Gonatocerus (Cosmocomoidea) ater Foerster, 1841 s

Triapitsyn, Serguei V., 2013, Review of Gonatocerus (Hymenoptera: Mymaridae) in the Palaearctic region, with notes on extralimital distributions, Zootaxa 3644 (1), pp. 1-178 : 119-136

Gonatocerus (Cosmocomoidea) ater Foerster, 1841 s . l.

( Figs 206–247 )

Gonatocerus ater Foerster 1841: 45 .

Type locality (of the lectotype designated here): Aachen, North Rhine-Westphalia, Germany.

Gonatocerus ater Foerster [or Förster]: Walker 1846: 53 (English translation of the original description); Kirchner 1867: 201 (catalog); Dalla Torre 1898: 429 (catalog); Hellén 1974: 13 (diagnosis, distribution); Matthews 1986: 221 –222 (synonymy, diagnosis, member of the ater species group ); Donev 1987: 73 –74 (distribution); Zeya & Hayat 1995: 70 –72 (synonymy, references on the Indian synonyms, redescription, host association, distribution) + 129, 137–138 (illustrations); Donev 1988d: 194 (distribution); Donev 1988e: 203 (distribution); Donev 2005: 383 (diagnosis, distribution); Pricop 2010b: 112 –113 (diagnosis, distribution), 114 (illustrations); Anwar & Zeya 2012: 52 (distribution in India); Zeya & Khan 2012: 57 (distribution).

Rachistus ater ( Foerster): Foerster 1847: 206–207 (diagnosis), 232 (list, distribution).

Gonatocerus pannonicus Soyka 1946: 39 .

Type locality: Hundsheim, Lower Austria, Austria. Synonymized under G. ater by Matthews 1986: 221.

Lymaenon ater (Förster) : Debauche 1948: 81 (list); Viggiani & Jesu 1988: 1023 (polytypic species based on the study of the type material).

Lymaenon schmitzi Debauche 1948: 86–88 , plate IX (illustrations).

Type locality: Heverlee (as Héverlé in the original description), Leuven, Flemish Brabant, Belgium. Synonymized under G. ater by Matthews 1986: 221.

Lymaenon indicus Subba Rao & Kaur 1959: 229–231 .

Holotype female [ IARI] (not examined). Type locality: New Delhi, Delhi, India. Synonymized under G. ater by Zeya & Hayat 1995: 70.

Lymaenon nigroides Narayanan & Subba Rao 1961: 656–657 .

Holotype female [lost from IARI ( Zeya & Hayat 1995)] (not examined). Type locality: Delhi, India. Synonymized under G. ater by Zeya & Hayat 1995: 70.

Lymaenon schmitzi Debauche : Boţoc 1962: 108 (short diagnosis); Viggiani 1969: 39 (member of the longicauda species group of Lymaenon ).

Lymaenon intermedius Boţoc 1962: 108–110 .

Type status not indicated, five females of the type series mentioned in the original description (although Matthews (1986) incorrectly designated a holotype), all of which were syntypes) [lost together with the entire M. Boţoc’s personal collection of the Romanian Mymaridae ( Pricop 2010b) ] (not examined). Type locality: “Aluviunea Someşului” (Someş [river] deposits—i.e., Someş River flood plain), near Cluj-Napoca, Cluj County, Romania. Synonymized under G. ater by Matthews 1986: 222 but treated as a valid species by Pricop (2010b), see “Comments” below.

Lymaenon empoascae Subba Rao 1966: 190 (illustrations [plate III, under Stethynium empoascae Subba Rao !]), 195–196. Holotype female [IARI] (not examined).

Type locality: “Jullunder” (Jalandhar), Punjab, India [or Jullundhur ( Subba Rao et al. 1968), although Zeya & Hayat (1995) indicated it as Jallunder (p. 70) but under “Specimens examined” (p. 72) they indicated it as “ INDIA: Delhi”]. Synonymized under G. ater by Zeya & Hayat 1995: 70.

Lymaenon populi Viggiani 1969: 40–44 .

Type locality: Rome, Lazio, Italy (Gennaro Viggiani, personal communication; the collecting locality of the holotype was not indicated either in the original description or on the two holotype slides). Synonymized under G. ater by Matthews 1986: 222.

Gonatocerus populi ( Viggiani): Graham 1973: 48 (record from Ireland); Pricop 2010b: 113 (possible synonym of G. intermedius (Boţoc) —see “Comments” below).

Gonatocerus empoascae ( Subba Rao): Subba Rao & Hayat 1983: 135 (catalog).

Gonatocerus indicus (Subba Rao & Kaur) : Subba Rao & Hayat 1983: 135 (catalog).

Gonatocerus nigriodes [sic] (Narayanan & Subba Rao): Subba Rao & Hayat 1983: 136 (catalog).

Lymaenon populi Viggiani : Viggiani 1988: 561 (illustration); Viggiani & Jesu 1988: 1023 (host association, valid species separate from G. ater ).

Lymaenon cicadellae Viggiani : Viggiani 1988: 563 (misspelling of L. populi ).

