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212:. Some forms of warning coloration provide this distance dependent camouflage by having an effective pattern and color combination that do not allow for easy detection by a predator from a distance, but are warning-like from a close proximity, allowing for an advantageous balance between camouflage and aposematism. Warning coloration evolves in response to background, light conditions, and predator vision. Visible signals may be accompanied by odors, sounds or behavior to provide a multi-modal signal which is more effectively detected by predators.
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by predators. Concurrent reciprocal selection (CRS) may entail learning by predators or it may give rise to unlearned avoidances by them. Aposematism arising by CRS operates without special conditions of the gregariousness or the relatedness of prey, and it is not contingent upon predator sampling of prey to learn that aposematic cues are associated with unpalatability or other unprofitable features.
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384:) that lack these toxic alkaloids. Although these frogs display an extensive array of coloration and toxicity, there is very little genetic difference between the species. Evolution of their conspicuous coloration is correlated to traits such as chemical defense, dietary specialization, acoustic diversification, and increased body mass.
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avoid conspicuously coloured objects, as demonstrated using mealworms painted yellow and black to resemble wasps, with dull green controls. This implies that warning coloration works at least in part by stimulating the evolution of predators to encode the meaning of the warning signal, rather than by
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Wallace replied the next day with the suggestion that since some caterpillars "...are protected by a disagreeable taste or odour, it would be a positive advantage to them never to be mistaken for any of the palatable catterpillars , because a slight wound such as would be caused by a peck of a bird's
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will endeavor to avoid a repetition of the experience. As a consequence, aposematic species are often gregarious. Before the memory of a bad experience attenuates, the predator may have the experience reinforced through repetition. Aposematic organisms are often slow-moving, as they have little need
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molluscs are the most commonly cited examples of aposematism in marine ecosystems, but the evidence for this has been contested, mostly because (1) there are few examples of mimicry among species, (2) many species are nocturnal or cryptic, and (3) bright colours at the red end of the colour spectrum
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A theory for the evolution of aposematism posits that it arises by reciprocal selection between predators and prey, where distinctive features in prey, which could be visual or chemical, are selected by non-discriminating predators, and where, concurrently, avoidance of distinctive prey is selected
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conducted experiments with caterpillars and birds in his aviary, and in 1869 he provided the first experimental evidence for warning coloration in animals. The evolution of aposematism surprised 19th-century naturalists because the probability of its establishment in a population was presumed to be
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The function of aposematism is to prevent attack, by warning potential predators that the prey animal has defenses such as being unpalatable or poisonous. The easily detected warning is a primary defense mechanism, and the non-visible defenses are secondary. Aposematic signals are primarily visual,
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Aposematic species do not need to hide or stay still as cryptic organisms do, so aposematic individuals benefit from more freedom in exposed areas and can spend more time foraging, allowing them to find more and better quality food. They may make use of conspicuous mating displays, including vocal
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It has been proposed that aposematism and mimicry is less evident in marine invertebrates than terrestrial insects because predation is a more intense selective force for many insects, which disperse as adults rather than as larvae and have much shorter generation times. Further, there is evidence
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to form clusters tight enough to enhance the warning signal. If the species was already unpalatable, predators might learn to avoid the cluster, protecting gregarious individuals with the new aposematic trait. Gregariousness would assist predators to learn to avoid unpalatable, gregarious prey.
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Many species of bee and wasp that occur together are Müllerian mimics. Their similar coloration teaches predators that a striped pattern is associated with being stung. Therefore, a predator which has had a negative experience with any such species will likely avoid any that resemble it in the
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Further, birds recall and avoid objects that are both conspicuous and foul-tasting longer than objects that are equally foul-tasting but cryptically coloured. This suggests that
Wallace's original view, that warning coloration helped to teach predators to avoid prey thus coloured, was correct.
659:& put a difficulty before him, which he could not answer, & as on some former similar occasion, his first suggestion was, 'you had better ask Wallace'. My difficulty is, why are caterpillars sometimes so beautifully & artistically coloured?" Darwin was puzzled because his theory of
805:. If so, predators would learn to associate the colour with unpalatability from males with the trait, while heterozygous females carry the trait until it becomes common and predators understand the signal. Well-fed predators might also ignore aposematic morphs, preferring other prey species.
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terms, as it makes individuals conspicuous to predators, so they may be killed and the trait eliminated before predators learn to avoid it. If warning coloration puts the first few individuals at such a strong disadvantage, it would never last in the species long enough to become beneficial.
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Once aposematic individuals reach a certain threshold population, for whatever reason, the predator learning process would be spread out over a larger number of individuals and therefore is less likely to wipe out the trait for warning coloration completely. If the population of aposematic
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cells. However, if they are provoked, they quickly change colour, becoming bright yellow with each of the 50-60 rings flashing bright iridescent blue within a third of a second. It is often stated this is an aposematic warning display, but the hypothesis has rarely if ever been tested.
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The existence of aposematism in marine ecosystems has been debated. Many marine organisms, particularly those on coral reefs, are brightly coloured or patterned, including sponges, corals, molluscs, and fish, with little or no connection to chemical or physical defenses. Caribbean reef
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could result in aposematic males having higher reproductive success than non-aposematic males if they can survive long enough to mate. Sexual selection is strong enough to allow seemingly maladaptive traits to persist despite other factors working against the trait.
862:, a British naturalist who studied Amazonian butterflies in the second half of the 19th century. Batesian mimicry is frequency dependent: it is most effective when the ratio of mimic to model is low; otherwise, predators will encounter the mimic too often.
339:, red and black birds whose toxic feathers and skin apparently comes from the poisonous beetles they ingest, could be included. It has been proposed that aposematism played a role in human evolution, body odour carrying a warning to predators of large
56:
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individuals all originated from the same few individuals, the predator learning process would result in a stronger warning signal for surviving kin, resulting in higher inclusive fitness for the dead or injured individuals through
207:
The most common and effective colors are red, yellow, black, and white. These colors provide strong contrast with green foliage, resist changes in shadow and lighting, are highly chromatic, and provide distance dependent
2740:
Williams, B. L. (2011). "Chemical defense in pelagic octopus paralarvae: Tetrodotoxin alone does not protect individual paralarvae of the greater blue-ringed octopus (Hapalochlaena lunulata) from common reef predators".
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and non-deceptively mimic each other, to the benefit of both species, since fewer individuals of either species need to be attacked for predators to learn to avoid both of them. This form of mimicry is known as
129:, where species with strong defences evolve to resemble one another. By mimicking similarly coloured species, the warning signal to predators is shared, causing them to learn more quickly at less of a cost.
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bill almost always I believe kills a growing catterpillar. Any gaudy & conspicuous colour therefore, that would plainly distinguish them from the brown & green eatable catterpillars, would enable
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requiring each new generation to learn the signal's meaning. All of these results contradict the idea that novel, brightly coloured individuals would be more likely to be eaten or attacked by predators.
765:, in which predators avoid new prey because it is an unknown quantity; this is a long-lasting effect. Dietary conservatism has been demonstrated experimentally in some species of birds and fish.
703:
1803:(2016). "A Review of Chemical Defense in Poison Frogs (Dendrobatidae): Ecology, Pharmacokinetics, and Autoresistance". In Schulte, Bruce A.; Goodwin, Thomas E.; Ferkin, Michael H. (eds.).
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of noxious prey, because conspicuousness evolves in tandem with noxiousness. Thus, the brighter and more conspicuous the organism, the more toxic it usually is. This is in contrast to
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is a deceptive mimic of the yellowjacket wasp; it resembles the wasp, but has no sting. A predator which avoids the wasp will to some degree also avoid the moth. This is known as
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Richards, E. Loys; Alexander, Lucille G.; Snellgrove, Donna; et al. (February 2014). "Variation in the expression of dietary conservatism within and between fish species".
2521:
Pawlik, J. R.; et al. (1988). "Defensive chemicals of the
Spanish Dancer nudibranch, Hexabranchus sanguineus, and its egg ribbons: Macrolides derived from a sponge diet".
2494:
Pawlik, J. R.; et al. (1988). "Defensive chemicals of the
Spanish Dancer nudibranch, Hexabranchus sanguineus, and its egg ribbons: Macrolides derived from a sponge diet".
517:
with weak defences can gain a measure of protection from their resemblance to aposematic species. Other studies have concluded that nudibranchs such as the slugs of the family
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Rudh, A.; Rogell, B.; Håstad, O.; et al. (2011). "Rapid population divergence linked with co-variation between coloration and sexual display in strawberry poison frogs".
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892:) which has several morphs throughout its natural geographical range, each of which looks very similar to a different species of poison frog which lives in that area.
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284:), which serve to reinforce the warning. Among mammals, predators can be dissuaded when a smaller animal is aggressive and able to defend itself, as for example in
3064:"Discussion [Wallace's explanation of brilliant colors in caterpillar larvae, and others' comments thereon, presented at the ESL meeting of 4 March 1867]"
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Lim, Ayh; Chan, Izw; Carrasco, Lr; Todd, Pa (21 November 2019). "Aposematism in pink warty sea cucumbers: independent effects of chromatic and achromatic cues".
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for speed and agility. Instead, their morphology is frequently tough and resistant to injury, thereby allowing them to escape once the predator is warned off.
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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,
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663:(where females choose their mates based on how attractive they are) could not apply to caterpillars since they are immature and hence not sexually active.
