Gonatocerus (Gonatocerus) aegyptiacus Soyka, 1950

(Figs 1–7)

Gonatocerus aegyptiacus Soyka 1950: 125 –126 .

Holotype female [lost from PPDD] (not examined). Type locality: Shareh ElHaram, Giza, Egypt.

Lymaenon saipanensis Doutt 1955: 13 –15 .

Holotype female [USNM] (not examined). Type locality: Saipan Island, Northern Mariana Islands. Syn. n.

Lymaenon tarae Narayanan & Subba Rao 1961: 657 –659 .

Holotype female [IARI] (not examined). Type locality: Delhi, India. Syn. n.

Gonatocerus sp. r: Sahad 1982a: 246, 257 (mentioned).

Gonatocerus miurai Sahad 1982c: 195 –198 (= G. sp. r of Sahad 1982a).

Holotype female [KUEC] (not examined, but a photograph is available at: http://konchudb.agr.agr.kyushu-u.ac.jp/elkutype/exec/refile.cgi?&lang=en&no=2386&tax = Gonatocerus %20 miurai %20Sahad). Type locality: Matsue, Shimane Prefecture (Honshū Island), Japan. Synonymized under G. tarae (Narayanan & Subba Rao) by Zeya & Hayat 1995: 84. Syn. n.

Gonatocerus tarae (Narayanan & Subba Rao): Subba Rao & Hayat 1983: 137 (catalog); Zeya & Hayat 1995: 84 –88 (synonymy, redescription, distribution, host associations, important comments) + 143 (illustrations); Zeya & Khan 2011: 11 –12 (distribution in India); Guo et al. 2011: 55 (key); Anwar & Zeya 2012: 52 –53 (distribution in India).

Gonatocerus alami Shamim & Shafee 1984: 623 –624 .

Holotype female (see Zeya & Hayat (1995) for details on the possible holotype) [ZDAMU] (not examined). Type locality: Aligarh, Uttar Pradesh, India. Synonymized under G. tarae (Narayanan & Subba Rao) by Zeya & Hayat 1995: 84. Syn. n.

Gonatocerus sulphuripes (Förster): Sahad & Hirashima 1984: 3 (list), 22–25 (redescription, illustrations, distribution) [misidentification; Zeya & Hayat (1995) commented that Sahad & Hirashima’s record could be due to a misidentification of G. tarae (Narayanan & Subba Rao)].

Gonatocerus miurai Sahad: Sahad & Hirashima 1984: 3 (list), 34–37 (redescription, illustrations, distribution, host association).

Gonatocerus minor Matthews 1986: 220 (member of the sulphuripes species group).

Type locality: Hatfield Forest, Essex Co., England, UK. Syn. n.

Gonatocerus minor Matthews: Baquero & Jordana 2003: 16 (diagnosis, redescription, distribution); Donev 2005: 379 –380 (diagnosis, distribution).

Gonatocerus aegyptiacus Soyka: Huber et al. 2009: 271 (list).

Type material examined. Gonatocerus aegyptiacus Soyka: paratypes [NHMW]: 1 Ƥ on slide labeled: 1. “Giza Panicum colonum Nov. 1931 ”, 2. “Para-Type”, 3. “ Gonatocerus aegyptiacus (Soyka) Ƥ”, 4. “16”; 3 Ƥ on individual slides, same data except lacking labels # 2 and # 4; 1 Ƥ on slide labeled: 1. “Egypt Piramids’ Road on citrus 4.XI.31 ”, 2. “ Gonatocerus aegyptiacus (Soyka) Ƥ” .

Lymaenon saipanensis Doutt: 3 3 paratypes [EMEC] on individual slides labeled: 1. “by sweeping 3 Saipan, M.I. July 26, 1948 R.L. Doutt”; 2. “ Lymaenon saipanensis Doutt Paratype”; 2 Ƥ, 2 3 on individual slides labeled: 1. “by sweeping grass [relevant Ƥ or 3 symbol] Saipan, M.I. Nov. 11, 1948 R.L. Doutt”; 2. “ Lymaenon saipanensis Doutt Paratype”. The specimens are poorly mounted (uncleared). There was also one male paratype (collected 11.xi.1948) (Doutt 1955) that I have not seen.

