Ring species

From Wikipedia, the free encyclopedia
In a ring species, gene flow occurs between neighbouring populations of a species, but at the ends of the "ring", the populations don't interbreed.
The coloured bars show natural populations (colours), varying along a cline. Such variation may occur in a line (e.g. up a mountain slope) as in A, or may wrap around as in B.
Where the cline bends around, populations next to each other on the cline can interbreed, but at the point that the beginning meets the end again, as at C, the differences along the cline prevent interbreeding (gap between pink and green). The interbreeding populations are then called a ring species.

In biology, a ring species is a connected series of neighbouring populations, each of which interbreeds with closely sited related populations, but for which there exist at least two "end populations" in the series, which are too distantly related to interbreed, though there is a potential gene flow between each "linked" population and the next.[1] Such non-breeding, though genetically connected, "end populations" may co-exist in the same region (sympatry) thus closing a "ring". The German term Rassenkreis, meaning a circle of races, is also used.

Ring species represent speciation and have been cited as evidence of evolution. They illustrate what happens over time as populations genetically diverge, specifically because they represent, in living populations, what normally happens over time between long-deceased ancestor populations and living populations, in which the intermediates have become extinct. The evolutionary biologist Richard Dawkins remarks that ring species "are only showing us in the spatial dimension something that must always happen in the time dimension".[2]

Formally, the issue is that interfertility (ability to interbreed) is not a transitive relation; if A breeds with B, and B breeds with C, it does not mean that A breeds with C, and therefore does not define an equivalence relation. A ring species is a species with a counterexample to the transitivity of interbreeding.[3] However, it is unclear whether any of the examples of ring species cited by scientists actually permit gene flow from end to end, with many being debated and contested.[4]


The classic ring species is the Larus gull. In 1925 Jonathan Dwight found the genus to form a chain of varieties around the Arctic Circle. However, doubts have arisen as to whether this represents an actual ring species.[5] In 1938, Claud Buchanan Ticehurst argued that the greenish warbler had spread from Nepal around the Tibetan Plateau, while adapting to each new environment, meeting again in Siberia where the ends no longer interbreed.[6] These and other discoveries led Mayr to first formulate a theory on ring species in his 1942 study Systematics and the Origin of Species. Also in the 1940s, Robert C. Stebbins described the Ensatina salamanders around the Californian Central Valley as a ring species;[7][8] but again, some authors such as Jerry Coyne consider this classification incorrect.[4] Finally in 2012, the first example of a ring species in plants was found in a spurge, forming a ring around the Caribbean Sea.[9]


The biologist Ernst Mayr championed the concept of ring species, stating that it unequivocally demonstrated the process of speciation.[10] A ring species is an alternative model to allopatric speciation, "illustrating how new species can arise through 'circular overlap', without interruption of gene flow through intervening populations…"[11] However, Jerry Coyne and H. Allen Orr point out that rings species more closely model parapatric speciation.[4]

Ring species often attract the interests of evolutionary biologists, systematists, and researchers of speciation leading to both thought provoking ideas and confusion concerning their definition.[1] Contemporary scholars recognize that examples in nature have proved rare due to various factors such as limitations in taxonomic delineation[12] or, "taxonomic zeal"[10]—explained by the fact that taxonomists classify organisms into "species", while ring species often cannot fit this definition.[1] Other reasons such as gene flow interruption from "vicariate divergence" and fragmented populations due to climate instability have also been cited.[10]

Ring species also present an interesting case of the species problem for those seeking to divide the living world into discrete species. All that distinguishes a ring species from two separate species is the existence of the connecting populations; if enough of the connecting populations within the ring perish to sever the breeding connection then the ring species' distal populations will be recognized as two distinct species. The problem is whether to quantify the whole ring as a single species (despite the fact that not all individuals interbreed) or to classify each population as a distinct species (despite the fact that it interbreeds with its near neighbours). Ring species illustrate that species boundaries arise gradually and often exist on a continuum.[10]


Ensatina salamanders example of ring species
Speculated evolution and spread of the greenish warbler, Phylloscopus trochiloides.
  P. t. trochiloides
  P. t. obscuratus
  P. t. plumbeitarsus
  P. t. ludlowi
  P. t. viridanus
Note: The P. t. nitidus in Caucasus Mountains not shown

