Belongs within: Vespina.
Contains: Ichneumonidae, Urosigalphini, Brachistinae, Macrocentrinae, Orgilinae, Euphorinae, Sigalphinae, Agathidinae, Cheloninae, Cardiochilinae, Microgastrinae, Aphidiinae, Rhyssalinae, Doryctinae.
The Ichneumonoidea are a hyperdiverse group of parasitoid wasps, most of which are more or less slender-bodied with long, multi-segmented antennae bearing a short scape. Synapomorphies are modification of the pronotum with the prepectus fused to the posterior margin, and division of the second metasomal sternite between a strongly sclerotised anterior portion and membranous posterior portion. The costal and subcostal veins of the fore wing are usually fused though a minute costal cell may remain adjacent to the parastigma in some members of the family Braconidae. The ovipositor is often strongly exserted (Naumann 1991).
Living species of the Ichneumonoidea are mostly divided between the two families Ichneumonidae and Braconidae, with members of the Braconidae having the second recurrent vein present in the fore wing, cell R in the hind wing shorter than the submarginal vein or absent, and metasomal tergites 3 and 4 rigidly joined. Praeichneumon townesi, a fossil species from the Early Cretaceous of Mongolia, may represent a basal representative of the Ichneumonoidea. The costal cell is present but narrowed, and the antenna includes more than fifteen segments (Grimaldi & Engel 2005). The Eoichneumonidae are more definitive ichneumonoids known from the Early Cretaceous of Asia and Australia (Grimaldi & Engel 2005).
The braconids are currently divided between about forty subfamilies. Depending on the subfamily, species may be either ectoparasitic or endoparasitic as larvae. Females typically attack host larvae though some may oviposit on eggs with development delayed until the host larva is mature (Naumann 1991). Achterberg (1993) regarded ectoparasitism on wood-boring beetles as the basal life habit for the family, but this remains open to contention. The Rhyssalinae and Doryctinae belong to a group known as the ‘cyclostome’ braconids, in which the face is marked by a deep, wide hypoclypeal depression, with the middle of the apparent ventral margin of the clypeus distinctly above the upper level of the mandibular bases (Achterberg 1993).
The Apozyginae are a small subfamily, known only from Chile, which retain certain wing veins usually absent in other braconids (2m-cu in the forewing and 2-CU in the hind wing), and have the second and third metasomal sternites fused into a single plate (Achterberg 1993). Some authors have historically treated the apozygines as a distinct family within the Ichneumonoidea. The Microtypinae are a small, cosmopolitan subfamily of endoparasites on larval Lepidoptera that have the second submarginal cell of the fore wing small and triangular or a narrow trapezoid, and the dorsal carinae of the first metasomal tergite weakly developed or absent basally (Achterberg 1993).
The Helconinae are a group of braconid wasps endoparasitic on larval Coleoptera. Their fore wings contain two submarginal cells and vein r-m is present (Achterberg 1993). The Asian and Australasian Xiphozelinae have the lateropes round, deep and far removed from the base of the first metasomal tergite, and vein cu-a of the hind wing very long and strongly reclivous (Achterberg 1993). A sclerotised vein 2-CU is retained in the hind wing of Trachypetinae, Pselaphaninae, Meteorideinae and Sigalphinae. The Australian Cercobarconini have a peculiar flattened area ventrally on the hind tibiae, and the hind spurs are flattened and lamelliform. The Neotropical Pselaphaninae have fore wings with vein r-m inclivous and SR1 long (Achterberg 1993).
Multifarities most horrid
Published 15 June 2009
We all know that J. B. S. Haldane is supposed to have remarked that God seemed to have an “extraordinary fondness for beetles”. What Haldane may not have realised was the possibility that the beetles were just a means to an end. As the current rate of taxonomic description is considered, some researchers have come to the suspicion that the true objects of the Creator’s affection are not beetles, but parasitoid wasps*. Which, when you consider the natures of parasitoid wasps, kind of explains some things about life.
*Personally, I’m still taking the long odds and backing the nematodes.
The Braconidae are just one of the stupidly diverse lineages of Hymenoptera. According to ToLWeb (in 2004), there are some 12,000 described species of braconids, with estimates of up to 50,000 in total. Braconids form the living sister group to the similarly diverse Ichneumonidae, though braconids tend to be smaller in size (still, some of them are more than big enough). Braconids include both exoparasitic and endoparasitic taxa, parasitoids of eggs, larvae or adult insects, and a small number of gall-forming plant-parasitic taxa for added variety. The usual opinion is that the exoparasitic taxa represent the ancestral lifestyle for the family, but the actual phylogeny of the family is still being hammered out (and the “usual opinion” may yet turn out to be the wrong opinion). About forty subfamilies are currently recognised, but most authors (e.g. Shi et al. 2005) divide those subfamilies between three main lineages, the cyclostomes, microgastroids and helcionoids, with some subfamilies of uncertain position relative to the three. The microgastroids and helcionoids are all koinobiont endoparasitoids (after the wasp has laid its eggs in the host, the host continues to grow and develop), while the cyclostomes include both exoparasitoids and endoparasitoids, with exoparasitoids usually paralysing the host before laying their eggs (Wharton 1993). The microgastroids are fastidious in their tastes, restricting their diet to Lepidoptera (Murphy et al. 2008), while helcionoids attack a wide variety of hosts, including hemimetabolous as well as holometabolous insects. Early phylogenetic studies suggested that the cyclostomes (which possess a distinctive mouthpart morphology) were paraphyletic with regard to the other braconids, but more recent studies support a monophyletic cyclostome clade (Shi et al. 2005). The monophyly of a microgastroid + helcionoid clade is supported by molecular data (Shi et al. 2005) but remains short on morphological support (Quicke et al. 1999).
