Revealing the colourful side of birds: spatial distribution of conspicuous plumage colours on the body of Australian birds

In many species of birds, different body parts often display very different colours. This spatial distribution of coloured plumage patches may be determined, among other factors, by the balance between being cryptic to predators, and conspicuous to intended receivers. If this is the case, ventral and anterior body parts in birds – which are less visible to predators but more prominent to conspecifics – should present more conspicuous and sexually dichromatic plumage colours. Here, I test these predictions using reflectance spectrometric measurements of standardised plumage patches across males and females for nearly an entire avifauna (Australian landbirds, n = 538 species). My data show that, as predicted, conspicuous and sexually dichromatic colours are mainly located near the head, while the plumage of the back is the most cryptic. One clear exception to this pattern is the conspicuous rump coloration. In many species, this patch can be concealed by wings, and therefore exposed only when necessary. In addition, conspicuous rump coloration could deflect or confuse predators in case of attack. However, there is considerable variation across species, and this makes position on the body a very poor predictor of plumage elaboration (R² < 0.02). Future studies should try to determine whether differences between species in the distribution of colours across the plumage are due to variation in ecological factors (predation risk, habitat, etc.).     

Interactions of environmental factors influencing pupal coloration in swallowtail butterfly Papilio xuthus

The swallowtail butterfly Papilio xuthus Linné [Lepidoptera: Papilionidae] exhibits pupal protective color polyphenism. Interactions of various environmental factors on pupal coloration were analyzed in non-diapausing individuals. Under sufficient light (200lux), most pupating larvae became green pupae when the surface of the pupation site was smooth, while they became brown when the surface was rough. Tactile signals are the positive environmental factors causing induction of the brown pupal coloration. In dark boxes, the induction of the brown pupal coloration was easily induced even on a smooth surface, suggesting that light suppresses induction of brown coloration. Different colors of pupation sites did not affect pupal coloration under sufficient light. Environmental factors received during a critical period both before girdling and after girdling affected pupal coloration. When tactile signals received from rough surfaces reach threshold levels during pupation, brown pupal coloration is determined. Larvae reared under a daily periodicity of natural light formed a girdle at midnight, subsequently, the prepupae received strong daylight the following day. Under natural light most larvae produced brown pupae on rough surfaces and green pupae on smooth surfaces.     

Predator preference for brightly colored males in the guppy: a viability cost for a sexually selected trait

Although conspicuous visual sexual signals, such as bright colors,in males serve to attract females in numerous species, theymay also attract the attention of potential predators and thusmay be costly in terms of increasing individual risk of mortalityto predation. Most models of the evolution of extravagant malesexual traits and female preferences for them assume that thesexually preferred male trait is costly to produce and maintain.However, there is surprisingly little empirical evidence fordirect fitness costs associated with sexually selected visualtraits that enhance male mating success. In the present study,we report a direct fitness cost for sexually selected, brightbody-color patterns in males in the form of an associated greaterrisk of mortality to predation. By using the guppy (Poeciliareticulata) and the blue acara cichlid fish (Aequidens pulcher)as a model prey–predator system, we demonstrate experimentallythat individual cichlids preferentially and consistently approached,attacked, and captured the more brightly colored of two size-matchedmale guppies presented simultaneously in staged encounters.This resulted in the brightly colored male incurring, on average,a significantly higher risk of mortality given an encounterwith the predator than with the drabber male in matched pairs.Our results constitute strong behavioral evidence for a directviability cost associated with bright coloration in male guppies,and they corroborate the generally accepted paradigm that directionalpredation by visual fish predators against brightly colored,adult male guppies underlies the evolution of the known divergentcolor patterns in natural guppy populations that experiencedifferent intensities of predation. The viability cost associatedwith bright conspicuous coloration in male guppies potentiallyreinforces for females the reliability of this sexually selectedtrait as an indicator trait of male quality.     

