Gonatopus bicuspis (Olmi, 1993) Xu & Olmi & He, 2013Gonatopus lankae (Ponomarenko, 1981) Xu & Olmi & He, 2013Gonatopus malesiae ( Olmi, 1984 ) Xu & Olmi & He, 2013Gonatopus nepalensis ( Olmi, 1986 ) Xu & Olmi & He, 2013Gonatopus sarawakensis ( Olmi, 1984 ) Xu & Olmi & He, 2013Gonatopus validus ( Olmi, 1984 ) Xu & Olmi & He, 2013Gonatopus bengalensis ( Olmi, 1984 ) Xu & Olmi & He, 2013Gonatopus borneanus ( Olmi, 1984 ) Xu & Olmi & He, 2013Gonatopus insularis ( Olmi, 1984 ) Xu & Olmi & He, 2013Gonatopus pajanensis (Olmi, 1989) Xu & Olmi & He, 2013Gonatopus pyrillae ( Mani, 1942 ) Xu & Olmi & He, 2013

Xu, Zaifu, Olmi, Massimo & He, Junhua, 2013, <strong> Dryinidae of the Oriental region (Hymenoptera: Chrysidoidea) </ strong>, Zootaxa 3614 (1), pp. 1-460 : 347-388

1. Haplogonatopus apicalis R. Perkins, 1905

( Plates 103A, B , 104A, B )

Haplogonatopus apicalis R. Perkins 1905: 39 ; Olmi 1982: 305; Olmi 1984: 1417; Xu & He 1996b: 128; Guglielmino & Olmi

1997: 181–252.; Olmi 1998f: 163; Olmi 2000a: 94, 109; Barrion 2002: 71; He & Xu 2002: 310; Guglielmino & Olmi

2006: 51; Xu et al. 2012a: 13; Mita et al. 2012: 50. Haplogonatopus moestus R. Perkins 1905: 40 (synonymized by Olmi 1984); type locality: Cairns (Queensland, Australia). Haplogonatopus brevicornis R. Perkins 1906a: 489 (synonymized by Olmi 1984); type locality: Condong (Queensland,

Australia). Haplogonatopus orientalis Rohwer 1920: 159 (synonymized by Olmi 1984); type locality: Bilaspur (Chattisgarh, India); Otake et al. 1976: 289; Otake 1977: 46; Olmi & Currado 1979b: 43. Haplogonatopus japonicus Esaki & Hashimoto 1931: 25 (synonymized by Olmi 1984); type locality: Fukuoka (Kyushu,

Japan); Chiu 1979: 339; Chen & Yang 1987: 237; Chen 1988: 44. Haplogonatopus sp. near americanus R. Perkins : Pagden 1934: 2. Haplogonatopus sp. 1 : Chandra 1978a: 21; Chandra 1978b: 5; Chandra 1980a: 128; Chandra 1980b: 167. Monogonatopus orientalis (Rohwer) : Currado & Olmi 1979: 181. Haplogonatopus apicalis R. Perkins partim: Olmi & Currado 1979b: 42. Haplogonatopus fuscus Xu & He 1995b: 594 (synonymized by Olmi 1998f); type locality: Pingyang (Zhejiang, China); Olmi

1998f: 163; He & Xu 2002: 308. nec Haplogonatopus apicalis R. Perkins : Sugiura et al. 2004: 11.

Description of female ( Plate 103A ). Apterous; length 2.1–3.7 mm. Head yellow-testaceous; occasionally head brown or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 and 10 yellow; mesosoma yellow-testaceous; petiole black; metasoma testaceous, or reddish, occasionally more or less darkened, or brown; legs yellow-testaceous. Antenna clavate; antennal segments in following proportions: 19:11:22:12:10:10:10:10:10:19. Head excavated, shiny, hairless, granulated; occipital carina incomplete, only shortly present on sides of posterior ocelli; frontal line complete; POL = 5; OL = 5; OOL = 15. Palpal formula 2/1. Pronotum shiny, smooth, hairless, unsculptured, not crossed by strong transverse impression. Scutum shiny, sculptured by some longitudinal keels. Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 10:2:3:7:11. Enlarged claw ( Plate 103B ) with one subapical tooth and one row of three–six lamellae. Segment 5 of protarsus ( Plate 103B ) with two rows of seven–ten lamellae; distal apex with about two–seven lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 104A ). Fully winged; length 2.0– 2.2 mm. Head black or brown, except mandible yellow; antenna brown, occasionally with segments 1–2 yellow; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments in following proportions: 7:7:12:11:11:10:9:9:9:12; antennal segment 3 more than three times as long as broad (about 4.0–4.5). Head shiny, hairy, weakly granulated; frontal line absent; occipital carina absent; POL = 9; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OOL (4:3); temple distinct; occiput concave. Palpal formula 2/1. Scutum hairy, dull or shiny, granulated. Notauli complete, posteriorly strongly converging, joint or almost joint. Scutellum and metanotum shiny, hairy, finely punctate. Propodeum reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 104B ) long, slender, distally broadened and serrate. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of H. apicalis : AUSTRALIA: Queensland, Bundaberg, 1904, reared, dissected 16.II.1905 ( BPBM); ♀ lectotype (designated by Olmi, 1982) of H. moestus : AUSTRALIA: Queensland, Cairns, Mulgrave, VIII.1904, reared from Hadeodelphax nymph ( BPBM); ♀ holotype of H. brevicornis : AUSTRALIA: New South Wales, Tweed River Queensland side, Condong, 22.XII.1904, A. Koebele leg., reared from Liburnia (near Chloriona ) 6.I.1905, dissected 6.I.1905 ( BPBM); ♀ holotype of H. japonicus : JAPAN: Kyushu Island, Fukuoka Pref., Fukuoka, 16.IX.1930 ( ELKU); ♀ holotype of H. orientalis : INDIA: Chattisgarh, Janigir, Bilaspur, 14.X.1915, reared from Sogata sp. , on rice, C.S. Misra leg. ( USNM); ♀ holotype of H. fuscus : CHINA: Zhejiang, Pingyang County, 26°07'N 120°05'E, 2.VIII.1965, Junhua He leg. ( ZJUC). Paratypes of H. apicalis : same locality label as lectotype, 2♀♀ and 1♂ ( BPBM). Paratypes of H. orientalis : same locality label as holotype, 2♀♀ and 1♂ ( USNM). Paratypes of H. japonicus : same locality label as holotype, 15.XI.1930, 1♀ ( ELKU); same locality label as holotype, 16.IX.1930, 2♀♀ and ♂ allotype ( ELKU). Other material. CHINA: Fukien (= Fujian), Hotien, Changting, 24.VII.1940, T.C. Maa leg., 2♀♀ ( TARI); Fujian, Shaxian, reared from rice planthoppers, Changming Liu & Biyu Luo leg., 30♀♀ and 46♂ ( ZJUC); Fujian, Shaowu, 12.IV.1944, Xiufu Zhao leg., 1♀ ( ZJUC); Fujian, Mt. Wuyishan, 1983, Kechi Zhang leg., 1♂ ( ZJUC); Fujian, Fuzhou, 12.VI.1991, Changming Liu leg., 1♀ ( ZJUC); Fujian, Shouning, 4.VIII.1996, Changming Liu leg., 61♂ ( ZJUC); Fujian, Yunxiao, 1.VIII.1985, Bo Liu leg., 1♀ and 18♂ ( ZJUC); Guagdong, Shantao (=Shek Tung) ( BPBM); Guangdong, Yangjiang, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangzhou, Guangzhou Botanical Garden, parasitized host collected 13.VI.2002, dryinid cocooning 15.VI.2002, dryinid adult emerged 26.VI.2002, M. Olmi leg. ( MOLC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, reared from nymphs of unidentified Delphacids, M. Olmi reared, 3♀♀ ( MOLC); Guangdong, Conghua, VI.1978, Junhua He leg., 1♀ ( ZJUC); Guangdong, Zhaoqing, 1974, Ciwei Huang leg., 1♂ ( ZJUC); Guangdong, Sihui, 8.V.1998, Yufang Liu leg., 3♀♀ and 3♂ ( ZJUC); Mt. Nankunshan, parasitized host collected 2.VII.2005, dryinid cocooning 4.VII.2005, M. Olmi reared from adult of unidentified Delphacid, 1♀ ( MOLC); Guangdong, Mt. Yunjishan, 9.VI.2002, Zaifu Xu leg., 1♂ ( SCAU); Hainan, Qionghai, 4.V.1966, Junhua He leg., 4♂ ( ZJUC); Guangxi, Wuming, 31.V.1981, Guixian Su leg., 1♀ ( ZJUC); Guangxi, Nanning, 13.IX.1987, Junhua He leg., 4♀♀ ( ZJUC); Guangxi, Xilin, 5.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Tianlin, 30.V.1982, Junhua He leg., 1♂ ( ZJUC); Sichuan, Yuexi, 14.VIII.1980, Junhua He leg., 3♀♀ ( ZJUC); Guizhou, Sandu, IV–VII.1980, 7♀♀ and 9♂ ( ZJUC); Guizhou, Sinan, IX.1986, VIII.1991, 12♀♀ and 3♂ ( ZJUC); Shaanxi, Mt. Tiantaishan, VIII.1980, Jianhua Wei leg., 1♀ ( SCAU); Hubei, Zhuxi, 1981, 2♀♀ ( ZJUC); Hubei, Zhushan, 28.VIII.1982, Junhua He leg., 6♂ ( ZJUC); Hubei, Yunxian, 28.VIII.1982, Junhua He leg., 2♂ ( ZJUC); Hubei, Gucheng, 28.VIII.1982, Junhua He leg., 1♂ ( ZJUC); Hunan, Yueyang, 5.VII.1973, Binsen Wang leg., 1♂ ( ZJUC); Henan, Xinxiang, 11.X.1973, Weihua Li leg., 2♀♀ and 1♂ ( ZJUC); Taiwan, Changhua, Erh lin ( TARI); Taiwan, Chiayi ( TARI); Taiwan, Taichung, Tatu ( TARI); Taiwan, Tainan, Paiho ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, TARI ( TARI); Taiwan, Taoyuan, Chungli ( TARI); Taiwan, Kijin ( ELKU); Zhejjiang, Linan, 31.VII.1974, Junhua He leg., 3♂ ( ZJUC); Zhejjiang, Hangzhou, 20.IX.1952, 6.X.1965, VIII.1987, Junhua He leg., 2♀♀ and 1♂ ( ZJUC); Zhejjiang, Mt. West Tianmushan, 11.X.1982, Junhua He leg., 1♂ ( ZJUC); Zhejjiang, Tonglu, III.1966, Houliang Xu leg., 3♂ ( ZJUC); Zhejjiang, Shangyu, VIII–IX.1984, Haifu Wei leg., 45 ♂ ( ZJUC); Zhejjiang, Yuyao, VI.1972, Junhua He leg., 2♀♀ and 4♂ ( ZJUC); Zhejjiang, Dongyang, 1); Zhejjiang, Lishui, 4.IX.1975, IV–VI.1982, 18.VII.1984, Zhenfang Gu et al. leg., 40♂ ( ZJUC); Zhejjiang, Suichang, VIII.1974, Junhua He leg., 16♂ ( ZJUC); Zhejjiang, Wenzhou, 10.V.1981, Junhua He leg., 6♀♀ and 6♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg., 1♀ and 5♂ ( ZJUC). INDIA: Andhra Pradesh, Warangal, 15.IX.1986, paddy field, 3♀♀ ( BMNH); Bihar, Pusa (erroneously labelled as ♂ allotype of Pseudogonatopus sogatae Rohwer ) ( USNM); Chattisgarh, Janigir ( IARI); Chattisgarh, Raipur ( AMNH, GCC); Delhi, New Delhi ( IARI): Jharkhand, Ranchi, Kankey Farm ( IARI); Karnataka, Mandya, 22.XI.1976, ex Nilaparvata lugens , 1♀ ( BMNH); Karnataka, Chikkaballapura, IV.1917, T.V. Campbell leg., 1♀ ( BMNH); Madhya Pradesh, Javalpur, 12.IX.1969, ex Sogatella furcifera , 1♀ and 1♂ ( BMNH); Orissa, Cuttack, 1981, 2♀♀ and 1♂ ( BMNH); Orissa ( Kurian 1954a); Punjab, Ludhiana, 21.VIII.1990, K. Shukla reared from Sogatella furcifera (Horváth) , 8♂ ( BMNH). MALAYSIA: Malaya, Kuala Lumpur, 21.XII.1930, ex Sogata sp. , H.T. Pagden leg., 3♀♀ and 4♂ ( BMNH); Malaya, Kuala Kurau ( BMNH); Malaya, Serdang ( DBUS); Sarawak, Kuching, Tarat ( KYC). PHILIPPINES: Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 10.XII.1977, ex Nilaparvata lugens, A.T. Barrion leg., 1♀ ( BMNH); same locality label, further specimens ( GCC); Luzon Island, Batangas Prov., Tanauan ( GCC); Panay Island, Iloilo Prov., 2 Km NW Tigbauan, Bo. Bagumbayan, 10°41'N 122°24'E ( GCC); Mindoro Island, Mindoro Oriental, Baco, Alag River, 16.XI.1994, H. Zettel leg., 1♀ ( NHMW). SRI LANKA: Kandy District, Tismada ( USNM); Kandy District, Kalugomuwa (specimens described by Otake, Somasundaram & Abeykoon, 1976, now in AMNH) ( AMNH, BPBM). THAILAND: Tung Saew ( KYC); Pasang ( KYC); Tung Fabad ( KYC); Banmakok ( KYC); Chom-Thong ( KYC). Palaearctic: CHINA: Liaoning, Shenyang, 6.VI.1991, Juxian Lou leg., 1♂ ( ZJUC); Liaoning, Fuxin, 27.VII.1994, Juxian Lou leg., 2♀♀ ( ZJUC); Heilongjiang, Yichun, 24.VII.1994, Juxian Lou leg., 1♂ ( ZJUC). JAPAN: Shikoku, Kochi Pref., Kami-gun ( AMNH, ELKN). Australian region: AUSTRALIA: Queensland, Childers ( BPBM); Queensland, Brisbane ( BPBM); Queensland, W Gordonvale, Alley Creek, 500 m, 20.IV.1987, E.C. Dahms & G. Sarnes leg. ( QMC); Victoria, Shepparton ( DEUQ).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997) : in Australia: Liburnia sp. (near Chloriona ), Sardia sp. (= Hadeodelphax sp. ); in India: Nilaparvata lugens (Stål) , Sogata sp. , Sogatella furcifera (Horváth) : in Japan: Laodelphax striatellus (Fallén) , Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) (see remarks after the key to the Oriental species of Haplogonatopus ); in Malaysia: Sogata sp. , Sogatella furcifera (Horváth) ; in the Philippines: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) , Stenocranus sp. ; in Sri Lanka: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in China (Taiwan): Nilaparvata lugens (Stål) , Tarophagus proserpina (Kirkaldy) .The records of Cicadellidae by Chandra (1978a, 1978b, 1980a, 1980b) and He & Xu (2002) are erroneous (Chandra listed as hosts Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky)) , because Haplogonatopus apicalis parasitizes only Delphacidae .

Biology. Information on biology by Pagden (1934), Kitamura (1987a, 1987b, 1987c, 1988a, 1988b) and Kitamura & Iwami (1998). Gan et al. (2002) discovered in China that both females of H. apicalis and specimens of Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) and Laodelphax striatellus (Fallén) were infected by Wolbachia , a bacteria living in many insects. The parthenogenesis is thelytokous ( Olmi, 2000a).

Natural enemies. In Malaya ( Pagden, 1934; Olmi, 2000a) H. apicalis is parasitized by the following parasitoids: Ceraphronidae : Allomicrops (= Ceraphron ) abnormis R. Perkins ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp. In Thailand H. apicalis is parasitized by the following parasitoids: Pteromalidae : Trichomalus sp. 1 (maybe a new species reared in Taiwan, Taipei, 21.VII.1975; VIII.1975; V.1976; VIII.1976; this species was reared from cocoons of H. apicalis by K.S. Lin); Trichomalus sp. 2 (reared in Thailand, Prachin Buri, rice fields, 26.VII.1960; this species was reared from cocoons of H. apicalis by A.T. Barrion).

Distribution. China (Zhejiang, Fujian, Guangdong, Guangxi, Guizhou, Shaanxi, Hubei, Hunan, Henan, Taiwan, Jiangxi, Liaoning, Heilongjiang), Australia (Queensland, New South Wales, Victoria), Japan (Kyushu, Shikoku), India (Chattisgarh, Andhra Pradesh, Bihar, Chattisgarh, Delhi, Jharkhand, Karnataka, Madhya Pradesh, Orissa, Punjab), Malaysia (Malaya, Sarawak), Philippines (Luzon, Panay, Mindoro), Sri Lanka (Kandy), Thailand (Tung Saew, Pasang, Tung Fabad, Banmakok, Chom-Thong).

2. Haplogonatopus oratorius ( Westwood, 1833)

( Plates 104C, D , 105A, B )

Gonatopus oratorius Westwood 1833: 496 ; Richards 1935: 145. Gonatopus mayeti Kieffer in Kieffer & Marshall 1905: 103 (synonymized by Abdul-Nour 1974); type locality: Austria. Dicondylus oratorius (Westwood) : Kieffer 1907: 22. Haplogonatopus atratus Esaki & Hashimoto 1932: 25 (synonymized by Olmi 1993d); type locality: Utsunomiya ( Japan);

Kitamura 1982: 172; Olmi 1984: 1413; Chen 1988: 45; Olmi 1993d: 51. Monogonatopus oratorius (Westwood) : Richards 1939: 200; Raatikainen 1961: 131; Currado & Olmi 1972: 143; Abdul-Nour

1974: 1015. Haplogonatopus suchovi Ponomarenko 1970: 429 (synonymized by Olmi 1984); type locality: Omsk (Siberia, Russia). Haplogonatopus oratorius (Westwood) : Olmi & Currado 1979b: 39; Olmi 1984: 1409; Tussac 1988: 118; Olmi 1989a: 7; Olmi

1993d: 51; Olmi 1994: 65; Olmi 1995g: 5; Xu & He 1996b: 129; Guglielmino & Olmi 1997: 180–252; Tussac & Olmi

1998: 491; Olmi 1999a: 227; Olmi 1999b: 28; He & Xu 2002: 315; Guglielmino & Olmi 2006: 37, 50; Xu et al. 2012a: 13. Haplogonatopus katangae (Benoit) : Xu & He 1995b: 594 (misidentified); He & Xu 2002: 309 (misidentified). Haplogonatopus apicalis R. Perkins : Sugiura et al. 2004: 11 (misidentified). Haplogonatopus oratrius (sic) (Westwood): Mita et al. 2012: 50.

Description of female ( Plate 105A ). Apterous; length 2.0– 3.1 mm. Head brown or black, except anterior region of face, clypeus and mandible yellow; antenna brown, except segments 1–2 and 10 yellow; mesosoma reddish or testaceous; petiole and metasoma black; legs yellow-testaceous. Occasionally body totally black. Antenna clavate; antennal segments in following proportions: 7:5:9:5:5:5:4:4:5:8. Head excavated, shiny, weakly granulated; frontal line complete; occipital carina absent; POL = 2; OL = 2; OOL = 7; greatest breadth of posterior ocelli about as long as POL. Palpal formula 2/1. Pronotum shiny, smooth, unsculptured, not crossed by strong transverse impression. Scutum dull, hairless, sculptured by some longitudinal keels. Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface, metanotum, mesopleuron and metapleuron. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 13:2:3:8:13. Enlarged claw ( Plate 104D ) with one subapical tooth and one row of four–five lamellae. Segment 5 of protarsus ( Plate 104D ) with two rows of 6–12 lamellae; distal apex with about three–nine lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 105B ). Fully winged; length 2.2–2.5 mm. Head black, except mandible yellow-brown; antenna brown; mesosoma and metasoma black; legs brown, except tarsi testaceous. Antenna filiform; antennal segments in following proportions: 6:5:11:10:10:10:9:9:8:11; antennal segment 3 more than four times as long as broad (about 4.4). Head shiny, hairy, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli much longer than OOL (4:2); temple distinct; occiput concave. Palpal formula 2/1. Scutum dull, granulated. Notauli complete, posteriorly strongly converging, but running parallel near posterior margin of scutum. Scutellum and metanotum dull, finely punctate. Propodeum reticulate rugose, without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 104C ) long, slender, little distally broadened. Tibial spurs 1/1/2.

Material examined. Types: ♀ holotype of G. oratorius : UNITED KINGDOM: England, Surrey, Ripley ( HMO); ♀ holotype of G. mayeti : AUSTRIA ( ENSAM); ♀ lectotype (designated by Olmi, 1984) of H. atratus : JAPAN: Honshu, Tochigi Pref., Utsunomiya, Gunma, 6.VI.1931, E. Amano leg. ( ELKU); ♀ holotype of H. suchovi : RUSSIA: Western Siberia, Omsk, 1940, K.S. Suchov leg. ( ZMM) ( Ponomarenko , 1992). Paratype of H. atratus : JAPAN: same label locality as lectotype, 1♂ paralectotype ( ELKU). Paratypes of H. suchovi : same locality label as holotype, 12♀♀ and 5♂ ( ZMM). Other material. CHINA: Fujian ( Xu & He, 2003); Guangdong, Mt. Yunjishan, parasitized host collected 9.VI.2002, dryinid cocooning 11.VI.2002, dryinid adult emerged 26.VI.2002, reared from an unidentified species of Delphacid, M. Olmi leg., 1♂ ( MOLC); same locality label, parasitized host collected 9.VI.2002, 8♀♀ ( MOLC); Guangdong, Zhaoqing, 1 8.V.1998, Yufang Liu leg., 1♀ and 2♂ ( ZJUC); Guizhou, Sinan, IX.1986, VIII.1991, 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Shaanxi, Ningshan County, 6.V.2011, Chengyuan Jin leg., 1♀ and 1♂ ( SCAU); Zhejjiang, Hangzhou, Junhua He leg., 4♀♀ and 21♂ ( ZJUC); Zhejjiang, Lin'an, Junhua He leg., 15♂ ( ZJUC); Zhejjiang, Jiashan, Junhua He leg., 2♀♀ and 1♂ ( ZJUC); Zhejjiang, Shangyu, IX.1984, Haifu Wei leg., 1♀ ( ZJUC); Zhejjiang, Lishui, VI–VIII.1981, Junhua He leg., 1♀ and 1♂ ( ZJUC); Zhejiang, Wenzhou, Junhua He leg., 10♀♀ and 10♂ ( ZJUC); Zhejiang, Yiwu, 1–3.VIII.1986, Xiaoming Lou leg., 7♂ ( ZJUC). JAPAN: Ryukyu Islands, Iriomote-jima, 30.VI.2003, Sugiura leg., 1♀ ( Sugiura et al., 2004); Ogasawara (=Bonin) Islands (Mita et al., 2012). Palaearctic: H. oratorius is broadly spread in the Palaearctic, where it is known from Austria, China, France, Hungary, Italy, Japan, Lebanon, Romania, Russia (including Siberia), South Korea, Spain, Turkey and United Kingdom. In China it is known from the following locality: Inner Mongolia, Bayanzhouer, 15–22.VII.2008, Bo Qiu & Huayan Chen leg., 1♀ ( SCAU); Liaoning, Shenyang, 7.VI.1995, Juxian Lou leg., 1♀ ( ZJUC); Xinjiang, Mosuowan, 10.IX.1985, 1♀ ( ZJUC); Shandong, Weixian, V.1966, 1♀ ( ZJUC); Shandong, Shanghe, VII–VIII.2003, Jiahe Yan leg., 2♀♀ ( SCAU). Australian region: MARIANA ISLANDS: Guam, Talofofo ( BMNH); Guam, Dededo, ex cocoon on taro ( BPBM); Guam, Libugon, on taro ( AMNH).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China: Saccharosydne procerus (Metsumura) ( He & Xu, 2002) ; in France: Javesella pellucida (Fabricius) , Laodelphax striatellus (Fallén) ; in Guam ( Mariana Islands): Tarophagus proserpina (Kirkaldy) ; in Italy: Laodelphax striatellus (Fallén) , Megadelphax sordidulus (Stål) ; in Japan: Laodelphax striatellus (Fallén) ( Esaki & Hashimoto, 1932; Mita et al., 2012), Hosunka hakonensis (Matsumura) (Mita et al., 2012) , Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ( Kitamura, 1982) (see remarks after the key to the Oriental species of Haplogonatopus ); in South Korea: Laodelphax striatellus (Fallén) ; in Siberia ( Russia): Laodelphax striatellus (Fallén) ; in Turkey: Laodelphax striatellus (Fallén) , Toya propinqua (Fieber) ; in Ukraina: Laodelphax striatellus (Fallén) .

Biology. Information on the biology in Japan by Nishioka (1980), Kitamura (1982, 1983, 1985, 1986, 1987a, 1987b, 1987c, 1989a, 1989b), Yamada (1996), Yamada & Miyamoto (1998), Yamada & Kawamura (1999), Yamada & Kitashiro (2002), Yamada & Watanabe (2002), Yamada & Sugaura (2003) and Mita et al. (2012). The papers of Koyama (1989, 1991a, 1991b, 1993), Koyama et al. (1988, 1989), Koyama & Mitsuhashi (1989, 1990a, 1990b) and Abe & Koyama (1991) are of difficult interpretation, because the above authors mistook Haplogonatopus atratus Esaki & Hashimoto (= H. oratorius ) for Gonatopus lucens (Olmi) . Their data are related to G. lucens if the hosts are Cicadellidae , to H. atratus if the hosts are Delphacidae . This mistake is common to other authors. In the Oriental region, Chandra mistook Haplogonatopus apicalis R. Perkins (attacking Delphacidae ) for G. lucens (Olmi) (attacking Cicadellidae ). In the environments of Viterbo (Central Italy) H. oratorius overwinters as cocoon, rarely as larva in parasitized planthoppers. In Japan, it overwinters as larva in nymphs of L. striatellus (Mita et al., 2012) . In Japan, Kitamura (1989a) reported the presence of overwintering nymphs and a relatively high ratio of parasitism (9–34%) by H. oratorius from winter to spring in Shimane (Mita et al., 2012).

Natural enemies. According to Xu & He (2003) H. oratorius can be parasitized in China, Fujian, by Cheiloneurus exitiosus (R. Perkins) (= Echthrogonatopus lateocaudatus Xu & He, 2003 ).

Distribution. China (Fujian, Guangdong, Guizhou, Shaanxi, Zhejiang), Japan (Ryukyus, Ogasawara), in addition to Mariana Islands ( Guam) and many countries of Eastern (Inner Mongolia, Liaoning, Xinjiang, Shandong ( China); Honshu ( Japan); Siberia ( Russia); South Korea) and Western Palaearctic region (from Spain to European Russia).

18. Genus Gonatopus Ljungh, 1810

Gonatopus Ljungh 1810: 161 ; Richards 1937: 118; Olmi 1993b: 77; Olmi 1994: 66; Olmi 1995c: 787; Olmi 1995g: 5; Olmi 1996b: 182; Olmi 1998c: 72; Olmi 1998d: 177.; Olmi 1999a: 231; He & Xu 2002: 318.; Virla & Olmi 2008: 371.; Moya- Raygoza & Olmi 2010: 100.

Dicondylus Haliday in Curtis 1829 –1930: 110 (synonymized by Olmi, 1993b); type species: Dicondylus bicolor (Haliday in Curtis 1828), designated by Kieffer (1914b); Haliday 1833: 273; Walker 1837: 411; Kieffer 1914b: 72; Richards 1935: 147; Richards 1937: 118; Olmi 1984: 1374; Olmi 1993b: 78.

Labeo Haliday 1833: 273 (nom. praeocc., nec Cuvier 1817) (synonymized by Richards 1937); type species: Labeo vitripennis Haliday 1833 , by monotypy; Kieffer in Kieffer & Marshall 1905: 220; Walker 1837: 429; Dalla Torre 1898: 542.

nec Labea Blanchard 1840: 430 .

Labeola Haldeman 1842: 191 , new name for Labeo Haliday, 1833 (synonymized by Krombein 1979 with Dicondylus Haliday, 1829 –1930).

Pseudogonatopus R. Perkins 1905: 34 (synonymized by Olmi 1993b); type species: Pseudogonatopus kurandae R. Perkins, 1905 , by original designation; Kieffer in Kieffer & Marshall 1906: 488; Kieffer 1914b: 79; Richards 1939: 195; J. Perkins 1976: 32; Chandra 1980c: 192; Olmi 1984: 1191; Olmi 1993b: 78.

Paragonatopus R. Perkins 1905: 40 (synonymized by Olmi 1984); type species: Paragonatopus nigricans R. Perkins 1905 , by monotypy; Kieffer in Kieffer & Marshall 1906: 488; Kieffer 1914b: 87; Richards 1939: 195; Olmi 1979: 218.

Neogonatopus R. Perkins 1905: 42 (synonymized by Olmi 1984); type species: Neogonatopus ombrodes R. Perkins 1905 , by original designation.

Pachygonatopus R. Perkins 1905: 45 (synonymized by Olmi 1984); type species: Pachygonatopus melanias R. Perkins 1905 , by original designation.

Chalcogonatopus R. Perkins 1905: 46 (synonymized by Olmi 1984); type species: Chalcogonatopus gigas R. Perkins 1905 , by original designation.

Eugonatopus R. Perkins 1905: 46 (synonymized by Olmi 1984); type species: Chalcogonatopus (Eugonatopus) pseudochromus R. Perkins 1905 , by original designation.

Mercetia Kieffer in Kieffer & Marshall 1906: 495 (synonymized by Olmi 1996b); type species: Mercetia longicollis (Kieffer 1905) , by original designation; Olmi 1996b: 182; Kieffer 1914b: 19.

Platygonatopus Kieffer in Kieffer & Marshall 1906: 500 (synonymized by Olmi, 1984); type species: Platygonatopus planiceps (Kieffer 1904) , designated by Richards (1939).

Apterodryinus R. Perkins 1907: 14 (synonymized by Olmi 1993b); type species: Apterodryinus torvus R. Perkins 1907 , by monotypy; Olmi 1984: 1342; Olmi 1993b: 78.

Agonatopus R. Perkins 1907: 29 (synonymized by Olmi 1984); type species: Agonatopus pallidicornis R. Perkins 1907 , designated by Richards (1939).

Agonatopoides R. Perkins 1907: 33 (synonymized by Olmi 1993b); type species: Agonatopoides synchromus R. Perkins 1907 , by monotypy; Olmi 1984: 1310; Olmi 1993b: 78.

Cyrtogonatopus Kieffer 1907: 19 (synonymized by Olmi 1984); type species: Cyrtogonatopus breviforceps (Kieffer 1904) , monotypic.

nec Trichogonatopus Kieffer 1909: 336 .

Pristogonatopus Kieffer 1913c: 325 (synonymized by Olmi & Currado 1974); type species: Pristogonatopus dentatiforceps (Kieffer 1904) , by original designation; Kieffer 1914b: 71; Richards 1939: 195.

Digonatopus Kieffer 1913c: 325 (synonymized by Richards 1939); type species: Digonatopus javanus (R. Perkins 1912) , by monotypy.

Trigonatopus Kieffer 1913c: 325 (synonymized by Olmi 1984); type species: Trigonatopus bifarius (Kieffer in Kieffer & Marshall 1906), by original designation; Kieffer 1914b: 72.

Laberius Kieffer 1914b: 59 , new name for Labeo Haliday 1833 (synonymized by Richards 1937); Richards 1936: 169.

Trichogonatopus Hellén 1930: 2 (synonymized by Olmi 1984); type species: Trichogonatopus lindbergi Hellén 1930 , original designation.

Laberinus Ogloblin 1932: 266 (mispelling).

Metagonatopus Ogloblin 1932: 264 (synonymized by Olmi 1984); type species: Metagonatopus brasiliensis Ogloblin 1932 , by original designation.

Allogonatopus Haupt 1938: 27 (synonymized by Olmi 1984); type species: Allogonatopus procerus Haupt 1938 , by original designation.

Donisthorpina Richards 1939: 201 (synonymized by Olmi 1993b); type species: Donisthorpina formicicola Richards 1939 , by original designation; Olmi 1984: 1304; Olmi 1993b: 78.

Plectrogonatopus Richards 1939: 207 (synonymized by Olmi 1984); type species: Plectrogonatopus striatus (Kieffer 1905) , by original designation.

Tetrodontochelys Richards 1939: 217 (synonymized by Olmi 1993b); type species: Tetrodontochelys ljunghii ( Westwood 1833) , by original designation; Olmi 1984: 1434; Olmi 1993b: 78.

Epigonatopoides Richards 1939: 221 (synonymized by Olmi 1984); type species: Epigonatopoides pulicarius ( Klug 1810) , by original designation.

Idologonatopus Ogloblin 1953: 111 (synonymized by Olmi 1984); type species: Idologonatopus nigrithorax Ogloblin 1953 , by original designation.

Rhynchogonatopus Benoit 1954: 390 (synonymized by Olmi 1984); type species: Rhynchogonatopus meridionalis Benoit 1954 , by original designation.

Madecagonatopus Benoit 1954: 392 (synonymized by Olmi 1984); type species: Madecagonatopus ankaratrus ( Ceballos 1936) , by original designation.

Pseudogonatopoides J. Williams 1956: 90 (synonymized by Olmi 1984); type species: Pseudogonatopoides mauritianus J. Williams 1956 , by original designation.

Cyrtogonatopoides Ponomarenko 1966: 1579 (synonymized by Olmi 1984); type species: Cyrtogonatopoides borealis Ponomarenko 1966 , by original designation.

Plectrogonatopoides Ponomarenko 1975a: 318 (synonymized by Olmi 1993b); type species: Plectrogonatopoides chortinis Ponomarenko 1975 , by original designation; Olmi 1984: 1431; Olmi 1993b: 78.

Tetradontochelys J. Perkins 1976: 35 (injustified emendation).

Neogonatopoides Abdul-Nour 1976: 274 (synonymized by Olmi 1984); type species: Neogonatopoides claudiae Abdul-Nour 1976 , by monotypy.

Dolichochelys Olmi & Currado 1976: 177 (synonymized by Olmi 1984); type species: Dolichochelys longicornis (Kieffer 1904) , by original designation.

Megagonatopus Olmi & Currado 1976: 178 (synonymized by Olmi 1984); type species: Megagonatopus birmanus Olmi & Currado 1976 , by original designation.

Acrodontochelys Currado 1976: 13 (synonymized by Olmi 1993b); type species: Acrodontochelys bouceki Currado 1976 , by original designation; Olmi 1984: 1172; Olmi 1993b: 78.

Nogatopus Ponomarenko 1979: 357 (synonymized by Olmi 1984); type species: Nogatopus gobiensis Ponomarenko 1979 , by original designation.

Paradicondylus Olmi 1986: 102 (synonymized by Olmi 1998c); type species: Paradicondylus paulyi Olmi 1986 , by monotypy and original designation; Olmi 1993b: 78; Olmi 1998c: 72.

Type species. Gonatopus formicarius Ljungh, 1810 , by monotypy.

Diagnosis. Female: apterous ( Plates 106B , 126A ) or fully winged ( Plate 106A ); palpal formula 3/2, 4/2, 4/3, 5/ 2, 5/3, or 6/3; antenna with or without rhinaria; pronotum crossed or not by transverse furrow ( Plate 108C, D ); enlarged claw with distal apex pointed and with one large or small subapical tooth ( Plate 107B, C ) (occasionally subapical tooth absent, then enlarged claw with distal group of lamellae ( Plate 107A )); in fully winged forms, segment 5 of protarsus with more than 20 lamellae; tibial spurs 1/0/1. Male: fully winged ( Plate 110A, B ); occipital carina absent or incomplete (in this last case, only present behind and shortly on sides of posterior ocelli); occiput concave; temple present; palpal formula 3/2, 4/2, 4/3, 5/2, 5/3, or 6/3; tibial spurs 1/1/2.

Distribution. Worldwide.

Hosts. Acanaloniidae , Cicadellidae (except Typhlocybinae , Idiocerinae , Macropsinae ), Delphacidae , Flatidae , Issidae , Lophopidae , Tropiduchidae ( Guglielmino & Olmi, 1997, 2006, 2007).

World species. Four hundred and thirty nine species are known, fifty eight species in the Oriental region and one species incertae sedis.

Remarks. Larvae of Gonatopus were described by Ponomarenko (1975b), Carcupino et al. (1996, 1998), Guglielmino & Virla (1998), Virla & Mangione (2000), Guglielmino (2002), Mangione & Virla (2004) and Guglielmino et al. (2006, 2007, 2008). Gonatopus females were divided for convenience by Olmi (1993b) into eleven groups on the basis of the following characters:

Key to the groups of Gonatopus

1. Fully winged ( Plate 106A )....................................................................... Group 11

- Apterous ( Plate 106B )................................................................................. 2

2. Enlarged claw without subapical tooth ( Plate 107A ), or with one very small subapical tooth ( Plate 107B )............... 3

- Enlarged claw with one large subapical tooth ( Plate 107C )..................................................... 5

3. Distal apex of enlarged claw with group of lamellae ( Plate 107A )......................................... Group 8

- Distal apex of enlarged claw without lamellae ( Plate 107C, D )................................................. 4

4. Pronotum not crossed by transverse impression or slightly impressed ( Plate 108A )........................... Group 6

- Pronotum crossed by strong transverse impression ( Plate 108B ).......................................... Group 7

5. Pronotum not crossed by transverse impression or slightly impressed ( Plate 108C )................................. 6

- Pronotum crossed by strong transverse impression ( Plate 108D )................................................ 7

6. Labial palpi 3-segmented......................................................................... Group 9

- Labial palpi 2-segmented......................................................................... Group 5

7. Enlarged claw without lamellae, with or without bristles or peg-like hairs ( Plate 109B )........................ Group 1

- Enlarged claw with lamellae ( Plate 107C ).................................................................. 8

8. Labial palpi 2-segmented............................................................................... 9

- Labial palpi 3-segmented.............................................................................. 10

9. Maxillary palpi composed of two–four segments...................................................... Group 2

- Maxillary palpi 5-segmented..................................................................... Group 10

10. Maxillary palpi 6-segmented...................................................................... Group 4

- Maxillary palpi 5-segmented...................................................................... Group 3

The above groups are distributed in the zoogeographical regions as follows:

Palaearctic: groups 1, 2, 3, 4, 5, 6, 7, 10, 11.

Afrotropical: groups 1, 2, 3, 4, 5, 6, 7, 8, 11.

Oriental: groups 1, 2, 3, 4, 5, 6, 7, 10.

Nearctic: groups 1, 2, 3, 4, 5, 6, 7, 9.

Neotropical: groups 1, 2, 3, 4, 5, 6, 7, 10.

Australian: groups 1, 2, 4, 5, 6, 7, 9.

As far as, it is not possible divide also males into the above groups.

All Oriental species of Gonatopus can be attributed to one of the above eight female groups, except Gonatopus leurosipropodeus He & Xu, 2002 . The attribution of this species to a distinct group is impossible, because it was described on the basis of male specimens and its female is unknown. For this reason, it is attributed to an uncertain group (see “Uncertain group”).

Key to the males

(unknown for the species not present in the key) (males not divided into groups)

1. Notauli incomplete.................................................................................... 2

- Notauli complete..................................................................................... 4

2. Propodeum with dorsal surface dull, granulated, sculptured by irregular striae, and posterior surface shiny, smooth and unsculptured; dorsal surface of propodeum much shorter than posterior surface...................... G. maurus Kieffer

- Propodeum completely smooth and shiny, unsculptured; dorsal surface of propodeum much longer than posterior surface... 3

3. Dorsal process of paramere transverse ( Plate 122C ); clypeus testaceous.............................. G.lucens (Olmi)

- Dorsal process of paramere about parallel to penis, not transverse ( Plate 111A ); clypeus black....... G. nearcticus (Fenton)

4. Notauli posteriorly broadly separated..................................................................... 5

- Notauli posteriorly joint (or very close and almost joint)..................................................... 14

5. Dorsal process of paramere reduced to inner expansion of paramere ( Plate 122B )....... G. flavifemur (Esaki & Hashimoto)

- Dorsal process of paramere not reduced ( Plates 111B , 118D , 119D , 128A , 131A, B )................................ 6

6. Dorsal process of paramere much shorter than paramere ( Plates 128A , 131A, B )................................... 7

- Dorsal process of paramere as long as, or longer, or slightly shorter than paramere ( Plates 111B , 118D , 119D )............ 9

7. Head with POL more than four times as long as OOL; ocelli very large............................ G. rufoniger Olmi

- Head with POL less than three times as long as OOL; ocelli small............................................... 8

8. Dorsal process of paramere very slender ( Plate 131B ); clypeus completely black...................... G. superbus Olmi

- Dorsal process of paramere less slender ( Plate 128A ); clypeus medially black, laterally testaceous... G.javanus (R. Perkins)

9. Face completely or almost completely testaceous or yellow................................................... 10

- Face completely black or brown........................................................................ 11

10. Dorsal process of paramere longer than penis and paramere ( Plate 118D )......................... G. borneanus (Olmi)

- Dorsal process of paramere shorter than penis and paramere ( Plate 111B )...................... G. nigricans (R. Perkins)

11. Antennal segment 3 less than three times as long as broad; dorsal process of parameres longer than penis ( Plate 119D )............................................................................................ G. pyrillae (Mani)

- Antennal segment 3 more than three times as long as broad; dorsal process of parameres shorter than penis ( Plates 111B , 117C , 122D )............................................................................................. 12

12. Dorsal process of paramere almost completely joint to paramere ( Plate 111B ).................. G. nigricans (R. Perkins)

- Dorsal process of paramere completely separated from paramere ( Plates 117C , 122D ).............................. 13

13. Dorsal process of paramere slender ( Plate 122D )............................................. G. pedestris Dalman

- Dorsal process of paramere broader ( Plate 117C )............ G.malesiae (Olmi) type B or G.sarawakensis (Olmi) type B

14. Dorsal process of paramere very reduced ( Plates 117A , 122B ), reduced to inner expansion of paramere................ 15

- Dorsal process of paramere not reduced ( Plates 111B , 117B ).................................................. 16

15. Inner expansion of paramere with many irregular striae ( Plate 122B )................. G. flavifemur (Esaki & Hashimoto)

- Inner expansion of paramere without irregular striae ( Plate 117A )............................... G. nudus (R. Perkins)

16. Dorsal process of paramere much shorter than paramere ( Plates 111B , 131C )..................................... 17

- Dorsal process of paramere approximately as long as paramere ( Plates 117B , 131D )............................... 18

17. Dorsal process of paramere not transverse, with distal apex broad ( Plate 111B ); head and mesosoma mostly black or brown................................................................................... G. nigricans (R. Perkins)

- Dorsal process of paramere transverse, with distal apex pointed ( Plate 131C ); head and mesosoma mostly yellow-testaceous....................................................................................... G. asiaticus (Olmi)

18. Propodeum completely reticulate rugose; dorsal process of paramere broader ( Plate 117B )................................................................................ G.malesiae (Olmi) type A or G. sarawakensis (Olmi) type A

- Propodeum with posterior inclined region reticulate rugose and anterior flat region smooth and unsculptured; dorsal process of paramere slender ( Plate 131D ).................................................... G. leurosipropodeus He & Xu

Remarks. In the above key, there are two species, whose males are not distinguishable, G. malesiae (Olmi) and

G. sarawakensis (Olmi) ; the situation is explained in remarks related to G. sarawakensis (see Group 2).

Group 1

One species is known in the Oriental region.

1. Gonatopus nearcticus (Fenton, 1905)

( Plates 109A, B , 110A, B , 111A )

nec Pseudogonatopus americanus R. Perkins 1905: 37 .

Epigonatopus americanus Fenton 1921: 70 (preoccupied by Pseudogonatopus americanus R. Perkins 1905 ) (synonymized by Olmi 1993b); type locality: Ames (Iowa, USA).

Pachygonatopus nearcticus Fenton 1927: 6 .

Platygonatopus ugandanus Benoit 1951a: 300 (synonymized by Olmi 1993b); type locality: Bululo ( Uganda); Benoit 1951d: 19.

Rhynchogonatopus ugandanus (Benoit) : Olmi & Currado 1976: 173.

Acrodontochelys bouceki Currado 1976: 14 (synonymized by Olmi 1993b); type locality: Nerja ( Spain); Graham 1983: 1; Olmi 1984: 1174; Graham 1988: 79.

Acrodontochelys ugandanus (Benoit) : Olmi 1984: 1174.

Acrodontochelys sinensis Olmi 1984: 1180 (synonymized by Xu et al. 2012b); type locality: Macau ( China) .

Acrodontochelys americanus (Fenton) : Olmi 1984: 1182.

Gonatopus nearcticus (Fenton) : Olmi 1993b: 80, 83; Guglielmino & Olmi 1997: 194, 237; Tussac & Olmi 1998: 490; Olmi 1999a: 246; Olmi 2004a: 368; Olmi 2005b: 126; Olmi 2006: 44; Guglielmino & Olmi 2007: 126; Olmi 2009b: 459; Burn 2011: 88; Penati & Olmi 2012: 79; Xu et al. 2012b: 21.

Gonatopus sinensis (Olmi) : He & Xu 2002: 349.

Description of female ( Plate 109A ). Apterous; length 2.0–4.0 mm. Head black or brown, with mandible, clypeus and anterior region of face yellow or testaceous; antenna black or brown, with segments 1–3 yellow or testaceous; occasionally antenna testaceous-brown; mesosoma, petiole and metasoma usually black, except posterior apex of propodeum testaceous; occasionally mesosoma and metasoma brown-testaceous or testaceous-reddish, with petiole black; occasionally mesosoma brown, with scutum yellow; legs yellow or testaceous, with coxae and clubs of femora almost completely black, or darkened, or brown-reddish. Antenna clavate; antennal segments of holotype of P. nearcticus in following proportions: 4:4:9:5.5:5:5:4:4:4:7; antennal segments of holotype of A. sinensis in following proportions: 5:3:7:4:4:4:4:4:3.5:5. Head flat or slightly convex, dull, weakly or strongly granulated; frontal line complete; occipital carina absent; temple distinct; POL = 3; OL = 2; OOL = 5. Palpal formula 5/2, 4/2, 3/2, or 2/2. Pronotum crossed by strong transverse impression, shiny, weakly or strongly granulated. Scutum dull, granulated. Metanotum granulated and transversely striate, not hollow behind scutellum. Metathorax + propodeum dull, granulated, with posterior surface of propodeum transversely striate; occasionally posterior surface almost completely granulated and not transversely striate. Meso-metapleural suture obsolete. Mesopleuron and metapleuron granulated, with or without transverse striae. Protarsus with first segment about twice as long as segment 4 (14:7; or 15:7; or 12:7). Protarsal segments of A. sinensis holotype in following proportions: 15:2:3:7:13. Enlarged claw ( Plate 109B ) with one large subapical tooth and one row of three–six peg-like bristles. Segment 5 of protarsus ( Plate 109B ) with two rows of 6–22 lamellae; distal apex with about six–nine lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 110A, B ): fully winged; length 1.6–2.8 mm. Head black or brown, except mandible testaceous or yellow; antenna brown or black; mesosoma and metasoma black or brown; legs banded, brown or black, with trochanters, stalks of femora, extremities of tibiae and tarsi yellow. Antenna hairy, filiform; antennal segments of male paratype of A. bouceki in following proportions: 5:4:8:7:7:7:6:6:6:8; antennal segment 3 four or more than four times as long as broad. Antennal segments of specimen from China, Ruyang, in following proportions: 4:4:7:7:6.5:6.5:5.5:5:5:7. Head shiny, alutaceous, slightly punctate; vertex laterally without two shiny and smooth areas situated between posterior ocelli and eyes; frontal line incomplete, only present between antennal toruli; occipital carina absent; occiput concave; temple distinct; POL = 6; OL = 4; OOL = 2 (or POL = 6; OL = 2; OOL = 2; or POL = 7; OL = 3; OOL = 1.5); greatest breadth of posterior ocelli about twice as long as OOL. Palpal formula 3/2, 4/2. Scutum dull, hairy, slightly granulated. Notauli incomplete, reaching approximately 0.4–0.7 length of scutum. Scutellum and metanotum shiny, smooth, finely punctate, unsculptured among punctures. Propodeum completely shiny, smooth, unsculptured; dorsal surface of propodeum with or without median longitudinal furrow. Fore wing hyaline, without dark transverse bands; stigmal vein regularly curved, with distal part much longer than proximal part. Dorsal process of paramere ( Plate 111A ) more or less long and slender, with distal third parallel to penis. Tibial spurs 1/1/2.

Material examined. Types: ♀ holotype of E. americanus : USA: Iowa, Story County, Ames, 25.VII.1919, E.D. Ball leg. ( USNM); ♀ holotype of P. nearcticus : USA: Woodbury County, Sioux City, IX.1919, reared from a jassid, C.N. Ainslie leg. ( USNM); ♀ holotype of P. ugandanus : UGANDA: Bululo (=present Bulule?), 1909, E. Bayon leg. ( MCSNG); ♀ holotype of A. bouceki : SPAIN: Malaga, Nerja, 3.VII.1974, Z. Bouček leg. ( BMNH); ♀ holotype of A. sinensis : CHINA: Macau ( BPBM). Paratypes of E. americanus : same locality label as holotype, 4♀♀ ( USNM). Paratypes of A. bouceki : same locality label as holoype, 23.VI.1973, Z. Bouček leg., 14♀♀ ( BMNH, DIVAPRA, MRSNT, USNM); SPAIN: Granada, La Herradura, 24.VI.1973, Z. Bouček leg., 1♀ and 1♂ ( BMNH); SPAIN: Granada, Calahonda, 22.VI.1973, Z. Bouček leg., 2♀♀ ( BMNH); SPAIN: Barcelona, Calella de la Costa, VI.1974, 1♀ ( BMNH); SPAIN: Castellón, Benicasim, 13–15.VI.1973, 22–24.VI.1974, Z. Bouček leg., 1♀ and 1♂ ( BMNH). Paratypes of A. sinensis : same locality label as holotype, 3♀♀ ( AMNH, BPBM); CHINA: Chin San, 1♀ ( BPBM); VIET NAM: Lam Dong Prov., Di Linh, Plateau of Djiring, 1♀ ( AMNH). Other material. CHINA: Guangdong, Mt. Nankunshan, 8.VI.2002, M. Olmi leg., 1♀ ( MOLC); Foot of Nanling National Nature Reserve, Ruyang, reared from adult of unidentified Cicadellid, M. Olmi reared, parasitized host collect 12.IX.2004, dryinid cocooning 13.IX.2004, dryinid adult emerged X.2004, 1♀ and 3♂ ( MOLC); same locality label, parasitized host collect 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 2♂ ( MOLC); Macau, X.1906, R. Perkins collection, 1♀ and 1♂ ( USNM). INDIA: Karnataka, Bangalore, 3.XI.1979, Z. Bouček leg., 1♀ ( BMNH); SINGAPORE ( NMNH). THAILAND: Chumphon Prov., Chumplion, 9.II.2000, Ralf Jochmann leg., 1♀ ( MTC); Mae Hong Son Prov., Soppong, 19°27'N 98°20'E, 1500 m, 7–12.V.1996, S. Becvar leg., 1♀ ( OLL). Palaearctic: G. nearcticus is known from the following countries: Croatia, Cyprus, France, Greece, Italy, Montenegro, Portugal, Spain (including Canary Islands) and Turkey ( Olmi, 1999a). Afrotropical region: G. nearcticus is known from the following countries: Benin, Botswana, Ethiopia, Kenya, Mozambique, Namibia, South Africa, Swaziland, Uganda, Zimbabwe ( Olmi, 1984, 1999a). Nearctic region: G. nearcticus is known from the USA ( Olmi, 1984, 1999a).

Hosts. Cicadellidae ( Guglielmino & Olmi, 1997, 2007): in France: Balclutha frontalis (Ferrari) ; in Italy: Balclutha rosea (Scott) ; in Namibia: Paradorydium spatulatum (Naudé) ; in South Africa: Balclutha rosea (Scott) ; in the USA: Balclutha neglecta (DeLong & Davidson) , Balclutha impicta (Van Duzee) .

Distribution. China ( Macau, Guangdong), India (Karnataka), Thailand (Chumphon, Mae Hong Son), Viet Nam (Lam Dong), in addition to many countries of the Afrotropical, Nearctic and Western Palaearctic regions.

Group 2

Ten species are known in the Oriental region.

Key to the females

1. Mesosoma totally black or black-brown (at most distal apex of propodeum and posterior margin of pronotum testaceous or reddish)............................................................................................... 2

- Mesosoma at least partly testaceous-reddish, or testaceous-ferruginous, or brown-reddish, or yellow................... 4

2. Metanotum with sides protruding; protrusions rounded ( Plate 113G )............................. G. nepalensis (Olmi)

- Metanotum with sides rounded ( Plates 112A , 114A ).......................................................... 3

3. Species large (body length about 5.0 mm); body covered with long sparse hairs ( Plate 112A ); temple absent ( Plate 112A ); head more excavated....................................................................... G. capillus Xu & He

- Species small (body length less than 4.5 mm); body hairless or covered with very short hairs; temple distinct ( Plate 114A ); head less excavated ( Plate 114B )...................................................... G. nigricans (R. Perkins)

4. Metanotum with sides rounded ( Plates 114A , 115A ); meso-metapleural suture obsolete.............................. 5

- Metanotum with sides protruding ( Plate 117G ); protrusions rounded ( Plates 111C , 117G ) or pointed ( Plate 113C ); meso-metapleural suture at least partly distinct....................................................................... 6

5. Metanotum short, about as long as scutellum............................................... G. nudus (R. Perkins)

- Metanotum long, about twice as long as scutellum........................................ G. nigricans (R. Perkins)

6. Anterior surface of metathorax + propodeum with numerous longitudinal keels.................. G. sarawakensis (Olmi)

- Anterior surface of metathorax + propodeum smooth, with no keels............................................. 7

7. Metathorax + propodeum with slight or strong median furrow.................................................. 8

- Metathorax + propodeum without median furrow............................................................ 9

8. Metanotum with lateral protrusions more prominent ( Plate 113A ); metathorax + propodeum with deep median furrow; mesometapleural suture very strong and distinct............................................. G.lankae (Ponomarenko)

- Metanotum with lateral protrusions less prominent ( Plate 113E ); metathorax + propodeum with shallow median furrow; mesometapleural suture slightly distinct, only visible near lateral protrusions of metanotum.................. G. muiri (Olmi)

9. Metanotum with lateral protrusions less prominent ( Plate 117G )................................... G. validus (Olmi)

- Metanotum with lateral protrusions more prominent ( Plates 111C , 113C )....................................... 10

10. Lateral protrusions of metanotum at right angle ( Plate 111C ); mesopleuron and metapleuron transversely striate................................................................................................. G.bicuspis (Olmi)

- Lateral protrusions of metanotum not at right angle ( Plate 113C ); mesopleuron and metapleuron smooth, not transversely striate................................................................................ G. malesiae (Olmi)

1. Gonatopus bicuspis (Olmi, 1993) , comb. nov.

( Plate 111C, D )

Pseudogonatopus bicuspis Olmi 1993d: 50 .

