Braconidae

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Braconidae
Temporal range: Cretaceous–Present
Braconidae Richard Bartz.jpg
Atanycolus sp.
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Superfamily: Ichneumonoidea
Family: Braconidae
Latreille, 1829
Subfamilies

47, see text

Synonyms

Stenophasmidae Benoit, 1949

The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with about 17,000 recognized species and many thousands more undescribed. [1] One analysis estimated a total between 30,000 and 50,000, and another provided a narrower estimate between 42,000 and 43,000 species. [1]

Contents

A female tropical braconid ovipositing into dead wood

Classification

The Braconidae are currently divided into about 47 subfamilies and over 1000 genera, [2] which include Aerophilus , Aleiodes , Apanteles , Asobara , Bracon , Cenocoelius , Chaenusa , Chorebus , Cotesia , Dacnusa , Diachasma , Microgaster , Opius , Parapanteles , Phaenocarpa , Spathius , and Syntretus.

These fall into two major groups, informally called the cyclostomes and noncyclostomes. In cyclostome braconids, the labrum and the lower part of the clypeus are concave with respect to the upper clypeus and the dorsal margin of the mandibles. These groups may be clades that diverged early in the evolution of braconids. [3] Cyclostomes are monophyletic whereas noncyclostomes can be divided formally into microgastroids, sigalphoids, helconoids, and euphoroids. [4]

Subfamilies

Head of a cyclostome braconid showing circular opening above mandibles. Aleiodes shakirae head.jpg
Head of a cyclostome braconid showing circular opening above mandibles.
Head of a noncyclostome braconid. Cf Aphidiinae (16779450492).jpg
Head of a noncyclostome braconid.

Morphology

The morphological variation among braconids is notable. They are often black-brown (sometimes with reddish markings), though some species exhibit striking coloration and patterns, being parts of Müllerian mimicry complexes. They have one or no recurrent veins, unlike other members of the other family in Ichneumonoidea (Ichneumonidae), which usually have two. Wing venation patterns are otherwise highly variable. The antennae typically have 16 segments or more; the trochanters have two segments.

Females often have long ovipositors, an organ that largely varies interspecifically. This variation is closely related to the host species upon which the wasp deposits its egg. Species that parasitize microlepidopterans, for instance, have longer ovipositors, presumably to reach the caterpillar through layers of plant tissue. Some wasps also have long ovipositors to bypass caterpillar defense mechanisms such as spines or hairs, or to reach deeply-burrowed Coleoptera larvae in tree trunks. [5]

Life history

Cotesia ruficrus illustrated by Des Helmore HYME Braconidae Cotesia ruficrus.png
Cotesia ruficrus illustrated by Des Helmore

The larvae of most braconids are internal or external primary parasitoids of other insects, especially the larval stages of Coleoptera, Diptera, and Lepidoptera, but also some hemimetabolous insects such as aphids, Heteroptera, or Embiidina. Most species kill their hosts, though some cause the hosts to become sterile and less active. Parasitoidy on adult insects (particularly on Hemiptera and Coleoptera) also occurs. Members of two subfamilies, the Mesostoinae and Doryctinae are known to form galls on plants. [6] [7] Braconids are often used as biological pest control agents, especially against aphids. [8]

Examples of hosts

Thousands of species of insects are used as hosts by braconid wasps. A few notable examples are detailed here.

Some species of braconids are parasitoids of Ostrinia furnacalis (the Asian corn borer, a lepidopteran moth known for being a pest of maize in East Asia), the African sugarcane borer (a moth commonly found in sub-Saharan Africa), [9] the butterfly Danaus chrysippus in Ghana, [10] and Liriomyza trifolii (the American serpentine leafminer) and Manduca quinquemaculata (the tomato hornworm) in North America. [11] Braconids often will prey on fruit fly larvae like Anastrepha suspensa as well. [12]

Polydnaviruses

Endoparasitoid species often display elaborate physiological adaptations to enhance larval survival within the host, such as the co-option of endosymbiotic viruses for compromising host immune defenses. These bracoviruses are often used by the wasps instead of, or in addition to, a venom cocktail. The DNA of the wasp actually contains portions that are the templates for the components of the viral particles and they are assembled in an organ in the female's abdomen known as the calyx. [13] A 2009 study has traced the origins of these templates to a 100-million-year-old viral infection whose alterations to its host DNA provided the necessary basis for these virus-like "templates". [14]

These viruses suppress the immune system and allow the parasitoid to grow inside the host undetected. The exact function and evolutionary history of these viruses are unknown. Sequences of polydnavirus genes show the possibility that venom-like proteins are expressed inside the host caterpillar. Through the evolutionary history of being used by the wasps, these viruses apparently have become so modified, they appear unlike any other known viruses today. Because of this highly modified system of host immunosuppression, a high level of parasitoid-host specificity is not surprising.

