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Phylogeny and generic concepts of the parasitoid wasp
family Aulacidae (Hymenoptera: Evanioidea)
Giuseppe F. Turrisi
A,D
, John T. Jennings
B
and Lars Vilhelmsen
C
A
University of Catania, Department of Animal Biology ‘Marcello La Greca’, via Androne 81, I-95124, Catania, Italy.
B
Australian Centre for Evolutionary Biology and Biodiversity, and School of Earth and Environmental Sciences,
The University of Adelaide, SA 5005, Australia.
C
Zoological Museum, Natural History Museum of Denmark, Universitetsparken 15, DK-2100, Copenhagen,
Denmark.
D
Corresponding author. Email: turrisifabrizio@yahoo.it
Abstract. The results of the first phylogenetic investigation of members of the Aulacidae of the world are presented. The
main objective was to test the monophyly of the currently recognised genera. In total, 79 morphological characters were
scored for a substantial sample of the extant aulacid fauna, including 72 species, as well as 12 outgroup taxa belonging to
Evaniidae, Gasteruptiidae, Megalyridae, Trigonalidae, Braconidae and Stephanidae. All zoogeographic regions were
represented. The dataset was analysed under different conditions (ordered, unordered, equal and implied weighting).
The results under different weighting conditions are not fully congruent and many relationships remain unresolved. However,
the analyses demonstrate that the current generic classification of the Aulacidae is not a natural one. There is support for a very
large, monophyletic clade which includes all Pristaulacus Kieffer spp. + Panaulix Benoit spp. This suggests a wider generic
concept for Pristaulacus, which is redefined and rediagnosed here. As a consequence, Panaulix becomes a junior synonym of
Pristaulacus (syn. nov.), and the two described species of Panaulix are transferred to Pristaulacus:Pristaulacus rex (Benoit,
1984), comb. nov., and Pristaulacus irenae (Madl, 1990), comb. nov. The genus Aulacus Jurine was consistently
paraphyletic and is not valid as currently defined. Furthermore, we failed to retrieve a consistent topology among the
different clades of Aulacus. A satisfactory reclassification of Aulacus, however, requires a much more comprehensive taxon
sample and/or additional character data.
Introduction
Aulacidae is a relatively small group of parasitoid wasps which
includes 218 extant described species currently placed in three
genera (Smith 2001): Aulacus Jurine, 1807, with 73 species,
Pristaulacus Kieffer, 1900, with 143 species, and Panaulix
Benoit, 1984, with two species. They are found in all
zoogeographic regions except Antarctica (Kieffer 1912; Hedicke
1939; Turrisi 2000, 2004, 2005, 2006, 2007; Smith 2001, 2005a,
2005b, 2008; He et al. 2002; Jennings et al. 2004a, 2004b, 2004c;
Jennings and Austin 2006; Sun and Sheng 2007a, 2007b, see
Appendix 3). In addition, numerous undescribed species have
been identified for the Australasian (>50, J. T. Jennings, unpubl.
data), Neotropical (~30, D. R. Smith, unpubl. data), and Oriental
regions (~20, G. F. Turrisi, unpubl. data).
The Aulacidae is also known from the fossil record, with 36
described species (Basibuyuk et al. 2002; Jennings et al. 2004d;
Nel et al. 2004; Zhang and Rasnitsyn 2004). The oldest
representatives are from the Lower Cretaceous, although the
identifications are controversial (Nel et al. 2004). The main
problem is the unclear relationships among the Baissinae,
Aulacidae and other Evanioidea because of the poor state of
preservation of the fossils (Nel et al. 2004; Zhang and Rasnitsyn
2004). However, Hyptiogastrites electrinus Cockerell, 1917, from
the Lower Cretaceous (Upper Albian), might represent the sister-
taxon to Aulacidae (Jennings et al. 2004d). According to Engel
(2006), this fossil species belongs to a distinct ancestral lineage of
the Aulacidae, the Hyptiogastritinae, characterised by the presence
of a distinct occipital carina, and the lack of forewing veins 1rs-m,
2rs-m, and 2m-cu, as well as the presence of a hind coxal
ovipositor guide (Jennings et al. 2004d; Engel 2006).
Half of the described aulacid fossil species are from the
Cenozoic, with records from the Upper Eocene of the Isle of
Wight, Baltic, and Paris basin amber, and from the Oligocene of
North America (Nel et al. 2004). Of these fossil aulacids, three are
included in the genus Aulacus,five in Pristaulacus (Brues 1910,
1923, 1932; Cockerell 1916, 1922; Nel et al. 2004), and 10 are
assigned to monospecific genera (see Nel et al. 2004).
Aulacidae are parasitoids of wood-boring Hymenoptera and
Coleoptera, employing a koinobiont endophagous strategy
(Skinner and Thompson 1960; Whitfield 1998; Jennings and
Austin 2004). Hosts are not known for most species, but some
Aulacus and a few Pristaulacus are associated with species of
Xiphydria Latreille (Xiphydriidae). Other species of Aulacus and
most Pristaulacus are associated with wood-boring Coleoptera,
"CSIRO 2009 10.1071/IS08031 1445-5226/09/010027
CSIRO PUBLISHING
www.publish.csiro.au/journals/is Invertebrate Systematics, 2009, 23, 27–59
especially Buprestidae and Cerambycidae (Barriga 1990;
Visitpanich 1994; Turrisi 1999; Jennings and Austin 2004).
Other coleopteran families have been recorded as hosts:
Bostrychidae, Cleridae (Oehlke 1983, 1984; Pagliano 1986), and
Scolytidae (Muesebeck 1958), but these records are regarded as
doubtful. Host data are often obtained in the laboratory, rearing
wasps directly from wood samples. However, since hosts are not
isolated, generally there is no clear knowledge of the true host/
parasitoid relationship. Most of the data on hosts are only inferred
and these taxa should be considered as ‘probable hosts’(Turrisi
2004, 2007). A summary of the data on hosts of the world’s
Aulacidae, as well as host plants, is provided by Smith (2001),
and a revised overview of hosts of Palaearctic Pristaulacus
species has been provided by Turrisi (2007). The biology is
known in some detail for Aulacus striatus Jurine, 1807, which is
associated with xylophagous larvae of Xiphydria camelus
(Linnaeus, 1758) (Skinner and Thompson 1960). The biology of
the two described species of Panaulix is unknown (Benoit 1984;
Madl 1990).
Our knowledge of the systematics, phylogeny, distribution
and biology of members of the Aulacidae is unsatisfactory due to
the fact that they are not easily observed in their natural habitats
and are only rarely collected by the most common collecting
methods. Members of the Aulacidae are relatively uncommon in
collections, and many species are known from only one or a few
specimens. The scarcity of available material, especially from
subtropical and tropical regions, is one of the most significant
limitations to improving the knowledge of the systematics of
these wasps.
The world aulacid fauna was first revised by Kieffer (1912).
However, the generic concepts were inadequate since they were
partly based on highly variable wing venation characters.
After the publication of Kieffer’s monograph, Hedicke (1939)
provided the first catalogue of world species, and more recently
Smith (2001) catalogued all Aulacidae including data on type
depositories, distribution, known hosts, and literature.
In the present contribution we present the first cladistic
analyses of the Aulacidae based on a representative sample of
species from all zoogeographic regions. The main aim of our
investigation was to test the monophyly of the three currently
recognised genera of Aulacidae in an attempt to improve the
current classification of the family involving strictly phylogenetic
concepts. Moreover, we also tested the monophyly of Aulacidae
and the relationship of this family to the other Evanioidea.
Status of the classification of Evanioidea
The three extant families of Evanioidea (Aulacidae, Evaniidae
and Gasteruptiidae) share the synapomorphy of having the
metasoma inserted dorsally on the propodeum (Crosskey 1951,
1962; Königsmann 1978; Rasnitsyn 1988). This definition of
Evanioidea has remained unchanged since Hedicke (1930).
However, at various times, the placement of Evaniidae with the
other two families has been questioned by numerous authors
(Townes 1950; Crosskey 1951; Gibson 1985; Gauld and
Hanson 1995; Gauld and Bolton 1996), who considered that
the high insertion of the metasoma might have been acquired
independently in the Evaniidae and Aulacidae+Gasteruptiidae.
Moreover, several other parasitoid Hymenoptera, for example, a
few specialised and distinctive groups within Ichneumonoidea,
Chalcidoidea and Cynipoidea, have independently acquired
this feature (Townes 1950; van Achterberg 1993; Basibuyuk
et al. 2000a, 2000b; G. F. Turrisi, unpubl. data), albeit
with morphological differences in the propodeal/metasomal
articulation.
Nevertheless, the metasomal insertion remains the most
convincing putative autapomorphy of the Evanioidea. The loss
of functional spiracles from all metasomal segments except
the eighth is sometimes cited as additional support for the
monophyly of the superfamily (e.g. Gauld and Bolton 1996).
This condition is, however, present in many parasitic wasps
(e.g. female Orussidae and Stephanidae (in which the males
have no metasomal spiracles at all), Megalyridae, Chalcidoidea,
Cynipoidea, and Proctotrupoidea (see Gibson 1986 and Ronquist
et al. 1999), and thus cannot be unambiguously upheld as a
potential evanioid autapomorphy.
Once fossil taxa are taken into consideration, the Evanioidea
become more difficult to define. In addition to the extant families,
one extinct family, the Praeaulacidae (Rasnitsyn 1972; Zhang
and Zhang 2000; Zhang and Rasnitsyn 2007, 2008), has been
placed within Evanioidea as a putative stem-group (Rasnitsyn
1988; Jennings et al. 2004d; Zhang and Rasnitsyn 2008);
however, the only reliable synapomorphy of this extinct family
with extant Evanioidea is the dorsal articulation of the metasoma
on the propodeum. Other basal lineages of extinct evanioid
wasps of uncertain systematic affinity include: the enigmatic
Andreneliidae (Rasnitsyn and Martìnez-Delclòs 2000; Deans
et al. 2004; Zhang and Rasnitsyn 2007) (currently placed
within Evaniidae by Engel (2006)) and the composite
Baissidae (inclusive of the speciose genus Manlaya Rasnitsyn
1980; Zhang and Rasnitsyn 2004) placed as sister to the
remaining Aulacidae (Nel et al. 2004), or as basal sister-group
of the clade Aulacidae+Gasteruptiidae (Engel 2006) or as a
subfamily of Gasteruptiidae s.l. (Zhang and Rasnitsyn, 2007).
Zhang and Rasnitsyn (2007) treat the Aulacidae as a subfamily
(Aulacinae) of Gasteruptiidae, following Rasnitsyn (1988); the
Kotujellidae (Rasnitsyn 1975), are currently placed within
Aulacidae (Rasnitsyn 1980).
Several features of comparative morphology seem to strongly
support a close relationship among the Aulacidae with the
Gasteruptiidae (Jennings and Austin 2000, 2004; Jennings
and Deans 2005): 1, they have a similar conformation of the
multiporous plate sensilla on the antenna, without an encircling
groove (Basibuyuk and Quicke 1999); 2, a rigid association of
the pronotum and the mesepisternum without an interlocking
mechanism or a prepectus (Gibson 1985, 1999); 3, the
mesothoracic spiracle is externally visible between the
posterolateral margin of the pronotum and the anterior margin of
the mesepisternum (Gibson 1985, 1999); 4, the mesofurcal/
mesotrochanteral depressor muscle originates as a conical or fan-
shaped muscle from each lateral mesofurcal arm (Gibson 1985,
1999); 5, they have a similar configuration of the antennal cleaner
(Basibuyuk and Quicke 1995); 6, only a few (2–3)distal hamuli on
the hind wing (Basibuyuk and Quicke 1997); 7, a partial fusion of
the first and second metasomal segments (Naumann 1991); 8, a
cylindrical ovipositor (clearly a symplesiomorphy), with the
olistheters slightly diverging ventrally and with a medial
thickening of the ventral wall of the upper valves (which may be
28 Invertebrate Systematics G. F. Turrisi et al.
Table 1. Characters and character states for the parasitoid wasp family Aulacidae
General
(1) Body length (excluding ovipositor): 0, <20 mm; 1, >20 mm.
Head
(2) Clypeus,medial process: 0, absent; 1, present: tooth-shaped; 2, present: lamelliform. Ordered (Fig. 1A).
(3) Anterior margin of head (dorsal view): 0, rounded; 1, angulated (Fig. 2A).
(4) Posterior margin of head (dorsal view): 0, straight or weakly concave; 1, deeply concave; 2, medially grooved. Ordered (Figs 2A, 3A,B).
