Synoptic Lists of World Genera and Family-group Names

By David Wahl

 

 

Synoptic lists of genera and family-group names:

 

·         Downloadable pdf of the genera of Acaenitinae - Campopleginae is available here

·         Downloadable pdf of the genera of Collyriinae - Cylloceriinae is available here

·         Downloadable pdf of the genera of Diacritinae - Ichneumoninae is available here

·         Downloadable pdf of the genera of Labeninae - Poemeniinae is available here

·         Downloadable pdf of the genera of Rhyssinae - Xoridinae is available here

·         Downloadable pdf of an alphabetical list of World genera of Ichneumonidae is available here

·         Downloadable pdf of the fossil genera of Ichneumonidae is available here

·         Downloadable pdf of the references used for this page and the above lists is available here

·         Downloadable pdf of the family-group names of Ichneumonidae is available here

 

     With the exception of the Ichneumoninae, Townes provided a comprehensive higher-level classification of the Ichneumonidae in his four generic monographs (Townes, 1969, 1970a&b, 1971). The ensuing 43 years have seen many changes, both at the generic level and in the number of subfamilies, as well as the publication of several regional catalogues and treatments (Townes & Townes, 1973; Carlson, 1979; Gauld, 1984; Gupta, 1987). Yu & Horstmann’s (1997) world catalogue gave an excellent consensus classification to which most active specialists would subscribe. I believe, however, that a need exists for a readily accessible listing of subfamilies and genera that can be periodically updated. A Web site with downloadable and searchable PDF files seems to be the ideal solution.

 

     Subfamilies are arranged alphabetically, as are tribes (when present) and constituent genera. Genus groups have been used in the Campopleginae, Ophioninae, Orthocentrinae, and Pimplinae. Ease of use is the paramount consideration. Townes' arrangement of genera within his subfamilies is a linear listing of various groups of genera, and unless one is intimately familiar with the systematics of a given subfamily or tribe, there is no way to tell where one group ends and another begins. Moreover, his classification is based on overall resemblance. In the few instances in which cladistic analyses have been performed, generic relationships are quite different (for example, see Gauld (1985) for Ophioninae, and Wahl & Gauld (1998) and Gauld et al. (2002) for the Pimpliformes.)

    

     In addition to the subfamily entries, I've also provided an alphabetical listing of all generic names. Junior synonyms are cross-indexed to the senior name, which has information on distribution and subfamilial placement.

 

     Townes' idiosyncratic family-group nomenclature is not used. In the matter of the application of Pimpla, Ephialtes, and Ichneumon, I follow the International Commission on Zoological Nomenclature (ICZN) for the reasons given in Wahl & Mason (1995).

 

     The foundation of the lists is Townes' series of four subfamilial monographs (except for the Ichneumoninae, where the four regional catalogs of Townes and his collaborators are my guides, as well as my own experience with the subfamily). The anomalonine genus Gravenhorstia is a typical entry:

 

            Gravenhorstia Boie, 1856

              subgenus Erigorgus Förster, 1869; Holarctic, Neotropical, Oriental (Gauld, 1976; Dasch, 1979)

                 Sympratis Förster, 1869

                 Paranomalon Viereck, 1912

              subgenus Gravenhorstia Boie, 1856; Palearctic

                 Odontopsis Förster, 1869

              subgenus Kokujewiella Shestakov, 1926; Palearctic (Gauld, 1976)

                 Nenethes Ceballos, 1957 (Gauld, 1976; Atanasov, 1982)

              subgenus Ribasia Ceballos, 1921; Palearctic (Gauld, 1976; Atanasov, 1982)

 

     Erigorgus, Kokujewiella, and Ribasia were treated by Gauld (1976) as subgenera of Gravenhorstia; Townes (1971) placed Kokujewiella and Nenethes as junior synonyms of Erigorgus, and Ribasia was recognized as a separate genus. Dasch (1984) kept Erigorgus as a separate genus, and Atanasov (1982) treated Nenethes and Ribasia as separate genera as well. It will be noted that I follow Gauld's classification, which was the result of a comprehensive study that dealt with the world fauna, whereas Dasch's and Atanasov's are regional treatments. I will be the first to admit that I have made decisions based upon my systematic experience and prejudices. Differing viewpoints, however, are documented and may be investigated by the interested user.

