Homoplasy Explained

Homoplasy, in biology and phylogenetics, is the term used to describe a feature that has been gained or lost independently in separate lineages over the course of evolution. This is different from homology, which is the term used to characterize the similarity of features that can be parsimoniously explained by common ancestry.[1] Homoplasy can arise from both similar selection pressures acting on adapting species, and the effects of genetic drift.[2] [3]

Most often, homoplasy is viewed as a similarity in morphological characters. However, homoplasy may also appear in other character types, such as similarity in the genetic sequence,[4] life cycle types[5] or even behavioral traits.[6]

Etymology

The term homoplasy was first used by Ray Lankester in 1870.[7] The corresponding adjective is either homoplasic or homoplastic.It is derived from the two Ancient Greek words, meaning "similar, alike, the same", and, meaning "to shape, to mold".[8] [9] [10] [11]

Parallelism and convergence

Parallel and convergent evolution lead to homoplasy when different species independently evolve or gain apparently identical features, which are different from the feature inferred to have been present in their common ancestor. When the similar features are caused by an equivalent developmental mechanism, the process is referred to as parallel evolution.[12] [13] The process is called convergent evolution when the similarity arises from different developmental mechanisms.[14] These types of homoplasy may occur when different lineages live in comparable ecological niches that require similar adaptations for an increase in fitness. An interesting example is that of the marsupial moles (Notoryctidae), golden moles (Chrysochloridae) and northern moles (Talpidae). These are mammals from different geographical regions and lineages, and have all independently evolved very similar burrowing characteristics (such as cone-shaped heads and flat frontal claws) to live in a subterranean ecological niche.[15]

Reversion

In contrast, reversal (a.k.a. vestigialization) leads to homoplasy through the disappearance of previously gained features.[16] This process may result from changes in the environment in which certain gained characteristics are no longer relevant, or have even become costly.[17] This can be observed in subterranean and cave-dwelling animals by their loss of sight,[18] in cave-dwelling animals through their loss of pigmentation, and in both snakes and legless lizards through their loss of limbs.[19] [20]

Distinguishing homology from homoplasy

Homoplasy, especially the type that occurs in more closely related phylogenetic groups, can make phylogenetic analysis more challenging. Phylogenetic trees are often selected by means of parsimony analysis.[21] [22] These analyses can be done with phenotypic characters, as well as DNA sequences.[23] Using parsimony analysis, the hypothesis of relationships that requires the fewest (or least costly) character state transformations is preferred over alternative hypotheses. Evaluation of these trees may become a challenge when clouded by the occurrence of homoplasy in the characters used for the analysis. The most important approach to overcoming these challenges is to increase the number of independent (non-pleiotropic, non-linked) characteristics used in the phylogenetic analysis. Along with parsimony analysis, one could perform a likelihood analysis, where the most likely tree, given a particular model of evolution, is selected, and branch lengths are inferred.

According to the cladistic interpretation, homoplasy is invoked when the distribution of a character state cannot be explained parsimoniously (without extra inferred character state transformations between the terminals and their ancestral node) on a preferred phylogenetic hypothesis - that is, the feature in question arises (or disappears) at more than one point on the tree.

In the case of DNA sequences, homoplasy is very common due to the redundancy of the genetic code. An observed homoplasy may simply be the result of random nucleotide substitutions accumulating over time, and thus may not need an adaptationist evolutionary explanation.

Examples and applications of homoplasy

There are numerous documented examples of homoplasy within the following taxa:

The occurrence of homoplasy can also be used to make predictions about evolution. Recent studies have used homoplasy to predict the possibility and the path of extraterrestrial evolution. For example, Levin et al. (2017) suggest that the development of eye-like structures is highly likely, due to its numerous, independently evolved incidences on earth.[32]

Homoplasy vs. evolutionary contingency

In his book Wonderful Life, Stephen Jay Gould claims that repeating the evolutionary process, from any point in time onward, would not produce the same results.[33] The occurrence of homoplasy is viewed by some biologists as an argument against Gould's theory of evolutionary contingency. Powell & Mariscal (2015) argue that this disagreement is caused by an equivocation and that both the theory of contingency and homoplastic occurrence can be true at the same time.[34]

