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A thousand and one wasps: a 28S rDNA and morphological phylogeny of the
Ichneumonidae (Insecta: Hymenoptera) with an investigation into alignment
parameter space and elision
Donald L. J. Quicke ab; Nina M. Laurenne c; Mike G. Fitton b; Gavin R. Broad b
a Department of Biology, and Centre for Population Biology, Imperial College London, Ascot, Berkshire, UK b
Department of Entomology, Natural History Museum, London, UK c Finnish Museum of Natural History,
Zoological Museum, Finland
Online Publication Date: 01 June 2009
To cite this Article Quicke, Donald L. J., Laurenne, Nina M., Fitton, Mike G. and Broad, Gavin R.(2009)'A thousand and one wasps: a
28S rDNA and morphological phylogeny of the Ichneumonidae (Insecta: Hymenoptera) with an investigation into alignment parameter
space and elision',Journal of Natural History,43:23,1305 — 1421
To link to this Article: DOI: 10.1080/00222930902807783
URL: http://dx.doi.org/10.1080/00222930902807783
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Journal of Natural History
Vol. 43, Nos. 23–24, June 2009, 1305–1421
ISSN 0022-2933 print/ISSN 1464-5262 online
© 2009 Taylor & Francis
DOI: 10.1080/00222930902807783
http://www.informaworld.com
TNAH0022-29331464-5262Journal of Natural History, Vol. 1, No. 1, April 2008: pp. 0–0Journal of Natural History
A thousand and one wasps: a 28S rDNA and morphological phylogeny
of the Ichneumonidae (Insecta: Hymenoptera) with an investigation
into alignment parameter space and elision
Journal of Natural History
Donald L.J. Quickea,b*, Nina M. Laurennec, Mike G. Fittonb and Gavin R. Broadb
aDepartment of Biology, and Centre for Population Biology, Imperial College London, Silwood
Park Campus, Ascot, Berkshire SL5 7PY, UK; bDepartment of Entomology, Natural History
Museum, Cromwell Road, London SW7 5BD, UK; cFinnish Museum of Natural History,
Zoological Museum, Entomological Division, P.O. Box 17 (P. Rautatiekatu 13), FIN-00014
University of Helsinki, Finland
(Received 9 December 2008; final version received 30 March 2009)
The internal phylogeny of the Ichneumonidae is investigated using parsimony
analysis of a large data set including 1001 partial 28S ribosomal DNA sequences,
621 of which are newly reported, and a morphological data set of 162 characters
scored variously at subfamily, tribe, genus group and genus levels and including
only informative characters. The data set includes members of 630 named genera,
representing all currently recognized subfamilies, all but four tribes and all but
one of the taxa noted by Townes as being of uncertain placement. Sequences were
aligned using CLUSTAL X, and a sensitivity analysis was performed varying gap-
opening and gap-extension parameters. Alignments were appraised by reference
to their ability to recover a range of traditional and morphologically recognized
groups. Each alignment was analysed both independently and simultaneously
with the morphological data set, and also with gap characters treated as both
missing data and as informative. No single set of alignment parameters was found
to be markedly better by this criterion, and different ranges of parameters led to the
recovery of different recognized groups of taxa. Elision (combining all alignments
into a single analysis) was therefore used, both with and without morphology and
with both gap character treatments, to summarize the overall molecular signal.
Analysis of the morphological matrix alone produced a number of results that are
undoubtedly a consequence of convergence of morphological characters as the
result of parallel evolution of similar life histories. Simultaneous analysis of the
morphological data set with each of the 120 DNA alignments recovered most
accepted subfamilies as monophyletic. Several currently recognized subfamilies are
supported by most of the molecular analyses but some appear to be paraphyletic or
polyphyletic. The Ctenopelmatinae are paraphyletic with respect to the Metopiinae.
Robustly recovered results lead us to resurrect the Brachyscleromatinae to include
Brachyschleroma and the Erythrodolius group of Phrudinae. The Neorhacodinae and
the Phrudus group of Phrudinae are transferred to the Tersilochinae. Nonnus is trans-
ferred to the Nesomesochorinae. Hyperacmus is transferred to the Cylloceriinae.
The major groupings of subfamilies that have recently been proposed (i.e.
ichneumoniformes, pimpliformes and ophioniformes) were recovered as mono-
phyletic, but their exact limits remain in question.
Keywords: pimpliformes; ophioniformes; ichneumoniformes; life history; evolution
*Corresponding author. Email: d.quicke@imperial.ac.uk
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1306 D.L.J. Quicke et al.
Introduction
The Ichneumonidae is the largest family of Hymenoptera with more than 24,000
described species (Yu et al. 2005, together with species described subsequently), but
with an actual species richness ventured to be well in excess of the 60,000 estimated
by Townes (1969) (Gauld 1991). Virtually all species are parasitoids of immature
holometabolous insects or, less commonly, arachnids, and occasionally other insects
as hyperparasitoids, and a very few are at least partially phytophagous (Gauld and
Bolton 1988; Gauld 1991; Quicke 1997). They include both ectoparasitoids and
endoparasitoids, idiobionts; (i.e. those that prevent their host developing further after
attack) and koinobionts (i.e. those that allow the host to continue development for at
least a while); primary parasitoids and hyperparasitoids; egg–larval, larval and pupal
parasitoids, etc. Given their range of biologies, and the numerous transitions between
them, ichneumonids, along with their sister group the Braconidae, provide an excel-
lent opportunity for studying the evolution of parasitoid life histories (Gauld 1988;
Eggleton 1989; Gokhman 1992, 1995).
Townes made major steps forward in recognizing natural groups within the
family, and although his work was largely phenetic in nature, his classification (sum-
marized in Townes 1969, 1970a,b, 1971) has formed a robust framework for most
subsequent work. The Ichneumonidae is currently divided into 41 extant subfamilies
(Gauld 1991; Yu and Horstmann 1997; Porter 1998; Quicke et al. 2005; Laurenne
et al. 2006). However, the exact limits of many of these are poorly defined and some
are potentially paraphyletic or polyphyletic. Although some aberrant generic groups
have been recognized as subfamilies in their own right over the last 30 years
(for example, Cylloceriinae, Diacritinae, Poemeniinae, Rhyssinae, Tatogastrinae,
Oxytorinae, Brachycyrtinae, Pedunculinae, Claseinae, Nesomesochorinae and
Nonninae), the higher classification of the family is still in need of considerable work.
Identification of ichneumonids has always been difficult, requiring considerable
expertise and experience, and this has undoubtedly held back work on their biology
and has probably discouraged many people from taking a serious interest in the
group. Indeed they have been cited as an example of a taxon displaying extreme levels
of homoplasy (Gauld and Mound 1982) and as noted by Wahl (1986), “adult charac-
ters have been of limited use for elucidating relationships of the subfamilies.” It seems
likely that many of their morphological features may reflect multiple parallel evolution-
ary shifts in biology, as has been demonstrated for their apparent sister group the
Braconidae, and so can be very misleading about relationships (Quicke and Belshaw
1999; Belshaw and Quicke 2002). Some workers have therefore paid much attention to
the considerable variation observed in the final instar larval head capsule (e.g. Short
1957, 1959, 1970, 1978; Finlayson 1975, 1976; Wahl 1986, 1988, 1990) in the hope that
these will be informative though, of course, such features may be just as susceptible to
convergence as a result of parallel shifts in biology. By improving our phylogenetic
understanding of this huge and biologically diverse group of parasitoid wasps it will
become possible to make more detailed and extensive comparative studies of parasitoid
evolutionary biology (see e.g. Jervis et al. 2003; Laurenne et al. 2009). To achieve this it
is clear that additional, and ideally independent, data are needed, and so attention is
turning to what additional insights molecular (i.e. DNA sequence) data can provide.
The first molecular phylogenetic study including members of the Ichneumonidae
was by Derr et al. (1992a) but it subsequently transpired that some of the parasitic
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Journal of Natural History 1307
wasp sequences generated were actually vertebrate contaminants. Derr et al. (1992b)
generated four new ichneumonoid sequences and performed a re-analysis. The next,
more detailed, study was by Belshaw et al. (1998), which included 40 species repre-
senting 22 currently recognized subfamilies as part of a larger study including also the
Braconidae. In that study, the rooting of both families to one another was artificial
because the gene fragment used, the D2 region of the nuclear 28S ribosomal DNA
(rDNA) gene, is highly dissimilar between the families with a large shift in base composi-
tion having occurred in the Braconidae (excluding the putatively basal Trachypetinae)
(Quicke et al. 1999a,b). A slightly more detailed study specifically of ichneumonid
relationships was presented by Quicke et al. (2000) which included molecular,
morphological and simultaneous analyses on a data set comprising 61 genera and 31
subfamilies of Ichneumonidae. Although it represents the densest taxon-sampling to
date, this work included only a few representatives from most subfamilies and did not
include some subfamilies at all. As a consequence, apart from painting a broad pic-
ture of possible groupings, it was not sufficient to test subfamily monophyly or the
placements of numerous atypical taxa. Since then, the bulk of molecular work on the
superfamily has concentrated on the Braconidae (e.g. Wharton et al. 1992; Gimeno
et al. 1997; Quicke and Belshaw 1999; Belshaw et al. 2000; Dowton et al. 1998, 2002;
Dowton and Austin 2001; Banks and Whitfield 2006; Sharkey et al. 2006), perhaps
because the identification of Ichneumonidae has posed a greater problem. Nevertheless,
some works have used data from the same gene fragment to reassess the placements of a
few genera whose taxonomic positions had been questionable (Broad et al. 2004;
Quicke et al. 2005), Most recently, Laurenne et al. (2003, 2006) also used the 28S
D2 + D3 fragment to investigate relationships within the largest subfamily, the
Cryptinae, and between Cryptinae and some putatively related groups such as the
Alomyinae, Brachycyrtinae, Claseinae and Pedunculinae.
Morphological phylogenetic analyses of some individual subfamilies and a few
groups of subfamilies have been undertaken [Labeninae – Gauld 1983; Wahl 1993a;
Gauld and Wahl 2000a; Orthocentrinae (= most of Microleptinae+Orthocentrinae
sensu Townes 1971) – Wahl 1986; Ophioninae – Gauld 1985; pimpliformes – Wahl
1990, Wahl and Gauld 1998; Pimplinae (sensu lato) – Eggleton 1989; Pimplinae – Gauld
et al. 2002a; Campopleginae – Wahl 1991, Miah and Bhuiya 2001; Mesochorinae – Wahl
1993b; Agriotypinae – Bennett 2001; Eucerotinae – Gauld and Wahl 2002; an early
phenetic analysis of the Anomaloninae by Gauld 1976a], but these works may have
been hampered by an incomplete understanding of the structure of the family as a
whole, potentially resulting in recognition of non-monophyletic subfamilies. The
inconsistent application of family-group names has been another factor hindering the
proposal of a “natural” classification of the family. The varying composition and
nomenclature of the group of genera centred around Townes’s (1971) Microleptinae
are illustrative of the classification problems and the difficulties for the naïve in track-
ing the different usages of a name, or of knowing which name to apply to a group of
genera. The name Oxytorinae (= Microleptinae sensu Townes, as Townes did not
accept that family-level names followed the rules of priority) has been used in a
variety of senses. Wahl (1986) stated that “While there is agreement that most of the
genera form a natural group, certain genera such as Microleptes, Oxytorus [the type
genus], Hyperacmus and Tatogaster have only provisionally been considered mem-
bers of the subfamily (Townes 1971).” Later, Wahl (1990) completed the process of
dismantling the group and restricted Oxytorinae just to Oxytorus, moving Tatogaster
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1308 D.L.J. Quicke et al.
to its own subfamily, resurrecting the name Microleptinae (see Townes et al. 1961)
for Microleptes, and transferring other members to the Orthocentrinae. A conse-
quence of all this was that the Oxytorinae (as then defined) were included in Wahl’s
analysis (Wahl 1986) as part of the pimpliformes group of subfamilies (to which at
least the Orthocentrinae belong) whereas Oxytorus itself, based on DNA data, clearly
belongs in the ophioniformes (Belshaw and Quicke 2002), and indeed, as part of a
polyphyletic Ctenopelmatinae where they had traditionally been placed (e.g. Kerrich
1939), prior to Perkins (1959) and Townes (1971). Humala (2002) subsequently
expanded the concept of “Microleptinae” (incorrectly using Townes’s subfamilial
name, rather than the oldest available) to again include Orthocentrinae, Microleptes
and Cylloceriinae.
Here we provide the first complete formal cladistic analysis of the Ichneumonidae at
subfamily level, with dense and approximately proportionate taxon sampling. In addi-
tion to including representatives of all currently recognized subfamilies it also includes
nearly all those that have ever been recognized as such, all but four recognized extant
tribes [Ankylophonini (Tryphoninae), Xenothyrini (Labeninae) and Clypeodromini and
Goedartiini (Ichneumoninae); see Wahl, 1999] and all except one (Earobia) of the
19 genera whose subfamily level placement was considered dubious by Townes (1969).
Material and methods
Morphological data set
A data set comprising 162 informative morphological characters was scored as far as
possible for 88 terminal taxa. The characters used derived from traditional external
morphology with many character definitions drawn from other sources (all checked
by examination of specimens by G.R.B.; see Appendix 1 for details), details of the
internal female reproductive system and terminal tergite and sternite anatomy
(Figure 2), base of ovipositor sheaths (Figure 3), base of ovipositor (Figures 4–6),
Figure 1. Plot of log10 number of genera sequenced versus number of currently recognized
genera in each subfamily (number of recognized extant genera from Yu and Horstmann 1997;
Yu et al. 2005). The point at the origin represents 15 monotypic subfamilies.
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Journal of Natural History 1309
transverse light microscope sections through mid-region of ovipositor (Figures 7, 8),
venom apparatus (Figures 9–14), egg, and final larval instar head capsule. The char-
acter state definitions are given in Appendix 1 together with notes on previous use of
characters and exceptions, and the data matrix is given in Table 1.
Choice of taxa for morphological scoring
Subfamily units used here broadly agree with those of Gauld et al. (2002b), though
we recognize the Claseinae of Porter (1998), the Nesomesochorinae and Nonninae as
defined in Quicke et al. (2005) and the Alomyinae (see Laurenne et al. 2006). Unless
otherwise stated, the compositions of these groups are as in Townes’s generic classifi-
cations (1969, 1970a,b, 1971), where, however, they may have a different rank or
name. Our terminal taxa comprise a mixture of subfamilies, tribes (either in their
entirety or with particular exclusions; see below), genus groups and various individ-
ual genera. We have scored all tribes separately except for those of the Cryptinae and
Ichneumoninae. In both these cases the distinctions between the tribes are subtle.
Regarding the Cryptinae, the results of the molecular study by Laurenne et al. (2006)
suggest that although most genera fit into the three tribes as broadly recognized,
there is considerable uncertainty about the placements of some, particularly the more
basal taxa. In the case of the Ichneumoninae there is no general consensus: Wahl and
Mason (1995) and Sime and Wahl (2002) proposed a tribal classification which dif-
fers from the picture of tribal relationships proposed by, for example, Heinrich (1934,
1967), Townes et al. (1961) and Townes and Townes (1973). Where we have treated
particular genus level taxa separately it is because their inclusion in higher taxa is not
convincingly supported. The scoring of higher groups does not mean that we assume
Figure 2. Light photomicrograph of eighth metasomal tergite of female Pimpla rufipes (Miller)
following KOH (aq.) maceration and staining with chlorazol black showing “Y”-shaped
internal ridge.
