Gall wasp

{{Short description|Family of wasps}} {{Use dmy dates|date=November 2025}} {{Automatic taxobox | name = Gall wasp | image = Gall Wasp - Cynipidae family, Leesylvania State Park, Woodbridge, Virginia.jpg | image_caption = | display_parents = 3 | taxon = Cynipidae | authority = Latreille, 1802 | diversity_link = List of Cynipidae genera | diversity = at least 80 genera }} thumb|''Diastrophus nebulosus'' on a raspberry gall

'''Gall wasps''', also traditionally called '''gallflies''', or cynipids, 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 cynipids are known worldwide,<ref name="Abe Melika Stone 2007">{{Cite Q|Q56853717}}</ref> with about 360 species of 36 different genera in Europe and some 800 species in North America. Paraulacidae and Diplolepididae were traditionally included within the Cynipidae but have since been designated as their own families after recent phylogenetic studies.<ref>{{Cite journal |last1=Hearn |first1=Jack |last2=Gobbo |first2=Erik |last3=Nieves-Aldrey |first3=José Luis |last4=Branca |first4=Antoine |last5=Nicholls |first5=James A. |last6=Koutsovoulos |first6=Georgios |last7=Lartillot |first7=Nicolas |last8=Stone |first8=Graham N. |last9=Ronquist |first9=Fredrik |date=January 2024 |title=Phylogenomic analysis of protein-coding genes resolves complex gall wasp relationships |url=https://resjournals.onlinelibrary.wiley.com/doi/10.1111/syen.12611 |journal=Systematic Entomology |language=en |volume=49 |issue=1 |pages=110–137 |doi=10.1111/syen.12611 |bibcode=2024SysEn..49..110H |issn=0307-6970|doi-access=free }}</ref><ref>{{Cite journal |last1=Blaimer |first1=Bonnie B. |last2=Gotzek |first2=Dietrich |last3=Brady |first3=Seán G. |last4=Buffington |first4=Matthew L. |date=23 November 2020 |title=Comprehensive phylogenomic analyses re-write the evolution of parasitism within cynipoid wasps |journal=BMC Evolutionary Biology |volume=20 |issue=1 |page=155 |doi=10.1186/s12862-020-01716-2 |doi-access=free |issn=1471-2148 |pmc=7686688 |pmid=33228574 |bibcode=2020BMCEE..20..155B }}</ref>

== 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.

For macropterous species, wing venation is used to diagnose the superfamily Cynipoidea and to separate the family Cynipidae proper from similar families within Cynipoidea (e.g. Figitidae, Liopteridae).<ref>{{Cite journal |last1=Buffington |first1=Matthew L |last2=Forshage |first2=Mattias |last3=Liljeblad |first3=Johan |last4=Tang |first4=Chang-Ti |last5=van Noort |first5=Simon |date=1 July 2020 |editor-last=Hines |editor-first=Heather |title=World Cynipoidea (Hymenoptera): A Key to Higher-Level Groups |url=https://academic.oup.com/isd/article/doi/10.1093/isd/ixaa003/5865384 |journal=Insect Systematics and Diversity |language=en |volume=4 |issue=4 |article-number=1 |doi=10.1093/isd/ixaa003 |issn=2399-3421}}</ref>

The antennae are straight and consist of two or three segments. In many species, 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 ==

Gall wasp larvae typically develop as either gall inducers or as inquilines of other gall wasps, as is the case in the genus ''Synergus.''<ref>{{Cite journal |last1=Lobato-Vila |first1=Irene |last2=Caicedo |first2=Guadalupe |last3=Rodríguez |first3=Pedro A. |last4=Pujade-Villar |first4=Juli |date=16 January 2020 |title=The inquiline oak gall wasp (Hymenoptera: Cynipidae) fauna from Colombia: new data and species |url=https://www.cambridge.org/core/product/identifier/S0008347X19000774/type/journal_article |journal=The Canadian Entomologist |language=en |pages=1–14 |doi=10.4039/tce.2019.77 |issn=0008-347X|url-access=subscription }}</ref> Females lay eggs in various plant tissues from which the growth of the gall structure is induced. Larvae feed on the nutritive tissues of these galls before pupating and emerging from the galls as adults. Different gall wasp species are able to induce galls on a large variety of plant organs with species producing galls on leaves, petioles, buds, flowers, roots, fruits, and branches/stems.<ref name=":0">{{Cite journal |last1=Egan |first1=Scott P. |last2=Hood |first2=Glen R. |last3=Martinson |first3=Ellen O. |last4=Ott |first4=James R. |date=December 2018 |title=Cynipid gall wasps |url=https://linkinghub.elsevier.com/retrieve/pii/S0960982218313575 |journal=Current Biology |language=en |volume=28 |issue=24 |pages=R1370–R1374 |doi=10.1016/j.cub.2018.10.028 |bibcode=2018CBio...28R1370E }}</ref>

