Fig wasp

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Fig wasps
Blastophaga psenes.jpg
Blastophaga psenes female
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Suborder: Apocrita
Infraorder: Proctotrupomorpha
Superfamily: Chalcidoidea

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 the pollinators are in the family Agaonidae. While pollinating fig wasps are gall-makers, the remaining types either make their own galls or usurp the galls of other fig wasps; reports of their being parasitoids are considered dubious. [1]

Contents

History

Aristotle 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. [2]

Taxonomy

The fig wasps are a polyphyletic group, including several unrelated lineages whose similarities are based upon their shared association with figs; efforts are underway to resolve the matter, and remove a number of constituent groups to other families, particularly the Pteromalidae and Torymidae. Thus, the number of genera in the family is in flux. The family Agaonidae has been recently updated to include all the pollinating fig wasps [3] and the subfamily Sycophaginae. [4] The remaining taxa such as Epichrysomallinae, Sycoecinae, Otitesellinae, and Sycoryctinae should be included in the Pteromalidae. [4]

Morphological adaptations

Female (left, with long ovipositor) and male Blastophaga psenes Blastophaga psenes2.png
Female (left, with long ovipositor) and male Blastophaga psenes

Among the Agaonidae, the female is a normal insect, while the males are mostly wingless. The males' only tasks 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 Strepsiptera and the bagworm, where the male is a normal insect and the female never leaves the host. The non-pollinating wasps have developed impressive morphological adaptations in order to oviposit eggs inside the fig but from the outside: an extremely long ovipositor.[ citation needed ]

Most figs (more than 600 species) have syconia that contain three types of flowers: 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. [5] 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. [6]

Life cycle

Pleistodontes sp. female Pleistodontes sp-Female.jpg
Pleistodontes sp. female
Ceratosolen species are pollinators of the Sycomorus, Sycocarpus and Neomorphe sections of Ficus. Ceratosolen capensis op Ficus sur, Jan Celliers Park, a.jpg
Ceratosolen species are pollinators of the Sycomorus, Sycocarpus and Neomorphe sections of Ficus .
Non-pollinating parasitoid wasps Apocrypta ovipositing on Ficus sur in South Africa Apocrypta guineensis, volw-wyfie op F sur, Manie vd Schijff BT, d.jpg
Non-pollinating parasitoid wasps Apocrypta ovipositing on Ficus sur in South Africa

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 very similar.

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 larvae. 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. [8] [9] [10] Since then, cocladogenesis and coadaptation on a coarse scale between wasp genera and fig sections have been demonstrated by both morphological and molecular studies. [10] [11] This illustrates the tendency towards coradiation of figs and wasps. [10] Such strict cospeciation should result in identical phylogenetic trees for the two lineages [9] 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. [9]

Groups of genetically well-defined pollinator wasp species coevolve in association with groups of genetically poorly defined figs. [12] 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. [12]

Genera

Fig wasps genera and classification according to the various publications: [3] [4] [13]

Museum collections

One of the world's major fig wasp collections resides in Leeds Museums and Galleries' Discovery Centre, [14] and was collected by Dr. Steve Compton. [15] [16]

Related Research Articles

<span class="mw-page-title-main">Ovipositor</span> Anatomical structure for laying eggs

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 typically its form is adapted to functions such as preparing a place for the egg, transmitting the egg, and then placing it properly. For most insects, the organ is used merely to attach the egg to some surface, but for many parasitic species, it is a piercing organ as well.

<i>Ficus</i> Genus of flowering plants in the mulberry family Moraceae

Ficus 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 into the semi-warm temperate zone. The common fig (F. carica) is a temperate species native to southwest Asia and the Mediterranean region, which has been widely cultivated from ancient times for its fruit, also referred to as figs. The fruit of most other species are also edible though they are usually of only local economic importance or eaten as bushfood. However, they are extremely important food resources for wildlife. Figs are also of considerable cultural importance throughout the tropics, both as objects of worship and for their many practical uses.

<span class="mw-page-title-main">Chalcid wasp</span> Superfamily of wasps

Chalcid wasps are insects within the superfamily Chalcidoidea, part of the order Hymenoptera. The superfamily contains some 22,500 known species, and an estimated total diversity of more than 500,000 species, meaning the vast majority have yet to be discovered and described. The name "chalcid" is often confused with the name "chalcidid", though the latter refers strictly to one constituent family, the Chalcididae, rather than the superfamily as a whole; accordingly, most recent publications (e.g.,) use the name "chalcidoid" when referring to members of the superfamily.

