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{{Distinguish|Homoplasmy}}Homoplasy, in biology and phylogenetics, is when a trait has been gained or lost independently in separate lineages over the course of evolution. This is different from homology, which is the similarity of traits can be parsimoniously explained by common ancestry.JOURNAL, Torres-Montúfar A, Borsch T, Ochoterena H, When Homoplasy Is Not Homoplasy: Dissecting Trait Evolution by Contrasting Composite and Reductive Coding, Systematic Biology, 67, 3, 543–551, May 2018, 28645204, 10.1093/sysbio/syx053, Homoplasy can arise from both similar selection pressures acting on adapting species, and the effects of genetic drift.BOOK, Evolution: an introduction, Stearns SC, Hoekstra RF, Oxford University Press, 2005, 9780199255634, 2nd, Oxford, JOURNAL, 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, (File:Homoplasy.jpg|thumb|Homoplasy is the similarity in trait that is not parsimoniously explained by descent from a common ancestor.)Most often, homoplasy is viewed as a similarity in morphological traits. However, homoplasy may also appear in other trait types, such as similarity in the genetic sequence,BOOK, Homoplasy: The Recurrence of Similarity in Evolution, Sanderson MJ, Hufford L, Academic Press, Inc., 1996, 0-12-618030-X, San Diego, CA, life cycle types JOURNAL, 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, or even behavioral traits.JOURNAL, 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,


The term homoplasy was first used by Ray Lankester in 1870.JOURNAL, 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, 34–43, 10.1080/00222937008696201, The corresponding adjective is either homoplasic or homoplastic.It is derived from the two ancient Greek words {{wikt-lang|grc|ὁμός}} ({{grc-transl|ὁμός}}), meaning "similar, alike, the same", and {{wikt-lang|grc|πλάσσω}} ({{grc-transl|πλάσσω}}), meaning "to shape, to mold".BOOK, Abrégé du dictionnaire grec français, Bailly, Anatole, vanc, 1981-01-01, Hachette, 2010035283, Paris, 461974285, WEB,weblink Greek-french dictionary online, Bailly, Anatole, vanc,, October 25, 2018, WEB,weblink Systematic Biology - Dictionary of Terms: Homoplasy, Holt JR, Judica CA, February 4, 2014, September 21, 2018, BOOK, Campbell Biology, Reece JB, Urry LA, Cain ML, Wasserman SA, Minorsky PV, Jackson RB, Pearson, 2011, 9780321739759, 9th,

Parallelism and convergence

Parallel and convergent evolution lead to homoplasy when different species independently evolve or gain a comparable trait, which diverges from the trait inferred to have been present in their common ancestor. When the similar traits are caused by an equivalent developmental mechanism, the process is referred to as parallel evolution.JOURNAL, Archie JW, 1989, Homoplasy excess ratios: new indices for measuring levels of homoplasy in phylogenetic systematics and a critique of the consistency index, Systematic Biology, 38, 253–269, 10.2307/2992286, JOURNAL, 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, The process is called convergent evolution when the similarity arises from different developmental mechanisms.JOURNAL, Hodin J, 2000, Plasticity and constraints in development and evolution, Journal of Experimental Zoology, 288, 1–20, 10.1002/(SICI)1097-010X(20000415)288:13.0.CO;2-7, 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 a cone-shaped heads and flat frontal claws) to live in a subterranean ecological niche.JOURNAL, Nevo E, 1979, Adaptive convergence and divergence of subterranean mammals, Annual Review of Ecology and Systematics, 10, 269–308, 10.1146/,


In contrast, reversal (a.k.a. vestigialization) leads to homoplasy through the disappearance of previously gained traits.JOURNAL, 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, This process may result from changes in the environment in which certain gained traits are no longer relevant, or have even become costly.JOURNAL, Fong DW, Kane TC, Culver DC, 1995, Vestigialization and loss of nonfunctional characters, Annual Review of Ecology and Systematics, 26, 249–68, 10.1146/, This can be observed in subterranean and cave-dwelling animals by their loss of sight,JOURNAL, Jones R, Culver DC, EEvidence 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, in cave-dwelling animals through their loss of pigmentation, and in both snakes and legless lizards through their loss of limbs.JOURNAL, 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, JOURNAL, 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,

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 created by means of parsimony analysis.BOOK, Phylogenetics: Theory and Practice of Phylogenetic Systematics, Wiley EO, Lieberman BS, John Wiley & Sons, Inc., 2011, 9780470905968, Hoboken, NJ, BOOK, Biological Systematics: Principles and Applications, Schuh RT, Brower AV, Cornell University Press, 2009, 9780801462436, Ithaca, NY, These analyses can be done with phenotypic, as well as genetic traits (DNA sequences).BOOK, Inferring phylogenies, Felsenstein J, Sinauer, 2004, 978-0878931774, Using parsimony analysis, the hypothesis of relationships that requires the fewest evolutionary changes is preferred over alternative hypotheses. Construction of these trees may become a challenge when clouded by the occurrence of homoplasy in the traits 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 measured.According to cladistic interpretation, homoplasy can be identified when the distribution of a trait cannot be explained by descent from a common ancestor 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:
  • Eusiroidea (Crustaceans and Amphipoda)JOURNAL, 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,
  • Urticaceae,JOURNAL, 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,
  • Asteraceae,JOURNAL, 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,
  • Polypodioideae (Selligueoid Ferns),JOURNAL, 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, 237–249, 10.12705/672.1,
  • Ants JOURNAL, 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,
  • Merluccius capensis (Cape Hakes) JOURNAL, 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,
  • Tarantula spiders of the genus Bonnetina: nearly all morphological traits within this genus are homoplastic. Only sexual features were observed to not be homoplastic, suggesting that sexual selection may have been a driving force in the divergence of tarantulas.JOURNAL, 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,
  • gharials, with homoplasy between Tomistoma and true crocodiles, and between thoracosaurs and Gavialis.JOURNAL, 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, 10.1098/rspb.2018.1071,
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.JOURNAL, Levin SR, Scott TW, Cooper HS, West SA, 2017, Darwin's aliens, International Journal of Astrobiology, 1–9, 10.1017/S1473550417000362,

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.BOOK, Wonderful Life: The Burgess Shale and the Nature of History, Gould SJ, Vintage Books, 2000, 9780099273455, London, 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 as well as homoplastic occurrence can be true at the same time.JOURNAL, Powell R, Mariscal C, Convergent evolution as natural experiment: the tape of life reconsidered, Interface Focus, 5, 6, 20150040, December 2015, 26640647, 10.1098/rsfs.2015.0040,

See also



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