The reproduction of gall wasps is usually heterogonic, where life cycles feature both a parthenogenic generation, in which individuals reproduce asexually, and a dioecious generation, requiring both males and females for sexual reproduction.<ref name="Stone 2002">{{Cite journal |last1=Stone |first1=Graham N. |last2=Schönrogge |first2=Karsten |last3=Atkinson |first3=Rachel J. |last4=Bellido |first4=David |last5=Pujade-Villar |first5=Juli |date=1 January 2002 |title=The Population Biology of Oak Gall Wasps (Hymenoptera: Cynipidae) |url=https://www.annualreviews.org/content/journals/10.1146/annurev.ento.47.091201.145247 |journal=Annual Review of Entomology |language=en |volume=47 |issue= |pages=633–668 |doi=10.1146/annurev.ento.47.091201.145247 |issn=0066-4170|url-access=subscription }}</ref> Most species have alternating generations, with one sexual generation and one asexual parthenogenic generation annually. However, some species reproduce entirely parthenogenically. Both thelytoky and arrhenotoky have been observed among the heterogonic species.<ref name="auto">{{Cite journal |last1=Rokas |first1=A. |last2=Atkinson |first2=R. J. |last3=Nieves-Aldrey |first3=J.-L. |last4=West |first4=S. A. |last5=Stone |first5=G. N. |date=September 2002 |title=The incidence and diversity of Wolbachia in gallwasps (Hymenoptera; Cynipidae) on oak |journal=Molecular Ecology |volume=11 |issue=9 |pages=1815–1829 |doi=10.1046/j.1365-294x.2002.01556.x |issn=0962-1083 |pmid=12207731 |bibcode=2002MolEc..11.1815R }}</ref> The cause of the arisal of parthenogenesis among Cynipidae remains unclear. Though infection by endosymbiotic ''Wolbachia'' bacteria has been shown to cause parthenogenesis in some studies, the evidence is inconsistent among different tribes in the family.<ref>{{Cite journal |last1=Abe |first1=Yoshihisa |last2=Miura |first2=Kazuki |date=1 September 2002 |title=Doses Wolbachia Induce Unisexuality in Oak Gall Wasps? (Hymenoptera: Cynipidae) |url=https://academic.oup.com/aesa/article/95/5/583/28687 |journal=Annals of the Entomological Society of America |language=en |volume=95 |issue=5 |pages=583–586 |doi=10.1603/0013-8746(2002)095[0583:DWIUIO]2.0.CO;2 |issn=0013-8746 |archive-url=http://web.archive.org/web/20210308150657/https://academic.oup.com/aesa/article/95/5/583/28687 |archive-date=8 March 2021}}</ref><ref name="auto"/>

