Abstract
Tropical and subtropical regions of East Asia harbor enormous yet undiscovered diversity of velvet ants. Nevertheless, continental Southeast Asia has been less intensively studied compared to Northeast Asia and islands of the Pacific Ocean. In this paper, the velvet-ant fauna of Vietnam is reviewed based on primary literature and specimens obtained in recent field surveys and from various museum collections. Before this study, fifty species were recorded, mostly from the northern provinces. Here, we examined 734 specimens and the following twenty-seven species are newly recognized from this country: Pherotilla brothersi Zhou, Lelej & Xu, 2017 (Rhopalomutillinae), Hemutilla bifurcata (Chen, 1957), H. granulata Tu, Lelej & Chen, 2014 (Sphaeropthalminae), Bischoffitilla malica (Zavattari, 1914), B. mammallifera (Chen, 1957), B. perakensis (Pagden, 1934), B. tumidula (Mickel, 1933) (Myrmillinae), Kurzenkotilla harmandi (André, 1898) (Mutillinae: Mutillini), Andreimyrme davidi (André, 1898), Mickelomyrme exiloides (Magretti, 1892), M. isora (Cameron, 1900), M. puttasoki Williams, 2019, Promecilla decora (Smith, 1879), P. levinaris (Chen, 1957), Sinotilla ansula (Chen, 1957), S. hong Lelej, 1995, S. trisecta (Chen, 1957), Smicromyrme thaochani Williams, 2019 (Mutillinae: Smicromyrmini), Krombeinidia chang Williams, 2019, Kr. meeungensis (Cockerell, 1928), Neotrogaspidia recticarinata (Chen, 1957), Orientidia thoawanae Williams, 2019, Pagdenidia hymalajensis (Radoszkowski, 1885), Promecidia abnormis Lelej, 2016, Trogaspidia wilsoni Williams, 2019, Wallacidia retinula (Chen, 1957) (Mutillinae: Trogaspidiini), and Zeugomutilla pangi Lelej, 2017 (Mutillinae: Zeugomutillini). Additionally, 169 new provincial records are presented. As a supplement to the previously published works, specimen-based records are provided for the species recorded without reference to voucher material. Taxonomic history, geographic distribution, sex association, seasonal occurrence, and habitats are reviewed for all 77 species in 29 genera. The number of known species from each province was greatly variable, and 46.0% of the 63 provinces currently have no recorded velvet ants. Twenty-five of the 77 recorded species are described from both sexes. Hosts are known for only two species. These data highlight the need for intensive field surveys, specifically in the southern provinces, to elucidate the true biodiversity of velvet ants in this country, which is likely even higher than what is documented here.
References
- Allen, H.W. (1975) The genus Tiphia of the Indian Subcontinent. United States Department of Agriculture Technical Bulletin, 1509, 1–96, 6 pls.
- Allen, H.W. & Jaynes, H.A. (1930) Contribution to the taxonomy of Asiatic wasps of the genus Tiphia (Scoliidae). Proceedings of the United States National Museum, 76 (17), 1–105, 4 pls. https://doi.org/10.5479/si.00963801.76-2814.1
- André, E. (1896a) Notes pour servir à la connaissance de mutilles paléarctiques et description de quelques espèces nouvelles. Deuxieme partie. Mémoires de la Société Zoologique de France, 9, 261–277.
- André, E. (1896b) Étude sur les Mutillides existant dans les collections du Musée civique de Gênes. Annali del Museo Civico di Storia Naturale di Genova, 37, 66–104.
- André, E. (1898) Étude sur les Mutillides du Muséum de Paris. Annales de la Société Entomologique de France, 67, 1–79.
- André, E. (1899) Les types des mutillides de la collection O. Radoszkowski. Annales de la Société Entomologique de France, 68, 1–43. https://doi.org/10.1080/21686351.1899.12279726
- André, E. (1901–1902a) Matériaux pour servir à la connaissance des Mutillides d’Afrique. Zeitschrift für Systematische Hymenopterologie und Dipterologie, 1 & 2, 279–288 + 305–352 (1901) & 17–48 (1902a).
- André, E. (1902b [1903]) Family Mutillidae. In: Wytsman, P. (Ed.), Genera Insectorum. Fascicles 11. L. Desmet-Verteneuil, Bruxelles, pp. 1–77, 3 pls.
- André, E. (1904) Examen critique d’une nouvelle classification proposée par M. le Dr. W. H. Ashmead pour la famille des Mutillidae. Revue d’Entomologie, 23 (1), 27–41.
- André, E. (1905) Mutillides nouveaux ou peu connues de l’Ancien Monds et de l’Australie. Zeitschrift für systematische Hymenopterologie und Dipterologie, 5, 265–277.
- André, E. (1909) Mutillides nouveaux ou imparfaitement connus du Musée royal d’histoire naturelle de Leide. Notes from the Leyden Museum, 31, 171–180.
- Añino, Y.J., Cambra, R.A., Windsor, D.M., Williams, K.A., Bartholomay, P.R., Sánchez, V. & Quintero, D. (2020) Seasonal and annual abundance of Ephuta wasp (Hymenoptera: Mutillidae) in Panama. Revista de Biología Tropical, 68 (2), 573–579. https://doi.org/10.15517/rbt.v68i2.38876
- Aranda, R. & Graciolli, G. (2016) Environmental heterogeneity in parasitoid-host interaction for Mutillidae (Hymenoptera: Apocrita). Environmental Entomology, 45 (5), 1146–1153. https://doi.org/10.1093/ee/nvw098
- Ashmead, W.H. (1899) Superfamilies in the Hymenoptera and generic synopses of the families Thynnidae, Myrmosidae, and Mutillidae. Journal of the New York Entomological Society, 7, 45–60.
- Ashmead, W.H. (1900–1904) Classification of the fossorial, predaceous and parasitic wasps, or the superfamily Vespoidea. The Canadian Entomologist, 32, 34, 35 & 36, 145–155 + 185–188 + 295–296 (1900), 79–88 + 131–137 + 163–166 + 203–210 + 219–231 + 268–273 + 287–291 (1902), 3–8 + 39–44 + 95–107 + 155–158 + 199–205 + 303–310 + 323–332 (1903) & 5–9 (1904).
- Averyanov, L.V., Loc, P.K., Hiep, N.T. & Harder, D.K. (2003) Phytogeographic review of Vietnam and adjacent areas of eastern Indochina. Komarovia, 3, 1–83.
- Ball, L., Rodríguez-Machado, S., Paredes-Burneo, D., Rutledge, S., Boyd, D.A., Vander Pluym, D., Babb-Biernacki, S., Chipps, A.S., Öztürk, R.Ç., Terzi, Y. & Chakrabarty, P. (2025) What ‘unexplored’ means: mapping regions with digitized natural history records to look for ‘biodiversity blindspots’. PeerJ, 13, e18511. https://doi.org/10.7717/peerj.18511
- Baltazar, C.R. (1966) A catalogue of Philippine Hymenoptera (with a bibliography, 1758–1963). Pacific Insects Monograph, 8, 1–488.
- Barthélémy, C. & Guénard, B. (2025) An annotated and illustrated checklist of Hong Kong aculeate wasps (Hymenoptera: Apoid wasps; Pompilioidea; Scolioidea; Thynnoidea; Tiphioidea; Vespoidea; Chrysidoidea). Insecta Mundi, 1115, 1–230.
- Barthélémy, C., Lelej, A.S. & Loktionov, V.M. (2018) A contribution to the Mutillidae (Hymenoptera) of Hong Kong. Zootaxa, 4514 (2), 230–242. https://doi.org/10.11646/zootaxa.4514.2.6
- Bartholomay, P.R., Williams, K.A., Lopez, V.M. & Oliveira, M.L. (2019) Revision of the Traumatomutilla americana species group (Hymenoptera: Mutillidae). Zootaxa, 4608 (1), 1–34. https://doi.org/10.11646/zootaxa.4608.1.1
- Bingham, C.T. (1897) The fauna of British India including Ceylon and Burma: Hymenoptera, I. Wasps and bees. Taylor and Francis, London, 608 pp., 4 pls. https://doi.org/10.5962/bhl.title.100738
- Bingham, C.T. (1903) Diagnoses of aculeate Hymenoptera. Fasciculi Malayenses, Zoology, 1 (2), appendix, iii–vii.
