Molecular Phylogeny, Species Delimitation and Taxonomic Revision of the Australasian Spider Wasp Genus Heterodontonyx Haupt, 1935 (Hymenoptera: Pompilidae) ()
1. Introduction
Pompilidae Latreille, 1805, commonly known as spider wasps or spider-hunting wasps, are a family of the order Hymenoptera (Superfamily Pompiloidea).There are five subfamilies within pompilid wasps [1] three of which (Pompilinae, Ceropalinae and Pepsinae) have Australasian distribution. Pepsinae of Australia are divided into the tribes Pepsini and Ageniellini and the Pepsini genus Heterodontonyx Haupt, is distributed in New Guinea, Australia, New Caledonia, and New Zealand [2]. Recently, molecular, and morphological data suggested some taxonomic changes involving the genera Heterodontonyx Haupt and Cryptocheilus Panzer that warranted taxonomic revision of both genera (Chavoshi, M. et al, 2024, unpublished). In this study, we utilized both molecular and morphological data to examine phylogeny and specific diversity of Heterodontonyx species and revise the genus. There are certain evidence for host shifts in spider wasps which are diversification contributing factors in this group [3].
2. Material and Methods
2.1. Taxon Sampling, DNA Extraction, Library Preparation, Pooling, and Sequencing
A total of 30 specimens covering almost all Heterodontonyx species (H. solomonis Turner and H. erythroura (Cameron) were not included in the phylogeny due to DNA library sequencing failure) included as in group in addition to one out group (POM044 in Table S1 and Figure 1). The material was identified to species level using the identification keys developed based on the species descriptions and careful specimen examination. One to three legs of adult spider wasps were cleaned and homogenized for proteolysis on the first day of extraction. Silica-column (Zymo product-Spin Columns II C-Cat No: 1920-050) were used on the second day to extract genomic DNA using the QIAGENDNeasy Blood & Tissue Kit (Cat No: 69504) as per manufacturer’s instruction. The purified DNA was resuspended in 25 ul of Tris-HCl 1 mM and stored at 4˚C. DNA quantification was carried out using Invitrogen Qubit dsDNA HS Assay Kit with Fluorometer Denovix DS-II followed by determination of DNA fragment length using Agilent HS Genomic DNA Kit DNF-488 with 12 Capillary Fragment Analyzer System (Advanced Analytical Technologies, Inc.). DNA library building was achieved using NEBNex Ultra II DNA Library Prep Kit for Illumina (Cat No: E7645) with 3 - 15 amplification cycles depending on the DNA input. As the final step, libraries were eluted in Tris-HCl 1 mM to reach equimolar concentration of 2 nM followed by post-PCR clean-up of the pooled libraries using 1× magnetic bead ratio (NEBNext Sample Purification Beads). Lastly, 25 - 200 ul of the pooled, cleaned, single-indexed (using NEBNext Multiplex Oligos for Illumina) libraries was submitted for whole-genome shotgun sequencing in one lane of NextSeq 550 System 300 cycles (paired end reads) at the ACRF Biomolecular Resource Facility (https://jcsmr.anu.edu.au/).
2.2. Data Processing, Sequence Alignment, Phylogenetic Assessment, and Species Delimitation
Raw reads were cleaned and trimmed with Trimmomatic [4] and quality checked with FastQC (Babraham Bioinformatics). The resulting trimmed reads were de-novo assembled using Trinity v2.1.1 [5]. ITS1, 5.8S, ITS2 reference sequences were obtained by mapping POM334 to a 5.8S sequence (accession no. X89481) from [6] with 25 iterations and was then used to as the reference sequence to extract ITS1, 5.8S, and ITS2 from assembled contigs. Phylogenetic analyses were carried out on the ITS dataset under maximum likelihood (ML). For ML analyses, we used IQ-TREE 1.7.0 [7] with Model Finder [8] and ultrafast bootstrap [9]. The phylogenetic tree was visualized in Figtree v.1.4.1 (http://tree.bio.ed.ac.uk/software/Figtree/). ITS matrix was then used to examine delimitation scheme with PTP and bPTP on the server (https://species.h-its.org/). Sequence data deposited on DRYAD at: DOI: 10.5061/dryad.5qfttdzbj.
2.3. Imaging
Where possible, images of the Type material of each species were included and as many non-type material as possible were examined using a Leica S7 APO dissecting microscope. Images were taken at the Australian National Insect Collection (Canberra, ACT 2601 Australia) using a Canon EOS 7D Mark II DSLR with Visionary Digital Big Kahuna (Bk) Camera system using 65 mm (Canon lens MP-E65mmf/2.8 1-5x Macro) and 100 mm (Canon lens EF 100 mm f/8 L IS USM), processed using Helicon focus Stacker and edited with Photoshop CS5.
2.4. Repositories
Specimens were examined from the following collections: Australian Museum, Sydney, Australia (AM); Australian National Insect Collection (ANIC); Humbolt University Zoologischen Musuem (HUM); Museo Civico di Storia Naturale “Giacomo Doria”, (MCG); Museum Victoria (MV); Natural History Museum (BMNH); Queensland Museum (QM); South Australian Museum (SAM); and Western Australian Museum (WAM). Morphological terminology used here follows Harris 1987 [10] and wing terminology follows Day 1988 [11].
3. Results
3.1. Phylogenetic Analysis and Species Delimitation
The IQ-TREE analyses overall yielded well supported interspecific relationships within Heterodontonyx (Figure 1). Cryptocheilus fulvus was used as outgroup and the in group (node A) is split in two branches B and C. Based on bPTP results, branch B holds five species and branch C contains three species. ML tree topology suggests that branch B is sister to group C which proposes the following phylogenetic estimates: H. tuberculatus (shown in yellow in Figure 1) is sister to a group containing H. bicolor, H. fulvidorsalis, H. darwinii and a new species (node D in Figure 1). POM 168 on Figure 1, the molecular new species for which the morphological description follows in the next section, is sister to node E containing H. darwinii, H. fulvidorsalis and H. bicolor. H. darwinii is sister to H. fulvidorsalis + H. bicolor (node F in Figure 1). H. fulvidorsalis (shown in brown in Figure 1) is sister to H. bicolor (shown in red in Figure 1). In addition, the phylogeny suggests that H. australis (shown in blue in Figure 1) is sister to a group containing H. distinctus and H. praepositus (node G in Figure 1) and that H. distinctus (shown in green in Figure 1) is sister group to H. praepositus (shown in purple in Figure 1).
