Batesian mimicry

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Plate from Bates 1861, illustrating Batesian mimicry between Dismorphia species (top row and third row) and various Ithomiini (Nymphalidae) (second and bottom rows). A non-Batesian species, Pseudopieris nehemia, is in the centre. Batesplate ArM.jpg
Plate from Bates 1861, illustrating Batesian mimicry between Dismorphia species (top row and third row) and various Ithomiini (Nymphalidae) (second and bottom rows). A non-Batesian species, Pseudopieris nehemia , is in the centre.

Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, who worked on butterflies in the rainforests of Brazil.

Contents

Batesian mimicry is the most commonly known and widely studied of mimicry complexes, such that the word mimicry is often treated as synonymous with Batesian mimicry. There are many other forms however, some very similar in principle, others far separated. It is often contrasted with Müllerian mimicry, a form of mutually beneficial convergence between two or more harmful species. However, because the mimic may have a degree of protection itself, the distinction is not absolute. It can also be contrasted with functionally different forms of mimicry. Perhaps the sharpest contrast here is with aggressive mimicry where a predator or parasite mimics a harmless species, avoiding detection and improving its foraging success.

The imitating species is called the mimic, while the imitated species (protected by its toxicity, foul taste or other defenses) is known as the model. The predatory species mediating indirect interactions between the mimic and the model is variously known as the [signal] receiver, dupe or operator. By parasitising the honest warning signal of the model, the Batesian mimic gains an advantage, without having to go to the expense of arming itself. The model, on the other hand, is disadvantaged, along with the dupe. If impostors appear in high numbers, positive experiences with the mimic may result in the model being treated as harmless. At higher frequency there is also a stronger selective advantage for the predator to distinguish mimic from model. For this reason, mimics are usually less numerous than models, an instance of frequency-dependent selection. Some mimetic populations have evolved multiple forms (polymorphism), enabling them to mimic several different models and thereby to gain greater protection. Batesian mimicry is not always perfect. A variety of explanations have been proposed for this, including limitations in predators' cognition.

While visual signals have attracted most study, Batesian mimicry can employ deception of any of the senses; some moths mimic the ultrasound warning signals sent by unpalatable moths to bat predators, constituting auditory Batesian mimicry, while some weakly electric fish appear to mimic the electrolocation signals of strongly electric fish, probably constituting electrical mimicry.

Historical background

Henry Walter Bates described the form of mimicry that bears his name in 1861. Henry Walter Bates Maull & Fox BNF Gallica (cropped).jpg
Henry Walter Bates described the form of mimicry that bears his name in 1861.

Henry Walter Bates (1825–1892) was an English explorer-naturalist who surveyed the Amazon rainforest with Alfred Russel Wallace in 1848. While Wallace returned in 1852, Bates remained for over a decade. Bates's field research included collecting almost a hundred species of butterflies from the families Ithomiinae and Heliconiinae, as well as thousands of other insects specimens. In sorting these butterflies into similar groups based on appearance, inconsistencies began to arise. Some appeared superficially similar to others, so much so that even Bates could not tell some species apart based only on wing appearance. However, closer examination of less obvious morphological characters seemed to show that they were not even closely related. Shortly after his return to England, he read a paper on his theory of mimicry at a meeting of the Linnean Society of London on 21 November 1861, which was then published in 1862 as 'Contributions to an Insect Fauna of the Amazon Valley' in the society's Transactions . [1] He elaborated on his experiences further in The Naturalist on the River Amazons . [2]

Bates put forward the hypothesis that the close resemblance between unrelated species was an antipredator adaptation. He noted that some species showed very striking coloration and flew in a leisurely manner, almost as if taunting predators to eat them. He reasoned that these butterflies were unpalatable to birds and other insectivores, and were thus avoided by them. He extended that logic to forms that closely resembled such protected species and mimicked their warning coloration but not their toxicity. [1] [2]

This naturalistic explanation fitted well with the recent account of evolution by Wallace and Charles Darwin, as outlined in his famous 1859 book The Origin of Species . Because the Darwinian explanation required no supernatural forces, it met with considerable criticism from anti-evolutionists, both in academic circles and in the broader social realm. [3]

Aposematism

The yellow-banded poison dart frog (Dendrobates leucomelas) has conspicuous aposematic coloration. Yellow-banded.poison.dart.frog.arp.jpg
The yellow-banded poison dart frog ( Dendrobates leucomelas ) has conspicuous aposematic coloration.