Gall wasps are known from a large variety of plant species including many species of Fagaceae, Rosaceae, Asteraceae, Papaveraceae, Lamiaceae, and Caprifoliaceae, among the more common host families.<ref>{{Cite journal |last1=Ronquist |first1=Fredrik |last2=Liljeblad |first2=Johan |date=December 2001 |title=Evolution of the Gall Wasp-Host Plant Association |url=https://academic.oup.com/evolut/article/55/12/2503/6758107 |journal=Evolution |language=en |volume=55 |issue=12 |pages=2503–2522 |doi=10.1111/j.0014-3820.2001.tb00765.x |issn=0014-3820}}</ref><ref name="auto1">{{Cite journal |last1=Nastasi |first1=Louis F. |last2=Davis |first2=Charles K. |last3=Nieves-Aldrey |first3=José Luis |last4=Buffington |first4=Matthew L. |last5=Van Noort |first5=Simon |last6=Deans |first6=Andrew R. |date=16 July 2025 |title=Review of Herb Gall Wasp Tribes (Hymenoptera: Cynipidae: Aylacini Sensu lato), with an Updated Key to Cynipid Tribes and a Checklist of World Species |url=https://bioone.org/journals/proceedings-of-the-entomological-society-of-washington/volume-127/issue-1/0013-8797.127.1.84/Review-of-Herb-Gall-Wasp-Tribes-Hymenoptera--Cynipidae/10.4289/0013-8797.127.1.84.full |journal=Proceedings of the Entomological Society of Washington |volume=127 |issue=1 |doi=10.4289/0013-8797.127.1.84 |issn=0013-8797|url-access=subscription }}</ref> Host breadth is moderately conserved among gall wasps, with multiple repeated host shifts representing major divergences among the various lineages in the family.<ref>{{Cite journal |last1=Stone |first1=Graham N. |last2=Hernandez-Lopez |first2=Antonio |last3=Nicholls |first3=James A. |last4=di Pierro |first4=Erica |last5=Pujade-Villar |first5=Juli |last6=Melika |first6=George |last7=Cook |first7=James M. |date=April 2009 |title=Extreme Host Plant Conservatism During at Least 20 Million Years of Host Plant Pursuit by Oak Gallwasps |url=https://academic.oup.com/evolut/article/63/4/854/6854678 |journal=Evolution |language=en |volume=63 |issue=4 |pages=854–869 |doi=10.1111/j.1558-5646.2008.00604.x |bibcode=2009Evolu..63..854S |url-access=subscription }}</ref><ref>{{Cite journal |last1=Ronquist |first1=Fredrik |last2=Nieves-Aldrey |first2=José-Luis |last3=Buffington |first3=Matthew L. |last4=Liu |first4=Zhiwei |last5=Liljeblad |first5=Johan |last6=Nylander |first6=Johan A. A. |date=20 May 2015 |editor-last=López-Vaamonde |editor-first=Carlos |title=Phylogeny, Evolution and Classification of Gall Wasps: The Plot Thickens |journal=PLOS ONE |language=en |volume=10 |issue=5 |article-number=e0123301 |doi=10.1371/journal.pone.0123301 |doi-access=free |issn=1932-6203 |pmc=4439057 |pmid=25993346 |bibcode=2015PLoSO..1023301R }}</ref><ref name="auto1"/> In contrast, speciation events and divergence at lower taxonomic levels are more associated with shifts in plant organ use.<ref>{{Cite journal |last1=Cook |first1=James M. |last2=Rokas |first2=Antonis |last3=Pagel |first3=Mark |last4=Stone |first4=Graham N. |date=September 2002 |title=Evolutionary Shifts Between Host Oak Sections and Host-Plant Organs in Andricus Gallwasps |url=https://academic.oup.com/evolut/article/56/9/1821/6755971 |journal=Evolution |language=en |volume=56 |issue=9 |pages=1821–1830 |doi=10.1111/j.0014-3820.2002.tb00196.x |pmid=12389727 |issn=0014-3820}}</ref> Host breadth within a species is usually restricted to one or a few closely-related plant host species.<ref>{{Cite journal |last1=Fang |first1=Zhiqiang |last2=Tang |first2=Chang-Ti |last3=Sinclair |first3=Frazer |last4=Csóka |first4=György |last5=Hearn |first5=Jack |last6=McCormack |first6=Koorosh |last7=Melika |first7=George |last8=Mikolajczak |first8=Katarzyna M. |last9=Nicholls |first9=James A. |last10=Nieves-Aldrey |first10=José-Luis |last11=Notton |first11=David G. |last12=Radosevic |first12=Sara |last13=Bailey |first13=Richard I. |last14=Reiss |first14=Alexander |last15=Zhang |first15=Yuanmeng M. |date=November 2024 |title=Network structure and taxonomic composition of tritrophic communities of Fagaceae, cynipid gallwasps and parasitoids in Sichuan, China |url=https://resjournals.onlinelibrary.wiley.com/doi/10.1111/icad.12768 |journal=Insect Conservation and Diversity |language=en |volume=17 |issue=6 |pages=1046–1071 |doi=10.1111/icad.12768 |issn=1752-458X|doi-access=free }}</ref> However, the different generations within a species may exploit different sets of host species as well as different plant organs to complete their life cycles.<ref>{{Cite journal |last1=Hood |first1=Glen R |last2=Zhang |first2=Linyi |last3=Topper |first3=Leah |last4=Brandão-Dias |first4=Pedro F P |last5=Del Pino |first5=Gaston A |last6=Comerford |first6=Mattheau S |last7=Egan |first7=Scott P |date=30 April 2018 |title='Closing the Life Cycle' of Andricus quercuslanigera (Hymenoptera: Cynipidae) |url=https://academic.oup.com/aesa/article/111/3/103/4955773 |journal=Annals of the Entomological Society of America |language=en |volume=111 |issue=3 |pages=103–113 |doi=10.1093/aesa/say005 |issn=0013-8746|url-access=subscription }}</ref><ref>{{Cite journal |last1=Stone |first1=Graham |last2=Atkinson |first2=Rachel |last3=Rokas |first3=Antonis |last4=Csóka |first4=György |last5=Nieves-Aldrey |first5=José-Luis |date=March 2001 |title=Differential success in northwards range expansion between ecotypes of the marble gallwasp Andricus kollari: a tale of two lifecycles |url=https://onlinelibrary.wiley.com/doi/10.1046/j.1365-294x.2001.01211.x |journal=Molecular Ecology |language=en |volume=10 |issue=3 |pages=761–778 |doi=10.1046/j.1365-294x.2001.01211.x |pmid=11298986 |bibcode=2001MolEc..10..761S |issn=0962-1083|url-access=subscription }}</ref>

