Gonatocerus (Lymaenon) litoralis ( Haliday, 1833 )

Triapitsyn, Serguei V., 2013, Review of Gonatocerus (Hymenoptera: Mymaridae) in the Palaearctic region, with notes on extralimital distributions, Zootaxa 3644 (1), pp. 1-178 : 82-97

Gonatocerus (Lymaenon) litoralis ( Haliday, 1833)

( Figs 136–153 )

Ooctonus litoralis Haliday 1833: 342 (mentioned in footnote), 344.

Lectotype female [ OUMNH], designated by Graham 1982: 223 –224 (not examined). Type locality: near Holywood, Co. Down, Northern Ireland, UK.

Ooctonus litoralis Haliday : Curtis 1837: 134 (list).

Lymaenon litoralis (Haliday) : Walker 1846: 51 (diagnosis); Debauche 1948: 81 (list), 89–91 (redescription); Radu & Boţoc 1960: 324 –325 (redescription, illustrations); Mathot 1969: 2 (list, records from Belgium and Norway), 11 (key).

Rachistus litoralis (Haliday) : Foerster 1847: 204 –205 (diagnosis), 232 (list, distribution).

Alaptus fuscus Förster 1861: 43 .

Type locality: Val Roseg, Upper Engadin, Graubünden, Switzerland. Syn. n.

Gonatocerus exiguus Förster 1861: 43 .

Type locality: Val Roseg, Upper Engadin, Graubünden, Switzerland. Syn. n.

Gonatocerus litoralis (Haliday) : Kirchner 1867: 201 (catalog); Dalla Torre 1898: 429 (catalog); Soyka 1946: 37 –38 (diagnosis, records from Netherlands and Poland); Graham 1973: 47 (mentioned); Hellén 1974: 12 –13 (diagnosis, distribution); Graham 1982: 223 –224 (references, lectotype designation, record from Northern Ireland); Sahad 1982c: 198 (key); Sahad & Hirashima 1984: 3 (list), 15–17 (redescription, illustrations, distribution); Matthews 1986: 223 (member of the litoralis species group , synonymy, diagnosis, distribution); Donev 1990: 68 (records from Bulgaria); Baquero & Jordana 2003: 5 –7 (diagnosis, redescription, distribution, illustrations, host associations); Donev 2005: 382 –383 (diagnosis, distribution); Pricop 2009a: 73 (records from Romania), 74 (illustrations); Pricop 2009b: 125 (list, references on the records from Romania); Pricop 2010a: 81 –83 (records from Romania, illustrations); Pricop 2010c: 78 –79 (illustrations), 81 (record from Romania, diagnosis); Fallahzadeh & Huber 2011: 298 (host in Iran).

Gonatocerus brunneus Webster 1894: 117 [as Gonotocerus (sic) brunneus Ashmead ]. Nom. nud. ( Peck 1963: 21), who considered it to be conspecific with G. brunneus Girault , which is synonymized here under G. litoralis .

Alaptus fuscus Förster : Dalla Torre 1898: 428 (catalog).

Gonatocerus exiguus Förster : Dalla Torre 1898: 429 (catalog).

Gonatocerus anthonomi Girault 1905: 288 –289 .

Originally described from the type series of 3 female and 1 male specimens, although only 1 female and 1 male, which are syntypes (the other 2 females are paratypes), were designated by Girault as “Type” under USNM No. 8435 (male not examined); later Girault (1911: 260) mentioned, besides the two aforementioned “types” of each sex (both according to him were tag-mounted), also “4 paratypes ” (a least two of these in fact have no type status), reared at the same time with the “types” (A.A. Girault deposited one of those “ paratypes ” in INHS).

Type locality: Fort Valley, Peach Co., Georgia, USA. Syn. n.

Gonatocerus welateri Ashmead : Nason 1906: 8. Nom. nud. ( Girault 1911: 324; a misprint of W.H. Ashmead’s manuscript name G. “ websteri ” = G. anthonomi ).

Gonatocerus americanus Brues 1907: 109 –110 .

Type locality: Milwaukee, Milwaukee Co., Wisconsin, USA. Syn. n.

Gonatocerus anthonomi Girault: Girault 1911: 256–257 , 260–261 (comparison, type information, distribution, etc.), 274 (key), 324 (comment); Girault 1929: 26 (key, synonymy); Huber 1988: 31 (member of the litoralis species group).

Gonatocerus americanus Brues : Girault 1911: 256 –257, 261 (type information, diagnosis), 274 (key); Girault 1929: 27 (key); Huber 1988: 31 (member of the litoralis species group).

Gonatocerus brunneus Girault 1911: 261 –263 , 274 (key).

Originally described from 2 female and 2 male specimens, which are all syntypes, although 3 females were designated as “Types” (2 females from Indiana on the same slide, USNM Accession No. 13,803) or “Cotypes” (1 female from Urbana, Illinois, on a slide, INHS Accession No. 44,237). The other specimens from Illinois mentioned (3 females, 2 males) are paratypes because they also belong to the type series. Type locality (of the lectotype designated here): an unspecified locality in Indiana, USA. Syn. n.

Gonatocerus texanus Girault 1911: 270 –271 , 274 (key).

Originally described from 3 female specimens, although only 2 females were designated by Girault as “Types” (syntypes on a slide, USNM Accession No. 13,823). The other specimen mentioned in the original description (the same data, examined) is paratype because it belongs to the type series; it is mounted under the same coverslip with the two syntypes. Type locality: College Station, Brazos Co., Texas, USA. Synonymized under G. anthonomi Girault by Girault 1929: 26. Syn. n.

Gonatocerus maevius Girault 1911: 272 , 273 (key) [as maevius ].

Holotype female on a slide [ INHS Accession No. 1692] (lost according to Huber (1988), not examined). Type locality: Normal, McLean Co., Illinois, USA. Synonymized under G. brunneus Girault by Girault 1929: 26. Syn. n.

Gonatocerus illinoiensis Girault 1917: 91 –92 .

Type locality: Coulterville, Randolph Co., Illinois, USA. Syn. n.

Gonatocerus radiculatus Ahlberg 1925: 85 –86 .

3? syntype females [ Matthews (1986) mentioned a holotype (an invalid designation) without seeing the original type material, supposedly deposited in NHRS] (not examined). Type locality: near Spånga, Sweden. Synonymized under G. litoralis by Matthews 1986: 223.

Gonatocerus brunneus Girault: Girault 1929: 26 (key, synonymy); Huber 1988: 31 (member of the litoralis species group).

Gonatocerus illinoisensis Girault: Girault 1929: 27 (key, misspelling or invalid emendation).

Lymaenon effusi Bakkendorf 1934: 23 –29 .

Lectotype female [lost from ZMUC; according to John S. Noyes (personal communication), some of Bakkendorf’s type specimens of Gonatocerus were lost when they were returned to ZMUC from BMNH where they had been on loan to M.J. Matthews] (not examined), effectively designated by Matthews (1986) (Article 74.6, [ ICZN] 1999) because although Bakkendorf (1934) did not designate any type material of the species described in that publication, it is apparent from it that only one female of Lymaenon effusi , which emerged 16.v.1929 from the host eggs collected 26.xii.1928, was available to him, and that specimen was later examined by Matthews. Type locality: Dyrehaven (Jaegersborg Dyrehave, Zealand Island), Hovedstaden, Denmark. Synonymized under G. litoralis by Matthews 1986: 223.

Lymaenon exiguus (Förster) : Debauche 1948: 81 (list).

Lymaenon fuscus (Förster) : Debauche 1948: 81 (list).

Lymaenon radiculatus (Ahlberg) : Debauche 1948: 81 (list).

Lymaenon paludis Debauche 1948: 91 –93 + plate X (illustrations).

Type locality: Abbaye du Parc, Heverlee (as Héverlé in the original description), Leuven, Flemish Brabant, Belgium. Synonymized under G. litoralis by Matthews 1986: 223.

Lymaenon ater (Förster) : Debauche 1948: 93 –95 (misidentification), plate X (illustrations); Hincks 1960: 210 (misidentification, diagnosis); Mathot 1969: 11 (key, apparently based on misidentified specimens from Belgium).

Lymaenon rhacodes Debauche 1948: 97 –99 + plate XI (illustrations).

Type locality: Parc d’Arenberg (park at Kasteel van Arenberg), Heverlee (as Héverlé in the original description), Leuven, Flemish Brabant, Belgium. Synonymized under G. litoralis by Matthews 1986: 223.

Lymaenon littoralis [sic] (Haliday): Debauche 1949: 27.

Gonatocerus priesneri Soyka 1950: 128 –129 .

Holotype female [lost from PPDD] (not examined). Type locality: Wadi Aideb, Gebel Elba, Red Sea Governorate, Egypt. Syn. n.

Gonatocerus radiculatus Ahlberg : Nikol’skaya 1952: 538 (mentioned, host association).

Lymaenon “B” : Huffaker et al. 1954: 786 –787, 789 (host association, imported from Spain and released in California, USA).

Lymaenon americanus (Brues) : Burks 1958: 63 (catalog [not actually listed but referring to Peck (1951)]); Peck 1963: 20 –21 (catalog).

Lymaenon anthonomi (Girault) : Burks 1958: 63 (catalog [not actually listed but referring to Peck (1951)]); Peck 1963: 21 (catalog).

Lymaenon brunneus (Girault) : Burks 1958: 63 (catalog [not actually listed but referring to Peck (1951)]); Peck 1963: 21 –22 (catalog).

Lymaenon illinoiensis (Girault) : Burks 1958: 63 (catalog [not actually listed but referring to Peck (1951)]).

? Lymaenon ramakrishnai Subba Rao & Kaur 1959: 229 , 231 (illustrations), 232–234 [as Lymaenon ], 237 (key).

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

Lymaenon illinoisensis (Girault) : Peck 1963: 22 –23 (catalog, invalid emendation).

Lymaenon arduennae Mathot 1969: 2 (list), 5–6, 12 (key).

