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Aggressive mimicry

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.[2][3][4] 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 (see pseudocopulation),[5] 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 (after George and Elizabeth Peckham),[6][7][8] but it is seldom used.[a]

The humpback anglerfish uses a modified dorsal spine as a fishing rod with a bioluminescent lure to attract and capture prey.[1]
A camouflaged predator: snow leopard in Ladakh. The distinction between aggressive mimicry and predator camouflage depends on the signal given to the prey, not easily determined.

Aggressive mimicry is opposite in principle to defensive mimicry, where the mimic generally benefits from being treated as harmful. The mimic may resemble its own prey, or some other organism which is beneficial or at least not harmful to the prey. The model, i.e. the organism being 'imitated', may experience increased or reduced fitness, or may not be affected at all by the relationship. On the other hand, the signal receiver inevitably suffers from being tricked, as is the case in most mimicry complexes.

Aggressive mimicry often involves the predator employing signals which draw its potential prey towards it, a strategy which allows predators to simply sit and wait for prey to come to them. The promise of food or sex are most commonly used as lures. However, this need not be the case; as long as the predator's true identity is concealed, it may be able to approach prey more easily than would otherwise be the case. In terms of species involved, systems may be composed of two or three species; in two-species systems the signal receiver, or "dupe", is the model.

In terms of the visual dimension, the distinction between aggressive mimicry and camouflage is not always clear. Authors such as Wickler[8] have emphasized the significance of the signal to its receiver as delineating mimicry from camouflage. However, it is not easy to assess how 'significant' a signal may be for the dupe, and the distinction between the two can thus be rather fuzzy. Mixed signals may be employed: aggressive mimics often have a specific part of the body sending a deceptive signal, with the rest being hidden or camouflaged.

Contrast with defensive mimicry edit

 
Defensive Batesian mimics, like this bumblebee-mimicking hoverfly, are the antithesis of aggressive mimics.

Aggressive mimicry stands in semantic contrast with defensive mimicry, where it is the prey that acts as a mimic, with predators being duped. Defensive mimicry includes the well-known Batesian and Müllerian forms of mimicry, where the mimic shares outward characteristics with an aposematic or harmful model. In Batesian mimicry, the mimic is modeled on a dangerous (usually unpalatable) species, while in Müllerian mimicry both species are harmful, and act as comimics, converging on a common set of signals and sharing the burden of 'educating' their predators. Included in defensive mimicry is the lesser known Mertensian mimicry, where the mimic is more harmful than the model, and Vavilovian mimicry, where weeds come to mimic crops through unintentional artificial selection. In defensive mimicry, the mimic benefits by avoiding a harmful interaction with another organism that would be more likely to take place without the deceptive signals employed. Harmful interactions might involve being eaten, or pulled out of the ground as a weed. In contrast, the aggressive mimic benefits from an interaction that would be less likely to take place without the deception, at the expense of its target.[9]

Classification edit

Luring prey edit

In some cases the signal receiver is lured toward the mimic. This involves mimicry of a resource that is often vital to the prey's survival (or more precisely, the survival of its genes) such as nutrition or a mate. If the bait offered is of little value to prey they would not be expected to take such a risk. For example, in all known cases of sexual signal mimicry it is always the male sex that is deceived (in fact, it has been suggested that females of some species have evolved mimicry as a strategy to avoid unwanted matings).[10] In these cases the predator need not move about foraging for prey, but may simply stay still and allow prey to come to it. Some studies suggest that the northern shrike (Lanius excubitor) sings in winter often imitating small passerines that may be preyed upon when lured within reach.[11] There has been one report of a margay using mimicry of the cry of an infant pied tamarin to try to lure an adult tamarin within striking distance.[12]

Appearance of food edit

 
The alligator snapping turtle uses its tongue to lure fish.

Many aggressive mimics use the promise of nourishment as a way of attracting prey. The alligator snapping turtle (Macrochelys temminckii) is a well-camouflaged ambush predator. Its tongue bears a conspicuous pink extension that resembles a worm and can be wriggled around;[13] fish that try to eat the "worm" are themselves eaten by the turtle. Similarly, some snakes employ caudal luring (using the tail)[14][15] or lingual luring (using the tongue) to entice small vertebrates into striking range.[16][17]

 
Argiope argentata and its web

Aggressive mimicry is common amongst spiders, both in luring prey and stealthily approaching predators.[18] One case is the golden orb weaver (Nephila clavipes), which spins a conspicuous golden coloured web in well-lit areas. Experiments show that bees are able to associate the webs with danger when the yellow pigment is not present, as occurs in less well-lit areas where the web is much harder to see. Other colours too were learned and avoided, but bees seemed least able to effectively associate yellow pigmented webs with danger. Yellow is the colour of many nectar bearing flowers, however, so perhaps avoiding yellow is not worthwhile. Another form of mimicry is based not on colour but pattern. Species such as 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 bees' ability to remember web patterns. Bees are able to associate a certain pattern with a spatial location, meaning the spider must spin a new pattern regularly or suffer diminishing prey capture.[19]

 
The bright leaves of the Venus flytrap (Dionaea muscipula) attract insects in the same way as flowers.

