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Sympatric speciation

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In evolutionary biology and biogeography, sympatric and sympatry are terms referring to organisms whose ranges overlap or are even identical, so that they occur together at least in some places. If these organisms are closely related (e.g. sister species), such a distribution may be the result of sympatric speciation. Etymologically, sympatry is derived from the Greek roots συν (together, with) and πατρίς (homeland or fatherland).[1] The term was invented by Poulton in 1904, who explains the derivation.[2]

Sympatry is one of four theoretical models for the phenomenon of speciation. In contrast to allopatry, populations undergoing sympatric speciation are not geographically isolated by, for example, a mountain or a river.

In multicellular eukaryotic organisms, sympatric speciation is thought to be an uncommon but plausible process by which genetic divergence (through reproductive isolation) of various populations from a single parent species and inhabiting the same geographic region leads to the creation of new species. In bacteria, however, the analogous process (defined as "the origin of new bacterial species that occupy definable ecological niches" ) is more common and occurs through horizontal gene transfer.[3]

Evidence

Comparison of allopatric, peripatric, parapatric and sympatric speciation.

A number of models have been proposed to account for this mode of speciation. The most popular, which invokes the disruptive selection model, was first put forward by John Maynard Smith in 1966.[4] Maynard Smith suggested that homozygous individuals may, under particular environmental conditions, have a greater fitness than those with alleles heterozygous for a certain trait. Under the mechanism of natural selection, therefore, homozygosity would be favoured over heterozygosity, eventually leading to speciation. Sympatric divergence could also result from the sexual conflict[5].

Disruption may also occur in multiple-gene traits. The Medium Ground Finch (Geospiza fortis) is showing gene pool divergence in a population on Santa Cruz Island. Beak morphology conforms to two different size ideals, while intermediate individuals are selected against. Some characteristics (termed magic traits) such as beak morphology may drive speciation because they also affect mating signals. In this case, different beak phenotypes may result in different bird calls, providing a barrier to exchange between the gene pools.[6]

A well studied circumstance of sympatric speciation is when insects feed on more than one species of host plant. In this case insects become specialized as they struggle to overcome the various plants' defense mechanisms. (Drès and Mallet, 2002)[7]

Rhagoletis pomonella, the apple maggot, may be currently undergoing sympatric or, more precisely, heteropatric (see heteropatry) speciation. The apple feeding race of this species appears to have spontaneously emerged from the hawthorn feeding race in the 1800 - 1850 AD time frame, after apples were first introduced into North America. The apple feeding race does not now normally feed on hawthorns, and the hawthorn feeding race does not now normally feed on apples. This may be an early step towards the emergence of a new species. [8] [9] [10] Isolated and relatively homogeneous habitats such as crater lakes and islands are among the best geographical settings in which to demonstrate sympatric speciation. For example, Nicaragua crater lake cichlid fishes include at least one species that has evolved by sympatric speciation [11]

Allochrony offers some empirical evidence that sympatric speciation has taken place, as many examples exist of recently diverged (sister taxa) allochronic species.

Sympatric speciation events are vastly more common in plants, as they are prone to developing multiple homologous sets of chromosomes, resulting in a condition called polyploidy. The polyploidal offspring occupy the same environment as the parent plants (hence sympatry), but are reproductively isolated.

A rare example of sympatric speciation in animals is the divergence of "resident" and "transient" Orca forms in the northeast Pacific[12]. Resident and transient orcas inhabit the same waters, but avoid each other and do not interbreed. The two forms hunt different prey species and have different diets, vocal behaviour, and social structures. Some divergences between species could also result from contrasts in microhabitats.

The polecat Mustela putorius exhibited a rare dark phenotype similar to the European mink Mustela lutreola phenotype which is directely influenced by peculiarities of forest brooks[13].

Controversy

Debated almost since the beginning of popular evolutionary thought, sympatric speciation is still a highly contentious issue. By 1980 the theory was largely unfavourable given the void of empirical evidence available, and more critically the conditions scientists expect to be required. Ernst Mayr, one of the foremost thinkers on evolution, completely rejected sympatry outright, ushering in a climate of hostility towards the theory. While still debatable, well documented empirical evidence now exists, and the development of sophisticated theories incorporating multilocus genetics have followed.

See also

Look up sympatric in Wiktionary, the free dictionary.

References

  1. ^ http://www.greek-language.gr/greekLang/index.html
  2. ^ Poulton, E. B. 1904. What is a species? Proceedings of the Entomological Society of London 1903:lxxvii-cxvi.
  3. ^ King, Stansfield, Mulligan (2006). Dictionary of Genetics (7th ed.). Oxford University Press.{{cite book}}: CS1 maint: multiple names: authors list (link)
  4. ^ John Maynard Smith (1966). "Sympatric Speciation". American Naturalist. 100 (916): 637–650. [1]
  5. ^ Thierry Lodé "La guerre des sexes chez les animaux" Eds O Jacob, Paris, 2006
  6. ^ *Huber, SK; De León, LF; Hendry, AP; Bermingham, E; Podos, J (2007). "Reproductive isolation of sympatric morphs in a population of [[Darwin's finches]]". Proc. Biol. Sci. 274 (1619): 1709–14. doi:10.1098/rspb.2007.0224. PMC 2493575. PMID 17504742. {{cite journal}}: More than one of |given1= and |first1= specified (help); More than one of |given2= and |first2= specified (help); More than one of |given3= and |first3= specified (help); More than one of |given4= and |first4= specified (help); More than one of |given5= and |first5= specified (help); More than one of |surname1= and |last1= specified (help); More than one of |surname2= and |last2= specified (help); More than one of |surname3= and |last3= specified (help); More than one of |surname4= and |last4= specified (help); More than one of |surname5= and |last5= specified (help); URL–wikilink conflict (help)
  7. ^ Begon, Townsend, Harper: Ecology - From individuals to ecosystems, 4th ed., p.10
  8. ^ McPheron et al. 1988. Nature 336:64-66
  9. ^ Smith, D.C. 1988. Nature 336:66-67
  10. ^ Feder et al. 1988. Nature 336:61-64
  11. ^ Sympatric speciation in Nicaraguan crater lake cichlid fish. By: Barluenga, Marta; Stölting, Kai N.; Salzburger, Walter; Muschick, Moritz; Meyer, Axel. Nature, 2/9/2006, Vol. 439 Issue 7077, p719-723.
  12. ^ Hoetzel et al. 1998, Low genetic variation among killer whales (Orcinus orca) in the eastern North Pacific and genetic differentiation between foraging specialists, J Hered
  13. ^ Thierry Lodé "Genetic divergence without spatial isolation in polecat Mustela putorius populations". J Evol Biol 14:228-236, 2001
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