Doryctinae

Life-forms

Superregnum: Eukaryota
Cladus: Orthokaryota
Cladus: Neokaryota
Cladus: Scotokaryota
Cladus: Podiata
Cladus: Amorphea
Cladus: Obazoa
Cladus: Opisthokonta
Cladus: Holozoa
Cladus: Filozoa
Cladus: Choanozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: ParaHoxozoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Cladus: Mandibulata
Cladus: Pancrustacea
Cladus: Allotriocarida
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Holometabola
Superordo: Hymenopterida
Ordo: Hymenoptera
Subordo: Apocrita
Superfamilia: Ichneumonoidea

Familia: Braconidae
Subfamilia: Doryctinae
Tribus (16): Binareini – Doryctini – Ecphylini – Evaniodini – Hecabolini – Heterospilini – Holcobraconini – Labaniini – Leptospathiini – Mononeurini – Percnobraconini – Rhaconotini – Sericobraconini – Siragrini – Spathiini – Ypsistocerini
Overview of genera

Acanthodoryctes – Aivalykus – Allorhogas – Antidoryctes – Aptenobracon – Arhaconotus – Asiaheterospilus – Asiaontsira – Australospathius – Austrospathius – Barbalhoa – Binarea – Bolivar – Caenophanes – Cryptodoryctes – Cryptontsira – Cyphodoryctes – Donquickeia – Doryctes – Doryctinus – Doryctopambolus – Doryctopsis – Ecphylopsis – Ecphylus – Eodendrus – Evaniodes – Guaygata – Gymnobracon – Halycaea – Hecabolus – Heerz – Iare – Ipodoryctes – Ivondrovia – Johnsonius – Kauriphanes – Labania – Leptospathius – Lianus – Liobracon – Lissopsius – Mexiare – Monarea – Monitoriella – Monolexis – Neoheterospilus – Notiospathius – Ondigus – Ontsira – Oroceguera – Pambolidea – Parallorhogas – Parsteres – Percnobracon – Platyspathius – Psenobolus – Pseudorhaconotus – Pseudosyngaster – Rasnitsynoryctes – Ryukyuspathius – Schlettereriella – Sergey – Spathiomorpha – Spathius – Stephanospathius – Tuberatra – Venifurca – Wachsmannia – Whitfieldiellus – Zombrus – ...
Name

