The family Agaonidae is a group of pollinating and nonpollinating fig wasps. They spend their

larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...

l stage inside the fruits of

fig The fig is the edible fruit of ''Ficus carica'', a species of small tree in the flowering plant family Moraceae. Native to the Mediterranean and western Asia, it has been cultivated since ancient times and is now widely grown throughout the world ...

s. The pollinating wasps (

Agaoninae The family Agaonidae is a group of pollinating and nonpollinating fig wasps. They spend their larval stage inside the fruits of figs. The pollinating wasps ( Agaoninae, Kradibiinae, and Tetrapusiinae) are the mutualistic partners of the fig tre ...

Kradibiinae The family Agaonidae is a group of pollinating and nonpollinating fig wasps. They spend their larval stage inside the fruits of figs. The pollinating wasps (Agaoninae, Kradibiinae, and Tetrapusiinae) are the mutualistic partners of the fig tree ...

, and Tetrapusiinae) are the mutualistic partners of the

trees. The nonpollinating fig wasps are parasitic. Extinct forms from the Eocene and Miocene are nearly identical to modern forms, suggesting that the niche has been stable over geologic time.

Taxonomy

The family has changed several times since its taxonomic appearance after the work of Francis Walker in 1846 described from the wasp genus '' Agaon''. Previously the subfamilies Epichrysomallinae, Otitesellinae, Sycoecinae, Sycoryctinae,

Sycophaginae The family Agaonidae is a group of pollinating and nonpollinating fig wasps. They spend their larval stage inside the fruits of figs. The pollinating wasps (Agaoninae, Kradibiinae, and Tetrapusiinae) are the mutualistic partners of the fig trees ...

, and

were the subdivisions of the family. Recent works building strong molecular

phylogenies A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

with an extended sampling size have changed the composition of Agaonidae. First, the

paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...

groups have been excluded (Epichrysomallinae, Otitesellinae, Sycoecinae, and Sycoryctinae) and new subfamilies have been instated (Kradibiinae and Tetrapusiinae). Then the subfamily

have been placed within the family Agaonidae. Within the Sycophaginae, some changes were made after the molecular phylogeny of the subfamily: the genus ''

Apocryptophagus ''Sycophaga'' is a mainly Afrotropical gall wasp genus of the superfamily Chalcidoidea that live on the section ''Sycomorus'' of the monoecious fig subgenus, ''Sycomorus'', and one of several fig wasp genera to exploit its mutualism with ''Cer ...

'' has been

synonymed A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...

under the genus ''

Sycophaga ''Sycophaga'' is a mainly Afrotropical gall wasp genus of the superfamily Chalcidoidea that live on the section ''Sycomorus'' of the monoecious fig subgenus, ''Sycomorus'', and one of several fig wasp genera to exploit its mutualism with ''Cera ...

''.

Ecology

Wasps from the three subfamilies

and Tetrapusiinae are pollinating fig wasps. On the other hand,

are parasites of the ''

Ficus ''Ficus'' ( or ) is a genus of about 850 species of woody trees, shrubs, vines, epiphytes and hemiepiphytes in the family Moraceae. Collectively known as fig trees or figs, they are native throughout the tropics with a few species extendi ...

'', developing in the fruits after other wasps have pollinated them. Nevertheless, some species in the genus ''

'' have a controversial status; as they enter the fig by its

ostiole An ''ostiole'' is a small hole or opening through which algae or fungi release their mature spores. The word is a diminutive of "ostium", "opening". The term is also used in higher plants, for example to denote the opening of the involuted ...

, they possibly bring

pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...

inside the fig and might pollinate it.

Morphological adaptations

The pollinating female fig wasps are winged and in general dark, while the males are mostly wingless and whitish. This difference of color is probably due to a clear split in the gender role. Once they have mated, male and female fig wasps have different fates. In some fig species, such as '' Ficus subpisocarpa'' or ''

Ficus tinctoria ''Ficus tinctoria'', also known as dye fig, or humped fig is a hemiepiphytic tree of genus ''Ficus''. It is also one of the species known as '' strangler fig''. It is found in Asia, Malesia, northern Australia, and the South Pacific islands. ...