Most living things have predators and therefore are in a constant evolutionary arms race to develop antipredator adaptations, while the predator adapts to become more efficient at defeating the prey's adaptations. Some organisms have evolved to make detection less likely, for example by nocturnality and camouflage. Others have developed chemical defences such as the deadly toxins of certain snakes and wasps, or the noxious scent of the skunk. Such prey often send clear and honest warning signals to their attackers with conspicuous aposematic (warning) patterns. The brightness of such warning signs is correlated with the level of toxicity of the organism. [4]

In Batesian mimicry, the mimic effectively copies the coloration of an aposematic animal, known as the model, to deceive predators into behaving as if it were distasteful. [lower-alpha 1] The success of this dishonest display depends on the level of toxicity of the model and the abundance of the model in the geographical area. The more toxic the model is, the more likely it is that the predator will avoid the mimic. [6] The abundance of the model species is also important for the success of the mimic because of frequency-dependent selection. When the model is abundant, mimics with imperfect model patterns or slightly different coloration from the model are still avoided by predators. This is because the predator has a strong incentive to avoid potentially lethal organisms, given the likelihood of encountering one. [7] However, in areas where the model is scarce or locally extinct, mimics are driven to accurate aposematic coloration. This is because predators attack imperfect mimics more readily where there is little chance that they are the model species. [8] Frequency-dependent selection may also have driven Batesian mimics to become polymorphic in rare cases where a single genetic switch controls appearance, as in the swallowtail butterflies (the Papilionidae) such as the pipevine swallowtail, [9] and in the New Zealand stonefly Zelandoperla fenestrata . [10]

Batesian vs Mullerian mimicry: the former is deceptive, the latter honest. Batesian vs Mullerian mimicry.svg
Batesian vs Müllerian mimicry: the former is deceptive, the latter honest.

Classification and comparisons

Papilio polytes-Thekkady-2016-12-03-001.jpg
Common Rose (Pachliopta aristolochiae) W IMG 9133.jpg
A well-known mimic, Papilio polytes (top) resembles the unpalatable Pachliopta aristolochiae (bottom).

Batesian mimicry is a case of protective or defensive mimicry, where the mimic does best by avoiding confrontations with the signal receiver. It is a disjunct system, which means that all three parties are from different species. [11] An example would be the robber fly Mallophora bomboides , which is a Batesian mimic of its bumblebee model and prey, B. americanorum (now more commonly known as Bombus pensylvanicus ), which is noxious to predators due to its sting. [12]

Batesian mimicry stands in contrast to other forms such as aggressive mimicry, where the mimic profits from interactions with the signal receiver. One such case of this is in fireflies, where females of one species mimic the mating signals of another species, deceiving males to come close enough for them to eat. Mimicry sometimes does not involve a predator at all though. Such is the case in dispersal mimicry, where the mimic once again benefits from the encounter. For instance, some fungi have their spores dispersed by insects by smelling like carrion. In protective mimicry, the meeting between mimic and dupe is not such a fortuitous occasion for the mimic, and the signals it mimics tend to lower the probability of such an encounter. [3]

A case somewhat similar to Batesian mimicry is that of mimetic weeds, which imitate agricultural crops. In weed or Vavilovian mimicry, the weed survives by having seeds which winnowing machinery identifies as belonging to the crop. Vavilovian mimicry is not Batesian, because humans and crops are not enemies. [3] By contrast, a leaf-mimicking plant, the chameleon vine, employs Batesian mimicry by adapting its leaf shape and colour to match that of its host to deter herbivores from eating its edible leaves. [13]

