a Toya, but I have not found where this species may have been transferred.

Plant associations

Reported hosts are mostly grasses.

Toya bridwelli – Cynodon dactylon (L.) Pers., Pennisetum clandestinum Hochst. ex Chiov., Saccharum officinarum L., Setaria sphacelata (Schumach.) Stapf & C.E. Hubb. ex M.B. Moss (Poaceae; Bonfils et al. 1994)

Toya demophoon – Pennisetum clandestinum Hochst. ex Chiov., Setaria sphacelata (Schumach.) Stapf & C.E. Hubb. ex M.B. Moss (Poaceae; Bonfils et al. 1994)

Toya ibiturca – Cynodon dactylon (L.) Pers. (Bermudagrass)

Toya mastanabal – Pennisetum clandestinum Hochst. ex Chiov., Zea mays L. (Poaceae; by Bonfils et al. 1994)

Toya recurva – Sporobolus virginicus (L.) Kunth (Poaceae), by Gonzon & Bartlett (2007: 233)

Toya obtusangula – Cynodon dactylon

Toya thomasseti – Cynodon dactylon (L.) Pers., Hemarthria altissima (Poir.) Stapf & C.E. Hubb., Panicum maximum Jacq., Paspalum sp. , Pennisetum clandestinum Hochst. ex Chiov., Setaria sphacelata (Schumach.) Stapf & C.E. Hubb. ex M.B. Moss, Zea mays L. (ALL Poaceae, Bonfils et al. 1994, ET. CIT)

Toya tuberculosa – Cynodon dactylon (L.) Pers.; Panicum repens L.; Pennisetum clandestinum Hochst. ex Chiov.; Setaria sphacelata (Schumach.) Stapf & C.E. Hubb. ex M.B. Moss; Zea mays L. (all Poaceae; Bonfils et al. 1994)

Toya venilia – Sporobolus virginicus (L.) Kunth (seashore dropseed)

Toya dryope – Carrot (Daucus carota L., Apiaceae)

Toya sp. – Urochloa mutica (Forssk.) T.Q. Nguyen (as Brachiaria mutica) (Para grass; Poaceae).

Plant names according to the PLANTS Database (http://plants.usda.gov).  See Also FLOW for plant associations I may have missed. See Bonfils et al. 1994, Wilson et al. 1994, Gonzon and Bartlett 2008.

Economic importance

Toya species are potentially of economic importance.  Unspecified Toya species are reported on rice, although evidently not an important component of the rice planthopper fauna (specimens reported at Toya spp. could also be Metadelphax propinqua). Toya bridwelli has been reported from sugarcane, other Toya species from corn (see above).  Three Toya species reported from bermudagrass.

Greber (1981) listed Toya sp as a vector of maize stripe tenuivirus.

“Toya catilina” was given as the vector of African Cereal Streak (ACSV) (Harder and Baker 1973, Abo and Sy 1997), which appears to be a synonym of Maize streak monogeminivirus (MSV), although delphacids are not given as a vector of that disease.

Recognition

A difficult genus in many respects.  The genera and species are best separated by features of the male genitalia.  Females generally can not be identified with confidence, even to genus.  New World species were recently revised by Gonzon and Bartlett (2008).  Very similar to Metadelphax, Harmalia (now subsumed into Opiconsiva), and Syndelphax in particular.  All of these genera share several similarities including small and usually pale (straw) colored (interestingly, the type species, Toya attenuata, is dark-colored along with Toya nigra).  All three of these genera also have an expanded dorsocaudal region of the pygofer.  In Toya, the apex of the dorsal expansion of the pygofer is inflected medially (this is not true of Syndelphax), and the diaphragm armature is wider than tall (taller than wide in Metadelphax, Harmalia, and Syndelphax).  Metadelphax probably should be considered a subgenus of Toya, but as noted by Gonzon and Bartlett (2008), the 34 Old World Toya should first be investigated to define Toya in the strict sense and define subgenera among the remaining species.