- Bingham, C.T. (1905) Report on the aculeate Hymenoptera. Fasciculi Malayenses, Zoology, 1 (3), 19–60, pl. A.
- Bischoff, H. (1920–1921) Monographie der Mutilliden Afrikas. Archiv für Naturgeschichte, 86A (1–3 & 4–5), 1–480 (1920) & 481–830 (1921).
- Borries, H. (1892) Mutilla erythrocephala Fabr. som parasit hos Crabro (Solenius) rubicola D. & P. Entomologisk Tidskrift, 13, 247–249.
- Boutin, M. & Vilhelmsen, L. (2024 [2025]) Colour pattern diversity and evolution in Oriental velvet ants (Hymenoptera: Aculeata: Mutillidae). Biological Journal of the Linnean Society, 145 (1), blae064. https://doi.org/10.1093/biolinnean/blae064
- Bradley, J.C. & Bequaert, J. (1923) Studies in African Mutillidae. Revue Zoologique Africaine, 11 (3), 211–258.
- Bradley, J.C. & Bequaert, J. (1928) A synopsis of the Mutillidae of the Belgian Congo. Bulletin of the American Museum of Natural History, 58 (2), 63–122.
- Branstetter, M.G., Danforth, B.N., Pitts, J.P., Faircloth, B.C., Ward, P.S., Buffington, M.L., Gates, M.W., Kula, R.R. & Brady, S.G. (2017) Phylogenomic insights into the evolution of stinging wasps and the origin of ants and bees. Current Biology, 27 (7), 1019–1025. https://doi.org/10.1016/j.cub.2017.03.027
- Brothers, D.J. (1975) Phylogeny and classification of the aculeate Hymenoptera, with special reference to Mutillidae. The University of Kansas Science Bulletin, 50 (11), 483–648.
- Brothers, D.J. (1989) Alternative life-history styles of mutillid wasps (Insecta, Hymenoptera). In: Bruton, M.N. (Ed.), Alternative life-history styles of animals. Perspectives in vertebrate sciences 6. Springer, Dordrecht, pp. 279–291. https://doi.org/10.1007/978-94-009-2605-9_14
- Brothers, D.J. (1995) Mutillidae. In: Gauld, I. & Hansen, P. (Eds.), The Hymenoptera of Costa Rica. Oxford University Press, Oxford, pp. 541–548.
- Brothers, D.J. (2006) Familia Mutillidae. In: Fernández, F. & Sharkey, M.J. (Eds.), Introducción a los Hymenoptera de la Región Neotropical. Sociedad Colombiana de Entomología y Universidad Nacional de Colombia, Bogotá, D.C., pp. 577–594.
- Brothers, D.J. (2015) Revision of the Rhopalomutillinae (Hymenoptera, Mutillidae): 1, generic review with descriptions of three new genera. Journal of Hymenoptera Research, 46, 1–24. https://doi.org/10.3897/JHR.46.5733
- Brothers, D.J. (2022) Critical analysis of the type material of Mutillidae described from the Australasian Region (Hymenoptera). Zootaxa, 5140 (1), 1–215. https://doi.org/10.11646/zootaxa.5140.1.1
- Brothers, D.J. & Carpenter, J.M. (1993) Phylogeny of Aculeata: Chrysidoidea and Vespoidea (Hymenoptera). Journal of Hymenoptera Research, 2 (1), 227–302.
- Brothers, D.J. & Lelej, A.S. (2017) Phylogeny and higher classification of Mutillidae (Hymenoptera) based on morphological reanalyses. Journal of Hymenoptera Research, 60, 1–97. https://doi.org/10.3897/jhr.60.20091
- Brothers, D.J., Lelej, A.S. & Williams, K.A. (2019) Genus-group names of Mutillidae (Hymenoptera): corrections and updates since 2008. Zootaxa, 4651 (3), 578–588. https://doi.org/10.11646/zootaxa.4651.3.10
- Bunyavejchewin, S., Baker, P.J. & Davies, S.J. (2011) Seasonally dry tropical forests in continental Southeast Asia: Structure, composition, and dynamics. In: McShea, W.J., Davies, S.J. & Bhumpakphan, N. (Eds.), The ecology and conservation of seasonally dry forests in Asia. Smithsonian Institution Scholarly Press, Washington D.C., pp. 9–35.
- Cameron, P. (1900) Hymenoptera Orientalia, or contributions to the knowledge of the Hymenoptera of the Oriental zoological region. Part IX. The Hymenoptera of the Khasia Hills. Part II. Section I. Memoirs and Proceedings of the Manchester Literary and Philosophical Society, 44 (15), 1–114.
- Cameron, P. (1901) On the Hymenoptera collected during the “Skeat Expedition” to the Malay Peninsula, 1899–1900. Proceedings of the Zoological Society of London, 2, 16–44. https://doi.org/10.1111/j.1469-7998.1901.tb08163.x
- Cameron, P. (1904) On some new species of Hymenoptera from northern India. The Annals and Magazine of Natural History, Series 7, 13 (76), 281–289. https://doi.org/10.1080/00222930409487070
- Campbell, L., Behrens, K., Hesse, C. & Chaon, P. (2021) Habitats of the world. Princeton University Press, Princeton, New Jersey, 568 pp. https://doi.org/10.2307/j.ctv1jhvngg
- Chan, M.-L., Yeh, J.-Y., Jeng, M.-L., Tsai, J.-F., Chen, Y.-L., Chen, L.-H. & Lin, S.-R. (2025) Integrated insect types database of Taiwanese species. National Museum of Natural Science, Taichung. Available from: http://twinsecttype.nmns.edu.tw/ (accessed 30 April 2025)
- Chen, C.-w. (1957) A revision of the velvety ants or Mutillidae of China (Hymenoptera). Quarterly Journal of the Taiwan Museum, 10 (3–4), 135–224, 6 pls.
- Clausen, C.P., Gardner, T.R. & Sato, K. (1932) Biology of some Japanese and Chosenese grub parasites (Scoliidae). United States Department of Agriculture Technical Bulletin, 308, 1–27.
- Cockerell, T.D.A. (1928) Mutillidae from Siam. The Annals and Magazine of Natural History, Series 10, 2 (12), 596–600. https://doi.org/10.1080/00222932808672926
- Cockerell, T.D.A. (1929) The genus Megaustenia. The Nautilus, 43 (2), 51–54. [https://www.biodiversitylibrary.org/item/34845]
- Dalla Torre, C.G. de (1897) Catalogus Hymenopterorum hucusque Descriptorum Systematicus et Synonimicus. Volumen VIII: Fossores (Sphegidae). Guilelmi Engelmann, Lipsiae, [4] + viii + 750 pp. https://doi.org/10.5962/bhl.title.10348
- Debevec, A.H., Cardinal, S. & Danforth, B.N. (2012) Identifying the sister group to the bees: A molecular phylogeny of Aculeata with an emphasis on the superfamily Apoidea. Zoologica Scripta, 41 (5), 527–535. https://doi.org/10.1111/j.1463-6409.2012.00549.x
- de Bruyn, M., Stelbrink, B., Morley, R.J., Hall, R., Carvalho, G.R., Cannon, C.H., van den Bergh, G., Meijaard, E., Metcalfe, I., Boitani, L., Maiorano, L., Shoup, R. & von Rintelen, T. (2014) Borneo and Indochina are major evolutionary hotspots for Southeast Asian biodiversity. Systematic Biology, 63 (6), 879–901. https://doi.org/10.1093/sysbio/syu047
- Dinerstein, E., Olson, D., Joshi, A., Vynne, C., Burgess, N.D., Wikramanayake, E., Hahn, N., Palminteri, S., Hedao, P., Noss, R., Hansen, M., Locke, H., Ellis, E.C., Jones, B., Barber, C.V., Hayes, R., Kormos, C., Martin, V., Crist, E., Sechrest, W., Price, L., Baillie, J.E.M., Weeden, D., Suckling, K., Davis, C., Sizer, N., Moore, R., Thau, D., Birch, T., Potapov, P., Turubanova, S., Tyukavina, A., de Souza, N., Pintea, L., Brito, J.C., Llewellyn, O.A., Miller, A.G., Patzelt, A., Ghazanfar, S.A., Timberlake, J., Klöser, H., Shennan-Farpόn, Y., Kindt, R., Lillesø, J.B., van Breugel, P., Graudal, L. & Voge, M. (2017) An ecoregion-based approach to protecting half the terrestrial realm. BioScience, 67 (6), 534–545. https://doi.org/10.1093/biosci/bix014
- Dover, C. (1926) A contribution to a list of the aculeate Hymenoptera (excepting ants) of Hongkong. The China Journal of Science & Arts, 4, 233–235.