 

     DNA studies of ichneumonid phylogeny have been equivocal. D.L.J. Quicke and his collaborators (Belshaw et al., 1998; Quicke et al. 1999, 2000; Broad et al., 2005; Laurenne et al., 2006; Quicke et al., 2009) have used 28S rDNA either solo or in combination with morphology to explore various phylogenetic questions. The results are broadly suggestive, but

the studies vary so much in tree structure that definitive taxonomic decisions are nearly impossible to implement. Quicke et al. (2009) used 1001 species and is the most comprehensive analysis to date, but: 1) as with previous Quicke studies, only 28S is used and secondary structure is not taken into account; 2) the morphological characters are mostly coded at the subfamily or tribe levels, although individual species were entered (the drawbacks of this approach were discussed in Gauld et al. (2002) under the topic of 'generic abstractions vs. exemplar species'); 3) the set of morphological characters were weighted so that they had equal weight to the molecular data, taking into account the homoplasy of each data set as judged by their retention indices. In summation, the picture appears to be that: 1) the subfamily Xoridinae is the sister group to the rest of the family; 2) the subfamily  Labeninae is near the base of the family, 3) Ophioniformes/Ichneumoniformes/Pimpliformes are more or less recovered, depending how the groups have been reinterpreted (as with Ophioniformes in Quicke et al., 2000). Beyond that, it is hard to makes sense of most of the subfamilial groupings or have confidence in the sunderings of various subfamilies or tribes, especially those that seem to be morphologically well-established. It would appear that a 700 -base pair DNA segment is not sufficient to accurately reconstruct relationships, and that accurate analyses of ichneumonid DNA will have to wait for multigene studies that utilize far more data (as in the aculeate studies of Pilgrim et al. (2008) and Debevec et al. (2012)). Certain findings of the Quicke corpus seem justified, however, and they are mentioned below under the appropriate subfamilies.

 

     Finally, I might note that the list of fossil Ichneumonidae does not include the subfamily Tanychorinae or the genera included in this 'subfamily': Amplicella, Megachora, Paratanychora, Tanchora, and Tanychorella (Rasnitysn, 1980; Kopylov, 2010b). Sharkey & Wahl (1992) demonstrated that Tanychorinae are incertae sedis within Ichneumonoidea.

 

     What follows is a brief commentary on each subfamily, indicating broad changes since Townes' monographs.

 

ACAENITINAE

     No substantial changes since Townes (1971) except for the description of two new genera (Asperpunctatus Wang and Dentifemura Sheng & Sun) and the abolition of tribes (Wahl & Gauld, 1998).

 

AGRIOTYPINAE

     No changes since Townes (1969). Bennett (2001) provided keys to the world species.

 

ADELOGNATHINAE

     No changes since Townes (1969).

 

ANOMALONINAE (= Anomalinae of Townes)

     I have followed Gauld's (1976) generic revision, which is notable for: 1) synonymizing Ophionellini and Podogastrini under Gravenhorstiini (= Theriini of Townes), and 2) synonymizing a number of genera recognized by Townes.    Dasch's (1979) division of the subfamily into Anomaloninae s.s. and Gravenhorstiinae (= Theriinae of Dasch) is not recognized for the reasons given in Wahl (1991).    

     Seven new genera have been proposed since Townes (1971). Gauld (1976) provided a new key to the World genera; Dasch (1984) provided a new key to the Nearctic genera.

 

BANCHINAE

     No substantive changes since Townes (1970b) except for the description of some additional genera. Lissonotini is a junior synonym of Atrophini (Gauld, 1984). Townes & Townes (1978) provided a new key to the Nearctic Atrophini.

     Kasparyan (1993) erected the Townesioninae for Sachtlabenia (formerly in the Glyptini) and a new genus, Townesion; he believed it to be related to Lycorininae, Stilbopinae, and Banchinae. Gauld & Wahl (2000b) considered these genera to be derived Glyptini and sank Townesioninae into that tribe. Gauld and his collaborators (Gauld, 2002) described several new genera, synonymized others, and discussed the problems with previous generic limits in Atrophini that were based primarily upon Holarctic concepts.

