The Braconidae are a family of

parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...

wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...

s. After the closely related

Ichneumonidae The Ichneumonidae, also known as the ichneumon wasps, Darwin wasps, or ichneumonids, 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 cur ...

, braconids make up the second-largest family in the order

Hymenoptera Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typic ...

, with about 17,000 recognized species and many thousands more undescribed. One analysis estimated a total between 30,000 and 50,000, and another provided a narrower estimate between 42,000 and 43,000 species.

Classification

The Braconidae are currently divided into about 47 subfamilies and over 1000 genera, which include '' Aerophilus'', ''

Aleiodes ''Aleiodes'' (Greek "not", "smooth", "appearance") is a genus of the family Braconidae of parasitoid wasps described by Constantin Wesmael in 1838. They are also known as mummy-wasps. The female attacks caterpillars of various species, inc ...

'', ''

Apanteles ''Apanteles'' is a very large genus of braconid wasps, containing more than 600 described species found worldwide. There are no native species in New Zealand, and none have been recorded in the high arctic. See also * List of Apanteles species ...

'', '' Asobara'', '' Bracon'', '' Cenocoelius'', '' Chaenusa'', '' Chorebus'', ''

Cotesia ''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 toba ...

'', '' Dacnusa'', '' Diachasma'', ''

Microgaster ''Microgaster'' is a genus of braconid wasps in the family Braconidae 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 ...

'', '' Opius'', '' Parapanteles'', '' Phaenocarpa'', ''

Spathius ''Spathius'' is a genus of doryctine wasps. The larvae of this genus of wasps feed on beetle larvae. They act as controllers of the parasitic ''Hylurgopinus rufipes ''Hylurgopinus rufipes'', known as the native elm bark beetle, is a species ...

'', 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

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...

s that diverged early in the evolution of braconids. Cyclostomes are

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...

whereas noncyclostomes can be divided formally into microgastroids, sigalphoids, helconoids, and euphoroids.

Subfamilies

Acampsohelconinae 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'', ' ...

Agathidinae Agathidinae is a subfamily of braconid parasitoid wasps. Some species have been used in biological control programs. Description and distribution Agathidines are among the larger and more colorful braconids. Diurnal members of this subfamil ...

* Alysiinae * Amicrocentrinae *

Aphidiinae The Aphidiinae are a subfamily of tiny parasitoid wasps that use aphids as their hosts. Several species have been used in biological control programs of various aphids. Biology and distribution Aphidiines are koinobiont endoparasitoids of adu ...

* Apozyginae * Betylobraconinae * Blacinae * Brachistinae * Braconinae *

Cardiochilinae 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. Description and d ...

* Cenocoeliinae * Charmontinae *

Cheloninae The Cheloninae are a cosmopolitan subfamily of braconid parasitoid wasps. Description Most Cheloninae are small and uniformly colored. They have a characteristic metasomal carapace formed from the fusion of the first three tergites. Biology ...

* Dirrhopinae *

Doryctinae 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 othe ...

* Ecnomiinae *

Euphorinae The Euphorinae are a large subfamily of Braconidae parasitoid wasps. Some species have been used for biological pest control. They are sister group to the Meteorinae. Description and distribution Euphorines are small, usually dark colored w ...

* Exothecinae * Gnamptodontinae * Helconinae * Histeromerinae * Homolobinae * Hormiinae * Ichneutinae * Khoikhoiiinae * Lysiterminae * Macrocentrinae * Masoninae * Maxfischeriinae * Mendesellinae * Mesostoinae * Meteorideinae * Meteorinae * Microgastrinae * Microtypinae * Miracinae * Neoneurinae *

Opiinae The Opiinae are a subfamily of braconid parasitoid wasps with over 1300 described species. Several species have been used in biocontrol programs against fruit flies and Agromyzidae flies. They are closely related to the Alysiinae. Description a ...

* Orgilinae * Pambolinae * Pselaphaninae *† Protorhyssalinae * Rhysipolinae * Rhyssalinae * Rogadinae *† Seneciobraconinae * Sigalphinae * Telengaiinae * Trachypetinae * Vaepellinae * Xiphozelinae * Ypsistocerinae Aleiodes shakirae head

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 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 no ...

