A peer-reviewed open-access journal ZooKeys 130: 473-494 (2011) doi: 10.3897/zookeys. 130.1717 Z00Ke y S www.zookeys.o rg Launched to accelerate biodiversity research Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales, with description of two new genera and review of Tertiary amber genera John T. Huber''t, Dale Greenwalt?* I Natural Resources Canada, clo AAFC, K.W. Neatby Building, 960 Carling Ave., Ottawa, ON, KIA 0C6, Canada 2 Paleobiology Department, National Museum of Natural History, Washington, DC, 20013, USA T urn:lsid:zoobank. org:author:6BE7E99B-9297-437D-A14E-76FEF6011B10 * urn:lsid:zoobank. org:author:13918687-A71E-4F9A-A6D7-6E9791BEF751 Corresponding author: John T: Huber (John.Huber@agr.gc.ca) Academic editor: Michael Engel | Received 15 June 2011 | Accepted 1 July 2011 | Published 24 September 2011 urn:lsid:zoobank.org:pub:5E5101 CB-35C6-49F F-97A5-A5A254B3B35F Citation: Huber JT, Greenwalt D (2011) Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales, with description of two new genera and review of Tertiary amber genera. In: Shcherbakov DE, Engel MS, Sharkey MJ (Eds) Advances in the Systematics of Fossil and Modern Insects: Honouring Alexandr Rasnitsyn. ZooKeys 130: 473-494, doi: 10.3897/zookeys. 130.1717 Abstract Compression fossils of three genera and six species of Mymaridae (Hymenoptera: Chalcidoidea) are de- scribed from 46 million year old Kishenehn oil shales in Montana, USA. Two new genera are described: Eoeustochus Huber, gen. n., with two included species, E. kishenehn Huber (type species) and E. borchersi Huber, sp. n., and Eoanaphes., gen. n., with E. stethynioides Huber, sp. n. Three new species of Gona- tocerus are also described, G. greenwalti Huber, sp. n. , G. Rootenai Huber, sp. n., and G. rasnitsyni Huber, sp. n. Previously described amber fossil genera are discussed and five genera in Baltic amber are tentatively recorded as fossils: Anagroidea, Camptoptera, Dorya, Eustochus, and Mimalaptus. Keywords Mymaridae, Kishenehn Formation, compression fossils, Eoanaphes, Eoeustochus, Gonatocerus, Baltic amber. Copyright John T. Huber and Dale Greenwalt. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 474 John T: Huber & Dale Greenwalt / ZooKeys 130: 473-494 (2011) Introduction The family Mymaridae (Hymenoptera: Chalcidoidea) is represented by 103 genera and about 1400 nominal extant species in all terrestrial habitats and a few fresh water habitats (Huber 1986, Noyes 2010). Fossil Mymaridae have been described from am- ber inclusions from various localities, including five extinct genera and species from Cretaceous amber (Yoshimoto 1975, Huber and Poinar 2011) and eight genera (seven still extant, one extinct) from Tertiary amber, mostly from the Samland Peninsula, Kaliningrad district, Russia, and Chiapas, Mexico. Probably because of their small size, no compression fossils of Mymaridae have been discovered until now. Here we describe the first ones, representing three genera and six species. These and the Tertiary amber fossil genera are discussed and compared with extant genera. Methods A total of 17 specimens of fossil Mymaridae, all females, were collected in 2009 and 2010 at six sites (e.g. W113°42.173', N48°23.476') along the Middle Fork of the Flathead River in northwestern Montana between Paola and Coal Creeks in accord- ance with USFS Authorization HUN281. Fossil specimens were collected from the middle sequence of the Coal Creek member of the Kishenehn Formation, which has been estimated to be 46.2 +/- 0.4 myo (Lutetian) by “°Ar/*?Ar analysis and 43.5 +/- 4.9 myo by fission-track analysis (Constenius 1996). Although the paper/oil shales of the middle sequence are thin (<1 mm to several mm), they were often easily split into even thinner pieces so as to expose unweathered surfaces on which the fossil insects reside. The compression fossils were immersed in 95% ethanol for examination and pho- tography. For figures 1-10 and 13-18, specimens were photographed using a Zeiss AxioCam MRc5 digital CCD camera