<span class="mw-page-title-main">Agaonidae</span> Family of wasps

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 are the mutualistic partners of the fig trees. The non-pollinating fig wasps are parasitoids. Extinct forms from the Eocene and Miocene are nearly identical to modern forms, suggesting that the niche has been stable over geologic time.

<span class="mw-page-title-main">Leucospidae</span> Group of wasps

The Leucospidae are a specialized group of wasps within the superfamily Chalcidoidea, that are ectoparasitoids of aculeate wasps or bees. They are typically mimics of bees or stinging wasps, often black with yellow, red, or white markings, sometimes metallic, with a robust mesosoma and very strong sculpturing. The hind femora are often greatly enlarged, with a row of teeth or serrations along the lower margin as in Chalcididae. The wing has a longitudinal fold. The female ovipositor is sometimes short, but if not, it is recurved and lies along the dorsal side of the metasoma, a unique feature. The males are also unusual, in the fusion of many of the metasomal segments to form a capsule-like "carapace".

<i>Ficus pumila <span style="font-style:normal;">var.</span> awkeotsang</i> Variety of climbing fig

Ficus pumila var. awkeotsang, the jelly fig, is a variety of Ficus pumila, and a member of the fig family Moraceae. Awkeotsang is Taiwanese, named after the daughter (Aiyu) of the individual who discovered gel-forming properties in 1921.

<i>Ficus aurea</i> Species of strangler fig

Ficus aurea, commonly known as the Florida strangler fig, golden fig, or higuerón, is a tree in the family Moraceae that is native to the U.S. state of Florida, the northern and western Caribbean, southern Mexico and Central America south to Panama. The specific epithet aurea was applied by English botanist Thomas Nuttall who described the species in 1846.

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 species in section Americana of the subgenus Urostigma.

Ficus maxima is a fig tree which is native to Mexico, Central America, the Caribbean and South America south to Paraguay. Figs belong to the family Moraceae. The specific epithet maxima was coined by Scottish botanist Philip Miller in 1768; Miller's name was applied to this species in the Flora of Jamaica, but it was later determined that Miller's description was actually of the species now known as Ficus aurea. To avoid confusion, Cornelis Berg proposed that the name should be conserved for this species. Berg's proposal was accepted in 2005.

<i>Ficus americana</i> Species of fig tree native to the Neotropics

Ficus americana, commonly known as the West Indian laurel fig or Jamaican cherry fig, is a tree in the family Moraceae which is native to the Caribbean, Mexico in the north, through Central and South America south to southern Brazil. It is an introduced species in Florida, USA. The species is variable; the five recognised subspecies were previously placed in a large number of other species.

<i>Ficus obliqua</i> A tree, the small-leaved fig

Ficus obliqua, commonly known as the small-leaved fig, is a tree in the family Moraceae, native to eastern Australia, New Guinea, eastern Indonesia to Sulawesi and islands in the southwestern Pacific Ocean. Previously known for many years as Ficus eugenioides, it is a banyan of the genus Ficus, which contains around 750 species worldwide in warm climates, including the edible fig. Beginning life as a seedling, which grows on other plants (epiphyte) or on rocks (lithophyte), F. obliqua can grow to 60 m (200 ft) high and nearly as wide with a pale grey buttressed trunk, and glossy green leaves.

<i>Pleistodontes froggatti</i> Species of wasp

Pleistodontes froggatti is a species of fig wasp which is native to Australia. It has an obligate mutualism with the Moreton Bay Fig, Ficus macrophylla, the species it pollinates. Outside Australia, populations have become established in Hawaii and New Zealand where it was either accidentally introduced or arrived by long-distance dispersal.

Trisecodes is a genus of parasitic chalcid wasps of the family Systasidae. The genus was originally placed in Eulophidae, based on a number of morphological features, but molecular evidence suggests that the genus is more closely related to Systasis and Semiotellus. The type species is a parasitoid of a range of Agromyzid leaf-mining flies.

<i>Blastophaga psenes</i> Species of wasp

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 few days or weeks. They locate the fig they wish to pollinate through olfactory senses.

