Gonatocerus Nees

Huber, John T., 2015, World reclassification of the Gonatocerus group of genera (Hymenoptera: Mymaridae), Zootaxa 3967 (1), pp. 1-184 : 29-32

Gonatocerus Nees (= sulphuripes species group, of authors)

( Figs 109–120 )

Gonatocerus Nees, 1834: 192 ; Bouček & Graham, 1972: 125 (validity of genus name and identity of the type-species); Triapitsyn et al., 2010: 12 (subgenus under Gonatocerus , s. l.), 14 (key to Neotropical species); Triapitsyn, 2013a: 8 (subgenus under Gonatocerus , s. l.), 9 (key to Palaearctic species).

Type species. Gonatocerus longicornis Nees , by monotypy; recognized by Ashmead, 1904: 376 and most subsequent authors.

Diagnosis. Within Gonatocerini, specimens of Gonatocerus are distinguished by the following combination of features: pronotum with lobes abutting medially; fore wing relatively narrow with microtrichia extending to base of venation and as dense behind as beyond venation; dorsellum rhomboidal; propodeum usually without carinae.

Description. FEMALE. Occiput entire ( Fig. 112 ). Vertex with 2 setae between lateral ocelli ( Figs 110, 112 ). Malar sulcus distinct ( Figs 109, 111, 113 ). Pronotum longitudinally divided, the lobes abutting medially except sometimes slightly separated posterodorsally ( Fig. 116 ). Pronotal spiracle about same size pronotal spiracle ( Fig. 116 ). Fore wing relatively narrow, with microtrichia usually as dense behind venation as beyond venation ( Fig. 108 b) but sometimes sparse or absent behind parastigma, the cubital line of setae always present and complete to base of parastigma. Stigmal vein with apex oblique ( Fig. 108 b). Dorsellum rhomboidal ( Figs 6 , 115 ). Propodeum with faint, converging sublateral carinae or carinae absent ( Fig. 115 ). Ovipositor not produced anteriorly under mesosoma but sometimes exserted beyond gastral apex ( Fig. 120 ).

MALE. Radicle usually long (up to half length of scape), separated from scape by a line or sulcus. Scape several times as long as pedicel. Genitalia with aedeagal apodemes fused at junction with aedeagus and almost as long as apodeme of genital sternum.

Discussion. Gonatocerus is not as easy to characterize as other genera. It is most easily confused with some Cosmocomoidea species. Additional features that may help distinguish Gonatocerus species are: fl2 and fl3 are often the longest funicle segments and the apex of fl1 is often oblique. The dorsellum is often more distinctly triangular than in Cosmocomoidea and its margin may be lighter in colour than it is medially. Species such as G. stenopterus (Ogloblin) with a very large pronotal spiracle, but not as large as in some Gastrogonatocerus species, are placed in Gonatocerus based on the relatively narrow wings with fore wing microtrichia extending uniformly to the base of the venation.

A fossil in Baltic amber is described below. The relatively narrow fore wing, venation length (as a proportion of the fore wing length) the same as in most extant Gonatocerus species, microtrichia extending proximally past the base of the parastigma, and lack of a hair line or fold extending posteroapically from the stigma indicated that the species is better placed in Gonatocerus than in Archigonatocerus. The species is described because it represents the second species of the genus to be found in amber (the first was G. henneberti Meunier ) and the only one that can be described and illustrated in some detail.

Distribution. The genus occurs worldwide.

Hosts and habitat. Cicadellidae (Hemiptera) . A few Gonatocerus species have reliably recorded hosts, e.g., Exitianus obscurinervis (Stål) and Spangbergiella vulnerata lacerdae Signoret for G. californicus Girault (Albarracin & Triapitsyn 2012) , Nephotettix cincticeps Uhler for G. cincticipitis Sahad , and Cicadella viridis (L.) for G. cicadellae Nikol’skaya ( Sahad & Hirashima 1984). Habitats range from forests to grasslands and deserts.

Included species:

Gonatocerus aegyptiacus Soyka , comb. rev. from Lymaenon . Gonatocerus aegyptiacus Soyka, 1950: 125 ;

holotype ♀ in PPDD (lost). 1 ♀ and 2 ♂ paratypes in NHMW (examined). Heqvist, 1960: 430 (transferred to

Lymaenon ). TL: Egypt, Giza, Shareh El-Haram.

