A revision of the westwoodiine genus Pergaphaga ( Hymenoptera , Ichneumonidae , Ctenopelmatinae )

Pergaphaga Gauld, a genus of ctenopelmatine Ichneumonidae endemic to Australia, is revised. Th e only previously described species, Pergaphaga nigra Gauld, is redescribed. Th ree additional species are newly described: Pergaphaga iangauldi Cammack & Wharton, sp. n. from New South Wales, ACT, and Victoria, P. leaski Wharton, sp. n. from Victoria, and P. xanthops Wharton, sp. n. from New South Wales and ACT. Keys to species and a phylogenetic analysis are presented. Morphological terms are linked to the Hymenoptera Anatomy Ontology.


Introduction
described the tribe Westwoodiini as part of his revised classifi cation of the Ctenopelmatinae, and included four genera (Westwoodia Brullé, Scolobatina Roman, Hypopheltes Cushman, and Megaceria Szépligeti), at that time known only from Australia.Gauld (1984) listed four characteristics shared by members of the Westwoo-diini and the more widespread but extralimital Scolobatini (which included Scolobates Gravenhorst, Onarion Townes, and Physotarsus Townes).On the basis of these shared features, Gauld (1984) combined all of the genera in a single tribe, the Scolobatini.Gauld (1984) removed considerable confusion regarding the Australian fauna by describing three new genera within his expanded concept of Scolobatini, transferring Megaceria Szépligeti, which had been misidentifi ed by Townes (1970), to Euryproctini, and treating Scolobatina as a synonym of Westwoodia.Zhaurova and Wharton (2009) retained Westwoodiini and Scolobatini as separate tribes based on a reassessment of morphological characters, described one new genus, and removed Tasmabates Gauld from Westwoodiini.Th e Westwoodiini, as delimited by Zhaurova and Wharton (2009), is one of the smallest tribes of the ichneumonid subfamily Ctenopelmatinae, consisting of fi ve genera but only nine described species (Townes 1970, Gauld 1984, Gupta 1987, Wharton et al. 2008, Zhaurova and Wharton 2009).Gauld (1984) indicated that this group of genera was endemic to Australia.Zhaurova and Wharton (2009) subsequently recorded a single specimen of one of these genera, Hypopheltes, from southern Papua New Guinea, a logical range extension for westwoodiines.Host records are available for all fi ve genera, including Pergaphaga Gauld, and are summarized by Townes (1970), Gauld (1984), Wharton et al. (2008), and Zhaurova and Wharton (2009).All recorded hosts are in the subfamily Perginae of the sawfl y family Pergidae (Hymenoptera).
When Gauld (1984) transferred Megaceria to Euryproctini, he noted that material previously misidentifi ed as Megaceria represented an undescribed genus, which he then described as Pergaphaga.Townes (1970) and Short (1978), both using the name Megaceria, provided fi gures of the adult and larval mouthparts, respectively, of the type species of Pergaphaga, P. nigra Gauld. Carne (1969), under the name "?Hypopheltes," recorded P. nigra as a parasitoid of Perga affi nis Kirby.Elliot and Bashford (1995) reared an undetermined species of Pergaphaga from Pergagrapta bella (Newman) in Tasmania.Although the plant host was not identifi ed for this rearing record, Elliot and Bashford (1995) list Eucalyptus amygdalina Labill., E. paucifl ora Sieber ex Spreng., and E. viminalis Dehnh.as the only plant hosts of P. bella in Tasmania.Aside from host records from label data reported by Gauld (1984) and repeated in Zhaurova and Wharton (2009), these are the only published data on the biology of Pergaphaga of which we are aware.Pergaphaga is thus far known only from southeastern Australia.Gauld (1984) described only one species in Pergaphaga, but stated that he had seen three others that diff ered from the type species in coloration as well as venation.Gauld (1984) also stated that Pergaphaga is most closely related to Hypopheltes.Although Gauld (1984) did not provide any supporting data for this relationship, these two appear together in the last couplet of his key to Australian ctenopelmatine genera.Quicke et al. (2009) showed Pergaphaga as the sister-group to Hypopheltes in a larger analysis of ichneumonid relationships, but they did not have Dictyopheltes Gauld in their analysis.Zhaurova and Wharton (2009) noted that Pergaphaga more closely resembles Dictyopheltes than Hypopheltes, based on the more heavily sculptured body and the reduced glymma.Pergaphaga was paraphyletic relative to Dictyopheltes in the two strict consensus trees that Zhaurova and Wharton (2009) presented but they considered their analyses preliminary because so many species were undescribed.Our objective is to present comparative descriptions for all known Pergaphaga and thereby facilitate a better understanding of relationships among the Westwoodiini.

Materials and methods
Specimens.We borrowed specimens from the following institutions for use in this study: Australian National Insect Collection, Canberra (ANIC), Th e Natural History Museum, London (BMNH), Museum of Victoria, Melbourne (MVMA), and Queensland Museum, Brisbane (QMBA).
In the materials examined section under each species description, we record label data for the holotype exactly as they appear on the labels.We use a more standardized format for paratypes, additional specimens examined, and published data for specimens not examined.Detailed label data are available on the taxon pages via the website http://peet.tamu.edu/projects/8/public/site/ich/home/otus_by_taxon/27303.

Figures.
Most images were acquired digitally using Syncroscopy's AutoMontage® software, in combination with a ProgRes 3008 digital camera mounted on a Leica MZ APO dissecting microscope.Figures 1 and 2 are modifi ed from Townes (1969).All images were further processed using various minor adjustment levels in Adobe Photoshop® such as image cropping and rotation, adjustment of contrast and brightness levels, color saturation, and background enhancement.Automontage images are available in color and high resolution at http://peet.tamu.edu/projects/8/public/site/ich/home/otus_by_taxon/27303.Several of the images were previously published in a companion paper by Zhaurova and Wharton (2009).
