Fig wasps are

wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...

s of the superfamily Chalcidoidea which spend their larval stage inside fig syconia. Some are

pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female carpel, stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are ...

s but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, while the pollinators are in the family

Agaonidae 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 ...

. Pollinating fig wasps are all

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 ...

-makers, while non-pollinating fig wasps either make their own galls or usurp the galls of other fig wasps. The lifestyles of these fig wasps rely on the fruit of fig trees to reproduce, with pollinating fig wasps acting as mutualists, and non-pollinating fig wasps as

parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...

History

Aristotle Aristotle (; 384–322 BC) was an Ancient Greek philosophy, Ancient Greek philosopher and polymath. His writings cover a broad range of subjects spanning the natural sciences, philosophy, linguistics, economics, politics, psychology, a ...

recorded in his '' History of Animals'' that the fruits of the wild fig (the caprifig) contain ''psenes'' (fig wasps); these begin life as grubs (larvae), and the adult ''psen'' splits its "skin" (pupa) and flies out of the fig to find and enter a cultivated fig, saving it from dropping. He believed that the ''psen'' was generated spontaneously; he did not recognise that the fig was reproducing sexually and that the ''psen'' was assisting in that process.

Taxonomy

The fig wasps are a

polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...

group, including several lineages whose similarities are based upon their shared association with figs. In 2022, family

was updated to include only the pollinating fig wasps under a single

monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

clade. Other fig wasps are now included in the families Epichrysomallidae, Eurytomidae, Melanosomellidae, Ormyridae, Pteromalidae, and Torymidae.van Noort, S. & Rasplus, JY. 2024
Classification of fig wasps
www.figweb.org (Accessed on 19 Dec 2024) Non-pollinating fig wasps represent a much more diverse taxon, distantly related from their pollinating cousins.

Morphological adaptations

In the Agaonidae, the female (as in most Hymenoptera) has four wings, whereas the males are wingless. The primary functions of agaonid males are to mate with the females while still within the fig syconium (inverted flower) and to chew a hole for the females to escape from the fig interior. This is the reverse of sex-linked functions in Strepsiptera and bagworms, where the male has wings and the female never leaves the host. The non-pollinating fig wasps have developed several impressive morphological adaptations in order to oviposit eggs within the fig syconium. Many species have extremely long ovipositors, so that they can deposit eggs from the outside of the syconium (Subtribe Sycoryctina of Otitesellinivan Noort, S. & Rasplus, JY. 2024
Sycoryctina
www.figweb.org (Accessed on 19 Dec 2024) and Subfamily Sycophaginaevan Noort, S. & Rasplus, JY. 2024
Sycophaginae
www.figweb.org (Accessed on 19 Dec 2024)). Others have evolved to enter the syconium in the same way as the Agaonidae, and now resemble the pollinators morphologically (Subtribe Sycoecina of Otitesellini).van Noort, S. & Rasplus, JY. 2024
Sycoecina
www.figweb.org (Accessed on 19 Dec 2024) Less is known about the evolution of non-pollinating fig wasps who form different clades from various lineages, each independently colonized the syconium. These wasps work around the mutualistic relationship, exploiting fig fruits as parasitoids. Most figs (more than 600 species) have syconia that contain three types of

flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...

s: male, short female, and long female. Female fig wasps can reach the ovaries of short female flowers with their ovipositors, but not long female flowers. Thus, the short female flowers grow wasps, and the long flowers only seeds. Contrary to popular belief, ripe figs are not full of dead wasps and the "crunchy bits" in the fruit are only seeds. The fig actually produces an enzyme called ficain (also known as ficin) which digests the dead wasps and the fig absorbs the nutrients to create the ripe fruits and seeds. Several commercial and ornamental varieties of fig are parthenocarpic and do not require pollination to produce (sterile) fruits; these varieties need not be visited by fig wasps to bear fruit.

Life cycle

The life cycle of the fig wasp is closely intertwined with that of the fig tree it inhabits. The wasps that inhabit a particular tree can be divided into two groups; pollinating and non-pollinating. The pollinating wasps are part of an obligate nursery pollination mutualism with the fig tree, while the non-pollinating wasps feed off the plant without benefiting it. The life cycles of the two groups, however, are similar.van Noort, S. & Rasplus, JY. 2024
Interaction of figs and fig wasps
www.figweb.org (Accessed on 19 Dec 2024) Though the lives of individual species differ, a typical pollinating fig wasp life cycle is as follows. At the beginning of the cycle, a mated mature female pollinator wasp enters the immature "fruit" (actually a stem-like structure known as a syconium) through a small natural opening (the ostiole) and deposits her eggs in the cavity. Forcing her way through the ostiole, the mated mature female often loses her wings and most of her antennae. To facilitate her passage through the ostiole, the underside of the female's head is covered with short spines that provide purchase on the walls of the ostiole. In depositing her eggs, the female also deposits pollen she picked up from her original host fig. This pollinates some of the female flowers on the inside surface of the fig and allows them to mature. After the female wasp lays her eggs and follows through with pollination, she dies. After pollination, there are several species of non-pollinating wasps that deposit their eggs before the figs harden. These wasps act as parasites to either the fig or possibly the pollinating wasps. As the fig develops, the wasp eggs hatch and develop into

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 ...

e. After going through the pupal stage, the mature male’s first act is to mate with a female - before the female hatches. Consequently, the female will emerge pregnant. The males of many species lack wings and cannot survive outside the fig for a sustained period of time. After mating, a male wasp begins to dig out of the fig, creating a tunnel through which the females escape. Once out of the fig, the male wasps quickly die. The females find their way out, picking up pollen as they do. They then fly to another tree of the same species, where they deposit their eggs and allow the cycle to begin again.