Spiders can be the prey of aggressive mimics. The assassin bug Stenolemus bituberus preys on spiders, entering their web and plucking its silk threads until the spider approaches. This vibrational aggressive mimicry matches a general pattern of vibrations which spiders treat as prey, having a similar temporal structure and amplitude to leg and body movements of typical prey caught in the web.[20]

Larvae of the ground beetle Epomis move their mandibles one after another to lure amphibians toward them and then prey on them. Their body structure allows them to bite and feed on the amphibians even when they are ingested by larger prey such as frogs.[21]

Although plants are better known for defensive mimicry, there are exceptions. For example, many flowers use mimicry to attract pollinators, while others may trick insects into dispersing their seeds. Nonetheless, most mimicry in plants[b] would not be classified as aggressive, as although luring pollinators is similar to cases above, they are certainly not eaten by the plant. However some carnivorous plants may be able to increase their rate of capture through mimicry.[24] For example, some have patterns in the ultraviolet region of the electromagnetic spectrum, much like the spider webs described above.[25]

Bipolar mimicry systems edit

Mimicry systems involving only two species are known as bipolar.[9] Only one bipolar arrangement is possible here, namely where the dupe is itself the model.[c] There are two such variants on this arrangement of mimic imitating its target, in the first case, termed Batesian-Wallacian mimicry[9] after Henry Walter Bates[26] and Alfred Russel Wallace,[27] the model is the prey species. In the second case, the model is the host of a brood parasite.

Batesian-Wallacian or prey mimicry edit

 
Mastophora spider holding its bolas with a leg, attracting and capturing a male moth

In some cases of Batesian-Wallacian mimicry, the model is a sexually receptive female, which provides a strong attractive effect on males. Some spiders use chemical rather than visual means to ensnare prey. Female bolas spiders of the genus Mastophora lure male moth-flies (Diptera, true flies, but resembling moths) by producing analogues of the moth species' sex pheromones. Each species of spider appears to specialize in a particular species of prey in the family Psychodidae. Juveniles use their front pair of legs to capture prey, such as flies. Older spiders use a different strategy however, swinging a sticky ball known as a bolas suspended by a silk thread at moths. But both old and juvenile are able to lure prey via this olfactory signal; even young spiderlings have been shown to attract prey species.[28]

 
The spotted predatory katydid (Chlorobalius leucoviridis) is an acoustic aggressive mimic of cicadas.

The listroscelidine katydid Chlorobalius leucoviridis of inland Australia is capable of attracting male cicadas of the Tribe Cicadettini by imitating the species-specific reply clicks of sexually receptive female cicadas. This example of acoustic aggressive mimicry is similar to the Photuris firefly case in that the predator's mimicry is remarkably versatile – playback experiments show that C. leucoviridis is able to attract males of many cicada species, including Cicadettine cicadas from other continents, even though cicada mating signals are species-specific. The evolution of versatile mimicry in C. leucoviridis may have been facilitated by constraints on song evolution in duetting communication systems in which reply signals are recognizable only by their precise timing in relation to the male song (<< 100 ms reply latency).[29][30]

Female fireflies of the genus Photuris emit the same light signals that females of the genus Photinus use as a mating signal.[31] Male fireflies from several different genera are attracted to these mimics, and are subsequently captured and eaten. Female signals are based on that received from the male, each female having a repertoire of signals matching the delay and duration of the female of the corresponding species. This mimicry may have evolved from non-mating signals that have become modified for predation.[32]

Kirbyan or brood parasite mimicry edit

 
Brood parasitism: four clutches of reed warbler eggs, each containing one (larger) cuckoo egg

Host-parasite mimicry is a situation where a parasite mimics its own host. As with mimicry of the female sex outlined previously, only two species are involved, the model and mimic being of the same species. Brood parasitism, a form of kleptoparasitism where the mother has its offspring raised by another unwitting organism, is one such situation where host-parasite mimicry has evolved. Georges Pasteur[9] terms this form of aggressive-reproductive mimicry Kirbyan mimicry, after the English entomologist William Kirby.[33]

Wicklerian-Eisnerian or mimicry of harmless species edit

The prey does not have to be attracted towards the predator for the predator to benefit: it is sufficient for the predator simply not to be identified as a threat. Wicklerian-Eisnerian mimics may resemble a mutualistic ally, or a species of little significance to the prey such as a commensal.[9] For example, the hemipteran Arachnocoris berytoides resembles Faiditus caudatus, a spider commensal of ants.[34]

Mimicry of cleaner fish edit

 
Two bluestreak cleaner wrasse cleaning a potato grouper, Epinephelus tukula

Mimicry of mutualistic species is seen in coral reef fish, where the models, certain cleaner fish, are greatly disadvantaged by the presence of the mimic. Cleaner fish are mutually beneficial to many other species, which allows them to eat their parasites and dead skin. Some allow the cleaner to venture inside their mouths and gill cavities to hunt these parasites. However, one species of cleaner, the bluestreak cleaner wrasse (Labroides dimidiatus), is the model of a mimic, the sabre-toothed blenny (Aspidontus taeniatus). The cleaner wrasse, shown in the image cleaning a grouper of the genus Epinephelus, resides in coral reefs in the Indian and the Pacific Oceans, and is recognised by other fishes who allow it to clean them. The blenny lives in the Indian Ocean and not only looks like the cleaner wrasse in terms of size and coloration, but even mimics the cleaner wrasse's 'dance'. Having fooled its prey into letting its guard down, the sabre-toothed blenny bites it, tearing off scales or pieces of fin. Fish grazed upon in this fashion learn to distinguish mimic from model, but because of the similarity between the two, they become much more cautious of the model as well, such that both are affected. Due to victims' ability to discriminate between foe and helper, the blennies have evolved close similarity, down to the regional level.[35] Another aggressive mimic of the cleaner wrasse, the bluestriped fangblenny, has evolved an opioid-containing venom which dulls pain and lowers blood pressure, confusing the bitten host and giving the mimic time to escape.[36]

Mimesis edit

 
The zone-tailed hawk resembles the turkey vulture in flight.