Doryctinae Förster, 1862

References

Foerster, A. 1862. Synopsis der Familien und Gattungen der Braconen. Verhandlungen des naturhistorischen Vereines der preussischen Rheinlande und Westphalens 19: 225–288. BHL Reference page.
Belokobylskij, S.A. 2008: Rhyukyuspathius, a new peculiar genus of the tribe Spathiini (Hymenoptera: Braconidae: Doryctinae) from Japan. Annales zoologici, 58: 761–766.
Belokobylskij, S.A. 2008: A new genus of the tribe Doryctini (Hymenoptera, Braconidae, Doryctinae). Zoosystematica Rossica, 17(1): 123–130.
Belokobylskij, S.A. 2011: Review of the genus Stephanospathius Belokobylskij, 1992 (Hymenoptera, Braconidae), with discussion of their tribal position. ZooKeys 118: 1–17. DOI: 10.3897/zookeys.118.914 Open access.
Belokobylskij, S.A., van Achterberg C. 2021. Review of the braconid parasitoid subfamily Doryctinae (Hymenoptera, Braconidae) from the United Arab Emirates and Yemen. European Journal of Taxonomy. 765(1), 1–143. DOI: 10.5852/ejt.2021.765.1479 Open access Reference page.
Belokobylskij, S.A.; Iqbal, M.; Austin, A.D. 2004: Systematics, distribution and diversity of the Australasian doryctine wasps (Hymenoptera, Braconidae, Doryctinae). Records of the South Australian Museum monograph series, 8: 1–150.
Belokobylskij, S.A.; Maeto, K. 2009: Doryctinae (Hymenoptera: Braconidae) of Japan. Fauna mundi, 1 ISBN 978-83-918040-8-7 abstract only seen
Belokobylskij, S. A., Samaca-Sáenz, E. & Zaldívar-Riverón, A. 2015: Mexiare gen. nov., a new Doryctinae genus (Hymenoptera: Braconidae) from Mexico with fused first and second metasomal tergites. Zootaxa 3914(2): 122–130. DOI: 10.11646/zootaxa.3914.2.2. Reference page.
Belokobylskij, S.A. ; Tang, P. & Chen, X. 2013. Asiaontsira gen. nov., a new tropical genus of the subfamily Doryctinae (Hymenoptera: Braconidae) from Vietnam and South-East China. Entomological Science, 16 (3): 309–315. Abstract: DOI: 10.1111/ens.12010
Gadelha, S. De S., Nunes, J.F. & Oliveira, M.L. de 2016. New genera, species and records of Doryctinae (Hymenoptera: Braconidae) from Brazil. Zootaxa 4083(3): 359–370. DOI: 10.11646/zootaxa.4083.3.2.Reference page.
Gadelha, S. de S., Nunes, J.F., Zaldivar-Riveron, A. & de Oliveira, M.L. 2016. Venifurca, a new genus of neotropical Doryctinae (Hymenoptera: Braconidae), and its phylogenetic placement. Journal of Hymenoptera Research 51: 91–100. DOI: 10.3897/jhr.51.9634. Reference page.
Gomes, S.A.G. & A.M. Penteado-Dias, 2006: Description of a new genus of Doryctinae wasps (Hymenoptera: Braconidae) from Brazil. Zoologische Mededelingen 80-4 (7): 81–85. Full article: [1]
Gutiérrez-Arellano, D., Gutiérrez-Arellano, C.R. & Zaldívar-Riverón, A. 2015. DNA Barcoding of the parasitoid wasp subfamily Doryctinae (Hymenoptera: Braconidae) from Chamela, Mexico. Biodiversity Data Journal 3: e5109. DOI: 10.3897/BDJ.3.e5109. Reference page.
Long, K.D. & Belokobylskij, S.A. 2019. First record of the genus Rasnitsynoryctes Belokobylskij, 2011 (Hymenoptera, Braconidae, Doryctinae) in Vietnam, with the description of a new species. Zookeys, 854: 17–24. DOI: 10.3897/zookeys.854.34810 Reference page.
Marsh, P.M. 1993: Descriptions of new Western Hemisphere genera of the subfamily Doryctinae (Hymenoptera: Braconidae). Contributions of the American Entomological Institute, 28(1): 1–58. [not seen]
Marsh, P.M. 1997: Replacement names for Western Hemisphere genera of Doryctinae (Hymenoptera: Braconidae). Proceedings of the Entomological Society of Washington, 99(3): 586. BHL
Martinez, J.J., Lázaro, R.N.M., Pedraza-Lara, C. & Zaldivar-Riveron, A. 2016. Sergey gen. n., a new doryctine genus from temperate forests of Mexico and Cuba (Hymenoptera, Braconidae). ZooKeys 589: 143–164. DOI: 10.3897/zookeys.589.8291. Reference page.
Quicke, D.L.J. & Butcher, B.A. 2015. Preembobracon gen. nov. (Hymenoptera: Braconidae: Doryctinae: Ypsistocerini: Embobraconina) from Brazil. Zootaxa 4000(2): 275–280. DOI: 10.11646/zootaxa.4000.2.5 Preview (PDF) Reference page.
Zaldivar-Riverón, A.; Belokobylskij, S.A.; León-Regagnon, V.; Briceño-G., R.; Quicke, D.L.J. 2008: Molecular phylogeny and historical biogeography of the cosmopolitan parasitic wasp subfamily Doryctinae (Hymenoptera: Braconidae). Invertebrate systematics, 22: 345–363. DOI: 10.1071/IS07028
Zaldivar-Riverón, A.; Belokobylskij, S.A.; León-Regagnon, V.; Martínez, J.J.; Briceño, R.; Quicke, D.L.J. 2007: A single origin of gall association in a group of parasitic wasps with disparate morphologies. Molecular phylogenetics and evolution, 44: 981–992. DOI: 10.1016/j.ympev.2007.05.016
Zaldívar-Riverón, A., Martínez, J.J., Bolokobylskij, S.A., Pedraza-Lara, C., Shaw, S.R., Hanson, P.E., Varela-Hernández, F. 2014. Systematics and evolution of gall formation in the plant-associated genera of the wasp subfamily Doryctinae (Hymenoptera: Braconidae). Systematic Entomology 39(4): 633–659. DOI: 10.1111/syen.12078 Reference page.