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1310 D.L.J. Quicke et al.
that they are monophyletic, just that they are diagnosable. Hence, the taxa scored for
analyses include, with a very few exceptions, all of the groups recognized as
subfamilies by contemporary ichneumonid taxonomists.
Our guidance for singling out certain genera and genus groups for separate mor-
phological scoring comes from various sources. Townes (1969: p.31) lists 19 genera
(indicating some of these as comprising groups of genera) whose placements needed
more evidence: namely Alomya, Coleocentrus, Brachyscleroma, Collyria, Orthopelma,
Euceros, Neorhacodes, Stilbops, Melanodolius, Phrudus, Adelognathus, Lapton,
Bremiella, Agriotypus, Lycorina, Olethrodotis (as Taschenbergia), Earobia, Micro-
leptes and Procinetus. Subsequently in his treatment of the Metopiinae, Townes
(1971) noted that the systematic placements of two other genera, Apolophus
(= Scolomus) and Ischyrocemis that he included therein, were uncertain. Several of
these genera have now been placed in their own subfamilies (e.g. Adelognathinae,
Agriotypinae, Collyriinae, Eucerotinae, Lycorininae, Microleptinae and Stilbopinae)
Figure 3. Light photomicrographs of base of ovipositor sheath following maceration in KOH
(aq.) and light staining with chlorazol black. (A) Odontocolon (Xoridinae); (B) Phaenolobus
(Acaenitinae); (C) Nonnus (Nonninae); (D) Hymenoepimecis (Pimplinae); (E) Coleocentrus
(Acaenitinae).
A B C
ED
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Journal of Natural History 1311
with broad support. Only in the cases of Stilbops (and the wider Stilbopinae)
Coleocentrus and Procinetus (both included by Townes and subsequent authors in the
Acaenitinae) have subfamily placements been presented explicitly based on synapo-
morphies (Wahl 1988; Wahl and Gauld 1998), although on a reduced set of characters
for Procinetus, in the absence of larval information.
In addition to the taxa mentioned above, the placements of several other taxa
have been subject to doubt or conflicting opinion and we have therefore also treated
them separately in the morphological data set, all of which are explained below.
The genus Scolomus was originally described in the Tryphoninae but subse-
quently included in the ctenopelmatine tribe Pionini by Townes and Townes (1966),
probably largely because of the presence of a tooth at the apex of the fore-tibia and
its very narrow ovipositor. Townes (1971) subsequently described the genus
Apolophus in the Metopiinae; Broad and Shaw (2005) recorded the host association
of Scolomus borealis, considering it a genus of metopiine, under the name Apolophus.
Subsequently, Gauld and Wahl (2006) synonymized Apolophus with Scolomus and
also concluded that this genus is indeed best placed in the Metopiinae but noted that
monophyly of the Ctenopelmatinae (excluding Metopiinae) is subject to question.
Figure 4. Light photomicrographs of upper ovipositor valve bases following maceration in KOH
(aq.) and light staining with chlorazol black. (A) Labium (Labeninae); (B) Poemenia (Poemeniinae);
(C) Echthromorpha (Pimplinae: Pimplini); (D) Pimpla flavicoxis Thomson (Pimplinae: Pimplini);
(E) Limerodops (Ichnumoninae); (F) Oedemopsis (Tryphoninae: Oedemopsini).
A B
C D
E F
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1312 D.L.J. Quicke et al.
Alomya and Megalomya have variously been treated separately as the Alomyinae
(Perkins 1959, 1960) or within the ichneumonine tribe Phaeogenini (with Alomyini a
senior synonym thereof: Wahl and Mason 1995). The placement of Alomya was con-
sidered based on 28S data alone by Laurenne et al. (2006), who concluded that it was
not a derived ichneumonine [it neither appeared within the Ichneumoninae in the
most parsimonious trees (MPTs) obtained nor shared any obvious molecular synapo-
morphies with the Ichneumoninae or Phaeogenini taxa investigated].
Figure 5. Light photomicrographs of upper ovipositor valve bases following maceration in KOH
(aq.) and light staining with chlorazol black. (A) Netelia (Tryphoninae: Phytodietini); (B)
Metopius (Metopiinae); (C) Lissonota (Banchinae: Atrophini); (D) Astiphromma (Mesochorinae);
(E) Gravenhorstia picta Boie (Anomaloninae); (F) Dusona (Campopleginae).
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Journal of Natural History 1313
Townesion was described by Kasparyan (1993), who erected the Townesioninae
for it and Sachtlebenia. However, Gauld and Wahl (2000b) reappraised their position
and concluded that these genera were in fact a derived clade of Banchinae (tribe
Glyptini). Nevertheless, these are highly unusual genera and we scored Townesion
separately (no specimens of Sachtlebenia were available for sequencing).
Gauld (1984) united Scolobates with three genera of the ctenopelmatine tribe
Westwoodiini as a redefined Scolobatini but removed Megaceria to the Euryproctini. The
‘westwoodiine’ genera and the Scolobatini sensu Townes (1970b) are scored separately.
The three sequenced genera of Westwoodiini (Pergaphaga, Westwoodia and Hypopheltes)
differ from one another in a number of ways and so were scored separately.
Wahl (1988) transferred Panteles from the Banchinae to the Stilbopinae and pos-
tulated some potential but not very convincing apomorphies, though noting that “. . .
conflicting character states preclude any definite conclusions.” Quicke (2005)
described aspects of the biology of Panteles schuetzeanus and showed that the egg
Figure 6. Light photomicrographs of upper ovipositor valve bases following maceration in
KOH (aq.) and light staining with chlorazol black. (A) Phytodietus (Tryphoninae); (B)
Westwoodia (Ctenopelmatini); (C) Oxytorus (Oxytorinae); (D) Hybrizon (Hybrizontinae).
A B
C D
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1314 D.L.J. Quicke et al.
and final instar larval head capsule differ greatly from those of Stilbops. The only
other genus placed in the Stilbopinae is Notostilbops from Chile and unfortunately we
have not been able to obtain sequence data for this and its biology and larval features
are still unknown.
Rodrigama was described in the Poemeniinae by Gauld (1991), a grouping also
recovered by Wahl and Gauld (1998), but as a basal tribe within the Poemeniinae. As
the monophyly of Poemeniinae is therefore upheld by very few morphological char-
acters Rodrigama is coded separately. The genus Ganodes was recovered by Wahl and
Gauld (1998) as a basal genus of Poemeniini with rather few apomorphies defining
the tribes, therefore we coded Ganodes separately too.
Monophyly of the Phrudinae has long been debated, and in addition to treating
the Erythrodolius group separately (vide Townes 1971), we also treated the sequenced
genera of “microphrudinae” as three separate morphological terminals: Phaestacoenitus,
Pygmaeolus and Phrudus +Astrenis. We were unable to obtain sequence data for
Earobia, Notophrudus or Peucobius. Seleucus had also been treated as belonging to
the Phrudinae (e.g. Townes 1971; Gauld 1997), but was treated as a genus of
Mesoleiini (Ctenopelmatinae) by Kolarov (1987), and as constituting a tribe
(Seleucini) in its own right in the Ctenopelmatinae by Vikberg and Koponen (2000).
Figure 7. Drawings of transverse sections through mid-part of ovipositor valves. (A) Brachycyrtus,
dorsal valve only; (B) Grotea sp. (Labeninae); (C) Pseudorhyssa sternat Merrill (Poemeniinae
incertae sedis) [NB. virtually identical to Ganodes sp. (Poemeniinae), not illustrated]; (D)
Podoschistus sp. (Poemeniinae); (E) Coelichneumon sp. (Ichneumoninae); (F) Peucobius sp.
(Phrudinae); (G) Xenoschesis sp. (Ctenopelmatinae).
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Journal of Natural History 1315
However, Wahl (1999) states “I do not accept Kolarov’s (1987) transfer of Seleucus to the
Ctenopelmatinae,” so retaining it in Phrudinae. Townes (in Townes et al. 1961) erected
the Brachyscleromatinae for the genus Brachyscleroma but later included this genus in the
Phrudinae. Gupta (1994) suggested that the name Brachyscleromatinae could be revived
for the Erythrodolius-group of larger phrudines, but did not suggest any apomorphies for
such a grouping. Here we code the distinctive genus Brachyscleroma separately from the
Erythrodolius group (i.e. Erythrodolius, Icariomimus and Melanodolius).
Townes (1970b) included the genera Chriodes (including Klutiana), Nonnus,
Hellwigia and Skiapus as comprising two tribes of the Campopleginae, though these
are all highly different from the remaining campoplegines in various ways, and
Horstmann (1969) had treated Hellwigia as constituting the subfamily Hellwigiinae (as
had Constantineanu 1961). Wahl (1991), in a phylogenetic analysis of campoplegine
genus groups, effectively retained all these taxa within the Dusona group of genera
that comprised the bulk of the subfamily. Quicke et al. (2005) presented separate and
combined molecular and morphological analyses of these taxa and a selection of
Figure 8. Drawings of transverse sections through mid-part of ovipositor valves. (A) Apophua sp.
(Banchinae); (B) Euryproctus sp. (Ctenopelmatinae); (C) Trematopygus sp. (Ctenopelmatinae); (D)
Alexeter sp. (Ctenopelmatinae); (E) Lamachus eques (Hartig) (Ctenopelmatinae); (F) Hadrodactylu
s
sp. (Ctenopelmatinae); (G) Skiapus sp. (Ophioninae); (H) Enicospilus kauaiensis (Ashmead), uppe
r
valve only (Ophioninae); (I) Scolobates sp., lower valve only (Ctenopelmatinae).
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1316 D.L.J. Quicke et al.
other campoplegines and ophioniformes and concluded that Hellwigia and Skiapus
were most probably members of the Ophioninae and that Chriodes, Klutiana and
Nonnus were not assignable to any subfamilies recognized at that time and so ele-
vated them to subfamily status (Nesomesochorinae and Nonninae, respectively).
The Belesica group of Cremastinae (represented in our molecular data set by
Eurygenys) are highly aberrant within the subfamily so were scored separately, as a
test of the synapomorphic or homoplastic nature of the hind-tibial spurs character
(insertions into the tibia separated by a chitinous bridge) presently used to define the
Cremastinae.
Perithous and Delomerista (which together with Atractogaster comprise the pim-
pline tribe Delomeristini; Gauld et al. 2002a) were each scored separately because
they had previously been separated as two tribes (Wahl and Gauld 1998). However,
following Gauld et al. (2002a) we treated the Polysphincta group (koinobiont
ectoparasitoids of spiders) as Ephialtini as they are clearly just a derived lineage with
specialized biology.
Figure 9. Chitinous venom gland and reservoir intimas. (A) Ganodes sp. (Poemeniinae); (B)
Labena sp. (Labeninae); (C) Phaenolobus sp. (Acaenitinae); (D) Coleocentrus sp. (Acaenitinae);
(E) Euceros sp. (Eucerotinae); (F) Euceros frigidus Cresson.
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Journal of Natural History 1317
An area of particular classificatory confusion concerns the use of the names
Oxytorinae, Microleptinae, Helictinae and Orthocentrinae, and these are discussed
briefly below. The Helictes group was scored separately from the Orthocentrus group
of Orthocentrinae. As employed here, the Helictes group corresponds to the Micro-
leptinae sensu Townes (1971), minus Microleptes, Oxytorus, Tatogaster, Cylloceria,
Allomacrus, Hyperacmus and Acaenitellus, and to the Helictinae of Gauld (1991)
minus Hyperacmus. In addition, we scored Proclitus separately because its venom
apparatus differed greatly from that of other Orthocentrinae. The affinity of
Hyperacmus has been debated. It was placed with Microleptes in the Microleptinae
by Dasch (1992) but was transferred to the Orthocentrinae by Wahl and Gauld
(1998). Latterly, it was treated again as being close to Microleptes by Humala (2003)
but later Humala (2007) agreed with Broad (2004, using character systems being
published in this paper) that its affinities lie with the Orthocentrinae/Cylloceriinae.
The molecular data set
Apart from the Ichneumoninae, the number of taxa sequenced in each subfamily is
approximately proportional to the number of described species (Figure 1), though we
have endeavoured to maximize the taxonomic diversity of the sequenced taxa where
possible. The D2 + D3 region was sequenced for 92% of taxa, but some rare taxa
(<2% of total) that were sequenced from dry specimens have only partial D2
Figure 10. Chitinous venom gland and reservoir intimas. (A) Hyperacmus sp. (Orthocentrinae
s.l.); (B) Cylloceria sp. (Cylloceriinae); (C) Eusterinx sp., (Orthocentrinae); (D) Aperileptus sp.
(Orthocentrinae).
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1318 D.L.J. Quicke et al.
sequences. Taxa sequenced, their provenances, depositories and EMBL/GenBank
accession numbers are given in Appendix 2.
Molecular methods
The included sequence data have been assembled over a period of approximately
10 years and a variety of protocols have necessarily been employed. For most recent
protocols see Laurenne et al. (2006). Data come primarily from either of two primer
pairs: D2-only or D2–D3. Additional internal primers (Quicke et al. 2007; Table 2)
were used to sequence the D2 region in three fragments of c. 160–230 base pairs for
some species which were only available from old or poorly preserved specimens
whose DNA had partly degraded and failed to produce larger amplicons.
Sequenced taxa
Six hundred and fifteen of the sequences used have been generated for this paper, a
further 366 derive from works published by our laboratories (Belshaw et al. 1998;
Figure 11. Chitinous venom gland and reservoir intimas. (A) Schizopyga sp. (Pimplinae); (B)
Perithous sp. (Pimplinae); (C) Delomerista sp. (Pimplinae); (D) Pseudorhyssa alpestris (Poeme-
niinae); (E) Rhyssa persuasoria (L.) (Rhyssinae); (F) Exeristes sp. (Pimplinae); (G) Clasis sp.
(Claseinae).
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Journal of Natural History 1319
Belshaw and Quicke 2002; Broad et al. 2004; Laurenne et al. 2003, 2006). A small
number of sequences have been taken from works by other groups and have been
deposited on GenBank, some of which have been included in published molecular
studies (Chen et al. 2004; Banks and Whitfield 2006) and others not as far as we are
aware (Lee and Choi 2004). The sequenced taxa used in our analyses are listed alpha-
betically, by subfamily, tribe where applicable, genus and species in Appendix 2.
Sequence alignment
We have tried with virtually no success to analyse our sequence data using optimiza-
tion alignment as implemented in the program POY (Gladstein and Wheeler 2001) on
a number of supercomputers. Despite intensive effort, the implementations were
unable to cope with the size of the matrix and certainly it would not have been feasi-
ble to carry out any degree of sensitivity analysis with the data. We therefore used
CLUSTAL W (Thompson et al. 1994) to create multiple alignments, each of which were
then analysed using the program TNT (Goloboff et al. 2003) to obtain one or more
maximum parsimony trees.
Figure 12. Chitinous venom gland and reservoir intimas. (A) Ateleute sp.; (B) Ichneumon sp.;
(C) Alomya sp. (Alomyinae); (D) Osprynchotus sp. (Cryptinae); (E) Colpognathus sp.
(Ichneumoninae); (F) Hoplismenus sp. (Ichneumoninae); (G) Eusterinx sp. (Orthocentrinae).
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1320 D.L.J. Quicke et al.
All alignments were carried out with transition cost = transversion cost = 1.
