mimicry

File:Wasp mimicry.jpg (C, D, E) and the wasp beetle (F) are Batesian mimics of stinging wasps (A, B), which are Müllerian mimics of each other.]]

In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simplest case, as in Batesian mimicry, a mimic resembles a model, so as to deceive a dupe, all three being of different species. A Batesian mimic, such as a hoverfly, is harmless, while its model, such as a wasp, is harmful, and is avoided by the dupe, such as an insect-eating bird. Birds hunt by sight, so the mimicry in that case is visual, but in other cases mimicry may make use of any of the senses. Most types of mimicry, including Batesian, are deceptive, as the mimics are not harmful, but Müllerian mimicry, where different harmful species resemble each other, is honest, as when species of wasps and of bees all have genuinely aposematic warning coloration. More complex types may be bipolar, involving only two species, such as when the model and the dupe are the same; this occurs for example in aggressive mimicry, where a predator in wolf-in-sheep's-clothing style resembles its prey, allowing it to hunt undetected. Mimicry is not limited to animals; in Pouyannian mimicry, an orchid flower is the mimic, resembling a female bee, its model; the dupe is the male bee of the same species, which tries to copulate with the flower, enabling it to transfer pollen, so the mimicry is again bipolar. In automimicry, another bipolar system, model and mimic are the same, as when blue lycaenid butterflies have 'tails' or eyespots on their wings that mimic their own heads, misdirecting predator dupes to strike harmlessly. Many other types of mimicry exist.

Etymology

Use of the word mimicry dates to 1637. It derives from the Greek term mimetikos, "imitative", in turn from mimetos, the verbal adjective of mimeisthai, "to imitate". "Mimicry" was first used in zoology by the English entomologists William Kirby and William Spence in 1823.{{cite book |last1=Kirby |first1=William |author1-link=William Kirby (entomologist) |last2=Spence |first2=William |author2-link=William Spence (entomologist) |year=1823 |title=An Introduction to Entomology |volume=2 |publisher=Longman, Hurst, Rees, Orme & Brown |edition=3rd |page=405 |url=https://books.google.com/books?id=ItZjam0xBBYC}} Originally used to describe people, "mimetic" was used in zoology from 1851.{{cite web |title=Online Etymology Dictionary |url=http://www.etymonline.com/index.php?search=mimicry&searchmode=none |last=Harper |first=Douglas |access-date=23 February 2022}}

History

= Ancient =

Aristotle wrote in his History of Animals that partridges use a deceptive distraction display to lure predators away from their flightless young:Aristotle, History of Animals, [https://archive.org/details/historiaanimaliu00aris_0/page/n417/mode/2up book 9, chapter 8.]

{{blockquote|When a man comes by chance upon a young brood [of partridges], and tries to catch them, the hen-bird rolls in front of the hunter, pretending to be lame: the man every moment thinks he is on the point of catching her, and so she draws him on and on, until every one of her brood has had time to escape; hereupon she returns to the nest and calls the young back.|Aristotle, translated by D'Arcy Wentworth Thompson}}

The behaviour is recognised as a form of mimicry by biologists.

= 19th century =

File:Fritz Müller 1891 (cropped).jpg created the first mathematical model of mimicry to explain why distasteful species should evolve similar appearances.]]

In 1823, Kirby and Spence, in their book An Introduction to Entomology, used the term "mimicry" informally to depict the way that the structure and coloration of some insects resembled objects in their environments:

{{blockquote|A jumping bug, very similar to the one figured by Schellenberg, also much resembles the lichens of the oak on which I took it. The spectre tribe (Phasma) go still further in this mimicry, representing a small branch with its spray.}}

The English naturalist Henry Walter Bates worked for several years on butterflies in the Amazon rainforest. Returning home, he described multiple forms of mimicry in an 1862 paper at the Linnean Society in London,{{cite journal |last=Bates |first=Henry W. |author-link=Henry Walter Bates |year=1862 |title=Contributions to an insect fauna of the Amazon valley. Lepidoptera: Heliconidae |journal=Transactions of the Linnean Society |volume=23 |issue=3 |pages=495–566 |doi=10.1111/j.1096-3642.1860.tb00146.x |url=https://zenodo.org/record/1447544}} and then in his 1863 book The Naturalist on the River Amazons.{{cite book |last=Bates |first=Henry W. |author-link=Henry Walter Bates |year=1863 |title=The naturalist on the river Amazons |title-link=The Naturalist on the River Amazons |publisher=Murray}} The term "Batesian mimicry" has since been used in his honour, its usage becoming restricted to the situation in which a harmless mimic gains protection from its predators by resembling a distasteful model. Among the observations in Bates's 1862 paper is the statement:

{{blockquote|I was never able to distinguish the Leptalides from the species they imitated, although they belong to a family totally different in structure and metamorphosis from the Heliconidae, without examining them closely after capture.}}

The German naturalist Fritz Müller also spent many years studying butterflies in the Amazon rainforest. He first published a journal article on mimicry in German in 1878,{{cite journal |last=Müller |first=Fritz |author-link=Fritz Müller |year=1878 |title=Ueber die Vortheile der Mimicry bei Schmetterlingen |language=de |trans-title=On the Advantages of Mimicry in Butterflies |journal=Zoologischer Anzeiger |volume=1 |pages=54–55}} followed in 1879 by a paper to the Entomological Society of London (translated and presented by Ralph Meldola).{{cite journal |last=Müller |first=Fritz |author-link=Fritz Müller |translator=R. Meldola |year=1879 |title=Ituna and Thyridia; a remarkable case of mimicry in butterflies |journal=Proclamations of the Entomological Society of London |volume=1879 |pages=20–29 |url=https://www.ucl.ac.uk/taxome/lit/muller_1879.pdf |archive-url=https://web.archive.org/web/20240302102124/https://www.ucl.ac.uk/taxome/lit/muller_1879.pdf |archive-date=2 March 2024 |url-status=live}} He described a situation where different species were each unpalatable to predators, and shared similar, genuine, warning signals. Bates found it hard to explain why this should be so, asking why they should need to mimic each other if both were harmful and could warn off predators on their own. Müller put forward the first mathematical model of mimicry for this phenomenon: if a common predator confuses the two species, individuals in both those species are more likely to survive, as fewer individuals of either species are killed by the predator. The term Müllerian mimicry, named in his honour, has since been used for this mutualistic form of mimicry.{{cite web |last=Mallet |first=James |author-link=James Mallet |title=Fritz Müller in 1891 |url=http://www.ucl.ac.uk/taxome/jim/Mim/Muller.html |access-date=18 November 2017 |archive-url=https://web.archive.org/web/20240630163011/http://www.ucl.ac.uk/taxome/jim/Mim/Muller.html |archive-date=30 June 2024 |url-status=live}}{{cite journal |last=Sherratt |first=Thomas |author-link=Thomas N. Sherratt |title=The Evolution of Müllerian Mimicry |journal=Die Naturwissenschaften |date=2008 |pages=681–695 |doi=10.1007/s00114-008-0403-y |pmid=18542902 |volume=95 |issue=8 |bibcode=2008NW.....95..681S |pmc=2443389 }}

Müller wrote that

{{blockquote|The resemblance of the genera named [Ituna and Thyridia] is the more worthy of notice since it occurs between insects both belonging to the group of butterflies which are protected by distastefulness. The explanation which applies in ordinary cases of [Batesian] mimicry—and no other has, so far as I know, been offered—cannot obtain for this imitation among protected species.}}

File:Batesplate ArM.jpg|Plate from Henry Walter Bates's 1862 paper illustrating Batesian mimicry between harmless Dismorphia species (top and third row) and distasteful Ithomiini (Nymphalidae, second and bottom row).

