Müllerian mimicry is a natural phenomenon in which two or more poisonous or venomous species, that may or may not be closely related and share one or more common predators, have come to mimic each other's warning signals. It is named after the German naturalist Fritz Müller, who first proposed the concept in 1878.
The phenomenon can be understood by imagining two poisonous species that do not resemble one another and are also prey to a common predator. Occasionally, individuals of the predatory third species will encounter one or the other type of noxious prey, and thereafter avoid it. Predators that avoid only one or the other type of harmful species provide no benefit to individuals of the species that is not avoided. Therefore, there is an advantage to be gained in the appearance of the two prey species. This is because a predator that learns to avoid either species in a pair of species exhibiting Müllerian mimicry learns, in effect, to avoid both.
This strategy is usually contrasted with Batesian mimicry, in which one harmless species adopts the appearance of another, harmful species to gain the advantage of predators' avoidance. However, because comimics may have differing degrees of protection, the distinction between Müllerian and Batesian mimicry is not absolute, and there can be said to be a spectrum between the two forms. Additionally, a species may be a Batesian mimic to one predator and a Müllerian mimic to another. Viceroy butterflies and monarchs (admiral butterflies) have often been called Batesian mimics; however, this is not the case as both are poisonous which makes them Müllerian mimics. Extensive mitochondrial DNA analysis of admiral butterflies has led to the discovery that the viceroy is the basal lineage of two western sister species in North America. The variation in the wing patterns are thought to precede the evolution of toxicity therefore challenging the hypothesis that the toxicity of the admiral butterflies is a conserved characteristic from a common ancestor. This explanation suggests that because some of the species of admiral butterflies that evolved after the node split from the viceroy lineage are not poisonous but look similar to their ancestor, the propensity for chemical defense is an analogous adaptation (an example of convergent evolution) that evolved separately after the species developed different phenotypic wing patterns. While Batesian and Müllerian mimicry are commonly given examples of mimicry, there is often little or no mention of other forms. There are many other types of mimicry however, some very similar in principle, others far separated. For example in aggressive mimicry a predator might mimic the food of its prey, luring them towards it and improving its foraging success.
Müllerian mimicry need not involve visual mimicry; it may employ any of the senses. For example, many snakes share the same auditory warning signals, forming an auditory Müllerian mimicry ring. More than one common signal may show convergences by the parties. While model and mimic are often closely related species, Müllerian mimicry between very distantly related taxa also occurs. And according to recent research, there may be more than just taste and population size that affect Müllerian mimetic relationships. For example: co-mimicry, a mutualistic relationship that occurs when the mimicking population is polymorphic and resembles more than one model. This keeps the ratio of mimic to model individuals for any particular coloration low, increasing overall fitness for both parties. The negative correlation between the amount of mimics and the “survivability” of both species involved. This suggests that it is reproductively beneficial for both species if the number of models outnumber the mimics; this increases the negative interactions between predator and prey.
Some insight into the evolution of mimetic color mimicry in Lepidoptera in particular can be seen through the study of the Optix gene. The Optix gene is responsible for the Heliconius butterflies signature red wing patterns that help it signal to predators that it is toxic. By sharing this coloration with other poisonous red winged butterflies the predator may have pursued previously the Heliconius butterfly increases its chance of survival through association. By mapping the genome of many related species of Heliconius butterflies “show[s] that the cis-regulatory evolution of a single transcription factor can repeatedly drive the convergent evolution of complex color patterns in distantly related species…”. This suggests that the evolution of a non-coding piece of DNA that regulates the transcription of nearby genes can be the reason behind similar phenotypic coloration between distant species, making it hard to determine if the trait is homologous, meaning that it evolved in both species because it existed in a previous ancestor, or that it is convergent in that both species came to a similar phenotypic expression through their own means and mutations.
It is commonly believed that males would be more likely to co-mimic than females (generally being the choosier sex) but in the case of sexually dimorphic species, females are the ones that tend to be mimetic.
Müllerian mimicry was proposed by the German zoologist and naturalist Johann Friedrich Theodor Müller (1821–1897), always known as Fritz. An early proponent of evolution, Müller offered the first explanation for resemblance between certain butterflies that had puzzled the English naturalist Henry Walter Bates, who, like Müller, spent a significant part of his life in Brazil. Müller had also seen these butterflies first hand, and collected specimens like Bates.
