Blastophaga psenes

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Blastophaga psenes
Blastophaga psenes.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Agaonidae
Genus: Blastophaga
Species:
B. psenes
Binomial name
Blastophaga psenes
Synonyms [2]
  • Blastophaga grossorum Gravenhorst, 1827
  • Blastophaga vaidiJoseph, 1954
  • Cynips psenesLinnaeus, 1758
Winged female and wingless male Blastophaga psenes2.png
Winged female and wingless male

Blastophaga psenes is a wasp species in the genus Blastophaga . It pollinates the common fig Ficus carica and the closely related Ficus palmata . [3] These wasps breed in figs without the need for a colony or nest, and the adults live for only a few days or weeks. [4] They locate the fig they wish to pollinate primarily using through olfaction. [5]

Contents

Taxonomy and phylogenetics

Mutualism occurs between figs and fig wasps, which creates a need for specific species of figs to be pollinated by specific species of wasps. The origin of mutualism is also the beginning of the fig wasp phylogeny. In the phylogenetic tree, the genus of Blastophaga and Wiebesia are very similar. Both of these genera pollinate Ficus genus of figs. [6]

Description and identification

B. psenes are small wasps, approximately only 2 millimetres (0.079 in) in length. [7] The females are black and shiny, while the males are smaller than the females. [8] The males are wingless, whereas females have wings that are transparent and extremely thin. When the female wasps enter the opening of a fig, their wings and antennae detach. [9] Upon dissecting a fig, the wings of the wasps can be seen at the opening of the fig. Additionally, adult wasps, larvae, and eggs are found within the fig. [7] The wasps are free-living and their lifespan spans from a few days to weeks. [4]

Distribution and habitat

Because B. psenes relies on Ficus carica to breed, it is found in regions where this fig species grows. The wasp is native to the Palaearctic, including Southern Europe near the Mediterranean Basin. B. psenes has been widely introduced to facilitate Ficus carica fruit bearing. [3]

Life cycle

Adult B. psenes only live for a up to a couple of weeks. These wasps lay fertilized eggs in female flowers of the syconium of a F. carica fig. When the larvae hatch, they develop in the fig ovaries, creating a gall. The larvae become adults around the same time that male fig flowers are ready to produce pollen. When an adult wasp is mature, it mates with another wasp within the syconium. After mating, females emerge from the fig and search for a new nearby fig in which to lay their eggs. The female then oviposits into a new syconium. From there, B. psenes will find another fig to repeat the process again. [4]

Mutualism

Each species of fig is pollinated by a specific species of fig wasp. This kind of interaction between these two different organism is known as mutualism. [6] Both of them live in close physical association and benefit from one another. The fig serves as a "nest" for fig wasps, a place to lay and grow their eggs. For the fig, the fig wasps act as agents of pollination where pollen is carried to other plants for reproduction. Specifically, B. psenes has a mutualistic relationship with the fig species F. carica. This fig can only be pollinated by the symbiotic wasp that has retrieved pollen from another syconium. Female wasps oviposit in the syconium for hatching. When these larvae emerge as adults, they carry the pollen accumulated in the syconium out of the fig to another nearby syconium. Thus, when wasps lay their eggs, they are also actually pollinating that syconium. A major difference between male and female fig trees can also be observed. [4]

Male trees

Male trees contain female flowers with short styles. This is beneficial for wasps which do not have very long ovipositors and can only parasitize ovaries of female flowers with short styles found only on male trees. All female flowers on male trees with parasitized ovaries means the ovaries contain wasp eggs and produce larva instead of seeds. On the other hand, female flowers on male trees with non-parasitized ovaries will produce seeds, helping in pollination and reproduction of that flower. [4]

Female trees

Female flowers on female trees have long styles. [4] Therefore, wasps cannot parasitize these ovaries because their ovipositors are too short to reach the bottom of the syconium. Because wasp eggs cannot be laid in these female flowers, all female flowers on female trees produce seeds instead of larvae. This feature of female flowers on female trees is the explanation why it is nearly impossible for wasps to emerge from a female fig tree. Fig wasps cannot perform oviposition in there and they can be dangerously stuck in that syconium. [4]

Difference between winter and spring caprifigs

There is also a difference in winter and spring caprifigs (male figs) and their availability to receive eggs and become pollinated. [4] Spring caprifigs usually produce more wasps than winter caprifigs because of better and more resources available for them. This implies that the fig wasp population is much more active and larger in the springtime. The spring and winter caprifigs have a life cycle related to each other as to maximize resources and output of figs and wasps. Winter, or delayed, caprifigs are usually observed to occur on male trees. Spring, or undelayed, caprifigs usually occur on female trees. Because female trees are lethal, wasps prefer these delayed caprifigs of male trees. [4]

