Classification and Systematics of the Ichneumonidae (Hymenoptera) David Wahl American Entomological Institute

The list is arranged alphabetically for several reasons. Ease of use is paramount. Secondly, the "natural" arrangement in Townes' volumes is a linear listing of complex branching patterns, and unless one is intimately familiar with the systematics of a given subfamily or tribe, there is no way to tell where one cluster of genera ends and another begins. Thirdly, Townes' classifications are based on overall resemblance: in the few cases where cladistic analyses been performed, they do not accurately reflect relationships (for example, see Gauld (1985) for Ophioninae, Wahl (1991) for Campopleginae, Wahl (1993) for Mesochorinae, and Wahl & Gauld (1998) 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 my list is Townes' series of four subfamilial monographs (except for the Ichneumoninae, where the four regional catalogs of Townes and his collaborators are my guides). 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)

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

ACAENITINAE

No substantial changes since Townes (1971) except for the description of a new genus (Asperpunctatus Wang, 1989) and the abolition of tribes (Wahl & Gauld, 1998).

AGRIOTYPINAE

No changes since Townes (1969) except for the description of a new genus, Atopotypus Chao, 1992.

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).

BANCHINAE

No substantive changes since Townes (1971) except for the description of some additional genera. Lissonotini is a junior synonym of Atrophini (Gauld, 1984).

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. Although Gauld (1997) had not examined Townesion, he discussed his reasons for keeping Sachtlabenia in Banchinae. After examining both genera, I conclude that: 1) the two genera are related, and 2) they are most likely highly derived glyptines (it should be noted that while Kasparyan cited many autapomorphies for Townesioninae, he did not give characters that definitively excluded the two genera from Banchinae or showed relationships to other subfamilies. They are here treated as glyptines.

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 does not belong in the Brachycyrtinae. Gauld & Ward (in Gauld 2000) studied the situation and convincingly 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 change has been the proposal of five informal genus-groups in place of the tribes (Campoplegini, Cymodusini, Hellwigiini, Limneriini, Nesomesochorini) used by Townes and other workers (Wahl, 1991).

COLLYRIINAE

No changes since Townes (1969).

CREMASTINAE

No substantive changes since Townes (1971) but for the description of five new genera and the synonymization of two others.

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. However, even if Opinion 157 is considered 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). The recent issue of 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). Recently, Gauld (1995) made a substantial change in the Hemigastrini (= Echthrini of Townes). Gauld pointed out 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 Hemigastrini, I am not convinced that his actions represent an improvement. To the best of my knowledge, the tribe Cryptini is defined by the lost 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 Hemigastrini 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.

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, especially in the Cryptina (Porter, 1985, 1987). In the Phygadeuontini, a number of names treated as junior synonyms by Townes have been recognized as valid genera by Horstmann (1976, 1978, 1990, 1992).

CTENOPELMATINAE (= Scolobatinae of Townes)

No substantive changes have been made with the exception of the synonymization of Westwoodiini with the Scolobatini (Gauld, 1984).

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, Sweaterella, was described by Wahl & Gauld (1998).

DIACRITINAE (part of Ephialtinae of Townes)

Gauld (1991) raised this former pimpline tribe to subfamilial rank. Wahl & Gauld transferred 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 by Diller (1970, 1982, 1984).

EUCEROTINAE (part of Tryphoninae of Townes)

Barron (1976) raised this former tryphonine tribe to subfamilial rank.

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) 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 used the above dates in this listing, following the example of Townes & Townes (1973).

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).

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.

METOPIINAE

No substantive changes since Townes (1971).

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 (1970) 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.

OPHIONINAE

Gauld (1978, 1979, 1985) has greatly changed ophionine generic concepts since Townes (1971). After demonstrating the unsatisfactory nature of the two tribes used by Townes (1971), Gauld (1985) arranged the genera into five informal genus-groups. This classification is used here.

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.

PAXYLOMMATINAE

Townes did not include this group within the Ichneumonidae, treating it as a separate family (Townes & Townes, 1982). Most authors now consider it to be an ichneumonid subfamily (Rasnitsyn, 1980; Gauld, 1984; Sharkey & Wahl, 1991). Several fossil genera have been described by Kasparyan (1988).

PEDUNCULINAE

The history and composition of this subfamily are as given in the discussion of Brachycyrtinae.

PHRUDINAE

No substantive changes since Townes (1971). I do not accept Kolarov's (1987) transfer of Seleucus to the Ctenopelmatinae. The tapered cylindrical metasomal apex strongly suggests utilization of a soil-dwelling host, rather than a symphytan. The presence of a fore tibial tooth is found in five other phrudine genera and is widely scattered in ichneumonids (Tryphon, Ctenopelmatinae, Labeninae, Mesochorinae); its does not provide a compelling reason to move the genus.

PIMPLINAE (= Ephialtinae of Townes)

While little has changed at the generic level, the tribal classification has been greatly modified by Gauld (1991) and Wahl & Gauld (1997). 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.

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.

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).

TERSILOCHINAE

No substantive changes except for the description of new genera and subgenera by Horstmann (1971, 1981).

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.