Ninety-six gap-opening (GO) and gap-extension (GE) cost combinations were
explored. GO costs were increased incrementally from 1 to 12 and for each incre-
ment, GE costs were increased in increments from 1 to the GO cost. These values
cover those that have yielded taxonomically congruent results in previous analyses
(e.g. Sharkey et al. 2006). PAUP* (Swofford 2004) was used to calculate indices and
confirm tree lengths.
Partial sequences
The great majority of the species included in the molecular study were represented by
essentially the whole of the D2 region (c. 475 base pairs) or of the D2 + D3 region
(c. 670 base pairs). A few taxa of particular taxonomic interest were only available as
old (5–25 years old) dry specimens, but with the use of internal primers (Table 2), we
were in most cases able to sequence most of the D2 region as a set of three short frag-
ments, though sometimes without complete contiguity. Satisfactory alignment of
these incomplete fragments was achieved by entering the minimum number of
unknown bases (“N”s) that occur in the unreadable regions in other sequences, which
Figure 13. Chitinous venom gland and reservoir intimas. (A) Orthopelma mediator (Thunberg)
(Orthopelmatinae); (B) Westwoodia sp. (Ctenopelmatinae); (C) Xenoschesis sp. (Ctenopelmati-
nae); (D) Exenterus sp.; (E) Phytodietus (Neuchorus) sp. (Tryphoninae); (F) Phytodietus sp.
(Tryphoninae); (G) Lissonota sp. (Banchinae); (H) Pion sp. (Ctenopelmatinae).
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Journal of Natural History 1321
avoided forced additional gaps and, more importantly, generally produced CLUSTAL
outputs in which the ends of these fragments were appropriately aligned. Failure to
enter a sufficient number of “N”s in the unsequenced regions resulted in very conspic-
uous misalignments. Similarly, for taxa for which we had only the D2 region, a
number of “N”s corresponding to the minimum number found in the missing D3
region was added to the 3' end of the available sequence.
Phylogenetic analysis
All characters were treated as unordered (non-additive). Because of the large size of the
data set we used TNT to search for MPTs (Goloboff et al. 2003), with the limitation that
all polymorphisms involving multistate characters where a taxon may possess more
than one state were treated as either “N” when DNA or “?” when morphological.
Trees were treated as unrooted because of problems with alignment with braco-
nid sequences (see Results). Therefore, trees are drawn with a xoridine at the base
because xoridines are reasonable candidates for being close to the base of the tree,
based on both biology and morphology. For simplicity we retain the term
Figure 14. Chitinous venom gland and reservoir intimas: (A) Banchus sp. (Banchinae); (B) Eryth-
rodolius griffithsorum Gauld (Phrudinae); (C) Hellwigia elegans Gravenhorst (Ophioninae); (D)
Ophion sp. (Ophioninae); (E) Stilbops vetula (Gravenhorst) (Stilbopinae); (F) Tersilochus sp.
(Tersilochinae); (G) Mesochorus sp. (Mesochorinae); (H) unmacerated venom gland and reser-
voir of Hybrizon sp. (Hybrizontinae).
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1322 D.L.J. Quicke et al.
Table 1. Morphological data set.
10 20 30 40 50 60 70 80
Hybrizontinae 0000110211 0110020000 1000000001 002?000002 0010121??0 12?20002?1 0001000000 01001002A1
Tersilochinae 0000000011 0100000001 000001?000 0000A00110 0000111?01 A1020011?A 0001000000 0100001211
Ophioninae 00000A0000 0100000000 000001?000 00C000B010 A0100B1?11 C01A0000C1 000AA00000 0100001211
Tatogastrinae 0000000000 0100000010 0000000000 0000100020 00000100?1 200A100001 0000010000 0100001201
Skiapus 0000100211 0100000000 1000000011 0000002010 0010001?11 201A000021 1011111000 00?0001211
Hellwigia 0000000000 1100000000 000001?011 0000002020 0010121?11 22?A000020 1011101000 00?0001211
Pergaphaga 0002000000 0200000000 000001?100 0000000010 0010100001 00?0000020 0000010000 00?0001201
Westwoodia 0002000000 0200000000 000001?100 0000000010 00001200?1 0100000010 0000010000 00?0000000
Hypopheltes 0?02000000 0200000000 010001?100 0000100010 00101D0001 0000000010 0000010000 00?0000100
Campopleginae 0000000000 01100000A0 A000000011 000000D010 A0A0ACA001 C10100A0A1 00000A0000 0100001DA1
Nonninae 0000000011 0100000000 1100000000 0000002020 1010AB01?1 2A00000001 0000001100 0110100201
Nesomesochorinae 0000000001 0100000000 1000000010 0000012020 101000AA01 DA0A001001 0001000100 0110100211
Gravenhorstiini 0000000000 01100000A0 1100000011 001?00A021 A011121?A1 CB0010A0C1 0000010000 00?0A01211
Anomalonini 0000000001 01A0000010 1100000011 0000002021 00110B1?11 20111000?1 0001111010 0100001211
Belesica group 00000000CA 0100000010 0A00A1?000 0000002020 00100000?1 AA00000011 0000010001 0100001100
other Cremastinae 0000000000 0100000000 0000000011 0000002020 A0A000A001 C10100A0C1 0000000AA1 0100001DA1
Scolobatini s.s. 0001000000 0200000010 010001?001 0000000000 0012121?01 2101000001 0000010000 00?0000000
Scolomus 00000100AA 02A0000A00 00000A0000 000110A010 00000A000A AA0A001000 00000A0000 0100000100
Ischyrocnemis 0000H00000 0110000010 0000000001 0000000020 00101000?1 0A0A0010A1 0000010000 0100000000
Metopiinae 0AA0000000 1110000000 0000000001 10A0AA0120 0010A0A00A 0C0000A0CA 00111100A0 00?0000000
Mesochorinae 0000000000 0A000000A0 0000000000 00001A0020 00100000?A D00A10A001 0000010000 010000A10A
Lapton 0?00100000 1110000200 1000000001 0000000010 0010021?01 000A000021 0000010000 00?0000200
Bremiella 0000000000 0110000000 000001?001 0000000110 00100100?1 0100001001 0000010000 00?0000100
Atrophini 0010000000 01AA001000 0000010101 00AA000020 00A01BA00A C0001000CA 0000000000 0100A00000
Glyptini 0010000000 0100000000 0100000101 00A0000020 10A000A001 A101000001 0000000000 0100000000
Exetastes group 0010000000 0100000100 1000000101 0001000020 10101200?A 0100000020 0000000000 0100000B00
Banchus group 00A0B00000 0100001100 0000010101 00B0000020 10101200?1 000000002A A000000000 00?0A00000
Townesion 0010010000 000?001000 0000010101 000A000022 00121200?0 00?0?00001 0000000000 00?0000000
Tryphonini 0000000000 010000A001 0000000001 00B0001020 00A000A000 0A0000AACA 0000000000 0A00000000
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Journal of Natural History 1323
Exenterini 0000000000 0100000001 0000000001 000AA01020 00A00000?0 A000001001 0000000010 1100000000
Oedemopsini 0000000000 01000000A1 A100000000 0000A11020 000000A000 A101001000 0001000000 0100000000
Eclytini 0000000000 0100000000 0000000000 0001100021 000000A000 1101000000 0000000000 00?0000000
Sphinctini 0000000000 0000000010 100001?101 000A000020 00100000?0 100A000011 0000000000 1100000001
Idiogrammatini 0000000000 0100000201 0100000000 0000000020 00000000?0 0101001100 0001010000 0100000000
Phytodietini 0000000000 0100000000 000001?000 00HAA00010 00A0000000 D1AA0000B1 0000001000 00?0000000
Neorhacodinae 0000000111 0100000B01 000001?001 0000000020 00001A1??? 01?2001100 0001000000 00?0000000
Olethrodotini 0000000000 0100000000 1000000001 0000000010 001012A001 0A00000000 0000010000 0100000100
Ctenopelmatini 0000000000 0100002C00 1000000000 0000A00010 00101BA001 00000000CA 0000010000 00?0001101
Mesoleiini 0000000000 010000BC00 0000000001 0000000010 00A011A000 CA0100A0C0 0000010000 0100000000
Pionini 0000000000 01A000B000 000001?001 00A0100120 001011A001 CA?A00A0A0 0000010000 00?0000211
Perilissini 0000B00000 0100000000 0000000001 0000A01020 0000AC0A01 C10100A0B0 0000010000 0100000100
Euryproctini 0000B00000 0100000000 0000000000 00BA000020 00A01BA001 CA0A00A0C0 0000010000 0100000200
Oxytorinae 0000000000 0100001000 0000000001 0000000020 000001A001 0101001000 0000000000 0100000201
Seleucini 000?000000 0001010001 0000100000 0000001000 0000000A?1 00?1000000 0000010000 0100001200
Erythrodolius group 00000000AA 0A00000010 000011?000 0000A00020 0010A11?00 0D00000000 0000010000 00?0000100
Brachyscleroma 0000000000 0000000000 0000100000 000000A120 00100100?0 0101000001 0000000000 00?0001201
Phaestacoenitus 0?00100201 0?00000001 000011?000 0000000010 01001201?1 01010011?0 0021010000 0100000000
Pygmaeolus 0000000011 0010000001 000011?000 0000000010 0002?21?01 11010011?0 0011110000 00?0000000
Phrudus & Astrenis 0000000000 0000000001 0000100000 0000000000 0000A1A101 11010011?A 0001000000 0100000201
Lycorininae 0000000000 0000000000 000000010A 0000100020 00001D1?01 A10A000001 0001100000 00?0000000
Panteles 0?00010000 1100000000 0000010000 0000000020 00000101?1 0A01000001 0000000000 0100000000
Stilbops 0000010000 0100000000 100000000A 0000000020 100000AA00 01?100000A 0000000000 0100000000
Orthopelmatinae 0000000000 1000000200 1000000000 0000001120 0000011?00 A1010010?0 0000100000 00?0000201
Coleocentrus 0000000000 010A0010A0 0010000000 00D?100020 201010A000 0110100020 0000101000 00?0000000
Procinetus 0000000000 1000002000 0010000000 0000100010 00A0A200?0 0110000020 000001000A 00?0000000
other Acaenitinae 0000000000 A00A0010A0 0010000000 00A0A0A02A 2010A010A0 AD001000C0 1011110000 00?0000200
Microleptinae 0100A00B00 00000000A0 010001?000 0000001020 0000AA1001 A1?1001000 0011110000 0101000201
Orthocentrus group 0000A10D00 02A0010000 0010000001 00N0100010 200010A001 C101001A00 0001100000 0100000000
Proclitus 0B00010000 0000010200 0000000000 0000100010 2000101?00 2101001000 0001100000 0100100200
Helictes group 0C00010A00 0200010000 A010000000 0000100010 200010A000 C101001000 0001100000 010000A20A
(Continued)
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1324 D.L.J. Quicke et al.
Table 1. (Continued)
10 20 30 40 50 60 70 80
Diplazontinae 0A00200000 AA00000A00 001001?00A 0000000020 000010A001 C00A00A0C0 0001100000 00?0000000
Hyperacmus 0D00000000 0A01000000 0000000000 1030100020 2010021?00 0A0A0000A0 0000100000 00?0000000
Cylloceriinae 0C00000A00 010A000000 001000A000 0000100020 2000101000 010A00A0C0 0000000000 0100000000
Diacritinae 0000000000 0100002000 0010000000 0000001010 20001000?0 1101001000 0000000000 0100000001
Collyriinae 0000000000 0000000010 0000000000 1000100000 2010121?00 0A10000020 0100000000 00?0100200
Poemeniinae 0000A00B00 0100A00000 10A10A1000 102?100000 00A010A00A C10A100020 A000101000 00?0100200
Ganodes 0000100200 0100100000 1001001000 102?100000 00001000?0 0100101020 0000101000 00?0100210
Pseudorhyssini 0000000000 0100100010 1011001000 1101000020 00001000?0 0A00000020 0000101000 00?0000000
Rhyssinae 0000H00000 0100100010 0000000000 1100000110 001011A000 A110100020 0020100000 00?0100000
Rodrigama 0000000000 0100100000 1011001000 101?100012 00101000?0 0010000020 0000101000 00?0000201
Ephialtini 0000000000 0A01000100 A010000000 A0CA100000 00A01000?0 C10AA0A0C0 0100101000 00?0000000
Delomerista 0000000000 0100000100 0010000000 1000100010 00101000?0 0110000020 0000100000 00?0000000
Perithous 0000000000 0101000100 0010000000 1001100010 00101000?0 1111000000 0000100000 00?0000000
Pimplini 1000000000 0101000000 00A0000000 0000000020 00101000?0 0AA0000020 0A20100A00 00?0000000
Adelognathinae 0100000000 1100000B00 1100000000 000010A010 00001DA101 1101001000 0001100000 01000012AA
Alomyinae 0010000000 0100000000 000001?000 000010A020 00000001?0 0000100000 0000001000 0100000201
Ichneumoninae 0110A00000 01A00000A0 0A00000000 0000001020 00100001?0 0A00000000 0000001000 0100001201
Ateleute 0010000000 01000000A0 000001?000 00E00A2010 100010A101 1101001010 0010000000 0100000200
other Cryptinae A1A0000000 0A000000A0 AA00000000 000001D010 00A000A10A C1?100A0C0 0000000A00 010A0012A1
Pedunculinae 0010000000 0100000000 0000000000 100AA0A110 000000AA00 1A01001010 0000000000 010100A211
Claseinae 1000000000 1101002000 0000000000 0001000011 00101000?0 0A0A000AB0 00000A1100 0100001211
Eucerotinae 0000000000 01A0000AA0 01000A0000 0001000021 00101C1?01 000A100001 0001100000 0100A00000
Brachycyrtinae 0000B00000 0000000000 0000000000 1001001021 0010001100 AA110000C0 0000000000 0100001211
Agriotypinae 0000000000 0100000000 0000000000 0000000120 2000001001 01?1000000 0000000000 00?0000201
Labenini A000001000 0000000000 0000010000 0A00100022 01100000?A 0A10110020 00AA100A00 00?0010200
Groteini 0000000000 1100000200 A000010011 0000A00011 0A100000?A CA11A100C0 0010010000 00?0000211
Poecilocryptini 0000001001 0100000000 1000000100 0000010021 00100001?1 1A11010020 0010110000 00?0001201
Xoridinae 0000A01000 1000001000 0000000010 1000000010 A010001000 0A000000A0 0001101100 00?0000200
Polymorphisms are coded as follows: A = (01), B = (02), C = (012), D = (12), E = (23), F = (13), G = (0134), H = (03), I = (24), J = (013), K = (0135),
L = (04), M = (14), N = (023), O = (05) and P=(15).
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Journal of Natural History 1325
90 100 110 120 130 140 150 160
Hybrizontinae 1011000000 1000000??? 0000012011 042101???? 10000??011 010100??00 ?????????? ?????????? ??
Tersilochinae 000100000A 122000?111 0000000001 02D110030? 0000210100 01?11?BH0A 1101050?00 00010?0120 00
Ophioninae 00000A000A 12200001?0 0000012011 ?2210A1??? 00000??040 02?0012000 1101050100 0001110120 10
Tatogastrinae 1011?00001 1221?0?0?1 03000?200? ?21??????? ?0???????? ?????????? ?????????? ?????????? ??
Skiapus 0011?00000 1110000100 0000010011 142011??0? 0000210010 010011E200 ?????????? ?????????? ??
Hellwigia 0000000001 1110?00??0 00000?2011 122000010? ?0???????0 02?1002000 ?????????? ?????????? ??
