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January 2007 Phylogeny and Geological History of the Cynipoid (: ) Zhiwei Liu Eastern Illinois University, [email protected]

Michael S. Engel University of Kansas, Lawrence

David A. Grimaldi American Museum of Natural History

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Recommended Citation Liu, Zhiwei; Engel, Michael S.; and Grimaldi, David A., "Phylogeny and Geological History of the Cynipoid Wasps (Hymenoptera: Cynipoidea)" (2007). Faculty Research & Creative Activity. 197. http://thekeep.eiu.edu/bio_fac/197

This Article is brought to you for free and open access by the Biological Sciences at The Keep. It has been accepted for inclusion in Faculty Research & Creative Activity by an authorized administrator of The Keep. For more information, please contact [email protected]. PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3583, 48 pp., 27 figures, 4 tables September 6, 2007

Phylogeny and Geological History of the Cynipoid Wasps (Hymenoptera: Cynipoidea)

ZHIWEI LIU,1 MICHAEL S. ENGEL,2 AND DAVID A. GRIMALDI3

CONTENTS

Abstract ...... 1 Introduction ...... 2 Systematic Paleontology ...... 3 Superfamily Cynipoidea Latreille ...... 3 Protimaspidae Liu and Engel, new family ...... 4 Family Stolamissidae Liu and Engel, new family ...... 7 Family Ashmead ...... 8 Family Thomson...... 13 Family Cynipidae Latreille...... 23 Cynipoidea Incertae Sedis ...... 29 Cladistics...... 29 Methods...... 29 Results...... 31 Discussion ...... 37 Acknowledgments ...... 43 References ...... 43 Appendix 1: Described Cynipoidea...... 47

1 Department of Biological Sciences, Eastern Illinois University, 600 Lincoln Avenue, Charleston, IL 61920 ([email protected]). 2 Division of Invertebrate , American Museum of Natural History; Division of Entomology (Paleoentomology), Natural History Museum, and Department of Ecology & Evolutionary Biology, 1501 Crestline Drive–Suite 140, University of Kansas, Lawrence, KS 66049-2811 ([email protected]). 3 Division of Invertebrate Zoology, American Museum of Natural History ([email protected]).

Copyright E American Museum of Natural History 2007 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3583

ABSTRACT The geological history of the superfamily Cynipoidea is reviewed, with the description of various new taxa, being mostly in Late from New Jersey and Canada. The various fossil lineages are incorporated into a phylogenetic analysis of the superfamily, and their implications for understanding the evolution of the group are explored. The following new taxa or taxonomic changes are proposed (authorship of all taxa is Liu and Engel): Protimaspidae, new family; Stolamissidae, new family; Stolamissus, new ; Stolamissus mirabilis, new ; Proliopterinae, new ; Proliopteron, new genus; Proliopteron redactus, new species; Goeraniinae, new subfamily; Goerania, new genus; Goerania petiolata, new species; Micropresbyteria, new genus; Micropresbyteria caputipressa, new species; Anteucoila, new genus; Anteucoila delicia, new species; Jerseucoila, new genus; Jerseucoila plesiosoma, new species; Syneucoila, new genus; Syneucoila magnifica, new species; Tanaoknemus, new genus; Tanaoknemus ecarinatus, new species; Kinseycynips, new genus; Kinseycynips succinea (Kinsey), new combina- tion. The extinct family Rasnicynipidae is newly transferred to Figitidae and classified as a basal subfamily therein (Rasnicynipinae, status novus). The Gerocynipidae, its type genus Gerocynips, and the type species upon which they are founded, Gerocynips zherichini, are found to be nomenclaturally unavailable. Gerocynips zherichini is regarded as a nomen nudum; the genus as newly validated is Gerocynips, new genus (with G. siberica Kovalev as type species); and the family as validated is Gerocynipidae, new family. The fossil records of Cynipoidea are summarized.

INTRODUCTION syndrome of parasitizing wood-boring hosts and are instead of various holo- Wasps of the superfamily Cynipoidea metabolan larvae ranging from to lace- (: Proctotrupomorpha) are one of wings, and some even victimize cynipids. Still the more remarkable lineages among the other figitids are hyperparasitoids of braconid Hymenoptera. The superfamily and chalcidoid primary parasitoids of is relatively small by comparison to the and psyllids (Buffington et al., 2005). Most more diverse lineages of apocritan wasps, with remarkably, however, the Cynipoidea in- presently ca. 3000 described species (Ronquist, cludes, aside from these parasitic lineages, 1999). However, Nordlander (1984) has esti- secondarily phytophagous species, namely mated that there are may be as many as 20,000 the gall wasps (family Cynipidae). Indeed, cynipoid species, and intensive fieldwork and the gall wasps are one of the three most monographic studies are desperately needed, notable transitions within the Hymenoptera particularly in species-rich regions such as from a parasitoid or predatory life-history to Mexico. Basal lineages of Cynipoidea are a vegetarian habit (the other two being the fig generally parasitoids of wood-boring wasps () and the (Anthophila), larvae, such as immatures of various beetle although others do, of course, exist). Cynipids families, oecophorid moths, or siricid wood produce as larvae characteristic galls in wasps (Liu and Nordlander, 1994; Ronquist, angiosperm tissue, the most famous of which 1995a; Liu, 2001; Liu et al., 2007). This are the gall wasps developing in various primitive parasitoid biology is, indeed, the tissues of species of the genus Quercus plesiomorphic condition for the parasitoid (Fagaceae). All cynipids feed on dicotyledon- Hymenoptera (Euhymenoptera) as a whole ous angiosperms except for a single species, (Grimaldi and Engel, 2005). Not surprisingly, Diastrophus smilacis Ashmead, which induces the rather robust, parasitoid lineages of galls on the monocotyledonous Smilax Cynipoidea (a.k.a. ‘‘macrocynipoids’’) form (Krombein et al., 1979; Ronquist and a grade to the so-called ‘‘microcynipoids.’’ Liljeblad, 2001). Phytophagous of The microcynipoids consist of two extant other cynipids have also evolved within the families, the Figitidae and the Cynipidae. family (Ronquist, 1994). The Cynipoidea Figitids diverge biologically from the primitive clearly encompasses an amazing range of 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 3 biological variation and significant evolution- from Medicine Hat, Alberta, Canada ary transitions making them worthy of in- (Campanian). In addition, we provide critical vestigation. new data for Kinsey’s fossil species. Although The Cynipoidea have long been assumed to many of the cynipoid from Eurasia are be a natural group. Although not supported well preserved and suitable for comparison by any striking autapomorphy, recent studies with extant cynipoids, particularly those en- have provided morphological evidence for tombed in amber, these species were only monophyly. Ronquist (1999) listed the follow- recently placed into a phylogenetic framework ing features as synapomorphic for the super- (Ronquist, 1995b, 1999). None, however, was family: absence of radicle; forewing media (M) included in cladistic analyses with modern taxa, displaced anteriorly, approaching the posteri- and we therefore have for the first time or end of the marginal cell, distinctly angled attempted to incorporate such paleontological and not running parallel to the posterior wing data into analyses of higher level cynipoid margin; abdominal sterna II (petiolar) and III phylogeny. With this wealth of new informa- (first postpetiolar) abutting or fused; metaso- tion we provide a new overview of the ma distinctly laterally compressed; and fore- phylogeny and geological history of the wing costa (C) absent. Ronquist (1999) also Cynipoidea. discussed several other possible autapomor- Terminology for the descriptions follows phies for the superfamily while Ronquist that of Ronquist and Nordlander (1989) and (1995b, 1999), Ronquist and Nieves-Aldrey Ronquist (1995a), although wing vein termi- (2001), Va˚rdal et al. (2003), and Nieves-Aldrey nology follows that more widely used across et al. (2004) have summarized the phylogeny the Hymenoptera, and at large, so as and classification. to highlight homologies (e.g., ‘‘first free abscis- Fossils of Cynipoidea are, unfortunately, sa of M’’ is used in place of ‘‘basalis’’ or ‘‘basal rare and only a handful of species have been vein’’, although we denote the latter in paren- definitively assigned to the superfamily (see theses to avoid confusion). Throughout we appendix 1). The first fossils assigned to number segments according to their metasomal Cynipoidea were described by Dr. Alfred C. number rather than to abdominal count owing Kinsey from material preserved in Baltic and to the fusion of the first abdominal segment into the posterior of the thorax. Metrics are Canadian amber (Kinsey, 1919, 1937). The to be considered approximate because the latter fossil, Protimaspis costalis Kinsey, from optimal angle for specific measurements was the Late Cretaceous (Campanian) amber of not always attainable owing to the position Cedar Lake, Manitoba, Canada, was particu- of the specimen in the amber. The abbrevia- larly notable as it was hitherto the oldest, tions F, S, and T are occasionally used for definitive fossil cynipoid. For most of the 20th flagellomere, metasomal sternum, and metaso- century Kinsey’s species stood alone as the mal tergum, respectively, and AMNH, RTMP, geological evidence for the Cynipoidea. ROM, and CNCareusedforAmericanMuseum Relatively recently, however, Oleg V. Kovalev of Natural History (New York), Royal Tyrell (1994, 1995, 1996) described a small wealth of Museum of Paleontology (Drumheller, Canada), cynipoids from deposits of Eurasia, greatly Royal Ontario Museum (Toronto, Canada), and expanding the fossil record of the super- the Canadian National Collection (Ottawa, family. This work included several new Late Canada), respectively. Cretaceous species in mudstones of the Ola Formation (Cenomanian) along the banks of Obeshchayushchiy Creek in the Russian Far SYSTEMATIC PALEONTOLOGY East and in amber from the Kheta Formation SUPERFAMILY CYNIPOIDEA LATREILLE of the Taimyr Peninsula (Santonian), as well as additional taxa from (middle DIAGNOSIS: Small to medium-sized Eocene) (appendix 1). Herein we describe eight without metallic luster. filiform, new North American fossils of Cynipoidea usually with 11–12 and 12–13 flagellomeres from the Late Cretaceous amber of Middlesex in female and male, respectively; radicle County in central New Jersey (Turonian) and absent. Pronotum posteriorly reaching tegula; 4 AMERICAN MUSEUM NOVITATES NO. 3583 mesoscutellum in lateral view prominent. TABLE 1 Forewing usually present and with distinctive Hierarchical Classification of Superfamily triangular radial cell; C always absent; ptero- Cynipoidea stigma always absent except in Austrocynips; (Updated from Ronquist, 1999, and Ronquist and M displaced anteriorly, approaching posterior Nieves-Aldrey, 2001) end of marginal cell, distinctly angled and not running parallel to posterior wing margin. Superfamily CYNIPOIDEA Latreille, 1802 ‘‘macrocynipoids’’ [paraphyletic] Tarsi pentamerous (i.e., 5-segmented). Meta- Family AUSTROCYNIPIDAE Riek, 1971 soma almost always laterally compressed; Family Thomson, 1862 metasomal sterna I (petiolar) and II (first Subfamily Eileenellinae Kovalev, 1994 postpetiolar) abutting or fused. Subfamily Ibaliinae Thomson, 1862 COMMENTS: The superfamily Cynipoidea Family {PROTIMASPIDAE Liu and Engel, new family presently comprises five extant and three Family {STOLAMISSIDAE Liu and Engel, new family extinct families, two of which are newly Family LIOPTERIDAE Ashmead, 1895 described herein. A summary of the classifica- Subfamily {Proliopterinae Liu and Engel, new tion of Cynipoidea is provided in table 1. Weld subfamily (1952) provided the most recent monograph of Subfamily Mayrellinae Hedicke, 1922 Subfamily Dallatorrellinae Kieffer, 1911 the world genera, but it is now largely out of Subfamily {Goeraniinae Liu and Engel, new date for most groups. Several works have subfamily examined the higher classification, establishing Subfamily Liopterinae Ashmead, 1895 relationships and redefining the families, sub- Subfamily Oberthuerellinae Kieffer, 1903 families, and tribes (e.g., Ronquist, 1995b, ‘‘microcynipoids’’ 1999; Nieves-Aldrey et al., 2004). In addition, Family {GEROCYNIPIDAE Liu and Engel, new family some groups have been newly monographed to Family FIGITIDAE Thomson, 1862 the level of species – e.g., Liu and Nordlander Subfamily {Rasnicynipinae Kovalev, 1996, new (1992, 1994) and Liu (1998b) for Ibaliidae; status Ronquist (1995a), Liu (2001), and Liu et al. Subfamily {Palaeocynipinae Kovalev, 1994 Subfamily Parnipinae Ronquist and Nieves-Aldrey, (2007) for Liopteridae, the first for world 2001 genera, the latter two for species of Subfamily Thrasorinae Kovalev, 1994 Dallatorrellinae and part of Mayrellinae; Subfamily Charipinae Dalla Torre and Kieffer, 1910 Nieves-Aldrey (2001) for Cynipidae of Iberia; {Protocharipini Kovalev, 1994 and Ronquist and Nieves-Aldrey (2001) for Tribe Alloxystini Helle´n, 1931 Parnipinae (Figitidae). Tribe Charipini Dalla Torre and Kieffer, 1910 Subfamily Anacharitinae Thomson, 1862 Subfamily Figitinae Thomson, 1862 Protimaspidae Liu and Engel, new family Subfamily Eucoilinae Thomson, 1862 Subfamily Pycnostigminae Cameron, 1905 TYPE GENUS: Protimaspis Kinsey, 1937. Subfamily Aspicerinae Dalla Torre and Kieffer, 1910 DIAGNOSIS: The new family shares with Tribe {Palaeoaspicerini Kovalev, 1994 Ibaliidae two unique characters that are not Tribe Aspicerini Dalla Torre and Kieffer, 1910 found in any other family of Cynipoidea: (1) Subfamily Emargininae Kovalev, 1994 mesopleuron with an oblique, strongly im- Family CYNIPIDAE Latreille, 1802 pressed femoral groove; (2) metacoxa dorsally Subfamily {Hodiernocynipinae Kovalev, 1994 strongly impressed longitudinally, anterolat- Subfamily Latreille, 1802 erally with a triangular crest. The family can Tribe Aylacini Ashmead, 1903 be easily distinguished from the Ibaliidae by Tribe Latreille, 1802 the absence of a pair of submedian processes Tribe Eschatocerini Ashmead, 1903 posteriorly on the mesoscutellum; the relative- Tribe Pediaspidini Ashmead, 1903 ly small area of the pronotum behind the Tribe Latreille, 1802 Tribe Ashmead, 1896 anterior vertical plate; the metafemur distinct- ly longer than the metacoxa; the short marginal cell of the forewing, about four times as long as wide; and the female tergum VII not enlarged. 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 5

Fig. 1. Photomicrograph of holotype female of Protimaspis costalis Kinsey (ROM) in Canadian amber from Cedar Lake.

