Mimicry
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/* Batesian */
{{otheruses|Mimic (disambiguation)}}
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[[Image:Batesplate ArM.jpg|right|thumb|200px|[[Photographic plate|Plate]] from [[Henry Walter Bates]] (1862) illustrating Batesian mimicry between ''Dismorphia'' species (top row, third row) and various ''Ithomiini'' (Nymphalidae) (second row, bottom row).]]
[[Biology|Biological]] '''mimicry'''<ref>Less commonly known as ''mimetism''.</ref> occurs when a group of organisms,<ref>This 'group' is often a species, but can also be a subgroup such as one particular sex or morph</ref> the ''mimics'', have [[evolution|evolved]] to share common [[perception|perceived]] characteristics with another group, the ''models'',<ref>In its broadest definition mimicry can include non-living models.</ref> through the [[selection|selective]] action of a ''signal-receiver''<ref name="Wickler 1965">{{ cite journal | last =Wickler | first =W. | authorlink =Wolfgang Wickler | year =1965 | month = | title =Mimicry and the evolution of animal communication | journal = [[Nature (journal)|Nature]]| volume =208 | issue = | pages =519–21 | id = | url = | accessdate = | quote =| doi =10.1038/208519a0 }}</ref> or ''dupe''. Collectively this is known as a ''mimicry complex''.<ref name="Wickler 1965" /> The model is usually another species except in cases of [[#Automimicry|automimicry]]. The signal-receiver is typically another intermediate organism like the common [[predator]] of two species, but may actually be the model itself, such as a moth resembling its spider predator.<ref>{{cite web |url=http://neurophilosophy.wordpress.com/2006/12/22/the-moth-in-spiders-clothing/ |title=A moth in spider's clothing <<Neurophilosophy |accessdate=2008-06-07 |publisher= |date=2006-12-22}} (includes video)</ref> As an [[biological interaction|interaction]], mimicry is in most cases advantageous to the mimic and harmful to the receiver, but may increase, reduce or have no effect on the [[fitness (biology)|fitness]] of the model depending on the situation. Models themselves are difficult to define in some cases, for example eye spots may not bear resemblance to any specific organism's eyes, and camouflage often cannot be attributed to any particular model.
[[Image:Mimicry of Siphanta acuta edit1.jpg|thumb|left|A [[planthopper]] mimics a [[leaf]] (mimesis)]]
[[Camouflage]], in which a species appears similar to its surroundings, is essentially a form of visual mimicry, but usually is restricted to cases where the model is non-living or abiotic.<!-- Is it? Many camouflaged organisms mimic a biotic background, such as lichens or algae on a rock, or brown or green vegetative background --> In between camouflage and mimicry is '''mimesis''', in which the mimic takes on the properties of a specific object or organism, but one to which the dupe is indifferent.<ref name=Pasteur>{{ cite journal | last =Pasteur | first =G. | authorlink = | year =1982 | month = | title =A classificatory review of mimicry systems | journal = [[Annual Review of Ecology and Systematics]]| volume =13 | issue = | pages =169–199 | id = | url = | accessdate = | quote =| doi =10.1146/annurev.es.13.110182.001125 }}</ref> The lack of a true distinction between the two phenomena can be seen in animals that resemble twigs, bark, leaves or flowers, in that they are often classified as camouflaged (a plant constitutes its "surroundings"), but are sometimes classified as mimics (a plant is also an organism). [[Crypsis]] is a broader concept that encompasses all forms of detection evasion, such as mimicry, camouflage, hiding etc.<!-- Also need a reference here (crypsis article is in a terrible state) -->
Though mimicry is most obvious to humans in [[visual]] mimics, others senses such as [[olfaction]] (smell) or [[hearing (sense)|hearing]] may be involved, and more than one type of [[signalling theory|signal]] may be employed.<ref name="Wickler">Wickler, W. 1968. Mimicry in plants and animals. McGraw-Hill, New York</ref> Mimicry may involve [[morphology (biology)|morphology]], [[behavior]], and other properties. In any case, the signal always functions to deceive the receiver by preventing it from correctly identifying the mimic. In evolutionary terms, this phenomenon is a form of [[co-evolution]] usually involving an [[evolutionary arms race]], and should not be confused with [[convergent evolution]], which occurs when species come to resemble one another ''independently'' due to similar lifestyles.<!-- A reference here would be good too -->
Mimics may have multiple models during different stages of their [[Biological life cycle|life cycle]], or they may be [[Polymorphism (biology)|polymorphic]], with different individuals imitating different models. Models themselves may have more than one mimic, though [[frequency dependent selection]] favors mimicry where models outnumber mimics. Models tend to be relatively closely [[Common descent|related]] organisms,<ref name=Campbell /> but mimicry of vastly different species is also known. Most known mimics are [[insect]]s,<ref name="Wickler" /> though many other animal mimics including [[mammal]]s are known. [[Plant]]s and [[fungi]] may also be mimics, though less research has been carried out in this area.<ref>Boyden, T. C. (1980) Floral Mimicry by Epidendrum ibaguense (Orchidaceae) in Panama ''Evolution'' '''34''':135-136.</ref><ref>Roy, B. A. (1994) The Effects of Pathogen-Induced Pseudoflowers and Buttercups on Each Other's Insect Visitation ''Ecology'' '''75''':352-358.</ref><ref name=EB>Wickler, Wolfgang (1998). “Mimicry”. ''[[Encyclopædia Britannica]]'', 15th edition. Macropædia 24, 144–151. http://www.britannica.com/eb/article-11910</ref>
<!-- A little on history also needed, as well as a quick run through of the basic types? -->
==Etymology==
Use of the word [[wikt:mimicry|mimicry]] dates back to 1637. It is [[etymology|derived]] from the [[Greek language|Greek]] term ''mimetikos'', "imitative," in turn from ''mimetos'', the verbal adjective of ''mimeisthai'', "to imitate." Originally used to describe people, it was only applied to other forms of life after 1851.<ref>{{citeweb|title=Online Etymology Dictionary|url=http://www.etymonline.com/index.php?search=mimicry&searchmode=none|author=Douglas Harper|date=2007-10-06}}</ref>
==Classification==
<!-- {{Main|Classification of mimicry}} -->
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 (biology)|function]] with respect to the mimic (e.g. avoiding harm), though other parameters can also be used, and multidimensional classifications are required to understand the full picture. For this reason, 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).
