Mimicry

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Many insects including hoverflies (C, D, E) and the wasp beetle (F) are Batesian mimics of stinging wasps (A, B), which are Mullerian mimics of each other. Wasp mimicry.jpg
Many insects including hoverflies (C, D, E) and the wasp beetle (F) are Batesian mimics of stinging wasps (A, B), which are Müllerian mimics of each other.

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.

Contents

Etymology

Use of the word mimicry dates to 1637. It derives from the Greek term mimetikos, "imitative", in turn from mimetos, the verbal adjective of mimeisthai, "to imitate". [1] "Mimicry" was first used in zoology by the English entomologists William Kirby and William Spence in 1823. [2] [3] Originally used to describe people, "mimetic" was used in zoology from 1851. [1]

History

Ancient

Aristotle wrote in his History of Animals that partridges use a deceptive distraction display to lure predators away from their flightless young: [2] [4]

When a man comes by chance upon a young brood [of partridges], and tries to catch them, the hen-bird rolls in front of the hunter, pretending to be lame: the man every moment thinks he is on the point of catching her, and so she draws him on and on, until every one of her brood has had time to escape; hereupon she returns to the nest and calls the young back.

— Aristotle, translated by D'Arcy Wentworth Thompson [4]

The behaviour is recognised as a form of mimicry by biologists. [2]

19th century

In 1879, Fritz Muller created the first mathematical model of mimicry to explain why distasteful species should evolve similar appearances. Fritz Muller 1891 (cropped).jpg
In 1879, Fritz Müller created the first mathematical model of mimicry to explain why distasteful species should evolve similar appearances.

In 1823, Kirby and Spence, in their book An Introduction to Entomology, used the term "mimicry" informally to depict the way that the structure and coloration of some insects resembled objects in their environments: [2] [3]

A jumping bug, very similar to the one figured by Schellenberg, also much resembles the lichens of the oak on which I took it. The spectre tribe ( Phasma ) go still further in this mimicry, representing a small branch with its spray. [3]

The English naturalist Henry Walter Bates worked for several years on butterflies in the Amazon rainforest. Returning home, he described multiple forms of mimicry in an 1862 paper at the Linnean Society in London, [6] and then in his 1863 book The Naturalist on the River Amazons . [2] [7] The term "Batesian mimicry" has since been used in his honour, its usage becoming restricted to the situation in which a harmless mimic gains protection from its predators by resembling a distasteful model. [2] Among the observations in Bates's 1862 paper is the statement:

I was never able to distinguish the Leptalides from the species they imitated, although they belong to a family totally different in structure and metamorphosis from the Heliconidae, without examining them closely after capture. [6]

The German naturalist Fritz Müller also spent many years studying butterflies in the Amazon rainforest. He first published a journal article on mimicry in German in 1878, [8] followed in 1879 by a paper to the Entomological Society of London (translated and presented by Ralph Meldola). [9] He described a situation where different species were each unpalatable to predators, and shared similar, genuine, warning signals. Bates found it hard to explain why this should be so, asking why they should need to mimic each other if both were harmful and could warn off predators on their own. Müller put forward the first mathematical model of mimicry for this phenomenon: if a common predator confuses the two species, individuals in both those species are more likely to survive, as fewer individuals of either species are killed by the predator. The term Müllerian mimicry, named in his honour, has since been used for this mutualistic form of mimicry. [5] [10]

Müller wrote that

The resemblance of the genera named [ Ituna and Thyridia ] is the more worthy of notice since it occurs between insects both belonging to the group of butterflies which are protected by distastefulness. The explanation which applies in ordinary cases of [Batesian] mimicry—and no other has, so far as I know, been offered—cannot obtain for this imitation among protected species. [9]

Overview

Evolved resemblance

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.

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. Often, mimicry functions to protect from predators. [11] Mimicry systems have three basic roles: a mimic, a model, and a dupe. When these correspond to three separate species, the system is called disjunct; when the roles are taken by just two species, the system is called bipolar. [2] [12] Mimicry evolves if a dupe (such as a predator) perceives a mimic (such as a palatable prey) as a model (the organism it resembles), and is deceived to change its behaviour to the mimic's selective advantage. [13] The resemblances can be via any sensory modality, including any combination of visual, acoustic, chemical, tactile, or electric. [13] [14] Mimicry may be to the advantage of both organisms that share a resemblance, in which case it is mutualistic; or it can be to the detriment of one, making it parasitic or competitive. The evolutionary convergence between groups is driven by the selective action of a dupe. [2] [15] Birds, for example, use sight to identify palatable insects, [16] whilst avoiding noxious ones. Over time, palatable insects may evolve to resemble noxious ones, making them mimics and the noxious ones models. Models do not have to be more abundant than mimics. [17] In the case of mutualism, each model is also a mimic; all such species can be called "co-mimics". [17] Many harmless species such as hoverflies are Batesian mimics of strongly defended species such as wasps, while many such well-defended species form Müllerian mimicry rings of co-mimics. [2] In the evolution of wasp-like appearance, it has been argued that insects evolve to masquerade wasps since predatory wasps do not attack each other, and that this mimetic resemblance has had the useful side-effect of deterring vertebrate predators. [18]

Mimicry can result in an evolutionary arms race if mimicry negatively affects the model, in which case the model can evolve a different appearance from the mimic. [17] p161 Mimics may have different models for different life cycle stages, or they may be polymorphic, with different individuals imitating different models, as occurs in Heliconius butterflies. Models tend to be relatively closely related to their mimics, [19] but mimicry can be of vastly different species, for example when spiders mimic ants. Most known mimics are insects, [14] though many other examples including vertebrates, plants, and fungi exist. [20] [21] [22]

Evolutionary explanations

It is widely accepted that mimicry evolves as a positive adaptation. The lepidopterist and novelist Vladimir Nabokov however argued that although natural selection might stabilize a "mimic" form, it would not be necessary to create it. [23] The most widely accepted model used to explain the evolution of mimicry in butterflies is the two-step hypothesis. The first step involves mutation in modifier genes that regulate a complex cluster of linked genes that cause large changes in morphology. The second step consists of selections on genes with smaller phenotypic effects, creating an increasingly close resemblance. This model is supported by empirical evidence that suggests that a few single point mutations cause large phenotypic effects, while numerous others produce smaller effects. Some regulatory elements collaborate to form a supergene for the development of butterfly color patterns. The model is supported by computational simulations of population genetics. [24] The Batesian mimicry in Papilio polytes is controlled by the doublesex gene. [25]