COMMENTS: Ronquist (1999) mentioned DIAGNOSIS: As for the genus (see above). that the wing venation of Protimaspis is DESCRIPTION: The placement of this taxon ‘‘remarkably similar to that of Rasnicynips’’, has historically been hindered by several but this statement is obviously incorrect. The factors. For example, Kinsey (1937) in his vein Rs+M is directed to the posterior end of description of the species stated that the sex of the first free abscissa of M (5 basalis vein) in the fossil ‘‘is not quite clear’’, but we have no Rasnicynips (Kovalev, 1994: as Rasnitsynia doubt that this is a female. Certainly more therein), but to the middle of the latter vein in details of the specimen can be observed owing Protimaspis (Kinsey, 1937) (figs. 1, 2). to the advances in methods of preparation as well as in optical equipment available. Genus Protimaspis Kinsey Furthermore, Kinsey’s original description used characters that are now understood to Protimaspis Kinsey, 1937: 22. Type species: be superficial. We therefore provide the Protimaspis costalis Kinsey, 1937. following new description for the species. Female. Body length 1.9 mm; forewing DIAGNOSIS: As for the family (see above). length 1.8 mm. Antenna 15-segmented, with distinct placodeal sensilla on all flagellomeres Protimaspis costalis Kinsey except F1; F1 slightly bent medially. Frons figures 1, 2 flat, without carinate structures. Lateral part of cranium distinctly expanded behind com- Protimaspis costalis Kinsey, 1937: 22. pound eye; gena broad; occipital carina 6 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 2. Holotype female of Protimaspis costalis Kinsey (ROM). 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 7 absent. Median dorsal length of pronotum free abscissa of M (5 basal vein) at about the behind anterior vertical plate short; pronotal middle; and, (7) posterior margin of metaso- carina absent; pronotal crest low and not mal T2 not distinctly oblique. incised medially, and dorsal pronotal area present and short; posteroventral margin of Stolamissus Liu and Engel, new genus pronotum long and straight; lateral surface of pronotum glabrous. Mesonotum transversely TYPE SPECIES: Stolamissus mirabilis Liu and costate; median mesoscutal impression and Engel, new species. notauli absent; mesoscutellar suture trans- DIAGNOSIS: As for the family (see above). verse; mesoscutellum posteriorly sloped, with- ETYMOLOGY: The new genus-group name out projection or process; mesopleuron ven- is derived from the Latin stolo (meaning trally protruding; mesocoxa directed obliquely ‘‘branch’’) and amissus (meaning ‘‘lost’’), backward, inserted on separate, oblique pos- which together mean ‘‘lost branch’’ (a refer- terior area; median mesopleural impression ence to the fact that this clade was lost via and lateroventral mesopleural carina absent; extinction within Cynipoidea). The name is mesopleural triangle slightly depressed with masculine. longitudinal striation; mesofemural groove COMMENTS: The genus has several of the present, granulate. Metapleural sulcus abrupt- symplesiomorphies that are shared among the ly bent at middle; metepisternum subrectan- macrocynipoids, including marginal cell of gular. Propodeal spiracular opening not com- forewing relatively long and anteriorly closed, pletely covered by spiracular process, similar no bulla in Sc+R1, remnant of pterostigma to extant when observed laterally; short and thick, and second metasomal lateral propodeal carina present; nucha short. tergum (5 third abdominal tergum) large with Forewing lacking pterostigma; marginal cell succeeding abdominal terga narrow. In addi- closed, four times as long as wide; bulla in tion, the pronotum is distinctly raised dor- Sc+R1 present; 2r-rs directed obliquely, slop- soanteriorly, a condition similar to that of ing outward posteriorly; Rs+M with mesal end Ibaliidae and Liopteridae, and the well-de- meeting first free abscissa of M (5 basal vein) veloped lateral pronotal carina resembles that slightly behind the middle; areolet present. of the liopterids. On the other hand, it appears Metacoxa dorsally strongly depressed and that the mesocoxae are inserted vertically lateroanteriorly with a triangular process. downward, a feature shared by the micro- Petiolar annulus crescentic and glabrous; cynipoids, but not obliquely on a ventrally metasomal T4 largest among all terga, latero- protruding, separate, and oblique posterior posterior margin more or less vertical and area as in the macrocynipoids. The mesopec- straight; metasomal T3–6 subequal along tus appears not to protrude ventrally, a feature middorsal line. Male. Unknown. considered a synapomorphy for the micro- HOLOTYPE: Female, Late Cretaceous (Cam- cynipoids (Ronquist, 1995a, 1999); however, panian), Cedar Lake, Manitoba, Canada; de- we are somewhat cautious about this obser- positedintheRoyalOntarioMuseum. vation because of the presence of a bubble laterally above the mesopectal area, and our Stolamissidae Liu and Engel, new family interpretation of this feature may be in error.

TYPE GENUS: Stolamissus Liu and Engel, Stolamissus mirabilis Liu and Engel, new genus. new species DIAGNOSIS: This new family is distin- figures 3, 4 guished from other cynipoids by the apo- morphic combination of the following traits: Cynipidae sp.: Grimaldi et al., 2000: 73 [figured]. (1) mesosoma short and high in lateral view; (2) pronotal crest and dorsal pronotal area DESCRIPTION: Female. Body length 0.8 absent; (3) mesocoxa directed vertically and mm; forewing length 0.7 mm. Body entirely downward; (4) lateral pronotal carina distinct; black; antennae and legs dark brown; wings (5) pterostigma lacking (figs. 3, 4); (6) Rs+M hyaline, without any macula or band. Antenna of forewing with mesal end meeting the first 14-segmented; flagellum distinctly expanded 8 AMERICAN MUSEUM NOVITATES NO. 3583

area (the mesopectal area is partly obscured by a large bubble and therefore our interpre- tation of the condition should be considered tentative). Propodeum devoid of processes. Wings hyaline, surface covered with appressed setae, margins ciliate. Forewing with marginal cell closed, slightly more than twice as long as wide; bulla in Sc+R1 absent, 2r-rs oblique, sloping outward posteriorly; areolet present; Rs+M arising from about middle of first free abscissa of M (5 basal vein). Mesofemur expanded dorsoventrally; first metatarsomere much shorter than combined length of second through fifth metatarsomeres; metapretarsal claw with basal lobe. not distinctly compressed; metasomal T2 (5 third abdomi- nal tergum) relatively large, about one-third length of postpetiolar metasoma; metasomal T3–7 dorsally subequal. Male. Unknown. HOLOTYPE: Female, AMNH NJ-709, Late Cretaceous (Turonian), White Oak Pit, Sayreville, Middlesex County, New Jersey, Fig. 3. Photomicrograph of holotype female of coll. AMNH expedition, 1996; deposited in Stolamissus mirabilis Liu and Engel, new genus and the Amber Fossil Collection, Division of species (NJ-709). Invertebrate Zoology, AMNH. ETYMOLOGY: The specific epithet is the Latin word mirabilis, meaning ‘‘beautiful and toward apex; scape and pedicel subequal in good-looking’’. both length and thickness, slightly longer and much wider than F1; F1 distinctly longer than FAMILY LIOPTERIDAE ASHMEAD F2; apical and subapical flagellomeres some- what fused with each other, other flagello- DIAGNOSIS: Ronquist (1995a) recently re- meres subequal to F1; elongate placodeal vised the family and suggested the following sensilla distinctly present on F8–12. Frons, characters as apomorphies: lateral surface of vertex, and gena glabrous; median frontal pronotum and dorsal surface of mesoscutel- carina absent; gena not expanded behind lum foveate; acetabulum more or less vertical, compound eye; length of compound eye about divided into two furrows for procoxa by three times as long as malar space. Anterior a strong median keel; acetabular carina de- plate of pronotum with dense punctures and scribing a v-shape; metapleural sulcus reach- posteriorly delimited by prominent lateral ing anterior metapectal margin far above the pronotal carinae; lateral surfaces of pronotum midheight; intermetacoxal processes present; irregularly carinate in lower part and narrowly lateral pronotal carina reaching the raised bridged medially. Mesoscutum transversely ventral pronotal margin; laterodorsal process costate; median impression present in posteri- of mesoscutellum present; lateroventral carina or two-thirds; notauli percurrent and promi- of mesopectus present; nucha long; metatibia nent; mesoscutellar foveae transverse; meso- shorter than metafemur; petiolar annulus scutellum posteriorly sloped and without pro- complete, tergal and sternal parts fused cess; mesopleuron mostly glabrate, ventrally completely; occipital carina present; and depressed along lower margin; median impres- mesopleural impression present. sion and lateroventral carina absent; meso- COMMENTS: Where known, extant Liop- pectus not distinctly protruding ventrally, teridae are parasitoids of wood-boring beetles mesocoxa directed vertically downward and of families such as Buprestidae, Cerambycidae, not inserted on separate, oblique posterior and Curculionidae (Ronquist, 1995a; Liu et al., 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 9

Fig. 4. Holotype female of Stolamissus mirabilis Liu and Engel, new genus and species (NJ-709). 10 AMERICAN MUSEUM NOVITATES NO. 3583

2007). The genera of Liopteridae were revised cess, and the metatibial lobe. Some of the by Ronquist (1995a) and an analysis of their features of the genus are also shared by relationships was undertaken. The genera of the Mesocynips, including the lack of foveate family are presently segregated into four sub- sculpturing on the pronotum and mesoscutel- families: the Liopterinae (New World) and lum and the absence of the posterior mesos- Oberthuerellinae (African) are sisters, with cutellar process. Dallatorrellinae (Asian and Australian) as sister ETYMOLOGY: The genus-group name is to them, and the Mayrellinae (cosmopolitan a combination of pro- (Latin, meaning ‘‘be- except the Australian region) basal. fore’’) and Liopteron, the type genus of the According to our cladistic analyses the new family. The name is masculine. fossil genus Proliopteron is basal to a clade consisting of all other Liopteridae (see below). Proliopteron redactus Liu and Engel, In addition, the genus Goerania, described new species below, obviously belongs to the clade consist- figures 5, 6 ing of the Oberthuerellinae + Liopterinae, although we have decided not to DIAGNOSIS: As for the genus (see above). include it in the cladistic analysis owing to the DESCRIPTION: Female. Body length 1.0 limited number of observable characters. Both mm; forewing length 0.9 mm. Body mostly genera are characterized by an array of black to dark brown; antenna and legs dark distinct features and therefore we herein erect brown; wings hyaline, without any macula or two new subfamilies, as Proliopterinae and band. Antennae cylindrical and slightly ex- Goeraniinae, to accommodate them. panded toward apex, 12-segmented; pedicel almost spherical, half as long as scape; F1 as long as F2; other flagellomeres subequal to Proliopterinae Liu and Engel, new subfamily F1, except apical flagellomere is twice as long TYPE GENUS: Proliopteron Liu and Engel, as preceding one; elongate placodeal sensilla new genus. present on all flagellomeres. Frons, vertex, DIAGNOSIS: As for the genus (see below). and gena glabrous; lower face with flat but distinct median carina complete to clypeus, Proliopteron Liu and Engel, new genus otherwise without apparent sculpture; gena not expanded behind compound eye; com- TYPE SPECIES: Proliopteron redactus Liu pound eye longer than malar space by one- and Engel, new species. third. Lateral surfaces of pronotum glabrous; DIAGNOSIS: The genus can be easily distin- lateral pronotal carina dorsally reduced. guished from all other genera in the family by Mesoscutum glabrate, without transverse cos- a combination of: (1) characteristic liopterid- ta; mesoscutellum posteriorly sloped without like sculpture on gena and lateral surface of process; mesopleuron glabrous and ventrally pronotum absent, (2) lateral pronotal carina bordered by straight lateroventral carina; dorsally very reduced, (3) posterior mesoscu- mesopectus not protruding ventrally, meso- tellar processes absent, (4) metatibial lobe coxa directed vertically and not inserted on absent, (5) pubescence of wings reduced such separate, oblique posterior area. Propodeum that it is represented by mere dots, and (6) devoid of processes; lateral propodeal carina mesocoxa inserted to mesopectus vertically, dorsally raised into a distinct right-angular not on a separate, projected area (a few lobe. Wings entirely hyaline, all wing margins modern species of Paramblynotus also have ciliate except anterior margin of forewing; this feature). pubescence of wings reduced and represented COMMENTS: Proliopteron is the sister clade by mere dots. Forewing with marginal cell to all other liopterids. The genus lacks several closed, four times as long as wide; bulla in of the synapomorphic features shared by other Sc+R1 present; 2r-rs oblique, sloping outward liopterids, including the characteristic foveate posteriorly; areolet absent; Rs+M arising from integumental sculpturing of the pronotum somewhat anterior to middle of first free and mesoscutellum, the median mesopleural abscissa of M (5 basal vein). First metatar- impression, the posterior mesoscutellar pro- somere slightly shorter than combined length 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 11