Gonatocerus schmitzi (Debauche) : Huber 1988: 50 ( ater species group , quite similar to the Nearctic species G. latipennis ).

Gonatocerus pannonicus Soyka : Pricop 2010b: 112 –113 (possible synonym of G. ater ), 114 (illustrations).

Gonatocerus intermedius (Boţoc) : Pricop 2010b: 113 (valid species, diagnosis based on non-type specimens, records from Romania—see “Comments” below), 114–116 (illustrations).

Type material examined. Gonatocerus ater Foerster : lectotype female [ NHMW], here designated to avoid the existing confusion regarding the type specimens of this species, on slide ( Fig. 206 ) labeled [in Soyka’s handwriting]: 1. “ Gonatocerus Ƥ ater ”; 2. [an empty red label]; 3. [Soyka’s slide number] “742”; 4. “ Gonat. ater Förster Type Aachen Förster Coll. G. Mayr In Canadab. 1943”. The lectotype was poorly remounted by Soyka from a minuten pin, with several parts of the specimen separate under the coverslip and one hind wing in the excess balsam not covered by it (the other hind wing is missing) . Matthews (1986: 221) and Pricop (2010b: 112–114) erroneously mentioned this specimen (on Soyka’s slide No. 742) as holotype of G. ater but that was not a valid lectotype designation (Article 74.5, [ ICZN] 1999) because the original description mentioned an unspecified number of female and male specimens, all of which are syntypes. Paralectotypes: 1 Ƥ [ NHMW] on slide labeled: 1. [in Soyka’s handwriting] “ Gonatocerus Ƥ foresteri [sic, Soyka’s manuscript name, in pencil]”; 2. [an empty red label]; 3. [Soyka’s slide number] “20”; 4. [partially in?Foerster’s handwriting, partially printed] “ Gon. ater Förster, Type”; 5. [printed] “Collect. G. Mayr”; 6. [in Soyka’s handwriting] “In Canadab. 1943”. This specimen ( Figs 290–292 ) actually belongs to G. (Cosmocomoidea) oxypygus Foerster. 1 3 [ NHMW] on slide labeled: 1. [in Soyka’s handwriting] “ Gonatocerus 3 ater Förster Type”; 2. [red] “Allo-Type” [incorrectly labeled as such by Soyka]; 3. [Soyka’s slide number] “18”; 4. [in Soyka’s handwriting] “ Gonatocerus ater Förster Type Coll. G. Mayr In Canadab. 1943”. This specimen ( Fig. 213 ) is mounted laterally, so the propodeal submedian carinae are not fully visible; it belongs to G. ( Cosmocomoidea ) and may be or may not be conspecific with the female lectotype; Pricop (2010b) was of opinion that it resembles G. ovicenatus Leonard & Crosby. 1 3 [ NHMW] on slide labeled: 1. [in Soyka’s handwriting] “ Gonatocerus 3 ater foersteri [Soyka’s manuscript name]”; 2. [partially in?Foerster’s handwriting, partially printed] “ Gon. ater Förster, Type”; 3. [printed] “Aach. Först.”; 4. [in Soyka’s handwriting] “In Canadab. 1943”. This specimen actually belongs to G. acuminatus . Potential paralectotypes: 1 Ƥ, 1 3 [ MHNG] on minuten pins inserted in the same small balsa wood piece on a pin labeled: 1. [in A. Foerster’s handwriting] “ Gonatocerus ater Frst. ”; 2. “Not Rachistus ater Fst. 1847 W. D. Hincks ”; 3. [in blue ink] “ Gonatocerus ater F.”. At least the female actually belongs to G. oxypygus . Although Foerster (1847) indicated 2 females and 1 male of G. ater , actually 4 specimens (2 females and 2 males) in NHMW can be unambiguously attributed to the syntype series of this species, and these belong at least to 3 different taxa. In addition, the abovementioned female in MHNG could also be part of the syntype series although that is less likely. All Foerster’s specimens identified by him as G. ater more or less fit the vague original description even though they represent several different species; in this situation I follow Matthews (1986), the first reviser, and designate as lectotype the specimen he mentioned as “ holotype ”. The original description of G. ater ( Foerster 1841, p. 45) stated, as newly translated here from German: “2. Gon. ater . Black, shining, antennae brown, scape yellowish, legs black-brown, knees and apices of tibiae and tarsi yellow, fore tibiae completely yellow. 3. Ƥ. Lg. [Length] 2/5 Lin. [Linie (an old German measuring unit, usually = 1/ 12 inch, but could also = 1/ 10 inch)]”. Later, Foerster (1847, pp. 206–207) added the following to the diagnosis of G. ater [as Rachistus ater ] (partial translation from German): “Body coloration completely dark. Scape dark brown and rather wide. F1–F4 very short and of the same length and progressively thicker. F5 considerably longer and thicker than F6; F6–F8 almost the same length. F8 equals F 5 in thickness. Clava almost the same length as F6–F8 combined. 2 Ƥ, 1 3 from the same area [i.e., Aachen]”.