Some mimicry is imperfect. Natural selection drives mimicry only far enough to deceive predators. For example, when predators avoid a mimic that imperfectly resembles a coral snake, the mimic is sufficiently protected. [26] [27] [28]

Convergent evolution is an alternative explanation for why coral reef fish have come to resemble each other; [29] [30] the same applies to benthic marine invertebrates such as sponges and nudibranchs. [31]

Living and non-living models

In its broadest definition, mimicry can include non-living models. The specific terms masquerade and mimesis are sometimes used when the models are inanimate, and the mimicry's purpose is crypsis. [32] [14] [2] For example, animals such as flower mantises, planthoppers, comma and geometer moth caterpillars resemble twigs, bark, leaves, bird droppings or flowers. [14] [17] [33] [34] Many animals bear eyespots, which are hypothesized to resemble the eyes of larger animals. They may not resemble any specific organism's eyes, and whether or not animals respond to them as eyes is also unclear. [35] [36] [37] [38] The model is usually another species, except in automimicry, where members of the species mimic other members, or other parts of their own bodies, and in inter-sexual mimicry, where members of one sex mimic members of the other. [17]

Types

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 with respect to the mimic (e.g., avoiding harm). 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). The terminology used has been debated, as classifications have differed or overlapped; attempts to clarify definitions have led to the partial replacement of old terms with new ones. [2] [39] [40]

Some kinds of mimicry classified by Pasteur 1982 [2]
NameNo. of
spp.		FunctionDupe finds
Model		 Deception Description (mimic, model, dupe)
Aristotelian 2ProtectiveAgreeableDeceptiveBrooding bird mimics itself with broken wing, luring predator away from nest [41]
Automimicry 1 or 2ProtectiveAgreeableDeceptiveMultiple forms, e.g. one sex mimics the other, tail mimics head, etc.
Bakerian 2ReproductiveForbiddingDeceptiveFemale flower resembles male flower, cheating pollinator
Batesian 3ProtectiveForbiddingDeceptivePalatable mimic resembles distasteful model, deceives dupe
Browerian 2ProtectiveForbiddingDeceptivePalatable butterfly resembles toxic member of same species
Emsleyan 3ProtectiveForbiddingDeceptiveDeadly snake resembles less deadly species, predators get chance to learn to avoid them
Gilbertian 2ProtectiveForbiddingDeceptive Host/prey mimics and so repels parasite/predator
Kirbyan 2 Aggressive AgreeableDeceptive Brood parasite adult or egg mimics host which raises the young as its own
Müllerian 3 or moreProtectiveForbidding Honest Distasteful co-mimics resemble each other, aposematically warning off predators [42]
Pouyannian 2ReproductiveAgreeableDeceptivePlant mimic resembles female bee, deceives male, gets itself pollinated
Vavilovian 3ReproductiveAgreeableDeceptiveMimic resembles crop, deceives farmer
Wasmannian 2 Commensalist AgreeableDeceptiveMimic resembles and deceives ant, lives in ant nest
Wicklerian 2 Aggressive AgreeableDeceptive Predator or parasite resembles and attacks prey or host; parasite may get itself swallowed
Camouflage 2ProtectiveUninterestingDeceptiveMimic resembles background (plant parts, or inanimate)

Defensive

Mimicry is defensive or protective when organisms are able to avoid harmful encounters by deceiving enemies into treating them as something else.

Batesian

Macroxiphus, a harmless bush cricket, mimics a well-defended ant. Macroxiphus sp cricket.jpg
Macroxiphus , a harmless bush cricket, mimics a well-defended ant.

In Batesian mimicry, the mimic resembles the model, but does not have the attribute that makes it unprofitable to predators (e.g., unpalatability, or the ability to sting). In other words, a Batesian mimic is a sheep in wolf's clothing. Mimics are less likely to be found out (for example by predators) when in low proportion to their model. Such negative frequency-dependent selection applies in most forms of mimicry. Specifically, Batesian mimicry can only be maintained if the harm caused to the predator by eating a model outweighs the benefit of eating a mimic. The nature of learning is weighted in favor of the mimics, for a predator that has a bad first experience with a model tends to avoid anything that looks like it for a long time, and does not re-sample soon to see whether the initial experience was a false negative. However, if mimics become more abundant than models, then the probability of a young predator having a first experience with a mimic increases. Batesian systems are therefore most likely to be stable where the model is more abundant than the mimic. [43]

There are many Batesian mimics among butterflies and moths. Consul fabius and Eresia eunice imitate unpalatable Heliconius butterflies such as H. ismenius . [44] Limenitis arthemis imitate the poisonous pipevine swallowtail ( Battus philenor ). Several palatable moths produce ultrasonic click calls to mimic unpalatable tiger moths. [45] [46] [47] [48] Octopuses of the genus Thaumoctopus (the mimic octopus) are able to intentionally alter their body shape and coloration to resemble dangerous sea snakes or lionfish. [49] In the Amazon, the helmeted woodpecker ( Dryocopus galeatus ), a rare species which lives in the Atlantic Forest of Brazil, Paraguay, and Argentina, has a similar red crest, black back, and barred underside to two larger woodpeckers: Dryocopus lineatus and Campephilus robustus . This mimicry reduces attacks on D. galeatus. [50] Batesian mimicry occurs in the plant kingdom, where the chameleon vine adapts its leaf shape and colour to match that of the plant it is climbing. [51]

Müllerian

Comparison of Batesian and Mullerian mimicry, illustrated with a hoverfly, a wasp and a bee Batesian vs Mullerian Mimicry.svg
Comparison of Batesian and Müllerian mimicry, illustrated with a hoverfly, a wasp and a bee

In Müllerian mimicry, two or more species have similar warning or aposematic signals and both share genuine anti-predation attributes (e.g. being unpalatable), as first described in Heliconius butterflies. [52] 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. The signal receiver also benefits by this system, despite being deceived about species identity, as it is able to generalize the pattern to potentially harmful encounters. The distinction between mimic and model that is clear in Batesian mimicry is also blurred. 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 makes more sense to speak of each as a co-mimic than of distinct 'mimic' and 'model' species, as their warning signals tend to converge. [42] Also, the mimetic species may exist on a continuum from harmless to highly noxious, so Batesian mimicry grades smoothly into Müllerian convergence. [53] [54]

