Open-access, online, rapid taxonomy
ISSN: 2653-4649 (Online)

Australian Journal of

Taxonomy

This paper has been reviewed by two reviewers. As Subject Editor handling the paper, your task is now to review the reviewers' comments and add comments of your own as necessary, using the Comment buttons.

Please comment on issues with the text that you pick up that the reviewers have not commented on, or responses from you to the author based on the reviewer comments (for example, agreeing, disagreeing, or clarifying the reviewer comments).

The reviewers' recommendations are given at the bottom of the page. Please also add your own general comments to the author, as necessary, in the Editor comments box.

Remember, you need to assess each taxon (if any) as well as the main body of the paper.

You may save your comments at any stage using the Save button at the bottom of the page, and return to the review at a later time. If you plan to do your assessment over multiple sessions, you must Save the main page and each taxon page for which you add comments.

When both the main text and all the taxa are assessed, check the Done checkbox, and Submit.

Note: if you need to make edits to the paper, in your capacity as Subject Editor please do so using the button below. If you need to grant full edit access to the paper to the author, please right-click on the button, choose 'Copy link address', and send the link to the lead authors by email with instructions on what you need them to do.

Edit Paper

Please Wait...

THIS IS A REVIEW COPY OF AN UNPUBLISHED MANUSCRIPT. PLEASE DO NOT DISTRIBUTE OR SHARE

Molecular phylogeny, species delimitation and taxonomic revision of the Australian spider wasp genus Heterodontonyx Haupt, 1935 (Hymenoptera: Pompilidae)

Rev1:

Rev2:

Comment

You have completed and submitted this review. Thanks. It is now being revised by the author and is no longer accessible.

Mahin Sadat Chavoshi Jolfaei [1,2], Juanita Rodriguez [1]

1. Australian National Insect Collection, CSIRO, Acton 2601, Australian Capital Territory, Australia
2. Australian National University, Acton 2601, Canberra, Australian Capital Territory, Australia

Mahinsadat.Chavoshijolfaei@csiro.au

Rev1:

Rev2:

Comment

Abstract

Heterodontonyx (Pompilidae: Pepsinae) is an understudied genus with Australasian distribution and most species endemic to Australia. There have recently been some taxonomic changes involving the genera Heterodontonyx Haupt and Cryptocheilus Panzer, based on molecular and morphological evidence, however, the phylogenetic relationships within Heterodontonyx have not been determined, and species delimitation and a formal taxonomic revision have not been conducted. Here, we construct a Maximum Likelihood phylogeny estimate of Heterodontonyx based on a region of the nuclear ribosomal operon spanning the internal transcribed spacer (ITS) 1, 5.8S, and ITS2 for 31 samples representing seven Heterodontonyx morphospecies. A species delimitation analysis was performed in bPTP, which informed the taxonomic revision of the genus. Phylogenetic analyses strongly support the monophyly of Heterodontonyx and the species delimitation method recognizes eight species among which one is described as new, Heterodontonyx wahisi sp. nov.

Taxonomic revision includes 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: Isopedella leai (Hirst) based on the identification of the host by spider species author (Hirst).

urn: lsid:zoobank.org:pub:94EFABCD-CE9C-4F5B-9A9A-C8447CDFAF10

Rev1:

Rev2:

Comment

Introduction

There are five subfamilies within the family Pompilidae (Waichert et al. 2015), three of which occur in Australasia (Pompilinae, Ceropalinae, and Pepsinae). Pepsinae of Australia are divided into the tribes Pepsini and Ageniellini. Within the Pepsini, the genus Heterodontonyx Haupt, is distributed in Australia including Tasmania, New Guinea, New Caledonia, and New Zealand (Elliott, 2007). Recently, molecular, and morphological data provided a delineation of these genera, with some taxonomic changes involving the genera Heterodontonyx and Cryptocheilus Panzer that warranted taxonomic revision of both genera (Chavoshi et al, 2023, submitted). In this study, we utilised both molecular and morphological data to examine the phylogeny and specific diversity of Heterodontonyx species and provide a taxonomic revision of the genus.