(5) Occipital carina: 0, absent; 1, present (Figs 2A, 3C,D).
(6) Occipital carina configuration: 0, narrow (width <0.2!diameter of an ocellus); 1, wide (width at least 0.5!diameter of an ocellus) (Figs 3A, 3E).
(7) Occipital carina extension: 0, entire; 1, absent medially (Fig. 3E).
(8) Frons: 0, transverse–carinulate or rugulose; 1, smooth, at most punctate (Figs 1A,B, 3F).
(9) POL:OOL: 0, 0.6–0.8; 1, 0.9–1.9; 2, equal or >2.0. Ordered.
(10) Eye size and shape: 0, large and elliptical; 1, small and circular/subcircular (Fig. 3F).
(11) Subantennal grooves: 0, absent; 1, weak; 2, well defined. Ordered (Fig. 1A,B).
(12) Occipital margin: 0, sculptured (usually rugose or carinulate); 1, smooth (Fig. 3C,D).
(13) Mandibles: 0, not broadly overlapping; 1, broadly overlapping (Fig. 1B).
(14) Antennal insertion relative to eye: 0, about level with base of eyes or slightly below (Fig. 1A,B); 1, about half-way up eyes.
(15) A1 shape: 0, stocky or moderately elongate (length <2.0!width); 1, rather elongate (length ~3.0!width) (Figs 1A, 2B).
(16) A1 in lateral view: 0, medial part swollen, convex; 1, median part not swollen, parallel-sided (Figs 1A, 2B).
(17) Antennomere number (female): 0, 14 segments or more; 1, 13 segments.
(18) Antennomere sculpturing: 0, not carinate; 1, with a weak ventral carina (Fig. 2B).
(19) Apex of last antennomere: 0, acute; 1, rounded.
(20) Shape of last antennomere: 0, subcylindrical; 1, flattened.
Mesosoma (including appendages)
(21) Mesosomal setae,length: 0, short (less than 1!diameter of an ocellus); 1, long (more than 1!diameter of an ocellus).
(22) Mesosomal setae,curvature: 0, erect; 1, recumbent.
(23) Mesosomal setae,density: 0, scattered (cuticle visible beneath); 1, dense (cuticle not visible).
(24) Pronotal spiracle: 0, visible, located in a notch in the posterolateral margin of pronotum; 1, not visible, covered by the posterolateral margin
of pronotum (Fig. 6A–C).
(25) Pronotum length: 0, long dorsal to transverse sulcus; 1, short dorsal to transverse sulcus (Fig. 7A,B).
(26) Lateroventral margin of pronotum: 0, rounded; 1, anteriorly angulated (Figs 5A,C, 6A,B).
(27) Pronotal lateral processes: 0, absent; 1, present (Fig. 5D).
(28) Pronotum,anterior tooth-like process: 0, absent; 1, present (Fig. 5A,D).
(29) Pronotum,posterior tooth-like process: 0, absent; 1, present (Fig. 5D).
(30) Pronotum,lateral view: 0, short (height/length >1.0, at most as long as high); 1, long (height/length <1.0).
(31) Propleura,dorsal margins: 0, widely separated for most of their length; 1, abutting for at least half their length (Fig. 7C,D).
(32) Propleura,posterolateral corners: 0, not extended; 1, extended, covering fore coxae proximally (Fig. 7E,F).
(33) Propleura,sculpture: 0, sculptured; 1, smooth, at most punctate.
(34) Premesoscutum,delimitation: 0, anteroadmedian lines not convergent posterodorsally; 1, anteroadmedian lines convergent posterodorsally,
delimiting premesoscutum (Fig. 4A,B).
(35) Premesoscutum,concavity: 0, not concave; 1, concave (Fig. 4B).
(36) Premesoscutum,sculpture: 0, sculptured; 1, smooth (Fig. 4B).
(37) Mesoscutum,surface: 0, smooth, at most punctate; 1, weakly sculptured; 2, strongly sculptured. Ordered (Figs 5D, 6B).
(38) Mesoscutum,sculpture: 0, areolate–rugose or other than transverse–carinate; 1, transverse–carinate (Fig. 5A).
(39) Mesonotum,longitudinal subdivision of scutellum: 0, not subdivided; 1, subdivided by submedian longitudinal grooves (Fig. 8A,B).
(40) Mesonotum,notauli configuration: 0, subparallel or convergent, not joining before reaching transscutal fissure; 1, convergent medially, joining before
reaching transscutal fissure; 2, forming an U-shaped sulcus, not reaching transscutal fissure. Ordered (Figs 4C, 9A).
(41) Mesoscutum,antero-lateral processes: 0, absent; 1, present (Figs 5C, 9A).
(42) Mesoscutum,anterior margin (lateral view): 0, rounded; 1, acute; 2, acute, with upward directed process. Ordered (Figs 5A,C,D, 6C).
(43) Parascutal carina: 0, not expanded; 1, expanded; 2, expanded, with tooth-like lateral projection. Ordered (Fig. 5B,E).
(44) Metapleuron,articulation notch: 0, absent or weakly developed; 1, notch in foraminal rim at metacoxal lateral articulation well developed (Fig. 8C,D).
(45) Fore coxa: 0, sculptured; 1, smooth, at most with scattered punctures.
(46) Mid leg,coxo-trochanter articulation: 0, broad, trochanter cylindrical proximally; 1, narrow, trochanter with flattened disk proximally (Fig. 8E,F).
(47) Hind coxa,sculpture: 0, sculptured; 1, smooth, at most punctate (Fig. 10C).
(48) Hind coxal groove: 0, absent; 1, present (Fig. 10A,B).
(49) Hind coxal groove,orientation: 0, longitudinal; 1, transverse (Fig. 10A–C).
(50) Hind coxal transverse groove: 0, not complete; 1, complete (Fig. 10C).
(51) Hind coxal distal lobe: 0, absent; 1, present (Fig. 10A,B).
(52) Prefemur (trochantellus)on hind leg: 0, present; 1, absent.
(53) Hind tibia: 0, not flattened; 1, flattened.
(54) Hind tibia,shape: 0, subcylindrical, not enlarged; 1, moderately enlarged; 2, strongly enlarged. Ordered (Fig. 14).
(continued next page )
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 29
a symplesiomorphy) (Quicke et al. 1994); 9, they have a blocking
mechanism on the dorsal and ventral valves of the ovipositor (near
the tip of the valves) that allows steering of the ovipositor during
oviposition (Quicke and Fitton 1995), and 10, two large submedial
third phragmal lobes (Whitfield et al. 1989). In each of these
features, Aulacidae+Gasteruptiidae differ significantly from the
Evaniidae, and some of them (2,7, and 9) might be genuine
synapomorphies.
Recent studies on Apocritan phylogeny have been based on
morphological data (Rasnitsyn 1999, 2002; Sharkey and Roy
2002; Sharkey 2007), molecular data (Dowton and Austin 1994,
2001; Dowton et al. 1997; Castro and Dowton 2006), or a
combination of both (Carpenter and Wheeler 1999; Dowton
and Austin 2001), but these have not resolved the relationships
within Evanioidea, or the placement of Evanioidea in relation to
other Apocritan clades. Some molecular (Dowton and Austin
1994, 2001; Dowton et al. 1997) and morphological analyses
(Ronquist et al. 1999: fig. 6) support a sister-group relationship
between the Evaniidae and Gasteruptiidae. However, several of
these studies did not include the Aulacidae in their taxon sample
(Dowton and Austin 1994, 2001; Dowton et al. 1997).
The current classification of the family Aulacidae
The Aulacidae was first proposed as a family name by Shuckard
(1841), although he included some genera currently placed in the
family Trigonalidae. There are different opinions regarding the
systematic rank of aulacid wasps, with most authors regarding
them as a distinct family (Konishi 1990; Mason 1993; Gauld and
Hanson 1995; Jennings and Austin 2000, 2004; Smith 2001;
Turrisi 2004), although others consider them as a subfamily of
Evaniidae (Kieffer 1912) or Gasteruptiidae (Townes 1950;
Rasnitsyn 1988; Whitfield et al. 1989; Zhang and Rasnitsyn
2007). Jennings and Austin (2000) found strong support
for the monophyly of the Aulacidae, providing the first
phylogenetic framework.
Several genera have been recognised within the Aulacidae at
one time or another (for a summary, see Kieffer 1912; Hedicke
1939; Smith 2001). Currently, all species are assigned to only
three genera (Smith 2001). This classification reflects the opinion
of Townes (1950), who considered several previously proposed
genera as unnatural. He briefly discussed the main characters for
generic assessment of the group, including the presence/absence
of the occipital carina, wing venation, the number of teeth on the
tarsal claws, and the absence/presence of tooth/teeth along the
lateroventral margin of the pronotum. He considered wing
venation characters too variable among species, and not valid
for generic recognition. Other characters used by Kieffer (1912),
although stable within species, were considered insufficient to
subdivide the family into natural genera. Thus, he recognised only
two genera as valid (at that time Panaulix was unknown), namely
Aulacus and Aulacostethus (=Pristaulacus) (Townes 1950).
Pristaulacus was characterised by the presence of an occipital
carina, the presence of two or more distinct tooth-like processes
on the tarsal claws, and having the ovipositor guiding groove on
Table 1. (continued )
(55) Claws: 0, one small basal tooth-like process; 1, one apical/medial tooth-like process; 2, two well spaced tooth-like processes; 3, 3–6 tooth-like processes
(pectinate claws). Ordered (Fig. 11A–D).
(56) Forewing folding at rest: 0, not folded; 1, folded longitudinally.
(57) Forewing vein 2r-m: 0, absent; 1, present mostly spectral, recognisable only posteriorly; 2, tubular or mostly tubular. Ordered (Fig. 12A,B).
(58) Forewing vein 3r-m: 0, absent; 1, mostly spectral; 2, mostly tubular. Ordered (Fig. 12A,B).
(59) Forewing vein 2 m-cu: 0, absent; 1, present (Fig. 12A).
(60) Forewing vein 2Rs+M: 0, long; 1, short or absent (Fig. 12A,B).
(61) Propodeum,propodeal foramen anterior incurvation: 0, absent; 1, present (Fig. 8C).
(62) Propodeum,propodeal foramen shelf: 0, absent; 1, distinct shelf extending anteriorly into body cavity from foramen (Fig. 9C,D).
(63) Propodeum,position of propodeal foramen: 0, situated at level with metacoxal foramina; 1, approx. halfway between metacoxal foramina and
metapostnotum; 2, adjacent to antecostal sulcus, compressing sulcus medially. Ordered (Figs 8D, 9B).
Metasoma
(64) Metasoma,shape (lateral view): 0, ovoidal; 1, pyriform; 2, subcylindrical; 3, subcircular (Fig. 18A–D).
(65) Petiole,T2 and S2 configuration: 0, separate; 1, fused, at least anteriorly (Fig. 15A,B).
(66) Petiole,transverse carina anteriorly on S2: 0, absent; 1, present (as a transverse carina posterior to sensillar patches) (Fig. 15C,D).
(67) Petiole,shape: 0, stocky (length/width: 1.0–1.1); 1, slender (length/width: 2.0–4.5 or more) (Figs 16A,B, 18B,C).
(68) Petiole base: 0, sculptured; 1, smooth (Fig. 16A,B).
(69) T2–T5,punctures and setae: 0, absent; 1, present.
(70) T8: 0, striolate; 1, not striolate.
(71) Subgenital plate: 0, not notched; 1, notched (Fig. 17A,B).
(72) Ovipositor exsertion: 0, completely exserted; 1, not exserted, hidden by ovipositor sheaths; 2, completely internal. Ordered (Fig. 18A–C).
(73) Ovipositor length/forewing length: 0, <0.3; 1, 0.5–0.9; 2, 1.0–2.0; 3, >2.0. Ordered.
Colour
(74) Head: 0, black; 1, yellow, orange or red; 2, bicoloured.
(75) Pronotum: 0, black; 1, yellow, orange or red; 2, bicoloured.
(76) Mesoscutum: 0, black; 1, yellow, orange or red; 2, bicoloured.
(77) Forewing: 0, hyaline, without dark spots; 1, at least with one dark spot (usually at apex of wing); 2, largely infuscated; 3, broad stripe of across fore margin
(Figs 12A,C, 13).
(78) Hind tarsus: 0, white or cream; 1, yellow, orange or red; 2, brown or black; 3, bicoloured.
(79) Ovipositor valvula 3: 0, brown or black; 1, broadly ivory before tip.