Emsleyan/Mertensian

Emsleyan or Mertensian mimicry describes the unusual case where a deadly prey mimics a less dangerous species. [2] It was first proposed by M. G. Emsley in 1966 as a possible explanation for how a predator can learn to avoid a very dangerous aposematic animal, such as a coral snake, when the predator is very likely to die, making learning unlikely. [55] The theory was developed by the German biologist Wolfgang Wickler who named it after the German herpetologist Robert Mertens. [14] [56] [57] [58] The scenario is unlike Müllerian mimicry, where the most harmful species is the model. But if a predator dies on its first encounter with a deadly snake, it has no occasion to learn to recognize the snake's warning signals. There would then be no advantage for an extremely deadly snake in being aposematic: any predator that attacked it would be killed before it could learn to avoid the deadly prey, so the snake would be better off being camouflaged to avoid attacks. But if the predator first learnt to avoid a less deadly warning-coloured snake, the deadly species could profit by mimicking the less dangerous snake. [57] [58] Some harmless milk snake (Lampropeltis triangulum), the moderately toxic false coral snakes ( Erythrolamprus aesculapii ), and the deadly coral snakes ( Micrurus ) all have a red background color with black and white/yellow rings. In this system, both the milk snakes and the deadly coral snakes are mimics, while the false coral snakes are the model. [55]

Wasmannian

In Wasmannian mimicry, the mimic resembles a model that it lives along with in a nest or colony. Most of the models here are eusocial insects, principally ants. [59] [60]

Gilbertian

Some Passiflora flower species use Gilbertian mimicry, defending against being eaten by larvae of Heliconius butterflies with leaf stipules (not shown) that resemble the butterfly's eggs. Heliconiinae - Heliconius numata.JPG
Some Passiflora flower species use Gilbertian mimicry, defending against being eaten by larvae of Heliconius butterflies with leaf stipules (not shown) that resemble the butterfly's eggs.

Gilbertian mimicry is bipolar, involving only two species. The potential host (or prey) drives away its parasite (or predator) by mimicking it, the reverse of host-parasite aggressive mimicry. It was coined by Pasteur as a phrase for such rare mimicry systems, [2] and is named after the American ecologist Lawrence E. Gilbert  [ nl ] who described it in 1975. [61] The classical instance of Gilbertian mimicry is in the plant genus Passiflora , which is grazed by the micropredator larvae of some Heliconius butterflies. The host plants have evolved stipules that mimic mature Heliconius eggs near the point of hatching. The butterflies avoid laying eggs near existing ones, reducing intraspecific competition between caterpillars, which are also cannibalistic, so those that lay on vacant leaves provide their offspring with a greater chance of survival. The stipules thus appear to have evolved as Gilbertian mimics of butterfly eggs, under selection pressure from these caterpillars. [2] [19]

Browerian

Browerian mimicry, named after Lincoln P. Brower and Jane Van Zandt Brower who first described it in 1967, [62] [63] is a postulated 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. Examples include the monarch and the queen from the subfamily Danainae, which feed on milkweed species of varying toxicity. These species store toxins from its host plant, which are maintained even in the adult. As levels of toxin vary depending on diet, some individuals are more toxic than the rest, which profit from the toxicity of those individuals, just as hoverflies benefit from mimicking well-defended wasps. [2]

Misdirection by automimicry

Eyespots of foureye butterflyfish (Chaetodon capistratus) mimic its own eyes, deflecting attacks from the vulnerable head. Chaetodon capistratus2.jpg
Eyespots of foureye butterflyfish (Chaetodon capistratus) mimic its own eyes, deflecting attacks from the vulnerable head.

One form of automimicry is where one part of an organism's body resembles another part. For example, the tails of some snakes resemble their heads; they move backwards when threatened and present the predator with the tail, improving their chances of escape without fatal harm. Some fishes have eyespots near their tails, and when mildly alarmed swim slowly backwards, presenting the tail as a head. Some insects such as some lycaenid butterflies have tail patterns and appendages of various degrees of sophistication that promote attacks at the rear rather than at the head. Several species of pygmy owl bear "false eyes" on the back of the head, misleading predators into reacting as though they were the subject of an aggressive stare. [64] Many insects have filamentous "tails" at the ends of their wings and patterns of markings on the wings themselves. These combine to create a "false head". This misdirects predators such as birds and jumping spiders. Spectacular examples occur in the hairstreak butterflies; when perching on a twig or flower, they commonly do so upside down and shift their rear wings repeatedly, causing antenna-like movements of the "tails" on their wings. Studies of rear-wing damage support the hypothesis that this strategy is effective in deflecting attacks from the insect's head. [65] [66]

Aggressive

Predators

Aggressive mimicry is found in predators or parasites that share some of the characteristics of a harmless species, allowing them to avoid detection by their prey or host; the strategy resembles a wolf in sheep's clothing, though no conscious deceptive intent is involved. The mimic may resemble the prey or host itself, or another organism that does not threaten the prey or host. [67]

Several spiders use aggressive mimicry to lure prey. [68] Species such as the silver argiope ( 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 guides. Spiders change their web day to day, which can be explained by the ability of bees to remember web patterns. [69]

Another case is where males are lured towards what seems to be a sexually receptive female. The model in this situation is the same species as the dupe. Female fireflies of the genus Photuris emit light signals that mimic the mating signals of females of the genus Photinus . [70] Male fireflies from several different genera are attracted to these "femmes fatales", and are captured and eaten. Each female has a repertoire of signals matching the delay and duration of the flashes of the female of the corresponding species. [71]

Some carnivorous plants may be able to increase their rate of capturing insect prey through mimicry. [72]

Two bluestreak cleaner wrasse cleaning a potato grouper, Epinephelus tukula Epinephelus tukula is cleaned by two Labroides dimidiatus.jpg
Two bluestreak cleaner wrasse cleaning a potato grouper, Epinephelus tukula

A different aggressive strategy is to mimic a mutualistic symbiont of the prey. Cleaner fish eat parasites and dead skin from client fish. Some allow the cleaner to venture inside their body to hunt these parasites. However, the sabre-toothed blenny or false cleanerfish (Aspidontus taeniatus) mimics the bluestreak cleaner wrasse (Labroides dimidiatus), which is recognized by other fishes as a cleaner. The false cleanerfish resembles the cleaner, and mimics the cleaner's "dance". Once it is allowed close to the client, it attacks, biting off a piece of its fin before fleeing. Fish wounded in this fashion soon learn to distinguish mimic from model, but because the similarity is close they also become much more cautious of the model. [73]

