Editing Ficus (section)

==Ecology==
Figs are [[keystone species]] in many [[tropical forest]] [[ecosystem]]s. Their fruit are a key resource for [[frugivores]] including [[Megabat|fruit bats]], [[capuchin monkey]]s, [[Colobinae|langurs]], [[Nomascus|gibbons]], and [[mangabey]]s. They are even more important for [[bird]]s such as [[megalaimidae|Asian barbets]], [[Columbidae|pigeons]], [[hornbill]]s, [[Cyclopsittacini|fig-parrots]], and [[bulbul]]s, which may subsist almost entirely  on figs when these are plentiful. Many [[Lepidoptera]] [[caterpillar]]s feed on fig leaves, for example several ''[[Euploea]]'' species (crow butterflies), the [[Danaus chrysippus|plain tiger]] (''Danaus chrysippus''), the [[Papilio cresphontes|giant swallowtail]] (''Papilio cresphontes''), the [[Badamia exclamationis|brown awl]] (''Badamia exclamationis''), and ''[[Chrysodeixis eriosoma]]'', [[Choreutidae]] and [[Copromorphidae]] [[moth]]s. The larvae of the [[citrus long-horned beetle]] (''Anoplophora chinensis''), for example, feed on the [[wood]] of the fig tree; the species  can become a [[pest (organism)|pest]] in fig plantations. Similarly, the [[sweet potato whitefly]] (''Bemisia tabaci'') is frequently found as a pest on figs grown as [[houseplant|potted plant]]s and can be spread through the export of these plants to other localities. For a list of other diseases common to fig trees, see [[List of foliage plant diseases (Moraceae)]].

===Fig fruit and reproduction system===
{{anchor|Caprifig}}
{{See also|Fig}}
[[File:Fig.jpg|thumb|A [[common fig]] fruit]]
[[File:Fig (Ficus carica) fruit halved.jpg|right|thumb|Cut through of a ripe common fig]]

Many fig species are grown for their fruits, though only ''[[Common fig|Ficus carica]]'' is cultivated to any extent for this purpose.{{fact|date=July 2022}}{{disputed inline|date=July 2022}} A fig "fruit" is a type of [[multiple fruit]] known as a [[syconium]], derived from an arrangement of many small flowers on an inverted, nearly closed receptacle. The many small flowers are unseen unless the fig is cut open.{{cn|date=June 2023}}

The fruit typically has a bulbous shape with a small opening (the [[ostiole]]) at the outward end that allows access to [[pollinator]]s. The flowers are pollinated by [[Fig wasp|very small wasp]]s such as ''[[Pegoscapus]]'' that crawl through the opening in search of a suitable place to lay eggs. Without this pollinator service fig trees could not reproduce by seed. In turn, the flowers provide a safe haven and nourishment for the next generation of wasps. This accounts for the frequent presence of wasp larvae in the fruit, and has led to a [[coevolution]]ary relationship. Technically, a fig fruit proper would be only one of the many tiny matured, seed-bearing [[Gynoecium|gynoecia]] found inside one fig – if you cut open a fresh fig, individual fruit will appear as fleshy "threads", each bearing a single seed inside. The genus ''[[Dorstenia]]'', also in the fig family (Moraceae), exhibits similar tiny flowers arranged on a receptacle but in this case the receptacle is a more or less flat, open surface.{{cn|date=June 2023}}

Fig plants can be [[Monoecy|monoecious]] ([[Androdioecious|hermaphrodite]]){{clarification needed|reason=androdioecious is neither synonymous with Monoicy or hermaphroditism|date=September 2021}} or [[gynodioecious]] (hermaphrodite and female).<ref>{{cite web |url=http://waynesword.palomar.edu/dawkins.htm |last1=Armstrong |first1=Wayne P |first2=Steven |last2=Disparti |title=A Key to Subgroups of Dioecious* (Gynodioecious) Figs Based On Fig Wasp/Male Syconium Pollination Patterns |website=Wayne's Word |date=4 April 1998 |access-date=2012-01-05 |url-status=dead |archive-url=https://web.archive.org/web/20120202215506/http://waynesword.palomar.edu/dawkins.htm |archive-date=2012-02-02}}</ref> Nearly half of fig species are gynodioecious, and therefore have some plants with inflorescences (syconium) with long styled pistillate flowers, and other plants with staminate flowers mixed with short styled pistillate flowers.<ref>{{Cite book |title=Plant diversity and complexity patterns: local, regional, and global dimensions |url=https://books.google.com/books?id=1QwCQFbPsa4C&pg=RA1-PA427 |year=2005 |publisher=Kgl. Danske Videnskabernes Selskab |page=472 |isbn=978-87-7304-304-2 |last1=Friis |first1=Ib |last2=Balslev |first2=Henrik}}</ref> The long-styled flowers tend to prevent wasps from laying their eggs within the ovules, while the short-styled flowers are accessible for egg laying.<ref>{{cite journal |title=Sex Differences and Flowering Phenology in the Common Fig, Ficus carica L. |last1=Valdeyron |first1=Georges |last2=Lloyd |first2=David G. |jstor=2407790 |journal=Evolution |volume=33 |issue=2 |date=June 1979 |pages=673–685 |doi=10.2307/2407790 |pmid=28563939}}</ref>