- Fabricius, J.C. (1787) Mantissa insectorum sistens eorum species nuper detecta, adjectis characteribus genericis, differentiis specificis, emendationibus, observationibus. Tom. I. Impensis Christ. Gottl. Proft, Hafniae [Copenhagen], 348 pp. https://doi.org/10.5962/bhl.title.11657
- Fabricius, J.C. (1793) Entomologica Systematica emendata et aucta, secundum Classes, Ordines, Genera, Species adjectis Synonymis, Locis, Observationibus, Descriptionibus. Tom. II. Christ. Gottl. Proft, Hafniae [Copenhagen], viii + 519 pp. https://doi.org/10.5962/bhl.title.125869
- Fabricius, J.C. (1804) Systema Piezatorum secundum Ordines, Genera, Species adjectis Synonymis, Locis, Observationibus, Descriptionibus. Carolum Reichard, Brunsvigae, 470 pp. [pp. i–xiv + 15–440 + 1–30] https://doi.org/10.5962/bhl.title.10490
- Faith, D.P. (1992) Conservation evaluation and phylogenetic diversity. Biological Conservation, 61 (1), 1–10. https://doi.org/10.1016/0006-3207(92)91201-3
- Ferton, C. (1908) Notes détachées sur l’instinct des hyménoptères mellifères et ravisseurs (4e Série). Annales de la Société entomologique de France, 70, 535–586. https://doi.org/10.1080/21686351.1908.12279451
- Fox, W.J. (1894) A proposed classification of the fossorial Hymenoptera of North America. Proceedings of the Academy of Natural Sciences of Philadelphia, 1894, 292–307.
- Gall, B.G., Spivey, K.L., Chapman, T.L., Delph, R.J., Brodie, E.D. Jr. & Wilson, J.S. (2018) The indestructible insect: Velvet ants from across the United States avoid predation by representatives from all major tetrapod clades. Ecology and Evolution, 8 (11), 5852–5862. https://doi.org/10.1002/ece3.4123
- Giner Marí, J. (1944) Himenópteros de España. Fams. Apterogynidae y Mutillidae. Instituto Español de Entomologia, Madrid, 124 pp.
- Gôukon, K. (1993) Host records and some bionomical notes on mutillid wasps in Japan (Hymenoptera, Mutillidae). Chugoku Kontyu, 7, 35–40. [in Japanese]
- Hammer, K. (1949) Über einige von Kjell Kolthoff und anderen in China gesammelten Hymenoptera. Chrysididae, Cleptidae, Mutillidae. Arkiv för zoologi, 42A (8), 1–12.
- Hammer, K. (1962 [1960]) Mutilliden (Insecta: Hymenoptera) aus dem Indischen Museum in Calcutta. Records of the Indian Museum, 58 (1), 1–51. https://doi.org/10.26515/rzsi/v58/i1/1960/161983
- Hashimoto, Y. & Nakanishi, A. (1997) A list of the type-specimens of Hymenoptera described by K. Tsuneki in the Museum of Nature and Human Activities, Hyogo (MNHAH). Catalogue of collections in the Museum of Nature and Human Activities, Hyogo No. 2. Museum of Nature and Human Activities, Sanda, 47 pp., 8 pls.
- He, J.-H., Chen, X.-X., Fan, J.-J., Li, Q., Liu, C.-M., Lou, X.-M., Ma, Y., Wang, S.-F., Wu, Y.-R., Xu, Z.-H., Xu, Z.-F. & Yao, J. (2004) Hymenopteran insect fauna of Zhejiang. Science Press, Beijing, lvi + 1373 pp., xliii pls. [in Chinese]
- Helmus, M.R., Bland, T.J., Williams, C.K. & Ives, A.R. (2007) Phylogenetic measures of biodiversity. The American Naturalist, 169 (3), E68–E83. https://doi.org/10.1086/511334
- Hill, D.C. (1982) Hong Kong insects. Vol. 2. Urban Council Publication, Hong Kong, 144 pp.
- Hirayama, S. (1937) [The illustrated thousand insects of Japan. Vol. 2]. Sanseidô, Tokyo, 194 pp., 88 pls., 28 pp. + 3 + 40 pp. [in Japanese]
- Hortal, J., de Bello, F., Diniz-Filho, J.A.F., Lewinsohn, T.M., Lobo, J.M. & Ladle, R.J. (2015) Seven shortfalls that beset large-scale knowledge of biodiversity. Annual Review of Ecology, Evolution, and Systematics, 46, 523–549. https://doi.org/10.1146/annurev-ecolsys-112414-054400
- Hua, L.-z. (2006) List of Chinese insects Vol. IV. Sun Yat-sen University Press, Guangzhou, 540 pp.
- Hughes, A.C. (2017) Understanding the drivers of Southeast Asian biodiversity loss. Ecosphere, 8 (1), e01624. https://doi.org/10.1002/ecs2.1624
- Invrea, F. (1954) Mutillidi della Dalmazia e del litorale Montenegrino. Nota prima. Memorie della Societa Entomologica Italiana, 32, 143–159.
- Invrea, F. (1964) Mutillidae-Myrmosidae. Fauna d’Italia, 5. Edizioni Calderini, Bologna, xii + 302 pp., 1 p.
- Ito, R. (2018) A second record of Eotrogaspidia auroguttata (Smith, 1855) (Hymenoptera, Mutillidae, Mutillinae) from the Yaeyama Islands, Japan. Tsunekibachi, 32, 25–26. [in Japanese]
- Janzen, D.H. (1988) Tropical dry forests: The most endangered major tropical ecosystem. In: Wilson, E.O. (Ed.), Biodiversity. National Academy Press, Washington D.C., pp. 130–137.
- Jenkins, C.N., Pimm, S.L. & Joppa, L.N. (2013) Global patterns of terrestrial vertebrate diversity and conservation. Proceedings of the National Academy of Sciences, 110 (28), E2602–E2610. https://doi.org/10.1073/pnas.1302251110
- Kass, J.M., Guénard, B., Dudley, K.L., Jenkins, C.N., Azuma, F., Fisher, B.L., Parr, C.L., Gibb, H., Longino, J.T., Ward, P.S., Chao, A., Lubertazzi, D., Weiser, M., Jetz, W., Guralnick, R., Blatrix, R., Lauriers, J.D., Donoso, D.A., Georgiadis, C., Gomez, K., Hawkes, P.G., Johnson, R.A., Lattke, J.E., MacGown, J.A., Mackay, W., Robson, S., Sanders, N.J., Dunn, R.R. & Economo, E.P. (2022) The global distribution of known and undiscovered ant biodiversity. Science Advances, 8 (31), eabp9908. https://doi.org/10.1126/sciadv.abp9908
- Kato, M. (1934) Three colour illustrated insects of Japan. Fasc. X. Kôseikaku, Tokyo, 15 + 22 pp., 50 pls. [in Japanese]
- Kier, G., Kreft, H., Lee, T.M., Jetz, W., Ibisch, P.L., Nowicki, C., Mutke, J. & Barthlott, W. (2009) A global assessment of endemism and species richness across island and mainland regions. Proceedings of the National Academy of Sciences, 106 (3), 9322–9327. https://doi.org/10.1073/pnas.0810306106
- Krombein, K.V. (1940 [1939]) Studies in the Tiphiidae (Hymenoptera Aculeata) IV. A revision of the Myrmosinae of the New World with a discussion of the Old World species. Transactions of the American Entomological Society, 65 (4), 415–465. [https://www.jstor.org/stable/25077448]
- Krombein, K.V. (1976) Synonymical notes on two Palaearctic subgenera of Myrmosa Latreille. Polskie Pismo Entomologiczne, 46, 257–260.