A mechanism that does not involve any luring is seen in the zone-tailed hawk, which resembles the turkey vulture. It flies amongst the vultures, effectively camouflaged as a vulture which poses no threat to the hawk's prey. It hunts by suddenly breaking from the formation and ambushing its prey. [74]

Parasites

Parasites can be aggressive mimics, though the situation is somewhat different from those outlined previously. They can mimic their hosts' natural prey, allowing themselves to be eaten as a pathway into their host. Leucochloridium , a genus of flatworm, matures in the digestive system of songbirds, their eggs then passing out of the bird in the faeces. They are then taken up by Succinea , a terrestrial snail. The eggs develop in this intermediate host, and must then find a suitable bird to mature in. Since the host birds do not eat snails, the sporocyst has another strategy to reach its host's intestine. They are brightly coloured and move in a pulsating fashion. A sporocyst-sac pulsates in the snail's eye stalks, [75] [76] 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. [14] A nematode ( Myrmeconema neotropicum ) 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 (rear part) is held raised. This presumably increases the chances of the ant being eaten by birds. [77]

Reproductive

Reproductive mimicry occurs when the actions of the dupe directly aid in the mimic's reproduction. This is common in plants with deceptive flowers that do not provide the reward they seem to offer and it may occur in Papua New Guinea fireflies, in which the signal of Pteroptyx effulgens is used by P. tarsalis to form aggregations to attract females. [78] Other forms of mimicry have a reproductive component, such as Vavilovian mimicry involving seeds, vocal mimicry in birds, [79] [80] [81] and aggressive and Batesian mimicry in brood parasite-host systems. [82]

Bakerian and Dodsonian

Bakerian mimicry, named after Herbert G. Baker, [83] is a form of automimicry where female flowers 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. [84]

In Dodsonian mimicry, named after Calaway H. Dodson, the model belongs to a different species than the mimic. [85] By resembling the model, a flower can lure its pollinators without offering nectar. The mechanism occurs in several orchids, including Epidendrum ibaguense which mimics flowers of Lantana camara and Asclepias curassavica , and is pollinated by monarch butterflies and perhaps hummingbirds. [86]

Kirbyan mimicry, brood parasitism

Brood parasitism or Kirbyan mimicry is a two species system where a brood parasite mimics its host. Cuckoos are a canonical example; the female cuckoo has its offspring raised by a bird of a different species, cutting down the biological mother's parental investment. The ability to lay eggs that mimic the host eggs is the key adaptation. The adaptation to different hosts is inherited through the female line in so-called gentes (gens, singular). Intraspecific brood parasitism, where a female lays in a conspecific's nest, as illustrated by the goldeneye duck (Bucephala clangula), do not involve mimicry [87] The parasitic butterfly Phengaris rebeli parasitizes the ant species Myrmica schencki by releasing chemicals that fool the worker ants to believe that the caterpillar larvae are ant larvae. This enables the larvae to be brought directly into the ant's nest. [88]

Pouyannian

Dasyscolia ciliata, a scoliid wasp, attempting to copulate with a flower of the orchid Ophrys speculum Dasyscolia ciliata.jpg
Dasyscolia ciliata , a scoliid wasp, attempting to copulate with a flower of the orchid Ophrys speculum

In Pouyannian mimicry, a flower mimics a female of a certain insect species, inducing the males of that species to try to copulate with the flower. This is much like aggressive mimicry in fireflies, but with a more benign outcome for the pollinator. The mechanism is named after Maurice-Alexandre Pouyanne, who first described the phenomenon. [2] [91] [92] 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. Depending on the morphology of the flower, a pollen sac called a pollinium is attached to the head or abdomen of the male. This is then transferred to the stigma of the next flower the male tries to inseminate, resulting in pollination. The mimicry is a combination of visual, by olfaction, and by touch. [93]

Vavilovian

Rye is a secondary crop, originally being a mimetic weed of wheat. Secale cereale.jpg
Rye is a secondary crop, originally being a mimetic weed of wheat.

Vavilovian mimicry is found in weeds that come to share characteristics with a domesticated plant through artificial selection. [2] It is named after Russian botanist and geneticist Nikolai Vavilov. [94] Selection against the weed may occur either by manually killing the weed, or by separating its seeds from those of the crop by winnowing. Vavilovian mimicry illustrates unintentional selection by man. Weeders do not want to select weeds and their seeds that look increasingly like cultivated plants, yet there is no other option. For example, early barnyard grass, Echinochloa oryzoides , is a weed in rice fields and looks similar to rice; its seeds are often mixed in rice and have become difficult to separate through Vavilovian mimicry. [95] Vavilovian mimics may eventually be domesticated themselves, as in the case of rye in wheat; Vavilov called these weed-crops secondary crops. [94]

Inter-sexual mimicry

Inter-sexual mimicry (a type of automimicry, as it is within a single species) occurs when individuals of one sex in a species mimic members of the opposite sex to facilitate sneak mating. An example is the three male forms of the marine isopod Paracerceis sculpta . Alpha males are the largest and guard a harem of females. Beta males mimic females and manage to enter the harem of females without being detected by the alpha males allowing them to mate. Gamma males are the smallest males and mimic juveniles. This also allows them to mate with the females without the alpha males detecting them. [96] Similarly, among common side-blotched lizards, some males mimic the yellow throat coloration and even mating rejection behaviour of the other sex to sneak matings with guarded females. These males look and behave like unreceptive females. This strategy is effective against "usurper" males with orange throats, but ineffective against blue throated "guarder" males, which chase them away. [97] [98] Female spotted hyenas have pseudo-penises that make them look like males. [99]

See also

Notes

    Related Research Articles

    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">Batesian mimicry</span> Bluffing imitation of a strongly defended species

    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.

    <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">Flower mantis</span> Species of mantis camouflaged to resemble flowers to lure their prey

    Flower mantises are praying mantises that use a special form of camouflage referred to as aggressive mimicry, which they not only use to attract prey, but avoid predators as well. These insects have specific colorations and behaviors that mimic flowers in their surrounding habitats.