Rev1:

Rev2:

Comment

Materials & Methods

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) were included as ingroups and Cryptocheilus fulvus (Eversmann) was nominated as the outgroup (see Figure 1 and Table S1). The material was identified to species level using identification keys developed based on the species descriptions and careful specimen examination, including that of type specimens. One to three legs of adult spider wasps were cleaned and homogenized for proteolysis on the first day of the 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 QIAGEN DNeasy Blood & Tissue Kit (Cat No: 69504) as per the manufacturer’s instruction. The purified DNA was resuspended in 25ul of Tris-HCl 1mM 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.). The 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 1mM to reach an equimolar concentration of 2nM followed by post-PCR clean-up of the pooled libraries using 1x magnetic bead ratio (NEBNext Sample Purification Beads). Lastly, 25-200 ul of the pooled, cleaned, single-indexed (using NEBNext Multiplex Oligos for Illumina) libraries were submitted for whole-genome shotgun sequencing in one lane of NextSeq 550 System 300 cycles (paired-end reads) at the ACRF Biomolecular Resource Facility (jcsmr.anu.edu.au).

Data processing, sequence alignment phylogenetic assessment, and species delimitation

Raw reads were cleaned and trimmed with Trimmomatic (Bolger et al., 2014), and quality checked with FastQC (Babraham Bioinformatics). The resulting trimmed reads were de novo assembled using Trinity v2.1.1 (Grabherr et al., 2011). References for the internal transcriber spacer 1 (ITS1), 5.8S, and ITS2 were obtained using Geneious mapper by mapping POM334 (See Table S1) to a 5.8S sequence of Sphaerophthalma orestes (Mutillidae) (NCBI accession no.DQ408488) from (Pilgrim et al., 2002) with medium-low sensitivity and 25 iterations. The consensus was then used as the reference 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 (Nguyen et al., 2015) with Model Finder (Kalyaanamoorthy et al., 2017) and ultrafast Bootstrap (Hoang et al., 2018). 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 the delimitation scheme with PTP and bPTP on the server (https://species.h-its.org/). Data available at: https://doi.org/10.5061/dryad.5qfttdzbj

Taxonomic revision of Heterodontonyx species

Where possible, images of the type material of each species were included, and specimens were examined using a Leica S7 APO dissecting microscope. Images that were taken at the Australian National Insect Collection (Canberra, ACT 2601 Australia) used a Canon EOS 7D Mark II DSLR with Visionary Digital Big Kahuna (Bk) Camera system with 65 mm (Canon lens MP-E65mm f/2.8 1-5x Macro) and 100 mm (Canon lens EF 100mm f/8 L IS USM), processed using Helicon focus Stacker and edited with Photoshop CS5. Specimens were examined from the following collections: Australian Museum (AM); Australian National Insect Collection (ANIC); Humbolt University Zoologischen Museum (HUM); Museo Civico di Storia Naturale 'Giacomo Doria' (MCG); Museum Victoria (MV); Natural History Museum (NHM); Queensland Museum (QM); South Australian Museum (SAM); and Western Australian Museum (WAM). Morphological terminology used here follows Harris (1987) and wing terminology follows Day (1988).

Change "9 locations" to "nine locations" [numbers less than 10 are usually spelled in full in general English writing].

Please provide author & date for the outgroup species.

Rev1:

Change "9 locations" to "nine locations" [numbers less than 10 are usually spelled in full in general English writing].

Please provide author & date for the outgroup species.

Rev2:

Comment

Results

Phylogenetic analysis and species delimitation

The IQ-TREE analyses overall yielded well-supported interspecific relationships within Heterodontonyx (Fig.1). The ingroup (node A) is split into 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). The molecular new species for which the morphological description follows in the next section (POM 168, Figure 1), is sister to node E containing H. darwinii, H. fulvidorsalis, and H. bicolor. Heterodontonyx darwinii is sister to H. fulvidorsalis+H. bicolor (node F in Figure 1). Heterodontonyx 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).

