The Doryctinae or doryctine wasps are a large

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...

parasitoid wasp Parasitoid wasps are a large group of hymenopteran Superfamily (zoology), 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, ...

s within 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 17,000 recognized species and many thousands more undescribed. One analysis ...

Description and identification

Doryctine wasps are found across almost the entire size range of Braconidae, from 1 to 25 mm. In the small species the head tends to be relatively large and the body, while slender, remarkably elongated. Doryctines tend to be small-winged, with many having very much reduced wings. Numerous species in this family are unable to fly or even lack wings entirely. They have a characteristic row of stout spines running lengthwise along the foreleg

tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...

, and a

cyclostome Cyclostome is a biological term (from the Greek for "round mouth") used in a few different senses: * for the taxon Cyclostomi, which comprises the extant jawless fishes: the hagfish (Myxini) and the lampreys (Petromyzontidae). This was thought for ...

depression above the

mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...

Distribution

Doryctinae have a worldwide distribution except for the

polar region The polar regions, also called the frigid zones or polar zones, of Earth are Earth's polar ice caps, the regions of the planet that surround its geographical poles (the North and South Poles), lying within the polar circles. These high latitu ...

Biology

Some 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 similar to the

Mesostoinae The Mesostoinae is a subfamily of Braconidae, braconid wasps with a Gondwanan distribution. The members of this family display sexual dimorphism, males are Brachyptery, brachypterous, which means that they have reduced, non-functional wings. Bi ...

(a small subfamily

endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

). Some species in the genus '' Allorhogas'' feed on seeds. The large majority of doryctines are

idiobiont 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 ...

ectoparasitoids of the larvae of wood-boring

beetle 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 ...

s – such as

jewel beetle Buprestidae is a family of beetles known as jewel beetles or metallic wood-boring beetles because of their glossy iridescent colors. Larvae of this family are known as flatheaded borers. The family is among the largest of the beetles, with some ...

s (Buprestidae). Some species parasitize

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 ...

n or

symphyta Sawflies are wasp-like insects that are in the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plants ...

n larvae. The host is paralyzed by venom injected through the female's

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 ...

before an egg is laid. The spines present on the foreleg of the adult enable it to escape from the narrow wooden tunnels of the hosts.

Biological control

Doryctines have been used to control pests in Europe, Australia, and North America. Several doryctine wasps are of economic importance as

biocontrol Biological control or biocontrol is a method of pest control, controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or phytopathology, plants by bioeffector, using other organisms. It relies o ...

agents in Australia against

eucalyptus ''Eucalyptus'' () is a genus of more than 700 species of flowering plants in the family Myrtaceae. Most species of ''Eucalyptus'' are trees, often Mallee (habit), mallees, and a few are shrubs. Along with several other genera in the tribe Eucalyp ...

pests. The species ''

Spathius agrili ''Spathius agrili'' is a parasitic non-stinging wasp of family Braconidae which is native to North Asia. It is a parasitoid of the emerald ash borer (''Agrilus planipennis'' Fairmaire), an invasive species which has destroyed tens of millions of ...

'' has been introduced to the United States from China in an effort to control the

emerald ash borer The emerald ash borer (''Agrilus planipennis''), also known by the abbreviation EAB, is a green buprestid or jewel beetle native to north-eastern Asia that feeds on ash trees, ash species (''Fraxinus'' spp.). Females lay eggs in bark crevices o ...

Taxonomy and phylogeny

Doryctinae is considered to be part of the cyclostome group, a

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 ...

of subfamilies including

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 ...

, Betylobraconinae,

Braconinae The Braconinae are a large subfamily of Braconidae, braconid parasitoid wasps with more than 2,000 described species.Wharton, Robert A.; Marsh, Paul M.; Sharkey, Michael J. (1997)''Manual of the New World Genera of the Family Braconidae (Hymenop ...

, Gnamptodontinae, Exothecinae, Hormiinae, Lysiterminae,

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 ...