<i>Ceratosolen</i> Genus of wasps

Ceratosolen is an Old World wasp genus in the family Agaonidae. They are pollinators of the monoecious fig subsections Sycomorus and Sycocarpus, and the section Neomorphe, all belonging to the subgenus Sycomorus. The genus is native to the Palearctic, Afrotropical, Indomalayan and Australasian realms.

<i>Apocrypta</i> Genus of wasps

Apocrypta is an Old World genus of parasitic fig wasps in the family Pteromalidae. They are parasitoids of gall-wasps in the Sycophagini tribe, and especially Ceratosolen species, pollinators of the Sycomorus, Sycocarpus and Neomorphe sections of Ficus. They seem to be fig species-specific.

Robertsia is a genus of fig wasps in the family Pteromalidae, native to Papua New Guinea.

Reproductive coevolution in <i>Ficus</i>

The genus Ficus is composed of 800 species of vines, shrubs, and trees, defined by their syconiums, the fruit-like vessels that either hold female flowers or pollen on the inside. In addition to being cultivated by humans for thousands of years, Ficus is also known for their reproductive mutualism with the fig wasp.

Wiebesia pumilae is a fig wasp species in Genus Wiebesia, Family Agaonidae. W. pumilae is the pollinator of Ficus pumila var. awkeotsang and Ficus pumila var. pumila. The scientific name was first published as Blastophaga pumilae in 1967 by Hill.

Podagrion mantis was first described in 1886 by W.H. Ashmead, and was the first species of its genus to have been described from the United States. All species in the genus are parasitoid wasps known only to parasitize mantids and have been observed most often utilizing the egg case (ootheca) of Stagomantis carolina, but have also been reported to choose S.limbata or Tenodera augustipennis as hosts, showing a high degree of specialization. The ootheca is made up of tanning agents and structural proteins that, while being deposited, is liquidy and tacky, but soon hardens into a stiff foam. The female Podagrion can parasitize the ootheca in either the fresh or hardened state. The female uses her long ovipositor to deposit an egg into the ootheca; as the eggs are developing they consume the mantid larvae before emerging as an adult. The cumulative effect of several generations of parasitoid wasps on ootheca are estimated to cause over 30% loss of developing mantids and there is a clear negative correlation between parasitoid burden and mantid eclosure.

References

  1. Boucek, Z. 1988. Australasian Chalcidoidea (Hymenoptera): a biosystematic revision of genera of fourteen families, with a reclassification of species. C.A.B. International, Wallingford, England. 832 pp.
  2. Leroi, Armand Marie (2014). The Lagoon: How Aristotle Invented Science. Bloomsbury. pp. 244–247. ISBN   978-1-4088-3622-4.
  3. 1 2 Cruaud et al. 2010
  4. 1 2 3 Heraty et al. 2013
  5. "Figs Without Wasps?". 2012-10-11.
  6. Roy, D. (2019). Tropical/subtropical fruit crops: Fig. In Breeding of fruit crops (pp. 113-115). Alpha Science International Ltd.
  7. Zhen, Wen-Quan; Huang, Da-Wei; Xiao, Jin-Hua; Yang, Da-Rong; Zhu, Chao-Dong; Xiao, Hui (April 2005). "Ovipositor length of threeApocrypta species: Effect on oviposition behavior and correlation with syconial thickness" (PDF). Phytoparasitica. 33 (2): 113–120. doi:10.1007/BF03029967. S2CID   35479915 . Retrieved 5 October 2015.
  8. Machado et al. 2001
  9. 1 2 3 Cook & Rasplus 2003
  10. 1 2 3 Herre et al. (2008)
  11. Molbo et al. 2003
  12. 1 2 Machado et al. 2005
  13. Cruaud et al. 2011
  14. "Natural Science – Leeds Museums and Galleries". Leeds Museums and Galleries. Retrieved 19 August 2020.
  15. Compton, Steve. "Dr Steve Compton" . Retrieved 19 August 2020.
  16. Compton, S (2018). "Host‐parasitoid relationships within figs of an invasive fig tree: a fig wasp community structured by gall size" (PDF). Insect Conservation and Diversity. 11 (4): 341–351. doi:10.1111/icad.12282. S2CID   89701549.

Sources

Further reading