Conspicuousness,color resemblance,and toxicity in geographically diverging mimicry: The pan‐Amazonian frog Allobates femoralis

Predation risk is allegedly reduced in Batesian and Müllerian mimics, because their coloration resembles the conspicuous coloration of unpalatable prey. The efficacy of mimicry is thought to be affected by variation in the unpalatability of prey, the conspicuousness of the signals, and the visual system of predators that see them. Many frog species exhibit small colorful patches contrasting against an otherwise dark body. By measuring toxicity and color reflectance in a geographically variable frog species and the syntopic toxic species, we tested whether unpalatability was correlated with between‐species color resemblance and whether resemblance was highest for the most conspicuous components of coloration pattern. Heterospecific resemblance in colorful patches was highest between species at the same locality, but unrelated to concomitant variation in toxicity. Surprisingly, resemblance was lower for the conspicuous femoral patches compared to the inconspicuous dorsum. By building visual models, we further tested whether resemblance was affected by the visual system of model predators. As predicted, mimic‐model resemblance was higher under the visual system of simulated predators compared to no visual system at all. Our results indicate that femoral patches are aposematic signals and support a role of mimicry in driving phenotypic divergence or mimetic radiation between localities.     

Eye camouflage and false eyespots: chaetodontid responses to predators

Synopsis The roles of eye camouflage and eyespots are examined within the genusChaetodon as are the various theories explaining the evolutionary significance of the brilliant colors. While eye camouflage is not common among reef fishes, 91% of the 90 species ofChaetodon, have eyemasks (82) or black heads (4). Eye camouflage occurs concomitantly with diurnal false eyespots in 45.5% (41 of 90) of the species. Diurnal false eyespots serve to misdirect attacks by predators and/or to advertise unpalatability. False eyespots are located on areas of the body which allow escape and survival following an attack. Data suggesting that predators learn about the undesirability of butterflyfishes are presented. Butterflyfishes are inactive at night, forage during the day and spawn at dusk. It is unlikely that nocturnal color changes are useful in conspecific interactions and are therefore believed to provide visual cues to potential predators. Nocturnal eyespots probably function to intimidate potential predators but could also remind them of unpalatability. The aggression release hypothesis (Lorenz 1962, 1966) to explain the brilliant coloration of chaetodontids is not supported because butterflyfish coloration changes and few species are territorial. The species recognition hypothesis (Zumpe 1965) is not supported by results of field experiments. The disruptive coloration hypothesis (Longley 1917) is rejected as a general explanation for poster coloration but does explain the prevalence of eyebars ofChaetodon spp. The aposematic hypothesis (Gosline 1965) is supported by morphology, behavior, a lack of predation and field observations. The possibility of Mullerian mimicry is suggested. It is concluded that the primary selective force behind chaetodontid coloration, particularly eyespots, has been predation and color patterns have evolved to minimize this threat.     

NOT EVERYTHING IS BLACK AND WHITE: COLOR AND BEHAVIORAL VARIATION REVEAL A CONTINUUM BETWEEN CRYPTIC AND APOSEMATIC STRATEGIES IN A POLYMORPHIC POISON FROG

Aposematism and crypsis are often viewed as two extremes of a continuum of visual conspicuousness to predators. Theory predicts that behavioral and coloration conspicuousness should vary in tandem along the conspicuousness spectrum for antipredator strategies to be effective. Here we used visual modeling of contrast and behavioral observations to examine the conspicuousness of four populations of the granular poison frog, Oophaga granulifera, which exhibits almost continuous variation in dorsal color. The patterns of geographic variation in color, visual contrast, and behavior support a gradient of overall conspicuousness along the distribution of O. granulifera. Red and green populations, at the extremes of the color distribution, differ in all elements of color, contrast, and behavior, strongly reflecting aposematic and cryptic strategies. However, there is no smooth cline in any elements of behavior or coloration between the two extremes. Instead populations of intermediate colors attain intermediate conspicuousness by displaying different combinations of aposematic and cryptic traits. We argue that coloration divergence among populations may be linked to the evolution of a gradient of strategies to balance the costs of detection by predators and the benefits of learned aversion.     

Gregarious pupation act as a defensive mechanism against cannibalism and intraguild predation

Coccinellid pupae use an array of defensive strategies against their natural enemies. This study aims to assess the efficiency of gregarious pupation as a defensive mechanism against intraguild predators and cannibals in coccinellid. The study was designed specifically (i) to determine the natural occurrence of gregarious pupation in the field for different coccinellid species, and (ii) to evaluate the adaptive value of gregarious pupation as a defensive mechanism against 2 types of predators (i.e., cannibals and intraguild predators). In the field, gregarious pupation consisted of a group of 2–5 pupae. The proportion of gregarious pupation observed varied according to species, the highest rate being observed with Harmonia axyridis Pallas (Coccinellidae; 14.17%). Gregarious pupation had no impact on the probability that intraguild predators and cannibals locate pupae. Intraguild predation occurred more often in site with gregarious pupation, while cannibalism occurred as often in site with gregarious pupation as in site with isolated pupa. However, for a specific pupa, the mortality rate was higher for isolated pupae than for pupae located in a gregarious pupation site both in the presence of intraguild predators and in the presence of cannibals. The spatial location of pupae within the group had no impact on mortality rate. Since it reduces the risk of predation, it is proposed that gregarious pupation act as a defensive mechanism for H. axyridis pupae.     