===Defensive===
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Defensive or protective mimicry takes place when organisms are able to avoid an encounter that would be harmful to them by deceiving an enemy into treating them as something else. Four such cases are discussed here, the first three of which entail mimicry of an aposematic, harmful organism: Batesian mimicry, where a harmless mimic poses as harmful; Müllerian mimicry, where two harmful species share similar perceived characteristics; and Mertensian mimicry, where a deadly mimic resembles a less harmful but lesson-teaching model. Finally, Vavilovian mimicry, where weeds resemble crops, is discussed.
====Batesian====
{{Main|Batesian mimicry}}
[[Image:Wasp mimicry.jpg|thumb|right|Several species, including several [[hoverfly|hoverflies]], mimic stinging species of wasp.]]
In Batesian mimicry the mimic shares signals similar to the model, but does not have the attribute that makes it unprofitable to predators (e.g. unpalatability). In other words, a Batesian mimic is a [[The Wolf in Sheep's Clothing|sheep in wolf's clothing]]. It is named after [[Henry Walter Bates]], an English naturalist whose work on [[butterflies]] in the [[Amazon rainforest]] (including ''[[The Naturalist on the River Amazons]]'') was pioneering in this field of study.<ref>Bates H. W. 1863. ''The naturalist on the river Amazons''. Murray, London.</ref><ref>Bates, H. W. (1961) Contributions to an insect fauna of the Amazon valley. Lepidoptera: Heliconidae. ''Transactions of the Linnean Society''. '''23''':495-566.</ref> Mimics are less likely to be found out when in low proportion to their model, a phenomenon known as [[negative frequency dependent selection]] which applies in most other forms of mimicry as well. This is not the case in Müllerian mimicry however, which is described next.
Examples:
* [[Lepidoptera]]
** The [[Ash Borer]] (''Podosesia syringae''), a moth of the Clearwing family ([[Sesiidae]]), is a Batesian mimic of the [[Common wasp]] because it resembles the wasp, but is not capable of stinging. A predator that has learned to avoid the wasp would similarly avoid the Ash Borer.
** [[Danaus chrysippus#Mimicry|Plain Tiger]] (''Danaus chrysippus'') - an unpalatable model with a number of mimics.
** [[Euploea core#Mimicry|Common Crow]] (''Euploea core'') - an unpalatable model with a number of mimics. See also under ''Müllerian mimicry'' below.
** ''[[Consul fabius]]'' and ''[[Eresia eunice]]'' imitate unpalatable ''[[Heliconius]]'' butterflies such as ''[[Heliconius ismenius|H. ismenius]]''.<ref name="pinheiro" />
** Several palatable butterflies resemble different species from the highly noxious [[Papilioninae|papilionine]] genus ''[[Battus (butterfly)|Battus]]''.<ref name="pinheiro" />
** Several palatable moths produce ultrasonic click calls to mimic the unpalatable tiger moths.<ref name=tigermoth>Barber, J. R. and W. E. Conner. (2007) Acoustic mimicry in a predator–prey interaction. Proc. Nat. Acad. Sci. 104(22):9331-9334
[http://www.pnas.org/cgi/content/figsonly/104/22/9331]</ref>
* The [[False Cobra]] (''Malpolon moilensis'') is a mildly venomous but harmless [[colubrid]] snake which mimics the characteristic "hood" of an [[Indian cobra]]'s threat display. The [[Eastern Hognose Snake]] (''Heterodon platirhinos'') similarly mimics the threat display of venomous snakes.
<!-- * The [[Bush Viper]] (young) and the [[copperhead snake]]: The copperhead snake is extremely venomous. The Bush Viper is also extremely dangerous when older, but only has a very similar coloring when young. Hence, the viper is still dangerous when young, just not to the same extent as when it is older and changes colors.
[There are no Bush Vipers in America! This seems to be an erroneous vernacular name for some pit viper species or such] -->
* The [[milk snake]] resembles the deadly poisonous [[coral snake]].
* [[Octopuses]] of the genus ''Thaumoctopus'' (the [[Mimic Octopus]] and the "wunderpus") are able to intentionally alter their body shape and color so that they resemble dangerous [[sea snakes]] or [[lionfish]].<ref>[http://marinebio.org/species.asp?id=260 Mimic Octopus, Thaumoctopus mimicus at MarineBio.org<!-- Bot generated title -->]</ref>
====Müllerian====
{{Main|Müllerian mimicry}}
[[Image:Heliconius mimicry.png|right|thumb|The Heliconius butterflies from the tropics of the Western Hemisphere are the classical model for Müllerian mimicry.<ref>Meyer A (2006) Repeating Patterns of Mimicry. PLoS Biol 4(10): e341 doi:10.1371/journal.pbio.0040341</ref>]]
Müllerian mimicry describes a situation where two or more species have very similar warning or aposematic signals and both share genuine [[anti-predation]] attributes (e.g. being unpalatable). At first Bates could not explain why this should be so; if both were harmful why did one need to mimic another? The German naturalist [[Fritz Müller]] put forward the first explanation for this phenomenon: If two species were confused with one another by a common predator, individuals in both would be more likely to survive.<ref>Müller, Fritz (1878) Ueber die Vortheile der Mimicry bei Schmetterlingen. ''Zoologischer Anzeiger'' '''1''': 54–55.</ref><ref>Müller, F. (1879) ''Ituna'' and ''Thyridia''; a remarkable case of mimicry in butterflies. (R. Meldola translation) ''Proclamations of the Entomological Society of London'' 1879:20-29.</ref> 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]] in this respect. The signal receiver is also advantaged by this system, despite being deceived regarding species identity, as it avoids potentially harmful encounters. The usually clear identity of mimic and model are also blurred. In cases 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 is more realistic to speak of each as ''comimics'' than of a distinct 'mimic' and 'model' species, as their warning signals tend to converge toward something intermediate between the two.<ref>Flannery, T. F. (2007) "Community ecology: Mimicry complexes". ''[[Encyclopædia Britannica Online]]''. http://www.britannica.com/eb/article-9117280/community-ecology</ref> Another theoretical problem comes up when one considers that the two species may exist on a continuum from the harmless to the highly noxious, raising the question of where Batesian mimicry ends and Müllerian convergence begins.<ref>Huheey, James E. (1976) Studies in warning coloration and mimicry VII. Evolutionary consequences of a Batesian–Müllerian spectrum: A model for Müllerian mimicry. ''Evolution'' '''30''':86–93.</ref><ref>Benson, W. W. (1977) On the Supposed Spectrum Between Batesian and Mullerian Mimicry. ''Evolution''. '''31''':454-455.</ref>
Examples:
* Lepidoptera
** The [[Monarch Butterfly]] (''Danaus plexippus'') is a member of a Müllerian complex with the [[Viceroy butterfly]] (''Limenitis archippus'') in shared coloration patterns and display behavior. The Viceroy has [[subspecies]] with somewhat different coloration, each one very closely matching the local ''[[Danaus (genus)|Danaus]]'' species. E.g., in [[Florida]], the pairing is of the Viceroy and the [[Queen Butterfly]], and in [[Mexico]], the Viceroy resembles the [[Soldier Butterfly]]. Therefore, the Viceroy is a single species involved in three different Müllerian pairs.<ref name = Ritland1>Ritland, D.B. 1995. Comparative unpalatability of mimetic viceroy butterflies (''Limenitis archippus'') from four south-eastern United States populations. ''Oecologia'' 103: 327-336</ref> This example was long believed to be a case of Batesian mimicry, with the Viceroy being the mimic and the Monarch the model, but it was more recently determined that the Viceroy is actually the ''more'' unpalatable species, though there is considerable individual variation.<ref>{{cite journal |last=Ritland |first=D.|coauthors=L. P. Brower|year=1991 |month= |title=The viceroy butterfly is not a Batesian mimic |journal=[[Nature (journal)|Nature]] |volume=350 |issue= |pages=497–498 |doi =10.1038/350497a0 |url=http://www.nature.com/nature/journal/v350/n6318/abs/350497a0.html |accessdate= 2008-02-23 |quote=Viceroys are as unpalatable as monarchs, and significantly more unpalatable than queens from representative Florida populations.}}</ref> While ''L. archippus'' is really bad-tasting, ''Danaus'' species tend to be toxic rather than just repugnant, due to their different food plants.