Type locality: Ry de Grahais, Nafraiture, Vresse-sur-Semois, Namur, Belgium. Synonymized under G. litoralis by Matthews 1986: 223.

Lymaenon rhacodes Debauche : Mathot 1969: 11 (key).

Lymaenon paludis Debauche : Mathot 1969: 11 (key); Donev 1985a: 62 –63 (distribution).

Gonatocerus paludis (Debauche) : Graham 1973: 47 (comments); Donev 1988b: 181 [as Gomatocerus [sic] paludis ] (distribution); Donev 1988d: 195 (distribution); Donev 1990: 68 (records from Bulgaria); Dimitriu 2001: 25 (record from Romania).

Gonatocerus pulchellus Hellén 1974: 11 .

Holotype female [ FMNH] (not examined). Type locality: Hauho, Finland. Synonymized under G. litoralis by Matthews 1986: 223.

Gonatocerus “B” : Clausen 1978: 56 (host association, information on releases in California).

Gonatocerus (Gonatocerus) anthonomi Girault : De Santis 1979: 365 (catalog).

? Gonatocerus ramakrishnai (Subba Rao & Kaur) : Subba Rao & Hayat 1983: 136 (catalog); Zeya & Hayat 1995: 99 –100 (redescription, holotype information, short diagnosis), 146 (illustrations).

Lymaenon effusi Bakkendorf : Donev 1985a: 62 (distribution).

Gonatocerus effusi (Bakkendorf) : Donev 1987: 73 –74 (distribution); Donev 1988d: 194 (distribution); Donev 1988e: 204 (distribution).

Gonatocerus arduennae (Mathot) : Donev 1988c: 186 (records from Greece).

Gonatocerus illinoiensis Girault : Huber 1988: 31 (member of the litoralis species group ).

Gonatocerus litoraris [sic] (Haliday): Donev 1990: 68.

? Gonatocerus sp. 2 : Walker et al. 1997: 240 –241 (culture #23-GD, specimens of Iran origin).

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

Gonatocerus (Lymaenon) litoralis (Haliday) : Triapitsyn 2010: 9 ( mentioned from Austria); Triapitsyn et al. 2010: 48–53 (references, records from the Nearctic and Neotropical regions as well from Austria, Belgium, Bulgaria, and UK, redescription based on the Neotropical specimens, diagnosis, illustrations, distribution, host associations), 57.

Gonatocerus (Lymaenon) anthonomi Girault : Triapitsyn et al. 2010: 42, 53 (comments, possible conspecificity with G. litoralis ).

Type material examined. Alaptus fuscus Förster : lectotype female [ NHMW], here designated to avoid confusion about the status of the type specimen(s) of this species, on slide ( Fig. 136 ) labeled originally: 1. “ Alaptus . Ƥ fuscus m. [in black ink in A. Foerster’s handwriting]”; 2. [in black ink, illegible but apparently] “Rosegg Thal”; 3. “Först.”; 4. “Collect. G. Mayr”; 5. “ Al. fuscus [in black ink in A. Foerster’s handwriting] Förster, Type]; 6. [in W. Soyka handwriting] “( Alaptus fuscus Förster ) Gonatocerus (Canada balsam)”; 7. “21” [W. Soyka’s slide number]. The lectotype, remounted by Soyka from a minuten pin, is in poor condition, uncleared, dissected in many parts under the same coverslip: head and 1 antenna, the other antenna, mesosoma (dorsoventrally mounted) plus most of the legs and 1 fore wing and both hind wings, gaster (mounted laterally), the other fore wing, 2 tarsi. This seems to be an aberrant specimen of G. litoralis : that may be explained perhaps by an observation that mymarid specimens from high altitudes, besides being usually somewhat darker in color, sometimes show higher and weirder variation in some other morphological features than conspecific specimens from low elevation habitats.

Gonatocerus exiguus Förster : lectotype male [ NHMW], here designated to avoid confusion about the status of the type specimen(s) of this species, on a minuten pin labeled in black ink in A. Foerster’s handwriting: 1. “Rosseg”; 2. “ Gonatocerus 3 exiguus m”; 3. “Coll. Förster don. Mayr”.

Gonatocerus americanus Brues : lectotype female [ MCPM], effectively designated (Article 74.6, [ICZN] 1999) by Girault (1911), on slide ( Fig. 137 ) labeled: “Milw. Public Museum Cat. No. 22256 Sp. Gonatocerus americanus Order Hymen Fam. MYMARIDAE TYPE ”. The lectotype ( Fig. 138 ) is in poor condition, uncleared, mounted laterally, with F5–F8 and clava of both antennae and one hind wing missing. The apical parts of the antennae were probably broken off at the time when A.A. Girault remounted the type specimen from a tag ( Girault 1911); earlier, Brues (1907) described the antenna but did not mention presence or absence of longitudinal sensilla on the funicle segments; he did not designate any type(s). The label on the slide is glued over the earlier label; information on the original label of the tag-mounted specimen was provided by Girault (1911).

Gonatocerus anthonomi Girault : lectotype female [ USNM], here designated to avoid confusion about the status of the type specimens of this species, on point labeled: 1. “ Gonatocerus anthonomi Girault ”; 2. “Ƥ Type No. 8435 U.S.N.M.”. The lectotype lacks the pedicel and flagellum of both antennae, one hind wing, and some leg segments.

Gonatocerus brunneus Girault : lectotype female [USNM], here designated to avoid confusion about the status of the type specimens of this species, on slide ( Fig. 141 ) labeled originally: 1. “ Gonatocerus brunneus Girlt Ƥ. 13803 [in pencil] Ind. Types USNM.”; 2. [red] “Type no. 13803”. The lectotype (circled in black ink) is the upper of the two original syntypes mounted under the same coverslip, the one at the bottom is the paralectotype. The lectotype is complete, mounted laterally; the paralectotype lacks pedicel and flagellum of one antenna.

Gonatocerus texanus Girault : lectotype female [ USNM], here designated to avoid confusion about the status of the type specimens of this species, on slide ( Fig. 139 ) labeled: 1. “ Gonatocerus texanus Girault texanus 2Ƥ’s. Types Ƥ 13823 [in pencil]”; 2. “USNM. Spms. Re-mounted from tags “College Station, Tex. Sept. Banks”. “ Alaptus Ƥ”.”; 3. [red] “Type no. 13823”. The lectotype (circled in black ink) is the upper left, most complete specimen of the three females mounted under the same coverslip, two of which are the original syntypes; one of the other two females (it is impossible to figure out which as both are incomplete) is the paralectotype. The lectotype is in a very poor condition, with several body parts detached.

Gonatocerus illinoiensis Girault : holotype female [ USNM] on slide ( Fig. 140 ) labeled: 1. “ Gonatocerus ? n. sp. Beautiful when fresh + golden yellow [in pencil] Coulterville, [an illegible word, apparently a wrong locality name, crossed out], Ill. VI.10.1911. aag. Laundry window”; 2. [red] “ G. illinoisensis Gir. Ƥ 20631”. The holotype is complete but uncleared, the head is detached, and the rest of the body is mounted laterally.

Lymaenon paludis Debauche : holotype female [ ISNB] on slide labeled: 1. “Dr. H. DEBAUCHE det. Lymaenon paludis Deb. Ƥ 1943 Type.”; 2. “Héverlé 28.VI.41 – n o 142”. The holotype is complete, uncleared, mounted laterally. Paratypes: 2 females [ISNB] on individual slides, labeled identically as the holotype except “Para-type.” instead of “Type.” on the first label.

Lymaenon rhacodes Debauche : holotype female [ ISNB] on slide labeled: 1. “Dr. H. Debauche det. Lymaenon rhacodes Deb. Ƥ 1943 TYPE [on red rectangle glued onto the label]”; 2. “Héverlé 1.X.41 – 161 6 [in pencil]”. The holotype is complete, uncleared, and mounted laterally.

Gonatocerus priesneri Soyka : paratypes [both NHMW]: 1 Ƥ on slide labeled: 1. “Gabal [sic] Elba W. Aideb 1.2.33 Lycium sp.”, 2. “Co-Type”, 3. “Ƥ Gonatocerus priesneri (Soyka) Ƥ”, 4. “819”; 1 3 on slide labeled: 1. “Gabal [sic] Elba W. Aideb 1.2.33 Lycium sp.”, 2. “Co-Type”, 3. “818”, 4. “Mym. 3 Gonatocerus priesneri (Soyka) 3”.

Lymaenon arduennae Mathot : holotype female [ ISNB] on slide ( Fig. 142 ) labeled: 1. “Université de Louvain LAB. ENTOMOLOGIE Nafraiture Ry des Grahais 9.IX.48 no279”; 2. “Dr. H. DEBAUCHE det. Lymaenon arduennae Ƥ Type Deb.”. The holotype ( Fig. 143 ) is in fair condition although uncleared, lacking a pair of wings, and mounted laterally.