1414:"Rapid color evolution in an aposematic species: A phylogenetic analysis of colour variation in the strikingly polymorphic strawberry poison-dart frog"
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The second head (Sematic
Colours) includes Warning Colours and Recognition Markings: the former warn an enemy off, and are therefore called Aposematic
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1365:"Aposematism and Crypsis Combined as a Result of Distance Dependence: Functional Versatility of the Colour Pattern in the Swallowtail Butterfly Larva"
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Aposematism is a sufficiently successful strategy to have had significant effects on the evolution of both aposematic and non-aposematic species.
510:), among the largest of tropical marine slugs, potently chemically defended, and brilliantly red and white, is nocturnal and has no known mimics.
559:, chemicals which could function as an effective defence; this evidence is argued to be sufficient for such species to be considered aposematic.
350:). These neotropical anuran amphibians exhibit a wide spectrum of coloration and toxicity. Some species in this poison frog family (particularly
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advertise their foul-smelling chemical defences with sharply contrasting black-and-white patterns on their fur, while the similarly-patterned
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defensive animal spines are often conspicuous (shape and colour) and should be considered aposematic... Classic examples are the starfishes
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are aposematic, and that the chromatic and achromatic signals that they provide to predators both independently reduce the rate of attack.
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has been implicated in the coloration of some
Mediterranean nudibranchs, all of which derive defensive chemicals from their sponge diet.
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are brightly coloured, and many species are full of toxic chemicals, but there is no statistical relationship between the two factors.
436:, may similarly be acoustic aposematism, connected to and preceded by chemical defences. Similar acoustic defences exist in a range of
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A genuine aposematic signal that a species actually possesses chemical or physical defences is not the only way to deter predators. In
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may adapt to visual cues more rapidly than do birds, making aposematism less effective. However, there is experimental evidence that
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2302:
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Schmidt, J. O.; Blum, M. S. (1977). "Adaptations and
Responses of Dasymutilla occidentalis (Hymenoptera: Mutillidae) to Predators".
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are venomous. They spend much of their time hiding in crevices whilst displaying effective camouflage patterns with their dermal
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with contrasting patterns may also be aposematic, at least in females; but since male birds are often brightly coloured through
204:, which attempt to startle a predator with a threatening appearance but which are bluffing, unsupported by any strong defences.
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136:, a mimicking species resembles an aposematic model closely enough to share the protection, while many species have bluffing
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Rubino, Darrin L.; McCarthy, Brian C. "Presence of
Aposematic (Warning) Coloration in Vascular Plants of Southeastern Ohio"
3895:
Lee, T. J.; Marples, N. M.; Speed, M. P. (2010). "Can dietary conservatism explain the primary evolution of aposematism?".
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Maan, M. E.; Cummings, M. E. (2012). "Poison frog colors are honest signals of toxicity, particularly for bird predators".
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311:, advertising its powerful defences, scent glands near the tail, by raising its tail and displaying its warning coloration
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570:, provides both chromatic and achromatic signals to predators, and both of these independently have an aposematic effect.
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4084:"Mimicry on the edge: why do mimics vary in resemblance to their model in different parts of their geographical range?"
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94:, foul taste or smell, sharp spines, or aggressive nature. These advertising signals may take the form of conspicuous
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4146:(1878). "Ueber die Vortheile der Mimicry bei Schmetterlingen" [On the Advantages of Mimicry in Butterflies].
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Götmark, Frank (1994). "Are Bright Birds
Distasteful? A Re-Analysis of H. B. Cott's Data on the Edibility of Birds".
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characteristics. Aposematic signals are beneficial for both predator and prey, since both avoid potential harm.
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3407:"The expression of dietary conservatism in solitary and shoaling 3-spined sticklebacks Gasterosteus aculeatus"
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Miller, A. M.; Pawlik, J. R. (2013). "Do coral reef fish learn to avoid unpalatable prey using visual cues?".
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Haber, M.; et al. (2010). "Coloration and defense in the nudibranch gastropod
Hypselodoris fontandraui".
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Hristov, N. I.; Conner, W. E. (2005). "Sound strategy: acoustic aposematism in the bat–tiger moth arms race".
1269:
Sendova-Franks, Ana; Scott, Michelle
Pellissier (2015). "Featured Articles in This Month's Animal Behaviour".
3935:"The effects of predator learning, forgetting, and recognition errors on the evolution of warning coloration"
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Mappes, Johanna; Marples, Nicola; Endler, John A. (2005). "The complex business of survival by aposematism".
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noises which warn bats to avoid them, or by warning postures which expose brightly coloured body parts (see
423:, and their coloration is not correlated with edibility, it is unclear whether aposematism is significant.
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Merilaita, Sami; Kaitala, Veijo (2002). "Community structure and the evolution of aposematic coloration".
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advertise their sharp claws, powerful jaws, and aggressive natures. Some brightly coloured birds such as
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make them dangerous or painful to attack. Tiger moths advertise their unpalatability by either producing
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3658:"Aposematism and gregariousness: the combined effect of group size and coloration on signal repellence"
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are rapidly attenuated as a function of water depth. For example, the Spanish Dancer nudibranch (genus
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of unpalatable chemicals or physical defences such as prickled leaves or thorns. Many insects, such as
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2786:"The evolution of conspicuous facultative mimicry in octopuses: an example of secondary adaptation?"
1900:"Phylogenomic Reconstruction of the Neotropical Poison Frogs (Dendrobatidae) and Their Conservation"
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3277:; et al. (2003). "Dietary conservatism may facilitate the initial evolution of aposematism".
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Wallace coined the term "warning colours" in an article about animal coloration in 1877. In 1890
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is an acoustic form of aposematism. Sound production by the caterpillar of the Polyphemus moth,
1067:, pp. Foldout "The Colours of Animals Classified According to Their Uses", after page 339.
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3049:"Letter from Alfred Russel Wallace to Charles Robert Darwin dated 24 February [1867]"
3034:"Letter from Charles Robert Darwin to Alfred Russel Wallace dated 23 February [1867]"
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which may startle a predator long enough to enable an otherwise undefended prey to escape.
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The Colours of Animals, their meaning and use, especially considered in the case of insects
3580:
Speed, Michael P. (2001). "Can receiver psychology explain the evolution of aposematism?".
2200:"Clicking caterpillars: acoustic aposematism in Antheraea polyphemus and other Bombycoidea"
1680:"Sexual dimorphism and directional sexual selection on aposematic signals in a poison frog"
1644:"Are we chemically aposematic? Revisiting L. S. B. Leakey's hypothesis on human body odour"
1572:"Black, White and Stinky: Explaining Coloration in Skunks and Other Boldly Colored Animals"
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Aposematism could also be favoured in dense populations even if these are not gregarious.
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3358:"Predator experience on cryptic prey affects the survival of conspicuous aposematic prey"
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2939:"The dual benefits of aposematism: predator avoidance and enhanced resources collection"
2886:"The dual benefits of aposematism: predator avoidance and enhanced resources collection"
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among all living species. Within the same family, there are also cryptic frogs (such as
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Briolat, Emmanuelle S.; Burdfield-Steel, Emily R.; Paul, Sarah C.; et al. (2019).
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786:) long enough for them to become established, but this is likely to be only temporary.
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Santos, Juan C.; Baquero, Margarita; Barrio-Amorós, César; et al. (2014-12-07).
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Higashi, Masahiko; Yachi, Shigeo (August 1998). "The evolution of warning signals".
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1942:"Multiple, recurring origins of aposematism and diet specialization in poison frogs"
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1742:"The evolution of coloration and toxicity in the poison frog family (Dendrobatidae)"
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1514:
1290:
959:"Multiple, recurring origins of aposematism and diet specialization in poison frogs"
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Perhaps the most numerous aposematic vertebrates are the poison dart frogs (family:
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using bright colors and high-contrast patterns such as stripes. Warning signals are
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1941:
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717:, 1890, introduced a set of new terms for animal coloration including "aposematic".
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4051:"Contributions to an insect fauna of the Amazon valley (Lepidoptera: Heliconidae)"
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3451:
2675:
2434:"Defenses of Caribbean sponges against predatory reef fish: I. Chemical deterrenc"
1282:
4279:
Avoiding Attack: The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry
4022:"Contributions to an insect fauna of the Amazon valley. Lepidoptera: Heliconidae"
2699:"How does the blue-ringed octopus (Hapalochlaena lunulata) flash its blue rings?"
1898:
Guillory, Wilson X.; Muell, Morgan R.; Summers, Kyle; et al. (August 2019).
1812:
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future. Müllerian mimicry is found in vertebrates such as the mimic poison frog (
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Other explanations are possible. Predators might innately fear unfamiliar forms (
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3475:"Better to be bimodal: the interaction of color and odor on learning and memory"
2550:"Marine benthic invertebrates use multimodal cues for defense against reef fish"
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2711:(21). Bell, G. R., Kuzirian, A. M., Allen, J. J. and Hanlon, R. T.: 3752–3757.
1851:"Aposematism increases acoustic diversification and speciation in poison frogs"
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Proceedings of the National Academy of Sciences of the United States of America
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Richards, E. Loys; Thomas, Robert J.; Marples, Nicola M.; et al. (2011).
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Unpalatability, broadly understood, can be created in a variety of ways. Some
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3715:
3698:
3625:"Avoidance of aposematic prey in European tits (Paridae): learned or innate?"