Database management, digital dissemination, and ontology reference.Character by OTU matrices, illustrations, keys, and free-text diagnoses for morphospecies were assembled in mx, a web-based content management system that facilitates data management and dissemination for taxonomic and phylogenetic works (e.g.Yoder et al. 2006).Th e mx project is open source, with code and further documentation available at http://sourceforge.net/projects/mx-database/.Data pertinent to this work, including interactive matrices, specimen-level data, and a multiple entry key, are available at http://peet.tamu.edu/projects/8/public/site/ich/home/otus_by_taxon/27303.
Hyperlinks on terms reference anatomical entities in the Hymenoptera Anatomy Ontology (HAO, version "06:11:2009 15:48";Yoder et al. 2009, see http://hymao.org).Th e HAO is a hierarchy of logically related entities (morphological parts).It allows multiple labels or terms (e.g."propodeum") to point to the defi nition of a morphological feature.Th is allows synonymous or otherwise confusing labels to unambiguously reference a morphological feature.Note that hyperlinked terms may resolve to a feature that itself has another label.Th is does not imply synonymy or preference of a given label, it is simply how the internal logical structure of the ontology is managed.Hyperlinked terms point to the HAO as archived in BioPortal (Musen et al. 2008).Labels and terms have further meaning in the context of the larger ontology, as such their meaning is specifi c to the cited version.Th is versioning further allows for the precise encapsulation of meaning at a specifi c reference point.All versions of the HAO are archived at the OBO Foundry (http://obo.svn.sourceforge.net/viewvc/obo/ontologies/trunk/HAO/, Smith et al. 2007).

Phylogenetic analysis.
Based on Zhaurova and Wharton (2009), we used Dictyopheltes as the outgroup for analyzing relationships among the four known species of Pergaphaga.Two species of Dictyopheltes were used to encompass variation within the genus.Characters and character states are presented under the results and discussion section.Characters 4 (notauli), 9 (propodeum), and 14 (wing color) were included because of purported (Gauld 1984, Zhaurova andWharton 2009) or potential value in supporting monophyly of either Dictyopheltes or Pergaphaga.We employed alternative character state codings for the propodeal carination, as explained below, to explore the impact of diff erent hypotheses of character evolution on our overall assessment of relationships.We also explored the eff ect of adding and deleting a character as noted below under the results section.Th e morphological dataset was analyzed under parsimony with NONA using the WinClada interface (Nixon 2002).For comparison, we also used PAUP version 4.0b10.In all analyses, all multistate characters were treated as unordered.Bootstrap values were generated using 200 replicates, with 10 searches per replicate, holding 1 tree per search.1-4) is essentially that used in Wharton et al. (2008) and Zhaurova and Wharton (2009), and is generally adapted from Gauld (1991Gauld ( , 1997) ) and more selectively from Townes (1969Townes ( , 1970)).We provide brief explanations in the following paragraphs where usage varies from or is in addition to that of Gauld (1991), or where elaboration may be helpful, a hyperlinked reference to the HAO.

Terminology and measurements. Terminology (Figs
Body length and antennal length, as given in the descriptions, are approximations of total length because of varying positions of the head and postmortem diff erences in telescoping and position of the metasoma.Wing length is measured from the base of the Costa to the apex of the wing and thus does not include the humeral plate and tegula.Abscissae are measured from the middle of their junctions.Length of the fi rst fl agellomere does not include the basal annellus, and all widths (fl agellomeres and tarsomeres) are taken at midlength.Th e face is the area between the anterior margin of the toruli and the anterior tentorial pits.We treat the face as separate from the clypeus (unlike Gauld 1991), with an epistomal sulcus, extending between the anterior tentorial pits, dividing the two.Th e antennal sockets are called toruli and the antennal tyloid on the lateral part of fl agellomere 1 (Figs 14, 15) is a distinctive patch of placoid sensilla (= placode or multiporous plate sensilla) used by Gauld (1984) to characterize his Scolobatini s. l. (= West- woodiini + Scolobatini of Zhaurova and Wharton 2009).An interantennal process (in the form of an elevated fl ange) extends posteriorly onto the frons from between the antenna in many westwoodiines and in two of the four species of Pergaphaga.Names for propodeal carinae (Fig. 2) and wing veins (Figs 3, 4) follow Gauld (1991)  (1991, 1997) did not label the fi rst abscissa of Cu1 in the hind wing, but we have done so in Fig. 4 for clarifi cation since the length of this abscissa relative to the length of cu-a is used in the descriptions.Townes (1969) referred to the two abscissae combined as the nervellus.Th e mesopleuron in Pergaphaga and other westwoodiines has a broad, longitudinal impression for reception of the mid femora similar to the feature used by Townes (1970) to defi ne the perilissine genera Opheltes Holmgren and Metopheltes Uchida.Th is is diff erent from the sternaulus (Wharton 2006) and is referred to in the descriptions below as the mesopleural depression (best illustrated in Fig. 27, somewhat below the arrow), following the usage of Townes (1970).We treat the metasoma as consisting of the petiole (T1 + its sternite, S1) and the gaster (remainder of metasoma posteriorad petiole).Use of the term gaster follows Sharkey and Wharton (1997) and facilitates descriptions of color patterns.We refer to the apical, expanded part of T1 as the postpetiole following Gauld (1991).T2 and T3 refer to the second and third metasomal terga, respectively.We use the term epipleura for the enlarged, membranous region between the tergum and sternum on the fi rst two metasomal segments, as described in some detail in Zhaurova and Wharton (2009).Th e defi nition of the following terms can be found in the HAO by following the associated hyperlinks: malar space 1 , orbital bands 2 , occipital carina 3 , hypostomal carina 4 , epicnemial carina 5 , mesoscutum 6 , notauli 7 , scuto-scutellar groove 8 , fore wing areolet 9 , bulla 10 , basitarsus 11 , trochantellus 12 , and glymma 13 .Note that the HAO is a hierarchy of classes, or things, and that multiple labels (or terms, e.g."scuto-scutellar groove") can point to the same class, or defi nition of a part.For example, we prefer to use the label "scuto-scutellar groove" for the concept in the HAO which has the label "scutoscutellar sulcus".