Another analogous case within a single species has been termed Browerian mimicry [3] (after Lincoln P. Brower and Jane Van Zandt Brower [14] [15] ). This is a case of automimicry; [11] the model is the same species as its mimic. Equivalent to Batesian mimicry within a single species, it occurs when there is a palatability spectrum within a population of harmful prey. For example, monarch (Danaus plexippus) caterpillars feed on milkweed species of varying toxicity. Some feed on more toxic plants and store these toxins within themselves. The more palatable caterpillars thus profit from the more toxic members of the same species. [14] [16]

Another important form of protective mimicry is Müllerian mimicry, discovered by and named after the naturalist Fritz Müller. [17] [18] In Müllerian mimicry, both model and mimic are aposematic, so mimicry may be mutual, does not necessarily [lower-alpha 2] constitute a bluff or deception and as in the wasps and bees may involve many species in a mimicry ring. [19] [20]

Imperfect Batesian mimicry

The hoverfly Spilomyia longicornis is an imperfect Batesian mimic of wasps, lacking their long antennae and wasp waist. Syrphid - Spilomyia longicornis, Meadowwood Farm SRMA, Mason Neck, Virginia.jpg
The hoverfly Spilomyia longicornis is an imperfect Batesian mimic of wasps, lacking their long antennae and wasp waist.

In imperfect Batesian mimicry, the mimics do not exactly resemble their models. An example of this is the fly Spilomyia longicornis , which mimics vespid wasps. However, it is not a perfect mimic. Wasps have long black antennae and this fly does not. Instead, they wave their front legs above their heads to look like the antennae on the wasps. [21] Many reasons have been suggested for imperfect mimicry. Imperfect mimics may simply be evolving towards perfection. [22] They may gain advantage from resembling multiple models at once. [23] Humans may evaluate mimics differently from actual predators. [24] Mimics may confuse predators by resembling both model and nonmimic at the same time (satyric mimicry). [25] Kin selection may enforce poor mimicry. [26] The selective advantage of better mimicry may not outweigh the advantages of other strategies like thermoregulation or camouflage. [27]

Only certain traits may be required to deceive predators; for example, tests on the sympatry/allopatry border (where the two are in the same area, and where they are not) of the mimic Lampropeltis elapsoides and the model Micrurus fulvius showed that color proportions in these snakes were important in deceiving predators but that the order of the colored rings was not. [28]

Plants mimicking ants

The elongated spots on the reproductive organs of Passiflora incarnata may mimic ants to deter herbivores. Passiflora incarnata (detail).jpg
The elongated spots on the reproductive organs of Passiflora incarnata may mimic ants to deter herbivores.

Batesian mimicry of ants appears to have evolved in certain plants, as a visual anti-herbivory strategy, analogous to a herbivorous insect's mimicking a well-defended insect to deter predators. [30] Passiflora flowers of at least 22 species, such as P. incarnata , have dark dots and stripes on their flowers thought to serve this purpose. [29]

Acoustic mimicry

Tiger moths like Cycnia tenera are aposematic by sound, emitting ultrasonic warning signals. They are mimicked by pyralid moths, which are not foul-tasting but emit similar sounds. Cycnia teneraPCCP20030807-2447B.jpg
Tiger moths like Cycnia tenera are aposematic by sound, emitting ultrasonic warning signals. They are mimicked by pyralid moths, which are not foul-tasting but emit similar sounds.

Predators may identify their prey by sound as well as sight; mimics have accordingly evolved to deceive the hearing of their predators. Bats are nocturnal predators that rely on echolocation to detect their prey. [32] Some potential prey are unpalatable to bats, and produce an ultrasonic aposematic signal, the auditory equivalent of warning coloration. In response to echolocating red bats and big brown bats, tiger moths such as Cycnia tenera produce warning sounds. Bats learn to avoid the harmful moths, but similarly avoid other species such as some pyralid moths that produce such warning sounds as well. Acoustic mimicry complexes, both Batesian and Müllerian, may be widespread in the auditory world. [31]

Electrical mimicry

The electric eel, Electrophorus, is capable of delivering a powerful electric shock that can stun or kill its prey. Bluntnose knifefishes, Brachyhypopomus , create an electric discharge pattern similar to the low voltage electrolocation discharge of the electric eel. This is thought to be Batesian mimicry of the powerfully protected electric eel. [33]

See also

Notes

  1. ↑ This is often described as parasitizing the honest signals. [5]
  2. ↑ Müllerian mimicry in its simplest form is not a bluff at all, but since toxicity is relative, there is a spectrum of mimicry from Batesian to Müllerian. [19]

Related Research Articles

<span class="mw-page-title-main">Mimicry</span> Evolutionary strategy

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.

