Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
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: Cynipoidea
Familia: Cynipidae
Subfamiliae (1 + 1†): Cynipinae - †Hodiernocynipinae
Name
Cynipidae Latreille, 1802
References
Azmaz, M. & Katılmış, Y. 2017. Updated species list of Cynipidae (Hymenoptera) from Turkey. Zootaxa 4303(3): 361–378. DOI: 10.11646/zootaxa.4303.3.3. Reference page.
Askew, R.R. et al. 2013: Catalogue of parasitoids and inquilines in cynipid oak galls in the West Palaearctic. Zootaxa 3643(1): 1–133. DOI: 10.11646/zootaxa.3643.1.1 Reference page.
Gómez, J.F.; Nieves-Aldrey, J.L. 2012: Notes on the larval morphology of Pteromalidae (Hymenoptera: Chalcidoidea) species parasitoids of gall wasps (Hymenoptera: Cynipidae) in Europe. Zootaxa, 3189: 39–55. Preview
Kovalev 1994: Palaeontological history, phylogeny and the system of brachycleistogastromorphs and cynipomorphs (Hymenoptera, Brachycleistogastromorpha infraorder n., Cynipomorpha infraorder n.) with description of new fossil and recent families, subfamilies and genera. Entomologicheskoe obozrenie, 73(2): 385–426, 495.
Liljeblad, J.; Ronquist, F. 1998: A phylogenetic analysis of higher-level gall wasp relationships (Hymenoptera: Cynipidae). Systematic entomology, 23: 229–252. DOI: 10.1046/j.1365-3113.1998.00053.x
Melika, G. 2006: Gall wasps of Ukraine. Cynipidae. Vestnik zoologii, supplement 21 (2 volumes). ISSN: 0084-5604
Rokas, A.; Nylander, J.A.A.; Ronquist, F.; Stone, G.N. 2002: A maximum-likelihood analysis of eight phylogenetic markers in gallwasps (Hymenoptera: Cynipidae): implications for insect phylogenetic studies. Molecular phylogenetics and evolution, 22: 206–219. DOI: 10.1006/mpev.2001.1032
Ronquist, F.; Liljeblad, J. 2001: Evolution of the gall wasp–host plant association. Evolution, 55: 2503–2522. DOI: 10.1111/j.0014-3820.2001.tb00765.x PDF
Vernacular names
Deutsch: Gallwespen
English: Gall wasps
עברית: צרעת העפצים
日本語: タマバチ科
Nederlands: Echte galwespen
norsk: Gallvepser
polski: Galasówkowate
svenska: Gallsteklar
中文: 瘿蜂科, 癭蜂科
Gall wasps, also traditionally called gallflies, are hymenopterans of the family Cynipidae in the wasp superfamily Cynipoidea. Their common name comes from the galls they induce on plants for larval development. About 1,300 species of this generally very small creature (1–8 millimetres (0.039–0.315 in)) are known worldwide,[1] with about 360 species of 36 different genera in Europe and some 800 species in North America.
Features
Like all Apocrita, gall wasps have a distinctive body shape, the so-called wasp waist. The first abdominal tergum (the propodeum) is conjoined with the thorax, while the second abdominal segment forms a sort of shaft, the petiole. The petiole connects with the gaster, which is the functional abdomen in apocritan wasps, starting with the third abdominal segment proper. Together, the petiole and the gaster form the metasoma, while the thorax and the propodeum make up the mesosoma.
The antennae are straight and consist of two or three segments. In many varieties, the backside of the mesosoma appears longitudinally banded. The wings are typically simply structured. The female's egg-depositing ovipositor is often seen protruding from the tip of the metasoma.
Reproduction and development
The reproduction of gall wasps is usually partly parthenogenesis, in which a male is completely unnecessary, and partly two-sex propagation.[2] Most species have alternating generations, with one two-sex generation and one parthenogenic generation annually, whereas some species produce very few males and reproduce only by parthenogenesis,[2] possibly because of infection of the females' gametes by endosymbiotic Wolbachia bacteria.[3] The various generations differentiate both in their appearance and in the form of the plant galls they induce.
The larvae of most gall wasps develop in characteristic plant galls they induce themselves, but many species are instead inquilines of other gall wasps, such as those of the genus Synergus.
The plant galls mostly develop directly after the female insect lays the eggs. The inducement for the gall formation is largely unknown; discussion speculates as to chemical, mechanical, and viral triggers. The hatching larvae nourish themselves with the nutritive tissue of the galls, in which they are otherwise well-protected from external environmental effects. The host plants, and the size and shape of the galls are specific to the majority of gall wasps, with about 70% of the known species parasitizing various types of oak. Galls can be found on nearly all parts of such trees, including the leaves, buds, branches, and roots. Other species of gall wasps live in eucalyptus, maple, and many herbs. Species determination is usually much easier through observation of the galls produced rather than of the insect itself.