mounted on a Zeiss Discovery V20 microscope and Zeiss AxioVision EDF software in Ottawa, ON. Measurements were generated using Zeiss AxioVision software. For figures 11 and 12, specimens were photographed using an Olympus SZX12 microscope, DP-25 camera and DPM imaging software in Washington, DC, and measurements were taken with the DP2-BSW software. All measurements are in micrometers (mm). If a measurement could not be made (e.g., scape length) it is represented by a dash. Abbreviations used: fl = funicle segment x. Measurements were taken as accu- rately as possible but given that the beginning and end points of a structure were not always clear or were hidden, the measurements may not be accurate. Appendage meas- urements are the most accurate except the wing bases cannot be clearly determined. Consequently, wing lengths were taken from the visible edge of the mesosoma and therefore they and length/width ratios are slightly smaller than they should be. Fossils examined are in the following institutions: AMNH_ American Museum of Natural History. Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales... 475 IPMG6 Institute and Museum ftir Paléontologie, Georg-August University, Gét- tingen, Germany. NMNH_ National Museum of Natural History, Washington, D.C. OSU Oregon State University, G. Poinar, Jr. collection, Corvallis, Oregon, USA. SVT S. V. Triapitsyn private collection, California, USA. UCRC — University of California, Riverside, California, USA. ZMUC = Zoological Museum, University of Copenhagen, Denmark. The compression fossils are housed in NMNH. Baltic amber fossils were examined from the remaining institutions. Systematics Eoeustochus Huber, gen. n. urn:lsid:zoobank.org:act:25AE7CE1-EB1D-4B39-8D61-AA86366A4A35 http://species-id.net/wiki/Eoeustochus Figs 1-8 Description. Female. Body length 718—1133 (seven specimens in total, only four de- scribed and named to species). Head. Normal in shape, wider than long and about % as high as wide. Face slightly convex in lateral view; vertex flat and slightly sloping anter- iorly, forming a moderately sharp angle with occiput; back of head slightly concave. Eye higher than wide, about 2/3 head height; malar space about 1/3 eye height. Antenna. Funicle 6-segmented, with each funicle segment longer than wide; clava 3-segmented with the claval sutures perpendicular to claval length. Wings. Fore wing wide, symmet- rical, with evenly rounded apex (shape resembling that of the extant genus Eustochus); marginal fringe shorter than fore wing width. Venation about 0.4x fore wing length, with long marginal vein and short but distinct stigmal vein. Hind wing narrow; mar- ginal fringe much longer than wing width. Mesosoma. Shorter than gaster. Pronotum length at most about half length of mesoscutum. Mesoscutum length subequal to scutellum. Scutellum with frenum apparently entire, not divided longitudinally. Met- anotum much shorter than scutellum. Metasoma. Constricted at base, probably with short petiole. Gastral terga similar in length. Ovipositor moderately short, probably arising near midpoint of gaster and its apex not or barely exserted beyond gastral apex. Type species. Eoeustochus kishenehn Huber, sp. n. Derivation of generic name. The name is from eo-, Greek for early, and Eustochus, an extant genus. The gender is masculine. Discussion. Two species are described, each from two specimens. Three additional specimens (Kishenehn #30,356, 40,410, and 40,023) belong to Eoeustochus but are not included in the type series of either species because they are not in as good a condition. The relationships of Eoeustochus are with Eustochus, one extant species of which has a 3-segmented clava in females. Although thirteen extant genera have a definite 476 John T: Huber & Dale Greenwalt / ZooKeys 130: 473-494 (2011) 3-segmented clava (Allanagrus, Anneckia, Idiocentrus, Krokella, Nesopatasson/Nesomy- mar, Neostethynium, Notomymar, Paracmotemnus, Parastethynium, Polynemoidea, Pseu- docleruchus, Stethynium), the closest one is