Material examined. ARMENIA. SYUNIK: Lichk, 7.vi.1953, V.A. Trjapitzin [2 Ƥ, ZIN]. Megri, V.A. Trjapitzin : 9.vi.1953 (on tamarisk) [1 Ƥ, ZIN]; 9.vi.1953 (fruit orchard) [1 Ƥ, ZIN]; Megriget River bank, 23.vi.1953 ( Typha latifolia thicket) [1 Ƥ, ZIN]; 28.vi.1953 [2 Ƥ, ZIN]. AUSTRIA. BURGENLAND, Sankt Andrä am Zicksee , 11.viii.1942, S. Novicky [1 Ƥ, NHMW]. LOWER AUSTRIA: Hainburg an der Donau , 48°08’45’’N 16°55’31’’E, 142 m, 17.vi.2007, S.V. Triapitsyn, C. Thuróczy [3 Ƥ, UCRC]. 1 km W of Hollern , 48°04’22’’N 16°52’37’’E, 150 m, 16–17.vi.2007, S.V. Triapitsyn, C. Thuróczy [6 Ƥ, 1 3, UCRC]. Hundsheim : 19.v.1942, H. Bischoff [1 Ƥ, 1 3, NHMW]; 2.vi.1942, H. Bischoff [1 Ƥ, NHMW]; 8.vi.1942, H. Bischoff [1 Ƥ, NHMW]; 10.vi.1942, H. Bischoff [1 Ƥ, 1 3, NHMW] (male misidentified by W. Soyka as G. terebrator); 12.vii.1944, W. Soyka [1 Ƥ, NHMW]; 19.ix.1953, W. Soyka (from hay in the garden) [1 Ƥ, NHMW]; 20.ix.1953, W. Soyka (on window from hay in the garden) [3 Ƥ, NHMW]; 21.ix.1953, W. Soyka [1 3, NHMW]; 23.ix.1953, W. Soyka [1 3, NHMW]; 24.ix.1953, W. Soyka [2 Ƥ, 1 3, NHMW]; 27.ix.1953, W. Soyka [3 Ƥ, NHMW]; 22.ix.1954, W. Soyka [1 3, NHMW]; 10.x.1954, W. Soyka [1 Ƥ, NHMW]; 10.ix.1956, W. Soyka [1 Ƥ, NHMW]; ix.1960, W. Soyka [1 Ƥ, NHMW]; 48°07’14’’N 16°55’59’’E, 220 m, 16–17.vi.2007, S.V. Triapitsyn, C. Thuróczy [1 Ƥ, 4 3, UCRC]. Spitzerberg ( S slope, ca. 2.5 km S of Hundsheim ), 48°05’48’’N 16°56’29’’E, 190–250 m: 16.vi.2007, S.V. Triapitsyn, C. Thuróczy [8 Ƥ, 3 3, UCRC]; 17.vi.2007, C. Thuróczy, S.V. Triapitsyn [17 Ƥ, 2 3, UCRC]. TYROL: Gschnitztal , 16.ix.1948, E. Pechlaner [1 Ƥ, NHMW]. Innsbruck, Arzler Alm , 1200 m, 12.ix.1948, E. Pechlaner [1 Ƥ, NHMW]. Kematen , 9.x.1949, E. Pechlaner [1 Ƥ, NHMW]. Krössbach , W. Soyka: vii.1947 (on window) [1 Ƥ, NHMW]; ix.1957 [1 Ƥ, NHMW]. Stubaital , 2300 m, 13.ix.1951, E. Pechlaner [3 Ƥ, 1 3, NHMW]. [ Locality unclear], 4.ix.1949, E. Pechlaner [1 Ƥ, NHMW]. VIENNA, Vienna , 25.ix.1953, H.- J. Stammer [1 Ƥ, NHMW]. BELGIUM. FLEMISH BRABANT: Leuven: Heverlee , [H.R. Debauche]: 19.ix.1941 [2 Ƥ, ISNB] (one of them identified as Lymaenon paludis by H.R. Debauche ); 9.vii.1942 [4 Ƥ, ISNB]; 22.vii.1942 [4 Ƥ, ISNB]; 14.viii.1942 [1 Ƥ, ISNB]; 16.vii.1945 [1 Ƥ, ISNB]. Kessel-Lo , 27.viii.1945, [H.R. Debauche] [1 Ƥ, 1 3, ISNB]. Tervuren: 23.viii.1944, [H.R. Debauche] [5 Ƥ, ISNB] (one of them identified as Lymaenon rhacodes by H.R. Debauche, and also two identified as L. rhacodes and one as L. paludis apparently by G. Mathot); Étang du Merisier , 7.viii.1945, [H.R. Debauche] [1 Ƥ, ISNB] (identified as L. paludis by H.R. Debauche). LIÈGE, Wanze, Antheit , Corphalie, R. Detry : 16–30.vi.1989 [2 Ƥ, ISNB]; 1–14.vii.1989 [1 Ƥ, ISNB]; 14–28.vii.1989 [2 Ƥ, ISNB]; 28.vii–11.viii.1989 [4 Ƥ, ISNB]; 11–25.viii.1989 [5 Ƥ, ISNB]; 25.viii–8.ix.1989 [1 3, ISNB]; 8–22.ix.1989 [1 Ƥ, ISNB]; 6–20.x.1989 [2 Ƥ, ISNB]; 20.x–3.xi.1989 [25 Ƥ, ISNB]; 17.xi–1.xii1989 [1 Ƥ, ISNB]; 27.iv–11.v1990 [2 Ƥ, ISNB]; 25.v–8.vi.1990 [3 Ƥ, ISNB]; 8–22.vi.1990 [2 Ƥ, ISNB]; 28.vi–6.vii.1990 [2 Ƥ, ISNB]; 6–20.vii.1990 [3 Ƥ, ISNB]; 3–17.viii.1990 [2 Ƥ, ISNB]; 14–28.ix.1990 [9 Ƥ, ISNB]. WALLOON BRABANT, Waterloo , P. Dessart: 1.iv–13.v.1992 [1 Ƥ, ISNB]; 30.vi–5.vii.1992 [1 Ƥ, ISNB]; 26.vii–2.viii.1992 [1 Ƥ, ISNB]; 30.viii– 9.ix.1992 [1 Ƥ, ISNB]; 10–20.ix.1992 [16 Ƥ, 3 3, ISNB]; 20–27.ix.1992 [9 Ƥ, 1 3, ISNB]; 28.ix–4.x.1992 [1 Ƥ, ISNB]. BULGARIA. KYUSTENDIL, Kyustendil, 1928, L. Biró [1 Ƥ, NHMW / HNHM]. CHINA. BEIJING, Mentougou District: Liyan Ling , Linshan Mts. , 40°00.28’N 115°30.75’E, 1749 m, 2.viii.2002, G. Melika [4 Ƥ, UCRC]. Xiaolongmen Station , 39°59.22’N 115°31.48’E, 1095 m, 28.vii.2002, G. Melika [4 Ƥ, 1 3, UCRC]. CYPRUS . LIMASSOL, Saitta ( Saittas ), 1.xi.1967, G.P. Georghiou [1 Ƥ, UCRC]. CZECH REPUBLIC. KARLOVY VARY, Sokolov District, Dolní Rychnov , 50°09’18.902’’N 12°39’38.824’’E, 461 m, 27.viii.2007, J. Macek [1 Ƥ, CUPC]. ÚSTÍ NAD LABEM, České Švýcarsko National Park , 50°50’00.494’’N 14°19’49.508’’E, 597 m, 16.v.2007, J. Macek [2 Ƥ, CUPC]. DENMARK. HOVEDSTADEN: Dyrehaven ( Jaegersborg Dyrehave , Zealand Island), Fortunens Indelukke , 21.vii.1924, O. Bakkendorf [1 Ƥ, 1 3, ZMUC]. Freerslev (as “ Frerslev Forest ”), 8.vii.1954, O. Bakkendorf [1 Ƥ, ZMUC]. Lingby , 20.ix.1951, O. Bakkendorf [1 Ƥ, ZMUC]. MIDTJYLLAND: Glatved , 56°17’N 10°50’E, 18.viii.1997, T. Munk [2 females, ZMUC]. Sillerup (12 km SW of Silkeborg ): 13.x.1986, T. Munk [1 female, ZMUC]. Skramsø , 56°17’N 10°40’E, 11.viii.1997, T. Munk [1 female, ZMUC]. SJAELLAND: Liselund, 10.viii.1924, O. Bakkendorf [1 Ƥ, ZMUC]. Rude , 7.ix.1924, O. Bakkendorf [2 Ƥ, ZMUC]. FINLAND. [ No locality or other label data] [3 Ƥ, 1 3, NHMW]. FRANCE. AUDE, Lézignan-Corbières , 28.vi.2000, S.V. Triapitsyn (on grape) [1 Ƥ, UCRC]. GARD: Near Gardon River , 43°55’45’’N 4°23’25’’E, 10– 13.vi.2005, J. George [6 Ƥ, 2 3, UCRC]. Sainte-Eulalie (W of Uzès ), 43°59’16’’N 04°17’53’’E, 96 m, 10– 12.vi.2005, J. George [11 Ƥ, UCRC]. GIRONDE: Sainte Colombe , 44°54’N 00°02’W, M. van Helden: 2.vii.1998 [3 Ƥ, 2 3, UCRC]; 30.vii.1998 [5 Ƥ, 2 3, UCRC]; 13.viii.1998 [61 Ƥ, 2 3, UCRC]; 10.ix.1998 [2 Ƥ, UCRC]; 9.vii.1999 [4 Ƥ, UCRC]; 24.viii.2000 [2 Ƥ, UCRC]. Tourtirac , 44°53’57’’N 00°02’02’’W, 100 m: 27.viii.1998, M. van Helden [2 Ƥ, UCRC]; 26–27.vi.2000, S.V. Triapitsyn [5 Ƥ, 1 3, UCRC]. HÉRAULT, St. Clément de Rivière, 43°41’47’’N 03°51’13’’E, 29.vi.2000, S.V. Triapitsyn (on grape near Lez River ) [1 Ƥ, UCRC]. GEORGIA. ADJARA: Batumi region, Kakhaberi , 5.ix.1953, V.A. Trjapitzin ( Gruzbiolaboratoriya ) [1 3, ZIN]. Keda , V.A. Trjapitzin: 29.viii.1953 (edge of pine forest) [1 Ƥ, ZIN]; 29–30.viii.1953 [3 Ƥ, ZIN]; 7.ix.1953 [9 Ƥ, ZIN]. Khulo , 2.vii.1953, V.A. Trjapitzin [1 Ƥ, ZIN]. GERMANY. BADEN-WÜRTTEMBERG, Schliffkopf ( Schwarzwald , 1100 m), 27.viii.1959 [1 Ƥ, ISNB]. BAVARIA: Erlangen, H.- J. Stammer [2 Ƥ, NHMW]. Veitshöchheim , 22.vii.2003, S.V. Triapitsyn [13 Ƥ, UCRC]. MECKLENBURG-WESTERN POMERANIA, Jettchens Hof (near Malchin ): viii.1935, H.- J. Stammer [2 Ƥ, NHMW]; viii.1936, H.- J. Stammer [3 Ƥ, NHMW]. NORTH RHINEWESTPHALIA: Aachen : [2 Ƥ, 2 3, NHMW] (det. by A. Foerster as “ G. littoralis ”); no other data [1 Ƥ, NHMW]. [No locality indicated], ? Aachen [2 Ƥ, 1 3, NHMW] (both females det. by A. Foerster, one of them as “ G. littoralis ”, the male misidentified by him as “ longicornis ”). Arloff, 17.iii.1963, M. Boness [1 3, NHMW]. Burscheid , vii–viii.1959, M. Boness [3 Ƥ, NHMW]. Cologne , M. Boness: 4.x.1962 [1 Ƥ, NHMW]; 28.viii.1963 [1 Ƥ, NHMW]. Leverkusen , M. Boness: 25.vii–3.viii.1966 [1 Ƥ, NHMW]; 1.ix.1966 [2 Ƥ, NHMW]; 5.ix.1966 [2 Ƥ, NHMW]; 12–16.ix.1966 [1 Ƥ, NHMW]; 30.ix.1966 [1 Ƥ, NHMW]. GREECE. CENTRAL MACEDONIA, Lake Kerkini: Ecotourism site, 41°08’15.6’’N 23°13’01.2’’E, 65 m, G. Ramel: 9–15.v.2006 [2 Ƥ, UCRC]; 16– 22.v.2006 [1 Ƥ, UCRC]; 23–29.v.2006 [1 3, UCRC]; 30.v–5.vi.2006 [3 Ƥ, UCRC]; 13–19.vi.2006 [2 Ƥ, 13, UCRC]; 20–26.vi.2006 [5 Ƥ, BMNH, UCRC]. Kerkini Lake site, 41°09’06.5’’N 23°11’55.0’’E, 75 m, 11– 17.iv.2005, G. Ramel [3 Ƥ, UCRC]. Kerkini Marsh , 41°13’32.8’’N 23°05’04.2’’E, 45 m, G. Ramel: 28.iii– 3.iv.2007 [1 Ƥ, 1 3, UCRC]; 4–10.iv.2007 [2 Ƥ, UCRC]; 11–17.iv.2007 [1 Ƥ, UCRC]; 18–24.iv.2007 [3 Ƥ, UCRC]; 25.iv–1.v.2007 [2 Ƥ, UCRC]. Procom site, 41°22’38.1’’N 23°21’58.8’’E, 60 m, 20–26.vi.2007, G. Ramel [1 Ƥ, UCRC]. Pumping station, 41°12’48.7’’N 23°06’11.9’’E, 40 m, G. Ramel: 13–19.vi.2007 [2 Ƥ, UCRC]; 2– 8.v.2007 [3 3, UCRC]. PELOPONNESE, 5 km S of Monemvasia , 27.xi.1983, G. Christensen [1 Ƥ, ZMUC]. THESSALIA, Larissa Prefecture, Ayiokambos , 39°43’N 22°52’E, 20 m, 7.iii.2001, A. Kapranas [5 Ƥ, UCRC]. HUNGARY. BÁCS-KISKUN: Gara , 46.035°N 19.020°E, 87 m, 26–30.vi.2009, I. Mikó [1 Ƥ, UCRC]. Kalocsa , J. Erdös: 26.ix.1942 [1 Ƥ, NHMW / HNHM]; 3.v.1945 [1 Ƥ, NHMW / HNHM]. Tompa , J. Erdös: 18.ix.1948 [1 Ƥ, NHMW / HNHM]; 16.iii.1949 [1 Ƥ, NHMW / HNHM]. PEST, Szigetszentmiklós , ix.1911, L. Biró [1 3, NHMW / HNHM]. VAS, W of Köszeg , 47°23’09’’N 16°31’19’’E, 355 m, 16–20.vi.2009, I. Mikó [2 Ƥ, UCRC]. ITALY. CAMPANIA: Avellino Prov.: Montemarano , 40°54.235’N 15°00.435’E, 760 m, 6.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [2 Ƥ, UCRC]. Piana del Dragone , 40°52.553’N 14°58.375’E, 760 m, 6.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [1 Ƥ, UCRC]. Piana Lacerno , 40°49.049’N 15°05.904’E, 1080 m, 6.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [1 Ƥ, UCRC]. Benevento Prov., 1.8 km E of Faicchio , 41°16.329’N 14°29.884’E, 210 m, 7.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [2 Ƥ, UCRC]. Caserta Prov.: 8.5 km E of Capriati a Volturno , 41°28.35’N 14°11.84’E, 850 m, 8.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [1 Ƥ, UCRC]. SE end of Lago del Matese , 41°24.411’N 14°24.800’E, 1050 m, 8.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [1 Ƥ, UCRC]. LAZIO: Roma Prov.: Bosco di Manziana , 42°07.392’N 12°07.314’E, 400 m, 9.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [1 Ƥ, UCRC]. Caldara di Manziana , 42°05.607’N 12°05.906’E, 305 m, 9–10.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [13 Ƥ, UCRC]. Castelporziano Presidential Estate : coastal dunes in N corner, 41°41.954’N 12°21.060’E, 3 m, 12.vi.2003, J. Munro, A. Owen [2 Ƥ, UCRC]. Fosso di Trafusina , 41°46.670’N 12°24.751’E, 30 m, 11–12.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [21 Ƥ, UCRC]. Ponte Guidoni , 41°45.415’N 12°23.851’E, 80 m, 11.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [2 Ƥ, UCRC]. Mignone River near Rota , 42°09.197’N 12°00.605’E, 150 m, 9.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [11 Ƥ, UCRC]. 0.8 km W of Sasso , 42°02.209’N 12°02.209’E, 264 m, 9– 10.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [21 Ƥ, UCRC]. Viterbo Prov.: Ponte San Pietro , 42°31.669’N 11°36.353’E, 75 m, 10.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [12 Ƥ, UCRC]. Roccaccia , 42°19.809’N 11°45.671’E, 125 m, 10.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [10 Ƥ, UCRC]. San Giovenale, Vesca Creek , 42°13’34’’N 12°00’02’’E, 9.vi.2003, J. Munro et al. [1 Ƥ, UCRC]. MOLISE, Campobasso Prov., 2.5 km SW of Guardiaregia , 41°26.322’N 14°32.635’E, 860 m, 7.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [1 Ƥ, UCRC]. SICILY, Palermo, University of Palermo garden, 38°06’27.2’’N 13°21’02.4’’E, 41–43 m, 13–14.viii.2009, S.V. Triapitsyn [3 Ƥ, UCRC]. KYRGYZSTAN. CHUY: Karagajly-Bulak, 9 km W Ak-Tyuz , 42°52’47’’N 76°02’13’’E, 2180–3400 m, 26.vii.2000, C.H. Dietrich [2 Ƥ, UCRC]. Kashka-Suu Ravine, ca. 32 km S of Bishkek , 42°38’50’’N 74°30’50’’E, 1759 m, 12.viii.1998, C.H. Dietrich [1 Ƥ, UCRC]. Suusamyr Valley, W side of Kichi-Korumdy River , 42°13’28’’N 73°41’31’’E, 2291 m, C.H. Dietrich: 16.viii.1998 [2 Ƥ, UCRC]; 14.vi.1999 [3 Ƥ, 1 3, UCRC]. 