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Hybrid speciation

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Hybrid speciation is a form of speciation wherein hybridization between two different species leads to a new species, reproductively isolated from the parent species. From the 1940s, reproductive isolation between hybrids and their parents was thought to be particularly difficult to achieve and thus hybrid species were thought to be extremely rare. With DNA analysis becoming more accessible in the 1990s, hybrid speciation has been shown to be a fairly common phenomenon, particularly in plants.[1][2]

In botanical nomenclature, a hybrid species is also called a nothospecies.[3] Hybrid species are by their nature polyphyletic.[4]

Hybrid speciation ecology

A hybrid may have a distinct trait (phenotype). This phenotype may in very rare cases be better fitted to the local environment than the parental lineage and as such natural selection may favor these individuals. If reproductive isolation subsequently is achieved, it will lead to a separate species. The reproductive isolation may be genetic, ecological, behavioural, or spatial, or a combination of these.

If reproductive isolation fails to establish, the hybrid population may breed back and finally merge with either or both parent species. This will lead to an influx of foreign genes in the parent population, a situation called an introgression. Introgression is a source of genetic variation, and can in itself facilitate speciation. There is evidence that introgression is a ubiquitous phenomenon in plants, animals,[5][6] and even humans,[7] where genetic material from Neanderthals and Denisovans is responsible for much of the immune genes in non-African populations.[8][9]

Ecological constraints to hybrid speciation

For a hybrid form to persist, it will generally have to be able to exploit the available resources better than either parent species, which, in most cases, it will have to compete with. While grizzly bears and polar bears may have offspring, a grizzly–polar bear hybrid will likely be less suited in either of the ecological roles than the parents themselves. Although the hybrid is fertile, this poor adaptation would prevent the establishment of a permanent population.[10]

Likewise, lions and tigers have historically overlapped in a portion of their range and can theoretically produce wild hybrids: ligers, which are a cross between a male lion and female tiger, and tigons, which are a cross between a male tiger and a female lion; however, tigers and lions have thus far only hybridized in captivity.[11] In both ligers and tigons, the females are fertile and the males are sterile.[11] One of these hybrids (the tigon) carries growth-inhibitor genes from both parents and thus is smaller than either parent species[11] and might in the wild come into competition with smaller carnivores, e.g. the leopard. The other hybrid, the liger, ends up larger than either of its parents: about a thousand pounds (450 kilograms) fully-grown.[11] No tiger-lion hybrids are known from the wild, particularly because each species is confined to geographically separated ranges (tigers aren't found in Africa, and the Asiatic lion is only found in the Gir Forest National Park, where tigers also are absent).[12]

Some situations may favour hybrid population. One example is rapid turnover of available environment types, like the historical fluctuation of water level in Lake Malawi, a situation that generally favors speciation.[13] A similar situation can be found where closely related species occupy a chain of islands. This will allow any present hybrid population to move into a new unoccupied habitats, avoiding direct competition with parent species and giving a hybrid population time and space to establish.[14] Genetics too can occasionally favour hybrids. In the Amboseli National Park in Kenya, yellow baboons and anubis baboons regularly interbreed. The hybrid males reach maturity earlier than their pure bred cousins, setting up a situation where the hybrid population may over time replace one or both of the parent species in the area.[15]

Genetics of hybridization

Genetics are more variable and malleable in plants than in animals, probably reflecting the higher activity level in animals. Hybrids genetics will necessarily be less stable than those of species evolving through isolation, which explains why hybrid species appear more common in plants than in animals. Many agricultural crops are hybrids with double or even triple chromosome sets. Having multiple sets of chromosomes is called polyploidy (or polyploidity). Polyploidy is usually fatal in animals where extra chromosome sets upset fetal development, but is often found in plants.[16] A form of hybrid speciation that is relatively common in plants, occurs when an infertile hybrid becomes fertile after doubling of the chromosome number.

Hybridization without change in chromosome number is called homoploid hybrid speciation.[1] This is the situation found in most animal hybrids. For a hybrid to be viable, the chromosomes of the two organisms will have to be very similar, i.e., the parent species must be closely related, or the difference in chromosome arrangement will make mitosis problematic. With polyploid hybridization, this constraint is less acute.

Super-numerary chromosome numbers can be unstable, which can lead to instability in the genetics of the hybrid. The European edible frog appears to be a species, but is actually triploid semi-permanent hybrids between pool frogs and marsh frogs.[17] In most populations, the edible frog population is dependent on the presence of at least one of the parents species to be maintained as each individual need two gene sets from one parent species and one from the other. Also, the male sex determination gene in the hybrids is only found in the genome of the pool frog, further undermining stability.[18] Such instability can also lead to rapid reduction of chromosome numbers, creating reproductive barriers and thus allowing speciation.