Key to species of New World Toya species and allies (modified from Gonzon and Bartlett 2008) Figure numbers fromgonzon and Bartlett (2008) (avaliable here on JSTOR and PDF below)

1.  Genital diaphragm much wider than long, in caudal view concave or slightly convex (Figs. 2 A-J); aedeagus of most species slightly curved dorsad, bearing a lateral row of teeth near midlength on both sides plus several subapical dorsal teeth (Figs. 4 A-D)  …   Toya… 2

1’.- Genital diaphragm strongly projecting, much taller than wide (Figs. 3 A-J); aedeagus not as above … 8

2.  Inner angle of parameres well-developed, outer angle obscure, represented by broadly rounded projection (Figs. 9 E, 10 E) … 3

2’.- Parameres with inner and outer angles well developed (Figs. 6 E, 18 E, 21 E) … 4

3.  Armature of genital diaphragm with distinct lateral and median hump; parameres with outer angle strongly bulged (Figs. 10 D, F) … Toya goliai

3’ .- Armature of genital diaphragm with low lateral projections and obscure median hump; parameres with outer angles broadly curved (Figs. 9 D, F) …  Toya dietrichi

4.  Processes on segment X strongly upcurved (Fig. 13 H) … Toya recurva

4’.- Processes on segment X not, or very slightly, upcurved (Figs. 4 A-D) …  5

5.  Parameres with bump approximately at midlength on median margin, outer angle of apex broader and 2x as long as inner angle, inner angle directed rather medially (Fig. 12 E); processes on 10 closely approximated basally (Fig. 12 D) … Toya nigra

5’.- Parameres without median bump at midlength of ental surface, outer angle of apex less than 2x as long as inner angle, inner angle directed more dorsad (Fig. 10 E); processes on 10 approximated or separated … 6

6. Parameres without expanded basal angle, inner and outer angles developed approximately equally (Fig. 10 E); processes of segment X longer, basally approximated, and frequently arising ventrad of dorsocaudal angle, tightly approximated to 10 (Fig. 10 D); aedeagus strongly curved dorsad, bearing 10 or more teeth on each side in subparallel rows (Figs. 10 F, G) … Toya idonea

6’.- Parameres with expanded basal angle bearing well developed process, outer angles stouter than inner angles (Figs. 8 E, 14 E); processes of segment X relatively short, basally separated and in lateral view arising from dorsocaudal angles of 10 (Figs. 8 D, 14 D); aedeagus slightly curved dorsad, bearing 6 or fewer teeth on each side in parallel rows (Figs. 8 F-G, 14 F-G) …  7

7.  Parameres in widest view with outer angle broader and longer than inner angle (Fig. 8 E), aedeagus in ventral view with left and right rows of teeth forming lateral flanges (Figs. 8 F-G); northern South America, Central America, Caribbean, Florida … Toya boxi

7’.- Parameres in widest view with inner angles subequal to or longer than outer angles (Fig. 14 F), aedeagus with lateral rows of teeth not produced into a flange (Figs. 14 F-G); Caribbean, on Sporobolus grasses  … Toya venilia

8.  Dorsocaudal margin of pygofer greatly expanded and foliaceous, apex bent broadly and strongly medially, genital diaphragm strongly arrowhead-shaped (Figs. 3 I-J); parameres apically acute, without distinct outer or basal angles (Fig. 21 E); aedeagus flattened, bearing few subapical teeth (Fig. 21 F) … Hadropygos Gonzon and Bartlett, H. rhombos

8’.- Dorsocaudal margin of pygofer expanded to varying degrees, but not foliaceous, apex bent ventrad to mediad, or not at all, genital diaphragm bifid or rounded apically (Figs. 3 A-H); parameres usually with distinct outer, inner and basal lobes (Figs. 15 E, 20 E); aedeagus tubular, bearing few to many teeth, variously arranged (Figs. 15 F, 16 F, 17 E, 18 F, 19 F) … 9

9. Pygofer with dorsocaudal region slightly to broadly expanded in lateral view, expanded dorsal portion of pygofer in caudal view directed caudad, apices not inflected; genital diaphragm elongate, apically rounded, often narrowly spoon-shaped; brachypterous males often yellowish with strongly contrasting tegmina … Syndelphax

9’.- Expanded dorsal portion of pygofer in caudal view with apices inflected (usually medially or ventromedially directed); genital diaphragm elongate, apically bilobed or trilobed (except M. pero) (Figs. 3 A-H); brachypterous males without strongly contrasting tegmina … Metadelphax … 10

10. Armature of genital diaphragm apically convex, appearing rounded to trilobed (Figs. 3 A-B) … 11

10’.- Armature of genital diaphragm apically concave, appearing bifid (Figs. 3 C-H) …  12

11. Parameres with outer apical angle reduced, appearing curved medially (Figs. 17 F, G) (Cayman Islands) … M. pero

11’.- Parameres with outer angle well-developed, triangular in shape (Fig. 15 E) (South America) … M. argentinensis