Fig. 5. Photomicrograph of holotype female of Proliopteron redactus Liu and Engel, new genus and species (CAS-409). of second through fifth metatarsomeres; meta- Goeraniinae Liu and Engel, new subfamily prestaral claws simple, without basal lobe. TYPE GENUS: Goerania Liu and Engel, new Metasoma inserted high on propodeum, post- genus. subpleuron long; metasomal T2–4 (5 abdom- DIAGNOSIS: As for the genus (see below). inal T3–5) subequal in size, metasomal T5 largest; metasomal T7 exposed, not complete- ly covered by metasomal T6; metasomal S3–5 Goerania Liu and Engel, new genus (5 abdominal S4–6) exposed, not covered by TYPE SPECIES: Goerania petiolata Liu and metasomal S2; ovipositor exserted, apically Engel, new species. with ventral serrations (fig. 6). Male. DIAGNOSIS: Median flagellomere of anten- Unknown. na cylindrical, with very dense longitudinal HOLOTYPE: Female, CNC CAS-409, Late placodeal sensilla. Head distinctly impressed Cretaceous (Campanian), Medicine Hat, posteriorly in dorsal view; gena expanded Alberta, Canada, coll. J.F. McAlpine, 1971- behind compound eyes; vertex, gena, and VII-8–10 [8–10 July 1971]; deposited in the occiput glabrous. Anterior pronotal flange CNC. long; submedian depression of pronotum pres- ETYMOLOGY: The specific epithet is derived ent and open laterally; dorsal pronotal area from the Latin word redactus meaning ‘‘re- distinct; pronotal crest not observable; lateral duced’’ and refers to the reduced pubescence surface of pronotum sloped evenly, without on the wing membranes. foveate sculpture. Mesoscutum transversely 12 MRCNMSU NOVITATES MUSEUM AMERICAN O 3583 NO.

Fig. 6. Holotype female of Proliopteron redactus Liu and Engel, new genus and species (CAS-409). 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 13

costate; median impression and lateral notauli DESCRIPTION: Female. Body length distinct; lateral margin of axilla conspicuously 3.0 mm, forewing length 2.5 mm. Antenna raised anteriorly. Forewing without pterostig- filiform, 14-segmented. Anterior plate of ma; 2r-rs long and perpendicular to anterior pronotum transversely costate. Mesoscutum wing margin; marginal cell closed; bulla in transversely weakly costate; median impres- Sc+R1 present; Rs+M issuing from close to sion distinctly present in posterior one-third; posterior two-thirds of first free abscissa of M notauli percurrent; parascutal carina postero- (5 basal vein). Mesocoxa inserted vertically; laterally smooth and not raised; mesoscutel- metatibia distinctly longer than metafemur and lum foveate-reticulate, posteriorly broadly with a longitudinal carina on dorsal surface. rounded; dorsal lateral process present; scu- Metasoma strongly compressed laterally; nucha to-scutellar suture transverse and separated low; petiolar annulus positioned low, long; into two foveae by a median carina. Forewing petiole at least twice as long as wide, longitu- marginal cell closed, 3.8 times as long as wide; dinally carinate; posterior margin of petiole and Rs+M proximally directed toward posterior anterior margin of metasomal T1 abutting, two-thirds of basal vein; areolet distinct. First dorsal margins of the two more or less metatarsomere slightly shorter than combined continuous (i.e., without constriction between length of second through fifth metatarso- segments); metasomal T2–7 of female visible meres. Petiolar annulus long, median dorsal and subequal in length along middorsal line. length at least two times as long as minimum dorsal width (estimated from lateral view). ETYMOLOGY: The genus is named after Dr. Go¨ran Nordlander, Uppsala, Sweden, who Male. Unknown. has significantly contributed to our current HOLOTYPE: Female, CAS, Late Cretaceous understanding of cynipoid relationships and (Campanian), Medicine Hat, Alberta, Canada; mentored several of the most active cynipoid deposited in the CNC. researchers worldwide. ETYMOLOGY: The specific epithet is taken from Latin, petiolata, meaning ‘‘stalked’’, and COMMENTS: The new genus belongs to the clade consisting of the subfamilies Oberthuerel- is a reference to the long petiolar segment of the species. Indeed, this is the oldest species linae and Liopterinae. Although it is difficult to with an extended petiole in the Cynipoidea. collect sufficient characters to incorporate the species into the analysis of cynipoid phylogeny, several observable features strongly indicate its FAMILY FIGITIDAE THOMSON placement within this clade, including the DIAGNOSIS: Figitids are morphologically di- following: (1) lateral margin of axilla conspic- verse, making unifying traits a challenge to uously raised anteriorly; (2) petiolar annulus identify. Ronquist (1995b, 1999) suggested long, median dorsal length at least twice as long that the Figitidae are supported by two as minimum dorsal width (estimated from apomorphies: presence of a distinct point of lateral view); and (3) posterior margin of petiole weakness in the eighth metasomal tergum of (5 metasomal T1) and anterior margin of the female at the position of the gonoplac (5 metasomal T2 abutting, dorsal margins of the third valvula), and forewing Rs+M situated two more or less continuous (i.e., without close to the end of the first free abscissa of M constriction between segments). (5 basal vein). Further potential apomorphies Goerania can be easily separated from genera include a vertical posterior margin of metaso- in the Oberthuerellinae and Liopterinae by the mal T2 and the presence of a longitudinal absence of foveate sculpture on the lateral carina on the posterior surface of the metati- surface of the pronotum and the postpe- bia. Although each of the above characters tiolar terga of the metasoma being subequal except for the first has similar states in other in length. lineages of Cynipoidea, the unique combina- tion of these traits supports the separation of Goerania petiolata Liu and Engel, new species Figitidae from other cynipoids. figures 7, 8 COMMENTS: Species of Figitidae are rela- tively small and are primary parasitoids, DIAGNOSIS: As for the genus (see above). principally on Diptera, but hosts also include 14 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 7. Photomicrograph of holotype female of Goerania petiolata Liu and Engel, new genus and species (CAS). chrysopid and hemerobiid lacewings (Neu- more or less complete median carina. Anterior roptera) or Cynipidae. In addition, some pronotal flange short; anterior submedian species are hyperparasitoids of braconid and depression open laterally; dorsal pronotal area chalcidoid primary parasitoids of aphids and narrow but distinct; lateral pronotal carina psyllids. Ronquist (1995b, 1999) considerably present; lateral pronotal surface without fove- altered the concept of Figitidae. Ronquist was ate sculpture. Mesoscutum glabrous; median able to demonstrate from his analyses that longitudinal impression and lateral notauli figitids were paraphyletic, and he incorporated distinct; mesoscutellum posteriorly extended into Figitidae some groups that historically into a prominent horizontal process; lower had been regarded as distinct families. The mesopleuron ventrally strongly expanded into current classification of Figitidae is summa- a broad lobular structure, posteriorly covering rized in table 1. basal part of mesocoxa. Forewing with bulla present in Sc+R1;Rs+M issuing from close to SUBFAMILY INCERTAE SEDIS posterior end of first free abscissa of M (5 basal vein). Mesocoxa inserted almost verti- Micropresbyteria Liu and Engel, new genus cally; metacoxa distinctly dorsomedially swol- len and dorsolaterally distinctly depressed. TYPE SPECIES: Micropresbyteria caputipres- Petiolar annulus low and without longitudinal sa Liu and Engel, new species. carina; petiole slightly shorter than wide, with DIAGNOSIS: F1 of male antenna medially distinct longitudinal carinae. not flattened and not twisted. Head distinctly ETYMOLOGY: The specific epithet is derived compressed longitudinally and not impressed from the Greek words mikros (meaning posteriorly; lateral occipital carina laterally ‘‘little’’ or ‘‘small’’) and presbytrion (meaning strongly expanded into a broad lobular ‘‘an assemblage of elders’’). The name is structure; lower face glabrate, with a simple, feminine. 2007 I TA. VLTO FCYNIPOIDEA OF EVOLUTION AL.: ET LIU

Fig. 8. Holotype female of Goerania petiolata Liu and Engel, new genus and species (CAS). 15 16 AMERICAN MUSEUM NOVITATES NO. 3583

COMMENTS: The extensive lobular expan- pleuron ventrally expanded; mesosoma across sion of the upper part of the lateral occipital mesopleuron almost as high as long, excluding carina and massively expanded ventrolateral mesoscutellar process. Mesocoxa inserted al- carina of the mesopectus separate Micropres- most vertically, not obliquely at posterior byteria from all other cynipoids. The longitu- area. Propodeum devoid of processes; nucha dinally distinctly compressed head, the lack of short and broad in lateral view. Wings entirely distinct sculpture on the head and mesosoma, hyaline, with sparse pubescence; all wing the short anterior pronotal flange, and the margins ciliate. Forewing with marginal cell downward insertion of the meso- and meta- closed, about 2.6 times as long as wide; bulla coxae strongly indicate that the fossil belongs in Sc+R1 present, 2r-rs oblique, sloping out- to the microcynipoids. The presence of an ward posteriorly; areolet large; Rs+M nebu- obvious dorsal pronotal area, the position of lous, arising from posterior end of first free the proximal end of Rs+M close to the abscissa of M (5 basal vein). Anterior margin posterior end of the first free abscissa of M of hind wing with three hamuli. Metabasitar- (5 basal vein), and the presence of the sus shorter than combined lengths of second longitudinal carina dorsolaterally on the to fifth metatarsomeres; metacoxa dorsally metatibia further suggest that the genus distinctly depressed; metatibia dorsolaterally belongs to the family Figitidae. Because of with a distinct longitudinal carina; all pre- a lack of information for female characters, tarsal claws simple, without basal lobe. which are more crucial in cynipoid phylogeny, Petiolar annulus relatively long, attached low we only provisionally place the genus in to propodeum; segmentation of metasoma Figitidae. indistinct owing to imperfect preservation. Female. Unknown. Micropresbyteria caputipressa Liu and Engel, HOLOTYPE: Male, RTMP 96.9.170, Late new species Cretaceous (Campanian), Grassy Lake (110u figures 9, 10 409W, 49u459N), Alberta, Canada; deposited in RTMP. DIAGNOSIS: As for the genus (see above). ETYMOLOGY: The specific epithet is derived DESCRIPTION: Male. Body length 1.00 mm; from Latin caputa (meaning ‘‘head’’) and forewing length 0.90 mm. Body mostly black pressus (meaning ‘‘pressed’’). The name refers to dark brown; wings hyaline, without any to the longitudinally compressed head. macula or band. Antenna cylindrical, 14- segmented; pedicel almost spherical, one-half SUBFAMILY EUCOILINAE THOMSON as long as scape; F1 medially not flattened and not twisted as in male of most known species DIAGNOSIS: The subfamily is easily distin- of cynipoids, slightly longer than F2 (F2 guished from all other cynipoids by the seven-sixths length of F1) and much thicker presence of a mesoscutellar cup or plate with than the latter; elongate placodeal sensilla a deep depression centrally or posteriorly. In present on all flagellomeres. Head compressed addition, eucoiline females, except for two of longitudinally, attached high to mesosoma; the new fossil genera described below, have frons, vertex, gena, and malar space glabrous; the second through fourth metasomal terga (5 compound eye prominent, distinctly produced abdominal T3–5) fused, a feature shared only in front of gena, and vertically much extended; with the Pycnostigminae (Figitidae) among malar space reduced to narrow strip beneath cynipoids. compound eye; gena not expanded behind COMMENTS: The presence of the mesoscu- compound eye. Lateral surface of pronotum tellar plate undoubtedly supports the mono- glabrous. Mesoscutum curved dorsally in phyly of the subfamily, a feature exhibited by lateral view and glabrous, with sparse pubes- all three of the taxa treated herein. The new cence; without transverse costa; median im- genera Anteucoila and Jerseucoila (see below), pression and lateral notauli distinct; mesoscu- however, lack the fusion of metasomal T2–4 tellum flat dorsally and posteriorly projected and are therefore obviously basal to all other into a blunt process (an artifact of preserva- eucoilines, representing a stem group to the tion?); mesopleuron glabrous; lower meso- subfamily as it is understood based on modern 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 17