- Krombein, K.V. (1979) Biosystematic studies of Ceylonese wasps, IV. Kudakrumiinae, a new subfamily of primitive wasps (Hymenoptera, Mutillidae). Transactions of the American Entomological Society, 105 (1), 67–83. https://doi.org/10.5479/si.00810282.298
- Krombein, K.V. & Lelej, A.S. (1999) Biosystematic studies of Ceylonese wasps, XXII: Bethsmyrmilla, a new genus of mutillid wasps (Hymenoptera: Mutillidae: Myrmillinae). Proceedings of the Entomological Society of Washington, 101 (1), 143–148.
- Kumar, G.P., Lelej, A.S., Das, D., Raveendran, H.K.P. & Loktionov, V.M. (2019) Discovery of the genus Kudakrumia Krombein, 1979 (Hymenoptera: Mutillidae) in India and description of a new species. Zootaxa, 4612 (2), 260–266. https://doi.org/10.11646/zootaxa.4612.2.8
- Kumpanenko, A., Gladun, D. & Vilhelmsen, L. (2022) Morphology of the sting apparatus in velvet ants of the subfamilies Myrmosinae, Dasylabrinae, Myrmillinae and Mutillinae (Hymenoptera: Mutillidae). Zoomorphology, 141 (1), 81–94. https://doi.org/10.1007/s00435-022-00552-w
- Latreille, P.A. (1802) Histoire naturelle, générale et particulière des Crustacés et des Insectes. Tome troisiéme. F. Dufart, Paris, xii + 467 pp. https://doi.org/10.5962/bhl.title.15764
- Lelej, A.S. (1978) Superfamily Mutilloidea. In: Tobias, V.I. (Ed.), Keys to the insects of the European part of the USSR. Vol. 3. Hymenoptera. Part 1. Nauka, Leningrad, pp. 71–84. [in Russian]
- Lelej, A.S. (1979) A new genus of velvet ants (Hymenoptera, Mutillidae) from south-east Asia. Zoologicheskiy Zhurnal, 58 (7), 1065–1067. [in Russian]
- Lelej, A.S. (1983) Contributions to the knowledge of the velvet ants (Hymenoptera, Mutillidae) of Australia. Entomologicheskoe Obozrenie, 62 (3), 612–619. [in Russian, English translation (1984) in: Entomological Review, 62 (3), 152–160]
- Lelej, A.S. (1985) Osy-nemki (Hymenoptera, Mutillidae) Fauny SSSR i Sopredelnykh Stran [The velvet ants (Hymenoptera, Mutillidae) of the USSR and neighboring countries]. Nauka, Leningrad, 268 pp. [in Russian]
- Lelej, A.S. (1992) Two interesting species of velvet ants from China (Hymenoptera, Mutillidae). Entomofauna, 13 (19), 281–288.
- Lelej, A.S. (1995a) A review of Palaearctic and Oriental species of genus Nemka Lelej with description of Oriental genus Mickelomyrme gen. n. (Hymenoptera, Mutillidae). Far Eastern Entomologist, 6, 1–20.
- Lelej, A.S. (1995b) To the knowledge of East Asian species of the tribe Smicromyrmini Bischoff, 1920 (Hymenoptera, Mutillidae) with description of four new genera and eight new species. Far Eastern Entomologist, 13, 1–28.
- Lelej, A.S. (1996a) A review of the East Asian species of Orientilla (Hymenoptera, Mutillidae). Memoirs of the Entomological Society of Washington, 17, 103–107.
- Lelej, A.S. (1996b) To the knowledge of the East Asian species of the tribe Trogaspidiini Bischoff, 1920 (Hymenoptera, Mutillidae) with description of eight new genera and two new species. Far Eastern Entomologist, 30, 1–24.
- Lelej, A.S. (1996c) Mutillid wasps collected in Malaysia and Indonesia by Dr. Sk. Yamane (Hymenoptera, Mutillidae). Tropics, 6 (1–2), 91–104. https://doi.org/10.3759/tropics.6.91
- Lelej, A.S. (1996d) A review of the East Asian species of Mickelomyrme Lelej, 1995 (Hymenoptera, Mutillidae). Entomofauna, 17 (15), 277–292.
- Lelej, A.S. (2002) Catalogue of the Mutillidae (Hymenoptera) of the Palaearctic Region. Dalnauka, Vladivostok, 172 pp.
- Lelej, A.S. (2005) Catalogue of the Mutillidae (Hymenoptera) of the Oriental Region. Dalnauka, Vladivostok, 252 pp.
- Lelej, A.S. (2012) A new species of Rhopalomutilla André, 1901 (Hymenoptera: Mutillidae) from South Vietnam. Russian Entomological Journal, 21 (2), 143–145. https://doi.org/10.15298/rusentj.21.2.06
- Lelej, A.S. (2020) Three new species of velvet ants from the tribe Smicromyrmini (Hymenoptera: Mutillidae) with unusual color syndrome. Far Eastern Entomologist, 421, 1–9. https://doi.org/10.25221/fee.421.1
- Lelej, A.S. (2021) A new species of the genus Petersenidia Lelej (Hymenoptera: Mutillidae) from Indonesia, eastward of Wallace’s line. Zootaxa, 5006 (1), 101–105. https://doi.org/10.11646/zootaxa.5006.1.13
- Lelej, A.S. (2023) Review of the tribe Ctenotillini (Hymenoptera: Mutillidae) from Oriental and Palaearctic Regions. Far Eastern Entomologist, 480, 1–22. https://doi.org/10.25221/fee.480.1
- Lelej, A.S. (2024) Review of the velvet ants (Hymenoptera: Mutillidae) of Wallacea with special reference to Lombok, Indonesia. Far Eastern Entomologist, 504, 1–25. https://doi.org/10.25221/fee.504.1
- Lelej, A.S. & Brothers, D.J. (2008) The genus-group names of Mutillidae (Hymenoptera) and their type species, with a new genus, new name, new synonymies, new combinations and lectotypifications. Zootaxa, 1889 (1), 1–79. https://doi.org/10.11646/zootaxa.1889.1.1
- Lelej, A.S., Choi, J.Y. & Tripotin, P. (2001) Review of the mutillid wasps (Hymenoptera, Mutillidae) of Korea. Far Eastern Entomologist, 96, 1–15.
- Lelej, A.S. & Kabakov, O.N. (1980) On Bradynobaenidae and Mutillidae (Hymenoptera) of Afghanistan. Entomologicheskoe Obozrenie, 59 (1), 181–196. [in Russian, English translation (1981) in Entomological Review, 59 (1), 141–156]
- Lelej, A.S. & Krombein, K.V. (1999) Two remarkable new genera of mutillid wasps (Hymenoptera: Mutillidae, Sphaeropthalminae, Pseudomethocini) from Thailand. Far Eastern Entomologist, 79, 1–8.
- Lelej, A.S. & Nemkov, P.G. (1997) Phylogeny, evolution and classification of Mutillidae (Hymenoptera). Far Eastern Entomologist, 46, 1–24.
- Lelej, A.S., Ovchinnikov, S.V. & Inayatullah, M. (2013) Velvet ants of the subfamily Myrmillinae (Hymenoptera: Mutillidae) of Pakistan. Russian Entomological Journal, 22 (3), 197–199.
- Lelej, A.S. & Schmid-Egger, C. (2005) The velvet ants (Hymenoptera, Mutillidae) of Central Europe. Linzer biologische Beiträge, 37 (2), 1505–1543.
- Lelej, A.S., Terine, J.B., Kumar, G.P., Das, D. & Sureshan, P.M. (2020) Review of the genus Odontomutilla Ashmead, 1899 (Hymenoptera: Mutillidae) of the Indian subcontinent, with description of two new species. Zootaxa, 4822 (2), 191–208. https://doi.org/10.11646/zootaxa.4822.2.3
- Lelej, A.S. & Yamane, Sk. (1992) Interesting velvet ants (Hymenoptera, Mutillidae) from Japan and Korea. Japanese Journal of Entomology, 60 (3), 625–632.