30 Invertebrate Systematics G. F. Turrisi et al.
the hind coxa oriented transversely; Aulacus would, by
default, be those taxa that lack the two former traits and have
the hind coxal groove (when present) oriented longitudinally
or obliquely. Since Townes (1950), the only other genus to
be described in Aulacidae is Panaulix, comprising two
Afrotropical species (Benoit 1984; Madl 1990). Panaulix was
considered similar to Pristaulacus from which it was separated
by the shape of the head, in having pronounced malar area and
mandibles, the shape of the petiole and a very long ovipositor
(Benoit 1984). The generic concepts of Townes (1950) were
retained in the catalogue of Smith (2001).
With the exception of a preliminary cladistic analysis by
Turrisi (2004), there has been no attempt to test the generic
classification of Aulacidae. However, the recent description of
new species, especially from previously unstudied areas such as
New Guinea, New Caledonia (Australasia) and the Neotropics,
have convinced several authors about the inadequateness of this
classification. Jennings and Deans (2005) argued that there is little
morphological evidence that the two main genera of Aulacidae
(Aulacus and Pristaulacus) are monophyletic. Based on this
assumption, the current classification of Aulacidae is largely
inadequate and in need of revision based on cladistic analyses.
Materials and methods
Material examined
This study was based on examination of 72 species of Aulacidae, representing
~30% of the described extant species (Smith 2001; see Appendix 3). The
material examined belongs to many collections, listed below; the acronyms of
the museums are according to Evenhuis and Samuelson (2004).
Museums
AEIC American Entomological Institute, Gainesville,
Florida, USA.
ANIC Australian National Insect Collection, Canberra,
Australia.
BMNH The Natural History Museum, London, UK.
BPBM Bernice P. Bishop Museum, Honolulu, Hawaii, USA.
CAS California Academy of Sciences, San Francisco,
California, USA.
CNCI Canadian National Collection of Insects and
Arachnids, Ottawa, Ontario, Canada.
DBAC Dipartimento di Biologia Animale ‘Marcello La
Greca’, Università di Catania, Museo Zoologico,
‘Turrisi G.F. Collection’, Italy.
Fig. 1. Head, frontal view of Aulacidae. (A)Aulacus striatus, (B)
Pristaulacus gibbator.
Fig. 2. Head of Aulacidae. (A) Head, dorsal view of Aulacus striatus;
(B) head, frontal view of Pristaulacus rex.
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 31
DEI Deutsches Entomologisches Institut, Müncheberg,
Germany.
HNHM Hungarian Natural History Museum, Budapest,
Hungary.
IBLP Instytut Badawczy Lesnictwa, Warszawa, Poland.
ITLJ National Institute for Agro-Environmental Sciences,
Insect Systematic Laboratory, Tsukuba (Ibaraki),
Japan.
LACM Los Angeles County Museum of Natural History,
Los Angeles, California, USA.
MCFS Museo Civico di Storia Naturale, Ferrara, Italy.
MCNC Museo de Ciencias Naturales, Canary Islands:
Tenerife, Spain.
MCSN Museo Civico di Storia Naturale ‘G. Doria’, Genova,
Italy.
MFNB Museo Friulano di Storia Naturale, Udine, Italy.
MHNG Muséum d’Histoire Naturelle de la Ville de Genève,
Switzerland.
MNHN Muséum National d’Histoire Naturelle, Laboratoire
d’Entomologie, Paris, France.
MNMS Museo Nacional de Ciencias Naturales, Madrid,
Spain.
Fig. 3. Head of Aulacidae. (A) Head, dorsal view of Pristaulacus gibbator; (B) head, dorsal view of Pristaulacus comptipennis;
(C) head, posterolateral view of Aulacus striatus; (D) head, posterolateral view of Pristaulacus compressus; (E) head, dorsal view of
Pristaulacus compressus; (F) head, frontolateral view of Pristaulacus compressus.
32 Invertebrate Systematics G. F. Turrisi et al.
MRAC Musée Royal de l’Afrique Centrale, Tervuren,
Belgium.
MSNP Museo Civico di Storia Naturale di Calci, Pisa, Italy.
MRSN Museo Regionale di Storia Naturale, Torino, Italy.
MV Museum Victoria, Melbourne, Australia.
MZLU Museum of Zoology, Lund University, Lund,
Sweden.
NMW Naturhistorisches Museum, Wien, Austria.
OLML Oberösterreichisches Landesmuseum, Linz, Austria.
SAM South Australian Museum, Adelaide, Australia.
Fig. 4. Mesosoma of (A–B) Aulacidae and (C) Gasteruptiidae. (A) Anterior part of mesoscutum, dorsal view of
Aulacus striatus;(B)anterior part of mesoscutum, dorsal view of Pristaulacus africanus; (C) mesoscutum, dorsal
view of Gasteruption sp. Abbreviations: ms, mesoscutum; pms, premesoscutum; nt, notaulus; tsf, transscutal
fissure.
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 33
SAMC South African Museum, Cape Town, Republic of
South Africa.
USNM National Museum of Natural History, Smithsonian
Institution, Washington DC, USA.
WADA Western Australian Department of Agriculture Insect
Collection, Perth, Australia.
WAM Western Australian Museum, Perth, Australia.
WINC Waite Insect and Nematode Collection,
The University of Adelaide, Australia.
ZFMK Zoologisches Forschungsinsitut und Museum
A. Koenig, Bonn, Germany.
ZIN Zoological Institute of the Russian Academy of
Sciences, St. Petersburg, Russia.
ZMHB Museum für Naturkunde der Humboldt-Universität,
Berlin, Germany.
ZMUC Zoological Museum, Copenhagen University,
Denmark.
ZSMC Zoologische Staatssammlung München, Germany.
Private collections
CPTO Guido Pagliano collection, Torino, Italy.
GGCP Gianluca Governatori collection, Pordenone, Italy.
Fig. 5. Mesosoma of Aulacidae. (A) Mesosoma, lateral view of Pristaulacus kostylevi; (B) mesosoma,left half, dorsal view of Aulacus
striatus; (C) mesosoma, laterodorsal view of Pristaulacus ryukyuensis; (D) part of mesosoma, lateral view of Pristaulacus compressus;
(E) part of mesoscutum, dorsal view of Pristaulacus compressus. Abbreviation: ms, mesoscutum.
34 Invertebrate Systematics G. F. Turrisi et al.
GSCC Gianfranco Sama collection, Cesena, Italy.
STCC Salvatore Tomarchio collection, San Gregorio di
Catania, Catania, Italy.
PRCU Pierpaolo Rapuzzi collection, Udine, Italy.
Methods for examination
Specimens were studied using stereomicroscopy and scanning
electron microscopy (SEM). Measurements were taken with the
aid of an ocular scale. Scanning electron micrographs were
produced with a Philips XL-20 (http://www.fei.com/). Some
pinned and air-dried specimens were fixed with Leit-C-plast
on an object table and observed at 1.6 kV using a special low
voltage anode (spot size 4–5); other specimens were previously
sputtered with a Polaron SEM coating system and observed at
10 kV using a conventional high voltage anode (spot size 3–4).
High resolution images were taken using light
stereomicroscopy with a Wild M5A (Wild Heerbrugg,
Switzerland) using a Nikon Coolpix 4300, 4.0 megapixel
digital camera (Nikon, Tokyo). Final illustrations were
integrated from a series of images with different focal planes,
using the freeware CombineZM (Hadley 2008). The final
illustrations were retouched using Adobe Photoshop CS2
software in order to enhance clarity; photographic tables were
assembled using CorelDraw X3.
Morphological terminology and abbreviations
The nomenclature for morphology follows Crosskey (1951),
Huber and Sharkey (1993), and Gauld and Bolton (1996).
Terminology for surface sculpture follows Harris (1979).
Regarding the number of tooth-like processes on the inner
margin of the claw, the apex is not included, since, in a strictly
morphological sense, it represents the tip of the claw (Turrisi
2007).
In the text, the following abbreviations are used for some
morphological structures: A, antennomere; OOL, distance
between posterior ocellus and eye; POL, distance between
posterior ocelli; T, tergite, and S, sternite.
Selection of taxa
The ingroup species were chosen to represent all the three
currently recognised genera of the Aulacidae, as well as a
diverse sampling from all zoogeographic regions (Appendix 1):
25 Aulacus (11 Australasian, six Neotropical, five Nearctic,
three Palaearctic); 45 Pristaulacus (eight Australasian, six
Neotropical, seven Oriental, four Afrotropical, 10 Nearctic,
10 Palaearctic), and both species of Panaulix (Afrotropical).
The outgroups include the two other recognised families of the
Evanioidea: Evaniidae (Brachygaster minuta (Olivier, 1792),
Evania appendigaster (Linnaeus, 1758), Szepligetella sp., and
Zeuxevania splendidula (Costa, 1884)) and Gasteruptiidae
(Gasteruption sp. 1 and 2, Hyptiogaster pinjarregaensis
Jennings and Austin, 1997, and Pseudofoenus crosskeyi
Jennings and Austin 2002). Also, members of four additional
families: Megalyridae (Megalyra fasciipennis Westwood, 1832)
and Trigonalidae (Orthogonalys pulchella (Cresson, 1867))
(which have been regarded as closely related to Evanioidea
Fig. 6. Mesosoma of (A–B) Aulacidae and (C) Gasteruptiidae. (A) Pronotum, lateral view of Pristaulacus strangaliae;
(B) mesosoma, lateral view of Aulacus striatus; (C) pronotum and mesoscutum, lateral view of Gasteruption sp.
Abbreviations: ms, mesoscutum; sp, spiracle; pt, pronotum.
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 35
(all are members of the clade Evaniomorpha as defined by
Rasnitsyn 1988)), Braconidae (Doryctes erythromelas (Brullé,
1846)), which is a parasite of woodboring insects, and
Stephanus serrator (Fabricius, 1798), a representative of the
putatively basal apocritan lineage Stephanidae (Vilhelmsen
1997a, 2001, 2003; Vilhelmsen et al. in press) are included
as outgroups.
Selection of characters
Morphological examination was restricted to external characters.
Examination of internal characters would have been difficult or
impossible, as many of the taxa included were only represented by
the type specimen, thus making dissection undesirable.
Characters and characters states
For the phylogenetic analyses, 79 characters were selected,
consisting of 183 character states; 58 characters were coded as
binary, 16 characters were coded with three states, five characters
had four states. Continuous characters were divided into discrete
binary or ternary states to minimise hierarchical linkage and the
problem of scoring non-applicable states (see Jennings and
Austin 2002).
All images showing character states reported in the present
paper have been deposited in the open web archive ‘Morphbank’
(http://www.morphbank.net) covering the numbers 465 131–
465 187, to give free access for future comparisons.
The characters and character states are listed in Table 1.
Fig. 7. Mesosoma of (A,C,E) Evaniidae: Evaniella semaeoda and (B,D,F) Gasteruptiidae: Pseudofoenus sp. (A,B) Pronotum, dorsal
view; (C,D) propleura, dorsal view; (E,F) posterolateral corner of propleura. Abbreviations: pt, pronotum; sc, sulcus; pl, propleuron;
cx1, fore coxa.
36 Invertebrate Systematics G. F. Turrisi et al.
Phylogenetic analyses
The character matrix for cladistic analyses (see Appendix 1) was
constructed and edited using Mesquite version 2.01 (Maddison
and Maddison 2007). Unknown character states were coded with
‘?’, and inapplicable characters were coded with ‘–’. The coded
data were exported in a TNT spreadsheet format before
performing parsimony analyses using TNT software (Goloboff
et al. 2003). All characters were given equal weight; multistate
characters were treated as additive only if the recognised
states appeared to form a linear transformation series based on
similarity. Characters were coded mainly from females, although
comparisons with males of the same species, when known
and/or available, were made to distinguish sexually dimorphic
characters.
A‘traditional search’option was instituted to find the most
parsimonious trees, using a memory set to hold 1 000 000 trees
and 1000 replicates with tree-bisection-reconnection (TBR)
branch swapping and saving 1000 trees per replicate. Separate
analyses were conducted with the same setting options but using
implied weights (Goloboff et al. 2003) with k-values from 1 to 20;
this was done in order to explore the variation in tree topology
over a range of different weighting conditions. Trees were rooted
on Stephanus. Character changes and a list of apomorphies for
each node were produced using TNT. Node support was assessed
by calculating Bremer support values (Bremer 1994) in TNT by
searching for suboptimal trees using the trees obtained by the
equal weights analyses as starting point. Suboptimal trees of up to
20 steps longer than the shortest trees were looked for, the search
Fig. 8. Mesosoma of (A) Aulacidae, (B,D,E) Gasteruptiidae and (C,F) Evaniidae. (A)Pristaulacus strangaliae, (B,D)Gasteruption
sp., (C,F)Evaniella semaeoda, (E)Pseudofoenus sp. (A,B) Mesoscutellum, dorsal view; (C,D) propodeum, dorsal view; (E,F) coxo-
trochanter articulation. Abbreviations: sc, scutellum; pm, propodeum; pmf, propodeum foramen; mtf, metacoxal foramen; cx, coxa.