Gonatocerus alami Shamim & Adam Shafee, 1984: 623 ; holotype ♀ in ZDAMU. TL: India, Uttar Pradesh,

Aligarh. Synonymy under G. tarae by Zeya & Hayat, 1995: 84. Synonymy under aegyptiacus by

Triapitsyn, 2013a: 9.

Gonatocerus minor Matthews, 1986: 220 ; holotype ♀ in BMNH (examined). TL: England, Essex, Hatfield Forest. Synonymy under G. aegyptiacus by Triapitsyn, 2013a: 9.

Gonatocerus miurai Sahad, 1982b: 195 ; holotype ♀ in KUEC. TL: Japan, Shimane, Matsue. Synonymy under G. tarae by Zeya & Hayat, 1995: 84. Synonymy under G. aegyptiacus by Triapitsyn, 2013a: 9. Gonatocerus saipanensis (Doutt) . Lymaenon saipanensis Doutt, 1955: 13 ; holotype ♀ in USNM (examined). TL: Mariana Islands, Saipan. Synonymy under G. aegyptiacus by Triapitsyn, 2013: 9.

Gonatocerus tarae (Narayanan & Subba Rao) . Lymaenon tarae Narayanan & Subba Rao, 1961: 657 ; holotype ♀ in IARI. Subba Rao & Hayat, 1983: 137 (transferred to Gonatocerus ). TL: India, Delhi. Synonymy under G. aegyptiacus by Triapitsyn, 2013a: 9.

Gonatocerus aequatorianus Ogloblin. Lymaenon aequatorianus Ogloblin, 1959a: 58 ; holotype ♂ in MLPA. De Santis, 1979: 12, 364 (transferred to Gonatocerus ); Triapitsyn et al., 2010: 15 [placed in Gonatocerus (Gonatocerus) ]. TL: Ecuador: Esmeraldas, San Mateo.

Gonatocerus alberti (Debauche) , comb. n. from Lymaenon . Lymaenon alberti Debauche, 1949: 56 ; holotype ♀ in MRAC (lost). TL: Congo, Parc National Albert, Kabasha escarpment.

Gonatocerus antillensis Dozier, 1937: 132 ; holotype ♀ in USNM (examined). TL: Puerto Rico, Mayaguez.

Gonatocerus appendiculatus (Ogloblin) . Lymaenon appendiculatus Ogloblin, 1939: 239 ; lectotype ♀ in MLPA (examined). De Santis, 1967: 10, 103 (transferred to Gonatocerus ). TL: Argentina: Misiones, Loreto.

Gonatocerus bonaerensis (Ogloblin) . Lymaenon bonaerensis Ogloblin, 1939: 246 ; lectotype ♀ in MLPA (examined). De Santis, 1967: 10, 105 (transferred to Gonatocerus ). TL: Argentina: Buenos Aires, José C. Paz. Gonatocerus ogloblini De Santis, 1967: 105 . Unnecessary replacement name for Gonatocerus bonaerensis ( Ogloblin, 1939) , not bonariensis ( Brèthes, 1922) [now in Cosmocomoidea ].

Gonatocerus bukashka Triapitsyn. Gonatocerus (Gonatocerus) bukashka Triapitsyn, 2013a: 15 ; holotype ♀ in CAS. TL: Russia, Sakhalinskaya Oblast’, Sakhalin Island, ~ 6 km E. Sokol.

Gonatocerus californicus Girault, 1911 [96]: 271; holotype ♀ in USNM (lost). TL: USA, California, Siskiyou Co., no locality given.

Gonatocerus utahensis Girault, 1917 [331]: 446; lectotype ♀ in USNM (examined). TL: USA, Utah, Salt Lake City. Synonymy under G. californicus by Triapitsyn, 2013b: 212.

Gonatocerus titillatus Girault, 1917 [336]: 87; holotype ♀ in USNM (examined). TL: USA, Utah, Salt Lake City. Synonymy under G. utahensis by Huber, 1988: 37.

Gonatocerus cincticipitis Sahad, 1982b: 192 ; holotype ♀ in KUEC. TL: Japan, Shimame, Matsue.

Gonatocerus comptei Girault, 1912 [120]: 135; 3 ♀ syntypes in QMBA. The original number and sex of the syntypes is unclear from the original description. According to Dahms (1983: 193) only 1 ♀ still exists. TL: Australia, Queensland, Nelson (= Gordonvale).

Gonatocerus edentulus Zeya in Zeya & Hayat, 1995: 78; holotype ♀ in BMNH (examined). TL: India, Bihar, Ranchi.