Rev1:

Rev2:

Comment

Discussion

We present a phylogeny, molecular species delimitation, and systematic revision of the genus Heterodontonyx. 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).

Rev1:

Rev2:

Comment

Figure 1. Maximum Likelihood tree reconstruction of Heterodontonyx based on molecular data (ITS1, 5.8S, ITS2 region from the ribosomal operon). Statistical support values of Ultra-Fast Bootstraps are shown on each node. The suggested delimitation scheme of molecular units produced by bPTP is shown on the right panel next to morphological identification.

Rev1:

Rev2:

Comment

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: 2mm, b, d: 1mm.

Image credit: Lars Krogmann from HUM.

Rev1:

Rev2:

Comment

Rev1:

Rev2:

Comment

Rev1:

Rev2:

Comment

Rev1:

Rev2:

Comment

Table content

I agree with the reviewers that the Key needs to be more tightly edited, especially by placing a capital letter for the first word.

Rev1:

I agree with the reviewers that the Key needs to be more tightly edited, especially by placing a capital letter for the first word.

Rev2:

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.

Key title

Rev1:

Rev2:

Comment

Taxonomy

Taxonomic revision

Heterodontonyx Haupt, 1935. Type species: Heterodontonyx basalis Haupt, 1935 (Figure 2) by subsequent designation, see Banks, 1941:234 and Pate, 1946: 90 (=Sphex bicolor, synonymy by Wahis (Wahis, 2008).

Diagnosis

Black and orange wasps of small to large size (10mm-40mm) with golden-yellow wings; clypeus medium to large and convex, with labrum concealed to completely exposed; antennal segments slender and elongated; 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, the ratio of the inter-tegular distance to the length of the second abscissa of the forewing vein Rs (ITD:SaRs) ranges from 1.5 to 2.5. Female metatibia serrated with spines.

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 is not distinctly dimorphic.

1. a. Head, mesosoma and metasoma black . H. solomonis Turner (Fig. 10)

b. Body black and orange/yellow/reddish-yellow. 2

2. a. Sides of propodeum rounded; body size small (10–20 mm) .3

b. Full tubercles or half tubercles on sides of propodeum; body size large (20mm-40mm). 4

3. a. Body with black setae; size 10-15 mm.5

b. Body bare, without long setae; second metasomal segment sometimes with apical black band. H. australis (Guérin–Méneville, 1838) (Fig. 3)

4. a. Full tubercles on sides of propodeum.6

b. Half tubercles on sides of propodeum. H. fulvidorsalis(Turner, 1910) (Fig. 8)

5. a. Female head entirely orange, some males with pronotum golden yellow-orange, body size 10-15mm.H. distinctus (Smith, 1868) (Fig. 6)

b. Colouration variable, metasomal segment sometimes with an apical black band; body size 10-15mm.H. praepositus (Haupt, 1935) (Fig. 9)

6. a. Clypeus long and narrow (length: width: 1.5-1.75); labrum completely exposed.7

b. Clypeus short and wide (length: width: 2-2.5); labrum slightly to moderately exposed. 8

7. a. Pubescence moderate; antennae very long and slender; head, pronotum, and mesoscutum reddish-yellow brown; slight compression between first and second metasomal segments. H. darwinii (Turner, 1910) (Fig. 5)

b. Body covered with dense, black, and golden setae; pronotum orange, first metasomal segment, and apical second metasomal segment black, remaining metasomal segments orange.H. wahisi sp. nov (Fig. 12)

8. a. Mesosoma entirely black, second metasomal segment orange.H. tuberculatus (Smith, 1855) (Fig. 11)

b. Mesosoma entirely black, second metasomal segment apically or entirely black.9

9. a. Second metasomal segment entirely black. H. erythroura (Cameron 1906) (Fig. 7)

b. Second metasomal segment with black basal band. Heterodontonyx bicolor (Fabricius, 1775) (Fig. 4)

Rev1:

Rev2:

Comment

Acknowledgments

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, and Olivia Evangelista for technical help.