** Unpalatable ''[[Euploea]]'' species look very similar. See also under ''Batesian mimicry'' above.
** The genus ''[[Morpho]]'' is palatable but are very strong fliers; birds - even species which are specialized for catching butterflies on the wing - find it very hard to catch them. The conspicuous blue coloration shared by most ''Morpho'' species seems to be a case of Müllerian mimicry.<ref name="pinheiro" />
**The "orange complex" of species, including the [[heliconiine]]s ''[[Agraulis vanillae]]'', ''[[Dryadula phaetusa]]'', and ''[[Dryas iulia]]'' which all taste bad.<ref name="pinheiro">Pinheiro, Carlos E. G. (1996) Palatability and escaping ability in Neotropical butterflies: tests with wild kingbirds (''Tyrannus melancholicus'', Tyrannidae). ''[[Biological Journal of the Linnean Society|Biol. J. Linn. Soc.]]'' '''59'''(4): 351–365. [http://www.ingentaconnect.com/content/ap/bj/1996/00000059/00000004/art00069 HTML abstract]</ref>
**Many different tiger moths make ultrasonic clicking calls to warn bats that they are unpalatable. Presumably a bat may learn to avoid ''any'' signalling moths, which would make this an example of Müllerian mimicry.<ref name=tigermoth />
* Various [[bee]]s and numerous [[vespid]] and [[sphecoidea|sphecoid]] wasps: These animals are examples of Müllerian mimics because they have the [[aposematic]] yellow and black stripes (sometimes black and red, or black and white). Females of most of these species are potentially harmful to predators, fulfilling the second requirement of Müllerian mimicry. However, in essentially all such species, the males are harmless, and can thus be considered automimics of their conspecific females (see below). There are also many genera in these groups where the females are not capable of stinging, and yet still possess aposematic coloration (e.g., the wasp genus ''[[Cerceris]]''), so they are considered Batesian mimics.
====Emsleyan/Mertensian====
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[[Image:Micrurus_tener.jpg|thumb|left|Texas Coral Snake, ''Micrurus tener'']]
''Emsleyan''<ref name=Pasteur /> or ''Mertensian mimicry'' describes unusual cases where deadly prey mimic a less dangerous species. It was first proposed by Emsley<ref>Emsley, M. G. (1966) The mimetic significance of ''Erythrolamprus aesculapii ocellatus'' Peters from Tobago. ''Evolution'' '''20''':663-64.</ref> as a possible answer for the problem of [[Coral Snake]] mimicry in the New World. It was elaborated on by the German biologist [[Wolfgang Wickler]] in a chapter of ''Mimicry in Plants and Animals'',<ref name="Wickler" /> who named it after the German [[herpetologist]] [[Robert Mertens]]<ref>Mertens, R. (1956) Das Problem der Mimikry bei Korallenschlangen. Zool. Jahrb. Syst. '''84''':541-76.</ref> (but see Sheppard (1969)<ref>But Sheppard points out that Hecht and Marien put forward a similar hypothesis ten years earlier (Hecht, M. K. & Marien, D. (1956) The coral snake mimic problem: A reinterpretation. ''Journal of Morphology''. '''98''':335-365), see Sheppard, P. M. (1969) Review of Mimicry in Plants and Animals by Wolfgang Wickler ''The Journal of Animal Ecology'' '''38''': 243.</ref>). This scenario is a little more difficult to understand, as it is usually the most harmful species that is the model. If a predator dies, it cannot [[learning|learn]] to recognize a warning signal, e.g. bright colors in a certain pattern. In other words, there is no advantage in being aposematic if an organism will kill any predators it succeeds in poisoning. It would then be better off camouflaged instead, so as to avoid encounters altogether. If, however, there is another species that is harmful but not deadly, the predator may learn to avoid it. Provided it results in less encounters than camouflage, the deadly species can then profit by mimicking this aposematic organism.