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 37
continuing until the number of trees checked reached the value of
100 000. Absolute Jacknife support was computed in TNT using
default settings and running 10 000 replications.
Results of cladistic analyses
Equal weighting
Analysis of the data matrix (Appendix 1), using unweighted and
some additive character options, resulted in 18 000 equally
parsimonious trees producing the strict consensus tree
presented in Fig. 19. The shortest trees are 593 steps long and
were retrieved in 443 replications out of 1000. Bremer and
Jacknife support values are displayed on Fig. 19. The
Aulacidae were retrieved in the equal weights analysis, but
there was little resolution within the family, and both Aulacus
and Pristaulacus were paraphyletic.
Implied weighting
The analysis under implied weights options, using k-values
varying from 1 to 20, produced one to four trees for each
k-value. The rate of retrieval of the shortest tree(s) was
extremely low for the implied weights analyses, between one
and 24 out of 1000 replications. The topology of the outgroup
taxa varied little among the trees retrieved with different
k-values, except the placement of D. erythromelas. In contrast,
there was some variation in the topology of the Aulacidae,
especially among the basal lineages within the family. The
Aulacidae were retrieved in all the implied weights analyses,
Fig. 9. Mesosoma of Aulacidae (A–C) and Evaniidae (D). (A)Pristaulacus gibbator, (B,C)Pristaulacus strangaliae, (D)Evaniella
semaeoda. (A) Mesoscutum, dorsal view; (B) metapostnotum and propodeum, dorsal view; (C,D) propodeal foramen. Abbreviations: nt,
notaulus; tsf, transscutal fissure; mtp, metapostnotum; pm, propodeum; pmf, propodeum foramen; mtf, metacoxal foramen.
38 Invertebrate Systematics G. F. Turrisi et al.
and always well resolved, but Aulacus and Pristaulacus were
still paraphyletic. The analyses with k-values between 9 and
11 retrieved very similar results, the tree obtained with a
k-value of nine being congruent with the consensus trees
produced by analyses with k = 10 or 11. The k = 9 tree
(Fig. 20) was chosen for character mapping and discussion.
Fig. 10. Hind coxae of Aulacidae. (A) Hind coxae, ventral view of Aulacus emineo; (B) hind coxae, ventral view of Pristaulacus
gibbator; (C) right hind coxa, inner view of Pristaulacus compressus. Abbreviations: cx2, mid coxa; cx3, hind coxa.
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 39
Discussion
In the following discussion, unequivocal character state changes
are displayed in parantheses, i.e. (character): (state), with
reference to Figs 19 and 20.
Monophyly of Aulacidae and relationships
with other Evanioidea
Aulacidae is consistently retrieved as monophyletic under all
parsimony-based settings (Figs 19 and 20). Only one putative
autapomorphy for Aulacidae was identified (node 4, Fig. 20):
mesoscutum transverse to carinate, with a few scattered reversals
within Aulacidae (38: 1). Two characters might be retained
plesiomorphies within Aulacidae: 1, forewing vein 3r-m
present (tubular or mostly spectral) (58: 1, 2); 2, forewing vein
2 m-cu present (59: 1). In addition, the presence of a hind coxal
groove (48: 1) represents a synapomorphy for most Aulacidae
except the basalmost species Aulacus wau and Aulacus sp. 1.
Jennings and Austin (2000) suggested the following putative
autapomorphies for the Aulacidae: 1, antennae inserted level with
base of eyes (character 14 in the present analysis); 2, antennal
sockets widely separated (not included in the present analysis);
3, eye small and circular/subcircular (character 10 in the present
analysis);4, notauli percurrent, Y-shaped (perhaps corresponding
to character 40 in the present analysis); 5, propleuron not elongate
(not included in the present analysis, perhaps corresponding to
character 31; according to the scoring presented in Jennings and
Austin 2000 (appendix 2, character 30), this character should
not optimise as an aulacid autapomorphy on their cladogram);
6, propodeum not pyramidial (not included in the present
analysis, perhaps corresponding to character 63); 7, presence
of a groove on the inner surface of the hind coxae of females
(character 48 in the present analysis). In the present study, only
one of these autapomorphies is confirmed; two have not been
confirmed (1,3), whereas the remaining character states have not
been included in the present analyses or have been coded
Fig. 11. Claws of Aulacidae. (A)Aulacus striatus, (B)Pristaulacus kostylevi, (C)Pristaulacus gibbator,
(D)Pristaulacus chlapowskii.
Fig. 12. Wings of Aulacidae. (A)Pristaulacus compressus, (B)Aulacus
krahmeri, (C)Aulacus braconiformis.
Fig. 13. Wings of Aulacidae: Pristaulacus gibbator.
Fig. 14. Hind tibia and tarsus of Gasteruptiidae: Gasteruption sp.
40 Invertebrate Systematics G. F. Turrisi et al.
differently and thus not properly tested. We are unable to confirm
any of the autapomorphies for Aulacidae suggested by Jennings
and Austin (2000). This might be partly due to differences in the
taxon sample (Jennings and Austin (2000) included only three
exemplars each of Aulacus and Pristaulacus, as their main focus
was on Gasteruptiidae), or to differences in coding and scoring
characters (see above).
The results of the cladistic analyses corroborate the placement
of the Gasteruptiidae as the sister-group of the Aulacidae,
which was previously based on only a few morphological
similarities in some pre-cladistic studies (e.g. Townes
1950; Crosskey 1962). Aulacidae shares the following
synapomorphies with Gasteruptiidae (node 2, Fig. 20):
1, dorsal margins of propleura abutting for at least half their
length (this character state was previously recognised as the
so-called ‘neck’, see Crosskey 1951) (31: 1); 2, metapleuron
with notch in foraminal rim at metacoxal lateral articulation well
developed (44: 1); 3, propodeum with propodeal foramen
anteriorly incurved (61: 1). The monophyly of Gasteruptiidae
is supported by three putative autapomorphies (node 3): 1, notauli
configuration on mesoscutum, forming an U-shaped sulcus, not
reaching transscutal fissure (40: 2); 2, forewings folded at rest
Fig. 15. Basal part of metasoma of (A,C) Aulacidae, (B) Gasteruptiidae and (D) Evaniidae. (A) Petiole, lateral view of Aulacus striatus;
(B) petiole, lateral view of Pseudofoenus sp.; (C) S2 of Pristaulacus strangaliae; (D) S2 of Evaniella semaeoda.
Fig. 16. Propodeum and petiole lateral view of Aulacidae. (A)Aulacus striatus, (B)Pristaulacus compressus. Abbreviations: pm,
propodeum; pe, petiole; cx3, hind coxa.
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 41
(56: 1); 3, propodeal foramen adjacent to antecostal sulcus,
compressing it medially (63: 2).
Several striking autapomorphies were identifed for Evaniidae
(node 1): 1, 13 antennomeres in females (17: 1); 2, mesosomal
setae dense (although a similar condition is known for some
Gasteruptiidae and Aulacidae) (23: 1); 3, coxo-trochanter
articulation of midlegs narrow, trochanter with flattened disk
proximally (46: 1); 4, propodeal foramen with distinct shelf
extending anteriorly into body cavity from foramen (62: 1);
5, petiole with transverse carina on S2, posterior to sensillar
patches (66: 1).
The status of Evanioidea as monophyletic, and thus as a valid
superfamily, still remains a matter of discussion. We did not
retrieve the Evanioidea as monophyletic. Trigonalidae were
always placed as sister to the Evaniidae, and Megalyra to
Aulacidae + Gasteruptiidae. Our taxon and character samples
are probably not sufficient to reliably resolve the relationships
between the outgroup taxa. However, our analyses show a sister
relationship of Gasteruptiidae with Aulacidae, confirming
previous morphological evidence (Crosskey 1951, 1962;
Naumann 1991; Quicke et al. 1994; Gauld and Bolton 1996).
Remarks on the genus-level phylogeny
and classification of Aulacidae
Ideally, a revised generic classification of the Aulacidae should
recognise only monophyletic genera. Of the three previously
recognised genera, only Panaulix is confirmed as monophyletic.
Pristaulacus is rendered paraphyletic by Panaulix, and Aulacus
comprises several lineages, many of them monotypic, which are
placed as a basal grade within the family.
Panaulix forms a strongly supported clade deeply nested
within Pristaulacus. The clade comprising Pristaulacus and
Panaulix is strongly supported by six unambiguous
autapomorphies (node 6): 1, prothoracic spiracle visible,
located in a notch in the posterolateral margin of pronotum
(24: 0); 2, mesoscutum with notauli convergent medially to
join prior reaching transscutal fissure (40: 1); 3, parascutal
carina expanded to partially cover the tegula, with or without
tooth-like lateral projection (43: 1–2); 4, claws pectinate, with
two well spaced or three to six tooth-like processes along inner
margin (55: 2–3); 5, forewing vein 2r-m absent (57: 0) (albeit
with some reversals); 6, forewing vein 2 Rs+M short (60: 1)
(with some reversals).
As mentioned in the Introduction, three characters have
primarily been used to diagnose genera in the Aulacidae
(cfr. Townes 1950; Oehlke 1983, 1984; Pagliano 1986; Konishi
1990; Smith 1996, 2005a, 2005b). We have traced them onto the
preferred topology (implied weights, k=9) to evaluate their
usefulness in defining generic boundaries within the family. The
presenceof theoccipitalcarina(5:1) hastraditionallybeenregarded
as diagnostic for Pristaulacus. However, at least Pristaulacus
ornatus has no occipital carina, and Aulacus grossi has the
carina present (Fig. 21A). Furthermore, it is evident that the
absence of the carina is a ground plan feature of the Aulacidae,
not just Aulacus. The orientation of the hind coxal groove is even
more variable (Fig. 21B). Some Aulacus spp. have no groove at
all, and less than half the species examined here have the groove
oriented longitudinally (49: 0) –the supposed diagnostic
feature of the genus. Indeed, the longitudinal orientation seems
to have been derived two times independently, the transverse
orientation (49: 1) being the ground plan feature of the family.
Finally, the configuration of the tarsal claws has been regarded as
important, with two or more tooth-like processes being diagnostic
forPristaulacus(55:2–3).Thisisupheldbyour analysis(Fig.21C),
even if the exact number of tooth-like processes within the
genus varies. However, having only one small basal tooth-like
Fig. 17. Female metasoma of Gasteruptiidae, ventral view, showing subgenital plate (sgp). (A)
Pseudofoenus sp., (B)Gasteruption sp.
42 Invertebrate Systematics G. F. Turrisi et al.
process (55: 0) is a plesiomorphic feature and does not support
the monophyly of Aulacus. In conclusion, the traditional
diagnostic characters used to separate Aulacus and Pristaulacus
at best provides support for only the latter, at worst for neither
genus.
Relationships within ‘Aulacus’Jurine
The genus Aulacus as previously recognised is clearly not
monophyletic, and the relationships at the base of the Aulacidae
clade are poorly resolved. However, ‘Aulacus’includes some
monophyletic groups that might be recognised in a revised
classification should they prove robust in further testing.
The occurrence and configuration of the hind coxal groove
is highly variable within ‘Aulacus’(see above). The presence of
a longitudinal groove, usually in conjunction with the presence
of a distal lobe on the hind coxa, is widespread. This character
state (49: 0) apparently represents a synapomorphic feature for a
very distinctive Holoarctic group of ‘Aulacus’spp. (including
Fig. 18. Female metasoma, lateral view of (A) Evaniidae, (B,C) Aulacidae and (D) Gasteruptiidae. (A)Zeuxevania splendidula,
(B)Aulacus japonicus, (C)Pristaulacus sp., (D)Gasteruption sp. Abbreviations: pm, propodeum; pe, petiole; me, metasoma;
ov, ovipositor.