== Natural enemies ==

Despite being concealed within galls, gall wasp larvae and pupae are the target of many parasitoid species, particularly wasps in the superfamily Chalcidoidea.<ref>{{Cite journal |last1=Janšta |first1=Petr |last2=Cruaud |first2=Astrid |last3=Delvare |first3=Gérard |last4=Genson |first4=Guénaëlle |last5=Heraty |first5=John |last6=Křížková |first6=Barbora |last7=Rasplus |first7=Jean-Yves |date=December 2018 |title=Torymidae (Hymenoptera, Chalcidoidea) revised: molecular phylogeny, circumscription and reclassification of the family with discussion of its biogeography and evolution of life-history traits |url=https://onlinelibrary.wiley.com/doi/10.1111/cla.12228 |journal=Cladistics |language=en |volume=34 |issue=6 |pages=627–651 |doi=10.1111/cla.12228 |pmid=34706481 |issn=0748-3007}}</ref><ref>{{Cite journal |last1=Askew |first1=Richard R. |last2=Melika |first2=George |last3=Pujade-Villar |first3=Juli |last4=SchöNrogge |first4=Karsten |last5=Stone |first5=Graham N. |last6=Nieves-Aldrey |first6=José Luis |date=30 April 2013 |title=Catalogue of parasitoids and inquilines in cynipid oak galls in the West Palaearctic |url=https://mapress.com/zt/article/view/zootaxa.3643.1.1 |journal=Zootaxa |volume=3643 |issue=1 |doi=10.11646/zootaxa.3643.1.1 |pmid=25340198 |bibcode=2013Zoot.36433.1.1A |issn=1175-5334}}</ref><ref>{{Cite journal |last1=Askew |first1=Richard R. |last2=Plantard |first2=Olivier |last3=GóMez |first3=José F. |last4=Nieves |first4=Maria Hernandez |last5=Nieves-Aldrey |first5=José Luis |date=24 August 2006 |title=Catalogue of parasitoids and inquilines in galls of Aylacini, Diplolepidini and Pediaspidini (Hym., Cynipidae) in the West Palaearctic |url=https://mapress.com/zt/article/view/zootaxa.1301.1.1 |journal=Zootaxa |volume=1301 |issue=1 |doi=10.11646/zootaxa.1301.1.1 |bibcode=2006Zoot.13011.1.1A |issn=1175-5334|url-access=subscription }}</ref> These parasitoids penetrate gall tissues with their ovipositors to lay eggs on or in the host gall wasp. Gall wasps may also be accidentally or facultatively killed by inquilines which accidentally injure the gall wasp while feeding on surrounding plant gall tissue or by causing changes to gall morphology.<ref name=":1">{{Cite journal |last1=PÉNZES |first1=ZSOLT |last2=MELIKA |first2=GEORGE |last3=BOZSÓKI |first3=ZOLTÁN |last4=BIHARI |first4=PÉTER |last5=MIKÓ |first5=ISTVÁN |last6=TAVAKOLI |first6=MAJID |last7=PUJADE-VILLAR |first7=JULI |last8=FEHÉR |first8=BALÁZS |last9=FÜLÖP |first9=DÁVID |last10=SZABÓ |first10=KRISZTIÁN |last11=BOZSÓ |first11=MIKLÓS |last12=SIPOS |first12=BOTOND |last13=SOMOGYI |first13=KÁLMÁN |last14=STONE |first14=GRAHAM N. |date=16 September 2009 |title=Systematic re-appraisal of the gall-usurping wasp genus ''Synophrus'' Hartig, 1843 (Hymenoptera: Cynipidae: Synergini) |journal=Systematic Entomology |volume=34 |issue=4 |pages=688–711 |doi=10.1111/j.1365-3113.2009.00482.x |bibcode=2009SysEn..34..688P |issn=0307-6970}}</ref>