File:Müllerian mimicry.jpg|Mutual Müllerian mimicry in distasteful Heliconius butterflies. Plate LXII from Müller's collected writings, 1881

File:Darwinism 1889 page 259 image of mimicry.jpg|Page from Alfred Russel Wallace's 1889 book Darwinism, showing a beetle (below) mimicking a wasp

Overview

= Evolved resemblance =

File:Batesian vs Müllerian mimicry.svg, the latter honest.]]

Mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry functions to protect from predators.{{cite book |last1=King |first1=R. C. |last2=Stansfield |first2=W. D. |last3=Mulligan |first3=P. K. |year=2006 |title=A dictionary of genetics |url=https://archive.org/details/dictionarygeneti02king |url-access=limited |edition=7th |publisher=Oxford University Press |page=[https://archive.org/details/dictionarygeneti02king/page/n290 278] |isbn=978-0-19-530762-7}} Mimicry systems have three basic roles: a mimic, a model, and a dupe. When these correspond to three separate species, the system is called disjunct; when the roles are taken by just two species, the system is called bipolar.{{cite journal |last1=Kikuchi |first1=D. W. |last2=Pfennig |first2=D. W. |date=2013 |title=Imperfect Mimicry and the Limits of Natural Selection |journal=The Quarterly Review of Biology |volume=88 |issue=4 |pages=297–315 |doi=10.1086/673758|pmid=24552099 |s2cid=11436992 |url=https://cdr.lib.unc.edu/downloads/ms35tj93n }} Mimicry evolves if a dupe (such as a predator) perceives a mimic (such as a palatable prey) as a model (the organism it resembles), and is deceived to change its behaviour to the mimic's selective advantage.{{Cite journal |last1=Dalziell |first1=Anastasia H. |last2=Welbergen |first2=Justin A. |date=27 April 2016 |title=Mimicry for all modalities |journal=Ecology Letters |volume=19 |issue=6 |pages=609–619 |doi=10.1111/ele.12602 |pmid=27117779 |bibcode=2016EcolL..19..609D}} The resemblances can be via any sensory modality, including any combination of visual, acoustic, chemical, tactile, or electric.{{cite book |title=Mimicry in plants and animals |url=https://archive.org/details/mimicryinplantsa00wick |url-access=registration |last=Wickler |first=Wolfgang |author-link=Wolfgang Wickler |publisher=McGraw-Hill |year=1968}} Mimicry may be to the advantage of both organisms that share a resemblance, in which case it is mutualistic; or it can be to the detriment of one, making it parasitic or competitive. The evolutionary convergence between groups is driven by the selective action of a dupe.{{cite journal |last=Wickler |first=Wolfgang |author-link=Wolfgang Wickler |year=1965 |title=Mimicry and the Evolution of Animal Communication |journal=Nature|volume=208 |issue=5010 |pages=519–21 |doi=10.1038/208519a0 |bibcode=1965Natur.208..519W |s2cid=37649827}} Birds, for example, use sight to identify palatable insects,{{cite journal |last1=Radford |first1=Benjamin |author1-link=Benjamin Radford |last2=Frazier |first2=Kendrick |author2-link=Kendrick Frazier |title=Cheats and Deceits: How Animals and Plants Exploit and Mislead |journal=Skeptical Inquirer |date=January 2017 |volume=41 |issue=1 |page=60}} whilst avoiding noxious ones. Over time, palatable insects may evolve to resemble noxious ones, making them mimics and the noxious ones models. Models do not have to be more abundant than mimics. In the case of mutualism, each model is also a mimic; all such species can be called "co-mimics". Many harmless species such as hoverflies are Batesian mimics of strongly defended species such as wasps, while many such well-defended species form Müllerian mimicry rings of co-mimics. In the evolution of wasp-like appearance, it has been argued that insects evolve to masquerade wasps since predatory wasps do not attack each other, and that this mimetic resemblance has had the useful side-effect of deterring vertebrate predators.{{Cite journal |last1=Boppré |first1=Michael |last2=Vane-Wright |first2=Richard I. |last3=Wickler |first3=Wolfgang |author-link=Wolfgang Wickler |date=2017-01-01 |title=A hypothesis to explain accuracy of wasp resemblances |journal=Ecology and Evolution |volume=7 |issue=1 |pages=73–81 |doi=10.1002/ece3.2586 |pmc=5214283 |pmid=28070276 |bibcode=2017EcoEv...7...73B}}

Mimicry can result in an evolutionary arms race if mimicry negatively affects the model, in which case the model can evolve a different appearance from the mimic.^p161 Mimics may have different models for different life cycle stages, or they may be polymorphic, with different individuals imitating different models, as occurs in Heliconius butterflies. Models tend to be relatively closely related to their mimics, but mimicry can be of vastly different species, for example when spiders mimic ants. Most known mimics are insects, though many other examples including vertebrates, plants, and fungi exist.{{cite journal |last=Boyden |first=T. C. |year=1980 |title=Floral mimicry by Epidendrum ibaguense (Orchidaceae) in Panama |journal=Evolution |volume=34 |issue=1 |pages=135–136 |doi=10.2307/2408322 |pmid=28563205 |jstor=2408322}}{{cite journal |last1=Roy |first1=B. A. |year=1994 |title=The effects of pathogen-induced pseudoflowers and buttercups on each other's insect visitation |journal=Ecology |volume=75 |issue=2 |pages=352–358 |doi=10.2307/1939539 |jstor=1939539 |bibcode=1994Ecol...75..352R}}{{cite book |url=https://books.google.com/books?id=gztuDQAAQBAJ&q=floral+mimicry+2016&pg=PP1 |title=Floral Mimicry |last1=Johnson |first1=Steven D. |last2=Schiestl |first2=Florian P. |date=2016 |publisher=Oxford University Press |isbn=978-0-19-104723-7}}

= Evolutionary explanations =

It is widely accepted that mimicry evolves as a positive adaptation. The lepidopterist and novelist Vladimir Nabokov however argued that although natural selection might stabilize a "mimic" form, it would not be necessary to create it.{{cite journal |last1=Alexander |first1=Victoria N. |year=2002 |title=Nabokov, Teleology and Insect Mimicry |journal=Nabokov Studies |volume=7 |pages=177–213 |doi=10.1353/nab.2010.0004|s2cid=42675699}} The most widely accepted model used to explain the evolution of mimicry in butterflies is the two-step hypothesis. The first step involves mutation in modifier genes that regulate a complex cluster of linked genes that cause large changes in morphology. The second step consists of selections on genes with smaller phenotypic effects, creating an increasingly close resemblance. This model is supported by empirical evidence that suggests that a few single point mutations cause large phenotypic effects, while numerous others produce smaller effects. Some regulatory elements collaborate to form a supergene for the development of butterfly color patterns. The model is supported by computational simulations of population genetics.{{cite journal |last1=Holmgren |first1=N. M. A. |last2=Enquist |first2=M. |year=1999 |title=Dynamics of mimicry evolution |url=http://www.comp.leeds.ac.uk/biosystems/reading/paper/dynamics.pdf |journal=Biological Journal of the Linnean Society |volume=66 |issue=2 |pages=145–158 |doi=10.1111/j.1095-8312.1999.tb01880.x |doi-access=free}} The Batesian mimicry in Papilio polytes is controlled by the doublesex gene.{{cite journal |last1=Kunte |first1=K. |last2=Zhang |first2=W. |last3=Tenger-Trolander |first3=A. |last4=Palmer |first4=D. H. |last5=Martin |first5=A. |last6=Reed |first6=R. D. |last7=Mullen |first7=S. P. |last8=Kronforst |first8=M. R. |title=doublesex is a mimicry supergene |journal=Nature |volume=507 |issue=7491 |date=2014 |pages=229–232 |doi=10.1038/nature13112 |pmid=24598547 |bibcode=2014Natur.507..229K |s2cid=4448793}}