Fig. 9. Photomicrograph of holotype male of Micropresbyteria caputipressa Liu and Engel, new genus and species (RTMP-96-9-170).

taxa. However, as already noted, Anteucoila Anteucoila Liu and Engel, new genus and Jerseucoila possess a well-developed mesoscutellar cup, indicative of their place- TYPE SPECIES: Anteucoila delicia Liu and ment as eucoilines. The third genus, Engel, new species. Syneucoila, is a typical crown-group eucoiline DIAGNOSIS: The new genus is easily distin- and is, in fact, apparently related to the clade guished from all other members of the sub- consisting of the Zaeucoila generic group and family (except Jerseucoila, see below) by the core + higher eucoilines (Fontal-Cazalla et absence of the characteristic fusion of meta- al., 2002; see below). somal T2–4 and by the postpetiolar terga of The Eucoilinae is the most species-rich about equal length along the middorsal figitid subfamily, currently with about 80 line. The only other genus exhibiting these genera and 1000 species. Where known, plesiomorphies is Jerseucoila, from which eucoilines are restricted to hosts of the Anteucoila can be differentiated by the strong- cyclorrhaphan Diptera (Ronquist, 1999, and ly sculptured mesoscutellar plate (weakly references therein). The other Cretaceous carinate and weakly granulose in Jerseu- genus, Syneucoila (see below), is more typical coila), straight path of Rs on the apical border of modern Eucoilinae (yet still rather plesio- of the marginal cell (distinctly arched apically morphic for its clade), attesting to the in Jerseucoila), the shorter metabasitarsus antiquity of the lineage. (elongate in Jerseucoila), the shorter second 18 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 10. Holotype male of Micropresbyteria caputipressa Liu and Engel, new genus and species (RTMP- 96-9-170). metasomal tergum (more elongate in macula or band. Antenna cylindrical, 12-seg- Jerseucoila), and the narrower petiole mented, with distal three flagellomeres conspic- (fig. 11) (broader in Jerseucoila). uously expanded; pedicel almost spherical, two- ETYMOLOGY: The new genus-group name thirds as long as scape; F1 distinctly longer than is a combination of ante- (Latin, meaning F2; elongate placodeal sensilla distinct on at ‘‘before’’) and Eucoila, type genus of the least distal flagellomeres. Frons, vertex, gena, subfamily. The name is feminine. and malar space glabrous; lower face somewhat elevated in upper part; gena not expanded Anteucoila delicia Liu and Engel, new species behind compound eye; compound eye longer, figures 11, 12 slightly more than twice length of malar space. Lateral surface of pronotum glabrous; lateral DIAGNOSIS: As for the genus (see above). pronotal carina present. Mesoscutum slightly DESCRIPTION: Female. Body length 0.63 curved dorsally in lateral view; mesoscutellum mm; forewing length 0.6 mm. Body mostly posteriorly sloped and dorsally with a central black to dark brown; wings hyaline, without plate; mesopleuron glabrous; mesopectus not 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 19

Fig. 11. Photomicrograph of holotype female of Anteucoila delicia Liu and Engel, new genus and species (RTMP-96-9-785).

protruding ventrally, mesocoxa directed verti- vein). Metabasitarsus about half as long as cally and not inserted on separate, oblique combined lengths of second to fifth metatar- posterior area. Propodeum devoid of processes; someres; pretarsal claws simple, without basal lateral propodeal carina distinct. Wings hyaline, lobe. Postpetiolar metasomal terga not fused, all with sparse pubescence; all wing margins, subequal in length along middorsal line; hypo- especially outer margin, ciliate. Forewing with pygium of female long and slender. Male. marginal cell closed, about three times as long as Unknown. wide; bulla in Sc+R1 absent; 2r-rs oblique, HOLOTYPE: Female, RTMP 96.9.785, Late sloping outward posteriorly; areolet absent; Cretaceous (Campanian), Grassy Lake (110u Rs+M reduced, hardly traceable, arising from 409W, 49u459N), Alberta, Canada; deposited posterior end of first free abscissa of M (5 basal in RTMP. 20 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 12. Holotype female of Anteucoila delicia Liu and Engel, new genus and species (RTMP-96-9-785); note that the right antenna (lower one in image) is partially pulled from the head capsule and so the basalmost sclerites depicted are not portions of the antenna but are instead sclerotic debris. 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 21

Fig. 13. Photomicrograph of holotype female of Jerseucoila plesiosoma Liu and Engel, new genus and species (NJ-1006).

ETYMOLOGY: The specific epithet is derived amber deposit in which it was discovered) and from the Latin word delicia (meaning ‘‘favor- Eucoila, type genus of the subfamily. The ite’’ or ‘‘lovely’’). name is feminine.

Jerseucoila Liu and Engel, new genus Jerseucoila plesiosoma Liu and Engel, new species TYPE SPECIES: Jerseucoila plesiosoma Liu figures 13, 14 and Engel, new species. DIAGNOSIS: Like Anteucoila (see above) Cynipoid sp.: Grimaldi and Engel, 2005: 422 [figured]. with freely articulated metasomal T2–4, this new genus can be readily excluded from DIAGNOSIS: As for the genus (see above). crown-group Eucoilinae. The presence of DESCRIPTION: Female. Body length 0.60 mm; a mesoscutellar plate, as in the former genus, forewing length 0.51 mm. Body mostly black; supports its position as a stem-group eucoi- wings hyaline, without macula or band. line. Jerseucoila differs from Anteucoila by the Antenna cylindrical, with distalmost flagello- apically less expanded antenna, the weakly meres more elongate than preceding flagello- carinate and sculptured mesoscutellar plate, meres; pedicel almost spherical, two-thirds as the arching of the distal abscissa of Rs, the long as scape; F1 about as long as F2; elongate metabasitarsus, the larger second elongate placodeal sensilla distinct on nearly metasomal tergum, and the broader petiole all flagellomeres except basal two. Frons, (fig. 13) (refer also to the Diagnosis for vertex, gena, and malar space apparently Anteucoila). glabrous (difficult to integument in holotype ETYMOLOGY: The new genus-group name owing to layer of microscopic bubbles); gena is a combination of Jersey (a reference to the not expanded behind compound eye; com- 22 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 14. Holotype female of Jerseucoila plesiosoma Liu and Engel, new genus and species (NJ-1006). pound eye much longer than malar space, Propodeum devoid of processes. Wings hya- nearly five times longer than malar length. line, with scattered pubescence; all wing Lateral surface of pronotum apparently margins, especially posterior margins, ciliate. glabrous. Mesoscutum weakly curved dorsal- Forewing with marginal cell closed, about ly in lateral view; mesoscutellum slightly three times as long as wide; bulla in Sc+R1 sloped posteriorly and with weakly defined absent; 2r-rs oblique, sloping outward central plate; mesopleuron apparently gla- posteriorly; areolet absent; Rs+M reduced, brous; mesocoxa directed vertically and not scarcely traceable, apparently arising from inserted on separate, oblique posterior area. posterior end of first free abscissa of M (5 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 23 basal vein). Metabasitarsus nearly as long as malar space with distinct sulcus; lower face combined lengths of second to fifth metatar- somewhat elevated in upper part; gena not someres; pretarsal claws simple, without expanded behind compound eye; compound eye basal lobe. Postpetiolar metasomal terga not nearly three times longer than malar space. fused, all subequal in length along middorsal Lateral surface of pronotum glabrous. line except metasomal T2 more elongate; Mesoscutum curved dorsally in lateral view; hypopygium of female long and slender. mesoscutellum posteriorly sloped and dorsal- Male. Unknown. ly with a central plate; mesopleuron glabrous, HOLOTYPE: Female, AMNH NJ-1006, subalar pit well developed; mesopectus not Late Cretaceous (Turonian), White Oak Pit, protruding ventrally, mesocoxa directed ver- Sayreville, Middlesex County, New Jersey; tically and not inserted on separate, oblique deposited in the Amber Fossil Collection, posterior area. Propodeum devoid of pro- Division of Invertebrate Zoology, AMNH. cesses; lateral propodeal carina distinct. ETYMOLOGY: The specific epithet is a com- Wings hyaline, with sparse pubescence; all bination of the Greek words plesios (meaning wing margins ciliate, ciliae particularly elon- ‘‘near’’) and soma (meaning ‘‘body’’). gate along apical posterior margin of fore- wing. Forewing with marginal cell, triangular closed, about 2.5 times as long as wide; bulla Syneucoila Liu and Engel, new genus in Sc+R1 present; 2r-rs short, projecting TYPE SPECIES: Syneucoila magnifica Liu straight posteriorly; areolet absent; Rs+M and Engel, new species. reduced, hardly traceable, apparently arising DIAGNOSIS: Typical eucoiline with postpet- from posterior end of first free abscissa of iolar metasomal terga fused but exhibiting M(5 basal vein). Metabasitarsus slightly a relatively shortened head (not elongate), more than one-half as long as combined possessing an anterior metepimeral impres- lengths of second to fifth metatarsomeres; sion, and with a well-developed subalar pit pretarsal claws simple, without basal lobe. and subalar area. Additional features defining Postpetiolar metasomal T2–4 fused. Male. the genus include: broad metasubpleural de- Unknown. pression anterior to metacoxal foramen pres- TYPE MATERIAL: Female, AMNH NJ- ent; metacoxa without setal patches; forewing 1075, Late Cretaceous (Turonian), White Oak with marginal cell closed; R1 complete and Pit, Sayreville, Middlesex County, New Jersey; elongate, surpassing marginal cell apex; mar- deposited in the Amber Fossil Collection, ginal ciliae of wing distinctly elongate along Division of Invertebrate Zoology, AMNH. posterior margin of forewing. ETYMOLOGY: The specific epithet is taken ETYMOLOGY: The new genus-group name from the Latin word magnificus (meaning is a combination of syn- (Greek, meaning ‘‘splendid’’) and is a reference to the esthetic ‘‘together’’; a reference to the fused metasomal beauty of the holotype. terga relative to other Cretaceous eucoilines) and Eucoila, type genus of the subfamily. The FAMILY CYNIPIDAE LATREILLE name is feminine. COMMENTS: Although the family Cyni- Syneucoila magnifica Liu and Engel, pidae is universally accepted as monophyletic, new species satisfactory character evidence readily defin- ing the family is lacking except for its figures 15, 16 phytophagous habit. Numerous characters DIAGNOSIS: As for the genus (see above). have been put forward as putative synapo- DESCRIPTION: Female. Body length 0.73 mm; morphies for the family, including the lack of forewing length 0.65 mm. Body mostly dark the lateral pronotal carina and an open brown; wings hyaline, without macula or band. marginal cell in forewing (Liljeblad and Antenna cylindrical; pedicel almost spherical; F1 Ronquist, 1998), as well as a medially nar- distinctly longer than F2; elongate placodeal rowed dorsellum (Ronquist, 1999). However, sensilla distinct on at least distal flagellomeres. all of these putative synapomorphies have Frons, vertex, gena, and malar space glabrous; exceptions both within Cynipidae (as rever- 24 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 15. Photomicrograph of holotype female of Syneucoila magnifica Liu and Engel, new genus and species (NJ-1075).

sals) and in Figitidae (as parallelisms) (e.g., behind compound eye; vertex, gena, malar Ronquist, 1999). Nonetheless, the family can space, and face glabrous; lower face with still be reasonably defined based on a combi- a simple, more or less complete median carina. nation of morphological attributes. In addi- Pronotum dorsomedially strongly extended tion to the aforementioned characters, the anteriorly; anterior pronotal flange short; following traits can be useful in the diagnosis lateral pronotal carina present; lateral prono- of cynipids: (1) dorsal pronotal area absent; tal surface transversely costate on upper half, (2) metasoma of female strongly compressed; and without foveate sculpture. Mesoscutum (3) Rs+M arising from middle of the first free glabrous; lateral notauli distinct; mesoscutel- abscissa of M (5 basal vein); (4) 2r-rs about lum posteriorly truncate, without process; 0.5–0.65 times as long as Sc+R1; (5) R1 lateral mesopleural impression anteriorly present, of 2r-rs directed distinctly obliquely laterally convergent to a longitudinal impression along rather than more or less perpendicular to ventral margin of lower mesopleuron toward anterior wing margin; and (6) R+Rs and posterior two-thirds and divided into several Sc+R1 smoothly continuous, not forming a dis- foveae by vertical carinae; lateroventral carina tinct angle (see Liljeblad and Ronquist, 1998). of mesopectus present and complete; lower mesopleuron ventrally not expanded. Fore- SUBFAMILY INCERTAE SEDIS wing with bulla in Sc+R1 present; Rs+M issuing from middle of the first free abscissa Tanaoknemus Liu and Engel, new genus of M (5 basal vein). Mesocoxa inserted TYPE SPECIES: Tanaoknemus ecarinatus Liu vertically and not obliquely at separate poste- and Engel, new species. rior area; metacoxa dorsally distinctly de- DIAGNOSIS: Male antenna 14-segmented, pressed and with a rounded, anterolateral with F1 distinctly excavated laterally. Com- triangular crest; metatibia longer than meta- pound eye prominent; gena not expanded femur by one-third. Petiolar annulus low, 2007 I TA. VLTO FCYNIPOIDEA OF EVOLUTION AL.: ET LIU