Gonatocerus minor Matthews: holotype female [BMNH] on slide labeled: 1. “Holotype Ƥ Gonatocerus minor sp. nov. det. Matthews 1984 Holo-type [in red circle]”; 2. “ENGLAND: ESSEX Hattfield Forest 23.viii.1975 Noyes BM 1975–265 26 Nov 1983 M1022/6”. The holotype is in good condition, dissected under 5 coverslips. Also examined was the entire paratype series (all from England, UK), as listed by Matthews (1986).

Material examined. AUSTRIA. VIENNA, Vienna: 9.vii.1952 [1 Ƥ, 2 3, NHMW] [misidentified by W. Soyka as G. litoralis (Haliday)]; 25.ix.1953, H.-J. Stammer [2 3, NHMW]. CHINA. BEIJING, Mentougou District, Xiaolongmen Station, 39°59.22’N 115°31.48’E, 1095 m, 28.vii.2002, G. Melika [8 3, UCRC]. DENMARK. HOVEDSTADEN, Dyrehaven ( Jaegersborg Dyrehave, Zealand Island), Fortunens Indelukke, 4.vii.1925, J.P. Kryger [1 Ƥ, ZMUC]. EGYPT. GIZA, Giza, xi.1930, H. Priesner (on Echinochloa colona [as “ Panicum coloxynum ”]) [3 Ƥ, NHMW] (identified by W. Soyka as G. aegyptiacus but not listed in the paratype series). Also formally not part of the published paratype series are 2 males on slides [NHMW] labeled: 1. “Giza Panicum colonum Nov. 30 ”, 2. “Co-Type”, 3. “736”, 4. “3 Gonatocerus aegyptiacus (Soyka) 3”; and 1. “ October 31 Pyramids Road on Grasses ”, 2. “Co-Type”; 3. “737”; 4. “Mym: 3 Gonatocerus aegyptiacus (Soyka) 3”. FRANCE. GARD, near Gard River, 43°55’45’’N 4°23’25’’E, 10–13.vi.2005, J. George [2 Ƥ, UCRC]. GIRONDE, Sainte Colombe, 44°54’N 00°02’W, M. van Helden: 2.vii.1998 [1 Ƥ, UCRC]; 30.vii.1998 [2 Ƥ, 1 3, UCRC]; 13.viii.1998 [20 Ƥ, 7 3, UCRC]; 9.vii.1999 [1 Ƥ, UCRC]; 17.viii.2000 [1 Ƥ, UCRC]. GREECE. CENTRAL MACEDONIA, Lake Kerkini, Ecotourism site, 41°08’15.6’’N 23°13’01.2’’E, 65 m, 13–19.vi.2006, G. Ramel [1 Ƥ, UCRC]. HUNGARY. BÁCS-KISKUN, Tompa, 28.vii.1949, J. Erdös [1 Ƥ, NHMW / HNHM]. ITALY. LAZIO, Viterbo Prov., 5.5 km E of Monte Romano, 42°15.284’N 11°57.315’E, 207 m, 9.