The exception here, ignoring any chance of animals [[observational learning|learning by watching]] a conspecific die (see Jouventin ''et al.'' for a discussion of observational learning and mimicry),<ref>Jouventin, P.; G. Pasteur; J. P. Cambefort (1977) Observational Learning of Baboons and Avoidance of Mimics: Exploratory Tests ''Evolution'' '''31''':214-218.</ref> is the possibility of not having to learn that it is harmful in the first place: [[instinct]]ive genetic programming to be wary of certain signals. In this case, other organisms could benefit from this programming, and Batesian or Müllerian mimics of it could potentially evolve. In fact, it has been shown that some species do have an innate recognition of certain aposematic warnings. Hand-reared [[Turquoise-browed Motmot]]s (''Eumomota superciliosa''), avian predators, instinctively avoid snakes with red and yellow rings.<ref>Smith, S. M. (1975) Innate Recognition of Coral Snake Pattern by a Possible Avian Predator. ''[[Science (journal)|Science]]''. '''187''':759-760.</ref> Other colors with the same pattern, and even red and yellow ''stripes'' with the same width as rings, were tolerated. However, models with red and yellow rings were feared, with the birds flying away and giving [[alarm call]]s in some cases. This provides one alternative explanation to Mertensian mimicry. See Greene and McDiarmid for a review of the subject.<ref>Greene, H. W., McDiarmid, R. W. (1981) Coral snake mimicry: Does it occur? ''Science'' '''213''':1207-12.</ref>
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Examples:
* Some [[Milk Snake]] (''Lampropeltis triangulum'') subspecies (harmless), the moderately toxic [[False Coral Snakes]] (genus ''Erythrolamprus''), and the deadly [[Coral Snakes]] all have a red background color with black and white/yellow stripes. In this system, both the milk snakes and the deadly coral snakes are mimics, whereas the false coral snakes are the model.<!-- Merge into text -->
====Wasmannian====
<!-- Pasteur classifies this as mutualistic/commensalist. Not sure where to put it. -->
Wasmannian mimicry refers to cases where the mimic resembles a model along with which it lives ([[inquiline|inquiline]]) in a nest or colony. Most of the models here are [[eusociality|social]] insects such as ants, termites, bees and wasps.<ref>Wasmann, E. 1894. Kritisches Verzeichniss der myrmecophilin und termitophilen Arthropoden. Felix Dames, Berlin xi + 231 pp. </ref>
====Mimetic weeds====
{{Main|Vavilovian mimicry}}
<!-- ! Pasteur classifies this as 'aggressive/mutualistic/reproductive', and definitely not defensive -->
[[Image:Secale cereale.jpg|right|thumb|[[Rye]] is a secondary crop, originally being a mimetic weed of [[wheat]].]]
Vavilovian mimicry describes [[weed]]s which comes to share characteristics with a [[Crop (agriculture)|domesticated plant]] through [[artificial selection]].<ref name=Pasteur /> It is named after Russian [[botanist]] and [[geneticist]] [[Nikolai Vavilov]].<ref>Vavilov, N. I. (1951) The origin, variation, immunity and breeding of cultivated plants. (Translation by K. S. Chester) ''Chronica Botanica'' 13:1-366.</ref> Selection against the weed may occur either by manually killing the weed, or separating its seeds from those of the crop. The latter process, known as [[winnowing]], can be done manually or by a machine.
Vavilovian mimicry presents an illustration of unintentional (or rather 'anti-intentional') selection by man. While some cases of artificial selection go in the direction desired, such as [[selective breeding]], this case presents the opposite characteristics. Weeders do not want to select weeds that look increasingly like the cultivated plant, yet there is no other option. A similar problem in agriculture is [[pesticide resistance|pesticide]]. Vavilovian mimics may eventually be domesticated themselves, and Vavilov called these weeds-come-crops ''secondary crops''.
It can be classified as defensive mimicry in that the weed mimics a protected species. This bears strong similarity to Batesian mimicry in that the weed does not share the properties that give the model its protection, and both the model and the dupe (in this case people) are both harmed by its presence. There are some key differences, though; in Batesian mimicry the model and signal receiver are enemies (the predator would eat the protected species if could), whereas here the crop and its human growers are in a mutualistic relationship: the crop benefits from being dispersed and protected by people, despite being eaten by them. In fact, the crop's only 'protection' relevant here is its usefulness to humans. Secondly, the weed is not eaten, but simply destroyed. The only motivation for killing the weed is its effect on crop yields. Finally, this type of mimicry does not occur in ecosystems unaltered by humans.
One case is ''[[Echinochloa oryzoides]]'', a species of grass which is found as a weed in [[rice]] (''Oryza sativa'') fields. The plant looks similar to rice and its seeds are often mixed in rice and difficult to separate. This close similarity was enhanced by the weeding process which is a selective force that increases the similarity of the weed in each subsequent generation.<ref>Barrett, S. (1983) Mimicry in Plants ''[[Scientific American]]''. '''257''': 76-83.</ref>
====Protective egg decoys====
<!-- Need an image here! Worthy of its own article? Could expand on in an article on plant mimicry perhaps? Also add this to relevant articles (Heliconius, passiflora) and include other example species. -->
Unlike the above forms of mimicry, ''Gilbertian mimicry'' involves only two species. The potential host/prey drives away its parasite/predator by mimicking it, the reverse of host-parasite aggressive mimicry. It was coined by Pasteur as a term for such rare mimicry systems,<ref name=Pasteur /> and is named after the American [[ecologist]] Lawrence E. Gilbert.<ref>Gilbert, L. E. (1975) Ecological consequences of a coevolved mutualism between butterflies and plants. In L. E. Gilbert, P. H. Raven (eds.) ''Coevolution of Animal and Plants'' pp. 210-40. Austin & London: University of Texas Press</ref> <!--It should not to be confused with the [[Gilbert and Sullivan]] sense of the word, meaning absurd or comic. -->
This form of protective mimicry occurs in the genus ''[[Passiflora]]''. The leaves of this plant contain toxins which deter herbivorous animals, however some ''[[Heliconius]]'' butterfly larvae have evolved enzymes which break down these toxins, allowing them to [[specialist species|specialize]] on this genus. This has created further selection pressure on the host plants, which have evolved [[stipules]]<!-- Only stipules? --> that mimic mature ''Heliconius'' eggs near the point of hatching. These butterflies tend to avoid laying eggs near each existing ones, which helps avoid exploitative [[intraspecific competition]] between caterpillars—those that lay on vacant leaves provide their offspring with a greater chance of survival. Additionally, most ''Heliconius'' larvae are [[cannibalism (zoology)|cannibalistic]], meaning those leaves with older eggs will hatch first and eat the new arrivals. Thus, it seems such plants have evolved egg dummies due to these grazing herbivore enemies. The decoy eggs are also [[nectaries]] though, attracting predators of the caterpillars such as ants and wasps. The extent of their mimetic function is therefore slightly more difficult to assess.<ref name=Campbell>[[Campbell, N. A.]] (1996) Biology (4th edition), Chapter 50. Benjamin Cummings, New York ISBN 0-8053-1957-3</ref>
The use of eggs is not essential to this system, only the species composition and protective function. Many other forms of mimicry also involve eggs, such as cuckoo eggs mimicking those of their host (the reverse of this situation), or plants seeds being dispersed by ants, who treat them as they would their own eggs.