Description of Female. Apterous; length 3.2 mm. Testaceous-ferruginous, except petiole black, antennal segment 8 testaceous and antennal segments 9–10 whitish. Antenna clavate; antennal segments in following proportions: 10:6:15:8:6:6:5.5:5:5:8.5. Head very excavated, shiny, smooth, unsculptured; frontal line absent; occipital carina absent; temple absent; POL = 0.3; OL = 3; OOL = 6. Palpal formula 3/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Metanotum flat, transversely striate, with two lateral pointed apophyses ( Plate 111C ). Metathorax + propodeum shiny, smooth, unsculptured; posterior surface of propodeum transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 13:3:4.5:15.5:25. Enlarged claw ( Plate 111D ) with one large subapical tooth and one row of six lamellae + one bristle. Segment 5 of protarsus ( Plate 111D ) with two rows of 16 lamellae; distal apex with about 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: PHILIPPINES: Negros Island , Camp, 16.V.1987 ( AMNH) .

Hosts. Unknown.

Distribution. Philippines (Negros).

2. Gonatopus capillus Xu & He, 1998

( Plate 112A, B )

Gonatopus capillus Xu & He 1998d: 306 ; He & Xu 2002: 351; Guglielmino & Olmi 2006: 49.

Description of female ( Plate 112A ). Apterous; length 5.0 mm. Head black, except mandible testaceous; antenna black, except segment 1 partly testaceous and partly brown and segment 10 yellow-whitish; mesosoma, petiole, metasoma and legs black. Body completely covered with long hairs (as in some species of Trichogonatopus Kieffer ). Antenna clavate; antennal segments in following proportions: 6:3.6:14.5:9:8.5:8:6:6:6:8. Head shiny, smooth, unsculptured, except ocellar region granulated; frontal line incomplete; occipital carina absent; temple absent; POL = 1; OL = 0.5; OOL = 5. Palpal formula 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum and metanotum without lateral pointed apophyses. Metanotum flat, not hollow behind scutellum. Metathorax + propodeum with anterior region smooth, disc reticulate rugose and posterior region transversely striate; anterior region and disc with track of long median longitudinal furrow. Mesopleura and metapleura transversely striate. Meso-metapleural suture almost completely absent, only present on sides of metanotum. Fore tarsal segments in following proportions: 10:1.5:3:10:15.5. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 112B ) with one large subapical tooth and one row of 15 lamellae. Segment 5 of protarsus ( Plate 112B ) with two rows of 12 + 14 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: CHINA: Guizhou, Sinan , IX.1986, Yuxiang Chen leg. ( ZJUC) . Paratype: same label as holotype, 1♀ ( ZJUC) .

Hosts. Delphacidae ( Guglielmino & Olmi, 2006) : in China: Nilaparvata lugens (Stål) ( Xu & He, 1998d) .

Distribution. China (Guizhou).

3. Gonatopus lankae (Ponomarenko, 1981) , comb. nov.

( Plate 113A, B )

Pseudogonatopus lankae Ponomarenko 1981b: 881 ; Olmi 1984: 1249.

Description of Female. Apterous; length 3.2–3.3 mm. Brown-reddish or testaceous-dark; antenna usually darkened, except segment 10 yellow; legs testaceous, except trochanters and procoxa partly whitish, hind side of metafemur whitish, distal apex of propodeum whitish. Antenna clavate; antennal segments in following proportions: 14:8:14:9:9:6:6:6:6:11. Head excavated, dull, alutaceous or weakly granulated; frontal line complete; occipital carina absent; temple prominent; POL = 2; OL = 3; OOL = 10. Palpal formula 4/2 or 5/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Scutellum shiny, smooth, unsculptured. Metanotum flat, shiny, unsculptured, weakly hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded. Metathorax + propodeum ( Plate 113A ) shiny, smooth, unsculptured or granulated; posterior surface of propodeum transversely striate. Disc of metathorax + propodeum with strong median furrow. Mesopleura and metapleura dull, weakly granulated, not transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 15:4:6:20:31. Enlarged claw ( Plate 113B ) with one large subapical tooth and one row of six–eight lamellae. Segment 5 of protarsus ( Plate 113B ) with two rows of 19–21 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Ratnapura District , Gilimale Induruwa Jungle, 13.III.1979, K. Krombein leg. ( USNM) . Other material. SRI LANKA: Ratnapura District ., Sinharaja Jungle, Waturawa Forest ( AMNH) ; Matale District , Sigiriya, 07°57'N 80°46'E, 18.III.1999, C.J. Burwell leg. ( QMC) .

Hosts. Unknown.

Distribution. Sri Lanka (Ratnapura, Matale).

Remarks. The female specimen from Sri Lanka, 07°57'N 80°46'E, has 5-segmented maxillary palps; the palpal formula 5/2 is typical of the group 10 (former genus Donisthorpina Richards ); for this reason, G. lankae is also included into group 10.

4. Gonatopus malesiae ( Olmi, 1984) , comb. nov.

( Plate 113C, D )

Pseudogonatopus malesiae Olmi 1984: 1248 .

Description of Female. Apterous; length 2.8–3.6 mm. Usually testaceous-ferruginous, with petiole black; antenna usually testaceous, with segments 9–10 whitish (occasionally antennal segment 9 partly whitish). In specimen from Myanmar, 17°38.9'N 94°38.1'E, antenna testaceous-brown, except segments 1 and 10 whitish. In specimen from China, Mt. Jinggangshan, antenna brown, except rhinaria areas testaceous, and mesosoma brown-testaceous. Antenna clavate; antennal segments of holotype in following proportions: 16:13:38:23:18:12:12:13:12:20. Head very excavated, shiny, smooth, unsculptured, hairless; frontal line complete; occipital carina absent; POL = 0.3; OL = 2; OOL = 8.5. Palpal formula 3/2 or 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat, without lateral pointed apophyses. Metanotum shiny, unsculptured, not hollow behind scutellum, with sides protruding; lateral protrusions pointed ( Plate 113C ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum without median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture very slightly distinct. In specimen from Myanmar, 17°38.9'N 94°38.1'E, meso-metapleural suture partly and slightly visible (not visible on sides of metanotum). Segment 1 of protarsus usually shorter than segment 4 (13:16). Enlarged claw ( Plate 113D ) with one large subapical tooth and one row of seven–ten lamellae. Segment 5 of protarsus ( Plate 113D ) with two rows of 12–17 lamellae; distal apex with at least 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown (see Gonatopus sarawakensis (Olmi)) .

Material examined. Type: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu National Park , 5.VIII.1978, P.M. Hammond & J.E. Marshall leg. ( BMNH) . Other material. CHINA: Jiangxi, Mt. Jinggangshan , 13.VIII.2007, Junhua He leg., 1♀ ( SCAU) . MALAYSIA: Malaya, Fraser's Hill , 4200' ( AMNH) . MYANMAR: Rakhine State, NE of Gwa, Elephant Camp in Elephant Sanctuary , 17°38.9'N 94°38.1'E, 3.XII.2000, leaf litter in secondary forest, K.J. Ribardo leg., 2♀♀ ( CAS) .

Hosts. Unknown.

Distribution. China (Jiangxi), Malaysia ( Sarawak, Malaya), Myanmar (Rakhine).

5. Gonatopus muiri ( Olmi, 1984)

( Plate 113E, F )

Pseudogonatopus muiri Olmi 1984: 1251 ; Xu et al. 2011g (Suppl.): 20.

Description of Female. Apterous; length 2.5–3.0 mm. Holotype testaceous–reddish, except metasoma brown and antenna brown, with segments 1–2 testaceous (specimen from China, Bawangling National Nature Reserve, testaceous, except antenna brown, with segments 1–2 testaceous and segment 10 whitish, petiole black and metasoma brown). Antenna clavate; antennal segments of a specimen from China, Bawangling National Nature Reserve, in following proportions: 8:4.5:10:6:5:5:4.5:4.5:4:8. Head excavated; head of holotype dull and slightly granulated; head of specimen from China, Bawangling National Nature Reserve, shiny and unsculptured; frontal line of specimen from China, Bawangling National Nature Reserve, complete; occipital carina absent; head of specimen from China, Bawangling National Nature Reserve, with POL = 0.5; OL = 1.5; OOL = 6. Palpal formula 3/ 2 in holotype, 4/ 2 in specimen from China, Bawangling National Nature Reserve. Pronotum crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat. Metanotum shiny, unsculptured, not hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded ( Plate 113E ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum with slight median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture weakly distinct, in holotype only visible near lateral protrusions of metanotum. Meso-metapleural suture of specimen from China, Bawangling National Nature Reserve, visible only between mesopleuron and metapleuron, not visible near lateral protrusions of metanotum. Protarsal segments of specimen from China, Bawangling National Nature Reserve, in following proportions: 10:2.5:4:10:17; in holotype segment 1 of protarsus as long as segment 4. Enlarged claw ( Plate 113F ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 113F ) with two rows of 12–17 lamellae; distal apex with 11–12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Java, Pekalongan, F. Muir leg. ( AMNH) . Other material. CHINA: Hainan, Bawangling National Nature Reserve, Reserve House , about 1000 m, parasitized host collected 19.VIII.2000, dryinid cocooning 22.VIII.2000, M. Olmi reared from an unidentified species of Delphacidae , 1♀ ( MOLC) .

Hosts. In China (Hainan), unidentified species of Delphacidae .

Distribution. China (Hainan), Indonesia (Java).

6. Gonatopus nepalensis ( Olmi, 1986) , comb. nov.

( Plate 113G, H )

Pseudogonatopus nepalensis Olmi 1986: 80 .

Description of Female. Apterous; length 4.8–5.0 mm. Head black or brown, except mandible, clypeus and anterior region of face testaceous; antenna brown, except segments 1–2 testaceous and 10 whitish; occasionally antennal segment 9 whitish; mesosoma and metasoma black; legs black or brown, except chela and meso- and metatarsus testaceous. Antenna clavate; antennal segments in following proportions: 12:6:18:10:7:7:7:6.5:6.5:10.5. Head excavated, dull, granulated; frontal line complete; occipital carina absent. Palpal formula 4/2. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum shiny, sculptured by longitudinal striae. Metanotum transversely striate, with sides protruding; protrusions rounded ( Plate 113G ). Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface and metapleura. Mesopleuron shiny, smooth, unsculptured, not transversely striate. Meso-metapleural suture partly distinct, only present between mesopleuron and metapleuron, not present dorsally on sides of metanotum. Protarsal segments in following proportions: 18:4:8:22:35. Enlarged claw ( Plate 113G ) with one large subapical tooth and one row of seven lamellae. Segment 5 of protarsus ( Plate 113G ) with two rows of about 21 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: NEPAL: Kosi, S of Mangsingma Forest ( MHNG). Paratypes: same locality label as holotype, 1♀ ( AMNH); E Nepal, Kosi, Induwa Kola Valley, 1♀ ( MHNG). Other material. NEPAL: Khala Valley, 2050 m ( PMA). Palaearctic: according to Ponomarenko (1992), G. nepalensis is also present in the following localities: RUSSIA: Transbaikalia, 5.VI.1963, 10.VII.1963, A. Emelianov leg., 2♀♀ ( Ponomarenko , 1992); Fra East, Primorskiy Krai, Gornotayezhnoe, 19.VI.1983, Sciarkov leg., 1♀ ( Ponomarenko , 1992). This quotation is doubtful.

Hosts. Unknown.

Distribution. Nepal (Kosi), Russia (Transbaicalia, Primorskiy Krai).

7. Gonatopus nigricans (R. Perkins, 1905)

( Plates 111B , 114A – D )

Paragonatopus nigricans R. Perkins 1905: 41 ; Kieffer 1914b: 87; Olmi 1979: 218.

Pseudogonatopus melanacrias R. Perkins 1906a: 487 (synonymized by R. Perkins 1912); type locality: Suva ( Viti Levu Island,

Fiji Islands). Gonatopus fulgori Nakagawa 1906: 163 ; type locality: Japan; Guglielmino & Olmi 1997: 181, 184; syn. nov. Pseudogonatopus hospes R. Perkins 1912: 13 (synonymized by He & Xu 2002); type locality: China; Kieffer 1914b: 81;

Swezey 1928: 670; F. Williams 1931: 247; Chiu 1979: 339; Móczár 1979: 77; Olmi 1984: 1290; Olmi 2000a: 101, 111. Gonatopus sauteri Strand 1913: 209 (synonymized by Olmi 1984); type locality: Tainan ( Taiwan, China). Dicondylus sauteri (Strand) : Kieffer 1914b: 77. Pseudogonatopus sogatea Rohwer 1920: 160 (printing-error; holotype labelled sogatae ); type locality: Pusa (Bihar, India);

Kurian 1954b: 417; Olmi 1984: 1254; syn. nov. Haplogonatopus fulgori (Nakagawa) : Esaki & Hashimoto 1930: 21. Pseudozonatopus (sic) hospes R. Perkins : Pagden 1934: 2. Paragonatopus fulgori (Nakagawa) : Mochizuki 1943: 99; Chen 1988: 44. nec Pseudogonatopus nr. hospes R. Perkins : Chandra 1978b: 5. Pseudogonatopus fulgori (Nakagawa) : Olmi 1984: 1194. Pseudogonatopus pusanus Olmi 1984: 1255 ; type locality: Pusa (Bihar, India); He & Xu 2002: 355; syn. nov. Pseudogonatopus nigricans (R. Perkins) : Olmi 1984: 1287; Olmi 1990b: 789; Olmi 1991: 319. Gonatopus nigricans (R. Perkins) : Guglielmino & Olmi 1997: 165, 224–226, 250–252; Olmi 1999b: 28; Olmi 2000a: 94; He &

Xu 2002: 364; Guglielmino & Olmi 2006: 49. Gonatopus sogatae Rohwer : Guglielmino & Olmi 1997: 225, 226; He & Xu 2002: 355. Gonatopus insulanus He & Xu in He et al. 1998: 172; type locality: Qionghai (Hainan, China); syn. nov.

Description of female ( Plate 114A ): apterous; length 2.4–4.4 mm. Head brown-reddish, or brown, or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 or 1–3 yellow; mesosoma black or brown; occasionally posterior margin of pronotum and posterior apex of propodeum testaceous; metasoma black or brown; legs yellow; occasionally clubs of femora and part of coxae brown. In G. pusanus the colour is different: head partly or almost totally brown; antenna brown, except segments 1–2 testaceous; mesosoma testaceous-reddish, except sides of scutum darkened; occasionally part of pronotum darkened; metasoma brown; petiole black; legs testaceous; occasionally club of profemur darkened. In G. sogatae the colour is the following: head brown, except mandible and clypeus yellow; antenna brown, except segment 1 yellow and 2–3 brown-testaceous; mesosoma brown, except scutum reddish and distal apex of propodeum yellow; petiole black; metasoma brown-dark; legs brown-yellow. Intermediate colours are known. Antenna clavate; antennal segments in following proportions: 19:19:52:25:22:22:18:17:15:26. Head ( Plate 114B ) excavated, shiny, granulated or unsculptured; occiput and anterior region of face always granulated; POL = 5; OL = 5; OOL = 14. Palpal formula 2/2, 3/2, 4/2. Pronotum granulated, crossed by strong transverse impression. Scutum granulated, without lateral pointed apophyses. Mesopleura transversely striate. Metanotum flat, very long, about as long as scutum, transversely striate, not hollow behind scutellum ( Plate 114C ), with sides rounded ( Plate 114A ). Metathorax + propodeum shiny, indistinctly sculptured, with or without track of median longitudinal furrow, with posterior surface transversely striate. Metapleuron transversely striate. Meso-metapleural suture obsolete. Protarsal segment 4 shorter than segment 1 (40:47; or 20:27). Enlarged claw ( Plate 114D ) with one large subapical tooth and one row of three–nine lamellae. Segment 5 of protarsus ( Plate 114D ) with two rows of 9–25 lamellae; distal apex with 4–10 lamellae; occasionally lamellae extended continuously to distal apex. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9–2.4 mm. Head brown or black (occasionally brown-reddish), except mandible testaceous; occasionally part of face testaceous; antenna brown (in type of G. insulanus , testaceous-dark); mesosoma and metasoma black or brown (in type of G. insulanus , mesosoma reddish-dark, with propodeum and pleura darker); legs yellow. Antenna filiform; antennal segments in following proportions: 5:5:9:8:8:9:7:8:7:10 (in other specimens 6:5.5:10:9:8:9:8:8:7:10); antennal segment 3 more than three times as long as broad (4.0; 4.3; 5.0). Head shiny, hairy, granulated; frontal line absent; temple distinct; occipital carina absent; POL = 8; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OL (4:3) (in other specimens POL = 7; OL = 3; OOL = 3.5; greatest breadth of posterior ocelli longer than OL (3.5:3)). Palpal formula 2/2, 3/2, 4/2. Scutum shiny, granulated. Notauli complete, posteriorly joint or almost joint in small specimens, separated in large specimens; in very large specimens, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2.5:4; or 2:4). Scutellum and metanotum shiny, finely punctate, unsculptured among punctures. Propodeum reticulate rugose; dorsal surface with median longitudinal furrow and occasionally with two lateral smooth areas. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 111B ) much shorter than paramere. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nigricans : AUSTRALIA: Queensland, Bundaberg, 19.XI.1904, reared ( BPBM); ♀ lectotype (designated by Olmi, 1982) of P. melanacrias : FIJI ISLANDS: Viti Levu Island, Suva, III.1905 , reared from delphacid near Stenocranus , on Zoysia pungens (BPBM) ; ♀ lectotype (designated by Olmi, 1982) of P. hospes : CHINA: no localities, F. Muir leg., reared in Honolulu from Perkinsiella saccharicida from Chinese stock, IV–V.1907 ( BPBM); ♀ holotype of G. sauteri : CHINA: Taiwan, Tainan, IV.1912, H. Sauter leg. ( DEI); ♀ holotype of P. pusanus : INDIA: Bihar, Pusa ( IARI); ♀ holotype of P. sogatae : INDIA: Bihar, Pusa, 6.VIII.1915, reared from Sogata sp. , C.S. Misra leg. ( USNM); ♂ holotype of G. insulanus : CHINA: Hainan, Qionghai, 4.V.1964, Junhua He leg. ( ZJUC). Type material of G. fulgori probably lost. Paratypes of P. nigricans : AUSTRALIA: same locality label as lectotype, 1♀ lebelled for mistake Fiji Islands ( BPBM); same locality label as holotype, parasitized host collected 1.X.1904, dryinid adult emerged 21.X.1904, R. Perkins leg., 1♀ “ paratype ” identified by the same R. Perkins ( BMNH); AUSTRALIA: Queensland, Bundaberg, Sandhills, X.1904, bred, 1♀ ( ANIC). Paratypes of P. melanacrias : same locality label as lectotype, 2♀♀ ( BPBM). Paratypes of P. hospes : same locality label as lectotype, 3♀♀ ( BPBM). Paratype of P. pusanus : same locality label as holotype, 1♀ ( IARI). Paratype of P. sogatae : same locality label as holotype, misidentified ♂ allotype not belonging to the same species of holotype. Other material. CHINA: Fujian, Shaxian, reared from rice planthoppers, 1979–1983, Changming Liu leg., 5♀♀ and 1♂ ( ZJUC); Fujian, IV.1987, 31.VII.1985, Changming Liu leg., 6♀♀ and 10♂ ( ZJUC); Guangdong, Yangjiang, 1979, 1981, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangdong, Xingfeng County, Mt. Yunjinshan, 9.VI.2002, 18.VII.2003, 19.VII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); same locality label, 20.VII.2003, Ping Li leg., 2♀♀ ( ZJUC); same locality label, Zaifu Xu leg., many ♀♀ ( ZJUC, AMNH); Guangdong, Shixing County, Chebaling National Nature Reserve, 21.VIII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, M. Olmi reared from adult of unidentified Delphacid, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); Guangdong, Mt. Nankunshan, 2.VII.2005, Zaifu Xu leg., 1♀ ( ZJUC); Guangdong, Yangjiang, Hailing Town, 1980, Renhuan Hou leg., 2♀♀ and 2♂ ( ZJUC); Guangdong, Huaiji County, 21–22.X.2007, Zaifu Xu leg., 1♀ ( SCAU); Guangdong, Conghua, 28.VI.1978, Junhua He leg., 1♀ ( SCAU); Guangdong, Zhaoqing, V.1998, Yufang Liu leg., 2♀♀ ( SCAU); Guizhou, Sinan, IX.1986, VIII.1991, 5♀♀ ( ZJUC); Guizhou, Sandu, VI–VII.1981, Dongxiang Xie leg., 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Hunan, Yueyang, 15.VIII.1975, Binsen Wang leg., 1♀ ( ZJUC); Guangxi, Luchuan, 30.VII.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Bama, 1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Luocheng, 18.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Yaxian, VI.1966, Junhua He leg., 1♀ ( ZJUC); Hong Kong, Tai Lung Farm, IX–XI.1965, swept above rice, I.W.B. Thornton leg., 1♀ ( BMNH); Macau ( BPBM); same locality label, R. Perkins collection, 2♀♀ ( USNM); Taiwan, Kaohsiung, Kosen ( AMNH, ELKU); Taiwan, Changhus, Erlin ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, Kungkuan ( TARI); Taiwan, Taihoku Pref. (=Taipei), 27.IX.1930, J. Sonan leg., 1♀ ( TARI); Taiwan, Tainan, ShinShi, 11.IX.1979, paddy field, Hirashima leg., 1♀ ( ELKU); Taiwan, Gakko ( ELKU); Itahashi (= Taiwan), 16.IX.1930, J. Sonan leg., 5♀♀ ( TARI); Takeyama (=Jhushan), 6.X.1937, J. Sonan leg., 3♀♀ ( TARI); Inrin (=Yuanlin), 8.X.1937, J. Sonan leg., 39♀♀ ( TARI). INDIA: Chhattisgarh, Raipur ( GCC, AMNH); Delhi, New Delhi ( IARI); Orissa, Cuttack, 1981, 1♀ ( BMNH); Orissa, Puri, 28.XI.1987, S.V. Fowler leg., 1♀ ( BMNH); Tamil Nadu, Tanjavur (=Tanjore) District, 28.IX.1987, ex Nilaparvata lugens on rice, C. Peter leg., 3♀♀ ( BMNH); West Bengal, Hooghly District, Khanakul, 12.XII.1986, in paddy seedbed, 1♀ ( BMNH). INDONESIA: Java, Pekalongan ( BPBM); Java, Dieng Plateau ( BPBM); Borneo, Kalimantan, Paloh ( BMNH). MALAYSIA: Malaya, Selangor, 13.VIII.1928, H.T. Pagden leg., 1♀ ( BMNH); Malaya, Selangor, Kuala Lumpur, 21.XII.1930, H. T. Pagden leg., 1♀ ( BMNH); same locality label, many ♀♀ and ♂ ( NMNH, KYC); Malaya, Seberang Perai ( MARDI); Malaya, Serdang ( DBUS); Sabah, Tuaran ( KYC); Sarawak, 1st. Division, Payah Paloh, 28.I.1965, G.H.L. Rotschild leg., 1♀ ( BMNH). THAILAND: Songkhla, Hadd Tai ( KYC); Non Bua Lampoo, Na Kam Hai ( KYC); Tung Saew ( KYC); Tung Fabad ( KYC); Chainat ( KYC, AMNH); Banmakok ( KYC); Mae Tang ( KYC); Lampamg ( ELKU); Mae-Theng ( ELKU); Hot ( ELKU); Chieng Dao ( ELKU); Thailand, 15.IX.1999, reared from Nilaparvata lugens (Stål) , Sawai leg., 4♂ ( BMNH). Australian region: G. nigricans is known from Australia (Queensland), Fiji Islands, Solomon Islands and USA (Hawaii) ( Olmi, 1984). Introduced in the Hawaii 1906 and 1907 from China, Guangdong, for biological control of Perkinsiella saccharicida Kirkaldy in sugarcane fields ( Olmi, 2000a).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in Australia: Liburnia sp. , Stenocranus sp. ; in China: Laodelphax striatellus (Fallén) ( Yang et al., 1982; He & Xu, 2002), Perkinsiella saccharicida Kirkaldy ; in Malaysia: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in Thailand: Nilaparvata lugens (Stål) ( Chiu, 1979) ; in the USA (Hawaii): Perkinsiella saccharicida Kirkaldy (introduced for biological control).

Biology. Information by Pagden (1934); the parthenogenesis is thelytokous.

Natural enemies. In the USA (Hawaii) ( Swezey, 1919; Timberlake, 1919; Muir, 1921; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Encyrtidae : Helegonatopus pseudophanes R. Perkins, 1906 ; Cheiloneurus flaccus (Walker 1847) ( Guerrieri & Viggiani, 2005; Guerrieri, 2006); Aphelinidae : Centrodora xiphidii (R. Perkins, 1906) . In Malaya ( Pagden, 1934; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Ceraphronidae : Allomicrops abnormis (R. Perkins, 1910) ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp.

Distribution. China ( Taiwan, Hainan, Fujian, Guangdong, Guizhou, Hubei, Hunan, Guangxi, Hong Kong, Macau), Australia (Queensland), Fiji Islands ( Viti Levu), India (Bihar, Chhattisgarh, Delhi, Orissa, Tamil Nadu), Indonesia (Java, Pekalongan, Borneo), Malaysia (Malaya, Sabah, Sarawak), Thailand (Songkhla, Non Bua Lampoo, Tung Saew, Tung Fabad, Chainat, Banmakok, Mae Tang, Lampamg, Mae-Theng, Chieng Dao).

Remarks. Gonatopus insulanus He & Xu is here synonymized with G. nigricans , because of the study of its type and comparison with male specimens of G. nigricans . There are no differences.

Gonatopus nigricans (= Pseudogonatopus hospes R. Perkins ) was established in the Hawaii from cocoons sent from China ( Perkins, 1912). This species was considered extinct in the Hawaii ( Olmi, 1999a). However, in 2010 one of the authors (Olmi) saw a male specimen collected in Oahu Island, Mt. Olympus, in April 1981 by a pan trap. Probably so this species is always present at least in Oahu Island, although rare. Gonatopus pusanus and G. sogatae are here synonymized with G. nigricans because the only differences concern the colour of mesosoma (black in nigricans , testaceous to reddish, with some darkened areas, in pusanus ; brown, with some areas reddish or yellow, in sogatae ). In the collections, there are also many specimens with intermediate colours, so that the synonymy is evident. Gonatopus fulgori is here synonymized with G. nigricans , because the original description is like that of G. nigricans (the type material of G. fulgori is lost). In the past these two species were separated, because attributed to two different zoogeographic regions ( fulgori in Palaearctic, nigricans in Oriental and Australian regions).

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

2. Haplogonatopus oratorius ( Westwood, 1833)

( Plates 104C, D , 105A, B )

Gonatopus oratorius Westwood 1833: 496 ; Richards 1935: 145. Gonatopus mayeti Kieffer in Kieffer & Marshall 1905: 103 (synonymized by Abdul-Nour 1974); type locality: Austria. Dicondylus oratorius (Westwood) : Kieffer 1907: 22. Haplogonatopus atratus Esaki & Hashimoto 1932: 25 (synonymized by Olmi 1993d); type locality: Utsunomiya ( Japan);

Kitamura 1982: 172; Olmi 1984: 1413; Chen 1988: 45; Olmi 1993d: 51. Monogonatopus oratorius (Westwood) : Richards 1939: 200; Raatikainen 1961: 131; Currado & Olmi 1972: 143; Abdul-Nour

1974: 1015. Haplogonatopus suchovi Ponomarenko 1970: 429 (synonymized by Olmi 1984); type locality: Omsk (Siberia, Russia). Haplogonatopus oratorius (Westwood) : Olmi & Currado 1979b: 39; Olmi 1984: 1409; Tussac 1988: 118; Olmi 1989a: 7; Olmi

1993d: 51; Olmi 1994: 65; Olmi 1995g: 5; Xu & He 1996b: 129; Guglielmino & Olmi 1997: 180–252; Tussac & Olmi

1998: 491; Olmi 1999a: 227; Olmi 1999b: 28; He & Xu 2002: 315; Guglielmino & Olmi 2006: 37, 50; Xu et al. 2012a: 13. Haplogonatopus katangae (Benoit) : Xu & He 1995b: 594 (misidentified); He & Xu 2002: 309 (misidentified). Haplogonatopus apicalis R. Perkins : Sugiura et al. 2004: 11 (misidentified). Haplogonatopus oratrius (sic) (Westwood): Mita et al. 2012: 50.

Description of female ( Plate 105A ). Apterous; length 2.0– 3.1 mm. Head brown or black, except anterior region of face, clypeus and mandible yellow; antenna brown, except segments 1–2 and 10 yellow; mesosoma reddish or testaceous; petiole and metasoma black; legs yellow-testaceous. Occasionally body totally black. Antenna clavate; antennal segments in following proportions: 7:5:9:5:5:5:4:4:5:8. Head excavated, shiny, weakly granulated; frontal line complete; occipital carina absent; POL = 2; OL = 2; OOL = 7; greatest breadth of posterior ocelli about as long as POL. Palpal formula 2/1. Pronotum shiny, smooth, unsculptured, not crossed by strong transverse impression. Scutum dull, hairless, sculptured by some longitudinal keels. Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface, metanotum, mesopleuron and metapleuron. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 13:2:3:8:13. Enlarged claw ( Plate 104D ) with one subapical tooth and one row of four–five lamellae. Segment 5 of protarsus ( Plate 104D ) with two rows of 6–12 lamellae; distal apex with about three–nine lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 105B ). Fully winged; length 2.2–2.5 mm. Head black, except mandible yellow-brown; antenna brown; mesosoma and metasoma black; legs brown, except tarsi testaceous. Antenna filiform; antennal segments in following proportions: 6:5:11:10:10:10:9:9:8:11; antennal segment 3 more than four times as long as broad (about 4.4). Head shiny, hairy, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli much longer than OOL (4:2); temple distinct; occiput concave. Palpal formula 2/1. Scutum dull, granulated. Notauli complete, posteriorly strongly converging, but running parallel near posterior margin of scutum. Scutellum and metanotum dull, finely punctate. Propodeum reticulate rugose, without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 104C ) long, slender, little distally broadened. Tibial spurs 1/1/2.

Material examined. Types: ♀ holotype of G. oratorius : UNITED KINGDOM: England, Surrey, Ripley ( HMO); ♀ holotype of G. mayeti : AUSTRIA ( ENSAM); ♀ lectotype (designated by Olmi, 1984) of H. atratus : JAPAN: Honshu, Tochigi Pref., Utsunomiya, Gunma, 6.VI.1931, E. Amano leg. ( ELKU); ♀ holotype of H. suchovi : RUSSIA: Western Siberia, Omsk, 1940, K.S. Suchov leg. ( ZMM) ( Ponomarenko , 1992). Paratype of H. atratus : JAPAN: same label locality as lectotype, 1♂ paralectotype ( ELKU). Paratypes of H. suchovi : same locality label as holotype, 12♀♀ and 5♂ ( ZMM). Other material. CHINA: Fujian ( Xu & He, 2003); Guangdong, Mt. Yunjishan, parasitized host collected 9.VI.2002, dryinid cocooning 11.VI.2002, dryinid adult emerged 26.VI.2002, reared from an unidentified species of Delphacid, M. Olmi leg., 1♂ ( MOLC); same locality label, parasitized host collected 9.VI.2002, 8♀♀ ( MOLC); Guangdong, Zhaoqing, 1 8.V.1998, Yufang Liu leg., 1♀ and 2♂ ( ZJUC); Guizhou, Sinan, IX.1986, VIII.1991, 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Shaanxi, Ningshan County, 6.V.2011, Chengyuan Jin leg., 1♀ and 1♂ ( SCAU); Zhejjiang, Hangzhou, Junhua He leg., 4♀♀ and 21♂ ( ZJUC); Zhejjiang, Lin'an, Junhua He leg., 15♂ ( ZJUC); Zhejjiang, Jiashan, Junhua He leg., 2♀♀ and 1♂ ( ZJUC); Zhejjiang, Shangyu, IX.1984, Haifu Wei leg., 1♀ ( ZJUC); Zhejjiang, Lishui, VI–VIII.1981, Junhua He leg., 1♀ and 1♂ ( ZJUC); Zhejiang, Wenzhou, Junhua He leg., 10♀♀ and 10♂ ( ZJUC); Zhejiang, Yiwu, 1–3.VIII.1986, Xiaoming Lou leg., 7♂ ( ZJUC). JAPAN: Ryukyu Islands, Iriomote-jima, 30.VI.2003, Sugiura leg., 1♀ ( Sugiura et al., 2004); Ogasawara (=Bonin) Islands (Mita et al., 2012). Palaearctic: H. oratorius is broadly spread in the Palaearctic, where it is known from Austria, China, France, Hungary, Italy, Japan, Lebanon, Romania, Russia (including Siberia), South Korea, Spain, Turkey and United Kingdom. In China it is known from the following locality: Inner Mongolia, Bayanzhouer, 15–22.VII.2008, Bo Qiu & Huayan Chen leg., 1♀ ( SCAU); Liaoning, Shenyang, 7.VI.1995, Juxian Lou leg., 1♀ ( ZJUC); Xinjiang, Mosuowan, 10.IX.1985, 1♀ ( ZJUC); Shandong, Weixian, V.1966, 1♀ ( ZJUC); Shandong, Shanghe, VII–VIII.2003, Jiahe Yan leg., 2♀♀ ( SCAU). Australian region: MARIANA ISLANDS: Guam, Talofofo ( BMNH); Guam, Dededo, ex cocoon on taro ( BPBM); Guam, Libugon, on taro ( AMNH).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China: Saccharosydne procerus (Metsumura) ( He & Xu, 2002) ; in France: Javesella pellucida (Fabricius) , Laodelphax striatellus (Fallén) ; in Guam ( Mariana Islands): Tarophagus proserpina (Kirkaldy) ; in Italy: Laodelphax striatellus (Fallén) , Megadelphax sordidulus (Stål) ; in Japan: Laodelphax striatellus (Fallén) ( Esaki & Hashimoto, 1932; Mita et al., 2012), Hosunka hakonensis (Matsumura) (Mita et al., 2012) , Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ( Kitamura, 1982) (see remarks after the key to the Oriental species of Haplogonatopus ); in South Korea: Laodelphax striatellus (Fallén) ; in Siberia ( Russia): Laodelphax striatellus (Fallén) ; in Turkey: Laodelphax striatellus (Fallén) , Toya propinqua (Fieber) ; in Ukraina: Laodelphax striatellus (Fallén) .

Biology. Information on the biology in Japan by Nishioka (1980), Kitamura (1982, 1983, 1985, 1986, 1987a, 1987b, 1987c, 1989a, 1989b), Yamada (1996), Yamada & Miyamoto (1998), Yamada & Kawamura (1999), Yamada & Kitashiro (2002), Yamada & Watanabe (2002), Yamada & Sugaura (2003) and Mita et al. (2012). The papers of Koyama (1989, 1991a, 1991b, 1993), Koyama et al. (1988, 1989), Koyama & Mitsuhashi (1989, 1990a, 1990b) and Abe & Koyama (1991) are of difficult interpretation, because the above authors mistook Haplogonatopus atratus Esaki & Hashimoto (= H. oratorius ) for Gonatopus lucens (Olmi) . Their data are related to G. lucens if the hosts are Cicadellidae , to H. atratus if the hosts are Delphacidae . This mistake is common to other authors. In the Oriental region, Chandra mistook Haplogonatopus apicalis R. Perkins (attacking Delphacidae ) for G. lucens (Olmi) (attacking Cicadellidae ). In the environments of Viterbo (Central Italy) H. oratorius overwinters as cocoon, rarely as larva in parasitized planthoppers. In Japan, it overwinters as larva in nymphs of L. striatellus (Mita et al., 2012) . In Japan, Kitamura (1989a) reported the presence of overwintering nymphs and a relatively high ratio of parasitism (9–34%) by H. oratorius from winter to spring in Shimane (Mita et al., 2012).

Natural enemies. According to Xu & He (2003) H. oratorius can be parasitized in China, Fujian, by Cheiloneurus exitiosus (R. Perkins) (= Echthrogonatopus lateocaudatus Xu & He, 2003 ).

Distribution. China (Fujian, Guangdong, Guizhou, Shaanxi, Zhejiang), Japan (Ryukyus, Ogasawara), in addition to Mariana Islands ( Guam) and many countries of Eastern (Inner Mongolia, Liaoning, Xinjiang, Shandong ( China); Honshu ( Japan); Siberia ( Russia); South Korea) and Western Palaearctic region (from Spain to European Russia).

18. Genus Gonatopus Ljungh, 1810

Gonatopus Ljungh 1810: 161 ; Richards 1937: 118; Olmi 1993b: 77; Olmi 1994: 66; Olmi 1995c: 787; Olmi 1995g: 5; Olmi 1996b: 182; Olmi 1998c: 72; Olmi 1998d: 177.; Olmi 1999a: 231; He & Xu 2002: 318.; Virla & Olmi 2008: 371.; Moya- Raygoza & Olmi 2010: 100.

Dicondylus Haliday in Curtis 1829 –1930: 110 (synonymized by Olmi, 1993b); type species: Dicondylus bicolor (Haliday in Curtis 1828), designated by Kieffer (1914b); Haliday 1833: 273; Walker 1837: 411; Kieffer 1914b: 72; Richards 1935: 147; Richards 1937: 118; Olmi 1984: 1374; Olmi 1993b: 78.

Labeo Haliday 1833: 273 (nom. praeocc., nec Cuvier 1817) (synonymized by Richards 1937); type species: Labeo vitripennis Haliday 1833 , by monotypy; Kieffer in Kieffer & Marshall 1905: 220; Walker 1837: 429; Dalla Torre 1898: 542.

nec Labea Blanchard 1840: 430 .

Labeola Haldeman 1842: 191 , new name for Labeo Haliday, 1833 (synonymized by Krombein 1979 with Dicondylus Haliday, 1829 –1930).

Pseudogonatopus R. Perkins 1905: 34 (synonymized by Olmi 1993b); type species: Pseudogonatopus kurandae R. Perkins, 1905 , by original designation; Kieffer in Kieffer & Marshall 1906: 488; Kieffer 1914b: 79; Richards 1939: 195; J. Perkins 1976: 32; Chandra 1980c: 192; Olmi 1984: 1191; Olmi 1993b: 78.

Paragonatopus R. Perkins 1905: 40 (synonymized by Olmi 1984); type species: Paragonatopus nigricans R. Perkins 1905 , by monotypy; Kieffer in Kieffer & Marshall 1906: 488; Kieffer 1914b: 87; Richards 1939: 195; Olmi 1979: 218.

Neogonatopus R. Perkins 1905: 42 (synonymized by Olmi 1984); type species: Neogonatopus ombrodes R. Perkins 1905 , by original designation.

Pachygonatopus R. Perkins 1905: 45 (synonymized by Olmi 1984); type species: Pachygonatopus melanias R. Perkins 1905 , by original designation.

Chalcogonatopus R. Perkins 1905: 46 (synonymized by Olmi 1984); type species: Chalcogonatopus gigas R. Perkins 1905 , by original designation.

Eugonatopus R. Perkins 1905: 46 (synonymized by Olmi 1984); type species: Chalcogonatopus (Eugonatopus) pseudochromus R. Perkins 1905 , by original designation.

Mercetia Kieffer in Kieffer & Marshall 1906: 495 (synonymized by Olmi 1996b); type species: Mercetia longicollis (Kieffer 1905) , by original designation; Olmi 1996b: 182; Kieffer 1914b: 19.

Platygonatopus Kieffer in Kieffer & Marshall 1906: 500 (synonymized by Olmi, 1984); type species: Platygonatopus planiceps (Kieffer 1904) , designated by Richards (1939).

Apterodryinus R. Perkins 1907: 14 (synonymized by Olmi 1993b); type species: Apterodryinus torvus R. Perkins 1907 , by monotypy; Olmi 1984: 1342; Olmi 1993b: 78.

Agonatopus R. Perkins 1907: 29 (synonymized by Olmi 1984); type species: Agonatopus pallidicornis R. Perkins 1907 , designated by Richards (1939).

Agonatopoides R. Perkins 1907: 33 (synonymized by Olmi 1993b); type species: Agonatopoides synchromus R. Perkins 1907 , by monotypy; Olmi 1984: 1310; Olmi 1993b: 78.

Cyrtogonatopus Kieffer 1907: 19 (synonymized by Olmi 1984); type species: Cyrtogonatopus breviforceps (Kieffer 1904) , monotypic.

nec Trichogonatopus Kieffer 1909: 336 .

Pristogonatopus Kieffer 1913c: 325 (synonymized by Olmi & Currado 1974); type species: Pristogonatopus dentatiforceps (Kieffer 1904) , by original designation; Kieffer 1914b: 71; Richards 1939: 195.

Digonatopus Kieffer 1913c: 325 (synonymized by Richards 1939); type species: Digonatopus javanus (R. Perkins 1912) , by monotypy.

Trigonatopus Kieffer 1913c: 325 (synonymized by Olmi 1984); type species: Trigonatopus bifarius (Kieffer in Kieffer & Marshall 1906), by original designation; Kieffer 1914b: 72.

Laberius Kieffer 1914b: 59 , new name for Labeo Haliday 1833 (synonymized by Richards 1937); Richards 1936: 169.

Trichogonatopus Hellén 1930: 2 (synonymized by Olmi 1984); type species: Trichogonatopus lindbergi Hellén 1930 , original designation.

Laberinus Ogloblin 1932: 266 (mispelling).

Metagonatopus Ogloblin 1932: 264 (synonymized by Olmi 1984); type species: Metagonatopus brasiliensis Ogloblin 1932 , by original designation.

Allogonatopus Haupt 1938: 27 (synonymized by Olmi 1984); type species: Allogonatopus procerus Haupt 1938 , by original designation.

Donisthorpina Richards 1939: 201 (synonymized by Olmi 1993b); type species: Donisthorpina formicicola Richards 1939 , by original designation; Olmi 1984: 1304; Olmi 1993b: 78.

Plectrogonatopus Richards 1939: 207 (synonymized by Olmi 1984); type species: Plectrogonatopus striatus (Kieffer 1905) , by original designation.

Tetrodontochelys Richards 1939: 217 (synonymized by Olmi 1993b); type species: Tetrodontochelys ljunghii ( Westwood 1833) , by original designation; Olmi 1984: 1434; Olmi 1993b: 78.

Epigonatopoides Richards 1939: 221 (synonymized by Olmi 1984); type species: Epigonatopoides pulicarius ( Klug 1810) , by original designation.

Idologonatopus Ogloblin 1953: 111 (synonymized by Olmi 1984); type species: Idologonatopus nigrithorax Ogloblin 1953 , by original designation.

Rhynchogonatopus Benoit 1954: 390 (synonymized by Olmi 1984); type species: Rhynchogonatopus meridionalis Benoit 1954 , by original designation.

Madecagonatopus Benoit 1954: 392 (synonymized by Olmi 1984); type species: Madecagonatopus ankaratrus ( Ceballos 1936) , by original designation.

Pseudogonatopoides J. Williams 1956: 90 (synonymized by Olmi 1984); type species: Pseudogonatopoides mauritianus J. Williams 1956 , by original designation.

Cyrtogonatopoides Ponomarenko 1966: 1579 (synonymized by Olmi 1984); type species: Cyrtogonatopoides borealis Ponomarenko 1966 , by original designation.

Plectrogonatopoides Ponomarenko 1975a: 318 (synonymized by Olmi 1993b); type species: Plectrogonatopoides chortinis Ponomarenko 1975 , by original designation; Olmi 1984: 1431; Olmi 1993b: 78.

Tetradontochelys J. Perkins 1976: 35 (injustified emendation).

Neogonatopoides Abdul-Nour 1976: 274 (synonymized by Olmi 1984); type species: Neogonatopoides claudiae Abdul-Nour 1976 , by monotypy.

Dolichochelys Olmi & Currado 1976: 177 (synonymized by Olmi 1984); type species: Dolichochelys longicornis (Kieffer 1904) , by original designation.

Megagonatopus Olmi & Currado 1976: 178 (synonymized by Olmi 1984); type species: Megagonatopus birmanus Olmi & Currado 1976 , by original designation.

Acrodontochelys Currado 1976: 13 (synonymized by Olmi 1993b); type species: Acrodontochelys bouceki Currado 1976 , by original designation; Olmi 1984: 1172; Olmi 1993b: 78.

Nogatopus Ponomarenko 1979: 357 (synonymized by Olmi 1984); type species: Nogatopus gobiensis Ponomarenko 1979 , by original designation.

Paradicondylus Olmi 1986: 102 (synonymized by Olmi 1998c); type species: Paradicondylus paulyi Olmi 1986 , by monotypy and original designation; Olmi 1993b: 78; Olmi 1998c: 72.

Type species. Gonatopus formicarius Ljungh, 1810 , by monotypy.

Diagnosis. Female: apterous ( Plates 106B , 126A ) or fully winged ( Plate 106A ); palpal formula 3/2, 4/2, 4/3, 5/ 2, 5/3, or 6/3; antenna with or without rhinaria; pronotum crossed or not by transverse furrow ( Plate 108C, D ); enlarged claw with distal apex pointed and with one large or small subapical tooth ( Plate 107B, C ) (occasionally subapical tooth absent, then enlarged claw with distal group of lamellae ( Plate 107A )); in fully winged forms, segment 5 of protarsus with more than 20 lamellae; tibial spurs 1/0/1. Male: fully winged ( Plate 110A, B ); occipital carina absent or incomplete (in this last case, only present behind and shortly on sides of posterior ocelli); occiput concave; temple present; palpal formula 3/2, 4/2, 4/3, 5/2, 5/3, or 6/3; tibial spurs 1/1/2.

Distribution. Worldwide.

Hosts. Acanaloniidae , Cicadellidae (except Typhlocybinae , Idiocerinae , Macropsinae ), Delphacidae , Flatidae , Issidae , Lophopidae , Tropiduchidae ( Guglielmino & Olmi, 1997, 2006, 2007).

World species. Four hundred and thirty nine species are known, fifty eight species in the Oriental region and one species incertae sedis.

Remarks. Larvae of Gonatopus were described by Ponomarenko (1975b), Carcupino et al. (1996, 1998), Guglielmino & Virla (1998), Virla & Mangione (2000), Guglielmino (2002), Mangione & Virla (2004) and Guglielmino et al. (2006, 2007, 2008). Gonatopus females were divided for convenience by Olmi (1993b) into eleven groups on the basis of the following characters:

Key to the groups of Gonatopus

1. Fully winged ( Plate 106A )....................................................................... Group 11

- Apterous ( Plate 106B )................................................................................. 2

2. Enlarged claw without subapical tooth ( Plate 107A ), or with one very small subapical tooth ( Plate 107B )............... 3

- Enlarged claw with one large subapical tooth ( Plate 107C )..................................................... 5

3. Distal apex of enlarged claw with group of lamellae ( Plate 107A )......................................... Group 8

- Distal apex of enlarged claw without lamellae ( Plate 107C, D )................................................. 4

4. Pronotum not crossed by transverse impression or slightly impressed ( Plate 108A )........................... Group 6

- Pronotum crossed by strong transverse impression ( Plate 108B ).......................................... Group 7

5. Pronotum not crossed by transverse impression or slightly impressed ( Plate 108C )................................. 6

- Pronotum crossed by strong transverse impression ( Plate 108D )................................................ 7

6. Labial palpi 3-segmented......................................................................... Group 9

- Labial palpi 2-segmented......................................................................... Group 5

7. Enlarged claw without lamellae, with or without bristles or peg-like hairs ( Plate 109B )........................ Group 1

- Enlarged claw with lamellae ( Plate 107C ).................................................................. 8

8. Labial palpi 2-segmented............................................................................... 9

- Labial palpi 3-segmented.............................................................................. 10

9. Maxillary palpi composed of two–four segments...................................................... Group 2

- Maxillary palpi 5-segmented..................................................................... Group 10

10. Maxillary palpi 6-segmented...................................................................... Group 4

- Maxillary palpi 5-segmented...................................................................... Group 3

The above groups are distributed in the zoogeographical regions as follows:

Palaearctic: groups 1, 2, 3, 4, 5, 6, 7, 10, 11.

Afrotropical: groups 1, 2, 3, 4, 5, 6, 7, 8, 11.

Oriental: groups 1, 2, 3, 4, 5, 6, 7, 10.

Nearctic: groups 1, 2, 3, 4, 5, 6, 7, 9.

Neotropical: groups 1, 2, 3, 4, 5, 6, 7, 10.

Australian: groups 1, 2, 4, 5, 6, 7, 9.

As far as, it is not possible divide also males into the above groups.

All Oriental species of Gonatopus can be attributed to one of the above eight female groups, except Gonatopus leurosipropodeus He & Xu, 2002 . The attribution of this species to a distinct group is impossible, because it was described on the basis of male specimens and its female is unknown. For this reason, it is attributed to an uncertain group (see “Uncertain group”).

Key to the males

(unknown for the species not present in the key) (males not divided into groups)

1. Notauli incomplete.................................................................................... 2

- Notauli complete..................................................................................... 4

2. Propodeum with dorsal surface dull, granulated, sculptured by irregular striae, and posterior surface shiny, smooth and unsculptured; dorsal surface of propodeum much shorter than posterior surface...................... G. maurus Kieffer

- Propodeum completely smooth and shiny, unsculptured; dorsal surface of propodeum much longer than posterior surface... 3

3. Dorsal process of paramere transverse ( Plate 122C ); clypeus testaceous.............................. G.lucens (Olmi)

- Dorsal process of paramere about parallel to penis, not transverse ( Plate 111A ); clypeus black....... G. nearcticus (Fenton)

4. Notauli posteriorly broadly separated..................................................................... 5

- Notauli posteriorly joint (or very close and almost joint)..................................................... 14

5. Dorsal process of paramere reduced to inner expansion of paramere ( Plate 122B )....... G. flavifemur (Esaki & Hashimoto)

- Dorsal process of paramere not reduced ( Plates 111B , 118D , 119D , 128A , 131A, B )................................ 6

6. Dorsal process of paramere much shorter than paramere ( Plates 128A , 131A, B )................................... 7

- Dorsal process of paramere as long as, or longer, or slightly shorter than paramere ( Plates 111B , 118D , 119D )............ 9

7. Head with POL more than four times as long as OOL; ocelli very large............................ G. rufoniger Olmi

- Head with POL less than three times as long as OOL; ocelli small............................................... 8

8. Dorsal process of paramere very slender ( Plate 131B ); clypeus completely black...................... G. superbus Olmi

- Dorsal process of paramere less slender ( Plate 128A ); clypeus medially black, laterally testaceous... G.javanus (R. Perkins)

9. Face completely or almost completely testaceous or yellow................................................... 10

- Face completely black or brown........................................................................ 11

10. Dorsal process of paramere longer than penis and paramere ( Plate 118D )......................... G. borneanus (Olmi)

- Dorsal process of paramere shorter than penis and paramere ( Plate 111B )...................... G. nigricans (R. Perkins)

11. Antennal segment 3 less than three times as long as broad; dorsal process of parameres longer than penis ( Plate 119D )............................................................................................ G. pyrillae (Mani)

- Antennal segment 3 more than three times as long as broad; dorsal process of parameres shorter than penis ( Plates 111B , 117C , 122D )............................................................................................. 12

12. Dorsal process of paramere almost completely joint to paramere ( Plate 111B ).................. G. nigricans (R. Perkins)

- Dorsal process of paramere completely separated from paramere ( Plates 117C , 122D ).............................. 13

13. Dorsal process of paramere slender ( Plate 122D )............................................. G. pedestris Dalman

- Dorsal process of paramere broader ( Plate 117C )............ G.malesiae (Olmi) type B or G.sarawakensis (Olmi) type B

14. Dorsal process of paramere very reduced ( Plates 117A , 122B ), reduced to inner expansion of paramere................ 15

- Dorsal process of paramere not reduced ( Plates 111B , 117B ).................................................. 16

15. Inner expansion of paramere with many irregular striae ( Plate 122B )................. G. flavifemur (Esaki & Hashimoto)

- Inner expansion of paramere without irregular striae ( Plate 117A )............................... G. nudus (R. Perkins)

16. Dorsal process of paramere much shorter than paramere ( Plates 111B , 131C )..................................... 17

- Dorsal process of paramere approximately as long as paramere ( Plates 117B , 131D )............................... 18

17. Dorsal process of paramere not transverse, with distal apex broad ( Plate 111B ); head and mesosoma mostly black or brown................................................................................... G. nigricans (R. Perkins)

- Dorsal process of paramere transverse, with distal apex pointed ( Plate 131C ); head and mesosoma mostly yellow-testaceous....................................................................................... G. asiaticus (Olmi)

18. Propodeum completely reticulate rugose; dorsal process of paramere broader ( Plate 117B )................................................................................ G.malesiae (Olmi) type A or G. sarawakensis (Olmi) type A

- Propodeum with posterior inclined region reticulate rugose and anterior flat region smooth and unsculptured; dorsal process of paramere slender ( Plate 131D ).................................................... G. leurosipropodeus He & Xu

Remarks. In the above key, there are two species, whose males are not distinguishable, G. malesiae (Olmi) and

G. sarawakensis (Olmi) ; the situation is explained in remarks related to G. sarawakensis (see Group 2).

Group 1

One species is known in the Oriental region.

1. Gonatopus nearcticus (Fenton, 1905)

( Plates 109A, B , 110A, B , 111A )

nec Pseudogonatopus americanus R. Perkins 1905: 37 .