Eustochus, based on the apparent strong constriction in lateral view between meso- and metasoma (none in Al/anagrus), trans- verse sutures between claval segments (sutures oblique in Stethynium), shape of fully developed wings (wings almost absent in Nesopatasson! Nesomymar and Notomymar, and of different shape in Anneckia, Neostethynium, Parastethynium and Pseudocleruchus) venation shorter than half fore wing length (venation longer in Krokella and Parac- motemnus), and ovipositor not distinctly exserted beyond apex of gaster (ovipositor strongly exserted in Polynemoidea) or projecting anteriorly under mesosoma (oviposi- tor strongly projecting anteriorly in [diocentrus). Eoeustochus kishenehn Huber, sp. n. urn:lsid:zoobank.org:act:5\CE2A3C6-9203-4A2A-8B2F-4FECC0937342 http://species-id.net/wiki/Eoeustochus_kishenehn Figs 1-4 Type material. Holotype female (NMNH), labelled “Eoeustochus kishenehn Huber Holotype female #543757”. Paratype female (NMNH), labelled as for holotype but “paratype #543758”. Description. Female. Colour brown, head dark brown, fore wing thinly margined with brown. Holotype (Fig. 1) measurements as follows. Body length 909. Antenna (Fig. 2) with total funicle length 317; scape 95, pedicel 70, fl, 52, fl, 63, fl, G3 stl Sls fl. AT, fl. 45; clava 180, 0.56x funicle length. FP and fl, the longest segments and fl. the shortest (Fig. 2). Mesosoma length 481. Fore wing (Fig. 3) length 912, width 271, length/width 3.36, longest marginal setae 160, venation length 355. Hind wing width 26, longest marginal setae 88. Metasoma length 340. Ovipositor length 290, appar- ently slightly exserted beyond apex of gaster. Paratype (Fig. 4) measurements as follows. Body length 966, head width 234, height 132. Derivation of species name. Named after the Kishenehn Formation shale in which the fossils were found. Eoeustochus borchersi Huber, s urn:lsid:zoobank.org:act: BOGEABBE. 964F-4D92-AF7A-D3DE8EBB780B http://species-id.net/wiki/Eoeustochus_borchersi Figs 5—8 Type material. Holotype female (NMNH_ ) labelled “Eoeustochus borchersi Huber Holotype female #543759” Paratype female (NMNH), labelled as for holotype but “paratype #543760”. Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales... 500 pm 2 geass agi ay << B i. a ae = Sts Figures 1-2. Eoeustochus kishenehn, holotype | habitus lateral 2 mesosoma, head, antennae. 477 478 John T: Huber & Dale Greenwalt / ZooKeys 130: 473-494 (2011) plead Ui a . e | . * + S oe Figures 3-4. Eveustochus kishenehn 3 wings, holotype 4 paratype, habitus dorsal. Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales... ; 500 um & Figures 5-6. Eoeustochus borchersi, holotype 5 habitus lateral 6 head and antennae. 480 John T. Huber & Dale Greenwalt / ZooKeys 130: 473-494 (2011) ee ; = fie Figures 7-8. Eveustochus borchersi, holotype 7 wings 8 mesosoma and base of wings. Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales... 481 Description. Female. Colour dark brown, fore wing thinly margined with brown. Holotype (Fig. 5) measurements as follows. Body length 1094. Antenna (Fig. 6) with total funicle length 432; scape -, pedicel 58, fl, 69, fl, 78, fl, 71, fl, 64, fl. 56, fl. 54; clava 186, 0.47x funicle length. Fl, distinctly the longest segment and fl, the shortest. Fore wing (Fig. 7) length 948, width 312, length/width 2.72, longest marginal setae 188, venation (Fig. 8) length 395. Hind wing length 781, width 32. Ovipositor length 292, not exserted beyond apex of gaster. Comments. This species differs from E. kishenehn by the longer funicle segments (funicle 2.31x clava length cf. 1.83x in E. kishenehn) and slightly wider fore wing (length/width at most 3.49 instead of at least 3.64 in E. kishenehn). Derivation of species name. Named after Harold Borchers, professor of ento- mology and early mentor to the junior author at Bemidji State University, Bemidji, Minnesota. Gonatocerus Nees Figs 9-16 Gonatocerus is a worldwide group with numerous described extant species classified in several subgenera (Triapitsyn et al. 