Effect of sex and bright coloration on survival and predator‐induced wing damage in an aposematic lantern fly with startle display

1. Aposematic coloration in prey promotes its survival by conspicuously advertising unpalatability to predators. Although classical examples of aposematic signals involve constant presentation of a signal at a distance, some animals suddenly display warning colours only when they are attacked. 2. Characteristics of body parts suddenly displayed, such as conspicuous coloration or eyespot pattern, may increase the survival of the prey by startling the predator, and/or by signalling unpalatability to the predators at the moment of attack. 3. The adaptive value of such colour patterns suddenly displayed by unpalatable prey has not been studied. We experimentally blackened the red patch in the conspicuous red–white–black hindwing pattern displayed by an unpalatable insect Lycorma delicatula White (Hemiptera: Fulgoridae) in response to predator's attack. 4. There was no evidence that the presence of the red patch increased prey survival over several weeks. We hypothesise that predators generalised from the red–white–black patches on the hindwings of unpalatable L. delicatula to any similar wing display as a signal of unpalatability. Because a higher proportion of males than females stay put at their resting sites, displaying their wings in response to repeated attacks by predators, wing damage was more frequent in males than in females. 5. To our knowledge, this is the first experimental test of an adaptive role of aposematic signals presented by unpalatable prey during sudden displays triggered by direct predatory attack.     

Predation on snakes of Argentina: Effects of coloration and ring pattern on coral and false coral snakes

The occurrence of coral snake coloration among unrelated venomous and non‐venomous snake species has often been explained in terms of warning coloration and mimicry. In Argentina, no field tests have been conducted to confirm this mimetic association between one venomous coral species (Micrurus phyrrocryptus, Elapidae) and two non‐venomous snake species with a similar color pattern (Lystrophis pulcher and Oxyrhopus rhombifer, Colubridae). The aims of this work were to test for the possible aposematic or cryptic function of the ring pattern and coloration of coral snakes and false coral snakes from central Argentina, and to analyse whether the pattern is effective throughout the year. Predation on snakes was estimated by using non‐toxic plasticine replicas of ringed venomous and non‐venomous snakes and unbanded green snakes placed along transects in their natural habitat during the dry and rainy season. Ringed color pattern was attacked by predators despite the background color. One of the replica types was attacked more than expected during the dry season, suggesting that both shape and width of rings may influence the choice by predators. The reaction of predators towards replicas that mimic snake species with ringed patterns is independent of the geographical region, and we can conclude that mimicry characteristics are quite general when the true models are present in the area.     

Pupal colour polymorphism in Papilio machaon L. and the survival in the field of cryptic versus non-cryptic pupae

• 1 A field experiment was carried out in a natural habitat of Papilio machaon L. in southern Sweden to assess the evolutionary significance of pupal colour polymorphism.
• 2 Cryptic and non-cryptic pupae were planted in pairs in the vegetation, and exposed to predators.
• 3 The protective coloration conferred a selective advantage approximating 1.5 on the cryptic pupae of the summer generation. In the overwintering generation no difference could be detected between the predation of cryptic and non-cryptic pupae.
• 4 The adaptive fitness of protective coloration, as determined by the different rates of elimination of the colour morphs, was greater for the green pupae than for the brown ones.
• 5 Natural selection favours the evolution of a seasonal difference in the proportion of green and brown pupae in the summer and hibernating generations of P.machaon.