Rev1:

Rev2:

Comment

Supplementary Material

Table S1, ITS phylogeny and tree file of Heterodontonyx species deposited in Dryad: https://doi.org/10.5061/dryad.5qfttdzbj

Rev1:

Rev2:

Comment

References

Banks, N. (1941) Psammocharidae from the Solomon Islands, Prince of Wales Island and New Caledonia. Occasional Papers of the Bernice Pauahi Bishop Museum, 16(10), 231–245.

Bolger, A. M., Lohse, M., & Usadel, B. (2014). Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics, 30(15), 2114-2120.

Cameron, P. (1906a) Hymenoptera I (all families except Apidae and Formicidae). Nova Guinea, 5. Zoologie, 41–65.

Day, M. C. (1988). Handbooks for the Identification of British Insects, SPIDER WASPS, HYMENOPTERA: POMPILIDAE. ROYAL ENTOMOLOGICAL SOCIETY OF LONDON

Elliott, M. G. (2007). Annotated catalogue of the Pompilidae (Hymenoptera) of Australia. Zootaxa(1428), 1-83.

Evans, H. E. (1953). Comparative Ethology and the Systematics of Spider Wasps. Systematic Zoology, 2(4), 155-172.

Evans, Y. (1962). The ecology and nesting behavior of the pompilidae (Hymenoptera) of the Northeastern United States. Entomological Society of America.

Evans, H. E. (1963). Predatory wasps. Scientific America, 208(4), 145-154.

Evans, H. E. (1966). A revision of the Mexican and Central American spider wasps of the subfamily Pompilinae (Hymenoptera: Pompilidae) (Vol. 20). Philadelphia, American Entomological Society,1916. (Memoirs of the American Entomological Society)

Evans, H. E., & Shimizu, A. (1996). The evolution of nest building and communal nesting in Ageniellini (Insecta: Hymenoptera: Pompilidae). Journal of Natural History, 30(11), 1633-1648.

Evans, H.E. & Matthews, R.W. (1973) Behavioural observations on some Australian spider wasps (Hymenoptera, Pompilidae). Transactions of the Entomological Society of London, 125(1), 45–55.

Evans, H.E. (1981) The genus Microphadnus Cameron in Australia (Hymenoptera, Pompilidae). Entomological News, 92(2), 81–83.

Fabricius, J. C. (1775). Systema Entomologicae, sistens insectorum classes, ordines, genera, species, synonyma auctorum, loca natalia, metamorphosis adiectis synonimis, locis, descriptionibus, observationibus. Flensbvrgi et Lipsiae.

Fabricius, J.C. (1781) Species Insectorum exhibentes eorum differentias specificas, synonyma auctorum, loca natalia, metamorphosis adiectis observationibus, desciptionibus. C.E. Bohni, Hamburgi er Lionii [=Hamburg & Cologne], iii–viii + 552 pp. [In Latin]

Fabricius, J.C. (1787) Mantissa insectorum sistens species nuper detectas adiectis characteribus genericis, differentiis specificis, emendationibus, observationibus. Tome I. Proft, Hafniae [=Copenhagen], 348 pp. [In Latin]

Fabricius, J.C. (1793) Entomologia Systematica Emendata et Aucta. Secundum classes, ordines, genera, species. Adjectis synonimis, locis, observationibus, descriptionibus. Tome II. C.G. Proft, Hafniae [=Copenhagen], viii + 519 pp. [In Latin]

Fabricius, J.C. (1804) Systema Piezatorum. Carolum Reichard, Brunsvigae, 439 pp. [In Latin]

Forster, P.I. (1994) Diurnal Insects Associated with the Flowers of Gomphocarpus physocarpus E. Mey. (Asclepiadaceae), an Introduced Weed in Australia. Biotropica, 26(2), 214–217

Froggatt, W.W. (1892) Catalogue of the described Hymenoptera of Australia. Part 2. Proceedings of the Linnean Society of New South Wales (2)7, 205–248.

