
Gall wasps, also
traditionally called gallflies, are
hymenoptera
Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic.
Females typi ...
ns of the family Cynipidae in the wasp superfamily
Cynipoidea
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, espec ...
. Their common name comes from the
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 they induce on
plant
Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
s for larval development. About 1,300 species of this generally very small creature () are known worldwide,
with about 360 species of 36 different genera in Europe and some 800 species in North America.
Features
Like all
Apocrita
Apocrita is a suborder of insects in the order Hymenoptera. It includes wasps, bees, and ants, and consists of many families. It contains the most advanced hymenopterans and is distinguished from Symphyta by the narrow "waist" ( petiole) formed ...
, gall wasps have a distinctive body shape, the so-called
wasp waist
Wasp waist is a women's fashion silhouette, produced by a style of corset and girdle, that has experienced various periods of popularity in the 19th and 20th centuries. Its primary feature is the abrupt transition from a natural-width rib cag ...
. The first abdominal
tergum
A ''tergum'' (Latin for "the back"; : ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod
Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton wi ...
(the
propodeum
Propodeum labelled within the alitrunk of an ant worker
The propodeum is a term that can refer to unrelated structures in insects or in mollusks.
Insects
The first abdominal segment in Apocrita Hymenoptera (wasps, bees and ants) is known a ...
) is conjoined with the
thorax
The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen.
In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...
, while the second abdominal segment forms a sort of shaft, the
petiole. The petiole connects with the
gaster, which is the functional
abdomen
The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
in apocritan wasps, starting with the third abdominal segment proper. Together, the petiole and the gaster form the
metasoma
The metasoma is the posterior part of the body, or tagma (biology), tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the mesosoma. In insects, it contains most of the digestive tract, respiratory sy ...
, while the thorax and the
propodeum
Propodeum labelled within the alitrunk of an ant worker
The propodeum is a term that can refer to unrelated structures in insects or in mollusks.
Insects
The first abdominal segment in Apocrita Hymenoptera (wasps, bees and ants) is known a ...
make up the
mesosoma
The mesosoma is the middle part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the metasoma. It bears the legs, and, in the case of winged insects, the wings.
Wasps, bees and a ...
.
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
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 ...
is often seen protruding from the tip of the metasoma.
Reproduction and development
The reproduction of gall wasps is usually partly
parthenogenesis
Parthenogenesis (; from the Greek + ) is a natural form of asexual reproduction in which the embryo develops directly from an egg without need for fertilization. In animals, parthenogenesis means the development of an embryo from an unfertiliz ...
, in which a male is completely unnecessary, and partly two-sex propagation.
[The population biology of oak gall wasps (Hymenoptera: Cynipidae)](_blank)
Stone et al. (2002) Annual Review of Entomology Vol. 47: 633-668 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,
possibly because of infection of the females'
gamete
A gamete ( ) is a Ploidy#Haploid and monoploid, haploid cell that fuses with another haploid cell during fertilization in organisms that Sexual reproduction, reproduce sexually. Gametes are an organism's reproductive cells, also referred to as s ...
s by
endosymbiotic
An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), which live in the root ...
''
Wolbachia
''Wolbachia'' is a genus of gram-negative bacteria infecting many species of arthropods and filarial nematodes. The symbiotic relationship ranges from parasitism to obligate mutualism. It is one of the most common parasitic microbes of arthrop ...
'' bacteria.
The various generations differentiate both in their appearance and in the form of the
plant galls they induce.
The
larvae
A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
of most gall wasps develop in characteristic plant galls they induce themselves, but many species are instead
inquiline
In zoology, an inquiline (from Latin ''inquilinus'', "lodger" or "tenant") is an animal that lives commensally in the nest, burrow, or dwelling place of an animal of another species. For example, some organisms, such as insects, may live in the ...
s 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
An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...
, inducing
oak galls
An oak apple or oak gall is a large, round, vaguely apple-like gall commonly found on many species of oak. Oak apples range in size from in diameter and are caused by chemicals injected by the larva of certain kinds of gall wasp in the family ...
. 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
''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 ...
,
maple
''Acer'' is a genus of trees and shrubs commonly known as maples. The genus is placed in the soapberry family Sapindaceae.Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 9, June 2008 nd more or less continuously updated si ...
, and many herbs. Species determination is usually much easier through observation of the galls produced rather than of the insect itself.
Parasitism
A
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 ...
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
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 ...
) to bore into the gall. These parasitoids may, in turn, be preyed upon by other wasps,
hyperparasitoids.
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.
File:Andricus foecundatrix Quercus01.jpg,
File:Gallwespe bedient sich Eichel2.jpg,
File:Diplolepis Quercus02.jpg,
File:Eikengallen op mannelijke bloeiwijze.jpg,
File:Oak Gall.jpg,
File:Cynips longiventris gall.jpg,
File:Oak Gall Wasp.jpeg,
File:Acorn Plum Gall.jpg, Acorn Plum Gall
File:Acorn Plum Gall, cut in half, with larva.jpg, Acorn Plum Gall, cut in half, with larva
File:Common Spangle gall.JPG,
File:Hopea Ponga 21.JPG, Galls on '' Hopea ponga'' at Peravoor
File:Diastrophus nebulosus gall.jpg, Raspberry gall made by '' Diastrophus nebulosus'', larvae inside gall
These turn reddish in the fall and are commonly known as
oak apple
An oak apple or oak gall is a large, round, vaguely apple-like gall commonly found on many species of oak. Oak apples range in size from in diameter and are caused by chemicals injected by the larva of certain kinds of gall wasp in the famil ...
s. 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.
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.
Internal phylogeny
The internal phylogeny of gall wasps in the cladogram is based on the molecular phylogenetic analysis of Hearn et al. 2023.
Taxonomy
The Cynipidae contains two subfamilies, one extinct and one extant:
*
Cynipinae
*
Hodiernocynipinae†
The Cynipinae consists of nine tribes:
*
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
Synergini is a tribe (biology), tribe of gall wasps in the subfamily Cynipinae.
Genera
The following genera are generally accepted within Synergini:
* ''Agastoroxenia''
* ''Lithosaphonecrus''
* ''Saphonecrus''
* ''Rhoophilus''
* ''Synergus ...
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
Human sexuality is the way people experience and express themselves sexually. This involves biological, psychological, physical, erotic, emotional, social, or spiritual feelings and behaviors. Because it is a broad term, which has varied ...
,
Alfred Kinsey was known for his study of gall wasps.
* Galls formed on oak trees are one of the main ingredients in
iron gall ink
Iron gall ink (also known as common ink, standard ink, oak gall ink or iron gall nut ink) is a purple-black or brown-black ink made from iron salts and tannic acids from vegetable sources. It was the standard ink formulation used in Europe for t ...
.
See also
*
Oak apple
An oak apple or oak gall is a large, round, vaguely apple-like gall commonly found on many species of oak. Oak apples range in size from in diameter and are caused by chemicals injected by the larva of certain kinds of gall wasp in the famil ...
*
Oak marble gall
*
Knopper gall
*
Common spangle gall
*
Silk button gall
References
Further reading
*
*
*
External links
*
''Andricus quercusclavigera'' and ''A. quercuscornigera''on the
UF /
IFAS Featured Creatures Web site
*
Wikispecies entry
Family Cynipidaea
BugGuide
{{Taxonbar, from=Q640774
Gall-inducing insects
Asexual reproduction in animals