Some mimicry is imperfect. Natural selection drives mimicry only far enough to deceive predators. For example, when predators avoid a mimic that imperfectly resembles a coral snake, the mimic is sufficiently protected.{{cite journal |last1=Wilson |first1=J. |last2=Jahner |first2=J. |last3=Williams |first3=K. |last4=Forister |first4=M. |date=2013 |title=Ecological and Evolutionary Processes Drive the Origin and Maintenance of Imperfect Mimicry |journal=PLOS ONE |volume=8 |issue=4 |pages=e61610 |doi=10.1371/journal.pone.0061610 |pmid=23593490 |pmc=3625143 |bibcode=2013PLoSO...861610W |doi-access=free}}{{cite journal |last1=Kikuchi |first1=D. |last2=Pfenning |first2=D. |date=2010 |title=Predator Cognition Permits Imperfect Coral Snake Mimicry |journal=The American Naturalist |volume=176 |issue=6 |pages=830–834 |doi=10.1086/657041 |pmid=20950143|s2cid=35411437}}{{cite journal |last1=Howse |first1=P. E. |last2=Allen |first2=J. A. |date=1994 |title=Satyric Mimicry: The Evolution of Apparent Imperfection |journal=Proceedings of the Royal Society B |volume=257 |issue=1349 |pages=111–114 |doi=10.1098/rspb.1994.0102 |bibcode=1994RSPSB.257..111H |s2cid=84458742}}

Convergent evolution is an alternative explanation for why coral reef fish have come to resemble each other;{{cite journal |title=Who resembles whom? Mimetic and coincidental look-alikes among tropical reef fishes |last=Robertson |first=D. Ross |date=2013 |journal=PLOS ONE |doi=10.1371/journal.pone.0054939 |pages=e54939 |volume=8 |issue=1 |pmid=23372795 |pmc=3556028|bibcode=2013PLoSO...854939R |doi-access=free}}{{cite journal |title=Coincidental resemblances among coral reef fishes from different oceans |last=Robertson |first=D. Ross |journal=Coral Reefs |date=2015 |page=977 |volume=34 |issue=3 |doi=10.1007/s00338-015-1309-8|bibcode=2015CorRe..34..977R |doi-access=free}} the same applies to benthic marine invertebrates such as sponges and nudibranchs.{{cite book |title=Antipredatory defensive roles of natural products from marine invertebrates |last=Pawlik |first=J.R. |publisher=Springer |year=2012 |isbn=978-90-481-3833-3 |pages=677–710 |editor-first=E. |editor-last=Fattorusso |editor-last2=Gerwick |editor-first2=W.H. |editor-first3=O. |editor-last3=Taglialatela-Scafati |chapter=12}}

= Living and non-living models =

In its broadest definition, mimicry can include non-living models. The specific terms masquerade and mimesis are sometimes used when the models are inanimate, and the mimicry's purpose is crypsis.{{cite journal |title=The Evolution and Ecology of Masquerade |last1=Skelhorn |first1=John |last2=Rowland |first2=Hannah M. |last3=Ruxton |first3=Graeme D. |author3-link=Graeme D. Ruxton |journal=Biological Journal of the Linnean Society|year=2010 |volume=99 |pages=1–8 |doi=10.1111/j.1095-8312.2009.01347.x |doi-access=free}}{{cite journal |last=Pasteur |first=G. |date=1982 |title=A Classificatory Review of Mimicry Systems |journal=Annual Review of Ecology and Systematics |volume=13 |pages=169–199 |doi=10.1146/annurev.es.13.110182.001125 |jstor=2097066}} For example, animals such as flower mantises, planthoppers, comma and geometer moth caterpillars resemble twigs, bark, leaves, bird droppings or flowers.{{cite book |last1=Ruxton |first1=Graeme D. |author1-link=Graeme Ruxton |last2=Sherratt |first2=Thomas N. |author2-link=Thomas N. Sherratt |last3=Speed |first3=M. P. |date=2004 |title=Avoiding Attack: the Evolutionary Ecology of Crypsis, Warning Signals, and Mimicry |publisher=Oxford University Press}}{{Cite journal |last1=Wiklund |first1=Christer |last2=Tullberg |first2=Birgitta S. |date=September 2004 |title=Seasonal polyphenism and leaf mimicry in the comma butterfly |journal=Animal Behaviour |volume=68 |issue=3 |pages=621–627 |doi=10.1016/j.anbehav.2003.12.008 |s2cid=54270418}}{{cite journal |last1=Endler |first1=John A. |title=An Overview of the Relationships Between Mimicry and Crypsis |journal=Biological Journal of the Linnean Society |date=August 1981 |volume=16 |issue=1 |pages=25–31 |doi=10.1111/j.1095-8312.1981.tb01840.x}} In addition, predators may make use of resemblance to harmless objects in aggressive masquerade, to enable them to approach prey. This wolf in sheep's clothing strategy differs from the more specific resemblance to the prey in aggressive mimicry, where the prey is both model and dupe.{{cite journal |last=Pembury Smith |first=Matilda Q. R. |last2=Ruxton |first2=Graeme D. |title=Camouflage in predators |journal=Biological Reviews |volume=95 |issue=5 |date=2020 |doi=10.1111/brv.12612 |doi-access=free |pages=1325–1340|hdl=10023/19948 |hdl-access=free }}

Many animals bear eyespots, which are hypothesized to resemble the eyes of larger animals. They may not resemble any specific organism's eyes, and whether or not animals respond to them as eyes is also unclear.{{cite journal |last1=Stevens |first1=Martin |author1-link=Martin Stevens (biologist) |last2=Hopkins |first2=Elinor |last3=Hinde |first3=William |last4=Adcock |first4=Amabel |last5=Connolly |first5=Yvonne |last6=Troscianko |first6=Tom |last7=Cuthill |first7=Innes C. |author7-link=Innes Cuthill |title=Field Experiments on the effectiveness of 'eyespots' as predator deterrents |journal=Animal Behaviour |date=November 2007 |volume=74 |issue=5 |pages=1215–1227 |doi=10.1016/j.anbehav.2007.01.031 |s2cid=53186893}}{{cite journal |last1=Stevens |first1=Martin |author1-link=Martin Stevens (biologist) |title=Predator perception and the interrelation between different forms of protective coloration |journal=Proceedings of the Royal Society B: Biological Sciences |date=22 June 2007 |volume=274|issue=1617 |pages=1457–1464 |doi=10.1098/rspb.2007.0220 |pmid=17426012 |pmc=1950298}}{{cite journal |last1=Stevens |first1=Martin |author1-link=Martin Stevens (biologist) |last2=Stubbins |first2=Claire L. |last3=Hardman |first3=Chloe J. |title=The anti-predator function of 'eyespots' on camouflaged and conspicuous prey |journal=Behavioral Ecology and Sociobiology |date=30 May 2008 |volume=62 |issue=11 |pages=1787–1793 |doi=10.1007/s00265-008-0607-3|s2cid=28288920}}{{cite journal |last1=Hossie |first1=Thomas John |last2=Sherratt |first2=Thomas N. |author2-link=Thomas N. Sherratt |title=Defensive posture and eyespots deter avian predators from attacking caterpillar models |journal=Animal Behaviour |date=August 2013|volume=86 |issue=2 |pages=383–389 |doi=10.1016/j.anbehav.2013.05.029 |s2cid=53263767}} The model is usually another species, except in automimicry, where members of the species mimic other members, or other parts of their own bodies, and in inter-sexual mimicry, where members of one sex mimic members of the other.