10 km N of Telek , 40°31’20’’N 74°03’55’’E, 570 m, C.H. Dietrich: 13.viii.1998 [1 Ƥ, 1 3, UCRC]; 11.vi.1999 [3 Ƥ, UCRC]. DZHALAL-ABAD: 7 km ESE of Ak-Tash River , 40°41’31’’N 70°42’07’’E, 1390 m, 21.vi.1999, C.H. Dietrich [1 Ƥ, UCRC]. Chandalash River , 41°44’19’’N 70°52’22’’E, 1630 m, 20–21.vi.1999, C.H. Dietrich [18 Ƥ, 1 3, UCRC]. 10 km W of Dzhalal-abad, 40°51’57’’N 72°53’35’’E, 27.vi.1999, C.H. Dietrich [1 Ƥ, UCRC]. Kara-Kysmak Ravine , 42°06’49’’N 71°33’28’’E, 2500 m, 18.vi.1999, C.H. Dietrich [8 Ƥ, 1 3, UCRC]. Near jct. Kara-Kysmak and Chatkal Rivers , 42°04’00’’N 71°35’41’’E, 2240 m, 18–19.vi.1999, C.H. Dietrich [8 Ƥ, UCRC]. 18 km WSW of Kazarman , 41°22’01’’N 73°48’37’’E, 1550 m, 15.vii.2000, C.H. Dietrich [4 Ƥ, UCRC]. Jct. of Kokerim and Kugart Rivers , 41°26’32’’N 73°57’07’’E, 1247 m, C.H. Dietrich: 28.viii.1998 [1 Ƥ, UCRC]; 29.vi.1999 [3 Ƥ, UCRC]. Teke-Uyuk Ravine , 41°29’12’’N 74°35’50’’E, 1850 m, 30.vi.1999, C.H. Dietrich [1 Ƥ, UCRC]. ISSYKKUL: Barskaun Ravine : 42°07’13’’N 77°35’46’’E, 1890 m, 5.vii.1999, C.H. Dietrich [1 Ƥ, UCRC]. 16 km of S Barskaun , 42°02’47’’N 77°35’52’’E, 2320 m, C.H. Dietrich: 4.ix.1998 [1 Ƥ, UCRC]; 15.vii.1999 [1 Ƥ, UCRC]. S Shore of Lake Issyk-kul, 10 km E of Kadzhi-Saj , 42°10’33’’N 77°18’55’’E, 1675 m, C.H. Dietrich: 5.ix.1998 [2 Ƥ, UCRC]; 2–6.vii.1999 [15 Ƥ, 3 3, UCRC]; 22.vii.2000 [1 Ƥ, UCRC]. 5 km W of Karasaj , 42°33’58’’N 77°51’50’’E, 3300 m, 3.ix.1998, C.H. Dietrich [1 Ƥ, UCRC]. NARYN: Alabuga River, 25 km W of Baetov , 41°17’47’’N 74°39’20’’E, 1700 m, C.H. Dietrich: 29.viii.1998 [1 Ƥ, 3 3, UCRC]; 16.vii.2000 [8 Ƥ, INHS, UCRC]. Dzhaman-Davan River near Saz , 41°17’31’’N 74°42’29’’E, 1826 m, 29.viii.1998, C.H. Dietrich [4 Ƥ, INHS, UCRC]. Moldo-Too Ridge, E of Kara-Go Pass , 41°30’22’’N 74°44’11’’E, 2260 m, 30.vi.1999, C.H. Dietrich [3 Ƥ, UCRC]. OSH: Gultcha Ravine, 50 km SSW of Gultcha , 39°52’17’’N 73°21’26’’E, 2530 m, 6.vii.2000, C.H. Dietrich [2 Ƥ, 1 3, UCRC]. Karakuldzha, Lajsu Ravine , 40°31’20’’N 73°37’10’’E, 1815 m, C.H. Dietrich: 25.vi.1998 [3 Ƥ, INHS, UCRC]; 25.vi.1999 [13 Ƥ, 9 3, INHS, UCRC]. TALAS: Near Boo-Terek , 42°35’15’’N 71°45’49’’E, 1000 m, 15.vi.1999, C.H. Dietrich [16 Ƥ, 1 3, INHS, UCRC]. Kara Buura Ravine, 20 km S Kyzyl-Adyr , 42°26’23’’N 71°33’16’’E, 1300 m, 15.vi.1999, C.H. Dietrich [2 Ƥ, UCRC]. Talas Valley, Kirov Reservoir , 42°39’19’’N 71°35’44’’E, 930 m, 15.vi.1999, C.H. Dietrich [4 Ƥ, UCRC]. 18 km WSW of Taldy Bulak , 42°26’31’’N 72°49’12’’E, 1930 m, 15–16.vi.1999, C.H. Dietrich [9 Ƥ, 1 3, UCRC]. MONGOLIA. ÖMNÖGOVI: Naran Bulag , 43°27’N 100°27’E, 1405 m, 17–20.vii.1994, J. Carpenter [26 Ƥ, UCRC]. Ukhaa Tolgod , 43°31’N 101°32’E, 1517 m, 6–16.vii.1994, J. Carpenter et al. [1 Ƥ, UCRC]. NETHERLANDS. LIMBURG: Epen , 23.v.1959, B. Petersen [1 3, ZMUC]. Valkenburg , W. Soyka: x.1931 [1 3, NHMW]; 28.vi.1932 [1 3, NHMW]; 28.vi.1932 (on window, Ignatiuskolleg ) [1 Ƥ, NHMW]. POLAND. LOWER SILESIA: Lenartowice (labeled as “ Leonhardwitz , Schlesien ”), vii.1934, W. Soyka [1 Ƥ, NHMW]. Wrocław : 28.viii.1933 [1 Ƥ, NHMW]; 30.iv.1978, G. Gordh, W. Pulawski [1 Ƥ, UCRC]. Near Wrocław (various localities with old, pre-1937 German names in former Schlesien, Germany): i. ix.1933, H.- J. Stammer [1 Ƥ, NHMW]; vii.1934, W. Soyka [6 Ƥ, NHMW]; viii.1934, W. Soyka [1 Ƥ, NHMW]. LUBUSZ, Sława Lake (near Sława; on the original label as “Schlawa-See, Schlesien ”, now Jezioro Sławskie ), 22.vii.1934, H.- J. Stammer [1 Ƥ, 1 3, ISNB; 10 Ƥ, 2 3, NHMW]. REPUBLIC OF KOREA. GYEONGGI-DO, Suwon-si, Seodun-dong: Seoul National University : 10.x.1997, J.- Y. Choi [1 Ƥ, UCRC]; 15.ix.2001, J.- W. Kim [1 Ƥ, UCRC]. Yeogisan , 7x.1997, J.- Y. Choi [6 Ƥ, UCRC]. ROMANIA. CLUJ, Apahida , 26.vii.1943, J. Erdös [2 Ƥ, NHMW / HNHM]. RUSSIA. KARACHAYCHERKESSIA (KARACHAY-CHERKESS REPUBLIC), Karachayevsk, Dombay , ca. 1650 m, 19.vii.2003, E.V. Khomchenko [2 Ƥ, 1 3, UCRC]. KRASNODARSKIY KRAY, Krasnodar, All-Russian Research Institute of Biological Plant Protection , V.V. Kostjukov: 10.viii–4.ix.2001 [35 Ƥ, 17 3, UCRC, ZIN]; 23.vii.2002 [3 Ƥ, UCRC]; 31.viii.2003, [61 Ƥ, UCRC, ZIN]. LENINGRADSKAYA OBLAST’, 69-km Railway Station (near Sosnovo ), 6.ix.1987, V.A. Trjapitzin [1 Ƥ, ZIN]. MOSCOVSKAYA OBLAST’: Noginskiy rayon, Fryazevo : 17.vi.1996, S.V. Triapitsyn [1 Ƥ, UCRC]; 2–15.vi.2000, M.E. Tretiakov [3 Ƥ, UCRC]; 25.vi–2.vii.2000, M.E. Tretiakov [7 Ƥ, 1 3, UCRC]; 3–8.vii.2000, M.E. Tretiakov [4 Ƥ, UCRC]; 7–15.vii.2000, M.E. Tretiakov [14 Ƥ, UCRC, ZIN]; 24.vii–14.viii.2000, M.E. Tretiakov [10 Ƥ, UCRC, ZIN]; 15–25.viii.2000, M.E. Tretiakov [14 Ƥ, UCRC, ZIN]; 25–31.viii.2000, M.E. Tretiakov [5 Ƥ, UCRC]; 8.vii.2001, M.E. Tretiakov [1 Ƥ, UCRC]; 20.vii.2001, M.E. Tretiakov [2 Ƥ, UCRC]; 13.viii.2001, M.E. Tretiakov [4 Ƥ, UCRC]; 13.vii.2002, S.V. Triapitsyn [1 Ƥ, UCRC]; 14.vii.2002, M.E. Tretiakov [6 Ƥ, UCRC]; 25.vii.2002 [32 Ƥ, UCRC, ZIN]; 1.viii.2002, M.E. Tretiakov [11 Ƥ, UCRC]. Pushkinskiy rayon, Pushkino , Mamontovka, E. Ya. Shuvakhina : 10–20.vii.2000 [3 Ƥ, UCRC, ZIN]; 20–31.vii.2000 [1 Ƥ, UCRC]; 20–31.viii.2000 [2 Ƥ, UCRC]. PRIMORSKIY KRAY: Terneyskiy rayon, Mel’nichnyi, M.V. Michailovskaya: 1–5.vi.2001 [3 Ƥ, IBPV, UCRC]; 26.vi.2001 [9 Ƥ, IBPV, UCRC]; 29.vi.–1.vii.2001 [3 Ƥ, UCRC, ZIN]. Ussuriyskiy rayon, Gornotayozhnoye , M.V. Michailovskaya: 27.v.1999 [1 Ƥ, UCRC]; 6.vi.1999 [1 Ƥ, 1 3, UCRC]; 10–14.vi.1999 [2 Ƥ, UCRC]; 17–27.vi.1999 [1 Ƥ, 1 3, UCRC]; 11– 14.vii.1999 [4 Ƥ, IBPV, UCRC]; 19–22.vii.1999 [2 Ƥ, UCRC]; 24.vii–1.viii.1999 [8 Ƥ, IBPV, UCRC, ZIN]; 1– 4.viii.1999 [2 Ƥ, 1 3, UCRC]; 5–11.viii.1999 [5 Ƥ, UCRC]; 12–17.viii.1999 [19 Ƥ, IBPV, UCRC, ZIN]; 22– 28.viii.1999 [1 Ƥ, UCRC]; 28.viii–5.ix.1999 [5 Ƥ, UCRC]; 6–14.ix.1999 [8 Ƥ, UCRC]; 15–26.ix.1999 [2 Ƥ, UCRC]; 21–28.x.1999 [1 Ƥ, UCRC]; 15–31.v.2000 [2 Ƥ, UCRC]; 11–21.vi.2000 [9 Ƥ, UCRC]; 21–30.vi.2000 [6 Ƥ, 1 3, IBPV]; 1–10.vii.2000 [6 Ƥ, 1 3, ZIN]; 10–20.vii.2000 [3 Ƥ, 1 3, UCRC]; 1–10.viii.2000 [11 Ƥ, 1 3, UCRC]; 21–26.viii.2000 [4 Ƥ, 1 3, UCRC]; 5–8.x.2000 [2 Ƥ, UCRC]; 9–12.x.2000 [3 Ƥ, UCRC]; 17.viii.2001 [11 Ƥ, UCRC]; 17–31.viii.2001 [2 Ƥ, UCRC]; ix–xi.2001 [5 Ƥ, UCRC]; 1–10.x.2001 [3 Ƥ, UCRC]; 1–10.v.2002 [1 Ƥ, UCRC]; 10–20.v.2002 [4 Ƥ, UCRC]; 20–31.v.2002 [3 Ƥ, UCRC]; 1–10.vi.2002 [5 Ƥ, UCRC]; 1–11.ix.2002 [1 Ƥ, UCRC]; 24.ix–5.x.2002 [5 Ƥ, IBPV, UCRC, ZIN]. SAKHALINSKAYA OBLAST’, Sakhalin Island: 28 km W of Poronaysk , 4.viii.2001, D.J. Bennett [1 Ƥ, 1 3, CAS]. 6 km E of Sokol, near Belaya River , 16.viii.2001, D.J. Bennett, T. Anderson [2 Ƥ, CAS]. SAMARSKAYA OBLAST’, Zhigulevskiy zapovednik ( Zhiguli National Park ), Strel’naya Mt., 17.vii.1985, V.A. Trjapitzin, E.Ya. Shuvakhina [1 Ƥ, ZIN]. STAVROPOL’SKIY KRAY: Achikulak , V.V. Kostjukov: 4–5.vi.2002 [4 Ƥ, UCRC]; 20.vi.2002 [1 Ƥ, UCRC]; 21.viii.2002 [1 Ƥ, UCRC]; 26.viii.2002 [31 Ƥ, UCRC, ZIN]. Apanasenkovskiy rayon, 15 km N of Kievka , “ Dundinskoye ” hunting establishment, 4.vii.2003, E.V. Khomchenko [5 Ƥ, 3 3, UCRC]. Georgievskiy rayon, Nezlobnenskiy (stanitsa Nezlobnaya ), 26.viii.2002, V.V. Kostjukov [1 Ƥ, UCRC]. Mikhaylovskoye, “ Aviator ” farm near Stavropol’ airport, 22.viii.2002, E.V. Khomchenko [10 Ƥ, 5 3, UCRC]. Novozavedennoye, 5.v.2002, V.V. Kostjukov [1 Ƥ, UCRC]. Prietokskiy , V.V. Kostjukov: 29.viii.2002 [3 Ƥ, UCRC]; 7.ix.2002 [24 Ƥ, 5 3, UCRC]; 13.vii.2003 [22 Ƥ, 1 3, UCRC]; 14.vii.2003 [6 Ƥ, 2 3, UCRC]; 27.vii.2003 [1 Ƥ, UCRC]; 7.viii.2003 [15 Ƥ, UCRC]; 12.viii.2003 [36 Ƥ, UCRC, ZIN]; 14.viii.2003 [1 Ƥ, UCRC]. Stavropol’, Russkiy Les, “ Besputskaya Polyana ” Botanical Sanctuary , 18.v.2003, E.V. Khomchenko [9 Ƥ, 1 3, UCRC]. TAMBOVSKAYA OBLAST’, Inzhavinskiy rayon, Talinka (7 km S of Pavlovka ), 26–27.v.2000, M.E. Tretiakov [2 Ƥ, UCRC]. SERBIA. Mt. Avala (near Belgrade), H.- J. Stammer: 28.viii.1934 [1 Ƥ, NHMW]; viii.1934 [1 Ƥ, NHMW]. SLOVAKIA. BRATISLAVA, Jurský Šúr Nature Reserve , 48°14’03’’N 17°12’47’’E, 133 m, 8.viii.2008, B.V. Brown (alder forest) [1 Ƥ, UCRC]. SPAIN. ANDALUCÍA, Fuengirola, 18.viii.1952, J.K. Holloway (on Heliotropium sp.) [5 Ƥ, EMEC] (according to the unpublished University of California, Berkeley ( UCB) quarantine records [Kent M. Daane, personal communication], these emerged from eggs of Neoaliturus (Circulifer) tenellus ( Baker) and were received 25.viii. 1952 in UCB quarantine under their Shipper/Receiver ( SR) No . 52-29). MADRID, Aranjuez, 3.viii.1952, J.K. Holloway (on Portulaca sp.) [3 Ƥ, 1 3, EMEC] (according to the unpublished UCB quarantine records, these were reared from eggs of N. tenellus and received 11.viii. 1952 in UCB quarantine under SR No . 52-23). SWITZERLAND. ST. GALLEN, Rheintal , 1934, H. Kutter (in pea field) [1 Ƥ, NHMW]. TURKMENISTAN. AHAL: Babadurmaz , 18–20.vi.1997, V.V. Berezovskiy [1 Ƥ, UCRC]. Central Kopet Dag Mts., Chuli Canyon , 11.vi.1992, S.V. Triapitsyn [1 Ƥ, UCRC]. Enev , S.N. Myartseva (in beet field): 5.vii.1993 [3 Ƥ, UCRC]; 14.vii.1993 [1 Ƥ, UCRC]. [Old] Nisa (in fortress ruins), S.V. Triapitsyn: 9.vi.1992 [1 Ƥ, UCRC]; 15.vi.1992 (emerged 30.vi. 1992 in University of California, Riverside , quarantine from Atriplex sp. plant material, S & R # 92- 26-8). ASHGABAT: Ashgabat, Karakum Canal bank, 10.vi.1992, V.A. Trjapitzin [2 Ƥ, UCRC]. UK. ENGLAND: Cheshire Co., Rostherne , 20.viii.1933, H. Britten [1 Ƥ, MMUE]. Cumbria Co.: Newton Reigny Moss , 5.ix.1945, H. Britten [1 Ƥ, MMUE]. Skirwith, 23.ix.1933, H. Britten [1 Ƥ, MMUE]. Dorset Co., Bournemouth , 8.x.1981, S.G.C. Brown [3 females, BMNH]. East Riding of Yorkshire Co., Allerthorpe Common, 12.ix.1950, W.D. Hincks [1 Ƥ, MMUE] (misidentified as Lymaenon ater ( Foerster) by W.D. Hincks). Greater Manchester Co., Metropolitan Borough of Trafford , Dunham Massey , Dunham Park ( S of Dunham Town ), 27.ix.1947, H. Britten [1 Ƥ, MMUE]. London Borough of Richmond upon Thames, Richmond Park , J.S. Noyes: 28.vii.1995 [1 Ƥ, 1 3, CNCI]; 15.viii.1997 [1 Ƥ, CNCI]. Hampshire Co., Matley Bog , 15.vi.1952, W.D. Hincks [1 Ƥ, MMUE]. Merseyside Co., Freshfield , 19.ix.1959, A. Brindle [1 Ƥ, MMUE]. North Yorkshire Co., Thornton-le-Dale ( Thornton Dale ), 12.ix.1959, W.D. Hincks (on window) [1 Ƥ, MMUE]. Surrey Co.: Barnes Common, 8.vii.1995, J.S. Noyes [2 Ƥ, CNCI]. Dorking , White Downs , 21.ix.1986, J.S. Noyes [8 Ƥ, CNCI]. WALES: Bridgend Co. Borough, Kenfig Pool National Nature Reserve , J.S. Noyes: 6.viii.1988 [2 Ƥ, CNCI]; 31.viii.1995 [3 Ƥ, CNCI]. City and Co. of Swansea: Oxwich National Nature Reserve , 5.viii.1994, J.S. Noyes [1 Ƥ, CNCI]. Whiteford Burrows National Nature Reserve , 2.viii.1988, J.S. Noyes [3 Ƥ, CNCI]. Vale of Glamorgan Co. Borough, Pendoylan , Hensol , Llanerch Vineyard , 9.ix.1999, S.V. Triapitsyn [2 Ƥ, UCRC].