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [1 3, UCRC]. JAPAN. SHIMANE, Matsue, Honjo Farm, 20.ix.1979, K.A. Sahad (ex. Nephotettix cincticeps Uhler on Oryza sativa) [2 Ƥ, UCRC]. KYRGYZSTAN. CHUY, Bishkek, Thor-Aryk (Boz-Peldek Mt.), 42°46’55’’N 74°34’11’’E, 1094 m, 14.viii.1998, C.H. Dietrich [2 Ƥ, 23, UCRC]. DZHALAL-ABAD: near jct. Kara Kysmak and Chatkal River, 42°04’00’’N 71°35’41’’E, 2240 m, 18.vi.1999, C.H. Dietrich [1 Ƥ, UCRC]. 18 km WSW of Kazaeman, 41°21’01’’N 73°48’37’’E, 1550 m, 15.vii.2000, C.H. Dietrich [1 Ƥ, 1 3, UCRC]. NARYN, Dzhumgal-Too Ridge, Seok River E ravine, 42°13’13’’N 75°00’55’’E, 2620 m, 24.vii.2000, C.H. Dietrich [1 Ƥ, UCRC]. OSH, Karakuldzha, Lajsu Ravine, 40°31’20’’N 73°37’10’’E, 1815 m, 25.viii.1998, C.H. Dietrich [1 Ƥ, 1 3, UCRC]. TALAS, 18 km WSW of Taldy Bulak, 42°26’31’’N 72°49’12’’E, 1930 m, 15.vi.1999, C.H. Dietrich [1 Ƥ, UCRC]. MONGOLIA. ÖMNÖGOVI, Naran Bulag, 43°27’N 100°27’E, 1405 m, 17–20.vii.1994, J. Carpenter [1 Ƥ, UCRC]. RUSSIA. KRASNODARSKIY KRAY, Krasnodar, Krasnodar, All-Russian Research Institute of Biological Plant Protection, V.V. Kostjukov: 30.viii.2001 [1 Ƥ, UCRC]; 31.viii.2003 [1 Ƥ, UCRC]. MOSKOVSKAYA OBLAST’, Noginskiy rayon, Fryazevo, 7–15.vii.2000, M.E. Tretiakov [1 Ƥ, UCRC]. STAVROPOL’SKIY KRAY: Achikulak, 23.viii.2002, V.V. Kostjukov [2 3, UCRC, ZIN]. Prietokskiy, 2.ix.2002, V.V. Kostjukov [2 3, UCRC]. PRIMORSKIY KRAY: Ussuriyskiy rayon, Gornotayozhnoye, M.V. Michailovskaya: 12–17.viii.1999 [2 Ƥ, UCRC, ZIN]; 29–31.v.2000 [1 3, UCRC]; 11– 12.vi.2000 [1 Ƥ, UCRC]. UK. ENGLAND, London Borough of Richmond upon Thames, Richmond Park, J.S. Noyes: 18.vii.1996 [1 Ƥ, 1 3, CNCI]; 15.viii.1997 [1 3, CNCI].