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 43
Stephanus serrator
Doryctes erythromelas
Hyptiogaster pinjarregaensis
Aulacus braconiformis
Aulacus burquei
Aulacus grossi
Pristaulacus africanus
Pristaulacus com
p
ressus
Orthogonalys pulchella
Megalyra fasciipennis
Szepligetella
Zeuxevania splendidula
Brachygaster minuta
Evania appendigaster
Gasteruption
Gasteruption
Pseudofoenus crosskeyi
Aulacus
Aulacus
Aulacus wau
Aulacus sedlaceki
Aulacus impolitus
Aulacus elegans
Aulacus brevicaudus
Aulacus
Aulacus
Aulacus krahmeri
Aulacus emineo
Aulacus burwelli
Aulacus houstoni
Aulacus douglasi
Aulacus belairensis
Aulacus enaratodi
Aulacus coracinus
Aulacus striatus
Aulacus schoenitzeri
Aulacus japonicus
Aulacus pallipes
Aulacus digitalis
Pristaulacus davisi
Pristaulacus lateritius
Pristaulacus ornatus
Pristaulacus albosignatus
Pristaulacus mouldsi
Pristaulacus fuscocostalis
Pristaulacus curryi
Pristaulacus kiunga
Pristaulacus fasciatus
Pristaulacus brasiliensis
Pristaulacus
Pristaulacus
Pristaulacus thoracicus
Pristaulacus stephanoides
Pristaulacus smithi
Pristaulacus signatus
Pristaulacus sexdentatus
Pristaulacus ryukyuensis
Pristaulacus rufitarsis
Pristaulacus rufipes
Pristaulacus rubriventer
Pristaulacus resutorivorus
Pristaulacus pilatoi
Pristaulacus pacificus
Pristaulacus occidentalis
Pristaulacus niger
Pristaulacus montanus
Pristaulacus kostylevi
Pristaulacus insularis
Pristaulacus gibbator
Pristaulacus galitae
Pristaulacus foxleei
Pristaulacus flavicrurus
Pristaulacus editus
Pristaulacus comptipennis
Pristaulacus chlapowskii
Pristaulacus boninensis
Pristaulacus barbeyi
Panaulix irenae
Panaulix rex
Pristaulacus
Pristaulacus emarginaticeps
Pristaulacus nigripes
Pristaulacus iridipennis
Pristaulacus lindae
sp.
sp. 2
sp. 1
sp. 2
sp. 1
sp. 4
sp. 3
sp. 2
sp. 1
sp. 3
Evaniidae
Gasteruptiidae
Stephanidae
Trigonalidae
Megalyridae
Braconidae
Aulacidae
7/99 3/73
9/95
3/68
5/-
2/-
2/-
1/-
1/-
2/-
3/-
1/- 2/-
2/-
5/64
1/-
2/-
2/-
10/99
2/-
2/60
1/-
2/-
5/-
Fig. 19. Phylogenetic relationships of Aulacidae (equal weighting analysis). Strict consensus of 18 000 equally parsimonious trees of 593
steps each (see text for detail). Bremer/Jacknife values are indicated above the branches (missing values as ‘–’).
44 Invertebrate Systematics G. F. Turrisi et al.
Stephanus serrator
Doryctes erythromelas
Orthogonalys pulchella
Szepligetella
Brachygaster minuta
Zeuxevania splendidula
Evania appendigaster
Megalyra fasciipennis
Gasteruption
Gasteruption
Pseudofoenus crosskeyi
Hyptiogaster pinjarregaensis
Aulacus wau
Aulacus sp. 1
Aulacus sp. 2
Aulacus sedlaceki
Aulacus
Aulacus
Aulacus krahmeri
Aulacus braconiformis
Aulacus impolitus
Aulacus brevicaudus
Aulacus burwelli
Aulacus emineo
Aulacus striatus
Aulacus schoenitzeri
Aulacus japonicus
Aulacus pallipes
Aulacus digitalis
Aulacus burquei
Aulacus elegans
Aulacus houstoni
Aulacus belairensis
Aulacus douglasi
Aulacus enaratodi
Aulacus coracinus
Aulacus grossi
Pristaulacus barbeyi
Pristaulacus rufitarsis
Pristaulacus kostylevi
Pristaulacus editus
Pristaulacus davisi
Pristaulacus rubriventer
Pristaulacus lateritius
Pristaulacus ornatus
Pristaulacus albosignatus
Pristaulacus fuscocostalis
Pristaulacus mouldsi
Pristaulacus curryi
Pristaulacus fasciatus
Pristaulacus kiunga
Pristaulacus brasiliensis
Pristaulacus pacificus
Pristaulacus occidentalis
Pristaulacus gibbator
Pristaulacus foxleei
Pristaulacus resutorivorus
Pristaulacus africanus
Pristaulacus thoracicus
Pristaulacus pilatoi
Pristaulacus stephanoides
Pristaulacus
Pristaulacus sexdentatus
Pristaulacus ryukyuensis
Pristaulacus signatus
Pristaulacus nigripes
Pristaulacus iridipennis
Pristaulacus smithi
Pristaulacus
Panaulix irenae
Panaulix rex
Pristaulacus rufipes
Pristaulacus flavicrurus
Pristaulacus niger
Pristaulacus chlapowskii
Pristaulacus montanus
Pristaulacus galitae
Pristaulacus lindae
Pristaulacus compressus
Pristaulacus
Pristaulacus emarginaticeps
Pristaulacus insularis
Pristaulacus comptipennis
Pristaulacus boninensis
sp.
sp. 1
sp. 2
sp. 4
sp. 3
sp. 2
sp. 1
sp. 3
Evaniidae
Gasteruptiidae
Stephanidae
Trigonalidae
Megalyridae
Braconidae
Aulacidae
1
3
2
4
5
6
7
8
9
10
Fig. 20. Phylogenetic relationships of Aulacidae (implied weighting analysis). Selected tree obtained with k =9 (see text for detail).
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 45
the type species of Aulacidae, A. striatus) (node 5). However, it
is also found in two species from New Caledonia (A. emineo)
and New Guinea (A. sedlaceki); the presence in the former is
probably convergent with the Holarctic clade (Jennings et al.
2004c). No other Australasian species possess the longitudinal
type of hind coxal groove (Jennings et al. 2004c). A few species
Stephanidae
Braconidae
Trigonalidae
Evaniidae
Megalyridae
Gasteruptiidae
Aulacus wau
Aulacus sp. 1
Aulacus braconiformis + 7 spp.
Aulacus burwelli + 7 spp.
Aulacus elegans
Aulacus belairensis + 4 spp.
Aulacus grossi
Pristaulacus barbeyi + 3 spp.
Pristaulacus africanus+ 37 spp.
Pristaulacus davisi
Pristaulacus rubriventer
Pristaulacus lateritius
Pristaulacus albosignatus
Pristaulacus ornatus
Stephanidae
Braconidae
Trigonalidae
Evaniidae
Megalyridae
Gasteruptiidae
Aulacus wau
Aulacus sp. 2
Aulacus sedlaceki
Aulacus braconiformis + 5 spp.
Aulacus burwelli
Aulacus emineo
Aulacus pallipes
Aulacus burquei
Aulacus digitalis
Aulacus striatus
Aulacus japonicus
Aulacus schoenitzeri
Aulacus elegans
Aulacus belairensis + 4 spp.
Aulacus grossi
Pristaulacus spp.
Stephanidae
Braconidae
Trigonalidae
Evaniidae
Megalyridae
Gasteruptiidae
Aulacus wau
Aulacus sp. 1
Aulacus braconiformis + 7 spp.
Aulacus burquei + 7 spp.
Aulacus elegans
Aulacus belairensis + 4 spp.
Aulacus grossi
Pristaulacus barbeyi + 3 spp.
Pristaulacus albosignatus + 4 spp.
Pristaulacus curryi + 2 spp.
Pristaulacus fasciatus
Pristaulacus kiunga
Pristaulacus brasiliensis
Pristaulacus pacificus
Pristaulacus occidentalis
Pristaulacus gibbator
Pristaulacus foxleei
Pristaulacus resutorivorus
Pristaulacus africanus
Pristaulacus boninensis + 23 spp.
Pristaulacus pilatoi + 1 sp.
Aulacus sp. 1
A B
C
Fig. 21. Diagnostic characters of Townes (1950) for Aulacus and Pristaulacus traced onto the preferred topology (implied weighting, k = 9). Clades
containing taxa with identical scorings for the character in question have been collapsed into single terminals. (A) Occipital carina (character 5): absent
(white) or present (black); (B) orientation of hind coxal groove (character 49): longitudinal (white) or transverse (black); (C) number and configuration
of tarsal claws (character 55): one small basal tooth present (white); one apical/medial tooth present (dark grey; only in Evaniidae and Trigonalidae);
two well spaced teeth present (light grey; only in some Pristaulacus spp.); three to six teeth present (black). The most recent common ancestor of
Pristaulacus is not unambiguously resolved.
46 Invertebrate Systematics G. F. Turrisi et al.
of Aulacus have a transverse hind coxal groove, as have all
Pristaulacus, thus this character is putatively homoplastic.
The genus Pristaulacus Kieffer
Two of the most consistent results of our analyses are the
placement of Panaulix spp. deeply within Pristaulacus, and
the retrieval of a well supported clade comprising all members
of the two genera. Pristaulacus is thus rendered paraphyletic.
The simplest solution to this quandary is to sink Panaulix
into Pristaulacus, thus making the latter monophyletic. As
consequence of these systematic changes, Pristaulacus must
be rediagnosed and redescribed. This is done in Appendix 2,
which also deals with the systematic changes to Panaulix.
Within the redefined Pristaulacus it is possible to recognise
some clades that are recoveredas monophyletic. A very large clade
(node 7), including 27 species (from nearly all zoogeographic
regions, excluding Australasia), is well supported by one
unambiguous autapomorphy: having the lateroventral margin of
the pronotum anteriorly angulated (26: 1). Within this clade, a
large subclade (node 8) is supported by the presence of an anterior
tooth-like process onthe pronotum (28: 1); a fewspecies within the
clade have an additional posterior process (29: 1).
Two distinct subgroups can be recognised within this large
clade. The first (node 9) contains the two species formerly assigned
to Panaulix, well supported by three unique autapomorphies: 1,
a large body length (more than 20 mm, excluding ovipositor)
(1: 1); 2, the presence of a lamelliform medial process on
the clypeus (2: 2); 3, having the antennomeres ventrally carinate
(18: 1). Another feature, the presence of a flattened hind tibia
(53: 1), is characteristic of Panaulix although also present in
Pristaulacus sp. 1, and thus should be considered properly as a
synapomorphy of Panaulix+Pristaulacus sp. 1. Other characters
are quite distinctive for the Panaulix clade, but apparently
represent plesiomorphic or homoplastic characters within
Evanioidea and Aulacidae: 1, A1 rather elongate, with length
~3.0!width (15: 1) (presumably the groundplan of Evanioidea);
2, mesosomal setae long (21: 1) (homoplastic with another small
clade within Pristaulacus); 3, mesosomal setae recumbent
(22: 1) (homoplastic with another small clade within
Pristaulacus); 4, mesosomal setae dense (23: 1) (homoplastic
with a few scattered clades within Aulacus and Pristaulacus);
5, mesoscutum areolate–rugose (38: 0) (presumably a retained
plesiomorphic character within Aulacidae, homoplastic in a few
scattered clades within Aulacus and Pristaulacus).
The other very distinctive subgroup (node 10), including
five Oriental and eastern Palaearctic species, the comptipennis-
group of Turrisi (2007) and Turrisi and Pilato (2008), is
well supported by the presence of a more or less deep and
wide medial groove on the posterior margin of the head (4: 2)
(Fig. 3B).
The results of our cladistic analyses do not warrant the
subdivision of Pristaulacus into formal species-groups or
subgenera; this must await a revision of the entire genus.
Functional morphology and homoplasy in Aulacidae
Recent studies on the functional morphology of some parasitoid
Hymenoptera have pointed out the possible adaptive value of
numerous features of the imago to the life-style as a parasitoid of
xylophagous insects (Quicke 1997; Vilhelmsen 1997b, 2003;
Vilhelmsen et al. 2001, 2008); some papers specifically treat
such features in Aulacidae (Quicke and Fitton 1995; Turrisi 2004;
Turrisi and Pilato 2004; G. F. Turrisi et al. unpubl. data). Several
morphological traits are believed to facilitate oviposition into
a woody gallery to reach the host larva (e.g. the length of the
ovipositor, and the presence of hind coxal grooves), whereas
others are involved in the emergence of the imago from the gallery
after completing the larval development (e.g. prominent sculpture
on the head and mesosoma).