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Figure 1. Maximum Likelihood tree reconstruction of Heterodontonyx based on the molecular information of ITS1, 5.8S, ITS2 region. Statistical support values of Ultra-Fast Bootstraps are shown on each node. bPTP suggested delimitation scheme of molecular units shown on the right panel next to their morphological identification.
3.2. Taxonomic Revision of the Australian Spider Wasp Genus Heterodontonyx Haupt, 1935 with Description of a New Species, Revised Key, New Records and a Host Association
Systematics of the genus Heterodontonyx Haupt 1935.
Heterodontonyx Haupt, 1935. Type species: Heterodontonyx basalis Haupt, 1935 (Figure 2) by subsequent designation, see Banks, 1941:234 and Pate, 1946: 90 [2] (=Sphex bicolor, synonymy by Wahis [12].
Diagnosis
Black and orange wasps of small to large size (10 - 40 mm) with golden-yellow wings; clypeus medium to large and convex, with labrum concealed to completely exposed; antennal segments slender and elongate; propodeum black or reddish-yellow, rugose, with or without tubercules; vein 2rs-m straight to slightly curved, 1m-cu vein reaching the second submarginal cell (SMC2) slightly to moderately beyond its middle; marginal cell elongated, ratio of the inter-tegular distance to the length of the second abscissa of forewing vein Rs (ITD: SaRs) ranges from 1.5 to 2.5. Female metatibia serrated with spines.
Figure 2. (a)-(d) Heterodontonyx basalis female lectotype (a) dorsal view (b) head, frontal view, (c) lateral view, left side (d) head, dorsal view, scale bars: (a), (c): 2 mm, (b), (d): 1 mm. Images taken by Lars Krogmann from HUM.
Key to Heterodontonyx Species
The key is based on both sexes, but caution should be taken as in a few cases only one sex is known (H. erythroura, H. solomonis and H. wahisi). Despite measurable sexual dimorphism in some Pompilidae (e.g., Aplochares imitator (Smith)), our observations suggest that, apart from genitalia, body size, size of black area on the head, relative length of first to second flagellar segment and occasional bifid tarsal claws in males, the sexes of Heterodontonyx are not distinctly dimorphic.
1. a. Head, mesosoma and metasoma blackH. solomonis Turner (Figure 10)
b. Body black and orange/yellow/reddish-yellow2
2. a. Sides of propodeum rounded; body size small (10 - 20 mm)3
b. Full tubercles or half tubercles on sides of propodeum4
3. a. Body with black setae; size 10 - 15 mm5
b. Body bare, without long setae; second metasomal segment sometimes with apical black bandH. australis (Guérin-Méneville, 1838) (Figure 3)
4. a. Full tubercles on sides of propodeum6
b. Half tubercles on sides of propodeum; prothorax and mesothorax brown yellowH. fulvidorsalis (Turner, 1910) (Figure 8)
5. a. Female head entirely orange, some males with pronotum golden yellow orange, body size 10 - 15 mmH. distinctus (Smith, 1868) (Figure 6)
b. Body size 10 - 15 mm, second metasomal segment sometimes with an apical black bandH. praepositus (Haupt, 1935) (Figure 9)
6. a. Clypeus long and narrow (length: width: 1.5 - 1.75); labrum completely Exposed7
b. Clypeus short and wide (length: width: 2 - 2.5); labrum slightly to moderately exposed8
7. a. Pubescence moderate; antennae very long and slender; head, pronotum and mesoscutum reddish-yellow brown; slight compression between first and second metasomal segmentsH. darwinii (Turner, 1910) (Figure 5)
b. Body covered with dense, black, and golden setae; pronotum orange, first metasomal segment and apical second metasomal segment black, remaining metasomal segments orangeH. wahisi sp. nov (Figure 12)
8. a. Mesosoma entirely black, second metasomal segment orangeH. tuberculatus (Smith, 1855) (Figure 11)
b. Mesosoma entirely black, second metasomal segment apically or entirely black9
9. a. Second metasomal segment entirely blackH. erythroura (Cameron 1906) (Figure 7)
b. Second metasomal segment with black basal bandHeterodontonyx bicolor (Fabricius, 1775) (Figure 4)
Heterodontonyx australis (Guérin-Méneville, 1838) (Figure 3(a)-(h))
Pompilus australis Guérin–Méneville, 1838: 260. Type data: syntypes MCG one male and one female. Type locality: NSW, Sydney (as Port Jackson, Nouvelle-Hollande); Froggatt, 1892: 212 [cat.]; Guiglia, 1948: 178; Guiglia & Pasteels, 1961: 19 [notes on the type (inItalian)]. Cryptochilus (Prionocnemis) australis Schulz, 1906: 166 [comb. nov., status as senior secondary homonym of Cryptochilus (Prionocnemis) australis Holmberg, 1903; Schulz, 1911: 115, 207 [note.].
Heterodontonyx australis Haupt, 1935: 309 [comb. nov., desc., Figure.].
Cryptocheilus australis Harris, 1987: 25 [comb. nov., descr., Figures., biol., dist. (New Zealand), notes]; Forster, 1994: 215 [note]; Harris, 1999: 143–158, 223, 226, 229–232 [notes, list, biol., Figures.]. Elliot, 2007: 20 [cat.].
Prey records: Dolomedes minor Koch, 1872; Dolomedes aquaticus Goyen, 1888; Dolomedes spp. (Pisauridae) (Harris, 1999: 157)
Adult food sources: Leptospermum scoparium Forst. & Forst. (Myrtaceae); Pastinaca sativa L. (Apiaceae) (Harris, 1987: 28 and Harris, 1999: 157); Gomphocarpus physocarpus E. Mey. (Asclepiadaceae) (Forster, 1994: 215).
Material examined.
Holotype image:
The male lectotype is from the Guérin collection and determined by Raymond Wahis in 1995. The material is pinned on a paper card and in a fair condition, some legs and antennal segments are missing from both sides. The female paralecto type is from the Guérin collection and determined by Raymond Wahis in 1995. The material is pinned on a paper card and in a fair condition, both eyes, distal segments from left antenna and legs are missing and wings also are slightly torn.
Other material (See Table 1):
Figure 3. (a)-(d) Heterodontonyx australis female paralectotype (a) dorsal view (b) lateral view, right side (c) head, frontal view (d) lateral view, left side (e)-(h) Pompilus australis male lectotype (e) dorsal view (f) lateral view, right side (g) head, frontal view (h) lateral view, left side. Images taken by Maria Tavano from MCG.
