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.

Batesian mimicry is a form of

rainforests
of Brazil.

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

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.

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

social realm.[3]

Aposematism

Main article: Aposematism
aposematic
coloration.

Most living things have predators and therefore are in a constant

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.

Papilionidae) such as the pipevine swallowtail,[9] and in the New Zealand stonefly Zelandoperla fenestrata.[10]

Classification and comparisons

Further information:
Cleaner fish mimicry
Common Mormon (Papilio polytes)
Common rose (Pachliopta aristolochiae)
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

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 man and crop 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

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[b] 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.

In imperfect Batesian mimicry, the mimics do not exactly resemble their models. An example of this is the fly

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

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

Further information: Ant mimicry
The elongated spots on the reproductive organs of Passiflora incarnata may mimic ants to deter herbivores.[29]

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.[31]

Predators may identify their prey by sound as well as sight; mimics have accordingly evolved to deceive the

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

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]

References

  1. ^
    doi:10.1111/j.1095-8312.1981.tb01842.x
    .
  2. ^ a b Bates, Henry Walter (1863). The Naturalist on the River Amazons. John Murray.
  3. ^
    doi:10.1146/annurev.es.13.110182.001125
    .
  4. PMID 11934367
    .
  5. PMID 20539774
    .
  6. doi:10.1093/czoolo/60.1.16
    .
  7. PMID 19955153
    .
  8. PMID 24919704
    .
  9. PMID 21238394
    .
  10. PMID 37503654
    .
  11. ^
    doi:10.1111/j.1095-8312.1976.tb00240.x
    .
  12. S2CID 83531239
    .
  13. PMID 24768053
    .
  14. ^ a b 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.
  15. PMID 5231352
    .
  16. ISBN 978-1-4899-3368-3
    . [Consider the case where one monarch caterpillar is feeding on cardenolide-containing milkweed, the other not], with one being completely potent with regard to cardiac glycoside toxicity, the second not. The first will fit all of the characteristics for warning coloration, the second not. In fact, the second butterfly is a harmless Batesian mimic of the first, even though both belong to the same species. L. Brower, J. Brower, and Corvino (1967) have termed this phenomenon automimicry, though others have suggested that Browerian mimicry would be a better term (Pasteur, 1972; Bees, 1977; Rothschild, 1979). Note that all of the antagonisms raised by Batesian mimicry will arise, but now the model and the mimic are conspecific.
  17. ^ Müller, Fritz (1878). "Ueber die Vortheile der Mimicry bei Schmetterlingen". Zoologischer Anzeiger. 1: 54–55.
  18. ^ 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.
  19. ^
    S2CID 45185502
    .
  20. PMID 19487663
    .
  21. ^ Curran, C. H. (1951). Synopsis of the North American species of Spilomyia (Syrphidae, Diptera). American Museum Novitates. http://digitallibrary.amnh.org/bitstream/handle/2246/2364//v2/dspace/ingest/pdfSource/nov/N1492.pdf?sequence=1&isAllowed=y.
  22. PMID 11886630
    .
  23. doi:10.1111/j.1095-8312.2000.tb01234.x
    .
  24. S2CID 84467142
    .
  25. S2CID 84458742
    .
  26. S2CID 4424680
    .
  27. PMID 17567563
    .
  28. S2CID 35411437
    .
  29. ^ a b Lev-Yadun, Simcha (2009). "Ant mimicry by Passiflora Flowers?". Israel Journal of Entomology. 39: 159–163.
  30. ^ Weins, D. (1978). "Mimicry in plants". Evolutionary Biology. 11: 365–403.
  31. ^
    PMID 17517637
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  32. ISBN 978-0-393-31570-7
    .
  33. S2CID 204994529
    .

Further reading

External links

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