Fig. 16. Holotype female of Syneucoila magnifica Liu and Engel, new genus and species (NJ-1075). 25 26 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 17. Photomicrograph of holotype male of Tanaoknemus ecarinatus Liu and Engel, new genus and species (CAS-78).

postsubpleuron short; petiole slightly longer Tanaoknemus ecarinatus Liu and Engel, than wide and longitudinally carinate. new species COMMENTS: The presence of a unique me- figures 17, 18 sopleural impression and unusually long tibia separate the genus from all other cynipoids. DIAGNOSIS: As for the genus (see above). The lack of the lateral pronotal carina, DESCRIPTION: Male. Body length 1.20 mm; length of forewing about 1.00 mm (tip of wing presence of a bulla in Sc+R1, and Rs+M issuing from the middle of the first free not well preserved). Body mostly black; wings abscissa of M (i.e., basal vein) suggest affinity hyaline, without any macula or band. Antenna with the family Cynipidae. Although the cylindrical, 14-segmented (10-segmented on relatively long petiole is a feature often found the other side, but that is apparently an in Figitidae and is not known in extant abnormality); F1 laterally excavated; elongate Cynipidae, the trait has evolved independently placodeal sensilla present on all flagellomeres. in many Hymenopteran lineages, including Head compressed longitudinally, attached several times within Cynipoidea. We therefore high to mesosoma; frons, vertex, gena, and interpret its presence in Tanaoknemus as an malar space glabrous; compound eye promi- autapomorphy, and the genus is provisionally nent, vertically more than twice as high as placed as a basal clade of Cynipidae. malar space; gena not expanded behind ETYMOLOGY: The new genus-group name compound eye. Lateral surface of pronotum is a combination of the Greek words tanao- transversely weakly costate in upper half; (meaning ‘‘long’’ or ‘‘outstretched’’) and mesoscutum dorsally slightly curved in lateral knem- (meaning ‘‘the leg between the knee view, glabrous with sparse pubescence, with- and ankle’’). The name is a reference to out transverse costa; median impression and the unusually long tibia of the new genus lateral notauli percurrent; mesoscutellum dor- relative to other cynipoids. The name is sally flat and irregularly sculptured, without masculine. posterior processes; mesoscutellar sulcus me- 2007 -.,_\\ . - I TA. VLTO FCYNIPOIDEA OF EVOLUTION AL.: ET LIU \· ·~ \ ~ ,_ I ! ~- 27 Fig. 18. Holotype male of Tanaoknemus ecarinatus Liu and Engel, new genus and species (CAS-78). 28 AMERICAN MUSEUM NOVITATES NO. 3583 dially divided into two large foveae by radiating strigae reaching lower margin of a longitudinal carina; mesopleuron glabrous. compound eye and medially glabrous. Mesocoxa inserted vertically beneath meso- Pronotum long medially, ratio of median to pleuron. Propodeum with no processes; nucha posterior distance between dorsal and ventral short; lateral propodeal carina percurrent and margins about 0.56; lateral pronotal carina dorsally not curved in lateral view. Wings absent; pronotal surface longitudinally stri- entirely hyaline and pubescent; wing margins gate; mesoscutum strongly convex and gla- ciliate. Forewing with marginal cell closed, brous, with percurrent notauli; mesopleuron about three times as long as wide; bulla in including mesopleural triangle glabrous; me- Sc+R1 present; 2r-rs oblique, sloping outward soscutellum convex, dorsally with broadly posteriorly; areolet large; Rs+M nebulous, spaced costae, posteriorly sloped gradually; arising from the middle of the first free mesoscutellar sulcus medially subdivided into abscissa of M (5 basal vein). First metatar- two foveae by a longitudinal carina; mesopec- somere slightly shorter than combined length tus ventrally not projected; mesocoxa directed of second through fifth metatarsomeres; me- vertically downward; metapleural sulcus ante- tacoxa dorsally distinctly depressed, antero- riorly ends high, distance between upper laterally with a rounded triangular crest; metapectal margin and its anterior end about metatibia dorsolaterally with a distinct longi- half the distance between its anterior end and tudinal carina; all pretarsal claws simple, anteroventral margin of episternum; metacox- without basal lobe. Petiolar annulus slightly al foramen close to anterior margin of longer than wide and longitudinally carinate, metepisternum. Forewing with marginal cell attached low to propodeum; all metasomal closed anteriorly; R1 laterad of 2r-rs directed terga subequal along middorsal line in lateral strongly obliquely laterally; Rs+M arising from view, each with posterior margin oblique and just posterior of middle of first free abscissa of almost straight. Female. Unknown. M; areolet present. (Note that we were unable HOLOTYPE: Male, CNC CAS-78, Late to observe the venation clearly in the holotype. Cretaceous (Campanian), Medicine Hat, This may be due to deterioration of the Alberta, Canada; deposited in CNC. specimen in a relatively exposed area through ETYMOLOGY: The specific epithet is derived time. Thus, the above description is largely from Latin and is a combination of e- based on Kinsey’s original illustration, assum- (meaning ‘‘without’’) and carinatus (meaning ing that his observations and interpretations of ‘‘keeled’’). The name describes the lack of the wing were correct.) Pretarsal claws of all a lateral pronotal carina in the new species. legs simple, without tooth. Nucha low and short, about one-eighth length of metacoxa, SUBFAMILY CYNIPINAE LATREILLE dorsally glabrous. Petiole short and inconspic- TRIBE AYALCINI ASHMEAD uous, dorsal part crescent-shaped, and gla- brous; metasoma laterally strongly com- Kinseycynips Liu and Engel, new genus pressed, as long as head and mesosoma TYPE SPECIES: Aulacidea succinea Kinsey, combined; metasomal T2–3 not fused, about 1919. half as long as postpetiolar metasoma; ventral DIAGNOSIS: Antenna of female long and spine of metasomal S6 short, slightly separated slender, reaching beyond posterior end of from and extending beyond apex of lateral flap. mesosoma when in repose and 14-segmented; ETYMOLOGY: The new genus is named F1 slightly curved and slightly shorter than F2 after Dr. Alfred C. Kinsey (1894–1956) for (50:55); placodeal sensilla present on all his significant contributions to the study of flagellomeres. Compound eyes prominent gall wasps. His collection of gall wasps is and long, about twice as long as malar space; estimated to amount to five million specimens gena and vertex glabrous; clypeus ventrally and two million galls, and is now deposited in projecting over mandibles, anterior margin the AMNH. In addition to the description of trapezoidal; epistomal sulcus absent; facial about 400 new species of gall wasps, he strigae radiating from clypeus and reaching pioneered the search for phylogeny. compound eye; lower face laterally with Dr. Kinsey studied gall wasps for more than 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 29

20 years, but later turned his interest to, and rose family. However, in the absence of became more widely known for, studies of a cladistic analysis we cannot exclude the human sexual behavior. possibility that Kinseycynips was a galler of COMMENTS: Aulacidea succinea is consid- herbaceous species of the rose family. ered here to belong to the gall wasp family Cynipidae owing to the absence of the lateral Kinseycynips succinea (Kinsey), pronotal carina, a synapomorphy for the new combination cynipids, and to the complete absence of figure 19 synapomorphies defining the sister family Figitidae (e.g., Rs+M arising from distinctly Aulacidea succinea Kinsey, 1919: 48. after middle of the first free abscissa of M, metasomal T2 with distinctly oblique posteri- DIAGNOSIS: As for the genus (see above). or margin). Ronquist (1999) suggested that A. HOLOTYPE: Female, Eocene (Lutetian), succinea belonged to the complex of Baltic amber; deposited in the Amber Fossil Synergus (Synergini). However, the following Collection, Division of Invertebrate Zoology, characters clearly separate the species from the AMNH. Kinsey (1919) stated, ‘‘Type: a single latter: lower face not covered with radiating specimen from collection of the Ko¨nigsberg strigae, but with a medial glabrous area; dorsal Museum, and temporarily at the Bussey part of petiole small, glabrous, and crescent- Institution, of Harvard University.’’ Once shaped; and metasomal T2–3 not fused to form thought to be lost, the specimen was recently a large tergite covering more than half of the rediscovered in the Kinsey Collection at the postpetiolar metasoma. Furthermore, the gla- AMNH. brous and asetose mesoscutum of A. succinea differs from all inquilines except Synophro- CYNIPOIDEA INCERTAE SEDIS morpha. Indeed, the fossil is further separated from all inquilines by several features, in- Two additional specimens of cynipoid wasps cluding toothless pretarsal claws; a clypeus were recognized, but owing to poor preserva- projecting over the mandibles, with its anterior tion and obscured views through the amber margin trapezoidal; and absence of the episto- they were not identifiable beyond the level of mal sulcus. Indeed, the fossil can be easily superfamily. There was one specimen each in identified as Aylacini using the key to tribes of Canadian and New Jersey . We provide Neotropical Cynipidae provided by Buffington here their accession numbers so that future et al. (2005). workers, with new questions and techniques for Within Aylacini, the new genus more closely old material, might know of their existence and resembles those genera that produce galls perhaps seek them for study: NJ-1069 in the on Rubus spp. and Potentilla spp. (i.e., AMNH and CAS-282 in the CNC. Xestophanes and Diastrophus) in that all share glabrous integument on the vertex, mesoscu- CLADISTICS tum, and mesopleuron. However, Kinseycy- METHODS nips can be easily distinguished from these by its toothless claws, closed marginal cell, and 14- The character matrix from a recent cladistic segmented antenna in the female. Kinseycynips analysis of the higher-level phylogeny of is also similar to Aulacidea, the genus where Cynipoidea (Ronquist, 1995b) was supple- Kinsey (1919) originally placed his species, but mented with data on six of the more plesio- Aulacidea females have only 13-segmented morphic, fossil taxa described in this mono- antennae, the vertex and mesoscutum coria- graph; i.e., Protimaspis, Stolamissus, Micro- ceous or otherwise sculptured (but never presbyteria, Tanaoknemus, Proliopteron, and glabrous), and the mesopleuron longitudinally Anteucoila (for the purposes of the higher-level striate. Because the new genus shares more analysis, Anteucoila and Jerseucoila are iden- similarities with those aylacine genera galling tical and so only one has been coded for on rosaceous, mostly woody hosts, we think study). The six genera were included because that it was associated with a woody host of the each is well preserved and allows detailed 30 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 19. Photomicrograph of holotype female of Kinseycynips succinea (Kinsey). morphological study and thereby meaningful 112. Shape of metacoxa: 0, dorsally comparison to modern taxa. Furthermore, depressed or not, but never with ante- each of these genera possesses a unique rolateral crest; 1, strongly depressed combination of plesiomorphic features mak- dorsally to form a longitudinal furrow ing their assignment to higher groups in the with a more or less triangular, anterolat- absence of a cladistic analysis more sub- eral crest. jective (in contrast to taxa like, Syneucoila and Kinseycynips, which are distinctly more Although the six fossil genera that are modern and exhibit a number of synapomor- included in the matrix are relatively well phies for their respective families and sub- preserved and we were able to code them for families). Ronquist (1995b) coded 110 ex- most of the 112 characters (table 3), two of the ternal morphological characters for his cla- genera (i.e., Tanaoknemus and Micropres- distic analysis of higher-level relationships in byteria) are only known from males. Because the Cynipoidea. For descriptions of charac- cladistic reconstruction of cynipoid phylogeny ters and the complete character matrix for depends disproportionately on attributes of modern taxa refer to Ronquist (1995b). Two female morphology, the two genera were new characters are supplemented in our initially excluded to avoid negative perturba- present re-analysis (codings for the extant tion to the analysis by an excessive amount of taxa for these two characters are provided in missing data (Schuh, 2000). Nonetheless, we table 2): included the two genera in further analyses to explore, albeit in a limited fashion, whether 111. Shape of remnant of pterostigma: the absence of data indeed had an overall 0, short and thick; 1, long and thin. effect. 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 31