865:
A second form of mimicry occurs when two aposematic organisms share the same
399:
caterpillars, acquire toxic chemicals from their host plants. Among mammals,
4831:
4664:
4620:
4600:
3641:
3624:
3493:
3474:
3423:
3406:
1991:
1704:
1620:
1039:
1012:
975:
850:
the conspicuous markings of their aposematic counterparts. For example, the
843:
798:
783:
749:
553:
552:, has conspicuous coloration and conspicuous long, sharp spines, as well as
416:
392:
324:
235:
87:
4204:
4117:
4099:
3960:
3836:
3817:
3772:
3754:
3673:
3601:
3593:
3561:
3391:
3373:
3339:
3250:
3208:
3007:
2964:
2911:
2726:
2610:
2419:
Antipredatory defensive roles of natural products from marine invertebrates
2367:
2284:
2227:
2184:
2165:
2090:
2072:
2010:
1967:"Phenotypic integration emerges from aposematism and scale in poison frogs"
1884:
1866:
1785:
1766:
1723:
1506:
1441:
1398:
1380:
1349:
1331:
1255:
1237:
1194:
994:
4307:
3739:"Aggregation, defense, and warning signals: the evolutionary relationship"
2849:"On the perception, production and function of blue coloration in animals"
2602:
2473:
Edmunds, Malcolm (1991). "Does warning coloration occur in nudibranchs?".
1628:
1048:
606:
of the would-be predator; a bird that has once experienced a foul-tasting
55:
4961:
4698:
2052:
576:
369:
231:
48:
44:
4852:
3356:
Lindstrom, Leena; Altalo, Rauno V.; Lyytinen, Anne; et al. (2001).
2241:
Bura, Veronica L.; Kawahara, Akito Y.; Yack, Jayne E. (11 August 2016).
695:
low, since a conspicuous signal suggested a higher chance of predation.
5215:
5173:
5016:
5008:
4971:
4760:
4733:
4460:
4450:
4318:
2717:
2698:
2574:
2549:
2458:
2433:
2218:
2199:
2125:
2028:
847:
834:
769:
However, some birds (inexperienced starlings and domestic chicks) also
556:
340:
336:
320:
3699:"Effects of novelty and gregariousness in survival of aposematic prey"
3189:
2865:
2848:
2648:
2266:
2243:"A Comparative Analysis of Sonic Defences in Bombycoidea Caterpillars"
2198:
Brown, Sarah G.; Boettner, George H.; Yack, Jayne E. (15 March 2007).
2061:
Philosophical Transactions of the Royal Society B: Biological Sciences
1926:
1916:
1899:
1186:
1177:
808:
A further explanation is that females might prefer brighter males, so
453:
3534:"Perspective: The Evolution of Warning Coloration is Not Paradoxical"
3314:"Perspective: The evolution of warning coloration is not paradoxical"
1807:. Vol. 13. Springer International Publishing. pp. 305–337.
1593:
Dumbacher, J. P.; Beehler, B. M.; Spande, T. F.; et al. (1992).
770:
603:
494:
408:
332:
280:
both have bright colors and produce audible noises when grabbed (via
227:
4476:
2117:
1595:"Homobatrachotoxin in the genus Pitohui: chemical defense in birds?"
3532:
Marples, Nicola M.; Kelly, David J.; Thomas, Robert J. (May 2005).
1168:
4786:
3135:
2784:
Huffard, C. L.; Saarman, N.; Hamilton, H.; Simison, W. B. (2010).
702:
626:
561:
527:
400:
388:
300:
239:
214:
103:
99:
91:
54:
31:
3623:
Exernova, Alice; Stys, Pavel; Fucikova, Eva; et al. (2007).
3171:"Diversity in warning coloration: selective paradox or the norm?"
2749:(3). Lovenburg, V., Huffard, C. L. and Caldwell, R. L.: 131–141.
1455:
MacAuslane, Heather J. (2008). "Aposematism". In Capinera (ed.).
368:) are conspicuously coloured and sequester one of the most toxic
75:
through its aggressive temperament and its sharp teeth and claws.
1316:"Linking the evolution and form of warning coloration in nature"
904:
A model (to be mimicked), the venomous and genuinely aposematic
686:
Since Darwin was enthusiastic about the idea, Wallace asked the
328:
4981:
4856:
4480:
4322:
3312:
Marples, Nicola M.; Kelly, David J.; Thomas, Robert J. (2005).
3084:
The Heretic in Darwin's Court:The Life of Alfred Russel Wallace
4582:
2630:
2628:
1117:"The Colours of Animals and Plants. I.—The Colours of Animals"
3068:
Journal of Proceedings of the Entomological Society of London
2819:
A Review of Blue-ringed Octopus Conservation (Masters thesis)
1222:"Warning displays may function as honest signals of toxicity"
532:
Iridescent blue rings on the mantles of the venomous octopus
797:
Another possibility is that a gene for aposematism might be
459:
There is evidence that a range of marine animals, including
521:
from Indo-Pacific coral reefs are aposematically coloured.
4977:
3791:"Maternal effects and the evolution of aposematic signals"
2417:
Pawlik, J. R. (2012). Fattorusso, E.; et al. (eds.).
957:
Santos, J. C.; Coloma, Luis A.; Cannatella, D. C. (2003).
86:
by an animal, whether terrestrial or marine, to potential
2333:"Conspicuous and aposematic spines in the animal kingdom"
2139:
Meik, Jesse M.; Pires-daSilva, André (10 February 2009).
170:'sign', referring to signs that warn other animals away.
148:
The term aposematism was coined by the English zoologist
878:, a German naturalist who studied the phenomenon in the
4088:
Proceedings of the Royal Society B: Biological Sciences
3662:
Proceedings of the Royal Society B: Biological Sciences
1855:
Proceedings of the Royal Society B: Biological Sciences
1320:
Proceedings of the Royal Society B: Biological Sciences
3974:
Weldon, P.J. (December 2013). "Chemical aposematism".
1470:
1468:
1013:"Toxicity, Odor Aversion, and 'Olfactory Aposematism'"
735:. He described the derivation of the term as follows:
690:
to test the hypothesis. In response, the entomologist
675:
a kind not fit for food, & thus they would escape
242:
produces a noxious odor, and the poison glands of the
315:
Aposematism is widespread in insects, but less so in
4178:
Proclamations of the Entomological Society of London
3656:
Gamberale, Gabriella; Tullberg, Birgitta S. (1998).
1965:
Santos, Juan C.; Cannatella, David C. (2011-04-12).
1844:
1842:
1840:
1150:
1148:
5147:
5109:
5015:
4970:
4890:
4779:
4691:
4663:
4581:
4512:
4404:
4356:
3473:Siddall, Emma C.; Marples, Nicola M. (2008-01-22).
1363:Tullberg, B. S.; Merilaita S.; Wiklund, C. (2005).
1139:
4230:
2523:Journal of Experimental Marine Biology and Ecology
2496:Journal of Experimental Marine Biology and Ecology
2141:"Evolutionary morphology of the rattlesnake style"
1673:
1671:
1574:. University of Massachusetts Amherst. 27 May 2011
789:Alternatively, prey animals might be sufficiently
27:Honest signalling of an animal's powerful defences
3789:Brodie, Edmund D. III; Agrawal, Anell F. (2001).
274:. Velvet ants (actually parasitic wasps) such as
4176:; a remarkable case of mimicry in butterflies".
3351:
3349:
3087:. New York: Columbia University Press. pp.
4082:Harper, G. R.; Pfennig, D. W (22 August 2007).
3796:Proceedings of the National Academy of Sciences
2057:"Contrasting coloration in terrestrial mammals"
2027:, Vol. 131, No. 3 (Jul-Sep 2004), pp. 252-256.
1971:Proceedings of the National Academy of Sciences
1746:Proceedings of the National Academy of Sciences
1302:
1300:
963:Proceedings of the National Academy of Sciences
737:
319:, being mostly confined to a smaller number of
2412:
2410:
1107:
1105:
331:species, and some foul-smelling or aggressive
4993:
4868:
4492:
4334:
3784:
3782:
3268:
3266:
3264:
3262:
3260:
3222:
3220:
3218:
1740:Summers, Kyle; Clough, Mark E. (2001-05-22).
1735:
1733:
761:There is evidence for explanations involving
8:
3575:
3573:
3571:
3521:. Reed Business Information. pp. 50–52.
3506:
3504:
238:contain bitter-tasting chemicals, while the
3697:Mappes, Johanna; Alatalo, Rauno V. (1996).
5000:
4986:
4978:
4875:
4861:
4853:
4499:
4485:
4477:
4341:
4327:
4319:
391:are thought to employ aposematism to warn
5194:Coloration evidence for natural selection
4364:Coloration evidence for natural selection
4107:
4066:
4055:Biological Journal of the Linnean Society
3950:
3826:
3816:
3762:
3714:
3681:
3640:
3551:
3492:
3422:
3381:
3329:
3198:
3188:
2954:
2901:
2864:
2801:
2790:Biological Journal of the Linnean Society
2716:
2573:
2548:Ritson-Williams, R.; Paul, V. J. (2007).
2457:
2274:
2217:
2174:
2164:
2080:
2000:
1990:
1925:
1915:
1874:
1775:
1765:
1713:
1703:
1659:
1648:Biological Journal of the Linnean Society
1431:
1388:
1339:
1245:
1176:
1038:
984:
974:
2832:Hanlon, R. T.; Messenger, J. B. (1998).
2331:Inbar, Moshe; Lev-Yadun, Simcha (2005).