= Types =

Many types of mimicry have been described. An overview of each follows, highlighting the similarities and differences between the various forms. Classification is often based on function with respect to the mimic (e.g., avoiding harm). Some cases may belong to more than one class, e.g., automimicry and aggressive mimicry are not mutually exclusive, as one describes the species relationship between model and mimic, while the other describes the function for the mimic (obtaining food). The terminology used has been debated, as classifications have differed or overlapped; attempts to clarify definitions have led to the partial replacement of old terms with new ones.{{cite journal |last=Endler |first=John A. |author-link=John Endler |title=An overview of the relationships between mimicry and crypsis |journal=Biological Journal of the Linnean Society |volume=16 |pages=25–31 |year=1981 |doi=10.1111/j.1095-8312.1981.tb01840.x}}{{cite journal |last1=Allen |first1=J. A. |last2=Cooper |first2=J. M. |title=Crypsis and masquerade |journal=Journal of Biological Education |volume=19 |issue=4 |page=268 |year=2010 |doi=10.1080/00219266.1985.9654747}}

Name	No. of spp.	Function	Dupe finds Model	Deception	Description (mimic, model, dupe)
class="wikitable sortable" style="margin: 1em auto;" \|+ Some kinds of mimicry classified by Pasteur 1982
Aristotelian	2	Protective	Agreeable	Deceptive	Brooding bird mimics itself with broken wing, luring predator away from nestPasteur cites Aristotle, History of Animals, book 9, chapter 9.
Automimicry	1 or 2	Protective	Agreeable	Deceptive	Multiple forms, e.g. one sex mimics the other, tail mimics head, etc.
Bakerian	2	Reproductive	Forbidding	Deceptive	Female flower resembles male flower, cheating pollinator
Batesian	3	Protective	Forbidding	Deceptive	Palatable mimic resembles distasteful model, deceives dupe
Browerian	2	Protective	Forbidding	Deceptive	Palatable butterfly resembles toxic member of same species
Emsleyan	3	Protective	Forbidding	Deceptive	Deadly snake resembles less deadly species, predators get chance to learn to avoid them
Gilbertian	2	Protective	Forbidding	Deceptive	Host/prey mimics and so repels parasite/predator
Kirbyan	2	Aggressive	Agreeable	Deceptive	Brood parasite adult or egg mimics host which raises the young as its own
Müllerian	3 or more	Protective	Forbidding	Honest	Distasteful co-mimics resemble each other, aposematically warning off predators
Pouyannian	2	Reproductive	Agreeable	Deceptive	Plant mimic resembles female bee, deceives male, gets itself pollinated
Vavilovian	3	Reproductive	Agreeable	Deceptive	Mimic resembles crop, deceives farmer
Wasmannian	2	Commensalist	Agreeable	Deceptive	Mimic resembles and deceives ant, lives in ant nest
Wicklerian	2	Aggressive	Agreeable	Deceptive	Predator or parasite resembles and attacks prey or host; parasite may get itself swallowed
Camouflage	2	Protective	Uninteresting	Deceptive	Mimic resembles background (plant parts, or inanimate)

Defensive

Mimicry is defensive or protective when organisms are able to avoid harmful encounters by deceiving enemies into treating them as something else.

= Batesian =

File:Macroxiphus sp cricket.jpg, a harmless bush cricket, mimics a well-defended ant. ]]

File:Bumblebee mimic.webm

In Batesian mimicry, the mimic resembles the model, but does not have the attribute that makes it unprofitable to predators (e.g., unpalatability, or the ability to sting). In other words, a Batesian mimic is a sheep in wolf's clothing. Mimics are less likely to be found out (for example by predators) when in low proportion to their model. Such negative frequency-dependent selection applies in most forms of mimicry. Specifically, Batesian mimicry can only be maintained if the harm caused to the predator by eating a model outweighs the benefit of eating a mimic. The nature of learning is weighted in favor of the mimics, for a predator that has a bad first experience with a model tends to avoid anything that looks like it for a long time, and does not re-sample soon to see whether the initial experience was a false negative. However, if mimics become more abundant than models, then the probability of a young predator having a first experience with a mimic increases. Batesian systems are therefore most likely to be stable where the model is more abundant than the mimic.{{cite book |last1=Stearns |first1=S. C. |last2=Hoekstra |first2=Rolf F. |title=Evolution: An Introduction |year=2000 |edition=5th |publisher=Oxford University Press |page=464 |isbn=978-0-19-854968-0}}

There are many Batesian mimics among butterflies and moths. Consul fabius and Eresia eunice imitate unpalatable Heliconius butterflies such as H. ismenius.{{cite journal |last1=Pinheiro |first1=Carlos E. G. |year=1996 |title=Palatability and escaping ability in Neotropical butterflies: tests with wild kingbirds (Tyrannus melancholicus, Tyrannidae) |journal=Biological Journal of the Linnean Society |volume=59 |issue=4 |pages=351–365 |doi=10.1111/j.1095-8312.1996.tb01471.x |doi-access=free}} Limenitis arthemis imitate the poisonous pipevine swallowtail (Battus philenor). Several palatable moths produce ultrasonic click calls to mimic unpalatable tiger moths.{{cite journal |last1=Barber |first1=J. R. |last2=Conner |first2=W. E. |year=2007 |title=Acoustic mimicry in a predator–prey interaction |journal=PNAS |volume=104 |issue=22 |pages=9331–9334 |doi=10.1073/pnas.0703627104 |pmid=17517637 |pmc=1890494 |bibcode=2007PNAS..104.9331B |doi-access=free}}{{Cite journal |last1=Barber |first1=Jesse R. |last2=Chadwell |first2=Brad A. |last3=Garrett |first3=Nick |last4=Schmidt-French |first4=Barbara |last5=Conner |first5=William E. |date=July 2009 |title=Naïve bats discriminate arctiid moth warning sounds but generalize their aposematic meaning |url=https://pubmed.ncbi.nlm.nih.gov/19561203/ |journal=The Journal of Experimental Biology |volume=212 |issue=Pt 14 |pages=2141–2148 |doi=10.1242/jeb.029991 |issn=0022-0949 |pmid=19561203 |s2cid=1303252}}{{Cite journal |last1=Barber |first1=Jesse R. |last2=Plotkin |first2=David |last3=Rubin |first3=Juliette J. |last4=Homziak |first4=Nicholas T. |last5=Leavell |first5=Brian C. |last6=Houlihan |first6=Peter R. |last7=Miner |first7=Krystie A. |last8=Breinholt |first8=Jesse W. |last9=Quirk-Royal |first9=Brandt |last10=Padrón |first10=Pablo Sebastián |last11=Nunez |first11=Matias |last12=Kawahara |first12=Akito Y. |display-authors=5 |date=2022-06-21 |title=Anti-bat ultrasound production in moths is globally and phylogenetically widespread |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=119 |issue=25 |pages=e2117485119 |doi=10.1073/pnas.2117485119 |doi-access=free |pmc=9231501 |pmid=35704762|bibcode=2022PNAS..11917485B}}{{Cite journal |last1=Kawahara |first1=Akito Y. |last2=Barber |first2=Jesse R. |date=2015-05-19 |title=Tempo and mode of antibat ultrasound production and sonar jamming in the diverse hawkmoth radiation |journal=Proceedings of the National Academy of Sciences |volume=112 |issue=20 |pages=6407–6412 |doi=10.1073/pnas.1416679112 |pmc=4443353 |pmid=25941377 |bibcode=2015PNAS..112.6407K |doi-access=free}} Octopuses of the genus Thaumoctopus (the mimic octopus) are able to intentionally alter their body shape and coloration to resemble dangerous sea snakes or lionfish.{{Cite web |url=http://marinebio.org/species.asp?id=260 |title=Mimic Octopus, Thaumoctopus mimicus at MarineBio.org |access-date=2007-06-09 |archive-url=https://web.archive.org/web/20170718195726/http://marinebio.org/species.asp?id=260 |archive-date=2017-07-18}} In the Amazon, the helmeted woodpecker (Dryocopus galeatus), a rare species which lives in the Atlantic Forest of Brazil, Paraguay, and Argentina, has a similar red crest, black back, and barred underside to two larger woodpeckers: Dryocopus lineatus and Campephilus robustus. This mimicry reduces attacks on D. galeatus.{{Cite web |title=Deceptive Woodpecker Uses Mimicry to Avoid Competition |url=http://www.amnh.org/explore/news-blogs/research-posts/deceptive-woodpecker-uses-mimicry-to-avoid-competition |website=AMNH |access-date=12 August 2015}} Batesian mimicry occurs in the plant kingdom, where the chameleon vine adapts its leaf shape and colour to match that of the plant it is climbing.{{cite journal |last=Gianoli |first=Ernesto |year=2014 |title=Leaf Mimicry in a Climbing Plant Protects against Herbivory |journal=Cell |volume=24 |issue=9 |pages=984–987 |doi=10.1016/j.cub.2014.03.010 |pmid=24768053 |doi-access=free}}