 

BRACHYCYRTINAE (part of Labiinae of Townes)

     At the time a tribe in Labeninae, Gauld (1983) changed the group's definition and composition by the removal of Poecilocryptus to its own tribe in the Labeninae. A new genus, Monganella Gauld, was described in 1984. Wahl (1993) removed the tribe from the Labeninae and elevated it to subfamilial rank. Porter (1998) elevated Pedunculus to subfamily, but listed only autapomorphies and gave no cogent reason why the genus did not belong in the Brachycyrtinae. Gauld & Ward (Gauld, 2000) studied the situation and argued that: a) Brachycyrtus and Pedunculus belong in separate subfamilies, and b) Adelphion and Monganella are best placed with Pedunculus in the Pedunculinae. Brachycyrtinae is thus restricted to Brachycyrtus.

 

CAMPOPLEGINAE (= Porizontinae of Townes)

     While some new genera have been erected and others synonymized, the biggest changes were: 1) the proposal of informal genus-groups in place of the tribes used by Townes and other workers (Wahl, 1991), 2) the elevation of Nesomesochorini to subfamily status (Miah & Bhuiya, 2001), and 3) the removal of Hellwigia and Skiapus (Quicke et al., 2005)- see the Ophioninae section, below.

 

COLLYRIINAE

     Two additional genera, Aubertiella Kuslitzky & Kasparyan and Bicurta Sheng, Broad, & Sun, have been described.

 

CREMASTINAE

     No substantive changes since Townes (1971) but for the description of new genera and the synonymization of others. Dasch (1979) provided a new key to Nearctic genera.

 

CRYPTINAE (= Phygadeuontinae and Hemitelinae of authors, Gelinae of Townes)

     The most important changes have been at the tribal level, both in nomenclature and composition. The name of the subfamily has been controversial for many years due to uncertainties about the application of Cryptus. Townes refused to accept the validity of Opinion 157 of ICZN (which placed Cryptus Fabricius, 1804 on the Official List of Generic Names in Zoology) and used Itamoplex for what was hitherto known as Cryptus; under his idiosyncratic system of nomenclature, Gelinae became the name of the subfamily. Although Opinion 157 is valid (Wahl & Mason, 1996), the ICZN failed to suppress Cryptus Panzer, 1804, resulting in Phygadeuontinae as the correct name for the subfamily (Fitton & Gauld, 1978). Opinion 1757 (ICZN, 1994) validated Cryptus Fabricius, 1804 by suppressing Cryptus Panzer, 1804, thereby changing the subfamily’s name to Cryptinae (and Cryptini in place of Mesostenini). The best comment on the matter is Gauld’s (1995: 415): "However, using their plenary powers, the International Commission on Zoological Nomenclature recently overturned good scholarship and the strict application of their own rules, preferring instead to validate names used by a few workers who had earlier chosen to ignore established ICZN rules of nomenclature ... Unfortunately, there is no appeal against such arbitrary abuse of plenary power by those charged with supposedly conserving nomenclatural stability, so I am here reluctantly adopting the use of Cryptinae for the group I have, in previous publications, referred to as the Phygadeuontinae."

     Claseini was transferred from the Labeninae to the Cryptinae by Gauld (1983). Gauld (1995) made a substantial change in the Hemigasterini (= Echthrini of Townes). He noted that the tribe included two dissimilar groups: the Hemigaster genus-group (Hemigaster, Litochila, Mansa) and the Aptesis genus-group (the remaining genera). As the two groups do not share any synapomorphies, Gauld transferred the Hemigaster genus-group to the Cryptini (with Hemigastrini becoming a junior synonym), leaving the Aptesis genus-group as the Aptesini. While Gauld is certainly correct in expressing dissatisfaction with the present composition of the Hemigasterini, I am not convinced that his actions represent an improvement. To the best of my knowledge, the tribe Cryptini is defined by the loss of the triangular extensions of the metanotum. Hemigaster and its related genera do possess the extensions: placing them in the Cryptini effectively destroys its monophyly unless one can come up with arguments that the extensions represent reversals in the Hemigaster genus-group. As for the Aptesini, removal of the Hemigaster genus-group still leaves it non-monophyletic. While many of these genera attack sawflies, a number of genera also parasitize Coleoptera and Lepidoptera. Lacking a cladistic analysis of the group, no decision can be made what represents the ground-plan biology. Although a heterogeneous, non-monophyletic Hemigasterini is unsatisfactory, Gauld's changes still leave a large, non-monophyletic group (Aptesini) and obfuscates that status of a previously monophyletic group (Cryptini). His changes are hence not followed. Laurenne et al. (2006) adds nothing of substance to the debate, as the results are a strange mix of familiar groupings with often wildly disparate elements.