Table 1. Material examined.
| Gender |
Accession # |
Collection date |
Locality |
Latitude |
Longitude |
Collected by |
| Female |
ANIC32_101009 |
20 Jun. 1929 |
NT |
18S |
135.5E |
Campbell, T.G. |
| Female |
ANIC32_100927 |
16 Feb. 1969 |
ACT |
35.35S |
148.8E |
Upton, M.S. |
| Female |
ANIC32_101089 |
4. Oct. 1969 |
WA |
24.8159S |
113.6298E |
R.W. Matthews |
| Female |
ANIC 32_099228 |
Oct. 1972 |
Qld. |
24.87S |
152.35E |
H. Frauca |
| Female |
ANIC32_101024 |
12. Nov. 1987 |
SA |
34.01S |
139.49E |
I. Naumann, J. Cardale |
| Male |
ANIC32_115588 |
22 Feb. 1957 |
New Caledonia |
21.2107S |
165.8517E |
J. Rageau |
| Male |
ANIC32_101063 |
12 Feb. 1970 |
ACT |
35.4735S |
149.0124E |
Evans, H. & Matthews, R.W. |
| Male |
ANIC32_099227 |
3 Jun. 1982 |
NSW |
33.71S |
150.38E |
Unknown |
| Male |
TMAG-F6673 |
18 Dec. 2013 |
Tas. |
42.8680S |
147.3854E |
S .J. Grove |
| Male |
ANIC-PO1507 |
5 Dec. 2017 |
Qld. |
26.1.15S |
153.051E |
Xuankun Li, David Yeates |
| Male |
ANIC-PO1508 |
19 Feb. 2018 |
NSW |
33.31.17S |
150.21.29E |
Xuankun Li, David Yeates |
| Male |
ANIC-PO1506 |
26 Feb. 2018 |
NSW |
33.31.17S |
150.21.29E |
Xuankun Li, Alan Lanford |
| Male |
ANIC-PO1509 |
26 Feb. 2018 |
NSW |
33.31.17S |
150.21.29E |
Xuankun Li, Alan Lanford |
Length 10 - 20 mm; colour black and orange; body not heavily covered with setae, almost bare; head of female with face and vertex yellow orange, except for a black band as wide as ocellar triangle passing through ocelli and reaching compound eyes; head of male with vertex and disc of frons black; propodeum without tubercles; second metasomal segment sometimes with a very narrow, apical black band.
Redescription
Female
Coloration. Head orange yellow except for a narrow area between the eyes, the end of the mandible, and the occipital region, which are dark/black; mesosoma black except for tegula which is orange; tibiae and tarsi orange, femora black basally, orange apically; first metasomal segment black, second metasomal segment orange, sometimes with narrow apical black band, remainder yellow orange; wings yellow orange, with a narrow apical darkening.
Head. Head wider than long, TFD/FD = 1.2; posterior ocelli closer to each other than to the compound eye, POL: OOL = 0.75; frons with median line reaching median ocellus; apical margin of clypeus slightly concave; clypeus rather short, clypeus length: width = 2.27; labrum on same plain as clypeus moderately exposed beneath; malar space very short, almostnon-existent; first flagellar segment slightly longer than the second. Mesosoma.
Mesosoma. Polished with few setae, pronotum relatively short, with a very weak median line, its posterior margin strongly angulate; metanotum knobbed; metapostnotum suppressed; propodeum slightly wrinkled across with median line, without tubercules; dorsal edge of hind tibia with carina bearing spines; tarsal claws dentate. Metasoma. Metasoma polished, pygidium covered by short, abundant setae, metasoma 1.21× length of mesosoma.
Male
Similar to female except differences described here: vertex and most of the frons black; clypeus relatively long; hind tibiae without serration with short spines; body size smaller.
Distribution: NSW, New Zealand (North Island). Tas., WA, Qld. (Elliot, 2007) Newrecords: NT, SA, New Caledonia.
Heterodontonyx bicolor (Fabricius, 1775) (Figures 4(a)-(d))
Sphex bicolor Fabricius, 1775: 352. Type data: type BMNH female. Type locality: Australia (as nova Hollandia); Fabricius, 1781: 450 [descr. (in Latin), dist.]; Fabricius, 1787: 277 [descr. (in Latin)]; Gmelin, 1790: 2732 [descr. (in Latin, dist.]; Fabricius, 1793: 217 [descr. (in Latin); dist.]; Zimsen, 1964: 387 [note on type]. Pompilus bicolor Fabricius, 1804: 198 [comb. nov.; descr. (in Latin); dist.]; Smith, 1863b: 132 [dist.]; Smith, 1863a: 29 [note, dist.]; Smith, 1871: 353 [cat., dist.]; Schulz, 1906: 169 [as senior secondary homonym of Pompilus bicolor (Spinola, 1808)]. Salius bicolor Tillyard, 1926: 292 [note]; McKeown, 1942: 185 [note]. Cryptocheilus bicolor Evans & Matthews, 1973: 46 [comb. nov., biol.]; Evans 1981: 9 [prey]; Hirst 1992: 365 [prey record]; Williams & Adam 1995: 54 [list]. Elliot, 2007: 20 [cat.].
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Figure 4. (a)-(c) Heterodontonyx bicolor female holotype (a) Dorsal view (b) Lateral view, left side (c) Head and fronto-lateral view (d) Non-type male, dorsal view. Scale bars: 1 mm. Images (a)-(c) taken by Xuankun Li in BMNH.
Prey records: Isopedellapessleri (Thorell, 1870) (as Isopedapessleri (Thorell)); Isopeda sp. (Sparassidae) (Evans & Matthews, 1973: 46-47), Heteropoda jugulans Koch, 1876; Isopedaleishmani Hogg, 1903 (as Isopoda leishmani Hogg) (Sparassidae); Lycosa sp. (Lycosidae) (Evans 1981: 9-10), Isopedacanberrana Hirst, 1992 (Heteropodidae) (Hirst, 1992).
Material examined
Holotype image
Type accession number: BMNH (E) #668328, NHMUK 013380164 holotype is from the Banks collection and is in good condition except that the last tarsal segment of hind legs are missing.