Mimesis or cryptic aggressive mimicry is where the predator mimics an organism that its prey is indifferent to. Unlike in all cases above, the predator is ignored by the prey, allowing it to avoid detection until the prey are close enough for the predator to strike. This is effectively a form of camouflage. The zone-tailed hawk (Buteo albonotatus), which resembles the turkey vulture (Cathartes aura), may provide one such example. It flies amongst them, suddenly breaking from the formation and ambushing its prey.[37] There is some controversy over whether this is a true case of mimicry.[38]

Parasites mimicking host prey edit

Just as predators such as angler fish have a structure that lures prey, so some parasites mimic their host's natural prey, but with roles reversed; the parasite gets eaten by the host. This deception provides the parasite easy entry into the host, which they can then feed upon, allowing them to continue their life cycle. Researchers may be able to predict the host of such parasites based on their appearance and behavior.[39]

One such case is a genus of mussel, Lampsilis, which feeds on the gills of fish in the larval stage of their development. Once they mature, they leave the fish as adult mollusc. Gaining entry into the host is not an easy task though, despite the fact that several hundred thousand larvae are released at once. This is especially the case in flowing water bodies such as streams, where they cannot lie on the substrate and wait to be taken up in the course of foraging. Female Lampsilis have evolved a special technique for delivering their offspring into a suitable host, however. Structures on the edge of the mantle are able to capture the interest of fish. Some resemble small fish themselves, with eye spots, a "tail" and horizontal stripes, and may even move in a similar fashion, as if facing the current (rheotaxis). When overshadowed by a fish, the larvae are forcefully expelled, becoming ecto-parasites on their unsuspecting host.[8] Some species of Lampsilis, notably Lampsilis ovata, attract fish in the genus Micropterus, Villosa has fish-like mantle lures that attract predatory fish Percina.[40]

Cercaria mirabilis, a trematode, has an especially large larval stage, a cercaria, which looks much like a small crustacean or mosquito larva. It mimics the locomotory behavior of such animals, allowing it to be eaten by predaceous fish.[39]

Another parasitic trematode example is seen in a terrestrial setting. Leucochloridium is a genus of flatworm (phylum Platyhelminthes) which matures in the intestine of songbirds. Their eggs pass out of the bird in the feces and are then taken in by Succinea, a terrestrial snail that lives in moist environments. The eggs develop into larvae inside this intermediate host, and then must find their way into the digestive system of a suitable bird. The problem here is that these birds do not eat snails, so the sporocyst must find some way of manipulating its future host into eating it. Unlike related species, these parasites are brightly colored and able to move in a pulsating manner. A sporocyst sac forces its way into the snail's eye stalks, and pulsates at high speed, enlarging the tentacle in the process.[41] It affects the host's behavior: the snail moves towards light, which it usually avoids. These combined factors make the sporocysts highly conspicuous, such that they are soon eaten by a hungry songbird. The snail then regenerates its tentacles, and Leucochloridium carries on with its life cycle.[8]

Wolf in sheep's clothing edit

 
Wolf in Sheep's Clothing by Józef Mehoffer, 1903. Detail of album cover

Zoologists have repeatedly compared aggressive mimicry to the wolf in sheep's clothing strategy of fable, including when describing jumping spiders,[2][3] lacewings,[42] ant-mimicking aphids,[43] hemipteran bugs mimicking chrysomelid beetles,[44] bird-dropping spiders,[4] orchid mantises,[4] cichlid fish,[45][46] and the zone-tailed hawk which flies with vultures, enabling it to approach terrestrial prey.[47][48] These animals have evolved to deceive their prey by appearing as other prey, or like angler fish[47] and snapping turtles[47] lure the prey by appearing as the prey's prey.

See also edit

Notes edit

  1. ^ Pasteur (1982) describes the term as redundant and states that there are many different forms of aggressive mimicry. The term was used earlier by Bates (1862) and Kirby & Spence (1823).
  2. ^ For an overview of mimicry in plants, see Wiens, 1978. Some plants mimic inanimate objects such as stones, as in Mesembryanthemum, clearly not aggressive. Some entomophilous plants such as the bee orchid attract pollinators by mimicking female insects, the males attempting to mate with the flower, but the duped insects are not eaten, and the mimicry is thus not aggressive. In Vavilovian mimicry, weeds of crops have evolved seeds similar to those of the crop, enabling the weed to be propagated by being planted as crop seed. But again, this cannot be called aggressive.[22][23]
  3. ^ The only theoretical possibilities outside this scope are a) a two species system with a model-mimic - perhaps a predator pretending to be a sleeping or dead predator (this stretches the usual scope of mimicry somewhat); and b) a cannibalistic species where a cannibalistic organism individual mimics another species.