Results and discussion
Characters and character states.Th e following characters and character states were used to assess relationships among the species of Pergaphaga in the phylogenetic analyses presented in Figs 47-48.Brief explanations are provided in most cases after descriptions of the character states.
Th e diff erence between the two character states is subtle, and diffi cult to assess without side by side comparisons.Th e diffi culty is compounded by the fact that the face is entirely black in P. nigra but has yellow orbital bands, often contrasting with darker coloration medially, in the other three species of Pergaphaga.
2. Frons laterally.(0) distinctly elevated (Fig. 24); (1) fl at or nearly so.Th e anterior portion of the frons adjacent the compound eye is distinctly elevated in some species, with the elevation delimited medially by a weak to strong postantennal depression.Th is character co-varies with character 1 in most species, but not in P. nigra.
In some species of Pergaphaga, as in many other westwoodiines, an interantennal process in the form of an elevated median fl ange extends between the antennae onto the median part of the frons, where it usually bifurcates or trifurcates posteriorly.In westwoodiines, maximum elevation is usually immediately posteriorad the posterior margins of the toruli.In one of the species of Dictyopheltes, the median part of the fl ange is nearly absent, but the bifurcating arms are well developed (Zhaurova and Wharton 2009) and thus both species of Dictyopheltes have been coded as having strongly elevated fl anges.
Th e mesopleural depression is usually extensively punctate in Pergaphaga, with the punctures tending to coalesce anteroventrally and the sculpture often becoming rugulose posteroventrally.Th e ventral part of the mesopleural depression is more heavily sculptured throughout in P. nigra (Fig. 29) but is clearly and distinctly punctate medially in the other three species (e. g.Fig. 30), with the punctures discrete but with punctation varying in density among species.
Th e pleural carina is always well developed anteriorly in Pergaphaga.In most species of Pergaphaga, however, it is greatly reduced posteriorly, either completely lost, replaced by a shallow groove, or present in the normal position as a very faint carina.In some specimens, the pleural carina appears to fuse with the lateral-most portion of the anterior transverse carina, curving medially around the ventral side of the spiracle to join the lateral longitudinal carina.Th is short transverse section is shown in Fig. 2 connecting the pleural and lateral longitudinal carinae, but the pattern of propodeal carinae in Pergaphaga is otherwise very diff erent than what is shown in this fi gure.A clear carina is present in one of the three specimens of P. xanthops Wharton, sp.n., and it has been coded as variable for this reason.
Th e distinctly elevated anterior transverse carina of P. nigra is variously replaced by weaker, irregular, transverse ridges in the other three species of Pergaphaga.In P. iangauldi Cammack and Wharton, sp.n., however, traces of a more distinct carina are often visible, and the basal median depression, in particular, is usually margined posteriorly by a carina.We coded P. iangauldi separately (state 1, Table 1) for our initial analysis, but also performed an analysis with this species coded either as in P. nigra or as in P. xanthops + P. leaski Wharton, sp.n.
Examination of a series of specimens from Murrumbeena (see remarks under P. iangauldi), suggests that in Pergaphaga, state 2 is derived from state 1.No such intermediates were apparent in Dictyopheltes (the outgroup), and character states were left unordered in all analyses.
Pergaphaga nigra and the species of Dictyopheltes tend to have darker legs than the other three species of Pergaphaga, with the pale basal ring thus strongly contrasting.
Th e hind tibia is yellow to orange in the other three species of Pergaphaga (Figs 42-44), though not always uniformly so.
In P. nigra, the petiole and all terga of the gaster are somewhat variable, but usually very dark brown to black anteriorly grading to dark reddish brown posteriorly, with pale apical margin, this margin tending to be broader in females than males.In the other three species of Pergaphaga, the gaster and parts of the petiole (especially postpetiole) are much lighter in coloration for the most part (Fig. 43).Pergaphaga iangauldi is slightly more variable (Fig. 42), and we have attempted to refl ect this variation in the coding for this species.In those species for which more than one male was available for examination, males were somewhat more variable in coloration than females, but our sample is very small.14.Color pattern of fore wing.(0) weakly infumate anteriorly, fading to hyaline posteriorly, without infumate apical spot (Fig. 36); (1) hyaline with infumate apical spot (Fig. 39).
Th e infumate apical spot is present in all known specimens of Pergaphaga, as best illustrated in Fig. 39.Th e spot does not show up as prominently in some of the other fi gures because of background contrast.