====Protective mimicry within a species====
[[Image:Monarch Butterfly Danaus plexippus Caterpillar 2000px.jpg|right|thumb|Monarch caterpillars, shown feeding, vary in toxicity depending on their diet.]]
''Browerian mimicry''<ref name=Pasteur />, named after Lincoln P. Brower and Jane Van Zandt Brower,<ref>Brower, L. P. (1970) Plant poisons in a terrestrial food chain and implications for mimicry theory. In K. L. Chambers (ed) ''Biochemical Coevolution'' Corvallis, OR: Oregon State Univ. pp. 69-82.</ref><ref>Brower, L. P., Brower, J. V. Z., Corvino, J. M. (1967) Plant poisons in a terrestrial food chain. ''Proceedings of the National Academy of Sciences USA'' '''57''':893-98.</ref> is a 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. One example is Monarch Butterflies (''Danaus plexippus''), which feed on [[milkweed]] species of varying toxicity. This species stores toxins from its host plant, which are maintained even in the adult ([[imago]]) form. As the levels of toxin will vary depending on diet during the larval stage, some individuals will be more toxic than others. The less palatable organisms will therefore be mimics of the more dangerous individuals, with their likeness already perfected. This need not be the case however; in sexually dimorphic species one sex may be more of a threat than the other, which could mimic the protected sex. Evidence for this possibility is provided by the behavior of a monkey from [[Gabon]], which regularly ate male moths of the genus ''Anaphe'', but promptly stopped after it tasted a noxious female.<ref>Bigot, L., Jouventin, P. (1974) Quelques expériences de comestibilité de Lépidoptères gabonais faites avec le mandrill, le cercocèbe à joues grises et legarde-boeufs. ''Terre Vie'' '''28''':521-43.</ref>
===Aggressive===
{{Main|Aggressive mimicry}}
Aggressive mimicry describes predators (or [[parasite]]s) which share the same characteristics as a harmless species, allowing them to avoid detection by their prey (or [[host (biology)|host]]). It is less often known as ''Peckhamian mimicry'' after [[George and Elizabeth Peckham]].<ref>Peckham, E. G. (1889) Protective resemblances of spiders. ''Occasional Papers of Natural History Society of Wisconsin'' '''1''':61-113.</ref><ref>Peckham, E. G. & G. W. Peckham (1892) Ant-like spiders of the family Attidae. ''Occasional Papers of Natural History Society of Wisconsin'' '''2''':1-84.</ref> The mimic may resemble the prey or host itself, or another organism which is either neutral or beneficial to the signal receiver. In this class of mimicry the model may be affected negatively, positively or not at all. Just as parasites can be treated as a form of predator,<ref name=Ecology>Begon, M., Townsend, C., Harper, J. (1996) ''[[Ecology: Individuals, populations and communities]]'' (Third edition) Blackwell Science, London</ref> host-parasite mimicry is treated here as a subclass of aggressive mimicry.
The mimic may have a particular significance for duped prey. <!-- Replace the following with anglerfish example (See Wickler and also The Compleat Angler: Aggressive Mimicry in an Antennariid Anglerfish
Theodore W. Pietsch; David B. Grobecker
Science > New Series, Vol. 201, No. 4353 (Jul., 1978), pp. 369-370 --> One such case is [[spider]]s, amongst which aggressive mimicry is quite common in both in luring prey and stealthily approaching predators.<ref>Jackson, R. R. (1995) Eight-legged tricksters: Spiders that specialize at catching other spiders. ''BioScience'' '''42''':590–98.</ref> One case is the [[Golden Orb Weaver]] (''Nephila clavipes''), which spins a conspicuous golden colored web in well-lit areas. Experiments show that bees are able to associate the webs with danger when the yellow pigment is not present, as occurs in less well-lit areas where the web is much harder to see. Other colors were also learned and avoided, but bees seemed least able to effectively associate yellow pigmented webs with danger. Yellow is the color of many nectar bearing flowers, however, so perhaps avoiding yellow is not worth while. Another form of mimicry is based not on [[color]] but pattern. Species such as ''[[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 guide]]s. Spiders change their web day to day, which can be explained by bee's ability to remember web patterns. Bees are able to associate a certain pattern with a spatial location, meaning the spider must spin a new pattern regularly or suffer diminishing prey capture.<ref>Craig, C. L. (1995) Webs of Deceit. ''[[Natural History (magazine)|Natural History]]'' '''104''' (3): 32-35.</ref>
Another case is where males are lured towards what would seem to be a sexually receptive female; the model in this situation being the same species as the dupe. Beginning in the 1960s, James E. Lloyd's investigation of female [[firefly|fireflies]] of the genus ''[[Photuris (genus)|Photuris]]'' revealed they emit the same light signals that females of the genus ''[[Photinus (genus)|Photinus]]'' use as a mating signal.<ref>Lloyd, J. E. (1965) Aggressive Mimicry in Photuris: Firefly Femmes Fatales Science 149:653-654.</ref> Further research showed male fireflies from several different [[genus|genera]] are attracted to these "[[femmes fatales]]", and are subsequently captured and eaten. Female signals are based on that received from the male, each female having a repertoire of signals matching the delay and duration of the female of the corresponding species. This mimicry may have evolved from non-mating signals that have become modified for predation.<ref>Lloyd, J. E. (1975) Aggressive Mimicry in Photuris Fireflies: Signal Repertoires by Femmes Fatales. ''Science''. '''187''':452-453.</ref>
Some [[carnivorous plant]]s may also be able to increase their rate of capture through mimicry.<ref>Moran, Jonathan A. (1996) Pitcher dimorphism, prey composition and the mechanisms of prey attraction in the pitcher plant ''Nepenthes rafflesiana'' in Borneo. ''Journal of Ecology'' '''84''':515–525.</ref>
[[Image:Epinephelus tukula is cleaned by two Labroides dimidiatus.jpg|left|thumb|Two Bluestreak cleaner wrasse cleaning a Potato grouper, ''Epinephelus tukula'']]
Luring is not a necessary condition however, as the predator will still have a significant advantage by simply not being identified as such. They may resemble a mutualistic [[symbiont]] or a species of little relevance to the prey.
