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

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

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http://dx.doi.org/10.11646/zootaxa.3644.1.1

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scientific name

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

( Figs 206–247 View FIGURES 206 – 208 View FIGURES 209 – 212 View FIGURE 213 View FIGURES 214, 215 View FIGURE 216 View FIGURES 217 – 219 View FIGURE 220 View FIGURES 221 – 223 View FIGURES 224 – 226 View FIGURES 227 – 229 View FIGURES 230 – 232 View FIGURES 233, 234 View FIGURES 235, 236 View FIGURES 237 – 240 View FIGURES 241 – 243 View FIGURES 244 – 246 View FIGURE 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 1988 d: 194 (distribution); Donev 1988 e: 203 (distribution); Donev 2005: 383 (diagnosis, distribution); Pricop 2010 b: 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 2010 b)] (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 (2010 b), 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 2010 b: 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 2010 b: 112 –113 (possible synonym of G. ater   ), 114 (illustrations).

Gonatocerus intermedius (Boţoc)   : Pricop 2010 b: 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 View FIGURES 206 – 208 ) 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 (2010 b: 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 View FIGURES 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 View FIGURE 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 (2010 b) 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. F 1 –F 4 very short and of the same length and progressively thicker. F 5 considerably longer and thicker than F 6; F 6 –F 8 almost the same length. F 8 equals F 5 in thickness. Clava almost the same length as F 6 –F 8 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 F 5 –F 8 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 View FIGURES 214, 215 ) labeled: 1. “Héverlé 1.VI. 41 — no 140 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 View FIGURE 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 no 183 ”, 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, 1988 e, 2005), Finland ( Hellén 1974), Greece ( Donev 1988 c, 2005), Ireland ( Graham 1973) [as G. populi   ], Italy ( Viggiani 1969) [as G. populi   ], Netherlands ( Noyes 2012), Romania ( Boţoc 1962 [as L. schmitzi   ]; Pricop 2009 b; Pricop 2010 b [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 View FIGURES 206 – 208 ) about as wide as mesosoma. Antenna ( Figs 207 View FIGURES 206 – 208 , 209 View FIGURES 209 – 212 , 217– 218 View FIGURES 217 – 219 ) 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 F 1; F 1 and F 2 subequal in length and the shortest funicle segments, F 4 slightly shorter than F 3 and shorter than following funicle segments, F 5 –F 8 more or less subequal in length when F 6 bears 1 or 2 mps except F 8 slightly shorter, but if F 6 lacks mps then F 6 about as long as F 8 or slightly shorter; mps on F 5 (2), F 6 (0, 1, or 2), F 7 (2), and F 8 (2); clava with 8 mps, 2.1–2.4 × as long as wide, almost as long as combined length of F 6 –F 8 or a little shorter.

Mesosoma ( Fig. 210 View FIGURES 209 – 212 ) about as long as gaster ( Fig. 211 View FIGURES 209 – 212 ) or a little shorter. Propodeum ( Figs 208 View FIGURES 206 – 208 , 219 View FIGURES 217 – 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 View FIGURES 206 – 208 ) or fading at dorsellum ( Fig. 219 View FIGURES 217 – 219 ). Fore wing ( Figs 212 View FIGURES 209 – 212 , 220 View FIGURE 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 View FIGURE 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 View FIGURES 209 – 212 ) 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; F 1 36; F 2 36; F 3 41; F 4 40; F 5 57; F 6 49; F 7 55; F 8 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 View FIGURES 221 – 223 ) with scape 1.5–1.8 × as long as wide, F 1 wider than other flagellomeres. Fore wing ( Fig. 222 View FIGURES 221 – 223 ) 2.5–2.7 × as long as wide. Genitalia as in Fig. 223 View FIGURES 221 – 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 View FIGURES 206 – 208 ) or fading at dorsellum ( Fig. 219 View FIGURES 217 – 219 ). The scape minus radicle of the female antenna ( Figs 217–218 View FIGURES 217 – 219 ) is 2.1–2.4 × as long as wide, and F 3 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 View FIGURES 209 – 212 ).

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 View FIGURES 214, 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 View FIGURE 216 ) is more or less similar to that of the lectotype of G. ater   : one antenna of the former has 2 mps on F 6 and the other lacks mps, and in the latter one antenna has 1 mps on F 6 and the other lacks mps; both lack mps on F 1 –F 4 and have 2 mps on F 5, F 7, and F 8. 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. F 3 of its antenna is notably longer than F 1, F 2, or F 4 and subequal to F 5 (both are the longest funicle segments); mps are on F 3 (1 or? 2), F 4 (0), F 5 (2), F 6 (1), F 7 (? 1 or 2), and F 8 (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 (2010 b) 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 (2010 b) 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 (2010 b), 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 (2010 b) 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 (2010 b) 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 View FIGURES 224 – 226 , 229 View FIGURES 227 – 229 ); other important features such as the antenna ( Figs 224 View FIGURES 224 – 226 , 227 View FIGURES 227 – 229 ), the propodeal submedian carinae ( Fig. 228 View FIGURES 227 – 229 ), and the wings ( Fig. 226 View FIGURES 224 – 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 F 3 and/or F 6.

