Abstract
Velvet ants (Hymenoptera: Mutillidae and Myrmosidae) are recognized as model organisms for studies in Müllerian mimicry and most velvet ant females exhibit aposematic color patterns. Mimicry rings, however, are inadequate for comparing coloration across different regions and among cryptically colored insects. Based on photographs of over 2800 velvet ant females, new terminology is developed for color syndromes. This can provide a starting point for future mimicry studies in various parts of the World and will simplify discussions of color similarities across geographical and phylogenetic space. Finally, this paper will be a useful reference for velvet ant identification, as dorsal habitus photographs are provided of expertly identified velvet ants representing more than half of the World’s species known from females.
Twenty-one new combinations are proposed: Darditilla albovittata (André, 1898), comb. nov. (from Mutilla Linnaeus, 1758); Darditilla curiosa (Cresson, 1902), comb. nov. (from Mutilla); Pseudomethoca argyrospila (Gerstaecker, 1874), comb. nov. (from Mutilla); Pseudomethoca bilineata (Klug, 1821), comb. nov. (from Mutilla); Pseudomethoca blandina (Smith, 1879), comb. nov. (from Mutilla); Pseudomethoca boliviana (André, 1906), comb. nov. (from Ephuta Say, 1836); Pseudomethoca confinis (Gerstaecker, 1874), comb. nov. (from Mutilla); Pseudomethoca extinctor (Cameron, 1894), comb. nov. (from Sphaerophthalma[!] Blake, 1871); Pseudomethoca hilaris (Gerstaecker, 1874), comb. nov. (from Mutilla); Pseudomethoca imbecilla (Gerstaecker, 1874), comb. nov. (from Mutilla); Pseudomethoca lycimnia (Cameron, 1894), comb. nov. (from Sphaerophthalma[!]); Pseudomethoca macropis (Gerstaecker, 1874), comb. nov. (from Mutilla); Pseudomethoca minutoria (Cameron, 1895), comb. nov. (from Sphaerophthalma[!]); Pseudomethoca orithyia (Cameron, 1894), comb. nov. (from Sphaerophthalma[!]); Pseudomethoca paulula (Cresson, 1902), comb. nov. (from Mutilla); Pseudomethoca polyspila (Gerstaecker, 1874), comb. nov. (from Mutilla); Pseudomethoca pumila (Burmeister, 1854), comb. nov. (from Mutilla); Pseudomethoca tantula (Cresson, 1902), comb. nov. (from Mutilla); Pseudomethoca turnalis (Cresson, 1902), comb. nov. (from Mutilla); Pseudomethoca vulnerithorax (André, 1906), comb. nov. (from Ephuta); and Xystromutilla leucogramma (Gerstaecker, 1874), comb. nov. (from Mutilla).
References
- André, E. (1905) Nouvelles espèces de Mutillides d’Amérique. (Hym.). Zeitschrift für Systematische Hymenopterologie und Dipterologie, 5 (1–6), 361–376.
- Bartholomay, P.R., Williams, K.A., Luz, D.R. & Morato, E.F. (2015) Frigitilla gen. nov., a new genus of Amazonian Mutillidae (Hymenoptera). Zootaxa, 3957 (1), 49–58. https://doi.org/10.11646/zootaxa.4433.2.7
- Bartholomay, P.R., Williams, K.A., Luz, D.R. & Oliveira, M.L. (2018) New species of Traumatomutilla André in the T. tabapua and T. integella species‐groups (Hymenoptera, Mutillidae). Zootaxa, 4433 (2), 361–385. https://doi.org/10.11646/zootaxa.4433.2.7
- Bartholomay, P.R., Williams, K.A., Cambra, R.A. & Oliveira, M.L. (2019) Does the genus Dasymutilla Ashmead occur in South America? The new genus Quwitilla, new combinations, and new distribution records from Neotropical velvet ants (Hymenoptera: Mutillidae). Zootaxa, 4623 (2), 261–282. https://doi.org/10.11646/zootaxa.4623.2.3
- Bartholomay, P.R., Williams, K.A., Cambra, R.A. & Oliveira, M.L. (2020) Revision of the Traumatomutilla juvenilis species‐group (Hymenoptera, Mutillidae). Journal of Natural History, 53 (43–44), 2639–2683. https://doi.org/10.1080/00222933.2020.1715501
- Bartholomay, P.R., Williams, K.A., Cambra, R.A. & Oliveira, M.L. (2021) Revision of the Traumatomutilla gemella species‐group (Hymenoptera, Mutillidae) with the description of its hitherto unknown males. Zoosystema, 43 (1), 1–28. https://doi.org/10.5252/zoosystema2021v43a1