     A few Indian authors (Gupta, 1970, 1986; Jonathan & Gupta, 1973) have elevated the cryptine tribes to subfamilies. The claim that "… Mesostenini as defined by Townes is sufficiently large and distinctive to merit a subfamily rank and therefore it seems advisable to raise it to the subfamily level …" has to date not convinced the ichneumonological community

     Aside from these higher-level issues, there has been the usual description and synonymization of genera.

     Taxapad 2012 (Yu et al., 2012) users will find a confusing situation with regards to the genera of Cryptina. Townes used Mesostenini in place of Cryptini, and early Townes publications (such as Townes et al. (1961)) had most genera in the subtribe Mesostenina. The Mesostenina were drastically redefined in Townes (1970a) and most genera of Mesostenina sensu lato were apportioned to other subtribes, especially Ischnina. The Ischnina of Townes (1970a) is properly known as Cryptina under the International Code of Zoological Nomenclature (ICZN, 1999). Taxapad 2012 apparently works under the guideline of a genus staying within a higher taxon unless its removal is explicitly stated in print. Thus, genera of Cryptina should be correctly placed into Taxapad 2012 subtribes:

Cryptina: Buathra, Caenocryptus, Camera, Chromocryptus, Cryptus, Diplohimas, Dotocryptus, Ferrocryptus, Joppidium, Meringopus, Myrmeleonostenus, Nelophia, Odontocryptus, Reptatrix, Trachysphyrus, Xenarthron.

Ischnina: Biconus, Caenocryptoides, Caenopelte, Compsocryptus,  Cyanodolius, Cyanopelor, Cyclaulus, Dihelus, Distictus, Dochmidium, Enclisis, Etha, Gessia, Glabridorsum, Gyropyga, Hedycryptus, Ischnus, Lanugo, Leptarthron, Mesophragis, Monothela, Nebostenus, Nedodontocryptus, Nippocryptus, Palmerella, Rhynchocryptus, Synechocryptus, Tricentrum, Trihapsis, Xylophrurus, Zonocryptus.

Mesostenina: Aeglocryptus, Aeliopotes, Aglaodina, Anacis, Araucacis, Chilecryptus, Cosmiocryptus, Hypsanacis, Itamuton, Mrymecacis, Neocryptopteryx, Nothischnus, Oecetiplex, Periplasma, Phycitiplex, Picrocryptoides, Sciocryptus, Xiphonychidion, Xylacis, Xylostenus.

 

CTENOPELMATINAE (= Scolobatinae of Townes)

     No substantive changes at the tribal or generic level have been made with the exception of the synonymization of Westwoodiini with Scolobatini by Gauld (1984), and the subsequent re-establishment of both as valid tribes by Zhaurova & Wharton (2009). Gauld & Wahl (2005) suggested that ctenopelmatine monophyly is doubtful and that the metopiines may have arisen within Ctenopelmatinae. Both Quicke et al. (2009) and the more detailed unpublished work by R.A. Wharton and his collaborators (http://mx.speciesfile.org/projects/8/public/public_content) indicate that ctenopelmatines are a paraphyletic group within the Ophioniformes, with Metopiinae and Mesochorinae closely related to one or more ctenopelmatine subgroups.

 

CYLLOCERIINAE (part of Microleptinae of Townes, Oxytorinae of authors)

     Allomacrus and Cylloceria were removed from the Oxytorinae and placed in their own subfamily by Wahl (1990). A new genus, Rossemia, was described by Humala (1997); Wahl & Gauld (1998), using the junior synonym Sweaterella, gave a more comprehensive description and analysis of relationships.

 

DIACRITINAE (part of Ephialtinae of Townes)

     Gauld (1991) raised this former pimpline tribe to subfamilial rank. Wahl & Gauld transferred Daschiana (as Cressonia) from Orthocentrinae to this subfamily.

 

DIPLAZONTINAE

     The only substantive changes have been the substitution of Syrphoctonus Förster, 1869 for Homotropus Förster, 1869, and Woldstedtius Carlson, 1979 for Syrphoctonus Förster, 1869 sensu Dasch, 1964 (Carlson, 1979), plus the description of several new genera.