Although the biology and the parasitoid/host relationship is
known for only some species, the Aulacidae are believed to be
parasitoids of wood-boring insects, mainly the Cerambycidae and
Buprestidae (Coleoptera). It is likely that many of the characters
included in the present study are adaptive with regard to the wood-
living lifestyle (see above). They might thus be prone to parallel
evolution because of the substantial selection pressures involved
in surviving and reproducing in this physically demanding
habitat. This indeed seems to be the case with at least some of
the characters studied here, for example, the sculpture on the frons
and mesosoma, the presence of tooth-like processes on the
mesosoma, and the orientation of the hind coxal groove
(Fig. 21B). If parallel evolution in morphological characters is
rampant in the Aulacidae, it makes the task of unravelling their
phylogenetic relationships even more difficult.
Concluding remarks
The main conclusions derived from the present cladistic study
of relationships within the Aulacidae are: 1, the existence of a
strongly supported clade corresponding to Pristaulacus s.l.
(including Panaulix spp.). We have dealt with this by
including Panaulix in Pristaulacus (see Appendix 2); 2, the
existence of a paraphyletic basal grade comprising the species
currently placed within Aulacus. In order to obtain a stable
classification of ‘Aulacus’, it will have to be split into several
genera. However, to do this on the basis of our current analysis
would entail the recognition of many monotypic or poorly
supported taxa. Furthermore, given that we have included
fewer than half the described species of ‘Aulacus’and the
Aulacidae as a whole in the present study, and also excluded
many undescribed species known to us, we deem a
reclassification of ‘Aulacus’to be premature. To obtain a
stable generic classification, a larger taxon and character
sample is essential. Thus, we recommend retaining Aulacus as
a taxonomic entity for the time being.
Acknowledgements
The Willi Hennig Society is kindly acknowledged for access to TNT software.
We are grateful to Prof. Dr Klaus Schönitzer (Zoologische Staatssammlung
München, Germany) and to Prof. Dr Matthias Starck (Zoological Institute of
Ludwig-Maximilians-Universität, Munich, Germany) for allowing us to
produce the SEM micrographs. We are also grateful to the curators of the
numerous museums and colleagues who provided material for this study
(see list in the section ‘Materials and methods’). Two anonymous referees
provided useful comments on an earlier versionof this paper. This research has
been partly supported by the University of Catania, Fondo Ricerca d’Ateneo
(ex 60%) to G. F. T. and an Australian Biological Resources Study grant to
J. T. J.
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 47
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Manuscript received 19 August 2008, accepted 28 January 2009
50 Invertebrate Systematics G. F. Turrisi et al.
Appendix 1. Data matrix for the parasitoid wasp family Aulacidae
–= unknown, A = polymorphic state 0 & 1
Taxa Origin Characters
0000000001 1111111112 2222222223 3333333334 4444444445 5555555556 6666666667 777777777
1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 123456789
Stephanus serrator Palaearctic 0000100020 0000100000 0001000001 0000––0000 0–00–000–– 0002000001 0001101000 002000230
Doryctes erythromelas Nearctic 0000100110 0101000001 0001100001 011–––0001 00001010–– 0000002000 0001000101 002000220
Megalyra fasciipennis Australasian 0010110100 2010010010 A000100001 011001100– 10001010–– 000000000– 0–00000111 103000121
Orthogonalys pulchella Nearctic 0000100120 0111000000 0001100001 0010010000 00001010–– 0000101210 0–00000111 02022200?
Hyptiogaster pinjarregaensis Australasian 0000110010 0111110000 0000100001 1000002012 00010000–– 0102010000 1022101111 101111010
Pseudofoenus crosskeyi Australasian 0001110010 0111110000 0001100001 1000000012 00010000–– 0102010000 1022101111 010111010
Gasteruption sp. 1 Palaearctic 0000100011 0101010000 0001100001 1000000012 00010000–– 0002010000 1022101111 1000000?1
Gasteruption sp. 2 Australasian 0001100000 0111110000 0001100001 1000002012 00010000–– 0002010000 1022101111 100020031
Evania appendigaster Palaearctic 0000100111 0101111000 0111000000 01100–0001 00001110–– 0000100001 0113111111 020000020
Brachygaster minuta Palaearctic 0000100021 0101111000 0011000000 01100–0001 00001110–– 0000100001 0113111111 020000020
Zeuxevania splendidula Palaearctic 0000100011 0101111000 0111000000 01100–0001 00001100–– 0000100101 0113111111 020000020
Szepligetella sp. Australasian 0000110010 0001111000 0011000000 00100–0000 00001100–– 0000100001 0113111111 020000020
Aulacus belairensis Australasian 01000––011 1010000000 0001100001 1001102010 0001000111 0000000111 1012000001 101100110
Aulacus braconiformis Neotropical 01100––000 21100000?? 0001100000 1001010000 0001000110 0000002210 1011000111 102000220
Aulacus brevicaudus Nearctic 01100––101 0010000000 0001100000 1011002110 00010000–– 0000000110 1010000110 101000010
Aulacus burquei Nearctic 01100––001 0011000000 0001100001 1000001110 000100010– 1000000110 1010000001 101111010
Aulacus burwelli Australasian 01000––020 0011000011 0001100001 1000012000 0001000111 0000001210 1010001001 102012020
Aulacus coracinus Australasian 00000––010 0011000011 0011100000 1001102010 0001000111 0000001210 1012001101 101000120
Aulacus digitalis Nearctic 01100––001 0011000000 0001100000 1000001100 000100010– 1000001210 1010000001 101200010
Aulacus douglasi Australasian 01000––010 21110000?? 0001100000 1001102010 0001000111 0000001111 1012000101 101012310
Aulacus elegans Australasian 00000––010 1010000010 0001100001 1001102100 0101000111 0000001110 1010000101 102100110
Aulacus emineo Australasian 01000––010 1010000011 0001100000 1000012100 020100010– 1000000110 1010000001 102011110
Aulacus enaratodi Australasian 01000––010 0011000010 0001100001 1001102010 0001000111 0000001210 1012001101 101000120
Aulacus grossi Australasian 0100101020 1010000011 0001100001 1001102100 0001000111 0000001110 1011000101 101011010
Aulacus houstoni Australasian 01000––000 2110000010 0001100000 1001102010 0001000111 0000001110 1011000001 101102120
Aulacus impolitus Nearctic 01100––000 1010000000 0001000000 1001001000 00010000–– 0000001110 1010000111 101111010
Aulacus japonicus Palaearctic 01100––001 1010000000 0001100000 1000001110 000100010– 1000001210 101000000? 101000020
Aulacus krahmeri Neotropical 01100––010 1010000011 0001100000 1001001100 0001000110 0000002210 1011001011 102000131
Aulacus pallipes Nearctic 01100––001 1010000000 0001100000 100000?110 000100010– 1000001110 101100000? 101222010
Aulacus schoenitzeri Palaearctic 01100––001 ?010000000 0001100000 1000001110 000100010– 1000001210 1010000010 101000010
Aulacus sedlaceki Australasian 00000––110 1010000000 0001100001 1010112110 100100010– 0000000210 1011000001 103111100
Aulacus striatus Palaearctic 01100––001 1010000000 0001100000 1000001110 000100010– 1000001110 1010000000 101000010
Aulacus wau Australasian 00000––100 00100??0?? 0001100001 1000011100 00010000–– 0000000110 1010000101 101011020
Aulacus sp. 1 Neotropical 00000––111 1110000000 0001100001 1010001100 0001100011 0000001110 1012001101 102111110
Aulacus sp. 2 Neotropical 01000––110 1110000001 0001100001 1010001010 0001000111 0000001110 1012000101 103111100
Aulacus sp. 3 Neotropical 01100––110 00100000?? 0001100001 1010011110 0001101110 0000001110 1011001111 102011111
Aulacus sp. 4 Neotropical 01100––110 1111000000 0001100001 1010011100 0001101110 0000002110 1011001111 103222110
Pristaulacus africanus Afrotropical 0100101110 1110000011 0000110000 1011011101 0021100111 0000300210 1011000011 102000210
Pristaulacus albosignatus Australasian 0001101110 0110000010 0000100001 1001101101 1121000111 0000300111 1012000101 102212130
Pristaulacus barbeyi Palaearctic 0100100010 1010000000 0000100000 1001101101 0021100111 0000200111 1010000001 102000020
Pristaulacus boninensis Palaearctic 0102110110 2110000011 0000110100 1011102101 0021100111 0000300211 1011001111 102000110
(continued next page )
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 51
Appendix 1. (continued )
Taxa Origin Characters
0000000001 1111111112 2222222223 3333333334 4444444445 5555555556 6666666667 777777777
1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 123456789
Pristaulacus brasiliensis Neotropical 0100100111 1110000011 1100100001 1000012101 1221101111 0000200110 1012000101 102111121
Pristaulacus chlapowskii Palaearctic 0100110110 2110000011 0000110100 1011102101 0021100111 0000300211 1011001111 102000120
Pristaulacus compressus Palaearctic 0100110110 2110000011 0000110110 1011102101 0021100111 0000301211 1011001111 102000110
Pristaulacus comptipennis Palaearctic-Oriental 0102110110 2110000011 0000110100 1011102101 0021100111 0000300211 1011001111 102000120
Pristaulacus curryi Australasian 0100110020 1010000010 0000100001 1001002100 1121000111 0000200111 1012000101 102000020
Pristaulacus davisi Australasian 0100111110 20100000?? 0000100001 1001102101 0011000111 0000300111 1011000101 1010111?0
Pristaulacus editus Nearctic 0100100010 11100000?? 0000100000 1011102110 0221100111 0000200111 1010000111 102000010
Pristaulacus emarginaticeps Oriental 0102110110 2110000010 0000011110 1011102101 1121100111 0000300111 1011001111 102000220
Pristaulacus fasciatus Nearctic 0100100111 11100000?? 1000100001 1001112101 1011000111 0000300111 1010000101 102000220
Pristaulacus flavicrurus Nearctic 0100100110 2110000011 0000110100 1011102101 0021100111 0000300111 1011001111 102000110
Pristaulacus foxleei Nearctic 0101100100 2110000011 0000100000 1011111101 0011101111 0000300110 1011001111 102111110
Pristaulacus fuscocostalis Australasian 0101110010 0110000010 0000000101 1011002111 1011000111 0000200111 1012001101 101000120
Pristaulacus kiunga Australasian 0101100111 2110000010 1110000110 1001012101 1111000111 0000300111 1012000101 102000120
Pristaulacus galitae Palaearctic 0100110110 2110000011 0000110100 1011102101 0021100111 0000300211 1011001111 102000120
Pristaulacus gibbator Palaearctic 0100100011 1110000011 0000100000 1011001101 0011000111 0000300110 1011001111 102000010
Pristaulacus insularis Palaearctic 0102110110 2110000011 0000010101 1011102101 0021100111 0000300211 1011001111 102000120
Pristaulacus iridipennis Oriental 0100100110 0110000011 0000111100 1011102101 1121100111 0000300211 1011001111 102000120
Pristaulacus kostylevi Palaearctic 0100100110 1110000011 0000100000 1011102101 0221100111 0000200111 1010000111 102000010
Pristaulacus lateritius Australasian 0100101010 1110000010 0010100011 1001112101 1121000111 0000300111 1011000101 101111110
Pristaulacus lindae Palaearctic 0100110110 2110000011 0000110110 1011102101 0021100111 0000301211 1011001111 102000120
Pristaulacus montanus Nearctic 0100110110 2111000011 0000110100 1011102101 0021100111 0000300211 1011001111 102000120
Pristaulacus mouldsi Australasian 01001–0011 2010000010 0000100001 1001002101 1111000111 0000200111 1011000101 102000220
Pristaulacus niger Nearctic 0100110110 1110000011 0000110100 1011102101 0011100111 0000300111 1011001111 102000120
Pristaulacus nigripes Oriental 0100100110 0110000011 0000111101 1011102101 1121100111 0000300211 1011001111 102000120
Pristaulacus occidentalis Nearctic 0100110001 1110000011 0000100000 1011012101 0011101111 0000300111 1011000001 101112210
Pristaulacus ornatus Australasian 01010––111 1010000011 0000100000 1001102101 1211000111 0000300110 1011000101 102222030
Pristaulacus pacificus Nearctic 0100100010 21100000?? 0000100000 1001002101 0011100111 0000300111 1011001101 102000110
Pristaulacus pilatoi Afrotropical 0100101110 1110000011 0000110000 1011011001 0021100111 0000200211 1011001111 102000220
Pristaulacus resutorivorus Nearctic 0100110010 2010000011 0000100000 1011111101 0011100111 0000300210 1010000111 102000120
Pristaulacus rubriventer Neotropical 0100101110 2010000010 0000100001 1011112101 0221100111 0001300211 1011000001 102000120
Pristaulacus rufipes Oriental 0100100110 11100000?? 0000110101 1011102101 0021100111 0000300111 1011001111 102000110
Pristaulacus rufitarsis Nearctic 0100100110 2110000010 0000100000 1001102101 0221100111 0000201211 1011000111 102000010
Pristaulacus ryukyuensis Palaearctic 0100100110 2110000011 0000111101 1011102101 0121100111 0000300211 1011001111 102000120
Pristaulacus sexdentatus Neotropical 0100110120 2110000011 0000111100 1011102111 1121100111 0000300210 1011001111 102111121
Pristaulacus signatus Oriental 0100100110 2110000011 0000111100 1011102101 1121100111 0000300211 1011001111 102000120
Pristaulacus smithi Afrotropical 0100100120 1110000000 0000110101 1011002101 0021100111 0001300110 1011001111 101010110
Pristaulacus stephanoides Neotropical 0100100110 21100000?? 0000110101 1011102101 1021101111 0000300210 1012001101 103111111
Pristaulacus thoracicus Afrotropical 0100101111 21100000?? 0000110000 1011012101 0021100111 0000200210 101100111? 10200103?