Gonatocerus pannonicus Soyka : holotype female [ NHMW] on slide labeled: 1. “ Gonatocerus pannonicus Ƥ ( Soyka ) Type det. W. Soyka ”, 2. [red] “Type”, 3. “809”, 4. “ Hundsheim 9 Sept 1940 in Canadabalsam ”. The collecting date on the holotype slide does not match the one (August 1941) indicated in the original description of G. pannonicus by Soyka (1946). The holotype is mounted laterally, insufficiently cleared, lacking F5–F8 and clava of one antenna.

Lymaenon populi Viggiani : holotype female [ DEZA (current depository), although Viggiani (1969) indicated that the holotype female was to be deposited in the collection of the Center of the Identification of Entomophagues of the International Organization for Biological Control in Geneva, Switzerland (at MHNG)] on 2 slides, as follows: slide 1 (head, one antenna, and 1 fore wing), labeled: “ Coll. O. I. L. B. 27.67/2 1 Ƥ Lymaenon populi n. sp. olotipo det. G. Viggiani ’69”; slide 1 (remainder of the specimen), labeled: “ Coll. O. I. L. B. 27.67/ 2 Ƥ Lymaenon populi n. sp. olotipo det. G. Viggiani ’69”. Paratypes [all DEZA]: 1 Ƥ on slide labeled: “ Lymaenon populi Vigg. paratipo 1 Ƥ Roma , VI.68 ex uova Cicadella viridis su pioppo”; 1 Ƥ on slide labeled: “ Lymaenon populi Vigg. paratipo Ƥ Roma , VI.68 ex uova Cic. viridis ”; 1 Ƥ on slide labeled: “ Lymaenon populi Vigg. paratipo Ƥ Roma , VI/68 ex uova Cic. viridis ”; 1 3 on slide labeled: “ Lymaenon populi Vigg. 3 paraallotipo de uova cicadellide su pioppo Roma , VI/68 leg. Cavascasolle ”. All the paratypes are dissected in several body parts.

Lymaenon schmitzi Debauche : holotype female [ ISNB] on slide ( Fig. 214 ) labeled: 1. “ Héverlé 1.VI.41 — no140 1 [the last number in pencil]”; 2. “Dr. H. DEBAUCHE det. Lymaenon schmitzi Deb. 1943 Ƥ TYPE [the latter glued on a red triangle onto the right label]”. The holotype ( Fig. 216 ) is in fair condition although uncleared, complete, and mounted dorsoventrally. Paratypes [both ISNB]: 13 (the allotype) on slide labeled: 1. “ Lab. D’Entomologie de l’Université Louvain Eegenhoven 11.V.42. 180”, 2. “Dr. H. DEBAUCHE det. Lymaenon schmitzi Deb. 1943 3 ALLO TYPE [sic, the latter glued on a red triangle onto the right label]”; 1 3 on slide labeled: 1. “ Lab. D’Entomologie de l’Université Louvain Eegenhoven 18.V.42 no183”, 2. “Dr. H. DEBAUCHE det. Lymaenon schmitzi Deb. 1943 3 PARA TYPE [sic, the latter glued on a red triangle onto the right label]”. Both paratypes are uncleared and mounted laterally so that the propodeal carinae are not visible.

Material examined. Gonatocerus ater s. str. (i.e., specimens that more or less agree with the lectotype, particularly in the shape of the propodeal carinae). CZECH REPUBLIC. HRADEC KRÁLOVÉ, Orlické Mountains, Kačerov Nature Reserve , 50°14’25.241’’N 16°23’07.139’’E, 690 m, 20.x.2008, J. Hájek [1 Ƥ, CUPC]. RUSSIA. MOSCOVSKAYA OBLAST’: Noginskiy rayon, Fryazevo, M.E. Tretiakov : 1–18.v.2000 [1 Ƥ, UCRC]; 2.vi.2002 [1 Ƥ, UCRC]; 9.vii.2002 [1 Ƥ, UCRC]. Pushkinskiy rayon, Pushkino , Mamontovka , Sosnovka , 5– 16.v.2001, E.Ya. Shuvakhina [2 Ƥ, UCRC, ZIN]. SAKHALINSKAYA OBLAST’, Sakhalin Island, 2 km E Sokol , D.J. Bennett, T. Anderson, 21.vii.2001 [1 Ƥ, CAS]. TAMBOVSKAYA OBLAST’, Inzhavinskiy rayon, Talinka (7 km S of Pavlovka ), 26–27.v.2000, M.E. Tretiakov [1 3, UCRC]. UK. ENGLAND: Berkshire Co., Ascot, Silwood Park , 11.vi.1994, J.S. Noyes [4 3, CNCI]. Cheshire Co., Lymm , 28.v.1949, W.D. Hincks [1 Ƥ, MMUE]. WALES, Bridgend Co. Borough, Kenfig Pool National Nature Reserve , 4.viii.1994, J.S. Noyes [1 3, CNCI].