The evolutionary pressure caused by these natural enemies provides the basis for one of the explanations for the evolution of galling behaviour across multiple galling insect lineages. The Enemy Hypothesis states that galls and their diverse morphologies evolved as defenses against parasitoids and other natural enemies.<ref>{{Cite journal |last1=Bailey |first1=Richard |last2=Schönrogge |first2=Karsten |last3=Cook |first3=James M. |last4=Melika |first4=George |last5=Csóka |first5=György |last6=Thuróczy |first6=Csaba |last7=Stone |first7=Graham N. |date=25 August 2009 |editor-last=Agrawal |editor-first=Anurag A. |title=Host Niches and Defensive Extended Phenotypes Structure Parasitoid Wasp Communities |journal=PLOS Biology |language=en |volume=7 |issue=8 |article-number=e1000179 |doi=10.1371/journal.pbio.1000179 |doi-access=free |issn=1545-7885 |pmc=2719808 |pmid=19707266}}</ref><ref>{{Cite journal |last1=Baine |first1=Quinlyn |last2=Hughes |first2=Daniel W. W. |last3=Casares |first3=Emily E. |last4=Martinson |first4=Ellen O. |last5=Martinson |first5=Vincent G. |date=December 2024 |title=External insect gall morphology influences the functional guilds of natural enemy communities |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=291 |issue=2036 |article-number=20242424 |doi=10.1098/rspb.2024.2424 |issn=1471-2954 |pmc=11631491 |pmid=39657810}}</ref> This hypothesis explains the wide variety in gall surface ornaments, in internal gall morphology, and in the overall shape of the galls produced by gall wasps. However, the large number of parasitoids present across the Cynipidae indicate that the parasitoids are able to adapt to these defenses and that some form of coevolution is present in this biological system.<ref>{{Cite journal |last1=Hayward |first1=Alex |last2=Stone |first2=Graham N. |date=October 2005 |title=Oak gall wasp communities: Evolution and ecology |url=https://linkinghub.elsevier.com/retrieve/pii/S1439179105000666 |journal=Basic and Applied Ecology |language=en |volume=6 |issue=5 |pages=435–443 |doi=10.1016/j.baae.2005.07.003 |bibcode=2005BApEc...6..435H |doi-access=free }}</ref>

Documenting and studying the community of natural enemies exploiting cynipids and their galls remains an active research avenue in ecology and evolutionary biology. However, due to the concealed nature of this community, it is often difficult to designate emerging insects as hosts, parasitoids of the hosts, inquilines, parasitoids of the inquilines, or hyperparasitoids.<ref name="Piper 2007">{{cite book |last=Piper |first=Ross |author-link=Ross Piper |year=2007 |chapter=Galls |title=Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals |url=https://archive.org/details/extraordinaryani0000pipe |url-access=registration |publisher=Greenwood Press |isbn=978-0-313-33922-6 }}</ref><ref name=":1" />