A case of the former situation is a species of [[cleaner fish]] and its mimic, though in this example the model is greatly disadvantaged by the presence of the mimic. Cleaner fish are the allies of many other species, which allow them to eat their parasites and dead skin. Some allow the cleaner to venture inside their body to hunt these parasites. However, one species of cleaner, the [[Bluestreak cleaner wrasse]] (''Labroides dimidiatus''), is the unknowing model of a mimetic species, the Sabre-toothed blenny (''Aspidontus taeniatus''). This [[wrasse]], shown to the left cleaning a [[grouper]] of the genus ''[[Epinephelus]]'', resides in [[coral reef]]s in the Indian and the Pacific Oceans, and is recognized by other fishes who then allow it to clean them. Its imposter, a species of [[blenny]], lives in the [[Indian Ocean]] and not only looks like it in terms of size and [[animal coloration|coloration]], but even mimics the cleaner's 'dance'. Having fooled its prey into letting its guard down, it then bites it, tearing off a piece of its fin before fleeing the scene. Fish [[grazing|grazed]] upon in this fashion soon learn to distinguish mimic from model, but because the similarity is close between the two they become much more cautious of the model as well, such that both are affected. Due to victim's ability to discriminate between foe and helper, the blennies have evolved close similarity, right down to the regional level.<ref>Wickler, W. (1966) Mimicry in Tropical Fishes. ''Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences''. '''251''':473-474.</ref>
Another interesting example that does not involve any luring is the [[Zone-tailed Hawk]], which resembles the [[Turkey Vulture]]. It flies amongst the vultures, suddenly breaking from the formation and ambushing its prey.<ref>Willis, E. O. (1963) Is the Zone-Tailed Hawk a Mimic of the Turkey Vulture? ''The Condor'' '''65''':313-317.</ref> Here the hawk's presence is of no evident significance to the vultures, affecting them neither negatively or positively.
====Parasites====
Parasites can also be aggressive mimics, though the situation is somewhat different than those outlined above.
Some of the predators described have a feature that draws prey, and parasites can also mimic their host's natural prey, but are eaten themselves, a pathway into their host. ''[[Leucochloridium]]'', a genus of [[flatworm]], matures in the digestive system of [[songbird]]s, their eggs then passing out of the bird via the [[feces]] . They are then taken up by ''[[Succinea]]'', a terrestrial snail. The eggs develop in this [[intermediate host]], and then must find of a suitable bird to mature in. Host birds do not eat snails though, so the sporocyst must find some strategy to reach its host's intestine. For this function, they are brightly colored and move in a pulsating fashion. A sporocyst-sac pulsates in the snail's eye stalks,<ref>See [http://www.semioticon.com/seo/M/images/mimicry_2.jpg here] for a photo.</ref><ref>Moore, J. 2002. Parasites and the behavior of animals. Oxford University
Press, Oxford.</ref> 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.<ref name=Wickler /> A nematode (''Myrmeconema'') 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 is held raised and this possibly increases the chances of the ant being eaten by birds. The droppings of birds are collected by other ants and fed to their brood, thereby helping to spread the nematode.<ref>Yanoviak, S. P., M. Kaspari, R. Dudley, and G. Poinar, Jr. 2008. Parasite-induced fruit mimicry in a tropical canopy ant. The American Naturalist 171: 536-544. [http://www.canopyants.com/2008_AmNat.pdf PDF]</ref>
In an unusual case, [[planidium]] larvae of some beetles of the genus ''[[Meloe]]'' will form a group and produce a [[pheromone]] that mimics the sex attractant of its host [[Apidae|bee]] species; when the male bee arrives and attempts to mate with the mass of larvae, they climb onto his abdomen, and from there transfer to a female bee, and from there to the bee nest to parasitize the bee larvae.<ref>Leslie Saul-Gershenz 2007. Bee nest parasites (''Meloe franciscanus'') use sexual deception to obtain transport to host bee (''Habropoda pallida'') nest. ESA Annual Meeting, 2007 [http://esa.confex.com/esa/2007/techprogram/paper_32157.htm Abstract]</ref>
Host-parasite mimicry is a two species system where a parasite mimics its own host.<!-- Limited to two species cases? --> [[Cuckoo]]s are a canonical example of [[brood parasitism]], a form of [[kleptoparasitism]] where the mother has its offspring raised by another unwitting organism, cutting down its the biological mother's [[parental investment]] in the process. Cases of ''intraspecific'' brood parasitism, where a female lays in conspecific's nest, as illustrated by the [[Goldeneye (duck)|Goldeneye]] duck (''Bucephala clangula''),<ref>Andersson, M. & Eriksson, M.O.G. 1982 Nest parasitism in goldeneyes ''Bucephala clangula'': some evolutionary aspects. American Naturalist 120, 1-16 (1982)</ref> do not represent a case of mimicry.<!-- Expand on cuckoo example -->
===Reproductive===
Reproductive mimicry occurs when the actions of the dupe directly aid in the mimic's [[reproduction]]. This is common in plants, which may have deceptive flowers that do not provide the reward they would seem to. Other forms of mimicry have a reproductive component, such as Vavilovian mimicry involving seeds, and brood parasitism, which also involves aggressive mimicry.