    <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">Gilbertian mimicry</span>

    In evolutionary biology, Gilbertian mimicry is a rare type of mimicry involving only two species, a host or prey animal which is the mimic, and its parasite or predator, which is both the model for the mimicry, and the dupe that is deceived by it. The mechanism provides a measure of protection for the mimic, as parasites and predators rarely attack their own species.

    <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.

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

    In evolutionary biology, mimicry in plants is where a plant evolves to resemble another organism physically or chemically, increasing the mimic's Darwinian fitness. Mimicry in plants has been studied far less than mimicry in animals, with fewer documented cases and peer-reviewed studies. However, it may provide protection against herbivory, or may deceptively encourage mutualists, like pollinators, to provide a service without offering a reward in return.

    Locomotor mimicry is a subtype of Batesian mimicry in which animals avoid predation by mimicking the movements of another species phylogenetically separated. This can be in the form of mimicking a less desirable species or by mimicking the predator itself. Animals can show similarity in swimming, walking, or flying of their model animals.

    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.

    <span class="mw-page-title-main">Chemical communication in insects</span>

    Chemical communication in insects is social signalling between insects of the same or different species, using chemicals. These chemicals may be volatile, to be detected at a distance by other insects' sense of smell, or non-volatile, to be detected on an insect's cuticle by other insects' sense of taste. Many of these chemicals are pheromones, acting like hormones outside the body.