Gonatocerus excisus (Ogloblin) . Lymaenon excisus Ogloblin, 1936: 53 ; lectotype ♀ in MLPA (examined). De Santis, 1967: 10, 104 (transferred to Gonatocerus ). TL: Argentina: Misiones, Loreto.

Gonatocerus floridensis Huber, 1988: 46 ; holotype ♀ in CNC (examined). TL: USA, Florida, Collier Co., Collier Seminole State Park.

Gonatocerus fuscicornis (Walker) . Lymaenon fuscicornis Walker, 1846: 51 ; lectotype ♀ in MVMA. Dalla Torre, 1898: 429 (transferred to Gonatocerus ). TL: probably England.

Gonatocerus alecto (Debauche) . Lymaenon alecto Debauche, 1948: 105 ; holotype ♀ in IRSNB (examined). Matthews, 1986: 218 (transferred to Gonatocerus ). TL: Belgium, Eegenhoven. Synonymy under G. sulphuripes by Matthews, 1986: 218. Synonymy under G. fuscicornis by Triapitsyn, 2013a: 17. Gonatocerus crassipes (Debauche) . Lymaenon crassipes Debauche, 1948: 109 ; holotype ♀ in IRSNB (examined). Matthews, 1986: 218 (transferred to Gonatocerus ). TL: Belgium, Eegenhoven. Synonymy under G. s ulphuripes by Matthews, 1986: 218. Synonymy under G. fuscicornis by Triapitsyn, 2013a: 17. Gonatocerus pictosimilis Soyka, 1946: 38 ; holotype ♀ in NHMW. TL: Germany, Ramischau near Breslau. Synonymy under G. sulphuripes by Matthews, 1986: 218. Synonymy under G. fuscicornis by Triapitsyn, 2013a: 17.

Gonatocerus sulphuripes (Foerster) . Rachistus sulphuripes Foerster, 1847: 204 ; lectotype ♀ in NHMW (examined). Dalla Torre, 1898: 430 (transferred to Gonatocerus ); Bakkendorf, 1934: 29 (transferred to Lymaenon ); Thompson, 1958: 571 (transferred to Gonatocerus ); Triapitsyn, 2013a: 19 (lectotype designation and discussion). TL: Germany, Aachen. Synonymy under G. fuscicornis by Triapitsyn, 2013a: 17.

Gonatocerus synaptus (Debauche) . Lymaenon synaptus Debauche, 1948: 102 ; holotype ♀ in IRSNB (examined). Matthews, 1986: 218 (transferred to Gonatocerus ). TL: Belgium, Eegenhoven. Synonymy under G. sulphuripes by Matthews, 1986: 218. Synonymy under G. fuscicornis by Triapitsyn, 2013a: 17.

Gonatocerus granulosus (Ogloblin) . Lymaenon (Lymaenon) granulosus Ogloblin, 1959b: 189 ; holotype ♂ [on same slide as a ♀] in MLPA (examined). De Santis, 1967: 10, 104 (transferred to Gonatocerus ). TL: Argentina: Misiones, 25 de Mayo. [This species may better be placed in Cosmocomoidea .]

Gonatocerus hallami Girault, 1920 [353]: 99;?15 ♀ and ♂ syntypes in QMBA (examined). TL: Australia, Queensland, Nelson (= Gordonvale).

† Gonatocerus henneberti Meunier, 1905: 2 ; holotype ♀ in Klebs Collection, Provincial Museum of Königsberg (now Kaliningrad) (lost, presumed destroyed). TL: Baltic amber (perhaps from Samland Peninsula), mid- Eocene at the earliest ( Weitschat & Wichard 2010).

Gonatocerus huberi Zeya in Zeya & Hayat, 1995: 83; holotype ♀ in BMNH (examined). TL: India, Kerala, Mukkal.

Gonatocerus inaequalis (Debauche) , comb. n. from Lymaenon . Lymaenon inaequalis Debauche, 1949: 54 ; holotype ♀ in MRAC (lost). TL: Congo, Kivu, Rutshuru, river Fuku.

Gonatocerus ipswichia Girault, 1922 [361]: 104;? holotype ♀ in QMBA (lost). TL: Australia, Queensland, Ipswich.

Gonatocerus ipswichia variguttus Girault, 1938 [452]: 393; holotype ♀ in QMBA (examined). TL: Australia, Queensland, Ashgrove.

† Gonatocerus janzeni Huber ; holotype ♀ in AMNH. TL: Eocene Baltic amber. No present day locality given.

Gonatocerus koebelei Perkins, 1912: 22 ; lectotype ♀ in BPBM (examined). TL: Mexico, Veracruz, Orizaba.