Understanding Müllerian mimicry is impossible without first understanding aposematism, or warning signals. Dangerous organisms with these aposematic signals are avoided by predators, who quickly learn after a bad experience not to pursue the same prey again. Learning is not actually necessary for animals which instinctively avoid certain prey; however, learning from experience is much more common. The underlying concept with predators that learn is that the warning signal makes the harmful organism easier to remember than if it remained as cryptic as possible (e.g. being still and silent, providing no scent, and blending in with the background). Aposematism and crypsis are in this way opposing concepts, but this does not mean they are mutually exclusive. Many animals remain inconspicuous until threatened, then suddenly employ warning signals, such as startling eyespots, bright colors on their undersides or loud vocalizations. In this way, they enjoy the best of both strategies. These strategies may also be employed differentially throughout stages of development. For instance, Large White butterflies are aposematic in color as larvae, but then display Müllerian mimicry once they emerge from the final stages of development as butterflies.
Many different prey of the same predator may employ separate warning colors, but this makes no sense for any party. Surely if they could all get together and agree on a common warning signal, the predator would have fewer detrimental experiences, and the prey would lose fewer individuals educating it. But no such conference needs to take place, as a prey species that just so happens to look a little like a harmful species will be safer than its conspecifics, leading to a tendency toward a single warning language. This can lead to the evolution of both Batesian and Müllerian mimicry, depending on whether the prey is harmful, as well, or just a free-rider. Multiple species can join this protective cooperative, expanding the mimicry ring.
Müller thus provided an explanation for 'Bates' paradox'; the mimicry was not a case of exploitation by one species, but rather a mutualistic arrangement.
A plausible explanation of the occurrence of Müllerian mimicry is known as the "two step hypothesis." This concept states that first, a large mutational leap occurs naturally and spontaneously to establish an approximate resemblance of one species to another species (the “model”). For this occurrence to be considered a part of a Müllerian mimicry model, the mimic and model species must both separately cause an unfavorable reaction in their shared predator. Once this leap has occurred, fine tuning then proceeds somewhat gradually in the mimic species so as to approach greater similarity to the model, albeit very slowly. This aposematic model (distinctive patterning linked to undesirable trait) must already exist in the model species. Both species will share in the benefits of a superior defense mechanism by way of patterning changes; generalized distrust of this pattern develops quickly in predators and has a large basis of reinforcement. The extent of a species' likelihood of being mimicked is considerably higher when there is only one key feature involved in a predator's generalization of undesirability. Once this trait (feature) spontaneously mutates in an individual it must also be able to be passed on to subsequent generations. After a significantly large number of generations have passed, it should eventually become fixated. In contrast, “non-feature traits,” (traits that do not play a role in predator recognition and distinction) do not affect the predator's selection of the mimic. Because individuals with mutated non-feature traits do not share a significant resemblance to the model in a way that is distinctive to the predator, the mimic containing this type of mutation is more likely to be eaten than mutants containing feature-trait resembling mutations. Therefore, the genetic mutation for the non-feature trait will likely not be passed on even if it confers an evolutionary advantage (unrelated to predator selection). The greatest selective pressure in the system is the propensity of the predators to avoid certain recognized patterns and when the mimic species eventually presents the trait in an individual, this trait is significantly more likely to be conserved and passed on to the individual's progeny. The strength of the influence of predator selection will remain strong and is much more likely to continue to permit the passing along of a feature-trait; although it is in no way guaranteed, it is highly plausible that eventually the whole species comes to share this trait which defines them as mimics.
Notes and references
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- For example, aggressive mimicry is not mentioned in Poulton, E. B., Ford, E. B. 1968. Mimcry. In Encyclopædia Britannica, vol. 15, pp. 487-89. Chicago: William Benton, bicentennial ed.
- Ihalainen, E.; Lindstrèom, L.; Mappes, J.; Puolakkainen, S. (January 1, 2008). "Butterfly effects in mimicry? Combining signal and taste can twist the relationship of Mèullerian co-mimics". Behavioral Ecology and Sociobiology 62 (8): 1267–1276. doi:10.1007/s00265-008-0555-y.
- Ihalainen, E.; Lindstreom, L.; Mappes, J.; Puolakkaien, S. (2008). "Butterfly effects in mimicry? Combining signal and taste can twist the relationship of Meullerian co-mimics.". Behavioral Ecology and Sociobiology 62 (8): 1267–1276. doi:10.1007/s00265-008-0555-y.
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Viceroys are as unpalatable as monarchs, and significantly more unpalatable than queens from representative Florida populations.
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- Feltwell, John (1982). Large White Butterfly: The Biology, Biochemistry, and Physiology of Pieris Brassicae (Linnaeus). The Hague: W. Junk. ISBN 90-6193-128-2.
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- Wickler, W. (1968) Mimicry in Plants and Animals (Translated from the German) McGraw-Hill, New York. ISBN 0-07-070100-8 Especially chapters 7 and 8.
- 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 0-19-852860-4 Chapter 9 and 11 provide an overview of current understanding