2326:
2324:
2296:
2294:
1216:Blount, Jonathan D.; Speed, Michael P.;
615:signals, which may then develop through
538:are considered by some to be aposematic.
485:spines may warn of strong toxins within.
343:able to defend themselves with weapons.
177:
4130:
3156:
2039:
2025:Journal of the Torrey Botanical Society
1530:Entomologia Experimentalis et Applicata
1077:
1075:
1073:
1064:
949:
894:
602:The mechanism of defence relies on the
655:wrote, "On Monday evening I called on
4683:Evolution of color vision in primates
1799:Santos, Juan C.; Tarvin, Rebecca D.;
1678:Maan, M. E.; Cummings, M. E. (2009).
7:
2029:https://www.jstor.org/stable/4126955
1412:Wang, I. J.; Shaffer, H. B. (2008).
4195:Twomey, Evan; Brown, Jason (1986).
4027:Transactions of the Linnean Society
2704:The Journal of Experimental Biology
2432:Pawlik, J. R.; et al. (1995).
2205:The Journal of Experimental Biology
631:Gregarious nymphs of an aposematic
222:, an aposematically coloured beetle
5134:Pouyannian (with pseudocopulation)
4184:. Translated by R. Meldola: 20–29.
4068:10.1111/j.1095-8312.1981.tb01842.x
4040:10.1111/j.1096-3642.1860.tb00146.x
3952:10.1111/j.0014-3820.2000.tb00077.x
3553:10.1111/j.0014-3820.2005.tb01032.x
3331:10.1111/j.0014-3820.2005.tb01032.x
1805:Chemical Signals in Vertebrates 13
1642:Weldon, Paul J. (18 August 2018).
1550:10.1111/j.1570-7458.1977.tb02663.x
1226:Proceedings of the Royal Society B
842:Non-aposematic species have often
25:
3513:"All things bright and poisonous"
2933:Speed, M. P.; Brockhurst, M. A.;
2880:Speed, M. P.; Brockhurst, M. A.;
1140:Ruxton, Sherratt & Speed 2004
5260:
5259:
5100:
5035:Aristotelian/Distraction display
4572:
4306:
4263:Kegan Paul, Trench & Trübner
3874:10.1046/j.1461-0248.2002.00362.x
3743:Proceedings of the Royal Society
3362:Proceedings of the Royal Society
3299:10.1034/j.1600-0706.2003.12061.x
3000:10.1111/j.1558-5646.2010.01210.x
2956:10.1111/j.1558-5646.2009.00931.x
2903:10.1111/j.1558-5646.2009.00931.x
2803:10.1111/j.1095-8312.2010.01484.x
2305:. Shedd Aquarium. Archived from
1461:. Vol. 4. pp. 239–242.
1433:10.1111/j.1558-5646.2008.00507.x
1369:Proceedings: Biological Sciences
912:
897:
474:
452:
3273:Thomas, R. J.; Marples, N. M.;
3230:Trends in Ecology and Evolution
688:Entomological Society of London
113:The term was coined in 1877 by
2637:Marine Ecology Progress Series
2554:Marine Ecology Progress Series
2438:Marine Ecology Progress Series
748:Aposematism is paradoxical in
546:, like other starfish such as
426:The sound-producing rattle of
125:. Aposematism is exploited in
1:
5226:Frequency-dependent selection
3909:10.1016/j.anbehav.2009.10.004
3511:Roper, Tim J. (9 July 1987).
3452:10.1016/j.anbehav.2013.11.009
2836:. Cambridge University Press.
2676:10.1016/j.anbehav.2012.11.002
2421:. Springer. pp. 677–710.
1283:10.1016/j.anbehav.2014.12.013
566:The pink warty sea cucumber,
513:Mimicry is to be expected as
2535:10.1016/0022-0981(88)90225-0
2508:10.1016/0022-0981(88)90225-0
1813:10.1007/978-3-319-22026-0_21
671:to recognise them easily as
575:that fish predators such as
4611:Simple eye in invertebrates
1086:. Apheloria. Archived from
882:in the late 19th century.
158:. He based the term on the
36:The bright colours of this
5338:
4807:Infrared sensing in snakes
3243:10.1016/j.tree.2005.07.011
2402:, which have red spines...
2303:"Crown of Thorns Sea Star"
832:
188:
5254:
5098:
4678:Evolution of color vision
4570:
4229:Edmunds, Malcolm (1974).
3996:10.1007/s00049-013-0140-3
2763:10.1007/s00049-011-0075-5
2360:10.1007/s00114-005-0608-2
1661:10.1093/biolinnean/bly109
1499:10.1007/s00114-005-0611-7
1458:Encyclopedia Entomologica
679:which is as bad as being
297:In terrestrial ecosystems
185:colours warn of toxicity
5302:Antipredator adaptations
5155:Anti-predator adaptation
4417:Anti-predator adaptation
3047:Wallace, Alfred Russel.
2145:BMC Evolutionary Biology
2106:Journal of Avian Biology
867:anti-predator adaptation
651:dated 23 February 1867,
581:pink warty sea cucumbers
544:crown-of-thorns starfish
483:crown-of-thorns starfish
277:Dasymutilla occidentalis
73:ability to defend itself
4391:Paradox of the plankton
4283:Oxford University Press
4277:; Speed, M. P. (2004).
4253:Poulton, Edward Bagnall
2817:Lambert, W. A. (2011).
2697:Mäthger, L. M. (2012).
2301:Shedd, John G. (2006).
1992:10.1073/pnas.1010952108
1705:10.1073/pnas.0903327106
1621:10.1126/science.1439786
1040:10.1126/science.7244647
1011:; Grant, R. P. (1981).
976:10.1073/pnas.2133521100
481:Conspicuous colours of
5307:Evolution by phenotype
4898:Advertising in biology
4100:10.1098/rspb.2007.0558
3818:10.1073/pnas.141075998
3755:10.1098/rspb.2006.3570
3716:10.1093/beheco/8.2.174
3674:10.1098/rspb.1998.0374
3594:10.1006/anbe.2000.1558
3374:10.1098/rspb.2000.1377
3081:Slotten, Ross (2004).
2847:Umbers, K. D. (2013).
2166:10.1186/1471-2148-9-35
2073:10.1098/rstb.2008.0221
1867:10.1098/rspb.2014.1761
1767:10.1073/pnas.101134898
1381:10.1098/rspb.2005.3079
1332:10.1098/rspb.2011.1932
1238:10.1098/rspb.2008.1407
1220:; et al. (2009).
1113:Wallace, Alfred Russel
778:Alternative hypotheses
757:Supported explanations
741:
732:The Colours of Animals
723:Edward Bagnall Poulton
718:
714:The Colours of Animals
709:Edward Bagnall Poulton
639:
571:
539:
535:Hapalochlaena lunulata
312:
223:
186:
155:The Colours of Animals
150:Edward Bagnall Poulton
115:Edward Bagnall Poulton
76:
67:makes it conspicuous,
65:reverse countershading
52:
5246:Underwater camouflage
5025:Aggressive/Wicklerian
4884:Signalling in biology
4812:Monocular deprivation
4771:Underwater camouflage
4766:Structural coloration
4744:Disruptive coloration
4357:Patterns of evolution
4197:"Ranitomeya imitator"
4148:Zoologischer Anzeiger
3642:10.1093/beheco/arl061
3494:10.1093/beheco/arm155
3424:10.1093/beheco/arr047
2603:10.1086/BBLv218n2p181
833:Further information:
706:
649:Alfred Russel Wallace
630:
623:Origins of the theory
587:Blue-ringed octopuses
565:
531:
304:
218:
189:Further information:
183:Flamboyant cuttlefish
181:
119:Alfred Russel Wallace
58:
35:
5297:Animal communication
5221:Evolutionary ecology
5206:Deception in animals
5200:Dazzled and Deceived
5160:Animal communication
4792:Blindness in animals
4724:Counter-illumination
4673:Evolution of the eye
4369:Convergent evolution
4350:Evolutionary ecology
4315:at Wikimedia Commons
2834:Cephalopod Behaviour
1801:O'Connell, Lauren A.
1122:Macmillan's Magazine
763:dietary conservatism
725:renamed the concept
444:In marine ecosystems
433:Antheraea polyphemus
38:granular poison frog
5067:Emsleyan/Mertensian
4947:Distraction display
4903:Agonistic behaviour
4442:Distraction display
4386:Divergent evolution
4094:(1621): 1955–1961.
4047:Bates, Henry Walter
4018:Bates, Henry Walter
3988:2013Checo..23..201W
3866:2002EcolL...5..495M
3809:2001PNAS...98.7884B
3749:(1600): 2417–2424.
3735:Sherratt, Thomas N.
3291:2003Oikos.101..458T
3159:, pp. 337–338.
3128:1998Natur.394..882Y
2821:. Prescott College.
2755:2011Checo..21..131W
2591:Biological Bulletin
2566:2007MEPS..340...29R
2450:1995MEPS..127..183P
2352:2005NW.....92..170I
2340:Naturwissenschaften
2309:on 22 February 2014
2259:2016NatSR...631469B
2157:2009BMCEE...9...35M
2042:, pp. 199–201.
1983:2011PNAS..108.6175S
1758:2001PNAS...98.6227S
1696:2009PNAS..10619072M
1690:(45): 19072–19077.
1613:1992Sci...258..799D
1542:1977EEApp..21...99S
1491:2005NW.....92..164H
1478:Naturwissenschaften
1375:(1570): 1315–1321.