====Mimicry of flowers====
<!-- Could expand on these two in an article on plant or floral mimicry perhaps? -->
<!--[[Image:Cardinal flower20010810.PNG|right|thumb|''[[Lobelia cardinalis]]'' is believed to be a Bakerian mimic, attracting [[humming bird]]s without producing [[nectar]].<ref>Williamson, G. B.; E. M. Black (1981) Mimicry in Hummingbird-Pollinated Plants? ''Ecology'' '''62''':494-496 </ref>]] Is this correct? Or is it a Dodsonian mimic? -->
''Bakerian mimicry'', named after [[Herbert G. Baker]],<ref>Baker H. 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. ''Variation, breeding, and conservation of tropical trees''. Academic Press, London, U.K.</ref> is a form of automimicry where female [[flower]]s 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]].<ref>Bawa, K. S. (1980) Mimicry of male by female flowers and intrasexual competition for pollinators in ''Jacaratia dolichaula'' (D. Smith) Woodson (Caricaceae). ''[[Evolution (journal)|Evolution]]'' '''34''':467-74.</ref>
Like Bakerian mimicry, ''Dodsonian mimicry'' is a form of reproductive floral mimicry, but the model belongs to a different species than the mimic. The name refers to [[Calaway H. Dodson]].<ref>Dodson, C. H., Frymire, G. P. (1961) Natural pollination of orchids. ''Missouri Botanical Garden Bulletin'' '''49''':133-39.</ref> By providing similar sensory signals as the model flower, it can lure its pollinators. Like Bakerian mimics, no nectar is provided. ''[[Epidendrurn ibaguense]]'' of the family [[Orchidaceae]] resembles flowers of ''[[Lantana camara]]'' and ''[[Asclepias curassavica]]'', and is pollinated by Monarch Butterflies and perhaps [[hummingbird]]s.<ref>Boyden, T. C. (1980) Floral mimicry by Epidendrurn ibaguense (Orchidaceae) in Panama. ''Evolution'' '''34''':135-36.</ref> Similar cases are seen in some other species of the same family. The mimetic species may still have pollinators of its own though, for example a [[lamellicorn beetle]] which usually pollinates correspondingly colored ''[[Cistus]]'' flowers is also known to aid in pollination of ''[[Ophrys]]'' species that are normally pollinated by bees.<ref>Kullenberg, B. (1961) Studies in ''Ophrys'' pollination. ''Zool. Bidr. Uppsala'' '''34''':1-340.</ref>
====Pseudocopulation====
[[Image:Ophrys insectifera Saarland 05.jpg|right|thumb|The [[Fly Orchid]] (''Ophrys insectifera'').]]
{{Further|[[Pseudocopulation]]}}
''Pseudocopulation'' occurs when a flower mimics a female of a certain insect species, the males of which try to copulate with it. This is much like the aggressive mimicry in fireflies described above, but with a much more benign outcome for the pollinator. This form of mimicry has been called ''Pouyannian mimicry'',<ref name=Pasteur /> after <!-- Can't find his first name -->Pouyanne, who first described the phenomenon.<ref>Correvon H., Pouyanne A. (1916) Uncurieux cas de mimetisme chez les Ophrydees. ''J. Soc. Nat. Hortic. Fr.'' '''17''':29–31, 41–42, 84.</ref><ref>Pouyanne, M. (1917) La fécondation des ''Ophrys'' par les insectes. ''Bull. Soc. Hist. Nat. Afr. Nord'' '''8''':1-2.</ref> 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.<ref name=Orchid>van der Pijl, L., Dodson, C. H. (1966) ''Orchid Flowers; Their Pollination and Evolution''. Coral Gables, FL: Univ. Miami Press</ref> Depending on the morphology of the flower, a pollen sac called a [[pollinia]] is attached to the head or abdomen of the male. This is then transferred to the [[Stigma (flower)|stigma]] of the next flower the male tries to inseminate, resulting in pollination. Visual mimicry is the most obvious sign of this deception for humans, but the visual aspect may be minor or non-existent. It is the senses of [[touch]] and [[olfaction]] that are most important.<ref name=Orchid />
===Automimicry===
<!-- {{Main|Automimicry mimicry}}
A picture is requested here, also need to discuss self-mimicry, auto-mimesis -->
''Automimicry'' or ''intraspecific mimicry'' occurs within a single species, one case being where one part of an organism's body resembles another part. Examples include snakes in which the tail resembles the head and show behavior such as moving backwards to confuse predators and insects and fishes with [[eyespot (mimicry)|eyespot]]s on their hind ends to resemble the head. The term is also used when the mimic imitates other morphs within the same species. When males mimic females or ''vice versa'' this may be referred to as [[sexual mimicry]].
Examples:<!-- Merge into text -->
* Many insects have filamentous "tails" at the ends of their wings which are combined with patterns of markings on the wings themselves to create a "false head" which misdirects predators (e.g., [[hairstreak]] butterflies).
* Several [[pygmy owl]]s bear "false eyes" on the back of their head to fool predators into believing the owl is alert to their presence.
* The yellow throated males of the [[Common Side-blotched Lizard]] use a 'sneaking' strategy in [[mating]]. They look and behave like unreceptive females. This [[evolutionary game theory|strategy]] is effective against 'usurper' males with orange throats, but ineffective against blue throated 'guarder' males, which will chase them away. <ref>Sinervo B.; Miles D.B.; Frankino W.A.; Klukowski M.; DeNardo D.F. (2000) Testosterone, Endurance, and Darwinian Fitness: Natural and Sexual Selection on the Physiological Bases of Alternative Male Behaviors in Side-Blotched Lizards. ''Hormones and Behavior''. '''38''':222-233.</ref>
*Female [[hyena]]s have [[pseudo-penis]]es which make them look like males.<ref>Muller, M. N.; Wrangham, R. (2002) Sexual Mimicry in Hyenas ''The Quarterly Review of Biology'' '''77''':3-16.</ref>
===Other===
[[Image:FinnBirdMimic.jpg|thumb|Some [[Hierococcyx varius|hawk-cuckoos]] resemble [[Accipiter badius|hawks]] like the [[Shikra]].<ref>{{cite journal|journal=Proceedings of the Royal Society B: Biological Sciences
|title=Cuckoo–hawk mimicry? An experimental test
|volume=275
|issue=1644
|pages=1817–1822
|year=2008
|author=Davies, NB & JA Welbergen
|doi=10.1098/rspb.2008.0331
}}</ref>
]]
Some forms of mimicry do not fit easily within the classification given above.
[[Owl butterflies]] (genus ''Caligo'') bear eye-spots on the underside of their wings; if turned upside-down, their undersides resemble the face of an [[owl]] (such as the [[Short-eared Owl]] or the [[Tropical Screech Owl]]) for which in turn the butterfly predators - small [[lizard]]s and birds - would be fooled.<ref>See [http://www.livewild.org/CostaRica/Pics/a6024.jpg here] for a photo</ref> Thus it has been supposed that the eye-spots are a form of Batesian mimicry. However, the pose in which the butterfly resembles an owl's head is not normally adopted in life. Recently zoologists have shown experimentally that eye-spots are not a form of mimicry and do not deter predators because they look like eyes, rather patterns on moth wings deter predators due to conspicuousness. <ref>Conspicuousness, not eye mimicry, makes ‘‘eyespots’’ effective antipredator signals (Martin Stevens, Chloe J. Hardman, and Claire L. Stubbins) Behavioral Ecology doi:10.1093/beheco/arm162</ref>
Another case is floral mimicry induced by the [[discomycete fungus]] ''[[Monilinia vaccinii-corymbosi]]''.<ref>Batra L. R.; Batra, S. (1985) Floral Mimicry Induced by Mummy-Berry Fungus Exploits Host's Pollinators as Vectors ''Science'' '''228''':1011-1013.</ref> In this unusual case, a fungal [[plant pathogen]] infects [[leaf|leaves]] of [[blueberries]], causing them to secrete sugary substances including glucose and fructose, in effect mimicking the [[nectar]] of flowers. To the naked eye the leaves do not look like flowers, yet strangely they still attract pollinating insects like bees. As it turns out, the sweet secretions are not the only cues—the leaves also reflect [[ultraviolet]], which is normally absorbed by the plant's leaves. Ultraviolet light is also employed by the host's flowers as a signal to insects, which have visual systems quite capable of picking up this low wavelength (300-400nm) radiation. The fungus is then transferred to the ovaries of the flower where it produces mummified, inedible berries, which overwinter before infecting new plants. This case is unusual in that the fungus benefits from the deception, but it is the leaves which act as mimics, being harmed in the process. It bears similarity to host-parasite mimicry, but the host does not receive the signal. It also has a little in common with automimicry, but the plant does not benefit from the mimicry, and the action of the pathogen is required to produce it.