     

Selective predation favouring cryptic individuals of marine snails (Littorina)

Many gastropods have inherited conspicuous shell colour polymorphisms. A challenging question is, are colour frequencies under selection or is polymorphism owing to random evolutionary processes? The intertidal species Littorina saxatilis (a rock‐dweller) and L. obtusata (confined to macroalgae) both have genetically determined shell colour variation. In Iceland, Littorina obtusata are mostly cryptic on brown macroalgae by having brown or yellow shells (~95% of the snails), while Littorina saxatilis often appears conspicuous to the background of dark rocks owing to non‐cryptic colours (15–20%). This difference may be due to selective elimination of conspicuously coloured L. obtusata by visual predators, while L. saxatilis, largely living in another habitat, is not under a similarly intense colour selection. To test this hypothesis we increased the frequencies of conspicuous L. saxatilis in experimental populations (from <12 to 55%) and placed these in the seaweed zone, the main habitat of L. obtusata. Fifteen populations were released on isolated spots of seaweed and three of these were covered by net cages to exclude bird predators. One month later, yellow snails had increased in frequency within the patches, and to our surprise the result did not differ between bare and caged patches. This suggests selection favouring a colour that matches the background of fucoid seaweeds by visual predators able to enter the cages. Birds acted as important predators by picking 16% of the experimental snails in the uncaged spots, but were unable to enter the caged spots. However, the bird predation was non‐selective with respect to snail colour. For various reasons the most likely predators able to enter the cages were intertidal fish, these were thus responsible for the selection of non‐cryptic snails. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 76 , 137–144.     

Conspicuous male coloration impairs survival against avian predators in Aegean wall lizards,Podarcis erhardii

Animal coloration is strikingly diverse in nature. Within‐species color variation can arise through local adaptation for camouflage, sexual dimorphism and conspicuous sexual signals, which often have conflicting effects on survival. Here, we tested whether color variation between two island populations of Aegean wall lizards (Podarcis erhardii) is due to sexual dimorphism and differential survival of individuals varying in appearance. On both islands, we measured attack rates by wild avian predators on clay models matching the coloration of real male and female P. erhardii from each island population, modeled to avian predator vision. Avian predator attack rates differed among model treatments, although only on one island. Male‐colored models, which were more conspicuous against their experimental backgrounds to avian predators, were accordingly detected and attacked more frequently by birds than less conspicuous female‐colored models. This suggests that female coloration has evolved primarily under selection for camouflage, whereas sexually competing males exhibit costly conspicuous coloration. Unexpectedly, there was no difference in avian attack frequency between local and non‐local model types. This may have arisen if the models did not resemble lizard coloration with sufficient precision, or if real lizards behaviorally choose backgrounds that improve camouflage. Overall, these results show that sexually dimorphic coloration can affect the risk of predator attacks, indicating that color variation within a species can be caused by interactions between natural and sexual selection. However, more work is needed to determine how these findings depend on the island environment that each population inhabits.     

Asymmetry in size, shape, and color impairs the protective value of conspicuous color patterns

The received view of protective coloration in animals is thatconspicuous colors and patterns have evolved because they elicitavoidance behavior in potential predators. In the present study,we examine the spontaneous response of naive predators (Gallusgallus domesticus) to artificial prey to test the hypothesisthat deviations from bilateral symmetry of signaling patternelements may negatively influence the avoidance-inducing effectof conspicuous color patterns. Chicks displayed stronger aversionsto artificial "butterfly" prey items possessing symmetric colorpattern elements than to those possessing asymmetric signalswith pattern elements of different color or shape. Althoughthey attacked signals with a size asymmetry of 5% at the samerate as symmetric signals, signals with a size asymmetry of7.5% or more were attacked more often than were symmetric signals.These results suggest that the protective value of conspicuouscolor patterns is impaired by asymmetry in color, shape, andsize of color pattern elements. Our findings also argue againstthe notion that animals have inherent preferences for symmetricover asymmetric objects, and demonstrate the existence of athreshold for asymmetry detection, beyond which further incrementsin asymmetry have no influence on signal efficacy.     