Chemoattraction

Once fig wasps emerge from the syconium, they must determine the fig in which they will deposit their eggs. [5] They need to make sure that the fig they find is available and acceptable for breeding. In the case of B. psenes, olfactory stimuli guide the wasp to a fig that is available to receive wasp eggs. This olfactory stimulus is specific for the wasps' host fig (in this case, F. carica) and enables the wasps to distinguish between their host and other fig species. [5] Figs in their receptive phase emit a compound called pentane that can attract B. psenes from at least 5 meters away. Upon sensing these signals from a specific syconium, the wasp will approach that fig. Before entering, the wasps will assess the fig by holding up their heads and antennae next to the opening of the syconium (the ostiole—where the actual attractive substances come from). If a wasp detects the signal, it will lower its antennae and search for the entrance to the fig. By using its sense of smell and taste, the wasp can trace the actual entrance into a fig once the desired fig has been located. If the wasp does not detect a signal, it will not enter the fig. Instead, it will move on and search for another receptive fig. Due to the difference between male and female trees, male tree figs are more attractive than female tree figs.[ citation needed ]

Mating

Males emerge first from their cocoon and begin searching for females to mate with. Sometimes mating occurs before the female has finished emerging from its cocoon. Males then start enlarging the fig's opening. Some fall from the fig to the ground. They have no wings and die shortly after. The enlarged opening enables the females to leave the syconium in search of a new one in which to oviposit. Mating occurs within the syconium and laying eggs occurs in the synconium of a different fig. [1]

Kin selection

Kin

The wasps breed inside the fig. Later, the female lays its eggs in the ovaries of another fig by sticking its ovipositor in each flower's style. This can lead to some flowers not being pollinated because some styles are too long. Each larva from a deposited egg destroys a female flower when it feeds on its growing seed. When wasps emerge from the syconium, they rush to the nearest syconia. This rush creates a large number of wasps all competing to enter an adjacent syconium. Due to this rush, pollination will become less effective as more pollen falls off of the wasp bodies. [4] Offspring number depends on number of pollinator wasps per syconium. The number of offspring is low when the entry number of wasps in a syconium is high. [9] On average, each wasp has 3 broods a year; one for each of the different seasonal caprifigs. [7] Also, reproductive success depends heavily on transmission of strong signals by plant. [5]

Interactions with other species

Diet

When they are hatched, B. psenes larvae feed on hyperplastic floret tissue. The mother produces this hyperplastic tissue when she lays the eggs in the syconium. [1]

Predation

One of the main predators of these wasps is ants. [10] Ants find these wasps using chemical signals such as odor cues. The ants use the fig-fig wasp mutualism to find the fig wasps by detecting an odor that comes from the figs of the male trees. They know that most fig wasps are located on male fig trees, so they use that relationship to prey on wasps. This concept is called associative learning of odor because the ants are indirectly finding these wasps by associating the smell of the fig with the wasps. However, some ants do not respond to the odor of figs for different reasons. For instance, the fig could be a non-pollinator and therefore not release any chemical substance. In the case where the ants cannot detect odors, the wasps will not be predated upon. [10]

Parasitism

B. psenes are parasitized by a nematode Schistonchus caprifici. [1] These parasites are carried in the hemocoel of the female wasp. When a B. psenes wasp oviposits her egg inside the syconium, nematodes are also deposited.These nematodes then invade, feed, and reproduce inside the floret tissues. Larvae finish development with nematode still inside the hemocoel. After fertilization, females emerge from a syconium with nematodes still in hemocoel along with pollen flakes along her body. Because this nematode is primarily found in the hemocoel of a female wasp, males are not associated with nematodes. B. psenes is a very efficient host of these nematodes. [1]

Interactions with kleptoparasitic species

B. psenes also is sometimes associated with, Philotrypesis caricae, a non-fig pollinating wasp, which is sometimes referred to as a kleptoparasite of B. psenes. [11] Unlike B. psenes which oviposit in the syconium, P. caricae oviposit at the outside of the florets and travel to multiple figs. Larvae of P. caricae also eat the galled fig tissue, so it is not clear if this interaction is a true kleptoparasitic relationship. [11] [12]