= Müllerian =

File:Batesian vs Müllerian Mimicry.svg, a wasp and a bee]]

In Müllerian mimicry, two or more species have similar warning or aposematic signals and both share genuine anti-predation attributes (e.g. being unpalatable), as first described in Heliconius butterflies.{{cite journal |last=Meyer |first=A. |year=2006 |title=Repeating Patterns of Mimicry |journal=PLOS Biology |volume=4 |issue=10|page=e341 |doi=10.1371/journal.pbio.0040341 |pmid=17048984 |pmc=1617347 |doi-access=free}} This type of mimicry is unique in several respects. Firstly, both the mimic and the model benefit from the interaction, which could thus be classified as mutualism. The signal receiver also benefits by this system, despite being deceived about species identity, as it is able to generalize the pattern to potentially harmful encounters. The distinction between mimic and model that is clear in Batesian mimicry is also blurred. Where one species is scarce and another abundant, the rare species can be said to be the mimic. When both are present in similar numbers, however, it makes more sense to speak of each as a co-mimic than of distinct 'mimic' and 'model' species, as their warning signals tend to converge.{{cite journal |last1=Rowland |first1=Hannah M. |last2=Ihalainen |first2=Eira |last3=Lindström |first3=Leena |last4=Mappes |first4=Johanna |last5=Speed |first5=Michael P. |title=Co-mimics have a mutualistic relationship despite unequal defences |journal=Nature |volume=448 |issue=7149 |date=2007 |issn=0028-0836 |doi=10.1038/nature05899 |pages=64–67}} Also, the mimetic species may exist on a continuum from harmless to highly noxious, so Batesian mimicry grades smoothly into Müllerian convergence.{{cite journal |last=Huheey |first=James E. |year=1976 |title=Studies in warning coloration and mimicry VII — Evolutionary consequences of a Batesian–Müllerian spectrum: A model for Müllerian mimicry |journal=Evolution |volume=30 |issue=1|pages=86–93 |doi=10.2307/2407675 |pmid=28565050 |jstor=2407675}}{{cite journal |last=Benson |first=W. W. |year=1977 |title=On the Supposed Spectrum Between Batesian and Mullerian Mimicry |journal=Evolution |volume=31 |issue=2 |pages=454–455 |doi=10.2307/2407770 |pmid=28563231 |jstor=2407770}}

= Emsleyan/Mertensian =

Emsleyan or Mertensian mimicry describes the unusual case where a deadly prey mimics a less dangerous species. It was first proposed by M. G. Emsley in 1966 as a possible explanation for how a predator can learn to avoid a very dangerous aposematic animal, such as a coral snake, when the predator is very likely to die, making learning unlikely.{{cite journal |last1=Emsley |first1=M. G. |year=1966 |title=The mimetic significance of Erythrolamprus aesculapii ocellatus Peters from Tobago |journal=Evolution |volume=20 |issue=4 |pages=663–64 |doi=10.2307/2406599 |jstor=2406599 |pmid=28562911}} The theory was developed by the German biologist Wolfgang Wickler who named it after the German herpetologist Robert Mertens.{{cite journal |last1=Mertens |first1=Robert |author-link=Robert Mertens |year=1956 |title=Das Problem der Mimikry bei Korallenschlangen |language=de |journal=Zool. Jahrb. Syst |volume=84 |pages=541–76}}{{cite journal |last1=Hecht |first1=M. K. |last2=Marien |first2=D. |year=1956 |title=The coral snake mimic problem: a reinterpretation |journal=Journal of Morphology |volume=98 |issue=2 |pages=335–365 |doi=10.1002/jmor.1050980207 |s2cid=83825414}}{{cite journal |last1=Sheppard |first1=P. M. |last2=Wickler |first2=Wolfgang |author2-link=Wolfgang Wickler |year=1969 |title=Review of Mimicry in plants and animals by Wolfgang Wickler |journal=Journal of Animal Ecology |volume=38 |issue=1 |page=243 |doi=10.2307/2762 |jstor=2762}} The scenario is unlike Müllerian mimicry, where the most harmful species is the model. But if a predator dies on its first encounter with a deadly snake, it has no occasion to learn to recognize the snake's warning signals. There would then be no advantage for an extremely deadly snake in being aposematic: any predator that attacked it would be killed before it could learn to avoid the deadly prey, so the snake would be better off being camouflaged to avoid attacks. But if the predator first learnt to avoid a less deadly warning-coloured snake, the deadly species could profit by mimicking the less dangerous snake. Some harmless milk snakes (Lampropeltis triangulum), the moderately toxic false coral snakes (Erythrolamprus aesculapii), and the deadly coral snakes (Micrurus) all have a red background color with black and white/yellow rings. In this system, both the milk snakes and the deadly coral snakes are mimics, while the false coral snakes are the model.

File:Micrurus tener.jpg|The deadly Texas coral snake,
Micrurus tener,
(the Emsleyan/Mertensian mimic)

File:Erythrolamprus aesculapii (cropped).jpg|The moderately toxic
Erythrolamprus aesculapii
(the model for both types of mimicry)

File:Lampropeltis triangulum annulata.jpg|The harmless Mexican milk snake,
Lampropeltis triangulum annulata
(the Batesian mimic)

= Wasmannian =

In Wasmannian mimicry, the mimic resembles a model that it lives along with in a nest or colony. Most of the models here are eusocial insects, principally ants.{{cite book |last=Wasmann |first=Erich |author-link=Erich Wasmann |year=1894 |title=Kritisches Verzeichniss der myrmecophilin und termitophilen Arthropoden |language=de |trans-title=Critical Inventory of Myrmecophile and Termitophile Arthropods |publisher=Felix Dames |location=Berlin}}{{cite book |last1=Hölldobler |first1=Bert |last2=Wilson |first2=Edward O. |author2-link=Edward O. Wilson |title=The Ants |url=https://books.google.com/books?id=ljxV4h61vhUC&pg=PA512 |year=1990 |publisher=Harvard University Press |isbn=978-0-674-04075-5 |pages=511–514}}

= Gilbertian =

File:Heliconiinae - Heliconius numata.JPG flower species use Gilbertian mimicry, defending against being eaten by larvae of Heliconius butterflies with leaf stipules (not shown) that resemble the butterfly's eggs.]]