The Aphidiinae are the largest group of braconids to not fit comfortably within the three-way division. Aphidiinae are parasitoids of aphids. Morphological data supports a relationship between aphidiines and the cyclostome group (Quicke et al. 1999), but the molecular analysis of Shi et al. (2005) suggested a relationship between the Aphidiinae and the Euphorinae, members of the helcionoids. The intriguing feature of this result is that Aphidiinae and Euphorinae both have the unusual characteristic (for insect parasitoids) of parasitising adult hosts rather than larvae—albeit with different host ranges in the two subfamilies. Euphorinae were probably originally parasitoids of beetles but some species have since become parasitoids of hosts as diverse as grasshoppers or Psocoptera. A more recent combined morphological and molecular study whose authors argued against an Aphidiinae-Euphorinae relationship in favour of an Aphidiinae-cyclostome connection (Zaldivar-Riverón et al. 2006) actually did not test anything either way, because the authors’ choice of taxa and outgroup effectively forced an a priori cyclostome position.
Finally, some would think it rather remiss of me to write about braconids without making some mention of polydnaviruses, but I don’t see why I should when a much better description of such things than I could produce has already been written by Merry Youle at Small Things Considered (Update: Unfortulately, it looks like this page may no longer be available). After you read the main article there, though, make sure you scroll down the comments to Merry’s description of the differences between polydnaviruses in Braconidae and Ichneumonidae suggesting the independent origins of the polydnavirus system in the two families. As well as the differences described by Mary, it also turns out that polydnaviruses are not characteristic of braconids as a whole, but are in fact only found within the microgastroid clade (Wharton 1993), so an independent origin from ichneumonid polydnaviruses has phylogenetic as well as biochemical support.
Systematics of Ichneumonoidea
<==Ichneumonoidea
|--Praeichneumon [Praeichneumonidae]GE05
| `--P. townesiRJ93
`--+--IchneumonidaeGE05
`--+--Eoichneumon Jell & Duncan 1986H02 [EoichneumonidaeGE05]
| `--E. duncanae Jell & Duncan 1986RJ93
`--Braconidae [Anomalidae, Branconidae]GE05
| i. s.: Pseudobathystomus (Atlantobathystomus Belokobylskij & Koponen 2004)F05
| `--P. (*A.) macaronesius Belokobylskij & Koponen 2004F05
| Chaenon ancepsMS01
| Diaeretus essigellaePK17
| Rhetinorhyssalus morticinusPK17
| Fushunobracon Hong 2002H02
| `--*F. orientalis Hong 2002H02
| Sinobracon Hong 1974H02
| `--*S. speciosus Hong 1974H02
| Eobraconus inopinatusRJ93
| PromicrogasterD96
| Sycosoter lavagneiRD77
| AnacanthobraconP92
| AspicolpusP92
| ChelonohelconP92
| CoeloreuteusP92
| DigastrothecaP92
| DiodontogasterP92
| DiospilitesP92
| DoryctomorphaP92
| ElasmosomitesP92
| ElectrohelconP92
| EocardiochilesP92
| EumacrocentrusP92
| HormiellusP92
| MeteoritesP92
| MiracoidesP92
| Neoblacus facialis Brues 1937P92
| OnychouraP92
| PalaeorhyssalusP92
| ParasyrrhizusP92
| PolystenusP92
| ProchremylusP92
| PromonolexisP92
| SemirhytusP92
| TaphaeusP92
| Dendrosotinus insularis Belokobylskij, Iqbal & Austin 2004JG19
| Neoheterospilus insularis Belokobylskij 2006JG19
| Apozyx [Apozygidae, Apozyginae]A93
| `--A. penyaiA93
| MicrotypinaeA93
| |--Microtypus wesmaeliiA93
| `--Neomicrotypus penaiA93
| Acaelius cadmium Papp 2003F04
| Antrusa curtitempus Fischer, Tormos et al. 2004F04
| GildoriaF04
| |--G. elongata van Achterberg 2003F04
| `--G. iberica van Achterberg 2003F04