- Bartholomay, P.R., Williams, K.A., Cambra, R.A. & Oliveira, M.L. (2022) Revision of the Traumatomutilla indica species‐group (Hymenoptera, Mutillidae). Zootaxa, 5108 (1), 1–97. https://doi.org/10.11646/zootaxa.5108.1.1
- Bartholomay, P.R., Cambra, R.A., Ramos, C.W., Wilkie, E. & Añino, Y.J. (2024) Kevinilla, a new velvet ant genus in the Sphaeropthalminae (Hymenoptera: Mutillidae). European Journal of Entomology, 121, 182–196. https://doi.org/10.14411/eje.2024.019
- Bartholomay, P.R., Williams, K.A., Cambra, R.A., Lopez, V.M. & Oliveira, M.L. (2025) Revision of the Traumatomutilla quadrinotata species group (Hymenoptera, Mutillidae, Sphaeropthalminae): new synonyms, sex associations, and a new species from the Brazilian Atlantic Forest. European Journal of Taxonomy, 995 (1), 1–75. https://doi.org/10.5852/ejt.2025.995.2913
- Bergamaschi, A.C., Cambra, R.A., Brothers, D.J. & Melo, G.A. (2012) Lynchiatilla Casal, 1963 (Hymenoptera: Mutillidae): a new species from Brazil associated with Paroxystoglossa spiloptera Moure (Hymenoptera: Apidae: Halictinae), and notes on other species. Zootaxa, 3548 (1), 55–64. https://doi.org/10.11646/zootaxa.3548.1.3
- Bingham, C.T. (1897) The fauna of British India including Ceylon and Burma: Hymenoptera, I. Wasps and Bees. Taylor and Francis, London, 608 pp. https://doi.org/10.5962/bhl.title.100738
- Bischoff, H. (1920–1921) Monographie der Mutilliden Afrikas. Archiv für Naturgeschichte, 86A (1–3 & 4–5), 1–480 (1920), 1 map & 481–830 (1921).
- Bogusch, P. (2006) The velvet ants (Hymenoptera: Mutillidae) of the Czech Republic and Slovakia: an identification key and annotated checklist. Acta Musei Moraviae, Scientiae biologicae, 91, 103–148.
- Boutin, M. & Vilhelmsen, L. (2024) 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
- Brabant, C.M. (2015) Taxonomic Revision and Cladistic Analysis of the Neotropical genus Tallium André,1902 (Hymenoptera: Mutillidae: Sphaeropthalminae). University of Wisconsin-Madison, Madison, Wisconsin, 362 pp.
- Bradley, J.C. & Bequert, J. (1928) A synopsis of the Mutillidae of the Belgian Congo. Bulletin of the American Museum of Natural History, 58 (2), 63–122.
- Brothers, D.J. (1971) Ascetotilla, a new genus of Mutillidae from New Guinea (Hymenoptera). Pacific Insects, 13 (3–4), 471–485.
- Brothers, D.J. (1994) A new genus and four new species of Mutillidae associated with Brachyponera lutea Mayr (Formicidae) in Western Australia (Hymenoptera). Journal of the Australian Entomological Society, 33, 143–152. https://doi.org/10.1111/j.1440-6055.1994.tb00941.x
- Brothers, D.J. (2006) Capítulo 14.4. Familia Mutillidae. In: Hanson, P.E. & Gauld, I.D. (Eds.), Hymenoptera de la Región Neotropical. The American Entomological Institute, Gainesville, Florida, pp. 586–593.
- Brothers, D.J. (2012) The new genus Ancistrotilla n. gen., with new species from Vanuatu and New Caledonia (Hymenoptera, Mutillidae). Zoosystema, 34 (2), 223–251. https://doi.org/10.5252/z2012n2a2
- 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. & 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
- Brothers, D.J., Lelej, A.S. & Yanega, D. (2025) Re‐evaluation of the grammatical genders of the names “Smicromyrme” and its derivatives, and “Ephutomma” (Hymenoptera: Mutillidae). Zootaxa, 5665, 277–280. https://doi.org/10.11646/zootaxa.5665.2.8
- Cambra, R.A. (1996) Sistemática y Biología del Género Lophomutilla Mickel (Hymenoptera: Mutillidae). M.Sc. Thesis, Universidad de Panama, Panama City, 99 pp.
- Cambra, R.A. & Quintero, A.D. (1996) The Mexican and Central American Species of Lophostigma Mickel, Including Records, and Taxonomic Notes for the Genus (Hymenoptera: Mutillidae). Pan-Pacific Entomologist, 72 (2), 92.