Epigonatopus americanus Fenton 1921: 70 (preoccupied by Pseudogonatopus americanus R. Perkins 1905 ) (synonymized by Olmi 1993b); type locality: Ames (Iowa, USA).

Pachygonatopus nearcticus Fenton 1927: 6 .

Platygonatopus ugandanus Benoit 1951a: 300 (synonymized by Olmi 1993b); type locality: Bululo ( Uganda); Benoit 1951d: 19.

Rhynchogonatopus ugandanus (Benoit) : Olmi & Currado 1976: 173.

Acrodontochelys bouceki Currado 1976: 14 (synonymized by Olmi 1993b); type locality: Nerja ( Spain); Graham 1983: 1; Olmi 1984: 1174; Graham 1988: 79.

Acrodontochelys ugandanus (Benoit) : Olmi 1984: 1174.

Acrodontochelys sinensis Olmi 1984: 1180 (synonymized by Xu et al. 2012b); type locality: Macau ( China) .

Acrodontochelys americanus (Fenton) : Olmi 1984: 1182.

Gonatopus nearcticus (Fenton) : Olmi 1993b: 80, 83; Guglielmino & Olmi 1997: 194, 237; Tussac & Olmi 1998: 490; Olmi 1999a: 246; Olmi 2004a: 368; Olmi 2005b: 126; Olmi 2006: 44; Guglielmino & Olmi 2007: 126; Olmi 2009b: 459; Burn 2011: 88; Penati & Olmi 2012: 79; Xu et al. 2012b: 21.

Gonatopus sinensis (Olmi) : He & Xu 2002: 349.

Description of female ( Plate 109A ). Apterous; length 2.0–4.0 mm. Head black or brown, with mandible, clypeus and anterior region of face yellow or testaceous; antenna black or brown, with segments 1–3 yellow or testaceous; occasionally antenna testaceous-brown; mesosoma, petiole and metasoma usually black, except posterior apex of propodeum testaceous; occasionally mesosoma and metasoma brown-testaceous or testaceous-reddish, with petiole black; occasionally mesosoma brown, with scutum yellow; legs yellow or testaceous, with coxae and clubs of femora almost completely black, or darkened, or brown-reddish. Antenna clavate; antennal segments of holotype of P. nearcticus in following proportions: 4:4:9:5.5:5:5:4:4:4:7; antennal segments of holotype of A. sinensis in following proportions: 5:3:7:4:4:4:4:4:3.5:5. Head flat or slightly convex, dull, weakly or strongly granulated; frontal line complete; occipital carina absent; temple distinct; POL = 3; OL = 2; OOL = 5. Palpal formula 5/2, 4/2, 3/2, or 2/2. Pronotum crossed by strong transverse impression, shiny, weakly or strongly granulated. Scutum dull, granulated. Metanotum granulated and transversely striate, not hollow behind scutellum. Metathorax + propodeum dull, granulated, with posterior surface of propodeum transversely striate; occasionally posterior surface almost completely granulated and not transversely striate. Meso-metapleural suture obsolete. Mesopleuron and metapleuron granulated, with or without transverse striae. Protarsus with first segment about twice as long as segment 4 (14:7; or 15:7; or 12:7). Protarsal segments of A. sinensis holotype in following proportions: 15:2:3:7:13. Enlarged claw ( Plate 109B ) with one large subapical tooth and one row of three–six peg-like bristles. Segment 5 of protarsus ( Plate 109B ) with two rows of 6–22 lamellae; distal apex with about six–nine lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 110A, B ): fully winged; length 1.6–2.8 mm. Head black or brown, except mandible testaceous or yellow; antenna brown or black; mesosoma and metasoma black or brown; legs banded, brown or black, with trochanters, stalks of femora, extremities of tibiae and tarsi yellow. Antenna hairy, filiform; antennal segments of male paratype of A. bouceki in following proportions: 5:4:8:7:7:7:6:6:6:8; antennal segment 3 four or more than four times as long as broad. Antennal segments of specimen from China, Ruyang, in following proportions: 4:4:7:7:6.5:6.5:5.5:5:5:7. Head shiny, alutaceous, slightly punctate; vertex laterally without two shiny and smooth areas situated between posterior ocelli and eyes; frontal line incomplete, only present between antennal toruli; occipital carina absent; occiput concave; temple distinct; POL = 6; OL = 4; OOL = 2 (or POL = 6; OL = 2; OOL = 2; or POL = 7; OL = 3; OOL = 1.5); greatest breadth of posterior ocelli about twice as long as OOL. Palpal formula 3/2, 4/2. Scutum dull, hairy, slightly granulated. Notauli incomplete, reaching approximately 0.4–0.7 length of scutum. Scutellum and metanotum shiny, smooth, finely punctate, unsculptured among punctures. Propodeum completely shiny, smooth, unsculptured; dorsal surface of propodeum with or without median longitudinal furrow. Fore wing hyaline, without dark transverse bands; stigmal vein regularly curved, with distal part much longer than proximal part. Dorsal process of paramere ( Plate 111A ) more or less long and slender, with distal third parallel to penis. Tibial spurs 1/1/2.

Material examined. Types: ♀ holotype of E. americanus : USA: Iowa, Story County, Ames, 25.VII.1919, E.D. Ball leg. ( USNM); ♀ holotype of P. nearcticus : USA: Woodbury County, Sioux City, IX.1919, reared from a jassid, C.N. Ainslie leg. ( USNM); ♀ holotype of P. ugandanus : UGANDA: Bululo (=present Bulule?), 1909, E. Bayon leg. ( MCSNG); ♀ holotype of A. bouceki : SPAIN: Malaga, Nerja, 3.VII.1974, Z. Bouček leg. ( BMNH); ♀ holotype of A. sinensis : CHINA: Macau ( BPBM). Paratypes of E. americanus : same locality label as holotype, 4♀♀ ( USNM). Paratypes of A. bouceki : same locality label as holoype, 23.VI.1973, Z. Bouček leg., 14♀♀ ( BMNH, DIVAPRA, MRSNT, USNM); SPAIN: Granada, La Herradura, 24.VI.1973, Z. Bouček leg., 1♀ and 1♂ ( BMNH); SPAIN: Granada, Calahonda, 22.VI.1973, Z. Bouček leg., 2♀♀ ( BMNH); SPAIN: Barcelona, Calella de la Costa, VI.1974, 1♀ ( BMNH); SPAIN: Castellón, Benicasim, 13–15.VI.1973, 22–24.VI.1974, Z. Bouček leg., 1♀ and 1♂ ( BMNH). Paratypes of A. sinensis : same locality label as holotype, 3♀♀ ( AMNH, BPBM); CHINA: Chin San, 1♀ ( BPBM); VIET NAM: Lam Dong Prov., Di Linh, Plateau of Djiring, 1♀ ( AMNH). Other material. CHINA: Guangdong, Mt. Nankunshan, 8.VI.2002, M. Olmi leg., 1♀ ( MOLC); Foot of Nanling National Nature Reserve, Ruyang, reared from adult of unidentified Cicadellid, M. Olmi reared, parasitized host collect 12.IX.2004, dryinid cocooning 13.IX.2004, dryinid adult emerged X.2004, 1♀ and 3♂ ( MOLC); same locality label, parasitized host collect 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 2♂ ( MOLC); Macau, X.1906, R. Perkins collection, 1♀ and 1♂ ( USNM). INDIA: Karnataka, Bangalore, 3.XI.1979, Z. Bouček leg., 1♀ ( BMNH); SINGAPORE ( NMNH). THAILAND: Chumphon Prov., Chumplion, 9.II.2000, Ralf Jochmann leg., 1♀ ( MTC); Mae Hong Son Prov., Soppong, 19°27'N 98°20'E, 1500 m, 7–12.V.1996, S. Becvar leg., 1♀ ( OLL). Palaearctic: G. nearcticus is known from the following countries: Croatia, Cyprus, France, Greece, Italy, Montenegro, Portugal, Spain (including Canary Islands) and Turkey ( Olmi, 1999a). Afrotropical region: G. nearcticus is known from the following countries: Benin, Botswana, Ethiopia, Kenya, Mozambique, Namibia, South Africa, Swaziland, Uganda, Zimbabwe ( Olmi, 1984, 1999a). Nearctic region: G. nearcticus is known from the USA ( Olmi, 1984, 1999a).

Hosts. Cicadellidae ( Guglielmino & Olmi, 1997, 2007): in France: Balclutha frontalis (Ferrari) ; in Italy: Balclutha rosea (Scott) ; in Namibia: Paradorydium spatulatum (Naudé) ; in South Africa: Balclutha rosea (Scott) ; in the USA: Balclutha neglecta (DeLong & Davidson) , Balclutha impicta (Van Duzee) .

Distribution. China ( Macau, Guangdong), India (Karnataka), Thailand (Chumphon, Mae Hong Son), Viet Nam (Lam Dong), in addition to many countries of the Afrotropical, Nearctic and Western Palaearctic regions.

Group 2

Ten species are known in the Oriental region.

Key to the females

1. Mesosoma totally black or black-brown (at most distal apex of propodeum and posterior margin of pronotum testaceous or reddish)............................................................................................... 2

- Mesosoma at least partly testaceous-reddish, or testaceous-ferruginous, or brown-reddish, or yellow................... 4

2. Metanotum with sides protruding; protrusions rounded ( Plate 113G )............................. G. nepalensis (Olmi)

- Metanotum with sides rounded ( Plates 112A , 114A ).......................................................... 3

3. Species large (body length about 5.0 mm); body covered with long sparse hairs ( Plate 112A ); temple absent ( Plate 112A ); head more excavated....................................................................... G. capillus Xu & He

- Species small (body length less than 4.5 mm); body hairless or covered with very short hairs; temple distinct ( Plate 114A ); head less excavated ( Plate 114B )...................................................... G. nigricans (R. Perkins)

4. Metanotum with sides rounded ( Plates 114A , 115A ); meso-metapleural suture obsolete.............................. 5

- Metanotum with sides protruding ( Plate 117G ); protrusions rounded ( Plates 111C , 117G ) or pointed ( Plate 113C ); meso-metapleural suture at least partly distinct....................................................................... 6

5. Metanotum short, about as long as scutellum............................................... G. nudus (R. Perkins)

- Metanotum long, about twice as long as scutellum........................................ G. nigricans (R. Perkins)

6. Anterior surface of metathorax + propodeum with numerous longitudinal keels.................. G. sarawakensis (Olmi)

- Anterior surface of metathorax + propodeum smooth, with no keels............................................. 7

7. Metathorax + propodeum with slight or strong median furrow.................................................. 8

- Metathorax + propodeum without median furrow............................................................ 9

8. Metanotum with lateral protrusions more prominent ( Plate 113A ); metathorax + propodeum with deep median furrow; mesometapleural suture very strong and distinct............................................. G.lankae (Ponomarenko)

- Metanotum with lateral protrusions less prominent ( Plate 113E ); metathorax + propodeum with shallow median furrow; mesometapleural suture slightly distinct, only visible near lateral protrusions of metanotum.................. G. muiri (Olmi)

9. Metanotum with lateral protrusions less prominent ( Plate 117G )................................... G. validus (Olmi)

- Metanotum with lateral protrusions more prominent ( Plates 111C , 113C )....................................... 10

10. Lateral protrusions of metanotum at right angle ( Plate 111C ); mesopleuron and metapleuron transversely striate................................................................................................. G.bicuspis (Olmi)

- Lateral protrusions of metanotum not at right angle ( Plate 113C ); mesopleuron and metapleuron smooth, not transversely striate................................................................................ G. malesiae (Olmi)

1. Gonatopus bicuspis (Olmi, 1993) , comb. nov.

( Plate 111C, D )

Pseudogonatopus bicuspis Olmi 1993d: 50 .

Description of Female. Apterous; length 3.2 mm. Testaceous-ferruginous, except petiole black, antennal segment 8 testaceous and antennal segments 9–10 whitish. Antenna clavate; antennal segments in following proportions: 10:6:15:8:6:6:5.5:5:5:8.5. Head very excavated, shiny, smooth, unsculptured; frontal line absent; occipital carina absent; temple absent; POL = 0.3; OL = 3; OOL = 6. Palpal formula 3/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Metanotum flat, transversely striate, with two lateral pointed apophyses ( Plate 111C ). Metathorax + propodeum shiny, smooth, unsculptured; posterior surface of propodeum transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 13:3:4.5:15.5:25. Enlarged claw ( Plate 111D ) with one large subapical tooth and one row of six lamellae + one bristle. Segment 5 of protarsus ( Plate 111D ) with two rows of 16 lamellae; distal apex with about 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: PHILIPPINES: Negros Island , Camp, 16.V.1987 ( AMNH) .

Hosts. Unknown.

Distribution. Philippines (Negros).

2. Gonatopus capillus Xu & He, 1998

( Plate 112A, B )

Gonatopus capillus Xu & He 1998d: 306 ; He & Xu 2002: 351; Guglielmino & Olmi 2006: 49.

Description of female ( Plate 112A ). Apterous; length 5.0 mm. Head black, except mandible testaceous; antenna black, except segment 1 partly testaceous and partly brown and segment 10 yellow-whitish; mesosoma, petiole, metasoma and legs black. Body completely covered with long hairs (as in some species of Trichogonatopus Kieffer ). Antenna clavate; antennal segments in following proportions: 6:3.6:14.5:9:8.5:8:6:6:6:8. Head shiny, smooth, unsculptured, except ocellar region granulated; frontal line incomplete; occipital carina absent; temple absent; POL = 1; OL = 0.5; OOL = 5. Palpal formula 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum and metanotum without lateral pointed apophyses. Metanotum flat, not hollow behind scutellum. Metathorax + propodeum with anterior region smooth, disc reticulate rugose and posterior region transversely striate; anterior region and disc with track of long median longitudinal furrow. Mesopleura and metapleura transversely striate. Meso-metapleural suture almost completely absent, only present on sides of metanotum. Fore tarsal segments in following proportions: 10:1.5:3:10:15.5. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 112B ) with one large subapical tooth and one row of 15 lamellae. Segment 5 of protarsus ( Plate 112B ) with two rows of 12 + 14 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: CHINA: Guizhou, Sinan , IX.1986, Yuxiang Chen leg. ( ZJUC) . Paratype: same label as holotype, 1♀ ( ZJUC) .

Hosts. Delphacidae ( Guglielmino & Olmi, 2006) : in China: Nilaparvata lugens (Stål) ( Xu & He, 1998d) .

Distribution. China (Guizhou).

3. Gonatopus lankae (Ponomarenko, 1981) , comb. nov.

( Plate 113A, B )

Pseudogonatopus lankae Ponomarenko 1981b: 881 ; Olmi 1984: 1249.

Description of Female. Apterous; length 3.2–3.3 mm. Brown-reddish or testaceous-dark; antenna usually darkened, except segment 10 yellow; legs testaceous, except trochanters and procoxa partly whitish, hind side of metafemur whitish, distal apex of propodeum whitish. Antenna clavate; antennal segments in following proportions: 14:8:14:9:9:6:6:6:6:11. Head excavated, dull, alutaceous or weakly granulated; frontal line complete; occipital carina absent; temple prominent; POL = 2; OL = 3; OOL = 10. Palpal formula 4/2 or 5/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Scutellum shiny, smooth, unsculptured. Metanotum flat, shiny, unsculptured, weakly hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded. Metathorax + propodeum ( Plate 113A ) shiny, smooth, unsculptured or granulated; posterior surface of propodeum transversely striate. Disc of metathorax + propodeum with strong median furrow. Mesopleura and metapleura dull, weakly granulated, not transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 15:4:6:20:31. Enlarged claw ( Plate 113B ) with one large subapical tooth and one row of six–eight lamellae. Segment 5 of protarsus ( Plate 113B ) with two rows of 19–21 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Ratnapura District , Gilimale Induruwa Jungle, 13.III.1979, K. Krombein leg. ( USNM) . Other material. SRI LANKA: Ratnapura District ., Sinharaja Jungle, Waturawa Forest ( AMNH) ; Matale District , Sigiriya, 07°57'N 80°46'E, 18.III.1999, C.J. Burwell leg. ( QMC) .

Hosts. Unknown.

Distribution. Sri Lanka (Ratnapura, Matale).

Remarks. The female specimen from Sri Lanka, 07°57'N 80°46'E, has 5-segmented maxillary palps; the palpal formula 5/2 is typical of the group 10 (former genus Donisthorpina Richards ); for this reason, G. lankae is also included into group 10.

4. Gonatopus malesiae ( Olmi, 1984) , comb. nov.

( Plate 113C, D )

Pseudogonatopus malesiae Olmi 1984: 1248 .

Description of Female. Apterous; length 2.8–3.6 mm. Usually testaceous-ferruginous, with petiole black; antenna usually testaceous, with segments 9–10 whitish (occasionally antennal segment 9 partly whitish). In specimen from Myanmar, 17°38.9'N 94°38.1'E, antenna testaceous-brown, except segments 1 and 10 whitish. In specimen from China, Mt. Jinggangshan, antenna brown, except rhinaria areas testaceous, and mesosoma brown-testaceous. Antenna clavate; antennal segments of holotype in following proportions: 16:13:38:23:18:12:12:13:12:20. Head very excavated, shiny, smooth, unsculptured, hairless; frontal line complete; occipital carina absent; POL = 0.3; OL = 2; OOL = 8.5. Palpal formula 3/2 or 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat, without lateral pointed apophyses. Metanotum shiny, unsculptured, not hollow behind scutellum, with sides protruding; lateral protrusions pointed ( Plate 113C ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum without median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture very slightly distinct. In specimen from Myanmar, 17°38.9'N 94°38.1'E, meso-metapleural suture partly and slightly visible (not visible on sides of metanotum). Segment 1 of protarsus usually shorter than segment 4 (13:16). Enlarged claw ( Plate 113D ) with one large subapical tooth and one row of seven–ten lamellae. Segment 5 of protarsus ( Plate 113D ) with two rows of 12–17 lamellae; distal apex with at least 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown (see Gonatopus sarawakensis (Olmi)) .

Material examined. Type: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu National Park , 5.VIII.1978, P.M. Hammond & J.E. Marshall leg. ( BMNH) . Other material. CHINA: Jiangxi, Mt. Jinggangshan , 13.VIII.2007, Junhua He leg., 1♀ ( SCAU) . MALAYSIA: Malaya, Fraser's Hill , 4200' ( AMNH) . MYANMAR: Rakhine State, NE of Gwa, Elephant Camp in Elephant Sanctuary , 17°38.9'N 94°38.1'E, 3.XII.2000, leaf litter in secondary forest, K.J. Ribardo leg., 2♀♀ ( CAS) .

Hosts. Unknown.

Distribution. China (Jiangxi), Malaysia ( Sarawak, Malaya), Myanmar (Rakhine).

5. Gonatopus muiri ( Olmi, 1984)

( Plate 113E, F )

Pseudogonatopus muiri Olmi 1984: 1251 ; Xu et al. 2011g (Suppl.): 20.

Description of Female. Apterous; length 2.5–3.0 mm. Holotype testaceous–reddish, except metasoma brown and antenna brown, with segments 1–2 testaceous (specimen from China, Bawangling National Nature Reserve, testaceous, except antenna brown, with segments 1–2 testaceous and segment 10 whitish, petiole black and metasoma brown). Antenna clavate; antennal segments of a specimen from China, Bawangling National Nature Reserve, in following proportions: 8:4.5:10:6:5:5:4.5:4.5:4:8. Head excavated; head of holotype dull and slightly granulated; head of specimen from China, Bawangling National Nature Reserve, shiny and unsculptured; frontal line of specimen from China, Bawangling National Nature Reserve, complete; occipital carina absent; head of specimen from China, Bawangling National Nature Reserve, with POL = 0.5; OL = 1.5; OOL = 6. Palpal formula 3/ 2 in holotype, 4/ 2 in specimen from China, Bawangling National Nature Reserve. Pronotum crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat. Metanotum shiny, unsculptured, not hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded ( Plate 113E ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum with slight median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture weakly distinct, in holotype only visible near lateral protrusions of metanotum. Meso-metapleural suture of specimen from China, Bawangling National Nature Reserve, visible only between mesopleuron and metapleuron, not visible near lateral protrusions of metanotum. Protarsal segments of specimen from China, Bawangling National Nature Reserve, in following proportions: 10:2.5:4:10:17; in holotype segment 1 of protarsus as long as segment 4. Enlarged claw ( Plate 113F ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 113F ) with two rows of 12–17 lamellae; distal apex with 11–12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Java, Pekalongan, F. Muir leg. ( AMNH) . Other material. CHINA: Hainan, Bawangling National Nature Reserve, Reserve House , about 1000 m, parasitized host collected 19.VIII.2000, dryinid cocooning 22.VIII.2000, M. Olmi reared from an unidentified species of Delphacidae , 1♀ ( MOLC) .

Hosts. In China (Hainan), unidentified species of Delphacidae .

Distribution. China (Hainan), Indonesia (Java).

6. Gonatopus nepalensis ( Olmi, 1986) , comb. nov.

( Plate 113G, H )

Pseudogonatopus nepalensis Olmi 1986: 80 .

Description of Female. Apterous; length 4.8–5.0 mm. Head black or brown, except mandible, clypeus and anterior region of face testaceous; antenna brown, except segments 1–2 testaceous and 10 whitish; occasionally antennal segment 9 whitish; mesosoma and metasoma black; legs black or brown, except chela and meso- and metatarsus testaceous. Antenna clavate; antennal segments in following proportions: 12:6:18:10:7:7:7:6.5:6.5:10.5. Head excavated, dull, granulated; frontal line complete; occipital carina absent. Palpal formula 4/2. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum shiny, sculptured by longitudinal striae. Metanotum transversely striate, with sides protruding; protrusions rounded ( Plate 113G ). Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface and metapleura. Mesopleuron shiny, smooth, unsculptured, not transversely striate. Meso-metapleural suture partly distinct, only present between mesopleuron and metapleuron, not present dorsally on sides of metanotum. Protarsal segments in following proportions: 18:4:8:22:35. Enlarged claw ( Plate 113G ) with one large subapical tooth and one row of seven lamellae. Segment 5 of protarsus ( Plate 113G ) with two rows of about 21 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: NEPAL: Kosi, S of Mangsingma Forest ( MHNG). Paratypes: same locality label as holotype, 1♀ ( AMNH); E Nepal, Kosi, Induwa Kola Valley, 1♀ ( MHNG). Other material. NEPAL: Khala Valley, 2050 m ( PMA). Palaearctic: according to Ponomarenko (1992), G. nepalensis is also present in the following localities: RUSSIA: Transbaikalia, 5.VI.1963, 10.VII.1963, A. Emelianov leg., 2♀♀ ( Ponomarenko , 1992); Fra East, Primorskiy Krai, Gornotayezhnoe, 19.VI.1983, Sciarkov leg., 1♀ ( Ponomarenko , 1992). This quotation is doubtful.

Hosts. Unknown.

Distribution. Nepal (Kosi), Russia (Transbaicalia, Primorskiy Krai).

7. Gonatopus nigricans (R. Perkins, 1905)

( Plates 111B , 114A – D )

Paragonatopus nigricans R. Perkins 1905: 41 ; Kieffer 1914b: 87; Olmi 1979: 218.

Pseudogonatopus melanacrias R. Perkins 1906a: 487 (synonymized by R. Perkins 1912); type locality: Suva ( Viti Levu Island,

Fiji Islands). Gonatopus fulgori Nakagawa 1906: 163 ; type locality: Japan; Guglielmino & Olmi 1997: 181, 184; syn. nov. Pseudogonatopus hospes R. Perkins 1912: 13 (synonymized by He & Xu 2002); type locality: China; Kieffer 1914b: 81;

Swezey 1928: 670; F. Williams 1931: 247; Chiu 1979: 339; Móczár 1979: 77; Olmi 1984: 1290; Olmi 2000a: 101, 111. Gonatopus sauteri Strand 1913: 209 (synonymized by Olmi 1984); type locality: Tainan ( Taiwan, China). Dicondylus sauteri (Strand) : Kieffer 1914b: 77. Pseudogonatopus sogatea Rohwer 1920: 160 (printing-error; holotype labelled sogatae ); type locality: Pusa (Bihar, India);

Kurian 1954b: 417; Olmi 1984: 1254; syn. nov. Haplogonatopus fulgori (Nakagawa) : Esaki & Hashimoto 1930: 21. Pseudozonatopus (sic) hospes R. Perkins : Pagden 1934: 2. Paragonatopus fulgori (Nakagawa) : Mochizuki 1943: 99; Chen 1988: 44. nec Pseudogonatopus nr. hospes R. Perkins : Chandra 1978b: 5. Pseudogonatopus fulgori (Nakagawa) : Olmi 1984: 1194. Pseudogonatopus pusanus Olmi 1984: 1255 ; type locality: Pusa (Bihar, India); He & Xu 2002: 355; syn. nov. Pseudogonatopus nigricans (R. Perkins) : Olmi 1984: 1287; Olmi 1990b: 789; Olmi 1991: 319. Gonatopus nigricans (R. Perkins) : Guglielmino & Olmi 1997: 165, 224–226, 250–252; Olmi 1999b: 28; Olmi 2000a: 94; He &

Xu 2002: 364; Guglielmino & Olmi 2006: 49. Gonatopus sogatae Rohwer : Guglielmino & Olmi 1997: 225, 226; He & Xu 2002: 355. Gonatopus insulanus He & Xu in He et al. 1998: 172; type locality: Qionghai (Hainan, China); syn. nov.

Description of female ( Plate 114A ): apterous; length 2.4–4.4 mm. Head brown-reddish, or brown, or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 or 1–3 yellow; mesosoma black or brown; occasionally posterior margin of pronotum and posterior apex of propodeum testaceous; metasoma black or brown; legs yellow; occasionally clubs of femora and part of coxae brown. In G. pusanus the colour is different: head partly or almost totally brown; antenna brown, except segments 1–2 testaceous; mesosoma testaceous-reddish, except sides of scutum darkened; occasionally part of pronotum darkened; metasoma brown; petiole black; legs testaceous; occasionally club of profemur darkened. In G. sogatae the colour is the following: head brown, except mandible and clypeus yellow; antenna brown, except segment 1 yellow and 2–3 brown-testaceous; mesosoma brown, except scutum reddish and distal apex of propodeum yellow; petiole black; metasoma brown-dark; legs brown-yellow. Intermediate colours are known. Antenna clavate; antennal segments in following proportions: 19:19:52:25:22:22:18:17:15:26. Head ( Plate 114B ) excavated, shiny, granulated or unsculptured; occiput and anterior region of face always granulated; POL = 5; OL = 5; OOL = 14. Palpal formula 2/2, 3/2, 4/2. Pronotum granulated, crossed by strong transverse impression. Scutum granulated, without lateral pointed apophyses. Mesopleura transversely striate. Metanotum flat, very long, about as long as scutum, transversely striate, not hollow behind scutellum ( Plate 114C ), with sides rounded ( Plate 114A ). Metathorax + propodeum shiny, indistinctly sculptured, with or without track of median longitudinal furrow, with posterior surface transversely striate. Metapleuron transversely striate. Meso-metapleural suture obsolete. Protarsal segment 4 shorter than segment 1 (40:47; or 20:27). Enlarged claw ( Plate 114D ) with one large subapical tooth and one row of three–nine lamellae. Segment 5 of protarsus ( Plate 114D ) with two rows of 9–25 lamellae; distal apex with 4–10 lamellae; occasionally lamellae extended continuously to distal apex. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9–2.4 mm. Head brown or black (occasionally brown-reddish), except mandible testaceous; occasionally part of face testaceous; antenna brown (in type of G. insulanus , testaceous-dark); mesosoma and metasoma black or brown (in type of G. insulanus , mesosoma reddish-dark, with propodeum and pleura darker); legs yellow. Antenna filiform; antennal segments in following proportions: 5:5:9:8:8:9:7:8:7:10 (in other specimens 6:5.5:10:9:8:9:8:8:7:10); antennal segment 3 more than three times as long as broad (4.0; 4.3; 5.0). Head shiny, hairy, granulated; frontal line absent; temple distinct; occipital carina absent; POL = 8; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OL (4:3) (in other specimens POL = 7; OL = 3; OOL = 3.5; greatest breadth of posterior ocelli longer than OL (3.5:3)). Palpal formula 2/2, 3/2, 4/2. Scutum shiny, granulated. Notauli complete, posteriorly joint or almost joint in small specimens, separated in large specimens; in very large specimens, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2.5:4; or 2:4). Scutellum and metanotum shiny, finely punctate, unsculptured among punctures. Propodeum reticulate rugose; dorsal surface with median longitudinal furrow and occasionally with two lateral smooth areas. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 111B ) much shorter than paramere. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nigricans : AUSTRALIA: Queensland, Bundaberg, 19.XI.1904, reared ( BPBM); ♀ lectotype (designated by Olmi, 1982) of P. melanacrias : FIJI ISLANDS: Viti Levu Island, Suva, III.1905 , reared from delphacid near Stenocranus , on Zoysia pungens (BPBM) ; ♀ lectotype (designated by Olmi, 1982) of P. hospes : CHINA: no localities, F. Muir leg., reared in Honolulu from Perkinsiella saccharicida from Chinese stock, IV–V.1907 ( BPBM); ♀ holotype of G. sauteri : CHINA: Taiwan, Tainan, IV.1912, H. Sauter leg. ( DEI); ♀ holotype of P. pusanus : INDIA: Bihar, Pusa ( IARI); ♀ holotype of P. sogatae : INDIA: Bihar, Pusa, 6.VIII.1915, reared from Sogata sp. , C.S. Misra leg. ( USNM); ♂ holotype of G. insulanus : CHINA: Hainan, Qionghai, 4.V.1964, Junhua He leg. ( ZJUC). Type material of G. fulgori probably lost. Paratypes of P. nigricans : AUSTRALIA: same locality label as lectotype, 1♀ lebelled for mistake Fiji Islands ( BPBM); same locality label as holotype, parasitized host collected 1.X.1904, dryinid adult emerged 21.X.1904, R. Perkins leg., 1♀ “ paratype ” identified by the same R. Perkins ( BMNH); AUSTRALIA: Queensland, Bundaberg, Sandhills, X.1904, bred, 1♀ ( ANIC). Paratypes of P. melanacrias : same locality label as lectotype, 2♀♀ ( BPBM). Paratypes of P. hospes : same locality label as lectotype, 3♀♀ ( BPBM). Paratype of P. pusanus : same locality label as holotype, 1♀ ( IARI). Paratype of P. sogatae : same locality label as holotype, misidentified ♂ allotype not belonging to the same species of holotype. Other material. CHINA: Fujian, Shaxian, reared from rice planthoppers, 1979–1983, Changming Liu leg., 5♀♀ and 1♂ ( ZJUC); Fujian, IV.1987, 31.VII.1985, Changming Liu leg., 6♀♀ and 10♂ ( ZJUC); Guangdong, Yangjiang, 1979, 1981, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangdong, Xingfeng County, Mt. Yunjinshan, 9.VI.2002, 18.VII.2003, 19.VII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); same locality label, 20.VII.2003, Ping Li leg., 2♀♀ ( ZJUC); same locality label, Zaifu Xu leg., many ♀♀ ( ZJUC, AMNH); Guangdong, Shixing County, Chebaling National Nature Reserve, 21.VIII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, M. Olmi reared from adult of unidentified Delphacid, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); Guangdong, Mt. Nankunshan, 2.VII.2005, Zaifu Xu leg., 1♀ ( ZJUC); Guangdong, Yangjiang, Hailing Town, 1980, Renhuan Hou leg., 2♀♀ and 2♂ ( ZJUC); Guangdong, Huaiji County, 21–22.X.2007, Zaifu Xu leg., 1♀ ( SCAU); Guangdong, Conghua, 28.VI.1978, Junhua He leg., 1♀ ( SCAU); Guangdong, Zhaoqing, V.1998, Yufang Liu leg., 2♀♀ ( SCAU); Guizhou, Sinan, IX.1986, VIII.1991, 5♀♀ ( ZJUC); Guizhou, Sandu, VI–VII.1981, Dongxiang Xie leg., 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Hunan, Yueyang, 15.VIII.1975, Binsen Wang leg., 1♀ ( ZJUC); Guangxi, Luchuan, 30.VII.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Bama, 1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Luocheng, 18.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Yaxian, VI.1966, Junhua He leg., 1♀ ( ZJUC); Hong Kong, Tai Lung Farm, IX–XI.1965, swept above rice, I.W.B. Thornton leg., 1♀ ( BMNH); Macau ( BPBM); same locality label, R. Perkins collection, 2♀♀ ( USNM); Taiwan, Kaohsiung, Kosen ( AMNH, ELKU); Taiwan, Changhus, Erlin ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, Kungkuan ( TARI); Taiwan, Taihoku Pref. (=Taipei), 27.IX.1930, J. Sonan leg., 1♀ ( TARI); Taiwan, Tainan, ShinShi, 11.IX.1979, paddy field, Hirashima leg., 1♀ ( ELKU); Taiwan, Gakko ( ELKU); Itahashi (= Taiwan), 16.IX.1930, J. Sonan leg., 5♀♀ ( TARI); Takeyama (=Jhushan), 6.X.1937, J. Sonan leg., 3♀♀ ( TARI); Inrin (=Yuanlin), 8.X.1937, J. Sonan leg., 39♀♀ ( TARI). INDIA: Chhattisgarh, Raipur ( GCC, AMNH); Delhi, New Delhi ( IARI); Orissa, Cuttack, 1981, 1♀ ( BMNH); Orissa, Puri, 28.XI.1987, S.V. Fowler leg., 1♀ ( BMNH); Tamil Nadu, Tanjavur (=Tanjore) District, 28.IX.1987, ex Nilaparvata lugens on rice, C. Peter leg., 3♀♀ ( BMNH); West Bengal, Hooghly District, Khanakul, 12.XII.1986, in paddy seedbed, 1♀ ( BMNH). INDONESIA: Java, Pekalongan ( BPBM); Java, Dieng Plateau ( BPBM); Borneo, Kalimantan, Paloh ( BMNH). MALAYSIA: Malaya, Selangor, 13.VIII.1928, H.T. Pagden leg., 1♀ ( BMNH); Malaya, Selangor, Kuala Lumpur, 21.XII.1930, H. T. Pagden leg., 1♀ ( BMNH); same locality label, many ♀♀ and ♂ ( NMNH, KYC); Malaya, Seberang Perai ( MARDI); Malaya, Serdang ( DBUS); Sabah, Tuaran ( KYC); Sarawak, 1st. Division, Payah Paloh, 28.I.1965, G.H.L. Rotschild leg., 1♀ ( BMNH). THAILAND: Songkhla, Hadd Tai ( KYC); Non Bua Lampoo, Na Kam Hai ( KYC); Tung Saew ( KYC); Tung Fabad ( KYC); Chainat ( KYC, AMNH); Banmakok ( KYC); Mae Tang ( KYC); Lampamg ( ELKU); Mae-Theng ( ELKU); Hot ( ELKU); Chieng Dao ( ELKU); Thailand, 15.IX.1999, reared from Nilaparvata lugens (Stål) , Sawai leg., 4♂ ( BMNH). Australian region: G. nigricans is known from Australia (Queensland), Fiji Islands, Solomon Islands and USA (Hawaii) ( Olmi, 1984). Introduced in the Hawaii 1906 and 1907 from China, Guangdong, for biological control of Perkinsiella saccharicida Kirkaldy in sugarcane fields ( Olmi, 2000a).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in Australia: Liburnia sp. , Stenocranus sp. ; in China: Laodelphax striatellus (Fallén) ( Yang et al., 1982; He & Xu, 2002), Perkinsiella saccharicida Kirkaldy ; in Malaysia: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in Thailand: Nilaparvata lugens (Stål) ( Chiu, 1979) ; in the USA (Hawaii): Perkinsiella saccharicida Kirkaldy (introduced for biological control).

Biology. Information by Pagden (1934); the parthenogenesis is thelytokous.

Natural enemies. In the USA (Hawaii) ( Swezey, 1919; Timberlake, 1919; Muir, 1921; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Encyrtidae : Helegonatopus pseudophanes R. Perkins, 1906 ; Cheiloneurus flaccus (Walker 1847) ( Guerrieri & Viggiani, 2005; Guerrieri, 2006); Aphelinidae : Centrodora xiphidii (R. Perkins, 1906) . In Malaya ( Pagden, 1934; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Ceraphronidae : Allomicrops abnormis (R. Perkins, 1910) ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp.

Distribution. China ( Taiwan, Hainan, Fujian, Guangdong, Guizhou, Hubei, Hunan, Guangxi, Hong Kong, Macau), Australia (Queensland), Fiji Islands ( Viti Levu), India (Bihar, Chhattisgarh, Delhi, Orissa, Tamil Nadu), Indonesia (Java, Pekalongan, Borneo), Malaysia (Malaya, Sabah, Sarawak), Thailand (Songkhla, Non Bua Lampoo, Tung Saew, Tung Fabad, Chainat, Banmakok, Mae Tang, Lampamg, Mae-Theng, Chieng Dao).

Remarks. Gonatopus insulanus He & Xu is here synonymized with G. nigricans , because of the study of its type and comparison with male specimens of G. nigricans . There are no differences.

Gonatopus nigricans (= Pseudogonatopus hospes R. Perkins ) was established in the Hawaii from cocoons sent from China ( Perkins, 1912). This species was considered extinct in the Hawaii ( Olmi, 1999a). However, in 2010 one of the authors (Olmi) saw a male specimen collected in Oahu Island, Mt. Olympus, in April 1981 by a pan trap. Probably so this species is always present at least in Oahu Island, although rare. Gonatopus pusanus and G. sogatae are here synonymized with G. nigricans because the only differences concern the colour of mesosoma (black in nigricans , testaceous to reddish, with some darkened areas, in pusanus ; brown, with some areas reddish or yellow, in sogatae ). In the collections, there are also many specimens with intermediate colours, so that the synonymy is evident. Gonatopus fulgori is here synonymized with G. nigricans , because the original description is like that of G. nigricans (the type material of G. fulgori is lost). In the past these two species were separated, because attributed to two different zoogeographic regions ( fulgori in Palaearctic, nigricans in Oriental and Australian regions).

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

18. Genus Gonatopus Ljungh, 1810

Gonatopus Ljungh 1810: 161 ; Richards 1937: 118; Olmi 1993b: 77; Olmi 1994: 66; Olmi 1995c: 787; Olmi 1995g: 5; Olmi 1996b: 182; Olmi 1998c: 72; Olmi 1998d: 177.; Olmi 1999a: 231; He & Xu 2002: 318.; Virla & Olmi 2008: 371.; Moya- Raygoza & Olmi 2010: 100.

Dicondylus Haliday in Curtis 1829 –1930: 110 (synonymized by Olmi, 1993b); type species: Dicondylus bicolor (Haliday in Curtis 1828), designated by Kieffer (1914b); Haliday 1833: 273; Walker 1837: 411; Kieffer 1914b: 72; Richards 1935: 147; Richards 1937: 118; Olmi 1984: 1374; Olmi 1993b: 78.

Labeo Haliday 1833: 273 (nom. praeocc., nec Cuvier 1817) (synonymized by Richards 1937); type species: Labeo vitripennis Haliday 1833 , by monotypy; Kieffer in Kieffer & Marshall 1905: 220; Walker 1837: 429; Dalla Torre 1898: 542.

nec Labea Blanchard 1840: 430 .

Labeola Haldeman 1842: 191 , new name for Labeo Haliday, 1833 (synonymized by Krombein 1979 with Dicondylus Haliday, 1829 –1930).

Pseudogonatopus R. Perkins 1905: 34 (synonymized by Olmi 1993b); type species: Pseudogonatopus kurandae R. Perkins, 1905 , by original designation; Kieffer in Kieffer & Marshall 1906: 488; Kieffer 1914b: 79; Richards 1939: 195; J. Perkins 1976: 32; Chandra 1980c: 192; Olmi 1984: 1191; Olmi 1993b: 78.

Paragonatopus R. Perkins 1905: 40 (synonymized by Olmi 1984); type species: Paragonatopus nigricans R. Perkins 1905 , by monotypy; Kieffer in Kieffer & Marshall 1906: 488; Kieffer 1914b: 87; Richards 1939: 195; Olmi 1979: 218.

Neogonatopus R. Perkins 1905: 42 (synonymized by Olmi 1984); type species: Neogonatopus ombrodes R. Perkins 1905 , by original designation.

Pachygonatopus R. Perkins 1905: 45 (synonymized by Olmi 1984); type species: Pachygonatopus melanias R. Perkins 1905 , by original designation.

Chalcogonatopus R. Perkins 1905: 46 (synonymized by Olmi 1984); type species: Chalcogonatopus gigas R. Perkins 1905 , by original designation.

Eugonatopus R. Perkins 1905: 46 (synonymized by Olmi 1984); type species: Chalcogonatopus (Eugonatopus) pseudochromus R. Perkins 1905 , by original designation.

Mercetia Kieffer in Kieffer & Marshall 1906: 495 (synonymized by Olmi 1996b); type species: Mercetia longicollis (Kieffer 1905) , by original designation; Olmi 1996b: 182; Kieffer 1914b: 19.

Platygonatopus Kieffer in Kieffer & Marshall 1906: 500 (synonymized by Olmi, 1984); type species: Platygonatopus planiceps (Kieffer 1904) , designated by Richards (1939).

Apterodryinus R. Perkins 1907: 14 (synonymized by Olmi 1993b); type species: Apterodryinus torvus R. Perkins 1907 , by monotypy; Olmi 1984: 1342; Olmi 1993b: 78.

Agonatopus R. Perkins 1907: 29 (synonymized by Olmi 1984); type species: Agonatopus pallidicornis R. Perkins 1907 , designated by Richards (1939).

Agonatopoides R. Perkins 1907: 33 (synonymized by Olmi 1993b); type species: Agonatopoides synchromus R. Perkins 1907 , by monotypy; Olmi 1984: 1310; Olmi 1993b: 78.

Cyrtogonatopus Kieffer 1907: 19 (synonymized by Olmi 1984); type species: Cyrtogonatopus breviforceps (Kieffer 1904) , monotypic.

nec Trichogonatopus Kieffer 1909: 336 .

Pristogonatopus Kieffer 1913c: 325 (synonymized by Olmi & Currado 1974); type species: Pristogonatopus dentatiforceps (Kieffer 1904) , by original designation; Kieffer 1914b: 71; Richards 1939: 195.

Digonatopus Kieffer 1913c: 325 (synonymized by Richards 1939); type species: Digonatopus javanus (R. Perkins 1912) , by monotypy.

Trigonatopus Kieffer 1913c: 325 (synonymized by Olmi 1984); type species: Trigonatopus bifarius (Kieffer in Kieffer & Marshall 1906), by original designation; Kieffer 1914b: 72.

Laberius Kieffer 1914b: 59 , new name for Labeo Haliday 1833 (synonymized by Richards 1937); Richards 1936: 169.

Trichogonatopus Hellén 1930: 2 (synonymized by Olmi 1984); type species: Trichogonatopus lindbergi Hellén 1930 , original designation.

Laberinus Ogloblin 1932: 266 (mispelling).

Metagonatopus Ogloblin 1932: 264 (synonymized by Olmi 1984); type species: Metagonatopus brasiliensis Ogloblin 1932 , by original designation.

Allogonatopus Haupt 1938: 27 (synonymized by Olmi 1984); type species: Allogonatopus procerus Haupt 1938 , by original designation.

Donisthorpina Richards 1939: 201 (synonymized by Olmi 1993b); type species: Donisthorpina formicicola Richards 1939 , by original designation; Olmi 1984: 1304; Olmi 1993b: 78.

Plectrogonatopus Richards 1939: 207 (synonymized by Olmi 1984); type species: Plectrogonatopus striatus (Kieffer 1905) , by original designation.

Tetrodontochelys Richards 1939: 217 (synonymized by Olmi 1993b); type species: Tetrodontochelys ljunghii ( Westwood 1833) , by original designation; Olmi 1984: 1434; Olmi 1993b: 78.

Epigonatopoides Richards 1939: 221 (synonymized by Olmi 1984); type species: Epigonatopoides pulicarius ( Klug 1810) , by original designation.

Idologonatopus Ogloblin 1953: 111 (synonymized by Olmi 1984); type species: Idologonatopus nigrithorax Ogloblin 1953 , by original designation.

Rhynchogonatopus Benoit 1954: 390 (synonymized by Olmi 1984); type species: Rhynchogonatopus meridionalis Benoit 1954 , by original designation.

Madecagonatopus Benoit 1954: 392 (synonymized by Olmi 1984); type species: Madecagonatopus ankaratrus ( Ceballos 1936) , by original designation.

Pseudogonatopoides J. Williams 1956: 90 (synonymized by Olmi 1984); type species: Pseudogonatopoides mauritianus J. Williams 1956 , by original designation.

Cyrtogonatopoides Ponomarenko 1966: 1579 (synonymized by Olmi 1984); type species: Cyrtogonatopoides borealis Ponomarenko 1966 , by original designation.

Plectrogonatopoides Ponomarenko 1975a: 318 (synonymized by Olmi 1993b); type species: Plectrogonatopoides chortinis Ponomarenko 1975 , by original designation; Olmi 1984: 1431; Olmi 1993b: 78.

Tetradontochelys J. Perkins 1976: 35 (injustified emendation).

Neogonatopoides Abdul-Nour 1976: 274 (synonymized by Olmi 1984); type species: Neogonatopoides claudiae Abdul-Nour 1976 , by monotypy.

Dolichochelys Olmi & Currado 1976: 177 (synonymized by Olmi 1984); type species: Dolichochelys longicornis (Kieffer 1904) , by original designation.

Megagonatopus Olmi & Currado 1976: 178 (synonymized by Olmi 1984); type species: Megagonatopus birmanus Olmi & Currado 1976 , by original designation.

Acrodontochelys Currado 1976: 13 (synonymized by Olmi 1993b); type species: Acrodontochelys bouceki Currado 1976 , by original designation; Olmi 1984: 1172; Olmi 1993b: 78.

Nogatopus Ponomarenko 1979: 357 (synonymized by Olmi 1984); type species: Nogatopus gobiensis Ponomarenko 1979 , by original designation.

Paradicondylus Olmi 1986: 102 (synonymized by Olmi 1998c); type species: Paradicondylus paulyi Olmi 1986 , by monotypy and original designation; Olmi 1993b: 78; Olmi 1998c: 72.

Type species. Gonatopus formicarius Ljungh, 1810 , by monotypy.

Diagnosis. Female: apterous ( Plates 106B , 126A ) or fully winged ( Plate 106A ); palpal formula 3/2, 4/2, 4/3, 5/ 2, 5/3, or 6/3; antenna with or without rhinaria; pronotum crossed or not by transverse furrow ( Plate 108C, D ); enlarged claw with distal apex pointed and with one large or small subapical tooth ( Plate 107B, C ) (occasionally subapical tooth absent, then enlarged claw with distal group of lamellae ( Plate 107A )); in fully winged forms, segment 5 of protarsus with more than 20 lamellae; tibial spurs 1/0/1. Male: fully winged ( Plate 110A, B ); occipital carina absent or incomplete (in this last case, only present behind and shortly on sides of posterior ocelli); occiput concave; temple present; palpal formula 3/2, 4/2, 4/3, 5/2, 5/3, or 6/3; tibial spurs 1/1/2.

Distribution. Worldwide.

Hosts. Acanaloniidae , Cicadellidae (except Typhlocybinae , Idiocerinae , Macropsinae ), Delphacidae , Flatidae , Issidae , Lophopidae , Tropiduchidae ( Guglielmino & Olmi, 1997, 2006, 2007).

World species. Four hundred and thirty nine species are known, fifty eight species in the Oriental region and one species incertae sedis.

Remarks. Larvae of Gonatopus were described by Ponomarenko (1975b), Carcupino et al. (1996, 1998), Guglielmino & Virla (1998), Virla & Mangione (2000), Guglielmino (2002), Mangione & Virla (2004) and Guglielmino et al. (2006, 2007, 2008). Gonatopus females were divided for convenience by Olmi (1993b) into eleven groups on the basis of the following characters:

Key to the groups of Gonatopus

1. Fully winged ( Plate 106A )....................................................................... Group 11

- Apterous ( Plate 106B )................................................................................. 2

2. Enlarged claw without subapical tooth ( Plate 107A ), or with one very small subapical tooth ( Plate 107B )............... 3

- Enlarged claw with one large subapical tooth ( Plate 107C )..................................................... 5

3. Distal apex of enlarged claw with group of lamellae ( Plate 107A )......................................... Group 8

- Distal apex of enlarged claw without lamellae ( Plate 107C, D )................................................. 4

4. Pronotum not crossed by transverse impression or slightly impressed ( Plate 108A )........................... Group 6

- Pronotum crossed by strong transverse impression ( Plate 108B ).......................................... Group 7

5. Pronotum not crossed by transverse impression or slightly impressed ( Plate 108C )................................. 6

- Pronotum crossed by strong transverse impression ( Plate 108D )................................................ 7

6. Labial palpi 3-segmented......................................................................... Group 9

- Labial palpi 2-segmented......................................................................... Group 5

7. Enlarged claw without lamellae, with or without bristles or peg-like hairs ( Plate 109B )........................ Group 1

- Enlarged claw with lamellae ( Plate 107C ).................................................................. 8

8. Labial palpi 2-segmented............................................................................... 9

- Labial palpi 3-segmented.............................................................................. 10

9. Maxillary palpi composed of two–four segments...................................................... Group 2

- Maxillary palpi 5-segmented..................................................................... Group 10

10. Maxillary palpi 6-segmented...................................................................... Group 4

- Maxillary palpi 5-segmented...................................................................... Group 3

The above groups are distributed in the zoogeographical regions as follows:

Palaearctic: groups 1, 2, 3, 4, 5, 6, 7, 10, 11.

Afrotropical: groups 1, 2, 3, 4, 5, 6, 7, 8, 11.

Oriental: groups 1, 2, 3, 4, 5, 6, 7, 10.

Nearctic: groups 1, 2, 3, 4, 5, 6, 7, 9.

Neotropical: groups 1, 2, 3, 4, 5, 6, 7, 10.

Australian: groups 1, 2, 4, 5, 6, 7, 9.

As far as, it is not possible divide also males into the above groups.

All Oriental species of Gonatopus can be attributed to one of the above eight female groups, except Gonatopus leurosipropodeus He & Xu, 2002 . The attribution of this species to a distinct group is impossible, because it was described on the basis of male specimens and its female is unknown. For this reason, it is attributed to an uncertain group (see “Uncertain group”).

Key to the males

(unknown for the species not present in the key) (males not divided into groups)

1. Notauli incomplete.................................................................................... 2

- Notauli complete..................................................................................... 4

2. Propodeum with dorsal surface dull, granulated, sculptured by irregular striae, and posterior surface shiny, smooth and unsculptured; dorsal surface of propodeum much shorter than posterior surface...................... G. maurus Kieffer

- Propodeum completely smooth and shiny, unsculptured; dorsal surface of propodeum much longer than posterior surface... 3

3. Dorsal process of paramere transverse ( Plate 122C ); clypeus testaceous.............................. G.lucens (Olmi)

- Dorsal process of paramere about parallel to penis, not transverse ( Plate 111A ); clypeus black....... G. nearcticus (Fenton)

4. Notauli posteriorly broadly separated..................................................................... 5

- Notauli posteriorly joint (or very close and almost joint)..................................................... 14

5. Dorsal process of paramere reduced to inner expansion of paramere ( Plate 122B )....... G. flavifemur (Esaki & Hashimoto)

- Dorsal process of paramere not reduced ( Plates 111B , 118D , 119D , 128A , 131A, B )................................ 6

6. Dorsal process of paramere much shorter than paramere ( Plates 128A , 131A, B )................................... 7

- Dorsal process of paramere as long as, or longer, or slightly shorter than paramere ( Plates 111B , 118D , 119D )............ 9

7. Head with POL more than four times as long as OOL; ocelli very large............................ G. rufoniger Olmi

- Head with POL less than three times as long as OOL; ocelli small............................................... 8

8. Dorsal process of paramere very slender ( Plate 131B ); clypeus completely black...................... G. superbus Olmi

- Dorsal process of paramere less slender ( Plate 128A ); clypeus medially black, laterally testaceous... G.javanus (R. Perkins)

9. Face completely or almost completely testaceous or yellow................................................... 10

- Face completely black or brown........................................................................ 11

10. Dorsal process of paramere longer than penis and paramere ( Plate 118D )......................... G. borneanus (Olmi)

- Dorsal process of paramere shorter than penis and paramere ( Plate 111B )...................... G. nigricans (R. Perkins)

11. Antennal segment 3 less than three times as long as broad; dorsal process of parameres longer than penis ( Plate 119D )............................................................................................ G. pyrillae (Mani)

- Antennal segment 3 more than three times as long as broad; dorsal process of parameres shorter than penis ( Plates 111B , 117C , 122D )............................................................................................. 12

12. Dorsal process of paramere almost completely joint to paramere ( Plate 111B ).................. G. nigricans (R. Perkins)

- Dorsal process of paramere completely separated from paramere ( Plates 117C , 122D ).............................. 13

13. Dorsal process of paramere slender ( Plate 122D )............................................. G. pedestris Dalman

- Dorsal process of paramere broader ( Plate 117C )............ G.malesiae (Olmi) type B or G.sarawakensis (Olmi) type B

14. Dorsal process of paramere very reduced ( Plates 117A , 122B ), reduced to inner expansion of paramere................ 15

- Dorsal process of paramere not reduced ( Plates 111B , 117B ).................................................. 16

15. Inner expansion of paramere with many irregular striae ( Plate 122B )................. G. flavifemur (Esaki & Hashimoto)

- Inner expansion of paramere without irregular striae ( Plate 117A )............................... G. nudus (R. Perkins)

16. Dorsal process of paramere much shorter than paramere ( Plates 111B , 131C )..................................... 17

- Dorsal process of paramere approximately as long as paramere ( Plates 117B , 131D )............................... 18

17. Dorsal process of paramere not transverse, with distal apex broad ( Plate 111B ); head and mesosoma mostly black or brown................................................................................... G. nigricans (R. Perkins)

- Dorsal process of paramere transverse, with distal apex pointed ( Plate 131C ); head and mesosoma mostly yellow-testaceous....................................................................................... G. asiaticus (Olmi)

18. Propodeum completely reticulate rugose; dorsal process of paramere broader ( Plate 117B )................................................................................ G.malesiae (Olmi) type A or G. sarawakensis (Olmi) type A

- Propodeum with posterior inclined region reticulate rugose and anterior flat region smooth and unsculptured; dorsal process of paramere slender ( Plate 131D ).................................................... G. leurosipropodeus He & Xu

Remarks. In the above key, there are two species, whose males are not distinguishable, G. malesiae (Olmi) and

G. sarawakensis (Olmi) ; the situation is explained in remarks related to G. sarawakensis (see Group 2).

Group 1

One species is known in the Oriental region.