1157:American Naturalist
1031:1981Sci...213..476E
969:(22): 12792–12797.
889:Ranitomeya imitator
801:and located on the
69:honestly signalling
5312:Warning coloration
5211:Deimatic behaviour
4952:Handicap principle
4942:Deimatic behaviour
4739:Deimatic behaviour
4437:Deimatic behaviour
4396:Predator satiation
4381:Parallel evolution
4313:Warning coloration
4233:Defence in Animals
4207:on 25 October 2016
3703:Behavioral Ecology
3629:Behavioral Ecology
3480:Behavioral Ecology
3411:Behavioral Ecology
3176:Biological Reviews
2853:Journal of Zoology
2718:10.1242/jeb.076869
2575:10.3354/meps340029
2459:10.3354/meps127183
2400:Metrodira subulata
2396:Acanthaster planci
2247:Scientific Reports
2219:10.1242/jeb.001990
1861:(1796): 20141761.
1142:, pp. 82–103.
939:Handicap principle
927:of the coral snake
860:Henry Walter Bates
719:
640:
572:
549:Metrodira subulata
540:
466:Phyllidia varicosa
313:
260:black widow spider
224:
198:honest indications
187:
123:warning coloration
77:
53:
5292:Signalling theory
5279:
5278:
5273:
5272:
5241:Signalling theory
5216:Mimicry#Evolution
5189:Community ecology
5184:Animal coloration
5030:Ant/Myrmecomorphy
4935:Courtship display
4850:
4849:
4842:Visual perception
4837:Underwater vision
4802:Feature detection
4797:Eyespot apparatus
4756:Eyespot (mimicry)
4704:Animal coloration
4507:Vision in animals
4474:
4473:
4412:Signalling theory
4311:Media related to
4292:978-0-19-852859-3
4271:Ruxton, Graeme D.
4244:978-0-582-44132-3
3803:(14): 7884–7887.
3731:Ruxton, Graeme D.
3668:(1399): 889–894.
3368:(1465): 357–361.
3275:Cuthill, Innes C.
3190:10.1111/brv.12460
3122:(6696): 882–884.
3098:978-0-231-13010-3
2866:10.1111/jzo.12001
2649:10.3354/meps13159
2267:10.1038/srep31469
2067:(1516): 537–548.
1977:(15): 6175–6180.
1917:10.3390/d11080126
1822:978-3-319-22026-0
1752:(11): 6227–6232.
1607:(5083): 799–801.
1426:(11): 2742–2759.
1326:(1728): 417–426.
1232:(1658): 871–877.
1218:Ruxton, Graeme D.
872:Müllerian mimicry
707:First edition of
568:Cercodemas anceps
523:Müllerian mimicry
469:, are aposematic.
308:Mephitis mephitis
202:deimatic displays
191:signalling theory
174:Defense mechanism
152:in his 1890 book
138:deimatic displays
127:Müllerian mimicry
16:(Redirected from
5329:
5322:Chemical ecology
5265:Category mimicry
5263:
5262:
5104:
5002:
4995:
4988:
4979:
4877:
4870:
4863:
4854:
4822:Palpebral (bone)
4655:Schizochroal eye
4576:
4501:
4494:
4487:
4478:
4343:
4336:
4329:
4320:
4310:
4296:
4266:
4248:
4236:
4217:
4216:
4214:
4212:
4203:. Archived from
4192:
4186:
4185:
4162:
4156:
4155:
4140:
4134:
4128:
4122:
4121:
4111:
4079:
4073:
4072:
4070:
4043:
4014:
4008:
4007:
3971:
3965:
3964:
3954:
3927:
3921:
3920:
3897:Animal Behaviour
3892:
3886:
3885:
3847:
3841:
3840:
3830:
3820:
3786:
3777:
3776:
3766:
3727:
3721:
3720:
3718:
3694:
3688:
3687:
3685:
3653:
3647:
3646:
3644:
3620:
3614:
3613:
3582:Animal Behaviour
3577:
3566:
3565:
3555:
3529:
3523:
3522:
3508:
3499:
3498:
3496:
3470:
3464:
3463:
3440:Animal Behaviour
3435:
3429:
3428:
3426:
3402:
3396:
3395:
3385:
3353:
3344:
3343:
3333:
3309:
3303:
3302:
3270:
3255:
3254:
3224:
3213:
3212:
3202:
3192:
3166:
3160:
3154:
3148:
3147:
3109:
3103:
3102:
3078:
3072:
3071:
3059:
3053:
3052:
3044:
3038:
3037:
3026:
3020:
3019:
2994:(5): 1271–1282.
2983:
2977:
2976:
2958:
2949:(6): 1622–1633.
2930:
2924:
2923:
2905:
2896:(6): 1622–1633.
2877:
2871:
2870:
2868:
2844:
2838:
2837:
2829:
2823:
2822:
2814:
2808:
2807:
2805:
2781:
2775:
2774:
2737:
2731:
2730:
2720:
2694:
2688:
2687:
2664:Animal Behaviour
2659:
2653:
2652:
2632:
2623:
2622:
2586:
2580:
2579:
2577:
2545:
2539:
2538:
2518:
2512:
2511:
2491:
2485:
2484:
2470:
2464:
2463:
2461:
2429:
2423:
2422:
2414:
2405:
2404:
2391:
2390:
2384:
2378:. Archived from
2337:
2328:
2319:
2318:
2316:
2314:
2298:
2289:
2288:
2278:
2238:
2232:
2231:
2221:
2195:
2189:
2188:
2178:
2168:
2136:
2130:
2129:
2101:
2095:
2094:
2084:
2049:
2043:
2037:
2031:
2021:
2015:
2014:
2004:
1994:
1962:
1956:
1955:
1953:
1952:
1938:
1932:
1931:
1929:
1919:
1895:
1889:
1888:
1878:
1846:
1835:
1834:
1796:
1790:
1789:
1779:
1769:
1737:
1728:
1727:
1717:
1707:
1675:
1666:
1665:
1663:
1639:
1633:
1632:
1590:
1584:
1583:
1581:
1579:
1568:
1562:
1561:
1525:
1519:
1518:
1472:
1463:
1462:
1452:
1446:
1445:
1435:
1409:
1403:
1402:
1392:
1360:
1354:
1353:
1343:
1304:
1295:
1294:
1271:Animal Behaviour
1266:
1260:
1259:
1249:
1213:
1207:
1206:
1180:
1152:
1143:
1137:
1131:
1130:
1109:
1100:
1099:
1097:
1095:
1079:
1068:
1062:
1053:
1052:
1042:
1005:
999:
998:
988:
978:
954:
916:
901:
856:Batesian mimicry
810:sexual selection
692:John Jenner Weir
661:sexual selection
617:sexual selection
478:
456:
421:sexual selection
244:poison dart frog
166:'away' and σῆμα
134:Batesian mimicry
21:
5337:
5336:
5332:
5331:
5330:
5328:
5327:
5326:
5282:
5281:
5280:
5275:
5274:
5269:
5250:
5143:
5105:
5096:
5011:
5006:
4966:
4886:
4881:
4851:
4846:
4775:
4687:
4659:
4577:
4568:
4508:
4505:
4475:
4470:
4400:
4352:
4347:
4303:
4293:
4275:Sherratt, T. N.
4269:
4251:
4245:
4228:
4225:
4220:
4210:
4208:
4201:Dendrobates.org
4194:
4193:
4189:
4164:
4163:
4159:
4142:
4141:
4137:
4129:
4125:
4081:
4080:
4076:
4045:
4016:
4015:
4011:
3973:
3972:
3968:
3931:Servedio, M. R.
3929:
3928:
3924:
3894:
3893:
3889:
3853:Ecology Letters
3849:
3848:
3844:
3788:
3787:
3780:
3729:
3728:
3724:
3696:
3695:
3691:
3655:
3654:
3650:
3622:
3621:
3617:
3579:
3578:
3569:
3531:
3530:
3526:
3510:
3509:
3502:
3472:
3471:
3467:
3437:
3436:
3432:
3404:
3403:
3399:
3355:
3354:
3347:
3311:
3310:
3306:
3272:
3271:
3258:
3237:(11): 598–603.
3226:
3225:
3216:
3168:
3167:
3163:
3155:
3151:
3111:
3110:
3106:
3099:
3080:
3079:
3075:
3061:
3060:
3056:
3046:
3045:
3041:
3030:Darwin, Charles
3028:
3027:
3023:
2985:
2984:
2980:
2932:
2931:
2927:
2879:
2878:
2874:
2846:
2845:
2841:
2831:
2830:
2826:
2816:
2815:
2811:
2783:
2782:
2778:
2739:
2738:
2734:
2696:
2695:
2691:
2661:
2660:
2656:
2634:
2633:
2626:
2588:
2587:
2583:
2547:
2546:
2542:
2520:
2519:
2515:
2493:
2492:
2488:
2472:
2471:
2467:
2431:
2430:
2426:
2416:
2415:
2408:
2388:
2386:
2382:
2335:
2330:
2329:
2322:
2312:
2310:
2300:
2299:
2292:
2240:
2239:
2235:
2212:(6): 993–1005.