TABLE 2 1997) with the options ‘‘hold 10,000; mult Partial Data Matrix for Cynipoidea 5,000’’, and character optimization and tree (New characters 111 and 112 for Recent and visualization were performed using WinClada fossil genera) (Nixon, 2002). Statistics from individual analyses are provided in the corresponding 11 figure captions. 11 12 RESULTS Outgroup 00 Austrocynips 00 Parsimony analysis of the data, with Eileenella 00 Tanaoknemus and Micropresbyteria excluded, Ibalia 01 and with no constraints resulted in 11 equally Heteribalia 00 Kiefferiella 00 parsimonious topologies (fig. 20). According Paramblynotus 00 to the analysis, Protimaspis is one of the Mesocynips 00 basalmost clades among cynipoids except Dallatorrella 00 Austrocynips, but its position relative to Xenocynips 00 Ibaliidae and the stem taxa of microcynipoids Tessmannella 00 and liopterids is unresolved. The genera Oberthuerella 00 Stolamissus, Proliopteron, and Anteucoila Liopteron 00 form an unresolved cluster with the stem Peras 00 Pseudibalia 00 species of the microcynipoids and the liopter- Isocolus 10 ids (fig. 20). When the same matrix was Diastrophus 10 analyzed with the position of Anteucoila with Euceroptres 10 Figitidae constrained (once again, reflecting Melanips 10 the placement of Eucoilinae within this Stolamissus 00 family), a single topology was recovered that Protimaspis 01 was only one step longer than the first Proliopteron 10 analysis, but also with slightly better tree Anteucoila 00 Presbyfigites 00 statistics (fig. 21) (since two nodes were Micropresbyteria 01 actually constrained, their support is low). Interestingly, the overall topology from this analysis matches Ronquist’s original tree, with Among the fossil genera considered herein, the obvious difference of the addition of the Anteucoila, Jerseucoila, and Syneucoila pos- fossil taxa. The single tree from the con- sess a mesoscutellar cup or plate, a feature strained analysis is therefore preferred as the unique to the subfamily Eucoilinae (Figitidae). most plausible phylogeny for the Cynipoidea Therefore, the presence of this feature in over the suite of trees resulting from the these taxa solidly places them in Eucoilinae. unconstrained analysis. Given that eucoilines According to Ronquist (1995b, 1999), the were not originally included, we think this to monophyly of the Figitidae including Eucoi- be a safe assertion. According to this analysis linae is well established. The character matrix (fig. 21), Austrocynipidae, Ibaliidae, Proti- he constructed (Ronquist, 1995b) is intended maspis, and Stolamissus form a basal grade, for analysis of cynipoid phylogeny at the each being progressively more related to familial level and did not include any eucoi- Liopteridae + microcynipoids. Proliopteron is lines. Thus, the data matrix updated herein supported as sister to the remainder of the was analyzed with the position of Anteucoila Liopteridae (fig. 21). within Figitidae constrained as well as un- Both unconstrained (fig. 22) and con- constrained. Anteucoila was selected to repre- strained, with respect to Anteucoila (fig. 23), sent these fossils as it is the more plesio- analyses of the data matrix with the addition morphic and complete of the three. of Tanaoknemus and Micropresbyteria re- Cladistic analyses were conducted using parsi- sulted in two equally parsimonious topologies mony as implemented in NONA (Goloboff, each. The two trees resulting from the un- 2AEIA UEMNVTTSN.3583 NO. NOVITATES MUSEUM AMERICAN 32

TABLE 3 Partial Data Matrix for Cynipoidea (Codings for new fossil genera only; refer to Ronquist, 1995b, for complete explanation of characters 1–110 and codings for extant groups; codings for the new characters, 111 and 112, are presented in table 2 for all Recent and fossil genera.)

11111111112222222222333333333344444444445555555555666666666677777777778 12345678901234567890123456789012345678901234567890123456789012345678901234567890

Stolamissus 0?00??1?0??20???0001-000110??101?1???111000010101?100?1110?000010000000-0?000??? Protimaspis ??00??0?0??20???00000010200??110?0010111000110101?1??01110?000000000?00-0?000??? Proliopteron 1110??0?0??1101000?011??010??101??120?21000110100?10000?10?0000?0?00?00-0?001??? Anteucoila 0-10??0?0??21???0001-00201011101?1120111000001100?0000?110?000000000000-00000??? Micropresbyteria 1110???0?1?210??0001-002010?1101?11?01100000111?0?100000?????0??00000?0-0?001??? Tanaoknemus ??10???0?0?12???00?1-0120?0?1101?1120111000?10100?1000?0?????0??00000?0-0?011???

11111111111 888888888999999999900000000001 123456789012345678901234567890

Stolamissus 100??0000000??00?00?000?0?0000 Protimaspis 000??0000000??00000?000?0?0000 Proliopteron ?00?0100?0000-000000000?0?0000 Anteucoila 000?00000000??000000000?0?0000 Micropresbyteria ?00?000000000-0??00???0??0?0?? Tanaoknemus 100??0000000??0??00???00?0?0?? 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 33

Fig. 20. Strict consensus of 11 shortest trees (L 5 227, CI 5 56, RI 5 79) resulting from parsimony analysis of Ronquist’s (1995b) data matrix, supplemented with paleontological data for several Late Cretaceous amber fossils described herein. We have added to the original matrix two additional characters (refer to section on Cladistics) and four fossil taxa (i.e., Protimaspis, Stolamissus, Anteucoila, and Proliopteron). No constraints were applied to the search which consisted of: hold 10,000; mult 5,000; wh*; max* in NONA (Goloboff, 1997). Values above branches are the number of unambiguous character changes supporting particular nodes, while those below are Jackknife and Bootstrap values. constrained analysis were 235 steps (fig. 22), However, the clade consisting of (Tanao- whereas the two resulting from the con- knemus (Micropresbyteria (Proliopterion + strained analysis were 239 steps (fig. 23). The extant liopterids))) is ‘‘supported’’ by merely cladograms of the two analyses are essentially a single female character (i.e., character 56: identical except for the position of Anteucoila, petiole is at least partially longitudinally and they are the same as the constrained carinate) (fig. 24). This is obviously a tenuous analysis without the inclusion of Tanaoknemus position given that the Tanaoknemus and and Micropresbyteria. Tanaoknemus and Micropresbyteria are known only from males! Micropresbyteria are basal, from the base We therefore think that this is not an entirely upward, to Proliopteron and extant liopterids. reliable homology statement and consider the 34 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 21. Preferred set of cladistic relationships among Cynipoidea; strict consensus of the two shortest tree (L 5 228, CI 5 57, RI 5 80) resulting from analysis of the data matrix (as described in the text and in fig. 20), with the phylogenetic position of Anteucoila with the two representative figitids constrained. cladograms with respect to placing these taxa family Protocharipinae (Charipidae sensu dubious at best (at least until such time as Kovalev, 1994). Ronquist (1999) transferred females are discovered for both and a new P. costalis to Rasnicynipidae and kept P. analysis can be undertaken). Accordingly, we evenhuisi as a tribe of Charipinae (then in have chosen to concentrate our attention on Figitidae), pointing out that the actions were the few preserved features (i.e., actual, observ- only tentative in the absence of supporting able characters rather than unknown female synapomorphies. However, Protimaspis pos- attributes) of the two genera, in particular those sesses several plesiomorphies that are found traits that have been previously identified as only in Austrocynipidae and Ibaliidae, in- being synapomorphies of individual groups. We cluding the venation and the manner by which have used such characters for tentative place- the mesocoxa is inserted into the mesopectus. ments of the two genera within the overall The cladistic analysis presented herein sug- phylogenetic hierarchy of Cynipoidea. gests that Protimaspis represents a clade basal Protimaspis costalis was considered by to all cynipoids except Austrocynipidae and Kinsey (1937) to be closely related to the tribe Ibaliidae (figs. 21, 25), and it is therefore Aylacini of the Cynipidae. Kovalev (1994) accorded familial status as the Protimaspidae. included it with another fossil species of As discussed above, the family was probably slightly younger age (i.e., Protocharips even- a parasitoid of wood-boring insect larvae, as is huisi Kovalev: appendix 1) in his new sub- the case for modern macrocynipoids. 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 35

Fig. 22. Strict consensus tree of the two shortest trees (L 5 235, CI 5 54, RI 5 79) resulting from analyses of the data matrix (as described in the text and in fig. 20), with the inclusion of two additional Cretaceous amber taxa known only from males (i.e., Tanaoknemus and Micropresbyteria) and no constraints.

Above Protimaspidae on the tree is in a basal subfamily. The genus Goerania can Stolamissus, representing the sister group of be placed well within Liopteridae with cer- liopterids and microcynipoids. As already tainty, even though it was not included in the mentioned, the family has several symplesio- analyses. As already noted, the genus was morphies shared with the macrocynipoids as excluded from the analyses since few of the well as the derived feature of the mesocoxa characters in the matrix could be readily inserted downward on the mesopectus observed on the fossil, resulting in a disturb- (fig. 25). The family is accordingly treated ingly large number of missing data points. herein as the sole member of a new family, the Nonetheless, a few of the preserved features Stolamissidae. permit us to identify it as belonging to the Two groups are allied to, or fall within, the clade consisting of the Gondwanan subfami- Liopteridae. Proliopteron represents the sister lies Liopterinae and Oberthuerellinae. As clade to Liopteridae, with the relationship noted, these synapomorphies include: lateral being particularly supported by two synapo- margin of axilla conspicuously raised anteri- morphies: nucha long and metasosoma in- orly; petiolar annulus long, median dorsal serted high on propodeum (fig. 25). In addi- length at least twice as long as minimum tion, the enlarged fifth metasomal tergum (5 dorsal width; and posterior margin of petiole sixth abdominal) further suggests liopterid and anterior margin of metasomal T2 abut- affinity. The concept of the family is here ting, with dorsal margins of the two more or expanded to include the genus, which is placed less continuous. We can also positively ex- 36 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 23. Strict consensus tree of the two shortest trees (L 5 239, CI 5 55, RI 5 79) resulting from analysis of the data matrix from figure 22 (i.e., with Tanaoknemus and Micropresbyteria included), with the phylogenetic relationship of Anteucoila with the two representative figitids constrained.

clude Goerania from crown-group Ober- to resolve the cladistic position of these genera thuerellinae as well as from crown-group with other cynipoids owing to the unavailabil- Liopterinae by the absence of foveate sculp- ity of females. The longitudinally distinctly ture on the lateral surface of the pronotum compressed head and the lack of distinct and by the postpetiolar terga of the metasoma sculpture on head and mesosoma, short being subequal in length. However, we are anterior pronotal flange, and the downward uncertain whether these characters unite insertion of the meso- and metacoxae indicate Liopterinae and Oberthuerellinae relative to that Micropresbyteria is a microcynipoid. The Goeraniinae, and we prefer to consider the presence of an obvious dorsal pronotal area, subfamily in a trichotomy with the modern the position of the proximal end of Rs+M lineages. close to the posterior end of the first free As discussed earlier, we think that abscissa of M (i.e., the basal vein), and the Micropresbyteria and Tanaoknemus can be presence of the longitudinal carina dorsolat- provisionally placed in Figitidae and Cyni- erally on the metatibia strongly suggest that pidae, respectively, although we were unable the genus belongs to the Figitidae. Conversely, 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 37

Fig. 24. Character optimization showing unambiguous character changes on internodes of one of the single shortest trees from figure 23 (the two minimal length topologies differ only in relationships within Liopteridae, and thus the subfamilies of this group are collapsed into a single terminal for the purposes of the figure, with those unambiguous character optimizations for the modern liopterid clade being noted). Solid circles indicate unambiguous character transformations; open circles show homoplastic character transformations.

the lack of the lateral pronotal carina, Cynipoidea support the notion of an origin presence of a bulla in Sc+R1, and Rs+M and initial divergence of the superfamily in the issuing from the middle of the first free (Ronquist, 1995a, 1995b; Nordlander abscissa of M (i.e., the basal vein) in et al., 1996; Liu, 1998a, 1998b, 2001). Tanaoknemus suggest affinity with the Rasnitsyn (1988, 2002) placed the origin of Cynipidae. The relatively long petiole of the Cynipoidea in the Late Jurassic based Tanaoknemus, a feature often found in mainly on the existence of Early Cretaceous Figitidae, is not found in the extant members fossils of the extinct family Archaeocynipidae, of Cynipidae. We prefer to tentatively include which he thought to be stem-group cynipoids the genus in Cynipidae based on the afore- (Rasnitsyn and Kovalev, 1988; Rasnitsyn, mentioned three characters. 1988; Kovalev, 1994, 1995). Kovalev (1994) also placed great emphasis on the characters DISCUSSION embodied in Archaeocynipidae for under- standing the early evolution of the Cyni- The Cynipoidea has been widely regarded as poidea. Indeed, this author placed the Cyni- being an ancient lineage within Hymenoptera poidea into its own infraorder, Cynipomor- (e.g., Rasnitsyn, 1988; Ronquist et al., 1999). pha, and created a second superfamily, Biogeographic patterns in the basal lineages of Archaeocynipoidea, to include, aside from 38 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 25. Preferred cladogram of cynipoid relationships; character optimization showing unambiguous character changes on internodes of the single shortest tree from figure 21 (the two minimal length topologies differ only in relationships within Liopteridae, and thus the subfamilies of this group are collapsed into a single terminal for the purposes of the figure, with those unambiguous character optimizations for the modern liopterid clade being noted). Solid circles indicate unambiguous character transformations; open circles show homoplastic character transformations.