References edit

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  43. ^ Salazar, Adrián; Fürstenau, Benjamin; Quero, Carmen; Pérez-Hidalgo, Nicolás; Carazo, Pau; Font, Enrique; Martínez-Torres, David (2015). "Aggressive mimicry coexists with mutualism in an aphid". Proceedings of the National Academy of Sciences. 112 (4): 1101–1106. Bibcode:2015PNAS..112.1101S. doi:10.1073/pnas.1414061112. PMC 4313836. PMID 25583474. The dual strategy developed by the aphid P. cimiciformis outlines a complex evolutionary scenario. On the one hand, the round morph and the ants, engaged in a trophobiotic relationship, should be subjected to the conflicts of interest typical of mutualism, with selection driving each partner to maximize its benefit by giving the least of its own energy and resources. On the other hand, the flat morph and the ants can be expected to be engaged in an arms race, with selection favoring improved deceiving abilities in the aphid and increasingly finer discrimination abilities to detect noncolony members in the ants. ... We believe that, beyond providing an unusual case of a 'wolf in sheep's clothing,' this system opens up a host of interesting and potentially novel questions about the evolution of cooperation and exploitation.
  44. ^ Jolivet, P.; Petitpierre, E.; Hsiao, T.H. (2012). Biology of Chrysomelidae. Springer. p. 276. ISBN 978-94-009-3105-3.
  45. ^ "Wolf in Sheep's Clothing: How Scale-Eating Cichlid Fish Trick Their Prey". University of Basel. 23 September 2015. Retrieved 2 February 2018. The results reveal the complexity of this so-called 'aggressive mimicry': the scale-eaters are actually imitating several blue and white striped species at once, in order to trick an entire natural community. The leader of the study, Prof. Walter Salzburger, summarizes the findings thus: 'The scale-eater pursues the strategy of a wolf that dresses up as a sheep only to then go for goats and cows.'
  46. ^ Boileau, Nicolas; Cortesi, Fabio; Egger, Bernd; Muschick, Moritz; Indermaur, Adrian; Theis, Anya; Büscher, Heinz H.; Salzburger, Walter (2015). "A complex mode of aggressive mimicry in a scale-eating cichlid fish". Biology Letters. 11 (9): 20150521. doi:10.1098/rsbl.2015.0521. PMC 4614428. PMID 26399975.
  47. ^ a b c Smith, William John (2009). The Behavior of Communicating: an ethological approach. Harvard University Press. p. 381. ISBN 978-0-674-04379-4. Others rely on the technique adopted by a wolf in sheep's clothing—they mimic a harmless species. ... Other predators even mimic their prey's prey: angler fish (Lophiiformes) and alligator snapping turtles Macroclemys temmincki can wriggle fleshy outgrowths of their fins or tongues and attract small predatory fish close to their mouths.
  48. ^ Stolesen, Scott H.; Sadoti, Giancarlo (2010). "Chapter 17: Zone-tailed Hawk (Buteo albonotatus)". In Cartron, Jean-Luc E. (ed.). Raptors of New Mexico (PDF). University of New Mexico Press. pp. 297–313. Willis (1963) postulated that the Zone-tail's strong physical resemblance to the Turkey Vulture may be a form of aggressive mimicry, which allows the hawk to closely approach potential prey that are habituated to the presence of the ubiquitous vultures (but see Mueller 1972). Snyder and Snyder (1991) report the capture success rate of Zone-tails in Arizona was significantly greater when soaring with vultures (30% successful) than when flying alone (7% successful), based on a sample of 55 observations. It is noteworthy that once a Zone-tail flying among vultures has spotted potential prey (as indicated by its locking its gaze on one spot on the ground), it often continues soaring past until well beyond the intended victim, often beyond some cover, at which point it stoops back at an acute angle in a surprise attack (Snyder and Glinski 1988; SHS).

Further reading edit

  • Wickler, W. (1968). Mimicry in Plants and Animals. McGraw-Hill. pp. 123–220. ISBN 0-07-070100-8.
  • Pietsch, T. W.; Grobecker, D. B. (1978). "The Compleat Angler: Aggressive Mimicry in an Antennariid Anglerfish". Science. 201 (4353): 369–370. Bibcode:1978Sci...201..369P. doi:10.1126/science.201.4353.369. PMID 17793734. S2CID 45349001.
  • Lloyd, J. E. (1981). "Mimicry in the sexual signals of fireflies". Scientific American. 245 (1): 110–111. Bibcode:1981SciAm.245a.138L. doi:10.1038/scientificamerican0781-138.
  • Nicolai, J. (October 1974). "Mimicry in parasitic birds". Scientific American. 231: 93–98.

External links edit

  • Feeding behavior of the frogfishes (Antennariidae) Description, images and video of aggressive mimicry in frogfish
  • Acoustic aggressive mimicry of cicadas by an Australian predatory katydid