- Cambra, R.A. & Quintero, A.D. (1997) A revision of Protophotopsis Schuster (Hymenoptera: Mutillidae). Journal of Hymenoptera Research, 6 (2), 263–272.
- Cambra, R.A. & Quintero, A.D. (2008) New species and new distribution records for Pseudomethoca Ashmead (Hymenoptera: Mutillidae) from Central America. Transactions of the American Entomological Society, 134 (1), 69–86. https://doi.org/10.3157/0002-8320(2008)134[69:NSANDR]2.0.CO;2
- Cambra, R.A. & Quintero, A.D. (2010) Review of the Neotropical genus Nanotopsis Schuster (Hymenoptera: Mutillidae: Sphaeropthalminae). Transactions of the American Entomological Society, 136 (3 + 4), 289–298. https://doi.org/10.3157/061.136.0312
- Cambra, R.A., Brothers, D.J. & Quintero, D. (2016) Review of Frigitilla Williams in Bartholomay et al., 2015, with description of a new species from Panama (Hymenoptera: Mutillidae). Zootaxa, 4189 (2), 395–400. https://doi.org/10.11646/zootaxa.4189.2.13
- Cambra, R.A., Quintero, D. & Pagliano, G. (2017) Three new species of Ephutini (Hymenoptera: Mutillidae) from Venezuela and Ecuador, including taxonomic changes for seven Neotropical species of Ephuta. Il Naturalista Valtellinse—Atti Museo Civico di Storia Naturale di Morbegno, 28, 47–64.
- Cambra, R.A., Waldren, G.C., Bartholomay, P.R., Williams, K.A. & Quintero, D. (2018) Taxonomic revision of the genus Timulla Ashmead, 1899 of Panama (Hymenoptera: Mutillidae). Tecnociencia, 20 (2), 23–57.
- Cambra, R.A., Brothers, D.J. & Quintero, A.D. (2020) Review of Calomutilla Mickel, 1952, a new species, and comparison with Pertyella Mickel, 1952 (Hymenoptera: Mutillidae). Zootaxa, 4789 (2), 466–480. https://doi.org/10.11646/zootaxa.4789.2.6
- Cambra, R.A., Williams, K.A. & Bartholomay, P.R. (2022) Two new species of Horcomutilla Casal, 1962 (Hymenoptera: Mutillidae), description of hitherto unknown males, and an illustrated key to all known species in the genus. Neotropical Entomology, 51 (3), 427–446. https://doi.org/10.1007/s13744-022-00957-8
- Cambra, R.A., Williams, K.A., Ramos, R.L., Bartholomay, P.R., Trindade-Santos, M.E. & de Oliveira, F.F. (2024) Taxonomic notes on Pertyella Mickel, 1952 (Hymenoptera; Mutillidae): Morphological characters rediscussion, new species, new combinations, new records, and species grouping. Revista peruana de biología, 31 (2), 1–10. https://doi.org/10.15381/rpb.v31i2.26220
- Cameron, P. (1894–1900) Fam. Mutillidae. In: Godman, F.D. & Salvin, O. (Eds.), Biologia Centrali-Americana. Insecta. Hymenoptera. (Fossores.) II. R.H. Porter, London, pp. 259–328 (1894) + 329–376 (1895) + 377–395 (1896) + 404 (1899) + iii + v + x–xi + 405–413 (1900), pls. 13–14.
- Casal, O.H. (1962) Mutillidae neotropicales XI. (Hymenoptera). Vianatilla, nuevo género de Sphaeropthalminae. Neotropica, 8, 77–78.
- Casal, O.H. (1963) Mutillidae neotropicales XIX. (Hymenoptera). Las especies del nuevas género Lynchiatilla. Acta Zoologica Lilloana, 19, 343–383.
- Casal, O.H. (1968a) Aportaciones para el conocimiento de las Mutillidae de la República Argentina. I. Las hembras del género Darditilla (Hymenoptera). Revista de la Sociedad Entomológica Argentina, 30, 83–96.
- Casal, O.H. (1968b) Comentarios sobre Reedomutilla Mickel, 1964 con la descripcion de Suareztilla gen. nov. (Hymenoptera, Mutillidae). Neotropica, 14 (44), 75–80.
- Casal, O.H. (1969a) Sobre Traumatomutilla André (Hymenoptera, Mutillidae). Physis, 28 (77), 279–298.
- Casal, O.H. (1969b) Las hembras de Ephutini de la Republica Argentina (Hymenoptera: Mutillidae). Revista de la Sociedad Entomológica Argentina, 31, 33–41.
- Casal, O.H. (1969c) Sobre Xystromutilla Andre (Hymenoptera: Mutillidae). Physis, 29 (78), 47–50.