Other material (See Table 2):
Length 20 - 40 mm; colour black and orange; body heavily covered in dense black setae; head of female with face and vertex orange, except for a black band
Table 2. Material examined.
| Gender |
Accession # |
Collection date |
Locality |
Latitude |
Longitude |
Collected by |
| Female |
QM.No.ENT19.08 |
10 Jan. 1966 |
PNG |
9.4814S |
147.1461 E |
E. Mann |
| Female |
ANIC32_100861 |
5 May. 1981 |
Vic. |
37.9145S |
145.1350 E |
M. Kotzman |
| Female |
ANIC32_100890 |
10-12 May. 1981 |
Qld. |
15.29S |
145.16E |
I. D. Naumann |
| Female |
ANIC32_100862 |
20 Aug. 1982 |
NT |
18.6S |
136E |
R. Patterson 4340 |
| Female |
ANIC32_100853 |
22 Aug. 1989 |
SA |
27.54S |
135.49E |
I. BUNIC |
| Female |
ANIC32_100848 |
17 Feb. 1986 |
NSW |
33.4622S |
151.2561 E |
D. B. Mc Corqundale |
| Male |
ANIC32_100837 |
5 Apr. 1971 |
WA |
21.34 S |
117.03E |
E. F. Riek |
| Male |
ANIC32_100882 |
3-14 Feb. 1977 |
NT |
11.01S |
136.45E |
T. A. Weir |
between ocelli; head of male with vertex and disc of frons black; propodeum with two distinct tubercles; second metasomal segment with a wide apical black band.
Redescription
Female
Coloration. Head orange yellow except for a narrow area between the eyes, the end of the mandible, and the occipital region, which are dark/black; mesosoma black; tibiae and tarsi orange, femora black basally, orange at apex; first metasomal segment black, second metasomal segment orange with apical black band, remainder yellow orange; wings yellow orange, with a narrow apical and basal darkening. Head. Head wider than long, TFD/FD = 1.16; posterior ocelli closer to each other than to the compound eye, POL: OOL = 0.72; frons with median line reaching median ocellus; apical margin of clypeus straight; clypeus rather short, clypeus length: width = 2.2; labrum on same plain as clypeus concealed to slightly exposed beneath; malar space very short, almost non-existent; first flagellar segment clearly longer than the second.
Mesosoma. Setose, pronotum relatively short, with a very weak median line, its posterior margin strongly angulate; metanotum knobbed; metapostnotum suppressed; propodeum wrinkled across with a median line which is more prominent at the posterior ridge, tuberculate; dorsal edge of hind tibia with two rows of spine-bearing scale like elevations which become less dense towards tarsus, tarsal claws dentate.
Metasoma. Metasoma punctate, pygidium covered by short, abundant setae, metasoma 1.25× length of mesosoma.
Male
Similar to female except differences described here: vertex and disc of frons black; clypeus relatively long; first flagellar segment slightly longer than the second; hind tibiae without serration with short spines; body size smaller.
Remarks
Original description of this species by Fabricius, 1775 was based on one sex only (female).
Distribution: ACT, NSW, QLD, WA, Indonesia, New Zealand (Elliot, 2007). New records: NT, SA, PNG, Vic.
New host association: Isopedellaleai (Hirst) loan from SAM accession number: 32-42650.
Heterodontonyx darwinii (Turner, 1910) (Figure 5(a)-(d))
Cryptocheilus darwinii Turner, 1910: 318. Type data: type BMNH one male and one female.
Type locality: NT, Darwin. Elliot, 2007: 21 [cat.]
Material examined
Holotype co-type male (accession number: NHMUK 013380165) and holotype female. The male cotype is pinned on a paper card and in a very good condition. The holotype material is also pinned and is in good condition except for the broken left antennae.
Other material (See Table 3):
Figure 5. (a)-(d) Cryptocheilus darwinii (a) Male co-type, dorsal view (b) Female holotype, dorsal view (c) Male co-type, lateral view, left side (d) Male co-type, head and fronto-dorsal view. Scale bars: 1 mm. Images (a), (c), (d) taken by Xuankun Li in BMNH.
Table 3. Material examined.
| Gender |
Accession # |
Collection date |
Locality |
Latitude |
Longitude |
Collected by |
| Female |
QMENT19.08 |
Unknown |
Qld. |
16.9203S |
145.7710E |
F. P. Dodd |
| Male |
ANIC |
Oct. 1980 |
Qld. |
14.17S |
144.12E |
L. Taylor |
Length 30 - 40 mm; colour black and brown-yellow; body not heavily covered with setae; head of female brown-yellow; head of male with vertex backwards black; pronotum and mesoscutum brown-yellow; propodeum with two distinct tubercles; first metasomal segment black with apical constriction.
Redescription
Female
Coloration. Head brown-yellow; mesosoma black except for tegula, pronotum and mesoscutum which are brown-yellow; tibiae and tarsi brown-yellow, femora black baslly, brown-yellow apically; first metasomal segment black, second metasomal segment onwards brown-yellow; wings yellow orange, with narrow apical and basal darkening.
Head. Almost the same width as length, TFD/FD = 1.05; posterior ocelli closer to each other than to the compound eye, POL: OOL = 0.77; frons with median line reaching median ocellus; apical margin of clypeus slightly concave; clypeus rather long, clypeus length: width = 1.25; labrum on same plain as clypeus moderately exposed beneath; malar space very short, almost nonexistent; antenna very long and slender; first flagellar segment slightly longer than the second.
Mesosoma. Mesosoma with moderate pubescence; pronotum relatively short, with an obvious median line, its posterior margin strongly angulate; metanotum knobbed; metapostnotum suppressed; propodeum wrinkled across with a median sulcus, tuberculate. spine-bearing dorsal edge of hind tibia moderately serrated; tarsal claws dentate.
Metasoma. Metasoma polished, softly punctate, pygidium covered by short, golden set ae, metasoma 1.14× length of mesosoma.
Male
Similar to female except differences described here: vertex black; clypeus longer; first flagellar segment slightly longer than the second; hind tibiae without serration with short spines; body size slightly smaller.
Distribution: NT. New records: QLD
Heterodontonyx distictus (Smith, 1868) (Figure 6(a)-(d))
Pompilus distinctus Smith, 1868: 242. Type data: type BMNH, male. Type locality: A, Champion Bay; Froggatt, 1892: 212 [cat.].
Cryptocheilus distinctus Evans et al. 1981: 10 [prey and nest]. Elliot, 2007: 21 [cat.].