Extralimital records. AMERICAN SAMOA. Tutuila Island, Mapusaga, 13–20.i.2002, M. Schmaedick [1 Ƥ, 1 3, UCRC]. AUSTRALIA. QUEENSLAND: Acacia Ridge (near Brisbane): 1–6.ix.1980, G. Gordh [2 3, UCRC]; 27.ix.1980, G. Gordh, E.C. Dahms [5 Ƥ, 1 m, UCRC]. Biggenden (32 km SE of Munna Creek), 24.ix.1995, J.D. Pinto [1 Ƥ, 8 3, UCRC]. Blackbutt Creek (9 km E of Blackbutt), 22.ix.1995, J.D. Pinto [1 3, UCRC]. Brisbane Forest Park, 27°24.05’S 152°48.11’E, 13–19.xii.2002, J. George, J. Munro, A. Owen [1 Ƥ, UCRC]. Cooloola State Forest, Camp Milo, 5.ix.1979, G. Gordh, E.C. Dahms [1 3, UCRC]. 9.3 km N of Ellis Beach, 30.iv.1990, J.M. Heraty [3 Ƥ, UCRC]. 5 km E of Gordonvale, 28.iv.1990, J.M. Heraty [2 Ƥ, 1 3, UCRC]. Great Sandy National Park, 26°00.62’S 153°02.80’E, 16–17.xii.2002, J. Munro, A. Owen [4 Ƥ, 1 3, UCRC]. Indooroopilly, 3.ix.1980, G. Gordh [1 Ƥ, UCRC]. Nambour, 18–22.xi.2000, C. Freebairn [2 Ƥ, UCRC]. 19 km S of Port Douglas, 19.xi.1979, E.C. Dahms, J.B. Woolley, J. LaSalle [1 Ƥ, 1 3, UCRC]. FIJI. Viti Levu, 25 km N of Sigatoka, 17.iv.1990, J.M. Heraty [1 3, UCRC]. INDIA. GOA, Bambolim Beach (S of Panaji), 15°26’46’’N 73°51’19’’E, 5 m, 15.xi.2003, J.M. Heraty [2 Ƥ, UCRC]. KARNATAKA: near Dandeli Reserve, 15°20’36’’N 74°37’19’’E, 530 m, 17.xi.2003, J.M. Heraty [1 Ƥ, UCRC]. E of Hassan, 12°58’36’’N 76°14’34’’E, 923 m, 26.xi.2003, J.M. Heraty [2 Ƥ, UCRC]. Kathagal, 14°30’41’’N 74°31’49’’E, 44 m, 18.xi.2003, J.M. Heraty [1 Ƥ, UCRC]. Mudigere, 13°07’09’’N 75°37’41’’E, 994 m, 24–25.xi.2003, J.M. Heraty [1 Ƥ, UCRC]. W of Mudigere, J.M. Heraty: 13°07’05’’N 75°30’20’’E, 850–912 m, 24.xi.2003 [1 Ƥ, UCRC]; 13°07’08’’N 75°30’37’’E, 950 m, 25.xi.2003 [1 Ƥ, UCRC]. [NATIONAL CAPITAL TERRITORY OF] DELHI, New Delhi, Indian Agricultural Research Institute, 28°37’51’’N 77°09’50’’E, 220 m, 5–6.xi.2003, J.M. Heraty [3 Ƥ, UCRC]. NEPAL. Kathmandu, 4500’, 19.xi.1961, E.S. Ross, D.Q. Cavagnaro [1 Ƥ, 1 3, CAS]. PAKISTAN. PUNJAB, Faisalabad, University of Agriculture Faisalabad, Horticulture Fruit Garden, 31°25.844’N 73°03.665’E, 184 m, 1–8.x.2011, M.S. &. C.D. Hoddle, citrus orchard [1 Ƥ, UCRC]. REPUBLIC OF SOUTH AFRICA. MPUMALANGA, Sabie, Bridal Veil, 11.xii.2003, I. Mikó, G. Melika [2 Ƥ, UCRC]. THAILAND. CHIANG MAI, 1–10.ii.1998, S. Sonthichai [1 Ƥ, UCRC]. UNITED ARAB EMIRATES. ABU DHABI, Al Ajban, 24°36’N 55°01’E, 9.xi– 7.xii.2005, A. van Harten [1 Ƥ, CNCI]. FUJAIRAH, Wadi Maidaq, 25°18’N 56°07’E, 26.x–9.xi.2006, A. van Harten [1 Ƥ, 1 3, CNCI]. RAS AL-KHAIMAH, Wadi Shawkah, 31.x–27.xi.2006, A. van Harten [3 Ƥ, 1 3, CNCI]. SHARJAH, Sharjah Desert Park, 6–28.xii.2006, A. van Harten [7 Ƥ, 1 3, CNCI]. USA. NEW YORK, Seneca Co., 4.5 mi. SW of Lodi, Silver Thread Vineyard, 42°33’45.5’’N 76°52’27.2’’W, 202 m, 24.v.2011, S.V. Triapitsyn, G. Loeb [1 3, UCRC].