- Casal, O.H. (1969d) Las hembras de Ephutini del Estado de Santa Catarina, Brazil. Revista de la Sociedad Entomológica Argentina, 31, 97–105.
- Casal, O.H. (1970) Las hembras de Ephutini del Estado de Rio de Janeiro, Brazil. Physis, 29 (79), 297–310.
- Chen, C. (1957) A revision of the velvety ants or Mutillidae of China. Quarterly Journal of the Taiwan Museum, 10 (3–4), 135–224, 6 pls.
- Fritz, M.A. (1990) Notas taxonómicas sobre los géneros Lophomitulla y Pertyella (Hymenoptera: Mutillidae). Revista de la Sociedad Entomológica Argentina, 48 (1–4), 129–142.
- Fritz, M.A. & Pagliano, G. (1993) Sobre Lophomutilla Mickel y Pertyella Mickel (Hymenoptera Multillidae [sic!]). Bollettino della Società Entomologica Italiana, Genova, 124 (3), 209–220.
- Garcia, E.Q., Cambra, R. & Melo, G.A.R. (2006) Sexual associations for two species of mutillid wasps (Hymenoptera, Mutillidae), with the description of a new species of Anomophotopsis. Revista Brasileira de Entomologia, 50 (3), 379–384. https://doi.org/10.1590/S0085-56262006000300007
- Gogala, A. (2017) Ose mravljarice Slovenije (Hymenoptera: Mutillidae). Velvet Ants of Slovenia (Hymenoptera: Mutillidae). Scopolia, 90, 1–39.
- Hammer, K. (1955) Contributions à l’étude de la faune entomologique du Ruanda-Urundi (Mission P. Basilewsky 1953). 76. Hymenoptera Mutillidae. Annales du Musée Royal du Congo Belge, Tervuren, (Série in 8º) Sciences Zoologiques, 40, 397–403.
- Klug, J.C.F. (1821) Entomologie brasilianae specimen. Nova Acta Academica Caesareae Leopoldino-Carolinae Germanicae Nature Curiosorum, 10 (2), 305–324.
- Krombein, K.V. (1940) Studies in the Tiphiidae (Hym. 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, 415–465 + pl. 24.
- Krombein, K.V. (1971) A monograph of the Mutillidae of New Guinea, Bismarck Archipelago and Solomon Islands. Part 1: Mutillinae (Hymenoptera Aculeata). In: Entomological Essays to Commemorate the Retirement of Professor K. Yasumatsu. Hokuryukan, Tokyo, pp. 25–60.
- Krombein, K.V. (1972) Monograph of the Madagascan Mutillidae (Hymenoptera). Part I: Myrmillini, Mutillini and Smicromyrmini. Annales, Musée Royal de l’Afrique Centrale, Série in 8°, Sciences Zoologiques, 199, i–x + 1–61.
- Krombein, K.V. (1979) Superfamily Scolioidea. In: Krombein, K.V., Hurd, P.D.Jr., Smith, D.R. & Burks, B.D. (Eds.), Catalog of Hymenoptera in America North of Mexico. Vol. 2. Apocrita (Aculeata). Smithsonian Institution Press, Washington, pp. 1253–1321.
- Lelej, A.S. (1976) Wasps of the genus Dasylabris Rad. (Hymenoptera, Mutillidae) of the fauna of the USSR. – Entomologicheskoe Obozrenie, 55 (1), 189–204, 19 figs. [in Russian, English translation (1979) in Entomological Review, 55, 136–146]
- Lelej, A.S. (1980) The genus Pseudophotopsis André, 1896 (Hymenoptera, Mutillidae) from the USSR and neighbouring countries. Entomologicheskoe Obozrenie, 59 (3), 634–649. [in Russian, English translation (1982) in Entomological Review, 59 (3), 112–126]
- 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. (1995) 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. (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) 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. (2020a) 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. (2020b) To the knowledge of velvet ants of the genera Artiotilla Invrea, Radoszkowskitilla Lelej and Taiwanomyrme Tsuneki (Hymenoptera: Mutillidae). Far Eastern Entomologist, 410, 1–10. https://doi.org/10.25221/fee.410.1
- 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. (2024a) 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. (2024b) Revision of the Palaearctic genus Skorikovia Ovtchinnikov, 2002 (Hymenoptera: Mutillidae). Zootaxa, 5529 (1), 52–72. https://doi.org/10.11646/zootaxa.5529.1.2
- Lelej, A.S. (2025) Revision of the genus Dentilla Lelej in Lelej et Kabakov, 1980 (Hymenoptera: Mutillidae). Zootaxa, 5633 (2), 288–315. https://doi.org/10.11646/zootaxa.5633.2.4
- 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. & van Harten, A. (2006) A review of the Mutillidae (Hymenoptera) of Yemen. Zootaxa, 1226 (1), 1–50. https://doi.org/10.11646/zootaxa.1226.1.1
- 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., 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., 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
- Lelej, A.S., Proshchalykin, M.Y. & Maharramov, M.M. (2022) A review of the Mutillidae (Hymenoptera) of Azerbaijan. Zootaxa, 5155 (1), 61–86. https://doi.org/10.11646/zootaxa.5155.1.3
- 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., 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
- Linnaeus, C. (1758) Systema Naturae per Regna Tria Naturae, secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. Editio Decima Reformata. Laurentius Salvius, Holmia [Stockholm], 823 pp. https://doi.org/10.5962/bhl.title.542
- Lo Cascio, P., Romano, M. & Grita, F. (2012) New species and new records of mutillid wasps from the Socotra Archipelago (Hymenoptera: Mutillidae). Acta Entomologica Musei Nationalis Pragae, 52 (Supplement 2), 525–544.