== Gall morphology ==

The mechanism underlying gall development has historically been one of the main challenges in understanding the biology of gall wasps. Female wasps lay their eggs within the meristematic tissue of the host plant and secretes chemicals inducing apoptosis of surrounding plant tissue.<ref name=":0" /> The metabolome of the local plant tissue changes, causing the development of the abnormal gall growth.<ref>{{Cite journal |last1=Harper |first1=L. J. |last2=SchöNrogge |first2=K. |last3=Lim |first3=K. Y. |last4=Francis |first4=P. |last5=Lichtenstein |first5=C. P. |date=February 2004 |title=Cynipid galls: insect-induced modifications of plant development create novel plant organs |url=https://onlinelibrary.wiley.com/doi/10.1046/j.1365-3040.2004.01145.x |journal=Plant, Cell & Environment |language=en |volume=27 |issue=3 |pages=327–335 |doi=10.1046/j.1365-3040.2004.01145.x |bibcode=2004PCEnv..27..327H |issn=0140-7791}}</ref><ref>{{Cite journal |last1=Markel |first1=Kasey |last2=Novak |first2=Vlastimil |last3=Bowen |first3=Benjamin P |last4=Tian |first4=Yang |last5=Chen |first5=Yi-Chun |last6=Sirirungruang |first6=Sasilada |last7=Zhou |first7=Andy |last8=Louie |first8=Katherine B |last9=Northen |first9=Trent R |last10=Eudes |first10=Aymerick |last11=Scheller |first11=Henrik V |last12=Shih |first12=Patrick M |date=30 April 2024 |title=Cynipid wasps systematically reprogram host metabolism and restructure cell walls in developing galls |url=https://academic.oup.com/plphys/article/195/1/698/7571489 |journal=Plant Physiology |language=en |volume=195 |issue=1 |pages=698–712 |doi=10.1093/plphys/kiae001 |pmid=38236304 |issn=0032-0889|pmc=11181936 }}</ref> These changes are modulated by the gall wasp and thus, galls represent an extended phenotype of the wasp.<ref name=":2">{{Cite journal |last1=Hearn |first1=Jack |last2=Blaxter |first2=Mark |last3=Schönrogge |first3=Karsten |last4=Nieves-Aldrey |first4=José-Luis |last5=Pujade-Villar |first5=Juli |last6=Huguet |first6=Elisabeth |last7=Drezen |first7=Jean-Michel |last8=Shorthouse |first8=Joseph D. |last9=Stone |first9=Graham N. |date=4 November 2019 |editor-last=Copenhaver |editor-first=Gregory P. |title=Genomic dissection of an extended phenotype: Oak galling by a cynipid gall wasp |journal=PLOS Genetics |language=en |volume=15 |issue=11 |article-number=e1008398 |doi=10.1371/journal.pgen.1008398 |doi-access=free |issn=1553-7404 |pmc=6855507 |pmid=31682601}}</ref> Although past hypotheses have suggested mutualisms with viruses or virus-like particles, there is little evidence for these as effectors in gall development.<ref name=":2" /><ref>{{Cite journal |last=Cornell |first=Howard V. |date=1983 |title=The Secondary Chemistry and Complex Morphology of Galls Formed by the Cynipinae (Hymenoptera): Why and How? |journal=The American Midland Naturalist |volume=110 |issue=2 |pages=225–234 |doi=10.2307/2425263 |jstor=2425263 |issn=0003-0031}}</ref>