Prey records: Eriophora biapicata (Koch, 1871) (Araneidae); Lycosalaeta Koch, 1877 (Lycosidae) (Evans, 1981: 10).
Material examined
Figure 6. (a), (b) Pompilus distinctus male holotype (a) Dorsal view (b) Head, frontal view (c), (d) Female non-type (c) lateral view, right side (d) head, frontal view. Scale bars: 1 mm.
Holotype
Type accession number: B.M. TYPE HYM. 19.156, NHMUK010577193, 68-6, The type is determined by R. Wahis in 1995, is pinned and is in fair condition, lower right part of the right forewing is torn, and the right hind wing is broken and missing for the most part, the lasttarsal segment of the right hind leg is missing, and left antennae has some distal broken segments.
Other material (See Table 4):
Length 10 - 15 mm; colour black and orange; body moderately setose; head of female entirely orange; head of male with vertex and disc of frons black; some males with pronotum golden; propodeum without tubercles; second metasomal segment without any apical black band.
Redescription
Female
Coloration. Head orange, mesosoma black; tibiae and tarsi orange, femora black basally, orange at apex; first metasomal segment black, second metasomal segment onward orange; wings yellow orange, with a narrow apical darkening.
Head. Head wider than long, TFD/FD = 1.2; posterior ocelli closer to each other than to the compound eye, POL: OOL = 0.86; frons with median line reaching median ocellus; apical margin of clypeus straight; clypeus medium, clypeus length: width = 2; labrum on same plain as clypeus slightly exposed beneath; malar space very short, almost non-existent; first flagellar segment about the same length as the second.
Mesosoma. Bears black setae; pronotum relatively short, with a median line, its posterior margin angulate; metanotum knobbed; metapostnotum suppressed; propodeum wrinkled across with a median line, without tubercules; dorsal edge of hind tibia serrate with spines; tarsal claws dentate.
Metasoma. Metasoma polished, sparsely punctate, pygidium covered by short, abundant golden setae, metasoma 1.16× length of mesosoma.
Male
Similar to female except differences described here: vertex and most of the frons black; hind tibiae without serration with short spines; body size slightly smaller.
Remarks
Table 4. Material examined.
| Gender |
Accession # |
Collection date |
Locality |
Latitude |
Longitude |
Collected by |
| Female |
MUS.VICENTO2019-HYM-62732 |
27 Mar. 1949 |
VIC |
35.5789S |
143.7749E |
A. N. Burns |
| Female |
ANIC32_101162 |
18-22 Feb. 1970 |
VIC |
35.7678S |
142.0545E |
H. E. Evans, R. W. Matthews |
| Female |
ANIC32_101166 |
Oct. 1971 |
QLD |
24.87S |
152.3489E |
H. Frauca |
| Male |
ANIC32_101154 |
3-6 Feb. 1970 |
QLD |
23.1335S |
150.7374E |
H. E. Evans |
| Male |
ANIC32101060 |
21-22 Feb. 1970 |
VIC |
36.1S |
142E |
Evans, H., Matthews, R. W. |
| Male |
ANIC32_101134 |
Oct. 1971 |
QLD |
24.87S |
152.35E |
Frauca, H. |
| Male |
WAMReg. No. E101033 |
1 Apr. 1980 |
SA |
31.82S |
138.59E |
Bohart, R. M. |
Original description of this species by Smith, 1868 was based on one sex only (male).
Distribution
NSW, WA. (Elliot, 2007). New records: Qld., Vic., SA
Heterodontonyx erythroura (Cameron 1906) (Figure 7(a)-(c))
Saliuserythroura Cameron 1906. Type data: Type depository unknown, female, Type locality: Lake Sentani
Material examined
One non-type female specimen studied Accession number: BM O.I.E.Coll.A 5723; collected in 07. Oct. 1968 by F.R.Wylie in Bulolo Morabe province, PNG Diagnosis (modified from Cameron 1906).
Length 35 mm; colour black and orange; body heavily setose; head of female with vertex black and disc of frons orange; propodeum with two distinct tubercles; second metasomal segment black.
Redescription
Female
Coloration. Head orange except for vertex backwards and the end of the mandible which are dark/black; mesosoma black; tibiae and tarsi orange; femora
Figure 7. (a)-(c) Heterodontonyx erythroura female non-type (a) Dorsal view (b) Head, frontal view (c) Lateral view, right side. Scale bars: 1 mm.
black basally, orange apically; first and second metasomal segments black; third metasomal segment onward orange; wings yellow orange, with a narrow apical darkening. Head. Head slightly wider than long, TFD/FD = 1.1; posterior ocelli closer to each other than to the compound eye, POL: OOL = 0.75; frons with median line reaching median ocellus; apical margin of clypeus straight; clypeus slightly long, clypeus length: width = 1.94; labrum on same plain as clypeus moderately exposed beneath; malar space very short, almost non-existent; first flagellar segment clearly longer the second. Mesosoma. Setose, pronotum relatively short, with a very weak median line, its posterior margin convergent towards the median line; metanotum knobbed; metapostnotum suppressed; propodeum wrinkled across without a median line, tuberculate; dorsal edge of hind tibia with spine-bearing elevations; tarsal claws dentate.
Metasoma. Metasoma setose, punctate, pygidium covered by short, abundant setae, metasoma 1.2× length of mesosoma.
Male
Unknown.
Distribution: Bulolo, PNG.
Heterodontonyx fulvidorsalis (Turner, 1910) (Figure 8(a)-(f))
Cryptocheilus fulvidorsalis Turner, 1910:319. Type data: type BMNH, female. Type locality: QLD, Mackay. Elliot, 2007: 21 [cat.].
Figure 8. (a)-(c) Cryptocheilus fulvidorsalis female holotype (a) Dorsal view (b) Head, frontal view (c) Left fore and hind wings (d)-(f) Non-type male (d) Dorsal view (e) Head, fronto-dorsal view (f) Lateral view, right side. Scale bars: 1 mm.
Material examined
Holotype
Type accession number: B.M. TYPE HYM. 19.155, NHMUK010577201 from Turner Collection with the accession number 1910-7. Type is female, pinned, in very good Condition.
Other material (See Table 5):
Length 15 - 25 mm; colour brown-yellow and black; body moderately covered in golden setae; head of female entirely brown-yellow; head of male brown yellow with a black area in vertex; pronotum and mesoscutum brown-yellow; propodeum with two half-tubercles; propodeum black in female, brown-yellow in male; first metasomal segment black in female, brown-yellow with apical darkening in male.