See also

Notes and References

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  2. Book: Evolution: an introduction. Stearns SC, Hoekstra RF . Oxford University Press . 2005 . 9780199255634 . 2nd. Oxford.
  3. Hall AR, Colegrave N . Decay of unused characters by selection and drift . Journal of Evolutionary Biology . 21 . 2 . 610–7 . March 2008 . 18081745 . 10.1111/j.1420-9101.2007.01473.x . 11165522 . free .
  4. Book: Homoplasy: The Recurrence of Similarity in Evolution. Sanderson MJ, Hufford L . Academic Press, Inc.. 1996. 0-12-618030-X. San Diego, CA.
  5. Silberfeld T, Leigh JW, Verbruggen H, Cruaud C, de Reviers B, Rousseau F . A multi-locus time-calibrated phylogeny of the brown algae (Heterokonta, Ochrophyta, Phaeophyceae): Investigating the evolutionary nature of the "brown algal crown radiation" . Molecular Phylogenetics and Evolution . 56 . 2 . 659–74 . August 2010 . 20412862 . 10.1016/j.ympev.2010.04.020 .
  6. de Queiroz A, Wimberger PH . The usefulness of behavior for phylogeny estimation: levels of homoplasy in behavioral and morphological characters . Evolution; International Journal of Organic Evolution . 47 . 1 . 46–60 . February 1993 . 28568085 . 10.1111/j.1558-5646.1993.tb01198.x . 205778379 .
  7. Lankester ER . 1870. On the use of the term homology in modern zoology, and the distinction between homogenetic and homoplastic agreements.. Annals and Magazine of Natural History. 6. 31. 34–43. 10.1080/00222937008696201.
  8. Book: Bailly, Anatole . Abrégé du dictionnaire grec français . vanc . 1981-01-01 . Hachette . 2010035283 . Paris . 461974285 .
  9. Web site: Greek-french dictionary online . Bailly . Anatole . vanc. www.tabularium.be . October 25, 2018.
  10. Web site: Systematic Biology - Dictionary of Terms: Homoplasy. Holt JR, Judica CA. February 4, 2014. September 21, 2018. July 1, 2017. https://web.archive.org/web/20170701234612/https://comenius.susqu.edu/biol/202/dictionary%20of%20terms/h/homoplasy.htm. dead.
  11. Book: Campbell Biology. Reece JB, Urry LA, Cain ML, Wasserman SA, Minorsky PV, Jackson RB. Pearson. 2011. 9780321739759. 9th. registration.
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  13. Wake DB . September 1991 . Homoplasy: The Result of Natural Selection, or Evidence of Design Limitations? . The American Naturalist. en. 138. 3. 543–567. 10.1086/285234 . 85043463 .
  14. Hodin J . 2000. Plasticity and constraints in development and evolution. Journal of Experimental Zoology. 288. 1. 1–20. 10.1002/(SICI)1097-010X(20000415)288:1<1::AID-JEZ1>3.0.CO;2-7. 10750048.
  15. Nevo E . 1979. Adaptive convergence and divergence of subterranean mammals. Annual Review of Ecology and Systematics. 10. 269–308. 10.1146/annurev.es.10.110179.001413.
  16. Wake DB, Wake MH, Specht CD . Homoplasy: from detecting pattern to determining process and mechanism of evolution . Science . 331 . 6020 . 1032–5 . February 2011 . 21350170 . 10.1126/science.1188545 . 2011Sci...331.1032W . 26845473 .
  17. Fong DW, Kane TC, Culver DC . 1995. Vestigialization and loss of nonfunctional characters. Annual Review of Ecology and Systematics. 26. 249–68. 10.1146/annurev.es.26.110195.001341.
  18. Jones R, Culver DC . Evidence for selection on sensory structures in a cave population of Gammarus minus (Amphipoda) . Evolution; International Journal of Organic Evolution . 43 . 3 . 688–693 . May 1989 . 28568387 . 10.1111/j.1558-5646.1989.tb04267.x . 32245717 . free .