2010). Its members are often the most commonly collected Mymaridae in almost any habitat so it seems surprising that it has been re- corded only once as a fossil, in Baltic amber by Meunier (1905). The three species described below definitely belong to Gonatocerus based on the entire clava, 8-segmented funicle, similar sized gastral segments, short and probably narrow petiole, and 5-segmented tarsi (at least in the one species where they can be counted). Because they all appear to have a rhomboidal dorsellum they would be clas- sified either in G. (Gonatocerus) or in G. (Cosmocomoidea), but not G. (Lymaenon), the most common extant subgenus because it has a narrow, strap-shaped dorsellum or G. (Gastrogonatocerus) because the ovipositor does not project forward under the mesosoma. Gonatocerus kootenai Huber, sp. n. urn:lsid:zoobank.org:act:5 FFOE145-6ABD-43 1A-A540-8FEC3 130E934 http://species-id.net/wiki/Gonatocerus_kootenai Figs 9-11 Type material. Holotype female (NMNH) labelled “Gonatocerus kootenai Huber Hol- otype female #543761”. Description. Female. Colour dark brown except pedicel laterally, pronotum, tro- chanters, base and apex of femora, base and apex of tibiae (at least of fore and middle legs) and tarsomeres except apical tarsomere of all legs whitish. Holotype (Fig. 9) meas- urements as follows. Body length 1270. Head height 246, length 157. Antenna (Fig. 482 John T: Huber & Dale Greenwalt / ZooKeys 130: 473-494 (2011) Figures 9-10. Gonatocerus kootenai, holotype 9 habitus lateral 10 head and antennae. Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales... 483 10) with total funicle length 476; scape -, pedicel 57, fl, 54, fl, 57, fl, D 25h 59; fl. 69, fl. 64, fl 6l, fl, 60, clava 120. Mesosoma length 462. Fore wing length 1101, width 309, length/width 3.56, venation length 259, longest marginal setae 95. Hind wing length 729, width 35, longest marginal setae 67. Metasoma (Fig. 11) length 658. Ovi- positor length 588. Comments. Gonatocerus kootenai differs from the following two species by the relatively narrower wings and shorter funicle segments. I tentatively place it in G. (Gonatocerus) because of the relatively narrow fore wing and fairly long and relatively slender fl,—fl, ‘The reverse side of the shale piece contains a specimen of Trichoptera. Derivation of species name. Named after the Kootenai tribe of the Flathead Na- tion in northwestern Montana, site of the Kishenehn shales. Gonatocerus rasnitsyni Huber, sp. n. urn:lsid:zoobank.org:act:7 F2CF9FB-5 B06-4ABF-B62B-6043AB4179C7 http://species-id.net/wiki/Gonatocerus_rasnitsyni Figs 12-13 Type material. Holotype female (NMNH) labelled “Gonatocerus rasnitsyni Huber Holotype female #543762”. Description. Female. Colour dark brown except middle leg (others less clearly vis- ible) with coxa, trochanter, apex of femur and base of tibia, and basal 4 tarsal segments yellowish. Holotype (Fig. 12) measurements as follows (measured in NMNH only). Body length 1046. Antenna (Fig. 13) with total funicle length 414: scape length and width 144/46, pedicel -, fl, 44, fl, 43, fl, 55, fl, 51, A, 65, fl, 56, fl, 58, fl, 50, clava 113. At least on one antenna it appears that fl, is wider than fl, and fl, Mesosoma length (excluding pronotum) 375. Fore wing length 789 (775 from margin of mesosoma), width 243, length/width 3.25. Comments. Gonatocerus rasnitsyni differs from G. kootenai by the relatively smaller size, shorter and wider fore wing, and colour of the antennal pedicel and legs. I tenta- tively place it in G. (Cosmocomoidea) because of the relatively wider fl, (on one antenna at least, suggesting it bears multiporous plate sensilla) compared to the slightly nar- rower fl, and fl, This resembles several extant members of this subgenus that also have alternately wider and narrower basal funicle segments. Derivation of species name. Named in honour of A. P. Rasnitsyn, the world’s foremost Hymenoptera palaeontologist, on the occasion of his 75" birthday. 