Extralimital records. ARGENTINA. BUENOS AIRES, Luján, Universidad Nacional de Luján , 34°35'07"S 59°04'45"W, 32 m, C. Coviella: 3.xi.2006 [2 Ƥ, UCRC]; 22.xii.2006 [1 Ƥ, UCRC]; 2.ii.2007 [1 Ƥ, UCRC]. BERMUDA ( BERMUDA ISLANDS). Bermuda Island, Southampton Parish , Munro Beach Cottages , 32°15’34’’N 64°52’39’’W, 2–16.iv.2002, J. Munro [1 Ƥ, UCRC]. CAMBODIA. SIEM REAP, Angkor, Preah Khan Temple , 12.v.2006, O. Yothin [1 Ƥ, ISNB]. CANADA. NORTHWEST TERRITORIES, km 240 Dempster Highway , 68.23°N 133.32°W, 10.viii.2000, B.V. Brown, D. Currie [3 Ƥ, 2 3, UCRC]. SASKATCHEWAN, Eb’s Trails (14 km N of Duck Lake ), 16.vi.2004, M. Yoder [1 Ƥ, UCRC]. CHILE. REGIÓN IX, Rancho Flor del Lago (11 km NE of Villarica ), 39°12.5’S 72°08.1’W, 290 m, 12.xii.2001, E. Arias (canopy fogging of Nothophagus obliqua ) [1 Ƥ, UCDC]. GREENLAND. Kap Farvel-området, Pamiagdluk , Anordliuitsoq , 29.viii.1970, J. Bocher [1 Ƥ, CNCI]. Kuussuaq , 19–23.viii.1982, P. Nielsen [1 Ƥ, CNCI]. INDIA. [NATIONAL CAPITAL TERRITORY OF] DELHI, New Delhi, Indian Agricultural Research Institute : 28°37’51’’N 77°09’50’’E, 220 m, 5–6.xi.2003, J.M. Heraty [4 Ƥ, UCRC]; 28°38’00’’N 77°09’53’’E, 227 m, 3.xi.2003, J.M. Heraty [1 Ƥ, UCRC]. KARNATAKA, Mudigere , 13°07’09’’N 75°37’41’’E, 994 m, 26.xi.2003, J.M. Heraty [1 Ƥ, UCRC]. MAURITIUS. Rodrigues Island, Canyon Tyeul , 19°44’32’’S 63°22’30’E, 35 m, 14–28.ix.2007, A. Meunier [2 Ƥ, CAS]. MEXICO. No locality indicated, intercepted with plants by customs (Fouts) at Nogales, Santa Cruz Co., Arizona, USA, 12.ii.1955 [1 Ƥ, USNM] (identified as Lymaenon brunneus by B.D. Burks). USA. CALIFORNIA: Alameda Co., Albany, University of California, Berkeley ( UCB) insectary/quarantine: 29.viii.1952 (“ Reared in quarantine. Ex Circulifer tenellus on sugar beet. Stock from Spain ”) [1 Ƥ, EMEC]; 31.vii.1953, R.L. Doutt (“1st generation ex C. tenellus Lymaenon B ”) [6 Ƥ, EMEC]; 11.xi.1953, R.L. Doutt (insectary culture of Lymaenon “ B ” from N. tenellus eggs on sugar beet) [14 Ƥ, 6 3, EMEC]; 27.i.1954, G.L. Finney (insectary culture of Lymaenon “ B ” from N. tenellus eggs on sugar beet) [10 Ƥ, 3 3, EMEC]; 10.i.1955, G.L. Finney (insectary culture, “Sample from cage of Lymaenon “ B ”, ex eggs of Circulifer tenellus on sugar beet ”) [10 Ƥ, EMEC]; originally from: SPAIN. [According to the unpublished quarantine records at UCB ( Kent M. Daane , personal communication), the originators of the successfully established colonies of Lymaenon sp. “A” and “B” were collected by J.K. Holloway in Spain likely in various localities during 1952 and 1953 (the host was N. tenellus on various plant material) and received in UCB quarantine under several Shipper/Receiver numbers, but the exact localities and collecting dates for the original stocks of these colonies are now impossible to figure out]. Kern Co., Ave. A & 100th St. W (near North Antelope Valley and Rosamond), 26.iv.2000, I.M. Bayoun (from leafhopper eggs on Salsola sp.) [1 Ƥ, UCRC]. Riverside Co., Hemet (along Soboba St. ), 9.vi.1999, I.M. Bayoun (from sentinel eggs of N. tenellus in sugar beet leaves) [2 Ƥ, UCRC]. San Bernardino Co., S of Barton Flats , 34°09’42’’N 116°52’23’’W, 2090 m, 19– 26.vi.2007, F. Reuter [13 Ƥ, 1 3, UCRC]. Stanislaus Co., Frank Raines Regional Park, Ranger Station , 37°25.294’N 121°22.666’W, 350 m, 20.viii–18.ix.2011, R.L. Zuparko [1 Ƥ, EMEC]. ILLINOIS: Champaign Co., Urbana , 1.vii.1910 [5 Ƥ, USNM] (identified by A.A. Girault as G. anthonomi). Marion Co., Centralia , 25.viii.1909 [1 Ƥ, INHS] (identified by A.A. Girault as G. anthonomi). MINNESOTA, Clay Co., ca. 4 mi SEE of Glyndon, Bluestem Prairie , 46.85521°N 96.47353°W, 1–3.vi.2004, R.A. Rakitov [1 Ƥ, 1 3, UCRC]. OREGON: Jackson Co., 2 mi S of Exit #6 on I5, 42.0477°N 122.6052°W, 4.v.2005, R.A. Rakitov [1 Ƥ, UCRC]. Lincoln Co., Waldport , 44°25’36’’N 124°03’05’’W, 56 m, 16–31.vii.2010, J.D. Pinto [1 Ƥ, UCRC].