Evolutionary history

The family seems to date from early Cretaceous (provided that Eobracon is properly assigned to this family). It underwent extensive diversification from mid or late Cretaceous to early Cenozoic, correlating with the radiation of flowering plants and associated insect herbivores, the main hosts of braconids.

Differentiation from Ichneumonidae

Ichneumonidae wing morphology Ichneumonidae.svg
Ichneumonidae wing morphology
Braconidae wing morphology Braconidae g1.svg
Braconidae wing morphology

Braconids are distinguished from their sister group Ichneumonidae by these character combinations. In Braconidae, vein 2m-cu of the forewing is absent except in the Chilean species Apozyx penyai – this vein is present in 95% of Ichneumonidae. Vein 1/Rs+M of the forewing is 85% present in Braconidae, but absent in all Ichneumonidae. Vein 1r-m of the hind wing is in 95% of Braconidae basal to the separation of R1 and Rs (it is opposite or apical in Ichneumonidae). In Braconidae, metasomal tergum 2 is fused with tergum 3, (secondarily flexible in Aphidiinae) – 90% of Ichneumonidae have a flexible suture. [15]

Other characteristics

The species Microplitis croceipes possesses an extremely accurate sense of smell and can be trained for use in narcotics and explosives detection. [16]

At least some braconids appear to be very resistant to ionizing radiation. While a dose of 400 to 1000 rads can kill an average human, a dose of 180,000 rads was required to kill a braconid of genus Habrobracon in an experiment. [17]

Related Research Articles

<span class="mw-page-title-main">Parasitoid</span> Organism that lives with its host and kills it

In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionary strategies within parasitism, distinguished by the fatal prognosis for the host, which makes the strategy close to predation.

<span class="mw-page-title-main">Ichneumonoidea</span> Superfamily of wasps

The superfamily Ichneumonoidea contains one extinct and three extant families, including the two largest families within Hymenoptera: Ichneumonidae and Braconidae. The group is thought to contain as many as 100,000 species, many of which have not yet been described. Like other parasitoid wasps, they were long placed in the "Parasitica", variously considered as an infraorder or an unranked clade, now known to be paraphyletic.

<span class="mw-page-title-main">Ichneumonidae</span> Family of wasps

The Ichneumonidae, also known as ichneumon wasps, ichneumonid wasps, ichneumonids, or Darwin wasps, are a family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 25,000 species described as of 2016. However, this likely represents less than a quarter of their true richness as reliable estimates are lacking, along with much of the most basic knowledge about their ecology, distribution, and evolution. It is estimated that there are more species in this family than there are species of birds and mammals combined. Ichneumonid wasps, with very few exceptions, attack the immature stages of holometabolous insects and spiders, eventually killing their hosts. They thus fulfill an important role as regulators of insect populations, both in natural and semi-natural systems, making them promising agents for biological control.

<i>Polydnaviriformidae</i> Family of viruses

Polydnaviriformidae ( PDV) is a family of insect viriforms; members are known as polydnaviruses. There are two genera in the family: bracoform and Ichnoviriform. Polydnaviruses form a symbiotic relationship with parasitoid wasps. Ichnoviriforms (IV) occur in Ichneumonid wasps and Bracoviriforms (BV) in Braconid wasps. The larvae of wasps in both of those groups are themselves parasitic on Lepidoptera, and the polydnaviruses are important in circumventing the immune response of their parasitized hosts. Little or no sequence homology exists between BV and IV, suggesting that the two genera have been evolving independently for a long time.

<span class="mw-page-title-main">Parasitoid wasp</span> Group of wasps

Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causing the death of these hosts. Different species specialise in hosts from different insect orders, most often Lepidoptera, though some select beetles, flies, or bugs; the spider wasps (Pompilidae) exclusively attack spiders.

<span class="mw-page-title-main">Microgastrinae</span> Subfamily of wasps

Microgastrinae is a subfamily of braconid wasps, encompassing almost 3,000 described species, with an estimated 30,000–50,000 total species. This makes it one of the richest subfamilies with the most species of parasitoid wasps.