Gmelin, J.F. (1790) Caroli a Linné Systema Naturae. Vol. 1. Part 5. Georg Emanuel Beer, Lipsiae, pp. 2225–3020. [In Latin]

Grabherr, M. G., Haas, B. J., Yassour, M., Levin, J. Z., Thompson, D. A., Amit, I., Adiconis, X., Fan, L., Raychowdhury, R., Zeng, Q. D., Chen, Z. H., Mauceli, E., Hacohen, N., Gnirke, A., Rhind, N., di Palma, F., Birren, B. W., Nusbaum, C., Lindblad-Toh, K., Friedman, N., & Regev, A. (2011). Full-length transcriptome assembly from RNA-Seq data without a reference genome. Nature Biotechnology, 29(7), 644-U130.

Guérin–Méneville, F.É. (1838) Chapitre XIII. Insectes. pp. 57–302 in Duperrey, L.I. Voyage Autour du Monde, exécuté par ordre du Roi, sur la corvette de S.M. la Coquille, pendant les anées 1822–1825. Zoologie. Vol. 2 Part 2. Arthus Bertrand, Paris, 319 pp [dated 1830, issued 1838].

Guiglia, D. (1948) I tipi di Imenotteri Guerin esistenti nelle collezioni del Museo di Genova. Annali Museo Civico di Storia Naturale Genova, 63, 175–191. [In Italian]

Guiglia, D. & Pasteels, J. (1961) Aggiunte ed osservazioni all'elenco delle species di imenotteri descritte da Guerin–Meneville che si trovano nelle collezioni del museu di Genova. Annali Museo Civico di Storia Naturale Genova, 72, 17–20. [In Italian]

Harris, A. C. (1987). Fauna of New Zealand (Vol. 12). DSIR Science Information Publishing Centre.

Harris, A.C. (1999) The life histories and nesting behaviour of the Pompilidae (Hymenoptera) in New Zealand, a comparative study. Species Diversity, 4, 143–235.

Haupt. H. (1935a) Psammocharidae. pp. 306–321 In: Schulthess Rechberg, A. Hymenoptera aus den Sundainseln und Nordaustralien (mit Ausschluss der Blattwespen, Schlupfwespen und Ameisen). Revue Suisse de Zoologie, 42(9), 293–323. [In German].

Haupt, H. (1935b) Psammocharidae mediterraneae. II. Bollettino del Laboratorio di Entomologia del R. Istituto Superiore Agrario di Bologna, 7, 263–302. [In German]

Hirst, D.B. (1992) Revision of the Genus Isopeda Koch (Heteropodidae, Araneae) in Australia. Invertebrate Taxonomy 6, 337–387.

Hoang, D. T., Chernomor, O., von Haeseler, A., Minh, B. Q., & Vinh, L. S. (2018). UFBoot2: Improving the Ultrafast Bootstrap Approximation. Molecular Biology and Evolution,35(2), 518-522.

Kalyaanamoorthy, S., Minh, B. Q., Wong, T. K. F., von Haeseler, A., & Jermiin, L. S. (2017). ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods, 14(6), 587-+.

McKeown, K.C. (1942) Australian Insects. An introductory Handbook. Royal Zoological Society of New South Wales, Sydney, 304 pp.

Nguyen, L. T., Schmidt, H. A., von Haeseler, A., & Minh, B. Q. (2015). IQ-TREE: A Fast and Effective Stochastic Algorithm for Estimating Maximum-Likelihood Phylogenies. Molecular Biology and Evolution, 32(1), 268-274.