The Doryctinae or doryctine wasps are a large subfamily of parasitoid wasps within the family Braconidae.[1]
Description and identification
Heterospilus eurostae, male

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, and a cyclostome depression above the mandibles.[2]
Distribution

Doryctinae have a worldwide distribution except for the polar regions.[2]
Biology

Some Doryctinae are known to form galls on plants similar to the Mesostoinae (a small subfamily endemic to Australia).[3] Some species in the genus Allorhogas feed on seeds.[2] The large majority of doryctines are idiobiont ectoparasitoids of the larvae of wood-boring beetles – such as jewel beetles (Buprestidae). Some species parasitize lepidopteran or symphytan larvae. The host is paralyzed by venom injected through the female's ovipositor before an egg is laid.[2] 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 agents in Australia against eucalyptus pests.[4] The species Spathius agrili has been introduced to the United States from China in an effort to control the emerald ash borer.[5]
Taxonomy and phylogeny

Doryctinae is considered to be part of the cyclostome group, a clade of subfamilies including Alysiinae, Betylobraconinae, Braconinae, Gnamptodontinae, Exothecinae, Hormiinae, Lysiterminae, Opiinae, Pambolinae, Rogadinae, Rhysipolinae, and Rhyssalinae.[6]

16 tribes of Doryctinae have been designated.[1][7][8] The former subfamily Ypsistocerinae has been subsumed under Doryctinae as one of these tribes, Ypsistocerini.[9][8]

Doryctinae comprises over 2000 species in 198 genera, with new taxa continuing to be described.[1][10] 182 of the genera 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 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[11]
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</small
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[12]
Canchim Barbalho & Penteado-Dias, 1999
Dapsilitas Braet, Barbalhoa & van Achterberg, 2003
Ecphylopsis Ashmead, 1900
Heterospilus Haliday, 1836[12]
Jataiella Barbalho & Penteado-Dias, 1999
Neoheterospilus Belokobylskij, 2006[12]
Paraheterospilus Marsh, 2013[12]
Pioscelus Muesebeck & Walkley 1951[12]
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[7]
Leptospathius Szépligeti, 1902[7]
Oroceguera Seltmann & Sharkey, 2007[7]
Schlettereriella Szépligeti, 1904[7]
Stephanospathius Belokobylskij, 1992[7]

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[13]
Aptenobracon Marsh, 1965[13]
Arhaconotus Belokobylskij, 2001
Bathycentor Saussure, 1892[13]
Grangerdoryctes Belokobylskij, 2004[13]
Ipodoryctes Granger, 1949[13]
Leptorhaconotus Granger, 1949[13]
Mimipodoryctes Belokobylskij, 2000[13]
Neorhaconotus Belokobylskij, Iqbal & Austin, 2004[13]
Platyspathius Viereck, 1911
Pseudorhaconotus van Achterberg, 2009[13]
Rhaconotus Ruthe, 1854[13]
Rhacontsira Belokobylskij, 1998[13]

Tribe Sericobraconini

Sericobracon Shaw & Edgerly, 1985

Tribe Siragrini

Arkoola 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[14]
Sisupala Nixon, 1943
Spathiomorpha Tobias, 1976
Spathioplites Fischer
Spathius Nees, 1818[14]
Termitospathius Belokobylskij, 2002
Toka Nixon, 1943
Trigonophasmus Enderlein, 1912

Tribe Ypsistocerini

Embobracon van Achterberg, 1995[9]
Preembobracon Quicke & Butcher, 2015[9]
Termitobracon Brues, 1923[9]
Ypsistocerus Cushman, 1923[9]