Gonatocerus ater s. l. (i.e., specimens that either do not agree with the lectotype in the shape of the propodeal carinae but fit Matthews’ (1986) and Zeya & Hayat’s (1995) concepts of the species, or for which the shape of the propodeal carinae is not known or has not been recorded for various reasons). BELGIUM. LIÈGE, Wanze, Antheit , Corphalie , 27.iv–11.v.1990, R. Detry [1 Ƥ, ISNB]. WALLOON BRABANT, Waterloo , 26.vii– 2.viii.1992, P. Dessart [1 Ƥ, [ ISNB]. GERMANY. NORTH RHINE-WESTPHALIA: Cologne , 6.viii.1962, M. Boness [3 Ƥ, NHMW]. Leverkusen , 3.vii.1963, M. Boness [1 Ƥ, NHMW]. GREECE. CENTRAL MACEDONIA, Lake Kerkini, Kerkini Marsh , 41°13’32.8’’N 23°05’04.2’’E, 45 m, 11–17.iv.2007, G. Ramel [1 Ƥ, 3 3, UCRC]. RUSSIA. KRASNODARSKIY KRAY, Krasnodar, 19–20.viii.2001, V.V. Kostjukov [1 3, UCRC]. KALUZHSKAYA OBLAST’, Sivkovo , 18.viii.1978, V.A. Trjapitzin [1 Ƥ, ZIN]. PRIMORSKIY KRAY, Ussuriyskiy rayon, Gornotayozhnoye, M.V. Michailovskaya : 11–12.vi.1999 [1 Ƥ, UCRC]; 23–24.vi.1999 [1 Ƥ, UCRC]; 1–2.vii.1999 [1 Ƥ, UCRC]; 11–14.vii.1999 [2 Ƥ, 1 3, UCRC]; 24.vii–1.viii.1999 [2 Ƥ, IBPV, UCRC]; 27.vii–1.viii.1999 [1 Ƥ, UCRC]; 4–5.viii.1999 [1 Ƥ, UCRC]; 5–11.viii.1999 [1 Ƥ, UCRC]; 12–17.viii.1999 [2 Ƥ, UCRC, ZIN]; 22–28.viii.1999 [1 Ƥ, UCRC]; viii.1999 [4 Ƥ, IBPV, UCRC, ZIN]; viii–ix.1999 [1 3, UCRC]; 10– 15.ix.1999 [3 Ƥ, UCRC]; 25–26.ix.1999 [1 Ƥ, UCRC]; ix.1999 [1 Ƥ, UCRC]; 8–11.x.1999 [1 Ƥ, UCRC]; 1– 10.vii.2000 [1 Ƥ, UCRC]; 10–20.vii.2000 [1 Ƥ, UCRC]; 21–26.viii.2000 [2 Ƥ, UCRC, ZIN]; viii.2000 [1 Ƥ, UCRC]. SAKHALINSKAYA OBLAST’, Sakhalin Island, 6 km E of Sokol, 16.viii.2001 [1 3, CAS]. STAVROPOL’SKIY KRAY, Prietokskiy , V.V. Kostjukov: 29.viii.2002 [1 Ƥ, UCRC]; 7.viii.2003 [4 Ƥ, UCRC]; 12.viii.2003 [1 Ƥ, UCRC]; 14.viii.2003 [3 Ƥ, UCRC]. UK. ENGLAND: Cheshire Co., Middlewood , 6.vii.1948, H. Britten [1 Ƥ, MMUE]. Surrey Co., Dorking, J.S. Noyes: Leith Hill , 26.viii.1984 [1 Ƥ, BMNH] (det. M.J. Matthews); White Downs, 21.ix.1986 [1 Ƥ, CNCI]. Country or locality not indicated (most likely Aachen area, North Rhine-Westphalia , GERMANY): 2 Ƥ [ MHNG] on minuten pins inserted in the same small balsa wood piece on a pin labeled: 1. [in A. Foerster’s handwriting] “ Gonatocerus ecaudatus Frst. ” [Foerster’s manuscript name]; 2. [in blue ink] “ Gonatocerus ecaudatus F.” (mounted together, but on separate minuten pins, with a female of G. oxypygus ).

Distribution. PALAEARCTIC: Gonatocerus ater s. str.: Belgium, Czech Republic, Germany, Russia *, and UK ( England, and Wales *). Records of this species from Austria ( Soyka 1946) [as G. pannonicus ], Bulgaria ( Donev 1986 [also as G. populi ], 1987, 1988d, 1988e, 2005), Finland ( Hellén 1974), Greece ( Donev 1988c, 2005), Ireland ( Graham 1973) [as G. populi ], Italy ( Viggiani 1969) [as G. populi ], Netherlands ( Noyes 2012), Romania ( Boţoc 1962 [as L. schmitzi ]; Pricop 2009b; Pricop 2010b [as G. intermedius ], and Sweden ( Hedqvist 2003) need confirmation. NEARCTIC*: USA * ( G. ater s. l., see “Comments” below for Lymaenon populi ). ORIENTAL: India ( Zeya & Hayat 1995; Anwar & Zeya 2012; Zeya & Khan 2012) ( G. ater s. l., records need confirmation).