Pergaphaga 0011?00000 1110???11? 000A0?2??? ?22??????? ?????????? ?????????? ?????????? ?????????? ??
Westwoodia 0101100000 0110?0?011 00000020?? ?22??????? ?0???????? ?????????? ?????????? ?????????? ??
Hypopheltes 0001100000 12?0?????? 0??10?20?? ?22??????? ?0???????? ?????????? ????050000 00010?0121 00
Campopleginae 0001000000 1110001111 00000A00?1 02C1100100 00102101D0 0D?0113000 1A01050100 0001110120 00
Nonninae 1001010000 1100000111 0000010101 0211100200 000020112? ???00020?? ?????????? ?????????? ??
Nesomesochorinae 100A000000 111000???? 0000010001 02101002?0 ?0?0A????? 0????????? ?????????? ?????????? ??
Gravenhorstiini A011?00000 12200?0100 0000010011 02211?0100 0000200??0 02?1003J0A ?1010P0001 00?10?0120 00
Anomalonini 1011?00000 1110000100 0000010001 0211100100 0010200121 021011?000 ?1??000001 00?10?0120 00
Belesica group 0A00000000 0110000??0 00000?0011 022??????? ?0???????? ?????????? ?????????? ?????????? ??
other Cremastinae A0A1000000 111000?100 0000000101 02A1000200 0A?02101D0 0210012000 ?101050100 0001A10120 00
Scolobatini s.s. 0101100000 0000000010 010101201A 1221000100 00001??12? 0210011000 ?101050000 00010?0121 00
Scolomus 0011?00000 0000?0??0? 0?01A????? ?32??????? ?????????? ?????????? ?????????? ?????????? ??
Ischyrocnemis 00AA000000 0110?????? 01000?1??? ?220?????? ?????????? ?????????? ?????????? ?????????? ??
Metopiinae A000000001 0000100111 000000201A ?22101???? 00001???I0 02?0013000 A101050A01 ?0?10?012A 0A
Mesochorinae 000A000000 1AA0000111 0101002A0A ?32D001??? 00?01??0?? 0D11000000 11010501?? 00010?012A 00
Lapton 0001000000 1110?0???? 0?00002??? ??2??????? ?????????? ?????????? ?????????? ?????????? ??
Bremiella 0011?00000 0110????00 00000?2??? ??2??????? ?0???????? ?????????? ?????????? ?????????? ??
Atrophini 0000000000 1000?0?111 0001110000 02A2100300 0000201120 0210012000 ?1010511?0 0001A0012A 00
Glyptini 0000000100 0000000111 00011100?0 02A2100301 001020112A 12?0012000 ?101051100 00010?0120 10
Exetastes group 0000000000 1100000111 0001111000 02D2100300 00001??02? 0210112000 ?101020000 000110012A 10
Banchus group 0000000000 1110000111 0001102000 122101???? 0000200120 ?1?0?12200 ?101050100 0001AA012A 00
(Continued)
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1326 D.L.J. Quicke et al.
Table 1. (Continued)
90 100 110 120 130 140 150 160
Townesion 0001000100 0000?????? 00000????? ?32??????? ?0???????? ?????????? ?????????? ?????????? ??
Tryphonini 0000000000 010000?111 000000100A A42111???? 10001??020 0D01013001 000A0O0000 00001001A0 00
Exenterini 0000000000 000000011? 000001200B 142B0????? ?0??????20 1?01002001 000A050000 00001A01AA 00
Oedemopsini 0000000000 000000001A 1000010000 04D11?0100 ?0??0??02? 1A0100??01 ?000000000 00000?0110 00
Eclytini 0000000000 0000?0?001 00000120?? ?42??????? ?0???????? ?????????1 ?000000000 00000?010? 00
Sphinctini 0000000000 0000?0?111 00000002?? ?420?????? ?0??????1? ?10??????1 ?000000000 0000100111 00
Idiogrammatini 0001000000 111000?1?1 0000010200 0102100201 0010211120 0?0011A001 ?200050100 00000?0110 00
Phytodietini 0001000000 010000?011 0000010100 11DD100300 0000201120 0200113101 ?00A000A00 0000A001C0 00
Neorhacodinae 00010000A0 0000100??? 0000000001 0212?00201 00001??0?? ?111001??0 ?000050100 00010?0101 00
Olethrodotini 0011?00000 0110?0?100 0?000100?0 02A0?????? ?0???????? ?????????? ?????????? ?????????? ??
Ctenopelmatini 00A1000000 0100?0?1?1 00000020?? ?220?00100 00001????0 02?0??1000 ?101050100 0001100121 00
Mesoleiini 000100000A 010000?011 0000002011 1220100100 00001???20 0D00012000 1101050A00 00?1AA0121 00
Pionini 0011?00000 000000?1?1 0000010?00 1E2A0A1??? 00001??AD0 1D10100000 1101050100 0001A00121 00
Perilissini 000110000A 0100000101 000000201A 1D2???020? 00001???40 0200012B00 1101050A00 000AAA012A 00
Euryproctini 0011?00001 01000??1?1 0000002??A ?I2101???? 00001???I0 ?D?00100?0 1101050A00 000AAA01DA 00
Oxytorinae 0001000001 0100000101 0000002010 1221100001 00100??020 ?201001000 ?????????? ?????????? ??
Seleucini 0011?00001 0210?0???1 00000?20?? ?22??????? ?0???????? ?????????? ?????????? ?????????? ??
Erythrodolius group 0000000000 0220?0??11 0100010A?0 140??00300 000120112? ???001300? ?????????? ?????????? ??
Brachyscleroma 0011?00001 1220?0??1? 1000000001 03A20003?? ?0??2???00 1??0?0330? ?????????? ?????????? ??
Phaestacoenitus 0001000000 0220?????? 00000?0??? ?02??????? ?????????? ?????????? ?????????? ?????????? ??
Pygmaeolus 0001000000 0220?0???? 00000?0??? ?41??????? ?????????? ?????????? ?????????? ?????????? ??
Phrudus & Astrenis 0001000000 0221?????? 00000?0??? ?42??????? ?????????? ?????????? ?????????? ?????????? ??
Lycorininae 0000000200 0000000?10 0001010000 0112000300 1010200111 0111100001 ?201000100 00010?013? 00
Panteles 0001000000 0000?0?0?0 0?01110000 0002?00300 000120112? 1???????02 11A100??00 000?10011? 00
Stilbops 000A000000 0000000011 0001A10000 0ED2000300 0000210011 000001??00 ?100000?00 00000?0110 00
Orthopelmatinae 0001000000 0010000010 0000002000 1310101??? 00000??002 1111012000 ?101040001 ?1?10?002? 00
Coleocentrus 00001A0100 010001?000 1102000200 04A??00000 10010??042 ?001210000 ?101050100 10100?0021 00
Procinetus 0001100000 0210????0? 01011?0??? ?40??????? ?????????? ?????????? ?????????? ?????????? ??
other Acaenitinae 0000100100 1110100010 0102000200 04A0000000 1A010??0E? 1A?1201000 ??01?50A00 10110?0020 00
Microleptinae 0000000000 0110000000 00000?2010 122101???? 00001001?2 011?????00 ??01050?00 00010?01?? 00
Orthocentrus group 000000000A 01101?0010 00100?B01A ?4D101???? 00000??0B0 1D01001000 ?11?150110 20110?0010 00
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Journal of Natural History 1327
Proclitus 0000000000 0100????1? 010000???? ?21??????? ?0??????10 101??0??00 ?????????? ?????????? ??
Helictes group 0000A00000 00A010?01? 010000020A AID001???? 000?A??010 101110??00 ?101150110 20110?00C0 00
Diplazontinae A001A00000 A1A0100?10 0200002010 022A11???? 00000??02? 1A1?????00 ?11?1501?1 21?10000D2 00
Hyperacmus 0000100000 0110000??? 01000?2000 022101??1? ?0000????? ???010002? ?????????? ?????????? ??
Cylloceriinae 000A100A00 01A000?1A1 0A00000200 02A1010010 00001??0D1 ?201103320 ?101150?00 1010??0021 00
Diacritinae 0000A10000 0000100010 0100000200 04A1000000 11010??0?2 0001102000 ?????????? ?????????? ??
Collyriinae 0010?00000 0110000010 1000001000 0310100000 00000??022 10???????0 1101150111 ?1?10?1012 0?
Poemeniinae 0000110000 000000A000 1100000200 04A0000000 10110??0I2 100100??00 ?000010A00 10100?0010 00
Ganodes 0000110000 1000?00110 1100000200 0400000000 10010??042 1???10101? ?????????? ?????????? ??
Pseudorhyssini 0000110100 0000001011 2100000000 0000000001 10110??022 10???01000 00000A0000 00100?0010 00
Rhyssinae 100011000A 000001011? 2100000000 0000000000 10010??042 ?A01111000 0000010100 00100?0020 0A
Rodrigama 0000110200 0000?11100 11000002?? ?400?????? ?1???????? ?????????? ?????????? ?????????? ??
Ephialtini A000111110 0000101A1? 0100000A00 0MA000001? 1A110??0I2 AA??A?H000 ?00A0A0000 00100?00C? 00
Delomerista 0001110110 000000101? 0110000100 011100010? ?0?10???22 10?1101000 ?000010100 00100?0000 00
Perithous 0001100110 000000101? 0110000100 00010000?? ?1?10???22 1??0001000 ?000010100 00100?0010 00
Pimplini 0000111110 00001011A1 01A0000000 00D000001? 10110??022 1A?100FA00 ?A0A0A0AA0 001A0?00C0 0A
Adelognathinae 0001010000 0220000011 0000000000 042001???? 10100??022 ?20??022?0 ?100000100 0001110110 00
Alomyinae 1011000000 0000000011 00000?2000 002001???? 10100??0D? 0??1100000 ?101050001 00?10?012D 0A
Ichneumoninae 0001010000 0000000111 00000020?A 0LD0000000 ?0??0??020 ?0A101J?00 ?101050001 01?10??121 0A
Ateleute 0000000000 0120001??? 0000000000 0420000011 10100??0?? ??01000000 ?????????? ?????????? ?0
other Cryptinae 000A01000A 01100001?1 0000000000 ?GC0000011 1?110??022 00A1A0H000 0A0A0K0A00 00000?01CA 00
Pedunculinae 0001010000 0A1000?000 0000000000 01110000?? 10010????? ???100200? ?????????? ?????????? ??
Claseinae 00A1000000 1000000010 0000010000 011100000? ?0??0???2? 0??100E00? ?????????? ?????????? ??
Eucerotinae 0000000000 0000000111 0000002012 ?421?1???? 00000??03? 10110133?2 0000050000 0001100110 00
Brachycyrtinae 0001000000 0001000000 0000000000 0411000100 ?0??????20 0?0???20?0 ?1??000?00 00000?0110 00
Agriotypinae 0011?00000 02200?0011 00000?2010 0420000011 10110??0?1 0??1?00000 ?100000000 00000?00?? 00
Labenini 0000000000 000111?0A0 1000010100 00A0000000 1A010??0I2 100110N000 ?0000A0A00 00000?0010 00
Groteini 0000000000 1001?01011 100001B10A 04C0000011 10110??022 1A0?????00 ?101000000 00000?0010 00
Poecilocryptini 0000000000 000100?000 0000010200 00100000?0 10010??0?? ???????0?? ?000050000 00000?0020 00
Xoridinae 000000000A 0000000110 10000002?1 00A0000000 10010??022 000100H200 ?000030100 00000?0110 00
Polymorphisms are coded as follows: A = (01), B = (02), C = (012), D = (12), E = (23), F = (13), G = (0134), H = (03), I = (24), J = (013), K = (0135),
L = (04), M = (14), N = (023), O = (05) and P=(15).
Downloaded By: [Imperial College] At: 12:43 29 May 2009
1328 D.L.J. Quicke et al.
“monophyly” when referring to groups, although strictly we are referring to coherent
groupings on our unrooted trees.
Elision of molecular alignments
Given that no single set of alignment parameters appeared to be justifiably preferable
to any others (see Results and Figure 16) in terms of external criteria, such as recovery
of generally accepted groups, we chose to apply the elision strategy (Wheeler et al.
1995), in which all alignments are combined to form a single data set. Groups recovered
when the elided matrix is analysed should be those that are robust to varying the align-
ment parameters (Hedin and Maddison 2001). With 96 alignments (for 1001 terminal
taxa), analysing the elided matrix was computationally challenging, and the implemen-
tation of TNT was liable to crash when default parameters were markedly increased.
Therefore we carried out multiple searches (at least five) using default search parame-
ters. Most of these independent searches yielded either trees of the same length or dif-
fering by only a few steps. Visual examination of the trees obtained suggest that they
were all finding very similar relationships for a given data set, although it is quite pos-
sible that the trees may not be the shortest. We carried out analyses of the elision data
set with gaps treated both as uninformative (missing) and informative.
Combining morphological and elided molecular data
As we prefer a total evidence approach and viewed the elision results as the ones least
influenced by arbitrary parameter selection, it was decided to carry out combined
Table 2. 28S ribosomal DNA primers used in the study, including two sets of internal primer
pairs used to amplify the D2 variable region in three shorter fragments each.
Primer Forward primer (5–3) Reverse primer (5–3)
Approximate
amplicon size
(base pairs)
D2-only AGA GAG AGT TCA AGA
GTA CGT G
TTG GTC CGT GTT TCA
AGA CGG G
490
D2-D3A GCG AAC AAG TAC CGT
GAG GG
TAG TTC ACC ATC TTT
CGG GTC
710
D2-D3B AGA GAG AGA GTT CAA
GAG TAC GTG
CGC TAC GGA CCT
CCA TCA
660
RJF_int1 AGA GAG AGA GTT CAA
GAG TAC GTG
CGG GTC GCG ACG TCC
TAC TA
200
RJF_int2 TAG TAG GAC GTC
GCG ACC CG
ATA CCG TGC GRR TAC
CGC C
140
RJF_int3 GGC GGT AYY CGC
ACG GTA T
CCG TGT TTC AAG
ACG GGT
180
JM_int1 AGA GAG AGA GTT CAA
GAG TAC GTG
GGT CAG ACA ACC
GRGGGTCTG
270
JM_int2 CAG ACC CYC GGT TGT
CTG ACC
CCA ACA GCY GRC
CAG GCC CG
150
JM_int3 CGG GCC TGG YCR GCT
GTT GG
GGT CCG TGT TTC
AAG ACG GGT CCT G
100
Downloaded By: [Imperial College] At: 12:43 29 May 2009
Journal of Natural History 1329
morphological and molecular analyses with the elided molecular data sets. However,
the vastly greater size of the elided data set would mean that if each molecular
character was assigned the same weight as each morphological one then the resulting
MPTs would undoubtedly be influenced almost entirely by the molecular data. There-
fore, to assign the elided molecular data partition and the morphological partition the
same effective weight, the latter was up-weighted based both on the numbers of inform-
ative characters in the two partitions and their phylogenetic signal as reflected by their
retention index on the MPTs from separate analyses. In each case there were 162
morphological characters (all informative) with an retention index of 0.507. For
analyses in which gaps were treated as uninformative, there were 53,247 informative
bases and their retention index on the MPT from their analysis alone was 0.685. There-
fore, for combined analysis of morphology and elided molecular data, the former were
up-weighted by (0.685/0.507)*(53,247/162) = 444.24. The corresponding calculation for
gaps treated as informative gave an up-weighting factor of 483.14.