Gilbertian mimicry is bipolar, involving only two species. The potential host (or prey) drives away its parasite (or predator) by mimicking it, the reverse of host-parasite aggressive mimicry. It was coined by Pasteur as a phrase for such rare mimicry systems, and is named after the American ecologist Lawrence E. Gilbert who described it in 1975.{{cite book |last=Gilbert |first=Lawrence E. |year=1975 |chapter=Ecological consequences of a coevolved mutualism between butterflies and plants |editor1=L. E. Gilbert |editor2=P. H. Raven |title=Coevolution of Animals and Plants |pages=210–240 |publisher=University of Texas Press |oclc=636384400}} The classical instance of Gilbertian mimicry is in the plant genus Passiflora, which is grazed by the micropredator larvae of some Heliconius butterflies. The host plants have evolved stipules that mimic mature Heliconius eggs near the point of hatching. The butterflies avoid laying eggs near existing ones, reducing intraspecific competition between caterpillars, which are also cannibalistic, so those that lay on vacant leaves provide their offspring with a greater chance of survival. The stipules thus appear to have evolved as Gilbertian mimics of butterfly eggs, under selection pressure from these caterpillars.{{cite book |last=Campbell |first=N. A. |year=1996 |title=Biology |edition=4th |at=Chapter 50 |publisher=Benjamin Cummings |isbn=0-8053-1957-3}}

= Browerian =

Browerian mimicry, named after Lincoln P. Brower and Jane Van Zandt Brower who first described it in 1967,{{cite book |last=Brower |first=Lincoln P. |author-link=Lincoln P. Brower |year=1970 |chapter=Plant poisons in a terrestrial food chain and implications for mimicry theory |editor-first=K. L. |editor-last=Chambers |title=Biochemical Coevolution |location=Corvallis, Oregon, USA |publisher=Oregon State Univ. |pages=69–82}}{{cite journal |last1=Brower |first1=Lincoln P. |author-link=Lincoln P. Brower |last2=Van Brower |first2=J. V. Z. |last3=Corvino |first3=J. M. |year=1967 |title=Plant poisons in a terrestrial food chain |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=57 |issue=4|pages=893–98 |doi=10.1073/pnas.57.4.893 |pmid=5231352 |pmc=224631 |bibcode=1967PNAS...57..893B|doi-access=free}} is a postulated form of automimicry; where the model belongs to the same species as the mimic. This is the analogue of Batesian mimicry within a single species, and occurs when there is a palatability spectrum within a population. Examples include the monarch and the queen from the subfamily Danainae, which feed on milkweed species of varying toxicity. These species store toxins from its host plant, which are maintained even in the adult. As levels of toxin vary depending on diet, some individuals are more toxic than the rest, which profit from the toxicity of those individuals, just as hoverflies benefit from mimicking well-defended wasps.

= Misdirection by automimicry =

File:Chaetodon capistratus2.jpg of foureye butterflyfish (Chaetodon capistratus) mimic its own eyes, deflecting attacks from the vulnerable head.]]

One form of automimicry is where one part of an organism's body resembles another part. For example, the tails of some snakes resemble their heads; they move backwards when threatened and present the predator with the tail, improving their chances of escape without fatal harm. Some fishes have eyespots near their tails, and when mildly alarmed swim slowly backwards, presenting the tail as a head. Some insects such as some lycaenid butterflies have tail patterns and appendages of various degrees of sophistication that promote attacks at the rear rather than at the head. Several species of pygmy owl bear "false eyes" on the back of the head, misleading predators into reacting as though they were the subject of an aggressive stare.{{cite web |title=Northern Pygmy Owl (Glaucidium californicum) |url=http://www.owlinstitute.org/northern-pygmy-owl.html |publisher=Owl Research Institute |access-date=23 August 2015 |archive-date=28 December 2015 |archive-url=https://web.archive.org/web/20151228222405/http://www.owlinstitute.org/northern-pygmy-owl.html |url-status=dead}} Many insects have filamentous "tails" at the ends of their wings and patterns of markings on the wings themselves. These combine to create a "false head". This misdirects predators such as birds and jumping spiders. Spectacular examples occur in the hairstreak butterflies; when perching on a twig or flower, they commonly do so upside down and shift their rear wings repeatedly, causing antenna-like movements of the "tails" on their wings. Studies of rear-wing damage support the hypothesis that this strategy is effective in deflecting attacks from the insect's head.Sourakov, Andrei (2013): Two heads are better than one: false head allows Calycopis cecrops (Lycaenidae) to escape predation by a Jumping Spider, Phidippus pulcherrimus (Salticidae), Journal of Natural History, 47:15-16, 1047–1054Robbins, Robert K. The "False Head" Hypothesis: Predation and Wing Pattern Variation of Lycaenid Butterflies. The American Naturalist Vol. 118, No. 5 (Nov., 1981), pp. 770–775

Aggressive

= Predators =

Aggressive mimicry is found in predators or parasites that share some of the characteristics of a harmless species, allowing them to avoid detection by their prey or host; the strategy resembles a wolf in sheep's clothing, though no conscious deceptive intent is involved. The mimic may resemble the prey or host itself, or another organism that does not threaten the prey or host. Begon, M.; Townsend, C.; Harper, J. (1996) Ecology: Individuals, populations and communities (third edition) Blackwell Science, London

Several spiders use aggressive mimicry to lure prey.{{cite journal |last1=Jackson |first1=R. R. |year=1995 |title=Eight-legged tricksters: Spiders that specialize at catching other spiders |journal=BioScience |volume=42 |issue=8 |pages=590–98 |jstor=1311924 |doi=10.2307/1311924}} Species such as the silver argiope (Argiope argentata) employ prominent patterns in the middle of their webs, such as zigzags. These may reflect ultraviolet light, and mimic the pattern seen in many flowers known as nectar guides. Spiders change their web day to day, which can be explained by the ability of bees to remember web patterns.{{cite journal |last1=Craig |first1=C. L. |year=1995 |title=Webs of Deceit |journal=Natural History |volume=104 |issue=3 |pages=32–35}}

Another case is where males are lured towards what seems to be a sexually receptive female. The model in this situation is the same species as the dupe. Female fireflies of the genus Photuris emit light signals that mimic the mating signals of females of the genus Photinus.Lloyd, J. E. (1965) Aggressive Mimicry in Photuris: Firefly Femmes Fatales Science 149:653–654. Male fireflies from several different genera are attracted to these "femmes fatales", and are captured and eaten. Each female has a repertoire of signals matching the delay and duration of the flashes of the female of the corresponding species.{{cite journal |last=Lloyd |first=J. E. |year=1975 |title=Aggressive Mimicry in Photuris Fireflies: Signal Repertoires by Femmes Fatales |journal=Science |volume=187 |issue=4175 |pages=452–453 |doi=10.1126/science.187.4175.452 |pmid=17835312 |bibcode=1975Sci...187..452L |s2cid=26761854}}

Some carnivorous plants may be able to increase their rate of capturing insect prey through mimicry.{{cite journal |last1=Moran |first1=Jonathan A. |year=1996 |title=Pitcher dimorphism, prey composition and the mechanisms of prey attraction in the pitcher plant Nepenthes rafflesiana in Borneo |journal=Journal of Ecology |volume=84 |issue=4 |pages=515–525 |doi=10.2307/2261474 |jstor=2261474 |bibcode=1996JEcol..84..515M}}

File:Epinephelus tukula is cleaned by two Labroides dimidiatus.jpg cleaning a potato grouper, Epinephelus tukula]]

A different aggressive strategy is to mimic a mutualistic symbiont of the prey. Cleaner fish eat parasites and dead skin from client fish. Some allow the cleaner to venture inside their body to hunt these parasites. However, the sabre-toothed blenny or false cleanerfish (Aspidontus taeniatus) mimics the bluestreak cleaner wrasse (Labroides dimidiatus), which is recognized by other fishes as a cleaner. The false cleanerfish resembles the cleaner, and mimics the cleaner's "dance". Once it is allowed close to the client, it attacks, biting off a piece of its fin before fleeing. Fish wounded in this fashion soon learn to distinguish mimic from model, but because the similarity is close they also become much more cautious of the model.{{cite journal |last=Wickler |first=W. |author-link=Wolfgang Wickler |year=1966 |title=Mimicry in Tropical Fishes |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |volume=251 |issue=772 |pages=473–474 |doi=10.1098/rstb.1966.0036 |bibcode=1966RSPTB.251..473W |s2cid=83609965}}