'', the males have to chew a hole for the females to leave their natal fig. The winged female wasps can fly over long distances before finding another fig to oviposit in it, while the male dies after chewing a hole. As the fig is closed by a tight

, the female wasps have developed adaptations to enter. First, the mandibles of the female wasps have developed specialized mandibular appendages to help them crawl into the figs. These appendages are adapted to the host fig species, with for instance spiraled ostioles matched by spiral mandibular appendages. The nonpollinating wasps also have developed impressive morphological adaptations to deposit eggs inside the fig from the outside, in the form of an extremely long

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 typical ...

Subfamilies and genera

Agaoninae The family Agaonidae is a group of pollinating and nonpollinating fig wasps. They spend their larval stage inside the fruits of figs. The pollinating wasps ( Agaoninae, Kradibiinae, and Tetrapusiinae) are the mutualistic partners of the fig tre ...

* '' Agaon'' Dalman, 1818 * '' Alfonsiella'' Waterston, 1920 * '' Allotriozoon'' Grandi, 1916 * ''

Blastophaga ''Blastophaga'' is a wasp genus in the family Agaonidae (fig wasps) which pollinate figs or are otherwise associated with figs, a coevolutional relationship that has been developing for at least 80 million years. Pollinating fig wasps are specifi ...

'' Gravenhorst, 1829 ** ''

Blastophaga psenes ''Blastophaga psenes'' is a wasp species in the genus ''Blastophaga''. It pollinates the common fig ''Ficus carica'' and the closely related '' Ficus palmata''. Without a colony or nest, these wasps breed in figs and the adults live for only a f ...

'' Linnaeus ( syn. ''Cynips psenes'' Linnaeus, 1758) * '' Courtella'' Kieffer, 1911 * '' Deilagaon'' Wiebes, 1977 * '' Dolichoris'' Hill, 1967 * '' Elisabethiella'' Grandi, 1928 * '' Eupristina'' Saunders, 1882 ** '' Eupristina verticillata'' Waterston, 1921 * '' Nigeriella'' Wiebes 1974 * '' Paragaon'' Joseph, 1959 * '' Pegoscapus'' Cameron, 1906 * '' Platyscapa'' Motschoulsky, 1863 * '' Pleistodontes'' Saunders, 1882 * '' Waterstoniella'' * ''

Wiebesia ''Wiebesia'' is a genus in the family Agaonidae (fig wasps). The scientific name of this genus was first published in 1988 by Boucek. Pollinating fig wasps are specific to specific figs. '' Ficus pumila var. awkeotsang,'' the source of aiyu jell ...

'' Boucek, 1988

Kradibiinae The family Agaonidae is a group of pollinating and nonpollinating fig wasps. They spend their larval stage inside the fruits of figs. The pollinating wasps (Agaoninae, Kradibiinae, and Tetrapusiinae) are the mutualistic partners of the fig tree ...

* '' Ceratosolen'' Mayr, 1885 * '' Kradibia'' Saunders, 1883 (syn. ''Liporrhopalum'' Waterston, 1920)

Sycophaginae The family Agaonidae is a group of pollinating and nonpollinating fig wasps. They spend their larval stage inside the fruits of figs. The pollinating wasps (Agaoninae, Kradibiinae, and Tetrapusiinae) are the mutualistic partners of the fig trees ...

* '' Anidarnes'' * '' Eukoebelea'' * '' Idarnes'' * '' Pseudidarnes'' * ''

Tetrapusiinae

* '' Tetrapus''

Extinct genera

* †'' Archaeagaon'' Insect Limestone, United Kingdom,

Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...

(

Priabonian The Priabonian is, in the ICS's geologic timescale, the latest age or the upper stage of the Eocene Epoch or Series. It spans the time between . The Priabonian is preceded by the Bartonian and is followed by the Rupelian, the lowest stage ...

) ** †''Archaeagaon minutum'' (Donisthorpe)Universal Chalcidoidea Database – Archaeagaon
Natural History Museum, London

References

{{Taxonbar, from=Q995960 Apocrita families