Outgroups and rooting problems
The Braconidae is uncontroversially considered to be the immediate extant sister
group of Ichneumonidae (Sharkey and Wahl 1992; Quicke et al. 1999a). The aculeates
(Chrysidoidea, Apoidea and Vespoidea) are considered the sister group of
Ichneumonidae + Braconidae, though support is weak and the only convincing char-
acter is the presence of valvilli in the ovipositor/sting apparatus (Mason 1983; Quicke
et al. 1992). Within the Braconidae, there is strong evidence from morphology and
DNA that the endemic Australian Trachypetinae is the most basal taxon, and might
possibly constitute a sister group to the ichneumonids (Quicke et al. 1999a,b; Gillespie
et al. 2005). Having said that, the issue of rooting the ichneumonid tree is not trivial
because of conflict between the perceived wisdom, based on paradigms such as the
ancestral taxa being idiobiont ectoparasitoids of concealed hosts (Gauld 1988), and the
results of 28S rDNA sequence analyses. The former would place the roots of the
Braconidae and Ichneumonidae either within groups with that biology, or between
them and the large, entirely koinobiont clade within each family (Quicke and van
Achterberg 1990). However, the percentage GC content of the 28S gene differs mark-
edly between the ichneumonids and braconids, the former typically having 59–63% AT
and the latter 30–52% (Belshaw et al. 1998). The exception is the Trachypetinae (59%),
but in this case their 28S sequence shows large autapomorphic inserts and alignment of
large parts of the gene between families has been considered almost impossible, leaving
little that is obviously homologous while simultaneously being informative about root-
ing (Belshaw et al. 1998). Quicke et al. (1999b) investigated the basal ichneumonoids
using the 28S gene with two aculeates used for rooting and obtained trees in which
Xorides (Xoridinae) was basal to the ichneumonids but, almost certainly erroneously,
with Hybrizon as sister to the Braconidae (including Trachypetinae). Nevertheless,
Labena, Euceros and Brachycyrtus were also recovered close to the bases of the two
major ichneumonid subgroups (i.e. an expanded ophioniformes and a combined ich-
neumoniformes plus pimpliformes, sensu Quicke et al. 2000) in some trees. Therefore,
here we have chosen to present trees rooted using the Xoridinae as they appear at least
to be a relatively basal subfamily and this maintains consistency with Quicke et al.
(2000), though we have no further evidence than that already published as to whether
they are derived more basally than the Labeninae, for example.
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1330 D.L.J. Quicke et al.
Tree statistics for the various analysed data combinations and molecular gas
treatment conditions are presented in Table 3.
Results
Morphological phylogeny
Parsimony analysis of the morphological data matrix comprising all terminal taxa
yielded five trees of length 1291 (Figure 15). While some of the relationships are
consistent with widely held views, other placements are very much at odds with them.
Most notable is that three phrudine taxa (Phrudus group, Pygmaeolus and
Phaestacoenitus) were recovered with the Cryptinae and Agriotypinae, and that the
ichneumoniformes (Cryptinae, Ichneumoninae, Alomyinae and Adelognathinae) was
not recovered. Labeninae (i.e. Labenini + Groteini + Poecilocryptini) was sister to
almost all the pimpliformes taxa as recognized recently, i.e. including Pimplinae,
Poemeniinae, Rhyssinae, Orthocentrinae, Diplazontinae, Cylloceriinae, Acaenitinae
and Collyriinae, but excluding the Diacritinae (Quicke et al. 2000; Laurenne et al.
2006). Pseudorhyssa appeared with the Pimplinae rather than with the Poemeniinae.
The Acaenitinae was rendered polyphyletic because Procinetus was recovered sepa-
rately from Coleocentrus and the remaining acaenitines. The tryphonine tribes were
divided into two groups, a monophyletic Idiogrammatini + Phytodietini, and the
others paraphyletic with respect to three other aberrant subfamilies, but including
Lycorininae, which have recently also been shown to possess anchored eggs
(Coronado-Rivera et al. 2004; Shaw 2004). The two included stilbopine genera,
Stilbops and Panteles, were separated, probably reflecting both the lack of any
obvious synapomorphy (see Wahl 1988) and their very different final instar larval
head capsule skeletons (Short 1957; Quicke 2005). The ctenopelmatine tribes plus
Mesochorinae and Tatogastrinae, but excluding Olethrodotini, formed a grade lead-
ing to Banchinae (monophyletic with Townesion and together sister to Metopiinae),
Lapton, Bremiella, Ischyrocnemis and a clade comprising the Ophioninae (including
Skiapus and Hellwigia), Anomaloninae, Nonninae, Nesomesochorinae, Cremastinae
and Campopleginae.
T
a
bl
e
3
.
S
ummary stat
i
st
i
cs
f
or e
lid
e
d
mo
l
ecu
l
ar an
d
com
bi
ne
d
ana
l
yses.
Data set
Gap
treatment
Morphological
partition
weighting
Number of
informative
characters*
Length of most
parsimonious
tree
Retention
index
Morphology 162 1291 0.507
Elided DNA missing 53,247 1,215,031 0.6852
Elided DNA 5th base 60,260 1,691,800 0.6585
Morphology+ elided
DNA
missing 444 125,175 1,823,755 0.888
Elided DNA plus
morphology
5th base 483 138,506 2,360,406 0.871
* Effective number, i.e. with morphological characters given their up-weighted value.
Downloaded By: [Imperial College] At: 12:43 29 May 2009
Journal of Natural History 1331
Figure 15. Strict consensus of most parsimonious trees from analysis of morphological data
alone (length 1291; RI 0.507; CI 0.166).
Nonninae
Nesomesochorinae
Campopleginae
other Cremastinae
Skiapus
Hellwigia
Ophioninae
Gravenhorstini
Anomalonini
Belesica group
Ischyrocnemis
Olethrodotini
Lapton
Bremiella
Glyptini
Exetastes group
Atrophini
Banchus group
Townesion
Metopiinae
Pergaphaga
Hypopheltes
Westwoodia
Scolobatini
Ctenopelmatini
Mesoleini
Perilissini
Euryproctini
Seleucini
Tatogastrinae
Mesochorinae
Pionini
Tersilochinae
Neorhacodinae
Orthopelmatinae
Microleptinae
Oxytorinae
Alomyinae
Ichneumoninae
Lycorininae
Eucerotinae
Hybrizontinae
Sphinctini
Tryphonini
Exenterini
Eclytini
Oedemopsini
Erythrodolius group
Brachyscleroma
Panteles
Idiogrammatini
Phytodietini
Scolomus
Stilbops
Phaestacaenitus
Pygmaeolus
Phrudus and Astrenis
Adelognathinae
Ateleute
other Cryptinae
Agriotypinae
Diacritinae
Pedunculinae
Claseinae
Brachycyrtinae
Proclitus
Helictes group
Orthocentrus group
Hyperacmus
Collyrinae
Diplazontinae
Cylloceriinae
Coleocentrus
Acaenitini
Procinetus
Ephialtini
Pimpla group
Pseudorhyssini
Delomerista
Perithous
Poemeniinae
Ganodes
Rodrigama
Rhyssinae
Labenini
Groteini
Poecilocryptini
Xoridinae
Downloaded By: [Imperial College] At: 12:43 29 May 2009
1332 D.L.J. Quicke et al.
Molecular phylogeny
Recovery of independently supported groups
Analyses of individual molecular alignments yielded between one and 15 MPTs,
mostly towards the lower end of this range. Relatively few currently accepted taxa,
from large genera to tribe and subfamily level, were recovered in all analyses (i.e. for
all combinations of GO : GE values or for analyses in which gaps were treated as
informative or not). Figure 16 summarizes, for 12 such taxa of various taxonomic
levels, whether or not they were recovered as monophyletic under different alignment
parameters. Yellow indicates that the group was recovered only when gaps were
treated as informative, blue only when gaps were treated as missing data, and black
indicates that the group was recovered when gaps were treated in both ways. The
numbers of recovered, generally accepted groups varied little between the different
alignments so these results do not suggest any particular region of GO : GE parame-
ter space for detailed consideration. Furthermore, no consistent trend was apparent
in the recovery of groups with GO : GE values. What is quite noticeable, however, is
that for groups that were recovered over a large area of parameter space the way in
which gaps were treated was an important factor in their recovery. For example,
Pimpla, Enicospilus (including Pycnophion: Bennett 2008), Exenterini (Tryphoninae),
Phaeogenini (Ichneumoninae) and Oxytorinae were recovered in many cases only
when gaps were treated as uninformative. In contrast, the Phytodietini (Tryphoni-
nae), Scolobatini (Ctenopelmatinae) and Mesochorinae were recovered in more cases
when gaps were treated as informative.
Several generally accepted subfamilies, tribes and genus groups were recovered as
monophyletic across the whole range of alignment parameter space investigated (e.g.
Brachycyrtinae, Labeninae, Orthocentrus group, Campopleginae), and there were
several accepted groups that were never recovered as monophyletic (e.g. Banchinae,
Tryphoninae). Other traditional higher groups were only recovered in some analyses,
either perfectly or occasionally with the inclusion of one or more genera of uncertain
affinity or with one species absent (though in these cases the excluded taxon was
often one for which we only obtained a partial sequence, e.g. the ophionine
Laticoleus infumatus). The Xoridinae, and even Xorides (Figure 16), were frequently
recovered in disparate places, with Xorides or some species thereof often recovered
among the Ophioniformes next to Lycorina and/or Brachyscleroma.
The Ichneumoninae was only recovered as monophyletic with GO : GE of 1 : 1;
with all other parameter combinations the Hybrizontinae were recovered within
them, often as the sister group of Lusius, and often leading Lusius to be separated
from the rest of the Phaeogenini. This strange and highly unlikely association was
probably recovered because the Lusius 28S sequence has numerous deletions (see
Laurenne et al. 2006). The Ophioninae (in the sense of Quicke et al. 2005, i.e. includ-
ing the genera Skiapus and Hellwigia) were recovered as monophyletic with only a
few intermediate GO and GE values; with lower values they fall into two groups
(more or less equivalent to the old tribes Ophionini and Enicospilini), whereas at
higher values, two species of Ophion were recovered separately. The Rhyssinae were
never recovered as monophyletic because Rhyssa was always included among a clade
comprising most Poemeniinae.
A number of higher and lower level relationships were consistently recovered
including a monophyletic pimpliformes clade, comprising Pimplinae, Rhyssinae,
Downloaded By: [Imperial College] At: 12:43 29 May 2009
Journal of Natural History 1333
Figure 16. Summary of recovery of selected genera and higher taxa as monophyletic in most
parsimonious trees from analysis of CLUSTAL-aligned molecular data alone, showing effects o
f
different gap-opening and gap-extension values and treatment of gaps as informative or unin-
formative. Groups are coloured if they were recovered in any of the most parsimonious trees
obtained from a particular combination of parameters.
1 12
gaps
uninformative
only
gaps
informative
only
recovered
with both
never
recovered
Exenterini Phytodietini
Scolobatini
Oxytorinae
+
Westwoodiini
Ophioninae
incl.
Cremastinae
Xoridinae Pimplini
Mesochorinae
grp
Diacritinae Phaeogenini
grp Atrophini
Downloaded By: [Imperial College] At: 12:43 29 May 2009
1334 D.L.J. Quicke et al.
Poemeniinae, Orthocentrinae, Microleptinae, Cylloceriinae, Diplazontinae, Acaenitinae,
Collyriinae and Ischyrocnemis, though sometimes (with very low GO : GE values)
nested within the ichneumoniformes. Several other consistent relationships were
found, including a sister group relationship between Scolobates and the Cremastinae;
monophyly of a clade comprising Tersilochinae, Mesochorinae and the Erythrodolius
genus group (of the Phrudinae); Oxytorus among the Ctenopelmatinae complex; and
a monophyletic Pedunculinae plus Claseinae. Contrary to the results of Quicke et al.
(2000), the Orthopelmatinae were nearly always recovered either within the Tryphon-
inae or Banchinae or Banchinae + Tryphoninae, or occasionally with members of the
Erythrodolius group of genera of the Phrudinae.
Cylloceria and Hyperacmus were recovered as sister groups, especially with
higher gap costs when gaps were treated as either missing or informative, and over a
somewhat larger area of parameter space when gaps were uninformative. Despite
their marked difference in habitus, Hyperacmus and Cylloceria share many features,
most notable of which is the probably uniquely completely divided venom reservoir
(Figure 10A,B). In addition, the venation is similar, males of both have concave
tyloids on rather basal flagellomeres, and in both genera the propodeum is elongated
with strong latero-median carinae (Wahl and Gauld 1998).
Combined molecular and morphological phylogenies
The addition of morphological data radically improved the probabilities of a given
subfamily being recovered as monophyletic and it seems that the molecular signal is
generally rather weak. For example, the Polysphincta group of the Pimplinae are
never recovered as monophyletic in the molecular analyses, where the pimpliform
genera did not form many suprageneric groupings that would be recognized on
morphological grounds. However, despite the Ephialtini being coded uniformly for
morphology, the Polysphincta group were recovered as a clade.
Recovery of independently supported groups
Despite including no autapomorphies for our terminal suprageneric taxa in the
morphological matrix, virtually all such groups were recovered as monophyletic in
the combined analyses. The recoveries of selected relationships are summarized in
Figures 17–20. The basic picture shown by most analyses agreed well with that found
by Quicke et al. (2000), with three large groupings apparent: ichneumoniformes,
pimpliformes and ophioniformes.
The ichneumoniformes, in a narrow sense comprising just the Ichneumoninae,
Alomyinae, Adelognathinae and Cryptinae, were recovered in the great majority of
analyses, though sometimes also with the Agriotypinae as sister to the
Cryptinae + Adelognathinae. All trees included Alomya and Ichneumoninae as sister
taxa. Most parameter combinations produced a clade comprising Cryptinae plus
Adelognathinae, though often the former was paraphyletic with respect to the latter.
When the ichneumoniformes were not recovered this was almost always because a
clade comprising Microleptinae + Eucerotinae, with or without Agriotypinae, were
recovered as sister to the Cryptinae. With high GO costs, the Cryptinae were some-
times recovered as paraphyletic with respect to the Ichneumoninae + Alomyinae.
Downloaded By: [Imperial College] At: 12:43 29 May 2009
Journal of Natural History 1335
The Claseinae + Pedunculinae always formed a monophyletic group, and usually
they were the sister group to the Brachycyrtinae, sometimes also with the Eucerotinae
and Microleptinae. These five small subfamilies were usually placed basal to the strict
ichneumoniformes group, but sometimes within it, and sometimes basal to the
ichneumoniformes + pimpliformes.
Figure 17. Summary of recovery of selected basal and Pimpliformes clades as monophyletic in
most parsimonious trees from analysis of combined morphological and CLUSTAL-aligned
molecular data showing effects of different gap-opening and gap-extension values and treat-
ment of gaps as informative or uninformative. Groups are coloured if they were recovered in
any of the most parsimonious trees obtained from a particular combination of parameters.