Redescription. FEMALE (lectotype of G. ater , 5 non-type specimens of G. ater s. str. from Moskovskaya oblast’, Russia, 1 non-type specimen from Kačerov Nature Reserve, Czech Republic, and 1 non-type specimen from Lymm, England, that agree with the lectotype). Body length 840–860 µm (dry-mounted specimens from Kačerov Nature Reserve and Lymm, respectively). Body and appendages mostly dark brown, legs light to dark brown.

Head ( Fig. 207 ) about as wide as mesosoma. Antenna ( Figs 207 , 209 , 217–218 ) with radicle 0.24–0.29× total length of scape, rest of scape 2.1–2.4× as long as wide; pedicel much longer than F1; F1 and F2 subequal in length and the shortest funicle segments, F4 slightly shorter than F3 and shorter than following funicle segments, F5–F8 more or less subequal in length when F6 bears 1 or 2 mps except F8 slightly shorter, but if F6 lacks mps then F6 about as long as F8 or slightly shorter; mps on F5 (2), F6 (0, 1, or 2), F7 (2), and F8 (2); clava with 8 mps, 2.1–2.4× as long as wide, almost as long as combined length of F6–F8 or a little shorter.

Mesosoma ( Fig. 210 ) about as long as gaster ( Fig. 211 ) or a little shorter. Propodeum ( Figs 208 , 219 ) with fine, usually complete submedian carinae that narrow from propodeal posterior margin (their widest point) and either join together anteriorly at propodeal anterior margin at apex of dorsellum (as in the lectotype, Fig. 208 ) or fading at dorsellum ( Fig. 219 ). Fore wing ( Figs 212 , 220 ) 2.6–2.8× as long as wide; longest marginal seta 0.19– 0.2× maximum wing width; disc with a slight brownish tinge and bare behind venation except for 3 or more setae behind stigmal vein, and densely setose elsewhere. Hind wing ( Fig. 220 ) 12–14× as long as wide; disc unevenly setose, with a slight brownish tinge; longest marginal seta 1.6–1.7× maximum wing width.

Metasoma. Petiole short, about 2.2× as wide as long; ovipositor ( Fig. 211 ) not or at most barely exserted beyond apex of gaster, 1.1–1.4× as long as mesotibia.

Measurements (µm) of the lectotype. Head (as height: width) 264:283; mesosoma 474; ovipositor 412. Antenna: radicle 42; rest of scape 124; pedicel 64; F1 36; F2 36; F3 41; F4 40; F5 57; F6 49; F7 55; F8 49; clava 142. Fore wing 1236:474; longest marginal seta 91. Hind wing 947:81. Mesotibia 354.

MALE (non-type specimens from England and Russia). Body length 1130–1250 µm (slide-mounted specimens). Similar to female except for normal sexually dimorphic features and the following. Antenna ( Fig. 221 ) with scape 1.5–1.8× as long as wide, F1 wider than other flagellomeres. Fore wing ( Fig. 222 ) 2.5–2.7× as long as wide. Genitalia as in Fig. 223 .

Diagnosis. Gonatocerus ater s. str. is characterized mainly by the fine, usually complete submedian carinae that narrow from propodeal posterior margin (their widest point) and either joining together anteriorly at propodeal anterior margin at apex of dorsellum ( Fig. 208 ) or fading at dorsellum ( Fig. 219 ). The scape minus radicle of the female antenna ( Figs 217–218 ) is 2.1–2.4× as long as wide, and F3 normally lacks mps (although not observed in the specimens studied, presumably F 3 may occasionally have a mps on one or both antennae, particularly in large specimens). The ovipositor is short (1.1–1.4× as long as mesotibia) and not or at most barely exserted beyond apex of gaster ( Fig. 211 ).

Hosts. Unknown for G. ater s. str. Amrasca biguttula (Ishida) [as Empoasca devastans Distant ] for Lymaenon empoascae ( Subba Rao 1966; Subba Rao et al. 1968) and Cicadella viridis (Linnaeus) for Lymaenon populi ( Viggiani 1969) (Cicadellidae) . However, later Viggiani (1988) [as Lymaenon cicadellae Viggiani] and Viggiani & Jesu (1988) indicated Rhytidodus decimaquartus (Schrank) [as R. decimusquartus ] as the host of L. populi in Italy.

Comments. In the holotype of L. schmitzi the submedian carinae on the propodeum are more or less clearly visible ( Fig. 215 ); they appear to be subparallel and fading anteriorly, not extending to the anterior margin of the propodeum and thus not joining at the apex of dorsellum. The antenna ( Fig. 216 ) is more or less similar to that of the lectotype of G. ater : one antenna of the former has 2 mps on F6 and the other lacks mps, and in the latter one antenna has 1 mps on F6 and the other lacks mps; both lack mps on F1–F4 and have 2 mps on F5, F7, and F8. Also, in the holotype of L. schmitzi the scape minus radicle is about 2.3× as long as wide, the fore wing is 2.7× as long as wide, and the ovipositor is about 1.4× mesotibia length. Taking all this into consideration, I accept (although not confidently) the synonymy of L. schmitzi under G. ater . More, fresh, and properly prepared material from or near the type localities of both nominal taxa is needed to assess variation, particularly of the shape of the propodeal submedian carinae.