Many examples have been documented in nature. Debate exists concerning much of the research, with some authors citing evidence against their existence entirely.[4][13][self-published source?] The following examples provide evidence that—despite the limited number of concrete, idealized examples in nature—continuums of species do exist and can be found in biological systems.[10] This is often characterized by sub-species level classifications such as clines, ecotypes, complexes, and varieties. Many examples have been disputed by researchers, and equally "many of the [proposed] cases have received very little attention from researchers, making it difficult to assess whether they display the characteristics of ideal ring species."[1]

The following list gives examples of ring species found in nature. Some of the examples such as the Larus gull complex, the greenish warbler of Asia, and the Ensatina salamanders of America, have been disputed.[13][14][15][16]

See also[edit]


  1. ^ a b c d e f g h i j k l m n o p q r Irwin, Darren E.; Irwin, Jessica H.; Price, Trevor D. (2001). "Ring species as bridges between microevolution and speciation". Genetica. 112/113: 223–243. doi:10.1023/A:1013319217703. PMID 11838767. S2CID 7108000.
  2. ^ Dawkins, Richard (2004). The Ancestor's Tale. Houghton Mifflin. p. 303. ISBN 0-618-00583-8.
  3. ^ Brown, Rob. "'Same Species' vs. 'Interfertile: concise wording can avoid confusion when discussing evolution".
  4. ^ a b c d Coyne, Jerry A.; Orr, H. Allen (2004). Speciation. Sinauer Associates. pp. 102–105. ISBN 0-87893-091-4.
  5. ^ Liebers, D.; De Knijff, P.; Helbig, A. J. (2004). "The herring gull complex is not a ring species". Proceedings of the Royal Society B: Biological Sciences. 271 (1542): 893–901. doi:10.1098/rspb.2004.2679. PMC 1691675. PMID 15255043.
  6. ^ Irwin, Darren. "The greenish warbler ring species".
  7. ^ "A closer look at a classic ring species: The work of Tom Devitt". Understanding Evolution. 29 April 2021.
  8. ^ This species ring forms the subject of "The Salamander's tale" in Richard Dawkins' The Ancestor's Tale, 2004.
  9. ^ a b Cacho, N. Ivalú; Baum, David A. (7 September 2012). "The Caribbean slipper spurge Euphorbia tithymaloides: the first example of a ring species in plants". Proceedings of the Royal Society B: Biological Sciences. 279 (1742): 3377–3383. doi:10.1098/rspb.2012.0498. PMC 3396892. PMID 22696529.
  10. ^ a b c d e f Pereira, Ricardo J.; Wake, David B. (November 2015). "Ring species as demonstrations of the continuum of species formation". Molecular Ecology. 24 (21): 5312–5314. doi:10.1111/mec.13412. PMID 26509692. S2CID 206182763.
  11. ^ Helbig, A J (August 2005). "Evolutionary genetics: A ring of species". Heredity. 95 (2): 113–114. doi:10.1038/sj.hdy.6800679. PMID 15999143. S2CID 29782163.
  12. ^ Irwin, Darren E (December 2012). "A novel approach for finding ring species: look for barriers rather than rings". BMC Biology. 10 (1): 21. doi:10.1186/1741-7007-10-21. PMC 3299606. PMID 22410355.
  13. ^ a b Coyne, Jerry (16 July 2014). "There are no ring species". Why Evolution is True.
  14. ^ a b Alcaide, Miguel; Scordato, Elizabeth S. C.; Price, Trevor D.; Irwin, Darren E. (July 2014). "Genomic divergence in a ring species complex". Nature. 511 (7507): 83–85. Bibcode:2014Natur.511...83A. doi:10.1038/nature13285. hdl:10261/101651. PMID 24870239. S2CID 4458956.
  15. ^ Liebers, Dorit; Knijff, Peter de; Helbig, Andreas J. (2004). "The herring gull complex is not a ring species". Proc Biol Sci. 271 (1542): 893–901. doi:10.1098/rspb.2004.2679. PMC 1691675. PMID 15255043.
  16. ^ Highton, R. (1998). "Is Ensatina eschscholtzii a ring species?". Herpetologica. 54 (2): 254–278. JSTOR 3893431.
  17. ^ Simpson, K.; Day, N.; Trusler, P. (1999), Birds of Australia (6 ed.), Princeton University Press