As indicated below in the redescription of P. nigra, there is some evidence for sexual dimorphism in proportions of the hind tibia, but the sample size is small.Assessment of relationships.All initial analyses produced a single tree (length = 21, CI = 90, RI = 86) with P. nigra as the sister group to the remaining species of Pergaphaga (Figs 47,48).Heuristic and exact searches produced trees of the same length.Th e clade comprising P. xanthops + (P.iangauldi + P. leaski) is characterized by weaker mesopleural and propodeal sculpture, longer hind basitarsus, and distinctly paler coloration.Loss of the interantennal fl ange and slight diff erences in facial features unite P. leaski and P. iangauldi relative to P. xanthops within this clade.Th e results suggest a single loss of the interantennal fl ange, but a complete loss followed by a gain of the fore wing areolet.Th e homoplasy resulting from minor incongruencies among characters is refl ected in the bootstrap values shown in Fig. 48.Bootstrap values shown in Fig.48 were generated using WinClada.Th ose generated via PAUP were equal to or very slightly higher for the P. xanthops + (P.iangauldi + P. leaski) clade and the P. iangauldi + P. leaski clade.Th ere was virtually no eff ect when changing the coding of the propodeum in P. iangauldi to match that of P. xanthops and P. leaski.When the coding was changed so that P. iangauldi had the same character state as P. nigra, the relationships remained the same, but bootstrap support values for the P. iangauldi + P. leaski clade not surprisingly dropped.
We were hesitant about using the epipleura as a character in our analyses despite the possibility that it may be phylogenetically informative (Townes 1970, Gauld 1984, 1997, Zhaurova and Wharton 2009).As noted by Zhaurova and Wharton (2009), there are problems in interpretation of this feature because its appearance varies with state of preservation.Th e epipleura appear to be better developed in P. nigra and P. iangauldi than in the other two species, though this is not evident in all specimens.In P. leaski and P. xanthops, which are smaller species, the epipleura are generally not apparent, or in some specimens only weakly indicated, but we are unable to determine if this is simply a preservation artifact.For these reasons, we excluded the epipleura from our initial analyses and delineation of characters and character states.We did, however, explore the eff ect this character might have on assessment of relationships within Pergaphaga by coding it for a separate PAUP analysis, assuming no preservation artifacts.Relationships remained the same as those presented in Fig. 48, but support for the P. iangauldi + P. leaski clade was greatly reduced.
Th ere is relatively weak support for the monophyly of Pergaphaga, though bootstrap values were distinctly higher (75 vs. 58) in the PAUP analysis.However, demonstration of the monophyly of Pergaphaga was not a goal of this analysis.Monophyly of Pergaphaga and Dictyopheltes is treated in the next section, under the diagnosis for Pergaphaga.

Pergaphaga Gauld, 1984
Pergaphaga Gauld, 1984: 15).Occipital and hypostomal carinae meeting ventrally well before base of mandible (Fig. 25).Mesoscutum densely punctate throughout; notauli deep throughout, including at anterior margin (Fig. 27), V-shaped, converging posteriorly in broad, shallow depression separated from scuto-scutellar groove by a weak elevation; mesopleural depression broad and distinct.Propodeum varying from partially and weakly to extensively and strongly rugose (Figs 33-34), but always with some carination visible.Inner hind tibial spur almost twice length of outer spur.Fore wing 2m-cu with a single bulla.Petiolar tergosternal sulcus located ventrally when viewed in profi le.Glymma represented by a shallow dorsal depression anteriorly (Fig. 45).S1 extending more than half distance to spiracle, usually to or nearly to level of spiracle.Metasomal segments 1-2 and more rarely 3 with epipleura bare and membranous when visible, often collapsed and extending outwardly as fl eshy protrusions in dried specimens.Female gaster laterally compressed apically from middle of T3.In lateral view, cerci attached ventrad middle of posterior aspect of gaster.All four of the known species have hyaline wings with an infumate spot at the tip of the fore wing.
Pergaphaga can be diff erentiated from other Ctenopelmatinae by a combination of characters specifi c to Westwoodiini, including fore wing RS+2r arising from or near the base of the stigma , cerci of females ventrally displaced, and fi rst fl agellomere with a large tyloid laterally.Th e tyloid is in the form of a bare patch of numerous (> 20), irregularly arranged placoid sensilla .
As noted by Zhaurova and Wharton (2009), Pergaphaga is perhaps the least readily characterized of the known westwoodiines, and the species most closely resemble those currently placed in Dictyopheltes.Th e species of Pergaphaga and Dictyopheltes are characterized relative to other westwoodiine genera by the shallow to indistinct glymma (Fig. 45), with the petiole thus resembling that of certain Euryproctini.Gauld (1984), in fact, stated that Pergaphaga and Dictyopheltes lacked a glymma, and he used this feature in his key to the Australian genera of ctenopelmatines.A glymma-like depression is present in nearly all individuals that we examined, but is much diff erent in appearance than the deeper glymma of Westwoodia, Gauldia, and Hypopheltes.It is narrow, shallow, and almost slit-like in some specimens, and never as distinct as it is in these other genera.
The sternite of the petiole is longer in Pergaphaga relative to that in Dictyopheltes.Additionally, the species of Pergaphaga are less heavily sculptured than those of Dictyopheltes, and retain at least some visible carination on the propodeum .Th e most distinctive feature of the propodeum is the presence of a straight to very weakly arched transverse carina or series of low, transverse ridges extending between the propodeal spiracles that is not found in other westwoodiines, and which is variously developed in the new species described below.Th e monophyly of Pergaphaga is thus largely supported by the patterns of propodeal carination and wing coloration.
Biology.Th e species of Pergaphaga have been reared from pergine sawfl ies feeding on Eucalyptus, and specifi cally from Perga affi nis (Carne 1969), Pergagrapta bella (Elliot and Bashford 1995) and Pergagrapta gravenhorstii (Westwood).Th e latter record is based solely on label data from the type series of P. leaski.Th ere are collection or rearing records from every month except September and November, but the vast majority are from February through June.