<span class="mw-page-title-main">Agathidinae</span> Subfamily of wasps

Agathidinae is a subfamily of braconid parasitoid wasps. Some species have been used in biological control programs.

<span class="mw-page-title-main">Doryctinae</span> Subfamily of wasps

The Doryctinae or doryctine wasps are a large subfamily of braconid parasitic wasps (Braconidae). Numerous genera and species formerly unknown to science are being described every year. This subfamily is presumably part of a clade containing otherwise any or all of the Alysiinae, Braconinae, Gnamptodontinae, Opiinae and Ypsistocerinae, and might be most closely related to the last one of these. Whether the Rogadinae are also part of this group is not known.

<i>Cotesia glomerata</i> Species of wasp

Cotesia glomerata, the white butterfly parasite, is a small parasitoid wasp belonging to family Braconidae. It was described by Carl Linnaeus in his 1758 publication 10th edition of Systema Naturae.

<i>Cotesia</i> Genus of wasps

Cotesia is a genus of braconid wasps first described by Peter Cameron in 1891. Some species parasitize caterpillars of species considered pests, and are used as biocontrol agents. Cotesia congregata parasitizes the tomato and the tobacco hornworms. C. glomerata and C. rubecula feed on the cabbage white and other white butterfly caterpillars. C. gonopterygis and C. risilis are host-specific and parasitize the common brimstone.

Bracovirus is a genus of viruses, in the family Polydnaviridae. Bracoviruses are an ancient symbiotic virus contained in parasitic braconid wasps that evolved off of the nudivirus about 190 million years ago and has been evolving at least 100 million years. It is one of two genera belonging to the Polydnaviridae family, Ichnovirus being the other genus. There are 32 species in this genus.

<i>Dinotrema</i> Genus of wasps

Dinotrema is a genus of wasps in the family Braconidae. Species are amongst the largest parasitoid wasps in the tribe Alysiini (Alysiinae). There are approximately 350 species described around worldwide.

<span class="mw-page-title-main">Rogadinae</span> Subfamily of wasps

The Rogadinae are a large subfamily of braconid parasitoid wasps. Several Rogadinae species parasitize pest caterpillars and are important for naturally occurring biological control.

<span class="mw-page-title-main">Cenocoeliinae</span> Subfamily of wasps

The Cenocoeliinae are a subfamily of braconid parasitoid wasps.

The Cardiochilinae are a subfamily of braconid parasitoid wasps. This subfamily has been treated as a tribe of Microgastrinae in the past. Some species including Toxoneuron nigriceps have been used in biocontrol programs.

The Homolobinae are a subfamily of braconid parasitoid wasps. The subfamily Charmontinae was previously included within Homolobinae as the tribe Charmontini.

The Acampsohelconinae are a subfamily of braconid parasitoid wasps. Extant members of this subfamily were previously included in the Helconinae, Blacinae, or Sigalphinae. The four genera included are †Acampsohelcon, Afrocampsis, Canalicephalus, and Urosigalphus.

The Amicrocentrinae are a subfamily of braconid parasitoid wasps. Members of this subfamily were previously included in the Macrocentrinae.

Hormius is a genus of parasitoid wasp in the subfamily Hormiinae first described by Christian Gottfried Daniel Nees von Esenbeck in 1818.