Known cases of hybrid speciation

Closely related Heliconius species
Saxifraga osloensis a natural tetraploid hybrid species

Animals

Hybrid speciation in animals is primarily homoploid. While not very common, a few animal species have been recognized as being the result of hybridization, mostly insects and fish.[14] The Lonicera fly is an example of a novel animal species that resulted from natural hybridization. The great skua has a surprising genetic similarity to the physically dissimilar pomarine skua, and most ornithologists now assume the great skua is a hybrid species between the pomarine skua and one of the northern skua species.[19]

Rapidly diverging species can sometimes form multiple hybrid species, giving rise to a species complex, like several physically divergent but closely related genera of cichlid fishes in Lake Malawi.[13] The duck genus Anas (mallards and teals) has a very recent divergence history, many of the species are inter-fertile and quite a few of them are thought to be hybrids.[20] While hybrid species generally appear rare in mammals,[14] the American red wolf too appear to be a hybrid species of the Canis species complex, between gray wolf and coyote.[21] This is also hypothesized to have led to the species rich Heliconius butterflies,[22] though the conclusion has been criticized.[23]

The first known instance of hybrid speciation in marine mammals was discovered by scientists in 2014. The clymene dolphin (Stenella clymene), also known in older texts as the short-snouted spinner dolphin, is a hybrid of the spinner dolphin and striped dolphin. All three species live in the Atlantic Ocean and often interact peacefully.[24]

Plants

With plants being more tolerant of polyploidity, hybrid species are more common in plants than in animals. Estimates indicate as much as 2–4% of all flowering plants and 7% of all fern species are the results of polyploid hybridization.[25] Many of the crop species are hybrids,[25] and hybridization is an important factor in speciation in some plant groups.[26] Hybrids of the flower genus Saxifraga is commonly used in gardening, and a tetraploid natural hybrid, Saxifraga osloenis is estimated to have formed at the end of the last ice age.[27][28]

Homoploid speciation has given rise to several species of sunflower.[29][30]