Galls produced by cynipid gall wasps bear a striking diversity in external and internal morphology. The presence of varying levels of anthocyanins lead to a diversity of colours and colour patterns on galls which is hypothesized to be an aposematic adaptation among galls more broadly.<ref>{{Citation |last1=de Oliveira |first1=Denis Coelho |title=The Physiological and Ecological Processes of Anthocyanins Accumulation and Red Gall Coloration |date=2025 |work=Plant Galls |pages=427–441 |editor-last=Oliveira |editor-first=Denis Coelho de |url=https://link.springer.com/10.1007/978-3-031-80064-1_19 |access-date=24 November 2025 |place=Cham |publisher=Springer Nature Switzerland |language=en |doi=10.1007/978-3-031-80064-1_19 |isbn=978-3-031-80063-4 |last2=Inbar |first2=Moshe |last3=Lev-Yadun |first3=Simcha |editor2-last=Isaias |editor2-first=Rosy Mary dos Santos|url-access=subscription }}</ref><ref>{{Cite journal |last1=Inbar |first1=M. |last2=Izhaki |first2=I. |last3=Koplovich |first3=A. |last4=Lupo |first4=I. |last5=Silanikove |first5=N. |last6=Glasser |first6=T. |last7=Gerchman |first7=Y. |last8=Perevolotsky |first8=A. |last9=Lev-Yadun |first9=S. |date=March 2010 |title=Why do many galls have conspicuous colors? A new hypothesis |url=http://link.springer.com/10.1007/s11829-009-9082-7 |journal=Arthropod-Plant Interactions |language=en |volume=4 |issue=1 |pages=1–6 |doi=10.1007/s11829-009-9082-7 |bibcode=2010APInt...4....1I |issn=1872-8855|url-access=subscription }}</ref> Galls may bear various ornaments such as hairs, hooks, spines, pegs, and thorns of varying lengths and densities. Some species produce sticky substances on the gall surface, which is hypothesized to deter parasitoids, either by making it more difficult for parasitoids to walk on gall surfaces, or through myrmecophily by recruiting ants to defend the gall.<ref>{{Cite journal |last=Abe |first=Yoshihisa |date=1 February 1992 |title=The advantage of attending ants and gall aggregation for the gall wasp Andricus symbioticus (Hymenoptera: Cynipidae) |journal=Oecologia |language=en |volume=89 |issue=2 |pages=166–167 |doi=10.1007/BF00317214 |pmid=28312869 |bibcode=1992Oecol..89..166A |issn=1432-1939}}</ref> Internally, galls can also possess various empty cavities or hairs which may also render parasitization more difficult.<ref>{{Cite journal |last1=Ide |first1=Tatsuya |last2=Koyama |first2=Asuka |date=30 October 2023 |title=The formation of a rolling larval chamber as the unique structural gall of a new species of cynipid gall wasps |journal=Scientific Reports |language=en |volume=13 |issue=1 |page=18149 |doi=10.1038/s41598-023-43641-6 |pmid=37903850 |pmc=10616116 |bibcode=2023NatSR..1318149I |issn=2045-2322}}</ref><ref>{{Cite journal |last1=Melika |first1=G. |last2=Equihua-MartíNez |first2=A. |last3=Estrada-Venegas |first3=E. G. |last4=CibriáN-Tovar |first4=D. |last5=CibriáN-Llanderal |first5=V. D. |last6=Pujade-Villar |first6=J. |date=22 November 2011 |title=New Amphibolips gallwasp species from Mexico (Hymenoptera: Cynipidae) |url=https://mapress.com/zt/article/view/zootaxa.3105.1.2 |journal=Zootaxa |volume=3105 |issue=1 |doi=10.11646/zootaxa.3105.1.2 |issn=1175-5334|url-access=subscription }}</ref>

Below are some selected images of various galls produced by cynipid gall wasps:

<gallery mode="nolines"> File:Diplolepis Quercus02.jpg|{{center|''Andricus quercuscalicis'' parthenogenetic generation gall}} File:Cynips longiventris gall.jpg|{{center|''Cynips longiventris'' parthenogenetic generation gall}} File:Hopea Ponga 21.JPG|Galls on ''Hopea ponga'' at Peravoor File:Translucent Oak Gall.jpg|''Amphibolips nubilipennis'' gall from Pennsylvania, USA File:Common Spangle gall - detail.JPG|''Neuroterus quercusbaccarum'' gall from Scotland, UK File:Galle Aulacidea hieracii.jpg|''Aulacidea hieracii'' gall File:Andricus foecundatrix gall on Quercus, Corwen, North Wales, Aug 2020.jpg|''Andricus foecundatrix'' gall from Wales, UK </gallery>

== 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.<ref name=Peters>{{Cite journal |last1=Peters |first1=Ralph S. |last2=Krogmann |first2=Lars |last3=Mayer |first3=Christoph |last4=Donath |first4=Alexander |last5=Gunkel |first5=Simon |last6=Meusemann |first6=Karen |last7=Kozlov |first7=Alexey |last8=Podsiadlowski |first8=Lars |last9=Petersen |first9=Malte |title=Evolutionary History of the Hymenoptera |journal=Current Biology |volume=27 |issue=7 |pages=1013–1018 |doi=10.1016/j.cub.2017.01.027 |pmid=28343967 |year=2017 |doi-access=free |bibcode=2017CBio...27.1013P |hdl=2434/801122 |hdl-access=free }}</ref>