Distribution. PALAEARCTIC: Armenia *, Austria ( Kirchner 1867), Belgium, Bulgaria ( Hellén 1974; Donev 1986 [also as G. effusi ], 1987 [also as G. effusi ], 1988d [also as G. effusi and G. paludis ], 1988e [also as G. effusi ], 1990 [also as G. paludis ], 2005), China *, Czech Republic, Cyprus *, Denmark, Finland ( Hellén 1974), France *, Georgia *, Germany, Greece ( Donev 1985a [as Lymaenon effusi and L. paludis ], 1988c [also as G. arduennae and G. effusi ], 2003, 2005), Hungary *, Iran ( Fallahzadeh & Huber 2011), Ireland ( Graham 1973 [as G. paludis ]; Matthews 1986), Italy *, Japan ( Sahad & Hirashima 1984), Kyrgyzstan *, Macedonia ( Donev 1988 a, 2005), Mongolia *, Netherlands ( Soyka 1946), Norway ( Mathot 1969; Hellén 1974), Poland ( Soyka 1946), Republic of Korea *, Romania ( Radu & Boţoc 1960; Pricop 2009b, 2010a, c), Russia ( Hellén 1974), Serbia ( Donev 1985b, 1988b [as G. paludis ]), Slovakia, Spain ( Arnaldos et al. 2003; Baquero & Jordana 2003), Sweden, Turkey ( Donev 2001, 2005), Turkmenistan *, UK ( England, Northern Ireland, Scotland, Wales). AFROTROPICAL*: Mauritius *. NEARCTIC: Canada *, Bermuda * ( Bermuda Islands), Greenland *, Mexico, and USA ( Triapitsyn et al. 2010). NEOTROPICAL: Argentina, Bahamas, Brazil, and Chile ( Triapitsyn et al. 2010). ORIENTAL*: Cambodia *, and India *. Sahad & Hirashima (1984) listed G. litoralis from Australia (most likely by mistake), but without references to any specimens or published records; Lin et al. (2007) did not include it in their list of the Australian species of Gonatocerus .