4 Archaeocynipidae, the extinct families Gero- The name Gerocynipidae as established by Kovalev 4 5 (1994) has, quite unfortunately, some serious nomencla- cynipidae, Rasnicynipidae, and Palaeocy- tural problems and is in fact unavailable. The trouble nipidae. Under this scheme the archaeocyni- stems from the fact that the species Gerocynips zherichini poids were a grade from which Cynipoidea Kovalev is a nomen nudum. Kovalev (1994), when describing the family, refers to an earlier article establish- ing the genus Gerocynips and its type species, G. zherichini. nomenclatural difficulty by newly validating Gerocyni- The article is cited as appearing as a chapter in Price et al. pidae and Gerocynips, but leaving G. zherichini as a nomen (1994), which represents the results of a meeting in nudum and instead designating one of the two actually Krasnoyarsk, Siberia (August 9–13, 1993) under the described species as the type. The nomenclatural details are auspices of the International Union of Forestry as follows: Gerocynipidae Liu and Engel, new family (type Research Organizations (IUFRO) and the Institute of genus: Gerocynips Liu and Engel, new genus), diagnosis as Forest and Wood, Siberian Branch of the Russian that provided by Kovalev (1994); Gerocynips Liu and Engel, Academy of Sciences. In examining a copy of this new genus (type species: G. siberica Kovalev, 1994), publication, however, no article by Kovalev could be diagnosis as that provided by Kovalev (1994). found. It therefore appears as if Gerocynips were first 5Originally proposed as Rasnitsyniidae (Kovalev, 1994), established in Kovalev (1994). Unfortunately, the ICZN the family was subsequently renamed Rasnicynipidae (1999: art. 13) dictates that all new taxa must have an (Kovalev, 1996) owing to homonymy of the type genus, available species explicitly designated as type species. Rasnitsynia, with a generic name established in Braco- Although Kovalev (1994) does indeed designate G. nidae (Pagliano and Scaramozzino, 1989). Interestingly, zherichini as the type species, this species is not described Kovalev (1995), in a key referring to the same material of therein (again referring back to the elusive article in Price 1994, used the replacement name as ‘‘Rasnicynipidae fam. et al., 1994). Thus, Gerocynips Kovalev, 1994 is an nov.’’ Although a description was provided (in the key unavailable name and G. zherichini a nomen nudum. couplet), it was not based on an available genus-group Similarly, the family-group name Gerocynipidae Kovalev, name at that time (the replacement name Rasnicynips was 1994 is rendered unavailable since it is based on an not proposed until the 1996 paper) and thus unavailable type genus (it is not subsequently made Rasnicynipidae was not validated until later in Kovalev available by Kovalev, 1995). We herein correct this (1996). 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 39 ~ ~ ~ trj '"'0 Cj p.. · 0 - . . wptennae Oberthuerellinae CYNIPIDAE AUSTROCYNIPIDAE FIGITIDAE Mayrellinae Dallatorrellinae IBALIIDAE L e n Neoge I I I I - "Tj ..... r I I I I l.l. ~iO .-.~ (1q ~ ..., o I I I I - T - ..... - ~- ttl 0 3 DAE I I I I - :::::. I a ttl (') Paleogene CENOZOIC erinae IDAE TIMASPI eraniinae ian I I I I ~PRO :Proliopt 0 - I ~Go Senon I I I I I I .... T - ~ cr-:; · I I I I T - ~ ~STOLAMISSIDAE I I I I l I ~ ~ ~GEROCYNIP i ...... - ...... Gallic - - CRETACEOUS Neocomian

Fig. 26. Phylogeny of the Cynipoidea summarizing relationships among major lineages. Fossil records are tabulated in appendix 1. Abbreviations for particular deposits are: Rott (Rott, Germany), Fr (Cantal, France), Wig (Isle of Wight, England), Flor (Florissant, Colorado), Biamo (Biamo, today Bol’shaya Svetlovodnaya), Baltic (Baltic amber), Can (Canadian amber), Sib (Siberian amber of the Taimyr Peninsula), NJ (New Jersey amber), and Obe (Obeshchayushchiy Creek). 40 AMERICAN MUSEUM NOVITATES NO. 3583

Fig. 27. Phylogeny of Eucoilinae after Fontal-Cazalla et al. (2002) with the putative positions of the three Cretaceous amber taxa (Anteucoila, Jerseucoila, and Syneucoila) being noted (support for clades is outlined in table 4). arose. Unfortunately, although Gerocynipi- cynipoid in the Early Cretaceous amber of dae, Rasnicynipidae, and Palaeocynipidae are Lebanon would be particularly revealing. indeed cynipoids (table 1), the Archaeocyni- While the position of archaeocynipids in pidae do not belong to the Cynipoidea and are Cynipoidea has eroded, the Gerocynipidae, instead more closely allied to the Rasnicynipidae, and Palaeocynipidae were (Ronquist, 1999). As such, to date there are no indeed correctly assigned to the superfamily. cynipoid fossils from the Jurassic or Early Ronquist (1999) considered Rasnicynipidae to Cretaceous, and we therefore cannot directly be the most basal among the microcynipoids, substantiate a Jurassic origin of the superfam- and Gerocynipidae as the next most basal ily. However, the presence in early stages of branch of the stem lineage leading to the the Late Cretaceous of various macrocyni- extant microcynipoid families Cynipidae and poids and, more significantly, a relatively Figitidae. Ronquist arrived at this conclusion derived liopterid (Goerania) along with de- mainly because both Rasnicynipidae and finitive eucoilines (Figitidae) indicate that the Gerocynipidae, among several features, have more basal divergences within Liopteridae and meso- and metacoxae that are directed down- across basal Cynipoidea must have predated ward (Kovalev, 1994; Ronquist, 1999), which the middle Cretaceous. The phylogenetic is the putatively derived state by contrast to placement of these fossils therefore suggests that seen in Austrocynipidae and Ibaliidae an origin of the Cynipoidea sometime in at (also in Protimaspis: see above) where the least the Early Cretaceous (fig. 26). As noted, meso- and metacoxae are directed obliquely however, a Late Jurassic origin, as suggested backward (Ronquist, 1995a). The illustration by biogeographic patterns (Nordlander et al., of the forewing of Rasnicynips, the sole genus 1996; Ronquist, 1995b; Liu, 1998a, 1998b, of Rasnicynipidae, as presented by Kovalev 2001), cannot be excluded, and exploration of (1994: fig. 19 and under the name Rasnitsynia) Early Cretaceous deposits might significantly shows the proximal end of Rs+M directed extend the ages of various cynipoid families. A toward the posterior end of the first free 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 41