1 12
12
1
gaps
uninformative
only
gaps
informative
only
recovered
with both
never
recovered
Gap extension cost
Gap extension cost
Xoridinae
Labeninae
the rest
Pimplinae+Pseudorhyssa+
Rhyssinae+Poemeniinae
Diacritinae
Acaenitinae (mono
or paraphyletic or
nested within)
Diacritinae
'Pimpliformes'
Orthocentrinae+
Diplazontinae+Coll.+
Hyperac.+Ischyr.+
Cylloceriinae
Orthoc.+Diplazont.+
Collyria+Cyllocer.+
Ischyrocnemis
Poemeniinae
other Pimplinae
Rhyssinae
Pseudorhyssa
Delomeristini
Poemeniinae
other Pimplinae
Rhyssinae
Pseudorhyssa
Delomeristini
Poemeniinae
other Pimplinae
Rhyssinae
Pseudorhyssa
Delomeristini Delomeristini Delomeristini
Poemeniinae
other Pimplinae
Rhyssinae
Pseudorhyssa
Poemeniinae
other Pimplinae
Rhyssinae
Pseudorhyssa
Acaenitinae Acaenitinae
Pimplinae+
Poemeniinae+
Rhyssinae
Diacritinae
Pimplinae+Poemeniinae+
Rhyssinae+Acaenitinae
Orthocentrus group
other Orthocentrinae
Diplazontinae
Collyria+Hyperacmus
Orthocentrus group
other Orthocentrinae*
Diplazontinae
Orthocentrus group
other Orthocentrinae*
Diplazontinae
Cylloceria
Allomacrus
Ischyrocnemis
Ischyrocnemis+Allomacrus+
Cylloceria+Hyperacmus
Downloaded By: [Imperial College] At: 12:43 29 May 2009
1336 D.L.J. Quicke et al.
A pimpliformes group comprising all members of the Pimplinae, Poemeniinae,
Rhyssinae, Orthocentrinae, Diplazontinae, Acaenitinae (though not always monophyletic;
Figure 17), Diacritinae, Collyriinae, Cylloceriinae and, surprisingly, Ischyrocnemis, was
recovered in virtually all analyses, and, in nearly all cases (Figure 17), the first three
formed a clade. In slightly less than half the analyses, the Diacritinae formed a sister
group to the remaining pimpliformes (Figure 17), especially when gaps were treated
as informative, and in several others it was sister to the pimpliformes excluding
Figure 18. Summary of recovery of selected ichneumoniformes and basal “ophioniformes” clades
as monophyletic in most parsimonious trees from analysis of combined morphological and
CLUSTAL-aligned molecular data showing effects of different gap-opening and gap-extension values
and treatment of gaps as informative or uninformative. Groups are coloured if they were recovered
in any of the most parsimonious trees obtained from a particular combination of parameters.
1 12
12
1
gaps
uninformative
only
gaps
informative
only
recovered
with both
never
recovered
Gap extension cost
Gap extension cost
Cryptinae
Adelognathus
Ichneumoninae
Alomya
Cryptinae
Ichneumoninae
Alomya
Cryptinae+Adelognathus
Ichneumoninae
Alomya
Adelognathus
Ichneumoninae
Alomya
Cryptinae
Cryptinae+Adelo.
Agriotypus
Ichneumoninae
Alomya
'Ophioniformes'+/-
Lycorina
Orthopelma
Tryphoninae
other 'Ophioniformes'
Orthopelma
Tryphoninae part
Tryphoninae part
Stilbops +/- Panteles
Stilbopinae
Idiogramma
Phytodietini
other Tryphoninae
Tryphoninae+
Stilbops+/-Panteles
Phrudinae
Tersilochinae
Neorhacodinae
Phaestac.+Pygmaeolus
Tersilochinae
Tersilochinae Neorhacodinae
Neorhacodinae Astrenis+Phrudus
Astrenis+Phrudus Erythrodolius grp
Phaestac.+
Pygmaeolus
Phaestac.+
Pygmaeolus
Metopiinae+/-Bremiella
Scolomus
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Journal of Natural History 1337
Pimplinae, Poemeniinae and Rhyssinae. Rodrigama always formed the sister group
to the other Poemeniinae excluding Pseudorhyssa, and the Pimplini was always
monophyletic with the Ephialtini. The relationships of Pseudorhyssa, and of the
Delomeristini (Perithous and Delomerista), varied greatly depending on alignment
parameters, and the configurations illustrated in Figure 17 are not exclusive. Almost
all possible topologies were recovered under some combinations of parameters. The
Orthocentrus group was monophyletic in nearly all analyses (> 95% of parameter
Figure 19. Summary of recovery of selected ophioniformes clades as monophyletic in most
parsimonious trees from analysis of combined morphological and CLUSTAL-aligned molecular
data showing effects of different gap-opening and gap-extension values and treatment of gaps
as informative or uninformative. Groups are coloured if they were recovered in any of the most
parsimonious trees obtained from a particular combination of parameters.
1 12
12
1
gaps
uninformative
only
gaps
informative
only
recovered
with both
never
recovered
Gap extension cost
Gap extension cost
Westwoodiini
Scolobatini
Banchinae +Townesion
other Banchinae
Banchus grp part
Banchus grp part
Townesion
Westwoodiini
Scolobatini
Banchinae
Metopiinae+Bremiella
Ophioninae+Skiapus
Ophioninae+/-Skiapus
Tatogaster
Mesochorinae
Tatogaster
Cremastinae
Campopleginae
other Anomaloninae
Anomalon
other Anomaloninae
Anomalon
Skiapus
Hybrizontinae
other Anomaloninae
Nonninae
Neomesochorinae
Anomalon
other Anomaloninae
Nonninae
Neomesochorinae
Anomalon
Skiapus
Hybrizontinae
Campopleginae+Cremastinae+
Anomaloninae+Nonninae+
Neomesochorinae
+/-Hybrizon+/-Skiapus
Ophioninae+Skiapus
Nonninae+Nesomeso.
other Anomaloninae
Anomalon
Downloaded By: [Imperial College] At: 12:43 29 May 2009
1338 D.L.J. Quicke et al.
combinations) but Orthocentrus was always paraphyletic with respect to all the other
genera within the group.
The Acaenitinae (when recovered as monophyletic) formed a sister group to either a
subset of or all of the pimpliformes. The Diplazontinae nearly always formed a clade with
the Orthocentrinae but sometimes also included Hyperacmus, Collyriinae, Cylloceriinae
and Ischyrocnemis. Hyperacmus was never recovered with the Orthocentrinae except,
less commonly, when a clade (Hyperacmus +Ischyrocnemis + Cylloceriinae) was
interspersed between the Orthocentrus group and the remaining Orthocentrinae. In
the molecular trees it was often recovered as sister to Cylloceria, which is in agree-
ment with the morphology of their venom reservoirs, which in both cases are com-
pletely longitudinally divided (Figure 10A,B). In the combined analyses Hyperacmus
was recovered with Collyria, in agreement with the morphology-only analysis.
Agriotypus was either recovered within the ichneumoniformes (sometimes as sister
group to Cryptinae + Adelognathinae) or appeared basal to the ichneumoniformes,
usually as sister group to the Brachycyrtinae.
The orthopelmatiformes was proposed by Quicke et al. (2000), containing just
Orthopelma, because in their analyses the group appeared as sister to the ophioniformes
Figure 20. Summary of recovery of “higher ophioniformes” group of subfamilies as mono-
phyletic in most parsimonious trees from analysis of combined morphological and CLUSTAL-
aligned molecular data showing effects of different gap-opening and gap-extension values and
treatment of gaps as informative or uninformative. Groups are coloured if they were recovered
in any of the most parsimonious trees obtained from a particular combination of parameters.
1 12
12
1
gaps
uninformative
only
gaps
informative
only
recovered
with both
never
recovered
Gap extension cost
Gap extension
cost
Campopleginae+Cremastinae+
Ophioninae+Tatogastrinae+
Anomaloninae+Hybrizontinae+
Skiapus+Nesomesochorinae+
Nonninae+Mesochorinae
Eucerotinae
+Claseinae
+Brachycyrtinae
+Pedunculinae
Ophioninae+Mesochorinae+
+/-Hybrizontinae+/-Skiapus
+/-Lapton
Campopleginae+Cremastinae+
Ophioninae+Tatogastrinae+
Anomaloninae+Skiapus+
Nesomesochorinae+Nonninae
+Mesochorinae
Eucerotinae +
Microleptinae
Ophioninae+Campopleginae
+/-Tatogastrinae+/-Skiapus
+/-Hybrizontinae +/-Lapton
Downloaded By: [Imperial College] At: 12:43 29 May 2009
Journal of Natural History 1339
but with no apparent morphological synapomorphies. This sister-group relationship
was recovered here with many parameter combinations. When it was not, the reason
was usually because Orthopelma was included in a clade, often basal to the other
ophioniformes, with some combination of Townesion, Lycorina, Brachyscleroma,
Erythrodolius group or Hybrizon.
The ophioniformes is taken here to comprise the bulk of the remaining koinobiont
subfamilies, namely a set that we refer to as the higher ophioniformes (comprising
Campopleginae, Cremastinae, Anomaloninae, Mesochorinae, Nesomesochorinae, Non-
ninae, Ophioninae and Tatogastrinae), and a basal grade of Banchinae, Ctenopelmatinae,
Metopiinae, Neorhacodinae, Oxytorinae, Phrudinae, Stilbopinae, Tersilochinae and
Tryphoninae, not all of which were recovered as monophyletic.
The Banchinae excluding Townesion were always recovered as monophyletic, and
in many analyses Townesion was recovered as their sister group, or nested basally
among the Banchus group of genera (Figure 19), especially when gaps were treated as
uninformative. Hence, the combined data support the conclusion of Gauld and Wahl
(2000b) that Townesion is best regarded as a member of the Banchinae and not separ-
ate “among the primitive subfamilies” as suggested by Kasparyan (1993), although
our data do not support their hypothesis of a close relationship between Townesion
and Sjostedtiella. The Banchus group of genera was never recovered as monophyletic.
The two sequenced stilbopine genera, Panteles and Stilbops, were nearly always
recovered either adjacently as a grade or as a monophyletic group, and with the great
majority of alignment parameters, were either sister to the Tryphoninae, interspersed
among them, or next to them as a grade (Figure 18).
The Ctenopelmatinae were never recovered as monophyletic, either alone or with
the Metopiinae or Mesochorinae included. Most consistently separate were the
Scolobatini and Westwoodiini, which very frequently formed a clade (Figure 19) or
were placed together as a grade.
The Phrudinae (even excluding Brachyscleroma and Seleucus) were only recovered
as monophyletic in one analysis (Figure 18). However, four genera of small phrudine
wasps (referred to here as “microphrudines” and comprising Astrenis, Phrudus,
Pygmaeolus and Phaestacoenitus), Neorhacodinae and Tersilochinae formed a very
consistent clade. The Erythrodolius group was sometimes included in the clade.
The Campopleginae were most often recovered with the Cremastinae, or in a
larger clade with the Cremastinae, Anomaloninae, Nesomesochorinae and Nonninae
(with or without Hybrizon).
The Ophioninae were nearly always recovered as monophyletic, with the aberrant
genus Skiapus usually recovered either as its sister group or included within it, often as the
sister group to Dicamptus. However, in a large minority of analyses Skiapus instead formed
a clade with Anomalon and Hybrizon, almost certainly as a result of long-branch attraction.
The placements of some taxa were highly inconsistent, notably the Lycorininae and
Hybrizontinae. Lycorina and Hybrizon were frequently associated with one another, or
the Lycorininae was sometimes recovered within the Banchinae as sister to Lissocaulus,
but the latter has a highly aberrant sequence. The placements of these taxa in particular
seem likely to be the result of long-branch attraction, as was indicated by Gillespie et al.
(2005) for Hybrizon, and appeared to effect results in the cases of several other ichneu-
monids based on 28S sequence data in the study of cryptines and other ichneumonids
by Laurenne et al. (2006). The presence in Lycorina of a reflexed dorsal ovipositor valve
base and an aulaciform rod (characters 114 and 118), as found in some members
Downloaded By: [Imperial College] At: 12:43 29 May 2009
1340 D.L.J. Quicke et al.
of Tryphoninae (Idiogrammatini and Phytodietini), Banchinae, Cremastinae and
Stilbopinae, strongly suggest that it belongs to the ophioniformes.
Little can be said about the relationships of the four genera currently placed with
uncertainty in the Metopiinae, namely Ischyrocnemis, Scolomus, Lapton and Bremiella.
The first of these was consistently recovered within the pimpliformes (Figure 17),
Scolomus was recovered with (as sister to) the Metopiinae in only a few analyses
(Figure 18), as was Bremiella. The placement of Lapton was very inconsistent, though in
several analyses it was recovered with a group of “higher ophioniformes” including
Ophioninae, Campopleginae, Anomaloninae and Mesochorinae. With the exception of
Ischyrocnemis, these genera appeared associated with various Ctenopelmatinae.
Molecular elision trees
Contrary to expectations, analysis of the elided molecular data, with gaps treated
either as missing characters or informative, yielded trees with few generally accepted
groups, indeed fewer than in several of the individual analyses, and therefore they are
not considered further here.
Combined weighted morphology and molecular elision trees
The combined morphological data and molecular elision trees are presented in
Figures 21 and 22 for gaps treated as missing and informative, respectively.
Relationships at genus level were often completely congruent between the two
analyses. The general picture of relationships obtained was also very similar to those
from combined analyses of individual alignments. Whereas the Labeninae and Xoridinae
formed a basal grade in only some of the individual analyses (Figure 17), they did form a
basal grade in both combined elision trees. In both cases, when rooted with the Xoridinae
as depicted, the Labeninae then formed a sister group to pimpliformes +
(ichneumoniformes + ophioniformes), each in a broad sense. The pimpliformes
included all the taxa placed in it by Wahl (1990) but with the addition of Collyriinae.
The ichneumoniformes in these trees includes the Ichneumoninae, Alomyinae,
Cryptinae, Adelognathinae and Agriotypinae. When gaps were treated as informative a
principally Gondwanan clade of Brachycyrtinae + Claseinae + Pedunculinae formed the
sister group to the ichneumoniformes, whereas when gaps were treated as missing the
Gondwanan clade was sister group to the ichneumoniformes + ophioniformes.
The pimpliformes was consistent between both analyses (Figures 21, 22) with
Diacritinae + (Procinetus + ((other Acaenitinae + (Cylloceriinae + ((Diplazontinae +
Collyriinae + Hyperacmus) + +Orthocentrinae))) (Rhyssinae + (Poemeniinae + Pim
plinae)))). Whereas the Acaenitinae were monophyletic in many of the individual
combined analyses (Figure 17), Procinetus was separated from the others in both
combined elision analyses (Figures 21, 22).
Several potentially important differences between the combined elided trees and the
individual combined analyses should be noted. Unlike all individual combined analyses
in which the Eucerotinae and Microleptinae, usually as a monophyletic clade, were recov-
ered with the ichneumoniformes or very basal (usually as sister group to the ichneumoni-
formes and pimpliformes combined), in the combined elision trees the Eucerotinae
appeared within the Tryphoninae and Microleptes as sister to Orthopelma within the
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Journal of Natural History 1341
Figure 21. Strict consensus most parsimonious tree of analysis of elided molecular data set
(with gaps treated as missing data) combined with morphological data up-weighted 444 times
to give equivalent total clade support (length 1,823,755; RI 0.888). Coloured bars indicate
current subfamily classification.