This is arguably the most common and widespread Palaearctic species of Gonatocerus , so that it is just a matter of time until G. litoralis would be recorded from most if not all the European countries. Bayoun et al. (2008) reported specimens tentatively identified as G. litoralis from California, USA, and here I confirm that identification and further document its presence in the Nearctic region in addition to the records provided by Triapitsyn et al. (2010).

Redescription. FEMALE (holotypes of Lymaenon arduennae , L. paludis and L. rhacodes , and non-type specimens from the Palaearctic region). Body length 600–1000 µm (dry-mounted specimens). Body generally brown to dark brown except mesosoma sometimes light brown along edges of midlobe of mesoscutum and scutellum in teneral adults, gaster mostly yellow-brown or light brown (particularly basally) to brown; appendages light brown to brown.

Antenna ( Figs 144, 145 ) with radicle 3.0–3.8× as long as wide, 0.31–0.38× total length of scape, rest of scape 3.2–3.8× as long as wide, faintly striate; pedicel longer than F1; F1–F4 subequal in length, shorter than following funicle segments; F5, F7, and F8 subequal in length, F6 a little shorter; F1–F4 without mps, typically mps on F5 (1), F7 (2), and F8 (2) but rarely F5 with 2 mps or sometimes (often in small specimens) without mps, F6 with 1 mps in a few, mostly large specimens (such as the holotype of Lymaenon rhacodes ), and occasionally F7 with just 1 mps; clava with 10 mps (4 in the middle and 6 subapical), 2.9–4.1× as long as wide, from a little shorter to a little longer than combined length of F6–F8.

Mesosoma ( Fig. 146 ). Propodeum ( Fig. 147 ) with faint submedian lines, the distance between them shorter anteriorly than posteriorly. Fore wing ( Figs 148, 150 ) 2.8–4.1× as long as wide; longest marginal seta 0.28– 0.52× maximum wing width. Fore wing disc almost hyaline (with a slight, uniform brownish tinge), almost bare behind submarginal vein except for a few setae behind its apex, more or less densely setose behind marginal vein and densely setose elsewhere. Hind wing ( Figs 149, 150 ) 17–24× as long as wide; disc setose and slightly infumate; longest marginal seta 2.3–4.0× maximum wing width.

Metasoma ( Fig. 146 ) a little longer than mesosoma. Petiole 1.6–2.1× as wide as long. Ovipositor occupying 0.6–0.8× length of gaster, not or barely exserted beyond its apex; ovipositor length: mesotibia length ratio 0.9–1.8:1.

MALE (non-type specimens from the Palaearctic region). Body length 600–800 µm (dry-mounted specimens). Similar to female except for normal sexually dimorphic features and the following. Antenna ( Fig. 151 ) with scape plus radicle 2.6–2.7× as long as wide, notably striate on lateral surface. Fore wing ( Fig. 152 ) 2.8–3.6× as long as wide. Genitalia as in Fig. 153 .

Diagnosis. Gonatocerus litoralis is a highly variable and thus difficult to diagnose species recognizable by the female antenna ( Figs 144, 145 ) typically having 1 mps on F5 and 2 mps on F7 and F8 but rarely with 2 mps on F5 or sometimes, often in small specimens, without mps on F5 and/or just 1 mps on F7, and without mps on F6 (rarely with 1 mps in large specimens), and 10 mps on the clava; a narrow to broad (2.8–4.1× as long as wide) fore wing ( Figs 148, 150 ) with a relatively densely setose area on the disc between the marginal vein and the cubital row of setae thus leaving no bare area, the ovipositor not or barely exserted beyond the gastral apex; and generally brown to dark brown body except for the gaster, which is mostly light brown (particularly basally) to brown.

Hosts. Aphrodes sp. ( Bakkendorf 1934) [as Acocephalus sp. for Lymaenon effusi ], Macrosteles sexnotatus (Fallén) ( Ahlberg 1925; Nikol’skaya 1952) [as Cicadula sexnotata Fallén for G. radiculatus ], Neoaliturus (Circulifer) tenellus ( Baker) ( Huffaker et al. 1954 [as Lymaenon “B”]; Bayoun et al. 2008) [as Circulifer tenellus ( Baker) ], and Zyginidia sohrab Zachvatkin ( Fallahzadeh & Huber 2011) (Cicadellidae) . Girault (1905) indicated that G. anthonomi had been supposedly bred from Anthonomus quadrigibbus Say ( Coleoptera : Curculionidae ) but that record was almost certainly erroneous. Also almost certainly incorrect were the records of G. brunneus (the identifications also need confirmation) from eggs of Microlarinus lypriformis (Wollaston) (Curculionidae) by Stegmaier (1973) as well as from Aphis pomi De Geer and Rhopalosiphum padi (Linnaeus) ( Hemiptera : Aphididae ), as listed by Peck (1963) and Thompson (1958), respectively.

