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 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 ...

, 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 typi ...

, 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, in ...

'', '' Apanteles'', '' Asobara'', '' Bracon'', '' Cenocoelius'', '' Chaenusa'', '' Chorebus'', '' Cotesia'', '' Dacnusa'', '' Diachasma'', ''

Dimeris ''Dimeris'' is a genus of insects belonging to 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 17,0 ...

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

Spathius ''Spathius'' is a genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV class ...

'', and ''

Syntretus ''Syntretus'' is a genus 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 ...

.'' 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 In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

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

monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

whereas noncyclostomes can be divided formally into microgastroids, sigalphoids, helconoids, and euphoroids. The rare Australian subfamily

Trachypetinae Trachypetinae is a subfamily of the wasp family Braconidae, comprising three rare Australian genera of large parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the ...

was raised to family rank in 2020 (as "Trachypetidae"), but was reduced back to a subfamily in 2022.

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 Agathidines are among the larger braconids, and diurnal members of this subfamily are often brightly patterned. ...

Alysiinae The Alysiinae are a subfamily of braconid parasitoid wasps with over 1000 described species. Several species have been used in Biological pest control, biocontrol programs. They are closely related to the Opiinae. Description and distribution ...

* Amicrocentrinae * Aphidiinae * Apozyginae * Betylobraconinae * Brachistinae * Braconinae * Cardiochilinae * Cenocoeliinae * Charmontinae * Cheloninae * Dirrhopinae * Doryctinae * Ecnomiinae * Euphorinae * Exothecinae * Gnamptodontinae * Helconinae * Histeromerinae * Homolobinae * Hormiinae * Ichneutinae * Khoikhoiiinae * Lysiterminae * Macrocentrinae * Masoninae * Maxfischeriinae * Mendesellinae * Mesostoinae * Meteorideinae * Meteorinae *

Microgastrinae 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. Genera These ...

* Microtypinae * Miracinae * Neoneurinae * Opiinae * Orgilinae * Pambolinae * Pselaphaninae *† Protorhyssalinae * Rhysipolinae * Rhyssalinae * Rogadinae *† Seneciobraconinae * Sigalphinae * Telengaiinae *

* 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 not ...

(

), which usually have two. Wing

venation Venation may refer to: * Venation (botany), the arrangement of veins in leaves * Wing venation, the arrangement of veins in insect wings See also * * Vernation Vernation or leafing is the formation of new leaves or fronds. In plant anatomy, it ...

patterns are otherwise highly variable. The antennae typically have 16 segments or more; the trochanters 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 typica ...

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, so are harder to iden ...

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 Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...

larvae in tree trunks.

Life history

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

insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...

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 advance ...

, and

Lepidoptera Lepidoptera ( ) or lepidopterans is an order (biology), order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organ ...

, but also some hemimetabolous insects such as

aphid Aphids are small sap-sucking insects in the Taxonomic rank, family Aphididae. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white Eriosomatinae, woolly ...

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

sterile Sterile or sterility may refer to: *Asepsis, a state of being free from biological contaminants * Sterile (archaeology), a sediment deposit which contains no evidence of human activity *Sterilization (microbiology), any process that eliminates or ...

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

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. Plant galls are abnormal outgrowths of plant tissues, similar to benign tumors or war ...

s on plants. Braconids are often used as

biological pest control Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or ot ...

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

maize Maize (; ''Zea mays''), also known as corn in North American English, is a tall stout grass that produces cereal grain. It was domesticated by indigenous peoples in southern Mexico about 9,000 years ago from wild teosinte. Native American ...

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

Ghana Ghana, officially the Republic of Ghana, is a country in West Africa. It is situated along the Gulf of Guinea and the Atlantic Ocean to the south, and shares borders with Côte d’Ivoire to the west, Burkina Faso to the north, and Togo to t ...

, and '' Liriomyza trifolii'' (the American serpentine leafminer) and '' Manduca quinquemaculata'' (the tomato hornworm) in North America. Braconids often will prey on fruit fly larvae like '' Anastrepha suspensa'' 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 react ...

, 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 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...

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

to early

Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...

, correlating with the radiation of flowering plants and associated insect

herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...

s, the main hosts of braconids. 73 individual Braconid

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...

s of the

Königsberg Königsberg (; ; ; ; ; ; , ) is the historic Germany, German and Prussian name of the city now called Kaliningrad, Russia. The city was founded in 1255 on the site of the small Old Prussians, Old Prussian settlement ''Twangste'' by the Teuton ...

Amber Collection were destroyed during

World War II World War II or the Second World War (1 September 1939 – 2 September 1945) was a World war, global conflict between two coalitions: the Allies of World War II, Allies and the Axis powers. World War II by country, Nearly all of the wo ...

as they were "stored in a salt-mine about 40 km from

Göttingen Göttingen (, ; ; ) is a college town, university city in Lower Saxony, central Germany, the Capital (political), capital of Göttingen (district), the eponymous district. The River Leine runs through it. According to the 2022 German census, t ...

" with petrol. They were set on fire by the

Allies An alliance is a relationship among people, groups, or states that have joined together for mutual benefit or to achieve some common purpose, whether or not an explicit agreement has been worked out among them. Members of an alliance are calle ...

. This has arguably slowed the study of the

evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...

of the family.

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'', "I make numb") originally referred medically to any psychoactive compound with numbing or paralyzing properties. In the United States, it has since become associated with opiates ...

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

ionizing radiation Ionizing (ionising) radiation, including Radioactive decay, nuclear radiation, consists of subatomic particles or electromagnetic waves that have enough energy per individual photon or particle to ionization, ionize atoms or molecules by detaching ...

. 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.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 spongy moth caterpillar File: Braconid parasitoid wasp Apanteles sp eggs & Lime Butterfly (Papilio demoleus) cat W IMG 2862.jpg, '' Apanteles'' sp. cocoons on '' Papilio demoleus'' File: Braconidae - Microgastrinae wasp empty cocoons.JPG, Braconid wasp empty cocoons File:Aphis nerii parasitized.jpg, ''

Aphis nerii ''Aphis nerii'' is an aphid of the Family (biology), family Aphididae. Its common names include oleander aphid, milkweed aphid, sweet pepper aphid, and nerium aphid. Distribution The oleander aphid is widespread in regions with tropical and Med ...

'' parasitized by Aphidiinae, possibly '' Lysiphlebus''. File:Cotesia adult.jpg, '' Cotesia'' 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.
''

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

University of Florida The University of Florida (Florida or UF) is a public university, public land-grant university, land-grant research university in Gainesville, Florida, United States. It is a senior member of the State University System of Florida and a preem ...

Institute of Food and Agricultural Sciences An institute is an organizational body created for a certain purpose. They are often research organisations (research institutes) created to do research on specific topics, or can also be a professional body. In some countries, institutes ca ...

: *
''Cotesia marginiventris''
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