Disease

Some wasps carry a disease that is carried by F. carica trees. [9] This disease is a fungus called Fusarium moniliforme , or fig endosepsis. The wasps carry this disease on their wings and body. Because the fungus grows on the ostiole, the fungus is transmitted to the wasps' bodies when the wasp emerges from the syconium through the ostiole. Fig endosepsis is not transmitted transovarily by the fig wasp. The wasps become contaminated with spores of the fungus as they contact plant surfaces upon emergence. Studies show that wasps on upper surfaces of the leaves were infected with this fungus in higher levels than other wasps. Wasps who were higher up in the tree or further out on a branch also showed more fungus on their wings and bodies. This led to the conclusion that contamination increases as the wasps walk on leaves, petioles, and fruits before they reach the opening to the syconium. This fungus affects both males and females. The fungus shows to be more evident in spring caprifigs that are pollinated with 5 to 10 winter caprifigs than when spring caprifigs are pollinated with only one winter caprifig. Also, the incidences[ spelling? ] of this fungus are higher when there is a high population of wasps with limited figs. The more wasps that pass through one ostiole, the more likely the wasp will contract F. moniliforme. [9]

Related Research Articles

<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">Fig wasp</span> Group of mostly pollinating insects whose larvae live in figs

Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside figs. Some 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.

<i>Ficus rubiginosa</i> Species of flowering plant in the family Moraceaea native to eastern Australia

Ficus rubiginosa, the rusty fig or Port Jackson fig, is a species of flowering plant native to eastern Australia in the genus Ficus. Beginning as a seedling that grows on other plants (hemiepiphyte) or rocks (lithophyte), F. rubiginosa matures into a tree 30 m (100 ft) high and nearly as wide with a yellow-brown buttressed trunk. The leaves are oval and glossy green and measure from 4 to 19.3 cm long and 1.25 to 13.2 cm wide.

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

The family Agaonidae is a group of pollinating fig wasps. They spend their larval stage inside the fruits of figs. The pollinating wasps are the mutualistic partners of the fig trees. Extinct forms from the Eocene and Miocene are nearly identical to modern forms, suggesting that the niche has been stable over geologic time.

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Ficus citrifolia, also known as the shortleaf fig, giant bearded fig, Jagüey, wild banyantree and Wimba tree, is a species of banyan native to southern Florida, the Caribbean, Mexico, Central America, and northern South America south to Paraguay. It is distinguished from the closely related Florida strangler fig mainly by the finer veining in the leaves.

<span class="mw-page-title-main">Syconium</span> Type of fruit produced by plants of the genus Ficus

Syconium is the type of fruit borne by figs, 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 considered both a multiple and accessory fruit.

<span class="mw-page-title-main">Entomophily</span> Form of pollination by insects

Entomophily or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, sometimes with conspicuous patterns leading to rewards of pollen and nectar; they may also have an attractive scent which in some cases mimics insect pheromones. Insect pollinators such as bees have adaptations for their role, such as lapping or sucking mouthparts to take in nectar, and in some species also pollen baskets on their hind legs. This required the coevolution of insects and flowering plants in the development of pollination behaviour by the insects and pollination mechanisms by the flowers, benefiting both groups. Both the size and the density of a population are known to affect pollination and subsequent reproductive performance.

<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.

<i>Pegoscapus</i> Genus of wasps

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 insipida</i> Species of fig tree from the Neotropics

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<span class="mw-page-title-main">Fig</span> Species of flowering plant known as the common 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 region, together with western and southern Asia. It has been cultivated since ancient times and is now widely grown throughout the world. Ficus carica is the type species of the genus Ficus, containing over 800 tropical and subtropical plant species.

<i>Ficus hispida</i> Species of tropical fig tree

Ficus hispida, also known as the opposite leaf Fig, is a small tree in the family Moraceae, with a distribution ranging from India and southern China southwards to northern Australia. It is morphologically gynodioecious, but functionally dioecious. Male trees are hermaphrodites with both staminate flowers that produce pollen and pistillate flowers that produce almost no seeds but can form galls containing pollinator wasp larvae. Female trees have pistillate flowers that do produce seeds but are inhospitable to pollinator wasp larvae.

<i>Blastophaga</i> Genus of wasps

Blastophaga is a wasp genus in the family Agaonidae 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 specific to specific figs. The common fig Ficus carica is pollinated by Blastophaga psenes.

Schistonchus is a genus of plant-parasitic nematodes in the family Aphelenchoididae.

Schistonchus caprifici is a plant parasitic nematode in the genus Schistonchus parasitizing the caprifig. It is found in Spain and Italy.