Frequency-dependent selection is an evolutionary process by which the fitness of a phenotype or genotype depends on the phenotype or genotype composition of a given population.

<span class="mw-page-title-main">Anti-predator adaptation</span> Defensive feature of prey for selective advantage

Anti-predator adaptations are mechanisms developed through evolution that assist prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, namely by avoiding detection, warding off attack, fighting back, or escaping when caught.

<span class="mw-page-title-main">Viceroy (butterfly)</span> Species of butterfly

The viceroy is a North American butterfly. It was long thought to be a Batesian mimic of the monarch butterfly, but since the viceroy is also distasteful to predators, it is now considered a Müllerian mimic instead.

<span class="mw-page-title-main">Müllerian mimicry</span> Mutually beneficial mimicry of strongly defended species

Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Müllerian mimics is that predators only need one unpleasant encounter with one member of a set of Müllerian mimics, and thereafter avoid all similar coloration, whether or not it belongs to the same species as the initial encounter. It is named after the German naturalist Fritz Müller, who first proposed the concept in 1878, supporting his theory with the first mathematical model of frequency-dependent selection, one of the first such models anywhere in biology.

<span class="mw-page-title-main">Aposematism</span> Honest signalling of an animals powerful defences

Aposematism is the advertising by an animal, whether terrestrial or marine, to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or smell, sharp spines, or aggressive nature. These advertising signals may take the form of conspicuous coloration, sounds, odours, or other perceivable characteristics. Aposematic signals are beneficial for both predator and prey, since both avoid potential harm.

<span class="mw-page-title-main">Ant mimicry</span> Animals that resemble ants

Ant mimicry or myrmecomorphy is mimicry of ants by other organisms; it has evolved over 70 times. Ants are abundant all over the world, and potential predators that rely on vision to identify their prey, such as birds and wasps, normally avoid them, because they are either unpalatable or aggressive. Some arthropods mimic ants to escape predation, while some predators of ants, especially spiders, mimic them anatomically and behaviourally in aggressive mimicry. Ant mimicry has existed almost as long as ants themselves; the earliest ant mimics in the fossil record appear in the mid-Cretaceous alongside the earliest ants.

<span class="mw-page-title-main">Automimicry</span> Mimicry of part of own body, e.g. the head

In zoology, automimicry, Browerian mimicry, or intraspecific mimicry, is a form of mimicry in which the same species of animal is imitated. There are two different forms.

<i>Heliconius</i> Genus of brush-footed butterflies

Heliconius comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America as far north as the southern United States. The larvae of these butterflies eat passion flower vines (Passifloraceae). Adults exhibit bright wing color patterns which signal their distastefulness to potential predators.

<span class="mw-page-title-main">Aggressive mimicry</span> Deceptive mimicry of a harmless species by a predator

Aggressive mimicry is a form of mimicry in which predators, parasites, or parasitoids share similar signals, using a harmless model, allowing them to avoid being correctly identified by their prey or host. Zoologists have repeatedly compared this strategy to a wolf in sheep's clothing. In its broadest sense, aggressive mimicry could include various types of exploitation, as when an orchid exploits a male insect by mimicking a sexually receptive female, but will here be restricted to forms of exploitation involving feeding. For example, indigenous Australians who dress up as and imitate kangaroos when hunting would not be considered aggressive mimics, nor would a human angler, though they are undoubtedly practising self-decoration camouflage. Treated separately is molecular mimicry, which shares some similarity; for instance a virus may mimic the molecular properties of its host, allowing it access to its cells. An alternative term, Peckhamian mimicry, has been suggested, but it is seldom used.

