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''Citrus ryukyuensis'' is a newly characterized wild
citrus ''Citrus'' is a genus of flowering plant, flowering trees and shrubs in the rue family, Rutaceae. Plants in the genus produce citrus fruits, including important crops such as Orange (fruit), oranges, Lemon, lemons, grapefruits, pomelos, and lim ...
species native to the
Ryukyus The , also known as the or the , are a chain of Japanese islands that stretch southwest from Kyushu to Taiwan: the Ōsumi, Tokara, Amami, Okinawa, and Sakishima Islands (further divided into the Miyako and Yaeyama Islands), with Yonaguni ...
and adjacent islands, most closely related to the mainland
mandarin orange The mandarin orange (''Citrus reticulata''), also known as the mandarin or mandarine, is a small citrus tree fruit. Treated as a distinct species of orange, it is usually eaten plain or in fruit salads. Tangerines are a group of orange-coloured ...
, ''C. reticulata''. As with other citrus, it is a member of the
Rutaceae The Rutaceae is a family, commonly known as the rueRUTACEAE
in BoDD – Botanical Derm ...
family. The Ryukyu and mainland species have diverged for more than 2 million years, and unlike the mainland mandarin, the Ryukyu mandarin reproduces sexually. Its hybridization with the mainland species has given rise to the unique mandarin hybrids of the islands.


Discovery and characterization

Previous genetic studies that characterized the mandarin oranges of the Asian mainland left unexplained the unusual genetic characteristics of those from Japan and nearby islands. A followup study designed at filling this gap in understanding produced an unexpected result, the identification of a pure wild species of mandarin, the hybridization of which with the mainland species has given rise to multiple distinct mandarin orange hybrid clusters. Because the identified specimens were native to the Ryukyu islands, the species was christened ''C. ryukyuensis''. Comparison with ''C. reticulata'' showed a calculated divergence time from mainland citrus between 2.2 and 2.8 million years ago, roughly corresponding to a
Pleistocene era The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed in ...
sea level rise Globally, sea levels are rising due to human-caused climate change. Between 1901 and 2018, the globally averaged sea level rose by , or 1–2 mm per year on average.IPCC, 2019Summary for Policymakers InIPCC Special Report on the Ocean and Cry ...
that would have cut off the islands, providing the isolation necessary for
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
. As a consequence of this isolation, the Ryukyu mandarin continued to reproduce sexually, while the mainland species experienced a genetic mutation that eventually spread throughout the mainland species and allowed them to reproduce asexually by
apomixis In botany, apomixis is asexual reproduction without fertilization. Its etymology is Greek for "away from" + "mixing". This definition notably does not mention meiosis. Thus "normal asexual reproduction" of plants, such as propagation from cuttin ...
. and Supplement


Hybrids

''C. ryukyuensis'' persists to the present, but is also of importance due to its contribution to Japanese and neighboring island mandarin hybrids.


Tachibana

Beginning between 40,000 and 200,000 years ago, either a subsequent sea level fall or
oceanic dispersal Oceanic dispersal is a type of biological dispersal that occurs when terrestrial organisms transfer from one land mass to another by way of a sea crossing. Island hopping is the crossing of an ocean by a series of shorter journeys between island ...
brought the Ryukyu species back into contact with mainland mandarins, and they interbred, producing different groupings of related hybrids. In one case, a mainland mandarin hybrid with similar contribution from the northern and southern mainland subspecies, and already containing some
introgressed Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Intr ...
DNA from the Ryukyu mandarin but no introgressed
pomelo The pomelo ( ), ''Citrus maxima'', is the largest citrus fruit from the family Rutaceae and the principal ancestor of the grapefruit. It is a natural, non-hybrid, citrus fruit, native to Southeast Asia. Similar in taste to a sweet grapefr ...
DNA like that found in most mainland mandarins, would form numerous crosses with the Ryukyu species. The mainland orange passed to its progeny the mutation allowing asexual reproduction, producing a cluster of similar clonal
Tachibana orange The tachibana orange (''Citrus tachibana,'' or ''Citrus reticulata tachibana'') is a variety of mandarin orange, a citrus fruit. They grow wild in the forests of Japan and are referred to in the poetry of the early Japanese and Ryukyu Islands ki ...
cultivars.


Shiikuwasha

Separately from the hybridizations that produced the tachibana, a mainland clonal lineage that can still be found on
Okinawa is a prefecture of Japan. Okinawa Prefecture is the southernmost and westernmost prefecture of Japan, has a population of 1,457,162 (as of 2 February 2020) and a geographic area of 2,281 km2 (880 sq mi). Naha is the capital and largest city ...
, similar to the Chinese
Sun Chu Sha mandarin The Sun is the star at the center of the Solar System. It is a nearly perfect ball of hot plasma, heated to incandescence by nuclear fusion reactions in its core. The Sun radiates this energy mainly as light, ultraviolet, and infrared radia ...
but with added pomelo introgression, interbred with the Ryukyu species to produce the shiikuwasha. It again is a cluster of similar hybrids and not a single uniform cultivar. Like the tachibana, these lines inherited the ability to propagate clonally from their mainland parent, but while the tachibana all have the mainland mandarin as seed parent, different shiikuwasha clones have each of the parent species as their seed parent. The diversity of both groups of clones, the fact that they are both found growing in the wild, and the mention of both in the early poetry of the Japanese and Ryukyu kingdoms suggests that these hybridization events occurred spontaneously in nature.


Other hybrids

The third important group of Ryukyu mandarin clones, referred to as ''yukunibu'', derive instead from a cross with a '' kunenbo''-type mandarin orange with higher levels of pomelo introgression, probably introduced agriculturally from South-East Asia or Indonesia in historical times. The Ryukyu mandarin has also contributed to more complex citrus hybrids, such as the ''rokugatsu'', a
bitter orange Bitter orange, Seville orange, bigarade orange, or marmalade orange is the citrus tree ''Citrus'' × ''aurantium'' and its fruit. It is native to Southeast Asia and has been spread by humans to many parts of the world. It is probably a cross bet ...
/Ryukyu mandarin cross, and the ''deedee'', a ''shiikuwasha''/pomelo cross.


References

{{Taxonbar, from=Q108055981 ryukyuensis Flora of the Ryukyu Islands