"Ring Species and Speciation"
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Ring Species and Advanced article Speciation Article Contents . What is a Ring Species? Heath Blackmon, The University of Texas at Arlington, Arlington, Texas, USA . Examples of Ring Species . Song Sparrow Jeffery P Demuth, The University of Texas at Arlington, Arlington, Texas, USA . African Acacia . Why Are Ring Species Rare? Based in part on the previous version of this eLS article ‘Ring Species and Speciation’ (2005) by Jeffery P Demuth and Loren H Rieseberg. Relevance to How Geographic Variation Promotes Speciation Online posting date: 15th October 2012 A ring species is a monophyletic group whose range has intergrading populations except in one area, where a sharp extended around a geographic barrier to produce a ring- morphological discontinuity is found. However, closer shaped distribution. The populations that make up the scrutiny almost inevitably leads to the discovery of gaps in ring should be contiguous and without barriers to gene the distribution of populations around the ring. Moreover, deoxyribonucleic acid (DNA) and protein analyses often flow anywhere in the ring except where the terminal detect cryptic barriers to gene flow, suggesting that speci- populations are sympatric but reproductively isolated ation is complete despite continuous morphological grad- from one another. Few, if any of the species that have been ation. Mayr (1963) recognised that many examples of ring described as ring species, actually meet all of these species did not fit his definition ‘in the most diagrammatic requirements. The best documented example of a species manner’. However, he maintained that it was the ability to that exhibits all of these traits is the greenish warbler follow the process of speciation step by step that made ring complex (Phylloscopus trochiloides). However, even species species such a convincing demonstration of geographic like the herring gull complex or Ensatina salamanders that speciation. See also: Species and Speciation: An Overview fail to exhibit some of the characteristics of a true ring The term ring species persists primarily because most species offer opportunities to study important evo- authors have adopted an implicit definition of ring species lutionary processes. These species are particularly helpful that describes taxa in which intergrading geographic var- ieties are distributed in a ring with overlapping ends. In the in understanding how microevolutionary changes can area of overlap, two sympatric forms maintain their dis- create two unique species, how speciation can occur in tinct phenotypes. This is similar to the definition of Mayr spite of gene flow, and how geographic speciation with or (1963) but allows for more than one species to be recog- without adaptive divergence can occur. nised in the ring, and for discontinuity in the distribution of populations. Ring species is often used synonymously with ‘circular overlap’ in the literature. The term ‘rassenkreis’ is What is a Ring Species? also sometimes used in the context of ring species (Stebbins, 1949); however, rassenkreis is more appropriately syn- The perfect demonstration of speciation is presented by the onymous with ‘polytypic species’, which does not imply situation in which a chain of intergrading subspecies forms anything pertaining to geographic distribution. Finally, it a loop or overlapping circle of which the terminal links deserves emphasis that the importance of studying ring have become sympatric without interbreeding, even species is not that they demonstrate a particular mode of though they are connected by a complete chain of inter- speciation such as allopatric or parapatric. Rather, they grading or interbreeding populations (i.e. ‘ring species’) provide natural evidence that speciation has occurred and (Mayr, 1963). offer a means for reconstructing the process through study Most organisms designated as ring species do not of extant members of the ring. See also: Speciation: adhere to the rigorous definition suggested by Mayr. If Introduction; Sympatric Speciation examined on a coarse geographical scale, they typically do have the requisite circular distribution of morphologically Examples of Ring Species eLS subject area: Evolution & Diversity of Life In the following section the authors describe some classic How to cite: examples of ring species and some more recent ones. The Blackmon, Heath; and Demuth, Jeffery P (October 2012) Ring Species cases reviewed include classic examples outlined by Mayr and Speciation. In: eLS. John Wiley & Sons, Ltd: Chichester. (1963) in Animal Species and Evolution, newer examples DOI: 10.1002/9780470015902.a0001751.pub3 reviewed by Irwin et al. (2001), and the first example of a eLS & 2012, John Wiley & Sons, Ltd. www.els.net 1 Ring