|--+--+--AcampsohelconinaeA02
| | | |--UrosigalphiniBS23
| | | |--Afrocampsis van Achterberg & Quicke 1990 [Afrocampsini]A02
| | | | `--A. griseosetosusA02
| | | `--Acampsohelcon Tobias 1987 [Acampsohelconini]A02
| | | `--A. rasnitzini Tobias 1987A02
| | `--+--+--MeteorideinaeA93
| | | | |--Pronkia [Pronkiini]A93
| | | | | `--P. antefurcalisA93
| | | | `--Meteoridea [Meteorideini]A93
| | | | `--M. hutsoniA93
| | | `--BrachistinaeBS23
| | `--+--HelconinaeA93
| | | |--TrichiohelconN91
| | | |--AustrohelconR70
| | | |--HelconideaR70
| | | |--Brulleia [Brulleiini]A93
| | | | `--B. melanocephalaA93
| | | |--HelconBS23 [incl. GymnoscelusP92; HelconiniA93]
| | | | |--H. nunciatorA93
| | | | `--H. tardator [=Alysia (Helcon) tardator]G20
| | | `--Wroughtonia ligatorBS23, HR11
| | `--+--+--Amicrocentrum [Amicrocentrinae]A93
| | | | `--A. concolorA93
| | | `--+--MacrocentrinaeA93
| | | `--Charmon [Charmontinae]BS23
| | | `--C. cruentatusA93
| | `--+--+--Xiphozele [Xiphozelinae]A93
| | | | `--X. compressiventrisA93
| | | `--HomolobinaeA93
| | | |--Westwoodiella [Westwoodiellini]A93
| | | | `--W. bicolorA93
| | | |--Homolobus [Homolobini]A93
| | | | |--H. huddlestoniA93
| | | | `--H. infumatorA93
| | | `--ExasticolusBS23
| | `--OrgilinaeBS23
| `--+--+--CenocoeliinaeA93
| | | |--CapitoniusBS23
| | | | |--C. bifasciatusA93
| | | | `--C. chontalensisBS23
| | | |--CenocoeliusPS05
| | | | |--C. analisA93
| | | | `--C. huggerti Pitz & Sharkey 2005PS05
| | | `--Aulacodes nigriventrisBS23
| | `--EuphorinaeBS23
| `--+--+--+--IchneutinaeBS23
| | | | | i. s.: Anaprixia Mason 1991A93
| | | | |--+--ParoligoneurusBS23
| | | | | `--Oligoneurus [Muesebeckiini]BS23
| | | | | `--O. concolorA93
| | | | `--Ichneutes Nees 1816BS23, HP-W05 [IchneutiniA93]
| | | | |--I. brevisA93
| | | | `--I. reunitorA93
| | | `--Dirrhope [Dirrhopinae]A93
| | | `--D. rufaA93
| | `--+--+--SigalphinaeBS23
| | | `--TrachypetinaeA93
| | | |--Cercobarcon [Cercobarconinae, Cercobarconini]A93
| | | | `--C. riekiA93
| | | `--Trachypetus [Trachypetini]A93
| | | `--T. clavatusA93
| | `--+--AgathidinaeBS23
| | `--Pselaphanus [Pselaphaninae]A93
| | `--P. trogoidesA93
| `--+--Proterops [Proteropinae]BS23
| | |--P. nigripennisA93
| | `--P. proteroptoidesBS23
| `--+--+--Ecnomios [Ecnomiinae]A93
| | | `--E. papuensisA93
| | `--CheloninaeBS23
| `--+--Neoneurus [Neoneurinae]A93
| | `--N. auctusA93
| `--+--+--MiracinaeA93
| | | |--Centistidea ectoedemiaeA93
| | | `--MiraxBS23
| | | |--M. insularis Muesebeck 1937M37
| | | |--M. leucopterae Wilkinson 1936M37
| | | `--M. texanaM37
| | `--+--Khoikhoia [Khoikhoiinae]A93
| | | `--K. lissionA93
| | `--CardiochilinaeBS23
| `--MicrogastrinaeBS23
`--+--AphidiinaeBS23
`--+--RhyssalinaeBS23
`--DoryctinaeBS23
*Type species of generic name indicated
References
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Murphy, N., J. C. Banks, J. B. Whitfield & A. D. Austin. 2008. Phylogeny of the parasitic microgastroid subfamilies (Hymenoptera: Braconidae) based on sequence data from seven genes, with an improved time estimate of the origin of the lineage. Molecular Phylogenetics and Evolution 47 (1): 378–395.
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Hi Christopher! The blog post you linked doesn’t appear to be accessible on Small Things Considered nor on the Wayback Machine. Any chance you know of a different way to access it?
Unfortunately, it looks like anything from earlier than 2019 may have disappeared from the Small Things Considered website. I looked around and couldn’t find any alternative source; if I do, I’ll update the post. Thank you for pointing this out to me, Jason.