1. Gonatopus nearcticus (Fenton, 1905)

( Plates 109A, B , 110A, B , 111A )

nec Pseudogonatopus americanus R. Perkins 1905: 37 .

Epigonatopus americanus Fenton 1921: 70 (preoccupied by Pseudogonatopus americanus R. Perkins 1905 ) (synonymized by Olmi 1993b); type locality: Ames (Iowa, USA).

Pachygonatopus nearcticus Fenton 1927: 6 .

Platygonatopus ugandanus Benoit 1951a: 300 (synonymized by Olmi 1993b); type locality: Bululo ( Uganda); Benoit 1951d: 19.

Rhynchogonatopus ugandanus (Benoit) : Olmi & Currado 1976: 173.

Acrodontochelys bouceki Currado 1976: 14 (synonymized by Olmi 1993b); type locality: Nerja ( Spain); Graham 1983: 1; Olmi 1984: 1174; Graham 1988: 79.

Acrodontochelys ugandanus (Benoit) : Olmi 1984: 1174.

Acrodontochelys sinensis Olmi 1984: 1180 (synonymized by Xu et al. 2012b); type locality: Macau ( China) .

Acrodontochelys americanus (Fenton) : Olmi 1984: 1182.

Gonatopus nearcticus (Fenton) : Olmi 1993b: 80, 83; Guglielmino & Olmi 1997: 194, 237; Tussac & Olmi 1998: 490; Olmi 1999a: 246; Olmi 2004a: 368; Olmi 2005b: 126; Olmi 2006: 44; Guglielmino & Olmi 2007: 126; Olmi 2009b: 459; Burn 2011: 88; Penati & Olmi 2012: 79; Xu et al. 2012b: 21.

Gonatopus sinensis (Olmi) : He & Xu 2002: 349.

Description of female ( Plate 109A ). Apterous; length 2.0–4.0 mm. Head black or brown, with mandible, clypeus and anterior region of face yellow or testaceous; antenna black or brown, with segments 1–3 yellow or testaceous; occasionally antenna testaceous-brown; mesosoma, petiole and metasoma usually black, except posterior apex of propodeum testaceous; occasionally mesosoma and metasoma brown-testaceous or testaceous-reddish, with petiole black; occasionally mesosoma brown, with scutum yellow; legs yellow or testaceous, with coxae and clubs of femora almost completely black, or darkened, or brown-reddish. Antenna clavate; antennal segments of holotype of P. nearcticus in following proportions: 4:4:9:5.5:5:5:4:4:4:7; antennal segments of holotype of A. sinensis in following proportions: 5:3:7:4:4:4:4:4:3.5:5. Head flat or slightly convex, dull, weakly or strongly granulated; frontal line complete; occipital carina absent; temple distinct; POL = 3; OL = 2; OOL = 5. Palpal formula 5/2, 4/2, 3/2, or 2/2. Pronotum crossed by strong transverse impression, shiny, weakly or strongly granulated. Scutum dull, granulated. Metanotum granulated and transversely striate, not hollow behind scutellum. Metathorax + propodeum dull, granulated, with posterior surface of propodeum transversely striate; occasionally posterior surface almost completely granulated and not transversely striate. Meso-metapleural suture obsolete. Mesopleuron and metapleuron granulated, with or without transverse striae. Protarsus with first segment about twice as long as segment 4 (14:7; or 15:7; or 12:7). Protarsal segments of A. sinensis holotype in following proportions: 15:2:3:7:13. Enlarged claw ( Plate 109B ) with one large subapical tooth and one row of three–six peg-like bristles. Segment 5 of protarsus ( Plate 109B ) with two rows of 6–22 lamellae; distal apex with about six–nine lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 110A, B ): fully winged; length 1.6–2.8 mm. Head black or brown, except mandible testaceous or yellow; antenna brown or black; mesosoma and metasoma black or brown; legs banded, brown or black, with trochanters, stalks of femora, extremities of tibiae and tarsi yellow. Antenna hairy, filiform; antennal segments of male paratype of A. bouceki in following proportions: 5:4:8:7:7:7:6:6:6:8; antennal segment 3 four or more than four times as long as broad. Antennal segments of specimen from China, Ruyang, in following proportions: 4:4:7:7:6.5:6.5:5.5:5:5:7. Head shiny, alutaceous, slightly punctate; vertex laterally without two shiny and smooth areas situated between posterior ocelli and eyes; frontal line incomplete, only present between antennal toruli; occipital carina absent; occiput concave; temple distinct; POL = 6; OL = 4; OOL = 2 (or POL = 6; OL = 2; OOL = 2; or POL = 7; OL = 3; OOL = 1.5); greatest breadth of posterior ocelli about twice as long as OOL. Palpal formula 3/2, 4/2. Scutum dull, hairy, slightly granulated. Notauli incomplete, reaching approximately 0.4–0.7 length of scutum. Scutellum and metanotum shiny, smooth, finely punctate, unsculptured among punctures. Propodeum completely shiny, smooth, unsculptured; dorsal surface of propodeum with or without median longitudinal furrow. Fore wing hyaline, without dark transverse bands; stigmal vein regularly curved, with distal part much longer than proximal part. Dorsal process of paramere ( Plate 111A ) more or less long and slender, with distal third parallel to penis. Tibial spurs 1/1/2.

Material examined. Types: ♀ holotype of E. americanus : USA: Iowa, Story County, Ames, 25.VII.1919, E.D. Ball leg. ( USNM); ♀ holotype of P. nearcticus : USA: Woodbury County, Sioux City, IX.1919, reared from a jassid, C.N. Ainslie leg. ( USNM); ♀ holotype of P. ugandanus : UGANDA: Bululo (=present Bulule?), 1909, E. Bayon leg. ( MCSNG); ♀ holotype of A. bouceki : SPAIN: Malaga, Nerja, 3.VII.1974, Z. Bouček leg. ( BMNH); ♀ holotype of A. sinensis : CHINA: Macau ( BPBM). Paratypes of E. americanus : same locality label as holotype, 4♀♀ ( USNM). Paratypes of A. bouceki : same locality label as holoype, 23.VI.1973, Z. Bouček leg., 14♀♀ ( BMNH, DIVAPRA, MRSNT, USNM); SPAIN: Granada, La Herradura, 24.VI.1973, Z. Bouček leg., 1♀ and 1♂ ( BMNH); SPAIN: Granada, Calahonda, 22.VI.1973, Z. Bouček leg., 2♀♀ ( BMNH); SPAIN: Barcelona, Calella de la Costa, VI.1974, 1♀ ( BMNH); SPAIN: Castellón, Benicasim, 13–15.VI.1973, 22–24.VI.1974, Z. Bouček leg., 1♀ and 1♂ ( BMNH). Paratypes of A. sinensis : same locality label as holotype, 3♀♀ ( AMNH, BPBM); CHINA: Chin San, 1♀ ( BPBM); VIET NAM: Lam Dong Prov., Di Linh, Plateau of Djiring, 1♀ ( AMNH). Other material. CHINA: Guangdong, Mt. Nankunshan, 8.VI.2002, M. Olmi leg., 1♀ ( MOLC); Foot of Nanling National Nature Reserve, Ruyang, reared from adult of unidentified Cicadellid, M. Olmi reared, parasitized host collect 12.IX.2004, dryinid cocooning 13.IX.2004, dryinid adult emerged X.2004, 1♀ and 3♂ ( MOLC); same locality label, parasitized host collect 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 2♂ ( MOLC); Macau, X.1906, R. Perkins collection, 1♀ and 1♂ ( USNM). INDIA: Karnataka, Bangalore, 3.XI.1979, Z. Bouček leg., 1♀ ( BMNH); SINGAPORE ( NMNH). THAILAND: Chumphon Prov., Chumplion, 9.II.2000, Ralf Jochmann leg., 1♀ ( MTC); Mae Hong Son Prov., Soppong, 19°27'N 98°20'E, 1500 m, 7–12.V.1996, S. Becvar leg., 1♀ ( OLL). Palaearctic: G. nearcticus is known from the following countries: Croatia, Cyprus, France, Greece, Italy, Montenegro, Portugal, Spain (including Canary Islands) and Turkey ( Olmi, 1999a). Afrotropical region: G. nearcticus is known from the following countries: Benin, Botswana, Ethiopia, Kenya, Mozambique, Namibia, South Africa, Swaziland, Uganda, Zimbabwe ( Olmi, 1984, 1999a). Nearctic region: G. nearcticus is known from the USA ( Olmi, 1984, 1999a).

Hosts. Cicadellidae ( Guglielmino & Olmi, 1997, 2007): in France: Balclutha frontalis (Ferrari) ; in Italy: Balclutha rosea (Scott) ; in Namibia: Paradorydium spatulatum (Naudé) ; in South Africa: Balclutha rosea (Scott) ; in the USA: Balclutha neglecta (DeLong & Davidson) , Balclutha impicta (Van Duzee) .

Distribution. China ( Macau, Guangdong), India (Karnataka), Thailand (Chumphon, Mae Hong Son), Viet Nam (Lam Dong), in addition to many countries of the Afrotropical, Nearctic and Western Palaearctic regions.

Group 2

Ten species are known in the Oriental region.

Key to the females

1. Mesosoma totally black or black-brown (at most distal apex of propodeum and posterior margin of pronotum testaceous or reddish)............................................................................................... 2

- Mesosoma at least partly testaceous-reddish, or testaceous-ferruginous, or brown-reddish, or yellow................... 4

2. Metanotum with sides protruding; protrusions rounded ( Plate 113G )............................. G. nepalensis (Olmi)

- Metanotum with sides rounded ( Plates 112A , 114A ).......................................................... 3

3. Species large (body length about 5.0 mm); body covered with long sparse hairs ( Plate 112A ); temple absent ( Plate 112A ); head more excavated....................................................................... G. capillus Xu & He

- Species small (body length less than 4.5 mm); body hairless or covered with very short hairs; temple distinct ( Plate 114A ); head less excavated ( Plate 114B )...................................................... G. nigricans (R. Perkins)

4. Metanotum with sides rounded ( Plates 114A , 115A ); meso-metapleural suture obsolete.............................. 5

- Metanotum with sides protruding ( Plate 117G ); protrusions rounded ( Plates 111C , 117G ) or pointed ( Plate 113C ); meso-metapleural suture at least partly distinct....................................................................... 6

5. Metanotum short, about as long as scutellum............................................... G. nudus (R. Perkins)

- Metanotum long, about twice as long as scutellum........................................ G. nigricans (R. Perkins)

6. Anterior surface of metathorax + propodeum with numerous longitudinal keels.................. G. sarawakensis (Olmi)

- Anterior surface of metathorax + propodeum smooth, with no keels............................................. 7

7. Metathorax + propodeum with slight or strong median furrow.................................................. 8

- Metathorax + propodeum without median furrow............................................................ 9

8. Metanotum with lateral protrusions more prominent ( Plate 113A ); metathorax + propodeum with deep median furrow; mesometapleural suture very strong and distinct............................................. G.lankae (Ponomarenko)

- Metanotum with lateral protrusions less prominent ( Plate 113E ); metathorax + propodeum with shallow median furrow; mesometapleural suture slightly distinct, only visible near lateral protrusions of metanotum.................. G. muiri (Olmi)

9. Metanotum with lateral protrusions less prominent ( Plate 117G )................................... G. validus (Olmi)

- Metanotum with lateral protrusions more prominent ( Plates 111C , 113C )....................................... 10

10. Lateral protrusions of metanotum at right angle ( Plate 111C ); mesopleuron and metapleuron transversely striate................................................................................................. G.bicuspis (Olmi)

- Lateral protrusions of metanotum not at right angle ( Plate 113C ); mesopleuron and metapleuron smooth, not transversely striate................................................................................ G. malesiae (Olmi)

1. Gonatopus bicuspis (Olmi, 1993) , comb. nov.

( Plate 111C, D )

Pseudogonatopus bicuspis Olmi 1993d: 50 .

Description of Female. Apterous; length 3.2 mm. Testaceous-ferruginous, except petiole black, antennal segment 8 testaceous and antennal segments 9–10 whitish. Antenna clavate; antennal segments in following proportions: 10:6:15:8:6:6:5.5:5:5:8.5. Head very excavated, shiny, smooth, unsculptured; frontal line absent; occipital carina absent; temple absent; POL = 0.3; OL = 3; OOL = 6. Palpal formula 3/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Metanotum flat, transversely striate, with two lateral pointed apophyses ( Plate 111C ). Metathorax + propodeum shiny, smooth, unsculptured; posterior surface of propodeum transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 13:3:4.5:15.5:25. Enlarged claw ( Plate 111D ) with one large subapical tooth and one row of six lamellae + one bristle. Segment 5 of protarsus ( Plate 111D ) with two rows of 16 lamellae; distal apex with about 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: PHILIPPINES: Negros Island , Camp, 16.V.1987 ( AMNH) .

Hosts. Unknown.

Distribution. Philippines (Negros).

2. Gonatopus capillus Xu & He, 1998

( Plate 112A, B )

Gonatopus capillus Xu & He 1998d: 306 ; He & Xu 2002: 351; Guglielmino & Olmi 2006: 49.

Description of female ( Plate 112A ). Apterous; length 5.0 mm. Head black, except mandible testaceous; antenna black, except segment 1 partly testaceous and partly brown and segment 10 yellow-whitish; mesosoma, petiole, metasoma and legs black. Body completely covered with long hairs (as in some species of Trichogonatopus Kieffer ). Antenna clavate; antennal segments in following proportions: 6:3.6:14.5:9:8.5:8:6:6:6:8. Head shiny, smooth, unsculptured, except ocellar region granulated; frontal line incomplete; occipital carina absent; temple absent; POL = 1; OL = 0.5; OOL = 5. Palpal formula 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum and metanotum without lateral pointed apophyses. Metanotum flat, not hollow behind scutellum. Metathorax + propodeum with anterior region smooth, disc reticulate rugose and posterior region transversely striate; anterior region and disc with track of long median longitudinal furrow. Mesopleura and metapleura transversely striate. Meso-metapleural suture almost completely absent, only present on sides of metanotum. Fore tarsal segments in following proportions: 10:1.5:3:10:15.5. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 112B ) with one large subapical tooth and one row of 15 lamellae. Segment 5 of protarsus ( Plate 112B ) with two rows of 12 + 14 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: CHINA: Guizhou, Sinan , IX.1986, Yuxiang Chen leg. ( ZJUC) . Paratype: same label as holotype, 1♀ ( ZJUC) .

Hosts. Delphacidae ( Guglielmino & Olmi, 2006) : in China: Nilaparvata lugens (Stål) ( Xu & He, 1998d) .

Distribution. China (Guizhou).

3. Gonatopus lankae (Ponomarenko, 1981) , comb. nov.

( Plate 113A, B )

Pseudogonatopus lankae Ponomarenko 1981b: 881 ; Olmi 1984: 1249.

Description of Female. Apterous; length 3.2–3.3 mm. Brown-reddish or testaceous-dark; antenna usually darkened, except segment 10 yellow; legs testaceous, except trochanters and procoxa partly whitish, hind side of metafemur whitish, distal apex of propodeum whitish. Antenna clavate; antennal segments in following proportions: 14:8:14:9:9:6:6:6:6:11. Head excavated, dull, alutaceous or weakly granulated; frontal line complete; occipital carina absent; temple prominent; POL = 2; OL = 3; OOL = 10. Palpal formula 4/2 or 5/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Scutellum shiny, smooth, unsculptured. Metanotum flat, shiny, unsculptured, weakly hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded. Metathorax + propodeum ( Plate 113A ) shiny, smooth, unsculptured or granulated; posterior surface of propodeum transversely striate. Disc of metathorax + propodeum with strong median furrow. Mesopleura and metapleura dull, weakly granulated, not transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 15:4:6:20:31. Enlarged claw ( Plate 113B ) with one large subapical tooth and one row of six–eight lamellae. Segment 5 of protarsus ( Plate 113B ) with two rows of 19–21 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Ratnapura District , Gilimale Induruwa Jungle, 13.III.1979, K. Krombein leg. ( USNM) . Other material. SRI LANKA: Ratnapura District ., Sinharaja Jungle, Waturawa Forest ( AMNH) ; Matale District , Sigiriya, 07°57'N 80°46'E, 18.III.1999, C.J. Burwell leg. ( QMC) .

Hosts. Unknown.

Distribution. Sri Lanka (Ratnapura, Matale).

Remarks. The female specimen from Sri Lanka, 07°57'N 80°46'E, has 5-segmented maxillary palps; the palpal formula 5/2 is typical of the group 10 (former genus Donisthorpina Richards ); for this reason, G. lankae is also included into group 10.

4. Gonatopus malesiae ( Olmi, 1984) , comb. nov.

( Plate 113C, D )

Pseudogonatopus malesiae Olmi 1984: 1248 .

Description of Female. Apterous; length 2.8–3.6 mm. Usually testaceous-ferruginous, with petiole black; antenna usually testaceous, with segments 9–10 whitish (occasionally antennal segment 9 partly whitish). In specimen from Myanmar, 17°38.9'N 94°38.1'E, antenna testaceous-brown, except segments 1 and 10 whitish. In specimen from China, Mt. Jinggangshan, antenna brown, except rhinaria areas testaceous, and mesosoma brown-testaceous. Antenna clavate; antennal segments of holotype in following proportions: 16:13:38:23:18:12:12:13:12:20. Head very excavated, shiny, smooth, unsculptured, hairless; frontal line complete; occipital carina absent; POL = 0.3; OL = 2; OOL = 8.5. Palpal formula 3/2 or 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat, without lateral pointed apophyses. Metanotum shiny, unsculptured, not hollow behind scutellum, with sides protruding; lateral protrusions pointed ( Plate 113C ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum without median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture very slightly distinct. In specimen from Myanmar, 17°38.9'N 94°38.1'E, meso-metapleural suture partly and slightly visible (not visible on sides of metanotum). Segment 1 of protarsus usually shorter than segment 4 (13:16). Enlarged claw ( Plate 113D ) with one large subapical tooth and one row of seven–ten lamellae. Segment 5 of protarsus ( Plate 113D ) with two rows of 12–17 lamellae; distal apex with at least 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown (see Gonatopus sarawakensis (Olmi)) .

Material examined. Type: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu National Park , 5.VIII.1978, P.M. Hammond & J.E. Marshall leg. ( BMNH) . Other material. CHINA: Jiangxi, Mt. Jinggangshan , 13.VIII.2007, Junhua He leg., 1♀ ( SCAU) . MALAYSIA: Malaya, Fraser's Hill , 4200' ( AMNH) . MYANMAR: Rakhine State, NE of Gwa, Elephant Camp in Elephant Sanctuary , 17°38.9'N 94°38.1'E, 3.XII.2000, leaf litter in secondary forest, K.J. Ribardo leg., 2♀♀ ( CAS) .

Hosts. Unknown.

Distribution. China (Jiangxi), Malaysia ( Sarawak, Malaya), Myanmar (Rakhine).

5. Gonatopus muiri ( Olmi, 1984)

( Plate 113E, F )

Pseudogonatopus muiri Olmi 1984: 1251 ; Xu et al. 2011g (Suppl.): 20.

Description of Female. Apterous; length 2.5–3.0 mm. Holotype testaceous–reddish, except metasoma brown and antenna brown, with segments 1–2 testaceous (specimen from China, Bawangling National Nature Reserve, testaceous, except antenna brown, with segments 1–2 testaceous and segment 10 whitish, petiole black and metasoma brown). Antenna clavate; antennal segments of a specimen from China, Bawangling National Nature Reserve, in following proportions: 8:4.5:10:6:5:5:4.5:4.5:4:8. Head excavated; head of holotype dull and slightly granulated; head of specimen from China, Bawangling National Nature Reserve, shiny and unsculptured; frontal line of specimen from China, Bawangling National Nature Reserve, complete; occipital carina absent; head of specimen from China, Bawangling National Nature Reserve, with POL = 0.5; OL = 1.5; OOL = 6. Palpal formula 3/ 2 in holotype, 4/ 2 in specimen from China, Bawangling National Nature Reserve. Pronotum crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat. Metanotum shiny, unsculptured, not hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded ( Plate 113E ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum with slight median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture weakly distinct, in holotype only visible near lateral protrusions of metanotum. Meso-metapleural suture of specimen from China, Bawangling National Nature Reserve, visible only between mesopleuron and metapleuron, not visible near lateral protrusions of metanotum. Protarsal segments of specimen from China, Bawangling National Nature Reserve, in following proportions: 10:2.5:4:10:17; in holotype segment 1 of protarsus as long as segment 4. Enlarged claw ( Plate 113F ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 113F ) with two rows of 12–17 lamellae; distal apex with 11–12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Java, Pekalongan, F. Muir leg. ( AMNH) . Other material. CHINA: Hainan, Bawangling National Nature Reserve, Reserve House , about 1000 m, parasitized host collected 19.VIII.2000, dryinid cocooning 22.VIII.2000, M. Olmi reared from an unidentified species of Delphacidae , 1♀ ( MOLC) .

Hosts. In China (Hainan), unidentified species of Delphacidae .

Distribution. China (Hainan), Indonesia (Java).

6. Gonatopus nepalensis ( Olmi, 1986) , comb. nov.

( Plate 113G, H )

Pseudogonatopus nepalensis Olmi 1986: 80 .

Description of Female. Apterous; length 4.8–5.0 mm. Head black or brown, except mandible, clypeus and anterior region of face testaceous; antenna brown, except segments 1–2 testaceous and 10 whitish; occasionally antennal segment 9 whitish; mesosoma and metasoma black; legs black or brown, except chela and meso- and metatarsus testaceous. Antenna clavate; antennal segments in following proportions: 12:6:18:10:7:7:7:6.5:6.5:10.5. Head excavated, dull, granulated; frontal line complete; occipital carina absent. Palpal formula 4/2. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum shiny, sculptured by longitudinal striae. Metanotum transversely striate, with sides protruding; protrusions rounded ( Plate 113G ). Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface and metapleura. Mesopleuron shiny, smooth, unsculptured, not transversely striate. Meso-metapleural suture partly distinct, only present between mesopleuron and metapleuron, not present dorsally on sides of metanotum. Protarsal segments in following proportions: 18:4:8:22:35. Enlarged claw ( Plate 113G ) with one large subapical tooth and one row of seven lamellae. Segment 5 of protarsus ( Plate 113G ) with two rows of about 21 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: NEPAL: Kosi, S of Mangsingma Forest ( MHNG). Paratypes: same locality label as holotype, 1♀ ( AMNH); E Nepal, Kosi, Induwa Kola Valley, 1♀ ( MHNG). Other material. NEPAL: Khala Valley, 2050 m ( PMA). Palaearctic: according to Ponomarenko (1992), G. nepalensis is also present in the following localities: RUSSIA: Transbaikalia, 5.VI.1963, 10.VII.1963, A. Emelianov leg., 2♀♀ ( Ponomarenko , 1992); Fra East, Primorskiy Krai, Gornotayezhnoe, 19.VI.1983, Sciarkov leg., 1♀ ( Ponomarenko , 1992). This quotation is doubtful.

Hosts. Unknown.

Distribution. Nepal (Kosi), Russia (Transbaicalia, Primorskiy Krai).

7. Gonatopus nigricans (R. Perkins, 1905)

( Plates 111B , 114A – D )

Paragonatopus nigricans R. Perkins 1905: 41 ; Kieffer 1914b: 87; Olmi 1979: 218.

Pseudogonatopus melanacrias R. Perkins 1906a: 487 (synonymized by R. Perkins 1912); type locality: Suva ( Viti Levu Island,

Fiji Islands). Gonatopus fulgori Nakagawa 1906: 163 ; type locality: Japan; Guglielmino & Olmi 1997: 181, 184; syn. nov. Pseudogonatopus hospes R. Perkins 1912: 13 (synonymized by He & Xu 2002); type locality: China; Kieffer 1914b: 81;

Swezey 1928: 670; F. Williams 1931: 247; Chiu 1979: 339; Móczár 1979: 77; Olmi 1984: 1290; Olmi 2000a: 101, 111. Gonatopus sauteri Strand 1913: 209 (synonymized by Olmi 1984); type locality: Tainan ( Taiwan, China). Dicondylus sauteri (Strand) : Kieffer 1914b: 77. Pseudogonatopus sogatea Rohwer 1920: 160 (printing-error; holotype labelled sogatae ); type locality: Pusa (Bihar, India);

Kurian 1954b: 417; Olmi 1984: 1254; syn. nov. Haplogonatopus fulgori (Nakagawa) : Esaki & Hashimoto 1930: 21. Pseudozonatopus (sic) hospes R. Perkins : Pagden 1934: 2. Paragonatopus fulgori (Nakagawa) : Mochizuki 1943: 99; Chen 1988: 44. nec Pseudogonatopus nr. hospes R. Perkins : Chandra 1978b: 5. Pseudogonatopus fulgori (Nakagawa) : Olmi 1984: 1194. Pseudogonatopus pusanus Olmi 1984: 1255 ; type locality: Pusa (Bihar, India); He & Xu 2002: 355; syn. nov. Pseudogonatopus nigricans (R. Perkins) : Olmi 1984: 1287; Olmi 1990b: 789; Olmi 1991: 319. Gonatopus nigricans (R. Perkins) : Guglielmino & Olmi 1997: 165, 224–226, 250–252; Olmi 1999b: 28; Olmi 2000a: 94; He &

Xu 2002: 364; Guglielmino & Olmi 2006: 49. Gonatopus sogatae Rohwer : Guglielmino & Olmi 1997: 225, 226; He & Xu 2002: 355. Gonatopus insulanus He & Xu in He et al. 1998: 172; type locality: Qionghai (Hainan, China); syn. nov.

Description of female ( Plate 114A ): apterous; length 2.4–4.4 mm. Head brown-reddish, or brown, or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 or 1–3 yellow; mesosoma black or brown; occasionally posterior margin of pronotum and posterior apex of propodeum testaceous; metasoma black or brown; legs yellow; occasionally clubs of femora and part of coxae brown. In G. pusanus the colour is different: head partly or almost totally brown; antenna brown, except segments 1–2 testaceous; mesosoma testaceous-reddish, except sides of scutum darkened; occasionally part of pronotum darkened; metasoma brown; petiole black; legs testaceous; occasionally club of profemur darkened. In G. sogatae the colour is the following: head brown, except mandible and clypeus yellow; antenna brown, except segment 1 yellow and 2–3 brown-testaceous; mesosoma brown, except scutum reddish and distal apex of propodeum yellow; petiole black; metasoma brown-dark; legs brown-yellow. Intermediate colours are known. Antenna clavate; antennal segments in following proportions: 19:19:52:25:22:22:18:17:15:26. Head ( Plate 114B ) excavated, shiny, granulated or unsculptured; occiput and anterior region of face always granulated; POL = 5; OL = 5; OOL = 14. Palpal formula 2/2, 3/2, 4/2. Pronotum granulated, crossed by strong transverse impression. Scutum granulated, without lateral pointed apophyses. Mesopleura transversely striate. Metanotum flat, very long, about as long as scutum, transversely striate, not hollow behind scutellum ( Plate 114C ), with sides rounded ( Plate 114A ). Metathorax + propodeum shiny, indistinctly sculptured, with or without track of median longitudinal furrow, with posterior surface transversely striate. Metapleuron transversely striate. Meso-metapleural suture obsolete. Protarsal segment 4 shorter than segment 1 (40:47; or 20:27). Enlarged claw ( Plate 114D ) with one large subapical tooth and one row of three–nine lamellae. Segment 5 of protarsus ( Plate 114D ) with two rows of 9–25 lamellae; distal apex with 4–10 lamellae; occasionally lamellae extended continuously to distal apex. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9–2.4 mm. Head brown or black (occasionally brown-reddish), except mandible testaceous; occasionally part of face testaceous; antenna brown (in type of G. insulanus , testaceous-dark); mesosoma and metasoma black or brown (in type of G. insulanus , mesosoma reddish-dark, with propodeum and pleura darker); legs yellow. Antenna filiform; antennal segments in following proportions: 5:5:9:8:8:9:7:8:7:10 (in other specimens 6:5.5:10:9:8:9:8:8:7:10); antennal segment 3 more than three times as long as broad (4.0; 4.3; 5.0). Head shiny, hairy, granulated; frontal line absent; temple distinct; occipital carina absent; POL = 8; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OL (4:3) (in other specimens POL = 7; OL = 3; OOL = 3.5; greatest breadth of posterior ocelli longer than OL (3.5:3)). Palpal formula 2/2, 3/2, 4/2. Scutum shiny, granulated. Notauli complete, posteriorly joint or almost joint in small specimens, separated in large specimens; in very large specimens, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2.5:4; or 2:4). Scutellum and metanotum shiny, finely punctate, unsculptured among punctures. Propodeum reticulate rugose; dorsal surface with median longitudinal furrow and occasionally with two lateral smooth areas. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 111B ) much shorter than paramere. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nigricans : AUSTRALIA: Queensland, Bundaberg, 19.XI.1904, reared ( BPBM); ♀ lectotype (designated by Olmi, 1982) of P. melanacrias : FIJI ISLANDS: Viti Levu Island, Suva, III.1905 , reared from delphacid near Stenocranus , on Zoysia pungens (BPBM) ; ♀ lectotype (designated by Olmi, 1982) of P. hospes : CHINA: no localities, F. Muir leg., reared in Honolulu from Perkinsiella saccharicida from Chinese stock, IV–V.1907 ( BPBM); ♀ holotype of G. sauteri : CHINA: Taiwan, Tainan, IV.1912, H. Sauter leg. ( DEI); ♀ holotype of P. pusanus : INDIA: Bihar, Pusa ( IARI); ♀ holotype of P. sogatae : INDIA: Bihar, Pusa, 6.VIII.1915, reared from Sogata sp. , C.S. Misra leg. ( USNM); ♂ holotype of G. insulanus : CHINA: Hainan, Qionghai, 4.V.1964, Junhua He leg. ( ZJUC). Type material of G. fulgori probably lost. Paratypes of P. nigricans : AUSTRALIA: same locality label as lectotype, 1♀ lebelled for mistake Fiji Islands ( BPBM); same locality label as holotype, parasitized host collected 1.X.1904, dryinid adult emerged 21.X.1904, R. Perkins leg., 1♀ “ paratype ” identified by the same R. Perkins ( BMNH); AUSTRALIA: Queensland, Bundaberg, Sandhills, X.1904, bred, 1♀ ( ANIC). Paratypes of P. melanacrias : same locality label as lectotype, 2♀♀ ( BPBM). Paratypes of P. hospes : same locality label as lectotype, 3♀♀ ( BPBM). Paratype of P. pusanus : same locality label as holotype, 1♀ ( IARI). Paratype of P. sogatae : same locality label as holotype, misidentified ♂ allotype not belonging to the same species of holotype. Other material. CHINA: Fujian, Shaxian, reared from rice planthoppers, 1979–1983, Changming Liu leg., 5♀♀ and 1♂ ( ZJUC); Fujian, IV.1987, 31.VII.1985, Changming Liu leg., 6♀♀ and 10♂ ( ZJUC); Guangdong, Yangjiang, 1979, 1981, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangdong, Xingfeng County, Mt. Yunjinshan, 9.VI.2002, 18.VII.2003, 19.VII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); same locality label, 20.VII.2003, Ping Li leg., 2♀♀ ( ZJUC); same locality label, Zaifu Xu leg., many ♀♀ ( ZJUC, AMNH); Guangdong, Shixing County, Chebaling National Nature Reserve, 21.VIII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, M. Olmi reared from adult of unidentified Delphacid, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); Guangdong, Mt. Nankunshan, 2.VII.2005, Zaifu Xu leg., 1♀ ( ZJUC); Guangdong, Yangjiang, Hailing Town, 1980, Renhuan Hou leg., 2♀♀ and 2♂ ( ZJUC); Guangdong, Huaiji County, 21–22.X.2007, Zaifu Xu leg., 1♀ ( SCAU); Guangdong, Conghua, 28.VI.1978, Junhua He leg., 1♀ ( SCAU); Guangdong, Zhaoqing, V.1998, Yufang Liu leg., 2♀♀ ( SCAU); Guizhou, Sinan, IX.1986, VIII.1991, 5♀♀ ( ZJUC); Guizhou, Sandu, VI–VII.1981, Dongxiang Xie leg., 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Hunan, Yueyang, 15.VIII.1975, Binsen Wang leg., 1♀ ( ZJUC); Guangxi, Luchuan, 30.VII.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Bama, 1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Luocheng, 18.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Yaxian, VI.1966, Junhua He leg., 1♀ ( ZJUC); Hong Kong, Tai Lung Farm, IX–XI.1965, swept above rice, I.W.B. Thornton leg., 1♀ ( BMNH); Macau ( BPBM); same locality label, R. Perkins collection, 2♀♀ ( USNM); Taiwan, Kaohsiung, Kosen ( AMNH, ELKU); Taiwan, Changhus, Erlin ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, Kungkuan ( TARI); Taiwan, Taihoku Pref. (=Taipei), 27.IX.1930, J. Sonan leg., 1♀ ( TARI); Taiwan, Tainan, ShinShi, 11.IX.1979, paddy field, Hirashima leg., 1♀ ( ELKU); Taiwan, Gakko ( ELKU); Itahashi (= Taiwan), 16.IX.1930, J. Sonan leg., 5♀♀ ( TARI); Takeyama (=Jhushan), 6.X.1937, J. Sonan leg., 3♀♀ ( TARI); Inrin (=Yuanlin), 8.X.1937, J. Sonan leg., 39♀♀ ( TARI). INDIA: Chhattisgarh, Raipur ( GCC, AMNH); Delhi, New Delhi ( IARI); Orissa, Cuttack, 1981, 1♀ ( BMNH); Orissa, Puri, 28.XI.1987, S.V. Fowler leg., 1♀ ( BMNH); Tamil Nadu, Tanjavur (=Tanjore) District, 28.IX.1987, ex Nilaparvata lugens on rice, C. Peter leg., 3♀♀ ( BMNH); West Bengal, Hooghly District, Khanakul, 12.XII.1986, in paddy seedbed, 1♀ ( BMNH). INDONESIA: Java, Pekalongan ( BPBM); Java, Dieng Plateau ( BPBM); Borneo, Kalimantan, Paloh ( BMNH). MALAYSIA: Malaya, Selangor, 13.VIII.1928, H.T. Pagden leg., 1♀ ( BMNH); Malaya, Selangor, Kuala Lumpur, 21.XII.1930, H. T. Pagden leg., 1♀ ( BMNH); same locality label, many ♀♀ and ♂ ( NMNH, KYC); Malaya, Seberang Perai ( MARDI); Malaya, Serdang ( DBUS); Sabah, Tuaran ( KYC); Sarawak, 1st. Division, Payah Paloh, 28.I.1965, G.H.L. Rotschild leg., 1♀ ( BMNH). THAILAND: Songkhla, Hadd Tai ( KYC); Non Bua Lampoo, Na Kam Hai ( KYC); Tung Saew ( KYC); Tung Fabad ( KYC); Chainat ( KYC, AMNH); Banmakok ( KYC); Mae Tang ( KYC); Lampamg ( ELKU); Mae-Theng ( ELKU); Hot ( ELKU); Chieng Dao ( ELKU); Thailand, 15.IX.1999, reared from Nilaparvata lugens (Stål) , Sawai leg., 4♂ ( BMNH). Australian region: G. nigricans is known from Australia (Queensland), Fiji Islands, Solomon Islands and USA (Hawaii) ( Olmi, 1984). Introduced in the Hawaii 1906 and 1907 from China, Guangdong, for biological control of Perkinsiella saccharicida Kirkaldy in sugarcane fields ( Olmi, 2000a).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in Australia: Liburnia sp. , Stenocranus sp. ; in China: Laodelphax striatellus (Fallén) ( Yang et al., 1982; He & Xu, 2002), Perkinsiella saccharicida Kirkaldy ; in Malaysia: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in Thailand: Nilaparvata lugens (Stål) ( Chiu, 1979) ; in the USA (Hawaii): Perkinsiella saccharicida Kirkaldy (introduced for biological control).

Biology. Information by Pagden (1934); the parthenogenesis is thelytokous.

Natural enemies. In the USA (Hawaii) ( Swezey, 1919; Timberlake, 1919; Muir, 1921; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Encyrtidae : Helegonatopus pseudophanes R. Perkins, 1906 ; Cheiloneurus flaccus (Walker 1847) ( Guerrieri & Viggiani, 2005; Guerrieri, 2006); Aphelinidae : Centrodora xiphidii (R. Perkins, 1906) . In Malaya ( Pagden, 1934; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Ceraphronidae : Allomicrops abnormis (R. Perkins, 1910) ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp.

Distribution. China ( Taiwan, Hainan, Fujian, Guangdong, Guizhou, Hubei, Hunan, Guangxi, Hong Kong, Macau), Australia (Queensland), Fiji Islands ( Viti Levu), India (Bihar, Chhattisgarh, Delhi, Orissa, Tamil Nadu), Indonesia (Java, Pekalongan, Borneo), Malaysia (Malaya, Sabah, Sarawak), Thailand (Songkhla, Non Bua Lampoo, Tung Saew, Tung Fabad, Chainat, Banmakok, Mae Tang, Lampamg, Mae-Theng, Chieng Dao).

Remarks. Gonatopus insulanus He & Xu is here synonymized with G. nigricans , because of the study of its type and comparison with male specimens of G. nigricans . There are no differences.

Gonatopus nigricans (= Pseudogonatopus hospes R. Perkins ) was established in the Hawaii from cocoons sent from China ( Perkins, 1912). This species was considered extinct in the Hawaii ( Olmi, 1999a). However, in 2010 one of the authors (Olmi) saw a male specimen collected in Oahu Island, Mt. Olympus, in April 1981 by a pan trap. Probably so this species is always present at least in Oahu Island, although rare. Gonatopus pusanus and G. sogatae are here synonymized with G. nigricans because the only differences concern the colour of mesosoma (black in nigricans , testaceous to reddish, with some darkened areas, in pusanus ; brown, with some areas reddish or yellow, in sogatae ). In the collections, there are also many specimens with intermediate colours, so that the synonymy is evident. Gonatopus fulgori is here synonymized with G. nigricans , because the original description is like that of G. nigricans (the type material of G. fulgori is lost). In the past these two species were separated, because attributed to two different zoogeographic regions ( fulgori in Palaearctic, nigricans in Oriental and Australian regions).

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

1. Gonatopus nearcticus (Fenton, 1905)

( Plates 109A, B , 110A, B , 111A )

nec Pseudogonatopus americanus R. Perkins 1905: 37 .

Epigonatopus americanus Fenton 1921: 70 (preoccupied by Pseudogonatopus americanus R. Perkins 1905 ) (synonymized by Olmi 1993b); type locality: Ames (Iowa, USA).

Pachygonatopus nearcticus Fenton 1927: 6 .

Platygonatopus ugandanus Benoit 1951a: 300 (synonymized by Olmi 1993b); type locality: Bululo ( Uganda); Benoit 1951d: 19.

Rhynchogonatopus ugandanus (Benoit) : Olmi & Currado 1976: 173.

Acrodontochelys bouceki Currado 1976: 14 (synonymized by Olmi 1993b); type locality: Nerja ( Spain); Graham 1983: 1; Olmi 1984: 1174; Graham 1988: 79.

Acrodontochelys ugandanus (Benoit) : Olmi 1984: 1174.

Acrodontochelys sinensis Olmi 1984: 1180 (synonymized by Xu et al. 2012b); type locality: Macau ( China) .

Acrodontochelys americanus (Fenton) : Olmi 1984: 1182.

Gonatopus nearcticus (Fenton) : Olmi 1993b: 80, 83; Guglielmino & Olmi 1997: 194, 237; Tussac & Olmi 1998: 490; Olmi 1999a: 246; Olmi 2004a: 368; Olmi 2005b: 126; Olmi 2006: 44; Guglielmino & Olmi 2007: 126; Olmi 2009b: 459; Burn 2011: 88; Penati & Olmi 2012: 79; Xu et al. 2012b: 21.

Gonatopus sinensis (Olmi) : He & Xu 2002: 349.

Description of female ( Plate 109A ). Apterous; length 2.0–4.0 mm. Head black or brown, with mandible, clypeus and anterior region of face yellow or testaceous; antenna black or brown, with segments 1–3 yellow or testaceous; occasionally antenna testaceous-brown; mesosoma, petiole and metasoma usually black, except posterior apex of propodeum testaceous; occasionally mesosoma and metasoma brown-testaceous or testaceous-reddish, with petiole black; occasionally mesosoma brown, with scutum yellow; legs yellow or testaceous, with coxae and clubs of femora almost completely black, or darkened, or brown-reddish. Antenna clavate; antennal segments of holotype of P. nearcticus in following proportions: 4:4:9:5.5:5:5:4:4:4:7; antennal segments of holotype of A. sinensis in following proportions: 5:3:7:4:4:4:4:4:3.5:5. Head flat or slightly convex, dull, weakly or strongly granulated; frontal line complete; occipital carina absent; temple distinct; POL = 3; OL = 2; OOL = 5. Palpal formula 5/2, 4/2, 3/2, or 2/2. Pronotum crossed by strong transverse impression, shiny, weakly or strongly granulated. Scutum dull, granulated. Metanotum granulated and transversely striate, not hollow behind scutellum. Metathorax + propodeum dull, granulated, with posterior surface of propodeum transversely striate; occasionally posterior surface almost completely granulated and not transversely striate. Meso-metapleural suture obsolete. Mesopleuron and metapleuron granulated, with or without transverse striae. Protarsus with first segment about twice as long as segment 4 (14:7; or 15:7; or 12:7). Protarsal segments of A. sinensis holotype in following proportions: 15:2:3:7:13. Enlarged claw ( Plate 109B ) with one large subapical tooth and one row of three–six peg-like bristles. Segment 5 of protarsus ( Plate 109B ) with two rows of 6–22 lamellae; distal apex with about six–nine lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 110A, B ): fully winged; length 1.6–2.8 mm. Head black or brown, except mandible testaceous or yellow; antenna brown or black; mesosoma and metasoma black or brown; legs banded, brown or black, with trochanters, stalks of femora, extremities of tibiae and tarsi yellow. Antenna hairy, filiform; antennal segments of male paratype of A. bouceki in following proportions: 5:4:8:7:7:7:6:6:6:8; antennal segment 3 four or more than four times as long as broad. Antennal segments of specimen from China, Ruyang, in following proportions: 4:4:7:7:6.5:6.5:5.5:5:5:7. Head shiny, alutaceous, slightly punctate; vertex laterally without two shiny and smooth areas situated between posterior ocelli and eyes; frontal line incomplete, only present between antennal toruli; occipital carina absent; occiput concave; temple distinct; POL = 6; OL = 4; OOL = 2 (or POL = 6; OL = 2; OOL = 2; or POL = 7; OL = 3; OOL = 1.5); greatest breadth of posterior ocelli about twice as long as OOL. Palpal formula 3/2, 4/2. Scutum dull, hairy, slightly granulated. Notauli incomplete, reaching approximately 0.4–0.7 length of scutum. Scutellum and metanotum shiny, smooth, finely punctate, unsculptured among punctures. Propodeum completely shiny, smooth, unsculptured; dorsal surface of propodeum with or without median longitudinal furrow. Fore wing hyaline, without dark transverse bands; stigmal vein regularly curved, with distal part much longer than proximal part. Dorsal process of paramere ( Plate 111A ) more or less long and slender, with distal third parallel to penis. Tibial spurs 1/1/2.

Material examined. Types: ♀ holotype of E. americanus : USA: Iowa, Story County, Ames, 25.VII.1919, E.D. Ball leg. ( USNM); ♀ holotype of P. nearcticus : USA: Woodbury County, Sioux City, IX.1919, reared from a jassid, C.N. Ainslie leg. ( USNM); ♀ holotype of P. ugandanus : UGANDA: Bululo (=present Bulule?), 1909, E. Bayon leg. ( MCSNG); ♀ holotype of A. bouceki : SPAIN: Malaga, Nerja, 3.VII.1974, Z. Bouček leg. ( BMNH); ♀ holotype of A. sinensis : CHINA: Macau ( BPBM). Paratypes of E. americanus : same locality label as holotype, 4♀♀ ( USNM). Paratypes of A. bouceki : same locality label as holoype, 23.VI.1973, Z. Bouček leg., 14♀♀ ( BMNH, DIVAPRA, MRSNT, USNM); SPAIN: Granada, La Herradura, 24.VI.1973, Z. Bouček leg., 1♀ and 1♂ ( BMNH); SPAIN: Granada, Calahonda, 22.VI.1973, Z. Bouček leg., 2♀♀ ( BMNH); SPAIN: Barcelona, Calella de la Costa, VI.1974, 1♀ ( BMNH); SPAIN: Castellón, Benicasim, 13–15.VI.1973, 22–24.VI.1974, Z. Bouček leg., 1♀ and 1♂ ( BMNH). Paratypes of A. sinensis : same locality label as holotype, 3♀♀ ( AMNH, BPBM); CHINA: Chin San, 1♀ ( BPBM); VIET NAM: Lam Dong Prov., Di Linh, Plateau of Djiring, 1♀ ( AMNH). Other material. CHINA: Guangdong, Mt. Nankunshan, 8.VI.2002, M. Olmi leg., 1♀ ( MOLC); Foot of Nanling National Nature Reserve, Ruyang, reared from adult of unidentified Cicadellid, M. Olmi reared, parasitized host collect 12.IX.2004, dryinid cocooning 13.IX.2004, dryinid adult emerged X.2004, 1♀ and 3♂ ( MOLC); same locality label, parasitized host collect 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 2♂ ( MOLC); Macau, X.1906, R. Perkins collection, 1♀ and 1♂ ( USNM). INDIA: Karnataka, Bangalore, 3.XI.1979, Z. Bouček leg., 1♀ ( BMNH); SINGAPORE ( NMNH). THAILAND: Chumphon Prov., Chumplion, 9.II.2000, Ralf Jochmann leg., 1♀ ( MTC); Mae Hong Son Prov., Soppong, 19°27'N 98°20'E, 1500 m, 7–12.V.1996, S. Becvar leg., 1♀ ( OLL). Palaearctic: G. nearcticus is known from the following countries: Croatia, Cyprus, France, Greece, Italy, Montenegro, Portugal, Spain (including Canary Islands) and Turkey ( Olmi, 1999a). Afrotropical region: G. nearcticus is known from the following countries: Benin, Botswana, Ethiopia, Kenya, Mozambique, Namibia, South Africa, Swaziland, Uganda, Zimbabwe ( Olmi, 1984, 1999a). Nearctic region: G. nearcticus is known from the USA ( Olmi, 1984, 1999a).

Hosts. Cicadellidae ( Guglielmino & Olmi, 1997, 2007): in France: Balclutha frontalis (Ferrari) ; in Italy: Balclutha rosea (Scott) ; in Namibia: Paradorydium spatulatum (Naudé) ; in South Africa: Balclutha rosea (Scott) ; in the USA: Balclutha neglecta (DeLong & Davidson) , Balclutha impicta (Van Duzee) .

Distribution. China ( Macau, Guangdong), India (Karnataka), Thailand (Chumphon, Mae Hong Son), Viet Nam (Lam Dong), in addition to many countries of the Afrotropical, Nearctic and Western Palaearctic regions.

Group 2

Ten species are known in the Oriental region.

Key to the females

1. Mesosoma totally black or black-brown (at most distal apex of propodeum and posterior margin of pronotum testaceous or reddish)............................................................................................... 2

- Mesosoma at least partly testaceous-reddish, or testaceous-ferruginous, or brown-reddish, or yellow................... 4

2. Metanotum with sides protruding; protrusions rounded ( Plate 113G )............................. G. nepalensis (Olmi)

- Metanotum with sides rounded ( Plates 112A , 114A ).......................................................... 3

3. Species large (body length about 5.0 mm); body covered with long sparse hairs ( Plate 112A ); temple absent ( Plate 112A ); head more excavated....................................................................... G. capillus Xu & He

- Species small (body length less than 4.5 mm); body hairless or covered with very short hairs; temple distinct ( Plate 114A ); head less excavated ( Plate 114B )...................................................... G. nigricans (R. Perkins)

4. Metanotum with sides rounded ( Plates 114A , 115A ); meso-metapleural suture obsolete.............................. 5

- Metanotum with sides protruding ( Plate 117G ); protrusions rounded ( Plates 111C , 117G ) or pointed ( Plate 113C ); meso-metapleural suture at least partly distinct....................................................................... 6

5. Metanotum short, about as long as scutellum............................................... G. nudus (R. Perkins)

- Metanotum long, about twice as long as scutellum........................................ G. nigricans (R. Perkins)

6. Anterior surface of metathorax + propodeum with numerous longitudinal keels.................. G. sarawakensis (Olmi)

- Anterior surface of metathorax + propodeum smooth, with no keels............................................. 7

7. Metathorax + propodeum with slight or strong median furrow.................................................. 8

- Metathorax + propodeum without median furrow............................................................ 9

8. Metanotum with lateral protrusions more prominent ( Plate 113A ); metathorax + propodeum with deep median furrow; mesometapleural suture very strong and distinct............................................. G.lankae (Ponomarenko)

- Metanotum with lateral protrusions less prominent ( Plate 113E ); metathorax + propodeum with shallow median furrow; mesometapleural suture slightly distinct, only visible near lateral protrusions of metanotum.................. G. muiri (Olmi)

9. Metanotum with lateral protrusions less prominent ( Plate 117G )................................... G. validus (Olmi)

- Metanotum with lateral protrusions more prominent ( Plates 111C , 113C )....................................... 10

10. Lateral protrusions of metanotum at right angle ( Plate 111C ); mesopleuron and metapleuron transversely striate................................................................................................. G.bicuspis (Olmi)

- Lateral protrusions of metanotum not at right angle ( Plate 113C ); mesopleuron and metapleuron smooth, not transversely striate................................................................................ G. malesiae (Olmi)

1. Gonatopus bicuspis (Olmi, 1993) , comb. nov.

( Plate 111C, D )

Pseudogonatopus bicuspis Olmi 1993d: 50 .

Description of Female. Apterous; length 3.2 mm. Testaceous-ferruginous, except petiole black, antennal segment 8 testaceous and antennal segments 9–10 whitish. Antenna clavate; antennal segments in following proportions: 10:6:15:8:6:6:5.5:5:5:8.5. Head very excavated, shiny, smooth, unsculptured; frontal line absent; occipital carina absent; temple absent; POL = 0.3; OL = 3; OOL = 6. Palpal formula 3/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Metanotum flat, transversely striate, with two lateral pointed apophyses ( Plate 111C ). Metathorax + propodeum shiny, smooth, unsculptured; posterior surface of propodeum transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 13:3:4.5:15.5:25. Enlarged claw ( Plate 111D ) with one large subapical tooth and one row of six lamellae + one bristle. Segment 5 of protarsus ( Plate 111D ) with two rows of 16 lamellae; distal apex with about 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: PHILIPPINES: Negros Island , Camp, 16.V.1987 ( AMNH) .

Hosts. Unknown.

Distribution. Philippines (Negros).

2. Gonatopus capillus Xu & He, 1998

( Plate 112A, B )

Gonatopus capillus Xu & He 1998d: 306 ; He & Xu 2002: 351; Guglielmino & Olmi 2006: 49.

Description of female ( Plate 112A ). Apterous; length 5.0 mm. Head black, except mandible testaceous; antenna black, except segment 1 partly testaceous and partly brown and segment 10 yellow-whitish; mesosoma, petiole, metasoma and legs black. Body completely covered with long hairs (as in some species of Trichogonatopus Kieffer ). Antenna clavate; antennal segments in following proportions: 6:3.6:14.5:9:8.5:8:6:6:6:8. Head shiny, smooth, unsculptured, except ocellar region granulated; frontal line incomplete; occipital carina absent; temple absent; POL = 1; OL = 0.5; OOL = 5. Palpal formula 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum and metanotum without lateral pointed apophyses. Metanotum flat, not hollow behind scutellum. Metathorax + propodeum with anterior region smooth, disc reticulate rugose and posterior region transversely striate; anterior region and disc with track of long median longitudinal furrow. Mesopleura and metapleura transversely striate. Meso-metapleural suture almost completely absent, only present on sides of metanotum. Fore tarsal segments in following proportions: 10:1.5:3:10:15.5. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 112B ) with one large subapical tooth and one row of 15 lamellae. Segment 5 of protarsus ( Plate 112B ) with two rows of 12 + 14 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: CHINA: Guizhou, Sinan , IX.1986, Yuxiang Chen leg. ( ZJUC) . Paratype: same label as holotype, 1♀ ( ZJUC) .

Hosts. Delphacidae ( Guglielmino & Olmi, 2006) : in China: Nilaparvata lugens (Stål) ( Xu & He, 1998d) .

Distribution. China (Guizhou).

3. Gonatopus lankae (Ponomarenko, 1981) , comb. nov.

( Plate 113A, B )

Pseudogonatopus lankae Ponomarenko 1981b: 881 ; Olmi 1984: 1249.

Description of Female. Apterous; length 3.2–3.3 mm. Brown-reddish or testaceous-dark; antenna usually darkened, except segment 10 yellow; legs testaceous, except trochanters and procoxa partly whitish, hind side of metafemur whitish, distal apex of propodeum whitish. Antenna clavate; antennal segments in following proportions: 14:8:14:9:9:6:6:6:6:11. Head excavated, dull, alutaceous or weakly granulated; frontal line complete; occipital carina absent; temple prominent; POL = 2; OL = 3; OOL = 10. Palpal formula 4/2 or 5/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Scutellum shiny, smooth, unsculptured. Metanotum flat, shiny, unsculptured, weakly hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded. Metathorax + propodeum ( Plate 113A ) shiny, smooth, unsculptured or granulated; posterior surface of propodeum transversely striate. Disc of metathorax + propodeum with strong median furrow. Mesopleura and metapleura dull, weakly granulated, not transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 15:4:6:20:31. Enlarged claw ( Plate 113B ) with one large subapical tooth and one row of six–eight lamellae. Segment 5 of protarsus ( Plate 113B ) with two rows of 19–21 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Ratnapura District , Gilimale Induruwa Jungle, 13.III.1979, K. Krombein leg. ( USNM) . Other material. SRI LANKA: Ratnapura District ., Sinharaja Jungle, Waturawa Forest ( AMNH) ; Matale District , Sigiriya, 07°57'N 80°46'E, 18.III.1999, C.J. Burwell leg. ( QMC) .