TABLE 4 the Cretaceous. Perhaps not surprisingly, the Clade Support for Phylogeny of Eucoilinae Cretaceous fauna includes two genera that are (Briefly summarized from Fontal-Cazalla et al., clearly stem groups of modern Eucoilinae. 2002, with fossil eucoilines added) While possessing the distinctive and unique mesoscutellar plate of the subfamily, Anteu- Subfamily Eucoilinae coila and Jerseucoila have the metasomal 1. Mesoscutellar cup present 2. Mesopleuron with longitudinal carina present segments primitively separated (figs. 12, 14), 3. Malar sulcus present (difficult to see in Jerseucoila the fusion of the second through fourth and Anteucoila) metasomal terga representing another feature of modern eucoilines (fig. 27, table 4). The Crown-group eucoilines 4. Metasomal T2–4 fused genus Syneucoila (fig. 16), on the other hand, is more intriguing in that it is not only a typical ‘‘Neo-eucoilines’’ crown-group eucoiline but it is apparently 5. Broad metasubpleural depression present intermediate between the basal ‘‘Gronotoma Zaeucoila group + core eucoilines group’’ (or tribe Diglyphosemini) and a clade 6. Subalar pit reduced consisting of the ‘‘Zaeucoila group’’ and core 7. Subalar area of mesopleuron reduced 8. Short vertical carina issuing from ventral margin of eucoilines (sensu Fontal-Cazalla et al., 2002) antennal torulus (fig. 27). The appearance of a ‘‘neo-eucoiline’’ as long ago as the Late Cretaceous further attests to the antiquity of the subfamily and abscissa of M (i.e., the basal vein), a derived thereby the figitids as a whole. character of the Figitidae not found in any Naturally, the cynipoids are most widely other cynipoids. We therefore transfer known by the gall wasps (Cynipidae). Rasnicynips to Figitidae, tentatively classified Although the secondary derivation of phyto- therein as a basal subfamily, although more phagy in the ancestor of cynipids from the work is required for a more definitive place- grade of parasitoids comprising the remainder ment (table 1 and appendix 1). Gerocynipidae of the superfamily is clear (e.g., Malyshev, has been more difficult to place, largely owing 1968; Roskam, 1992; Ronquist, 1995b, 1999; to the less perfect preservation as the family is and analyses herein), the age and means of this known only as compressions from the Late significant evolutionary transition remain un- Cretaceous of the Russian Far East (Kovalev, known. All Cretaceous cynipoid fossils dis- 1994). Kovalev (1994) concluded that the covered to date belong to parasitoid families. gerocynipids were gall inducers because only Although the Gerocynipidae could be gall- females were discovered and their large and makers as supposed by Kovalev (1994), this round resembled several archaic assertion is entirely speculative. The extinct lineages of modern gall-inducing cynipids. As cynipid subfamily Hodiernocynipinae, repre- correctly noted by Ronquist (1999), such sented solely by compressions (appendix 1), features would indicate that the gerocynipids was included in Cynipidae by Kovalev (1994), belong to Cynipidae. However, he also rightly but again, no synapomorphy of the family has noted the lack of any observable cynipid been identified in these specimens (Ronquist, synapomorphies in Gerocynipidae and ac- 1999) and they are perhaps stem-group cordingly placed them at the base of micro- cynipids at best. If such a placement of cynipids. Pending the discovery of new mate- hodiernocynipines is correct, it still leaves rial permitting a more definitive assignment, unresolved whether these taxa were gall- we follow Ronquist’s placement of the family. inducers, as the origin of phytophagy could It is particularly remarkable that the Late have taken place in the common ancestor of Cretaceous amber fauna of North America crown-group Cynipidae. However, phyto- should possess species of the subfamily phagy in Cynipidae is assuredly much older Eucoilinae. Eucoilines are perhaps related to than the hodiernocynipines, regardless of their the Pycnostigminae, and together are perhaps dubious phylogenetic position. Tanaoknemus, not basal within Figitidae, suggesting that the described herein, is very likely a true cynipid. radiation of this family took place well within If this is so, then phytophagy may have 42 AMERICAN MUSEUM NOVITATES NO. 3583 originated in the Late Cretaceous and around of the plant family Fagaceae. It is interesting the time of angiosperm diversification (Crane that all these fossils are representative of the and Lidgard, 1989, 1990; Crane et. al, 1995), derived ‘‘woody gallers’’, with the questionable unless, of course, Tanaoknemus is subsequent- position of A. vectensis being pushed momen- ly demonstrated to be basal among true tarily aside. The slightly older K. succinea,aswe cynipids, in which case the same difficulty discussed above, was almost assuredly a woody ascribed to Hodiernocynipinae applies. The rosaceous galler. Regardless, the earliest gallers oldest fossil that can be definitively placed were significantly older (by perhaps as much as in Cynipidae is K. succinea (see above). 45 Ma) than K. succinea. Kinseycynips is therefore the oldest, definitive Among the Cynipidae, the familiar oak gall gall wasp, and it probably induced galls on wasps (tribe Cynipini) are disproportionately herbaceous host plants belonging to the family more diverse compared to other lineages, with Rosaceae (refer to comments in generic de- about 1000 described species (Liljeblad and scription; see above). Considering that de- Ronquist, 1998). As their name suggests, finitive species of the Cynipidae have existed species of the tribe are almost exclusively since at least the middle Eocene (as evidenced associated with of the genus Quercus by Kinseycynips), we think it reasonable to (Fagaceae). It would be of particular interest hypothesize that the gall wasps likely origi- to find evidence on the age of such a diverse nated much earlier than the age of the known lineage and one with such conservatism in host body and trace fossils and that the discovery use (hence the interest in the identity of of more wasp fossils will eventually fill the Cockerell’s A. vectensis). Ronquist (1999) gap, likely extending definitive cynipids into considered the Cynipidae to be from at least the Cretaceous (as we suspect from the 80–85 Ma based on sister-group dating of putative position of Tanaoknemus as well as figitid fossils in Siberian amber (Kovalev, cynipid-like galls from the Late Cretaceous; 1994). Herein we consider the Cynipidae to Scott et al., 1994). be slightly older, perhaps extending to 90 Ma Liljeblad and Ronquist (1998) mapped cyni- (fig. 26). Certaintly, given the derived position pid hosts on a phylogeny of the family, noting of the oak gall wasps in the family, the that the inquilines formed a monophyletic Cynipini are undoubtedly much younger, group (tribe Synergini). The synergines, con- clearly having a Tertiary origin and radiation. sisting of about 200 species, do not induce galls The Tertiary age of Cynipini is largely themselves but instead live in galls started and dicatated by the fossil record of its hosts. maintained by other species. The Synergini, Although an exceedingly primitive and unde- nested among the gall-inducing cynipids, are scribed flower of Fagaceae has been discov- likely a relatively recent lineage although there ered in New Jersey amber (see Grimaldi and is no direct evidence of their age. Putatively Engel, 2005: fig 14.5), fossils of Fagaceae s. basal lineages of Cynipidae form galls on str. (i.e., excluding Nothofagaceae) are basi- herbaceous plants, especially species of Papa- cally confined to the Tertiary (Herendeen et veraceae and Lamiaceae (Ronquist and al., 1995). Megafossils of the modern genera Liljeblad, 2001). Presumably the ancestral habit Quercus, Castanea, Castanopsis, and Fagus are of Cynipidae was gall-inducing behavior on documented from as long ago as the middle such herbaceous plants, although recent molec- Eocene, and by the Eocene–Oligocene several ular data suggest the opposite, with woody rosid of the major lineages of oaks had originated gallers being more primitive (Nylander et al., (some represented by archaic forms), and the 2004). Cockerell (1921) described two fossils oak flora had clearly become prominent and from the Oligocene (appendix 1) as Rhodites diverse by the Miocene (e.g., Nixon, 1989, vetus and Andricus vectensis. Ronquist (1999) 1993; Herendeen et al., 1995; Manos et al., suggested that the former might well have been 1999, 2001; Manchester and Dillhoff, 2004). correctly placed in the tribe Diplolepidini, the Interestingly, the fossil record of cynipines is wasps that induce galls on Rosa, but he was not more thoroughly documented by paleobotan- certain about the placement of the other species ical evidence rather than by fossils of the oak in Cynipini, the tribe that induces galls on hosts gall wasps themselves (a situation similar to 2007 LIU ET AL.: EVOLUTION OF CYNIPOIDEA 43 the leaf-cutter bees, where the earliest records Ashmead, W.H. 1903. Classification of the gall- of the group come from the distinctive traces wasps and the parasitic cynipoids, or the they produced in fossilized leaves; e.g., Engel, superfamily Cynipoidea, IV. Psyche 10: 2004; Engel and Perkovsky, 2006). Fossils of 210–215. galls are quite diverse in the Tertiary (e.g., Brues, C.T. 1910. The parasitic Hymenoptera of the Tertiary of Florissant, Colorado. Bulletin of Larew, 1986, 1992). The earliest, definitive the Museum of Comparative Zoology 54: evidence of the association between Cynipini 3–125. and oaks are some fossil galls described from Buffington, M., Z. Liu, and F. Ronquist. 2005. middle Miocene (ca. 12.5–15 Ma) deposits of Cynipoidea. In F. Ferna´ndez and M. Sharkey the western United States (Waggoner and (editors), Introduccio´n a los Hymenoptera de la Poteet, 1996; Waggoner, 1999), as well as in regio´n Neotropical. Bogota´ D.C.: Sociedad younger deposits. These ichnofossils are well- Colombiana de Entomologı´a, xx + 870 pp. preserved leaf galls on Quercus hannibali Dorf, Cameron, P. 1905. A new species of Cynipidae from a fossil species morphologically similar to the South Africa, representing a new subfamily. Annals of Natural History, series 7, 16: 20–21. extant species Q. chrysolepis Liebman. It seems Cockerell, T.D.A. 1921. Fossil in the reasonable to assume that cynipines are perhaps British Museum, V. Annals and Magazine of as old as the Eocene, with their radiation Natural History, series 9, 7: 1–25. roughly corresponding with the Late Eocene Crane, P.R., E.M. Friis, and K.R. Pedersen. 1995. through Miocene diversification of oaks. The origin and early diversification of angio- sperms. Nature 374(6517): 27–33. ACKNOWLEDGMENTS Crane, P.R., and S. Lidgard. 1989. Angiosperm diversification and paleolatitudinal gradients in Cretaceous floristic diversity. Science We are grateful to G. Nordlander and F. 246(4930): 675–678. Ronquist for comments on a much earlier Crane, P.R., and S. Lidgard. 1990. Angiosperm version of the manuscript; to B.W.T. Coelho radiation and patterns of Cretaceous palyno- and E. Pen˜alver for critical reviews of the final logical diversity. In P.D. Taylor and G.P. draft; and to J.D. Garnder (RTMP) for the loan Larwood (editors), Major evolutionary radia- of specimens. Z.L.’s participation was partly tions: 377–407. Oxford: Clarendon Press, xi + supported by a Boyd Postdoctoral Research 437 pp. Fellowship (Field Museum of Natural History), Dalla Torre, K.W., and J.J. Kieffer. 1910. by a Kalbfleisch Postdoctoral Research Cynipidae. Das Tierreich 24: 1–891. Fellowship (AMNH), and by a PERT Engel, M.S. 2004 [2005]. Geological history of the bees (Hymenoptera: ). Revista de Postdoctoral Fellowship (Center for Insect Tecnologia e Ambiente 10(2): 9–33. Science, University of Arizona). Portions of Engel, M.S., and E.E. Perkovsky. 2006. An Eocene this work were supported by a Kansas in , Ukraine (Hymenoptera: Technology Enterprise Corporation/Kansas ). American Museum Novitates NSF-EPSCoR grant (KAN29503 to M.S.E.) 3506: 1–12. and by National Science Foundations grants Fontal-Cazalla, F.M., M.L. Buffington, G. EF-0341724 (to M.S.E.), DEB-0542909 (to Nordlander, J. Liljeblad, P. Ros-Farre, J.L. M.S.E.), and DEB-0542726 (to D.A.G.). This Nieves-Aldrey, J. 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APPENDIX 1 Described Fossil Cynipoidea The following table summarizes those fossil species described within the Cynipoidea. The classificatory structure follows that from table 1 in the text. Geological ages follow those of Gradstein et al. (2004). Various fossils originally described as cynipoids but subsequently removed are not included herein. The family {Archaeocynipidae (Rasnitsyn and Kovalev, 1988) was removed from the Cynipoidea by Ronquist (1999) and suggested to be more closely allied to the Diapriidae (). Similarly, various putative ‘‘cynipids’’ in Eocene amber from China (Hong, 2002) are not included as they appear to be misidentifications. In particular, {Sinocynipites fushunensis Hong and {Hemerocynipites wanghuacunensis Hong appear to be proctotrupoid wasps (Proctotrupoidea), {Eocynipites xilutianensis Hong is a male (: Formicidae), and {Eucynipites guchengzensis Hong and {Asiacynipites virides Hong cannot be determined to superfamily (Apocrita incertae sedis). Hong’s ability to grossly misidentify specimens and to oversplit taxa such that nearly everything resides in a new genus or higher taxonomic entity is infamous (all of this material needs to be reevaluated and revised). Lastly, the species {Gerocynips zherichini Kovalev is excluded as this is a nomen nudum (refer to footnote 4 in the text). Note also that Brues’ (1910) Andricus myrciae is in fact a synonym of Cecidomyia pontaniiformis Cockerell (Kinsey, 1919) and is therefore not included here.

Taxon Deposit Age

Family IBALIIDAE Thomson, 1862 Subfamily Incertae sedis Genus {Protoibalia Brues, 1910 {Protoibalia connexiva Brues, 1910 Florissant, Colorado Eocene–Oligocene Subfamily Ibaliinae Thomson, 1862 Genus Ibalia Latreille, 1802 {Ibalia sp. (Nel, 1996) Cantal, France Miocene Family {PROTIMASPIDAE Liu and Engel, n. fam. Subfamily {PROTIMASPINAE Liu and Engel, n. subfam. Genus {Protimaspis Kinsey, 1937 {Protimaspis costalis Kinsey, 1937 Canadian amber Campanian Family {STOLAMISSIDAE Liu and Engel, n. fam. Subfamily {STOLAMISSINAE Liu and Engel, n. subfam. Genus {Stolamissus Liu and Engel, n. gen. {Stolamissus mirabilis Liu and Engel, n. sp. New Jersey amber Turonian Family LIOPTERIDAE Ashmead, 1895 Subfamily {Proliopterinae Liu and Engel, n. subfam. Genus {Proliopteron Liu and Engel, n. gen. {Proliopteron redactus Liu and Engel, n. sp. Canadian amber Campanian Subfamily {Goeraniinae Liu and Engel, n. subfam. Genus {Goerania Liu and Engel, n. gen. {Goerania petiolata Liu and Engel, n. sp. Canadian amber Campanian Family {GEROCYNIPIDAE Liu and Engel, n. fam. Subfamily {GEROCYNIPINAE Liu and Engel, n. subfam. Genus {Gerocynips Liu and Engel, n. gen. {Gerocynips zherichini Kovalev, 1994 Obeshchayushchiy Creek Cenomanian {Gerocynips sibirica Kovalev, 1994 Obeshchayushchiy Creek Cenomanian {Gerocynips florenskayae Kovalev, 1994 Obeshchayushchiy Creek Cenomanian Genus {Antiquecynips Kovalev, 1994 {Antiquecynips orientalis Kovalev, 1994 Obeshchayushchiy Creek Cenomanian Genus {Arctogerocynips Kovalev, 1994 {Arctogerocynips magadanicus Kovalev, 1994 Obeshchayushchiy Creek Cenomanian Family FIGITIDAE Thomson, 1862 Subfamily Incertae sedis Genus {Micropresbyteria Liu and Engel, n. gen. {Micropresbyteria caputipressa Liu and Engel, n. sp. Canadian amber Campanian 48 AMERICAN MUSEUM NOVITATES NO. 3583

APPENDIX 1 (Continued)

Taxon Deposit Age

Subfamily {RASNICYNIPINAE Kovalev, 1996 Genus {Rasnicynips Kovalev, 1996 {Rasnicynips eximia (Kovalev, 1994) Taimyr amber Santonian Subfamily {PALAEOCYNIPINAE Kovalev, 1995 Genus {Palaeocynips Kovalev, 1994 {Palaeocynips arcticus Kovalev, 1994 Taimyr amber Santonian Genus {Palaeocynipiana Kovalev, 1994 {Palaeocynipiana santonica Kovalev, 1994 Taimyr amber Santonian Subfamily CHARIPINAE Dalla-Torre and Kieffer, 1910 Genus {Protocharips Kovalev, 1994 {Protocharips evenhuisi Kovalev, 1994 Taimyr amber Santonian Subfamily EUCOILINAE Thomson, 1862 Genus {Anteucoila Liu and Engel, n. gen. {Anteucoila delicia Liu and Engel, n. sp. Canadian amber Campanian Genus {Jerseucoila Liu & Engel, n. gen. {Jerseucoila plesiosoma Liu and Engel, n. sp. New Jersey amber Turonian Genus {Syneucoila Liu & Engel, n. gen. {Syneucoila magnifica Liu and Engel, n. sp. New Jersey amber Turonian Subfamily FIGITINAE Thomson, 1862 Genus Figites Latreille, 1802 {Figites solus Brues, 1910 Florissant, Colorado Eocene–Oligocene Genus {Palaeofigites Kovalev, 1995 {Palaeofigites balticus Kovalev, 1995 Baltic amber Eocene (Lutetian) Subfamily ASPICERINAE Dalla Torre and Kieffer, 1910 Genus {Palaeoaspicera Kovalev, 1994 {Palaeoaspicera orientalia Kovalev, 1994 Taimyr amber Santonian Family CYNIPIDAE Latreille, 1802 Subfamily Incertae Sedis Genus {Tanaoknemus Liu and Engel, n. gen. {Tanaoknemus ecarinatus Liu and Engel, n. sp. Canadian amber Campanian Subfamily {HODIERNOCYNIPINAE Kovalev, 1994 Genus {Hodiernocynips Kovalev, 1994 {Hodiernocynips primigenius Kovalev, 1994 Bol’shaya Svetlovodnaya Eocene–Oligocene (Biamo) {Hodiernocynips planus (Statz, 1938) Rott, Germany Miocene (Aquitanian) {Hodiernocynips rotundatus (Statz, 1938) Rott, Germany Miocene (Aquitanian) {Hodiernocynips spiniger (Statz, 1938) Rott, Germany Miocene (Aquitanian) {Hodiernocynips progenitrix (Kinsey, 1919) Florissant, Colorado Eocene–Oligocene {Hodiernocynips ampliforma (Kinsey, 1919) Florissant, Colorado Eocene–Oligocene Subfamily CYNIPINAE Latreille, 1802 Genus {Kinseycynips Liu and Engel, n. gen. {Kinscynips succinea (Kinsey, 1919) Baltic amber Eocene (Lutetian) Genus Geoffroy, 1762 {Diplolepis vetus (Cockerell, 1921) Isle of Wight, England Oligocene Genus Andricus Hartig, 1840 {Andricus vectensis Cockerell, 1921 Isle of Wight, England Oligocene Incertae Sedis {‘‘Cynips’’ succinea Presl, 1822 Baltic amber Eocene (Lutetian)