Pergaphaga is known only from southeastern Australia, ranging from South Australia, through Victoria and north about midway through New South Wales.In addition to the specimens recorded below under the species descriptions there are published records of Pergaphaga from Tasmania (Elliot and Bashford 1995), but we have not seen this material.Th e specimens of P. nigra labeled as South Australia (BMNH) have no additional data, and we are unable to pinpoint the locality further.
Remarks.Pergaphaga is similar in size and general appearance to species of Megaceria, a more commonly encountered ctenopelmatine in the tribe Euryproctini (Gauld 1984).Both are Australian endemics, and the two have been confused in the past, prior to the detailed study of the Australian fauna by Gauld (1984).Th e species of Megaceria lack the distinct tyloid at the base of the outer side of the fi rst fl agellomere.Th e fi rst fl agellomere also tends to be longer in Megaceria, with RS generally arising more distally from the stigma and terminating closer to the wing tip.Additionally, the petiole is more completely tubular in Megaceria, with no obvious sulcus between the fused tergum and sternum, the spiracle is more anteriorly displaced, and there is a broader, more distinct bridge separating the foramen of the petiole from the coxal cavities.Th e shape of the forewing areolet is distinctive in Megaceria, unlike the various forms seen in Pergaphaga.In Megaceria, ornamentation on the frons varies, as does the development of lateral carinae on the scutellum and there's an unusual amount of variation in the trough and associated carinae and projections along the margin of the propodeum and metanotum.According to Gauld (1984), there are three described species of Megaceria and at least 10 undescribed species.
Head.Clypeus (Fig. 17) 2.5-2.7 times as broad as long; slightly undulating; convex dorsomedially and along midline, impressed ventrolaterally, except raised and thickened at extreme lateral margin; ventral margin broadly truncate, slightly thickened medially; surface punctate and weakly rugulose on shagreened background, punctures coalescing, especially ventrally; epistomal sulcus indistinct.Malar space 0.6-0.7 times basal width of mandible, coarsely granular and shagreened.Lower gena deeply and densely punctate on fi nely shagreened background, becoming more sparsely punctate and polished dorsally, upper gena with punctures separated by 2-3 times their diameter, more densely punctate along occipital carina, more sparsely punctate adja- cent eye at mid eye height.Face (Fig. 17) deeply and densely punctate, varying from rugose punctate to more densely granular rugose medially, surface slightly undulating transversely, weakly elevated medially, distinctly depressed near ventrolateral margin of toruli, nearly fl at between anterior tentorial pit and eye.Frons with median, elevated fl ange (Fig. 21) extending posteriorly beyond posterior margin of toruli, bifurcating posteriorly, the resulting depressions between fl ange and toruli and between bifurcating arms and median ocellus polished and often weakly striate; frons densely granular on fl attened portion between ocellar fi eld and eye, rugose punctate on distinctly elevated portion between antenna and eye (Fig. 24).Antenna with 40-46 fl agellomeres; fi rst fl agellomere at most 1.2 times longer than second, 3.0-3.7 times longer than wide, second fl agellomere 2.5-3.1 times longer than wide, tenth 1.8-2.2times longer Mesosoma.Pronotum laterally densely punctate (Fig. 27).Mesopleural depression rugulose punctate ventrally (Fig. 29), the punctures large, deep, coalescing; rounded lobe forming anterodorsal margin of mesopleural depression very fi nely, densely punctate, the punctures discrete but often touching or nearly so, much smaller than punctures on ventral part of mesopleural depression.Scutellum coarsely punctate.Posteromedian plate of metanotum densely, coarsely punctate to rugose punctate, never polished.Metapleuron medially densely rugose to rugose punctate.Propodeum in profi le (Fig. 31) with anterior and sharply declivous posterior fi elds distinctly separated by prominent anterior transverse carina extending between spiracles, the carina somewhat irregular, often rugose medially; base of propodeum, on either side of median depression, heavily sculptured as in surrounding areas (Fig. 33); pleural carina complete (Fig. 31): strongly elevated anterior portion extending posteromedially toward spiracle, then angled posteriorly near spiracle, less strongly elevated posteriorly, never touching spiracle; lateral longitudinal carina distinct posteriorly, absent or diffi cult to distinguish anteriorly; lateromedian portions of posterior transverse carinae often weakly indicated as elevated rugosities, forming a low to prominent tubercle at  anterior margin.T3 and T4 often uniformly densely punctate and short-setose, occasionally with median bare, impunctate line on T3.
Color (Figs 5,9,13,17,41).Head and body black, antenna entirely black; mandibles dark basally and apically, often reddish brown medially; metasoma with petiole and terga of gaster black basally, becoming reddish brown apically in half specimens examined, posterior margin always pale yellow, with yellow markings expanded along midline on laterally compressed terga; all coxae black, trochanters and trochantelli nearly always black; femora transitioning gradually or somewhat abruptly from darker ventrally to paler mid dorsally as follows: from dark brown ventrally and orange brown dorsally to orange ventrally and yellow orange dorsally, rarely uniformly colored, fore and mid Male.Essentially as in female except as follows: Hind basitarsus often slightly narrower, 8.5-10.0times longer than wide; terminal segments of gaster not as laterally compressed; fore trochanter more frequently (30%) brownish than in female.
