Pseudocephalotilla beira, copyright Bernard Dupont.

Belongs within: Mutillinae.
Contains: Smicromyrme.

Carried on velvet wings
Published 12 January 2023
Female Smicromyrme rufipes, copyright Roman Providukhin.

In their recent reclassification of the velvet ants of the Mutillidae, Brothers & Lelej (2017) assigned thirty genera found in Eurasia and Africa to the tribe Smicromyrmini. As with other mutillids, smicromyrmins usually show significant sexual dimorphism between winged males and wingless females (though the males of two genera, Gynandrotilla and Indratilla, are wingless—Lelej 2005). Males are characterised by a deep notch on the inner margin of the eyes, a metasoma with the first segment much narrower than the second, and antennae with the first flagellar segment much shorter than the second. Females have hairless eyes, concave sides to the mid-thorax, a narrow first metasomal segment as in the male, and usually one or three spots of pale pubescence arranged transversely across the dorsum of the second metasomal segment (Lelej 2005). This last distinctive pattern of pubescence was identified as a potential synapomorphy of the clade by Brothers & Lelej (2017). Other potential synapomorphies identified by these authors for females included the presence of a scutellar scale, a protrusion or tubercle on the mid-line of the mesosoma above the posterior face. Potential synapomorphies for males included the presence of mandibles with three apical teeth and a large posteroventral tooth, and genitalia with a very short gonobase not reaching dorsally and a basal ventral lamellate expansion on the volsella (though this was secondarily lost in several sub-groups).

Male Smicromyrme suberrata, copyright Thijs Valkenburg.

Host records for Smicromyrmini include a wide range of aculeate wasps and bees with the majority being ground-nesting solitary wasps (Ronchetti & Polidori 2020). Males of smicromyrmins are significantly larger than females, a feature that correlates with sexual phoresy (Brothers 1989). The male velvet ant grasps the female around the neck region with his mandibles and then flies some distance while carrying her underneath his mesosoma. Mating typically takes place after the female has been given her joyride. Brothers (1989) suggested that this behaviour may assist with dispersal, allowing the female to be carried between patches of suitable habitat far more quickly, safely and efficiently than she could manage herself by walking. Obviously, this behaviour would not be possible for those species in which the males are wingless. So what are they up to?

Systematics of Smicromyrmini
|--Promecilla André 1902BL17, LB08
| |--*P. regia (Smith 1855) [=Mutilla regia]LB08
| |--P. decora (Smith 1879) [=Sinotilla decora]BL17
| `--P. unicingulataBM76
`--+--Pseudocephalotilla Bischoff 1920BL17, LB08
| |--*P. beirana Bischoff 1921LB08
| |--P. atroposBL17
| | |--P. a. atroposBL17
| | `--P. a. kalahariensis (Bischoff 1921) [=Smicromyrme kalahariensis]BL17
| |--P. beira (Péringuey 1914) [=Mutilla beira]BL17
| `--P. tettensisBL17
| |--P. t. tettensisBL17
| `--P. t. brunni (Bischoff 1921) [=Smicromyrme brunni]BL17
`--+--+--Dentilla Lelej 1980BL17
| | |--D. curtiventris (André 1901)BL17 (see below for synonymy)
| | |--D. dichroa (Sichel & Radoszkowski 1869)BL17
| | |--D. persica (Sichel & Radoszkowski 1869)BL17
| | `--D. saharica (Gimer Mari 1945)BL17
| `--+--Sulcotilla Bischoff 1920BL17, LB08
| | `--*S. sulcata (Magretti 1884) [=Mutilla sulcata]LB08
| `--Nemka Lelej 1985BL17, LB08 [incl. Horaia Tsuneki 1993 non Tonnoir 1930LB08]
| |--*N. viduata (Pallas 1773) (see below for synonymy)LB08
| | |--N. v. viduataBL17
| | |--N. v. bartholomaei (Radoszkowski 1865)BL17
| | `--N. v. insulae (Invrea 1940)BL17
| `--‘Smicromyrme (*Horaia)’ chihpenchius Tsuneki 1993LB08
| `--Mickelomyrme Lelej 1995BL17, LB08
| |--*M. hageni (Zavattari 1913) [=Mutilla hageni; incl. Mu. alberici Zavattari 1913]LB08
| |--M. exacta (Smith 1879)BL17
| |--M. kuznetsovi Lelej 1996BL17
| |--M. yunnanensis Lelej 1996BL17
| `--M. zebina (Smith 1860) [=Mutilla zebina, Smicromyrme zebina]B22
`--+--Ephutomma Ashmead 1899BL17, LB08 [=Eputhomma (l. c.)LB08, Eremomyrme Suárez 1965LB08]
| |--*E. turcestanica (de Dalla Torre 1897)BL17 (see below for synonymy)
| |--E. angustata (Skorikov 1935)BL17
| `--E. continuaE66
`--Physetopoda Schuster 1949BL17, LB08 [incl. Paramyrme Lelej 1977LB08]
|--‘Mutilla’ rubrocincta Lucas 1848E12 [incl. *Physetopoda insularis Schuster 1949LB08]
|--P. daghestanica (Radoszkowski 1885)BL17
|--P. halensis (Fabricius 1787)BL17 (see below for synonymy)
|--P. pierrei (Suárez 1958)BL17
|--P. punctata (Latreille 1792)BL17
|--P. portschinskii (Radoszkowski 1888)BL17
`--P. scutellaris (Latreille 1792)BL17