12. Aedeagus simple, straight, bearing few teeth near apex and in lateral row (teeth variable in development, usually weak) (Fig. 18 F); frons infuscated along carinae, intercarinal regions pale (rarely more broadly infuscate in macropters, or pale in brachypters) (Fig. 18 B); parameres weakly concave apically, with moderately developed basal angle (Fig. 18 E); expansion of pygofer with ventromedially directed apex (Figs. 3 E-F); common and widespread species …  M. propinqua

12’.- Aedeagus slightly bent ventrad near apex (Figs. 16 F, 20 F); frons with intercarinal region infuscated (Figs. 16 B, 20 B); parameres with basal angles obscure, apically concave or with dorsomedially directed with subapical medially directed tooth (Figs. 16 E, 20 E); expansion of pygofer bilobed apically, with medially and caudally directed apices (Figs. 3 C, G) … 11

13. Paramere apices concave, lacking medially directed subapical tooth (Fig. 20 E) … M. wetmorei

13’.- Parameres dorsomedially pointed with medially directed subapical tooth (Fig. 16 E) …  M. dentata

Toya attenuata – Lectotype

 

Toya bridwelli

Toya dryope

(Numbered figures, below, from Gonzon and Bartlett, 2008.

Fig. 1. Left forewing of Toya, Metadelphax, and Hadropygos (scale = 0.5 mm). A. Toya attenuata (holotype), B. Metadelphax propinqua, C. Hadropygos rhombos (holotype).

 

Online resources

3i.
EOL.
Bugguide.
Discover Life.
FLOW.
Hoppers of North Carolina. (link to Toya idonea)
Kunzweb Gallery. (To Delphacini, Europe)
American Insects.
BOLD.
GBIF.
Fauna Europaea. (Treats Metadelphax as a synonym of Toya)

Molecular resources

Molecular resources for Toya appear quite limited. As of this writing, BOLD lists barcode data for lists barcode data for Toya attenuata. Genbank has data for 2 Toya species, several undet. Toya, and Metadelphax propinqua (as of 25 Sept. 2017).

Selected references

[References below are intended to be expansive – including some Metadelphax references and most of those included in Gonzon & Bartlett 2008; intended to include all descriptive taxonomy, nomenclatural acts, geographic records and biology, but some may be inadvertantly omitted]

Abo, M. E. and A. A. Sy. 1997. Rice Virus Diseases: Epidemiology and Management Strategies. Journal of Sustainable Agriculture 11(2): 113-134.

Anufriev, G. A. 1977. Delphacids (Homoptera Auchenorrhyncha) of the Kurile Islands Fauna. Zoologicheskii Zhurnal 56(6): 855-869. [In Russian.]

Anufriev, A. F. and A. F. Emeljanov. 1988. [20. Order Homoptera]: pp. 9-11. In: P. A. Ler, ed. [Keys to the identification of insects of the Soviet Far East. Vol. 2: Homoptera and Heteroptera] Nauka, Leningrad. 972 pp. [In Russian.]

Asche, M. 1980. Zwei neue Arten der Gattung Toya Distant 1906, aus dem Mittelmeergebiet mit erganzenden Bemerkungen zu Toya hispidula (Lindberg 1953) und Toya obtusangula (Linnavuori 1957) (Homoptera Cicadina Delphacidae). Marburger Entomologische Publikationen 1(4): 1-36. [Toya hispijimena Asche, 1980]

Asche, M. 1982. Beitrage zur Delphaciden-Fauna der Turkei (Anatolien) (Homoptera Cicadina Delphacidae). Marburger Entomologische Publikationen 1(7): 71-98.

Asche, M. 1985. Zur Phylogenie der Delphacidae Leach, 1815 (Homoptera: Cicadina: Fulgoromorpha). Marburger Entomologische Publikationen 2(1), volume 1 pp. 1-398, volume 2 pp. 399-910.

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Asche, M. and H. Hoch. 1982.  Beitrage zur Delphaciden-Fauna Griechenlands II (Homoptera Cicadina Delphacidae). Marburger Entomologische Publikationen 1(7): 37-70.

Asche, M. and R. Remane. 1982.  Beitrage zur Delphaciden-Fauna Griechenlands I (Homoptera Cicadina Delphacidae) Marburger Entomologische Publikationen 1(6): 231-290.

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