A mechanism that does not involve any luring is seen in the zone-tailed hawk, which resembles the turkey vulture. It flies amongst the vultures, effectively camouflaged as a vulture which poses no threat to the hawk's prey. It hunts by suddenly breaking from the formation and ambushing its prey.{{cite journal |last=Willis |first=E. O. |year=1963 |title=Is the Zone-Tailed Hawk a Mimic of the Turkey Vulture? |journal=The Condor |volume=65 |issue=4 |pages=313–317 |doi=10.2307/1365357 |jstor=1365357}}

= Parasites =

Parasites can be aggressive mimics, though the situation is somewhat different from those outlined previously. They can mimic their hosts' natural prey, allowing themselves to be eaten as a pathway into their host. Leucochloridium, a genus of flatworm, matures in the digestive system of songbirds, their eggs then passing out of the bird in the faeces. They are then taken up by Succinea, a terrestrial snail. The eggs develop in this intermediate host, and must then find a suitable bird to mature in. Since the host birds do not eat snails, the sporocyst has another strategy to reach its host's intestine. They are brightly coloured and move in a pulsating fashion. A sporocyst-sac pulsates in the snail's eye stalks,[http://www.semioticon.com/seo/M/images/mimicry_2.jpg See here for a photo.]{{cite book |last=Moore |first=J. |date=2002 |title=Parasites and the behavior of animals |publisher=Oxford University Press}} coming to resemble an irresistible meal for a songbird. In this way, it can bridge the gap between hosts, allowing it to complete its life cycle. A nematode (Myrmeconema neotropicum) changes the colour of the abdomen of workers of the canopy ant Cephalotes atratus to make it appear like the ripe fruits of Hyeronima alchorneoides. It also changes the behaviour of the ant so that the gaster (rear part) is held raised. This presumably increases the chances of the ant being eaten by birds.{{cite journal |pmid=18279076 |year=2008 |last1=Yanoviak |first1=S. P. |last2=Kaspari |first2=M. |last3=Dudley |first3=R. |last4=Poinar |first4=G. Jr |title=Parasite-induced fruit mimicry in a tropical canopy ant |volume=171 |issue=4 |pages=536–44 |doi=10.1086/528968 |journal=The American Naturalist |s2cid=23857167 |url=http://www.canopyants.com/2008_AmNat.pdf}}

Reproductive

Reproductive mimicry occurs when the actions of the dupe directly aid in the mimic's reproduction. This is common in plants with deceptive flowers that do not provide the reward they seem to offer and it may occur in Papua New Guinea fireflies, in which the signal of Pteroptyx effulgens is used by P. tarsalis to form aggregations to attract females.{{cite book |last1=Ohba |first1=N. |last2=Shimoyama |first2=Ayu|title=Bioluminescence in Focus - a collection of illuminating essays|year=2009|editor=Meyer-Rochow, V. B. |publisher=Research Signpost; Trivandrum, Kerala, India |pages=229–242}} Other forms of mimicry have a reproductive component, such as Vavilovian mimicry involving seeds, vocal mimicry in birds,{{Cite journal |last1=Dalziell |first1=Anastasia H. |last2=Welbergen |first2=Justin A. |last3=Igic |first3=Branislav |last4=Magrath |first4=Robert D. |date=2014-07-30 |title=Avian vocal mimicry: a unified conceptual framework |journal=Biological Reviews |volume=90 |issue=2 |pages=643–668 |doi=10.1111/brv.12129 |pmid=25079896 |s2cid=207101926}}{{cite journal |date=1 September 2008 |title=Vocal mimicry in songbirds |journal=Animal Behaviour |volume=76 |issue=3 |pages=521–528 |doi=10.1016/j.anbehav.2008.04.012 |last1=Kelley |first1=Laura A. |last2=Coe |first2=Rebecca L. |last3=Madden |first3=Joah R. |last4=Healy |first4=Susan D. |s2cid=53192695}}{{cite journal |last1=Goller |first1=Maria |last2=Shizuka |first2=Daizaburo |date=22 June 2018 |title=Evolutionary origins of vocal mimicry in songbirds |journal=Evolution Letters |volume=2 |issue=4 |pages=417–426 |doi=10.1002/evl3.62 |pmc=6121844 |pmid=30283692}} and aggressive and Batesian mimicry in brood parasite-host systems.{{cite book |url=https://www.bloomsbury.com/uk/cuckoo-9781408856567/ |title=Cuckoo: Cheating by Nature |last=Davies |first=Nick |date=2015 |publisher=Bloomsbury |isbn=978-1-4088-5656-7 |access-date=8 November 2018 |archive-date=28 February 2021 |archive-url=https://web.archive.org/web/20210228094002/https://www.bloomsbury.com/uk/cuckoo-9781408856567/ |url-status=dead }}

= Bakerian and Dodsonian =

Bakerian mimicry, named after Herbert G. Baker,Baker, Herbert G. 1976. "Mistake" pollination as a reproductive system, with special reference to the Caricaceae. Pp 161–169 in J. Burley and B. T. Styles, eds. {{Clarify|date=July 2013}} Variation, breeding, and conservation of tropical trees. Academic Press, London, U.K. is a form of automimicry where female flowers mimic male flowers of their own species, cheating pollinators out of a reward. This reproductive mimicry may not be readily apparent as members of the same species may still exhibit some degree of sexual dimorphism. It is common in many species of Caricaceae.{{cite journal |last1=Bawa |first1=K. S. |year=1980 |title=Mimicry of male by female flowers and intrasexual competition for pollinators in Jacaratia dolichaula (D. Smith) Woodson (Caricaceae) |journal=Evolution |volume=34 |issue=3|pages=467–74 |doi=10.2307/2408216 |jstor=2408216 |pmid=28568703}}

In Dodsonian mimicry, named after Calaway H. Dodson, the model belongs to a different species than the mimic.{{cite journal |last1=Dodson |first1=C. H. |last2=Frymire |first2=G. P. |year=1961 |title=Natural pollination of orchids |journal=Missouri Botanical Garden Bulletin |volume=49 |pages=133–39}} By resembling the model, a flower can lure its pollinators without offering nectar. The mechanism occurs in several orchids, including Epidendrum ibaguense which mimics flowers of Lantana camara and Asclepias curassavica, and is pollinated by monarch butterflies and perhaps hummingbirds.{{cite journal |last1=Boyden |first1=T. C. |year=1980 |title=Floral mimicry by Epidendrurn ibaguense (Orchidaceae) in Panama |journal=Evolution |volume=34 |issue=1|pages=135–36 |doi=10.2307/2408322 |jstor=2408322 |pmid=28563205}}

= Kirbyan mimicry, brood parasitism =

Brood parasitism or Kirbyan mimicry is a two species system where a brood parasite mimics its host. Cuckoos are a canonical example; the female cuckoo has its offspring raised by a bird of a different species, cutting down the biological mother's parental investment. The ability to lay eggs that mimic the host eggs is the key adaptation. The adaptation to different hosts is inherited through the female line in so-called gentes (gens, singular). Intraspecific brood parasitism, where a female lays in a conspecific's nest, as illustrated by the goldeneye duck (Bucephala clangula), do not involve mimicry{{cite journal |last1=Andersson |first1=M. |last2=Eriksson |first2=M. O. G. |year=1982 |title=Nest parasitism in Goldeneyes Bucephala clangula: some evolutionary aspects |journal=American Naturalist |volume=120 |pages=1–16 |doi=10.1086/283965 |s2cid=86699716}} The parasitic butterfly Phengaris rebeli parasitizes the ant species Myrmica schencki by releasing chemicals that fool the worker ants to believe that the caterpillar larvae are ant larvae. This enables the larvae to be brought directly into the ant's nest.{{cite journal |last=Barbero |first=Francesca |author2=Thomas, J.A. |author3=Bonelli, S. |author4=Balletto, E. |author5=Schonrogge, K. |title=Acoustical mimicry in a predatory social parasite of ants |journal=Journal of Experimental Biology |year=2009 |volume=212 |pages=4084–4090 |url=https://www.researchgate.net/publication/40039224 |access-date=28 September 2013 |doi=10.1242/jeb.032912 |pmid=19946088 |issue=Pt 24 |doi-access=free}}

