Citrus Ryukyuensis
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''Citrus ryukyuensis'', also known as the tanibuta (), is a newly characterized wild
citrus ''Citrus'' is a genus of flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes. ''Citrus'' is nativ ...
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 Ryukyu Islands are divided into the Satsunan Islands ( Ōsumi, Tokara and Amami) and Okinawa Prefecture ( Daitō, Miyako, Y ...
and adjacent islands, most closely related to the mainland
mandarin orange A mandarin orange (''Citrus reticulata''), often simply called mandarin, is a small, rounded citrus tree fruit. Treated as a distinct species of orange, it is usually eaten plain or in fruit salads. The mandarin is small and oblate, unlike the ...
, ''C. reticulata''. As with other citrus, it is a member of the
Rutaceae The Rutaceae () is a family (biology), family, commonly known as the rueRUTACEAE
in Bo ...
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 named ''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
sea level rise The sea level has been rising from the end of the last ice age, which was around 20,000 years ago. Between 1901 and 2018, the average sea level rose by , with an increase of per year since the 1970s. This was faster than the sea level had e ...
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 development of seed or embryo without fertilization. However, other definitions include replacement of the seed by a plantlet or replacement of the flower by bulbils. Apomictically produced offspring are geneti ...
. 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 animal, 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 jour ...
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
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
from the Ryukyu mandarin but no introgressed
pomelo The pomelo ( ; or pummelo, ''Citrus maxima''), also known as a shaddock, is the largest citrus fruit. It is an ancestor of several cultivated citrus species, including the bitter orange and the grapefruit. It is a natural, non-hybrid citrus fr ...
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'' subsp. ''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 ...
cultivars.


Shiikuwasha

Separately from the hybridizations that produced the tachibana, a mainland clonal lineage that can still be found on
Okinawa most commonly refers to: * Okinawa Prefecture, Japan's southernmost prefecture * Okinawa Island, the largest island of Okinawa Prefecture * Okinawa Islands, an island group including Okinawa itself * Okinawa (city), the second largest city in th ...
, similar to the Chinese Sun Chu Sha mandarin 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 The bitter orange, sour orange, Seville orange, bigarade orange, or marmalade orange is the hybrid citrus tree species ''Citrus'' × ''aurantium'', and its fruit. It is native to Southeast Asia and has been spread by humans to many parts of th ...
/Ryukyu mandarin cross, and the ''deedee'', a ''shiikuwasha''/pomelo cross.


References

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