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===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}}
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