- Luz, D.R. (2015) Revisão taxonômica do gênero Hoplocrates Mickel (Hymenoptera, Mutillidae). Repositório Institucional da UFPR, 242 pp. Available from: http://hdl.handle.net/1884/49507 (accessed 27 May 2025)
- Manley, D.G. & Pitts, J.P. (2007) Tropical and subtropical velvet ants of the genus Dasymutilla Ashmead (Hymenoptera: Mutillidae) with descriptions of 45 new species. Zootaxa, 1487 (1), 1–128. https://doi.org/10.11646/zootaxa.1487.1.1
- Manley, D.G., Williams, K.A. & Pitts, J.P. (2020) Keys to Nearctic velvet ants of the genus Dasymutilla Ashmead (Hymenoptera: Mutillidae), with notes on taxonomic changes since Krombein (1979). Proceedings of the Entomological Society of Washington, 122 (2), 335–414. https://doi.org/10.4289/0013-8797.122.2.335
- Matias, R. (2024) Undescribed diversity in Iberian Mutillidae (Hymenoptera): a new species of Smicromyrme Thomson, 1870 from Portugal. Zootaxa, 5446 (3), 301–330. https://doi.org/10.11646/zootaxa.5446.3.1
- Mickel, C.E. (1933) 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. (1934) Mutillidae of the Philippine Islands (Hymenoptera). The Philippine Journal of Science, 54 (1), 91–219.
- 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. (1937) 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. (1938a) The Neotropical Mutillid Wasps of the Genus Dimorphomutilla (Hymenoptera). Revista de Entomologia, Rio de Janeiro, 9, 349–364.
- Mickel, C.E. (1938b) A synopsis of the neotropical mutillid genus Euspinolia Ashmead (Hymenoptera). Revista de Entomologia, Rio de Janeiro, 9, 53–74.
- Mickel, C.E. (1938c) 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. (1939a) A monograph of the Neotropical mutillid genus Hoplomutilla Ashmead (Hymenoptera: Mutillidae). Revista de Entomologia, Rio de Janeiro, 10 (2 & 3), 337–403 & 641–717.
- Mickel, C.E. (1939b) Monograph of a new neotropical mutillid genus, Pappognatha (Hymenoptera: Mutillidae). Annals of the Entomological Society of America, 32 (2), 329–343. https://doi.org/10.1093/aesa/32.2.329
- Mickel, C.E. (1941) Monograph of the South American Genus Hoplocrates Mickel (Hymenoptera: Mutillidae). Revista de Entomologia, Rio de Janeiro, 12, 341–414.
- Mickel, C.E. (1943) Monograph of the South American Genus Atillum André (Hymenoptera: Mutillidae). Revista de Entomologia, Rio de Janeiro, 14, 174–254.
- Mickel, C.E. (1952) The Mutillidae (wasps) of British Guiana. Zoologica, New York, 37 (3), 105–150. https://doi.org/10.5962/p.184691
- Mitchell, A. & Brothers, D.J. (1998) Revision and cladistic analysis of the Afrotropical genus Areotilla Bischoff (Hymenoptera: Mutillidae, Ticoplinae). African Entomology, 6 (2), 193–214.
- Mitchell, A. & Brothers, D.J. (2002) Phylogeny of the genera of Ticoplinae (Hymenoptera: Mutillidae). Journal of Hymenoptera Research, 11, 312–325.