Hosts. Unknown.

Distribution. Sri Lanka (Ratnapura, Matale).

Remarks. The female specimen from Sri Lanka, 07°57'N 80°46'E, has 5-segmented maxillary palps; the palpal formula 5/2 is typical of the group 10 (former genus Donisthorpina Richards ); for this reason, G. lankae is also included into group 10.

4. Gonatopus malesiae ( Olmi, 1984) , comb. nov.

( Plate 113C, D )

Pseudogonatopus malesiae Olmi 1984: 1248 .

Description of Female. Apterous; length 2.8–3.6 mm. Usually testaceous-ferruginous, with petiole black; antenna usually testaceous, with segments 9–10 whitish (occasionally antennal segment 9 partly whitish). In specimen from Myanmar, 17°38.9'N 94°38.1'E, antenna testaceous-brown, except segments 1 and 10 whitish. In specimen from China, Mt. Jinggangshan, antenna brown, except rhinaria areas testaceous, and mesosoma brown-testaceous. Antenna clavate; antennal segments of holotype in following proportions: 16:13:38:23:18:12:12:13:12:20. Head very excavated, shiny, smooth, unsculptured, hairless; frontal line complete; occipital carina absent; POL = 0.3; OL = 2; OOL = 8.5. Palpal formula 3/2 or 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat, without lateral pointed apophyses. Metanotum shiny, unsculptured, not hollow behind scutellum, with sides protruding; lateral protrusions pointed ( Plate 113C ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum without median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture very slightly distinct. In specimen from Myanmar, 17°38.9'N 94°38.1'E, meso-metapleural suture partly and slightly visible (not visible on sides of metanotum). Segment 1 of protarsus usually shorter than segment 4 (13:16). Enlarged claw ( Plate 113D ) with one large subapical tooth and one row of seven–ten lamellae. Segment 5 of protarsus ( Plate 113D ) with two rows of 12–17 lamellae; distal apex with at least 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown (see Gonatopus sarawakensis (Olmi)) .

Material examined. Type: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu National Park , 5.VIII.1978, P.M. Hammond & J.E. Marshall leg. ( BMNH) . Other material. CHINA: Jiangxi, Mt. Jinggangshan , 13.VIII.2007, Junhua He leg., 1♀ ( SCAU) . MALAYSIA: Malaya, Fraser's Hill , 4200' ( AMNH) . MYANMAR: Rakhine State, NE of Gwa, Elephant Camp in Elephant Sanctuary , 17°38.9'N 94°38.1'E, 3.XII.2000, leaf litter in secondary forest, K.J. Ribardo leg., 2♀♀ ( CAS) .

Hosts. Unknown.

Distribution. China (Jiangxi), Malaysia ( Sarawak, Malaya), Myanmar (Rakhine).

5. Gonatopus muiri ( Olmi, 1984)

( Plate 113E, F )

Pseudogonatopus muiri Olmi 1984: 1251 ; Xu et al. 2011g (Suppl.): 20.

Description of Female. Apterous; length 2.5–3.0 mm. Holotype testaceous–reddish, except metasoma brown and antenna brown, with segments 1–2 testaceous (specimen from China, Bawangling National Nature Reserve, testaceous, except antenna brown, with segments 1–2 testaceous and segment 10 whitish, petiole black and metasoma brown). Antenna clavate; antennal segments of a specimen from China, Bawangling National Nature Reserve, in following proportions: 8:4.5:10:6:5:5:4.5:4.5:4:8. Head excavated; head of holotype dull and slightly granulated; head of specimen from China, Bawangling National Nature Reserve, shiny and unsculptured; frontal line of specimen from China, Bawangling National Nature Reserve, complete; occipital carina absent; head of specimen from China, Bawangling National Nature Reserve, with POL = 0.5; OL = 1.5; OOL = 6. Palpal formula 3/ 2 in holotype, 4/ 2 in specimen from China, Bawangling National Nature Reserve. Pronotum crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat. Metanotum shiny, unsculptured, not hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded ( Plate 113E ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum with slight median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture weakly distinct, in holotype only visible near lateral protrusions of metanotum. Meso-metapleural suture of specimen from China, Bawangling National Nature Reserve, visible only between mesopleuron and metapleuron, not visible near lateral protrusions of metanotum. Protarsal segments of specimen from China, Bawangling National Nature Reserve, in following proportions: 10:2.5:4:10:17; in holotype segment 1 of protarsus as long as segment 4. Enlarged claw ( Plate 113F ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 113F ) with two rows of 12–17 lamellae; distal apex with 11–12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Java, Pekalongan, F. Muir leg. ( AMNH) . Other material. CHINA: Hainan, Bawangling National Nature Reserve, Reserve House , about 1000 m, parasitized host collected 19.VIII.2000, dryinid cocooning 22.VIII.2000, M. Olmi reared from an unidentified species of Delphacidae , 1♀ ( MOLC) .

Hosts. In China (Hainan), unidentified species of Delphacidae .

Distribution. China (Hainan), Indonesia (Java).

6. Gonatopus nepalensis ( Olmi, 1986) , comb. nov.

( Plate 113G, H )

Pseudogonatopus nepalensis Olmi 1986: 80 .

Description of Female. Apterous; length 4.8–5.0 mm. Head black or brown, except mandible, clypeus and anterior region of face testaceous; antenna brown, except segments 1–2 testaceous and 10 whitish; occasionally antennal segment 9 whitish; mesosoma and metasoma black; legs black or brown, except chela and meso- and metatarsus testaceous. Antenna clavate; antennal segments in following proportions: 12:6:18:10:7:7:7:6.5:6.5:10.5. Head excavated, dull, granulated; frontal line complete; occipital carina absent. Palpal formula 4/2. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum shiny, sculptured by longitudinal striae. Metanotum transversely striate, with sides protruding; protrusions rounded ( Plate 113G ). Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface and metapleura. Mesopleuron shiny, smooth, unsculptured, not transversely striate. Meso-metapleural suture partly distinct, only present between mesopleuron and metapleuron, not present dorsally on sides of metanotum. Protarsal segments in following proportions: 18:4:8:22:35. Enlarged claw ( Plate 113G ) with one large subapical tooth and one row of seven lamellae. Segment 5 of protarsus ( Plate 113G ) with two rows of about 21 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: NEPAL: Kosi, S of Mangsingma Forest ( MHNG). Paratypes: same locality label as holotype, 1♀ ( AMNH); E Nepal, Kosi, Induwa Kola Valley, 1♀ ( MHNG). Other material. NEPAL: Khala Valley, 2050 m ( PMA). Palaearctic: according to Ponomarenko (1992), G. nepalensis is also present in the following localities: RUSSIA: Transbaikalia, 5.VI.1963, 10.VII.1963, A. Emelianov leg., 2♀♀ ( Ponomarenko , 1992); Fra East, Primorskiy Krai, Gornotayezhnoe, 19.VI.1983, Sciarkov leg., 1♀ ( Ponomarenko , 1992). This quotation is doubtful.

Hosts. Unknown.

Distribution. Nepal (Kosi), Russia (Transbaicalia, Primorskiy Krai).

7. Gonatopus nigricans (R. Perkins, 1905)

( Plates 111B , 114A – D )

Paragonatopus nigricans R. Perkins 1905: 41 ; Kieffer 1914b: 87; Olmi 1979: 218.

Pseudogonatopus melanacrias R. Perkins 1906a: 487 (synonymized by R. Perkins 1912); type locality: Suva ( Viti Levu Island,

Fiji Islands). Gonatopus fulgori Nakagawa 1906: 163 ; type locality: Japan; Guglielmino & Olmi 1997: 181, 184; syn. nov. Pseudogonatopus hospes R. Perkins 1912: 13 (synonymized by He & Xu 2002); type locality: China; Kieffer 1914b: 81;

Swezey 1928: 670; F. Williams 1931: 247; Chiu 1979: 339; Móczár 1979: 77; Olmi 1984: 1290; Olmi 2000a: 101, 111. Gonatopus sauteri Strand 1913: 209 (synonymized by Olmi 1984); type locality: Tainan ( Taiwan, China). Dicondylus sauteri (Strand) : Kieffer 1914b: 77. Pseudogonatopus sogatea Rohwer 1920: 160 (printing-error; holotype labelled sogatae ); type locality: Pusa (Bihar, India);

Kurian 1954b: 417; Olmi 1984: 1254; syn. nov. Haplogonatopus fulgori (Nakagawa) : Esaki & Hashimoto 1930: 21. Pseudozonatopus (sic) hospes R. Perkins : Pagden 1934: 2. Paragonatopus fulgori (Nakagawa) : Mochizuki 1943: 99; Chen 1988: 44. nec Pseudogonatopus nr. hospes R. Perkins : Chandra 1978b: 5. Pseudogonatopus fulgori (Nakagawa) : Olmi 1984: 1194. Pseudogonatopus pusanus Olmi 1984: 1255 ; type locality: Pusa (Bihar, India); He & Xu 2002: 355; syn. nov. Pseudogonatopus nigricans (R. Perkins) : Olmi 1984: 1287; Olmi 1990b: 789; Olmi 1991: 319. Gonatopus nigricans (R. Perkins) : Guglielmino & Olmi 1997: 165, 224–226, 250–252; Olmi 1999b: 28; Olmi 2000a: 94; He &

Xu 2002: 364; Guglielmino & Olmi 2006: 49. Gonatopus sogatae Rohwer : Guglielmino & Olmi 1997: 225, 226; He & Xu 2002: 355. Gonatopus insulanus He & Xu in He et al. 1998: 172; type locality: Qionghai (Hainan, China); syn. nov.

Description of female ( Plate 114A ): apterous; length 2.4–4.4 mm. Head brown-reddish, or brown, or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 or 1–3 yellow; mesosoma black or brown; occasionally posterior margin of pronotum and posterior apex of propodeum testaceous; metasoma black or brown; legs yellow; occasionally clubs of femora and part of coxae brown. In G. pusanus the colour is different: head partly or almost totally brown; antenna brown, except segments 1–2 testaceous; mesosoma testaceous-reddish, except sides of scutum darkened; occasionally part of pronotum darkened; metasoma brown; petiole black; legs testaceous; occasionally club of profemur darkened. In G. sogatae the colour is the following: head brown, except mandible and clypeus yellow; antenna brown, except segment 1 yellow and 2–3 brown-testaceous; mesosoma brown, except scutum reddish and distal apex of propodeum yellow; petiole black; metasoma brown-dark; legs brown-yellow. Intermediate colours are known. Antenna clavate; antennal segments in following proportions: 19:19:52:25:22:22:18:17:15:26. Head ( Plate 114B ) excavated, shiny, granulated or unsculptured; occiput and anterior region of face always granulated; POL = 5; OL = 5; OOL = 14. Palpal formula 2/2, 3/2, 4/2. Pronotum granulated, crossed by strong transverse impression. Scutum granulated, without lateral pointed apophyses. Mesopleura transversely striate. Metanotum flat, very long, about as long as scutum, transversely striate, not hollow behind scutellum ( Plate 114C ), with sides rounded ( Plate 114A ). Metathorax + propodeum shiny, indistinctly sculptured, with or without track of median longitudinal furrow, with posterior surface transversely striate. Metapleuron transversely striate. Meso-metapleural suture obsolete. Protarsal segment 4 shorter than segment 1 (40:47; or 20:27). Enlarged claw ( Plate 114D ) with one large subapical tooth and one row of three–nine lamellae. Segment 5 of protarsus ( Plate 114D ) with two rows of 9–25 lamellae; distal apex with 4–10 lamellae; occasionally lamellae extended continuously to distal apex. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9–2.4 mm. Head brown or black (occasionally brown-reddish), except mandible testaceous; occasionally part of face testaceous; antenna brown (in type of G. insulanus , testaceous-dark); mesosoma and metasoma black or brown (in type of G. insulanus , mesosoma reddish-dark, with propodeum and pleura darker); legs yellow. Antenna filiform; antennal segments in following proportions: 5:5:9:8:8:9:7:8:7:10 (in other specimens 6:5.5:10:9:8:9:8:8:7:10); antennal segment 3 more than three times as long as broad (4.0; 4.3; 5.0). Head shiny, hairy, granulated; frontal line absent; temple distinct; occipital carina absent; POL = 8; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OL (4:3) (in other specimens POL = 7; OL = 3; OOL = 3.5; greatest breadth of posterior ocelli longer than OL (3.5:3)). Palpal formula 2/2, 3/2, 4/2. Scutum shiny, granulated. Notauli complete, posteriorly joint or almost joint in small specimens, separated in large specimens; in very large specimens, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2.5:4; or 2:4). Scutellum and metanotum shiny, finely punctate, unsculptured among punctures. Propodeum reticulate rugose; dorsal surface with median longitudinal furrow and occasionally with two lateral smooth areas. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 111B ) much shorter than paramere. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nigricans : AUSTRALIA: Queensland, Bundaberg, 19.XI.1904, reared ( BPBM); ♀ lectotype (designated by Olmi, 1982) of P. melanacrias : FIJI ISLANDS: Viti Levu Island, Suva, III.1905 , reared from delphacid near Stenocranus , on Zoysia pungens (BPBM) ; ♀ lectotype (designated by Olmi, 1982) of P. hospes : CHINA: no localities, F. Muir leg., reared in Honolulu from Perkinsiella saccharicida from Chinese stock, IV–V.1907 ( BPBM); ♀ holotype of G. sauteri : CHINA: Taiwan, Tainan, IV.1912, H. Sauter leg. ( DEI); ♀ holotype of P. pusanus : INDIA: Bihar, Pusa ( IARI); ♀ holotype of P. sogatae : INDIA: Bihar, Pusa, 6.VIII.1915, reared from Sogata sp. , C.S. Misra leg. ( USNM); ♂ holotype of G. insulanus : CHINA: Hainan, Qionghai, 4.V.1964, Junhua He leg. ( ZJUC). Type material of G. fulgori probably lost. Paratypes of P. nigricans : AUSTRALIA: same locality label as lectotype, 1♀ lebelled for mistake Fiji Islands ( BPBM); same locality label as holotype, parasitized host collected 1.X.1904, dryinid adult emerged 21.X.1904, R. Perkins leg., 1♀ “ paratype ” identified by the same R. Perkins ( BMNH); AUSTRALIA: Queensland, Bundaberg, Sandhills, X.1904, bred, 1♀ ( ANIC). Paratypes of P. melanacrias : same locality label as lectotype, 2♀♀ ( BPBM). Paratypes of P. hospes : same locality label as lectotype, 3♀♀ ( BPBM). Paratype of P. pusanus : same locality label as holotype, 1♀ ( IARI). Paratype of P. sogatae : same locality label as holotype, misidentified ♂ allotype not belonging to the same species of holotype. Other material. CHINA: Fujian, Shaxian, reared from rice planthoppers, 1979–1983, Changming Liu leg., 5♀♀ and 1♂ ( ZJUC); Fujian, IV.1987, 31.VII.1985, Changming Liu leg., 6♀♀ and 10♂ ( ZJUC); Guangdong, Yangjiang, 1979, 1981, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangdong, Xingfeng County, Mt. Yunjinshan, 9.VI.2002, 18.VII.2003, 19.VII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); same locality label, 20.VII.2003, Ping Li leg., 2♀♀ ( ZJUC); same locality label, Zaifu Xu leg., many ♀♀ ( ZJUC, AMNH); Guangdong, Shixing County, Chebaling National Nature Reserve, 21.VIII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, M. Olmi reared from adult of unidentified Delphacid, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); Guangdong, Mt. Nankunshan, 2.VII.2005, Zaifu Xu leg., 1♀ ( ZJUC); Guangdong, Yangjiang, Hailing Town, 1980, Renhuan Hou leg., 2♀♀ and 2♂ ( ZJUC); Guangdong, Huaiji County, 21–22.X.2007, Zaifu Xu leg., 1♀ ( SCAU); Guangdong, Conghua, 28.VI.1978, Junhua He leg., 1♀ ( SCAU); Guangdong, Zhaoqing, V.1998, Yufang Liu leg., 2♀♀ ( SCAU); Guizhou, Sinan, IX.1986, VIII.1991, 5♀♀ ( ZJUC); Guizhou, Sandu, VI–VII.1981, Dongxiang Xie leg., 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Hunan, Yueyang, 15.VIII.1975, Binsen Wang leg., 1♀ ( ZJUC); Guangxi, Luchuan, 30.VII.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Bama, 1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Luocheng, 18.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Yaxian, VI.1966, Junhua He leg., 1♀ ( ZJUC); Hong Kong, Tai Lung Farm, IX–XI.1965, swept above rice, I.W.B. Thornton leg., 1♀ ( BMNH); Macau ( BPBM); same locality label, R. Perkins collection, 2♀♀ ( USNM); Taiwan, Kaohsiung, Kosen ( AMNH, ELKU); Taiwan, Changhus, Erlin ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, Kungkuan ( TARI); Taiwan, Taihoku Pref. (=Taipei), 27.IX.1930, J. Sonan leg., 1♀ ( TARI); Taiwan, Tainan, ShinShi, 11.IX.1979, paddy field, Hirashima leg., 1♀ ( ELKU); Taiwan, Gakko ( ELKU); Itahashi (= Taiwan), 16.IX.1930, J. Sonan leg., 5♀♀ ( TARI); Takeyama (=Jhushan), 6.X.1937, J. Sonan leg., 3♀♀ ( TARI); Inrin (=Yuanlin), 8.X.1937, J. Sonan leg., 39♀♀ ( TARI). INDIA: Chhattisgarh, Raipur ( GCC, AMNH); Delhi, New Delhi ( IARI); Orissa, Cuttack, 1981, 1♀ ( BMNH); Orissa, Puri, 28.XI.1987, S.V. Fowler leg., 1♀ ( BMNH); Tamil Nadu, Tanjavur (=Tanjore) District, 28.IX.1987, ex Nilaparvata lugens on rice, C. Peter leg., 3♀♀ ( BMNH); West Bengal, Hooghly District, Khanakul, 12.XII.1986, in paddy seedbed, 1♀ ( BMNH). INDONESIA: Java, Pekalongan ( BPBM); Java, Dieng Plateau ( BPBM); Borneo, Kalimantan, Paloh ( BMNH). MALAYSIA: Malaya, Selangor, 13.VIII.1928, H.T. Pagden leg., 1♀ ( BMNH); Malaya, Selangor, Kuala Lumpur, 21.XII.1930, H. T. Pagden leg., 1♀ ( BMNH); same locality label, many ♀♀ and ♂ ( NMNH, KYC); Malaya, Seberang Perai ( MARDI); Malaya, Serdang ( DBUS); Sabah, Tuaran ( KYC); Sarawak, 1st. Division, Payah Paloh, 28.I.1965, G.H.L. Rotschild leg., 1♀ ( BMNH). THAILAND: Songkhla, Hadd Tai ( KYC); Non Bua Lampoo, Na Kam Hai ( KYC); Tung Saew ( KYC); Tung Fabad ( KYC); Chainat ( KYC, AMNH); Banmakok ( KYC); Mae Tang ( KYC); Lampamg ( ELKU); Mae-Theng ( ELKU); Hot ( ELKU); Chieng Dao ( ELKU); Thailand, 15.IX.1999, reared from Nilaparvata lugens (Stål) , Sawai leg., 4♂ ( BMNH). Australian region: G. nigricans is known from Australia (Queensland), Fiji Islands, Solomon Islands and USA (Hawaii) ( Olmi, 1984). Introduced in the Hawaii 1906 and 1907 from China, Guangdong, for biological control of Perkinsiella saccharicida Kirkaldy in sugarcane fields ( Olmi, 2000a).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in Australia: Liburnia sp. , Stenocranus sp. ; in China: Laodelphax striatellus (Fallén) ( Yang et al., 1982; He & Xu, 2002), Perkinsiella saccharicida Kirkaldy ; in Malaysia: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in Thailand: Nilaparvata lugens (Stål) ( Chiu, 1979) ; in the USA (Hawaii): Perkinsiella saccharicida Kirkaldy (introduced for biological control).

Biology. Information by Pagden (1934); the parthenogenesis is thelytokous.

Natural enemies. In the USA (Hawaii) ( Swezey, 1919; Timberlake, 1919; Muir, 1921; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Encyrtidae : Helegonatopus pseudophanes R. Perkins, 1906 ; Cheiloneurus flaccus (Walker 1847) ( Guerrieri & Viggiani, 2005; Guerrieri, 2006); Aphelinidae : Centrodora xiphidii (R. Perkins, 1906) . In Malaya ( Pagden, 1934; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Ceraphronidae : Allomicrops abnormis (R. Perkins, 1910) ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp.

Distribution. China ( Taiwan, Hainan, Fujian, Guangdong, Guizhou, Hubei, Hunan, Guangxi, Hong Kong, Macau), Australia (Queensland), Fiji Islands ( Viti Levu), India (Bihar, Chhattisgarh, Delhi, Orissa, Tamil Nadu), Indonesia (Java, Pekalongan, Borneo), Malaysia (Malaya, Sabah, Sarawak), Thailand (Songkhla, Non Bua Lampoo, Tung Saew, Tung Fabad, Chainat, Banmakok, Mae Tang, Lampamg, Mae-Theng, Chieng Dao).

Remarks. Gonatopus insulanus He & Xu is here synonymized with G. nigricans , because of the study of its type and comparison with male specimens of G. nigricans . There are no differences.

Gonatopus nigricans (= Pseudogonatopus hospes R. Perkins ) was established in the Hawaii from cocoons sent from China ( Perkins, 1912). This species was considered extinct in the Hawaii ( Olmi, 1999a). However, in 2010 one of the authors (Olmi) saw a male specimen collected in Oahu Island, Mt. Olympus, in April 1981 by a pan trap. Probably so this species is always present at least in Oahu Island, although rare. Gonatopus pusanus and G. sogatae are here synonymized with G. nigricans because the only differences concern the colour of mesosoma (black in nigricans , testaceous to reddish, with some darkened areas, in pusanus ; brown, with some areas reddish or yellow, in sogatae ). In the collections, there are also many specimens with intermediate colours, so that the synonymy is evident. Gonatopus fulgori is here synonymized with G. nigricans , because the original description is like that of G. nigricans (the type material of G. fulgori is lost). In the past these two species were separated, because attributed to two different zoogeographic regions ( fulgori in Palaearctic, nigricans in Oriental and Australian regions).

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

1. Gonatopus bicuspis (Olmi, 1993) , comb. nov.

( Plate 111C, D )

Pseudogonatopus bicuspis Olmi 1993d: 50 .

Description of Female. Apterous; length 3.2 mm. Testaceous-ferruginous, except petiole black, antennal segment 8 testaceous and antennal segments 9–10 whitish. Antenna clavate; antennal segments in following proportions: 10:6:15:8:6:6:5.5:5:5:8.5. Head very excavated, shiny, smooth, unsculptured; frontal line absent; occipital carina absent; temple absent; POL = 0.3; OL = 3; OOL = 6. Palpal formula 3/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Metanotum flat, transversely striate, with two lateral pointed apophyses ( Plate 111C ). Metathorax + propodeum shiny, smooth, unsculptured; posterior surface of propodeum transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 13:3:4.5:15.5:25. Enlarged claw ( Plate 111D ) with one large subapical tooth and one row of six lamellae + one bristle. Segment 5 of protarsus ( Plate 111D ) with two rows of 16 lamellae; distal apex with about 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: PHILIPPINES: Negros Island , Camp, 16.V.1987 ( AMNH) .

Hosts. Unknown.

Distribution. Philippines (Negros).

2. Gonatopus capillus Xu & He, 1998

( Plate 112A, B )

Gonatopus capillus Xu & He 1998d: 306 ; He & Xu 2002: 351; Guglielmino & Olmi 2006: 49.

Description of female ( Plate 112A ). Apterous; length 5.0 mm. Head black, except mandible testaceous; antenna black, except segment 1 partly testaceous and partly brown and segment 10 yellow-whitish; mesosoma, petiole, metasoma and legs black. Body completely covered with long hairs (as in some species of Trichogonatopus Kieffer ). Antenna clavate; antennal segments in following proportions: 6:3.6:14.5:9:8.5:8:6:6:6:8. Head shiny, smooth, unsculptured, except ocellar region granulated; frontal line incomplete; occipital carina absent; temple absent; POL = 1; OL = 0.5; OOL = 5. Palpal formula 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum and metanotum without lateral pointed apophyses. Metanotum flat, not hollow behind scutellum. Metathorax + propodeum with anterior region smooth, disc reticulate rugose and posterior region transversely striate; anterior region and disc with track of long median longitudinal furrow. Mesopleura and metapleura transversely striate. Meso-metapleural suture almost completely absent, only present on sides of metanotum. Fore tarsal segments in following proportions: 10:1.5:3:10:15.5. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 112B ) with one large subapical tooth and one row of 15 lamellae. Segment 5 of protarsus ( Plate 112B ) with two rows of 12 + 14 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: CHINA: Guizhou, Sinan , IX.1986, Yuxiang Chen leg. ( ZJUC) . Paratype: same label as holotype, 1♀ ( ZJUC) .

Hosts. Delphacidae ( Guglielmino & Olmi, 2006) : in China: Nilaparvata lugens (Stål) ( Xu & He, 1998d) .

Distribution. China (Guizhou).

3. Gonatopus lankae (Ponomarenko, 1981) , comb. nov.

( Plate 113A, B )

Pseudogonatopus lankae Ponomarenko 1981b: 881 ; Olmi 1984: 1249.

Description of Female. Apterous; length 3.2–3.3 mm. Brown-reddish or testaceous-dark; antenna usually darkened, except segment 10 yellow; legs testaceous, except trochanters and procoxa partly whitish, hind side of metafemur whitish, distal apex of propodeum whitish. Antenna clavate; antennal segments in following proportions: 14:8:14:9:9:6:6:6:6:11. Head excavated, dull, alutaceous or weakly granulated; frontal line complete; occipital carina absent; temple prominent; POL = 2; OL = 3; OOL = 10. Palpal formula 4/2 or 5/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Scutellum shiny, smooth, unsculptured. Metanotum flat, shiny, unsculptured, weakly hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded. Metathorax + propodeum ( Plate 113A ) shiny, smooth, unsculptured or granulated; posterior surface of propodeum transversely striate. Disc of metathorax + propodeum with strong median furrow. Mesopleura and metapleura dull, weakly granulated, not transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 15:4:6:20:31. Enlarged claw ( Plate 113B ) with one large subapical tooth and one row of six–eight lamellae. Segment 5 of protarsus ( Plate 113B ) with two rows of 19–21 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Ratnapura District , Gilimale Induruwa Jungle, 13.III.1979, K. Krombein leg. ( USNM) . Other material. SRI LANKA: Ratnapura District ., Sinharaja Jungle, Waturawa Forest ( AMNH) ; Matale District , Sigiriya, 07°57'N 80°46'E, 18.III.1999, C.J. Burwell leg. ( QMC) .

Hosts. Unknown.

Distribution. Sri Lanka (Ratnapura, Matale).

Remarks. The female specimen from Sri Lanka, 07°57'N 80°46'E, has 5-segmented maxillary palps; the palpal formula 5/2 is typical of the group 10 (former genus Donisthorpina Richards ); for this reason, G. lankae is also included into group 10.

4. Gonatopus malesiae ( Olmi, 1984) , comb. nov.

( Plate 113C, D )

Pseudogonatopus malesiae Olmi 1984: 1248 .

Description of Female. Apterous; length 2.8–3.6 mm. Usually testaceous-ferruginous, with petiole black; antenna usually testaceous, with segments 9–10 whitish (occasionally antennal segment 9 partly whitish). In specimen from Myanmar, 17°38.9'N 94°38.1'E, antenna testaceous-brown, except segments 1 and 10 whitish. In specimen from China, Mt. Jinggangshan, antenna brown, except rhinaria areas testaceous, and mesosoma brown-testaceous. Antenna clavate; antennal segments of holotype in following proportions: 16:13:38:23:18:12:12:13:12:20. Head very excavated, shiny, smooth, unsculptured, hairless; frontal line complete; occipital carina absent; POL = 0.3; OL = 2; OOL = 8.5. Palpal formula 3/2 or 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat, without lateral pointed apophyses. Metanotum shiny, unsculptured, not hollow behind scutellum, with sides protruding; lateral protrusions pointed ( Plate 113C ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum without median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture very slightly distinct. In specimen from Myanmar, 17°38.9'N 94°38.1'E, meso-metapleural suture partly and slightly visible (not visible on sides of metanotum). Segment 1 of protarsus usually shorter than segment 4 (13:16). Enlarged claw ( Plate 113D ) with one large subapical tooth and one row of seven–ten lamellae. Segment 5 of protarsus ( Plate 113D ) with two rows of 12–17 lamellae; distal apex with at least 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown (see Gonatopus sarawakensis (Olmi)) .

Material examined. Type: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu National Park , 5.VIII.1978, P.M. Hammond & J.E. Marshall leg. ( BMNH) . Other material. CHINA: Jiangxi, Mt. Jinggangshan , 13.VIII.2007, Junhua He leg., 1♀ ( SCAU) . MALAYSIA: Malaya, Fraser's Hill , 4200' ( AMNH) . MYANMAR: Rakhine State, NE of Gwa, Elephant Camp in Elephant Sanctuary , 17°38.9'N 94°38.1'E, 3.XII.2000, leaf litter in secondary forest, K.J. Ribardo leg., 2♀♀ ( CAS) .

Hosts. Unknown.

Distribution. China (Jiangxi), Malaysia ( Sarawak, Malaya), Myanmar (Rakhine).

5. Gonatopus muiri ( Olmi, 1984)

( Plate 113E, F )

Pseudogonatopus muiri Olmi 1984: 1251 ; Xu et al. 2011g (Suppl.): 20.

Description of Female. Apterous; length 2.5–3.0 mm. Holotype testaceous–reddish, except metasoma brown and antenna brown, with segments 1–2 testaceous (specimen from China, Bawangling National Nature Reserve, testaceous, except antenna brown, with segments 1–2 testaceous and segment 10 whitish, petiole black and metasoma brown). Antenna clavate; antennal segments of a specimen from China, Bawangling National Nature Reserve, in following proportions: 8:4.5:10:6:5:5:4.5:4.5:4:8. Head excavated; head of holotype dull and slightly granulated; head of specimen from China, Bawangling National Nature Reserve, shiny and unsculptured; frontal line of specimen from China, Bawangling National Nature Reserve, complete; occipital carina absent; head of specimen from China, Bawangling National Nature Reserve, with POL = 0.5; OL = 1.5; OOL = 6. Palpal formula 3/ 2 in holotype, 4/ 2 in specimen from China, Bawangling National Nature Reserve. Pronotum crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat. Metanotum shiny, unsculptured, not hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded ( Plate 113E ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum with slight median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture weakly distinct, in holotype only visible near lateral protrusions of metanotum. Meso-metapleural suture of specimen from China, Bawangling National Nature Reserve, visible only between mesopleuron and metapleuron, not visible near lateral protrusions of metanotum. Protarsal segments of specimen from China, Bawangling National Nature Reserve, in following proportions: 10:2.5:4:10:17; in holotype segment 1 of protarsus as long as segment 4. Enlarged claw ( Plate 113F ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 113F ) with two rows of 12–17 lamellae; distal apex with 11–12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Java, Pekalongan, F. Muir leg. ( AMNH) . Other material. CHINA: Hainan, Bawangling National Nature Reserve, Reserve House , about 1000 m, parasitized host collected 19.VIII.2000, dryinid cocooning 22.VIII.2000, M. Olmi reared from an unidentified species of Delphacidae , 1♀ ( MOLC) .

Hosts. In China (Hainan), unidentified species of Delphacidae .

Distribution. China (Hainan), Indonesia (Java).

6. Gonatopus nepalensis ( Olmi, 1986) , comb. nov.

( Plate 113G, H )

Pseudogonatopus nepalensis Olmi 1986: 80 .

Description of Female. Apterous; length 4.8–5.0 mm. Head black or brown, except mandible, clypeus and anterior region of face testaceous; antenna brown, except segments 1–2 testaceous and 10 whitish; occasionally antennal segment 9 whitish; mesosoma and metasoma black; legs black or brown, except chela and meso- and metatarsus testaceous. Antenna clavate; antennal segments in following proportions: 12:6:18:10:7:7:7:6.5:6.5:10.5. Head excavated, dull, granulated; frontal line complete; occipital carina absent. Palpal formula 4/2. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum shiny, sculptured by longitudinal striae. Metanotum transversely striate, with sides protruding; protrusions rounded ( Plate 113G ). Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface and metapleura. Mesopleuron shiny, smooth, unsculptured, not transversely striate. Meso-metapleural suture partly distinct, only present between mesopleuron and metapleuron, not present dorsally on sides of metanotum. Protarsal segments in following proportions: 18:4:8:22:35. Enlarged claw ( Plate 113G ) with one large subapical tooth and one row of seven lamellae. Segment 5 of protarsus ( Plate 113G ) with two rows of about 21 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: NEPAL: Kosi, S of Mangsingma Forest ( MHNG). Paratypes: same locality label as holotype, 1♀ ( AMNH); E Nepal, Kosi, Induwa Kola Valley, 1♀ ( MHNG). Other material. NEPAL: Khala Valley, 2050 m ( PMA). Palaearctic: according to Ponomarenko (1992), G. nepalensis is also present in the following localities: RUSSIA: Transbaikalia, 5.VI.1963, 10.VII.1963, A. Emelianov leg., 2♀♀ ( Ponomarenko , 1992); Fra East, Primorskiy Krai, Gornotayezhnoe, 19.VI.1983, Sciarkov leg., 1♀ ( Ponomarenko , 1992). This quotation is doubtful.

Hosts. Unknown.

Distribution. Nepal (Kosi), Russia (Transbaicalia, Primorskiy Krai).

7. Gonatopus nigricans (R. Perkins, 1905)

( Plates 111B , 114A – D )

Paragonatopus nigricans R. Perkins 1905: 41 ; Kieffer 1914b: 87; Olmi 1979: 218.

Pseudogonatopus melanacrias R. Perkins 1906a: 487 (synonymized by R. Perkins 1912); type locality: Suva ( Viti Levu Island,

Fiji Islands). Gonatopus fulgori Nakagawa 1906: 163 ; type locality: Japan; Guglielmino & Olmi 1997: 181, 184; syn. nov. Pseudogonatopus hospes R. Perkins 1912: 13 (synonymized by He & Xu 2002); type locality: China; Kieffer 1914b: 81;

Swezey 1928: 670; F. Williams 1931: 247; Chiu 1979: 339; Móczár 1979: 77; Olmi 1984: 1290; Olmi 2000a: 101, 111. Gonatopus sauteri Strand 1913: 209 (synonymized by Olmi 1984); type locality: Tainan ( Taiwan, China). Dicondylus sauteri (Strand) : Kieffer 1914b: 77. Pseudogonatopus sogatea Rohwer 1920: 160 (printing-error; holotype labelled sogatae ); type locality: Pusa (Bihar, India);

Kurian 1954b: 417; Olmi 1984: 1254; syn. nov. Haplogonatopus fulgori (Nakagawa) : Esaki & Hashimoto 1930: 21. Pseudozonatopus (sic) hospes R. Perkins : Pagden 1934: 2. Paragonatopus fulgori (Nakagawa) : Mochizuki 1943: 99; Chen 1988: 44. nec Pseudogonatopus nr. hospes R. Perkins : Chandra 1978b: 5. Pseudogonatopus fulgori (Nakagawa) : Olmi 1984: 1194. Pseudogonatopus pusanus Olmi 1984: 1255 ; type locality: Pusa (Bihar, India); He & Xu 2002: 355; syn. nov. Pseudogonatopus nigricans (R. Perkins) : Olmi 1984: 1287; Olmi 1990b: 789; Olmi 1991: 319. Gonatopus nigricans (R. Perkins) : Guglielmino & Olmi 1997: 165, 224–226, 250–252; Olmi 1999b: 28; Olmi 2000a: 94; He &

Xu 2002: 364; Guglielmino & Olmi 2006: 49. Gonatopus sogatae Rohwer : Guglielmino & Olmi 1997: 225, 226; He & Xu 2002: 355. Gonatopus insulanus He & Xu in He et al. 1998: 172; type locality: Qionghai (Hainan, China); syn. nov.

Description of female ( Plate 114A ): apterous; length 2.4–4.4 mm. Head brown-reddish, or brown, or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 or 1–3 yellow; mesosoma black or brown; occasionally posterior margin of pronotum and posterior apex of propodeum testaceous; metasoma black or brown; legs yellow; occasionally clubs of femora and part of coxae brown. In G. pusanus the colour is different: head partly or almost totally brown; antenna brown, except segments 1–2 testaceous; mesosoma testaceous-reddish, except sides of scutum darkened; occasionally part of pronotum darkened; metasoma brown; petiole black; legs testaceous; occasionally club of profemur darkened. In G. sogatae the colour is the following: head brown, except mandible and clypeus yellow; antenna brown, except segment 1 yellow and 2–3 brown-testaceous; mesosoma brown, except scutum reddish and distal apex of propodeum yellow; petiole black; metasoma brown-dark; legs brown-yellow. Intermediate colours are known. Antenna clavate; antennal segments in following proportions: 19:19:52:25:22:22:18:17:15:26. Head ( Plate 114B ) excavated, shiny, granulated or unsculptured; occiput and anterior region of face always granulated; POL = 5; OL = 5; OOL = 14. Palpal formula 2/2, 3/2, 4/2. Pronotum granulated, crossed by strong transverse impression. Scutum granulated, without lateral pointed apophyses. Mesopleura transversely striate. Metanotum flat, very long, about as long as scutum, transversely striate, not hollow behind scutellum ( Plate 114C ), with sides rounded ( Plate 114A ). Metathorax + propodeum shiny, indistinctly sculptured, with or without track of median longitudinal furrow, with posterior surface transversely striate. Metapleuron transversely striate. Meso-metapleural suture obsolete. Protarsal segment 4 shorter than segment 1 (40:47; or 20:27). Enlarged claw ( Plate 114D ) with one large subapical tooth and one row of three–nine lamellae. Segment 5 of protarsus ( Plate 114D ) with two rows of 9–25 lamellae; distal apex with 4–10 lamellae; occasionally lamellae extended continuously to distal apex. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9–2.4 mm. Head brown or black (occasionally brown-reddish), except mandible testaceous; occasionally part of face testaceous; antenna brown (in type of G. insulanus , testaceous-dark); mesosoma and metasoma black or brown (in type of G. insulanus , mesosoma reddish-dark, with propodeum and pleura darker); legs yellow. Antenna filiform; antennal segments in following proportions: 5:5:9:8:8:9:7:8:7:10 (in other specimens 6:5.5:10:9:8:9:8:8:7:10); antennal segment 3 more than three times as long as broad (4.0; 4.3; 5.0). Head shiny, hairy, granulated; frontal line absent; temple distinct; occipital carina absent; POL = 8; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OL (4:3) (in other specimens POL = 7; OL = 3; OOL = 3.5; greatest breadth of posterior ocelli longer than OL (3.5:3)). Palpal formula 2/2, 3/2, 4/2. Scutum shiny, granulated. Notauli complete, posteriorly joint or almost joint in small specimens, separated in large specimens; in very large specimens, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2.5:4; or 2:4). Scutellum and metanotum shiny, finely punctate, unsculptured among punctures. Propodeum reticulate rugose; dorsal surface with median longitudinal furrow and occasionally with two lateral smooth areas. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 111B ) much shorter than paramere. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nigricans : AUSTRALIA: Queensland, Bundaberg, 19.XI.1904, reared ( BPBM); ♀ lectotype (designated by Olmi, 1982) of P. melanacrias : FIJI ISLANDS: Viti Levu Island, Suva, III.1905 , reared from delphacid near Stenocranus , on Zoysia pungens (BPBM) ; ♀ lectotype (designated by Olmi, 1982) of P. hospes : CHINA: no localities, F. Muir leg., reared in Honolulu from Perkinsiella saccharicida from Chinese stock, IV–V.1907 ( BPBM); ♀ holotype of G. sauteri : CHINA: Taiwan, Tainan, IV.1912, H. Sauter leg. ( DEI); ♀ holotype of P. pusanus : INDIA: Bihar, Pusa ( IARI); ♀ holotype of P. sogatae : INDIA: Bihar, Pusa, 6.VIII.1915, reared from Sogata sp. , C.S. Misra leg. ( USNM); ♂ holotype of G. insulanus : CHINA: Hainan, Qionghai, 4.V.1964, Junhua He leg. ( ZJUC). Type material of G. fulgori probably lost. Paratypes of P. nigricans : AUSTRALIA: same locality label as lectotype, 1♀ lebelled for mistake Fiji Islands ( BPBM); same locality label as holotype, parasitized host collected 1.X.1904, dryinid adult emerged 21.X.1904, R. Perkins leg., 1♀ “ paratype ” identified by the same R. Perkins ( BMNH); AUSTRALIA: Queensland, Bundaberg, Sandhills, X.1904, bred, 1♀ ( ANIC). Paratypes of P. melanacrias : same locality label as lectotype, 2♀♀ ( BPBM). Paratypes of P. hospes : same locality label as lectotype, 3♀♀ ( BPBM). Paratype of P. pusanus : same locality label as holotype, 1♀ ( IARI). Paratype of P. sogatae : same locality label as holotype, misidentified ♂ allotype not belonging to the same species of holotype. Other material. CHINA: Fujian, Shaxian, reared from rice planthoppers, 1979–1983, Changming Liu leg., 5♀♀ and 1♂ ( ZJUC); Fujian, IV.1987, 31.VII.1985, Changming Liu leg., 6♀♀ and 10♂ ( ZJUC); Guangdong, Yangjiang, 1979, 1981, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangdong, Xingfeng County, Mt. Yunjinshan, 9.VI.2002, 18.VII.2003, 19.VII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); same locality label, 20.VII.2003, Ping Li leg., 2♀♀ ( ZJUC); same locality label, Zaifu Xu leg., many ♀♀ ( ZJUC, AMNH); Guangdong, Shixing County, Chebaling National Nature Reserve, 21.VIII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, M. Olmi reared from adult of unidentified Delphacid, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); Guangdong, Mt. Nankunshan, 2.VII.2005, Zaifu Xu leg., 1♀ ( ZJUC); Guangdong, Yangjiang, Hailing Town, 1980, Renhuan Hou leg., 2♀♀ and 2♂ ( ZJUC); Guangdong, Huaiji County, 21–22.X.2007, Zaifu Xu leg., 1♀ ( SCAU); Guangdong, Conghua, 28.VI.1978, Junhua He leg., 1♀ ( SCAU); Guangdong, Zhaoqing, V.1998, Yufang Liu leg., 2♀♀ ( SCAU); Guizhou, Sinan, IX.1986, VIII.1991, 5♀♀ ( ZJUC); Guizhou, Sandu, VI–VII.1981, Dongxiang Xie leg., 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Hunan, Yueyang, 15.VIII.1975, Binsen Wang leg., 1♀ ( ZJUC); Guangxi, Luchuan, 30.VII.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Bama, 1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Luocheng, 18.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Yaxian, VI.1966, Junhua He leg., 1♀ ( ZJUC); Hong Kong, Tai Lung Farm, IX–XI.1965, swept above rice, I.W.B. Thornton leg., 1♀ ( BMNH); Macau ( BPBM); same locality label, R. Perkins collection, 2♀♀ ( USNM); Taiwan, Kaohsiung, Kosen ( AMNH, ELKU); Taiwan, Changhus, Erlin ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, Kungkuan ( TARI); Taiwan, Taihoku Pref. (=Taipei), 27.IX.1930, J. Sonan leg., 1♀ ( TARI); Taiwan, Tainan, ShinShi, 11.IX.1979, paddy field, Hirashima leg., 1♀ ( ELKU); Taiwan, Gakko ( ELKU); Itahashi (= Taiwan), 16.IX.1930, J. Sonan leg., 5♀♀ ( TARI); Takeyama (=Jhushan), 6.X.1937, J. Sonan leg., 3♀♀ ( TARI); Inrin (=Yuanlin), 8.X.1937, J. Sonan leg., 39♀♀ ( TARI). INDIA: Chhattisgarh, Raipur ( GCC, AMNH); Delhi, New Delhi ( IARI); Orissa, Cuttack, 1981, 1♀ ( BMNH); Orissa, Puri, 28.XI.1987, S.V. Fowler leg., 1♀ ( BMNH); Tamil Nadu, Tanjavur (=Tanjore) District, 28.IX.1987, ex Nilaparvata lugens on rice, C. Peter leg., 3♀♀ ( BMNH); West Bengal, Hooghly District, Khanakul, 12.XII.1986, in paddy seedbed, 1♀ ( BMNH). INDONESIA: Java, Pekalongan ( BPBM); Java, Dieng Plateau ( BPBM); Borneo, Kalimantan, Paloh ( BMNH). MALAYSIA: Malaya, Selangor, 13.VIII.1928, H.T. Pagden leg., 1♀ ( BMNH); Malaya, Selangor, Kuala Lumpur, 21.XII.1930, H. T. Pagden leg., 1♀ ( BMNH); same locality label, many ♀♀ and ♂ ( NMNH, KYC); Malaya, Seberang Perai ( MARDI); Malaya, Serdang ( DBUS); Sabah, Tuaran ( KYC); Sarawak, 1st. Division, Payah Paloh, 28.I.1965, G.H.L. Rotschild leg., 1♀ ( BMNH). THAILAND: Songkhla, Hadd Tai ( KYC); Non Bua Lampoo, Na Kam Hai ( KYC); Tung Saew ( KYC); Tung Fabad ( KYC); Chainat ( KYC, AMNH); Banmakok ( KYC); Mae Tang ( KYC); Lampamg ( ELKU); Mae-Theng ( ELKU); Hot ( ELKU); Chieng Dao ( ELKU); Thailand, 15.IX.1999, reared from Nilaparvata lugens (Stål) , Sawai leg., 4♂ ( BMNH). Australian region: G. nigricans is known from Australia (Queensland), Fiji Islands, Solomon Islands and USA (Hawaii) ( Olmi, 1984). Introduced in the Hawaii 1906 and 1907 from China, Guangdong, for biological control of Perkinsiella saccharicida Kirkaldy in sugarcane fields ( Olmi, 2000a).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in Australia: Liburnia sp. , Stenocranus sp. ; in China: Laodelphax striatellus (Fallén) ( Yang et al., 1982; He & Xu, 2002), Perkinsiella saccharicida Kirkaldy ; in Malaysia: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in Thailand: Nilaparvata lugens (Stål) ( Chiu, 1979) ; in the USA (Hawaii): Perkinsiella saccharicida Kirkaldy (introduced for biological control).

Biology. Information by Pagden (1934); the parthenogenesis is thelytokous.

Natural enemies. In the USA (Hawaii) ( Swezey, 1919; Timberlake, 1919; Muir, 1921; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Encyrtidae : Helegonatopus pseudophanes R. Perkins, 1906 ; Cheiloneurus flaccus (Walker 1847) ( Guerrieri & Viggiani, 2005; Guerrieri, 2006); Aphelinidae : Centrodora xiphidii (R. Perkins, 1906) . In Malaya ( Pagden, 1934; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Ceraphronidae : Allomicrops abnormis (R. Perkins, 1910) ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp.

Distribution. China ( Taiwan, Hainan, Fujian, Guangdong, Guizhou, Hubei, Hunan, Guangxi, Hong Kong, Macau), Australia (Queensland), Fiji Islands ( Viti Levu), India (Bihar, Chhattisgarh, Delhi, Orissa, Tamil Nadu), Indonesia (Java, Pekalongan, Borneo), Malaysia (Malaya, Sabah, Sarawak), Thailand (Songkhla, Non Bua Lampoo, Tung Saew, Tung Fabad, Chainat, Banmakok, Mae Tang, Lampamg, Mae-Theng, Chieng Dao).

Remarks. Gonatopus insulanus He & Xu is here synonymized with G. nigricans , because of the study of its type and comparison with male specimens of G. nigricans . There are no differences.

Gonatopus nigricans (= Pseudogonatopus hospes R. Perkins ) was established in the Hawaii from cocoons sent from China ( Perkins, 1912). This species was considered extinct in the Hawaii ( Olmi, 1999a). However, in 2010 one of the authors (Olmi) saw a male specimen collected in Oahu Island, Mt. Olympus, in April 1981 by a pan trap. Probably so this species is always present at least in Oahu Island, although rare. Gonatopus pusanus and G. sogatae are here synonymized with G. nigricans because the only differences concern the colour of mesosoma (black in nigricans , testaceous to reddish, with some darkened areas, in pusanus ; brown, with some areas reddish or yellow, in sogatae ). In the collections, there are also many specimens with intermediate colours, so that the synonymy is evident. Gonatopus fulgori is here synonymized with G. nigricans , because the original description is like that of G. nigricans (the type material of G. fulgori is lost). In the past these two species were separated, because attributed to two different zoogeographic regions ( fulgori in Palaearctic, nigricans in Oriental and Australian regions).

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

2. Gonatopus capillus Xu & He, 1998

( Plate 112A, B )

Gonatopus capillus Xu & He 1998d: 306 ; He & Xu 2002: 351; Guglielmino & Olmi 2006: 49.

Description of female ( Plate 112A ). Apterous; length 5.0 mm. Head black, except mandible testaceous; antenna black, except segment 1 partly testaceous and partly brown and segment 10 yellow-whitish; mesosoma, petiole, metasoma and legs black. Body completely covered with long hairs (as in some species of Trichogonatopus Kieffer ). Antenna clavate; antennal segments in following proportions: 6:3.6:14.5:9:8.5:8:6:6:6:8. Head shiny, smooth, unsculptured, except ocellar region granulated; frontal line incomplete; occipital carina absent; temple absent; POL = 1; OL = 0.5; OOL = 5. Palpal formula 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum and metanotum without lateral pointed apophyses. Metanotum flat, not hollow behind scutellum. Metathorax + propodeum with anterior region smooth, disc reticulate rugose and posterior region transversely striate; anterior region and disc with track of long median longitudinal furrow. Mesopleura and metapleura transversely striate. Meso-metapleural suture almost completely absent, only present on sides of metanotum. Fore tarsal segments in following proportions: 10:1.5:3:10:15.5. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 112B ) with one large subapical tooth and one row of 15 lamellae. Segment 5 of protarsus ( Plate 112B ) with two rows of 12 + 14 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: CHINA: Guizhou, Sinan , IX.1986, Yuxiang Chen leg. ( ZJUC) . Paratype: same label as holotype, 1♀ ( ZJUC) .

Hosts. Delphacidae ( Guglielmino & Olmi, 2006) : in China: Nilaparvata lugens (Stål) ( Xu & He, 1998d) .

Distribution. China (Guizhou).

3. Gonatopus lankae (Ponomarenko, 1981) , comb. nov.

( Plate 113A, B )

Pseudogonatopus lankae Ponomarenko 1981b: 881 ; Olmi 1984: 1249.

Description of Female. Apterous; length 3.2–3.3 mm. Brown-reddish or testaceous-dark; antenna usually darkened, except segment 10 yellow; legs testaceous, except trochanters and procoxa partly whitish, hind side of metafemur whitish, distal apex of propodeum whitish. Antenna clavate; antennal segments in following proportions: 14:8:14:9:9:6:6:6:6:11. Head excavated, dull, alutaceous or weakly granulated; frontal line complete; occipital carina absent; temple prominent; POL = 2; OL = 3; OOL = 10. Palpal formula 4/2 or 5/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Scutellum shiny, smooth, unsculptured. Metanotum flat, shiny, unsculptured, weakly hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded. Metathorax + propodeum ( Plate 113A ) shiny, smooth, unsculptured or granulated; posterior surface of propodeum transversely striate. Disc of metathorax + propodeum with strong median furrow. Mesopleura and metapleura dull, weakly granulated, not transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 15:4:6:20:31. Enlarged claw ( Plate 113B ) with one large subapical tooth and one row of six–eight lamellae. Segment 5 of protarsus ( Plate 113B ) with two rows of 19–21 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Ratnapura District , Gilimale Induruwa Jungle, 13.III.1979, K. Krombein leg. ( USNM) . Other material. SRI LANKA: Ratnapura District ., Sinharaja Jungle, Waturawa Forest ( AMNH) ; Matale District , Sigiriya, 07°57'N 80°46'E, 18.III.1999, C.J. Burwell leg. ( QMC) .

Hosts. Unknown.

Distribution. Sri Lanka (Ratnapura, Matale).

Remarks. The female specimen from Sri Lanka, 07°57'N 80°46'E, has 5-segmented maxillary palps; the palpal formula 5/2 is typical of the group 10 (former genus Donisthorpina Richards ); for this reason, G. lankae is also included into group 10.

4. Gonatopus malesiae ( Olmi, 1984) , comb. nov.

( Plate 113C, D )

Pseudogonatopus malesiae Olmi 1984: 1248 .

Description of Female. Apterous; length 2.8–3.6 mm. Usually testaceous-ferruginous, with petiole black; antenna usually testaceous, with segments 9–10 whitish (occasionally antennal segment 9 partly whitish). In specimen from Myanmar, 17°38.9'N 94°38.1'E, antenna testaceous-brown, except segments 1 and 10 whitish. In specimen from China, Mt. Jinggangshan, antenna brown, except rhinaria areas testaceous, and mesosoma brown-testaceous. Antenna clavate; antennal segments of holotype in following proportions: 16:13:38:23:18:12:12:13:12:20. Head very excavated, shiny, smooth, unsculptured, hairless; frontal line complete; occipital carina absent; POL = 0.3; OL = 2; OOL = 8.5. Palpal formula 3/2 or 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat, without lateral pointed apophyses. Metanotum shiny, unsculptured, not hollow behind scutellum, with sides protruding; lateral protrusions pointed ( Plate 113C ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum without median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture very slightly distinct. In specimen from Myanmar, 17°38.9'N 94°38.1'E, meso-metapleural suture partly and slightly visible (not visible on sides of metanotum). Segment 1 of protarsus usually shorter than segment 4 (13:16). Enlarged claw ( Plate 113D ) with one large subapical tooth and one row of seven–ten lamellae. Segment 5 of protarsus ( Plate 113D ) with two rows of 12–17 lamellae; distal apex with at least 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown (see Gonatopus sarawakensis (Olmi)) .