Distribution and biology.Th is species is known only from southeastern Australia, with the known range of this species extending from South Australia through Remarks.Th e male specimen from Duntroon listed above as a paratype has a paratype label and an Ian Gauld det.label, but does not exactly match the information on paratypes provided in Gauld's (1984) original description.Th e specimen from Cookardinia, though listed by Gauld as a paratype, lacks a paratype label.Handwritten labels correctly give R.S. for initials of McInnes, but those typed labels with emergence dates of 1973 incorrectly give initials as R.B. Gauld (1984) lists an additional 10 female and 10 male paratypes from Murrumbateman (ANIC) that we did not see.Gauld also noted that one of the paratypes from Avoca bears a label indicating that it was the specimen fi gured by Townes (1970) as Megaceria.Observed variation was as great within populations as between them.Sculpture of the median part of the face varied from rugose punctate to more densely granular rugose and the hind tibia varied from black to orange in both Duntroon and Murrumbateman series.Th e female specimen from Duntroon shown in Fig. 5 illustrates the maximum extent of pale coloration on the apical margins of the terga among the material available for study.In one of the 10 females measured, the hind basitarsus was 10.0 times longer than wide; in one of the nine males examined, the areolet was not petiolate.
In addition to a generally more heavily sculptured propodeum relative to other species of Pergaphaga, P. nigra has the anterior transverse propodeal carina not only better developed but also generally more posteriorly displaced.
Head.Clypeus (Fig. 18) 2.6-2.7 times as broad as long; weakly and uniformly convex to nearly fl at in profi le, very weakly thickened along lateral margin; ventral margin broadly truncate to very weakly concave, very slightly thickened; surface punctate on weakly shagreened background, punctures deep, separated from one another by their diameter; epistomal sulcus weak medially but distinct.Malar space 0.45-0.55times basal width of mandible, strongly shagreened to fi nely granular-matt/punctate.Lower gena punctate, with punctures separated by 0.5-1.0 times their diameter, usually weakly shagreened, upper gena more polished and very slightly more sparsely punctate except more densely punctate along occipital carina.Face (Fig. 18) deeply and densely punctate, varying from punctate and weakly shagreened to more densely granular punctate medially, surface distinctly undulating transversely, elevated medially and laterally, distinctly depressed near ventrolateral margin of toruli, convex between anterior tentorial pit and eye (Fig 16).Frons lacking distinctly elevated interantennal fl ange of P. nigra, with at most a low, very short, weak, median carina (Fig. 23); frons distinctly elevated adjacent eye, the elevated area punctate and somewhat crescent-shaped, frons otherwise fl at with post-antennal area adjacent elevated lateral margin appearing weakly concave; surface often polished immediately behind scape, fi nely matt punctate posteriorly between posterior ocelli and eye, otherwise variously rugulose to rugose to densely granular.Antenna with 42-44 fl agellomeres; fi rst fl agellomere at most 1.2 times longer than second, 3.6-3.7 times longer than wide, second fl agellomere 2.8-3.1 times longer than wide, tenth 2.0-2.2 times longer than wide; tyloid of fi rst fl agellomere (Figs 14, 15) large, oval, extending 0.30-0.35length of fi rst fl agellomere.
Mesosoma.Pronotum laterally densely punctate.Mesopleural depression densely punctate to weakly rugulose or strigose punctate anteroventrally, the punctures large, deep, coalescing at least in part anteriorly but usually discrete though nearly adjacent medially; rounded lobe forming anterodorsal margin of mesopleural depression with similarly large, deep punctures, but with punctures often more widely spaced.Scutellum coarsely punctate.Posteromedian plate of metanotum varying from unsculptured to nearly so, polished.Metapleuron densely punctate with additional strigose sculpture in some individuals.Propodeum varying from weakly convex to nearly fl at in profi le, with very narrow anterior and elongate posterior fi elds often diff erentiated; base of propodeum, on either side of median depression, smooth, polished (as in Fig. 34); pleural carina nearly always incomplete: sharply defi ned anteriorly, touching ventrolateral corner of propodeal spiracle, discontinuous or nearly so between spiracle and weaker posterior fragment; anterior transverse carina often poorly diff erentiated among narrow band of transverse strigose sculpture, though sometimes distinct as a low ridge; median basal depression margined posteriorly with what may represent the median portion of anterior transverse carina; distinctly arched lateral portion of posterior transverse carina and posterior portion of lateral longitudinal carina forming large, polished, apicolateral area (Figs 32), junction of lateral longitudinal and posterior transverse carinae not tuberculate.Hind basitarsus 12.2-14.7 times longer than wide.Fore wing areolet very small, petiolate above, the stalk equal to or longer than both length and width of areolet (Fig. 38); 2m-cu variable, arising from middle (rarely) to extreme apex (more commonly) of areolet.Hind wing with 1st abscissa of Cu1 0.80-0.85times length of cu-a; cu-a reclivous.
Metasoma.Petiole with S1 not extending to level of spiracle.Postpetiole and T2-4 polished, almost completely bare and impunctate, with a few scattered punctures and setae, the latter concentrated laterally.
Color (Figs 6,10,16,18,42).Mostly black; mandible (except dark apical teeth), broad orbital bands on face extending onto anterior part of frons, clypeus (except small, irregular dark spot dorsomedially) and variously sized spot on lower gena bright yellow; scape, pedicel, and basal 19-21 fl agellomeres, tegula, legs from trochanter to apex, and most of gaster orange, with fore and mid legs often yellow orange; petiole completely, T2 extensively (holotype) to completely and base of T3 at least partially black in 3 specimens; gaster completely orange and petiole with irregular orange markings posteriorly in 2 specimens.
Male.Essentially as in female except as follows: Antenna with 41-43 fl agellomeres, fi rst fl agellomere 3.1-3.2times longer than wide; hind wing with 1st abscissa of Cu1 0.65-0.85times length of cu-a; anterior transverse carina less evident than in most females; terminal segments of gaster not laterally compressed; face medially and clypeus dorsomedially pale orange instead of black, basal 22-23 fl agellomeres orange, coxae, especially on fore leg, partly orange, gaster almost completely black in one specimen, almost completely orange in the other.
Distribution and biology.Known only from Victoria, ACT, and central New South Wales.Reared from an undetermined species of Perga (based on label data).