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

Edmardash, Yusuf A.; El-Ghiet, Usama M. Abu; Soliman, Ahmed M.; Al-Fifi, Zarrag I. A.; Gadallah, Neveen S. (2020). "First contribution to the doryctine fauna (Hymenoptera, Braconidae, Doryctinae) of Farasan Archipelago, Saudi Arabia, with new records and the description of a new species". ZooKeys (977): 41–74. Bibcode:2020ZooK..977...41E. doi:10.3897/zookeys.977.56314. PMC 7596022. PMID 33177942.
Marsh, Paul M. (1997). "Subfamily Doryctinae". In Wharton, Robert A.; Marsh, Paul M.; Sharkey, Michael J. (eds.). Manual of the New World Genera of the Family Braconidae (Hymenoptera) (PDF). Washington, D.C.: The International Society of Hymenopterists. p. 207-233. Archived from the original (PDF) on 3 Aug 2023.
Samacá-Sáenz, Ernesto; Meza-Lázaro, Rubi N.; Branstetter, Michael G.; Zaldívar-Riverón, Alejandro (2019-11-17). "Phylogenomics and mitochondrial genome evolution of the gall-associated doryctine wasp genera (Hymenoptera: Braconidae)". Systematics and Biodiversity. 17 (8): 731–744. Bibcode:2019SyBio..17..731S. doi:10.1080/14772000.2019.1685608. ISSN 1477-2000. S2CID 213816441.
Stevens, N.B.; Iqbal, M; Austin, A.D.; Jennings, J.T. (2008). "Subfamily Doryctinae". Australian Faunal Directory. Archived from the original on 24 May 2015.
Gould, Juli; Bauer, Leah, "Biological Control of Emerald Ash Borer (Agrilus planipennis)" (PDF), Animal and Plant Health Inspection Service (APHIS), United States Department of Agriculture, archived from the original (PDF) on 10 January 2011, retrieved 28 April 2011
Sharanowski, Barbara J.; Dowling, Ashley P. G.; Sharkey, Michael J. (2011). "Molecular phylogenetics of Braconidae (Hymenoptera: Ichneumonoidea), based on multiple nuclear genes, and implications for classification". Systematic Entomology. 36 (3): 549–572. Bibcode:2011SysEn..36..549S. doi:10.1111/j.1365-3113.2011.00580.x. S2CID 52226040.
Belokobylskij, Sergey A. (2011). "Review of the genus Stephanospathius Belokobylskij, 1992 (Hymenoptera, Braconidae), with discussion of their tribal position". ZooKeys (118): 1–17. Bibcode:2011ZooK..118....1B. doi:10.3897/zookeys.118.914. PMC 3175109. PMID 21998508.
"Taxa: Doryctinae". MBD Database. OSU. Retrieved 2024-04-01.
Quicke, Donald L.J.; Bitcher, Buntika A. (2015). "Preembobracon gen. nov. (Hymenoptera: Braconidae: Doryctinae: Ypsistocerini: Embobraconina) from Brazil". Zootaxa. 4000 (2): 275–280. doi:10.11646/zootaxa.4000.2.5. PMID 26623614.
Braet, Yves (2016). "Doryctinae". Key to the genera of Doryctinae of the world. Retrieved 2024-04-01.
Wang, Man-man; Tan, Jiang-li; He, Jun-Hua; Chen, Xue-xin (9 November 2009). "The genus Eodendrus Belokobylskij (Hymenoptera, Braconidae, Doryctinae) from China, with description of a new species". ZooKeys (27): 43–50. Bibcode:2009ZooK...27...43W. doi:10.3897/zookeys.27.252.
Marsh, Paul M.; Wild, Alexander L.; Whitfield, James B. (2013). "The Doryctinae (Braconidae) of Costa Rica: genera and species of the tribe Heterospilini". ZooKeys (347): 1–474. Bibcode:2013ZooK..347....1M. doi:10.3897/zookeys.347.6002. PMC 3822444. PMID 24222723.
Jasso-Martínez, Jovana M.; Belokobylskij, Sergey A.; Zaldívar-Riveron, Alejandro (2019). "Molecular phylogenetics and evolution of generic diagnostic morphological features in the doryctine wasp tribe Rhaconotini (Hymenoptera: Braconidae)". Zoologischer Anzeiger. 279: 164–171. Bibcode:2019ZooAn.279..164J. doi:10.1016/j.jcz.2019.02.002.
Belokoblylskij, Sergey A. (2008). "Rhyukyuspathius, A New Peculiar Genus of the Tribe Spathiini (Hymenoptera: Braconidae: Doryctinae) from Japan". Annales Zoologici. 58 (4): 761–766. doi:10.3161/000345408X396693.