  18. ^ Ward, David (December 2011). "Population differentiation in a purported ring species, Acacia karroo (Mimosoideae)". Biological Journal of the Linnean Society. 104 (4): 748–755. doi:10.1111/j.1095-8312.2011.01757.x.
  19. ^ Fuchs, Jérôme; Ericson, Per G.P.; Bonillo, Céline; Couloux, Arnaud; Pasquet, Eric (November 2015). "The complex phylogeography of the Indo-Malayan Alophoixus bulbuls with the description of a putative new ring species complex". Molecular Ecology. 24 (21): 5460–5474. doi:10.1111/mec.13337. PMID 26224534. S2CID 44672012.
  20. ^ Bowen, B. W.; Bass, A. L.; Rocha, L. A.; Grant, W. S.; Robertson, D. R. (2001). "Phytogeography of the Trumpetfishes (Aulostomus): Ring Species Complex on a Global Scale". Evolution. 55 (5): 1029–1039. doi:10.1554/0014-3820(2001)055[1029:pottar]2.0.co;2. PMID 11430639. S2CID 221735739.
  21. ^ Naidoo, Theshnie; Goodman, Steven M.; Schoeman, M. Corrie; Taylor, Peter J.; Lamb, Jennifer M. (1 January 2016). "Partial support for the classical ring species hypothesis in the Chaerephon pumilus species complex (Chiroptera: Molossidae) from southeastern Africa and western Indian Ocean islands". Mammalia. 80 (6). doi:10.1515/mammalia-2015-0062. S2CID 89517662.
  22. ^ Moritz, C.; Schneider, C. J.; Wake, D. B. (1 September 1992). "Evolutionary Relationships Within the Ensatina Eschscholtzii Complex Confirm the Ring Species Interpretation". Systematic Biology. 41 (3): 273–291. doi:10.1093/sysbio/41.3.273. S2CID 49267121.
  23. ^ Moritz, Craig; Schneider, Christopher J.; Wake, David B. (September 1992). "Evolutionary Relationships Within the Ensatina eschscholtzii Complex Confirm the Ring Species Interpretation". Systematic Biology. 41 (3): 273–291. doi:10.2307/2992567. JSTOR 2992567.
  24. ^ Päckert, Martin; Martens, Jochen; Eck, Siegfried; Nazarenko, Alexander A.; Valchuk, Olga P.; Petri, Bernd; Veith, Michael (27 September 2005). "The great tit (Parus major) - a misclassified ring species". Biological Journal of the Linnean Society. 86 (2): 153–174. doi:10.1111/j.1095-8312.2005.00529.x.
  25. ^ Kvist, Laura; Martens, Jochen; Higuchi, Hiroyoshi; Nazarenko, Alexander A.; Valchuk, Olga P.; Orell, Markku (22 July 2003). "Evolution and genetic structure of the great tit (Parus major) complex". Proceedings of the Royal Society of London. Series B: Biological Sciences. 270 (1523): 1447–1454. doi:10.1098/rspb.2002.2321. PMC 1691391. PMID 12965008.
  26. ^ Alström, Per (2006). "Species concepts and their application: insights from the genera Seicercus and Phylloscopus" (PDF). Acta Zoologica Sinica. 52 (Suppl): 429–434. S2CID 145032793. Archived from the original (PDF) on 21 July 2011.
  27. ^ Irwin, D. E.; Bensch, S; Irwin, JH; Price, TD (21 January 2005). "Speciation by Distance in a Ring Species". Science. 307 (5708): 414–416. Bibcode:2005Sci...307..414I. doi:10.1126/science.1105201. PMID 15662011. S2CID 18347146.
  28. ^ Liebers, Dorit; de Knijff, Peter; Helbig, Andreas J. (7 May 2004). "The herring gull complex is not a ring species". Proceedings of the Royal Society of London. Series B: Biological Sciences. 271 (1542): 893–901. doi:10.1098/rspb.2004.2679. PMC 1691675. PMID 15255043.
  29. ^ Caire, W.; Zimmerman, E. G. (1 March 1975). "Chromosomal and Morphological Variation and Circular Overlap in the Deer Mouse, Peromyscus Maniculatus, in Texas and Oklahoma". Systematic Biology. 24 (1): 89–95. doi:10.1093/sysbio/24.1.89.
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  34. ^ Bensch, Staffan; Grahn, Mats; Müller, Nils; Gay, Laurene; Åkesson, Susanne (July 2009). "Genetic, morphological, and feather isotope variation of migratory willow warblers show gradual divergence in a ring". Molecular Ecology. 18 (14): 3087–3096. doi:10.1111/j.1365-294X.2009.04210.x. PMID 19457197. S2CID 205361829.
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