Perspective: the evolution of warning coloration is not paradoxical

Animals that are brightly colored have intrigued scientists since the time of Darwin, because it seems surprising that prey should have evolved to be clearly visible to predators. Often this self-advertisement is explained by the prey being unprofitable in some way, with the conspicuous warning coloration helping to protect the prey because it signals to potential predators that the prey is unprofitable. However, such signals only work in this way once predators have learned to associate the conspicuous color with the unprofitability of the prey. The evolution of warning coloration is still widely considered to be a paradox, because it has traditionally been assumed that the very first brightly colored individuals would be at an immediate selective disadvantage because of their greater conspicuousness to predators that are naive to the meaning of the signal. As a result, it has been difficult to understand how a novel conspicuous color morph could ever avoid extinction for long enough for predators to become educated about the signal. Thus, the traditional view that the evolution of warning coloration is difficult to explain rests entirely on assumptions about the foraging behavior of predators. However, we review recent evidence from a range of studies of predator foraging decisions, which refute these established assumptions. These studies show that: (1) Many predators are so conservative in their food preferences that even very conspicuous novel prey morphs are not necessarily at a selective disadvantage. (2) The survival and spread of novel color morphs can be simulated in field and aviary experiments using real predators (birds) foraging on successive generations of artificial prey populations. This work demonstrates that the foraging preferences of predators can regularly (though not always) result in the increase to fixation of a novel morph appearing in a population of familiar-colored prey. Such fixation events occur even if both novel and familiar prey are fully palatable and despite the novel food being much more conspicuous than the familiar prey. These studies therefore provide strong empirical evidence that conspicuous coloration can evolve readily, and repeatedly, as a result of the conservative foraging decisions of predators.     

Carotenoid-based colour polyphenism in a moth species: search for fitness correlates

Carotenoid‐based integumental coloration is often associated with individual performance in various animals. This is because the limited amount of the pigment has to be allocated to different vital functions. However, most of the evidence for the carotenoid‐based trade‐off comes from vertebrate studies, and it is unclear if this principle can be applied to insects. This possibility was investigated in Orgyia antiqua L. (Lepidoptera: Lymantriidae). The larvae of this species are polyphenic in their coloration, varying from a highly conspicuous combination of yellow hair tufts on black background to cryptic appearance with brown hair tufts. The conspicuous larvae are aposematic, advertising their aversive hairiness. The maintenance of different colour morphs in O. antiqua requires explanation, as an aposematic signal is expected to evolve towards monomorphism. Chromatographic analysis showed that the yellow coloration of the hair is based on the carotenoid pigment lutein (α‐carotene‐3,3’‐diol). The colour of hair tufts was dependent on their carotenoid content. This justifies an expectation of carotenoid‐based physiological trade‐offs between aposematic coloration and individual performance. To test this hypothesis, we monitored life histories of differently coloured larvae reared on various host plants, recording their body sizes, growth rates, and mortalities in each instar. There was a significant but relatively low heritability of tuft coloration, which allowed us to expect environmental effects. We found no phenotypic associations between hair tuft colour and performance indices in O. antiqua larvae, neither did the quality of host plant affect the frequency of colour morphs. However, the frequency of colour morphs differed between larval instars. Our results suggest that carotenoid‐mediated physiological trade‐offs are not involved in the maintenance of colour morphs in O. antiqua larvae, and factors other than individual condition should be responsible for the observed variability.     

EVOLUTION OF GREGARIOUSNESS IN APOSEMATIC BUTTERFLY LARVAE: A PHYLOGENETIC ANALYSIS

Gregariousness ought to be disadvantageous for palatable organisms that live exposed and are relatively immobile and small in comparison to potential predators. Therefore, the idea that unpalatability generally evolves before egg clustering/larval gregariousness in butterflies was tested. Aposematic coloration in the larva was used as the criterion of unpalatability (it is argued that Batesian mimicry is rare in butterfly larvae), and the relative order of evolution of aposematism and gregariousness was inferred through phylogenetic analysis. Here, existing phylogenies were used, and the analysis was based on an assumption of a minimum number of evolutionary changes (parsimony). A total of 23 cases of independent evolution of gregariousness and 12 cases of independent evolution of aposematic coloration were found. In five cases, gregariousness evolved in cryptic species, the palatability of which is unknown. For lineages in which both unpalatability, as evidenced by aposematic coloration, and gregariousness were found and the two evolutionary events could be separated, unpalatability always preceded gregariousness: five cases of independent evolution of warning coloration were followed by a total of 15 cases of independent evolution of gregariousness. In no lineage did gregariousness evolve before warning coloration. It is thus concluded that unpalatability is an important predisposing factor for the evolution of egg clustering and larval gregariousness in butterflies. Insofar as kin selection is related to larval gregariousness, this study indicates that kin selection is of minor importance for the evolution of both unpalatability and warning coloration.     