, Pambolinae, Rogadinae, Rhysipolinae, and Rhyssalinae. 16 tribes of Doryctinae have been designated. The former subfamily Ypsistocerinae has been subsumed under Doryctinae as one of these tribes, Ypsistocerini. Doryctinae comprises over 2000 species in 198 genera, with new taxa continuing to be described. 182 of the

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

placed in the Doryctinae include:

Tribe Binareini

* '' Acanthodoryctes'' Turner, 1918 * '' Antidoryctes'' Belokobylskij & Quicke, 2000 * '' Binarea'' Brullé, 1846 * '' Dooliba'' Belokobylskij, Iqbal & Austin, 2004 * '' Liobracon'' Szépligeti, 1901

Tribe Doryctini

* '' Acanthorhogas'' Szepligeti, 1906 * †'' Anacanthobracon'' Brues, 1939 * '' Aphelopsia'' Marsh, 1993 * '' Asiaontsira'' Belokobylskij, Tang, & Chen, 2013 * '' Bracodoryctes'' Belokobylskij & Quicke, 2000 * '' Bracocesa'' Koçak & Kemal, 2008 * '' Bulbonervus'' Shenefelt, 1969 * '' Caenophanes'' Foerster, 1862 * '' Caingangia'' Marsh, 1993 * '' Cecidospathius'' Kieffer & Jörgensen, 1910 * '' Ceratodoryctes'' Belokobylskij, Iqbal & Austin, 2004 * '' Chelonodoryctes'' Belokobylskij & Quicke, 2000 * ''

Coiba Coiba () is the largest island in Central America, with an area of , off the Pacific coast of the Panamanian province of Veraguas. It is part of the Montijo District of that province. History Coiba separated from continental Panama between 1 ...

'' Marsh, 1993 * '' Cryptodoryctes'' Belokobylskij & Quicke, 2000 * '' Cryptontsira'' Belokobylskij, 2008 * '' Cyphodoryctes'' Marsh, 1993 * '' Dendrosoter'' Wesmael, 1838 * '' Dendrosotinus'' Telenga, 1941 * '' Dicarinoryctes'' Braet & van Achterberg, 2001 * '' Donquickeia'' Marsh, 1997 * '' Doryctes'' Haliday, 1836 * '' Doryctodes'' Hellén, 1927 * '' Doryctophasmus'' Enderlein, 1912 * '' Doryctopsis'' Belokobylskij, Iqbal & Austin, 2004 * '' Echinodoryctes'' Belokobylskij, Iqbal & Austin, 2004 * '' Eodendrus'' Belokobylskij, 2005 * '' Esterella'' Pagliano & Scaramozzino, 1990 * '' Euscelinus'' Westwood, 1882 * '' Guaygata'' Marsh, 1993 * '' Gymnobracon'' Szépligeti, 1902 * '' Halycaea'' Cameron, 1903 * '' Heerz'' Marsh, 1993 * '' Histeromeroides'' Marsh, 1993 * '' Hybodoryctes'' Szépligeti, 1906 * '' Hypodoryctes'' Kokujev, 1900 * '' Janzenia'' Marsh, 1993 * '' Johnsonius'' Marsh, 1993 * '' Kauriphanes'' Belokobylskij, Ceccarelli, Zaldívar-Riverón, 2012 * '' Lamquetia'' Braet, Barbalho & van Achterberg, 2003 * '' Latana'' Cameron, 1906 * '' Megaloproctus'' Schulz, 1906 * '' Neostaphius'' Braet, Barbalhoa & van Achterberg, 2003 * '' Neurocrassus'' Šnoflák, 1945 * '' Odontodoryctes'' Granger, 1949 * '' Ontsira'' Cameron, 1900 * '' Ontsirospathius'' Belokobylskij, Iqbal & Austin, 2004 * '' Osmophila'' Szépligeti, 1902 * †'' Palaeorhyssalus'' Brues, 1933 * '' Pedinotus'' Szépligeti, 1902 * '' Pirramurra'' Belokobylskij, Iqbal & Austin, 2004 * '' Priosphys'' Enderlein, 1920 * †'' Promonolexis'' Brues, 1933 * '' Pseudodoryctes'' Szépligeti, 1915 * '' Rhoptrocentrus'' Marshall, 1897 * '' Sharkeyella'' Marsh, 1993 * '' Shawius'' Marsh, 1993 * '' Sinaodoryctes'' Chen & Shi, 2004 * '' Sonanus'' Belokobylskij & Konishi, 2001 * '' Tarasco'' Marsh, 1993 * '' Verae'' Marsh, 1993 * '' Waitaca'' Marsh, 1993 * '' Whartonius'' Marsh, 1993 * '' Whitfieldiellus'' Marsh, 1997