- Muskovits, J. & Gyorgy, Z. (2011) Magyarország hangyadarazsai / Velvet ants of Hungary (Hymenoptera: Mutillidae). Kaposvár, Budapest, 198 pp. [Natura Somogyiensis, 18] https://doi.org/10.24394/NatSom.2011.18.2
- Nonveiller, G. (1973a) Recherches sur les Mutillides de l’Afrique (Hymenoptera, Mutillidae). I. Contribution à la connaisance du genre Trispilotilla Bischoff 1920 ♂♀. Annales de la Faculté des Sciences du Cameroun, 13, 77–134.
- Nonveiller, G. (1973b) Recherches sur les Mutillides de l’Afrique (Hymenoptera, Mutillidae). II. Carinotilla gen. nov. Annales de la Faculté des Sciences du Cameroun, 14, 47–102.
- Nonveiller, G. (1975) Recherches sur les Mutillides de l’Afrique (Hymenoptera, Mutillidae). VI. Les espèces actuellement connues du genre Clinotilla Arnold 1956 ♀♀. Annales de la Faculté des Sciences de Yaoundé, 19, 101–130.
- Nonveiller, G. (1977) Recherches sur les Mutillides de l’Afrique (Hymenoptera, Mutillidae). VII. Dentotilla gen. nov. Beiträge zur Entomologie, Berlin, 27 (2), 245–300.
- Nonveiller, G. (1979 [“1978”]) Recherches sur les Mutillides de l’Afrique (Hymenoptera, Mutillidae). VIII. Révision des genres Ctenotilla, Cephalotilla et Pseudocephalotilla sensu Bischoff. Memoires publies par l’Institut pour la Protection des Plantes, Beograd, 13, 1–184.
- Nonveiller, G. (1990) Catalogue of the Mutillidae, Myrmosidae and Bradynobaenidae of the Neotropical Region including Mexico (Insecta, Hymenoptera). Hymenopterorum Catalogus. Nova Editio. 18. SPB Academic Publishing, Den Haag, 150 pp.
- Nonveiller, G. (1994a) Description du nouveau genre Afrotropical Spinulomutilla et de onze espèces nouvelles (Hymenoptera: Mutillidae) (Première partie). Annales de la Societé Entomologique de France, New Series, 30 (3), 329–344. https://doi.org/10.1080/21686351.1994.12277711
- Nonveiller, G. (1994b) Description du nouveau genre Afrotropical Spinulomutilla et de onze espèces nouvelles (Hymenoptera: Mutillidae) (Deuxième partie). Annales de la Societé Entomologique de France, New Series, 30 (4), 367–389. https://doi.org/10.1080/21686351.1994.12277872
- Nonveiller, G. (1995) Recherches sur les Mutillides de l’Afrique XVII. Note pour servir à la connaissance du genre Pristomutilla Ashmead, 1903 (mâle, femelle) avec description du mâle encore inconnu du genre, d’espèce nouvelle du genre et des nouveaux sous-genres Diacanthotilla (femelles) et Acanthomutilla (femelles) (Hymenoptera, Mutillidae). Entomofauna, 16 (5), 29–119.
- Nonveiller, G. (1996a [“1995”]) Somaliatilla somalica n. gen. et n. sp., femelle (Hymenoptera, Mutillidae). Revue Française d’Entomologie, New Series, 17 (4), 125–128.
- Nonveiller, G. (1996d) Révision du genre Afrotropical Odontotilla Bischoff, 1920 (mâle et femelle) (Hymenoptera: Mutillidae). Annales de la Societé Entomologique de France, New Series, 32 (3), 299–357. https://doi.org/10.1080/21686351.1996.12278872
- Nonveiller, G. (1998 [“1997”]) Révision du genre Afrotropical Mimecomutilla Ashmead, 1903 (mâle et femelle) avec description d’espèces nouvelles et description du nouveau sous-genre Mimecotilla (Hymenoptera, Mutillidae). Annales de la Societé Entomologique de France, New Series, 33 (2), 447–485. https://doi.org/10.1080/21686351.1997.12279189
- Nonveiller, G. (1999) A revision of the genus Lobotilla Bischoff, 1920 (♂,♀) (Insecta: Hymenoptera: Mutillidae). Entomologische Abhandlungen Staatliches Museum für Tierkunde Dresden, 58 (14), 255–282.
- Nonveiller, G. & Ćetković, A. (1996) Seriatospidia n. gen. with a description of three new species (Hymenoptera Mutillidae). Tropical Zoology, 9, 305–314. https://doi.org/10.1080/03946975.1996.10539314
- Olsoufieff, G. (1938 [“1937”]) Revision systématique des Mutilles de Madagascar. Bulletin de l’Academie Malgache, New Series, 20, 171–217, pls. 27–28.