Evolution of color variation in dragon lizards: quantitative tests of the role of crypsis and local adaptation

Abstract Many animal species display striking color differences with respect to geographic location, sex, and body region. Traditional adaptive explanations for such complex patterns invoke an interaction between selection for conspicuous signals and natural selection for crypsis. Although there is now a substantial body of evidence supporting the role of sexual selection for signaling functions, quantitative studies of crypsis remain comparatively rare. Here, we combine objective measures of coloration with information on predator visual sensitivities to study the role of crypsis in the evolution of color variation in an Australian lizard species complex (Ctenophorus decresii). We apply a model that allows us to quantify crypsis in terms of the visual contrast of the lizards against their natural backgrounds, as perceived by potential avian predators. We then use these quantitative estimates of crypsis to answer the following questions. Are there significant differences in crypsis conspicuousness among populations? Are there significant differences in crypsis conspicuousness between the sexes? Are body regions “exposed” to visual predators more cryptic than “hidden” body regions? Is there evidence for local adaptation with respect to crypsis against different substrates? In general, our results confirmed that there are real differences in crypsis conspicuousness both between populations and between sexes; that exposed body regions were significantly more cryptic than hidden ones, particularly in females; and that females, but not males, are more cryptic against their own local background than against the background of other populations. Body regions that varied most in contrast between the sexes and between populations were also most conspicuous and are emphasized by males during social and sexual signaling. However, results varied with respect to the aspect of coloration studied. Results based on chromatic contrast (“hue’ of color) provided better support for the crypsis hypothesis than did results based on achromatic contrast (“brightness’ of color). Taken together, these results support the view that crypsis plays a substantial role in the evolution of color variation and that color patterns represent a balance between the need for conspicuousness for signaling and the need for crypsis to avoid predation.     

The interplay between multiple predators and prey colour divergence

Evolutionary divergence in the coloration of toxic prey is expected when geographic variation in predator composition and behavior favours shifts in prey conspicuousness. A fundamental prediction of predator‐driven colour divergence is that the local coloration should experience lower predation risk than novel prey phenotypes. The dorsal coloration of the granular poison frog varies gradually from populations of conspicuous bright red frogs to populations of dull green and relatively cryptic frogs. We conducted experiments with clay models in four populations to examine the geographic patterns of taxon‐specific predation. Birds avoided the local phenotype while lizards consistently selected for decreased conspicuousness and crab predation did not depend on frog coloration. Importantly, birds and lizards favoured low conspicuousness in populations where relatively cryptic green morphs have evolved. This study provides evidence for the interplay among distinct selective pressures, from multiple‐predator taxa, acting on the divergence in protective coloration of prey species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 580–589.     

The model of early evolution of aposematic coloration

First stages of evolution of aposematic coloration include a region of negative selection. During these stages, individuals with aberrant coloration remain to be rare, while predators are still not able to associate coloration with unpalatability. The simulation model is proposed, in which this "problematic zone" is overcome by individual selection for the increasing of unpalatable prey conspicuity in a small unisexual population. It is shown that under this assumption aposematic coloration develops within a wide range of parameters such as the cost of unpalatability, the cost of coloring, the survival rate of unpalatable prey after being attacked by na?ve predator, the probability of discovering of differently colored preys by predator as well as the predator's learning rate and memory depth. Thus, the early evolution ofaposematic coloration does not require any unusual or unique set of circumstances; aposematic coloration along with concomitant Bates mimicry inevitably evolve within a wide range of initial conditions. The loss of cryptic coloration by the original form (e.g., due to a change of food preferences, and thereby the structure of a background coloring, changes in habitat structure, color mutations etc.) is one such condition.     

Non-warning odors trigger innate color aversions--as long as they are novel

Warning signals made by unpalatable insects to potential predatorscommonly target more than one sense: such signals are "multimodal." Pyrazines are odors produced by warningly colored insects whenattacked, and have been shown to interact with food coloration,biasing avian predators against novel and typically aposematicfood. However, at present it is not known whether this is anadaptation by prey to exploit a general feature of avian psychology,or an evolutionary response by birds to enhance their avoidanceof unpalatable prey. Here we investigate the effect of otherodors on the innate responses of naive domestic chicks (Gallusgallus domesticus) to food that is of novel color, or of acolor that is associated with warning coloration, yellow. Inthe first experiment, we demonstrate that natural and artificialodors that have no association with aposematism in the wildcan produce biases against both novel colored foods and yellowcolored foods. In a second experiment, we also show that odor novelty is vital for eliciting such effects. These results supportthe idea that warning odors have evolved in response to preexistingpsychological biases against novel odors in predators, ratherthan predators evolving specific responses against odors associatedwith unpalatable prey.