Diagnosis.Th is species is most readily recognized by the distinctively bicolored antenna, which is pale basally and dark apically.Pergaphaga xanthops also has bicolored antennae, but the pattern is reversed.Pergaphaga iangauldi lacks the elevated median fl ange on the frons as found in P. nigra and P. xanthops, and is thus more similar to P. leaski in this regard.In addition to diff erences in antennal color pattern, P. iangauldi has a more heavily sculptured mesopleuron than P. leaski.
Remarks.Th e female specimen from Bright is larger than members of the type series, with 46 fl agellomeres, the fi rst 30 of which are orange.Th e fore wing areolet is also distinctly larger (roughly intermediate in size between that of P. nigra and those from the type series of P. iangauldi), and the metapleuron is distinctly strigose in this specimen.Although considered a member of this species, it is not included as a paratype because of this variation.Th e following additional variation was noted among members of the type series: in one female, the fl agellum is brown basally rather than orange, the epicnemial carina does not extend dorsally to the level of the ventral corner of the pronotum in several specimens, and in the left wing of one of the females, Cu1 is 1.15 times longer than cu-a.Th e female paratype from Canberra has a slightly diff erent color pattern than the series from Murrumbeena, but unlike the specimen from Bright, sculpture and venation are the same and it is therefore included in the paratype series to emphasize color variation in this species.In the Canberra specimen, all terga of the gaster are black medially, face and clypeus are brownish yellow medially, fore and mid coxae are orange ventrally, and fi rst 32 fl agellomeres are orange.
One male specimen lacks a fore wing areolet in both wings.In the left wing, it is easier to see that that the absence of an areolet is due to the fusion of 2rs-m and 3rs-m rather than the loss of one or the other of these.Th is suggests that within Pergaphaga, at least, there is a trend toward gradual loss of the areolet through reduction in size and fusion of the adjacent cross veins.
Th e fi rst two labels on the paratype from Windsor are handwritten.Th e fi rst is diffi cult to read, with the month, day, and locality legible, but the remainder diffi cult to decipher and possibly misinterpreted.Th e second label is completely illegible.Th is and the specimen from Bright bear Pergaphaga det.labels by I. D. Gauld dated 1984.Th e paratype from Canberra is also the voucher specimen for the Pergaphaga sequence reported in Quicke et al. (2009).
Mesosoma.Pronotum punctate laterally on shagreened background, punctation slightly less dense than in P. nigra and P. iangauldi, punctures separated by 1.0-1.5 times their diameter.Mesopleural depression more sparsely punctate ventrally than in P. nigra and P. iangauldi, punctures widely separated, not coalescing medially to form either strigose or rugulose sculpture; rounded lobe forming anterodorsal margin of mesopleural depression with similarly large, well-spaced punctures.Scutellum slightly more fi nely punctate than in P. nigra and P. iangauldi.Posteromedian plate of metanotum polished, varying from distinctly punctate anteriorly to sparsely, indistinctly punctate throughout.Metapleuron medially fi nely granular-matt and sometimes weakly punctate.Propodeum weakly convex in profi le, anterior and posterior fi elds not diff erentiated; base of propodeum, on either side of median depression, smooth, polished (Fig. 34); pleural carina incomplete: sharply defi ned anteriorly, extending to propodeal spiracle, usually absent posteriorly; anterior transverse carina not distinguishable from surrounding narrow band of transverse ridges; distinctly arched lateral portion of posterior transverse carina and posterior portion of lateral longitudinal carina forming apicolateral area similar to but usually weaker than in P. iangauldi, junction of lateral longitudinal and posterior transverse carinae not tuberculate.Hind basitarsus 10.6-12.0 times longer than wide.Fore wing areolet very small, triangular, petiolate above, the stalk equal to length and longer than width of areolet (Fig. 39); 2m-cu arising near but distad middle.Hind wing with 1st abscissa of Cu1 0.75-0.85times length of cu-a; cu-a reclivous.
Color (Figs 7,11,19,43).Mostly black, antenna dark brown to black; mandible (except dark apical teeth), broad orbital bands on face extending onto anterior part of frons, at least lateral corners of clypeus, lower gena, tibia and tarsi of fore and mid legs, and tarsomeres 2-4 of hind legs bright yellow; tegula, trochanters, trochantelli, and femora of fore and mid legs darker yellow; trochanter, trochantellus, femur, and apical tarsomere of hind leg orange, hind tibia orange brown basally becoming orange over distal 0.25, hind basitarsus orange brown basally, yellow over distal 0.25; postpetiole, except along lateral margins, and entire gaster orange; middle of gena with small, dark reddish brown spot posteriorly.
Male.About as in female, but only known specimen in poor condition.Distribution and biology.Known from a single series of specimens reared from Pergagrapta gravenhorstii (Westwood) in Ballarat, Victoria.
Diagnosis.Th is species is readily recognized by the combination of orange gaster and entirely dark antenna.As in the larger-bodied P. iangauldi, an elevated median fl ange is absent on the frons.As noted above, P. iangauldi also has a more heavily sculptured mesopleuron than P. leaski.
Remarks.Th ere is some minor variation in color in the type series.Th e clypeus is mostly reddish brown in the holotype but extensively yellow in the paratypes, with a small reddish brown spot dorsomedially.
Th e species is named for Maurice Leask, who reared numerous westwoodiines from various pergid sawfl ies, including all of the members of the type series.Th is species is referred to as Pergaphaga sp. 1 in Gauld (1984).Description.Female (Figs 8,12).Length of body (exclusive of antenna) 10.7-11.0mm; of fore wing 9.7-10.5mm; of antenna 12.0-12.8mm.