Labena A
Labena B
Certonotus annulatus
Apechoneura
Grotea cortesi
Labium
Grotea vanessae
Poecilocryptus nigromaculatus
Odontocolon dentipes
Odontocolon mellipes
Odontocolon punctulatum
Ischnocerus caligatus
Xorides maculiceps
Xorides propinquus grp
Xorides brookei grp
Xorides praecatorius
A
B
Eusterinx bispinosa
Eusterinx B
Eusterinx A
Kentrotryphon longicaudatus
Symplesis bicingulata
Symplesis
Megastylus B
Megastylus C
Megastylus A
Pantisarthrus inaequalis
Dialipsis exilis
Plectiscidea A
Plectiscidea B
Helictes conspicuus
Gnathochorisis crassulus
Gnathochorisis dentifer
Hemiphanes erratum
Aperileptus
Apoclima
Entypoma robustator
Proclitus
Neurateles A
Stenomacrus B
Orthocentrus A
Picrostigeus recticauda
Orthocentrus B
Stenomacrus A
Orthocentrus K
Orthocentrus N
Plectiscus
Orthocentrus H
Orthocentrus L
Orthocentrus F
Orthocentrus G
Orthocentrus D
Orthocentrus C
Chilocyrtus
Orthocentrus I
Orthocentrus E
Orthocentrus J
Neurateles B
Orthocentrus M
Diplazon tetragonus
Diplazon annulatus B
Diplazon laetatorius Z97925
Diplazon annulatus A
Campocraspedon annulitarsis
Tymmophorus obscuripes
Syrphophilus tricinctorius
Xestopelta gracillima
Enizemum ornata
Phthorima compressa
Syrphoctonus megaspis
Syrphoctonus longiventris
Syrphoctonus cultiformis
Woldstedtius cf biguttatus
Woldstedtius holarcticus
Syrphidepulo chaconi
Woldstedtius flavolineatus
Promethes melanaspis
Promethes
Sussaba cf flaviceps
Sussaba
Collyria coxator
Collyria
Hyperacmus crassicornis
Cylloceria melancholica
Cylloceria
Allomacrus arcticus
Jezarotes tamanukii
Spilopteron apicale
Yezo ce r yx
Phaenolobus
Phorotrophus
Ishigakia exetasea
Yamatarotes bicolor
Coleocentrus
Coleocentrus chipsani
Procinetus decimator
Diacritus aciculatus
Ortholaba tenuis
Flavopimpla A
Hemipimpla
Xanthophenax depressor
Flavopimpla B
Sericopimpla
Iseropus stercorator
Zaglyptus
Xanthophenax A
Xanthophenax B
Pimplaetus
Acropimpla
Scambus
Camptotypus
Dolichomitus annulicornis
Gregopimpla
Endromopoda arundinator
Scambus signatus grp
Clistopyga incitator
Clistopyga ? incitator
Ephialtes hokkaidonis
Ephialtes manifestator
Zonopimpla cf morai
Tromatobia variabilis
Dolichomitus agnoscendus
Liotryphon caudatus
Calliephialtes nr thurberiae
Anastelgis garciai
Dolichomitus imperator
Exeristes
Zatypota A
Zatypota C
Sinarachna nigricornis
Zatypota B
Acrodactyla degener
Eruga
Acrotaphus nr tibialis
Hymenoepimecis
Polysphincta ?purcelli
Inbioia
Piogaster
Schizopyga pictifrons
Schizopyga frigida
Brachyzapus unicarinatus
Pimpla luctuosa
Pimpla rufipes
Pimpla alboannulata
Pimpla disparis
Pimpla mahalensis
Pimpla A
Pimpla flavicoxa
Pimpla B
Apechthis zapoteca
Apechthis B
Apechthis A
Echthromorpha
Itoplectis naranyae
Itoplectis
Xanthopimpla A
Xanthopimpla B
Lissopimpla albopicta
Theronia Neotheronia
Theronia
Theronia Nomosphecia
Theronia Poecilopimpla
Theronia Epitheronia
Theronia Parema
Delomerista mandibularis
Perithous scurra
Deuteroxorides elevator
Neoxorides nitens
Ganodes matai
Eugalta
Cnastis
Poemenia hectica
Podoschistus scutellaris
Rodrigamma gamezi
Pseudorhyssa alpestris
Pseudorhyssa nigricornis
Myllenyxis
Triancyra
Epirhyssa (Sychnostigma)
Cyrtorhyssa
Megarhyssa
Lytarm es
Rhyssella approximator
Epirhyssa mexicana
Rhyssa amoena
Rhyssa
A
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1342 D.L.J. Quicke et al.
Ctenopelmatinae (gaps informative; Figure 22) or, also among the ctenopelmatine group,
but next to Tersilochinae + “microphrudines” + Neorhacodinae (gaps missing).
The Cremastinae was recovered as monophyletic in all the individual
analyses but in the elision tree with gaps treated as missing data, Eurygenys
Figure 21. (Continued)
Brachycyrtus convergens
Brachycyrtus A
Brachycyrtus B
Pedunculus
Adelphion
Clasis A
Clasis B
Macrojoppa blandita
Macrojoppa
Daggoo ?philoctetes
Diadromus troglodytes
Lusius
Lusius tenuissimus
Imeria seyrigi
Imeriella
Afrotrogus nr nigripedalis
Holchichneumon ?areolator
Compsophorus ?metallicus
Compsophorus seyrigi
Ulesta perspicua
Setanta
Virgichneumon maculicauda
Amblyjoppa
Coelichneumon ?orbitator
Protichneumon
Heresiarches endoxius
Heresiarchini gen sp indet
Cratichneumon
Homotherus locutor
Crytea nr immaculaticeps
Neotypus
Ctenichneumon funereus
Ichneumon
Achaius oratorius
Limerodops elongatus
Diphyus latebricola
Diphyus quadripunctorius
Amblyteles armatorius
Diphyus palliatorius
Eutanyacra glaucatoria
Spilothyrateles punctus
Crypteffiges albilarvatus
Ichneumon promissorius
Obtusodonta equitatoria
Ichneumon sarcitorius
Ceratojoppa seyrigi
Tricyphus
Ischnojoppa ?flavipennis
Ctenochares bicolorus
Pseudoamblyteles homocerus
Stenichneumon culpator
Gavrana maculipes grp
Uloola brevis
Joppa
Eccoptosage
Joppa dorsata
Listrodomus
Lophojoppa
Dilopharius
Anisobas hostilis
Neischnus oxypygus
Chasmias motatorius
Cratichneumon fabricator
?Gavrana
Leptogea
Phaesurella
Aethioplitops fulvator
Trogus lapidator
Joppocryptus
Magwengiella
Aethianoplis ?excavata
Stenobenyllus pictus
Vulgichneumon
Vulgichneumon saevus
Eurylabus torvus
Notosemus bohemani
Afrectopius
Platylabus ?decipiens
Hypomecus quadriannulatus
Cotiheresiarches dirus
Probolus concinnus
Cyclolabus nigricollis
Linycus exhortator
Eparces
Tyc he ru s
Phaeogenes
Aethecerus nitidus
Centeterus rubriginosus
Colpognathus
Oiorhinus pallipalpis
Oronatus binotatus
Baeosemus
Pseudalomya
Phaeogenini gen sp indet
Epitomus infuscatus
Hemichneumon subdolus
Misetus oculatus
Nematomicrus tenellus
Heterischnus
Notacma
Alomya debellator
Alomya semiflava
B
D
C
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Journal of Natural History 1343
(Belesica group) was separated from the rest (Figure 21E). In contrast, neither
the Banchus group of genera in the Banchinae nor the Polysphincta group in
Pimplinae were ever recovered as monophyletic in any of the individual
analyses but both were in the elision tree with gaps missing (Figure 21D),
Figure 21. (Continued)
Aconias
Parmortha
Colocnema rufina
Rhembobius perscrutator
Rhembobius
Cubocephalus anatorius
Demopheles corruptor
Aptesis nigrocincta
Mansa singularis
Stomacis
Meringops
Ateleute
Tamaulipeca
Ateleute linearis
Handaoia
Austriteles
Rothneya
Theroscopus
?Phygadeuon C
Phygadeuon A
Ceratophygadeuon
Phygadeuon B
Amydraulax
Mastrus
Cremnodes atricapillus
Orthizema hadrocerum
Charitopes
Lochetica westoni
Isadelphus gallicola
Dichrogaster longicaudata
Dichrogaster ?modesta
Mastrus deminuens
Tropistes
Gnotus chionops
Atractodes ?fittoni
Atractodes gravidus
Mesoleptus ?devotus
Stilpnus pavoniae
Mesoleptus A
Mesoleptus angustulus
Mesoleptus B
Gnotus
Hyparcha
Aclastus A
Nipponaetes haeussleri
Hemiteles ?maricesca
Cisaris
Chirotica A
Chirotica B
Gabia
Paraphylax B
Bentyra B
Astomaspis Caenopimpla
Paraphylax C
Asmenophylax
?Paraphylax D
Xenolytus bitinctus
Gelis B
Thaumatogelis audax grp
Grasseiteles puncta
Gelis A
Agasthenes varsitarsus
Encrateola B
Gelis rufogaster
Micromonodon tener
Palpostilpnus
Trachaner
Isdromas granulatus grp
Acrolyta
Isdromas A
Acrolyta marginata
Lysibia ceylonensis
Bodedia striata
Tricholinum ischnocerum
Diaglyptidea conformis
Xenolytus
Brachedra
?Stiboscopus
Distathma
Stenotes
Acidnus
Toechorychus nr abactus
Encrateola A
Eudelus ?simillimus
Paraphylax A
Isdromas B
Micraris
Zoophthorus nr bridgmani
Bilira
Idiolispa
Endasys
Bathythrix B
Chrysocryptus
Bathythrix laminata
Bathythrx collaris
Surculus A
Surculus B
Bathythrix pellucidator
Bathythrix A
Polyaulon paradoxus
Adelognathus
Agriotypus armatorius
Cryptanura spinaria
Cryptanura A
Cryptanura B
Iaria
Listrognathus
Buodias
Nematocryptus B
Owenus
Trafana A
Prionacis
Stiromesostenus
Fotsiforia
Mesostenus C
Larpelites
Mesostenus B
Stenarella A
Stenarella B
Frion a
?Baltazaria
Goryphus B
Goryphina gen sp indet
Goryphus A
Isotima
Picardiella
Irabatha cairnensis
Diloa antipodialis
Trafana B
Nematopodius debilis
Polycyrtus
Hemisphragia
Camera
Lanugo
Joppidium
?Bodedia
Diapetimorpha A
Dicamixus
Thrybius
Bathyzonus
Sphecophaga vesparum
Mesostenus A
Nematocryptus A
Trachysphyrus
Polytribax
Messatoporus
Photocryptus
Gnypetomorpha
Acroricnus stylator
Cestrus
?Dotocryptus
Gambrus
Hoplocryptus C
Hoplocryptus B
Thrybius brevispina
Odontocryptus ?spiniclunatus
Lamprocryptus
Bentyra A
Bicristella
Dotocryptus
Trychosis A
Trychosis B
Hidryta
Agrothereutes abbreviatus
Ischnus B
Aritranis director
Hoplocryptus A
Chromocryptus
Trachysphyrus agenor
Baryceros
Lymeon B
Lymeon C
Lymeon A
Acerastes
Diapetimorpha B
Priotomis
Platymystax
Anacis B
Aclastus B
Cryptinae gen sp indet B
Dinocryptus
Xoridesopus
Torbda
Xanthocryptus
Cryptinae gen sp indet B
Gabunia A
Gabunia B
Arhytis
Hadrocryptus
Microstenus canaliculatus
Agonocryptus chichimecus
Eurycryptus laticeps
Dagathia
Echthrus reluctator
Helcostizus restorator
Pterocryptus
Buathra laborator
Cryptus armator
Chlorocryptus
Coccygodes
Coesula
Ischnus A
Arthula
Latibulus ?argiolus
Glabridorsum stokesii
Ceratomansa prima
Ceratomansa
Cryptus viduatorius
Anacis A
Myrmelonostenus
C
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1344 D.L.J. Quicke et al.
though in that tree Townesion is sister to Lycorina and together these are sister
to the remaining Banchinae.
Ischyrocnemis was recovered, along with the Eucerotinae, among the Tryphoni-
nae when gaps were treated either as missing or informative (Figures 21D, 22D,
respectively). The association with the Eucerotinae and their recovery with them
among the tryphonines seem highly improbable.