- Lelej, A.S. & Williams, K.A. (2023) Review of the tribe Smicromyrmini Bischoff, 1920 (Hymenoptera: Mutillidae). Zootaxa, 5231 (2), 101–140. https://doi.org/10.11646/zootaxa.5231.2.1
- Lelej, A.S., Williams, K.A., Loktionov, V.M., Pang, H. & Xu, Z.-F. (2017) Review of the genus Zeugomutilla Chen, 1957 (Hymenoptera, Mutillidae, Mutillini), with description of two new species. Zootaxa, 4247 (1), 1–15. https://doi.org/10.11646/zootaxa.4247.1.1
- Lelej, A.S., Williams, K.A., Terine, J.B., Okayasu, J., Parikh, G. & Kumar, G.P. (2023) Review of the tribe Mutillini (Hymenoptera: Mutillidae) from the Oriental Region. Zootaxa, 5228 (4), 455–476. https://doi.org/10.11646/zootaxa.5228.4.5
- Lelej, A.S., Zhou, H.-T., Loktionov, V.M. & Xu, Z.-F. (2016) Review of the genus Promecidia Lelej, 1996, with description of two new species from China (Hymenoptera, Mutillidae, Trogaspidiini). ZooKeys, 641, 103–120. https://doi.org/10.3897/zookeys.641.10765
- Lepeletier de Saint-Fargeau, A. (1845) Histoire Naturelle des Insectes. Hyménoptères. Tome troisième. Roret, Paris, [4] + 646 pp. https://doi.org/10.5962/bhl.title.9005
- Linnaeus, C. (1758) Systema Naturae per Regna Tria Naturae, secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. Editio Decima Reformata. Laurentii Salvii, Holmiae [Stockholm], [4] + 823 + [5] pp. https://doi.org/10.5962/bhl.title.542
- Lo Cascio, P. (2015) Worldwide checklist of the island mutillid wasps (Hymenoptera Mutillidae). Biodiversity Journal, 6 (2), 529–592.
- Magretti, P. (1892) Viaggio di Leonardo Fea in Birmania e regioni vicine XLIII. Imenotteri. Parte prima. Mutillidei, Scoliidei, Tifiidei, Tinnidei colla descrizione di parecchie nuove specie. Annali del Museo Civico di Storia Naturale di Genova, Serie 2, 12 (32), 197–266, pl. V.
- Marshall, L., Leclercq, N., Carvalheiro, L.G., Dathe, H.H., Jacobi, B., Kuhlmann, M., Potts, S.G., Rasmont, P., Roberts, S.P.M. & Vereecken, N.J. (2024) Understanding and addressing shortfalls in European wild bee data. Biological Conservation, 290, 110455. https://doi.org/10.1016/j.biocon.2024.110455
- Matsumura, S. (1911) Thousand insects of Japan supplement III. Keiseisha-shoten, Tokyo, 147 pp., pls. 30–41. https://doi.org/10.5962/bhl.title.50811
- Matsumura, S. (1930) The illustrated thousand insects of Japan vol. 2 (Hymenoptera). Tôkô Shoin, Tokyo, 198 pp., 18 pls., 89 + 8 pp. [in Japanese]
- Matsumura, S. (1931) 6000 illustrated insects of Japan-Empire. Tôkô Shoin, Tokyo, 3 + 1497 + 23 pp., 10 pls. [in Japanese]
- Mergler, C.J. & Gall, B.G. (2020) Has the indestructible insect met its match? Velvet ants as prey to bufonid toads. Ethology Ecology & Evolution, 33 (1), 15–24. https://doi.org/10.1080/03949370.2020.1789747
- Mickel, C.E. (1933a) The Mutillidae of Formosa. Annals of the Entomological Society of America, 26 (2), 381–423. https://doi.org/10.1093/aesa/26.2.381
- Mickel, C.E. (1933b) A new species and subspecies of Mutillidae from the Orient. Lingnan Science Journal, 12 (3), 273–288.
- Mickel, C.E. (1933c) The Mutillidae of Eastern Asia. Lingnan Science Journal, 12 (3), 289–325.
- Mickel, C.E. (1934) Mutillidae of the Philippine Islands. Philippine Journal of Science, 54 (1), 91–219, 1 pl.
- Mickel, C.E. (1935) The mutillid wasps of the islands of the Pacific Ocean (Hymenoptera, Mutillidae). Transactions of the Royal Entomological Society of London, 83 (2), 177–312. https://doi.org/10.1111/j.1365-2311.1935.tb01207.x
- Mickel, C.E. (1937a) The mutillid wasps of the genus Timulla which occur in North America north of Mexico. Entomological Americana, 17 (1–2), 1–113, pls. 1–3.
- Mickel, C.E. (1937b) New species and records of Mutillidae (Hymenoptera) from Borneo and the Solomon Islands. The Annals and Magazine of Natural History, Series 10, 19, 441–456. https://doi.org/10.1080/00222933708655284
- Mickel, C.E. (1938) The Neotropical mutillid wasps of the genus Timulla Ashmead (Hymenoptera: Mutillidae). Transactions of the Royal Entomological Society of London, 87 (22), 529–679. https://doi.org/10.1111/j.1365-2311.1938.tb00728.x
- Mickel, C.E. (1939) Correction of type locality for two species of Mutillidae described by Frederick Smith (Hymenoptera). Proceeding of the Royal Entomological Society of London, Series B, 8, 192–194. https://doi.org/10.1111/j.1365-3113.1939.tb00514.x
- Mittermeier, R.A., Robles-Gil, P., Hoffmann, M., Pilgrim, J., Brooks, T., Mittermeier, C.G., Lamoreux, J. & da Fonseca, G.A.B. (2004) Hotspots revisited: Earth’s biologically richest and most endangered terrestrial ecoregions. CEMEX, Mexico City, 390 pp.
- Mooney, H.A., Bullock, S.H. & Medina, E. (1995) Introduction. In: Bullock, S.H., Mooney, H.A. & Medina, E. (Eds) Seasonally dry tropical forests. Cambridge University Press, Cambridge, pp. 1–8. https://doi.org/10.1017/CBO9780511753398.001
- Morawitz, F. (1890) Hymenoptera Fossoria Transcaspica nova. Horae Societatis Entomologicae Rossicae, 24, 570–645.
- Moura, M.R. & Jetz, W. (2021) Shortfalls and opportunities in terrestrial vertebrate species discovery. Nature Ecology & Evolution, 5, 631–639. https://doi.org/10.1038/s41559-021-01411-5
- Murota, T. (1973) Sphecidae, Mutillidae, Scoliidae and Chrysididae collected in Formosa in 1972. Life Study, 17 (3–4), 115–120. [in Japanese]
- Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca, G.A.B. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853–858. https://doi.org/10.1038/35002501
- Nambu, T. & Tano, T. (1983) [Wasps and ants collected in Taiwan in 1980]. Hymenopterists’ Communication, 16, 11–23. [in Japanese]
- National Museum of Natural History (2025) Department of Entomology Collections. National Museum of Natural History, Washington D.C. Available from: https://collections.nmnh.si.edu/search/ento/ (accessed 30 April 2025)
- Newman, M., Ketphanh, S., Svengsuksa, B., Thomas, P., Sengdala, K., Lamxay, V. & Armstrong, K. (2007) A checklist of the vascular plants of Lao PDR. Royal Botanic Garden Edinburgh, Edinburgh, vi + 394 pp.
- Nguyen, T.P.L. & Dang, T.H. (2024) [Biodiversity of sting bees and wasps (Hymenoptera: Aculeata) at Me Linh station for biodiversity, Vinh Phuc]. Publishing house for Science and Technology, Hanoi, 280 pp. [in Vietnamese]
- Nonveiller, G. (1980) Recherches sur les Mutillides de l’Afrique (Hymenoptera, Mutillidae) XI. Révision des mâles du genre Trogaspidia Ashmead 1899 (♂♀) apparentés ou semblables à medon (Smith) 1855 (♂) avec description du nouveau sous-genre Tuberocoxotilla (♂♂). Memoires publies par l’Institut pour la Protection des Plantes, Beograd, 14, 69–108.