    References

    1. 1 2 Harper, Douglas. "Online Etymology Dictionary" . Retrieved 23 February 2022.
    2. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Pasteur, G. (1982). "A Classificatory Review of Mimicry Systems". Annual Review of Ecology and Systematics . 13: 169–199. doi:10.1146/annurev.es.13.110182.001125. JSTOR   2097066.
    3. 1 2 3 Kirby, William; Spence, William (1823). An Introduction to Entomology. Vol. 2 (3rd ed.). Longman, Hurst, Rees, Orme & Brown. p. 405.
    4. 1 2 Aristotle, History of Animals , book 9, chapter 8.
    5. 1 2 Mallet, James. "Fritz Müller in 1891". Archived from the original on 30 June 2024. Retrieved 18 November 2017.
    6. 1 2 3 Bates, Henry W. (1862). "Contributions to an insect fauna of the Amazon valley. Lepidoptera: Heliconidae". Transactions of the Linnean Society. 23 (3): 495–566. doi:10.1111/j.1096-3642.1860.tb00146.x.
    7. ↑ Bates, Henry W. (1863). The naturalist on the river Amazons. Murray.
    8. ↑ Müller, Fritz (1878). "Ueber die Vortheile der Mimicry bei Schmetterlingen" [On the Advantages of Mimicry in Butterflies]. Zoologischer Anzeiger (in German). 1: 54–55.
    9. 1 2 Müller, Fritz (1879). "Ituna and Thyridia; a remarkable case of mimicry in butterflies" (PDF). Proclamations of the Entomological Society of London. 1879. Translated by R. Meldola: 20–29. Archived (PDF) from the original on 2 March 2024.
    10. ↑ Sherratt, Thomas (2008). "The Evolution of Müllerian Mimicry". Die Naturwissenschaften. 95 (8): 681–695. Bibcode:2008NW.....95..681S. doi:10.1007/s00114-008-0403-y. PMC   2443389 . PMID   18542902.
    11. ↑ King, R. C.; Stansfield, W. D.; Mulligan, P. K. (2006). A dictionary of genetics (7th ed.). Oxford University Press. p.  278. ISBN   978-0-19-530762-7.
    12. ↑ Kikuchi, D. W.; Pfennig, D. W. (2013). "Imperfect Mimicry and the Limits of Natural Selection". The Quarterly Review of Biology . 88 (4): 297–315. doi:10.1086/673758. PMID   24552099. S2CID   11436992.
    13. 1 2 Dalziell, Anastasia H.; Welbergen, Justin A. (27 April 2016). "Mimicry for all modalities". Ecology Letters . 19 (6): 609–619. Bibcode:2016EcolL..19..609D. doi:10.1111/ele.12602. PMID   27117779.
    14. 1 2 3 4 5 6 Wickler, Wolfgang (1968). Mimicry in plants and animals . McGraw-Hill.
    15. ↑ Wickler, Wolfgang (1965). "Mimicry and the Evolution of Animal Communication". Nature . 208 (5010): 519–21. Bibcode:1965Natur.208..519W. doi:10.1038/208519a0. S2CID   37649827.
    16. ↑ Radford, Benjamin; Frazier, Kendrick (January 2017). "Cheats and Deceits: How Animals and Plants Exploit and Mislead". Skeptical Inquirer. 41 (1): 60.
    17. 1 2 3 4 5 Ruxton, Graeme D.; Sherratt, Thomas N.; Speed, M. P. (2004). Avoiding Attack: the Evolutionary Ecology of Crypsis, Warning Signals, and Mimicry. Oxford University Press.
    18. ↑ Boppré, Michael; Vane-Wright, Richard I.; Wickler, Wolfgang (1 January 2017). "A hypothesis to explain accuracy of wasp resemblances". Ecology and Evolution. 7 (1): 73–81. Bibcode:2017EcoEv...7...73B. doi:10.1002/ece3.2586. PMC   5214283 . PMID   28070276.
    19. 1 2 3 Campbell, N. A. (1996). Biology (4th ed.). Benjamin Cummings. Chapter 50. ISBN   0-8053-1957-3.
    20. ↑ Boyden, T. C. (1980). "Floral mimicry by Epidendrum ibaguense (Orchidaceae) in Panama". Evolution. 34 (1): 135–136. doi:10.2307/2408322. JSTOR   2408322. PMID   28563205.
    21. ↑ Roy, B. A. (1994). "The effects of pathogen-induced pseudoflowers and buttercups on each other's insect visitation". Ecology. 75 (2): 352–358. Bibcode:1994Ecol...75..352R. doi:10.2307/1939539. JSTOR   1939539.
    22. ↑ Johnson, Steven D.; Schiestl, Florian P. (2016). Floral Mimicry. Oxford University Press. ISBN   978-0-19-104723-7.
    23. ↑ Alexander, Victoria N. (2002). "Nabokov, Teleology and Insect Mimicry". Nabokov Studies . 7: 177–213. doi:10.1353/nab.2010.0004. S2CID   42675699.
    24. ↑ Holmgren, N. M. A.; Enquist, M. (1999). "Dynamics of mimicry evolution" (PDF). Biological Journal of the Linnean Society. 66 (2): 145–158. doi: 10.1111/j.1095-8312.1999.tb01880.x .
    25. ↑ Kunte, K.; Zhang, W.; Tenger-Trolander, A.; Palmer, D. H.; Martin, A.; Reed, R. D.; Mullen, S. P.; Kronforst, M. R. (2014). "doublesex is a mimicry supergene". Nature. 507 (7491): 229–232. Bibcode:2014Natur.507..229K. doi:10.1038/nature13112. PMID   24598547. S2CID   4448793.
    26. ↑ Wilson, J.; Jahner, J.; Williams, K.; Forister, M. (2013). "Ecological and Evolutionary Processes Drive the Origin and Maintenance of Imperfect Mimicry". PLOS ONE. 8 (4): e61610. Bibcode:2013PLoSO...861610W. doi: 10.1371/journal.pone.0061610 . PMC   3625143 . PMID   23593490.
    27. ↑ Kikuchi, D.; Pfenning, D. (2010). "Predator Cognition Permits Imperfect Coral Snake Mimicry". The American Naturalist. 176 (6): 830–834. doi:10.1086/657041. PMID   20950143. S2CID   35411437.
    28. ↑ Howse, P. E.; Allen, J. A. (1994). "Satyric Mimicry: The Evolution of Apparent Imperfection". Proceedings of the Royal Society B. 257 (1349): 111–114. Bibcode:1994RSPSB.257..111H. doi:10.1098/rspb.1994.0102. S2CID   84458742.
    29. ↑ Robertson, D. Ross (2013). "Who resembles whom? Mimetic and coincidental look-alikes among tropical reef fishes". PLOS ONE. 8 (1): e54939. Bibcode:2013PLoSO...854939R. doi: 10.1371/journal.pone.0054939 . PMC   3556028 . PMID   23372795.
    30. ↑ Robertson, D. Ross (2015). "Coincidental resemblances among coral reef fishes from different oceans". Coral Reefs. 34 (3): 977. Bibcode:2015CorRe..34..977R. doi: 10.1007/s00338-015-1309-8 .
    31. ↑ Pawlik, J.R. (2012). "12". In Fattorusso, E.; Gerwick, W.H.; Taglialatela-Scafati, O. (eds.). Antipredatory defensive roles of natural products from marine invertebrates. Springer. pp. 677–710. ISBN   978-90-481-3833-3.
    32. ↑ Skelhorn, John; Rowland, Hannah M.; Ruxton, Graeme D. (2010). "The Evolution and Ecology of Masquerade". Biological Journal of the Linnean Society. 99: 1–8. doi: 10.1111/j.1095-8312.2009.01347.x .
    33. ↑ Wiklund, Christer; Tullberg, Birgitta S. (September 2004). "Seasonal polyphenism and leaf mimicry in the comma butterfly". Animal Behaviour . 68 (3): 621–627. doi:10.1016/j.anbehav.2003.12.008. S2CID   54270418.
    34. ↑ Endler, John A. (August 1981). "An Overview of the Relationships Between Mimicry and Crypsis". Biological Journal of the Linnean Society. 16 (1): 25–31. doi:10.1111/j.1095-8312.1981.tb01840.x.