Redescription
Female
Primarily based on holotype
Coloration. Head brown-yellow; mesosoma brown-yellow (pronotum, tegula and mesoscutum) and black (scutellum and propodeum); tibiae and tarsi brown-yellow, femora black basally, brown-yellow apically; first metasomal segment black, second metasomal segment onward brown-yellow.
Head. Head slightly wider than long, TFD/FD = 1.12; posterior ocelli closer to each other than to the compound eye, POL: OOL = 0.83; frons with median line reaching median ocellus; apical margin of clypeus slightly concave; clypeus medium, clypeus length: width = 2; labrum on same plain as clypeus slightly exposed beneath; malar space very short, almost non-existent; first flagellar segment slightly longer the second.
Mesosoma. Moderate pubescence, pronotum relatively short, with a clear median line, its posterior margin convergent towards median line; metanotum without knob; propodeum wrinkled across with a very weak median line, without any prominent tubercles (half-tuberculate); dorsal edge of hind tibia with spine-bearing tooth-like elevations; tarsal claws dentate.
Metasoma. Metasoma polished, sparsely punctate, pygidium covered by short, abundant golden setae, metasoma 1.05× length of mesosoma.
Male
Similar to female except differences described here: head bears black area between compound eyes; propodeum and first metasomal segment brown-yellow; hind tibiae without serration with short spines; tarsal claws bifid; body size slightly smaller.
Table 5. Material examined.
| Gender |
Accession # |
Collection date |
Locality |
Latitude |
Longitude |
Collected by |
| Female |
ANIC32_099267 |
24 May. 1972 |
QLD |
18.1311S |
139.5244E |
G. B & S. R. Monteith |
| Female |
ANIC32_099273 |
25 Oct. 1975 |
NT |
16.47S |
135.45E |
J. C. Cardale |
| Male |
ANIC99272 |
3 Oct. 1950 |
QLD |
19.568516S |
147.406387E |
E. F. Riek |
Remarks
Original description of this species by Turner, 1910 was based on one sex only (female).
Distribution: QLD. New records: NT.
Heterodontonyx prapositus (Haupt, 1935) (Figure 9(a)-(f))
Cryptochilus (Cryptochilus) praepositus Haupt, 1935b: 269. Type data: holotype HUM, Type locality: WA, SA, Tas., Celebes. Elliot, 2007: 21 [cat.].
Material examined
Holotype images
Holotype is female and from the Bingham collection. From the label, it seems that Haupt determined the specimen in 1920s, but formally described the species in 1935. Male specimen was also first studied and labelled by Haupt in 1920s and later in 1995 the “allotype” was designated as lectotype by Raymond Wahis. Both specimens are kept in HUM and are in good condition.
Other material (See Table 6):
Length 10 - 25 mm; colour black and orange; body moderately setose; head of female with face and vertex yellow orange, except for a black band as wide as ocellar triangle passing through ocelli and reaching compound eyes; head of male with vertex and disc of frons black; propodeum without tubercles; second metasomal segment sometimes with a very narrow, apical black band in male, sometimes without it in female.
Redescription
Female
Coloration. Head orange yellow except for a narrow area between the eyes, and the end of the mandible, which are dark/black; mesosoma black except for tegula which is orange; tibiae and tarsi orange, femora black basally, orange apically;
Figure 9. (a)-(c) Cryptochilus praepositus male lectotype (a) Dorsal view (b) Lateral view, left side (c) Head, frontal view (d) (f) Cryptochilus praepositus female holotype (d) Dorsal view (e) Lateral view, left side (f) Head, frontal view. Scale bars (a), (b), (d), (e): 2 mm, (c), (f): 5 mm. Images taken by Lars Krogmann from HUM.
Table 6. Material examined.
| Gender |
Accession # |
Collection date |
Locality |
Latitude |
Longitude |
Collected by |
| Female |
ANIC 32_101102 |
21 Oct. 1949 |
NSW |
32.03S |
147.98E |
Paramonov, S. J. |
| Female |
ANIC 32_099228 |
Oct. 1972 |
QLD |
24.87S |
152.35E |
H. Frauca |
| Female |
ANIC32_099287 |
2 Jan. 1987 |
NSW |
35.15S |
149.96E |
Naumann, I. D. |
| Male |
ANIC32_101057 |
23-27 Sept. 1969 |
WA |
26.65S |
120.25E |
H. Evans, R. W. Matthews |
| Male |
ANIC32_099249 |
24 May. 1972 |
QLD |
17.9S |
139.6E |
Monteith, G. B. & Monteith, S. R. |
| Male |
ANIC32099245 |
27-28.Sept. 1972 |
NT |
25.3S |
135.5E |
Z, Liepa |
| Male |
WAM. E101033 |
1 Apr. 1980 |
SA |
31.82S |
138.59E |
Bohart, R. M. |
| Male |
ANIC32_100972 |
29 Nov. - 2 Dec. 1981 |
NSW |
31.05S |
141.42E |
Cardale, J. C. |
| Male |
ANIC32_100975 |
29 Nov. - 2 Dec. 1981 |
NSW |
31.05S |
141.42E |
I. D. Naumann, Cardale, J. C. |
| Male |
ANIC32_101028 |
12 Nov. 1987 |
SA |
34.01S |
139.49E |
Naumann, I. & Cardale, J |
first metasomal segment black, second metasomal segment orange, sometimes with narrow apical black band, remainder yellow orange; wings yellow orange, with a narrow apical darkening.
Head. Head wider than long, TFD/FD = 1.21; posterior ocelli closer to each other than to the compound eye, POL: OOL = 0.66; frons with median line not reaching median ocellus; apical margin of clypeus straight; clypeus medium, clypeus length: width = 2.08; labrum on same plain as clypeus completely exposed beneath; malar space very short, almost non-existent; first flagellarsegment about the same length as the second.
Mesosoma. Mesosoma covered with blacksetae, pronotum relatively short, with a very weak median line, its posterior margin angulate; metanotum knobbed; metapostnotum suppressed; propodeum wrinkled across with noobvious median line, without tubercules; dorsal edge of hind tibia with spine-bearing scale like elevations which becomes less prominent towards tarsus; tarsal claws dentate.
Metasoma. Metasoma with black setae, sparsely punctate, pygidium covered by short, abundant setae, metasoma 1.17× length of mesosoma.