2197:
2196:
2192:
2138:
2137:
2133:
2118:10.2307/3677074
2103:
2102:
2098:
2051:
2050:
2046:
2038:
2034:
2022:
2018:
1964:
1963:
1959:
1950:
1948:
1940:
1939:
1935:
1897:
1896:
1892:
1848:
1847:
1838:
1823:
1798:
1797:
1793:
1739:
1738:
1731:
1677:
1676:
1669:
1641:
1640:
1636:
1592:
1591:
1587:
1577:
1575:
1570:
1569:
1565:
1527:
1526:
1522:
1474:
1473:
1466:
1454:
1453:
1449:
1411:
1410:
1406:
1362:
1361:
1357:
1306:
1305:
1298:
1268:
1267:
1263:
1215:
1214:
1210:
1154:
1153:
1146:
1138:
1134:
1129:(215): 384–408.
1111:
1110:
1103:
1093:
1091:
1081:
1080:
1071:
1063:
1056:
1007:
1006:
1002:
956:
955:
951:
947:
935:
928:
917:
908:
902:
837:
831:
780:
759:
746:
701:
647:In a letter to
645:
625:
600:
515:Batesian mimics
490:
489:
488:
487:
486:
479:
471:
470:
457:
446:
299:
294:
270:), or exposing
193:
176:
146:
28:
23:
22:
15:
12:
11:
5:
5335:
5333:
5325:
5324:
5319:
5314:
5309:
5304:
5299:
5294:
5284:
5283:
5277:
5276:
5271:
5270:
5268:
5267:
5255:
5252:
5251:
5249:
5248:
5243:
5238:
5233:
5228:
5223:
5218:
5213:
5208:
5203:
5196:
5191:
5186:
5181:
5176:
5167:
5162:
5157:
5151:
5149:
5148:Related topics
5145:
5144:
5142:
5141:
5136:
5131:
5126:
5121:
5115:
5113:
5107:
5106:
5099:
5097:
5095:
5094:
5089:
5087:In vertebrates
5084:
5079:
5074:
5069:
5064:
5059:
5054:
5053:
5052:
5042:
5037:
5032:
5027:
5021:
5019:
5013:
5012:
5007:
5005:
5004:
4997:
4990:
4982:
4976:
4974:
4968:
4967:
4965:
4964:
4959:
4954:
4949:
4944:
4939:
4938:
4937:
4927:
4925:Apparent death
4922:
4921:
4920:
4910:
4905:
4900:
4894:
4892:
4888:
4887:
4882:
4880:
4879:
4872:
4865:
4857:
4848:
4847:
4845:
4844:
4839:
4834:
4829:
4824:
4819:
4814:
4809:
4804:
4799:
4794:
4789:
4783:
4781:
4780:Related topics
4777:
4776:
4774:
4773:
4768:
4763:
4758:
4753:
4752:
4751:
4741:
4736:
4731:
4729:Countershading
4726:
4721:
4716:
4711:
4706:
4701:
4695:
4693:
4689:
4688:
4686:
4685:
4680:
4675:
4669:
4667:
4661:
4660:
4658:
4657:
4652:
4647:
4645:Holochroal eye
4642:
4641:
4640:
4635:
4625:
4624:
4623:
4613:
4608:
4603:
4598:
4593:
4587:
4585:
4579:
4578:
4571:
4569:
4567:
4566:
4565:
4564:
4559:
4554:
4544:
4539:
4534:
4529:
4524:
4518:
4516:
4510:
4509:
4506:
4504:
4503:
4496:
4489:
4481:
4472:
4471:
4469:
4468:
4463:
4458:
4453:
4448:
4446:
4445:
4444:
4439:
4434:
4432:Apparent death
4429:
4424:
4414:
4408:
4406:
4402:
4401:
4399:
4398:
4393:
4388:
4383:
4378:
4377:
4376:
4366:
4360:
4358:
4354:
4353:
4348:
4346:
4345:
4338:
4331:
4323:
4317:
4316:
4302:
4301:External links
4299:
4298:
4297:
4291:
4267:
4249:
4243:
4224:
4221:
4219:
4218:
4187:
4157:
4135:
4133:, p. 112.
4123:
4074:
4034:(3): 495–566.
4009:
3982:(4): 201–202.
3966:
3945:(3): 751–763.
3922:
3887:
3860:(4): 495–501.
3842:
3778:
3722:
3709:(2): 174–177.
3689:
3648:
3635:(1): 148–156.
3615:
3588:(1): 205–216.
3567:
3546:(5): 933–940.
3524:
3500:
3487:(2): 425–432.
3465:
3430:
3417:(4): 738–744.
3397:
3345:
3324:(5): 933–940.
3304:
3285:(3): 548–566.
3256:
3214:
3183:(2): 388–414.
3161:
3149:
3104:
3097:
3073:
3054:
3039:
3021:
2978:
2925:
2872:
2859:(4): 229–242.
2839:
2824:
2809:
2776:
2732:
2689:
2670:(2): 339–347.
2654:
2624:
2597:(2): 181–188.
2581:
2540:
2513:
2486:
2465:
2424:
2406:
2346:(4): 170–172.
2320:
2290:
2233:
2190:
2131:
2112:(3): 184–197.
2096:
2044:
2032:
2016:
1957:
1933:
1890:
1836:
1821:
1791:
1729:
1667:
1654:(2): 221–228.
1634:
1585:
1563:
1520:
1485:(4): 164–169.
1464:
1447:
1404:
1355:
1296:
1261:
1208:
1169:10.1086/663197
1144:
1132:
1101:
1090:on 8 July 2017
1069:
1054:
1000:
948:
946:
943:
942:
941:
934:
931:
930:
929:
925:Batesian mimic
921:red milk snake
918:
911:
909:
903:
896:
830:
827:
779:
776:
758:
755:
745:
742:
700:
697:
653:Charles Darwin
644:
641:
637:Lygaeus kalmii
624:
621:
599:
596:
480:
473:
472:
458:
451:
450:
449:
448:
447:
445:
442:
440:caterpillars.
298:
295:
293:
290:
220:Hycleus lugens
175:
172:
145:
142:
121:'s concept of
26:
24:
18:Warning signal
14:
13:
10:
9:
6:
4:
3:
2:
5334:
5323:
5320:
5318:
5315:
5313:
5310:
5308:
5305:
5303:
5300:
5298:
5295:
5293:
5290:
5289:
5287:
5266:
5257:
5256:
5253:
5247:
5244:
5242:
5239:
5237:
5234:
5232:
5229:
5227:
5224:
5222:
5219:
5217:
5214:
5212:
5209:
5207:
5204:
5202:
5201:
5197:
5195:
5192:
5190:
5187:
5185:
5182:
5180:
5177:
5175:
5171:
5168:
5166:
5163:
5161:
5158:
5156:
5153:
5152:
5150:
5146:
5140:
5137:
5135:
5132:
5130:
5127:
5125:
5122:
5120:
5117:
5116:
5114:
5112:
5108:
5103:
5093:
5090:
5088:
5085:
5083:
5080:
5078:
5075:
5073:
5070:
5068:
5065:
5063:
5060:
5058:
5055:
5051:
5048:
5047:
5046:
5043:
5041:
5038:
5036:
5033:
5031:
5028:
5026:
5023:
5022:
5020:
5018:
5014:
5010:
5003:
4998:
4996:
4991:
4989:
4984:
4983:
4980:
4975:
4973:
4969:
4963:
4960:
4958:
4957:Mobbing calls
4955:
4953:
4950:
4948:
4945:
4943:
4940:
4936:
4933:
4932:
4931:
4928:
4926:
4923:
4919:
4916:
4915:
4914:
4911:
4909:
4906:
4904:
4901:
4899:
4896:
4895:
4893:
4889:
4885:
4878:
4873:
4871:
4866:
4864:
4859:
4858:
4855:
4843:
4840:
4838:
4835:
4833:
4830:
4828:
4825:
4823:
4820:
4818:
4815:
4813:
4810:
4808:
4805:
4803:
4800:
4798:
4795:
4793:
4790:
4788:
4787:Animal senses
4785:
4784:
4782:
4778:
4772:
4769:
4767:
4764:
4762:
4759:
4757:
4754:
4750:
4747:
4746:
4745:
4742:
4740:
4737:
4735:
4732:
4730:
4727:
4725:
4722:
4720:
4719:Chromatophore
4717:
4715:
4712:
4710:
4707:
4705:
4702:
4700:
4697:
4696:
4694:
4690:
4684:
4681:
4679:
4676:
4674:
4671:
4670:
4668:
4666:
4662:
4656:
4653:
4651:
4648:
4646:
4643:
4639:
4636:
4634:
4631:
4630:
4629:
4626:
4622:
4619:
4618:
4617:
4616:Mammalian eye
4614:
4612:
4609:
4607:
4604:
4602:
4599:
4597:
4594:
4592:
4591:Arthropod eye
4589:
4588:
4586:
4584:
4580:
4575:
4563:
4560:
4558:
4555:
4553:
4550:
4549:
4548:
4545:
4543:
4540:
4538:
4535:
4533:
4530:
4528:
4525:
4523:
4520:
4519:
4517:
4515:
4511:
4502:
4497:
4495:
4490:
4488:
4483:
4482:
4479:
4467:
4464:
4462:
4459:
4457:
4454:
4452:
4449:
4447:
4443:
4440:
4438:
4435:
4433:
4430:
4428:
4425:
4423:
4420:
4419:
4418:
4415:
4413:
4410:
4409:
4407:
4403:
4397:
4394:
4392:
4389:
4387:
4384:
4382:
4379:
4375:
4372:
4371:
4370:
4367:
4365:
4362:
4361:
4359:
4355:
4351:
4344:
4339:
4337:
4332:
4330:
4325:
4324:
4321:
4314:
4309:
4305:
4304:
4300:
4294:
4288:
4284:
4280:
4276:
4272:
4268:
4264:
4260:
4259:
4254:
4250:
4246:
4240:
4235:
4234:
4227:
4226:
4222:
4206:
4202:
4198:
4191:
4188:
4183:
4179:
4175:
4171:
4167:
4166:Müller, Fritz
4161:
4158:
4153:
4150:(in German).
