Batesian mimicry

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. He elaborated on his experiences further in The Naturalist on the River Amazons. 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. == Aposematism ==

'') 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. 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. 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. 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. 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. 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, and in the New Zealand stonefly Zelandoperla fenestrata. : the former is deceptive, the latter honest. == Classification and comparisons ==

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. 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. 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. Another analogous case within a single species has been termed Browerian mimicry). This is a case of automimicry; Another important form of protective mimicry is Müllerian mimicry, discovered by and named after the naturalist Fritz Müller. In Müllerian mimicry, both model and mimic are aposematic, so mimicry may be mutual, does not necessarily constitute a bluff or deception and as in the wasps and bees may involve many species in a mimicry ring. == Imperfect Batesian mimicry ==

'' 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. Many reasons have been suggested for imperfect mimicry. Imperfect mimics may simply be evolving towards perfection. They may gain advantage from resembling multiple models at once. Humans may not evaluate mimics in the same way as actual predators. Mimics may confuse predators by resembling both model and nonmimic at the same time (satyric mimicry). Kin selection may enforce poor mimicry. The selective advantage of better mimicry may not outweigh the advantages of other strategies like thermoregulation or camouflage. 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. == Plants mimicking ants ==

may mimic ants to deter herbivores. Passiflora flowers of at least 22 species, such as P. incarnata'', have dark dots and stripes on their flowers thought to serve this purpose. == Acoustic mimicry ==

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. 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. == 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. == See also ==