- Nonveiller, G. (1995) A preliminary approach to a revision of the Afrotropical representatives of the genus Trogaspidia Ashmead, 1899 (sensu Bischoff, 1920) (Hymenoptera, Mutillidae). Studies on African Mutillidae: Note XXI. (Third contribution to the knowledge of the Afrotropical Trogaspidia). Annales de la Societé Entomologique de France, New Series, 31 (4), 349–368. https://doi.org/10.1080/21686351.1995.12277734
- Okayasu, J. (2016) [First record of Eotrogaspidia auroguttata (Smith) from Yaeyama Islands, Japan]. Gekkan-Mushi, 543, 61–62. [in Japanese]
- Okayasu, J. (2017) Two new species of the genus Sinotilla Lelej, 1995 (Hymenoptera: Mutillidae), with notes on taxonomic characters. Zootaxa, 4294 (2), 151–169. https://doi.org/10.11646/zootaxa.4294.2.1
- Okayasu, J. (2018) Taxonomic review of Chin-wen Chen’s species described in the genus Smicromyrme (Hymenoptera: Mutillidae: Smicromyrmini). Acta Entomologica Musei Nationalis Pragae, 58 (2), 479–494. https://doi.org/10.2478/aemnp-2018-0036
- Okayasu, J. (2020) Velvet ants of the tribe Smicromyrmini Bischoff (Hymenoptera: Mutillidae) of Japan. Zootaxa, 4723 (1), 1–110. https://doi.org/10.11646/zootaxa.4723.1.1
- Okayasu, J. (2021a) A new velvet ant genus Arkaditilla (Hymenoptera, Mutillidae, Trogaspidiini) from the Oriental Region, with review of species. Journal of Hymenoptera Research, 84, 145–168. https://doi.org/10.3897/jhr.84.68709
- Okayasu, J. (2021b) Exploring the final piece of Japanese mutillid fauna in Amami-Ôshima Island, the newest World Heritage site. Hamuli, 12, 25–29.
- Okayasu, J. (2023a) Love is in the air and beyond the ocean: Taxonomic review of Neotrogaspidia Lelej (Hymenoptera: Mutillidae: Trogaspidiini) in Northeast Asia highlights its unique distributional pattern. Entomological Science, 26 (1), e12532. https://doi.org/10.1111/ens.12532
- Okayasu, J. (2023b) Discovery of the velvet ant genus Orientilla Lelej from Laos (Hymenoptera, Mutillidae, Dasylabrinae), with description of a related new species from India. Journal of Hymenoptera Research, 96, 817–834. https://doi.org/10.3897/jhr.96.110590
- Okayasu, J. (2023c) Preliminary assessment of the velvet ant fauna in Nepal, with description of new species of the genera Strangulotilla Nonveiller and Zeugomutilla Chen (Hymenoptera: Mutillidae). Insecta matsumurana, New Series, 79, 1–27. [http://hdl.handle.net/2115/90642]
- Okayasu, J., Lelej, A.S. & Williams, K.A. (2021a) Review of Eotrogaspidia Lelej (Hymenoptera: Mutillidae: Trogaspidiini). Zootaxa, 4920 (1), 56–90. https://doi.org/10.11646/zootaxa.4920.1.2
- Okayasu, J., Williams, K.A. & Lelej, A.S. (2018) A remarkable new species of the genus Sinotilla Lelej (Hymenoptera: Mutillidae: Smicromyrmini) from Taiwan and an overview of color diversity in East Asian mutillid females. Zootaxa, 4446 (3), 301–324. https://doi.org/10.11646/zootaxa.4446.3.1
- Okayasu, J., Williams, K.A., Lelej, A.S. & Pham, T.H. (2021b) Review of female Andreimyrme Lelej (Hymenoptera: Mutillidae: Smicromyrmini). Zootaxa, 5061 (1), 1–38. https://doi.org/10.11646/zootaxa.5061.1.1
- Olivier, [A.G.] (1811) Mutille. Encyclopédie Methodique. Histoire Naturelle. Insectes, 8, 49–66.
- Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, G.V.N., Underwood, E.C., D’Amico, J.A., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt, T.F., Lamoreux, J.F., Ricketts, T.H., Itoua, I., Wettengel, W.W., Kura, Y., Hedao, P. & Kassem, K. (2001) Terrestrial ecoregions of the world: A new map of life on Earth. BioScience, 51 (11), 933–938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
- O’Toole, C. (1975) The systematics of Timulla oculata (Fabricius) (Hymenoptera, Mutillidae). Zoologica Scripta, 4 (1), 229–251. https://doi.org/10.1111/j.1463-6409.1975.tb00733.x
- Pagden, H.T. (1934a) New species of Mutillidae (Hymenoptera, Vespoidea) from the Malay Peninsula. Journal of the Federated Malay States Museums, 17, 419–457.
- Pagden, H.T. (1934b) Biological notes on some Malayan aculeate Hymenoptera I. (Sphecoidea and Vespoidea). Journal of the Federated Malay States Museums, 17 (3), 458–466.
- Pagden, H.T. (1949) Descriptions and records of Austro-Malaysian Methocidae and Mutillidae (Hymenoptera). Transactions of the Royal Entomological Society of London, 100 (8), 191–231. https://doi.org/10.1111/j.1365-2311.1949.tb01431.x
- Pagliano, G. (2011) I Mutillidae (Hymenoptera) della collezione Zavattari le collezioni del Museo Regionale di Scienze Naturali di Torino. Bolletino del Musei Regionale di Scienze Naturali di Torino, 28 (1), 37–60.
- Pagliano, G., Brothers, D.J., Cambra, R., Lelej, A.S., Lo Cascio, P., Matteini Palmerini, M., Scaramozzino, P.L., Williams, K.A. & Romano, M. (2020) Checklist of names in Mutillidae (Hymenoptera), with illustrations of selected species. Bollettino del Museo Regionale di Scienze Naturali di Torino, 36 (1–2), 5–425.
- Pagliano, G. & Scaramozzino, P.L. (2021) La collezione di Mutillidae e di Bradynobaenidae (Hymenoptera) di Guido Pagliano. Bollettino del Museo Regionale di Scienze Naturali di Torino, 38 (1–2), 63–147.
- Pallas, P.S. (1773) Reisen durch verschiedene Provinzen des Russischen Reichs. Zweiter Theil. Zweites Buch vom Jahr 1771. Kayserlichen Academie der Wissenschaften, St. Petersburg, [12] + 744 pp.
- Parejo-Pulido, D. & Romano, M. (2022) A new species of Krombeinella from Turkey (Hymenoptera: Mutillidae). Boletín de la Sociedad Entomológica Aragonesa, 70, 75–78.
- Pate, V.S.L. (1947 [1946]) A minute on Paramyrmosa Saussure (Hymenoptera, Tiphiidae). The Canadian Entomologist, 78 (11–12), 196–197. https://doi.org/10.4039/Ent78196-11
- Peel, M.C., Finlayson, B.L. & McMahon, T.A. (2007) Updated world map of the Köppen-Geiger climate classification. Hydrology and Earth System Sciences, 11 (5), 1633–1644. https://doi.org/10.5194/hess-11-1633-2007
- Pennington, R.T., Lehmann, C.E.R. & Rowland, L.M. (2018) Tropical savannas and dry forests. Current Biology, 28 (9), R541–R545. https://doi.org/10.1016/j.cub.2018.03.014
- Péringuey, L. (1898) Description of some new or little known South African Mutillidae in the collection of the South African Museum. Annals of the South African Museum, 1 (1), 33–94. https://doi.org/10.5962/bhl.part.13904
- Pilgrim, E.M., von Dohlen, C.D. & Pitts, J.P. (2008) Molecular phylogenetics of Vespoidea indicate paraphyly of the superfamily and novel relationships of its component families and subfamilies. Zoologica Scripta, 37 (5), 539–560. https://doi.org/10.1111/j.1463-6409.2008.00340.x
- Pitts, J.P., Wilson, J.S. & von Dohlen, C.D. (2010) Evolution of the nocturnal Nearctic Sphaeropthalminae velvet ants (Hymenoptera: Mutillidae) driven by Neogene orogeny and Pleistocene glaciation. Molecular Phylogenetics and Evolution, 56 (1), 134–145. https://doi.org/10.1016/j.ympev.2010.03.033
- Polidori, C., Mendiola, P., Asís, J.D., Tormos, J. & Selfa, J. (2010) Temporal asynchrony and spatial co-occurrence with the host: The foraging patterns of Nemka viduata, a parasitoid of digger wasps (Hymenoptera: Mutillidae and Crabronidae). Journal of Ethology, 28, 353–361. https://doi.org/10.1007/s10164-009-0196-2
- Radoszkowski, O. (1885) Revision des armures copulatrices des mâles de la famille de Mutillides. Horae Societatis Entomologicae Rossicae, 19 (1–2), 3–49 + 9 pls.