- 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. (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., 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., 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. & 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
- O’Toole, Ch. (1975) The Systematics of Timulla oculata (Fabricius) (Hymenoptera, Mutillidae). Zoologica Scripta, 4 (5–6), 229–251. https://doi.org/10.1111/j.1463-6409.1975.tb00733.x
- 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. & Matteini Palmerini, M. (2014) Contributo allo studio dei Mutillidi della Tunisia (Insecta Hymenoptera Mutillidae). Quaderno di studi e notizie di storia naturale della Romagna, 39, 175–209.
- Pagliano, G. & Strumia, F. (2007) Contributo alla conoscenza dei Mutillidae italiani (Hymenoptera, Scolioidea). Bollettino del Musei Regionale di Scienze Naturali di Torino, 24 (1), 25–110.
- Pan, A.D., Williams, K.A. & Wilson, J.S. (2017) Are diurnal iguanian lizards the evolutionary drivers of New World female velvet ant (Hymenoptera: Mutillidae) Müllerian mimicry rings? Biological Journal of the Linnean Society, 120, 436–447. https://doi.org/10.1111/bij.12894
- Parejo-Pulido, D. & Romano, M. (2022) A new species of Krombeinella from Turkey (Hymenoptera: Mutillidae). Boletín de la Sociedad Entomológica Aragonesa, 70, 75–78.
- Parejo-Pulido, D., Pérez-Gómez, Á., Obregón, R. & París, M. (2023) Catálogo de los mutílidos (Hymenoptera: Mutillidae) de la provincia de Cádiz (Andalucía, España). Revista de la Sociedad Gaditana de Historia Natural, 17, 1–28.
- Pitts, J.P. (2000) New species and a new record of Bordontilla (Hymenoptera: Mutillidae) from Chile. Entomological News, 111 (1), 32–38.
- Quintero, A.D. & Cambra, R.A. (2001) On the identity of Scaptopoda F. Lynch Arribálzaga, new taxonomic changes and new distribution records for Neotropical Mutillidae (Hymenoptera), with notes on their biology. Transactions of the American Entomological Society, 127 (3), 291–304.
- Sherratt, T.N. (2002) The evolution of imperfect mimicry. Behavioral Ecology, 13, 821–826. https://doi.org/10.1093/beheco/13.6.821
- Shorthouse, D.P. (2010) SimpleMappr, an online tool to produce publication-quality point maps. Available from: https://www.simplemappr.net (accessed 06 May 2025)
- Suárez, F.J. (1962) Datos sobre Mutílidos neotropicales. III. Un nuevo género de Centroamérica y paises noroccidentales de América del Sur (Hymenoptera). Eos, Revista Española de Entomología, Madrid, 38 (1), 115–146.
- Suárez, F.J. (1966 [“1965”]) Datos sobre Mutílidos neotropicales. VI. Dos nuevos géneros de Pseudomethocini de Méjico y Brasil (Hymenoptera). Eos, Revista Española de Entomología, Madrid, 41 (2–3), 471–482.
- Suárez, F.J. (1988) Mirmosidos de la Peninsula Iberica (Hymenoptera Myrmosidae). Graellsia, 44, 81–158.
- Terayama, M., Suda, H., Murota, T., Haneda, Y. & Tano, T. (2011) The velvet ants (Mutillidae) of Japan. Gekkan-Mushi, 481, 12–25. [in Japanese]
- Terine, J.B. & Kumar, G.P. (2024) Three new species of the genus Odontomutilla Ashmead, 1899 (Hymenoptera: Mutillidae) from India. Zootaxa, 5405 (1), 131–141. https://doi.org/10.11646/zootaxa.5405.1.6
- 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
- Tu, B.-B., Lelej, A.S. & Chen, X.-X. (2014a) 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
- Tu, B.-B., Lelej, A.S. & Chen, X.-X. (2014b) Five newly recorded species of Dasylabris (Hymenoptera: Mutillidae) from China. Entomotaxonomia, 34 (4), 300–310.