4149:
4145:
4144:Müller, Fritz
4139:
4136:
4132:
4127:
4124:
4119:
4115:
4110:
4105:
4101:
4097:
4093:
4089:
4085:
4078:
4075:
4069:
4064:
4060:
4056:
4052:
4048:
4041:
4037:
4033:
4029:
4028:
4023:
4019:
4013:
4010:
4005:
4001:
3997:
3993:
3989:
3985:
3981:
3977:
3970:
3967:
3962:
3958:
3953:
3948:
3944:
3940:
3936:
3932:
3926:
3923:
3918:
3914:
3910:
3906:
3902:
3898:
3891:
3888:
3883:
3879:
3875:
3871:
3867:
3863:
3859:
3855:
3854:
3846:
3843:
3838:
3834:
3829:
3824:
3819:
3814:
3810:
3806:
3802:
3798:
3797:
3792:
3785:
3783:
3779:
3774:
3770:
3765:
3760:
3756:
3752:
3748:
3744:
3740:
3736:
3732:
3726:
3723:
3717:
3712:
3708:
3704:
3700:
3693:
3690:
3684:
3679:
3675:
3671:
3667:
3663:
3659:
3652:
3649:
3643:
3638:
3634:
3630:
3626:
3619:
3616:
3611:
3607:
3603:
3599:
3595:
3591:
3587:
3583:
3576:
3574:
3572:
3568:
3563:
3559:
3554:
3549:
3545:
3541:
3540:
3535:
3528:
3525:
3520:
3519:
3518:New Scientist
3514:
3507:
3505:
3501:
3495:
3490:
3486:
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3466:
3461:
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3408:
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3379:
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3363:
3359:
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3350:
3346:
3341:
3337:
3332:
3327:
3323:
3319:
3315:
3308:
3305:
3300:
3296:
3292:
3288:
3284:
3280:
3276:
3269:
3267:
3265:
3263:
3261:
3257:
3252:
3248:
3244:
3240:
3236:
3232:
3231:
3223:
3221:
3219:
3215:
3210:
3206:
3201:
3196:
3191:
3186:
3182:
3178:
3177:
3172:
3165:
3162:
3158:
3153:
3150:
3145:
3141:
3137:
3136:10.1038/29751
3133:
3129:
3125:
3121:
3117:
3116:
3108:
3105:
3100:
3094:
3090:
3086:
3085:
3077:
3074:
3070:: lxxx–lxxxi.
3069:
3065:
3062:Anon (1867).
3058:
3055:
3050:
3043:
3040:
3035:
3031:
3025:
3022:
3017:
3013:
3009:
3005:
3001:
2997:
2993:
2989:
2982:
2979:
2974:
2970:
2966:
2962:
2957:
2952:
2948:
2944:
2940:
2936:
2935:Ruxton, G. D.
2929:
2926:
2921:
2917:
2913:
2909:
2904:
2899:
2895:
2891:
2887:
2883:
2882:Ruxton, G. D.
2876:
2873:
2867:
2862:
2858:
2854:
2850:
2843:
2840:
2835:
2828:
2825:
2820:
2813:
2810:
2804:
2799:
2795:
2791:
2787:
2780:
2777:
2772:
2768:
2764:
2760:
2756:
2752:
2748:
2744:
2736:
2733:
2728:
2724:
2719:
2714:
2710:
2706:
2705:
2700:
2693:
2690:
2685:
2681:
2677:
2673:
2669:
2665:
2658:
2655:
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2646:
2642:
2638:
2631:
2629:
2625:
2620:
2616:
2612:
2608:
2604:
2600:
2596:
2592:
2585:
2582:
2576:
2571:
2567:
2563:
2559:
2555:
2551:
2544:
2541:
2536:
2532:
2529:(2): 99–109.
2528:
2524:
2517:
2514:
2509:
2505:
2502:(2): 99–109.
2501:
2497:
2490:
2487:
2482:
2478:
2477:
2469:
2466:
2460:
2455:
2451:
2447:
2443:
2439:
2435:
2428:
2425:
2420:
2413:
2411:
2407:
2403:
2401:
2397:
2385:on 2015-12-16
2381:
2377:
2373:
2369:
2365:
2361:
2357:
2353:
2349:
2345:
2341:
2334:
2327:
2325:
2321:
2308:
2304:
2297:
2295:
2291:
2286:
2282:
2277:
2272:
2268:
2264:
2260:
2256:
2252:
2248:
2244:
2237:
2234:
2229:
2225:
2220:
2215:
2211:
2207:
2206:
2201:
2194:
2191:
2186:
2182:
2177:
2172:
2167:
2162:
2158:
2154:
2150:
2146:
2142:
2135:
2132:
2127:
2123:
2119:
2115:
2111:
2107:
2100:
2097:
2092:
2088:
2083:
2078:
2074:
2070:
2066:
2062:
2058:
2054:
2048:
2045:
2041:
2036:
2033:
2030:
2026:
2020:
2017:
2012:
2008:
2003:
1998:
1993:
1988:
1984:
1980:
1976:
1972:
1968:
1961:
1958:
1947:
1943:
1937:
1934:
1928:
1923:
1918:
1913:
1909:
1905:
1901:
1894:
1891:
1886:
1882:
1877:
1872:
1868:
1864:
1860:
1856:
1852:
1845:
1843:
1841:
1837:
1832:
1828:
1824:
1818:
1814:
1810:
1806:
1802:
1795:
1792:
1787:
1783:
1778:
1773:
1768:
1763:
1759:
1755:
1751:
1747:
1743:
1736:
1734:
1730:
1725:
1721:
1716:
1711:
1706:
1701:
1697:
1693:
1689:
1685:
1681:
1674:
1672:
1668:
1662:
1657:
1653:
1649:
1645:
1638:
1635:
1630:
1626:
1622:
1618:
1614:
1610:
1606:
1602:
1601:
1596:
1589:
1586:
1573:
1567:
1564:
1559:
1555:
1551:
1547:
1543:
1539:
1536:(2): 99–111.
1535:
1531:
1524:
1521:
1516:
1512:
1508:
1504:
1500:
1496:
1492:
1488:
1484:
1480:
1479:
1471:
1469:
1465:
1460:
1459:
1451:
1448:
1443:
1439:
1434:
1429:
1425:
1421:
1420:
1415:
1408:
1405:
1400:
1396:
1391:
1386:
1382:
1378:
1374:
1370:
1366:
1359:
1356:
1351:
1347:
1342:
1337:
1333:
1329:
1325:
1321:
1317:
1313:
1312:Ruxton, G. D.
1309:
1303:
1301:
1297:
1292:
1288:
1284:
1280:
1276:
1272:
1265:
1262:
1257:
1253:
1248:
1243:
1239:
1235:
1231:
1227:
1223:
1219:
1212:
1209:
1204:
1200:
1196:
1192:
1188:
1184:
1179:
1174:
1170:
1166:
1163:(1): E1–E14.
1162:
1158:
1151:
1149:
1145:
1141:
1136:
1133:
1128:
1124:
1123:
1118:
1114:
1108:
1106:
1102:
1089:
1085:
1084:"Aposematism"
1082:Marek, Paul.
1078:
1076:
1074:
1070:
1066:
1061:
1059:
1055:
1050:
1046:
1041:
1036:
1032:
1028:
1025:(4506): 476.
1024:
1020:
1019:
1014:
1010:
1004:
1001:
996:
992:
987:
982:
977:
972:
968:
964:
960:
953:
950:
944:
940:
937:
936:
932:
926:
922:
919:The harmless
915:
910:
907:
900:
895:
893:
891:
890:
883:
881:
877:
873:
868:
863:
861:
857:
853:
849:
845:
840:
836:
828:
826:
822:
820:
819:kin selection
814:
811:
806:
804:
800:
795:
792:
787:
785:
777:
775:
772:
766:
764:
756:
754:
751:
743:
740:
736:
734:
733:
728:
724:
716:
715:
710:
705:
699:Poulton, 1890
698:
696:
693:
689:
684:
682:
678:
674:
670:
664:
662:
658:
654:
650:
643:Wallace, 1867
642:
638:
634:
629:
622:
620:
618:
612:
609:
605:
597:
595:
592:
591:chromatophore
588:
584:
582:
578:
569:
564:
560:
558:
555:
551:
550:
545:
537:
536:
530:
526:
524:
520:
516:
511:
509:
508:
502:
498:
496:
484:
477:
468:
467:
462:
455:
443:
441:
439:
435:
434:
429:
424:
422:
418:
414:
410:
406:
402:
398:
397:cinnabar moth
394:
390:
385:
383:
382:
377:
376:
371:
367:
366:
361:
360:
355:
354:
349:
348:Dendrobatidae
344:
342:
338:
334:
330:
326:
322:
318:
310:
309:
303:
296:
291:
289:
287:
286:honey badgers
283:
279:
278:
273:
269:
265:
261:
257:
253:
249:
245:
241:
237:
233:
229:
221:
217:
213:
211:
205:
203:
199:
192:
184:
180:
173:
171:
169:
165:
161:
160:Ancient Greek
157:
156:
151:
143:
141:
139:
135:
130:
128:
124:
120:
116:
111:
109:
105:
101:
97:
93:
89:
85:
81:
74:
70:
66:
62:
57:
50:
46:
43:a warning to
42:
39:
34:
30:
19:
5236:Polymorphism
5231:Phagomimicry
5198:
5179:Co-evolution
5164:
4912:
4908:Alarm signal
4708:
4650:Parietal eye
4596:Compound eye
4426:
4422:Alarm signal
4278:
4257:
4232:
4209:. Retrieved
4205:the original
4200:
4190:
4181:
4177:
4173:
4169:
4160:
4151:
4147:
4138:
4131:Edmunds 1974
4126:
4091:
4087:
4077:
4061:(1): 41–54.