    35. ↑ Stevens, Martin; Hopkins, Elinor; Hinde, William; Adcock, Amabel; Connolly, Yvonne; Troscianko, Tom; Cuthill, Innes C. (November 2007). "Field Experiments on the effectiveness of 'eyespots' as predator deterrents". Animal Behaviour . 74 (5): 1215–1227. doi:10.1016/j.anbehav.2007.01.031. S2CID   53186893.
    36. ↑ Stevens, Martin (22 June 2007). "Predator perception and the interrelation between different forms of protective coloration". Proceedings of the Royal Society B: Biological Sciences. 274 (1617): 1457–1464. doi:10.1098/rspb.2007.0220. PMC   1950298 . PMID   17426012.
    37. ↑ Stevens, Martin; Stubbins, Claire L.; Hardman, Chloe J. (30 May 2008). "The anti-predator function of 'eyespots' on camouflaged and conspicuous prey". Behavioral Ecology and Sociobiology. 62 (11): 1787–1793. doi:10.1007/s00265-008-0607-3. S2CID   28288920.
    38. ↑ Hossie, Thomas John; Sherratt, Thomas N. (August 2013). "Defensive posture and eyespots deter avian predators from attacking caterpillar models". Animal Behaviour. 86 (2): 383–389. doi:10.1016/j.anbehav.2013.05.029. S2CID   53263767.
    39. ↑ Endler, John A. (1981). "An overview of the relationships between mimicry and crypsis". Biological Journal of the Linnean Society. 16: 25–31. doi:10.1111/j.1095-8312.1981.tb01840.x.
    40. ↑ Allen, J. A.; Cooper, J. M. (2010). "Crypsis and masquerade". Journal of Biological Education. 19 (4): 268. doi:10.1080/00219266.1985.9654747.
    41. ↑ Pasteur cites Aristotle, History of Animals , book 9, chapter 9.
    42. 1 2 Rowland, Hannah M.; Ihalainen, Eira; Lindström, Leena; Mappes, Johanna; Speed, Michael P. (2007). "Co-mimics have a mutualistic relationship despite unequal defences". Nature . 448 (7149): 64–67. doi:10.1038/nature05899. ISSN   0028-0836.
    43. ↑ Stearns, S. C.; Hoekstra, Rolf F. (2000). Evolution: An Introduction (5th ed.). Oxford University Press. p. 464. ISBN   978-0-19-854968-0.
    44. ↑ 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. 59 (4): 351–365. doi: 10.1111/j.1095-8312.1996.tb01471.x .
    45. ↑ Barber, J. R.; Conner, W. E. (2007). "Acoustic mimicry in a predator–prey interaction". PNAS. 104 (22): 9331–9334. Bibcode:2007PNAS..104.9331B. doi: 10.1073/pnas.0703627104 . PMC   1890494 . PMID   17517637.
    46. ↑ Barber, Jesse R.; Chadwell, Brad A.; Garrett, Nick; Schmidt-French, Barbara; Conner, William E. (July 2009). "Naïve bats discriminate arctiid moth warning sounds but generalize their aposematic meaning". The Journal of Experimental Biology. 212 (Pt 14): 2141–2148. doi:10.1242/jeb.029991. ISSN   0022-0949. PMID   19561203. S2CID   1303252.
    47. ↑ Barber, Jesse R.; Plotkin, David; Rubin, Juliette J.; Homziak, Nicholas T.; Leavell, Brian C.; et al. (21 June 2022). "Anti-bat ultrasound production in moths is globally and phylogenetically widespread". Proceedings of the National Academy of Sciences of the United States of America. 119 (25): e2117485119. Bibcode:2022PNAS..11917485B. doi: 10.1073/pnas.2117485119 . PMC   9231501 . PMID   35704762.
    48. ↑ Kawahara, Akito Y.; Barber, Jesse R. (19 May 2015). "Tempo and mode of antibat ultrasound production and sonar jamming in the diverse hawkmoth radiation". Proceedings of the National Academy of Sciences. 112 (20): 6407–6412. Bibcode:2015PNAS..112.6407K. doi: 10.1073/pnas.1416679112 . PMC   4443353 . PMID   25941377.
    49. ↑ "Mimic Octopus, Thaumoctopus mimicus at MarineBio.org". Archived from the original on 18 July 2017. Retrieved 9 June 2007.
    50. ↑ "Deceptive Woodpecker Uses Mimicry to Avoid Competition". AMNH. Retrieved 12 August 2015.
    51. ↑ Gianoli, Ernesto (2014). "Leaf Mimicry in a Climbing Plant Protects against Herbivory". Cell. 24 (9): 984–987. doi: 10.1016/j.cub.2014.03.010 . PMID   24768053.
    52. ↑ Meyer, A. (2006). "Repeating Patterns of Mimicry". PLOS Biology . 4 (10): e341. doi: 10.1371/journal.pbio.0040341 . PMC   1617347 . PMID   17048984.
    53. ↑ 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 (1): 86–93. doi:10.2307/2407675. JSTOR   2407675. PMID   28565050.
    54. ↑ Benson, W. W. (1977). "On the Supposed Spectrum Between Batesian and Mullerian Mimicry". Evolution. 31 (2): 454–455. doi:10.2307/2407770. JSTOR   2407770. PMID   28563231.
    55. 1 2 Emsley, M. G. (1966). "The mimetic significance of Erythrolamprus aesculapii ocellatus Peters from Tobago". Evolution. 20 (4): 663–64. doi:10.2307/2406599. JSTOR   2406599. PMID   28562911.
    56. ↑ Mertens, Robert (1956). "Das Problem der Mimikry bei Korallenschlangen". Zool. Jahrb. Syst (in German). 84: 541–76.
    57. 1 2 Hecht, M. K.; Marien, D. (1956). "The coral snake mimic problem: a reinterpretation". Journal of Morphology. 98 (2): 335–365. doi:10.1002/jmor.1050980207. S2CID   83825414.
    58. 1 2 Sheppard, P. M.; Wickler, Wolfgang (1969). "Review of Mimicry in plants and animals by Wolfgang Wickler". Journal of Animal Ecology. 38 (1): 243. doi:10.2307/2762. JSTOR   2762.
    59. ↑ Wasmann, Erich (1894). Kritisches Verzeichniss der myrmecophilin und termitophilen Arthropoden[Critical Inventory of Myrmecophile and Termitophile Arthropods] (in German). Berlin: Felix Dames.
    60. ↑ Hölldobler, Bert; Wilson, Edward O. (1990). The Ants. Harvard University Press. pp. 511–514. ISBN   978-0-674-04075-5.
    61. ↑ Gilbert, Lawrence E. (1975). "Ecological consequences of a coevolved mutualism between butterflies and plants". In L. E. Gilbert; P. H. Raven (eds.). Coevolution of Animals and Plants. University of Texas Press. pp. 210–240. OCLC   636384400.
    62. ↑ Brower, Lincoln P. (1970). "Plant poisons in a terrestrial food chain and implications for mimicry theory". In Chambers, K. L. (ed.). Biochemical Coevolution. Corvallis, Oregon, USA: Oregon State Univ. pp. 69–82.
    63. ↑ Brower, Lincoln P.; Van Brower, J. V. Z.; Corvino, J. M. (1967). "Plant poisons in a terrestrial food chain". Proceedings of the National Academy of Sciences of the United States of America. 57 (4): 893–98. Bibcode:1967PNAS...57..893B. doi: 10.1073/pnas.57.4.893 . PMC   224631 . PMID   5231352.
    64. ↑ "Northern Pygmy Owl (Glaucidium californicum)". Owl Research Institute. Archived from the original on 28 December 2015. Retrieved 23 August 2015.
    65. ↑ Sourakov, Andrei (2013): Two heads are better than one: false head allows Calycopis cecrops (Lycaenidae) to escape predation by a Jumping Spider, Phidippus pulcherrimus (Salticidae), Journal of Natural History, 47:15-16, 1047-1054