{{clade |label1=Hymenoptera |1={{clade |1=Sawflies (paraphyletic) 55px |label2=Apocrita |2={{clade |label1=Parasitoida |1={{clade |1={{clade |1=Ceraphronoidea |2=Ichneumonoidea 70px }} |label2=Proctotrupomorpha |2={{clade |label1=Cynipoidea |1={{clade |1='''Cynipidae''' 70px |2=other families }} |2=Chalcidoidea and other groups 70px }} }} |2={{clade |1={{clade |1=Evanioidea 90px |2=Stephanoidea }} |2={{clade |1=Trigonaloidea |2=Aculeata (stinging wasps, bees, ants) 70px }} }} }} }} }}

=== Internal phylogeny ===

The internal phylogeny of gall wasps in the cladogram is based on the molecular phylogenetic analysis of Hearn et al. 2023.<ref name="Hearn Gobbo Nieves-Aldrey 2023"/>

{{clade |label1='''Cynipidae''' |1={{clade |1={{clade |1=Eschatocerini |2={{clade |1={{clade |1=Phanacidini |2=Aulacideini }} |2={{clade |1={{clade |1=Qwaqwaiini |2=Synergini (inquiline gall wasps) }} |2={{clade |1=Diastrophini |2={{clade |1={{clade |1=Ceroptresini (inquiline oak gall wasps) |2=Aylacini }} |2={{clade |1=Cynipini (oak gall wasps) 70px }} }} }} }} }} }} }} }}

=== Taxonomy ===

The Cynipidae contains two subfamilies, one extinct and one extant:

* Cynipinae * Hodiernocynipinae†

The Cynipinae consists of nine tribes:<ref name="Hearn Gobbo Nieves-Aldrey 2023">{{Cite Q|Q123440111}}</ref>

* 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.<ref name="Yudell 1999">{{cite journal |last=Yudell |first=Michael |title=Kinsey's Other Report |url=http://findarticles.com/p/articles/mi_m1134/is_6_108/ai_55127889 |journal=Natural History |issn=0028-0712 |date=1 July 1999 |volume=108 |issue=6}}</ref> * Galls formed on oak trees are one of the main ingredients in iron gall ink. * Galls of some species have been used across various cultures in traditional medicine.<ref>{{Cite journal |last1=Elham |first1=Aliya |last2=Arken |first2=Miradel |last3=Kalimanjan |first3=Gulina |last4=Arkin |first4=Abdulaziz |last5=Iminjan |first5=Mubarak |date=12 June 2021 |title=A review of the phytochemical, pharmacological, pharmacokinetic, and toxicological evaluation of Quercus Infectoria galls |journal=Journal of Ethnopharmacology |volume=273 |article-number=113592 |doi=10.1016/j.jep.2020.113592 |issn=1872-7573 |pmid=33217520}}</ref>

== References ==

== Further reading ==

* {{cite book |last1=Gauld |first1=I. D. |last2=Bolton |first2=B. |year=1988 |title=The Hymenoptera |location=Oxford |publisher=Oxford University Press |isbn=978-0-19-858521-3 |ref=none}} * {{cite book |last1=Honomichl |first1=K. |last2=Bellmann |first2=H. |year=1994 |title=Biologie und Ökologie der Insekten |location=Stuttgart |publisher=Gustav Fischer |isbn=978-3-437-25020-0 |language=de |ref=none}} * {{cite book |last=Liljeblad |first=J. |year=2002 |title=Phylogeny and Evolution of Gall Wasps ({{lang|la|Hymenoptera: Cynipidae}}) |publisher=Department of Zoology, Stockholm University. 1–176. Doctoral thesis |isbn=978-91-7265-494-5 |ref=none}}

== External links ==

* {{Commons category-inline|Cynipidae}} * [http://entomology.ifas.ufl.edu/creatures/trees/Callirhytis_gall_wasps.htm ''Andricus quercusclavigera'' and ''A. quercuscornigera''] on the UF / IFAS Featured Creatures Web site * Wikispecies entry * [http://bugguide.net/node/view/14878/bgpage Family Cynipidae] at [http://bugguide.net/ BugGuide]

Category:Cynipidae Category:Gall-inducing insects Category:Asexual reproduction in animals Category:Taxa named by Pierre André Latreille