Pristaulacus sp. 1 Neotropical 0101100110 2110000011 0000110101 1011102101 0021100111 0011300111 1011001111 102011030
Pristaulacus sp. 2 Neotropical 0100110110 2110000011 0000110101 1011102101 1121101111 0000300210 1011001111 102000120
Pristaulacus sp. 3 Oriental 0102110110 2110000011 0000110111 1011102001 0121100111 0000301211 1011001111 102000220
Panaulix rex Afrotropical 1200100010 21101101?? 1110?10100 1011112001 0021101111 0011300210 1011001111 102002020
Panaulix irenae Afrotropical 1200100110 21101101?? 1110?10100 10?1112001 0021?01111 0011300210 101100111? 1021110?0
52 Invertebrate Systematics G. F. Turrisi et al.
Appendix 2. Diagnosis and redescription of the genus Pristaulacus Kieffer
Pristaulacus Kieffer, 1900
Aulacostethus Philippi, 1873: 302. Type species: A. rubriventer Philippi, by monotypy (preoccupied by Waterhouse, 1869, in Coleoptera and by Uhler, 1871,
in Heteroptera).
Aulacosthethus Schletterer, 1890: 523. Error.
Pristaulacus Kieffer, 1900: 813. Type species: P. chlapowskii Kieffer, designated by Kieffer, 1903: 455.
Deraiodontus Bradley, 1901a: 29 (no species included). Type species: Aulacus montanus Cresson, by subsequent designation. First species included
by Bradley, 1901b: 321.
Oleisoprister Bradley, 1901b: 324. Type species: Aulacus firmus Cresson, by original designation.
Aulacostathus Dalla Torre, 1902: 1062. Error.
Anaulacus Semenow, 1903: 173. Type species: Aulacus sibiricola Semenow, by subsequent designation of Bradley, 1908: 120. Preoccupied by MacLeay,
1825, in Coleoptera.
Semenowia Kieffer, 1903: 382. Nomen novum pro Anaulacus Semenow. Preoccupied by Weise, 1889, in Coleoptera.
Odontaulacus Kieffer, 1903: 382. Type species: Aulacus rufitarsis Cresson, designated by Bradley, 1908: 120.
Semenovius Bradley, 1908: 120, 123. Nomen novum pro Semenowia Kieffer.
Tropaulacus Bradley, 1908: 120. Type species: T. torridus Bradley, by original designation.
Pristaulacus (Neaulacus) Bradley, 1908: 121. Type species: Aulacus occidentalis Cresson, by original designation.
Interaulacus Bradley, 1908: 120. Type species: I. kiefferi Bradley, by original designation.
Tetraulacinus Kieffer, 1910: 350. No species included. Type species: Pristaulacus rufobalteatus Cameron, by subsequent monotypy. First species
included by Kieffer, 1911: 214.
Psilaulacus Kieffer, 1910: 350. No species included. Type species: P. annulatus Kieffer, by subsequent monotypy. First species included
by Kieffer, 1911: 215.
Aulacosthetus Kieffer, 1912: 370. Error.
Aulacites Cockerell, 1916: 102. Type species: Aulacites secundus Cockerell, by original designation.
Aulacomastus Muesebeck & Walkley, 1956: 333. Unnecessary nomen novum pro Aulacostethus Philippi.
Odontacolus Kozlov, 1988: 242. Error.
Panaulix Benoit, 1984: 800. Type species: Panaulix rex Benoit, 1984 by original designation (syn. nov.).
Pristaulacus was erected by Kieffer (1900) to include the species formerly included within the genus Aulacostethus Philippi, 1873, which was preoccupied
by genera of Heteroptera and Coleoptera, and thus invalid. Subsequently, several new genera of the Aulacidae allied to the genus Pristaulacus were
described, mainly on the basis of wing venation and of the number and shape of teeth on the inner margin of the claw. But these genera (see synonym list) were
considered as subgenera (Szépligeti 1903; Bradley 1908, 1926) or synonyms (Townes 1950) of Pristaulacus.
Diagnosis
The genus, based mainly on our phylogenetic investigations, is defined by the combination of the following characters, some of which are autapomorphies within
Aulacidae: 1, occipital carina present; 2, prothoracic spiracle visible, located in a notch in the posterolateral margin of pronotum; 3, prescutum defined;
4, mesoscutum with notauli convergent medially to join prior reaching transscutal fissure (the so called Y-shaped notauli, cfr. Crosskey, 1962); 5, parascutal carina
expanded to partially cover the tegula, mostly with tooth-like lateral projection; 6, claws pectinate, with two well spaced teeth or 3–6 teeth along inner margin;
7, forewing vein 2r-m mostly absent; 8, forewing vein 2-Rs+M mostly short; 9, transverse groove on each medial surface of hind coxa (forming together
the hind coxal ovipositor guide).
Redescription on a worldwide basis
Medium, mostly moderately large, or rarely large sized wasps, length (excluding ovipositor) up to 28 mm (e.g. in the Afrotropical P. rex).
Head. General shape variable, mostly large and massive; anterior margin regularly rounded; posterior margin almost straight or weakly concave, but with a
more or less deep and wide medial groove in a distinctive eastern Palaearctic and Oriental group (P. comptipennis group); occipital margin smooth; occipital carina
present, very narrow, cerciniform(at most 0.2!diameter of anocellus)to wide andlamelliform (0.5–1.5!diameter ofanocellus), entireormedially absent; frontal
area mostly smooth and punctate or less frequently transversely carinulate; POL:OOL 0.9–2.0 or more; eye mostly large and elliptical, or small and circular to
subcircular; subantennal groove present, weak or mostly well developed; antenna 14-segmented in female, 13-segmented in male; antennal insertion about level
with base of eye, weakly below, or about half-way; A1 mostly stocky or moderately elongate (length <2.0!width), to rather elongate (length >2.0!width) in the
formerPanaulix,in lateral view medially convex and swollen; antennomeres mostly not carinate, otherwise with a weak ventral longitudinal carina (only in the two
former Panaulix species); last antennomere flattened with apex rounded; clypeus mostly with medial process (apparently absent in one Australasian species),
tooth-like or lamelliform (in the two former Panaulix-species); mandibles robust and wide, with three apical teeth, broadly overlapping at rest. Setal configuration:
variable, short to moderately long, scattered to dense and mostly erect on top of head and on temple, semi erect on clypeus, recumbent on frontal and malar area.
Mesosoma. General shape variable, relatively robust and stocky to moderately elongate and weakly depressed; pronotum narrow (length/width >1.0, at most
as long as high) or wide (length/width <1.0), and short dorsally of transverse sulcus with lateroventral margin rounded or mostly angulated, without tooth or mostly
with one tooth, in a few species with two teeth, anterior and posterior; pronotal lateral process mostly absent; pronotal spiracle visible, located in a notch in the
posterolateral margin of pronotum; propleuron polished, at most punctate; anterior margin of mesoscutum (lateral view) rounded, acute or acute and upward
directed; anterolateral parts of mesoscutum normal or sometimes produced in two more or less developed horn-like processes; parascutal carina expanded, mostly
with a lateral tooth-like projection; mesoscutum surface mostly strongly transverse–carinate; latero-anteroadmedian lines convergent posterodorsally, delimiting
prescutum; prescutum not concave or more or less concave; mesonotum with scutellum not subdivided by longitudinal grooves; notauli convergent medially to
join prior reaching transscutal fissure, forming a Y-shaped line; hind coxa mostly transverse to carinate; hind coxal guide of ovipositor present, with each coxa
notched, medially grooved, groove transverse or oblique respect to the coxal axis; transverse medial groove of hind coxa complete; distal lobe of hind coxa absent;
hind tibia mostly subcylindrical or flattened, sometimes moderatelyenlarged toward apex (in the former Panaulix species); claw withtwo main configurations: two
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 53
wellspacedteeth (serrated pattern)orthree tosixteeth (strictly pectinatepattern);forewing vein2r-mabsent or present,but mostly spectral,recognizableonly inthe
hind part; forewing vein 3r-m present or mostly spectral; forewing vein 2-Rs+M mostly short (less than 0.5!length of Rs+M), sometimes long (length more than
0.5!length of Rs+M). Setal configuration: highly variable, short to very long, scattered to very dense, erect or semi erect to recumbent.
Metasoma. Mostly pyriform in lateral view, sometimes ovoidal; petiole mostly elongate (length/width: 2.0–4.5 or more), sometimes stocky (length/width:
1.0–1.1); base of petiole polished; segments 1–2 polished; T2–T5 with fine and dense punctures; ovipositor mostly long or very long, normally 1.0–2.0!length of
forewing. Setal configuration: segments 1–2 glabrous; T2–T5 with short, recumbent and mostly dense setae.
Colour patterns. Variable; head mostly blackish, sometimes partially or entirely red orange or reddish; antenna mostly entirely blackish, sometimes
bicoloured, with scape red orange; mesosoma mostly blackish, sometimes partially or entirely red orange or reddish; fore and hind wing hyaline to strongly
infuscated; forewing mostly with one or more dark brown spots (one frequently below stigma, and one on apex); legs variously coloured mostly blackish with tarsi
lighter; hind tarsus very variable, white or cream, yellow, orange or black, sometimes bicoloured; metasoma mostly variously bicoloured, blackish and red orange
or reddish, but sometimes entirely black or blackish, reddish, or red orange; valvula 3 mostly entirely blackish or dark brown, sometimes broadly ivory before tip.
Systematic changes
Panaulix Benoit, 1984 is here regarded as a synonym of Pristaulacus Kieffer, 1900 (syn. nov.). The two species previously placed in Panaulix are newly
combined as Pristaulacus rex (Benoit, 1984), comb. nov., and Pristaulacus irenae (Madl, 1990), comb. nov.