Smicromyrmini incertae sedis:
Andreimyrme Lelej 1995BL17
|--‘Smicromyrme’ tridentiens Chen 1957 [incl. *Andreimyrme long Lelej 1995]LB08
`--A. silvorientalis Okayasu in Okayasu, Williams et al. 2021B22
Antennotilla Bischoff 1920BL17
`--*A. phoebe (Péringuey 1899) [=Mutilla phoebe]LB08
Corytilla Arnold 1956BL17
`--*C. pallidipes Arnold 1956LB08
Ctenoceraea Nonveiller 1993BL17
`--*C. pectinella Nonveiller 1993LB08
Ephucilla Lelej 1995BL17
`--*E. naja (Zavattari 1913) [=Mutilla naja]LB08
Guineomutilla Suárez 1977BL17 [incl. Phalacromyrme Nonveiller 1975 (n. n.)LB08]
`--*G. mateui Suárez 1977LB08
Gynandrotilla Arnold 1946BL17
`--*G. ferruginea Arnold 1946LB08
Indratilla Lelej 1993BL17
`--*I. gynandromorpha Lelej 1993LB08
Karunaratnea Lelej 2005BL17
`--*K. palatupanae Lelej 2005LB08
Nordeniella Lelej 2005BL17
`--*N. thermophila (Turner 1911) [=Mutilla thermophila]LB08
Nuristanilla Lelej in Lelej & Kabakov 1980BL17
`--*N. kabakovi Lelej in Lelej & Kabakov 1980LB08
Paglianotilla Lelej in Lelej & van Harten 2006BL17
`--*P. hogenesi Lelej in Lelej & van Harten 2006LB08
Psammotherma Latreille 1825BL17
`--*P. flabellata (Fabricius 1804) [=Mutilla flabellata]LB08
Rasnitsynitilla Lelej in Lelej & van Harten 2006BL17
`--*R. brachyptera Lelej in Lelej & van Harten 2006LB08
Sinotilla Lelej 1995BL17
`--*S. boheana (Chen 1957) [=Smicromyrme boheanus]LB08
Skorikovia Ovtchinnikov 2002BL17
`--*S. elongata (Radoszkowski 1885) [=Mutilla elongata; incl. Smicromyrme ambiguus Skorikov 1935]LB08
Tsunekimyrme Lelej 1995BL17
`--*T. fluctuata (Smith 1865)LB08 [=Mutilla fluctuataLB08, Smicromyrme fluctuataB22]

Dentilla curtiventris (André 1901)BL17 [=Mutilla curtiventrisLB08; incl. M. erronea André 1902LB08, *Dentilla erroneaLB08]

*Ephutomma turcestanica (de Dalla Torre 1897)BL17 [=Mutilla turcestanicaLB08, *Eremomyrme turcestanicaLB08, M. incerta Radoszkowski 1877 non Spinola 1841LB08; incl. Smicromyrme semenovi Skorikov 1935LB08]

*Nemka viduata (Pallas 1773) [=Mutilla viduata, Smicromyrme (*Nemka) viduata; incl. M. coronata Fabricius 1793]LB08

Physetopoda halensis (Fabricius 1787)BL17 [=Mutilla halensisLB08; incl. M. ephippium Fabricius 1793LB08, Mu. (Myrmosa) ephippiumG20, M. montana Panzer 1805LB08, Physetopoda montanaLB08, Smicromyrme (*Paramyrme) montanaLB08]

*Type species of generic name indicated


[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

Brothers, D. J. 1989. Alternative life-history styles of mutillid wasps (Insecta, Hymenoptera). In: M. N. Bruton (ed.) Alternative Life-History Styles of Animals pp. 279–291. Kluwer Academic Publishers: Dordrecht.

[B22] Brothers, D. J. 2022. Critical analysis of the type material of Mutillidae described from the Australasian region (Hymenoptera). Zootaxa 5140 (1): 1–215.

[BL17] Brothers, D. J., & A. S. Lelej. 2017. Phylogeny and higher classification of Mutillidae (Hymenoptera) based on morphological reanalyses. Journal of Hymenoptera Research 60: 1–97.

[E66] Evans, H. E. 1966. The Comparative Ethology and Evolution of the Sand Wasps. Harvard University Press: Cambridge (Massachusetts).

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

Lelej, A. S. 2005. Catalogue of the Mutillidae (Hymenoptera) of the Oriental Region. Dalnauka: Vladivostok.

[LB08] Lelej, A. S., & D. J. Brothers. 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: 1–79.

Ronchetti, F., & C. Polidori. 2020. A sting affair: a global quantitative exploration of bee, wasp and ant hosts of velvet ants. PLoS One 15 (9): e0238888.

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