(

), which usually have two. Wing venation patterns are otherwise highly variable. The antennae typically have 16 segments or more; the

trochanter A trochanter is a tubercle of the femur near its joint with the hip bone. In humans and most mammals, the trochanters serve as important muscle attachment sites. Humans are known to have three trochanters, though the anatomic "normal" includes ...

s have two segments. Females often have long

ovipositor The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...

s, 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

microlepidoptera Microlepidoptera (micromoths) is an artificial (i.e., unranked and not monophyletic) grouping of moth families, commonly known as the 'smaller moths' (micro, Lepidoptera). These generally have wingspans of under 20 mm, and are thus harder to ...

ns, 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 Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describe ...

larvae in tree trunks.

Life history

The larvae of most braconids are internal or external primary parasitoids of other

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...

s, especially the larval stages of

Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...

, and

Lepidoptera Lepidoptera ( ) is an order of insects that includes butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 families and 46 superfamilies, 10 percent of the total described speci ...

, but also some hemimetabolous insects such as

aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...

Heteroptera The Heteroptera are a group of about 40,000 species of insects in the order Hemiptera. They are sometimes called "true bugs", though that name more commonly refers to the Hemiptera as a whole. "Typical bugs" might be used as a more unequivocal al ...

, or

Embiidina The order Embioptera, commonly known as webspinners or footspinners, are a small group of mostly tropical and subtropical insects, classified under the subclass Pterygota. The order has also been called Embiodea or Embiidina. More than 400 sp ...

. Most species kill their hosts, though some cause the hosts to become

sterile Sterile or sterility may refer to: *Asepsis Asepsis is the state of being free from disease-causing micro-organisms (such as pathogenic bacteria, viruses, pathogenic fungi, and parasites). There are two categories of asepsis: medical and surgi ...

and less active. Parasitoidy on adult insects (particularly on Hemiptera and Coleoptera) also occurs. Members of two subfamilies, the Mesostoinae and

are known to form

gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...

s on plants. Braconids are often used as

biological pest control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also i ...

agents, especially against aphids.

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 ''Ostrinia furnacalis'' is a species of moth in the family Crambidae, the grass moths. It was described by Achille Guenée in 1854 and is known by the common name Asian corn borer since this species is found in Asia and feeds mainly on corn cro ...

'' (the Asian corn borer, a lepidopteran moth known for being a pest of

maize Maize ( ; ''Zea mays'' subsp. ''mays'', from es, maíz after tnq, mahiz), also known as corn (North American English, North American and Australian English), is a cereal grain first domesticated by indigenous peoples of Mexico, indigenous ...

in East Asia), the African sugarcane borer (a moth commonly found in sub-Saharan Africa), the butterfly ''

Danaus chrysippus ''Danaus chrysippus'', also known as the plain tiger, African queen, or African monarch, is a medium-sized butterfly widespread in Asia, Australia and Africa. It belongs to the Danainae subfamily of the brush-footed butterfly family Nymphalidae. ...

'' in

Ghana Ghana (; tw, Gaana, ee, Gana), officially the Republic of Ghana, is a country in West Africa. It abuts the Gulf of Guinea and the Atlantic Ocean to the south, sharing borders with Ivory Coast in the west, Burkina Faso in the north, and Tog ...

, and ''

Liriomyza trifolii ''Liriomyza trifolii'', known generally as the American serpentine leafminer or celery leafminer, is a species of leaf miner fly in the family Agromyzidae. ''L. trifolii'' is a damaging pest, as it consumes and destroys produce and other plant p ...

'' (the American serpentine leafminer) and ''

Manduca quinquemaculata ''Manduca quinquemaculata'', the five-spotted hawkmoth, is a brown and gray hawk moth of the family Sphingidae. The caterpillar, often referred to as the tomato hornworm, can be a major pest in gardens; they get their name from a dark projection ...

'' (the tomato hornworm) in North America. Braconids often will prey on fruit fly larvae like ''

Anastrepha suspensa ''Anastrepha suspensa'', known as the Caribbean fruit fly, the Greater Antillean fruit fly, guava fruit fly, or the Caribfly, is a species of tephritid fruit fly. As the names suggest, these flies feed on and develop in a variety of fruits, prim ...

'' as well.