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 View FIGURES 230 – 232 , 236 View FIGURES 235, 236 ). In the holotype and female paratypes of L. populi   , the scape minus radicle ( Figs 230–231 View FIGURES 230 – 232 ) is rather wide (2.0– 2.1 × as long as wide, very similar to that in G. ater   s. str.), F 3 and F 6 bear 1 mps each and F 5, F 6 and F 7 bear 2 mps each, the clava has 8 mps, and the ovipositor ( Fig. 233 View FIGURES 233, 234 ) is short (1.1–1.2 × as long as mesotibia). Also illustrated here are the female wings ( Fig. 234 View FIGURES 233, 234 ) and the male antenna ( Fig. 235 View FIGURES 235, 236 ) of the paratypes of L. populi   as well as the antennae ( Figs 237–239 View FIGURES 237 – 240 ), wings ( Fig. 240 View FIGURES 237 – 240 ), body ( Fig. 241 View FIGURES 241 – 243 ), propodeum ( Fig. 242 View FIGURES 241 – 243 ), and metasoma ( Fig. 243 View FIGURES 241 – 243 ) of the female, and the antenna ( Fig. 244 View FIGURES 244 – 246 ), fore wing ( Fig. 245 View FIGURES 244 – 246 ), and genitalia ( Fig. 246 View FIGURES 244 – 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, F 3 and F 6 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 View FIGURE 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 F 3 and F 6) 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.

IARI

Indian Agricultural Research Institute

NHMW

Naturhistorisches Museum, Wien

MHNG

Museum d'Histoire Naturelle

DEZA

Dipartimento di Entomologia e Zoologia Agraria dell'Universita

UCRC

University of California, Riverside

ZIN

Russian Academy of Sciences, Zoological Institute, Zoological Museum

CAS

California Academy of Sciences

CNCI

Canadian National Collection Insects

MMUE

Museum of Manchester University

NEW

University of Newcastle

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Hymenoptera

Family

Mymaridae

Genus

Gonatocerus

Loc

Gonatocerus (Cosmocomoidea) ater Foerster, 1841 s

Triapitsyn, Serguei V. 2013
2013
Loc

Gonatocerus pannonicus

Pricop 2010: 112
2010
Loc

Gonatocerus intermedius (Boţoc)

Pricop 2010: 113
2010
Loc

Lymaenon populi

Viggiani 1988: 1023
1988
Loc

Gonatocerus schmitzi

Huber 1988: 50
1988
Loc

Gonatocerus empoascae (

Subba 1983: 135
1983
Loc

Gonatocerus indicus

Subba 1983: 135
1983
Loc

Gonatocerus nigriodes

Subba 1983: 136
1983
Loc

Gonatocerus populi (

Pricop 2010: 113
Viggiani 1973: 48
1973
Loc

Lymaenon populi

Matthews 1986: 222
Viggiani 1969: 40
1969
Loc

Lymaenon empoascae

Zeya 1995: 70
Subba 1966: 190
1966
Loc

Lymaenon schmitzi

Viggiani 1969: 39
Botoc 1962: 108
1962
Loc

Lymaenon intermedius Boţoc 1962 : 108

Matthews 1986: 222
Botoc 1962: 108
1962
Loc

Lymaenon nigroides

Zeya 1995: 70
Narayanan 1961: 656
1961
Loc

Lymaenon indicus

Zeya 1995: 70
Subba 1959: 229
1959
Loc

Lymaenon ater (Förster)

Viggiani 1988: 1023
Debauche 1948: 81
1948
Loc

Lymaenon schmitzi

Matthews 1986: 221
Debauche 1948: 86
1948
Loc

Gonatocerus pannonicus

Matthews 1986: 221
Soyka 1946: 39
1946
Loc

Rachistus ater (

Foerster 1847: 206
1847
Loc

Gonatocerus ater

Anwar 2012: 52
Zeya 2012: 57
Pricop 2010: 112
Donev 2005: 383
Zeya 1995: 70
Donev 1988: 194
Donev 1988: 203
Donev 1987: 73
Matthews 1986: 221
Hellen 1974: 13
Dalla 1898: 429
Kirchner 1867: 201
Walker 1846: 53
1846
Loc

Gonatocerus ater

Foerster 1841: 45
1841