==Evolution==
<!-- {{Main|Evolution of mimicry}} -->
It is widely accepted that mimicry [[evolution|evolves]] as a positive adaptation; that is, the mimic gains fitness ''via'' [[convergent evolution]] which results in resemblance to another species, though there are a few who have suggested that evolution is non-adaptive or merely a result of structural similarities. The lepidopterist (and sometime author) [[Vladimir Nabokov]] argued that much of insect mimicry, including the Viceroy/Monarch mimicry, resulted from the fact that coloration patterns in both species simply had a common structural basis, and thus the tendency for convergence by chance was high.<ref>Alexander, Victoria N. [http://www.dactyl.org/directors/vna/papers/InsectMimicry.pdf Nabokov and Insect mimicry]. ''Nabokov Studies''</ref> However, this very example provides evidence precisely to the contrary, as the viceroy's color pattern is completely unlike any of the species to which it is closely related, and the viceroy itself has three color forms, each adapted to resemble a different species of ''[[Danaus (genus)|Danaus]]''.<ref name = Ritland1/> Likewise, this example is based on two organisms that are indeed fairly similar in structure (both butterflies), while a great many cases of mimicry (especially in large Batesian/Mũllerian complexes) involve insects from multiple [[order (biology)|orders]] that share virtually no structural similarities whatsoever; [[beetle]]s, [[true bug]]s, [[moth]]s, [[wasp]]s, [[bee]]s, and [[fly|flies]] may all belong to a single mimetic complex, despite profound differences.<ref name="Wickler" />
<!-- There is a problem here about the idea of directed evolution. -->
The most widely accepted model used to explain the evolution of mimicry in butterflies is the two-step hypothesis. In this model the first step involves mutation in modifier genes that regulate a complex cluster of linked genes associated with large changes in morphology. The second step consists of selections on genes with smaller phenotypic effects and this leading to increasing closeness of resemblance. This model is supported by empirical evidence that suggests that there are only a few single point mutations that cause large phenotypic effects while there are numerous others that produce smaller effects. Some regulatory elements are now known to be involved in a [[supergene]] that is involved in the development of butterfly color patterns. Computational simulations of population genetics have also supported this idea.<ref>Holmgren, N. M. A and M. Enquist 1999. Dynamics of mimicry evolution. Biological Journal of the Linnean Society. 66:145–158. [http://www.comp.leeds.ac.uk/biosystems/reading/paper/dynamics.pdf PDF]</ref>
==See also==
*[[Biomimicry]]
*[[Community ecology]]
*[[Evolutionary ecology]]
*[[Fixed action pattern#Mimicry|Code-breaking]]
*[[Molecular mimicry]]
*[[Preadaptation]]
*[[Semiotics]]
*[[Thanatosis]] ('playing dead')
===Similar terms===
*''Mimetic'' is an adjective used to describe cases of mimicry, but is also used in mathematics (see [[mimetic]]). This should not be confused with [[memetics]], the scientific study of [[meme]]s.
*[[Mimesis]] also refers to [[imitation]], especially relating to the [[art]]s.
==Further reading==
*Vane-Wright RI. 1976. A unified classification of mimetic resemblances. ''Biol. J. Linn. Soc.'' 8:25-56
*Cott, H.B. (1940) ''Adaptive Coloration in Animals''. Methuen and Co, Ltd., London ISBN 0416300502
*[[Wolfgang Wickler|Wickler, W.]] (1968) ''Mimicry in Plants and Animals'' (Translated from the German) McGraw-Hill, New York. ISBN 0070701008
*Edmunds, M. 1974. ''Defence in Animals: A Survey of Anti-Predator Defences''. Harlow, Essex & NY: Longman 357 p. ISBN 0582441323
*Owen, D. (1980) ''Camouflage and Mimicry''. Oxford University Press ISBN 0192176838
*Pasteur, Georges (1982). “A classificatory review of mimicry systems”. ''Annual Review of Ecology and Systematics'' '''13''': 169–199.
*Brower, L. (ed.) (1988). ''Mimicry and the Evolutionary Process''. Chicago: The University of Chicago Press. ISBN 0226076083 (a supplement of volume 131 of the journal ''[[American Naturalist]]'' dedicated to [[E. B. Ford]].)
*[[Graeme Ruxton|Ruxton, G. D.]]; Speed, M. P.; Sherratt, T. N. (2004). ''Avoiding Attack. The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry''. Oxford: Oxford University Press. ISBN 0198528604
*Evans, M. A. (1965) Mimicry and the Darwinian Heritage ''Journal of the History of Ideas'' '''26''' (2): 211-220.
*Wiens, D. (1978) Mimicry in Plants. ''Evolutionary Biology''. '''11''':365–403.
*Dafni, A. (1984) Mimicry and Deception in Pollination ''Annual Review of Ecology and Systematics'' '''15''' : 259-278.
*An introductory book for a younger audience: Hoff, M. K. (2003) ''Mimicry and Camouflage''. Creative Education. Mankato, Minn. Great Britain. ISBN 1583412379
==References==
{{reflist|2}}
==External links==
{{Commonscat|Mimicry}}
* [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]
* [http://www.colostate.edu/Depts/Entomology/courses/en570/papers_1996/bernklau.html Chemical Mimicry in Pollination]
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{{evo ecol}}
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