The family Agaonidae is a group of pollinating and nonpollinating

fig wasp Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside figs. Most are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, while ...

s. 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. Th ...

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 tree ...

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 ...

, 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, 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 (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

groups have been excluded (Epichrysomallinae, Otitesellinae, Sycoecinae, and Sycoryctinae) and new subfamilies have been instated (Kradibiinae and Tetrapusiinae). Then the subfamily Sycophaginae have been placed within the family Agaonidae. Within the Sycophaginae, some changes were made after the molecular phylogeny of the subfamily: the genus '' Apocryptophagus'' has been synonymed 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, Sycophaginae 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 extending int ...

'', 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 ''Ficus subpisocarpa'' (called 笔管榕 in China and 雀榕 in Taiwan) is a species of small deciduous tree native to Japan, China, Taiwan and southeast Asia to the Moluccas (Ceram). Two subspecies are recognised. Terrestrial or hemiepiphytic, i ...

'' 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. It ...

'', 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 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 ...

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 In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...

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 typica ...

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 tree ...

* '' Agaon'' Dalman, 1818 * '' Alfonsiella'' Waterston, 1920 * '' Allotriozoon'' Grandi, 1916 * '' Blastophaga'' Gravenhorst, 1829 ** '' Blastophaga psenes'' Linnaeus (

syn. The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...

''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 ''Pleistodontes '' is a genus of fig wasps native to Australia and New Guinea, with one species (''P. claviger'') from Java. Fig wasps have an obligate mutualism with the fig species they pollinate. ''Pleistodontes '' pollinates species in sec ...

'' 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 je ...

'' 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 ...

* ''

Ceratosolen ''Ceratosolen'' is an Old World wasp genus in the family Agaonidae (fig wasps). They are pollinators of the monoecious fig subsections ''Sycomorus'' and ''Sycocarpus'', and the section ''Neomorphe'', all belonging to the subgenus ''Sycomorus''. ...

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

Sycophaginae

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

Tetrapusiinae

* ''

Tetrapus ''Tetrapus'' is a genus of fig wasp native to the Americas. Fig wasps have an obligate mutualism with the fig species they pollinate. ''Tetrapus'' pollinates figs in the subgenus ''Pharmacosycea ''Pharmacosycea'' is one of six subgenera ...

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