Because the holotype female of G. pannonicus is mounted laterally, it is impossible to see the propodeal submedian carinae in dorsal view. F3 of its antenna is notably longer than F1, F2, or F4 and subequal to F5 (both are the longest funicle segments); mps are on F3 (1 or?2), F4 (0), F5 (2), F6 (1), F7 (?1 or 2), and F8 (2). The ovipositor is short, about 0.7× length of the gaster, not exserted beyond its apex. I tentatively accept the synonymy of this species under G. ater by Matthews (1986) although at the same time agree with Pricop (2010b) that these two taxa may not be conspecific. Possibly, G. pannonicus is conspecific with L. intermedius and also with L. populi , as described and illustrated by Boţoc (1962) and Viggiani (1969), respectively, as all these taxa have similar female antennae and short ovipositors. The holotype of G. pannonicus needs to be carefully re-mounted dorsoventrally to be able to see its propodeal submedian carinae and thus determine its true identity.

Pricop (2010b) considered G. intermedius to be a valid species based on several specimens of both sexes collected in Romania but these are not from Cluj County where the unspecified type locality (near Cluj-Napoca) of Lymaenon intermedius was. According to the diagnosis and illustrations of the non-type specimens attributed to this species by Pricop (2010b), indeed these appear to fit well with the original description except for the markedly longer ovipositor (1.8× as long as mesotibia and slightly exserted beyond the gastral apex). According to Boţoc (1962), the ovipositor was short and not exserted in the syntypes of L. intermedius , as seen on the photograph of the female habitus (p. 109, her fig. 3). Collections should be made near Cluj-Napoca in June, and if a female that fits the original description of L. intermedius is found (it is important to have a short ovipositor!), a neotype needs to be designated meeting all the requirements of Article 75.3 ([ICZN] 1999). It is inadvisable to use any of the female specimens mentioned by Pricop (2010b) for such a designation, even though they might eventually turn out to be conspecific with L. intermedius , because they have markedly longer ovipositors and were collected significantly far away from the type locality. I have examined the following specimens that fit well the diagnosis and the illustrations of the form that was attributed by Pricop (2010b) to G. intermedius — AUSTRIA. LOWER AUSTRIA, 1 km W of Hollern , 48°04’22’’N 16°52’37’’E, 150 m, 16.vi.2007, C. Thuróczy, S.V. Triapitsyn [2 Ƥ, UCRC].

ITALY. LAZIO, 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 [3 Ƥ, UCRC]. RUSSIA. PRIMORSKIY KRAY, Ussuriyskiy rayon, Gornotayozhnoye , 28.viii–5.ix.1999, M.V. Michailovskaya [1 Ƥ, UCRC]. STAVROPOL’SKIY KRAY, Prietokskiy , V.V. Kostjukov: 14.vii.2003 [4 Ƥ, UCRC]; 12.viii.2003 [16 Ƥ, UCRC, ZIN]. SLOVAKIA. BRATISLAVA, Jurský Šúr Nature Reserve , 48°14’03’’N 17°12’47’’E, 133 m, 8.viii.2008, B.V. Brown (alder forest) [2 Ƥ, UCRC] . In these specimens, the ovipositor is 1.6–1.9× as long as mesotibia and slightly exserted beyond the gastral apex ( Figs 225 , 229 ); other important features such as the antenna ( Figs 224 , 227 ), the propodeal submedian carinae ( Fig. 228 ), and the wings ( Fig. 226 ) are identical to the ones in Pricop’s specimens from Romania and to the numerous specimens attributable to L. populi from several European countries, as discussed below. Thus, the length of the ovipositor relative to the length of the mesotibia in both forms seems to be slightly overlapping while gradually increasing in various specimens between 1.0× and 1.6× (in some individuals attributable to L. populi sensu Viggiani ) to 1.6–1.9× (in the specimens attributable to G. intermedius sensu Pricop ). However, the significance of that is not clear, and I would abstain from resurrecting or sinking nominal species in the G. ater complex based on such variable morphological features as the relative length of the ovipositor or presence/absence of mps on F3 and/or F6.

