Brachymeria (Brachymeria)

Brachymeria ucalegon, a parasitoid of blow fly larvae, photographed by C. Bento.

Belongs within: Chalcididae.

Brachymeria subgenus Brachymeria includes both primary parasitic and hyperparasitic wasps in pupae of holometabolous insects, particularly Lepidoptera (Bouček 1988).

Brachymeria perflavipes and beyond
Published 4 June 2012
Male of a Brachymeria species, possibly B. hammari, photographed by John Hallmén.

For the subject of today’s post, the wheel-spin brought up the parasitic wasp Brachymeria perflavipes. This is a species of the family Chalcididae described by our old friend Alexander Arsene Girault in 1913 from a female specimen collected in the Dandenong Mountains in Victoria, Australia. He described it under the name of Tumidicoxa flavipes; when that genus was later synonymised by Girault in 1915, the species name was replaced by perflavipes as there was already a Brachymeria flavipes. Girault’s description of this species was, thankfully, more detailed than some of his later productions, though the species was not illustrated (not an uncommon state of affairs at the time) and, to the best of my knowledge, never has been. Girault (1915) later also recorded a male from Brisbane in Queensland, collected from a lepidopteran host. And that, except for an extra host record by Bouček (1988), is pretty much the sum total of our knowledge of this species.

Brachymeria is a large genus of Chalcididae, with over a hundred species worldwide. The Chalcididae are, in my opinion, one of the more instantly recognisable families of Chalcidoidea, with hind femora distinctly larger than those on other legs and with a ventral row of teeth. These large hind femora enable the wasp to jump, and I can confirm from personal observation that they are adept pingers. Most Chalcididae are black with coarse punctation over most of the body, though an interesting find on one of our recent field trips was a bright electric blue chalcidid. However, despite their recognisability, a recent major molecular phylogenetic analysis of Chalcidoidea failed to recover the Chalcididae as monophyletic. Brachymeria is distinguished from other chalcidid genera by features such as a short petiole, short postmarginal vein (but even shorter stigmal vein), distinct malar suture, and hind tibia with a distinct apical spine (Bouček 1988). Attempts have been made to subdivide Brachymeria into subgenera but proposed subgeneric divisions have been based on species found within a restricted biogeographic area (e.g. Burks 1960; Bouček 1988) and a broad worldwide survey of Brachymeria still hasn’t been conducted.

Brachymeria ovata on pupa of crow butterfly Euploea core, from Brisbane Insects.

Most Brachymeria species (including B. perflavipes) are parasitoids of lepidopteran pupae, but some attack dipteran larvae or pupae (more on that in a moment). Though more than one egg may be laid within a single pupa, Dowden (1935) found that in the case of B. intermedia, a parasitoid of the gypsy moth Lymantria dispar, only one wasp would develop to maturity in a single host. Indications are that the wasp larvae themselves kill any would-be competitors within the host: the mandibles of the larva are strongly sclerotised even in the first instar, and Dowden found that dead larvae dissected out from a host invariably bore signs of injury from the mandibles. Larvae dissected out from a host and placed together would quickly attack each other.

Diagram of distinguishing characters of Brachymeria intermedia vs B. compsilurae, from Dowden (1935).

Brachymeria species also include a good example of the potential importance of reliable taxonomic identifications. The aforementioned B. intermedia has been the subject of deliberate introductions outside its native range in attempts to control its host, the gypsy moth (or, more accurately, one of its wide range of hosts). Another Brachymeria species, B. compsilurae, closely resembles B. intermedia and also appears to lay its eggs in gypsy moths. However, the target of B. compsilurae is not the gypsy moth itself, but the larvae of tachinid flies that are parasitising the moth and are themselves used in its control. Programs to introduce B. intermedia to New England had to be careful to eliminate B. compsilurae (Burks 1960), whose release would have the opposite effect of that intended.