 

EUCEROTINAE (part of Tryphoninae of Townes)

     Barron (1976) raised this former tryphonine tribe to subfamilial rank. Gauld & Wahl (2002) described a second genus, Barronia.

 

HYBRIZONTINAE (Paxylommatinae of authors)

     Townes did not include this group within the Ichneumonidae, treating it as a separate family (Townes & Townes, 1983). Most authors now consider it to be a subfamily of Ichneumonidae (Rasnitsyn, 1980; Gauld, 1984; Sharkey & Wahl, 1992). Several extant (Tobias, 1988) or fossil (Kasparyan, 1988) genera have been described.

 

ICHNEUMONINAE

     This subfamily was not monographed by Townes and thus there exists a wide array of opinions regarding tribal and generic limits. Based upon Townes' arrangement of the American Entomological Institute collection and my own studies, 15 tribes are recognized here (Wahl & Mason, 1996). I have used Townes' generic concepts as laid out in the regional catalogs produced by him and his collaborators (Townes et al. 1961; Townes et al., 1965; Townes & Townes, 1966, 1973). The observant user will realize there are often substantial differences of opinion between Heinrich and Townes, especially for the Ethiopian and Oriental faunas. Heinrich's generic concepts are used in Gupta (1987). Townes, in my opinion, was better acquainted with the world fauna and I have elected to follow his generic concepts. For the Oriental region, the junior synonyms in Townes et al. (1961) remain as such, even though Heinrich subsequently raised many of these back to generic status. Townes' synonymizations of Heinrich Ethiopian genera (Townes & Townes, 1973) are accepted.

     Ichneumonines are plagued by competing generic nomenclatures, due to Townes' refusal to accept the validity of Opinion 159. The nomenclature used here is that of the ICZN for the reasons put forth in Wahl & Mason (1996):

  Heresiarchini (= Ichneumonini of Townes, Protichneumonini of Heinrich)

     ICZN name                Townes name

             Coelichneumon        Ichneumon

  Ichneumonini (= Joppini of Townes)

      ICZN name             Townes name

       Ichneumon              Pterocormus

    

     Heinrich’s 5-volume magnum opus on Ethiopian ichneumonines is commonly cited as 1967-1968 (Heinrich, 1977; Yu & Horstmann, 1997; Yu et al., 2012) or 1967 (Gauld, 1984; Gupta, 1987). Townes & Townes (1973) cite the volumes as covering the span 1967-1969. Henry Townes’ copy of volume V in the American Entomological Institute library has the dates he received his copies written adjacent to the printed dates of issue on p. 1258. They are as follows (the printed dates of issue are in brackets):

     Volume I – December 20, 1967 [April 3, 1967]

     Volume II – December 20, 1967 [June 28, 1967]

     Volume III – June 18, 1968 [December 21, 1967]

     Volume IV – February 18, 1969 [June 20, 1968]

     Volume V – August 8, 1969 [November 10, 1968]

Townes wrote the following passage below the dates; it is reproduced here in full: “In August, 1969, I asked Hilda Heinrich why the above dates are so much earlier than the dates on which I received copies. Her reply indicates that the above dates are the ones on which the books were printed in Germany. After this, a copy of the books was sent to Gerd Heinrich & the rest came by ship, through customs, and to Farmington College. She believes that I am among the first subscribers to receive copies. This means that the publication dates are 7-10 days prior to the dates that I received my copies. H. Townes.  Aug. 1969.” I have therefore used the publication dates of Townes & Townes (1973).

     Taxapad 2012 placed the following genera in Alomyinae, when they should actually be in Phaeogenini: Liaodontus, Maxodontus, Mevesia, Phairichneumon, and Wahliodontus. Taxapad 2012 also treats six genera described by Porter (1998) as incertae sedis within Ichneumoninae: Barythixis, Chilelabus, Chilhoplites, Ithaechma, Notophasma, and Zophoplites. Examination of these genera show that they belong in the Ichneumonini sensu Heinrich.

 

LABENINAE (= Labiinae of Townes)

     The subfamily's composition was changed substantially since Townes (1969). The Claseini (Gauld, 1983) and Brachycyrtini (Wahl, 1993) have been removed, the Poecilocryptini enlarged (Gauld, 1984), and a separate tribe erected for Xenothyris (Wahl, 1996). In the Groteini, Macrogrotea was synonymized with Grotea (Wahl, 1993), and in the Labenini, Apechoneura and Asperellus were synonymized with Certonotus (Gauld, 1986; Wahl, 1993). Gauld & Wahl (2000a) reanalyzed relationships within the family and several new genera were described.