aggressive, mimicry, form, mimicry, which, predators, parasites, parasitoids, share, similar, signals, using, harmless, model, allowing, them, avoid, being, correctly, identified, their, prey, host, zoologists, have, repeatedly, compared, this, strategy, wolf,. 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 2 3 4 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 see pseudocopulation 5 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 after George and Elizabeth Peckham 6 7 8 but it is seldom used a The humpback anglerfish uses a modified dorsal spine as a fishing rod with a bioluminescent lure to attract and capture prey 1 A camouflaged predator snow leopard in Ladakh The distinction between aggressive mimicry and predator camouflage depends on the signal given to the prey not easily determined Aggressive mimicry is opposite in principle to defensive mimicry where the mimic generally benefits from being treated as harmful The mimic may resemble its own prey or some other organism which is beneficial or at least not harmful to the prey The model i e the organism being imitated may experience increased or reduced fitness or may not be affected at all by the relationship On the other hand the signal receiver inevitably suffers from being tricked as is the case in most mimicry complexes Aggressive mimicry often involves the predator employing signals which draw its potential prey towards it a strategy which allows predators to simply sit and wait for prey to come to them The promise of food or sex are most commonly used as lures However this need not be the case as long as the predator s true identity is concealed it may be able to approach prey more easily than would otherwise be the case In terms of species involved systems may be composed of two or three species in two species systems the signal receiver or dupe is the model In terms of the visual dimension the distinction between aggressive mimicry and camouflage is not always clear Authors such as Wickler 8 have emphasized the significance of the signal to its receiver as delineating mimicry from camouflage However it is not easy to assess how significant a signal may be for the dupe and the distinction between the two can thus be rather fuzzy Mixed signals may be employed aggressive mimics often have a specific part of the body sending a deceptive signal with the rest being hidden or camouflaged Contents 1 Contrast with defensive mimicry 2 Classification 2 1 Luring prey 2 1 1 Appearance of food 2 2 Bipolar mimicry systems 2 2 1 Batesian Wallacian or prey mimicry 2 2 2 Kirbyan or brood parasite mimicry 2 3 Wicklerian Eisnerian or mimicry of harmless species 2 4 Mimicry of cleaner fish 2 5 Mimesis 2 6 Parasites mimicking host prey 3 Wolf in sheep s clothing 4 See also 5 Notes 6 References 7 Further reading 8 External linksContrast with defensive mimicry edit nbsp Defensive Batesian mimics like this bumblebee mimicking hoverfly are the antithesis of aggressive mimics Aggressive mimicry stands in semantic contrast with defensive mimicry where it is the prey that acts as a mimic with predators being duped Defensive mimicry includes the well known Batesian and Mullerian forms of mimicry where the mimic shares outward characteristics with an aposematic or harmful model In Batesian mimicry the mimic is modeled on a dangerous usually unpalatable species while in Mullerian mimicry both species are harmful and act as comimics converging on a common set of signals and sharing the burden of educating their predators Included in defensive mimicry is the lesser known Mertensian mimicry where the mimic is more harmful than the model and Vavilovian mimicry where weeds come to mimic crops through unintentional artificial selection In defensive mimicry the mimic benefits by avoiding a harmful interaction with another organism that would be more likely to take place without the deceptive signals employed Harmful interactions might involve being eaten or pulled out of the ground as a weed In contrast the aggressive mimic benefits from an interaction that would be less likely to take place without the deception at the expense of its target 9 Classification editLuring prey edit In some cases the signal receiver is lured toward the mimic This involves mimicry of a resource that is often vital to the prey s survival or more precisely the survival of its genes such as nutrition or a mate If the bait offered is of little value to prey they would not be expected to take such a risk For example in all known cases of sexual signal mimicry it is always the male sex that is deceived in fact it has been suggested that females of some species have evolved mimicry as a strategy to avoid unwanted matings 10 In these cases the predator need not move about foraging for prey but may simply stay still and allow prey to come to it Some studies suggest that the northern shrike Lanius excubitor sings in winter often imitating small passerines that may be preyed upon when lured within reach 11 There has been one report of a margay using mimicry of the cry of an infant pied tamarin to try to lure an adult tamarin within striking distance 12 Appearance of food edit nbsp The alligator snapping turtle uses its tongue to lure fish Many aggressive mimics use the promise of nourishment as a way of attracting prey The alligator snapping turtle Macrochelys temminckii is a well camouflaged ambush predator Its tongue bears a conspicuous pink extension that resembles a worm and can be wriggled around 13 fish that try to eat the worm are themselves eaten by the turtle Similarly some snakes employ caudal luring using the tail 14 15 or lingual luring using the tongue to entice small vertebrates into striking range 16 17 nbsp Argiope argentata and its web Aggressive mimicry is common amongst spiders both in luring prey and stealthily approaching predators 18 One case is the golden orb weaver Nephila clavipes which spins a conspicuous golden coloured web in well lit areas Experiments show that bees are able to associate the webs with danger when the yellow pigment is not present as occurs in less well lit areas where the web is much harder to see Other colours too were learned and avoided but bees seemed least able to effectively associate yellow pigmented webs with danger Yellow is the colour of many nectar bearing flowers however so perhaps avoiding yellow is not worthwhile Another form of mimicry is based not on colour but pattern Species such as 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 bees ability to remember web patterns Bees are able to associate a certain pattern with a spatial location