Viggiani & Jesu (1988) did not accept the synonymy of Lymaenon populi under G. ater by Matthews (1986) and they might be right unless the shape of the propodeal submedian carinae and the relative length of the ovipositor vary significantly in G. ater . This is quite possible because forms with somewhat different shapes of the propodeal submedian carinae and/or different relative lengths of the ovipositor do occur in Austria (near Hollern), Italy (Ponte San Pietro), and Russia (Fryazevo, Prietokskiy) in the same place and at the same or about the same time: unlike in the lectotype of G. ater , the propodeal carinae in specimens of the type series of L. populi are more or less parallel to each other, not joining anteriorly at the posterior margin of the dorsellum and not extending (or sometimes almost extending) to it ( Figs 232 , 236 ). In the holotype and female paratypes of L. populi , the scape minus radicle ( Figs 230–231 ) is rather wide (2.0–2.1× as long as wide, very similar to that in G. ater s. str.), F3 and F6 bear 1 mps each and F5, F6 and F7 bear 2 mps each, the clava has 8 mps, and the ovipositor ( Fig. 233 ) is short (1.1–1.2× as long as mesotibia). Also illustrated here are the female wings ( Fig. 234 ) and the male antenna ( Fig. 235 ) of the paratypes of L. populi as well as the antennae ( Figs 237–239 ), wings ( Fig. 240 ), body ( Fig. 241 ), propodeum ( Fig. 242 ), and metasoma ( Fig. 243 ) of the female, and the antenna ( Fig. 244 ), fore wing ( Fig. 245 ), and genitalia ( Fig. 246 ) of the male to illustrate variation of the key morphological features in the non-type specimens attributable to this form, of which I have examined the following material— AUSTRIA. LOWER AUSTRIA, 1 km W of Hollern, 48°04’22’’N 16°52’37’’E, 150 m, 16.vi.2007, C. Thuróczy, S.V. Triapitsyn [1 Ƥ, UCRC]. GREECE. CENTRAL MACEDONIA, Lake Kerkini : Beles Mts., 41°17’19.5’’N 23°12’18.4’’E, 550 m, 9–15.v.2007, G. Ramel [1 Ƥ, UCRC]. Kerkini Marsh , 41°13’32.8’’N 23°05’04.2’’E, 45 m, 11–17.iv.2007, G. Ramel [1 Ƥ, UCRC]. ITALY. LAZIO, Roma Prov.: Castelporziano Presidential Estate , Ponte Guidoni , 41°45.415’N 12°23.851’E, 80 m, 11.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [1 Ƥ, UCRC]. Near Maccarese Cemetary , 41°52.836’N 12°16.190’E, 40 m, 11.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [1 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 [5 Ƥ, 1 3, UCRC]. Roccaccia , 42°19.809’N 11°45.671’E, 125 m, 10.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [5 Ƥ, UCRC]. San Giovenale , 42°13.568’N 12°00.039’E, 225 m, 9.vi.2003, M. Bologna, J. Munro, A. Owen, J.D. Pinto [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 [7 Ƥ, UCRC]. RUSSIA. KRASNODARSKIY KRAY, Krasnodar, 31.viii.2003, V.V. Kostjukov [1 Ƥ, UCRC]. MOSCOVSKAYA OBLAST’, Noginskiy rayon, Fryazevo, M.E. Tretiakov : 2–15.vi.2000 [1 Ƥ, [ UCRC]; 21.vi.2001 [1 Ƥ, [ UCRC]. PRIMORSKIY KRAY, Ussuriyskiy rayon, Gornotayozhnoye , 2–28.x.1999, M.V. Michailovskaya [1 Ƥ, UCRC]. STAVROPOL’SKIY KRAY: Achikulak , 26.viii.2002, V.V. Kostjukov [2 Ƥ, UCRC, ZIN]. Prietokskiy, V.V. Kostjukov: 14.vii.2003 [3 Ƥ, UCRC, ZIN]; 12.viii.2003 [1 Ƥ, UCRC]. UK. ENGLAND, East Sussex Co., Ashdown Forest , 28.vii.1982, J.S. Noyes [1 Ƥ, BMNH] (det. by M.J. Matthews). USA. CALIFORNIA, Orange Co., Irvine, Northwood Pointe , 33°43’18’’N 117°45’12’’W, 76 m, 7.viii.2011, S.V. Triapitsyn (on Lombardy poplar, Populus nigra ) [1 Ƥ, UCRC]. NEW YORK, Ontario Co., Geneva, 42°52’46’’N 77°00’40’’W, 185 m (on Lombardy poplar, Populus nigra , roadside of County Road 6): 3.viii.2010, S.V. Triapitsyn [4 Ƥ, 1 3, UCRC]; 23.ix.2010, S.V. Triapitsyn, G. Loeb [1 Ƥ, UCRC]. In these specimens, some of which are at least tentatively attributable to Viggiani’s species, F3 and F6 of the female antenna sometimes may lack mps (usually in small specimens) or F 6 may occasionally bear 2 mps in large specimens, occasionally the propodeal submedian carinae may slightly curve towards each other anteriorly (but not joining nor extending to the anterior margin of the propodeum), and the ovipositor length varies from usually 1.0–1.3× to sometimes up to 1.6× as long as mesotibia. I found the same form ( Fig. 247 ) on Lombardy poplars (non-native to North America, of the European origin) in Geneva, New York, USA, which was apparently unintentionally introduced there with its likely host, Rhytidodus decimaquartus , along with G. oxypygus . Both species are also known from Italy ( Viggiani 1969), apparently from the similar habitat. I also found this form on Lombardy poplars in southern California, USA; it differs from the native species G. (Cosmocomoidea) impar Huber by a shorter radicle and a relatively wider main body of the scape of the female antenna and a relatively wider hind wing.