Material examined. Type: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu National Park , 5.VIII.1978, P.M. Hammond & J.E. Marshall leg. ( BMNH) . Other material. CHINA: Jiangxi, Mt. Jinggangshan , 13.VIII.2007, Junhua He leg., 1♀ ( SCAU) . MALAYSIA: Malaya, Fraser's Hill , 4200' ( AMNH) . MYANMAR: Rakhine State, NE of Gwa, Elephant Camp in Elephant Sanctuary , 17°38.9'N 94°38.1'E, 3.XII.2000, leaf litter in secondary forest, K.J. Ribardo leg., 2♀♀ ( CAS) .

Hosts. Unknown.

Distribution. China (Jiangxi), Malaysia ( Sarawak, Malaya), Myanmar (Rakhine).

5. Gonatopus muiri ( Olmi, 1984)

( Plate 113E, F )

Pseudogonatopus muiri Olmi 1984: 1251 ; Xu et al. 2011g (Suppl.): 20.

Description of Female. Apterous; length 2.5–3.0 mm. Holotype testaceous–reddish, except metasoma brown and antenna brown, with segments 1–2 testaceous (specimen from China, Bawangling National Nature Reserve, testaceous, except antenna brown, with segments 1–2 testaceous and segment 10 whitish, petiole black and metasoma brown). Antenna clavate; antennal segments of a specimen from China, Bawangling National Nature Reserve, in following proportions: 8:4.5:10:6:5:5:4.5:4.5:4:8. Head excavated; head of holotype dull and slightly granulated; head of specimen from China, Bawangling National Nature Reserve, shiny and unsculptured; frontal line of specimen from China, Bawangling National Nature Reserve, complete; occipital carina absent; head of specimen from China, Bawangling National Nature Reserve, with POL = 0.5; OL = 1.5; OOL = 6. Palpal formula 3/ 2 in holotype, 4/ 2 in specimen from China, Bawangling National Nature Reserve. Pronotum crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat. Metanotum shiny, unsculptured, not hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded ( Plate 113E ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum with slight median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture weakly distinct, in holotype only visible near lateral protrusions of metanotum. Meso-metapleural suture of specimen from China, Bawangling National Nature Reserve, visible only between mesopleuron and metapleuron, not visible near lateral protrusions of metanotum. Protarsal segments of specimen from China, Bawangling National Nature Reserve, in following proportions: 10:2.5:4:10:17; in holotype segment 1 of protarsus as long as segment 4. Enlarged claw ( Plate 113F ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 113F ) with two rows of 12–17 lamellae; distal apex with 11–12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Java, Pekalongan, F. Muir leg. ( AMNH) . Other material. CHINA: Hainan, Bawangling National Nature Reserve, Reserve House , about 1000 m, parasitized host collected 19.VIII.2000, dryinid cocooning 22.VIII.2000, M. Olmi reared from an unidentified species of Delphacidae , 1♀ ( MOLC) .

Hosts. In China (Hainan), unidentified species of Delphacidae .

Distribution. China (Hainan), Indonesia (Java).

6. Gonatopus nepalensis ( Olmi, 1986) , comb. nov.

( Plate 113G, H )

Pseudogonatopus nepalensis Olmi 1986: 80 .

Description of Female. Apterous; length 4.8–5.0 mm. Head black or brown, except mandible, clypeus and anterior region of face testaceous; antenna brown, except segments 1–2 testaceous and 10 whitish; occasionally antennal segment 9 whitish; mesosoma and metasoma black; legs black or brown, except chela and meso- and metatarsus testaceous. Antenna clavate; antennal segments in following proportions: 12:6:18:10:7:7:7:6.5:6.5:10.5. Head excavated, dull, granulated; frontal line complete; occipital carina absent. Palpal formula 4/2. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum shiny, sculptured by longitudinal striae. Metanotum transversely striate, with sides protruding; protrusions rounded ( Plate 113G ). Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface and metapleura. Mesopleuron shiny, smooth, unsculptured, not transversely striate. Meso-metapleural suture partly distinct, only present between mesopleuron and metapleuron, not present dorsally on sides of metanotum. Protarsal segments in following proportions: 18:4:8:22:35. Enlarged claw ( Plate 113G ) with one large subapical tooth and one row of seven lamellae. Segment 5 of protarsus ( Plate 113G ) with two rows of about 21 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: NEPAL: Kosi, S of Mangsingma Forest ( MHNG). Paratypes: same locality label as holotype, 1♀ ( AMNH); E Nepal, Kosi, Induwa Kola Valley, 1♀ ( MHNG). Other material. NEPAL: Khala Valley, 2050 m ( PMA). Palaearctic: according to Ponomarenko (1992), G. nepalensis is also present in the following localities: RUSSIA: Transbaikalia, 5.VI.1963, 10.VII.1963, A. Emelianov leg., 2♀♀ ( Ponomarenko , 1992); Fra East, Primorskiy Krai, Gornotayezhnoe, 19.VI.1983, Sciarkov leg., 1♀ ( Ponomarenko , 1992). This quotation is doubtful.

Hosts. Unknown.

Distribution. Nepal (Kosi), Russia (Transbaicalia, Primorskiy Krai).

7. Gonatopus nigricans (R. Perkins, 1905)

( Plates 111B , 114A – D )

Paragonatopus nigricans R. Perkins 1905: 41 ; Kieffer 1914b: 87; Olmi 1979: 218.

Pseudogonatopus melanacrias R. Perkins 1906a: 487 (synonymized by R. Perkins 1912); type locality: Suva ( Viti Levu Island,

Fiji Islands). Gonatopus fulgori Nakagawa 1906: 163 ; type locality: Japan; Guglielmino & Olmi 1997: 181, 184; syn. nov. Pseudogonatopus hospes R. Perkins 1912: 13 (synonymized by He & Xu 2002); type locality: China; Kieffer 1914b: 81;

Swezey 1928: 670; F. Williams 1931: 247; Chiu 1979: 339; Móczár 1979: 77; Olmi 1984: 1290; Olmi 2000a: 101, 111. Gonatopus sauteri Strand 1913: 209 (synonymized by Olmi 1984); type locality: Tainan ( Taiwan, China). Dicondylus sauteri (Strand) : Kieffer 1914b: 77. Pseudogonatopus sogatea Rohwer 1920: 160 (printing-error; holotype labelled sogatae ); type locality: Pusa (Bihar, India);

Kurian 1954b: 417; Olmi 1984: 1254; syn. nov. Haplogonatopus fulgori (Nakagawa) : Esaki & Hashimoto 1930: 21. Pseudozonatopus (sic) hospes R. Perkins : Pagden 1934: 2. Paragonatopus fulgori (Nakagawa) : Mochizuki 1943: 99; Chen 1988: 44. nec Pseudogonatopus nr. hospes R. Perkins : Chandra 1978b: 5. Pseudogonatopus fulgori (Nakagawa) : Olmi 1984: 1194. Pseudogonatopus pusanus Olmi 1984: 1255 ; type locality: Pusa (Bihar, India); He & Xu 2002: 355; syn. nov. Pseudogonatopus nigricans (R. Perkins) : Olmi 1984: 1287; Olmi 1990b: 789; Olmi 1991: 319. Gonatopus nigricans (R. Perkins) : Guglielmino & Olmi 1997: 165, 224–226, 250–252; Olmi 1999b: 28; Olmi 2000a: 94; He &

Xu 2002: 364; Guglielmino & Olmi 2006: 49. Gonatopus sogatae Rohwer : Guglielmino & Olmi 1997: 225, 226; He & Xu 2002: 355. Gonatopus insulanus He & Xu in He et al. 1998: 172; type locality: Qionghai (Hainan, China); syn. nov.

Description of female ( Plate 114A ): apterous; length 2.4–4.4 mm. Head brown-reddish, or brown, or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 or 1–3 yellow; mesosoma black or brown; occasionally posterior margin of pronotum and posterior apex of propodeum testaceous; metasoma black or brown; legs yellow; occasionally clubs of femora and part of coxae brown. In G. pusanus the colour is different: head partly or almost totally brown; antenna brown, except segments 1–2 testaceous; mesosoma testaceous-reddish, except sides of scutum darkened; occasionally part of pronotum darkened; metasoma brown; petiole black; legs testaceous; occasionally club of profemur darkened. In G. sogatae the colour is the following: head brown, except mandible and clypeus yellow; antenna brown, except segment 1 yellow and 2–3 brown-testaceous; mesosoma brown, except scutum reddish and distal apex of propodeum yellow; petiole black; metasoma brown-dark; legs brown-yellow. Intermediate colours are known. Antenna clavate; antennal segments in following proportions: 19:19:52:25:22:22:18:17:15:26. Head ( Plate 114B ) excavated, shiny, granulated or unsculptured; occiput and anterior region of face always granulated; POL = 5; OL = 5; OOL = 14. Palpal formula 2/2, 3/2, 4/2. Pronotum granulated, crossed by strong transverse impression. Scutum granulated, without lateral pointed apophyses. Mesopleura transversely striate. Metanotum flat, very long, about as long as scutum, transversely striate, not hollow behind scutellum ( Plate 114C ), with sides rounded ( Plate 114A ). Metathorax + propodeum shiny, indistinctly sculptured, with or without track of median longitudinal furrow, with posterior surface transversely striate. Metapleuron transversely striate. Meso-metapleural suture obsolete. Protarsal segment 4 shorter than segment 1 (40:47; or 20:27). Enlarged claw ( Plate 114D ) with one large subapical tooth and one row of three–nine lamellae. Segment 5 of protarsus ( Plate 114D ) with two rows of 9–25 lamellae; distal apex with 4–10 lamellae; occasionally lamellae extended continuously to distal apex. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9–2.4 mm. Head brown or black (occasionally brown-reddish), except mandible testaceous; occasionally part of face testaceous; antenna brown (in type of G. insulanus , testaceous-dark); mesosoma and metasoma black or brown (in type of G. insulanus , mesosoma reddish-dark, with propodeum and pleura darker); legs yellow. Antenna filiform; antennal segments in following proportions: 5:5:9:8:8:9:7:8:7:10 (in other specimens 6:5.5:10:9:8:9:8:8:7:10); antennal segment 3 more than three times as long as broad (4.0; 4.3; 5.0). Head shiny, hairy, granulated; frontal line absent; temple distinct; occipital carina absent; POL = 8; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OL (4:3) (in other specimens POL = 7; OL = 3; OOL = 3.5; greatest breadth of posterior ocelli longer than OL (3.5:3)). Palpal formula 2/2, 3/2, 4/2. Scutum shiny, granulated. Notauli complete, posteriorly joint or almost joint in small specimens, separated in large specimens; in very large specimens, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2.5:4; or 2:4). Scutellum and metanotum shiny, finely punctate, unsculptured among punctures. Propodeum reticulate rugose; dorsal surface with median longitudinal furrow and occasionally with two lateral smooth areas. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 111B ) much shorter than paramere. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nigricans : AUSTRALIA: Queensland, Bundaberg, 19.XI.1904, reared ( BPBM); ♀ lectotype (designated by Olmi, 1982) of P. melanacrias : FIJI ISLANDS: Viti Levu Island, Suva, III.1905 , reared from delphacid near Stenocranus , on Zoysia pungens (BPBM) ; ♀ lectotype (designated by Olmi, 1982) of P. hospes : CHINA: no localities, F. Muir leg., reared in Honolulu from Perkinsiella saccharicida from Chinese stock, IV–V.1907 ( BPBM); ♀ holotype of G. sauteri : CHINA: Taiwan, Tainan, IV.1912, H. Sauter leg. ( DEI); ♀ holotype of P. pusanus : INDIA: Bihar, Pusa ( IARI); ♀ holotype of P. sogatae : INDIA: Bihar, Pusa, 6.VIII.1915, reared from Sogata sp. , C.S. Misra leg. ( USNM); ♂ holotype of G. insulanus : CHINA: Hainan, Qionghai, 4.V.1964, Junhua He leg. ( ZJUC). Type material of G. fulgori probably lost. Paratypes of P. nigricans : AUSTRALIA: same locality label as lectotype, 1♀ lebelled for mistake Fiji Islands ( BPBM); same locality label as holotype, parasitized host collected 1.X.1904, dryinid adult emerged 21.X.1904, R. Perkins leg., 1♀ “ paratype ” identified by the same R. Perkins ( BMNH); AUSTRALIA: Queensland, Bundaberg, Sandhills, X.1904, bred, 1♀ ( ANIC). Paratypes of P. melanacrias : same locality label as lectotype, 2♀♀ ( BPBM). Paratypes of P. hospes : same locality label as lectotype, 3♀♀ ( BPBM). Paratype of P. pusanus : same locality label as holotype, 1♀ ( IARI). Paratype of P. sogatae : same locality label as holotype, misidentified ♂ allotype not belonging to the same species of holotype. Other material. CHINA: Fujian, Shaxian, reared from rice planthoppers, 1979–1983, Changming Liu leg., 5♀♀ and 1♂ ( ZJUC); Fujian, IV.1987, 31.VII.1985, Changming Liu leg., 6♀♀ and 10♂ ( ZJUC); Guangdong, Yangjiang, 1979, 1981, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangdong, Xingfeng County, Mt. Yunjinshan, 9.VI.2002, 18.VII.2003, 19.VII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); same locality label, 20.VII.2003, Ping Li leg., 2♀♀ ( ZJUC); same locality label, Zaifu Xu leg., many ♀♀ ( ZJUC, AMNH); Guangdong, Shixing County, Chebaling National Nature Reserve, 21.VIII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, M. Olmi reared from adult of unidentified Delphacid, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); Guangdong, Mt. Nankunshan, 2.VII.2005, Zaifu Xu leg., 1♀ ( ZJUC); Guangdong, Yangjiang, Hailing Town, 1980, Renhuan Hou leg., 2♀♀ and 2♂ ( ZJUC); Guangdong, Huaiji County, 21–22.X.2007, Zaifu Xu leg., 1♀ ( SCAU); Guangdong, Conghua, 28.VI.1978, Junhua He leg., 1♀ ( SCAU); Guangdong, Zhaoqing, V.1998, Yufang Liu leg., 2♀♀ ( SCAU); Guizhou, Sinan, IX.1986, VIII.1991, 5♀♀ ( ZJUC); Guizhou, Sandu, VI–VII.1981, Dongxiang Xie leg., 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Hunan, Yueyang, 15.VIII.1975, Binsen Wang leg., 1♀ ( ZJUC); Guangxi, Luchuan, 30.VII.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Bama, 1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Luocheng, 18.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Yaxian, VI.1966, Junhua He leg., 1♀ ( ZJUC); Hong Kong, Tai Lung Farm, IX–XI.1965, swept above rice, I.W.B. Thornton leg., 1♀ ( BMNH); Macau ( BPBM); same locality label, R. Perkins collection, 2♀♀ ( USNM); Taiwan, Kaohsiung, Kosen ( AMNH, ELKU); Taiwan, Changhus, Erlin ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, Kungkuan ( TARI); Taiwan, Taihoku Pref. (=Taipei), 27.IX.1930, J. Sonan leg., 1♀ ( TARI); Taiwan, Tainan, ShinShi, 11.IX.1979, paddy field, Hirashima leg., 1♀ ( ELKU); Taiwan, Gakko ( ELKU); Itahashi (= Taiwan), 16.IX.1930, J. Sonan leg., 5♀♀ ( TARI); Takeyama (=Jhushan), 6.X.1937, J. Sonan leg., 3♀♀ ( TARI); Inrin (=Yuanlin), 8.X.1937, J. Sonan leg., 39♀♀ ( TARI). INDIA: Chhattisgarh, Raipur ( GCC, AMNH); Delhi, New Delhi ( IARI); Orissa, Cuttack, 1981, 1♀ ( BMNH); Orissa, Puri, 28.XI.1987, S.V. Fowler leg., 1♀ ( BMNH); Tamil Nadu, Tanjavur (=Tanjore) District, 28.IX.1987, ex Nilaparvata lugens on rice, C. Peter leg., 3♀♀ ( BMNH); West Bengal, Hooghly District, Khanakul, 12.XII.1986, in paddy seedbed, 1♀ ( BMNH). INDONESIA: Java, Pekalongan ( BPBM); Java, Dieng Plateau ( BPBM); Borneo, Kalimantan, Paloh ( BMNH). MALAYSIA: Malaya, Selangor, 13.VIII.1928, H.T. Pagden leg., 1♀ ( BMNH); Malaya, Selangor, Kuala Lumpur, 21.XII.1930, H. T. Pagden leg., 1♀ ( BMNH); same locality label, many ♀♀ and ♂ ( NMNH, KYC); Malaya, Seberang Perai ( MARDI); Malaya, Serdang ( DBUS); Sabah, Tuaran ( KYC); Sarawak, 1st. Division, Payah Paloh, 28.I.1965, G.H.L. Rotschild leg., 1♀ ( BMNH). THAILAND: Songkhla, Hadd Tai ( KYC); Non Bua Lampoo, Na Kam Hai ( KYC); Tung Saew ( KYC); Tung Fabad ( KYC); Chainat ( KYC, AMNH); Banmakok ( KYC); Mae Tang ( KYC); Lampamg ( ELKU); Mae-Theng ( ELKU); Hot ( ELKU); Chieng Dao ( ELKU); Thailand, 15.IX.1999, reared from Nilaparvata lugens (Stål) , Sawai leg., 4♂ ( BMNH). Australian region: G. nigricans is known from Australia (Queensland), Fiji Islands, Solomon Islands and USA (Hawaii) ( Olmi, 1984). Introduced in the Hawaii 1906 and 1907 from China, Guangdong, for biological control of Perkinsiella saccharicida Kirkaldy in sugarcane fields ( Olmi, 2000a).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in Australia: Liburnia sp. , Stenocranus sp. ; in China: Laodelphax striatellus (Fallén) ( Yang et al., 1982; He & Xu, 2002), Perkinsiella saccharicida Kirkaldy ; in Malaysia: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in Thailand: Nilaparvata lugens (Stål) ( Chiu, 1979) ; in the USA (Hawaii): Perkinsiella saccharicida Kirkaldy (introduced for biological control).

Biology. Information by Pagden (1934); the parthenogenesis is thelytokous.

Natural enemies. In the USA (Hawaii) ( Swezey, 1919; Timberlake, 1919; Muir, 1921; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Encyrtidae : Helegonatopus pseudophanes R. Perkins, 1906 ; Cheiloneurus flaccus (Walker 1847) ( Guerrieri & Viggiani, 2005; Guerrieri, 2006); Aphelinidae : Centrodora xiphidii (R. Perkins, 1906) . In Malaya ( Pagden, 1934; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Ceraphronidae : Allomicrops abnormis (R. Perkins, 1910) ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp.

Distribution. China ( Taiwan, Hainan, Fujian, Guangdong, Guizhou, Hubei, Hunan, Guangxi, Hong Kong, Macau), Australia (Queensland), Fiji Islands ( Viti Levu), India (Bihar, Chhattisgarh, Delhi, Orissa, Tamil Nadu), Indonesia (Java, Pekalongan, Borneo), Malaysia (Malaya, Sabah, Sarawak), Thailand (Songkhla, Non Bua Lampoo, Tung Saew, Tung Fabad, Chainat, Banmakok, Mae Tang, Lampamg, Mae-Theng, Chieng Dao).

Remarks. Gonatopus insulanus He & Xu is here synonymized with G. nigricans , because of the study of its type and comparison with male specimens of G. nigricans . There are no differences.

Gonatopus nigricans (= Pseudogonatopus hospes R. Perkins ) was established in the Hawaii from cocoons sent from China ( Perkins, 1912). This species was considered extinct in the Hawaii ( Olmi, 1999a). However, in 2010 one of the authors (Olmi) saw a male specimen collected in Oahu Island, Mt. Olympus, in April 1981 by a pan trap. Probably so this species is always present at least in Oahu Island, although rare. Gonatopus pusanus and G. sogatae are here synonymized with G. nigricans because the only differences concern the colour of mesosoma (black in nigricans , testaceous to reddish, with some darkened areas, in pusanus ; brown, with some areas reddish or yellow, in sogatae ). In the collections, there are also many specimens with intermediate colours, so that the synonymy is evident. Gonatopus fulgori is here synonymized with G. nigricans , because the original description is like that of G. nigricans (the type material of G. fulgori is lost). In the past these two species were separated, because attributed to two different zoogeographic regions ( fulgori in Palaearctic, nigricans in Oriental and Australian regions).

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

3. Gonatopus lankae (Ponomarenko, 1981) , comb. nov.

( Plate 113A, B )

Pseudogonatopus lankae Ponomarenko 1981b: 881 ; Olmi 1984: 1249.

Description of Female. Apterous; length 3.2–3.3 mm. Brown-reddish or testaceous-dark; antenna usually darkened, except segment 10 yellow; legs testaceous, except trochanters and procoxa partly whitish, hind side of metafemur whitish, distal apex of propodeum whitish. Antenna clavate; antennal segments in following proportions: 14:8:14:9:9:6:6:6:6:11. Head excavated, dull, alutaceous or weakly granulated; frontal line complete; occipital carina absent; temple prominent; POL = 2; OL = 3; OOL = 10. Palpal formula 4/2 or 5/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, smooth, unsculptured, with some transverse keels on lateral regions. Scutellum shiny, smooth, unsculptured. Metanotum flat, shiny, unsculptured, weakly hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded. Metathorax + propodeum ( Plate 113A ) shiny, smooth, unsculptured or granulated; posterior surface of propodeum transversely striate. Disc of metathorax + propodeum with strong median furrow. Mesopleura and metapleura dull, weakly granulated, not transversely striate. Meso-metapleural suture distinct and complete. Protarsal segments in following proportions: 15:4:6:20:31. Enlarged claw ( Plate 113B ) with one large subapical tooth and one row of six–eight lamellae. Segment 5 of protarsus ( Plate 113B ) with two rows of 19–21 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Ratnapura District , Gilimale Induruwa Jungle, 13.III.1979, K. Krombein leg. ( USNM) . Other material. SRI LANKA: Ratnapura District ., Sinharaja Jungle, Waturawa Forest ( AMNH) ; Matale District , Sigiriya, 07°57'N 80°46'E, 18.III.1999, C.J. Burwell leg. ( QMC) .

Hosts. Unknown.

Distribution. Sri Lanka (Ratnapura, Matale).

Remarks. The female specimen from Sri Lanka, 07°57'N 80°46'E, has 5-segmented maxillary palps; the palpal formula 5/2 is typical of the group 10 (former genus Donisthorpina Richards ); for this reason, G. lankae is also included into group 10.

4. Gonatopus malesiae ( Olmi, 1984) , comb. nov.

( Plate 113C, D )

Pseudogonatopus malesiae Olmi 1984: 1248 .

Description of Female. Apterous; length 2.8–3.6 mm. Usually testaceous-ferruginous, with petiole black; antenna usually testaceous, with segments 9–10 whitish (occasionally antennal segment 9 partly whitish). In specimen from Myanmar, 17°38.9'N 94°38.1'E, antenna testaceous-brown, except segments 1 and 10 whitish. In specimen from China, Mt. Jinggangshan, antenna brown, except rhinaria areas testaceous, and mesosoma brown-testaceous. Antenna clavate; antennal segments of holotype in following proportions: 16:13:38:23:18:12:12:13:12:20. Head very excavated, shiny, smooth, unsculptured, hairless; frontal line complete; occipital carina absent; POL = 0.3; OL = 2; OOL = 8.5. Palpal formula 3/2 or 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat, without lateral pointed apophyses. Metanotum shiny, unsculptured, not hollow behind scutellum, with sides protruding; lateral protrusions pointed ( Plate 113C ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum without median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture very slightly distinct. In specimen from Myanmar, 17°38.9'N 94°38.1'E, meso-metapleural suture partly and slightly visible (not visible on sides of metanotum). Segment 1 of protarsus usually shorter than segment 4 (13:16). Enlarged claw ( Plate 113D ) with one large subapical tooth and one row of seven–ten lamellae. Segment 5 of protarsus ( Plate 113D ) with two rows of 12–17 lamellae; distal apex with at least 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown (see Gonatopus sarawakensis (Olmi)) .

Material examined. Type: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu National Park , 5.VIII.1978, P.M. Hammond & J.E. Marshall leg. ( BMNH) . Other material. CHINA: Jiangxi, Mt. Jinggangshan , 13.VIII.2007, Junhua He leg., 1♀ ( SCAU) . MALAYSIA: Malaya, Fraser's Hill , 4200' ( AMNH) . MYANMAR: Rakhine State, NE of Gwa, Elephant Camp in Elephant Sanctuary , 17°38.9'N 94°38.1'E, 3.XII.2000, leaf litter in secondary forest, K.J. Ribardo leg., 2♀♀ ( CAS) .

Hosts. Unknown.

Distribution. China (Jiangxi), Malaysia ( Sarawak, Malaya), Myanmar (Rakhine).

5. Gonatopus muiri ( Olmi, 1984)

( Plate 113E, F )

Pseudogonatopus muiri Olmi 1984: 1251 ; Xu et al. 2011g (Suppl.): 20.

Description of Female. Apterous; length 2.5–3.0 mm. Holotype testaceous–reddish, except metasoma brown and antenna brown, with segments 1–2 testaceous (specimen from China, Bawangling National Nature Reserve, testaceous, except antenna brown, with segments 1–2 testaceous and segment 10 whitish, petiole black and metasoma brown). Antenna clavate; antennal segments of a specimen from China, Bawangling National Nature Reserve, in following proportions: 8:4.5:10:6:5:5:4.5:4.5:4:8. Head excavated; head of holotype dull and slightly granulated; head of specimen from China, Bawangling National Nature Reserve, shiny and unsculptured; frontal line of specimen from China, Bawangling National Nature Reserve, complete; occipital carina absent; head of specimen from China, Bawangling National Nature Reserve, with POL = 0.5; OL = 1.5; OOL = 6. Palpal formula 3/ 2 in holotype, 4/ 2 in specimen from China, Bawangling National Nature Reserve. Pronotum crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat. Metanotum shiny, unsculptured, not hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded ( Plate 113E ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum with slight median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture weakly distinct, in holotype only visible near lateral protrusions of metanotum. Meso-metapleural suture of specimen from China, Bawangling National Nature Reserve, visible only between mesopleuron and metapleuron, not visible near lateral protrusions of metanotum. Protarsal segments of specimen from China, Bawangling National Nature Reserve, in following proportions: 10:2.5:4:10:17; in holotype segment 1 of protarsus as long as segment 4. Enlarged claw ( Plate 113F ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 113F ) with two rows of 12–17 lamellae; distal apex with 11–12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Java, Pekalongan, F. Muir leg. ( AMNH) . Other material. CHINA: Hainan, Bawangling National Nature Reserve, Reserve House , about 1000 m, parasitized host collected 19.VIII.2000, dryinid cocooning 22.VIII.2000, M. Olmi reared from an unidentified species of Delphacidae , 1♀ ( MOLC) .

Hosts. In China (Hainan), unidentified species of Delphacidae .

Distribution. China (Hainan), Indonesia (Java).

6. Gonatopus nepalensis ( Olmi, 1986) , comb. nov.

( Plate 113G, H )

Pseudogonatopus nepalensis Olmi 1986: 80 .

Description of Female. Apterous; length 4.8–5.0 mm. Head black or brown, except mandible, clypeus and anterior region of face testaceous; antenna brown, except segments 1–2 testaceous and 10 whitish; occasionally antennal segment 9 whitish; mesosoma and metasoma black; legs black or brown, except chela and meso- and metatarsus testaceous. Antenna clavate; antennal segments in following proportions: 12:6:18:10:7:7:7:6.5:6.5:10.5. Head excavated, dull, granulated; frontal line complete; occipital carina absent. Palpal formula 4/2. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum shiny, sculptured by longitudinal striae. Metanotum transversely striate, with sides protruding; protrusions rounded ( Plate 113G ). Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface and metapleura. Mesopleuron shiny, smooth, unsculptured, not transversely striate. Meso-metapleural suture partly distinct, only present between mesopleuron and metapleuron, not present dorsally on sides of metanotum. Protarsal segments in following proportions: 18:4:8:22:35. Enlarged claw ( Plate 113G ) with one large subapical tooth and one row of seven lamellae. Segment 5 of protarsus ( Plate 113G ) with two rows of about 21 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: NEPAL: Kosi, S of Mangsingma Forest ( MHNG). Paratypes: same locality label as holotype, 1♀ ( AMNH); E Nepal, Kosi, Induwa Kola Valley, 1♀ ( MHNG). Other material. NEPAL: Khala Valley, 2050 m ( PMA). Palaearctic: according to Ponomarenko (1992), G. nepalensis is also present in the following localities: RUSSIA: Transbaikalia, 5.VI.1963, 10.VII.1963, A. Emelianov leg., 2♀♀ ( Ponomarenko , 1992); Fra East, Primorskiy Krai, Gornotayezhnoe, 19.VI.1983, Sciarkov leg., 1♀ ( Ponomarenko , 1992). This quotation is doubtful.

Hosts. Unknown.

Distribution. Nepal (Kosi), Russia (Transbaicalia, Primorskiy Krai).

7. Gonatopus nigricans (R. Perkins, 1905)

( Plates 111B , 114A – D )

Paragonatopus nigricans R. Perkins 1905: 41 ; Kieffer 1914b: 87; Olmi 1979: 218.

Pseudogonatopus melanacrias R. Perkins 1906a: 487 (synonymized by R. Perkins 1912); type locality: Suva ( Viti Levu Island,

Fiji Islands). Gonatopus fulgori Nakagawa 1906: 163 ; type locality: Japan; Guglielmino & Olmi 1997: 181, 184; syn. nov. Pseudogonatopus hospes R. Perkins 1912: 13 (synonymized by He & Xu 2002); type locality: China; Kieffer 1914b: 81;

Swezey 1928: 670; F. Williams 1931: 247; Chiu 1979: 339; Móczár 1979: 77; Olmi 1984: 1290; Olmi 2000a: 101, 111. Gonatopus sauteri Strand 1913: 209 (synonymized by Olmi 1984); type locality: Tainan ( Taiwan, China). Dicondylus sauteri (Strand) : Kieffer 1914b: 77. Pseudogonatopus sogatea Rohwer 1920: 160 (printing-error; holotype labelled sogatae ); type locality: Pusa (Bihar, India);

Kurian 1954b: 417; Olmi 1984: 1254; syn. nov. Haplogonatopus fulgori (Nakagawa) : Esaki & Hashimoto 1930: 21. Pseudozonatopus (sic) hospes R. Perkins : Pagden 1934: 2. Paragonatopus fulgori (Nakagawa) : Mochizuki 1943: 99; Chen 1988: 44. nec Pseudogonatopus nr. hospes R. Perkins : Chandra 1978b: 5. Pseudogonatopus fulgori (Nakagawa) : Olmi 1984: 1194. Pseudogonatopus pusanus Olmi 1984: 1255 ; type locality: Pusa (Bihar, India); He & Xu 2002: 355; syn. nov. Pseudogonatopus nigricans (R. Perkins) : Olmi 1984: 1287; Olmi 1990b: 789; Olmi 1991: 319. Gonatopus nigricans (R. Perkins) : Guglielmino & Olmi 1997: 165, 224–226, 250–252; Olmi 1999b: 28; Olmi 2000a: 94; He &

Xu 2002: 364; Guglielmino & Olmi 2006: 49. Gonatopus sogatae Rohwer : Guglielmino & Olmi 1997: 225, 226; He & Xu 2002: 355. Gonatopus insulanus He & Xu in He et al. 1998: 172; type locality: Qionghai (Hainan, China); syn. nov.

Description of female ( Plate 114A ): apterous; length 2.4–4.4 mm. Head brown-reddish, or brown, or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 or 1–3 yellow; mesosoma black or brown; occasionally posterior margin of pronotum and posterior apex of propodeum testaceous; metasoma black or brown; legs yellow; occasionally clubs of femora and part of coxae brown. In G. pusanus the colour is different: head partly or almost totally brown; antenna brown, except segments 1–2 testaceous; mesosoma testaceous-reddish, except sides of scutum darkened; occasionally part of pronotum darkened; metasoma brown; petiole black; legs testaceous; occasionally club of profemur darkened. In G. sogatae the colour is the following: head brown, except mandible and clypeus yellow; antenna brown, except segment 1 yellow and 2–3 brown-testaceous; mesosoma brown, except scutum reddish and distal apex of propodeum yellow; petiole black; metasoma brown-dark; legs brown-yellow. Intermediate colours are known. Antenna clavate; antennal segments in following proportions: 19:19:52:25:22:22:18:17:15:26. Head ( Plate 114B ) excavated, shiny, granulated or unsculptured; occiput and anterior region of face always granulated; POL = 5; OL = 5; OOL = 14. Palpal formula 2/2, 3/2, 4/2. Pronotum granulated, crossed by strong transverse impression. Scutum granulated, without lateral pointed apophyses. Mesopleura transversely striate. Metanotum flat, very long, about as long as scutum, transversely striate, not hollow behind scutellum ( Plate 114C ), with sides rounded ( Plate 114A ). Metathorax + propodeum shiny, indistinctly sculptured, with or without track of median longitudinal furrow, with posterior surface transversely striate. Metapleuron transversely striate. Meso-metapleural suture obsolete. Protarsal segment 4 shorter than segment 1 (40:47; or 20:27). Enlarged claw ( Plate 114D ) with one large subapical tooth and one row of three–nine lamellae. Segment 5 of protarsus ( Plate 114D ) with two rows of 9–25 lamellae; distal apex with 4–10 lamellae; occasionally lamellae extended continuously to distal apex. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9–2.4 mm. Head brown or black (occasionally brown-reddish), except mandible testaceous; occasionally part of face testaceous; antenna brown (in type of G. insulanus , testaceous-dark); mesosoma and metasoma black or brown (in type of G. insulanus , mesosoma reddish-dark, with propodeum and pleura darker); legs yellow. Antenna filiform; antennal segments in following proportions: 5:5:9:8:8:9:7:8:7:10 (in other specimens 6:5.5:10:9:8:9:8:8:7:10); antennal segment 3 more than three times as long as broad (4.0; 4.3; 5.0). Head shiny, hairy, granulated; frontal line absent; temple distinct; occipital carina absent; POL = 8; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OL (4:3) (in other specimens POL = 7; OL = 3; OOL = 3.5; greatest breadth of posterior ocelli longer than OL (3.5:3)). Palpal formula 2/2, 3/2, 4/2. Scutum shiny, granulated. Notauli complete, posteriorly joint or almost joint in small specimens, separated in large specimens; in very large specimens, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2.5:4; or 2:4). Scutellum and metanotum shiny, finely punctate, unsculptured among punctures. Propodeum reticulate rugose; dorsal surface with median longitudinal furrow and occasionally with two lateral smooth areas. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 111B ) much shorter than paramere. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nigricans : AUSTRALIA: Queensland, Bundaberg, 19.XI.1904, reared ( BPBM); ♀ lectotype (designated by Olmi, 1982) of P. melanacrias : FIJI ISLANDS: Viti Levu Island, Suva, III.1905 , reared from delphacid near Stenocranus , on Zoysia pungens (BPBM) ; ♀ lectotype (designated by Olmi, 1982) of P. hospes : CHINA: no localities, F. Muir leg., reared in Honolulu from Perkinsiella saccharicida from Chinese stock, IV–V.1907 ( BPBM); ♀ holotype of G. sauteri : CHINA: Taiwan, Tainan, IV.1912, H. Sauter leg. ( DEI); ♀ holotype of P. pusanus : INDIA: Bihar, Pusa ( IARI); ♀ holotype of P. sogatae : INDIA: Bihar, Pusa, 6.VIII.1915, reared from Sogata sp. , C.S. Misra leg. ( USNM); ♂ holotype of G. insulanus : CHINA: Hainan, Qionghai, 4.V.1964, Junhua He leg. ( ZJUC). Type material of G. fulgori probably lost. Paratypes of P. nigricans : AUSTRALIA: same locality label as lectotype, 1♀ lebelled for mistake Fiji Islands ( BPBM); same locality label as holotype, parasitized host collected 1.X.1904, dryinid adult emerged 21.X.1904, R. Perkins leg., 1♀ “ paratype ” identified by the same R. Perkins ( BMNH); AUSTRALIA: Queensland, Bundaberg, Sandhills, X.1904, bred, 1♀ ( ANIC). Paratypes of P. melanacrias : same locality label as lectotype, 2♀♀ ( BPBM). Paratypes of P. hospes : same locality label as lectotype, 3♀♀ ( BPBM). Paratype of P. pusanus : same locality label as holotype, 1♀ ( IARI). Paratype of P. sogatae : same locality label as holotype, misidentified ♂ allotype not belonging to the same species of holotype. Other material. CHINA: Fujian, Shaxian, reared from rice planthoppers, 1979–1983, Changming Liu leg., 5♀♀ and 1♂ ( ZJUC); Fujian, IV.1987, 31.VII.1985, Changming Liu leg., 6♀♀ and 10♂ ( ZJUC); Guangdong, Yangjiang, 1979, 1981, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangdong, Xingfeng County, Mt. Yunjinshan, 9.VI.2002, 18.VII.2003, 19.VII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); same locality label, 20.VII.2003, Ping Li leg., 2♀♀ ( ZJUC); same locality label, Zaifu Xu leg., many ♀♀ ( ZJUC, AMNH); Guangdong, Shixing County, Chebaling National Nature Reserve, 21.VIII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, M. Olmi reared from adult of unidentified Delphacid, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); Guangdong, Mt. Nankunshan, 2.VII.2005, Zaifu Xu leg., 1♀ ( ZJUC); Guangdong, Yangjiang, Hailing Town, 1980, Renhuan Hou leg., 2♀♀ and 2♂ ( ZJUC); Guangdong, Huaiji County, 21–22.X.2007, Zaifu Xu leg., 1♀ ( SCAU); Guangdong, Conghua, 28.VI.1978, Junhua He leg., 1♀ ( SCAU); Guangdong, Zhaoqing, V.1998, Yufang Liu leg., 2♀♀ ( SCAU); Guizhou, Sinan, IX.1986, VIII.1991, 5♀♀ ( ZJUC); Guizhou, Sandu, VI–VII.1981, Dongxiang Xie leg., 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Hunan, Yueyang, 15.VIII.1975, Binsen Wang leg., 1♀ ( ZJUC); Guangxi, Luchuan, 30.VII.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Bama, 1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Luocheng, 18.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Yaxian, VI.1966, Junhua He leg., 1♀ ( ZJUC); Hong Kong, Tai Lung Farm, IX–XI.1965, swept above rice, I.W.B. Thornton leg., 1♀ ( BMNH); Macau ( BPBM); same locality label, R. Perkins collection, 2♀♀ ( USNM); Taiwan, Kaohsiung, Kosen ( AMNH, ELKU); Taiwan, Changhus, Erlin ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, Kungkuan ( TARI); Taiwan, Taihoku Pref. (=Taipei), 27.IX.1930, J. Sonan leg., 1♀ ( TARI); Taiwan, Tainan, ShinShi, 11.IX.1979, paddy field, Hirashima leg., 1♀ ( ELKU); Taiwan, Gakko ( ELKU); Itahashi (= Taiwan), 16.IX.1930, J. Sonan leg., 5♀♀ ( TARI); Takeyama (=Jhushan), 6.X.1937, J. Sonan leg., 3♀♀ ( TARI); Inrin (=Yuanlin), 8.X.1937, J. Sonan leg., 39♀♀ ( TARI). INDIA: Chhattisgarh, Raipur ( GCC, AMNH); Delhi, New Delhi ( IARI); Orissa, Cuttack, 1981, 1♀ ( BMNH); Orissa, Puri, 28.XI.1987, S.V. Fowler leg., 1♀ ( BMNH); Tamil Nadu, Tanjavur (=Tanjore) District, 28.IX.1987, ex Nilaparvata lugens on rice, C. Peter leg., 3♀♀ ( BMNH); West Bengal, Hooghly District, Khanakul, 12.XII.1986, in paddy seedbed, 1♀ ( BMNH). INDONESIA: Java, Pekalongan ( BPBM); Java, Dieng Plateau ( BPBM); Borneo, Kalimantan, Paloh ( BMNH). MALAYSIA: Malaya, Selangor, 13.VIII.1928, H.T. Pagden leg., 1♀ ( BMNH); Malaya, Selangor, Kuala Lumpur, 21.XII.1930, H. T. Pagden leg., 1♀ ( BMNH); same locality label, many ♀♀ and ♂ ( NMNH, KYC); Malaya, Seberang Perai ( MARDI); Malaya, Serdang ( DBUS); Sabah, Tuaran ( KYC); Sarawak, 1st. Division, Payah Paloh, 28.I.1965, G.H.L. Rotschild leg., 1♀ ( BMNH). THAILAND: Songkhla, Hadd Tai ( KYC); Non Bua Lampoo, Na Kam Hai ( KYC); Tung Saew ( KYC); Tung Fabad ( KYC); Chainat ( KYC, AMNH); Banmakok ( KYC); Mae Tang ( KYC); Lampamg ( ELKU); Mae-Theng ( ELKU); Hot ( ELKU); Chieng Dao ( ELKU); Thailand, 15.IX.1999, reared from Nilaparvata lugens (Stål) , Sawai leg., 4♂ ( BMNH). Australian region: G. nigricans is known from Australia (Queensland), Fiji Islands, Solomon Islands and USA (Hawaii) ( Olmi, 1984). Introduced in the Hawaii 1906 and 1907 from China, Guangdong, for biological control of Perkinsiella saccharicida Kirkaldy in sugarcane fields ( Olmi, 2000a).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in Australia: Liburnia sp. , Stenocranus sp. ; in China: Laodelphax striatellus (Fallén) ( Yang et al., 1982; He & Xu, 2002), Perkinsiella saccharicida Kirkaldy ; in Malaysia: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in Thailand: Nilaparvata lugens (Stål) ( Chiu, 1979) ; in the USA (Hawaii): Perkinsiella saccharicida Kirkaldy (introduced for biological control).

Biology. Information by Pagden (1934); the parthenogenesis is thelytokous.

Natural enemies. In the USA (Hawaii) ( Swezey, 1919; Timberlake, 1919; Muir, 1921; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Encyrtidae : Helegonatopus pseudophanes R. Perkins, 1906 ; Cheiloneurus flaccus (Walker 1847) ( Guerrieri & Viggiani, 2005; Guerrieri, 2006); Aphelinidae : Centrodora xiphidii (R. Perkins, 1906) . In Malaya ( Pagden, 1934; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Ceraphronidae : Allomicrops abnormis (R. Perkins, 1910) ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp.

Distribution. China ( Taiwan, Hainan, Fujian, Guangdong, Guizhou, Hubei, Hunan, Guangxi, Hong Kong, Macau), Australia (Queensland), Fiji Islands ( Viti Levu), India (Bihar, Chhattisgarh, Delhi, Orissa, Tamil Nadu), Indonesia (Java, Pekalongan, Borneo), Malaysia (Malaya, Sabah, Sarawak), Thailand (Songkhla, Non Bua Lampoo, Tung Saew, Tung Fabad, Chainat, Banmakok, Mae Tang, Lampamg, Mae-Theng, Chieng Dao).

Remarks. Gonatopus insulanus He & Xu is here synonymized with G. nigricans , because of the study of its type and comparison with male specimens of G. nigricans . There are no differences.

Gonatopus nigricans (= Pseudogonatopus hospes R. Perkins ) was established in the Hawaii from cocoons sent from China ( Perkins, 1912). This species was considered extinct in the Hawaii ( Olmi, 1999a). However, in 2010 one of the authors (Olmi) saw a male specimen collected in Oahu Island, Mt. Olympus, in April 1981 by a pan trap. Probably so this species is always present at least in Oahu Island, although rare. Gonatopus pusanus and G. sogatae are here synonymized with G. nigricans because the only differences concern the colour of mesosoma (black in nigricans , testaceous to reddish, with some darkened areas, in pusanus ; brown, with some areas reddish or yellow, in sogatae ). In the collections, there are also many specimens with intermediate colours, so that the synonymy is evident. Gonatopus fulgori is here synonymized with G. nigricans , because the original description is like that of G. nigricans (the type material of G. fulgori is lost). In the past these two species were separated, because attributed to two different zoogeographic regions ( fulgori in Palaearctic, nigricans in Oriental and Australian regions).

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

4. Gonatopus malesiae ( Olmi, 1984) , comb. nov.

( Plate 113C, D )

Pseudogonatopus malesiae Olmi 1984: 1248 .

Description of Female. Apterous; length 2.8–3.6 mm. Usually testaceous-ferruginous, with petiole black; antenna usually testaceous, with segments 9–10 whitish (occasionally antennal segment 9 partly whitish). In specimen from Myanmar, 17°38.9'N 94°38.1'E, antenna testaceous-brown, except segments 1 and 10 whitish. In specimen from China, Mt. Jinggangshan, antenna brown, except rhinaria areas testaceous, and mesosoma brown-testaceous. Antenna clavate; antennal segments of holotype in following proportions: 16:13:38:23:18:12:12:13:12:20. Head very excavated, shiny, smooth, unsculptured, hairless; frontal line complete; occipital carina absent; POL = 0.3; OL = 2; OOL = 8.5. Palpal formula 3/2 or 4/2. Pronotum crossed by strong transverse impression, shiny, smooth, unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat, without lateral pointed apophyses. Metanotum shiny, unsculptured, not hollow behind scutellum, with sides protruding; lateral protrusions pointed ( Plate 113C ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum without median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture very slightly distinct. In specimen from Myanmar, 17°38.9'N 94°38.1'E, meso-metapleural suture partly and slightly visible (not visible on sides of metanotum). Segment 1 of protarsus usually shorter than segment 4 (13:16). Enlarged claw ( Plate 113D ) with one large subapical tooth and one row of seven–ten lamellae. Segment 5 of protarsus ( Plate 113D ) with two rows of 12–17 lamellae; distal apex with at least 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown (see Gonatopus sarawakensis (Olmi)) .

Material examined. Type: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu National Park , 5.VIII.1978, P.M. Hammond & J.E. Marshall leg. ( BMNH) . Other material. CHINA: Jiangxi, Mt. Jinggangshan , 13.VIII.2007, Junhua He leg., 1♀ ( SCAU) . MALAYSIA: Malaya, Fraser's Hill , 4200' ( AMNH) . MYANMAR: Rakhine State, NE of Gwa, Elephant Camp in Elephant Sanctuary , 17°38.9'N 94°38.1'E, 3.XII.2000, leaf litter in secondary forest, K.J. Ribardo leg., 2♀♀ ( CAS) .

Hosts. Unknown.

Distribution. China (Jiangxi), Malaysia ( Sarawak, Malaya), Myanmar (Rakhine).

5. Gonatopus muiri ( Olmi, 1984)

( Plate 113E, F )

Pseudogonatopus muiri Olmi 1984: 1251 ; Xu et al. 2011g (Suppl.): 20.

Description of Female. Apterous; length 2.5–3.0 mm. Holotype testaceous–reddish, except metasoma brown and antenna brown, with segments 1–2 testaceous (specimen from China, Bawangling National Nature Reserve, testaceous, except antenna brown, with segments 1–2 testaceous and segment 10 whitish, petiole black and metasoma brown). Antenna clavate; antennal segments of a specimen from China, Bawangling National Nature Reserve, in following proportions: 8:4.5:10:6:5:5:4.5:4.5:4:8. Head excavated; head of holotype dull and slightly granulated; head of specimen from China, Bawangling National Nature Reserve, shiny and unsculptured; frontal line of specimen from China, Bawangling National Nature Reserve, complete; occipital carina absent; head of specimen from China, Bawangling National Nature Reserve, with POL = 0.5; OL = 1.5; OOL = 6. Palpal formula 3/ 2 in holotype, 4/ 2 in specimen from China, Bawangling National Nature Reserve. Pronotum crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat. Metanotum shiny, unsculptured, not hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded ( Plate 113E ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum with slight median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture weakly distinct, in holotype only visible near lateral protrusions of metanotum. Meso-metapleural suture of specimen from China, Bawangling National Nature Reserve, visible only between mesopleuron and metapleuron, not visible near lateral protrusions of metanotum. Protarsal segments of specimen from China, Bawangling National Nature Reserve, in following proportions: 10:2.5:4:10:17; in holotype segment 1 of protarsus as long as segment 4. Enlarged claw ( Plate 113F ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 113F ) with two rows of 12–17 lamellae; distal apex with 11–12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Java, Pekalongan, F. Muir leg. ( AMNH) . Other material. CHINA: Hainan, Bawangling National Nature Reserve, Reserve House , about 1000 m, parasitized host collected 19.VIII.2000, dryinid cocooning 22.VIII.2000, M. Olmi reared from an unidentified species of Delphacidae , 1♀ ( MOLC) .

Hosts. In China (Hainan), unidentified species of Delphacidae .

Distribution. China (Hainan), Indonesia (Java).

6. Gonatopus nepalensis ( Olmi, 1986) , comb. nov.

( Plate 113G, H )

Pseudogonatopus nepalensis Olmi 1986: 80 .

Description of Female. Apterous; length 4.8–5.0 mm. Head black or brown, except mandible, clypeus and anterior region of face testaceous; antenna brown, except segments 1–2 testaceous and 10 whitish; occasionally antennal segment 9 whitish; mesosoma and metasoma black; legs black or brown, except chela and meso- and metatarsus testaceous. Antenna clavate; antennal segments in following proportions: 12:6:18:10:7:7:7:6.5:6.5:10.5. Head excavated, dull, granulated; frontal line complete; occipital carina absent. Palpal formula 4/2. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum shiny, sculptured by longitudinal striae. Metanotum transversely striate, with sides protruding; protrusions rounded ( Plate 113G ). Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface and metapleura. Mesopleuron shiny, smooth, unsculptured, not transversely striate. Meso-metapleural suture partly distinct, only present between mesopleuron and metapleuron, not present dorsally on sides of metanotum. Protarsal segments in following proportions: 18:4:8:22:35. Enlarged claw ( Plate 113G ) with one large subapical tooth and one row of seven lamellae. Segment 5 of protarsus ( Plate 113G ) with two rows of about 21 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: NEPAL: Kosi, S of Mangsingma Forest ( MHNG). Paratypes: same locality label as holotype, 1♀ ( AMNH); E Nepal, Kosi, Induwa Kola Valley, 1♀ ( MHNG). Other material. NEPAL: Khala Valley, 2050 m ( PMA). Palaearctic: according to Ponomarenko (1992), G. nepalensis is also present in the following localities: RUSSIA: Transbaikalia, 5.VI.1963, 10.VII.1963, A. Emelianov leg., 2♀♀ ( Ponomarenko , 1992); Fra East, Primorskiy Krai, Gornotayezhnoe, 19.VI.1983, Sciarkov leg., 1♀ ( Ponomarenko , 1992). This quotation is doubtful.

Hosts. Unknown.

Distribution. Nepal (Kosi), Russia (Transbaicalia, Primorskiy Krai).

7. Gonatopus nigricans (R. Perkins, 1905)

( Plates 111B , 114A – D )

Paragonatopus nigricans R. Perkins 1905: 41 ; Kieffer 1914b: 87; Olmi 1979: 218.

Pseudogonatopus melanacrias R. Perkins 1906a: 487 (synonymized by R. Perkins 1912); type locality: Suva ( Viti Levu Island,

Fiji Islands). Gonatopus fulgori Nakagawa 1906: 163 ; type locality: Japan; Guglielmino & Olmi 1997: 181, 184; syn. nov. Pseudogonatopus hospes R. Perkins 1912: 13 (synonymized by He & Xu 2002); type locality: China; Kieffer 1914b: 81;

Swezey 1928: 670; F. Williams 1931: 247; Chiu 1979: 339; Móczár 1979: 77; Olmi 1984: 1290; Olmi 2000a: 101, 111. Gonatopus sauteri Strand 1913: 209 (synonymized by Olmi 1984); type locality: Tainan ( Taiwan, China). Dicondylus sauteri (Strand) : Kieffer 1914b: 77. Pseudogonatopus sogatea Rohwer 1920: 160 (printing-error; holotype labelled sogatae ); type locality: Pusa (Bihar, India);

Kurian 1954b: 417; Olmi 1984: 1254; syn. nov. Haplogonatopus fulgori (Nakagawa) : Esaki & Hashimoto 1930: 21. Pseudozonatopus (sic) hospes R. Perkins : Pagden 1934: 2. Paragonatopus fulgori (Nakagawa) : Mochizuki 1943: 99; Chen 1988: 44. nec Pseudogonatopus nr. hospes R. Perkins : Chandra 1978b: 5. Pseudogonatopus fulgori (Nakagawa) : Olmi 1984: 1194. Pseudogonatopus pusanus Olmi 1984: 1255 ; type locality: Pusa (Bihar, India); He & Xu 2002: 355; syn. nov. Pseudogonatopus nigricans (R. Perkins) : Olmi 1984: 1287; Olmi 1990b: 789; Olmi 1991: 319. Gonatopus nigricans (R. Perkins) : Guglielmino & Olmi 1997: 165, 224–226, 250–252; Olmi 1999b: 28; Olmi 2000a: 94; He &

Xu 2002: 364; Guglielmino & Olmi 2006: 49. Gonatopus sogatae Rohwer : Guglielmino & Olmi 1997: 225, 226; He & Xu 2002: 355. Gonatopus insulanus He & Xu in He et al. 1998: 172; type locality: Qionghai (Hainan, China); syn. nov.