Comments. Following Matthews (1986), I treat G. litoralis in the broad sense thus including specimens in which F5 of the female antenna lacks mps (like for instance in G. texanus ). However, there are also small specimens in the Palaearctic region listed below which often have relatively somewhat shorter funicle segments and lack mps on F1–F7 of the female antenna ( Figs 154 , 156 ) or sometimes have 1 mps on F7 only on one antenna in a few specimens from Mongolia, which otherwise fit G. litoralis well including the typical wings ( Fig. 155 ). Particularly, they bear 2 mps on F8 and 10 mps on the clava and are not conspecific with the Nearctic species G. pygmaeus because they have F3 as short as F2 or F4 and have no distinct bare area between the marginal vein and the cubital row of setae on the fore wing disc. Such specimens can only tentatively be attributed to G. litoralis and perhaps should rather be regarded as an undescribed taxon within the G. litoralis complex unless they turn out to be within the broader variability range of G. litoralis than we currently recognize. Describing them as a new species, however, without having at hand results of thorough morphometric studies and biological and molecular data would be counterproductive at this time: perhaps the loss of mps on both F5 and F7 of the female antenna could be attributed to parasitization of eggs of smaller hosts. Material examined: GREECE. CENTRAL MACEDONIA, Lake Kerkini, pumping station, 41°12’48.7’’N 23°06’11.9’’E, 40 m, 13–19.vi.2007, G. Ramel [1 Ƥ, UCRC]. EGYPT. [Governorate unknown], El-Alag, 31.v.1954, C.B. Huffaker, on Chenopodium sp., received 3.vi.1954 at UC Berkeley (UCB) quarantine (Albany, California, USA, SR # 54-17), presumed host Neoaliturus tenellus ( Baker) [1 Ƥ, 1 3, EMEC]. GIZA, Giza,? v.1954, C.B. Huffaker, “on Chenopodium sp. at Pyramids”, emerged 4.vi.1954 at UCB quarantine (Albany, California, USA, SR # 54-17), presumed host Neoaliturus tenellus ( Baker) [5 Ƥ, 3 3, EMEC]. MONGOLIA. ÖMNÖGOVI: Naran Bulag, 43°27’N 100°27’E, 1405 m, 17–20.vii.1994, J. Carpenter [3 Ƥ, UCRC]. Ukhaa Tolgod, 43°31’N 101°32’E, 1517 m, 6–16.vii.1994, J. Carpenter et al. [6 Ƥ, UCRC]. RUSSIA. STAVROPOL’SKIY KRAY, Prietokskiy, 12.viii.2003, V.V. Kostjukov [1 Ƥ, UCRC]. TURKMENISTAN. ASHGABAT, Ashgabat, near Kurtlinskoye vodokhranilishche [reservoir] shore, 10.vi.1992, S.V. Triapitsyn (emerged 25.vi. 1992 in University of California, Riverside, quarantine from Atriplex sp. plant material, S&R # 92-25-9). USA. CALIFORNIA, Riverside Co., Riverside, University of California at Riverside (UCR), Department of Entomology Insectary & Quarantine facility, viii.1995, I.M. Bayoun (quarantine culture #36-GL [= Gonatocerus sp. 1A of Walker et al. (1997)], first generation on eggs of Neoaliturus tenellus ( Baker) on sugar beet) [3 Ƥ, UCRC], originally from: IRAN. RAZAVI KHORASAN, Atar (near old Neyshabour-Mashhad road and Zax Road), 7.vii.1995, N. Zareh (from eggs of N. tenellus on Kochia sp. leaves, UCR Quarantine S&R #95-40-36A).

Identification of other voucher material of Walker et al. (1997) is very difficult. Specimens of their Gonatocerus sp. 2 (culture #23-GD) fit G. litoralis in every regard except setae on the fore wing ( Fig. 157 ) disc between the marginal vein and the cubital line of setae are somewhat sparser (yet unlike in G. ( Lymaenon ) sp. 1 from Iran leaving no distinct bare area) than in the typical G. litoralis , and therefore they only tentatively are identified as such also considering that their culture was uniparental. Material examined: USA. CALIFORNIA, Riverside Co., Riverside, UCR Insectary & Quarantine facility: viii.1995, I.M. Bayoun (quarantine culture, first generation on eggs of N. tenellus on sugar beet) [2 Ƥ, UCRC], originally from: IRAN. RAZAVI KHORASAN, 35 km from Mashhad on the road to Ghouchan, 4.vii.1995, N. Zareh (from eggs of N. tenellus on Artemisia sp. and Salsola sp. leaves, mixed culture from UCR Quarantine S&R #95-40-11 and S&R #95-40-23); ix.2000, I.M. Bayoun (insectary culture #23-GD on eggs of N. tenellus on sugar beet) [5 Ƥ, UCRC], of the same origin but collected only on Artemisia sp., S&R #95-40-23.

The specimens from Belgium and Germany in the H.R. Debauche collection at ISNB were misidentified by H.R. Debauche and G. Mathot as “ Lymaenon ater (Foerster) ”.

From the redescription and illustrations of G. ramakrishnai in Zeya & Hayat (1995) I conclude that this Indian species might be conspecific with G. litoralis although the indicated number of mps (6, although in reality it is most likely more than that) on the clava on the holotype female does not fit that of G. litoralis (10). I examined several females of G. litoralis collected by John M. Heraty at IARI, the likely type locality of G. ramakrishnai (or in the general area of it), and they fit both the original description and Zeya & Hayat’s redescription of Subba Rao & Kaur’s species except for the fact that F 5 in these specimens bears 2 mps whereas the holotype of Lymaenon ramakrishnai bears only 1 mps on F5 ( Zeya & Hayat 1995), but that is within the known variability range of G. litoralis . However, I am reluctant to synonymize G. ramakrishnai under G. litoralis until the holotype of the former species is examined.

The numerous slide-mounted voucher specimens of Lymaenon spp. “A” and “B” of the study by Huffaker et al. (1954) are stored in EMEC. Most specimens labeled by R.L. Doutt as Lymaenon “B” have the typical arrangement of mps on the female antenna (F5 bearing a mps) and belong to G. litoralis but some (particularly those collected 10.i.1955 and marked as “Sample from cage of Lymaenon “B””) lack one and also have 8 mps on the clava, and are identical to the specimens of Lymaenon “A” (= G. kazak ), thus indicating to a possible contamination of the colony at the later stages of the rearing campaign. According to Huffaker et al. (1954), both forms were imported from Spain (the culture originators were collected there by J.K. Holloway during 1952 and 1953) and then reared in the University of California, Berkeley quarantine/insectary at Albany, California on eggs of the beet leafhopper, N. tenellus [as Circulifer tenellus ]. 112,455 individuals of Lymaenon “A” (8 sites) and 400 individuals of Lymaenon “B” (1 site) were released in California during 1953, and in addition also likely a good number of Lymaenon “A” were sent to Arizona, Idaho, and Utah ( Huffaker et al. 1954). Post-release data on these parasitoids are very sketchy; Huffaker et al. (1954) only mentioned that specimens of Lymaenon “A” were recovered in two release sites of Fresno and Kern Counties soon after the releases but not at a later time; Clausen (1978) later stated that none of these introduced and released species became established. Difficulties in their identification probably contributed to the apparent failure of that project in the field, and as we know now G. litoralis occurs naturally in the USA; in California it was reared from eggs of N. tenellus ( Bayoun et al. 2008) .

Girault (1911) considered G. americanus to be very close in structure to his G. anthonomi , which is synonymized here under G. litoralis based on examination of its lectotype and other, non-type, specimens identified as G. anthonomi by A.A. Girault himself and also on Girault’s descriptions (1905, 1911). Based on the habitus and other morphological features that can be observed on the poorly preserved lectotype, G. americanus is almost certainly conspecific with G. litoralis , hence its proposed synonymy under the latter. I believe that even though it is impossible to know the distribution of mps on the missing funicle segments in the lectotype of Brues’ species, which is known only from this single specimen, the synonymy is warranted. Brues’ (1907) description of the female antenna, particularly of the proportions of the funicle segments, is consistent with the female antenna of G. litoralis . From my own collecting, I found G. litoralis to be by far the most common species of G. ( Lymaenon ) in Illinois, USA, where diversity of this subgenus seems to be quite limited, and I expect that to be the case also in the nearby Milwaukee, Wisconsin, the type locality of G. americanus .

Figures

FIGURE 136. Gonatocerus (Lymaenon) litoralis Ƥ (lectotype of Alaptus fuscus Förster): slide.

FIGURES 137, 138. Gonatocerus (Lymaenon) litoralis Ƥ (lectotype of G. americanus Brues): 137, slide; 138, habitus.

FIGURES 139 – 141. Gonatocerus (Lymaenon) litoralis (primary type slides of the synonyms from the Nearctic region): 139, G. texanus Girault; 140, G. illinoiensis Girault; 141, G. brunneus Girault.

FIGURES 142, 143. Gonatocerus (Lymaenon) litoralis Ƥ (holotype of Lymaenon arduennae Mathot): 142, slide; 143, habitus.

FIGURES 144 – 147. Gonatocerus (Lymaenon) litoralis Ƥ (144 — Hensol, Pendoylan, Wales; 145 – 147 — Kenfig Pool National Nature Reserve, Wales): 144 – 145, antennae; 146, mesosoma and metasoma; 147, dorsellum and propodeum.

FIGURES 148 – 150. Gonatocerus (Lymaenon) litoralis Ƥ (148 – 149 — Kenfig Pool National Nature Reserve, Wales; 150 — Fryazevo, Moskovskaya oblast’, Russia): 148, fore wing; 149, hind wing; 150, wings.

FIGURES 151 – 153. Gonatocerus (Lymaenon) litoralis 3 (Waterloo, Belgium): 151, antenna; 152, fore wing; 153, genitalia.

FIGURES 154, 155. Gonatocerus (Lymaenon)? litoralis or G. (Lymaenon) sp. near litoralis Ƥ (Lake Kerkini, Greece): 154, antenna; 155, wings.

FIGURES 156, 157. Gonatocerus (Lymaenon) spp. near litoralis Ƥ (156 — Gonatocerus sp. “ 1 A ” [culture # 36 - GL] of Walker et al. (1997); 157 — Gonatocerus sp. “ 2 ” [culture # 23 - GD] of Walker et al. (1997); both of Iran origin): 156, antenna; 157, fore wing.