Pate, V.S.L. (1946) The generic names of spider wasps (Psammocharidae olim Pompilidae) and their type species (Hymenoptera Aculeata). Transactions of the American Entomological Society, 72, 65–137.

Pilgrim, E. M., Roush, S. A., & Krane, D. E. (2002). Combining DNA sequences and morphology in systematics: testing the validity of the dragonfly species Cordulegaster bilineata. Heredity, 89, 184-190.

Schulz, G.L. (1906) Spolia Hymenopterologica. Pederborn, Berlin, 355 pp. [In German]

Schulz, G.L. (1908) Fossores. In: W. Michaelsen & R. Hartmeyer (Eds.) Die Fauna Südwest–Australiens. Ergebnisse der

Hamburger südwest–australischen Forschungsreise. Vol. 1. Part 13. G. Fischer, Jena, pp. 447–488. [In German] Schulz, W.A. (1911) Zweihundert alte Hymenopteren. Zoologischer Anzeiger, 4, 1–220. [In German]

Smith, F. (1855) Catalogue of hymenopterous insects in the collection of the British Museum. Mutillidae and Pompilidae. Vol. 3. British Museum of Natural History, London, 206 pp.

Smith, F. (1863a) Catalogue of hymenopterous insects collected by Mr. A.R. Wallace in the islands of Mysol, Ceram, Waigiou, Bouru and Timor. Journal of the Proceedings of the Linnean Society of London, 7, 1–48.

Smith, F. (1863b) Notes on the geographical distribution of the aculeate Hymenoptera collected by Mr. A. R. Wallace in the Eastern Archipelago. Journal of the Proceedings of the Linnean Society of London, 7, 109–145.

Smith, F. (1868) Descriptions of aculeate Hymenoptera from Australia. Transactions of the Entomological Society of London, 1868, 231–258.

Smith, F. (1871) A Catalogue of the Aculeate Hymenoptera and Ichneumonidae of India and the Eastern Archipelago. Journal of the Linnaean Society of London. Zoology, 11, 285–415.

Turner (1910) Additions to our knowledge of the fossorial Hymenoptera of Australia. Proceedings of the Zoological Society of London, 1910, 253–356.

Tillyard, R.J. (1926) The insects of Australia and New Zealand. Angus and Robertson, Sydney, 560 pp.

Waichert, C., Rodriguez, J., Wasbauer, M. S., von Dohlen, C. D., & Pitts, J. P. (2015). Molecular phylogeny and systematics of spider wasps (Hymenoptera: Pompilidae): redefining subfamily boundaries and the origin of the family. Zoological Journal of the Linnean Society, 175(2), 271-287.

Williams, G. & Adam, P. (1995) Records of aculeate wasps from flowering subtropical rainforest trees. Australian Entomologist, 22(2), 51–58.

Zimsen, E. (1964) The Type Material of I.C. Fabricius. Munksgaard, Copenhagen, 656 pp.

Rev1:

Rev2:

Comment

Success message

Your decision

Review completed?

Done

Clicking Submit will send an email to the author.

Would you like to be cc'd?

Save

Success message

You have chosen to reject this paper. Please indicate your reasons for rejection in the General Comments text box above. These will be used to generate an email to the author after you click the OK button. (Note that you will be able to edit the content of that email before sending it.)

Allow or encourage resubmission

Disallow or discourage resubmission

Please choose a resubmission option

The email text below will be sent to the author. Please review it, amend as appropriate, then click the Send Email button. Doing so will set the status of the paper as Rejected.

Would you like to be cc'd?

Please choose a resubmission option

Enter the reasons for your rejection below. They will be stored, and added to an email that will be sent to the lead author

Open-access, online, rapid taxonomy ISSN: 2653-4649 (Online)

Australian Journal of

Taxonomy

Open-access, online, rapid taxonomy
ISSN: 2653-4649 (Online)