    66. ↑ Robbins, Robert K. The "False Head" Hypothesis: Predation and Wing Pattern Variation of Lycaenid Butterflies. The American Naturalist Vol. 118, No. 5 (Nov., 1981), pp. 770-775
    67. ↑ Begon, M.; Townsend, C.; Harper, J. (1996) Ecology: Individuals, populations and communities (third edition) Blackwell Science, London
    68. ↑ Jackson, R. R. (1995). "Eight-legged tricksters: Spiders that specialize at catching other spiders". BioScience. 42 (8): 590–98. doi:10.2307/1311924. JSTOR   1311924.
    69. ↑ Craig, C. L. (1995). "Webs of Deceit". Natural History . 104 (3): 32–35.
    70. ↑ Lloyd, J. E. (1965) Aggressive Mimicry in Photuris: Firefly Femmes Fatales Science 149:653–654.
    71. ↑ Lloyd, J. E. (1975). "Aggressive Mimicry in Photuris Fireflies: Signal Repertoires by Femmes Fatales". Science. 187 (4175): 452–453. Bibcode:1975Sci...187..452L. doi:10.1126/science.187.4175.452. PMID   17835312. S2CID   26761854.
    72. ↑ 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 (4): 515–525. Bibcode:1996JEcol..84..515M. doi:10.2307/2261474. JSTOR   2261474.
    73. ↑ Wickler, W. (1966). "Mimicry in Tropical Fishes". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 251 (772): 473–474. Bibcode:1966RSPTB.251..473W. doi:10.1098/rstb.1966.0036. S2CID   83609965.
    74. ↑ Willis, E. O. (1963). "Is the Zone-Tailed Hawk a Mimic of the Turkey Vulture?". The Condor. 65 (4): 313–317. doi:10.2307/1365357. JSTOR   1365357.
    75. ↑ See here for a photo.
    76. ↑ Moore, J. (2002). Parasites and the behavior of animals. Oxford University Press.
    77. ↑ Yanoviak, S. P.; Kaspari, M.; Dudley, R.; Poinar, G. Jr (2008). "Parasite-induced fruit mimicry in a tropical canopy ant" (PDF). The American Naturalist. 171 (4): 536–44. doi:10.1086/528968. PMID   18279076. S2CID   23857167.
    78. ↑ Ohba, N.; Shimoyama, Ayu (2009). Meyer-Rochow, V. B. (ed.). Bioluminescence in Focus - a collection of illuminating essays. Research Signpost; Trivandrum, Kerala, India. pp. 229–242.
    79. ↑ Dalziell, Anastasia H.; Welbergen, Justin A.; Igic, Branislav; Magrath, Robert D. (30 July 2014). "Avian vocal mimicry: a unified conceptual framework". Biological Reviews. 90 (2): 643–668. doi:10.1111/brv.12129. PMID   25079896. S2CID   207101926.
    80. ↑ Kelley, Laura A.; Coe, Rebecca L.; Madden, Joah R.; Healy, Susan D. (1 September 2008). "Vocal mimicry in songbirds". Animal Behaviour. 76 (3): 521–528. doi:10.1016/j.anbehav.2008.04.012. S2CID   53192695.
    81. ↑ Goller, Maria; Shizuka, Daizaburo (22 June 2018). "Evolutionary origins of vocal mimicry in songbirds". Evolution Letters. 2 (4): 417–426. doi:10.1002/evl3.62. PMC   6121844 . PMID   30283692.
    82. ↑ Davies, Nick (2015). Cuckoo: Cheating by Nature. Bloomsbury. ISBN   978-1-4088-5656-7. Archived from the original on 28 February 2021. Retrieved 8 November 2018.
    83. ↑ Baker, Herbert 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. [ clarification needed ]Variation, breeding, and conservation of tropical trees. Academic Press, London, U.K.
    84. ↑ Bawa, K. S. (1980). "Mimicry of male by female flowers and intrasexual competition for pollinators in Jacaratia dolichaula (D. Smith) Woodson (Caricaceae)". Evolution . 34 (3): 467–74. doi:10.2307/2408216. JSTOR   2408216. PMID   28568703.
    85. ↑ Dodson, C. H.; Frymire, G. P. (1961). "Natural pollination of orchids". Missouri Botanical Garden Bulletin. 49: 133–39.
    86. ↑ Boyden, T. C. (1980). "Floral mimicry by Epidendrurn ibaguense (Orchidaceae) in Panama". Evolution. 34 (1): 135–36. doi:10.2307/2408322. JSTOR   2408322. PMID   28563205.
    87. ↑ Andersson, M.; Eriksson, M. O. G. (1982). "Nest parasitism in Goldeneyes Bucephala clangula: some evolutionary aspects". American Naturalist. 120: 1–16. doi:10.1086/283965. S2CID   86699716.
    88. ↑ Barbero, Francesca; Thomas, J.A.; Bonelli, S.; Balletto, E.; Schonrogge, K. (2009). "Acoustical mimicry in a predatory social parasite of ants". Journal of Experimental Biology. 212 (Pt 24): 4084–4090. doi: 10.1242/jeb.032912 . PMID   19946088 . Retrieved 28 September 2013.
    89. ↑ Welbergen, Justin A.; Davies, Nicholas B. (2011). "A parasite in wolf's clothing: hawk mimicry reduces mobbing of cuckoos by hosts". Behavioral Ecology. 22 (3): 574–579. doi: 10.1093/beheco/arr008 .
    90. ↑ Davies, N. B.; Welbergen, J. A. (2008). "Cuckoo–hawk mimicry? An experimental test". Proceedings of the Royal Society B. 275 (1644): 1817–1822. doi:10.1098/rspb.2008.0331. PMC   2587796 . PMID   18467298.
    91. ↑ Correvon H., Pouyanne M. (1916) Un curieux cas de mimetisme chez les Ophrydées. J. Soc. Nat. Hortic. Fr.17: 29–31, 41–42, 84.
    92. ↑ Pouyanne, M.-A. (1917). "La fécondation des Ophrys par les insectes". Bulletin de la Société d'histoire naturelle de l'Afrique du Nord. 8: 1–2.
    93. ↑ Pramanik, Dewi; Dorst, Nemi; Meesters, Niels; et al. (2020). "Evolution and development of three highly specialized floral structures of bee-pollinated Phalaenopsis species". EvoDevo. 11 (1): 16. doi: 10.1186/s13227-020-00160-z . PMC   7418404 . PMID   32793330.
    94. 1 2 Vavilov, N. I. (1951). "The origin, variation, immunity and breeding of cultivated plants (translation by K. S. Chester)". Chronica Botanica. 13: 1–366.
    95. ↑ Barrett, S. (1983). "Mimicry in Plants". Scientific American . 257 (3): 76–83. Bibcode:1987SciAm.257c..76B. doi:10.1038/scientificamerican0987-76.
    96. ↑ Shuster, Stephen (May 1987). "Alternative Reproductive Behaviors: Three Discrete Male Morphs in Paracerceis sculpta, an Intertidal Isopod from the Northern Gulf of California". Journal of Crustacean Biology. 7 (2): 318–327. doi:10.2307/1548612. JSTOR   1548612.
    97. ↑ Sinervo, B.; C. M. Lively (1996). "The rock–paper–scissors game and the evolution of alternative male strategies". Nature. 380 (6571): 240–243. Bibcode:1996Natur.380..240S. doi:10.1038/380240a0. S2CID   205026253.
    98. ↑ 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 (4): 222–233. doi:10.1006/hbeh.2000.1622. PMID   11104640. S2CID   5759575.
    99. ↑ Muller, M. N.; Wrangham, R. (2002). "Sexual Mimicry in Hyenas". The Quarterly Review of Biology . 77 (1): 3–16. doi:10.1086/339199. PMID   11963460. S2CID   43440407.

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