484 John T: Huber & Dale Greenwalt / ZooKeys 130: 473-494 (2011) * a eee eS Ss Figs 11, 12. Gonatocerus spp. 11 G. kootenai holotype, posterior half of mesosoma and metasoma 12 G. rasnitsyni holotype, habitus lateral. Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales... 485 Ae ¢ a 3 vi o i Gi Oh, 1000 [im aes : , - 14 ————————————————— SSS Figs 13, 14. Gonatocerus spp. 13 G. rasnitsyni holotype, mesosoma, head, antennae 14 G. greenwalti holotype, habitus lateral. 486 John T: Huber & Dale Greenwalt / ZooKeys 130: 473-494 (2011) Gonatocerus greenwalti Huber, sp. n. urn:lsid:zoobank.org:act:957095F5-B63B-4FB5-9FF0-OACD 1B029730 http://species-id.net/wiki/Gonatocerus_greenwalti Figs 14-16 Type material. Holotype female (NMNH) labelled “Gonatocerus greenwalti Huber Holotype female #543763”. Description. Female. Colour dark brown except apex of pedicel and legs beyond coxae lighter (yellowish). Holotype (Fig. 14) measurements as follows. Body length 926. Antenna (Fig. 15) with total funicle length 398; scape -, pedicel 51, fl, 35, fl, 4D. fl, 57, fl 54, fl. 56, fl a8 fl 56, fl, 54, clava 116. Mesosoma length 410. Fore wing (Fig. 16) length 730, width 273, length/width 2.67, longest marginal setae 58. Fore wing seeming- ly bare (without microtrichia) behind and just apical to the venation. Hind wing length 458, width 23, longest marginal setae 73. Metasoma length 460. Ovipositor length 417. Comments. G. greenwalti differs from G. kootenai and G. rasnitsyni by the wider fore wing and thicker funicle segments. ‘The apparent absence of microtrichia behind the venation, the wide fore wing, and fairly uniformly thick funicle segments suggest that G. greenwalti should be classified in G. (Cosmocomoidea). The shale fragment in which the fossil occurs contains several aquatic insects (e.g., Notonectidae), an indication of the lacustrine environment in which the mymarid lived (though it is not aquatic itself). Derivation of species name. Named by the senior author in honour of the junior au- thor, Dale Greenwalt, who collected and curated the insect fossils from Kishenehn shale. Eoanaphes Huber, gen. n. urn:lsid:zoobank.org:act: 1 DF7117D-B881-487B-94D8-63BCAA207F8F http://species-id.net/wiki/Eoanaphes Figs 17, 18 Description. Female. Body about 700. Head. About as wide as high (measured in frontal view). Malar space long, almost eye height. Antenna. Funicle 6-segmented, with fl, very short and remaining segments longer than wide; clava 3-segmented with the claval sutures almost perpendicular to claval length. Wings. Fore wing wide, slight- ly asymmetrical, with somewhat truncate apex; marginal fringe much shorter than fore wing width. Venation almost 1/3 fore wing length, with marginal vein fairly short, about as long as stigmal vein. Hind wing narrow; marginal fringe much longer than wing width and along posterior margin extending to base of membrane. Mesosoma. About 0.75 x gaster length. Mesoscutum length shorter than scutellum. Metanotum much shorter than scutellum, apparently with strap-like dorsellum. Metasoma. Ap- parently slightly constricted at base, probably with fairly wide, short petiole. Gastral terga similar in length. Ovipositor moderately short, apparently arising near base of gaster and its apex not exserted beyond gastral apex. Type species. Eoanaphes stethynioides Huber, sp. n. Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales... 487 Figs 15, 16. Gonatocerus greenwalti holotype: antennae 16 wings. 