<span class="mw-page-title-main">Animal coloration</span> General appearance of an animal

Animal colouration is the general appearance of an animal resulting from the reflection or emission of light from its surfaces. Some animals are brightly coloured, while others are hard to see. In some species, such as the peafowl, the male has strong patterns, conspicuous colours and is iridescent, while the female is far less visible.

<span class="mw-page-title-main">Emsleyan mimicry</span> Mimicry of a less deadly species

Emsleyan mimicry, also called Mertensian mimicry, describes an unusual type of mimicry where a deadly prey mimics a less dangerous species.

<span class="mw-page-title-main">Chemical mimicry</span> Biological mimicry using chemicals

Chemical mimicry is a type of biological mimicry involving the use of chemicals to dupe an operator.

Many types of polymorphism can be seen in the insect order Lepidoptera. Polymorphism is the appearance of forms or "morphs" differing in color and number of attributes within a single species. In Lepidoptera, polymorphism can be seen not only between individuals in a population but also between the sexes as sexual dimorphism, between geographically separated populations in geographical polymorphism and also between generations flying at different seasons of the year. It also includes the phenomenon of mimicry when mimetic morphs fly alongside non-mimetic morphs in a population of a particular species. Polymorphism occurs both at a specific level with heritable variation in the overall morphological design of individuals as well as in certain specific morphological or physiological traits within a species.

<i>Adaptive Coloration in Animals</i> 1940 textbook on camouflage, mimicry and aposematism by Hugh Cott

Adaptive Coloration in Animals is a 500-page textbook about camouflage, warning coloration and mimicry by the Cambridge zoologist Hugh Cott, first published during the Second World War in 1940; the book sold widely and made him famous.

<span class="mw-page-title-main">Mimicry in plants</span> Evolutionary mechanism

In evolutionary biology, mimicry in plants is where a plant evolves to resemble another organism physically or chemically. Mimicry in plants has been studied far less than mimicry in animals. It may provide protection against herbivory, or may deceptively encourage mutualists, like pollinators, to provide a service without offering a reward in return.

The novel world method is a technique used in animal behaviour experiments that address questions on the evolution of warning signals that chemically defended prey use to deter predators, and also on warning signal mimicry.

<i>Animal Coloration</i> (book) 1892 book by Frank Evers Beddard

Animal Coloration, or in full Animal Coloration: An Account of the Principal Facts and Theories Relating to the Colours and Markings of Animals, is a book by the English zoologist Frank Evers Beddard, published by Swan Sonnenschein in 1892. It formed part of the ongoing debate amongst zoologists about the relevance of Charles Darwin's theory of natural selection to the observed appearance, structure, and behaviour of animals, and vice versa.

<span class="mw-page-title-main">Coloration evidence for natural selection</span> Early evidence for Darwinism from animal coloration

Animal coloration provided important early evidence for evolution by natural selection, at a time when little direct evidence was available. Three major functions of coloration were discovered in the second half of the 19th century, and subsequently used as evidence of selection: camouflage ; mimicry, both Batesian and Müllerian; and aposematism.

In evolutionary biology, mimicry in vertebrates is mimicry by a vertebrate of some model, deceiving some other animal, the dupe. Mimicry differs from camouflage as it is meant to be seen, while animals use camouflage to remain hidden. Visual, olfactory, auditory, biochemical, and behavioral modalities of mimicry have been documented in vertebrates.

References

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  31. 1 2 Barber, J. R.; Conner, W. E. (2007). "Acoustic mimicry in a predator prey interaction". Proceedings of the National Academy of Sciences of the United States of America. 104 (22): 9331–9334. Bibcode:2007PNAS..104.9331B. doi: 10.1073/pnas.0703627104 . PMC   1890494 . PMID   17517637.
  32. ↑ Dawkins, Richard (1986). The Blind Watchmaker. W. W. Norton. ISBN   978-0-393-31570-7.
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