4058:
4054:
4031:
4025:
4012:
3979:
3976:Chemoecology
3975:
3969:
3942:
3938:
3925:
3900:
3896:
3890:
3857:
3851:
3845:
3800:
3794:
3746:
3742:
3725:
3706:
3702:
3692:
3665:
3661:
3651:
3632:
3628:
3618:
3585:
3581:
3543:
3537:
3527:
3516:
3484:
3478:
3468:
3443:
3439:
3433:
3414:
3410:
3400:
3365:
3361:
3321:
3317:
3307:
3282:
3278:
3234:
3228:
3180:
3174:
3164:
3157:Poulton 1890
3152:
3119:
3113:
3107:
3083:
3076:
3067:
3057:
3042:
3024:
2991:
2987:
2981:
2946:
2942:
2928:
2893:
2889:
2875:
2856:
2852:
2842:
2833:
2827:
2818:
2812:
2796:(1): 68–77.
2793:
2789:
2779:
2746:
2743:Chemoecology
2742:
2735:
2708:
2702:
2692:
2667:
2663:
2657:
2640:
2636:
2594:
2590:
2584:
2557:
2553:
2543:
2526:
2522:
2516:
2499:
2495:
2489:
2480:
2474:
2468:
2441:
2437:
2427:
2418:
2399:
2395:
2393:
2387:. Retrieved
2380:the original
2343:
2339:
2311:. Retrieved
2307:the original
2253:(1): 31469.
2250:
2246:
2236:
2209:
2203:
2193:
2148:
2144:
2134:
2109:
2105:
2099:
2064:
2060:
2047:
2040:Edmunds 1974
2035:
2024:
2019:
1974:
1970:
1960:
1949:. Retrieved
1946:ResearchGate
1945:
1936:
1907:
1903:
1893:
1858:
1854:
1804:
1794:
1749:
1745:
1687:
1683:
1651:
1647:
1637:
1604:
1598:
1588:
1576:. Retrieved
1566:
1533:
1529:
1523:
1482:
1476:
1456:
1450:
1423:
1417:
1407:
1372:
1368:
1358:
1323:
1319:
1274:
1270:
1264:
1229:
1225:
1211:
1160:
1156:
1135:
1126:
1120:
1094:November 24,
1092:. Retrieved
1088:the original
1065:Poulton 1890
1022:
1016:
1003:
966:
962:
952:
887:
884:
876:Fritz Müller
864:
841:
838:
823:
815:
807:
803:X chromosome
796:
788:
781:
767:
760:
750:evolutionary
747:
738:
730:
729:in his book
726:
720:
712:
685:
680:
676:
672:
668:
665:
646:
636:
633:milkweed bug
613:
601:
585:
573:
567:
547:
541:
533:
519:Phyllidiidae
512:
507:Hexabranchus
505:
499:
491:
464:
431:
428:rattlesnakes
425:
413:honey badger
386:
379:
373:
363:
359:Epipedobates
357:
351:
345:
314:
306:
282:stridulation
275:
230:such as the
225:
219:
206:
194:
167:
163:
153:
147:
131:
122:
112:
79:
78:
61:honey badger
29:
5165:Aposematism
5040:Automimicry
4918:Unkenreflex
4913:Aposematism
4891:Non-mimicry
4827:Pseudopupil
4709:Aposematism
4628:Mollusc eye
4466:Unkenreflex
4427:Aposematism
4237:. Longman.
4044:; Reprint:
2643:: 157–164.
2476:Malacologia
2444:: 183–194.
1308:Stevens, M.
906:coral snake
852:hornet moth
727:aposematism
608:grasshopper
461:nudibranchs
438:Bombycoidea
381:Mannophryne
375:Colostethus
365:Phyllobates
353:Dendrobates
317:vertebrates
268:Unkenreflex
108:perceivable
106:, or other
84:advertising
80:Aposematism
5286:Categories
5170:Camouflage
5139:Vavilovian
5129:Gilbertian
5092:Wasmannian
5017:In animals
4817:Ommatidium
4749:coincident
4714:Camouflage
4692:Coloration
4633:cephalopod
4527:Chameleons
4456:Camouflage
4261:. London:
2483:: 241–255.
2389:2015-06-21
1951:2017-11-11
1927:10342/8273
1910:(8): 126.
1178:2152/31175
1009:Eisner, T.
945:References
791:gregarious
501:Nudibranch
417:passerines
393:herbivores
292:Prevalence
264:ultrasonic
256:neurotoxin
252:velvet ant
236:tiger moth
210:camouflage
162:words ἀπό
96:coloration
5124:Dodsonian
5111:In plants
5077:Müllerian
5050:Locomotor
4832:Rhopalium
4665:Evolution
4638:gastropod
4606:Eye shine
4601:Eagle eye
4532:Dinosaurs
4168:(1879). "
3939:Evolution
3903:: 63–74.
3539:Evolution
3446:: 49–56.
3318:Evolution
3144:204999972
2988:Evolution
2943:Evolution
2890:Evolution
2560:: 29–39.
2151:(1): 35.
2053:Caro, Tim
1904:Diversity
1419:Evolution
1277:: iii–v.
799:recessive
784:neophobia
744:Evolution
598:Behaviour
577:blueheads
554:cytolytic
370:alkaloids
325:amphibian
144:Etymology
88:predators
45:predators
5119:Bakerian
5062:Chemical
5045:Batesian
4962:Stotting
4699:Albinism
4374:examples
4255:(1890).
4174:Thyridia
4154:: 54–55.
4118:17567563
4049:(1981).
4020:(1861).
4004:27025142
3961:10937250
3933:(2000).
3917:54273453
3837:11416165
3773:16959629
3737:(2006).
3610:13434024
3602:11170710
3562:16136793
3460:53174146
3392:11270431
3340:16136793
3251:16701442
3209:30152037
3016:10785432
3008:21166789
2973:21509940
2965:20050915
2937:(2010).
2920:21509940
2912:20050915
2884:(2010).
2727:23053367
2684:43941481
2619:10948319
2611:20413794
2368:15761732
2285:27510510
2228:17337712
2185:19208237
2091:18990666
2055:(2009).
2011:21444790
1885:25320164
1831:85845409
1786:11353830
1724:19858491
1578:21 March
1558:83847876
1515:18306198
1507:15772807
1442:18764916
1399:16006332
1350:22113031
1314:(2012).
1291:53202282
1256:19019790
1195:22173468
1115:(1877).
995:14555763
933:See also
874:, after
858:, after
771:innately
557:saponins
405:zorillas
341:hominins
337:Pitohuis
272:eyespots
232:ladybird
49:toxicity
5317:Ecology
5174:Crypsis
5072:Eyespot
5009:Mimicry
4972:Mimicry
4930:Display
4761:Mimicry
4734:Crypsis
4547:Mammals
4461:Mimicry
4451:Crypsis
4405:Signals
4223:Sources
4109:2275182
3984:Bibcode
3882:7188102
3862:Bibcode
3805:Bibcode
3764:1634906
3683:1689056
3383:1088614
3287:Bibcode
3200:6446817
3124:Bibcode
3089:253–524
2771:9953958
2751:Bibcode
2562:Bibcode
2446:Bibcode
2376:8525349
2348:Bibcode
2313:21 June
2276:4980592
2255:Bibcode
2176:2645363
2153:Bibcode
2126:3677074
2082:2674080
2002:3076872
1979:Bibcode
1876:4213648
1754:Bibcode
1715:2776464
1692:Bibcode
1629:1439786
1609:Bibcode
1600:Science
1538:Bibcode
1487:Bibcode
1390:1560331
1341:3234570
1247:2664363
1203:1963316
1049:7244647
1027:Bibcode
1018:Science
844:evolved
835:Mimicry
829:Mimicry
677:seizure
495:sponges
333:mammals
321:reptile
305:Skunk,
228:insects
82:is the
47:of its
5258:
5082:Sexual
4552:horses
4514:Vision
4289:
4241:
4211:11 May
4116:
4106:
4002:
3959:
3915:
3880:
3835:
3825:
3771:
3761:
3680:
3608:
3600:
3560:
3458:
3390:
3380:
3338:
3249:
3207:
3197:
3142:
3115:Nature
3095:
3014:
3006:
2971:
2963:
2918:
2910:
2769:
2725:
2682:
2617:
2609:
2374:
2366:
2283:
2273:
2226:
2183:
2173:
2124:
2089:
2079:
2009:
1999:
1883:
1873:
1829:
1819:
1784:
1774:
1722:
1712:
1627:
1556:
1513:
1505:
1440:
1397:
1387:
1348:
1338:
1289:
1254:
1244:
1201:
1193:
1187:663197
1185:
1047:
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