Systematics of Brachymeria (Brachymeria)

Characters (from Bouček 1988): Malar suture between eye and mouth distinct, usually indicated by a carina; thorax rather densely punctured, dorsally convex; propodeum with rough alveolation reaching its anterior margin; propodeal spiracle elongate in oblique subhorizontal direction; postmarginal vein much shorter than marginal vein, often longer than stigmal vein, latter very short; hind legs at most moderately thickened; apex of hind tibia tapering into a strong spine extended far beyond insertion of tarsus; gaster attached at coxae; petiole very short, mostly concealed in dorsal view.

<==Brachymeria (Brachymeria) (see below for synonymy)B88
    |--*B. (B.) minuta (Linnaeus 1767) [=Vespa minuta, Chalcis minuta; incl. C. pusilla, Haltichella pusilla]B88
    |--B. (B.) albitegula (Girault 1915) [=Chalcis albitegula]B88
    |--B. (B.) aligherei (Girault 1927) [=Chalcis aligherei]B88
    |--B. (B.) anselmi (Girault 1926)B88 (see below for synonymy)
    |--B. (B.) aurea (Girault 1915) [=Chalcis aurea; incl. C. delii Girault 1924]B88
    |--B. (B.) australiensis (Girault 1913) [=Tumidicoxa (Tumidicoxella) australiensis]B88
    |--B. (B.) borealis (Girault 1915) (see below for synonymy)B88
    |--B. (B.) bouceki Narendran 1978B88
    |--B. (B.) brisbanensis (Girault 1915) [=Chalcis brisbanensis; incl. C. miltoni Girault 1926]B88
    |--B. (B.) browningi (Girault 1915) [=Chalcis browningi]B88
    |--B. (B.) caesar (Girault 1926) [=Chalcis caesar]B88
    |--B. (B.) cailliaudii (Montrousier 1864) [=Chalcis cailliaudii]B88
    |--B. (B.) confalonierii [=B. (*Neobrachymeria) confalonierii]B88
    |--B. (B.) corneillei (Girault 1924) [=Chalcis corneillei; incl. C. aequalipunctatus Girault 1924]B88
    |--B. (B.) cosmophilae (Girault 1925) [=Chalcis cosmophilae]B88
    |--B. (B.) curtisi (Girault 1915) [=Chalcis curtisi]B88
    |--B. (B.) decens (Girault 1929) [=Chalcis decens]B88
    |--B. (B.) edna (Girault 1927) [=Chalcis pomonae var. edna]B88
    |--B. (B.) euploeae (Westwood 1837) [=Chalcis (B.) euploeae]B88
    |--B. (B.) falsosa (Vachal 1907) [=Chalcis falsosa]B88
    |--B. (B.) froggatti (Cameron 1912) [=Chalcis froggatti]B88
    |--B. (B.) hercules (Girault 1913) [=Pseudopitelea (l. c.) hercules, Chalcis hercules]B88
    |--B. (B.) hypolycaenae (Girault 1915) [=Chalcis hypolycaenae]B88
    |--B. (B.) indignator (Walker 1862) [=Halticella indignator]B88
    |--B. (B.) internata (Walker 1862) [=Halticella internata]B88
    |--B. (B.) judaei (Girault 1937) [=Chalcis judaei]B88
    |--B. (B.) juno (Girault 1927) [=Chalcis juno; incl. C. leo Girault 1933, C. minerva Girault 1927]B88
    |--B. (B.) lasus (Walker 1841) (see below for synonymy)B88
    |--B. (B.) lugubris (Walker 1871) [=Chalcis lugubris; incl. C. atrata Kirby 1883]B88