All the native fig trees of the American continent are hermaphrodites, as well as species like [[Indian banyan]] (''F.&nbsp;benghalensis''), [[weeping fig]] (''F.&nbsp;benjamina''), [[Indian rubber plant]] (''F.&nbsp;elastica''), [[fiddle-leaved fig]] (''F.&nbsp;lyrata''), [[Moreton Bay fig]] (''F.&nbsp;macrophylla''), [[Chinese banyan]] (''F.&nbsp;microcarpa''), [[sacred fig]] (''F.&nbsp;religiosa'') and [[sycamore fig]] (''F.&nbsp;sycomorus'').{{sfnp|Berg|Corner|2005}} The common fig (''Ficus carica'') is a gynodioecious plant, as well as [[lofty fig]] or clown fig (''[[Ficus aspera|F.&nbsp;aspera]]''), [[Roxburgh fig]] (''[[Ficus auriculata|F.&nbsp;auriculata]]''), [[mistletoe fig]] (''[[Ficus deltoidea|F.&nbsp;deltoidea]]''), ''[[Ficus pseudopalma|F.&nbsp;pseudopalma]]'', [[creeping fig]] (''F.&nbsp;pumila'') and related species. The hermaphrodite common figs are called "inedible figs" or "caprifigs"; in traditional culture in the [[Mediterranean region]] they were considered food for [[goat]]s (''Capra aegagrus''). In the female fig trees, the male flower parts fail to develop; they produce the "'edible figs". [[Fig wasp]]s grow in common fig caprifigs but not in the female syconiums because the female flower is too long for the wasp to successfully lay her eggs in them. Nonetheless, the wasp pollinates the flower with pollen from the caprifig it grew up in. In many situations, the wasp pollinator is unable to escape and dies within the fruit. When the wasp dies, it is broken down by enzymes ([[Ficain]]) inside the fig. Fig wasps are not known to transmit any diseases harmful to humans.

When a caprifig ripens, another caprifig must be ready to be pollinated. In temperate climes, wasps hibernate in figs, and there are distinct crops. Caprifigs have three crops per year; common figs have two.<ref name="Sinha">{{cite book |doi=10.1016/B0-12-227055-X/00463-6 |chapter=FIGS |title=Encyclopedia of Food Sciences and Nutrition |year=2003 |last1=Sinha |first1=K.K. |pages=2394–2399 |isbn=978-0-12-227055-0}}</ref> The first crop ([[breba]]) is larger and juicier, and usually eaten fresh.<ref name=Sinha/>  In cold climates the breba crop is often destroyed by spring frosts.<ref>California Rare Fruit Growers, Inc. (1996): [http://www.crfg.org/pubs/ff/fig.html Fig] {{Webarchive |url=https://web.archive.org/web/20201031124540/http://www.crfg.org/pubs/ff/fig.html |date=2020-10-31 }}. Retrieved November 1, 2008.</ref> Some [[parthenocarpic]] [[cultivar]]s of common figs do not require pollination at all, and will produce a crop of figs (albeit [[sterility (physiology)|sterile]]) in the absence of caprifigs and fig wasps.

Depending on the species, each fruit can contain hundreds or even thousand of seeds.<ref>{{cite web |title=The Weird Sex Life of the Fig |url=http://figs4fun.com/Links/FigLink006a.pdf |website=Ray's Figs |access-date=2012-01-05}}</ref> Figs can be propagated by seeds, cuttings, air-layering or grafting. However, as with any plant, figs grown from seed are not necessarily genetically identical to the parent and are only propagated this way for breeding purposes.