54 Invertebrate Systematics G. F. Turrisi et al.
Appendix 3. Updated checklist of the world Aulacidae
Only the most recently proposed synonymies are reported. The symbol (†) refers to extinct species
Genus Aulacus Jurine, 1807
Palaearctic
Nearctic
Oriental
Afrotropical
Neotropical
Australasian
(1) A. albimanus (Kieffer, 1911) X
(2) A. amazonicus (Roman, 1917) X
(3) A. aneurus Walkley, 1952 X
(4) A. aroueti (Girault, 1926) X
(5) A. atriceps (Kieffer, 1911) X
(6) A. belairensis Jennings, Austin & Stevens, 2004 X
(7) A. biroi (Szépligeti, 1903) X
(8) A. bituberculatus Cameron, 1899 X
(9) A. braconiformis (Kieffer, 1911) X
(10) A. brasiliensis (Szépligeti, 1903) X
(11) A. braunsi (Kieffer, 1911) X
(12) A. brevicaudus (Cushman, 1929) X
(13) A. brevis Smith, 2005 X
(14) A. burquei (Provancher, 1882) X
(15) A. burwelli Jennings, Austin & Stevens, 2004 X
(16) A. cephalus Smith, 2005 X
(17) A. coracinus Jennings, Austin & Stevens, 2004 X
(18) A. costaricensis Smith, 2008 X
(19) A. costulatus (Kieffer, 1904) X
(20) A. digitalis Townes, 1950 X
(21) A. dispilis Townes, 1950 X
(22) A. douglasi Jennings, Austin & Stevens, 2004 X
(23) A. elegans (Kieffer, 1911) X
(24) A. elongatus Smith, 2008 X
(25) A. emineo Jennings, Austin & Stevens, 2004 X
(26) A. enarotadi Jennings & Austin, 2006 X
(27) (†)A. eocenicus Nel, Waller & Ploëg, 2004 X
(28) A. fascius Smith, 2008 X
(29) A. festivus (Kieffer, 1911) X
(30) A. flavicornis (Kieffer, 1911) X
(31) A. flavigenis Alekseyev, 1986 X
(32) A. flavimanus (Kieffer, 1911) X
(33) A. flindersbaudini Jennings, Austin & Stevens, 2004 X
(34) A. forus Smith, 2001 X
(35) (†)A. fritschii (Brues, 1933)
(36) A. fuscicornis Cameron, 1911 X
(37) A. fusiger Schletterer, 1890 X
(38) A. gaullei (Kieffer, 1904) X
(39) A. grossi Jennings, Austin & Stevens, 2004 X
(40) A. heredia Smith, 2008 X
(41) A. houstoni Jennings, Austin & Stevens, 2004 X
(42) A. hyalinipennis Westwood, 1841 X
(43) A. impolitus Smith, 1991 X
(44) A. japonicus Konishi, 1990 X
(45) A. jeoffreyi Alekseyev, 1993 X
(46) A. kohli Schletterer, 1890 X
(47) A. krahmeri Elgueta & Lanfranco, 1994 X
(48) A. leon Smith, 2008 X
(49) A. longiventris (Kieffer, 1911) X
(50) A. lovei (Ashmead, 1901) X
(51) A. maculosus Smith, 2008 X
(52) A. mcmillani Jennings, Austin & Stevens, 2004 X
(53) A. minutus Crosskey, 1953 X
(54) A. moerens Westwood, 1868 X
(55) A. nigriventris (Kieffer, 1911) X
(continued next page )
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 55
Appendix 3. (continued )
Genus Aulacus Jurine, 1807
Palaearctic
Nearctic
Oriental
Afrotropical
Neotropical
Australasian
(56) A. notatus (Szépligeti, 1903) X
(57) A. ochreus Smith, 2005 X
(58) A. pallidicaudis (Cameron, 1911) X
(59) A. pallipes Cresson, 1879 X
(60) A. pediculatus Schletterer, 1890 X
(61) A. philippinensis (Kieffer, 1916) X
(62) A. planiceps (Szépligeti, 1903) X
(63) (†)A. prisculus (Brues, 1933)
(64) A. pterostigmatus (Szépligeti, 1903) X
(65) A. salicius Sun & Sheng, 2007 X
(66) A. schiffiSmith, 1996 X
(67) A. schoenitzeri Turrisi, 2005 X
(68) A. sedlaceki Jennings & Austin, 2006 X
(69) A. sinensis He & Chen, 2007
(= A. erythrogaster He & Chen, 2002; nom. nov.)
X
(70) A. striatus Jurine, 1807 X
(71) A. truncatus (Kieffer, 1911) X
(72) A. veracruz Smith, 2008 X
(73) A. vespiformis (Kieffer, 1911) X
(74) A. wau Jennings & Austin, 2006 X
(75) A. whartoni Smith, 2008 X
(76) A. westwoodi Kieffer, 1868 X
Genus Pristaulacus Kieffer, 1900
(1) P. absens Smith, 2005 X
(2) P. acutipennis Kieffer, 1910 X
(3) P. africanus (Brues, 1924) X
(4) P. albitarsatus Sun & Sheng, 2007 X
(5) P. albosignatus (Kieffer, 1911) X
(6) P. ambiguus (Schletterer, 1890) X
(7) P. angularis (Crosskey, 1953) X
(8) P. annulatus (Kieffer, 1911) X
(9) P. anteala Smith, 2008 X
(10) P. aquilus Smith, 2008 X
(11) P. arcuatus Kieffer, 1904 X
(12) P. argutus Smith, 2008 X
(13) P. arizonicus (Townes, 1950) X
(14) P. auricomus Smith, 2008 X
(15) P. barbeyi (Ferrière, 1933) X
(16) P. bilobatus (Provancher, 1878) X
(17) P. boninensis Konishi, 1989 X
(18) (†)P. bradleyi (Brues, 1910) (Miocene) X
(19) P. brasiliensis (Kieffer, 1911) X
(20) P. californicus (Townes, 1950) X
(21) P. canadensis (Townes, 1950) X
(22) P. candidus Smith, 2008 X
(23) P. capitalis (Schletterer, 1890) X
(24) P. caudatus Szépligeti, 1903 X
(25) P. chlapowskii Kieffer, 1900 X
(26) P. cingulatus (Westwood, 1841) X
(27) P. colombianus Smith, 2005 X
(28) P. compressus (Spinola, 1808)
=Aulacus (Pristaulacus)holtzi Schulz, 1906
=Aulacus obscuripennis Westwood, 1841
X
(29) P. comptipennis Enderlein, 1912 X X
(30) P. concolor (Schletterer, 1890) X
(continued next page )
56 Invertebrate Systematics G. F. Turrisi et al.
Appendix 3. (continued )
Genus Aulacus Jurine, 1807
Palaearctic
Nearctic
Oriental
Afrotropical
Neotropical
Australasian
(31) P. cordatus (Schletterer, 1890) X
(32) P. cordiformis (Crosskey, 1953) X
(33) P. curryi Jennings, Austin & Stevens, 2004 X
(34) P. davisi Jennings, Austin & Stevens, 2004 X
(35) P. decemdentatus Kieffer, 1906 X
(36) P. decolorus Smith, 2008 X
(37) P. decorus Smith, 2008 X
(38) P. disjunctus Kieffer, 1904 X
(39) P. duporti Kieffer, 1921 X
(40) P. edoardoi Turrisi, 2007 X
(41) P. editus (Cresson, 1880) X
(42) P. emarginaticeps Turner, 1922 X
(43) P. erytrocephalus Cameron, 1905 X
(44) P. excisus Turner, 1922 X
(45) P. fasciatipennis Cameron, 1906 X
(46) P. fasciatus (Say, 1829) X
(47) P. femurrubrum Smith, 2005 X
(48) P. fiebrigi Brèthes, 1909 X
(49) P. flavicrurus (Bradley, 1901) X
(50) P. flavipennis (Cameron, 1887) X
(51) P. flavoguttatus (Westwood, 1851) X
(52) P. formosus (Westwood, 1868) X
(53) P. foxleei (Townes, 1950) X
(54) P. fulvus Turner, 1918 X
(55) P. fuscocostalis Turner, 1918 X
(56) P. galitae (Gribodo, 1879)
=Aulacus bimaculatus Kieffer, 1900
=P. immaculatus Kieffer, 1904
=P. bimaculatus arozarenae Ortega & Baez, 1985
X
(57) P. gibbator (Thunberg, 1822)
=Aulacus sibiricola Semenow, 1892
X
(58) P. gloriator (Fabricius, 1804)
=P. holzschuhi Madl, 1990
X
(59) P. guerini (Westwood, 1851) X
(60) P. haemorrhoidalis (Westwood, 1851) X
(61) P. haemorrhoidellus (Westwood, 1868) X
(62) P. hespenheidei Smith, 2008 X X
(63) P. insularis Konishi, 1990 X
(64) P. intermedius Uchida, 1932 X
(65) P. irenae (Madl, 1990) X
(66) P. iridipennis (Cameron, 1900) X
(67) P. karinulus Smith, 2001 X
(68) P. kiunga Jennings & Austin, 2006 X
(69) P. kostylevi (Alekseyev, 1986) X
(70) P. krombeini Smith, 1997 X
(71) P. laloki Jennings & Austin, 2006 X
(72) P. lateritius (Shuckard, 1841) X
(73) P. leviceps (Kieffer, 1911) X
(74) P. lindae Turrisi, 2000 X
(75) P. longicornis Kieffer, 1911 X
(76) P. maculatus (Schletterer, 1890) X
(77) P. major Szépligeti, 1903 X
(78) (†)P. mandibularis Brues, 1933 (Oligocene) X
(79) P. melleus (Cresson, 1879) X
(80) P. memnonius Sun & Sheng, 2007 X
(81) P. mexiuni Smith, 2008 X X
(continued next page )
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 57
Appendix 3. (continued )
Genus Aulacus Jurine, 1807
Palaearctic
Nearctic
Oriental
Afrotropical
Neotropical
Australasian
(82) P. minor (Cresson, 1880) X
(83) P. mouldsi Jennings, Austin, Stevens, 2004 X
(84) P. montanus (Cresson, 1879) X
(85) P. morawitzi (Semenow, 1892) X
(86) P. mourguesi Maneval,1935 X
(87) P. niger (Shuckard, 1841) X
(88) P. nigricoxae Smith, 2008 X
(89) P. nigripes Kieffer, 1911 X
(90) P. nobilis (Westwood, 1868) X
(91) P. occidentalis (Cresson, 1879) X
(92) P. omninoniger Smith, 2008 X
(93) P. oregonus (Townes, 1950) X
(94) P. ornatus Kieffer, 1913 X
(95) P. pacificus (Cresson, 1879) X
(96) P. paglianoi Turrisi, 2007 X
(97) P. parkeri Smith, 2008 X
(98) P. patrati (Audinet-Serville, 1833) X
(99) P. pieli Kieffer, 1924 X
(100) P. pilatoi Turrisi, 2006 X
(101) P. polychromus Kieffer, 1906 X
(102) P. porcatus Sun & Sheng, 2007 X
(103) P. postala Smith, 2008 X
(104) (†)P. praevolans (Brues, 1923) (Oligocene) X
(105) P. proximus Kieffer, 1906 X
(106) P. punctatus Smith, 2005 X
(107) P. punctum Smith, 2008 X
(108) P. resutorivorus (Westwood, 1851) X
(109) P. rex (Benoit, 1984) X
(110) (†)P. rohweri Brues, 1910 (Miocene) X
(111) P. rubidus (Schletterer, 1890) X
(112) P. rubriventer (Philippi, 1873) X
(113) P. ruficeps (Westwood, 1851)
(= P. bicornutus Schletterer, 1889)
X
(114) P. ruficollis (Cameron, 1887) X
(115) P. ruficruris Smith, 2008 X X
(116) P. rufipes Enderlein, 1912 X
(117) P. rufipilosus Uchida, 1932 X
(118) P. rufitarsis (Cresson, 1864) X
(119) P. rufobalteatus Cameron, 1907 X
(120) P. rufus (Westwood, 1841) X
(121) P. ryukyuensis Konishi, 1990 X
(122) (†)P. secundus (Cockerell, 1916) (Miocene) X
(123) P. sexdentatus Kieffer, 1904 X
(124) P. signatus (Shuckard, 1841) X
(125) P. singulus Smith, 2008 X
(126) P. smithi Turrisi, 2006 X
(127) P. spinifer (Westwood, 1868) X
(128) P. stangei Smith, 2008 X
(129) P. stephanoides (Westwood, 1841) X
(130) P. stigmaterus (Cresson, 1864) X
(131) P. stigmaticus (Westwood, 1868) X
(132) P. strangaliae Rohwer, 1917 X
(133) P. tamaulipas Smith, 2008 X
(134) P. tenuis Smith, 2008 X
(135) P. thoracicus (Westwood, 1841) X
(136) P. tonkinensis (Turner, 1919) X
(continued next page )
58 Invertebrate Systematics G. F. Turrisi et al.
Appendix 3. (continued )
Genus Aulacus Jurine, 1807
Palaearctic
Nearctic
Oriental
Afrotropical
Neotropical
Australasian
(137) P. torridus (Bradley, 1908) X
(138) P. totoferrugineus Smith, 2008 X
(139) P. townesi Smith, 2008 X
(140) P. tria Smith, 2008 X
(141) P. triclora Smith, 2008 X
(142) P. tricolor Szépligeti, 1903 X
(143) P. tridentatus Smith, 2005 X
(144) P. tuberculiceps (Turner, 1919) X
(145) P. unimacula Smith, 2008 X
(146) P. variegatus (Shuckard, 1841) X
(147) P. violaceus (Bradley, 1905) X
(148) P. virga Smith, 2008 X
(149) P. zhejiangensis He & Ma, 2002 X
(150) P. zonatipennis Roman, 1917 X
Systematics of the parasitoid wasp family Aulacidae Invertebrate Systematics 59
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