Gonatocerus bifasciatus Girault ( Viggiani 2005: 65) , who compared its male genitalia with those of “ G. prope populi Viggiani ” (two males from Williamsville, Wayne Co., Missouri, USA), is a nomen nudum ( Noyes 2012).

Because the proportions of funicle segments and presence/absence of mps (particularly on F3 and F6) are quite variable in specimens that can be attributed more or less confidently to the already described species within G. ater complex, in any possible combination with the shape of the propodeal submedian carinae and the length of the ovipositor relative to the length of the mesotibia (which is also quite variable), it is currently impossible to decide where the limits between the likely cryptic species are versus intraspecific variation. Until this complex is studied extensively using molecular methods and cross-breeding experiments (e.g., Triapitsyn et al. 2008) to go along a thorough morphometrical analysis, and until more freshly collected, preferably reared from known hosts, specimens are obtained in or near the type localities of the nominal species comprising it, the synonymies proposed by Matthews (1986) are accepted (including that of L. intermedius ) and I treat all the specimens discussed here as belonging to G. ater s. l.

Figures

FIGURE 213. Gonatocerus (Cosmocomoidea) ater 3 (paralectotype, may or may not belong to this species): habitus.

FIGURE 216. Gonatocerus (Cosmocomoidea) ater? s. str. Ƥ (holotype of Lymaenon schmitzi Debauche): habitus.

FIGURE 220. Gonatocerus (Cosmocomoidea) ater s. str. Ƥ (Mamontovka, Moskovskaya oblast’, Russia): wings.

FIGURE 247. Gonatocerus (Cosmocomoidea) ater s. l. [attributable to Lymaenon populi Viggiani] Ƥ (Geneva, New York, USA): habitus.

FIGURES 206 – 208. Gonatocerus (Cosmocomoidea) ater Ƥ (lectotype): 206, slide; 207, head and antennae; 208, dorsellum and propodeum.

FIGURES 209 – 212. Gonatocerus (Cosmocomoidea) ater Ƥ (lectotype): 209, antenna; 210, mesosoma; 211, gaster; 212, fore wing.

FIGURES 214, 215. Gonatocerus (Cosmocomoidea) ater? s. str. Ƥ (holotype of Lymaenon schmitzi Debauche): 214, slide; 215, dorsellum and propodeum.

FIGURES 217 – 219. Gonatocerus (Cosmocomoidea) ater s. str. Ƥ (Mamontovka, Moskovskaya oblast’, Russia): 217 – 218, antennae; 219, dorsellum and propodeum.

FIGURES 221 – 223. Gonatocerus (Cosmocomoidea) ater s. str. 3 (Silwood Park, Ascot, England): 221, antenna; 222, fore wing; 223, genitalia.

FIGURES 224 – 226. Gonatocerus (Cosmocomoidea) ater s. l. [= G. intermedius (Boţoc) sensu Pricop (2010 b)] Ƥ (224, 225 — 1 km W of Hollern, Austria; 226 — Ponte San Pietro, Lazio, Italy): 224, antenna; 225, dorsellum, propodeum, and metasoma; 226, wings.

FIGURES 227 – 229. Gonatocerus (Cosmocomoidea) ater s. l. [= G. intermedius (Boţoc) sensu Pricop (2010 b)] Ƥ (Ponte San Pietro, Lazio, Italy): 227, antenna; 228, dorsellum and propodeum; 229, gaster.

FIGURES 230 – 232. Gonatocerus (Cosmocomoidea) ater s. l. (paratypes of Lymaenon populi Viggiani) Ƥ: 230, head and antennae; 231, antenna; 232, mesosoma.

FIGURES 233, 234. Gonatocerus (Cosmocomoidea) ater s. l. (paratypes of Lymaenon populi Viggiani) Ƥ: 233, gaster; 234, wings.

FIGURES 235, 236. Gonatocerus (Cosmocomoidea) ater s. l. (paratype of Lymaenon populi Viggiani) 3: 235, antenna; 236, dorsellum and propodeum.

FIGURES 237 – 240. Gonatocerus (Cosmocomoidea) ater s. l. [attributable to Lymaenon populi Viggiani] Ƥ (237, 238, 240 — Ponte San Pietro, Lazio, Italy; 239 — near Maccarese Cemetary, Lazio, Italy): 237 – 239, antennae; 240, wings.

FIGURES 241 – 243. Gonatocerus (Cosmocomoidea) ater s. l. [attributable to Lymaenon populi Viggiani] Ƥ (241, 242 — Ponte San Pietro, Lazio, Italy; 243 — Krasnodar, Russia): 241, body; 242, dorsellum and propodeum; 243, propodeum and metasoma.

FIGURES 244 – 246. Gonatocerus (Cosmocomoidea) ater s. l. [attributable to Lymaenon populi Viggiani] 3 (Ponte San Pietro, Lazio, Italy): 244, antenna; 245, fore wing; 246, genitalia.

FIGURES 290 – 292. Gonatocerus (Cosmocomoidea) oxypygus Ƥ (paralectotype of G. a t e r Foerster): 290, antennae; 291, body; 292, wings.