 

LYCORININAE (part of Banchinae of Townes)

     This former tribe of Banchinae was elevated to subfamilial rank by Townes (Townes & Townes, 1973). Gonioglyphus and Toxophoroides were synonymized with Lycorina by Gauld (1984).

 

MESOCHORINAE

     Wahl's (1993) generic revision resulted in the addition of four new genera, and the synonymization of Oncocotta, Piestetron, Plectochorus, Rhaibaspis, and Stictopisthus with Mesochorus. Several genera have been described since then.

 

METOPIINAE

     Most of the generic definitions are unchanged since Townes & Townes (1959). Townes (1971) added Apolophus, Bremiella, Ischyrocnemis, and Lapton, commenting that "[n]one of these additions are unquestionably metopiines, but this subfamily seems to be the best home for them at the present". Aubert (2000) removed Bremiella and Ischyrocnemis to Ctenopelmatinae without comment. Gauld & Wahl (2005): 1) made Apolophus a junior synonym of the ctenopelmatine genus Scolomus, 2) concluded that Scolomus belongs in the Metopiinae as one of the basal genera, and 3) suggested that ctenopelmatine monophyly is doubtful and that the metopiines may have arisen within Ctenopelmatinae.

 

 

MICROLEPTINAE

     The subfamily was restricted to Microleptes by Wahl (1986). Dasch placed Hyperacmus and Cushmania in this subfamily; Wahl & Gauld (1998) synonymized Cushmania with Hyperacmus, and placed Hyperacmus back in Orthocentrinae.

 

NEORHACODINAE

     Although Townes (1970b) initially treated the group as a tribe of Banchinae, he later (Townes, 1971) raised it to subfamilial status. Kasparyan (1995) described a new genus, Eremura.  The proposal by Quicke et al. (2009) to merge Neorhacodinae and Phrudinae with Tersilochinae is not followed here.

 

NESOMESOCHORINAE

     Townes (1970b) placed the Old World genus Chriodes and the Neotropic genus Nonnus together as the Nonnini, a tribe he placed with some reservations in the Campopleginae. Fitton & Gauld (1976) established the correct name of the tribe as Nesomesochorini. Miah & Bhuiya (2001) elevated this tribe to subfamilial status. Quicke et al. (2009) confirmed that the three genera currently recognized in Nesomesochorinae (Chriodes, Klutiana, and Nonnus) did not belong in Campopleginae.

 

OPHIONINAE

     Gauld (1978, 1979, 1985) greatly changed Townesian generic concepts. In addition, he demonstrated the unsatisfactory nature of the two tribes used by Townes (1971) and arranged the genera into five informal genus-groups. This classification is used here. Hellwigia and Skiapus were transferred to the Ophioninae in Quicke et al. (2005) but only Hellwigia seems to belong here. Pending further investigation, Skiapus is treated as incertae sedis in Ichneumonidae.

 

ORTHOCENTRINAE (part of Microleptinae of Townes, plus Orthocentrinae s.s.)

     Wahl (1990) demonstrated that Microleptinae sensu Townes, after the removal of extraneous elements (Allomacrus, Cylloceria, Microleptes, Oxytorus, Tatogaster) was paraphyletic with respect to Orthocentrinae s.s. The two were merged, with the name Orthocentrinae having priority.

 

ORTHOPELMATINAE

     No changes since Townes (1971).

 

OXYTORINAE

     As restricted by Wahl (1990), the subfamily consists only of Oxytorus.

 

PEDUNCULINAE (part of Labiinae of Townes)

     See the above treatment of Brachycyrtinae for the taxonomic history of Pedunculinae; the subfamily currently consists of Adelphion, Monganella, and Pedunculus

 

PHRUDINAE

     After the establishment of the Sisyrostolinae (see discussion under that subfamily), the subfamily Phrudinae now consists of Astrenis, Earobia, Notophrudus, Peucobius, Phaestacoenitus, Phrudus, and Pygmaeolus. The proposal by Quicke et al. (2009) to merge Neorhacodinae and Phrudinae with Tersilochinae is not followed here.

