Kradibiinae
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The family Agaonidae is a group of pollinating
fig wasp Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside fig syconia. Some are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, ...
s. They spend their
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
l stage inside the fruits of
fig The fig is the edible fruit of ''Ficus carica'', a species of tree or shrub in the flowering plant family Moraceae, native to the Mediterranean region, together with western and southern Asia. It has been cultivated since ancient times and i ...
s. The pollinating wasps (
Agaoninae The family Agaonidae is a group of pollinating fig wasps. They spend their larval stage inside the fruits of Ficus, figs. The pollinating wasps (Agaoninae, Kradibiinae, and Tetrapusiinae) are the mutualism (biology), mutualistic partners of the ...
, Kradibiinae, and Tetrapusiinae) are the mutualistic partners of the
fig The fig is the edible fruit of ''Ficus carica'', a species of tree or shrub in the flowering plant family Moraceae, native to the Mediterranean region, together with western and southern Asia. It has been cultivated since ancient times and i ...
trees. Extinct forms from the Eocene and Miocene are nearly identical to modern forms, suggesting that the niche has been stable over geologic time. Females emerge from ripe figs and fly to another fig tree with developing
syconia Syconium (: syconia) is the type of fruit borne by figs (genus ''Ficus''), formed by an enlarged, fleshy, hollow receptacle with multiple ovaries on the inside surface. In essence, it is really a fleshy stem with a number of flowers, so it is co ...
(which contain the flowers). They enter the syconium via the
ostiole An ''ostiole'' is a small hole or opening through which algae or fungi release their mature spores. The word is a diminutive of wikt:ostium, "ostium", "opening". The term is also used in higher plants, for example to denote the opening of the ...
, pollinate the flowers, and lay their eggs in some of the ovules. The ovules containing eggs develop into galls that support the growth of the wasp larvae. Prior to the final ripening of the fig, wingless males emerge from the galls they developed in. The males enter the galls of their winged sibling females, mate with them and die within the fruit. The newly hatched females then make their way out of the fruit continuing this cycle.


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 Pteromalidae are a large family of wasps, the majority being parasitoids of other insects. They are found throughout the world in virtually all habitats, and many are important as biological control agents. The oldest known fossil is known fr ...
, and
Agaoninae The family Agaonidae is a group of pollinating fig wasps. They spend their larval stage inside the fruits of Ficus, figs. The pollinating wasps (Agaoninae, Kradibiinae, and Tetrapusiinae) are the mutualism (biology), mutualistic partners of the ...
were the subdivisions of the family. Recent works building strong molecular
phylogenies A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA. In o ...
with an extended sampling size have changed the composition of Agaonidae. The
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
groups have been excluded; Epichrysomallinae was raised to family status (
Epichrysomallidae Epichrysomallidae is a family of gall-forming wasps associated with fig trees (genus ''Ficus'') - they make galls in figs, or on leaves or twigs. Once considered a subfamily of Pteromalidae (Epichrysomallinae), this group of genera has been eleva ...
), whereas Otitesellinae, Sycoecinae, Sycophaginae, and Sycoryctinae were transferred to
Pteromalidae The Pteromalidae are a large family of wasps, the majority being parasitoids of other insects. They are found throughout the world in virtually all habitats, and many are important as biological control agents. The oldest known fossil is known fr ...
. New subfamilies have been instated (Kradibiinae and Tetrapusiinae).


Ecology

The relationship between fig trees and agaonid fig wasps is an
obligate mutualism Obligate mutualism is a special case of mutualism where an ecological interaction between species mutually benefits each other, and one or all species are unable to survive without the other. In some obligate relationships, only one species is de ...
that has evolved over a period of about sixty million years.


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'' 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, it reaches a height of . Ants predominantly of the ...
'' 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 typical ...
in it, while the male dies after chewing a hole. As the fig is closed by a tight
ostiole An ''ostiole'' is a small hole or opening through which algae or fungi release their mature spores. The word is a diminutive of wikt:ostium, "ostium", "opening". The term is also used in higher plants, for example to denote the opening of the ...
, the female wasps have developed adaptations to enter. First, the mandibles of the female wasps have developed specialized
mandibular In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
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.


Subfamilies and genera


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

* '' Agaon'' Dalman, 1818 * '' Alfonsiella'' Waterston, 1920 * '' Allotriozoon'' Grandi, 1916 * '' Blastophaga'' 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''. These wasps breed in figs without the need for a colony or nest, and the adults l ...
'' 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 ''Pegoscapus'' is a genus of fig wasp native to the Americas. They range from Florida and Mexico in the north to Argentina in the south. Fig wasps have an obligate mutualism with the fig species they pollinate. ''Pegoscapus'' pollinates spec ...
'' Cameron, 1906 * ''
Platyscapa ''Platyscapa'' is a genus of 19 species of pollinating fig wasps found in Africa and Madagascar, Middle East, southern Asia and the Indo-Pacific islands. They are pollinators of ''Ficus'' species belonging to subsections '' Conosycea'' and '' Ur ...
'' Motschoulsky, 1863 * '' Pleistodontes'' Saunders, 1882 * '' Waterstoniella'' * '' Wiebesia'' Boucek, 1988


Kradibiinae

* ''
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)


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''. ''Tetrapus'' appears to be the sole genus of ...
''


Extinct genera

* †'' Archaeagaon'' Insect Limestone, United Kingdom,
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
(
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

{{Authority control Apocrita families Insects described in 1848