File:European Cuckoo Mimics Sparrowhawk.jpg|Mimicry in a brood parasite: Cuckoo mimics sparrowhawk, alarming small birds enough to give time to lay eggs.{{Cite journal |last1=Welbergen |first1=Justin A. |last2=Davies |first2=Nicholas B. |date=2011 |title=A parasite in wolf's clothing: hawk mimicry reduces mobbing of cuckoos by hosts |journal=Behavioral Ecology |volume=22 |issue=3 |pages=574–579 |doi=10.1093/beheco/arr008|doi-access=free}}

File:FinnBirdMimic.jpg|Common hawk-cuckoo resembles a predator, the shikra.{{cite journal |last1=Davies |first1=N. B. |last2=Welbergen |first2=J. A. |title=Cuckoo–hawk mimicry? An experimental test |journal=Proceedings of the Royal Society B |volume=275 |issue=1644 |pages=1817–1822 |year=2008 |doi=10.1098/rspb.2008.0331 |pmid=18467298 |pmc=2587796}}

File:Cuckoo Eggs Mimicking Reed Warbler Eggs.JPG|Egg mimicry: cuckoo eggs (larger) mimic many species of host birds' eggs, in this case of reed warbler.

= Pouyannian =

File:Dasyscolia_ciliata.jpg, a scoliid wasp, attempting to copulate with a flower of the orchid Ophrys speculum]]

In Pouyannian mimicry, a flower mimics a female of a certain insect species, inducing the males of that species to try to copulate with the flower. This is much like aggressive mimicry in fireflies, but with a more benign outcome for the pollinator. The mechanism is named after Maurice-Alexandre Pouyanne, who first described the phenomenon.Correvon H., Pouyanne M. (1916) {{lang|fr|Un curieux cas de mimetisme chez les Ophrydées}}. J. Soc. Nat. Hortic. Fr. 17: 29–31, 41–42, 84.{{cite journal |last1=Pouyanne |first1=M.-A. |year=1917 |title=La fécondation des Ophrys par les insectes |journal=Bulletin de la Société d'histoire naturelle de l'Afrique du Nord |volume=8 |pages=1–2}} It is most common in orchids, which mimic females of the order Hymenoptera (generally bees and wasps), and may account for around 60% of pollinations. Depending on the morphology of the flower, a pollen sac called a pollinium is attached to the head or abdomen of the male. This is then transferred to the stigma of the next flower the male tries to inseminate, resulting in pollination. The mimicry is a combination of visual, by olfaction, and by touch.{{cite journal |last1=Pramanik |first1=Dewi |last2=Dorst |first2=Nemi |last3=Meesters |first3=Niels |last4=Spaans |first4=Marlies |last5=Smets |first5=Erik |last6=Welten |first6=Monique |last7=Gravendeel |first7=Barbara |display-authors=3 |title=Evolution and development of three highly specialized floral structures of bee-pollinated Phalaenopsis species |journal=EvoDevo |volume=11 |issue=1 |date=2020 |page=16 |pmid=32793330 |pmc=7418404 |doi=10.1186/s13227-020-00160-z |doi-access=free }}

= Vavilovian =

File:Secale cereale.jpg is a secondary crop, originally being a mimetic weed of wheat.]]

Vavilovian mimicry is found in weeds that come to share characteristics with a domesticated plant through unintentional selection. It is named after Russian botanist and geneticist Nikolai Vavilov.{{cite journal |last1=Vavilov |first1=N. I. |year=1951 |title=The origin, variation, immunity and breeding of cultivated plants (translation by K. S. Chester) |journal=Chronica Botanica |volume=13 |pages=1–366}} Selection against the weed may occur either by manually killing the weed, or by separating its seeds from those of the crop by winnowing. Vavilovian mimicry illustrates unintentional selection by man. Weeders do not want to select weeds and their seeds that look increasingly like cultivated plants, yet there is no other option. For example, early barnyard grass, Echinochloa oryzoides, is a weed in rice fields and looks similar to rice; its seeds are often mixed in rice and have become difficult to separate through Vavilovian mimicry.{{cite journal |last1=Barrett |first1=S. |year=1983 |title=Mimicry in Plants |journal=Scientific American |volume=257 |issue=3|pages=76–83 |doi=10.1038/scientificamerican0987-76 |bibcode=1987SciAm.257c..76B}} Vavilovian mimics may eventually be domesticated themselves, as in the case of rye in wheat; Vavilov called these weed-crops secondary crops.

= Inter-sexual mimicry =

Inter-sexual mimicry (a type of automimicry, as it is within a single species) occurs when individuals of one sex in a species mimic members of the opposite sex to facilitate sneak mating. An example is the three male forms of the marine isopod Paracerceis sculpta. Alpha males are the largest and guard a harem of females. Beta males mimic females and manage to enter the harem of females without being detected by the alpha males allowing them to mate. Gamma males are the smallest males and mimic juveniles. This also allows them to mate with the females without the alpha males detecting them.{{cite journal |last=Shuster |first=Stephen |date=May 1987 |title=Alternative Reproductive Behaviors: Three Discrete Male Morphs in Paracerceis sculpta, an Intertidal Isopod from the Northern Gulf of California |journal=Journal of Crustacean Biology |volume=7 |issue=2 |pages=318–327 |doi=10.2307/1548612 |jstor=1548612}} Similarly, among common side-blotched lizards, some males mimic the yellow throat coloration and even mating rejection behaviour of the other sex to sneak matings with guarded females. These males look and behave like unreceptive females. This strategy is effective against "usurper" males with orange throats, but ineffective against blue throated "guarder" males, which chase them away.{{cite journal |last=Sinervo |first=B. |author2=C. M. Lively |year=1996 |title=The rock–paper–scissors game and the evolution of alternative male strategies |journal=Nature |volume=380 |issue=6571 |pages=240–243 |doi=10.1038/380240a0 |bibcode=1996Natur.380..240S |s2cid=205026253}}{{cite journal |last1=Sinervo |first1=B. |last2=Miles |first2=D. B. |last3=Frankino |first3=W. A. |last4=Klukowski |first4=M. |last5=Denardo |first5=D. F. |year=2000 |title=Testosterone, Endurance, and Darwinian Fitness: Natural and Sexual Selection on the Physiological Bases of Alternative Male Behaviors in Side-Blotched Lizards |journal=Hormones and Behavior |volume=38 |issue=4|pages=222–233 |doi=10.1006/hbeh.2000.1622 |pmid=11104640 |s2cid=5759575}} Female spotted hyenas have pseudo-penises that make them look like males.{{cite journal |last1=Muller |first1=M. N. |last2=Wrangham |first2=R. |year=2002 |title=Sexual Mimicry in Hyenas |journal=The Quarterly Review of Biology |volume=77 |issue=1|pages=3–16 |doi=10.1086/339199 |pmid=11963460 |s2cid=43440407 |url=http://nrs.harvard.edu/urn-3:HUL.InstRepos:41467428|url-access=subscription }}

Notes

References

=Children's=

Hoff, M. K. (2003) Mimicry and Camouflage. Creative Education. Mankato, Minnesota, USA, Great Britain. {{ISBN|1-58341-237-9}}.

External links

[http://www.ucl.ac.uk/~ucbhdjm/courses/b242/Mimic/Mimic.html Warning colour and mimicry] • Lecture outline from University College London
[http://www.mprinstitute.org/vaclav/Camouflage.htm Camouflage and Mimicry in Fossils]

Mimicry

Category:Polymorphism (biology)

Category:Camouflage mechanisms

Category:Warning coloration