 

PIMPLINAE (= Ephialtinae of Townes)

     The tribal classification was greatly modified by Gauld (1991) and Wahl & Gauld (1998). Gauld (1991) elevated the Diacritini, Poemeniini, and Rhyssini to subfamilial status, and dismantled the Delomeristini (transferring Pseudorhyssa to the Poemeniinae, Theronia s.l. to the Pimplini, and the remaining genera to the Ephialtini). Wahl & Gauld (1998) resurrected the Delomeristini for Delomerista and Atractogaster, erected Perithoini for Perithous, and sank Polysphinctini into the Ephialtini. Gauld et al. (2002) transferred Perithous to the Delomeristini, and created genus-groups within the Ephialtini. Gauld & Dubois (2006) revised the genera of the Polysphincta genus-group.

 

POEMENIINAE (part of Ephialtinae of Townes; Neoxoridini of authors)

     Gauld (1991) raised this former pimpline tribe to subfamilial rank, at the same time incorporating Pseudorhyssa from the Pimplinae. Wahl & Gauld (1998) recognized three tribes: Pseudorhyssini, Rodrigamini, and Poemeniini.

 

RHYSSINAE (part of Ephialtinae of Townes)

     Gauld (1991) raised this former pimpline tribe to subfamilial rank.

 

STILBOPINAE (part of Banchinae of Townes)

     Townes elevated the tribe to subfamilial rank (Townes & Townes, 1978) but retained Panteles in the Banchinae. Wahl (1988) placed Panteles with the other stilbopine genera.

 

SISYROSTOLINAE (part of Phrudinae of Townes)

     Seyrig (1932) erected the tribe Sisyrostolini in the Pimplinae for Erythrodolius, Icariomimus, and Melandolius. Townes (1971) combined the Sisyrostolini and Townes' Brachyscleromatinae (consisting of Brachyscleroma (Townes et al., 1961)) with the Phrudinae, although he expressed doubt (p. 25) that the genera belonged together. Gupta (1994) suggested that the Brachyscleromatinae should be restored for the large-bodied tropical genera. Quicke et al. (2009) found that the Sisyrostolini and Brachyscleroma were not closely related to the Phrudinae s.s. and removed them to an expanded Brachyscleromatine. Bennett et al. (2013) pointed out that Sisyrostolini Seyrig, 1932 has priority over Brachyscleromatinae Townes, 1961 and that Sisyrostolinae is the correct name of the subfamly. The subfamily consists of the following genera: Brachyscerloma, Erythrodolius, Icariomimus, Laxiareola, Lygurus, and Melanodolius (Bennett et al., 2013).

 

TATOGASTRINAE (part of Microleptinae of Townes and Oxytorinae of authors)

     Tatogaster was removed from the Oxytorinae and placed in its own subfamily by Wahl (1990). Quicke et al. (2009) had Tatogaster as the sister-group to the Mesochorinae and suggested it could be a basal member of the subfamily, although they did allow that it lacked the critical mesochorine autapomorphies (enlarged triangular hypopygium, elongate gonoforceps, and needle-like ovipositor without a dorsal subapical notch). The presence of a rhombic areolet and an elongate metasoma led to a (hopefully whimsical) suggestion that the overall facies was similar to the mesochorine genus Lepidura. It is worth pointing out that Tatogaster has a short, laterally compressed ovipositor with a dorsal subapical notch (Wahl, 1990); although the wasp's biology is unknown, it obviously does not have the mesochorine biology of ovipositing into a parasitoid larva within a host.

 

TERSILOCHINAE

     No substantive changes except for the description of new genera and subgenera. The proposal by Quicke et al. (2009) to merge Neorhacodinae and Phrudinae with Tersilochinae is not followed here.

 

TRYPHONINAE

     Gauld (1984) described a new tribe, Anklyophionini, from Australia. Gupta (1988) restricted Eclytini to Eclytus, with the remaining genera placed in Oedemopsini. Acaenitellus has been transferred from Orthocentrinae to Oedemopsini (Gupta, 1988). Aside from these higher-level changes, little has changed since Townes (1969) except for the description of several new genera and subgenera by Kasparyan and Gauld.

 

XORIDINAE

     Wahl (1997) synonymized all subgenera of Xorides under that genus, recognizing instead species-groups.