- Reid, J.A. (1941) The thorax of the wingless and short-winged Hymenoptera. Transaction of the Royal Entomological Society of London, 91 (8), 367–446. https://doi.org/10.1111/j.1365-2311.1941.tb01046.x
- Roll, U., Feldman, A., Novosolov, M., Allison, A., Bauer, A.M., Bernard, R., Böhm, M., Castro-Herrera, F., Chirio, L., Collen, B., Colli, G.R., Dabool, L., Das, I., Doan, T.M., Grismer, L.L., Hoogmoed, M., Itescu, Y., Kraus, F., LeBreton, M., Lewin, A., Martins, M., Maza, E., Meirte, D., Nagy, Z.T., Nogueira, C. de C., Pauwels, O.S.G., Pincheira-Donoso, D., Powney, G.D., Sindaco, R., Tallowin, O.J.S., Torres-Carvajal, O., Trape, J.-F., Vidan, E., Uetz, P., Wagner, P., Wang, Y., Orme, C.D.L., Grenyer, R. & Meiri, S. (2017) The global distribution of tetrapods reveals a need for targeted reptile conservation. Nature Ecology & Evolution, 1, 1677–1682. https://doi.org/10.1038/s41559-017-0332-2
- de Saussure, H. (1867) Reise der österreichschen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859 unter den Befehlen des Commodore B. von Wüllerstorf-Urbair. Zoologischer Theil. Zweiter Band. Hymenoptera. Familien der Vespiden, Sphegiden, Pompiliden, Crabroniden und Heterogynen. C. Gerold’s Sohn, Wien, 138 pp. + 4 pls.
- de Saussure, H. (1880) Scolidae. In: Puteshestvie v Turkestan A.P. Fedtshenko [Voyage au Turkestan d’Alexis Fedtschenko]. Series 16. 2(5). Sankt-Petersburg, pp. 1–46, pls. 1–2. [in Russian]
- Sabi, K., Nozaka, C., Kurokawa, H. & Tano, T. (1977) [Insect collecting trip to Taiwan]. Hymenopterists’ Communication, 5, 34–42. [in Japanese]
- Say, T. (1836) Descriptions of new species of North American Hymenoptera, and observation on some already described. Boston Journal of Natural History, 1 (3), 209–305.
- Schröder, J.M., Rodríguez, L.P.Á. & Günter, S. (2021) Research trends: Tropical dry forests: The neglected research agenda? Forest Policy and Economics, 122, 102333. https://doi.org/10.1016/j.forpol.2020.102333
- Schuster, R.M. (1949) Contribution towards a monograph of the Mutillidae of the Neotropical region. III. A key to the subfamilies represented and descriptions of several new genera (Hymenoptera). Entomologica Americana, New Series, 29 (3–4), 59–140.
- Shorthouse, D.P. (2010) SimpleMappr, an online tool to produce publication-quality point maps. Available from: http://www.simplemappr.net (accessed 3 March 2025)
- Sichel, J. & Radoszkowski, O. (1869–1870) Essai d’une monographie des Mutilles de l’Ancion Continent. Horae Societatis Entomologicae Rossicae, 6 (3 & 4), 139–172 (1869) & 173–309, tabs. 6–11 (1870).
- Skorikov, A.S. (1935) Zur Mutilliden-Fauna Zentralasiens. Trudy tadzhikskoi basy Akademii Nauk SSSR, 5, 257–349, 7 pls. [in Russian]
- Smith, F. (1855) Catalogue of hymenopterous insects in the collection of the British Museum. Part III. Mutillidae and Pompilidae. Taylor and Francis, London, 206 pp., 5 pls. https://doi.org/10.5962/bhl.title.20858
- Smith, F. (1857–1858) Catalogue of the hymenopterous insects collected at Sarawak, Borneo; Mount Ophir, Malacca; and at Singapore, by A.R. Wallace. Journal of the Proceedings of the Linnean Society, Zoology, 2, 42–88, pls. 1–2 (1857), 89–130 (1858). https://doi.org/10.1111/j.1096-3642.1857.tb01759.x https://doi.org/10.1111/j.1096-3642.1858.tb02548.x
- Smith, F. (1879) Descriptions of new species of Hymenoptera in the collection of the British Museum. Taylor and Francis, London, XXI + 240 pp. https://doi.org/10.5962/bhl.title.60089
- Sodhi, N.S., Koh, L.P., Clements, R., Wanger, T.C., Hill, J.K., Hamer, K.C., Clough, Y., Tscharntke, T., Posa, M.R.C. & Lee, T.M. (2010) Conserving Southeast Asian forest biodiversity in human-modified landscape. Biological Conservation, 143 (10), 2375–2384. https://doi.org/10.1016/j.biocon.2009.12.029
- Starr, C.K. (2004) Nesting biology of the solitary wasp Pison argentatum (Hymenoptera: Sphecidae) in Borneo and the Philippines. Journal of the Kansas Entomological Society, 77 (4), 565–572. https://doi.org/10.2317/E32.1
- Sterling, E.J. & Hurley, M.M. (2005) Conserving biodiversity in Vietnam: Applying biogeography to conservation research. Proceedings of the California Academy of Sciences, 56 (Supplement 1), 98–118.
- Suárez, F.J. (1963) Diagnosis previas para una revision de las especies mediterráneas del genero Myrmosa Latreille (Hymenoptera, Myrmosidae). Bollettino della Società Entomologica Italiana, 93 (7–8), 113–117.
- Suárez, F.J. & Nonveiller, G. (1988) Mirmosidos de la Peninsula Iberica (Hymenoptera, Myrmosidae). Graellsia, 44, 81–158.
- Sutanto, D.D., Williams, K.A., Nugroho, H. & Lelej, A.S. (2021) To the knowledge of the velvet ant genus Bischoffitilla Lelej (Hymenoptera: Mutillidae) in Indonesia. Treubia, 48 (1), 1–12. https://doi.org/10.14203/treubia.v48i1.4011
- Suzuki, M. (1934) A list of Japanese Mutillidae. Kansai Konchu Zasshi, 2 (1), 23–25. [in Japanese]
- Terayama, M., Suda, H., Murota, T., Haneda, Y. & Tano, T. (2011) The velvet ants (Mutillidae) of Japan. Gekkan-Mushi, 481, 12–25. [in Japanese]
- Terayama, M., Suda, H., Murota, T. & Tano, T. (2016) Family Mutillidae. In: Terayama, M. & Suda, H. (Eds), A guide to the aculeate wasps of Japan. Tokai University Press, Hiratsuka, pp. 161–174. [in Japanese]
- Terine, J.B., Das, D. & Kumar, G.P. (2024) Occurrence of Andreimyrme substriolata (Chen, 1957) and Odontomutilla fletcheri Lelej, Terine & Girish Kumar, 2020 (Hymenoptera, Mutillidae) in India, with note on their geographical affinities. Journal of Insect Biodiversity and Systematics, 10 (1), 81–89. https://doi.org/10.61186/jibs.10.1.81
- Terine, J.B., Lelej, A.S. & Kumar, G.P. (2021) Discovery of the genus Andreimyrme Lelej, 1995 (Hymenoptera: Mutillidae) in India and description of a new species from the Western Ghats. Zootaxa, 5020 (1), 184–190. https://doi.org/10.11646/zootaxa.5020.1.10
- Thaochan, N., Williams, K.A., Thoawan, K., Jeenthong, T. & Sittichaya, W. (2022) Three new species and one new country record of velvet ants (Hymenoptera, Mutillidae) from Thailand. Journal of Hymenoptera Research, 93, 151–165. https://doi.org/10.3897/jhr.93.94727
- Thomson, C.G. (1870) Öfversigt af Sveriges rofsteklar. Opuscula Entomologica, Lund, 1 (2), 202–251.