Species and Speciation possible Coleopteran ring species. Of the original 22 pro- xanthoptica and (4) coastal and interior populations fol- posed ring species, 15 were birds. Mayr (1963) attributes lowed different patterns of divergence in response to this to the maturity of avian taxonomy. In recent years, the different ecological factors. bias has been towards mammals. This taxonomic skew is reflected in the examples discussed below. New evidence Ensatina salamanders Based on Stebbins’ observations, Dobzhansky (1958) concluded that virtually the entire process of speciation Early hypotheses was evident in Ensatina, but that completion of speciation The salamanders of the genus Ensatina are probably the was inhibited by continued gene flow through the con- best known and most thoroughly studied of all ring species. tinuous string of interbreeding subspecies. Morphological, Prior to 1949, Ensatina contained four species (Ensatina behavioural, biochemical and molecular measures from eschscholtzii, Ensatina sierrae, Ensatina croceater and extensively surveyed locations throughout the Ensatina Ensatina platensis) of fully terrestrial, lungless salamanders complex range have since shown that Stebbins’ (1949) (family Plethodontidae). Beginning with a comprehensive biogeographical hypothesis is too simple, and speciation is study of colouration and morphology by Stebbins (1949), no longer prevented by persistent gene flow (Brown, 1974; E. eschscholtzii has been subdivided into seven subspecies. Wake and Yanev, 1986; Moritz et al., 1992; Jackman and The seven subspecies encircle the Central Valley of Cali- Wake, 1994; Wake, 1997; Wake and Schneider, 1998; fornia. Three unblotched subspecies (oregonensis, xan- Pereira et al., 2011). However, Stebbins’ hypothesis has not thoptica and eschscholtzii) ring the coastal side of the valley, been falsified in a general sense. For example, allozyme three blotched subspecies (platensis, croceater and klau- variation is highest in the northern part of the range and beri) complete the ring along the interior Sierra Nevada genetic distances between pairs of populations from across Mountains and an intermediate ‘mottled’ subspecies the Central Valley generally increase from north to south (picta) is found north of the valley along the California– (Wake and Yanev, 1986), both of which are predicted by Oregon coast (Figure 1). In addition to the obvious mor- Stebbins’ hypothesis. Additionally, phylogenetic analysis phological gradation, extensive collecting (Stebbins, 1949) of cytochrome b (cyt b) haplotypes showed that croceater, demonstrated that subspecies freely interbreed in northern klauberi and southern populations of platensis form a California, but hybridise only occasionally where xan- monophyletic group (i.e. a group that includes all des- thoptica crosses the valley in central California (Figure 1). cendants of a common ancestor), as do xanthoptica and Where the two subspecies eschscholtzii and klauberi meet in eschscholtzii. Monophyly of these southern segments of the southern California they occur microsympatrically with- ring is consistent with independent southward dispersal out interbreeding. Based on this information, Stebbins along opposite sides of the valley (Moritz et al., 1992). (1949) hypothesised that (1) the species originated in forests See also: Biogeographical Regions; Speciation: Genetics near the California–Oregon border from a picta-like Although the predicted correlation of geographic dis- ancestor; (2) populations dispersed southwards along both tance with genetic and morphological divergence largely sides of the Central Valley during times of increased holds at a macrogeographic scale, exceptions are found at a humidity; (3) interior populations of xanthoptica were the finer scale (Jackman and Wake, 1994). Perhaps the most result of a Pleistocene transvalley invasion from coastal striking exception involves populations from the northern part of the platensis range. Specifically, in the area near Lassen Peak (Figure 1) there is a sharp transition in the genetic and morphological data between the unblotched and blotched forms. There is also a morphologically cryptic boundary to gene flow that separates northern platensis from southern platensis populations. Jackman and Wake (1994) interpret this genetic structure, which contradicts the classical model of steady north to south population expansion, as evidence that northern platensis is the prod- uct of introgression between southern platensis and ore- gonensis. In their view, the southern boundary is due to ancient vicariance; whereas, the presently distinct transi- tion between oregonensis and northern platensis is due to geologically recent volcanism and glaciations that caused repeated extinction and recolonisation. Historical biogeography also poses