Discussion
The general picture of relationships that has emerged from the current study is
largely in agreement with that proposed by Quicke et al. (2000) based on far sparser
taxon sampling and fewer morphological characters. Our summary conclusions are
presented in Table 4. The Ichneumonidae comprises three large monophyletic
groups, the ophioniformes in a much enlarged sense relative to Wahl (1991), forming
the sister group to the ichneumoniformes + pimpliformes, with the orthopelmatiformes
Figure 21. (Continued)
Lissocaulus ?emaceratus
Wahlamia phloridifobia
Tossinola
Alloplasta A
Alloplasta B
Cryptopimpla quadrilineata
Amphirhachis D
Stictolissonota
Amphyrhachis A
Amphirhachis B
Amphirhachis C
Arenetra pilosella
Lissonota A
Lissonota Isomeris D
Lissonota B
Diradops mexicana
Hadrostethus hansoni
Diradops
Hylesicida crassitarsus
Meniscomorpha kerna
Meniscomorpha sotoi
Meniscomorpha zacasta
Meniscomorpha albomaculata grp
Quilonota cangrejae
Lissonota C
Odinophora ?graecator
Loxodotus pallidus
Spilopimpla D
Spilopimpla B
Spilopimpla C
Spilopimpla A
Leptobatopsis indica
Leptobaptopsis
Atropha
Occia jereza
Occia
Syzeuctus A
Syzeuctus B
Syseuctus C
Atrophini gen sp indet
Mnioes albispina
Lissonota like E
gen nov nr Cordeleboea
Lissonota like D
Hapsinotus convexus
Podeleboea bifenestralis
Glypta wahl
Zaglyptomorpha auxiliadorae
Glypta
Glypta fumiferanae
Glypta altamira
Apophua bipunctoria
Sphelodon phoxopteridis
Australoglypta
Sjoestedtiella
Banchus falcatorius
Banchus volutatorius
Rhynchobanchus bicolor
Geraldus
Banchopsis crassicornis
Philogalleria bobbyi
Agathilla bradleyi
Exetastes adpressorius
Exetastes
Exetastes mexicanus
Tetractenion
Townesion ussuriensis
Townesion
Lycorina apicalis
Lycorina triangulifera
Netelia F
Netelia G
Netelia C
Netelia ?infractor
Netelia A
Netelia D
Netelia E
Netelia macrostigma
Netelia B
Netelia virgata
Netelia H
Netelia bicolor
Phytodietus gelitorius
Phytodietus B
Phytodietus A
Idiogramma
Erythrodolius A
Melanodolius
Erythrodolius B
Icariomimus
Erythrodolius griffithsorum
Brachyscleroma
Panteles schuetzeanus
Stilbops vetula
Stilbops limneriaeformis
Ctenochira marginata
Ctenochira
Ctenochira xanthopyga
Ctenochira angustata
Ctenochira sphaerocephala
Ctenochira propinqua
?Erromenus
Erromenus punctatus
Tryphon bidentulus
Tryphon thomsoni
Cosmoconus
Cosmoconus meridionator
Monoblastus ?brachyacanthus
Dyspetes luteomarginata
Polyblastus pedalis
Polyblastus varsitarsus
Polyblastus Labroctonus
Grypocentrus A
Grypocentrus B
Boethus forresti
Boethus thoracicus
Chiloplatys mexicanus
Ibornia
Euceros serricornis
Euceros madecassus
Euceros pruinosus
Ischyrocnemus
Sphinctus gastoni
Eridolius flavomaculatus
Eridolius pictus
Eridolius
Acrotomus succinctus
Kristotomus ridibundus
Exenterus confusus
Exenterus amictorius
Excavarus sibiricola
Exyston sibiricus
Exyston pratorum
Smicroplectus perkinsorum
Neliopisthus A
Neliopisthus B
Zagryphus zulaya
Thymaris A
Thymaris B
Oedemopsis scabricula
Oedemopsis
Oedemopsis ryitoni
Atopotrophos bucephalus
Hercus fontinalis
Eclytus exornatus
D
E + F
Downloaded By: [Imperial College] At: 12:43 29 May 2009
Journal of Natural History 1345
(i.e. Orthopelma) sister to or placed close to the base of the ophioniformes. The Labeninae
are often recovered basal with the Xoridinae in the combined analyses, but some-
times form the sister group to the ichneumoniformes + pimpliformes, or rarely to the
ophioniformes + orthopelmatiformes. Some groups, however, were very labile in
their placements and probably caused some disruption to other relationships. This group-
ing of subfamilies is somewhat different from the early attempts by Gauld (1991, 1997) at
an informal clustering, most notably in the splitting of his “Tryphoniformes”, which
he based on anchored eggs; however, the nature of the anchoring differs among his
Figure 21. (Continued)
Dusona B
Dusona E
Dusona F
Dusona G
Casinaria B
Dusona D
Phobocampe
Cymodusa A
Cymodusopsis
Cymodusa B
Casinaria petiolaris
Rhimphoctona grandis
Rhimphoctona A
Rhimphoctona B
Rhimphoctona obscuripes
Gonotypus melanostoma
Melanoplex bucculentus
Tranosema rostrale
Scirtetes robustus
Charops B
Charops C
Charops A
Casinaria A
Eucaphila
Campoletis sonorensis
Campoletis B
Leptocampoplex cremastoides
Porizon
Nemeritis
Sinophorus townesorum
Bathyplectes curculionis
Dusona A
?Enytus A
Nepiesta rufocincta
Campoletis A
Venturia
Campoplex deficiens
Venturia (Slenda) ocypeta
Venturia canescens
Callidora B
Echthronomas
Callidora A
Eriborus
?Diadegma C
Eriborus ?terebrans
Enytus neoapostata
Cryptophion manueli
Hyposoter A
Olesicampe
?Enytus B
Hyposoter B
Diadegma A
Diadegma B
Melalophacharops B
Hyposoter didymiator
Hyposoter exiguae
Xanthocampoplex B
Melalophacharops A
Diadegma mollipla
Lathrostizus ?lugens
Hyposotor fugitivus
Microcharops
Echthronomas facialis
Xanthocampoplex A
Xanthocampoplex C
Lemophagus errabundusm
Lemophagus pulcher
Prochas
Creagrura nigripes
Eutanygaster tabascensis
Xiphosomella nigroornata sp grp
Eiphosoma macrum sp grp
Eiphosoma B
Eiphosoma A
Xiphosomella
Pimplomorpha
Sustenus ridens
Temelucha B
Eucremastoides
Temelucha A
Cremastus spectator
Ptilobaptus cinctus
Dimophora
Pristomerus vulnerator
Trathala
Chriodes B
Chriodes A
Klutiana
Chriodes C
Nonnus
Ophion bicarinatus
Ophion obscuratus
Ophion costatus
Ophion mocsaryi
Ophion minutus
Ophion zerus
Ophion Platophion ocellaris
Rhopalophion discinervus
Ophion scutellaris
Afrophion hynnis
Xylophion ketus
Alophophion
Agathophiona fulvicornis
Ophion ventricosus
Dictyonotus purpurescens
Rhynchophion flammipennis
Euryophion latipennis
Thyreodon nr atriventris
Thyreodon laticinctus
Enicospilus melanocarpus
Enicospilus ramidulus
Enicospilus herero
Enicospilus umbratus
Pycnophion kauaiensis
Enicospilus equatus
Enicospilus biimpressus
Enicospilus congoensis
Dicamptus pellucidus
Dicamptus seyrigi
Hellwigiella dichromoptera
Pamophion sorus
Ophiogastrella maculithorax
Riekophion emandibulator
Leptophion anici
Barytatocephalus mocsaryi
Janzophion nebosus
Laticoleus unicolor
Staurpoctonus bombycivorus
Eremotylus marginatus
Simophion
Lepiscelus distans
Prethophion latus
Sicophion fenestralis
Hellwigia obscura
Encardia picta
Gravenhorstia
Aphanistes nocturnus
Aphanistes jozankeanus
Agrypon
Parania prima
Agrypon varitarsum
Perisphincter taloomi
Agrypon japonicum
Corsoncus magus
Corsoncus minori
Habronyx
Aphanistes klugii
Aphanistes ruficornis
Aphanistes bellicoides
Aphanistes guadamuzae
Barylypa
Clypeocampulum ?tibiale
Parania
Heteropelma amictum
Trichomma
Brachynervus
Bimentum
Ophiopterus cincticornis
Ophionellus albofascialis
Ophionellus B
Ophionellus A
Podogaster eldae
Podogaster tranae
Anomalon A
Anomalon B
Anomalon C
Hybrizon buccatus
Hybrizon sp
Hybrizon ghilarovi
Skiapus
Eurygenys
E
Downloaded By: [Imperial College] At: 12:43 29 May 2009
1346 D.L.J. Quicke et al.
included taxa (Tryphoninae, Eucerotinae and Adelognathinae) and is not homologous.
Instead, although their placement in the current study is highly problematic because of
their aberrant sequences, the Lycorininae may in fact be related to the Tryphoninae as
both possess eggs with chorionic anchors (Coronado-Rivera et al. 2004; Shaw 2004).
Figure 21. (Continued)
Mesochorus Q
Mesochorus R
Mesochorus J
Mesochorus T
Mesochorus Plectochorus
Mesochorus K
Mesochorus P
Mesochorus L
Mesochorus O
Mesochorus U
Mesochorus V
Cidaphus
Mesochorus S
Mesochorus H
Mesochorus I
Mesochorus M
Mesochorus F
Mesochorus A
Mesochorus C
Mesochorus G
Mesochorus N
Mesochorus E
Mesochorus B
Mesochorus D
Astiphromma
Tatogaster nigra
Sympherta A
Sympherta B
Rhaestus
Rhorus B
Rhorus austriator
Rhorus longicornis
Rhorus A
Hodostates brevis
Trematopygus
Lethades curvispina
Lethades cingulator
Glyptorhaestus
Cacomisthus
Pion fortipes
Pion
Pion nr fortipes
Syntactus
Asthenara socia
Scolomus A
Scolomus B
Phobetes floryae
Phobetes
Synomelix faciator
Zemiophora scutulata
Anisotacrus A
Hadrodactylus faciator
Hadrodactylus
Anisotacrus B
Mesoleptidea
Hypamblys
Pantorhaestus xanthostomus
Syndipnus
Gunomeria sordida
Euryproctus A
Euryproctus B
Cataphygma pectorale
Megaceria A
Megaceria B
Seleucus
Perilissus ?albitarsis
Perilissus albitarsis
Absyrtus vicinator
Absyrtus aff vicinator
Jorgeus jimenzi
Lathrolestes
Coelorhachis
Opheltes glaucopterus
Lathiponus semiluctuosus
Tetrambon
Priopoda
Neurogenia
Perilissus lutescens
Perilissus rufoniger
Nanium mairenai
Diaparsis B
Diaparsis G
Diaparsis E
Diaparsis F
Diaparsis C
Diaparsis D
Diaparsis A
Diaparsis jucunda
Aneuclis
Sathropterus
Barycnemis
Probles
Tersilochus heterocerus
Allophroides
Phradis morionellus
Allophrys
Phradis
Stethantyx A
Stethantyx B
Phaestacoenitus
Pygmaeolus nitidus
Neorhacodes enslini
Astrenis brunneofacies
Astrenis paradoxa
Phrudus badensis
Phrudus defectus
Orthopelma mediator
Orthopelma
Microleptes aquisgranensis
Microleptes
Oxytorus armatus
Oxytorus luridator
Oxytorus
Oxytorus knappe
Himerta
Lagarotis semicaligata
Alexeter A
Anoncus gracilicornis
Mesoleius
Barytarbes
Saotis
Mesoleius armillatorius
Campodorus
Rhinotorus similis
Alexeter nebulator
Alexeter gracilentus
Campodorus variegatus
Anoncus
Scopesis fruterna
Lamachus eques
Scopesis
Campodorus scapularis
Lamachus
Otlophorus
Rhinotorus
Alexeter B
Azelus erythropalpus
Forrestopius brenti
Leurus caeruliventris
Seticornuta albopilosa
Carria
Carria fortipes
Sciron
Periope ?aethiops
Drepanoctonus
Spudaeus
Trieces A
Trieces B
Trieces C
Chorinaeus
Hemimetopius
Acerataspis A
Aceratapsis C
Acerataspis B
Metopius A
Metopius B
Metopius C
Hypsicera A
Hypsicera C
?Hypsicera D
Hypsicera B
Exochus A
Exochus D
Exochus B
Exochus C
Triclistus B
Triclistus C
Triclistus A
Cubus
Colpotrocia ?fusca
Colpotrochia texana
Colpotrochia
Colpotrochia cincta
Xenoschesis fulvipes
Xenoschesis mordax
Xenosch ustulata
Homaspis
Ctenopelma ruficorne
Lapton femoralis
Bremiella pulchella
Olethrodotus modesta
Scolobates
Scolobates auriculatus
Physotarsus
Physotarsus adriani JM186
Pergaphaga
Hypopheltes
Westwoodia
F
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Journal of Natural History 1347
Figure 22. Strict consensus most parsimonious tree of analysis of elided molecular data set
(with gaps treated as informative) combined with morphological data up-weighted 483 times to
give equivalent total clade support (length 2,360,406; RI 0.871). Coloured bars indicate current
subfamily classification.
Labena A
Labena B
Certonotus annulatus
Apechoneura
Grotea cortesi
Labium
Grotea vanessae
Poecilocryptus
Odontocolon dentipes
Odontocolon mellipes
Odontocolon punctulatum
Ischnocerus caligatus
Xorides maculiceps
Xorides propinquus grp
Xorides brookei grp
Xorides praecatorius
Dialipsis exilis
Plectiscidea A
Plectiscidea B
Eusterinx A
Gnathochorisis crassulus
Gnathochorisis dentifer
Helictes conspicuus
Entypoma robustator
Aperileptus
Hemiphanes erratum
Apoclima
Megastylus A
Megastylus C
Megastylus B
Eusterinx bispinosa
Eusterinx B
Kentrotryphon longicaudatus
Symplesis bicingulata
Symplesis
Pantisarthrus inaequalis
Proclitus
Orthocentrus F
Orthocentrus G
Orthocentrus C
Chilocyrtus
Orthocentrus E
Orthocentrus I
Orthocentrus J
Orthocentrus K
Orthocentrus N
Plectiscus
Orthocentrus H
Orthocentrus L
Neurateles B
Orthocentrus D
Orthocentrus B
Stenomacrus A
Neurateles A
Stenomacrus B
Orthocentrus A
Picrostigeus recticauda
Orthocentrus M
Syrphophilus tricinctorius
Tymmophorus obscuripes
Campocraspedon annulitarsis
Xestopelta gracillima
Diplazon tetragonus
Diplazon annulatus B
Diplazon laetatorius Z97925
Diplazon annulatus A
Enizemum ornata
Phthorima compressa
Syrphoctonus megaspis
Syrphoctonus longiventris
Syrphoctonus cultiformis
Woldstedtius cf biguttatus
Woldstedtius holarcticus
Syrphidepulo chaconi
Woldstedtius flavolineatus
Promethes melanaspis
Promethes
Sussaba cf flaviceps
Sussaba
Collyria coxator
Collyria
Hyperacmus crassicornis
Cylloceria melancholica
Cylloceria
Allomacrus arcticus
Jezarotes tamanukii
Spilopteron apicale
Yezoceryx
Phorotrophus
Phaenolobus
Ishigakia exetasea
Yamatarotes bicolor
Coleocentrus
Coleocentrus chipsani
Procinetus decimator
Diacritus aciculatus
Ortholaba tenuis
Xanthophenax A
Xanthophenax B
Calliephialtes nr thurberiae
Dolichomitus agnoscendus
Liotryphon caudatus
Anastelgis garciai
Ephialtes hokkaidonis
Flavopimpla B
Tromatobia variabilis
Scambus signatus grp
Gregopimpla
Clistopyga ? incitator
Camptotypus
Flavopimpla A
Hemipimpla
Xanthophenax depressor
Dolichomitus annulicornis
Endromopoda arundinator
Acropimpla
Pimplaetus
Scambus
Iseropus stercorator
Zaglyptus
Sericopimpla
Dolichomitus imperator
Exeristes
Zonopimpla cf morai
Clistopyga incitator
Ephialtes manifestator
Sinarachna nigricornis
Zatypota A
Zatypota B
Zatypota C
Acrodactyla degener
Eruga
Acrotaphus nr tibialis
Hymenoepimecis
Polysphincta ?purcelli
Inbioia
Piogaster
Schizopyga pictifrons
Schizopyga frigida
Brachyzapus unicarinatus
Pimpla mahalensis
Pimpla A
Pimpla disparis
Pimpla flavicoxa
Pimpla B
Apechthis zapoteca
Apechthis B
Itoplectis naranyae
Itoplectis
Apechthis A
Pimpla luctuosa
Pimpla rufipes
Pimpla alboannulata
Echthromorpha
Xanthopimpla A
Xanthopimpla B
Lissopimpla albopicta
Theronia Neotheronia
Theronia Epitheronia
Theronia Parema
Theronia Poecilopimpla
Theronia Nomosphecia
Theronia
Delomerista mandibularis
Perithous scurra
Neoxorides nitens
Poemenia hectica
Deuteroxorides elevator
Ganodes matai
Eugalta
Cnastis
Podoschistus scutellaris
Rodrigama gamezi
Pseudorhyssa alpestris
Pseudorhyssa nigricornis
Cyrtorhyssa
Megarhyssa
Epirhyssa (Sychnostigma)
Triancyra
Myllenyxis
Rhyssella approximator
Epirhyssa mexicana
Lytarmes
Rhyssa amoena
Rhyssa
A
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1348 D.L.J. Quicke et al.
Implications for classification
The great variation in recovered relationships dependent on analysis observed here
means that only a few taxonomic changes can be made with reasonable confidence
based on current findings, though our results do indicate that it is highly probable
Figure 22. (Continued)
Agriotypus armatorius
Cratichneumon
Homotherus locutor
Crytea nr immaculaticeps
Ichneumon
Ctenichneumon funereus
Neotypus
Achaius oratorius
Limerodops elongatus
Diphyus latebricola
Diphyus quadripunctorius
Diphyus palliatorius
Eutanyacra glaucatoria
Amblyteles armatorius
Spilothyrateles punctus
Heresiarchini gen sp indet
Trogus lapidator
Ischnojoppa ?flavipennis
Imeria seyrigi
Imeriella
Afrotrogus nr nigripedalis
Holchichneumon ?areolator
Compsophorus ?metallicus
Compsophorus seyrigi
Heresiarches endoxius
Setanta
Virgichneumon maculicauda
Protichneumon
Amblyjoppa
Coelichneumon ?orbitator
Platylabus ?decipiens
Cotiheresiarches dirus
Probolus concinnus
Cyclolabus nigricollis
Linycus exhortator
Afrectopius
Hypomecus quadriannulatus
Notosemus bohemani
Vulgichneumon saevus
Stenobenyllus pictus
Vulgichneumon
?Gavrana
Chasmias motatorius
Cratichneumon fabricator
Ctenochares bicolorus
Pseudoamblyteles homocerus
Stenichneumon culpator