meaning the spider must spin a new pattern regularly or suffer diminishing prey capture 19 nbsp The bright leaves of the Venus flytrap Dionaea muscipula attract insects in the same way as flowers Spiders can be the prey of aggressive mimics The assassin bug Stenolemus bituberus preys on spiders entering their web and plucking its silk threads until the spider approaches This vibrational aggressive mimicry matches a general pattern of vibrations which spiders treat as prey having a similar temporal structure and amplitude to leg and body movements of typical prey caught in the web 20 Larvae of the ground beetle Epomis move their mandibles one after another to lure amphibians toward them and then prey on them Their body structure allows them to bite and feed on the amphibians even when they are ingested by larger prey such as frogs 21 Although plants are better known for defensive mimicry there are exceptions For example many flowers use mimicry to attract pollinators while others may trick insects into dispersing their seeds Nonetheless most mimicry in plants b would not be classified as aggressive as although luring pollinators is similar to cases above they are certainly not eaten by the plant However some carnivorous plants may be able to increase their rate of capture through mimicry 24 For example some have patterns in the ultraviolet region of the electromagnetic spectrum much like the spider webs described above 25 Bipolar mimicry systems edit Mimicry systems involving only two species are known as bipolar 9 Only one bipolar arrangement is possible here namely where the dupe is itself the model c There are two such variants on this arrangement of mimic imitating its target in the first case termed Batesian Wallacian mimicry 9 after Henry Walter Bates 26 and Alfred Russel Wallace 27 the model is the prey species In the second case the model is the host of a brood parasite Batesian Wallacian or prey mimicry edit nbsp Mastophora spider holding its bolas with a leg attracting and capturing a male moth In some cases of Batesian Wallacian mimicry the model is a sexually receptive female which provides a strong attractive effect on males Some spiders use chemical rather than visual means to ensnare prey Female bolas spiders of the genus Mastophora lure male moth flies Diptera true flies but resembling moths by producing analogues of the moth species sex pheromones Each species of spider appears to specialize in a particular species of prey in the family Psychodidae Juveniles use their front pair of legs to capture prey such as flies Older spiders use a different strategy however swinging a sticky ball known as a bolas suspended by a silk thread at moths But both old and juvenile are able to lure prey via this olfactory signal even young spiderlings have been shown to attract prey species 28 nbsp The spotted predatory katydid Chlorobalius leucoviridis is an acoustic aggressive mimic of cicadas The listroscelidine katydid Chlorobalius leucoviridis of inland Australia is capable of attracting male cicadas of the Tribe Cicadettini by imitating the species specific reply clicks of sexually receptive female cicadas This example of acoustic aggressive mimicry is similar to the Photuris firefly case in that the predator s mimicry is remarkably versatile playback experiments show that C leucoviridis is able to attract males of many cicada species including Cicadettine cicadas from other continents even though cicada mating signals are species specific The evolution of versatile mimicry in C leucoviridis may have been facilitated by constraints on song evolution in duetting communication systems in which reply signals are recognizable only by their precise timing in relation to the male song lt lt 100 ms reply latency 29 30 Female fireflies of the genus Photuris emit the same light signals that females of the genus Photinus use as a mating signal 31 Male fireflies from several different genera are attracted to these mimics and are subsequently captured and eaten Female signals are based on that received from the male each female having a repertoire of signals matching the delay and duration of the female of the corresponding species This mimicry may have evolved from non mating signals that have become modified for predation 32 Kirbyan or brood parasite mimicry edit Further information Brood parasitism nbsp Brood parasitism four clutches of reed warbler eggs each containing one larger cuckoo egg Host parasite mimicry is a situation where a parasite mimics its own host As with mimicry of the female sex outlined previously only two species are involved the model and mimic being of the same species Brood parasitism a form of kleptoparasitism where the mother has its offspring raised by another unwitting organism is one such situation where host parasite mimicry has evolved Georges Pasteur 9 terms this form of aggressive reproductive mimicry Kirbyan mimicry after the English entomologist William Kirby 33 Wicklerian Eisnerian or mimicry of harmless species edit The prey does not have to be attracted towards the predator for the predator to benefit it is sufficient for the predator simply not to be identified as a threat Wicklerian Eisnerian mimics may resemble a mutualistic ally or a species of little significance to the prey such as a commensal 9 For example the hemipteran Arachnocoris berytoides resembles Faiditus caudatus a spider commensal of ants 34 Mimicry of cleaner fish edit Further information Cleaning symbiosis nbsp Two bluestreak cleaner wrasse cleaning a potato grouper Epinephelus tukula Mimicry of mutualistic species is seen in coral reef fish where the models certain cleaner fish are greatly disadvantaged by the presence of the mimic Cleaner fish are mutually beneficial to many other species which allows them to eat their parasites and dead skin Some allow the cleaner to venture inside their mouths and gill cavities to hunt these parasites However one species of cleaner the bluestreak cleaner wrasse Labroides dimidiatus is the model of a mimic the sabre toothed blenny Aspidontus taeniatus The cleaner wrasse shown in the image cleaning a grouper of the genus Epinephelus resides in coral reefs in the Indian and the Pacific Oceans and is recognised by other fishes who allow it to clean them The blenny lives in the Indian Ocean and not only looks like the cleaner wrasse in terms of size and coloration but even mimics the cleaner wrasse s dance Having fooled its prey into letting its guard down the sabre toothed blenny bites it tearing off scales or pieces of fin Fish grazed upon in this fashion learn to distinguish