===Mutualism with the pollinating fig wasps===
{{Further|Reproductive coevolution in Ficus}}
[[File:Ficus exasperata by kadavoor.jpg|thumb|upright|''[[Ficus exasperata]]'', fruits]]

The unique fig [[pollination]] system involves tiny, highly specific wasps, known as [[fig wasp]]s, that enter via [[ostiole]] these subclosed inflorescences to both pollinate and lay their own eggs.{{sfnp|Rønsted ''et al.''|2005}} Each species of fig is pollinated by one or a few specialised wasp species, and therefore plantings of fig species outside of their native range results in effectively sterile individuals. For example, in [[Hawaii]], some 60 species of figs have been introduced, but only four of the wasps that fertilize them, so only those species of figs produce viable seeds there and can become [[invasive species]]. This is an example of [[Mutualism (biology)|mutualism]], in which each organism (fig plant and [[fig wasp]]) benefit each other, in this case reproductively.{{cn|date=June 2023}}

The intimate association between fig species and their wasp pollinators, along with the high incidence of a one-to-one plant-pollinator ratio have long led scientists to believe that figs and wasps are a clear example of [[coevolution]]. Morphological and reproductive behavior evidence, such as the correspondence between fig and wasp larvae maturation rates, have been cited as support for this hypothesis for many years.<ref name="Machado Jousselin Kjellberg Compton pp. 685–694">{{cite journal |last1=Machado |first1=C. A. |last2=Jousselin |first2=E. |last3=Kjellberg |first3=F. |last4=Compton |first4=S. G. |last5=Herre |first5=E. A. |title=Phylogenetic relationships, historical biogeography and character evolution of fig-pollinating wasps |journal=Proceedings of the Royal Society B: Biological Sciences |volume=268 |issue=1468 |date=7 April 2001 |doi=10.1098/rspb.2000.1418 |pmid=11321056 |pmc=1088657 |pages=685–694}}</ref> Additionally, recent genetic and molecular dating analyses have shown a very close correspondence in the character evolution and [[speciation]] phylogenies of these two clades.{{sfnp|Rønsted ''et al.''|2005}}

According to meta-analysis of molecular data for 119 fig species 35% (41) have multiple pollinator wasp species. The real proportion is higher because not all wasp species were detected.<ref>{{cite journal |last1=Yang |first1=Li-Yuan |last2=Machado |first2=Carlos A. |last3=Dang |first3=Xiao-Dong |last4=Peng |first4=Yan-Qiong |last5=Yang |first5=Da-Rong |last6=Zhang |first6=Da-Yong |last7=Liao |first7=Wan-Jin |title=The incidence and pattern of copollinator diversification in dioecious and monoecious figs |journal=Evolution |date=February 2015 |volume=69 |issue=2 |pages=294–304 |doi=10.1111/evo.12584 |pmid=25495152 |pmc=4328460}}</ref> On the other hand, species of wasps pollinate multiple host fig species.<ref name="Machado Robbins Gilbert Herre pp. 6558–6565">{{cite journal |last1=Machado |first1=C. A. |last2=Robbins |first2=N. |last3=Gilbert |first3=M. T. P. |last4=Herre |first4=E. A. |title=Critical review of host specificity and its coevolutionary implications in the fig/fig-wasp mutualism |journal=Proceedings of the National Academy of Sciences |volume=102 |issue=Supplement 1 |date=3 May 2005 |doi=10.1073/pnas.0501840102 |pmid=15851680 |pmc=1131861 |pages=6558–6565 |bibcode=2005PNAS..102.6558M |doi-access=free}}</ref> Molecular techniques, like [[microsatellite (genetics)|microsatellite]] markers and mitochondrial sequence analysis, allowed a discovery of multiple genetically distinct, [[cryptic species complex|cryptic]] wasp species. Not all these cryptic species are sister taxa and thus must have experienced a host fig shift at some point.<ref name="Molbo et al. 2003">{{cite journal |last1=Molbo |first1=D. |last2=Machado |first2=C.A. |last3=Sevenster |first3=J.G. |last4=Keller |first4=L. |last5=Herre |first5=E.A. |title=Cryptic species of fig-pollinating wasps: Implications for the evolution of the fig-wasp mutualism, sex allocation, and precision of adaptation |journal=Proceedings of the National Academy of Sciences |date=24 April 2003 |volume=100 |issue=10 |pages=5867–5872 |doi=10.1073/pnas.0930903100 |pmid=12714682 |pmc=156293 |bibcode=2003PNAS..100.5867M |doi-access=free}}</ref> These cryptic species lacked evidence of genetic [[introgression]] or [[backcrossing|backcrosses]] indicating limited fitness for [[hybrid (biology)|hybrid]]s and effective [[reproductive isolation]] and [[speciation]].<ref name="Molbo et al. 2003"/>

The existence of cryptic species suggests that neither the number of symbionts nor [[Reproductive coevolution in ficus|their evolutionary relationships]] are necessarily fixed ecologically.<ref name="Molbo et al. 2003"/><!-- supports this sentence because: host shift --> While the morphological characteristics that facilitate the fig-wasp mutualisms are likely to be shared more fully in closer relatives, the absence of unique pairings would make it impossible to do a one-to-one tree comparison and difficult to determine cospeciation.{{cn|date=June 2023}}