Male
Similar to female except differences described here: vertex and frons black; clypeus convex, apical margin straight, labrum completely exposed beneath; hind tibiae without serration with short spines; body size slightly smaller.
Distribution: SA, Tas., WA, Sulawesi (Celebes).
New records: NSW, Qld., NT.
Heterodontonyx solomonis Turner (Figure 10(a)-(c))
Type data: Holotype whereabouts unknown; probable type depository Natural History Museum of Denmark Banks, 1941: 233.
Material examined
One non-type female specimen studied from Solomon Islands; Accession number BM-048, collected by R.A. Lever and determined by R. Wahis in 1967.
Figure 10. (a)-(c) Heterodontonyx solomonis female non-type (a) Dorsal view (b) Head, frontal view (c) Fronto-lateral view, right side. Scale bars: 1 mm.
Diagnosis (modified from Banks, 1941)
Length 35 mm; colour black; body heavily covered with black setae; head of female black; mesosoma black except for brown wings and legs; propodeum with half-tubercles; metasomal segments black.
Redescription
Female
Coloration. Head black except for antenna which is dark brown; mesosoma black; tibiae and tarsi brown; femora black basally, brown apically; metasomal segments black; wings dark brown.
Head. Head wider than long, TFD/FD = 1.2; posterior ocelli closer to each other than to the compound eye, POL: OOL = 0.88; frons with median line not reaching median ocellus; clypeus rather short, clypeus length: width = 2.6; labrum on same plain as clypeus, slightly exposed beneath; malar space very short, almost non-existent; first flagellar segment longer than the second.
Mesosoma. Mesosoma setose; pronotum relatively short, with weak median line, its posterior margin angulate; metanotum without knob; propodeum wrinkled across with a median line; without any prominent tubrcles (half-tubercu- late); dorsal edge of hind tibia with spine-bearing scale-like elevations; tarsal claws dentate.
Metasoma. Metasoma setose, densely punctate, pygidium covered by short, abundant black setae, metasoma 1.13x length of mesosoma.
Male: Unknown
Distribution: Guadalcanar Island, Solomons
Heterodontonyx tuberculatus (Smith, 1855) (Figure 11(a)-(d))
Pompilus tuberculatus Smith, 1855: 166. Type data: type BMNH, female. Type locality: Western Australia (as New Holland (Houtman’s Abrolhos, W. Coast)); Froggatt, 1892: 214 [cat.]. Priocnemis tuberculatus (Smith, 1855) comb. nov. Eliot, 2007: 30 [cat.] Cryptochilus (Prionocnemis) tuberculatus Schulz, 1908:471 [comb. nov., descr., dist.]. Salius Tuberculatus Tillyard, 1926: 292 [note].
Material examined
Holotype
Holotype is female and kept in BMNH with accession number: 185510577164. The specimen is in fair condition; antennae tips and right hind tarsus broken.
Other material (See Table 7):
Figure 11. (a)-(c) Heterodontonyx tuberculatus female non-type (a) Dorsal view (b) Head, frontal view (c) Lateral view, left side (d) Male, non-type, dorsal view. Scale bars: 1 mm.
Table 7. Material examined.
| Gender |
Accession # |
Collection date |
Locality |
Latitude |
Longitude |
Collected by |
| Female |
ANIC32_096531 |
12 Nov. 1953 |
WA |
20.5167S |
115.5333E |
T. G. Capbell |
| Female |
ANIC32_096627 |
20 Dec. 1984 |
NSW |
34.51S |
150.44E |
D. B. McCorquodale |
| Female |
ANIC |
20 Mar. 1990 |
Qld. |
27.56S |
145.47E |
E. D. Edwards, J. H. Fisk |
| Female |
AM K551011 |
12 Apr. 2007 |
Qld. |
14.40S |
145.28E |
R. B. Lachlan |
| Female |
ANIC specimen |
19 Mar. 2016 |
SA |
27.44.99S |
138.14.42E |
D. K. Yeates, Y. N. Su & A. Lanford |
| Female |
ANIC |
21 Mar. 2017 |
NT |
19.4914S |
132.5510E |
Unknown |
| Male |
AM K513656 |
May 1927 |
PNG |
8.9353S |
148.1664E |
C. T. McNamara |
Length 20 - 40 mm; colour black and orange; body heavily covered with setae, head of female entirely orange, head of male orange except for a black band as wide as ocellar triangle passing through ocelli and reaching compound eyes; propodeum with two distinct tubercles; second metasomal segment orange, without apical black band.
Redescription
Female
Coloration. Head orange yellow except for the end of the mandible which is dark; mesosoma black; tibiae and tarsi orange, femora black basally, orange apically; first metasomal segment black, second metasomal segment onward orange; wings yellow orange, with a narrow apical darkening.
Head. Head wider than long, TFD/FD = 1.18; posterior ocelli closer to each other than to the compound eye, POL: OOL = 0.68; frons with median line reaching median ocellus; apical margin of clypeus straight; clypeus medium, clypeus length: width = 2; labrum on same plain as clypeus slightly exposed beneath; malar space very short, almost non-existent; first flagellar segment clearly longer than the second.
Mesosoma. Setose; pronotum relatively short, with a median line, its posterior margin angulate; metanotum knobbed; metapostnotum suppressed; propodeum wrinkled across with amedian line, tuberculate; dorsal edge of hind tibia with spine-bearing serration; tarsal claws dentate.
Metasoma. Metasoma with less setae than mesosoma, softly punctate, pygidium covered by short, golden setae, metasoma 1.15× length of mesosoma.
Male
Similar to female except differences described here: inter-ocellar line containing ocelli black; clypeus slightly longer with labrum more exposed beneath; first flagellar segment slightly longer than the second; hind tibiae without serration with short spines; tarsal claws bifid; body size smaller.
Distribution WA.
New records: Qld., NT, SA, NSW, PNG.
Heterodontonyx wahisi Chavoshi sp. nov. (Figures 12(a)-(e))
Holotype male from Australia: New South Wales: 5 km North Pooncarie; 27.11.1988.
N.W.Rodd; (AM-K551514)
ParatypesTwo male paratypes from SA and Qld.
Holotype and paratypes deposited in ANIC.
Diagnosis
Length 30 - 35 mm; colour black and orange; body heavily covered with black and golden setae; head of male with face and vertex yellow orange, except for a black band as wide as ocellar triangle passing through ocelli and reaching compound eyes; pronotum orange; propodeum with two distinct tubercles; second metasomal segment with apical black band.