- Tu, B.-B., Lelej, A.S. & Chen, X.-X. (2015) Review of the genus Taiwanomyrme Tsuneki, 1993 (Hymenoptera, Mutillidae, Mutillinae), with description of two new species from China. Zootaxa, 4020 (3), 588–600. https://doi.org/10.11646/zootaxa.4020.3.10
- Turrisi, G.F., Matteini Palmerini, M. & Brothers, D.J. (2015) Systematic revision and phylogeny of the genera Blakeius Ashmead, 1903 and Liomutilla André, 1907, with description of two new genera (Hymenoptera: Mutillidae, Myrmillinae). Zootaxa, 4010 (1), 1–78. https://doi.org/10.11646/zootaxa.4010.1.1
- van Noort, S. (2025) WaspWeb: Hymenoptera of the World. Available from: https://www.waspweb.org/ (accessed 8 July 2025)]
- Waldren, G.C., Sadler, E.A., Murray, E.A., Bossert, S., Danforth, B.N. & Pitts, J.P. (2023) Phylogenomics inference of the higher classification of velvet ants (Hymenoptera: Mutillidae). Systematic Entomology, 48 (3), 463–487. https://doi.org/10.1111/syen.12588
- Waldren, G.C., Williams, K.A., Cambra, R.A. & Pitts, J.P. (2020) Systematic revision of the North American velvet ant genus Invreiella Suárez (Hymenoptera: Mutillidae) with description of eleven new species. Zootaxa, 4894 (2), 151–205. https://doi.org/10.11646/zootaxa.4894.2.1
- Wasbauer, M.S. (1973) Some new taxa in the Myrmosinae with keys to the females in North America. The Pan-Pacific Entomologist, 49, 325–337.
- Williams, K.A., Bartholomay, P.R., Cambra, R.A. & Oliveira, M.L. (2019d) Review of Leucospilomutilla Ashmead and description of the new genus Atlantilla (Hymenoptera: Mutillidae). Neotropical Entomology, 49, 82–97. https://doi.org/10.1007/s13744-019-00723-3
- Williams, K.A., Bartholomay, P.R., Cambra, R.A., Pitts, J.P. & Oliveira, M.L. (2022) Revision of Cephalomutilla André (Hymenoptera: Mutillidae) including one new species, three new sex associations, and nine new synonymies. Austral Entomology, 61 (2), 137–155. https://doi.org/10.1111/aen.12596
- Williams, K.A., Bartholomay, P.R. & Oliveira, M.L. (2017) Species groups of Traumatomutilla André (Hymenoptera: Mutillidae). Insecta Mundi, 533, 1–33.
- Williams, K.A., Brothers, D.J. & Pitts, J.P. (2011) New species of Tobantilla Casal, 1965 and a new genus and species, Gogoltilla chichikovi gen. et sp. nov., from Argentina (Hymenoptera: Mutillidae). Zootaxa, 3064 (1), 41–68. https://doi.org/10.11646/zootaxa.3064.1.2
- Williams, K.A., Cambra, R.A., Bartholomay, P.R., Lopez, V.M., Brothers, D.J., Martins, H.O. & Guillermo-Ferreira, R. (2024b) Two new genera of Neotropical Dasymutillini (Hymenoptera, Mutillidae, Sphaeropthalminae): Goncharovtilla gen. nov. from Brazil and Dasyphuta gen. nov. from Panama. Zootaxa, 5538 (2), 127–152. https://doi.org/10.11646/zootaxa.5538.2.2
- 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., Cambra, R.A., Waldren, G.C., Quintero, D.A. & Pitts, J.P. (2019b [“2018”]) Silvorientilla, gen. nov. a new Neotropical velvet ant genus (Hymenoptera: Mutillidae). Folia Entomológica Mexicana, Nueva Serie, 4 (3), 91–109.
- Williams, K.A., Lelej, A.S., Okayasu, J., Borkent, C.J., Malee, R., Thoawan, K. & Thaochan, N. (2019c) 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., Manley, D.G., Pilgrim, E.K., von Dohlen, C.D. & Pitts, J.P. (2010) Multifaceted assessment of species validity in the Dasymutilla bioculata species group (Hymenoptera: Mutillidae). Systematic Entomology, 36, 180–191. https://doi.org/10.1111/j.1365-3113.2010.00555.x
- Williams, K.A., Pan, A.D. & Wilson, J.S. (2024a) Velvet ants of North America. Princeton University Press, Princeton and Oxford, 440 pp. https://doi.org/10.2307/jj.5104027
- Williams, K.A., Pitts, J.P., Parikh, G.R., Cambra, R.A., Zhang, Y. & Bartholomay, P.R. (2025) Diagnostic Review of the Sphaeropthalmini (Hymenoptera: Mutillidae) of Central and North America, Part 1: minor nocturnal genera. Zootaxa. [in press]
- 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) African velvet ant (Hymenoptera: Mutillidae) Müllerian mimicry rings reveal the depauperate nature of insectivorous lizards in Africa. PLoS One, 13 (1), e0189482, 1–15. https://doi.org/10.1371/journal.pone.0189482
- Wilson, J.S., Williams, K.A., Forister, M.L., von Dohlen, C.D. & Pitts, J.P. (2012) Repeated evolution in overlapping mimicry rings among North American velvet ants. Nature Communications, 3, 1272. https://doi.org/10.1038/ncomms2275
- Zhou, H.-T., Lelej, A.S. & Liu, J.-X. (2018) 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