    |--B. (B.) mariana (Girault 1927) [=Chalcis schuberti var. mariana]B88
    |--B. (B.) marmonti (Girault 1924) (see below for synonymy)B88
    |--B. (B.) megaspila (Cameron 1907) [=Chalcis megaspila; incl. C. koebelei Crawford 1910, C. poema Girault 1927]B88
    |--B. (B.) menoni Joseph, Narendran & Joy 1972B88
    |--B. (B.) minutissima (Girault 1926) [=Chalcis minutissimus]B88
    |--B. (B.) mochii Masi 1936B88
    |--*Tumidicoxa’ nigraB88
    |--B. (B.) nigricoxa (Girault 1913) [=Tumidicoxa (*Tumidicoxella) nigricoxa, Chalcis nigricoxa]B88
    |--B. (B.) nortia (Girault 1929) [=Chalcis plutellophaga var. nortia]B88
    |--B. (B.) ogyrisidis (Girault 1922) [=Chalcis ogyrisidis]B88
    |--B. (B.) opponens (Walker 1871) [=Chalcis opponens]B88
    |--B. (B.) pandani (Girault 1926) [=Chalcis pandani]B88
    |--B. (B.) parvicorpus (Girault 1925) [=Chalcis parvicorpus]B88
    |--B. (B.) perflavipes (Girault 1915) (see below for synonymy)B88
    |--B. (B.) persplendidipes (Girault 1926) [=Chalcis persplendidipes]B88
    |--B. (B.) phya (Walker 1838) [=Chalcis phya; incl. C. bachi Girault 1924]B88
    |--B. (B.) piratica (Brues 1918) [=Chalcis piraticus]B88
    |--B. (B.) plutellophaga (Girault 1922) [=Tumidicoxella plutellophaga]B88
    |--B. (B.) podagrica (Fabricius 1787) (see below for synonymy)B88
    |--B. (B.) pomonae (Cameron 1912) [=Chalcis pomonae]B88
    |--B. (B.) puella (Girault 1927) [=Chalcis puella]B88
    |--B. (B.) redia (Girault 1929) [=Chalcis redia]B88
    |--B. (B.) rubrifemur (Girault 1913) [=*Pseudepitelia rubrifemur, Chalcis rubrifemur; incl. C. ruskini Girault 1915]B88
    |--B. (B.) ruficornis (Girault 1913) [=Tumidicoxa ruficornis]B88
    |--B. (B.) rufifemur (Girault 1913) [=Tumidicoxa rufifemur]B88
    |--B. (B.) salomonis (Cameron 1911) [=Chalcis salomonis]B88
    |--B. (B.) sanguiniventris (Girault 1927) [=Chalcis calliphorae sanguiniventris]B88
    |--B. (B.) schuberti (Girault 1924) [=Chalcis schuberti]B88
    |--B. (B.) shakespearei (Girault 1926) [=Chalcis shakespearei]B88
    |--B. (B.) shellyi (Girault 1915) [=Chalcis shellyi]B88
    |--B. (B.) sidnica Holmgren 1868B88 (see below for synonymy)
    |--B. (B.) taiwana [=Chalcis taiwana, B. (*Matsumurameria) taiwana]B88
    |--B. (B.) terribilis (Brues 1928) [=Chalcis terribilis]B88
    |--B. (B.) teuta (Walker 1841) (see below for synonymy)B88
    |--B. (B.) ucalegon (Walker 1846) (see below for synonymy)B88
    |--B. (B.) unguttifemur (Girault 1926) [=Chalcis unguttifemur]B88
    |--B. (B.) vergilii (Girault 1927) [=Chalcis rubripes vergilii]B88
    |--B. (B.) veronesini (Girault 1924) [=Chalcis veronesini]B88
    |--B. (B.) vicaria (Walker 1861)B88
    |--B. (B.) victoria (Girault 1913) [=Tumidicoxa victoria, Chalcis victoria]B88
    `--B. (B.) yamalae (Girault 1924) [=Chalcis yamalae]B88