  19. Skinner A, Lee MS . 2009. Body-form evolution in the scincid lizard Lerista and the mode of macroevolutionary transitions. Evolutionary Biology. 36. 292–300. 10.1007/s11692-009-9064-9. 42060285.
  20. Skinner A, Lee MS, Hutchinson MN . Rapid and repeated limb loss in a clade of scincid lizards . BMC Evolutionary Biology . 8 . 310 . November 2008 . 19014443 . 10.1186/1471-2148-8-310 . 2596130 . free .
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  24. Verheye ML, Martin P, Backeljau T, D'Udekem D'Acoz C . 2015-12-22. DNA analyses reveal abundant homoplasy in taxonomically important morphological characters of Eusiroidea (Crustacea, Amphipoda) . Zoologica Scripta . 45. 3. 300–321. 10.1111/zsc.12153 . 86052388.
  25. Wu ZY, Milne RI, Chen CJ, Liu J, Wang H, Li DZ . Ancestral State Reconstruction Reveals Rampant Homoplasy of Diagnostic Morphological Characters in Urticaceae, Conflicting with Current Classification Schemes . PLOS ONE . 10 . 11 . e0141821 . 2015-11-03 . 26529598 . 4631448 . 10.1371/journal.pone.0141821 . 2015PLoSO..1041821W . free .
  26. Mejías JA, Chambouleyron M, Kim SH, Infante MD, Kim SC, Léger JF . 2018-07-19. Phylogenetic and morphological analysis of a new cliff-dwelling species reveals a remnant ancestral diversity and evolutionary parallelism in Sonchus (Asteraceae) . Plant Systematics and Evolution. en. 304. 8. 1023–1040. 10.1007/s00606-018-1523-2 . 49873212.
  27. He L, Schneider H, Hovenkamp P, Marquardt J, Wei R, Wei X, Zhang X, Xiang Q . 2018-05-09. A molecular phylogeny of selligueoid ferns (Polypodiaceae): Implications for a natural delimitation despite homoplasy and rapid radiation . Taxon. 67. 2. 237–249. 10.12705/672.1.
  28. Schär S, Talavera G, Espadaler X, Rana JD, Andersen Andersen A, Cover SP, Vila R . 2018-06-27. Do Holarctic ant species exist? Trans-Beringian dispersal and homoplasy in the Formicidae . Journal of Biogeography . 45. 8. 1917–1928 . 10.1111/jbi.13380 . 51832848 .
  29. Henriques R, von der Heyden S, Matthee CA . When homoplasy mimics hybridization: a case study of Cape hakes (Merluccius capensis and M. paradoxus) . PeerJ . 4 . e1827 . 2016-03-28 . 27069785 . 10.7717/peerj.1827 . 4824878 . free .
  30. Ortiz D, Francke OF, Bond JE . A tangle of forms and phylogeny: Extensive morphological homoplasy and molecular clock heterogeneity in Bonnetina and related tarantulas . Molecular Phylogenetics and Evolution . 127 . 55–73 . October 2018 . 29778724 . 10.1016/j.ympev.2018.05.013 . 29152043 .
  31. Lee MS, Yates AM . Tip-dating and homoplasy: reconciling the shallow molecular divergences of modern gharials with their long fossil record . Proceedings. Biological Sciences . 285 . 1881 . 20181071 . June 2018 . 30051855 . 6030529 . 10.1098/rspb.2018.1071 .
  32. Levin SR, Scott TW, Cooper HS, West SA . 2017. Darwin's aliens. International Journal of Astrobiology. 18. 1–9. 10.1017/S1473550417000362. free.
  33. Book: Wonderful Life: The Burgess Shale and the Nature of History. Gould SJ . Vintage Books. 2000. 9780099273455. London.
  34. Powell R, Mariscal C . Convergent evolution as natural experiment: the tape of life reconsidered . Interface Focus . 5 . 6 . 20150040 . December 2015 . 26640647 . 4633857 . 10.1098/rsfs.2015.0040 .

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