<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.

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.

Ficus amplissima, also known as the Indian bat tree, Indian bat fig, Pimpri, Pipri (Piparee), Pipali or Bilibasari mara is a tree species of flowering plants that belongs to Moraceae, the fig or mulberry family. It is native to Central and southern Peninsular India, Sri Lanka and Maldives, having a significant distribution throughout Western Ghats of India. It is most commonly planted to provide shade in coffee plantations due to its dense and wide foliage. The ripened figs attract many birds, especially during the spring.

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.

References

  1. 1 2 3 4 5 Vovlas, Nicola; Alessandra Larizza (1996). "Relationships of Schistonchus caprifici (Aphelenchoididae) with fig inflorescences, the fig pollinator Blastophaga psenes, and its cleptoparasite Philotrypesis caricae". Fundamental and Applied Nematology. 19 (5): 443–448.
  2. ↑ "Blastophaga psenes (Linnaeus, 1758)". GBIF.org. Retrieved 26 May 2017.
  3. 1 2 "Blastophaga psenes Linnaeus". Figweb. Archived from the original on 27 April 2015. Retrieved 20 April 2015.
  4. 1 2 3 4 5 6 7 8 9 10 L. F. Kjellberg; P.-H. Gouyon; M. Ibrahim; M. Raymond; G. Valdeyron (July 1987). "The Stability of the Symbiosis between Dioecious Figs and Their Pollinators: A Study of Ficus carica L. and Blastophaga psenes". Evolution. 41 (4): 693–704. doi:10.2307/2408881. JSTOR   2408881. PMID   28564365.
  5. 1 2 3 4 Hossaert-Mckey, M.; M. Gibernau; J. E. Frey (1994). "Chemosensory Attraction of Fig Wasps to Substances Produced by Receptive Figs". Entomologia Experimentalis et Applicata. 70 (2). The Netherlands Entomological Society: 185–91. doi:10.1007/BF02380526 . Retrieved 11 October 2014.
  6. 1 2 Machado, C. A.; E. Jousselin; F. Kjellberg; S. G. Compton; E. A. Herre (2001). "Phylogenetic Relationships, Historical Biogeography and Character Evolution of Fig-pollinating Wasps". Proceedings of the Royal Society B: Biological Sciences. 268 (1468). The Royal Society: 685–94. doi:10.1098/rspb.2000.1418. PMC   1088657 . PMID   11321056.
  7. 1 2 3 Eisen, Gustav (1891). "The Introduction of Blastophaga Psenes into California". Zoe: A Biological Journal. 2: 114–15.
  8. ↑ Prat, Roger; Rubinstein, Jean-Pierre (13 Feb 2014). "Arbres et Arbustes". Fig (Ficus carica, Moraceae). Biologie et Multimédia.
  9. 1 2 3 4 Michailides, T. J.; D. P. Morgan (1994). "Dynamics of Blastophaga psenes Populations, Availability of Caprifigs, and Fig Endosepsis Cause by Fusarium moniliforme" (PDF). Phytopathology. 84 (11). The American Phytopathological Society: 1254–263. doi:10.1094/Phyto-84-1254 . Retrieved 11 October 2014.
  10. 1 2 Schatz, Bertrand; Martine Hossaert-Mckey (2010). "Ants Use Odour Cues to Exploit Fig–fig Wasp Interactions". Acta Oecologica. 36 (1): 107–13. Bibcode:2010AcO....36..107S. doi:10.1016/j.actao.2009.10.008.
  11. 1 2 Jiang, Z. -F.; Huang, D. -W.; Chen, L.-L.; Zhen, W. -Q.; Fu, Y. -G.; Peng, Z. -Q. (2006-07-01). "Rampant host switching and multiple female body colour transitions in Philotrypesis (Hymenoptera: Chalcidoidea: Agaonidae)". Journal of Evolutionary Biology. 19 (4): 1157–1166. doi:10.1111/j.1420-9101.2006.01087.x. ISSN   1010-061X.
  12. ↑ Wang, Rong; Matthews, Ashley; Ratcliffe, Joanne; Barwell, Louise J.; Peng, Yan‐Qiong; Chou, Lien‐Siang; Yu, Hui; Yang, Hui‐Wen; Compton, Stephen G. (2014-05-30). "First record of an apparently rare fig wasp feeding strategy: obligate seed predation". Ecological Entomology. 39 (4): 492–500. doi:10.1111/een.12122. ISSN   0307-6946.
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