Pergaphaga xanthops
Head.Clypeus (Fig. 20) 2.5-2.6 times as broad as long; weakly and uniformly convex to nearly fl at in profi le; ventral margin broadly truncate to very weakly concave, broadly but weakly thickened medially; surface punctate on weakly shagreened background, punctures deep, separated from one another by their diameter dorsally, nearly coalescing ventrally; epistomal sulcus distinct.Malar space 0.40-0.45times basal width of mandible, shagreened or polished, punctate near mandible, weakly granular near eye.Gena weakly to distinctly shagreened and punctate ventrally, polished or nearly so dorsally, punctures separated by 0.5-1.0 times their diameter ventrally, more fi nely punctate dorsally with punctures 1-2 times their diameter.Face (Fig. 20) deeply punctate laterally, densely granular punctate to granular rugose medially, surface very slightly undulating transversely, weakly elevated medially, weakly depressed near ventrolateral margin of toruli, nearly fl at between anterior tentorial pit and eye.Frons with median fl ange (Fig. 22) extending posteriorly beyond posterior margin of toruli, indistinctly bifurcating posteriorly, the resulting depressions between fl ange and toruli polished; frons rugose between posterior end of fl ange and median ocellus, strigose and fi nely granular to partly polished on fl attened portion between ocellar fi eld and eye, punctate on weakly elevated portion between antenna and eye.Antennal fl agellum with 36 fl agellomeres; fi rst fl agellomere 1.1 times longer than second, 3.1 times longer than wide, second fl agellomere 2.6 times longer than wide, tenth 2.2 times longer than wide; tyloid of fi rst fl agellomere large, oval, extending 0.4 length of fi rst fl agellomere.
Mesosoma.Pronotum laterally densely punctate around margins, fi nely granularmatt to granular rugulose medially.Mesopleural depression more densely punctate ventrally (Fig. 30) than in P. leaski, but punctures discrete, not coalesing to form either strigose or rugulose sculpture as in P. nigra, and anteroventrally as in P. iangauldi; rounded lobe forming anterodorsal margin of mesopleural depression very fi nely, densely punctate, the punctures discrete but often touching or nearly so.Scutellum fi nely punctate.Posteromedian plate of metanotum sparsely punctate, polished.Metapleuron medially granular-matt.Propodeum weakly convex in profi le, anterior and posterior fi elds very weakly diff erentiated; base of propodeum, on either side of median depression, smooth, polished (as in Fig. 34); pleural carina incomplete in one specimen, complete in the other: sharply defi ned anteriorly, extending to propodeal spiracle, absent or very weak posteriorly; anterior transverse carina not readily distinguishable within band of transversely strigose sculpture; posterior portion of lateral longitudinal carina arched medially to form nearly complete apicolateral area with portions of posterior transverse carina, junction of lateral longitudinal and posterior transverse carinae not tuberculate.Hind basitarsus 11.6-14.0times longer than wide.Fore wing areolet absent (Fig. 40).Hind wing with 1st abscissa of Cu1 0.70 times length of cu-a; cu-a reclivous.
Color (Figs 8,12,20,44).Mostly dark brown to black; mandible (except dark apical teeth), face (except small triangle extending anteriorly from between antennae), clypeus, malar space, broad band on gena adjacent eye, fore and mid legs (except coxae and apical tarsomere), and subapical fl agellomeres 19-31 bright yellow; mid and hind coxae dark brown dorsally, brown ventrally, fore coxa partly yellow ventrally; hind legs distad coxa light yellow brown, a little darker dorsally, with hind basitarsus and apical tarsomeres on all legs a little darker; gaster and apical margin of petiole orange.
Male.Essentially as in female except as follows: Clypeus punctate, polished, without shagreened sculpture; antenna with 39 fl agellomeres; face with black vertical band, the band narrower than adjacent yellow orbital bands; frons more polished, with less extensive rugose sculpture medially between median ocellus and most elevated portion of fl ange; apical fl agellomere weakly infumate, otherwise yellow from fl agellomere 19 to apex.
Distribution and biology.Australia; known only from ACT and a nearby locality in NSW.Eucalyptus paucifl ora is listed on the label as the host plant for the male paratype from NSW. Morrow et al. (1976) collected four species of Perginae (Pergidae), representing four diff erent genera, from this plant host at this locality.It is unclear from the label data whether this specimen was reared or simply collected from this plant, but no parasitoid rearings were indicated in Morrow et al. (1976).
Diagnosis.Th is species is readily recognized by the absence of a fore wing areolet and the bicolored antenna which is dark basally and pale apically/subapically.As in P. nigra and unlike the other two species, there is an elevated, Y-shaped fl ange extending between the antenna and onto the frons in P. xanthops.
Remarks.Th e apical teeth of the mandible are slightly shorter and more bluntly rounded in this species relative to the other three.Th is may be due to wear, however, as indicated by slight variation in size and shape of the teeth in the longer series of P. nigra available for study.Additionally, the epipleura are not as fl eshy in appearance in this species as they are in P. nigra and P. iangauldi.It is diffi cult to determine whether this is a true diff erence or a preservation artifact.Th e epicnemial carina is also weaker in this species than in the other three, and the dorsal extent is particularly diffi cult to discern.Th e clypeus appears somewhat diff erent in shape when Fig. 20 is compared to the fi gures of the other three species shown on the same plate.Th is is almost entirely due to the angle of view, with the face angled more ventrally in Fig. 20.
Th e underlying sculpture of the female paratype is not as fi nely granular or shagreened as it is in the holotype.Th is is especially noticeable on the malar space, lower gena and area extending between the eye and ocellar fi eld.
Th e species name is in reference to the extensively yellow face of the females.