Distribution. PALAEARCTIC: Austria *, Bulgaria (Donev 2005 [as G. minor], China *, Denmark *, Egypt, France *, Greece (Donev 2003, 2005) [as G. minor], Hungary *, Italy *, Japan (Sahad 1982c [as G. miurai]; Sahad & Hirashima 1984 [as G. miurai and G. sulphuripes]), Kyrgyzstan *, Republic of Korea (Sahad & Hirashima 1984) [as G. miurai], Mongolia *, Russia *, Spain (Baquero & Jordana 2003) [as G. minor], and UK (England) (Matthews 1986) [as G. minor]. AFROTROPICAL*: Republic of South Africa *, and United Arab Emirates *. AUSTRALASIA*: Australia *. OCEANIA: American Samoa *, Fiji *, Northern Mariana Islands (Doutt 1955) [as G. saipanensis]. ORIENTAL: India (Zeya & Hayat 1995; Zeya & Khan 2011) [as G. tarae], Nepal *, Pakistan *, and Thailand *. NEARCTIC*: USA *.

Redescription. FEMALE (non-type specimens from the Palaearctic region). Body length 630–960 µm (dry-mounted specimens). Head dark brown; scape, pedicel, and usually F1–F4 light brown, F5–F8 and clava brown; mesosoma and metasoma normally mostly yellow or light brown with brown markings on mesoscutum, axillae, scutellum, propodeum, and apical gastral terga, but sometimes mesosoma mostly brown with yellow or light brown areas and apical gastral terga brown to dark brown; legs mostly light brown.

Antenna (Fig. 1) with radicle 0.32–0.4× total length of scape, rest of scape 2.1–3.1× as long as wide; pedicel longer than F1; F2 about as long as F3 and often F8 (the longest funicle segments), F4 the shortest funicle segment, F7 about as long as F8 when bearing mps but notably shorter when lacking mps; F1–F6 withouts mps, F7 with 1 or 2 mps but sometimes without mps, F8 with 2 mps; clava with 8 mps, 3.2–4.1× as long as wide, about as long as combined length of F5–F8 or slightly shorter when F7 lacks mps.

Mesosoma (Fig. 3) a little shorter than gaster. Propodeum (Fig. 2) with submedian lines often hardly visible. Fore wing (Fig. 4) 3.8–4.4× as long as wide; longest marginal seta 0.38–0.52× maximum wing width. Fore wing disc slightly infumate throughout, bare behind submarginal vein, setose behind and beyond marginal vein although occasionally setae sparse between marginal vein and cubital row of setae. Hind wing 26–30× as long as wide; disc bare except for rows of setae along margins and a few additional setae, slightly infumate throughout; longest marginal seta 3.3–4.3× maximum wing width.

Metasoma (Fig. 4). Petiole wider than long. Ovipositor 0.8–0.9× length of gaster, at most barely exserted beyond its apex; ovipositor length: mesotibia length ratio 1.4–1.5:1.

MALE (non-type specimens from the Palaearctic region, previously unknown from Europe). Body length 630–800 µm (dry-mounted specimens). Similar to female except for normal sexually dimorphic features and the following. Coloration of body and legs usually at least a little darker than in female; scape and pedicel light brown, flagellum brown. Antenna (Fig. 5) with scape minus radicle about 1.8× as long as wide. Fore wing (Fig. 6) 4.4– 4.7× as long as wide. Genitalia as in Fig. 7.

Diagnosis. Gonatocerus aegyptiacus may be difficult to distinguish from G. fuscicornis (especially the darker-colored specimens and females in which F7 of the antenna bears 2 mps) but in the latter species the discal setae originate behind about middle of the submarginal vein (Figs 20, 24, 26) whereas in the former the fore wing disc is bare behind the entire submarginal vein (Figs 4, 6). In most specimens from Egypt identified by W. Soyka as G. aegyptiacus, and also in some other, small, specimens from Europe F7 of the female antenna either lacks mps or bears just 1 mps.

Gonatocerus aegyptiacus is also very similar to G. (Gonatocerus) californicus Girault, known from the New World and the Hawaiian Islands (Triapitsyn et al. 2010; Luft Albarracin & Triapitsyn 2012), and G. (Gonatocerus) utahensis Girault, known from the Nearctic region (Triapitsyn et al. 2010), but females of the latter two species have the antenna with F1 deeply incised dorsoapically (Huber 1988).

Hosts. Nephotettix cincticeps Uhler (Cidadellidae) (Sahad 1982a [as G. sp. r], 1982c [as G. miurai]; Sahad & Hirashima 1984 [as G. miurai]), as well as Nilaparvata lugens (Stål) and Sogatella furcifera (Horváth) (Delphacidae) (Zeya & Hayat 1995) [as G. tarae].