See also

References

  1. ^ a b Arnold, M.L. (1996). Natural Hybridization and Evolution. New York: Oxford University Press. p. 232. ISBN 978-0-19-509975-1.
  2. ^ Wendel, J F. & Doyle, J.J. (1998): DNA Sequencing. In Molecular Systematics of Plants II. Editors: D.E. Soltis, P.S. Soltis, J.J. Doyle. Kluwer, Boston, pp. 265–296.
  3. ^ McNeill, J.; Barrie, F.R.; Buck, W.R.; Demoulin, V.; Greuter, W.; Hawksworth, D.L.; Herendeen, P.S.; Knapp, S.; Marhold, K.; Prado, J.; Prud'homme Van Reine, W.F.; Smith, G.F.; Wiersema, J.H.; Turland, N.J. (2012). International Code of Nomenclature for algae, fungi, and plants (Melbourne Code) adopted by the Eighteenth International Botanical Congress Melbourne, Australia, July 2011. Vol. Regnum Vegetabile 154. A.R.G. Gantner Verlag KG. ISBN 978-3-87429-425-6.{{cite book}}: CS1 maint: multiple names: authors list (link) Article H.1
  4. ^ Hörandl, E. & Stuessy, T.F. (2010). "Paraphyletic groups as natural units of biological classification". Taxon. 59 (6): 1641–1653.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  5. ^ Dowling T. E., Secor C. L. (1997). "The role of hybridization and introgression in the diversification of animals". Annual Review of Ecology and Systematics. 28: 593–619. doi:10.1146/annurev.ecolsys.28.1.593.
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  7. ^ Holliday T. W. (2003). "Species concepts, reticulations, and human evolution". Current Anthropology. 44 (5): 653–673. doi:10.1086/377663.
  8. ^ Mendez, F. L.; Watkins, J. C.; Hammer, M. F. (12 January 2013). "Neandertal Origin of Genetic Variation at the Cluster of OAS Immunity Genes". Molecular Biology and Evolution. 30 (4): 798–801. doi:10.1093/molbev/mst004.
  9. ^ Mendez, F.L. (2012). Archaic introgression and natural selection in the evolution of modern humans: A study of genetic variation at the loci containing the immune genes OAS1 and STAT2 (Phd thesis). University of Arizona. Retrieved 6 December 2013.
  10. ^ "Bear shot in N.W.T. was grizzly-polar hybrid". Cbc.ca. 2010-04-30. Archived from the original on July 5, 2010. Retrieved 2011-03-09. {{cite news}}: Unknown parameter |deadurl= ignored (|url-status= suggested) (help)
  11. ^ a b c d Mott, M. (2005, August 5). Retrieved February 13, 2013, from Liger Facts. Big Cat Rescue
  12. ^ "Frequently asked questions". University of Minnesota Lion Research Project. Retrieved 2011-06-28.
  13. ^ a b Genner, M.J.; Turner, G.F. (December 2011). "Ancient Hybridization and Phenotypic Novelty within Lake Malawi's Cichlid Fish Radiation". Molecular Biology and Evolution. 29 (Published online): 195–206. doi:10.1093/molbev/msr183. Retrieved 14 December 2011.
  14. ^ a b c Larsen, P.A.; Marchán-Rivadeneira, M.R.; Baker, R.J. (5 January 2010). "Natural hybridization generates mammalian lineage with species characteristics". Proceedings of the National Academy of Sciences of the United States of America. 107: 11447–11452. doi:10.1073/pnas.1000133107.
  15. ^ Charpentier & al. (2012). "Genetic structure in a dynamic baboon hybrid zone corroborates behavioural observations in a hybrid population". Molecular Ecology. 21 (3): 715–731. doi:10.1111/j.1365-294X.2011.05302.x. {{cite journal}}: |access-date= requires |url= (help)
  16. ^ von Wettstein, F. (1927). "Die Erscheinung der Heteroploidie, besonders im Pflanzenreich". Ergebnisse der Biologie. 2: 311–356. doi:10.1007/978-3-642-49712-4_5.
  17. ^ Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History. Number 297. New York. Issued March 15, 2006.
  18. ^ Guldager Christiansen, D. (2010): Genetic Structure and Dynamics of All-hybrid Edible Frog Populations. Doctoral dissertation for the University of Zurich. 140 pages
  19. ^ Furness, R.W.; Hamer, K. (2003). Christopher Perrins (ed.). Firefly Encyclopedia of Birds. Firefly Books. pp. 270–273. ISBN 1-55297-777-3.
  20. ^ A mid-sized species: Bernor, R.L.; Kordos, L. & Rook, L. (eds): Recent Advances on Multidisciplinary Research at Rudabánya, Late Miocene (MN9), Hungary: A compendium. Paleontographica Italiana 89: 3–36.
  21. ^ Esch, Mary (31 May 2011). "Study: Eastern wolves are hybrids with coyotes". The Huffington Post. Retrieved 1 June 201`. {{cite news}}: Check date values in: |accessdate= (help)
  22. ^ Mallet, J.; Beltrán, M.; Neukirchen, W.; Linares, M. (2007). "Natural hybridization in heliconiine butterflies: The species boundary as a continuum". BMC Evolutionary Biology. 7: 28–28. doi:10.1186/1471-2148-7-28. PMC 1821009. PMID 17319954.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  23. ^ Brower AVZ (2011). "Hybrid speciation in Heliconius butterflies? A review and critique of the evidence". Genetica. 139 (2): 589–609. doi:10.1007/s10709-010-9530-4.
  24. ^ Bhanoo, Sindya. "Scientists Find Rare Hybrid of Two Other Dolphin Species". The New York Times. Retrieved 20 January 2014.
  25. ^ a b Otto, S.; Witton, P.J. (2000). "Polyploid incidence and evolution" (PDF). Annual Review of Genetics. 34: 401–437. doi:10.1146/annurev.genet.34.1.401. PMID 11092833.
  26. ^ Linder, C.R.; Risenberg, L.H. (22 June 2004). "Reconstructing patterns of reticulate evolution in plants". American Journal of Botany. 91 (10): 1700–1708. doi:10.3732/ajb.91.10.1700. Retrieved 14 December 2011.
  27. ^ Knaben, G. (1934). "Saxifraga osloensis n. sp., a tetraploid species of the Tridactylites section". Nytt Magasin for Botanikk: 117–138.
  28. ^ Brochmann, C. (1998). "Molecular Evidence for Polyploid Origins in Saxifraga (Saxifragaceae): The Narrow Arctic Endemic S. svalbardensis and its Widespread Allies" (PDF). American Journal of Botany. 85 (1): 135–143. doi:10.2307/2446562. Retrieved 14 December 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  29. ^ Rieseberg, L.H.; Raymond, O.; Rosenthal, D.M.; Lai, Z.; Livingston, K.; Nakazato, T.; Durpy, J.L.; Schwarzbach, A.E.; Donovan, L.A.; Lexer, C. (2003). "Major Ecological Transitions in Wild Sunflowers Facilitated by Hybridization". Science. 301 (5637): 1211–1216. Bibcode:2003Sci...301.1211R. doi:10.1126/science.1086949. Retrieved 14 December 2011.
  30. ^ Welch, M.E.; Riesberg, L.H. (2002). "Habitat divergence between a homoploid hybrid sunflower species, Helianthus paradoxus (Asteraceae), and its progenitors". American Journal of Botany. 89 (3): 472–478. doi:10.3732/ajb.89.3.472.
  • Mavarez, J., Salazar, C.A., Bermingham, E., Salcedo, C., Jiggins, C.D., Linares, M. (2006) Speciation by hybridization in Heliconius butterflies. Nature.
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