Tribe Ecphylini

* '' Achterbergia'' Marsh, 1993 * '' Aivalykus'' Nixon, 1938 * '' Bohartiellus'' Marsh, 1983 * '' Ecphylus'' Foerster, 1862 * '' Sycosoter'' Picard & Lichtenstein, 1917

Tribe Evaniodini

* '' Evaniodes'' Szépligeti, 1901

Tribe Hecabolini

* '' Allorhogas'' Gahan, 1912 * '' Callihormius'' Ashmead, 1900 * '' Curtisella'' Spinola, 1851 * '' Doryctinus'' Roman, 1910 (= ''Acrophasmus'') * '' Glyptocolastes'' Ashmead, 1900 * '' Hecabolodes'' Wilkinson 1929 * '' Hecabolomorpha'' Belokobylskij & Chen, 2006 * '' Hecabolus'' Curtis, 1834 * '' Hemidoryctes'' Belokobylskij, 1993 (= ''Atopodoryctes'') * '' Leluthia'' Cameron, 1887 * '' Monolexis'' Foerster, 1862 * '' Pambolidea'' Ashmead, 1900 * '' Parallorhogas'' Marsh, 1993 * '' Pareucorystes'' Tobias, 1961 * '' Percnobraconoides'' Marsh, 1989 * '' Polystenus'' Foerster, 1862 * '' Semirhytus'' Szépligeti, 1902 * '' Spathiostenus'' Belokobylskij, 1993 * '' Stenocorse'' Marsh, 1968 * '' Terate'' Nixon, 1943 * '' Tripteria'' Enderlein, 1912

Tribe Heterospilini

* '' Asiaheterospilus'' Belokobylskij & Konishi, 2001 * ''

Canchim The Canchim is a Brazilian breed of beef cattle. Its origin is Central Brazil and it was developed by crossing the Indian Zebu cattle (''Bos taurus indicus'') and the European (''Bos taurus taurus'') Charolais. Compared to the very productiv ...

'' Barbalho & Penteado-Dias, 1999 * '' Dapsilitas'' Braet, Barbalhoa & van Achterberg, 2003 * '' Ecphylopsis'' Ashmead, 1900 * '' Heterospilus'' Haliday, 1836 * '' Jataiella'' Barbalho & Penteado-Dias, 1999 * '' Neoheterospilus'' Belokobylskij, 2006 * '' Paraheterospilus'' Marsh, 2013 * '' Pioscelus'' Muesebeck & Walkley 1951 * '' Synspilus'' Belokobylskij & Quicke, 2000

Tribe Holcobraconini

* '' Holcobracon'' Cameron, 1905 * '' Ivondrovia'' Shenefelt & Marsh, 1976 * '' Liodoryctes'' Szépligeti, 1906 * '' Odontobracon'' Cameron, 1887 * '' Monarea'' Szépligeti, 1904 * '' Nervellius'' Roman, 1924 * '' Zombrus'' Marshall, 1897

Tribe Labaniini

* '' Labania'' Hedqvist, 1963

Tribe Leptospathiini

* '' Australospathius'' Belokobylskij, Iqbal & Austin, 2004 * '' Leptospathius'' Szépligeti, 1902 * '' Oroceguera'' Seltmann & Sharkey, 2007 * '' Schlettereriella'' Szépligeti, 1904 * '' Stephanospathius'' Belokobylskij, 1992

Tribe Mononeurini

* '' Leptodoryctes'' Barbalho & Penteado-Dias, 1999 * '' Mononeuron'' Fischer, 1981 * '' Nipponecphylus'' Belokobylskij, 2001

Tribe Percnobraconini

* '' Fijibracon'' Belokobylskij, 1995 * '' Fritziella'' Marsh, 1993 * '' Masonius'' Marsh, 1993 * '' Micrommatus'' Marsh, 1993 * '' Percnobracon'' Kieffer & Jörgensen, 1910