488 John T. Huber & Dale Greenwalt / ZooKeys 130: 473-494 (2011) ie r ¥ ’ Waa Lyre sugmal ee He. A ty 60% of all the fossil insects are Corixidae and Chironomidae). Because the climate and habitat of both Baltic and Kishenehn fossils was apparently similar it is not surprising that the genera of Mymaridae found in each are similar. Meunier (1901) described most Baltic amber fossils of Mymaridae. Doutt (1973) described a few species from Mexican amber (Simojovel, Chiapas), dated as 15—20 my, and ‘Thuréczy (1983) described one species. There is no indication from morphology that any of these or the mymarid compression fossils form a link between Cretaceous and Tertiary/Quaternary genera and species. Instead, they add support to the analysis by Rasnitsyn and Kulicka (1990) that showed Hymenoptera assemblages from Baltic amber seem to be more similar to the extant fauna than to the Late Cretaceous fauna. One clear morphological tendency from Cretaceous to Tertiary and Quaternary (present) species can be seen — a reduction in the number of funicle or claval seg- ments in females. A 3-segmented clava occurs in three of five (60%) Cretaceous genera, four of ten (40%) previously reported Tertiary genera from amber and shale (reported above), and 13 of 103 (about 8%) currently recognized extant genera. An 8-segmented funicle occurs in three of five (60%) Cretaceous genera, one of the 10 (10%) previ- ously reported Tertiary genera, and five of the 103 (about 5%) extant genera. Except for Gonatocerus, which has eight funicle segments but an entire clava, all other Tertiary fossils have six (rarely fewer) funicle segments in females. Spahr (1987) catalogued the literature and listed 11 mymarid genera from amber, three of them Cretaceous and eight Tertiary. He followed previous authors by including genera and species now correctly classified in Mymarommatidae (Gibson et al. 2007). Witsack (1986) described Palaeopatasson for one species from Dominican amber. The senior author examined 16 Baltic amber pieces (SVT, UCRC) in 2005, 13 pieces (AMNH) in 2011, and three pieces IPMG6., ZMUC) in about 2005. Only a few specimens could be referred confidently to an extant genus. These were four Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales... 491 specimens of Gonatocerus, and (less confidently) three specimens of Anaphes and two of Stethynium. If correctly identified, five more genera, all known from the extant fauna, are reported here for the first time as amber fossils. Having examined these 32 addi- tional amber specimens and realizing that most of them cannot definitely be classified in an extant genus, the senior author has doubts about the correct generic placement of at least some of the specimens studied by past workers. None of those specimens were examined, however. Each Tertiary fossil genus is discussed briefly below. Anagroidea (1 female, C.V. Henningsen, B-1 1956, ZMUC, examined) is no long- er known from extant species in Europe and is rare in the Holarctic region (eastern Asia and south east USA). Alaptus is known from at least two species in Miocene amber (Doutt 1973) that are probably correctly identified to genus. Anaphes was known from three specimens in Baltic amber. Anaphes splendens Meunier is probably incorrectly classified. According to Meunier the ovipositor ex- tends appreciably beyond the apex of the metasoma, unlike any extant Anaphes. The fore wing with a slight but distinct ventral lobe, its overall shape, the venation with a distinct stigmal vein, and the antenna with fl, almost as long as fl, also do not fit the genus. Anaphes schellwieniens Meunier is described from a male so it is impossible to determine its placement. Doutt (1973) examined a male from Chiapas amber and tentatively placed it in Anaphes. Three specimens examined here are fairly confidently placed in Anaphes (1 female, UCRC; 2 females, SVT). Additional specimens (2 fe- males, SVT) examined are tentatively placed in Anaphes. Arescon is known from two species in amber (Meunier 1901, 1905) that are probably correctly placed because of their 5-segmented funicle, though fore wing venation length, which is important information for correct generic placement, was not given by Meunier. ?Camptoptera (1 female, SVT), examined in 2005, if correctly classified, would be the first fossil specimen for this worldwide genus. ?