- Tonnoir, A.L. (1930) Notes on Indian blepharocerid larvae and pupae with remarks on the morphology of Blepharocerid larvae and pupae in general. Records of the Indian Museum, 32, 161–214. https://doi.org/10.26515/rzsi/v32/i2/1930/162531
- Tordoff, A.W., Baltzer, M.C., Fellowes, J.R., Pilgrim, J.D. & Langhammer, P.F. (2012) Key biodiversity areas in the Indo-Burma hotspot: Process, progress and future directions. Journal of Threatened Taxa, 4 (8), 2779–2787. https://doi.org/10.11609/JoTT.o3000.2779-87
- Tsuneki, K. (1972a) Studies on the Mutillidae of Japan (Hymenoptera). Etizenia, 61, 1–26.
- Tsuneki, K. (1972b) Mutillidae collected in Formosa in 1966 and 68 (Hymenoptera). Etizenia, 64, 1–25.
- Tsuneki, K. (1993) On some Taiwanese Mutillidae, collected in 1976 by Mr. T. Murota, with description of new taxa (Hymenoptera). Special Publications of the Japan Hymenopterists Association, 41, 39–50.
- Tsuneki, K., Nozaka, C., Tano, T., Kurokawa, H. & Murota, T. (1993) Mutillidae recently collected in the Philippines (Hymenoptera). Special Publications of the Japan Hymenopterists Association, 41, 1–38.
- Tu, B.-B., Lelej, A.S. & Chen, X.-X. (2014) Review of the genus Cystomutilla André, 1896 (Hymenoptera: Mutillidae: Sphaeropthalminae: Sphaeropthalmini), with description of the new genus Hemutilla gen. nov. and four new species from China. Zootaxa, 3889 (1), 71–91. https://doi.org/10.11646/zootaxa.3889.1.4
- Waldren, G.C., Sadler, E.A., Murray, E.A., Bossert, S., Danforth, B.N. & Pitts, J. P. (2023) Phylogenomic inference of the higher classification of velvet ants (Hymenoptera: Mutillidae). Systematic Entomology, 48 (3), 463–487. https://doi.org/10.1111/syen.12588
- Wesmael, M. (1851) Revue critique des Hyménoptères fouisseurs de Belgique. Bulletin de l’Académie Royale des Sciences, des Lettres et des Beaux-Arts de Belgique, 18 (2), 362–384 + 451–493.
- Wikramanayake, E., Dinerstein, E., Loucks, C.J., Olson, D.M., Morrison, J., Lamoreux, J., McKnight, M. & Hedao, P. (2002) Terrestrial ecoregions of the Indo-Pacific: A conservation assessment. Island Press, Washington DC, xxix + 643 pp.
- Williams, K.A., Cambra, R.A., Bartholomay, P.R., Luz, D.R., Quintero A., D. & Pitts, J.P. (2019a) Review of the genus Lomachaeta Mickel, 1936 (Hymenoptera: Mutillidae) with new species and sex associations. Zootaxa, 4564 (1), 101–136. https://doi.org/10.11646/zootaxa.4564.1.4
- Williams, K.A., Lelej, A.S., Okayasu, J., Borkent, C.J., Malee, R., Thoawan, K. & Thaochan, N. (2019b) The female velvet ants (aka modkhong) of southern Thailand (Hymenoptera: Mutillidae), with a key to the genera of southeast Asia. Zootaxa, 4602 (1), 1–69. https://doi.org/10.11646/zootaxa.4602.1.1
- Williams, K.A., Lelej, A.S. & Thaochan, N. (2019c) New species of Myrmosinae (Hymenoptera: Mutillidae) from Southeastern Asia. Zootaxa, 4656 (3), 525–534. https://doi.org/10.11646/zootaxa.4656.3.9
- Williams, K.A., Pan, A.D. & Wilson, J.S. (2024) Velvet ants of North America. Princeton University Press, Princeton and Oxford, 440 pp. https://doi.org/10.2307/jj.5104027
- Wilson, J.S., Clark, S.L., Williams, K.A. & Pitts, J.P. (2012a) Historical biogeography of the arid-adapted velvet ant Sphaeropthalma arota (Hymenoptera: Mutillidae) reveals cryptic species. Journal of Biogeography, 39 (2), 336–352. https://doi.org/10.1111/j.1365-2699.2011.02580.x
- Wilson, J.S. & Pitts, J.P. (2011) Pleistocene connection between the Nearctic Mediterranean and desert regions in the Sphaeropthalma unicolor species-complex (Hymenoptera: Mutillidae). Insect Conservation and Diversity, 4 (3), 222–234. https://doi.org/10.1111/j.1752-4598.2010.00124.x
- Wilson, J.S. & Pitts, J.P. (2012) Identifying Pleistocene refugia in North American cold deserts using phylogeographic analyses and ecological niche modelling. Diversity and Distributions, 18 (11), 1139–1152. https://doi.org/10.1111/j.1472-4642.2012.00902.x
- Wilson, J.S., Williams, K.A., Forister, M.L., von Dohlen, C.D. & Pitts, J.P. (2012b) Repeated evolution in overlapping mimicry rings among North American velvet ants. Nature Communications, 3, 1272. https://doi.org/10.1038/ncomms2275
- Wilson, J.S., Jahner, J.P., Forister, M.L., Sheehan, E.S., Williams, K.A. & Pitts, J.P. (2015) North American velvet ants form one of the world’s largest known Müllerian mimicry complexes. Current Biology, 25, R704–R706. https://doi.org/10.1016/j.cub.2015.06.053
- Wilson, J.S., Pan, A.D., Limb, E.S. & Williams, K.A. (2018) Comparison of African and North American velvet ant mimicry complexes: Another example of Africa as the ‘odd man out’. PLoS ONE, 13 (1), e0189482. https://doi.org/10.1371/journal.pone.0189482
- Wu, C.F. (1941) Catalogus Insectorum Sinensium. Volume VI. The Department of Biology, Yenching University, Peiping, vi + 333 + xv pp
- Yasumatsu, K. (1938a) Les Mutillides de Coree et de la Mandschourie du Sud. Transactions of the Natural History Society of Formosa, 28 (175), 83–87.
- Yasumatsu, K. (1938b) Hyménoptères trouvées dans les environs de Nanking et de Changsha du bassin Yang-tse (Chine) par M. Kwo-Mo Ho. Transactions of the Natural History Society of Formosa, 28 (180–181), 345–347. [in Japanese]
- Yasumatsu, K. (1948) Records of East Chinese Mutillidae (Hymenoptera). Lingnan Science Journal, 22 (1–4), 71–73.
- Yasumatsu, K. (1951) Mutillidae of Shansi, North China (Hymenoptera). Mushi, 22 (12), 69–72.
- Yano, M. (1932) Mutillidae. In: Uchida, S. (Ed.), Iconographia Insectorum Japonicorum Editio Prima. Hokuryukan, Tokyo, pp. 325–327, 1 pl. [in Japanese]
- Zavattari, E. (1910) Catalogo delle Mutille del Museo Zoologico di Napoli. Annuario del Museo Zoologico dell’Università di Napoli, New Series, 3 (9), 1–16.
- Zavattari, E. (1913) H. Sauter’s Formosa-Ausbeute. Mutillidae (Hym.). Archiv für Naturgeschichte, 79A (3), 19–42. https://doi.org/10.5962/bhl.part.7709
- Zavattari, E. (1914) Mutille Austro-Malesi. Bolletino della Societa Entomologica Italiana, 45, 61–114.
- Zhou, H.-T., Lelej, A.S. & Liu, J.-X. (2018a) New records of velvet ants (Hymenoptera: Mutillidae) from China with taxonomic notes. Far Eastern Entomologist, 354, 23–28. https://doi.org/10.25221/fee.354.4
- Zhou, H.-t., Lelej, A.S., Williams, K.A. & Liu, J.-X. (2018b) Revision of the Oriental genus Zavatilla Tsuneki (Hymenoptera, Mutillidae, Trogaspidiini), with descriptions of two new species. Zootaxa, 4418 (2), 101–120. https://doi.org/10.11646/zootaxa.4418.2.1
- Zhou, H.-T., Lelej, A.S. & Xu, Z.-F. (2017) Discovery of the subfamily Rhopalomutillinae (Hymenoptera: Mutillidae) in China and description of Pherotilla brothersi sp. nov. Zootaxa, 4273 (4), 587–594. https://doi.org/10.11646/zootaxa.4273.4.8