Gall wasp | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Suborder: | Apocrita |
Infraorder: | Proctotrupomorpha |
Superfamily: | Cynipoidea |
Family: | Cynipidae Latreille, 1802 |
Diversity | |
at least 80 genera |
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 or 1⁄32–5⁄16 inch) are known worldwide, [1] with about 360 species of 36 different genera in Europe and some 800 species in North America.
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.
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, inducing oak galls. 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.
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 gall and/or larva(e) within. Some of these inquilines and parasitoids use their long, hardened egg-laying tube (ovipositor) to bore into the gall. These parasitoids may, in turn, be preyed upon by other wasps, hyperparasitoids. [4]
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]
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 |
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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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The Cynipidae contains two subfamilies, one extinct and one extant:
The Cynipinae consists of nine tribes: [7]
Diplolepis is a genus of approximately fifty species of gall-inducing wasps in the family Diplolepididae. The larvae induce galls on wild roses (Rosa), and rarely on domestic roses.
The Cynipoidea are a moderate-sized hymenopteran superfamily that presently includes seven extant families and three extinct families, though others have been recognized in the past. The most familiar members of the group are phytophagous, especially as gall-formers, though the actual majority of included species are parasitoids or hyperparasitoids. They are typically glossy, dark, smooth wasps with somewhat compressed bodies and somewhat reduced wing venation. It is common for various metasomal segments to be fused in various ways, and the petiole is very short, when present.
Andricus kollari, also known as the marble gall wasp, is a parthenogenetic species of wasp which causes the formation of marble galls on oak trees. Synonyms for the species include Cynips kollari, Andricus quercusgemmae, A. minor, A. indigenus and A. circulans.
Cynips is a genus of gall wasps in the tribe Cynipini, the oak gall wasps. One of the best known is the common oak gall wasp, which induces characteristic spherical galls about two centimeters wide on the undersides of oak leaves.
Andricus is a genus of oak gall wasps in the family Cynipidae.
Andricus grossulariae is a gall wasp species inducing agamic acorn cup galls on oak tree acorn cups and sexual phase galls on catkins. Synonyms include Andricus fructuum, Andricus gemellus, Andricus intermedius, Andricus mayri and Cynips panteli.
Cynipinae is a subfamily of gall wasps (Cynipidae). Many of the approximately 1,500 described species cause galls on oaks, but some induce galls on other plant species or are inquilines of the gall-inducing species. Species occur on all continents except Antarctica, with most found in the temperate regions of the northern hemisphere. All extant cynipid species are within Cynipinae since the only other recognized subfamily is Hodiernocynipinae which is based on the fossil genus Hodiernocynips.
Cynipini is a tribe of gall wasps. These insects induce galls in plants of the beech and oak family, Fagaceae. They are known commonly as the oak gall wasps. It is the largest cynipid tribe, with about 936 to 1000 recognized species, most of which are associated with oaks. The tribe is mainly native to the Holarctic.
Andricus quercuscalifornicus, or the California gall wasp, is a small wasp species that induces oak apple galls on white oaks, primarily the valley oak but also other species such as Quercus berberidifolia. The California gall wasp is considered an ecosystem engineer, capable of manipulating the growth of galls for their own development. It is found from Washington, Oregon, and California to northern regions of Mexico. Often multiple wasps in different life stages occupy the same gall. The induced galls help establish complex insect communities, promoting the diversification in niche differentiation. Furthermore, the adaptive value of these galls could be attributed their ecological benefits such as nutrition, provision of microenvironment, and enemy avoidance.
Atrusca is a genus of gall wasps in the family Cynipidae. It consists of approximately 50 species, and is found in North and Central America.
Bassettia is a genus of gall wasps found in North America.
Synergini is a tribe of gall wasps in the subfamily Cynipinae.
Disholcaspis is a genus of gall wasps in the family Cynipidae. There are more than 40 species described in the genus Disholcaspis. Some Disholcaspis species induce galls that produce honeydew, a sweet liquid that attracts yellow jackets, ants, and bees. These insects then protect the galls from parasitic wasps.
Andricus dimorphus, also called the clustered midrib gall wasp, is a species of oak gall wasp in the family Cynipidae. Galls in which the larvae live and feed are formed in clusters along the midrib on the underside of oak leaves.
Synergus japonicus is a species of gall wasp in the family Cynipidae. Whereas most gall wasps create the galls in which they live, Synergus japonicus is an inquiline species, living in the gall created by another species of wasp. It is native to Japan, China and Russia.
Ceroptresini is a tribe of oak gall wasps in the family Cynipidae, and includes two genera: Ceroptres and Buffingtonella. All but one of the 22 species currently recognized are in Ceroptres. Ceroptresini, containing only Ceroptres, was first proposed as a tribe in 2015, and Buffingtonella was included the tribe in 2019 when the genus was first described. Species in this tribe are believed to be inquilines in galls induced by other gall wasps but this has not been confirmed.
Kokkocynips is an American genus of gall wasps in the family Cynipidae. There are about 8 described species in the genus Kokkocynips with several others still undescribed.
Andricus hispanicus is a parthenogenetic species of wasp which causes the formation of marble galls on oak trees. The galls caused by the agamous generation are similar to the closely related Andricus kollari.
Diplolepididae is a family of small gall-inducing wasps. Until recently these wasps were included in the gall wasp family (Cynipidae) but were moved to their own family based on genetic and morphological features. It contains two subfamilies: Diplolepidinae and Pediaspidinae.