References

  1. 1 2 Jones, Owen R.; Purvis, Andy; Baumgart, Eligiusz; Quicke, Donald L.J. (2009). "Using taxonomic revision data to estimate the geographic and taxonomic distribution of undescribed species richness in the Braconidae (Hymenoptera: Ichneumonoidea)". Insect Conservation and Diversity. 2 (3): 204–212. doi: 10.1111/j.1752-4598.2009.00057.x . S2CID   86775770.
  2. ↑ Beyarslan, A. and M. Aydogdu. (2013). Additions to the rare species of Braconidae fauna (Hymenoptera: Braconidae) from Turkey. Mun Ent Zool 8(1) 369-74.
  3. ↑ Wharton, R. M. (2000). "Can braconid classification be restructured to facilitate portrayal of relationships?". In Austin, A. D.; Dowton, M. (eds.). Hymenoptera: evolution, biodiversity, and biological control. 4th. Vol. International Hymenopterists Conference. Collingwood, Victoria, Australia: Commonwealth Scientific and Industrial Research Organisation (CSIRO). pp. 143–153. ISBN   978-0-643-06610-6.
  4. ↑ Chen, Xue-xin; van Achterberg, Cornelis (January 2019). "Systematics, Phylogeny, and Evolution of Braconid Wasps: 30 Years of Progress". Annual Review of Entomology. 64: 335–358. doi:10.1146/annurev-ento-011118-111856. PMID   30332295. S2CID   52986629.
  5. ↑ Shaw, Scott; Edgerly-Rooks, Janice (1986). "A new braconid genus (Hymenoptera) parasitizing webspinners (Embiidina) in Trinidad". Psyche: A Journal of Entomology. 92 (4): 505–511. doi: 10.1155/1985/54285 . ISSN   0033-2615 – via Santa Clara University Scholar Commons.
  6. ↑ Centrella, Mary L.; Shaw, Scott R. (June 2010). "A new species of phytophagous braconid Allorhogas minimus (Hymenoptera: Braconidae: Doryctinae) reared from fruit galls on Miconia longifolia (Melastomataceae) in Costa Rica". International Journal of Tropical Insect Science. 30 (2): 101–107. doi:10.1017/S1742758410000147. ISSN   1742-7592. S2CID   85057439.
  7. ↑ Quicke, Donald L. J.; Huddleston, Tom (December 1989). "The Australian braconid wasp subfamily Mesostoinae (Hymenoptera: Braconidae) with the description of a new species of Mesostoa". Journal of Natural History. 23 (6): 1309–1317. doi:10.1080/00222938900770691. ISSN   0022-2933.
  8. ↑ Mahr, S. (February 1998). "Know Your Friends: Aphidius Wasps". Midwest Biological Control News Online. University of Wisconsin–Madison. Archived from the original on 2 January 2013. Retrieved 25 March 2013.
  9. ↑ Conling, D.E.; Graham, D.Y.; Hastings, H. (1 March 1988). "Notes on the natural host surveys and laboratory rearing of Goniozus natalensis Gordh (Hymenoptera: Bethylidae), a parasitoid of Eldana saccharina Walker (Lepidoptera: Pyralidae) larvae from Cyperus papyrus L. in Southern Africa". Journal of the Entomological Society of Southern Africa. 51 (1): 115–127. hdl:10520/AJA00128789_2538.
  10. ↑ Edmunds, Malcolm (1976-03-01). "Larval mortality and population regulation in the butterfly Danaus chrysippus in Ghana". Zoological Journal of the Linnean Society. 58 (2): 129–145. doi:10.1111/j.1096-3642.1976.tb00823.x. ISSN   0024-4082.
  11. ↑ Gray, Betty. "Beneficial insects in the garden: #04 Braconid Wasp on Hornworm (Cotesia congregatus)". aggie-horticulture.tamu.edu. Retrieved 2017-11-14.
  12. ↑ Núñez-Bueno, Ligia (1982). Trybliographa daci Weld (Hymenoptera: Cynipidae): Biology and aspects of the relationship with its host Anastrepha suspensa (Loew) (Diptera: Tephritidae) (PhD thesis). University of Florida. OCLC   9311697.
  13. ↑ Piper, R. (2007), Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals, Greenwood Press.
  14. ↑ BéZier, Annie; Annaheim, Marc; HerbinièRe, Juline; Wetterwald, Christoph; Gyapay, Gabor; Bernard-Samain, Sylvie; Wincker, Patrick; Roditi, Isabel; Heller, Manfred; Belghazi, Maya; Pfister-Wilhem, Rita; Periquet, Georges; Dupuy, Catherine; Huguet, Elisabeth; Volkoff, Anne-Nathalie; Lanzrein, Beatrice; Drezen, Jean-Michel (2009). "Polydnaviruses of Braconid Wasps Derive from an Ancestral Nudivirus". Science. 323 (5916): 926–930. Bibcode:2009Sci...323..926B. doi:10.1126/science.1166788. PMID   19213916. S2CID   6538583.
  15. ↑ Sharkey, M. J. Family Braconidae. pp. 362–94 In: Goulet, H. and J. Huber (eds.) Hymenoptera of the World, an Identification Guide to Families. Agriculture Canada Research Branch Monograph No. 1894E. 1993.
  16. ↑ Hall, M. Scientists recruit wasps for war on terror. USA Today December 26, 2005. Accessed June 19, 2012.
  17. ↑ Cockroaches and radiation. ABC Science. February 23, 2006.

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