Tribe Rhaconotini

* '' Antespathius'' Belokobylskij, 1995 * '' Aptenobracon'' Marsh, 1965 * '' Arhaconotus'' Belokobylskij, 2001 * '' Bathycentor'' Saussure, 1892 * '' Grangerdoryctes'' Belokobylskij, 2004 * '' Ipodoryctes'' Granger, 1949 * '' Leptorhaconotus'' Granger, 1949 * '' Mimipodoryctes'' Belokobylskij, 2000 * '' Neorhaconotus'' Belokobylskij, Iqbal & Austin, 2004 * '' Platyspathius'' Viereck, 1911 * '' Pseudorhaconotus'' van Achterberg, 2009 * '' Rhaconotus'' Ruthe, 1854 * '' Rhacontsira'' Belokobylskij, 1998

Tribe Sericobraconini

* '' Sericobracon'' Shaw & Edgerly, 1985

Tribe Siragrini

* ''

Arkoola ''Arkoola'' is a genus of fungus in the family Venturiaceae. This is a monotypic genus, containing the single species ''Arkoola nigra''. See also * List of soybean diseases Soybean plants (''Glycine max'') are subject to a variety of diseases ...

'' Belokobylskij, Iqbal & Austin, 2004 * '' Gurburra'' Belokobylskij, Iqbal & Austin, 2004 * '' Jarra'' Marsh & Austin, 1994 * '' Pseudosyngaster'' Belokobylskij, Iqbal & Austin, 2004 * '' Siragra'' Cameron, 1907 * '' Syngaster'' Brullé, 1846

Tribe Spathiini

* '' Afrospathius'' Belokobylskij & Quicke, 2000 * '' Celereon'' Say, 1836 * '' Ceylonspathius'' Belokobylskij * '' Concurtisella'' Roman, 1924 * '' Fijispathius'' Belokobylskij, Iqbal & Austin, 2004 * '' Hemispathius'' Belokobylskij & Quicke, 2000 * '' Notiospathius'' Matthews & Marsh * '' Parana'' Nixon, 1943 * '' Paraspathius'' Viereck, 1911 * '' Psenobolus'' Reinhard, 1885 * '' Ptesimogaster'' Marsh * '' Rhyukyuspathius'' Belokoblylskij, 2008 * '' Sisupala'' Nixon, 1943 * '' Spathiomorpha'' Tobias, 1976 * '' Spathioplites'' Fischer * ''

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 ...

'' Nees, 1818 * '' Termitospathius'' Belokobylskij, 2002 * '' Toka'' Nixon, 1943 * '' Trigonophasmus'' Enderlein, 1912

Tribe Ypsistocerini

* '' Embobracon'' van Achterberg, 1995 * '' Preembobracon'' Quicke & Butcher, 2015 * '' Termitobracon'' Brues, 1923 * '' Ypsistocerus'' Cushman, 1923

Tribal placement unknown

* '' Barbalhoa'' Marsh, 2002 * '' Bolivar'' Zaldívar-Riverón & Rodríguez-Jiménez, 2013 * '' Doryctopambolus'' Nunes, et al., 2012 * '' Iare'' Barbalho & Penteado-Dias, 2000 * '' Lianus'' Gomes & Penteado-Dias, 2006 * '' Lissopsius'' Marsh, 2002 * '' Mexiare'' Belokobylskij, Samaca-Sáenz, & Zaldívar-Riverón, 2015 * '' Mimodoryctes'' Zaldívar-Riverón & Rodríguez-Jiménez, 2013 * '' Rasnitsynoryctes'' Belokobylskij, 2011 * '' Sergey'' Martinez, Lazaro, Pedraza-Lara, & Zaldivar-Riveron, 2016 * '' Tuberatra'' Gadelha, Nunes, & de Oliveira, 2016 * '' Venifurca'' Gadelha, Nunes, Zaldivar-Riveron, & de Oliveira, 2016

References

External links

Photos on BugGuide

DNA barcodes on BOLD Systems
{{Taxonbar, from=Q602043 Braconidae Apocrita subfamilies