Coevolution

The fig–wasp mutualism originated between 70 and 90 million years ago as the product of a unique evolutionary event.Herre et al. (2008) Since then, cocladogenesis and coadaptation on a coarse scale between wasp genera and fig sections have been demonstrated by both morphological and molecular studies. This illustrates the tendency towards coradiation of figs and wasps. Such strict cospeciation should result in identical

phylogenetic tree 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 ...

s for the two lineages and recent work mapping fig sections onto molecular phylogenies of wasp genera and performing statistical comparisons has provided strong evidence for cospeciation at that scale. Groups of genetically well-defined pollinator wasp species coevolve in association with groups of genetically poorly defined figs.Machado C.A., Robbins N., Gilbert M.T.P., Herre E.A. Critical review of host specificity and this coevolutionary implications in the fig/fig-wasp mutualism. (2005). ''Proc. Of the National Acad. Of Sci. of the U.S.A''.102(1), 6558-6565. doi: 10.1073/pnas.0501840102 The constant hybridization of the figs promotes the constant evolution of new pollinator wasp species. Host switching and pollinator host sharing may contribute to the incredible diversity of figs and fig wasp species like '' Pegoscapus'' as they result in hybridization and introgression''.''

Conservation

Conservation efforts aim to control the populations often times targeting figs and fig wasps separately in order to develop strategies that are distinct for each species. Because many of these mutualist interactions are species specific it makes it difficult for conservationists to focus on the group at large, rather tackling individual populations with high concern. There is already heavily studied control mechanisms in figs that control for wasp populations. The current focus in the field is the conservation of fig wasp species as the role of

s is steadily declining with climate change. Because many of these species have coevolved together through generations the main aim of conservation strategies is that protection of one species in the mutualism in turn affects the other, so by developing strategies to protect threatened wasp populations, the species of fig associated with it will also be impacted. Many figs are also keystone species in their environment, being food sources and homes for a wide range of species. Fig wasps are obligate mutualists with their respective fig species, not being able to survive without each other. The loss of a pollinator wasp would result in the decline of a fig species, resulting in the general decline in the

habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...

. These reasons are why fig wasps have become a main focus among conservationists with the aim of protecting crucial keystone fig species.

Genera

Fig wasp genera and classification: *

** Agaoninae ***'' Agaon'' ***'' Alfonsiella'' ***'' Allotriozoon'' ***'' Courtella'' ***'' Elisabethiella'' ***'' Nigeriella'' ***'' Paragaon'' ***'' Pleistodontes'' ** Blastophaginae ***'' Blastophaga'' ***'' Valisia'' ** Ceratosolen ***'' Ceratosolen'' ***'' Rothropus'' ***'' Strepitus'' ** Deilagaon ** Dolichoris ** Eupristina ***'' Eupristina'' ***'' Parapristina'' ** Kradibiinae ***'' Kradibia'' ** Liporrhopalum ** Pegoscapus ** Platyscapa ** Waterstoniella ** Wiebesia ** Tetrapusiinae ***'' Tetrapus'' * Epichrysomallidae **'' Acophila'' **'' Asycobia'' **'' Camarothorax'' **'' Epichrysomalla'' **'' Eufroggattia'' **'' Herodotia'' **'' Lachaisea'' **'' Meselatus'' **'' Neosycophila'' **'' Odontofroggatia'' **'' Parapilkhanivora'' **'' Sycobia'' **'' Sycobiomorphella'' **'' Sycomacophila'' **'' Sycophilodes'' **'' Sycophilomorpha'' **'' Sycotetra'' * Pteromalidae ** Colotrechinae ***'' Podvina'' ** Pteromalinae ***'' Adiyodiella'' ***'' Apocrypta'' ***'' Arachonia'' ***'' Bouceka'' ***'' Comptoniella'' ***'' Critogaster'' ***'' Crossogaster'' ***'' Diaziella'' ***'' Dobunabaa'' ***'' Eujacobsonia'' ***'' Ficicola'' ***'' Gaudalia'' ***'' Grandiana'' ***'' Grasseiana'' ***'' Hansonita'' ***'' Lipothymus'' ***''

Marginalia Marginalia (or apostils) are marks made in the margin (typography), margins of a book or other document. They may be scribbles, comments, gloss (annotation), glosses (annotations), critiques, doodles, drolleries, or illuminated manuscript, ...

'' ***'' Micranisa'' ***'' Micrognathophora'' ***'' Otitesella'' ***'' Parasycobia'' ***'' Philocaenus'' ***'' Philosycus'' ***'' Philosycella'' ***'' Philotrypesis'' ***'' Philoverdance'' ***'' Robertsia'' ***'' Seres'' ***'' Sycoecus'' ***'' Sycoscapter'' ***'' Walkerella'' ***'' Watshamiella'' ** Sycophaginae ***'' Anidarnes'' ***'' Eukoebelea'' ***'' Idarnes'' ***'' Pseudidarnes'' ***'' Sycophaga'' * Ormyridae **'' Ormyrus'' * Eurytomidae **'' Bruchophagus'' **'' Eurytoma'' **'' Ficomila'' **'' Syceurytoma'' **'' Sycophila'' * Torymidae **'' Megastigmus'' **'' Physothorax'' **'' Torymus''

Museum collections

One of the world's major fig wasp collections resides in Leeds Museums and Galleries' Discovery Centre, and was collected by Dr. Steve Compton.

References

Sources

* * * * * * * *

External links

Images of fig wasps on Morphbank, a biological image database
{{Hymenoptera, 2 Agaonidae Chalcidoidea Insect common names