Brachymeria Westwood in Stephens 1829 (Brachymeria) [incl. Brachepitelia Girault 1913, Matsumurameria Habu 1960, Neobrachymeria Masi 1929, Pseudepitelia Girault 1913, Tumidicoxa Girault 1911, Tumidicoxella Girault 1913, Tumidicoxoides Girault 1913]B88

Brachymeria (Brachymeria) anselmi (Girault 1926)B88 [=Chalcis anselmiB88; incl. Tumidicoxoides pulchripes Girault & Dodd in Girault 1915 non Brachymeria pulchripes Holmgren 1868B88, C. pulchripesGM79, C. xenophoni Girault 1930B88]

Brachymeria (Brachymeria) borealis (Girault 1915) [=Chalcis robusta var. borealis; incl. C. epicteti Girault 1924, C. rex Girault 1924, C. robusta Girault 1915 non Cresson 1865]B88

Brachymeria (Brachymeria) lasus (Walker 1841) [=Chalcis lasus; incl. C. regina var. copernici Girault 1936, C. regina var. dentata Girault 1915, C. nitator Walker 1862, C. papuana Cameron 1913, Tumidicoxa regina Girault 1913, C. regina]B88

Brachymeria (Brachymeria) marmonti (Girault 1924) [=Chalcis marmonti; incl. B. koduvalliensis Joseph, Narendran & Joy 1973, C. wittei Schmitz 1946]B88

Brachymeria (Brachymeria) perflavipes (Girault 1915) [=Chalcis perflavipes, Tumidicoxa flavipes Girault 1925 non Chalcis flavipes Fabricius 1793]B88

Brachymeria (Brachymeria) podagrica (Fabricius 1787) [=Chalcis podagrica; incl. C. dipterophaga Girault & Dodd in Girault 1915, *Tumidicoxoides kurandaensis Girault 1913, Brachymeria kurandaensis, Chalcis kurandanensis, Tumidicoxoides paucipunctatus Girault 1915, C. vegai Girault 1924]B88

Brachymeria (Brachymeria) sidnica Holmgren 1868B88 [incl. Chalcis elijahi Girault 1927B88, C. multicolor Girault 1915 non Kieffer 1905B88, C. silvae Girault 1924B88, C. tegularis Cameron 1912 non Cresson 1872B88, Brachymeria tegularisBD17, C. thymus Girault 1917B88, Pseudepitelia tricolor Girault 1913 non Chalcis tricolor Förster 1859B88]

Brachymeria (Brachymeria) teuta (Walker 1841) [=Chalcis teuta; incl. *Brachepitelia rubripes Girault 1913, C. rubripes, Tumidicoxa (Tumidicoxella) tasmaniensis Girault 1913]B88

Brachymeria (Brachymeria) ucalegon (Walker 1846) [=Chalcis ucalegon; incl. C. calliphorae Froggatt 1916, C. rubriventris Girault 1915, Tumidicoxa rufiventris Girault 1913 non Chalcis rufiventris Kieffer 1905]B88

*Type species of generic name indicated

References

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[BD17] Branstetter, M. G., B. N. Danforth, J. P. Pitts, B. C. Faircloth, P. S. Ward, M. L. Buffington, M. W. Gates, R. R. Kula & S. G. Brady. 2017. Phylogenomic insights into the evolution of stinging wasps and the origins of ants and bees. Current Biology 27: 1019–1025.

Burks, B. D. 1960. A Revision of the genus Brachymeria Westwood in America north of Mexico (Hymenoptera: Chalcididae). Transactions of the American Entomological Society 86 (3): 225–273.

Dowden, P. B. 1935. Brachymeria intermedia (Nees), a primary parasite, and B. compsilurae (Cwfd.), a secondary parasite, of the gypsy moth. Journal of Agricultural Research 50 (6): 495–523.

Girault, A. A. 1913. On several new genera and species of Australian Hymenoptera Chalcidoidea. Canadian Entomologist 45: 101–106.

Girault, A. A. 1915. Australian Hymenoptera Chalcidoidea—XIV. The family Chalcididae with descriptions of new genera and species. Memoirs of the Queensland Museum 4: 314–365.

[GM79] Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 1–400.

Munro, J. B., J. M. Heraty, R. A. Burks, D. Hawks, J. Mottern, A. Cruaud, J.-Y. Rasplus & P. Jansta. 2011. A molecular phylogeny of the Chalcidoidea (Hymenoptera). PLoS One 6 (11): e27023. doi:10.1371/journal.pone.0027023.

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