Figure 12. (a)-(e) Heterodontonyx wahisi male holotype, (a) Dorsal view (b) Head, frontal view (c) Lateral view, right side (d) Subgenital plate (e) Genitalia, scale bars: 1 mm.
Description
Male
Primarily based on holotype
Coloration. Head orange yellow except for a narrow area between the eyes and the end of the mandible, which are dark/black; mesosoma black except for tegula and pronotum which are orange; tibiae and tarsi orange, femora black basally, orange at apex; first metasomal segment black, second metasomal segment orange with apical black band, remainder yellow orange; wings yellow orange, with apical and basal darkening.
Head. Head almost the same width as length, TFD/FD = 1.02; posterior ocelli as far apart from each other as from the compound eye, POL: OOL = 1; eyes slightly convergent toward the vertex; frons with median line reaching median ocellus; apical margin of clypeus slightly convex; clypeus ratherlong, clypeus length: width = 1.42; labrum on same plain as clypeus moderately exposedbeneath; malar space very short, almost non-existent; antenna long but not very slender; firstflagellar segment longer than the second.
Mesosoma. Setose; pronotum relatively short, with an obvious median line, its posterior margin angulate; metanotum knobbed; metapostnotum suppressed; propodeum weakly wrinkled across with no obvious median line, tuberculate; dorsal edge of hind tibia with spine-bearing scale-like elevations which becomes lessprominent towards tarsus; tarsal claws dentate.
Metasoma. Metasoma polished, sparsely punctate, pygidium covered by short, abundant golden setae, metasoma almost the samelength of mesosoma.
Genitalia. Parapenial lobe elongate and slender, 0.75× as long as entire genitalia; length 0.63× paramere length; setae long, thin abundant on external surface. Aedeagus almost the same length as parapenial lobes. Paramere length 0.83× total genitalia length; apex rounded; setae long, thick, covering 0.3 of length apically. Sub genital plate, wide, apex rounded, setae abundant apically, long, thin.
Etymology
This species is named after late Raymond Wahis, who, along with his long- standing work on Pompilidae taxonomy, has contributed to knowledge of Australian Heterodontonyx Distribution.
4. Discussion
This study was the first ever updated species-level taxonomy for the Australasian Heterodontonyx. Molecular clues and traditional morphological characters were useful in delimiting one new species from Heterodontonyx stored in museum collections. We present phylogeny, molecular species delimitation and systematic revision of the genus Heterodontonyx including diagnosis and description for 10 species. New geographic records are shown for seven species and undescribed sexes are described for the first time from identified material for four species: H. bicolor, H. distinctus, H. fulvidorsalis and H. tuberculatus. A new host-association is made for H. bicolor: Isopedellaleai (Hirst) based on the identification of the host by spider species author (Hirst). Due to difficult access to the type of material of H. tuberculatus (deposited in BMNH) we have used images of non-type material (Figure 11). We were also unable to find type depositories of two species: H. erythroura and H. solomonis and as such we imaged two non-type specimens and used their original description as well as geographic data as a means of identification verification. These two species were not catalogued in Elliot, 2007 since their distribution is outside Australia (PNG and Solomon Islands).
5. Conclusion
This study reports the first molecular phylogeny of Heterodontonyx and provides new insights into the evolution of diversity in this pompilid group using assemblies of whole-genome data of pinned museum specimens. Molecular species delimitation complemented with morphological examination recovers one new species, three new sex associations and seven new records.
Acknowledgements
We would like to thank Gavin Broad, Karin Koch, Lars Krogmann, Xuankun Li, Catriona McPhee, Benjamin Parslow, Matthew Shaw, Russell Cox, Derek Smith, Maria Tavano, Ken Walker, Susan Wright for loaning museum material or taking images, Olivia Evangelista for technical help and Ian Naumann for making corrections and comments to the text.
Appendix
Table S1. Voucher information
| Voucher ID |
ID- Species Name-Sex-Locality-Collection Year |
| POM003 |
POM003-H. bicolor-M-NT-1977 |
| POM006 |
POM006-H. distinctus-F-QLD-1972 |
| POM009 |
POM009-H. bicolor-F-SA-1989 |
| POM014 |
POM014-H. bicolor-F-NSW-1986 |
| POM015 |
POM015-H. bicolor-F-NT-1982 |
| POM044 |
POM044-C. fulvus-F-Burma-F-1924 |
| POM068 |
POM068-H. darwinii-M-QLD-1980 |
| POM128 |
POM128-H. praepositus-M-VIC-1970 |
| POM131 |
POM131-H. praepositus-M-QLD-1972 |
| POM133 |
POM133-H. praepositus-M-WA-1969 |
| POM134 |
POM134-H. praepositus-M1-NSW-1981 |
| POM135 |
POM135-H. praepositus-M2-NSW-1981 |
| POM138 |
POM138-H. praepositus-F-NSW |
| POM139 |
POM139-H. australis-M-ACT-1970 |
| POM140 |
POM140-H. australis-F-ACT-1969 |
| POM141 |
POM141-H. praepositus-F-NT-1929 |
| POM142 |
POM142-H. australis-F-NSW-1987 |
| POM144 |
POM144-H. distinctus-M-QLD-1971 |
| POM146 |
POM146-H. distinctus-M-QLD-1970 |
| POM148 |
POM148-H. distinctus-F-VIC-1970 |
| POM152 |
POM152-H. distinctus-F-WA-1983 |
| POM163 |
POM163-H. tuberculatus-F-NT-2017 |
| POM168 |
POM168-H. new species-M-NSW-1988 |
| POM192 |
POM192-H. australis-M-NSW-2003 |
| POM273 |
POM273-H. distinctus-F-VIC-1949 |
| POM308 |
POM308-H. fulvidorsalis-M-QLD-1950 |
| POM309 |
POM309-H. fulvidorsalis-F-QLD-1972 |
| POM312 |
POM312-H. fulvidorsalis-F-NT-1975 |
| POM329 |
POM329-H. distinctus-F-NSW-2017 |
| POM334 |
POM334-H. praepositus-F-SA-1987 |
| POM350 |
POM350-H. praepositus-F-WA-1979 |
Key: POM = Pompilidae; H = Heterodontonyx; M = Male; F = Female; NT = Northern Territory; QLD = Queensland; SA = South Australia; NSW = New South Wales; VIC = Victoria; ACT = Australian Capital Territory; WA = Western Australia.