mimic from model but because of the similarity between the two they become much more cautious of the model as well such that both are affected Due to victims ability to discriminate between foe and helper the blennies have evolved close similarity down to the regional level 35 Another aggressive mimic of the cleaner wrasse the bluestriped fangblenny has evolved an opioid containing venom which dulls pain and lowers blood pressure confusing the bitten host and giving the mimic time to escape 36 Mimesis edit nbsp The zone tailed hawk resembles the turkey vulture in flight Mimesis or cryptic aggressive mimicry is where the predator mimics an organism that its prey is indifferent to Unlike in all cases above the predator is ignored by the prey allowing it to avoid detection until the prey are close enough for the predator to strike This is effectively a form of camouflage The zone tailed hawk Buteo albonotatus which resembles the turkey vulture Cathartes aura may provide one such example It flies amongst them suddenly breaking from the formation and ambushing its prey 37 There is some controversy over whether this is a true case of mimicry 38 Parasites mimicking host prey edit Further information Parasitism Just as predators such as angler fish have a structure that lures prey so some parasites mimic their host s natural prey but with roles reversed the parasite gets eaten by the host This deception provides the parasite easy entry into the host which they can then feed upon allowing them to continue their life cycle Researchers may be able to predict the host of such parasites based on their appearance and behavior 39 One such case is a genus of mussel Lampsilis which feeds on the gills of fish in the larval stage of their development Once they mature they leave the fish as adult mollusc Gaining entry into the host is not an easy task though despite the fact that several hundred thousand larvae are released at once This is especially the case in flowing water bodies such as streams where they cannot lie on the substrate and wait to be taken up in the course of foraging Female Lampsilis have evolved a special technique for delivering their offspring into a suitable host however Structures on the edge of the mantle are able to capture the interest of fish Some resemble small fish themselves with eye spots a tail and horizontal stripes and may even move in a similar fashion as if facing the current rheotaxis When overshadowed by a fish the larvae are forcefully expelled becoming ecto parasites on their unsuspecting host 8 Some species of Lampsilis notably Lampsilis ovata attract fish in the genus Micropterus Villosa has fish like mantle lures that attract predatory fish Percina 40 Cercaria mirabilis a trematode has an especially large larval stage a cercaria which looks much like a small crustacean or mosquito larva It mimics the locomotory behavior of such animals allowing it to be eaten by predaceous fish 39 Another parasitic trematode example is seen in a terrestrial setting Leucochloridium is a genus of flatworm phylum Platyhelminthes which matures in the intestine of songbirds Their eggs pass out of the bird in the feces and are then taken in by Succinea a terrestrial snail that lives in moist environments The eggs develop into larvae inside this intermediate host and then must find their way into the digestive system of a suitable bird The problem here is that these birds do not eat snails so the sporocyst must find some way of manipulating its future host into eating it Unlike related species these parasites are brightly colored and able to move in a pulsating manner A sporocyst sac forces its way into the snail s eye stalks and pulsates at high speed enlarging the tentacle in the process 41 It affects the host s behavior the snail moves towards light which it usually avoids These combined factors make the sporocysts highly conspicuous such that they are soon eaten by a hungry songbird The snail then regenerates its tentacles and Leucochloridium carries on with its life cycle 8 Wolf in sheep s clothing editFurther information Wolf in sheep s clothing nbsp Wolf in Sheep s Clothing by Jozef Mehoffer 1903 Detail of album cover Zoologists have repeatedly compared aggressive mimicry to the wolf in sheep s clothing strategy of fable including when describing jumping spiders 2 3 lacewings 42 ant mimicking aphids 43 hemipteran bugs mimicking chrysomelid beetles 44 bird dropping spiders 4 orchid mantises 4 cichlid fish 45 46 and the zone tailed hawk which flies with vultures enabling it to approach terrestrial prey 47 48 These animals have evolved to deceive their prey by appearing as other prey or like angler fish 47 and snapping turtles 47 lure the prey by appearing as the prey s prey See also editAnti predator adaptation Defensive feature of prey for selective advantage Apparent death Behavior in which animals take on the appearance of being dead Wolf in sheep s clothing Idiom of Biblical origin to describe those playing a role contrary to their real characterNotes edit Pasteur 1982 describes the term as redundant and states that there are many different forms of aggressive mimicry The term was used earlier by Bates 1862 and Kirby amp Spence 1823 For an overview of mimicry in plants see Wiens 1978 Some plants mimic inanimate objects such as stones as in Mesembryanthemum clearly not aggressive Some entomophilous plants such as the bee orchid attract pollinators by mimicking female insects the males attempting to mate with the flower but the duped insects are not eaten and the mimicry is thus not aggressive In Vavilovian mimicry weeds of crops have evolved seeds similar to those of the crop enabling the weed to be propagated by being planted as crop seed But again this cannot be called aggressive 22 23 The only theoretical possibilities outside this scope are a a two species system with a model mimic perhaps a predator pretending to be a sleeping or dead predator this stretches the usual scope of mimicry somewhat and b a cannibalistic species where a cannibalistic organism individual mimics another species References edit Haddock Steven H D Moline Mark A Case James F 2010 Bioluminescence in the Sea Annual Review of Marine Science 2 443 493 Bibcode 2010ARMS 2 443H doi 10 1146 annurev marine 120308 081028 PMID 21141672 S2CID 3872860 a b Nelson X J Jackson R R 2009 Aggressive use of Batesian mimicry by an ant like jumping spider Biology Letters 5 6 755 757 doi 10 1098 rsbl 2009 0355 PMC 2827978 PMID 19570776 Cosmophasis bitaeniata like comparable examples from insects Eisner et al 1978 Lucas amp Brodeur 2001 can be