Gonatocerus koziavka Triapitsyn, 2013 . Gonatocerus (Gonatocerus) koziavka Triapitsyn, 2013a: 26 ; holotype ♀ in UCRC. TL: Kyrgyzstan, Osh, Karakuldzha, Lajsu Ravine.

Gonatocerus lissonotus Huber, 1988: 47 ; holotype ♀ in CNC. TL: Canada, Ontario, Oxford Mills.

Gonatocerus longicornis Nees, 1834: 127 ; neotype ♀ in BMNH (examined), designated by Bouček & Graham, 1972: 129. TL: Italy, Aosta Province, Quart.

Gonatocerus britteni (Hincks) . Lymaenon britteni Hincks, 1960: 211 ; holotype ♀ in MMUE. Bouček & Graham, 1978a: 109 (transferred to Gonatocerus ). TL: UK, Cumberland, Skirwith. Synonymy under G. longicornis by Bouček & Graham, 1978b: 235.

Gonatocerus cicadellae Nikol'skaya, 1951: 575; 30 ♀ and 10 ♂ syntypes in ZIN (lost). TL: Kyrgyzstan (then Kyrgyz SSR), Bishkek (then Frunze) region, Kyzyl-Asker district. Synonymy under G. longicornis by Triapitsyn, 2003: 211.

Gonatocerus longiventris (Boţoc) . Lymaenon longiventris Boţoc, 1963: 96 ;?1 ♀ and? 1 ♂ syntypes in Boţoc collection (lost). Matthews, 1986: 218 (transferred to Gonatocerus ). TL: Romania, Cluj environs. Synonymy under G. longicornis by Matthews, 1986: 218.

Gonatocerus shasthryi (Subba Rao & Kaur) . Lymaenon shasthryi Subba Rao & Kaur, 1959: 234 ; holotype ♀ in IARI. Zeya & Hayat, 1995: 80 (transferred to Gonatocerus ). TL: India, Karnataka, Mandya. Synonymy under G. longicornis by Zeya & Hayat, 1995: 80.

Gonatocerus terebrator (Foerster) . Rachistus terebrator Foerster, 1847: 203 ; lectotype ♀ in NMV. Dalla Torre, 1898: 430 (transferred to Gonatocerus ); Debauche, 1948: 81 (transferred to Lymaenon ); Bouček & Graham, 1972: 127 (transferred to Gonatocerus ). TL: Germany, Aachen. Synonymy under G. longicornis by Bouček & Graham, 1972: 127.

Gonatocerus uttarodeccanus Mani & Saraswat, 1973: 97 , holotype ♀ in USNM. TL: India, Tamil Nadu, Anamalai Hills, Munnar. Synonymy under G. longicornis by Zeya & Hayat, 1995: 81.

Gonatocerus longicrus Kieffer, 1913: 201 ; holotype ♀ in NZSI (lost). Zeya & Hayat, 1995: 125 (species incertae sedis). TL: India, Orissa, Puri.

Gonatocerus maga Girault, 1911 [96]: 267; holotype ♀ in INHS (lost). TL: USA, Illinois, Litchfield.

Gonatocerus mandyanus Zeya & Usman, 2014: 64 ; holotype ♀ in NBAII. TL: India, Karnataka, Mandya.

Gonatocerus meghalayanus Zeya, 2011: 33 . Replacement name for Gonatocerus orientalis Zeya, 1995 , not G.

orientalis (Girault, 1917) , which is now in Lymaenon .

Gonatocerus orientalis Zeya in Zeya & Hayat, 1995: 89; holotype ♀ in BMNH (examined). TL: India, Meghalaya [no town locality given], botanic garden.

Gonatocerus zeyai Özdikmen, 2011: 840 . Unnecessary replacement name for G. orientalis Zeya ; G. meghalayanus is a pre-existing replacement name.

Gonatocerus mexicanus Perkins, 1912: 21 ; lectotype ♀ in BPBM (examined). TL: Mexico, Districto Federal, Chapultepec.

Gonatocerus eximius Gahan, 1913: 441 ; lectotype ♀ in USNM (examined). TL: USA, Florida, Orlando. Synonymy under G. mexicanus by Girault, 1929 [428]: 254.

Gonatocerus gibsoni Crawford, 1915: 586 ; lectotype ♀ in USNM (examined). TL: USA, AZ, Tempe. Synonymy under G. mexicanus by Girault, 1929 [428]: 254.