>Dorya (1 female, UCRC) is known only from extant species in Australia and New Zealand. ?Eustochus (1 female, AMNH). The gaster appears to be petiolate. This suggests a Eustochus with a distinct 3-segmented clava (most have a 2-segmented clava) though the wing shape and antenna suggests Eoeustochus. ?Mimalaptus (3 females, SVT; 2 females, UCRC; 1 female, box G 3.910 Hymen- optera #BST03124, IPMG6.) is known from extant species in Australia and New Zea- land and possibly eastern Asia. Gonatocerus was known from only one fossil specimen, Gonatocerus henneberti (Me- unier 1905), which appears to belong to either G. (Gonatocerus) or G. (Lymaenon) as suggested by the narrow wings. However, the “écusson semilunaire” [semilunate shield], which I interpret from Meunier’s drawing as being the dorsellum, appears to be rhom- boidal in shape, which would eliminate G. (Lymaenon) as the correct subgenus. The small size of the specimen, only 1/3 mm long, is unusual; it is much smaller than the smallest extant Gonatocerus | have seen. A specimen of Gonatocerus (Cosmocomoidea) is reported here (1 male, Poinar collection, OSU) from Dominican amber, 15—20 my. 492 John T. Huber & Dale Greenwalt / ZooKeys 130: 473-494 (2011) Litus is represented by two fossil species. Litus elegans Meunier (1901) may or may not be correctly classified because the description and illustration are insufficient to place it. It apparently differs considerably from Litus mexicanus Doutt (1973), which is likely correctly classified. Extant Litus species tend to have a bi-geniculate funicle, unlike that illustrated by Meunier. Litus beneficus Meunier from copal (recent) from Madagascar also needs to be re-examined; it is unlikely to be classified correctly. Malfatia molitorae Meunier (1901) is based on a male and from the description alone it is impossible to determine how it relates to extant genera. Its status remains uncertain. Palaeopatasson grollei Witsack (1986) is based on a female and may be related to Anaphes. Polynemoidea mexicana Doutt (1973) from Chiapas amber was stated to be similar to P domestica Girault, but the latter species is likely incorrectly placed in Polynemoidea (Lin et al. 2010). Consequently, I suspect that P mexicana is also incorrectly classified; it does not have a strongly exserted ovipositor as in the type species of Polynemoidea. Stethynium (1 female, S. Triapitsyn collection), if correctly identified, confirms that the genus occurs in Baltic amber, as reported by Thuréczy (1983). Conclusions 1. Tertiary fossils of Mymaridae are clearly more related to the extant fauna than to the Cretaceous fauna. 2. Except for Gonatocerus and the two new genera described above, tertiary fossils of Mymaridae (except perhaps Anaphes and Stethynium) are for the most part doubt- fully assigned to extant genera. 3. Several Eocene amber fossils, if correctly identified to genus, represent genera that no longer occur in Europe (Anagroidea, Dorya, Mimalaptus) or are very poorly represented in most of the Holarctic region (Stethynium). These genera are now distributed mostly in the tropics or southern hemisphere, supporting observations that the Eocene climate (in parts of the Holarctic region at least) was considerably warmer than at present. 4. Ifcorrectly identified, Anagroidea, Eustochus, and perhaps Eoeustochus would be the first fossil records of genera belonging to the extant tribe Mymarini sensu Annecke and Doutt (1961), likely the most derived lineage in Mymaridae. Acknowledgements The authors thank the authorities of NMNH for permission to work on the fossils and use of their equipment. The senior author thanks S. Triapitsyn (UCRC) for letting him examine the fossils from his personal collection and the ones in UCRC during a visit to Riverside in 2005. J.W. Jansen (Sevetal, Germany) sent fossils from his private Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales... 493 collection for study of the senior author. These were then purchased by AMNH, where they will be deposited. References Annecke DP, Doutt RL (1961) The genera of the Mymaridae. Hymenoptera: Chalcidoidea. Entomology Memoirs. Department of Agricultural Technical Services, Republic of South Africa 5: 1-71. 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