likened to a wolf in sheep s clothing e g Eisner et al 1978 These predators practise aggressive mimicry by making it easy for prey to misidentify the predator as just another member of a prey group as though lulling the prey into a false sense of security a b Heneberg Petr Perger Robert Rubio Gonzalo D 2018 A wolf in sheep s clothing The description of a fly resembling jumping spider of the genus Scoturius Simon 1901 Araneae Salticidae Huriini PLOS ONE 13 1 e0190582 Bibcode 2018PLoSO 1390582P doi 10 1371 journal pone 0190582 PMC 5783343 PMID 29364905 a b c Levine Timothy R 2014 Encyclopedia of Deception SAGE Publications p 675 ISBN 978 1 4833 8898 4 In aggressive mimicry the predator is a wolf in sheep s clothing Mimicry is used to appear harmless or even attractive to lure its prey 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 Peckham Elizabeth G 1889 Protective resemblances of spiders PDF Occasional Papers of Natural History Society of Wisconsin 1 61 113 Peckham Elizabeth G Peckham George W 1892 Ant like spiders of the family Attidae PDF Occasional Papers of Natural History Society of Wisconsin 2 1 84 a b c d Wickler Wolfgang 1968 Mimicry in plants and animals McGraw Hill 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Casewell Nicholas R et al 2017 The Evolution of Fangs Venom and Mimicry Systems in Blenny Fishes Current Biology 27 8 1184 1191 doi 10 1016 j cub 2017 02 067 hdl 1887 115463 PMID 28366739 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 Clark William S 2004 Is the Zone tailed Hawk a Mimic Birding 36 5 495 498 a b Wickler Wolfgang 1998 Mimicry Encyclopaedia Britannica 15th edition Macropaedia 24 144 151 http www britannica com eb article 11910 Haag Wendell R Warren Melvin L Jr 1999 Mantle displays of freshwater mussels elicit attacks from fish Freshwater Biology 42 35 40 doi 10 1046 j 1365 2427 1999 00454 x S2CID 50529814 See here for a photo Eisner T Hicks K Eisner M Robson D S 1978 Wolf in Sheep s Clothing Strategy of a Predaceous Insect Larva Science 199 4330 790 794 Bibcode 1978Sci 199 790E doi 10 1126 science 199 4330 790 JSTOR 1745242 PMID 17836295 S2CID 11558335 Salazar Adrian Furstenau Benjamin Quero Carmen Perez Hidalgo Nicolas Carazo Pau Font Enrique Martinez Torres David 2015 Aggressive mimicry coexists with mutualism in an aphid Proceedings of the National Academy of Sciences 112 4 1101 1106 Bibcode 2015PNAS 112 1101S doi 10 1073 pnas 1414061112 PMC 4313836 PMID 25583474 The dual strategy developed by the aphid P cimiciformis outlines a complex evolutionary scenario On the one hand the round morph and the ants engaged in a trophobiotic relationship should be subjected to the conflicts of interest typical of mutualism with selection driving each partner to maximize its benefit by giving the least of its own energy and resources On the other hand the flat morph and the ants can be expected to be engaged in an arms race with selection favoring improved deceiving abilities in the aphid and increasingly finer discrimination abilities to detect noncolony members in the ants We believe that beyond providing an unusual case of a wolf in sheep s clothing this system opens up a host of interesting and potentially novel questions about the evolution of cooperation and exploitation Jolivet P Petitpierre E Hsiao T H 2012 Biology of Chrysomelidae Springer p 276 ISBN 978 94 009 3105 3 Wolf in Sheep s Clothing How Scale Eating Cichlid Fish Trick Their Prey University of Basel 23 September 2015 Retrieved 2 February 2018 The results reveal the complexity of this so called aggressive mimicry the scale eaters are actually imitating several blue and white striped species at once in order to trick an entire natural community The leader of the study Prof Walter Salzburger summarizes the findings thus The scale eater pursues the strategy of a wolf that dresses up as a sheep only to then go for goats and cows Boileau Nicolas Cortesi Fabio Egger Bernd Muschick Moritz Indermaur Adrian Theis Anya Buscher Heinz H Salzburger Walter 2015 A complex mode of aggressive mimicry in a scale eating cichlid fish Biology Letters 11 9 20150521 doi 10 1098 rsbl 2015 0521 PMC 4614428 PMID 26399975 a b c Smith William John 2009 The Behavior of Communicating an ethological approach Harvard University Press p 381 ISBN 978 0 674 04379 4 Others rely on the technique adopted by a wolf in sheep s clothing they mimic a harmless species Other predators even mimic their prey s prey angler fish Lophiiformes and alligator snapping turtles Macroclemys temmincki can wriggle fleshy outgrowths of their fins or tongues and attract small predatory fish close to their mouths Stolesen Scott H Sadoti Giancarlo 2010 Chapter 17 Zone tailed Hawk Buteo albonotatus In Cartron Jean Luc E ed Raptors of New Mexico PDF University of New Mexico Press pp 297 313 Willis 1963 postulated that the Zone tail s strong physical resemblance to the Turkey Vulture may be a form of aggressive mimicry which allows the hawk to closely approach potential prey that are habituated to the presence of the ubiquitous vultures but see Mueller 1972 Snyder and Snyder 1991 report the capture success rate of Zone tails in Arizona was significantly greater when soaring with vultures 30 successful than when flying alone 7 successful based on a sample of 55 observations It is noteworthy that once a Zone tail flying among vultures has spotted potential prey as indicated by its locking its gaze on one spot on the ground it often continues soaring past until well beyond the intended victim often beyond some cover at which point it stoops back at an acute angle in a surprise attack Snyder and Glinski 1988 SHS Further reading editWickler W 1968 Mimicry in Plants and Animals McGraw Hill pp 123 220 ISBN 0 07 070100 8 Pietsch T W Grobecker D B 1978 The Compleat Angler Aggressive Mimicry in an Antennariid Anglerfish Science 201 4353 369 370 Bibcode 1978Sci 201 369P doi 10 1126 science 201 4353 369 PMID 17793734 S2CID 45349001 Lloyd J E 1981 Mimicry in the sexual signals of fireflies Scientific American 245 1 110 111 Bibcode 1981SciAm 245a 138L doi 10 1038 scientificamerican0781 138 Nicolai J October 1974 Mimicry in parasitic birds Scientific American 231 93 98 External links editFeeding behavior of the frogfishes Antennariidae Description images and video of aggressive mimicry in frogfish Acoustic aggressive mimicry of cicadas by an Australian predatory katydid Retrieved from https en wikipedia org w index php title Aggressive mimicry amp oldid 1215286011 Parasites mimicking host prey, wikipedia, wiki, book, books, library,

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