Description of female ( Plate 114A ): apterous; length 2.4–4.4 mm. Head brown-reddish, or brown, or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 or 1–3 yellow; mesosoma black or brown; occasionally posterior margin of pronotum and posterior apex of propodeum testaceous; metasoma black or brown; legs yellow; occasionally clubs of femora and part of coxae brown. In G. pusanus the colour is different: head partly or almost totally brown; antenna brown, except segments 1–2 testaceous; mesosoma testaceous-reddish, except sides of scutum darkened; occasionally part of pronotum darkened; metasoma brown; petiole black; legs testaceous; occasionally club of profemur darkened. In G. sogatae the colour is the following: head brown, except mandible and clypeus yellow; antenna brown, except segment 1 yellow and 2–3 brown-testaceous; mesosoma brown, except scutum reddish and distal apex of propodeum yellow; petiole black; metasoma brown-dark; legs brown-yellow. Intermediate colours are known. Antenna clavate; antennal segments in following proportions: 19:19:52:25:22:22:18:17:15:26. Head ( Plate 114B ) excavated, shiny, granulated or unsculptured; occiput and anterior region of face always granulated; POL = 5; OL = 5; OOL = 14. Palpal formula 2/2, 3/2, 4/2. Pronotum granulated, crossed by strong transverse impression. Scutum granulated, without lateral pointed apophyses. Mesopleura transversely striate. Metanotum flat, very long, about as long as scutum, transversely striate, not hollow behind scutellum ( Plate 114C ), with sides rounded ( Plate 114A ). Metathorax + propodeum shiny, indistinctly sculptured, with or without track of median longitudinal furrow, with posterior surface transversely striate. Metapleuron transversely striate. Meso-metapleural suture obsolete. Protarsal segment 4 shorter than segment 1 (40:47; or 20:27). Enlarged claw ( Plate 114D ) with one large subapical tooth and one row of three–nine lamellae. Segment 5 of protarsus ( Plate 114D ) with two rows of 9–25 lamellae; distal apex with 4–10 lamellae; occasionally lamellae extended continuously to distal apex. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9–2.4 mm. Head brown or black (occasionally brown-reddish), except mandible testaceous; occasionally part of face testaceous; antenna brown (in type of G. insulanus , testaceous-dark); mesosoma and metasoma black or brown (in type of G. insulanus , mesosoma reddish-dark, with propodeum and pleura darker); legs yellow. Antenna filiform; antennal segments in following proportions: 5:5:9:8:8:9:7:8:7:10 (in other specimens 6:5.5:10:9:8:9:8:8:7:10); antennal segment 3 more than three times as long as broad (4.0; 4.3; 5.0). Head shiny, hairy, granulated; frontal line absent; temple distinct; occipital carina absent; POL = 8; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OL (4:3) (in other specimens POL = 7; OL = 3; OOL = 3.5; greatest breadth of posterior ocelli longer than OL (3.5:3)). Palpal formula 2/2, 3/2, 4/2. Scutum shiny, granulated. Notauli complete, posteriorly joint or almost joint in small specimens, separated in large specimens; in very large specimens, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2.5:4; or 2:4). Scutellum and metanotum shiny, finely punctate, unsculptured among punctures. Propodeum reticulate rugose; dorsal surface with median longitudinal furrow and occasionally with two lateral smooth areas. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 111B ) much shorter than paramere. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nigricans : AUSTRALIA: Queensland, Bundaberg, 19.XI.1904, reared ( BPBM); ♀ lectotype (designated by Olmi, 1982) of P. melanacrias : FIJI ISLANDS: Viti Levu Island, Suva, III.1905 , reared from delphacid near Stenocranus , on Zoysia pungens (BPBM) ; ♀ lectotype (designated by Olmi, 1982) of P. hospes : CHINA: no localities, F. Muir leg., reared in Honolulu from Perkinsiella saccharicida from Chinese stock, IV–V.1907 ( BPBM); ♀ holotype of G. sauteri : CHINA: Taiwan, Tainan, IV.1912, H. Sauter leg. ( DEI); ♀ holotype of P. pusanus : INDIA: Bihar, Pusa ( IARI); ♀ holotype of P. sogatae : INDIA: Bihar, Pusa, 6.VIII.1915, reared from Sogata sp. , C.S. Misra leg. ( USNM); ♂ holotype of G. insulanus : CHINA: Hainan, Qionghai, 4.V.1964, Junhua He leg. ( ZJUC). Type material of G. fulgori probably lost. Paratypes of P. nigricans : AUSTRALIA: same locality label as lectotype, 1♀ lebelled for mistake Fiji Islands ( BPBM); same locality label as holotype, parasitized host collected 1.X.1904, dryinid adult emerged 21.X.1904, R. Perkins leg., 1♀ “ paratype ” identified by the same R. Perkins ( BMNH); AUSTRALIA: Queensland, Bundaberg, Sandhills, X.1904, bred, 1♀ ( ANIC). Paratypes of P. melanacrias : same locality label as lectotype, 2♀♀ ( BPBM). Paratypes of P. hospes : same locality label as lectotype, 3♀♀ ( BPBM). Paratype of P. pusanus : same locality label as holotype, 1♀ ( IARI). Paratype of P. sogatae : same locality label as holotype, misidentified ♂ allotype not belonging to the same species of holotype. Other material. CHINA: Fujian, Shaxian, reared from rice planthoppers, 1979–1983, Changming Liu leg., 5♀♀ and 1♂ ( ZJUC); Fujian, IV.1987, 31.VII.1985, Changming Liu leg., 6♀♀ and 10♂ ( ZJUC); Guangdong, Yangjiang, 1979, 1981, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangdong, Xingfeng County, Mt. Yunjinshan, 9.VI.2002, 18.VII.2003, 19.VII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); same locality label, 20.VII.2003, Ping Li leg., 2♀♀ ( ZJUC); same locality label, Zaifu Xu leg., many ♀♀ ( ZJUC, AMNH); Guangdong, Shixing County, Chebaling National Nature Reserve, 21.VIII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, M. Olmi reared from adult of unidentified Delphacid, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); Guangdong, Mt. Nankunshan, 2.VII.2005, Zaifu Xu leg., 1♀ ( ZJUC); Guangdong, Yangjiang, Hailing Town, 1980, Renhuan Hou leg., 2♀♀ and 2♂ ( ZJUC); Guangdong, Huaiji County, 21–22.X.2007, Zaifu Xu leg., 1♀ ( SCAU); Guangdong, Conghua, 28.VI.1978, Junhua He leg., 1♀ ( SCAU); Guangdong, Zhaoqing, V.1998, Yufang Liu leg., 2♀♀ ( SCAU); Guizhou, Sinan, IX.1986, VIII.1991, 5♀♀ ( ZJUC); Guizhou, Sandu, VI–VII.1981, Dongxiang Xie leg., 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Hunan, Yueyang, 15.VIII.1975, Binsen Wang leg., 1♀ ( ZJUC); Guangxi, Luchuan, 30.VII.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Bama, 1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Luocheng, 18.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Yaxian, VI.1966, Junhua He leg., 1♀ ( ZJUC); Hong Kong, Tai Lung Farm, IX–XI.1965, swept above rice, I.W.B. Thornton leg., 1♀ ( BMNH); Macau ( BPBM); same locality label, R. Perkins collection, 2♀♀ ( USNM); Taiwan, Kaohsiung, Kosen ( AMNH, ELKU); Taiwan, Changhus, Erlin ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, Kungkuan ( TARI); Taiwan, Taihoku Pref. (=Taipei), 27.IX.1930, J. Sonan leg., 1♀ ( TARI); Taiwan, Tainan, ShinShi, 11.IX.1979, paddy field, Hirashima leg., 1♀ ( ELKU); Taiwan, Gakko ( ELKU); Itahashi (= Taiwan), 16.IX.1930, J. Sonan leg., 5♀♀ ( TARI); Takeyama (=Jhushan), 6.X.1937, J. Sonan leg., 3♀♀ ( TARI); Inrin (=Yuanlin), 8.X.1937, J. Sonan leg., 39♀♀ ( TARI). INDIA: Chhattisgarh, Raipur ( GCC, AMNH); Delhi, New Delhi ( IARI); Orissa, Cuttack, 1981, 1♀ ( BMNH); Orissa, Puri, 28.XI.1987, S.V. Fowler leg., 1♀ ( BMNH); Tamil Nadu, Tanjavur (=Tanjore) District, 28.IX.1987, ex Nilaparvata lugens on rice, C. Peter leg., 3♀♀ ( BMNH); West Bengal, Hooghly District, Khanakul, 12.XII.1986, in paddy seedbed, 1♀ ( BMNH). INDONESIA: Java, Pekalongan ( BPBM); Java, Dieng Plateau ( BPBM); Borneo, Kalimantan, Paloh ( BMNH). MALAYSIA: Malaya, Selangor, 13.VIII.1928, H.T. Pagden leg., 1♀ ( BMNH); Malaya, Selangor, Kuala Lumpur, 21.XII.1930, H. T. Pagden leg., 1♀ ( BMNH); same locality label, many ♀♀ and ♂ ( NMNH, KYC); Malaya, Seberang Perai ( MARDI); Malaya, Serdang ( DBUS); Sabah, Tuaran ( KYC); Sarawak, 1st. Division, Payah Paloh, 28.I.1965, G.H.L. Rotschild leg., 1♀ ( BMNH). THAILAND: Songkhla, Hadd Tai ( KYC); Non Bua Lampoo, Na Kam Hai ( KYC); Tung Saew ( KYC); Tung Fabad ( KYC); Chainat ( KYC, AMNH); Banmakok ( KYC); Mae Tang ( KYC); Lampamg ( ELKU); Mae-Theng ( ELKU); Hot ( ELKU); Chieng Dao ( ELKU); Thailand, 15.IX.1999, reared from Nilaparvata lugens (Stål) , Sawai leg., 4♂ ( BMNH). Australian region: G. nigricans is known from Australia (Queensland), Fiji Islands, Solomon Islands and USA (Hawaii) ( Olmi, 1984). Introduced in the Hawaii 1906 and 1907 from China, Guangdong, for biological control of Perkinsiella saccharicida Kirkaldy in sugarcane fields ( Olmi, 2000a).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in Australia: Liburnia sp. , Stenocranus sp. ; in China: Laodelphax striatellus (Fallén) ( Yang et al., 1982; He & Xu, 2002), Perkinsiella saccharicida Kirkaldy ; in Malaysia: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in Thailand: Nilaparvata lugens (Stål) ( Chiu, 1979) ; in the USA (Hawaii): Perkinsiella saccharicida Kirkaldy (introduced for biological control).

Biology. Information by Pagden (1934); the parthenogenesis is thelytokous.

Natural enemies. In the USA (Hawaii) ( Swezey, 1919; Timberlake, 1919; Muir, 1921; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Encyrtidae : Helegonatopus pseudophanes R. Perkins, 1906 ; Cheiloneurus flaccus (Walker 1847) ( Guerrieri & Viggiani, 2005; Guerrieri, 2006); Aphelinidae : Centrodora xiphidii (R. Perkins, 1906) . In Malaya ( Pagden, 1934; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Ceraphronidae : Allomicrops abnormis (R. Perkins, 1910) ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp.

Distribution. China ( Taiwan, Hainan, Fujian, Guangdong, Guizhou, Hubei, Hunan, Guangxi, Hong Kong, Macau), Australia (Queensland), Fiji Islands ( Viti Levu), India (Bihar, Chhattisgarh, Delhi, Orissa, Tamil Nadu), Indonesia (Java, Pekalongan, Borneo), Malaysia (Malaya, Sabah, Sarawak), Thailand (Songkhla, Non Bua Lampoo, Tung Saew, Tung Fabad, Chainat, Banmakok, Mae Tang, Lampamg, Mae-Theng, Chieng Dao).

Remarks. Gonatopus insulanus He & Xu is here synonymized with G. nigricans , because of the study of its type and comparison with male specimens of G. nigricans . There are no differences.

Gonatopus nigricans (= Pseudogonatopus hospes R. Perkins ) was established in the Hawaii from cocoons sent from China ( Perkins, 1912). This species was considered extinct in the Hawaii ( Olmi, 1999a). However, in 2010 one of the authors (Olmi) saw a male specimen collected in Oahu Island, Mt. Olympus, in April 1981 by a pan trap. Probably so this species is always present at least in Oahu Island, although rare. Gonatopus pusanus and G. sogatae are here synonymized with G. nigricans because the only differences concern the colour of mesosoma (black in nigricans , testaceous to reddish, with some darkened areas, in pusanus ; brown, with some areas reddish or yellow, in sogatae ). In the collections, there are also many specimens with intermediate colours, so that the synonymy is evident. Gonatopus fulgori is here synonymized with G. nigricans , because the original description is like that of G. nigricans (the type material of G. fulgori is lost). In the past these two species were separated, because attributed to two different zoogeographic regions ( fulgori in Palaearctic, nigricans in Oriental and Australian regions).

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

5. Gonatopus muiri ( Olmi, 1984)

( Plate 113E, F )

Pseudogonatopus muiri Olmi 1984: 1251 ; Xu et al. 2011g (Suppl.): 20.

Description of Female. Apterous; length 2.5–3.0 mm. Holotype testaceous–reddish, except metasoma brown and antenna brown, with segments 1–2 testaceous (specimen from China, Bawangling National Nature Reserve, testaceous, except antenna brown, with segments 1–2 testaceous and segment 10 whitish, petiole black and metasoma brown). Antenna clavate; antennal segments of a specimen from China, Bawangling National Nature Reserve, in following proportions: 8:4.5:10:6:5:5:4.5:4.5:4:8. Head excavated; head of holotype dull and slightly granulated; head of specimen from China, Bawangling National Nature Reserve, shiny and unsculptured; frontal line of specimen from China, Bawangling National Nature Reserve, complete; occipital carina absent; head of specimen from China, Bawangling National Nature Reserve, with POL = 0.5; OL = 1.5; OOL = 6. Palpal formula 3/ 2 in holotype, 4/ 2 in specimen from China, Bawangling National Nature Reserve. Pronotum crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum shiny, unsculptured, laterally without pointed apophyses. Scutellum shiny, smooth, flat. Metanotum shiny, unsculptured, not hollow behind scutellum; sides of metanotum protruding; lateral protrusions rounded ( Plate 113E ). Metathorax + propodeum shiny, hairless, with disc and anterior surface unsculptured; disc of metathorax + propodeum with slight median longitudinal furrow; posterior surface of propodeum transversely striate. Meso-metapleural suture weakly distinct, in holotype only visible near lateral protrusions of metanotum. Meso-metapleural suture of specimen from China, Bawangling National Nature Reserve, visible only between mesopleuron and metapleuron, not visible near lateral protrusions of metanotum. Protarsal segments of specimen from China, Bawangling National Nature Reserve, in following proportions: 10:2.5:4:10:17; in holotype segment 1 of protarsus as long as segment 4. Enlarged claw ( Plate 113F ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 113F ) with two rows of 12–17 lamellae; distal apex with 11–12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Java, Pekalongan, F. Muir leg. ( AMNH) . Other material. CHINA: Hainan, Bawangling National Nature Reserve, Reserve House , about 1000 m, parasitized host collected 19.VIII.2000, dryinid cocooning 22.VIII.2000, M. Olmi reared from an unidentified species of Delphacidae , 1♀ ( MOLC) .

Hosts. In China (Hainan), unidentified species of Delphacidae .

Distribution. China (Hainan), Indonesia (Java).

6. Gonatopus nepalensis ( Olmi, 1986) , comb. nov.

( Plate 113G, H )

Pseudogonatopus nepalensis Olmi 1986: 80 .

Description of Female. Apterous; length 4.8–5.0 mm. Head black or brown, except mandible, clypeus and anterior region of face testaceous; antenna brown, except segments 1–2 testaceous and 10 whitish; occasionally antennal segment 9 whitish; mesosoma and metasoma black; legs black or brown, except chela and meso- and metatarsus testaceous. Antenna clavate; antennal segments in following proportions: 12:6:18:10:7:7:7:6.5:6.5:10.5. Head excavated, dull, granulated; frontal line complete; occipital carina absent. Palpal formula 4/2. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum shiny, sculptured by longitudinal striae. Metanotum transversely striate, with sides protruding; protrusions rounded ( Plate 113G ). Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface and metapleura. Mesopleuron shiny, smooth, unsculptured, not transversely striate. Meso-metapleural suture partly distinct, only present between mesopleuron and metapleuron, not present dorsally on sides of metanotum. Protarsal segments in following proportions: 18:4:8:22:35. Enlarged claw ( Plate 113G ) with one large subapical tooth and one row of seven lamellae. Segment 5 of protarsus ( Plate 113G ) with two rows of about 21 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: NEPAL: Kosi, S of Mangsingma Forest ( MHNG). Paratypes: same locality label as holotype, 1♀ ( AMNH); E Nepal, Kosi, Induwa Kola Valley, 1♀ ( MHNG). Other material. NEPAL: Khala Valley, 2050 m ( PMA). Palaearctic: according to Ponomarenko (1992), G. nepalensis is also present in the following localities: RUSSIA: Transbaikalia, 5.VI.1963, 10.VII.1963, A. Emelianov leg., 2♀♀ ( Ponomarenko , 1992); Fra East, Primorskiy Krai, Gornotayezhnoe, 19.VI.1983, Sciarkov leg., 1♀ ( Ponomarenko , 1992). This quotation is doubtful.

Hosts. Unknown.

Distribution. Nepal (Kosi), Russia (Transbaicalia, Primorskiy Krai).

7. Gonatopus nigricans (R. Perkins, 1905)

( Plates 111B , 114A – D )

Paragonatopus nigricans R. Perkins 1905: 41 ; Kieffer 1914b: 87; Olmi 1979: 218.

Pseudogonatopus melanacrias R. Perkins 1906a: 487 (synonymized by R. Perkins 1912); type locality: Suva ( Viti Levu Island,

Fiji Islands). Gonatopus fulgori Nakagawa 1906: 163 ; type locality: Japan; Guglielmino & Olmi 1997: 181, 184; syn. nov. Pseudogonatopus hospes R. Perkins 1912: 13 (synonymized by He & Xu 2002); type locality: China; Kieffer 1914b: 81;

Swezey 1928: 670; F. Williams 1931: 247; Chiu 1979: 339; Móczár 1979: 77; Olmi 1984: 1290; Olmi 2000a: 101, 111. Gonatopus sauteri Strand 1913: 209 (synonymized by Olmi 1984); type locality: Tainan ( Taiwan, China). Dicondylus sauteri (Strand) : Kieffer 1914b: 77. Pseudogonatopus sogatea Rohwer 1920: 160 (printing-error; holotype labelled sogatae ); type locality: Pusa (Bihar, India);

Kurian 1954b: 417; Olmi 1984: 1254; syn. nov. Haplogonatopus fulgori (Nakagawa) : Esaki & Hashimoto 1930: 21. Pseudozonatopus (sic) hospes R. Perkins : Pagden 1934: 2. Paragonatopus fulgori (Nakagawa) : Mochizuki 1943: 99; Chen 1988: 44. nec Pseudogonatopus nr. hospes R. Perkins : Chandra 1978b: 5. Pseudogonatopus fulgori (Nakagawa) : Olmi 1984: 1194. Pseudogonatopus pusanus Olmi 1984: 1255 ; type locality: Pusa (Bihar, India); He & Xu 2002: 355; syn. nov. Pseudogonatopus nigricans (R. Perkins) : Olmi 1984: 1287; Olmi 1990b: 789; Olmi 1991: 319. Gonatopus nigricans (R. Perkins) : Guglielmino & Olmi 1997: 165, 224–226, 250–252; Olmi 1999b: 28; Olmi 2000a: 94; He &

Xu 2002: 364; Guglielmino & Olmi 2006: 49. Gonatopus sogatae Rohwer : Guglielmino & Olmi 1997: 225, 226; He & Xu 2002: 355. Gonatopus insulanus He & Xu in He et al. 1998: 172; type locality: Qionghai (Hainan, China); syn. nov.

Description of female ( Plate 114A ): apterous; length 2.4–4.4 mm. Head brown-reddish, or brown, or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 or 1–3 yellow; mesosoma black or brown; occasionally posterior margin of pronotum and posterior apex of propodeum testaceous; metasoma black or brown; legs yellow; occasionally clubs of femora and part of coxae brown. In G. pusanus the colour is different: head partly or almost totally brown; antenna brown, except segments 1–2 testaceous; mesosoma testaceous-reddish, except sides of scutum darkened; occasionally part of pronotum darkened; metasoma brown; petiole black; legs testaceous; occasionally club of profemur darkened. In G. sogatae the colour is the following: head brown, except mandible and clypeus yellow; antenna brown, except segment 1 yellow and 2–3 brown-testaceous; mesosoma brown, except scutum reddish and distal apex of propodeum yellow; petiole black; metasoma brown-dark; legs brown-yellow. Intermediate colours are known. Antenna clavate; antennal segments in following proportions: 19:19:52:25:22:22:18:17:15:26. Head ( Plate 114B ) excavated, shiny, granulated or unsculptured; occiput and anterior region of face always granulated; POL = 5; OL = 5; OOL = 14. Palpal formula 2/2, 3/2, 4/2. Pronotum granulated, crossed by strong transverse impression. Scutum granulated, without lateral pointed apophyses. Mesopleura transversely striate. Metanotum flat, very long, about as long as scutum, transversely striate, not hollow behind scutellum ( Plate 114C ), with sides rounded ( Plate 114A ). Metathorax + propodeum shiny, indistinctly sculptured, with or without track of median longitudinal furrow, with posterior surface transversely striate. Metapleuron transversely striate. Meso-metapleural suture obsolete. Protarsal segment 4 shorter than segment 1 (40:47; or 20:27). Enlarged claw ( Plate 114D ) with one large subapical tooth and one row of three–nine lamellae. Segment 5 of protarsus ( Plate 114D ) with two rows of 9–25 lamellae; distal apex with 4–10 lamellae; occasionally lamellae extended continuously to distal apex. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9–2.4 mm. Head brown or black (occasionally brown-reddish), except mandible testaceous; occasionally part of face testaceous; antenna brown (in type of G. insulanus , testaceous-dark); mesosoma and metasoma black or brown (in type of G. insulanus , mesosoma reddish-dark, with propodeum and pleura darker); legs yellow. Antenna filiform; antennal segments in following proportions: 5:5:9:8:8:9:7:8:7:10 (in other specimens 6:5.5:10:9:8:9:8:8:7:10); antennal segment 3 more than three times as long as broad (4.0; 4.3; 5.0). Head shiny, hairy, granulated; frontal line absent; temple distinct; occipital carina absent; POL = 8; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OL (4:3) (in other specimens POL = 7; OL = 3; OOL = 3.5; greatest breadth of posterior ocelli longer than OL (3.5:3)). Palpal formula 2/2, 3/2, 4/2. Scutum shiny, granulated. Notauli complete, posteriorly joint or almost joint in small specimens, separated in large specimens; in very large specimens, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2.5:4; or 2:4). Scutellum and metanotum shiny, finely punctate, unsculptured among punctures. Propodeum reticulate rugose; dorsal surface with median longitudinal furrow and occasionally with two lateral smooth areas. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 111B ) much shorter than paramere. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nigricans : AUSTRALIA: Queensland, Bundaberg, 19.XI.1904, reared ( BPBM); ♀ lectotype (designated by Olmi, 1982) of P. melanacrias : FIJI ISLANDS: Viti Levu Island, Suva, III.1905 , reared from delphacid near Stenocranus , on Zoysia pungens (BPBM) ; ♀ lectotype (designated by Olmi, 1982) of P. hospes : CHINA: no localities, F. Muir leg., reared in Honolulu from Perkinsiella saccharicida from Chinese stock, IV–V.1907 ( BPBM); ♀ holotype of G. sauteri : CHINA: Taiwan, Tainan, IV.1912, H. Sauter leg. ( DEI); ♀ holotype of P. pusanus : INDIA: Bihar, Pusa ( IARI); ♀ holotype of P. sogatae : INDIA: Bihar, Pusa, 6.VIII.1915, reared from Sogata sp. , C.S. Misra leg. ( USNM); ♂ holotype of G. insulanus : CHINA: Hainan, Qionghai, 4.V.1964, Junhua He leg. ( ZJUC). Type material of G. fulgori probably lost. Paratypes of P. nigricans : AUSTRALIA: same locality label as lectotype, 1♀ lebelled for mistake Fiji Islands ( BPBM); same locality label as holotype, parasitized host collected 1.X.1904, dryinid adult emerged 21.X.1904, R. Perkins leg., 1♀ “ paratype ” identified by the same R. Perkins ( BMNH); AUSTRALIA: Queensland, Bundaberg, Sandhills, X.1904, bred, 1♀ ( ANIC). Paratypes of P. melanacrias : same locality label as lectotype, 2♀♀ ( BPBM). Paratypes of P. hospes : same locality label as lectotype, 3♀♀ ( BPBM). Paratype of P. pusanus : same locality label as holotype, 1♀ ( IARI). Paratype of P. sogatae : same locality label as holotype, misidentified ♂ allotype not belonging to the same species of holotype. Other material. CHINA: Fujian, Shaxian, reared from rice planthoppers, 1979–1983, Changming Liu leg., 5♀♀ and 1♂ ( ZJUC); Fujian, IV.1987, 31.VII.1985, Changming Liu leg., 6♀♀ and 10♂ ( ZJUC); Guangdong, Yangjiang, 1979, 1981, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangdong, Xingfeng County, Mt. Yunjinshan, 9.VI.2002, 18.VII.2003, 19.VII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); same locality label, 20.VII.2003, Ping Li leg., 2♀♀ ( ZJUC); same locality label, Zaifu Xu leg., many ♀♀ ( ZJUC, AMNH); Guangdong, Shixing County, Chebaling National Nature Reserve, 21.VIII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, M. Olmi reared from adult of unidentified Delphacid, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); Guangdong, Mt. Nankunshan, 2.VII.2005, Zaifu Xu leg., 1♀ ( ZJUC); Guangdong, Yangjiang, Hailing Town, 1980, Renhuan Hou leg., 2♀♀ and 2♂ ( ZJUC); Guangdong, Huaiji County, 21–22.X.2007, Zaifu Xu leg., 1♀ ( SCAU); Guangdong, Conghua, 28.VI.1978, Junhua He leg., 1♀ ( SCAU); Guangdong, Zhaoqing, V.1998, Yufang Liu leg., 2♀♀ ( SCAU); Guizhou, Sinan, IX.1986, VIII.1991, 5♀♀ ( ZJUC); Guizhou, Sandu, VI–VII.1981, Dongxiang Xie leg., 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Hunan, Yueyang, 15.VIII.1975, Binsen Wang leg., 1♀ ( ZJUC); Guangxi, Luchuan, 30.VII.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Bama, 1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Luocheng, 18.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Yaxian, VI.1966, Junhua He leg., 1♀ ( ZJUC); Hong Kong, Tai Lung Farm, IX–XI.1965, swept above rice, I.W.B. Thornton leg., 1♀ ( BMNH); Macau ( BPBM); same locality label, R. Perkins collection, 2♀♀ ( USNM); Taiwan, Kaohsiung, Kosen ( AMNH, ELKU); Taiwan, Changhus, Erlin ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, Kungkuan ( TARI); Taiwan, Taihoku Pref. (=Taipei), 27.IX.1930, J. Sonan leg., 1♀ ( TARI); Taiwan, Tainan, ShinShi, 11.IX.1979, paddy field, Hirashima leg., 1♀ ( ELKU); Taiwan, Gakko ( ELKU); Itahashi (= Taiwan), 16.IX.1930, J. Sonan leg., 5♀♀ ( TARI); Takeyama (=Jhushan), 6.X.1937, J. Sonan leg., 3♀♀ ( TARI); Inrin (=Yuanlin), 8.X.1937, J. Sonan leg., 39♀♀ ( TARI). INDIA: Chhattisgarh, Raipur ( GCC, AMNH); Delhi, New Delhi ( IARI); Orissa, Cuttack, 1981, 1♀ ( BMNH); Orissa, Puri, 28.XI.1987, S.V. Fowler leg., 1♀ ( BMNH); Tamil Nadu, Tanjavur (=Tanjore) District, 28.IX.1987, ex Nilaparvata lugens on rice, C. Peter leg., 3♀♀ ( BMNH); West Bengal, Hooghly District, Khanakul, 12.XII.1986, in paddy seedbed, 1♀ ( BMNH). INDONESIA: Java, Pekalongan ( BPBM); Java, Dieng Plateau ( BPBM); Borneo, Kalimantan, Paloh ( BMNH). MALAYSIA: Malaya, Selangor, 13.VIII.1928, H.T. Pagden leg., 1♀ ( BMNH); Malaya, Selangor, Kuala Lumpur, 21.XII.1930, H. T. Pagden leg., 1♀ ( BMNH); same locality label, many ♀♀ and ♂ ( NMNH, KYC); Malaya, Seberang Perai ( MARDI); Malaya, Serdang ( DBUS); Sabah, Tuaran ( KYC); Sarawak, 1st. Division, Payah Paloh, 28.I.1965, G.H.L. Rotschild leg., 1♀ ( BMNH). THAILAND: Songkhla, Hadd Tai ( KYC); Non Bua Lampoo, Na Kam Hai ( KYC); Tung Saew ( KYC); Tung Fabad ( KYC); Chainat ( KYC, AMNH); Banmakok ( KYC); Mae Tang ( KYC); Lampamg ( ELKU); Mae-Theng ( ELKU); Hot ( ELKU); Chieng Dao ( ELKU); Thailand, 15.IX.1999, reared from Nilaparvata lugens (Stål) , Sawai leg., 4♂ ( BMNH). Australian region: G. nigricans is known from Australia (Queensland), Fiji Islands, Solomon Islands and USA (Hawaii) ( Olmi, 1984). Introduced in the Hawaii 1906 and 1907 from China, Guangdong, for biological control of Perkinsiella saccharicida Kirkaldy in sugarcane fields ( Olmi, 2000a).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in Australia: Liburnia sp. , Stenocranus sp. ; in China: Laodelphax striatellus (Fallén) ( Yang et al., 1982; He & Xu, 2002), Perkinsiella saccharicida Kirkaldy ; in Malaysia: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in Thailand: Nilaparvata lugens (Stål) ( Chiu, 1979) ; in the USA (Hawaii): Perkinsiella saccharicida Kirkaldy (introduced for biological control).

Biology. Information by Pagden (1934); the parthenogenesis is thelytokous.

Natural enemies. In the USA (Hawaii) ( Swezey, 1919; Timberlake, 1919; Muir, 1921; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Encyrtidae : Helegonatopus pseudophanes R. Perkins, 1906 ; Cheiloneurus flaccus (Walker 1847) ( Guerrieri & Viggiani, 2005; Guerrieri, 2006); Aphelinidae : Centrodora xiphidii (R. Perkins, 1906) . In Malaya ( Pagden, 1934; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Ceraphronidae : Allomicrops abnormis (R. Perkins, 1910) ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp.

Distribution. China ( Taiwan, Hainan, Fujian, Guangdong, Guizhou, Hubei, Hunan, Guangxi, Hong Kong, Macau), Australia (Queensland), Fiji Islands ( Viti Levu), India (Bihar, Chhattisgarh, Delhi, Orissa, Tamil Nadu), Indonesia (Java, Pekalongan, Borneo), Malaysia (Malaya, Sabah, Sarawak), Thailand (Songkhla, Non Bua Lampoo, Tung Saew, Tung Fabad, Chainat, Banmakok, Mae Tang, Lampamg, Mae-Theng, Chieng Dao).

Remarks. Gonatopus insulanus He & Xu is here synonymized with G. nigricans , because of the study of its type and comparison with male specimens of G. nigricans . There are no differences.

Gonatopus nigricans (= Pseudogonatopus hospes R. Perkins ) was established in the Hawaii from cocoons sent from China ( Perkins, 1912). This species was considered extinct in the Hawaii ( Olmi, 1999a). However, in 2010 one of the authors (Olmi) saw a male specimen collected in Oahu Island, Mt. Olympus, in April 1981 by a pan trap. Probably so this species is always present at least in Oahu Island, although rare. Gonatopus pusanus and G. sogatae are here synonymized with G. nigricans because the only differences concern the colour of mesosoma (black in nigricans , testaceous to reddish, with some darkened areas, in pusanus ; brown, with some areas reddish or yellow, in sogatae ). In the collections, there are also many specimens with intermediate colours, so that the synonymy is evident. Gonatopus fulgori is here synonymized with G. nigricans , because the original description is like that of G. nigricans (the type material of G. fulgori is lost). In the past these two species were separated, because attributed to two different zoogeographic regions ( fulgori in Palaearctic, nigricans in Oriental and Australian regions).

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

6. Gonatopus nepalensis ( Olmi, 1986) , comb. nov.

( Plate 113G, H )

Pseudogonatopus nepalensis Olmi 1986: 80 .

Description of Female. Apterous; length 4.8–5.0 mm. Head black or brown, except mandible, clypeus and anterior region of face testaceous; antenna brown, except segments 1–2 testaceous and 10 whitish; occasionally antennal segment 9 whitish; mesosoma and metasoma black; legs black or brown, except chela and meso- and metatarsus testaceous. Antenna clavate; antennal segments in following proportions: 12:6:18:10:7:7:7:6.5:6.5:10.5. Head excavated, dull, granulated; frontal line complete; occipital carina absent. Palpal formula 4/2. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum shiny, sculptured by longitudinal striae. Metanotum transversely striate, with sides protruding; protrusions rounded ( Plate 113G ). Metathorax + propodeum shiny, unsculptured, except transverse striae on posterior surface and metapleura. Mesopleuron shiny, smooth, unsculptured, not transversely striate. Meso-metapleural suture partly distinct, only present between mesopleuron and metapleuron, not present dorsally on sides of metanotum. Protarsal segments in following proportions: 18:4:8:22:35. Enlarged claw ( Plate 113G ) with one large subapical tooth and one row of seven lamellae. Segment 5 of protarsus ( Plate 113G ) with two rows of about 21 lamellae; distal apex with about 15 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: NEPAL: Kosi, S of Mangsingma Forest ( MHNG). Paratypes: same locality label as holotype, 1♀ ( AMNH); E Nepal, Kosi, Induwa Kola Valley, 1♀ ( MHNG). Other material. NEPAL: Khala Valley, 2050 m ( PMA). Palaearctic: according to Ponomarenko (1992), G. nepalensis is also present in the following localities: RUSSIA: Transbaikalia, 5.VI.1963, 10.VII.1963, A. Emelianov leg., 2♀♀ ( Ponomarenko , 1992); Fra East, Primorskiy Krai, Gornotayezhnoe, 19.VI.1983, Sciarkov leg., 1♀ ( Ponomarenko , 1992). This quotation is doubtful.

Hosts. Unknown.

Distribution. Nepal (Kosi), Russia (Transbaicalia, Primorskiy Krai).

7. Gonatopus nigricans (R. Perkins, 1905)

( Plates 111B , 114A – D )

Paragonatopus nigricans R. Perkins 1905: 41 ; Kieffer 1914b: 87; Olmi 1979: 218.

Pseudogonatopus melanacrias R. Perkins 1906a: 487 (synonymized by R. Perkins 1912); type locality: Suva ( Viti Levu Island,

Fiji Islands). Gonatopus fulgori Nakagawa 1906: 163 ; type locality: Japan; Guglielmino & Olmi 1997: 181, 184; syn. nov. Pseudogonatopus hospes R. Perkins 1912: 13 (synonymized by He & Xu 2002); type locality: China; Kieffer 1914b: 81;

Swezey 1928: 670; F. Williams 1931: 247; Chiu 1979: 339; Móczár 1979: 77; Olmi 1984: 1290; Olmi 2000a: 101, 111. Gonatopus sauteri Strand 1913: 209 (synonymized by Olmi 1984); type locality: Tainan ( Taiwan, China). Dicondylus sauteri (Strand) : Kieffer 1914b: 77. Pseudogonatopus sogatea Rohwer 1920: 160 (printing-error; holotype labelled sogatae ); type locality: Pusa (Bihar, India);

Kurian 1954b: 417; Olmi 1984: 1254; syn. nov. Haplogonatopus fulgori (Nakagawa) : Esaki & Hashimoto 1930: 21. Pseudozonatopus (sic) hospes R. Perkins : Pagden 1934: 2. Paragonatopus fulgori (Nakagawa) : Mochizuki 1943: 99; Chen 1988: 44. nec Pseudogonatopus nr. hospes R. Perkins : Chandra 1978b: 5. Pseudogonatopus fulgori (Nakagawa) : Olmi 1984: 1194. Pseudogonatopus pusanus Olmi 1984: 1255 ; type locality: Pusa (Bihar, India); He & Xu 2002: 355; syn. nov. Pseudogonatopus nigricans (R. Perkins) : Olmi 1984: 1287; Olmi 1990b: 789; Olmi 1991: 319. Gonatopus nigricans (R. Perkins) : Guglielmino & Olmi 1997: 165, 224–226, 250–252; Olmi 1999b: 28; Olmi 2000a: 94; He &

Xu 2002: 364; Guglielmino & Olmi 2006: 49. Gonatopus sogatae Rohwer : Guglielmino & Olmi 1997: 225, 226; He & Xu 2002: 355. Gonatopus insulanus He & Xu in He et al. 1998: 172; type locality: Qionghai (Hainan, China); syn. nov.

Description of female ( Plate 114A ): apterous; length 2.4–4.4 mm. Head brown-reddish, or brown, or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 or 1–3 yellow; mesosoma black or brown; occasionally posterior margin of pronotum and posterior apex of propodeum testaceous; metasoma black or brown; legs yellow; occasionally clubs of femora and part of coxae brown. In G. pusanus the colour is different: head partly or almost totally brown; antenna brown, except segments 1–2 testaceous; mesosoma testaceous-reddish, except sides of scutum darkened; occasionally part of pronotum darkened; metasoma brown; petiole black; legs testaceous; occasionally club of profemur darkened. In G. sogatae the colour is the following: head brown, except mandible and clypeus yellow; antenna brown, except segment 1 yellow and 2–3 brown-testaceous; mesosoma brown, except scutum reddish and distal apex of propodeum yellow; petiole black; metasoma brown-dark; legs brown-yellow. Intermediate colours are known. Antenna clavate; antennal segments in following proportions: 19:19:52:25:22:22:18:17:15:26. Head ( Plate 114B ) excavated, shiny, granulated or unsculptured; occiput and anterior region of face always granulated; POL = 5; OL = 5; OOL = 14. Palpal formula 2/2, 3/2, 4/2. Pronotum granulated, crossed by strong transverse impression. Scutum granulated, without lateral pointed apophyses. Mesopleura transversely striate. Metanotum flat, very long, about as long as scutum, transversely striate, not hollow behind scutellum ( Plate 114C ), with sides rounded ( Plate 114A ). Metathorax + propodeum shiny, indistinctly sculptured, with or without track of median longitudinal furrow, with posterior surface transversely striate. Metapleuron transversely striate. Meso-metapleural suture obsolete. Protarsal segment 4 shorter than segment 1 (40:47; or 20:27). Enlarged claw ( Plate 114D ) with one large subapical tooth and one row of three–nine lamellae. Segment 5 of protarsus ( Plate 114D ) with two rows of 9–25 lamellae; distal apex with 4–10 lamellae; occasionally lamellae extended continuously to distal apex. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9–2.4 mm. Head brown or black (occasionally brown-reddish), except mandible testaceous; occasionally part of face testaceous; antenna brown (in type of G. insulanus , testaceous-dark); mesosoma and metasoma black or brown (in type of G. insulanus , mesosoma reddish-dark, with propodeum and pleura darker); legs yellow. Antenna filiform; antennal segments in following proportions: 5:5:9:8:8:9:7:8:7:10 (in other specimens 6:5.5:10:9:8:9:8:8:7:10); antennal segment 3 more than three times as long as broad (4.0; 4.3; 5.0). Head shiny, hairy, granulated; frontal line absent; temple distinct; occipital carina absent; POL = 8; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OL (4:3) (in other specimens POL = 7; OL = 3; OOL = 3.5; greatest breadth of posterior ocelli longer than OL (3.5:3)). Palpal formula 2/2, 3/2, 4/2. Scutum shiny, granulated. Notauli complete, posteriorly joint or almost joint in small specimens, separated in large specimens; in very large specimens, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2.5:4; or 2:4). Scutellum and metanotum shiny, finely punctate, unsculptured among punctures. Propodeum reticulate rugose; dorsal surface with median longitudinal furrow and occasionally with two lateral smooth areas. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 111B ) much shorter than paramere. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nigricans : AUSTRALIA: Queensland, Bundaberg, 19.XI.1904, reared ( BPBM); ♀ lectotype (designated by Olmi, 1982) of P. melanacrias : FIJI ISLANDS: Viti Levu Island, Suva, III.1905 , reared from delphacid near Stenocranus , on Zoysia pungens (BPBM) ; ♀ lectotype (designated by Olmi, 1982) of P. hospes : CHINA: no localities, F. Muir leg., reared in Honolulu from Perkinsiella saccharicida from Chinese stock, IV–V.1907 ( BPBM); ♀ holotype of G. sauteri : CHINA: Taiwan, Tainan, IV.1912, H. Sauter leg. ( DEI); ♀ holotype of P. pusanus : INDIA: Bihar, Pusa ( IARI); ♀ holotype of P. sogatae : INDIA: Bihar, Pusa, 6.VIII.1915, reared from Sogata sp. , C.S. Misra leg. ( USNM); ♂ holotype of G. insulanus : CHINA: Hainan, Qionghai, 4.V.1964, Junhua He leg. ( ZJUC). Type material of G. fulgori probably lost. Paratypes of P. nigricans : AUSTRALIA: same locality label as lectotype, 1♀ lebelled for mistake Fiji Islands ( BPBM); same locality label as holotype, parasitized host collected 1.X.1904, dryinid adult emerged 21.X.1904, R. Perkins leg., 1♀ “ paratype ” identified by the same R. Perkins ( BMNH); AUSTRALIA: Queensland, Bundaberg, Sandhills, X.1904, bred, 1♀ ( ANIC). Paratypes of P. melanacrias : same locality label as lectotype, 2♀♀ ( BPBM). Paratypes of P. hospes : same locality label as lectotype, 3♀♀ ( BPBM). Paratype of P. pusanus : same locality label as holotype, 1♀ ( IARI). Paratype of P. sogatae : same locality label as holotype, misidentified ♂ allotype not belonging to the same species of holotype. Other material. CHINA: Fujian, Shaxian, reared from rice planthoppers, 1979–1983, Changming Liu leg., 5♀♀ and 1♂ ( ZJUC); Fujian, IV.1987, 31.VII.1985, Changming Liu leg., 6♀♀ and 10♂ ( ZJUC); Guangdong, Yangjiang, 1979, 1981, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangdong, Xingfeng County, Mt. Yunjinshan, 9.VI.2002, 18.VII.2003, 19.VII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); same locality label, 20.VII.2003, Ping Li leg., 2♀♀ ( ZJUC); same locality label, Zaifu Xu leg., many ♀♀ ( ZJUC, AMNH); Guangdong, Shixing County, Chebaling National Nature Reserve, 21.VIII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, M. Olmi reared from adult of unidentified Delphacid, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); Guangdong, Mt. Nankunshan, 2.VII.2005, Zaifu Xu leg., 1♀ ( ZJUC); Guangdong, Yangjiang, Hailing Town, 1980, Renhuan Hou leg., 2♀♀ and 2♂ ( ZJUC); Guangdong, Huaiji County, 21–22.X.2007, Zaifu Xu leg., 1♀ ( SCAU); Guangdong, Conghua, 28.VI.1978, Junhua He leg., 1♀ ( SCAU); Guangdong, Zhaoqing, V.1998, Yufang Liu leg., 2♀♀ ( SCAU); Guizhou, Sinan, IX.1986, VIII.1991, 5♀♀ ( ZJUC); Guizhou, Sandu, VI–VII.1981, Dongxiang Xie leg., 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Hunan, Yueyang, 15.VIII.1975, Binsen Wang leg., 1♀ ( ZJUC); Guangxi, Luchuan, 30.VII.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Bama, 1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Luocheng, 18.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Yaxian, VI.1966, Junhua He leg., 1♀ ( ZJUC); Hong Kong, Tai Lung Farm, IX–XI.1965, swept above rice, I.W.B. Thornton leg., 1♀ ( BMNH); Macau ( BPBM); same locality label, R. Perkins collection, 2♀♀ ( USNM); Taiwan, Kaohsiung, Kosen ( AMNH, ELKU); Taiwan, Changhus, Erlin ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, Kungkuan ( TARI); Taiwan, Taihoku Pref. (=Taipei), 27.IX.1930, J. Sonan leg., 1♀ ( TARI); Taiwan, Tainan, ShinShi, 11.IX.1979, paddy field, Hirashima leg., 1♀ ( ELKU); Taiwan, Gakko ( ELKU); Itahashi (= Taiwan), 16.IX.1930, J. Sonan leg., 5♀♀ ( TARI); Takeyama (=Jhushan), 6.X.1937, J. Sonan leg., 3♀♀ ( TARI); Inrin (=Yuanlin), 8.X.1937, J. Sonan leg., 39♀♀ ( TARI). INDIA: Chhattisgarh, Raipur ( GCC, AMNH); Delhi, New Delhi ( IARI); Orissa, Cuttack, 1981, 1♀ ( BMNH); Orissa, Puri, 28.XI.1987, S.V. Fowler leg., 1♀ ( BMNH); Tamil Nadu, Tanjavur (=Tanjore) District, 28.IX.1987, ex Nilaparvata lugens on rice, C. Peter leg., 3♀♀ ( BMNH); West Bengal, Hooghly District, Khanakul, 12.XII.1986, in paddy seedbed, 1♀ ( BMNH). INDONESIA: Java, Pekalongan ( BPBM); Java, Dieng Plateau ( BPBM); Borneo, Kalimantan, Paloh ( BMNH). MALAYSIA: Malaya, Selangor, 13.VIII.1928, H.T. Pagden leg., 1♀ ( BMNH); Malaya, Selangor, Kuala Lumpur, 21.XII.1930, H. T. Pagden leg., 1♀ ( BMNH); same locality label, many ♀♀ and ♂ ( NMNH, KYC); Malaya, Seberang Perai ( MARDI); Malaya, Serdang ( DBUS); Sabah, Tuaran ( KYC); Sarawak, 1st. Division, Payah Paloh, 28.I.1965, G.H.L. Rotschild leg., 1♀ ( BMNH). THAILAND: Songkhla, Hadd Tai ( KYC); Non Bua Lampoo, Na Kam Hai ( KYC); Tung Saew ( KYC); Tung Fabad ( KYC); Chainat ( KYC, AMNH); Banmakok ( KYC); Mae Tang ( KYC); Lampamg ( ELKU); Mae-Theng ( ELKU); Hot ( ELKU); Chieng Dao ( ELKU); Thailand, 15.IX.1999, reared from Nilaparvata lugens (Stål) , Sawai leg., 4♂ ( BMNH). Australian region: G. nigricans is known from Australia (Queensland), Fiji Islands, Solomon Islands and USA (Hawaii) ( Olmi, 1984). Introduced in the Hawaii 1906 and 1907 from China, Guangdong, for biological control of Perkinsiella saccharicida Kirkaldy in sugarcane fields ( Olmi, 2000a).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in Australia: Liburnia sp. , Stenocranus sp. ; in China: Laodelphax striatellus (Fallén) ( Yang et al., 1982; He & Xu, 2002), Perkinsiella saccharicida Kirkaldy ; in Malaysia: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in Thailand: Nilaparvata lugens (Stål) ( Chiu, 1979) ; in the USA (Hawaii): Perkinsiella saccharicida Kirkaldy (introduced for biological control).

Biology. Information by Pagden (1934); the parthenogenesis is thelytokous.

Natural enemies. In the USA (Hawaii) ( Swezey, 1919; Timberlake, 1919; Muir, 1921; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Encyrtidae : Helegonatopus pseudophanes R. Perkins, 1906 ; Cheiloneurus flaccus (Walker 1847) ( Guerrieri & Viggiani, 2005; Guerrieri, 2006); Aphelinidae : Centrodora xiphidii (R. Perkins, 1906) . In Malaya ( Pagden, 1934; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Ceraphronidae : Allomicrops abnormis (R. Perkins, 1910) ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp.

Distribution. China ( Taiwan, Hainan, Fujian, Guangdong, Guizhou, Hubei, Hunan, Guangxi, Hong Kong, Macau), Australia (Queensland), Fiji Islands ( Viti Levu), India (Bihar, Chhattisgarh, Delhi, Orissa, Tamil Nadu), Indonesia (Java, Pekalongan, Borneo), Malaysia (Malaya, Sabah, Sarawak), Thailand (Songkhla, Non Bua Lampoo, Tung Saew, Tung Fabad, Chainat, Banmakok, Mae Tang, Lampamg, Mae-Theng, Chieng Dao).

Remarks. Gonatopus insulanus He & Xu is here synonymized with G. nigricans , because of the study of its type and comparison with male specimens of G. nigricans . There are no differences.

Gonatopus nigricans (= Pseudogonatopus hospes R. Perkins ) was established in the Hawaii from cocoons sent from China ( Perkins, 1912). This species was considered extinct in the Hawaii ( Olmi, 1999a). However, in 2010 one of the authors (Olmi) saw a male specimen collected in Oahu Island, Mt. Olympus, in April 1981 by a pan trap. Probably so this species is always present at least in Oahu Island, although rare. Gonatopus pusanus and G. sogatae are here synonymized with G. nigricans because the only differences concern the colour of mesosoma (black in nigricans , testaceous to reddish, with some darkened areas, in pusanus ; brown, with some areas reddish or yellow, in sogatae ). In the collections, there are also many specimens with intermediate colours, so that the synonymy is evident. Gonatopus fulgori is here synonymized with G. nigricans , because the original description is like that of G. nigricans (the type material of G. fulgori is lost). In the past these two species were separated, because attributed to two different zoogeographic regions ( fulgori in Palaearctic, nigricans in Oriental and Australian regions).

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

7. Gonatopus nigricans (R. Perkins, 1905)

( Plates 111B , 114A – D )

Paragonatopus nigricans R. Perkins 1905: 41 ; Kieffer 1914b: 87; Olmi 1979: 218.

Pseudogonatopus melanacrias R. Perkins 1906a: 487 (synonymized by R. Perkins 1912); type locality: Suva ( Viti Levu Island,

Fiji Islands). Gonatopus fulgori Nakagawa 1906: 163 ; type locality: Japan; Guglielmino & Olmi 1997: 181, 184; syn. nov. Pseudogonatopus hospes R. Perkins 1912: 13 (synonymized by He & Xu 2002); type locality: China; Kieffer 1914b: 81;

Swezey 1928: 670; F. Williams 1931: 247; Chiu 1979: 339; Móczár 1979: 77; Olmi 1984: 1290; Olmi 2000a: 101, 111. Gonatopus sauteri Strand 1913: 209 (synonymized by Olmi 1984); type locality: Tainan ( Taiwan, China). Dicondylus sauteri (Strand) : Kieffer 1914b: 77. Pseudogonatopus sogatea Rohwer 1920: 160 (printing-error; holotype labelled sogatae ); type locality: Pusa (Bihar, India);

Kurian 1954b: 417; Olmi 1984: 1254; syn. nov. Haplogonatopus fulgori (Nakagawa) : Esaki & Hashimoto 1930: 21. Pseudozonatopus (sic) hospes R. Perkins : Pagden 1934: 2. Paragonatopus fulgori (Nakagawa) : Mochizuki 1943: 99; Chen 1988: 44. nec Pseudogonatopus nr. hospes R. Perkins : Chandra 1978b: 5. Pseudogonatopus fulgori (Nakagawa) : Olmi 1984: 1194. Pseudogonatopus pusanus Olmi 1984: 1255 ; type locality: Pusa (Bihar, India); He & Xu 2002: 355; syn. nov. Pseudogonatopus nigricans (R. Perkins) : Olmi 1984: 1287; Olmi 1990b: 789; Olmi 1991: 319. Gonatopus nigricans (R. Perkins) : Guglielmino & Olmi 1997: 165, 224–226, 250–252; Olmi 1999b: 28; Olmi 2000a: 94; He &

Xu 2002: 364; Guglielmino & Olmi 2006: 49. Gonatopus sogatae Rohwer : Guglielmino & Olmi 1997: 225, 226; He & Xu 2002: 355. Gonatopus insulanus He & Xu in He et al. 1998: 172; type locality: Qionghai (Hainan, China); syn. nov.