Parasitism
A gall protects the developing gall wasp for the most vulnerable stage of its life cycle, but many other wasps have found a way to penetrate this defence and parasitise the larva(e) within. Some of these parasitoids use their long, hardened egg-laying tube (ovipositor) to bore into the gall and lay an egg on the helpless gall maker. These parasitoids may, in turn, be preyed upon by other wasps, hyperparasitoids.[4][page needed]
Types
Most species of gall wasps live as gall-formers on oaks. One of the best-known is the common oak gall wasp (Cynips quercusfolii), which induces characteristic, 2-cm in diameter, spherical galls on the undersides of oak leaves.
These turn reddish in the fall and are commonly known as oak apples. Light lentiform galls on the undersides of the same leaves are induced by Neuroterus quercusbaccarum; darker ones with bulging edges are formed by Neuroterus numismalis. Also striking are the galls of Cynips longiventris, which likewise can be found on the undersides of leaves, and are recognizable for their spheroidal shape and irregular red streaks. The oak potato gall wasp (Biorrhiza pallida) has round galls that grow to about 4 cm. These are known colloquially as oak potatoes. The latter type of gall is induced by this type of wasp not on the leaves, but on the roots of the oak. On the buds of young oak twigs, one can often find the hard-shelled galls of Andricus kollari and Andricus quercustozae. Galls do not cause significant harm to oak trees.[5]
Evolution
External phylogeny
The external phylogeny of the Cynipidae is based on Peters et al 2017. The Apocrita is within the "Sawflies" which are shown separately for simplicity here.[6]
| Hymenoptera |
|
Internal phylogeny
The internal phylogeny of gall wasps in the cladogram is based on the molecular phylogenetic analysis of Hearn et al. 2023.[7]
| Cynipidae |
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Taxonomy
Further information: List of Cynipidae genera
The Cynipidae contains two subfamilies, one extinct and one extant:
Cynipinae
Hodiernocynipinae†
The Cynipinae consists of nine tribes:[7]
Aulacideini Nieves-Aldrey, Nylander & Ronquist, 2015.
Aylacini Ashmead, 1903.
Ceroptresini Nieves-Aldrey, Nylander & Ronquist, 2015.
Cynipini Billberg, 1820.
Diastrophini Nieves-Aldrey, Nylander & Ronquist, 2015.
Eschatocerini Ashmead, 1903.
Phanacidini Nieves-Aldrey, Nylander & Ronquist, 2015.
Qwaqwaiini Liljeblad, Nieves-Aldrey & Melika, 2011.
Synergini Ashmead, 1896.
In human culture
The galls of several species, especially Mediterranean variants, were once used as tanning agents.
Before his work in human sexuality, Alfred Kinsey was known for his study of gall wasps.[8]
Galls formed on oak trees are one of the main ingredients in iron gall ink.
See also
Oak apple
Oak marble gall
Knopper gall
Common spangle gall
Silk button gall
References
Yoshihisa Abe; George Melika; Graham N. Stone (January 2007). "The diversity and phylogeography of cynipid gallwasps (Hymenoptera: Cynipidae) of the Oriental and eastern Palearctic regions, and their associated communities". Oriental Insects. 41 (1): 169–212. doi:10.1080/00305316.2007.10417504. ISSN 0030-5316. Wikidata Q56853717.
The population biology of oak gall wasps (Hymenoptera: Cynipidae) Stone et al. (2002) Annual Review of Entomology Vol. 47: 633-668
Menno Schilthuizen; Richard Stouthamer (September 1998). "Distribution of Wolbachia among the guild associated with the parthenogenetic gall wasp Diplolepis rosae". Heredity. 81 (3): 270–274. doi:10.1046/J.1365-2540.1998.00385.X. ISSN 0018-067X. Wikidata Q56609382.
Piper, Ross (2007). Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals. Greenwood Press. ISBN 978-0313339226.
Kidd, Russell (September 10, 2012). "Tree galls are rarely cause for concern". Michigan State University Extension.
Peters, Ralph S.; Krogmann, Lars; Mayer, Christoph; Donath, Alexander; Gunkel, Simon; Meusemann, Karen; Kozlov, Alexey; Podsiadlowski, Lars; Petersen, Malte (2017). "Evolutionary History of the Hymenoptera". Current Biology. 27 (7): 1013–1018. doi:10.1016/j.cub.2017.01.027. hdl:2434/801122. PMID 28343967.
Jack Hearn; Erik Gobbo; José Luis Nieves-Aldrey; et al. (3 October 2023). "Phylogenomic analysis of protein-coding genes resolves complex gall wasp relationships". Systematic Entomology. doi:10.1111/SYEN.12611. ISSN 0307-6970. Wikidata Q123440111.
Yudell, Michael (July 1, 1999). "Kinsey's Other Report". Natural History. 108 (6). ISSN 0028-0712.
Further reading
Gauld, I. D.; Bolton, B. (1988). The Hymenoptera. Oxford: Oxford University Press. ISBN 978-0-19-858521-3.
Honomichl, K.; Bellmann, H. (1994). Biologie und Ökologie der Insekten (in German). Stuttgart: Gustav Fischer. ISBN 978-3-437-25020-0.
Liljeblad, J. (2002). Phylogeny and Evolution of Gall Wasps (Hymenoptera: Cynipidae). Department of Zoology, Stockholm University. 1–176. Doctoral thesis. ISBN 978-91-7265-494-5.
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