Gonatocerus petrarchi Girault, 1920 [353]: 99; 3 ♀ syntypes in QMBA (examined). TL: Australia, Queensland, Port Douglas, Nelson.

Gonatocerus pictus (Haliday) . Ooctonus pictus Haliday, 1833: 344 ; lectotype ♀ in NMID. Walker, 1846: 51 (transferred to Lymaenon ); Foerster, 1847: 205 (transferred to Rachistus ); Dalla Torre, 1898: 429 (transferred back to Gonatocerus ). TL:? Ireland, no locality given.

Gonatocerus flavus Förster, 1841: 45 ; 2 ♀ syntypes in NHMW (lost). Foerster, 1847: 205 (implied transfer to Rachistus ); Dalla Torre, 1898: 429 (transferred back to Gonatocerus ); Triapitsyn (2013a: 34) implied there was only one type specimen but Foerster (1847: 206) stated he had two specimens. TL: Germany,?Aachen area. Synonymy under G. pictus by Foerster, 1847: 205.

Gonatocerus orthopenitus Guo, Lin & Hu, 2011: 55 ; holotype ♀ in ICXU. TL: China Xinjiang, Uyghur Autonomous Region, Wujiaqu. Synonymy under G. pictus by Triapitsyn, 2013a: 34.

Gonatocerus risbeci (Heqvist) , comb. n. from Lymaenon . Lymaenon risbeci Heqvist, 1960: 430 . Replacement name for Gonatocerus africanus Risbec, 1956 , not G. africanus ( Soyka, 1950) , which is now in Lymaenon . Gonatocerus africanus Risbec , comb. rev. from Lymaenon . Gonatocerus africanus Risbec, 1956a: 153 ; 14 ♀ and 1 ♂ syntypes in MNHN (5 ♀ examined). Heqvist, 1960: 430 (transferred to Lymaenon ). TL: Cameroon, Garoua.

Gonatocerus camerounensis Özdikmen, 2011: 840 . Unnecessary replacement name for G. africanus Risbec ; G. risbeci is a pre-existing replacement name.

Gonatocerus rivalis Girault, 1911 [96]: 257; holotype ♀ in USNM (examined). TL: USA, Illinois, Butler.

Gonatocerus sarawakensis Sveum, 1982: 81 ; holotype ♀ in NTNU. TL: Malaysia, Sarawak, 20 km S. Kuching, Semengo.

Gonatocerus shamimi Subba Rao & Hayat, 1986: 187 . Replacement name for Gonatocerus terebrator Shamim & Adam Shafee, 1984 , not G. terebrator ( Foerster, 1847) , which is now a synonym of G. longicornis . Gonatocerus terebrator Shamim & Adam Shafee, 1984: 624 ; holotype ♀ in ZDAMU. TL: India, Uttar Pradesh, Aligarh.

Gonatocerus stenopterus (Ogloblin) . Lymaenon stenopterus Ogloblin, 1936: 33 ; lectotype ♀ in MLPA (examined). De Santis, 1967: 10, 105 (transferred to Gonatocerus ). TL: Argentina, Misiones, Loreto.

Gonatocerus urocerus Ogloblin, 1935: 72 ; lectotype ♀ in MLPA (examined). TL: Argentina: Buenos Aires, La Plata.

Gonatocerus utkalensis Subba Rao, 1989: 144 ; holotype ♀ in BMNH (examined). TL: India, Orissa, Cuttack, Central Rice Research Institute.

Figures

FIGURE 108. Gonatocerus janzeni, holotype in amber: a, habitus; b, wing showing venation details. Scale bars = 500 Μm.

FIGURES 109 – 114. Gonatocerus rivalis, head: 109, anterior; 110, dorsal; 111, lateral; 112, posterior; 113, ventral; 114, mouthparts, ventral. Scale bars = 50 Μm.

FIGURES 115 – 120. Gonatocerus rivalis: 115, mesosoma dorsal; 116, mesosoma lateral; 117, mesosoma ventral; 118, mesosoma anterior; 119, metasoma dorsal; 120, metasoma lateral. Scale bars = 100 Μm.

FIGURES 6 – 9. Dorsal views: 6, Gonatocerus sp., entire body; 7, Lymaenon sp., entire body; 8, Lymaenon sp., pronotum, the lateral lobes widely separated medially; 9, Cosmocomoidea sp., pronotum, the lateral lobes abutting medially. Maximum diameter across mesoscutum (just anterior to fore wing base): 8 = 300 Μm, 9 = 330 Μm.