Description of female ( Plate 114A ): apterous; length 2.4–4.4 mm. Head brown-reddish, or brown, or black, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 or 1–3 yellow; mesosoma black or brown; occasionally posterior margin of pronotum and posterior apex of propodeum testaceous; metasoma black or brown; legs yellow; occasionally clubs of femora and part of coxae brown. In G. pusanus the colour is different: head partly or almost totally brown; antenna brown, except segments 1–2 testaceous; mesosoma testaceous-reddish, except sides of scutum darkened; occasionally part of pronotum darkened; metasoma brown; petiole black; legs testaceous; occasionally club of profemur darkened. In G. sogatae the colour is the following: head brown, except mandible and clypeus yellow; antenna brown, except segment 1 yellow and 2–3 brown-testaceous; mesosoma brown, except scutum reddish and distal apex of propodeum yellow; petiole black; metasoma brown-dark; legs brown-yellow. Intermediate colours are known. Antenna clavate; antennal segments in following proportions: 19:19:52:25:22:22:18:17:15:26. Head ( Plate 114B ) excavated, shiny, granulated or unsculptured; occiput and anterior region of face always granulated; POL = 5; OL = 5; OOL = 14. Palpal formula 2/2, 3/2, 4/2. Pronotum granulated, crossed by strong transverse impression. Scutum granulated, without lateral pointed apophyses. Mesopleura transversely striate. Metanotum flat, very long, about as long as scutum, transversely striate, not hollow behind scutellum ( Plate 114C ), with sides rounded ( Plate 114A ). Metathorax + propodeum shiny, indistinctly sculptured, with or without track of median longitudinal furrow, with posterior surface transversely striate. Metapleuron transversely striate. Meso-metapleural suture obsolete. Protarsal segment 4 shorter than segment 1 (40:47; or 20:27). Enlarged claw ( Plate 114D ) with one large subapical tooth and one row of three–nine lamellae. Segment 5 of protarsus ( Plate 114D ) with two rows of 9–25 lamellae; distal apex with 4–10 lamellae; occasionally lamellae extended continuously to distal apex. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9–2.4 mm. Head brown or black (occasionally brown-reddish), except mandible testaceous; occasionally part of face testaceous; antenna brown (in type of G. insulanus , testaceous-dark); mesosoma and metasoma black or brown (in type of G. insulanus , mesosoma reddish-dark, with propodeum and pleura darker); legs yellow. Antenna filiform; antennal segments in following proportions: 5:5:9:8:8:9:7:8:7:10 (in other specimens 6:5.5:10:9:8:9:8:8:7:10); antennal segment 3 more than three times as long as broad (4.0; 4.3; 5.0). Head shiny, hairy, granulated; frontal line absent; temple distinct; occipital carina absent; POL = 8; OL = 3; OOL = 3; greatest breadth of posterior ocelli longer than OL (4:3) (in other specimens POL = 7; OL = 3; OOL = 3.5; greatest breadth of posterior ocelli longer than OL (3.5:3)). Palpal formula 2/2, 3/2, 4/2. Scutum shiny, granulated. Notauli complete, posteriorly joint or almost joint in small specimens, separated in large specimens; in very large specimens, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2.5:4; or 2:4). Scutellum and metanotum shiny, finely punctate, unsculptured among punctures. Propodeum reticulate rugose; dorsal surface with median longitudinal furrow and occasionally with two lateral smooth areas. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 111B ) much shorter than paramere. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nigricans : AUSTRALIA: Queensland, Bundaberg, 19.XI.1904, reared ( BPBM); ♀ lectotype (designated by Olmi, 1982) of P. melanacrias : FIJI ISLANDS: Viti Levu Island, Suva, III.1905 , reared from delphacid near Stenocranus , on Zoysia pungens (BPBM) ; ♀ lectotype (designated by Olmi, 1982) of P. hospes : CHINA: no localities, F. Muir leg., reared in Honolulu from Perkinsiella saccharicida from Chinese stock, IV–V.1907 ( BPBM); ♀ holotype of G. sauteri : CHINA: Taiwan, Tainan, IV.1912, H. Sauter leg. ( DEI); ♀ holotype of P. pusanus : INDIA: Bihar, Pusa ( IARI); ♀ holotype of P. sogatae : INDIA: Bihar, Pusa, 6.VIII.1915, reared from Sogata sp. , C.S. Misra leg. ( USNM); ♂ holotype of G. insulanus : CHINA: Hainan, Qionghai, 4.V.1964, Junhua He leg. ( ZJUC). Type material of G. fulgori probably lost. Paratypes of P. nigricans : AUSTRALIA: same locality label as lectotype, 1♀ lebelled for mistake Fiji Islands ( BPBM); same locality label as holotype, parasitized host collected 1.X.1904, dryinid adult emerged 21.X.1904, R. Perkins leg., 1♀ “ paratype ” identified by the same R. Perkins ( BMNH); AUSTRALIA: Queensland, Bundaberg, Sandhills, X.1904, bred, 1♀ ( ANIC). Paratypes of P. melanacrias : same locality label as lectotype, 2♀♀ ( BPBM). Paratypes of P. hospes : same locality label as lectotype, 3♀♀ ( BPBM). Paratype of P. pusanus : same locality label as holotype, 1♀ ( IARI). Paratype of P. sogatae : same locality label as holotype, misidentified ♂ allotype not belonging to the same species of holotype. Other material. CHINA: Fujian, Shaxian, reared from rice planthoppers, 1979–1983, Changming Liu leg., 5♀♀ and 1♂ ( ZJUC); Fujian, IV.1987, 31.VII.1985, Changming Liu leg., 6♀♀ and 10♂ ( ZJUC); Guangdong, Yangjiang, 1979, 1981, Renhuan Hou leg., many ♀♀ and ♂ ( ZJUC); Guangdong, Xingfeng County, Mt. Yunjinshan, 9.VI.2002, 18.VII.2003, 19.VII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); same locality label, 20.VII.2003, Ping Li leg., 2♀♀ ( ZJUC); same locality label, Zaifu Xu leg., many ♀♀ ( ZJUC, AMNH); Guangdong, Shixing County, Chebaling National Nature Reserve, 21.VIII.2003, Zaifu Xu leg., 3♀♀ ( ZJUC); Guangdong, Foot of Nanling National Nature Reserve, Ruyang, M. Olmi reared from adult of unidentified Delphacid, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 14.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); same locality label, parasitized host collected 12.IX.2004, dryinid cocooning 15.IX.2004, dryinid adult emerged X.2004, 1♀ ( MOLC); Guangdong, Mt. Nankunshan, 2.VII.2005, Zaifu Xu leg., 1♀ ( ZJUC); Guangdong, Yangjiang, Hailing Town, 1980, Renhuan Hou leg., 2♀♀ and 2♂ ( ZJUC); Guangdong, Huaiji County, 21–22.X.2007, Zaifu Xu leg., 1♀ ( SCAU); Guangdong, Conghua, 28.VI.1978, Junhua He leg., 1♀ ( SCAU); Guangdong, Zhaoqing, V.1998, Yufang Liu leg., 2♀♀ ( SCAU); Guizhou, Sinan, IX.1986, VIII.1991, 5♀♀ ( ZJUC); Guizhou, Sandu, VI–VII.1981, Dongxiang Xie leg., 3♀♀ ( ZJUC); Hubei, Zhuxi, 1981, 1♀ ( ZJUC); Hunan, Yueyang, 15.VIII.1975, Binsen Wang leg., 1♀ ( ZJUC); Guangxi, Luchuan, 30.VII.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Bama, 1980, Tiansheng Pu leg., 1♀ ( ZJUC); Guangxi, Luocheng, 18.VI.1980, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Yaxian, VI.1966, Junhua He leg., 1♀ ( ZJUC); Hong Kong, Tai Lung Farm, IX–XI.1965, swept above rice, I.W.B. Thornton leg., 1♀ ( BMNH); Macau ( BPBM); same locality label, R. Perkins collection, 2♀♀ ( USNM); Taiwan, Kaohsiung, Kosen ( AMNH, ELKU); Taiwan, Changhus, Erlin ( TARI); Taiwan, Taipei ( TARI); Taiwan, Taipei, Kungkuan ( TARI); Taiwan, Taihoku Pref. (=Taipei), 27.IX.1930, J. Sonan leg., 1♀ ( TARI); Taiwan, Tainan, ShinShi, 11.IX.1979, paddy field, Hirashima leg., 1♀ ( ELKU); Taiwan, Gakko ( ELKU); Itahashi (= Taiwan), 16.IX.1930, J. Sonan leg., 5♀♀ ( TARI); Takeyama (=Jhushan), 6.X.1937, J. Sonan leg., 3♀♀ ( TARI); Inrin (=Yuanlin), 8.X.1937, J. Sonan leg., 39♀♀ ( TARI). INDIA: Chhattisgarh, Raipur ( GCC, AMNH); Delhi, New Delhi ( IARI); Orissa, Cuttack, 1981, 1♀ ( BMNH); Orissa, Puri, 28.XI.1987, S.V. Fowler leg., 1♀ ( BMNH); Tamil Nadu, Tanjavur (=Tanjore) District, 28.IX.1987, ex Nilaparvata lugens on rice, C. Peter leg., 3♀♀ ( BMNH); West Bengal, Hooghly District, Khanakul, 12.XII.1986, in paddy seedbed, 1♀ ( BMNH). INDONESIA: Java, Pekalongan ( BPBM); Java, Dieng Plateau ( BPBM); Borneo, Kalimantan, Paloh ( BMNH). MALAYSIA: Malaya, Selangor, 13.VIII.1928, H.T. Pagden leg., 1♀ ( BMNH); Malaya, Selangor, Kuala Lumpur, 21.XII.1930, H. T. Pagden leg., 1♀ ( BMNH); same locality label, many ♀♀ and ♂ ( NMNH, KYC); Malaya, Seberang Perai ( MARDI); Malaya, Serdang ( DBUS); Sabah, Tuaran ( KYC); Sarawak, 1st. Division, Payah Paloh, 28.I.1965, G.H.L. Rotschild leg., 1♀ ( BMNH). THAILAND: Songkhla, Hadd Tai ( KYC); Non Bua Lampoo, Na Kam Hai ( KYC); Tung Saew ( KYC); Tung Fabad ( KYC); Chainat ( KYC, AMNH); Banmakok ( KYC); Mae Tang ( KYC); Lampamg ( ELKU); Mae-Theng ( ELKU); Hot ( ELKU); Chieng Dao ( ELKU); Thailand, 15.IX.1999, reared from Nilaparvata lugens (Stål) , Sawai leg., 4♂ ( BMNH). Australian region: G. nigricans is known from Australia (Queensland), Fiji Islands, Solomon Islands and USA (Hawaii) ( Olmi, 1984). Introduced in the Hawaii 1906 and 1907 from China, Guangdong, for biological control of Perkinsiella saccharicida Kirkaldy in sugarcane fields ( Olmi, 2000a).

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in Australia: Liburnia sp. , Stenocranus sp. ; in China: Laodelphax striatellus (Fallén) ( Yang et al., 1982; He & Xu, 2002), Perkinsiella saccharicida Kirkaldy ; in Malaysia: Nilaparvata lugens (Stål) , Sogatella furcifera (Horváth) ; in Thailand: Nilaparvata lugens (Stål) ( Chiu, 1979) ; in the USA (Hawaii): Perkinsiella saccharicida Kirkaldy (introduced for biological control).

Biology. Information by Pagden (1934); the parthenogenesis is thelytokous.

Natural enemies. In the USA (Hawaii) ( Swezey, 1919; Timberlake, 1919; Muir, 1921; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Encyrtidae : Helegonatopus pseudophanes R. Perkins, 1906 ; Cheiloneurus flaccus (Walker 1847) ( Guerrieri & Viggiani, 2005; Guerrieri, 2006); Aphelinidae : Centrodora xiphidii (R. Perkins, 1906) . In Malaya ( Pagden, 1934; Olmi, 2000a), G. nigricans can be parasitized by the following parasitoids: Ceraphronidae : Allomicrops abnormis (R. Perkins, 1910) ; Encyrtidae : Cheiloneurus exitiosus (R. Perkins, 1906) ( Guerrieri & Viggiani, 2005) ; Pteromalidae : Eupteromalus sp.

Distribution. China ( Taiwan, Hainan, Fujian, Guangdong, Guizhou, Hubei, Hunan, Guangxi, Hong Kong, Macau), Australia (Queensland), Fiji Islands ( Viti Levu), India (Bihar, Chhattisgarh, Delhi, Orissa, Tamil Nadu), Indonesia (Java, Pekalongan, Borneo), Malaysia (Malaya, Sabah, Sarawak), Thailand (Songkhla, Non Bua Lampoo, Tung Saew, Tung Fabad, Chainat, Banmakok, Mae Tang, Lampamg, Mae-Theng, Chieng Dao).

Remarks. Gonatopus insulanus He & Xu is here synonymized with G. nigricans , because of the study of its type and comparison with male specimens of G. nigricans . There are no differences.

Gonatopus nigricans (= Pseudogonatopus hospes R. Perkins ) was established in the Hawaii from cocoons sent from China ( Perkins, 1912). This species was considered extinct in the Hawaii ( Olmi, 1999a). However, in 2010 one of the authors (Olmi) saw a male specimen collected in Oahu Island, Mt. Olympus, in April 1981 by a pan trap. Probably so this species is always present at least in Oahu Island, although rare. Gonatopus pusanus and G. sogatae are here synonymized with G. nigricans because the only differences concern the colour of mesosoma (black in nigricans , testaceous to reddish, with some darkened areas, in pusanus ; brown, with some areas reddish or yellow, in sogatae ). In the collections, there are also many specimens with intermediate colours, so that the synonymy is evident. Gonatopus fulgori is here synonymized with G. nigricans , because the original description is like that of G. nigricans (the type material of G. fulgori is lost). In the past these two species were separated, because attributed to two different zoogeographic regions ( fulgori in Palaearctic, nigricans in Oriental and Australian regions).

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

8. Gonatopus nudus (R. Perkins, 1912)

( Plates 115A, B , 116A, B , 117A )

Pseudogonatopus nudus R. Perkins 1912: 12 ; Chandra 1978a: 20; Chandra 1978b: 3; Chandra 1980a: 128; Chandra 1980b: 163; Olmi 1984: 1245; Kim et al., 1987: 64; Barrion 2002: 71; Manickavasagam et al. 2006: 24.

nec Pseudogonatopus nudas (sic) Perkins: Kim, 1990: 165.

Pseudogonatopus sarawaki Móczár 1979: 77 (synonymized by Olmi 1992b); type locality: Payah Paloh ( Sarawak, Malaysia).

Pseudogonatopus ponomarenkoi Móczár 1979: 80 (synonymized by Olmi 1984); type locality: Payah Paloh ( Sarawak, Malaysia); Ponomarenko 1981b: 880.

Pseudogonatopus sarawaki Móczár : Olmi 1984: 1238; Fiori 1984: 4.

Gonatopus camelinus Kieffer : Xu & He 1996b: 126.

Gonatopus nudus (R. Perkins) : Xu & He 1996b: 127; Guglielmino & Olmi 1997: 224–231; Olmi 1998f: 164; Olmi 1999b: 28; He & Xu 2002: 354; Guglielmino & Olmi 2006: 49, 51; Xu et al. 2011g (Suppl.): 21.

Gonatopus australis Xu & He in Xu et al., 1997a: 153 (synonymized by Olmi 1998f); type locality: Wenzhou (Zhejiang, China); Olmi 1998f: 164.

Gonatopus oryzaetorae Xu & He in Xu et al., 1997a: 154 (synonymized by Olmi 1998f); type locality: Debao (Guangxi, China); Olmi 1998f: 164.

Gonatopus yangi He & Xu 1998: 172; type locality: Yacheng (Hainan, China); He & Xu 2002: 369; syn. nov.

Description of female ( Plate 115A ): apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, or 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, with sides rounded ( Plate 115A ). Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Fore tarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 115B ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 115B ) with two rows of 14–20 lamellae; distal apex with about three–eight lamellae. Tibial spurs 1/0/1.

Description of male ( Plate 116A, B ). Fully winged; length 2.2–2.6 mm. Head black, except mandible testaceous; antenna brown; mesosoma black; metasoma brown; legs yellow. In holotype of G. yangi , head and mesosoma brown-reddish. Antenna filiform; antennal segments in following proportions: 5:5:11:8.5:8:9:8:8.5:8:11; antennal segment 3 more than four times as long as broad (11:2). Head dull, granulated; frontal line absent; occipital carina absent; POL = 7; OL = 2; OOL = 2; ocelli very broad; greatest breadth of posterior ocelli longer than OOL (4:2). Palpal formula 3/2 or 4/2. Scutum dull, granulated. Notauli complete, posteriorly joint. Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without transverse or longitudinal keels. Fore wing hyaline, without dark transverse bands; distal part of stigmal vein longer than proximal part; stigmal vein regularly curved. Dorsal process of paramere ( Plate 117A ) very reduced, reduced to a small membranous inner band. Tibial spurs 1/1/2.

Material examined. Types: ♀ lectotype (designated by Olmi, 1982) of P. nudus : INDONESIA: Java, Pekalongan ( BPBM); ♀ holotype of P. sarawaki : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of P. ponomarenkoi : MALAYSIA: Sarawak, Payah Paloh ( NMNH); ♀ holotype of G. australis : CHINA: Zhejiang, Wenzhou, VI.1964, Junhua He leg. ( ZJUC); ♀ holotype of G. oryzaetorae : CHINA, Guangxi, Debao, 10.IX.1980, Jian Lu leg. ( ZJUC); ♂ holotype of G. yangi : CHINA: Hainan, Yacheng, 23.V.1966, Junhua He leg. ( ZJUC). Paratype of P. ponomarenkoi : same locality label as holotype, 1♀ ( NMNH). Paratypes of G. australis : CHINA: same locality label as holotype, 10♀♀ (nine in ZJUC, one in AMNH). Paratype of G. oryzaetorae : CHINA, Guangxi, Qinzhou, VII.1981, Shaolong Yang leg. ( ZJUC). Other material. CHINA: Fujian, Mt. Huanggangshan ( ZJUC); Fujian, Shaxian, reared from rice planthoppers, 24.V–2.VII.1982, Changming Liu leg., 3♂ ( ZJUC); Fujian, Ninghua, 21.VIII.1978, Guiyu Luo leg., 1♂ ( ZJUC); Jiangxi, Nanchang, 1978, Zhengxiang Ye leg. ( ZJUC); Guangdong, Mt. Nankunshan, M. Olmi reared from an adult of an unidentified Delphacid, parasitized host collected 3.VII.2005, dryinid cocooning 4.VII.2005, 1♀ ( MOLC); Guizhou, Sandu, VII.1981, ex Delphacid, Dongxiang Xie leg., 1♀ ( ZJUC); Guizhou, Sinan, IX.1986, ex Delphacid, Yuxiang Chen leg., 1♀ ( ZJUC); Hainan, Yaxian, 1966, Junhua He leg., 2♀♀ ( ZJUC); Guangxi, Wuming, VII.1982, Shaolong Yang leg., 1♀ ( ZJUC); Guangxi, Guixian, 7.XI.1979, Tiansheng Pu leg., 1♀ ( ZJUC); Hainan, Haikou, 24.V.1983, Z. Bouček leg., 1♀ ( BMNH); Taiwan, Taichung, Wanfeng ( TARI); Taiwan, Taipei, V.1972, K.S. Lin leg., 1♀ ( TARI); Taiwan, Kosen ( AMNH); Yunnan, Kunming, 6.X.1980, Luzhe Wang leg., 1♀ ( ZJUC). INDIA: Andhra Pradesh, Hyderabad, 20.III.1980, 20.X.1977, reared from Nilaparvata lugens (Stål) , 3♀♀ ( BMNH); Gujarat, Disa (=Deesa) ( CAS); Madhya Pradesh, Raipur ( GCC); Orissa, Cuttack, 1981, reared from Nilaparvata lugens (Stål) , 6♀♀ ( BMNH); Tamil Nadu, Chennai (=Madras) ( ZSM); Tamil Nadu, Tanjore ( GCC). INDONESIA: Java, Bogor, 20.XI.1975, G. van Vreden reared from Nilaparvata lugens (Stål) , 4♂ ( BMNH). MALAYSIA: Malaya, Selangor, Serdang, 1.II.1980, reared from Nilaparvata lugens (Stål) , 2♀♀ and 1♂ ( BMNH); same locality label, many ♀♀ ( DBUS); Malaya, Seberang Perai ( MARDI). PHILIPPINES: Luzon Island, Bataan Prov., Olongapo, Subic Base, Teich b. Gate, 3.XII.1993, H. Zettel leg., 1♀ ( NHMW); Luzon Island, Laguna Prov., 4 Km E Los Baños, 14°10'N 121°15'E, 20.IX.1977, ex Nilaparvata lugens, G. Chandra leg., 1♂ ( BMNH). SRI LANKA: North Western Prov., Kalugomuwa ( AMNH); Kandy District, Gannoruwa ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM); Kandy District, Tismada ( Ponomarenko , 1981b: as Pseudogonatopus ponomarenkoi Móczár ) ( USNM). THAILAND: Bangkok ( BMNH); Sawai, 22.IX.1999, reared from Nilaparvata lugens (Stål) , 1♀ and 1♂ ( BMNH). Palaearctic: according to Kim et al. (1987) G. nudus is present in South Korea. However, the authors never saw specimens of this species collected in South Korea. The quotation of G. nudas in South Korea by Kim (1990) is erroneous, because it is referred to Haplogonatopus sp. (figure 1 in fact shows a drawing of female clearly related to Haplogonatopus , because the pronotum is not crossed by a transverse furrow.

Hosts. Delphacidae ( Guglielmino & Olmi, 1997, 2006): in China (Zhejiang): Nilaparvata bakeri (Muir) (Xu et al., 1997a) ; in China (Taiwan): Sogatella furcifera (Horváth) ( Guglielmino & Olmi, 1997) ; in India: Nilaparvata lugens (Stål) ( Guglielmino & Olmi, 1997) ; in Indonesia: Dicranotropis sp. (R. Perkins, 1912); Nilaparvata lugens (Stål) ; in Malaysia: Nilaparvata lugens (Stål) ( Móczár, 1979; Fiori, 1984); Sogatella sp. ( Móczár, 1979); in the Philippines: Nilaparvata lugens (Stål) ( Chandra, 1978a) ; Sogatella furcifera (Horváth) ( Chandra, 1978a; Barrion, 2002); Toya attenuata Distant ( Olmi, 1984) ; Stenocranus sp. ( Chandra, 1978a); in South Korea: Nilaparvata lugens (Stål)( Kim et al., 1987) ; in Sri Lanka: Nilaparvata lugens (Stål) ( Olmi, 1984) ; Sogatella sp. ( Olmi, 1984); in Thailand: Nilaparvata lugens (Stål) ( Fiori, 1984) .

According to Chandra (1978a, 1978b, 1980a, 1980b) in the Philippines G. nudus parasitizes also Nephotettix nigropictus (Stål) , Nephotettix virescens (Distant) and Recilia dorsalis (Motschulsky) ( Cicadellidae ). These data, however, are surely erroneous, because G. nudus is not able to parasitize Cicadellidae .

Biology. According to Dr. Mohd Yusuf Omar (pers. comm.) in Malaya G. nudus has an arrhenotokous parthenogenesis.

Natural enemies. In India (Tamil Nadu), G. nudus is parasitized by Cheiloneurus exitiosus (R. Perkins) ( Encyrtidae ) ( Manickavasagam et al., 2006). According to Manickavasagam et al. (2006) the female of Ch. exitiosus lays its eggs in the dryinid larva, when it is in the cyst on the host body. The encyrtid female inserted its ovipositor in four different sites of the dryinid cyst. One day after parasitization, the matured dryinid larva came out of the sac and pupated in a cocoon. From a single cocoon, four encyrtid adults emerged. The total life cycle of the encyrtid took about 14 days, from egg laying through the dryinid sac until adult emergence.

Distribution. China (Zhejiang, Guangxi, Hainan, Fujian, Jiangxi, Guizhou, Guangdong, Taiwan, Yunnan), Indonesia (Java), Malaysia ( Sarawak, Malaya), India (Andhra Pradesh, Gujarat, Madhya Pradesh, Orissa, Tamil Nadu), Philippines (Luzon), Sri Lanka (Kalugomuwa, Kandy), Thailand (Bangkok).

Remarks. Gonatopus yangi He & Xu is here synonymized with G. nudus after study of its holotype nd comparison with males of G. nudus . No differences were observed, excluding the colour (black in G. nudus , brownreddish in G. yangi ). However, this difference of colour is common in males of other species, so that it is not considered important here. The genitalia are perfectly like.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

9. Gonatopus sarawakensis ( Olmi, 1984) , comb. nov.

( Plate 117D )

Pseudogonatopus sarawakensis Olmi 1984: 1241 .

Description of Female. Apterous; length 2.6–4.2 mm. Head brown-reddish, except mandible, clypeus and anterior region of face yellow; occasionally head mostly dark or black; antenna brown, except segments 1–2 yellow; mesosoma testaceous-yellow; occasionally scutum yellow; occasionally scutellum and lateral regions of metanotum darkened or blackish; occasionally mesopleuron and metapleuron darkened; petiole black; metasoma black, with reddish nuances; legs testaceous. Antenna clavate, with rhinaria on segments 9–10; antennal segments in following proportions: 10:7.5:27:11:10:10:9:8:8:12. Head excavated, shiny, smooth, weakly alutaceous; POL = 7; OL = 4; OOL = 12. Palpal formula 2/2, 3/2, 4/2. Pronotum crossed by strong transverse impression, shiny, weakly granulated. Metanotum shiny, transversely striate, laterally protruding; lateral protrusions rounded. Mesopleuron transversely striate. Metathorax + propodeum shiny, smooth, unsculptured, with numerous transverse striae on posterior surface and metapleuron; occasionally disc of metathorax + propodeum with slight track of median longitudinal furrow. Meso-metapleural suture obsolete. Protarsal segments in following proportions: 15:3:4:15:23. Enlarged claw ( Plate 117D ) with one large subapical tooth and one row of five–seven lamellae. Segment 5 of protarsus ( Plate 117D ) with two rows of 14–20 lamellae; distal apex with 3–8 lamellae. Tibial spurs 1/0/1.

Male. See Remarks.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , 17.IX–23.X.1977, D. Hollis leg. ( BMNH) . Paratype: same locality label as holotype, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak).

Remarks. In BMNH there is a series of Gonatopus males with palpal formula 3/2 or 4/2 collected in Malaysia, Sarawak, Gunung Mulu National Park. They may be the opposite sex of G. sarawakensis (Olmi) or G. malesiae (Olmi) , whose females were collected in the same site by the same collectors. The above males were divided by Olmi (1984) into two groups (A and B), according to the following characters: A: distal apex of paramere rounded; notauli posteriorly joint; B: distal apex of paramere pointed; notauli posteriorly separated. With no rearings and/or DNA studies, it is impossible to attribute the above males to one of the above two species, G. sarawakensis and G. malesiae . The descriptions of the above males is the following: fully winged; length 1.4–2.5 mm. Head black or brown; antenna brown or partly testaceous; mesosoma and metasoma black or brown; legs yellow. Antenna filiform; antennal segments of male A in following proportions: 5:5:9:7:7:7:8:7:6:10; antennal segments of male B in following proportions: 5:6:9:9:8:8:8:8:8:11; in both males, antennal segment 3 more than three times as long as broad (male A: 4.0; male B: 3.66). Head dull, hairy, rugose; occipital carina absent; temple short, distinct; POL = 5; OL = 2; OOL = 2.5; greatest breadth of posterior ocelli longer than OL (3:2). Palpal formula: 3/2, 4/2. Scutum dull, hairy, punctate. Notauli complete, posteriorly joint (male A) or separated (male B); in male B, minimum distance between notauli shorter than greatest breadth of posterior ocelli (2:3). Scutellum and metanotum shiny, smooth, hairy, punctate, unsculptured among punctures. Propodeum reticulate rugose. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 117B, C ) long and broad, with distal apex rounded (male A; plate 117B) or pointed (male B; Plate 117C ). Tibial spurs 1/1/2.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

10. Gonatopus validus ( Olmi, 1984) , comb. nov.

( Plate 117E – H )

Pseudogonatopus validus Olmi 1984: 1248 .

Description of Female. Apterous; length 2.5 mm. Holotype with head testaceous-reddish or brown-reddish, except mandible, clypeus and anterior region of face yellow; antenna brown, except segments 1–2 yellow; mesosoma reddish, or testaceous, or brown-reddish; petiole black; metasoma brown-reddish; legs yellow. In specimen from China, Mt. Yunjishan, body brown-testaceous, with antenna brown, except segments 1–2 testaceous and segment 10 whitish. In specimen from China, Fengxi, antenna testaceous-darkened, except segments 1–2 testaceous and segment 10 whitish. Antenna clavate; antennal segments of holotype in following proportions: 17:14:27:14:13:11:11:11:11:21. Antennal segments of specimen from China, Fengxi, in following proportions: 8:5:20:6.5:5:4.5:5:5:5:9. Head ( Plate 117E ) excavated, shiny, smooth, unsculptured. Palpal formula of holotype 4/ 2. In specimens from China, Fengxi, and China, Mt. Nankunshan, palpal formula 5/2. Pronotum ( Plate 117F ) crossed by strong transverse impression, shiny, smooth and unsculptured. Scutum and scutellum shiny, smooth, unsculptured. Mesopleuron transversely striate. Metanotum not transversely striate, with sides protruding; lateral protrusions of metanotum rounded ( Plate 117G ). Metathorax + propodeum smooth, shiny, unsculptured; posterior surface of propodeum and metapleuron transversely striate. Meso-metapleural suture only posteriorly distinct. Segment 1 of protarsus of holotype about as long as segment 4. In specimen from China, Mt. Yunjishan, segment 1 of protarsus slightly longer than segment 4 (13:12). Enlarged claw ( Plate 117H ) with one large subapical tooth and one row of five lamellae. Segment 5 of protarsus ( Plate 117H ) with two rows of 13 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: holotype ♀: MALAYSIA: Malaya, Kuala Lumpur, Spooner Road , 8.III.1936, H.T. Pagden leg. ( BMNH) . Paratype: SRI LANKA: Gampaha District , Yakkala , 1♀ ( AMNH) . Other material. CHINA: Guangdong, Mt. Yunjishan, M. Olmi reared from an unidentified Delphacid , parasitized host collected 9.VI.2002, dryinid cocooning 12.VI.2002, dryinid adult emerged 29.VI.2002 ( MOLC) ; Guangdong, Shixing County, Chebaling National Nature Reserve , 21–31.VIII.2003, Jujian Chen leg., 1♀ ( ZJUC); Guangdong, Meizhou, Fengxi , 30.VII.2003, Jingxian Liu leg., 1♀ ( ZJUC); Guangdong, Mt. Nankunshan , 27–29.IX.2011, YPT, Zaifu Xu leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Guangdong), Malaysia (Malaya), Sri Lanka (Gampaha).

Group 3

Five species are known.

Oriental Gonatopus group 3: key to the females

1. Disc of metathorax + propodeum sculptured by strong transverse keels.......................... G. bengalensis (Olmi)

- Disc of metathorax + propodeum not sculptured by strong transverse keels; transverse keels present only on posterior surface of propodeum; occasionally short keels are present on ground of median furrow................................... 2

2. Vertex of head and metathorax + propodeum very shiny, smooth, unsculptured, except transverse striae on posterior surface of propodeum, metapleuron and occasionally mesopleuron...................................................... 3

- Vertex of head and metathorax + propodeum dull, granulated.................................................. 4

3. Median furrow of metathorax + propodeum less deep ( Plate 118B ); lateral regions of scutum with one or two transverse keels ( Plate 118A )......................................................................... G. borneanus (Olmi)

- Median furrow of metathorax + propodeum deeper ( Plate 119A ); lateral regions of scutum with four transverse keels ( Plate 119C ).............................................................................. G. pajanensis (Olmi)

4. Antenna short, more thickened distally and with segment 9 slightly longer than broad................. G. insularis (Olmi)

- Antenna long, less thickened distally and with segment 9 about three times as long as broad............ G. pyrillae (Mani)

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

1. Gonatopus bengalensis ( Olmi, 1984) , comb. nov.

( Plate 107C )

Agonatopoides bengalensis Olmi 1984: 1333 .

Description of Female. Apterous; length 3.5–5.3 mm. Head testaceous; mesosoma brown-reddish; petiole black; metasoma and legs testaceous. Antenna of holotype broken, with only segment 1. Head excavated, shiny, hairless, unsculptured; frontal line complete; occipital carina absent; POL = 3; OL = 5; OOL = 21. Palpal formula 5/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression; lateral regions with few longitudinal keels. Scutum shiny, smooth, with few longitudinal keels, laterally with two rounded apophyses. Scutellum shiny, smooth, rectangular. Metanotum transversely striate, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum shiny, with deep median furrow; disc and posterior surface of metathorax + propodeum transversely striate. Meso-metapleural suture distinct. Mesopleuron and metapleuron not transversely striate. Protarsal segments in following proportions: 16:4:5:20:33. Enlarged claw ( Plate 107C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 107C ) with one row of 13 lamellae; distal apex with about 20 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDIA: Assam, Mishmi Hills, Delai Valley, Chanliang , 6500 ft., 2.XII.1936, M. Steele leg. ( BMNH) .

Hosts. Unknown.

Distribution. India (Assam).

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

2. Gonatopus borneanus ( Olmi, 1984) , comb. nov.

( Plate 118A–D )

Agonatopoides borneanus Olmi 1984: 1333 ; Olmi 1989b: 159.

Description of Female. Apterous; length 3.5–3.6 mm. Completely brown-testaceous. Occasionally totally testaceous, except ocellar triangle brown, petiole black and metasoma partly brown. Antenna weakly clavate; antennal segments in following proportions: 12:7:22:12:11:9:7:7:7:11. Head very excavated, shiny, smooth, unsculptured; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 10. Palpal formula 5/3. Pronotum hairless, unsculptured, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured; sides of scutum with two lateral points (corresponding to one lateral transverse keel). Scutellum shiny, smooth, unsculptured. Metanotum smooth, shiny, unsculptured, hollow behind scutellum ( Plate 118A ). Metathorax + propodeum with anterior surface unsculptured and posterior surface transversely striate; disc with deep median longitudinal furrow ( Plate 118B ). Meso-metapleural suture distinct and complete. Mesopleuron and metapleuron transversely striate. Protarsal segments in following proportions: 19:5:8:22:33. Enlarged claw ( Plate 118C ) with one large subdistal tooth and one row of five–eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118C ) with two rows of about 11 + 2 (proximal) lamellae (occasionally only one row of 11–12 lamellae); distal apex with about 18 lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 1.9 mm. Head and antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 6:6:12:8:8:8:8:8:7:10; antennal segment 3 more than four times as long as broad (about 4.8). Head shiny, hairy, alutaceous, except face weakly granulated; frontal line absent; occipital carina absent; temple very short, reduced; POL = 6; OL = 2.5; OOL = 2; greatest breadth of posterior ocelli slightly longer than OL (3:2.5). Palpal formula 5/3. Scutum shiny, hairy, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (2.5:3). Scutellum and metanotum shiny, smooth, unsculptured. Propodeum reticulate rugose; posterior surface without longitudinal keels. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 118D ) long, broad and distally pointed. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: MALAYSIA: Sarawak, Gunung Mulu , IV.1978, N.M. Collins leg. ( BMNH) . Paratype: same locality label as holotype, 1♂ ( BMNH) . Other material. MALAYSIA: Sabah, Danum Valley Field Camp , about 140 m ( RNHL) . SRI LANKA: Sabaragamuwa Prov., Kegalle District, Maha-Oya , 24.VIII.1963, on ground, R. Winney leg., University London Ceylon Expedition, 1♀ ( BMNH) .

Hosts. Unknown.

Distribution. Malaysia ( Sarawak, Sabah), Sri Lanka (Sabaragamuwa).

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

3. Gonatopus insularis ( Olmi, 1984) , comb. nov.

( Plate 118E )

Agonatopoides insularis Olmi 1984: 1334 .

Description of Female. Apterous; length 3.2 mm. Testaceous, except antennal segments 4–9 brown and petiole black. Antenna clavate; antennal segments in following proportions: 10:7:18:10:9:8:8:6:5.5:10; antennal segment 9 slightly longer than broad (5.5:4). Head very excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 12. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, smooth, unsculptured. Metanotum weakly transversely striate, hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum granulated; posterior surface transversely striate. Protarsal segments in following proportions: 16:4:6:19:30. Enlarged claw ( Plate 118E ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 118E ) with two rows of 12 + 3 (proximal) lamellae; distal apex with about 12 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: SRI LANKA: Badulla District , 15 mi. NNE of Mahiyangana, Ulhitiya Oya ( USNM) .

Hosts. Unknown.

Distribution. Sri Lanka (Badulla).

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

4. Gonatopus pajanensis (Olmi, 1989) , comb. nov.

( Plate 119A–C )

Agonatopoides pajanensis Olmi 1989b: 157 .

Description of Female. Apterous; length 6.6 mm. Testaceous-dark, except petiole black and tibiae darkened. Antenna clavate; antennal segments in following proportions: 14:7:37:19:17:14:10:9:9:13. Head excavated, shiny, smooth, alutaceous; ocellar triangle rugose; frontal line complete; occipital carina incomplete, only shortly present on sides of posterior ocelli; POL = 1; OL = 3; OOL = 11. Palpal formula 5/3. Pronotum crossed by strong transverse impression, with disc dull and alutaceous and anterior collar shiny and unsculptured. Scutum dull, irregularly striate; sides of scutum with four strong transverse keels. Scutellum shiny, smooth, unsculptured. Metanotum dull, irregularly striate, hollow behind scutellum ( Plate 119A ). Metathorax + propodeum with posterior surface transversely striate (striae also present on disc on sides of median longitudinal furrow); disc with deep median longitudinal furrow ( Plate 119B ). Meso-metapleural suture distinct and complete. Mesopleura and metapleura transversely striate. Protarsal segments in following proportions: 22:4:9:29:45. Enlarged claw ( Plate 119C ) with one large subdistal tooth and one row of eight lamellae + one bristle. Segment 5 of protarsus ( Plate 119C ) with two rows of about 32 lamellae; distal apex with about 25 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Borneo , Pajan River, XII.1925, Eric Mjöberg leg. ( MCZ) .

Hosts. Unknown.

Distribution. Indonesia (Borneo).

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

5. Gonatopus pyrillae ( Mani, 1942) , comb. nov.

( Plate 119D, E )

Pseudogonatopus pyrillae Mani 1942: 161 ; Kurian 1954a: 258.

Agonatopoides pyrillae (Mani) : Olmi 1984: 1336.

Description of Female. Apterous; length 4.2 mm. Testaceous-reddish, except metasoma black. Antenna clavate; antennal segments in following proportions: 11:8:30:20:15:13:10:10:9:11; antennal segment 3 about three times as long as broad (9:3). Head excavated, dull, granulated; frontal line complete; occipital carina absent; POL = 1; OL = 3; OOL = 14. Palpal formula 5/3. Pronotum shiny, weakly granulated, crossed by strong transverse impression. Scutum shiny, weakly granulated. Scutellum shiny, indistinct. Metanotum hollow behind scutellum, with sides protruding; lateral protrusions pointed. Metathorax + propodeum with anterior surface dull, granulated and posterior surface transversely striate; disc rugose, with deep median longitudinal furrow. Meso-metapleural suture distinct and complete. Mesopleuron not transversely striate. Metapleuron transversely striate. Protarsal segments in following proportions: 22:5:7:25:40. Segment 3 of protarsus produced into hook. Enlarged claw ( Plate 119E ) with one large subdistal tooth and one row of 11 lamellae + one bristle. Segment 5 of protarsus ( Plate 119E ) with one row of 15 lamellae; distal apex with about nine lamellae. Tibial spurs 1/0/1.

Description of Male. Fully winged; length 2.4 mm. Head brown, except mandible testaceous; antenna testaceous; mesosoma and metasoma brown; legs testaceous. Antenna filiform; antennal segments in following proportions: 7:6:8:7:7:6:6:6:6:9; antennal segment 3 less than three times as long as broad (8:3). Head dull, granulated; frontal line absent; occipital carina absent; temple distinct; POL = 8; OL = 4; OOL = 4; greatest breadth of posterior ocelli shorter than OL (3:4). Palpal formula 5/3. Scutum shiny, granulated. Notauli complete, posteriorly separated; minimum distance between notauli slightly shorter than greatest breadth of posterior ocelli (1.5:3). Scutellum and metanotum shiny, smooth, punctate, unsculptured among punctures. Propodeum dull, reticulate rugose, without longitudinal or transverse keels; dorsal surface with median narrow longitudinal furrow. Fore wing hyaline, without dark transverse bands. Dorsal process of paramere ( Plate 119D ) long, distally broad. Tibial spurs 1/1/2.

Material examined. Types: holotype ♀: INDIA: Punjab, Karnal , 7.X.1941, reared from Pyrilla sp. , Lakshmanan leg. ( IARI) . Paratype: INDIA: Uttar Pradesh, Meerut, Daurala Farm , 23.X.1938, reared from Pyrilla sp. , Lakshmanan leg., ♂ allotype ( IARI). According to Mani (1942) there are three female paratypes.

Hosts. Lophopidae ( Guglielmino & Olmi, 1997) : in India: Pyrilla sp. ( Mani, 1942).

Distribution. India (Punjab, Uttar Pradesh).

Group 4

Four species are known.

Oriental Gonatopus group 4: key to the females

1. Antenna very long, slender, with segment 4 approximately four times as long as segment 2................. G.asiae Olmi

- Antenna short, with segment 4 less than three times as long as segment 2......................................... 2

2. Meso-metapleural suture obsolete; metanotum not hollow behind scutellum........................... G. insulae Olmi

- Meso-metapleural suture distinct and complete; metanotum slightly hollow behind scutellum......................... 3

3. Anterior surface of metathorax + propodeum sculptured by many longitudinal striae....................... G.lini Olmi

- Anterior surface of metathorax + propodeum smooth, unsculptured, not sculptured by many longitudinal striae................................................................................... G.hagoromo Terayama & Ohbayashi

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

1. Gonatopus asiae Olmi, 1993

( Plate 120A )

nec Donisthorpina asiatica Olmi 1984: 1309 .

Apterodryinus asiaticus Olmi 1984: 1350 (preoccupied).

Gonatopus asiae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 6.0 mm. Yellow-ferruginous, except petiole black. Antenna clavate; antennal segments in following proportions: 9:8:40:32:25:17:11:8:7:10. Head excavated, shiny, unsculptured, except occiput and anterior region of face granulated; frontal line incomplete, absent near clypeus; occipital carina incomplete, shortly present on sides of posterior ocelli; POL = 6; OL = 6; OOL = 27. Palpal formula 6/3. Pronotum shiny, hairless, granulated, crossed by strong transverse impression. Scutum granulated, with two lateral pointed protrusions. Scutellum shiny, distinct. Metanotum transversely striate, not hollow behind scutellum, with sides not protruding. Metathorax + propodeum shiny, with disc, anterior and posterior surface transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture distinct and complete, very thin. Protarsal segments in following proportions: 57:8:17:75:104. Enlarged claw ( Plate 120A ) with one large subdistal tooth and one row of eight lamellae. Segment 5 of protarsus ( Plate 120A ) with two rows of 19 + 22 lamellae; distal apex with at least 30 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Type: holotype ♀: INDONESIA: Sulawesi, Tondano , VI.1912, Wallace leg. ( BMNH) .

Hosts. Unknown.

Distribution. Indonesia (Sulawesi).

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

2. Gonatopus hagoromo Terayama & Ohbayashi, 2003

( Plate 120B )

Gonatopus hagoromo Terayama & Ohbayashi 2003: 237 ; Guglielmino & Olmi 2006: 39.

Description of Female. Apterous; length 2.6–4.2 mm. Head brown, except anterior region of clypeus and mandible testaceous; antenna brown, except segments 1–2 and 8–10 whitish; mesosoma brown-testaceous; metasoma brown; forelegs brown, except chela testaceous, trochanter and part of coxa withish; mid- and hindlegs with tarsi and part of tibiae testaceous, part of coxae, trochanters and part of clubs of femora whitish. Antenna clavate; antennal segments of holotype in following proportions: 10:7:18.5:12:9.5:8:6:6:5:7.5. Head excavated, shiny, smooth, unsculptured in holotype. Head with frontal line complete, occipital carina absent; POL = 1.5; OL = 1.5; OOL = 11. Palpal formula 6/3. Pronotum shiny, smooth, unsculptured, crossed by strong transverse impression. Scutum dull, sculptured by longitudinal keels, without lateral pointes apophyses. Metanotum smooth, shiny, weakly hollow behind scutellum, with sides protruding; lateral protrusions rounded. Metathorax + propodeum of holotype with anterior surface shiny, smooth, unsculptured; posterior surface strongly transversely striate. Mesopleuron and metapleuron almost completely smooth, unsculptured, partly transversely striate. Mesometapleural suture distinct and complete. Protarsal segments of holotype in following proportions: 18:4:6:21:34. Enlarged claw ( Plate 120B ) with one large subdistal tooth and one row of 8–11 lamellae + one hair. Segment 5 of protarsus ( Plate 120B ) with two rows of 14–22 lamellae; distal apex with about 24 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Types: female holtoype: JAPAN: Ogasawara Islands (= Bonin Islands ), Chichi-jima Island, Komagari, 3.VII.1997, T. Ohbayashi leg. ( NIAES) . Paratypes: JAPAN: Ogasawara Islands (= Bonin Islands), same locality label as holotype, 10.VII.1997, 1♀ ( NIAES) ; Ogasawara Islands (= Bonin Islands ), Hahajima, Yake-yama, 20.IV.1997, K. Nishimoto leg., 1♀ ( NIAES) .

Hosts. Flatidae ( Guglielmino & Olmi, 2006) : in Japan (Ogasawara Islands): nymphs of Geisha distinctissima (Walker) ( Terayama & Ohbayashi, 2003) .

Distribution. Japan (Ogasawara).

Remarks. Gonatopus hagoromo was described by Terayama & Ohbayashi (2003) from the Ogasawara Islands, known also as Bonin Islands. This archipelago is part of Japan, but with a subtropical climate. Ogasawara Islands belong to the Oriental zoogeographical region ( Inoue & Miyatake, 2001).

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

3. Gonatopus insulae Olmi, 1993

( Plate 120C )

nec Agonatopoides insularis Olmi 1984: 1334 .

Apterodryinus insularis Olmi 1984: 1374 (preoccupied).

Gonatopus insulae Olmi 1993b: 80 (new name).

Description of Female. Apterous; length 2.5–3.0 mm. Testaceous-yellow, except petiole black and occasionally metasoma brownish. Antenna clavate. Antennal segments in following proportions: 8:5:11:6.5:5:4:5 (segments 8–10 missing in holotype). Head excavated, shiny, smooth, hairless, unsculptured, except occiput and anterior region of face granulated. Palpal formula 6/3. Pronotum shiny, unsculptured, crossed by strong transverse impression. Scutum shiny, unsculptured, without lateral pointed apophyses. Scutellum hardly visible. Metanotum not hollow behind scutellum. Metathorax + propodeum shiny, unsculptured, with posterior surface strongly transversely striate. Mesopleuron and metapleuron transversely striate. Meso-metapleural suture obsolete in holotype, with anterior third very slightly visible in specimen from China, Daheba. Segment 1 of protarsus of holotype as long as segment 4. Protarsal segments of specimen from China, Daheba, in following proportions: 15:3:5:18:27. Enlarged claw ( Plate 120C ) with one large subdistal tooth and one row of eight-ten lamellae. Segment 5 of protarsus ( Plate 120C ) with two rows of 15 lamellae; distal apex with at least 16 lamellae. Tibial spurs 1/0/1.

Male. Unknown.

Material examined. Australian region: INDONESIA (see remarks): Tanimbar Islands, Larat Island , holotype ♀ ( BPBM). Other material : CHINA: Gansu, Dangchang, Daheba , 2530 m, 31.VII.2004, Hongying Zhang leg., 1♀ ( SCAU) .

Hosts. Unknown.

Distribution. China (Gansu), Indonesia (Tanimbar).

Remarks. The Tanimbar Islands belong to the Australian zoogeographical region because they are located after the Wallace's line. However, G. insulae is here treated, because of the continuity with the other Indonesian islands. The Wallace's line in fact is only a conventional border.

Figures

PLATE 103. Haplogonatopus apicalis. A. Female from China, Guangdong. Length 2.5 mm. B. Chela of female from Sri Lanka (from Olmi 1984). Scale bar 0.16 mm for B.

PLATE 104. A. Dorsal view. B, C. Male genitalia. D. Chela. A. Haplogonatopus apicalis, Male. Length 2.0 mm. B. H. apicalis from Sri Lanka (from Olmi 1984). C. H. oratorius (=H. atratus) from Japan (from Olmi 1984). D. H. oratorius from Italy (from Olmi 1984). Scale bar 0.09 mm for B, 0.13 mm for C and 0.18 mm for D.

PLATE 105. A. Haplogonatopus oratorius, female from Italy, lateral view. Length 2.5 mm. B. H. oratorius, male from Italy, lateral view. Length 2.3 mm. (From Olmi 1984).

PLATE 106. A. Gonatopus longicollis, female from Spain. Length 4.8 mm (from Olmi 1999a). B. G. asiaticus, holotype. Length 3.0 mm (from Olmi 1984).

PLATE 107. Chelae, holotypes. A. Gonatopus paulyi (from Olmi 1986). B. G.abjectus (from Olmi 1984). C. G. bengalensis (from Olmi 1984). D. G. lucens (from Olmi 1984). Scale bar 0.15 mm for A, 0.14 mm for B, 0.21 mm for C and 0.20 mm for D.

PLATE 108. A. Gonatopus pedestris, female from Italy, lateral view. Length 3.0 mm. B. Mesosoma, lateral view of G. rufoniger (=G. indianus Olmi), from India. Length 2.0 mm. C. Female of G. bicolor, lateral view. Lenght 2.9 mm. D. G. acuminatus, holotype, lateral view. Length 3.12 mm. (From Olmi 1984)

PLATE 109. A. Gonatopus nearcticus (=Acrodontochelys bouceki Currado), female from Spain, Nerja (from Currado, 1976). Length 3.5 mm. B. Chela of G. nearcticus (=Acrodontochelys sinensis Olmi), female from China, Macau (from Olmi 1984). Scale bar 0.17 mm for B.

PLATE 110. Gonatopus nearcticus (=Acrodontochelys ugandanus (Benoit)), male from Uganda. Length 3.0 mm (from Olmi 1984). A. Dorsal view. B. Lateral view.

PLATE 111. A, B. Male genitalia. C. Scutum + metathorax + propodeum. D. Chela. A. Gonatopus nearcticus, from Greece. B. G. nigricans, from Malaya. Different shapes of dorsal process of paramere. C, D. G. bicuspis, holotype (from Olmi 1993d). Scale bar 0.14 mm for A and B, 0.40 mm for C and 0.12 mm for D.

PLATE 112. Gonatopus capillus, holotype (from Xu & He, 1998d). A. Dorsal view. Length 5.0 mm. B. Chela. Scale bar 0.26 mm for B.

PLATE 113. A, C, E. Scutum + metathorax + propodeum. G. Mesosoma. B, D, F, H. Chelae. A, B. Gonatopus lankae, holotype (from Olmi 1984). C, D. G.malesiae, female (from Olmi 1984). E, F. G. muiri, holotype (from Olmi 1984). G, H. G. nepalensis, holotype (from Olmi 1986). Scale bar 0.63 mm for A, 0.16 mm for B and D, 0.59 mm for C, 0.61 mm for E, 0.21 mm for F, 0.75 mm for G and 0.25 mm for H.

PLATE 114. Gonatopus nigricans, Female. A. From China, Guangdong. Length 2.5 mm. B. Head, from China, Macau (from Olmi 1984). C. Mesoma, lateral view, from China, Macau (from Olmi 1984). D. Chela, from China, Macau (from Olmi 1984). Scale bar 1.10 mm for B, 1.02 mm for C and 0.12 mm for D.

PLATE 115. Gonatopus nudus, Female. A. From China, Guangdong, Mt. Nankunshan. Length 3.0 mm. B. Chela of lectotype (from Olmi 1984). Scale bar 0.22 mm for B.

PLATE 116. Gonatopus nudus (=Pseudogonatopus sarawki Móczár), male from Sri Lanka. Length 2.2 mm (from Olmi 1984). A. Dorsal view. B. Lateral view.

PLATE 117. A–C. Male genitalia. D, H. Chelae, holotypse. E. Head. F, G. Mesosoma. A. Gonatopus nudus from Malaya. B. G. sarawakensis or malesiae, var. a. C. G. sarawakensis or malesiae, var. b. D. G. sarawakensis. E, F, H. G. validus. G. G. validus, female from China. Scale bar 0.24 mm for A, 0.27 mm for B, 0.29 mm for C, 0.16 mm for D, 0.12 mm for H. (From Olmi 1984).

PLATE 118. A. Scutum and metathorax + propodeum, lateral view (from Olmi 1989b). B. Furrow of metathorax + propodeum, frontal view (from Olmi 1989b). C, E. Chelae (from Olmi 1984). D. Male genitalia (from Olmi 1984). A–C. Gonatopus borneanus, holotype. D. G. borneanus, paratype. E. G. insularis, holotype. Scale bar 0.75 mm for A, 0.15 mm for C, 0.18 mm for D and 0.14 mm for E.

PLATE 119. A. Scutum and metathorax + propodeum, lateral view. B. Furrow of metathorax + propodeum, frontal view. C, E. Chelae, holotypes. D. Male genitalia. A–C. Gonatopus pajanensis (from Olmi 1989b). D, E. G. pyrillae (from Olmi 1984). Scale bar 0.29 mm for C, 0.19 mm for D and 0.16 mm for E.

PLATE 120. A–C, E. Chelae, holotypes. D. Scutum and metathorax + propodeum, holotype. A. Gonatopus asiae (=Apterodryinus asiaticus) (from Olmi 1984). B. G. hagoromo. C. G. insulae (=Apterodryinus insularis) (from Olmi 1984). D. G. lini (courtesy of Michael Sharkey). Length 1.15 mm. E. G. lini (from Olmi 1995a). Scale bar 0.29 mm for A, 0.22 mm for B, 0.12 mm for C and 0.19 mm for E.

PLATE 122. A. Male, lateral view. B–D. Male genitalia. E. Chela, holotype. A. Gonatopus flavifemur (=Dicondylus indianus Olmi) from India. Length 2.6 mm. B. G. flavifemur from Australia. C. G. lucens from Japan. D. G. pedestris from Finland. E. G. lucens. Scale bar 0.17 mm for B, 0.10 mm for C, 0.15 mm for D and 0.19 mm for E. (From Olmi 1984).

PLATE 126. A. Dorsal view. Length 7.5 mm. B, D, E. Heads. C. Mesosoma. F. Chela. A–C, F. Gonatopus cristatus (=Megagonatopus birmanus Olmi & Currado, 1976), female from Myanmar (from Olmi & Currado, 1976). D, E. Gonatopus daunus, holotype, frontal view (D) and dorsal view (E) (from Olmi 1984). Scale bar 2.27 mm for B, 1.55 mm for C, 0.94 mm for D and E, 0.36 mm for F.

PLATE 128. A, B. Male genitalia. C. Head, holotype. D, F, G. Chelae, holotypes. E. Scutum and metathorax + propodeum, holotype. A. Gonatopus javanus from Japan. B. G. maurus from China (from Xu et al. 2006c). C, D. G. lankanus (from Olmi 2003). E, F. G.lucidus (from Olmi 1984). G. G. maurus, holotype (from Olmi 1984). Scale bar 0.18 mm for A and G, 0.13 mm for B, 0.31 mm for D, 0.56 mm for E, 0.24 mm for F.

PLATE 131. A–D. Male genitalia. E,G, I. Mesosoma. F, H, J.Chelae, holotypes. A. Gonatopus rufoniger from China. B, I, J. G. superbus (from Olmi 1987c). C. G. asiaticus from Japan. D. G. leurosipropodeus, holotype (from He & Xu, 2002). E, F. G. schenklingi, holotype (from Olmi 1984). G, H. G. sitae (from Olmi 1984). Scale bar 0.19 mm for A, 0.18 mm for B, 0.15 mm for C, 0.17 mm for D, 0.21 mm for F, 0.34 mm for H, and 0.32 mm for J.