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. 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". 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 Immunosuppression is a reduction of the activation or efficacy of the immune system. Some portions of the immune system itself have immunosuppressive effects on other parts of the immune system, and immunosuppression may occur as an adverse reacti ...

, a high level of parasitoid-host specificity is not surprising.

Evolutionary history

The family seems to date from early

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...

(provided that '' Eobracon'' is properly assigned to this family). It underwent extensive diversification from mid or late

to early

Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configu ...

, correlating with the radiation of flowering plants and associated insect

herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...

s, the main hosts of braconids.

Differentiation from Ichneumonidae

Braconids are distinguished from their

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

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.

Other characteristics

The species '' Microplitis croceipes'' possesses an extremely accurate sense of smell and can be trained for use in

narcotics The term narcotic (, from ancient Greek ναρκῶ ''narkō'', "to make numb") originally referred medically to any psychoactive compound with numbing or paralyzing properties. In the United States, it has since become associated with opiate ...

and explosives detection. At least some braconids appear to be very resistant to

ionizing radiation Ionizing radiation (or ionising radiation), including nuclear radiation, consists of subatomic particles or electromagnetic waves that have sufficient energy to ionize atoms or molecules by detaching electrons from them. Some particles can travel ...

. While a dose of 400 to 1000

rads RAD or Rad may refer to: People * Robert Anthony Rad Dougall (born 1951), South African former racing driver * Rad Hourani, Canadian fashion designer and artist * Nickname of Leonardus Rad Kortenhorst (1886–1963), Dutch politician * Radley ...

can kill an average human, a dose of 180,000 rads was required to kill a braconid of genus '' Habrobracon'' in an experiment.Cockroaches and radiation.
ABC Science. February 23, 2006.

Gallery

File: Cotesia.melanoscela.-.lindsey.jpg, '' Cotesia melanoscela '' File: Aleiodes indiscretus wasp parasitizing gypsy moth caterpillar.jpg, '' Aleiodes indiscretus'' ovipositing in its host, a gypsy moth caterpillar File: Braconid parasitoid wasp Apanteles sp eggs & Lime Butterfly (Papilio demoleus) cat W IMG 2862.jpg, ''

'' sp. cocoons on ''

Papilio demoleus ''Papilio demoleus'' is a common and widespread swallowtail butterfly. The butterfly is also known as the lime butterfly, lemon butterfly, lime swallowtail, and chequered swallowtail. These common names refer to their host plants, which are usu ...

'' File: Braconidae - Microgastrinae wasp empty cocoons.JPG, Braconid wasp empty cocoons File:Aphis nerii parasitized.jpg, '' Aphis nerii'' parasitized by

, possibly '' Lysiphlebus''. File:Cotesia adult.jpg, ''

'' adult

References

External links

Braconidae.
Tree of Life.
Braconidae.
Encyclopedia of Life. * Sharkey, M. J. (2004)
Synopsis of the Agathidinae (Hymenoptera: Braconidae) of America north of Mexico.
''Proceedings of the Russian Entomological Society'' 75(1), 134–52. * Ghahari, H., et al
Bibliography of the family Braconidae (Hymenoptera: Ichneumonoidea) (1964–2003).
NNM Technical Bulletin 8. 2006. ISSN 1387-0211 * van Achterberg, C. and C. O'Toole. (1993)
Annotated catalogue of the types of Braconidae (Hymenoptera) in the Oxford University Museum.
''

Zoologische Verhandelingen ''Zoologische Verhandelingen'' was a Dutch scientific journal covering research in zoology. It was published between 1948 and 2002 by the Rijksmuseum van Natuurlijke Historie in Leiden Leiden (; in English and archaic Dutch also Leyden) ...

'' 287(1) 1–43. * Species profiles from the

University of Florida The University of Florida (Florida or UF) is a public land-grant research university in Gainesville, Florida. It is a senior member of the State University System of Florida, traces its origins to 1853, and has operated continuously on its ...

Institute of Food and Agricultural Sciences The University of Florida Institute of Food and Agricultural Sciences (UF/IFAS) is a teaching, research and Extension scientific organization focused on agriculture and natural resources. It is a partnership of federal, state, and county governmen ...

: *
''Cotesia marginiventris''
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