Prey from the eyes of predators: Color discriminability of aposematic and mimetic butterflies from an avian visual perspective

Predation exerts strong selection on mimetic butterfly wing color patterns, which also serve other functions such as sexual selection. Therefore, specific selection pressures may affect the sexes and signal components differentially. We tested three predictions about the evolution of mimetic resemblance by comparing wing coloration of aposematic butterflies and their Batesian mimics: (a) females gain greater mimetic advantage than males and therefore are better mimics, (b) due to intersexual genetic correlations, sexually monomorphic mimics are better mimics than female‐limited mimics, and (c) mimetic resemblance is better on the dorsal wing surface that is visible to predators in flight. Using a physiological model of avian color vision, we quantified mimetic resemblance from predators’ perspective, which showed that female butterflies were better mimics than males. Mimetic resemblance in female‐limited mimics was comparable to that in sexually monomorphic mimics, suggesting that intersexual genetic correlations did not constrain adaptive response to selection for female‐limited mimicry. Mimetic resemblance on the ventral wing surface was better than that on the dorsal wing surface, implying stronger natural and sexual selection on ventral and dorsal surfaces, respectively. These results suggest that mimetic resemblance in butterfly mimicry rings has evolved under various selective pressures acting in a sex‐ and wing surface‐specific manner.     

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.     

Alternative prey can change model–mimic dynamics between parasitism and mutualism

Classical (conventional) Müllerian mimicry theory predicts that two (or more) defended prey sharing the same signal always benefit each other despite the fact that one species can be more toxic than the other. The quasi‐Batesian (unconventional) mimicry theory, instead, predicts that the less defended partner of the mimetic relationship may act as a parasite of the signal, causing a fitness loss to the model. Here we clarify the conditions for parasitic or mutualistic relationships between aposematic prey, and build a model to examine the hypothesis that the availability of alternative prey is crucial to Müllerian and quasi‐Batesian mimicry. Our model is based on optimal behaviour of the predator. We ask if and when it is in the interest of the predator to learn to avoid certain species as prey when there is alternative (cryptic) prey available. Our model clearly shows that the role of alternative prey must be taken into consideration when studying model–mimic dynamics. When food is scarce it pays for the predator to test the models and mimics, whereas if food is abundant predators should leave the mimics and models untouched even if the mimics are quite edible. Dynamics of the mimicry tend to be classically Müllerian if mimics are well defended, while quasi‐Batesian dynamics are more likely when they are relatively edible. However, there is significant overlap: in extreme cases mimics can be harmful to models (a quasi‐Batesian case) even if the species are equally toxic. A crucial parameter explaining this overlap is the search efficiency with which indiscriminating vs. discriminating predators find cryptic prey. Quasi‐Batesian mimicry becomes much more likely if discrimination increases the efficiency with which the specialized predator finds cryptic prey, while the opposite case tends to predict Müllerian mimicry. Our model shows that both mutualistic and parasitic relationship between model and mimic are possible and the availability of alternative prey can easily alter this relationship.     

Cryptic differences in colour among Müllerian mimics: how can the visual capacities of predators and prey shape the evolution of wing colours?

Antagonistic interactions between predators and prey often lead to co‐evolution. In the case of toxic prey, aposematic colours act as warning signals for predators and play a protective role. Evolutionary convergence in colour patterns among toxic prey evolves due to positive density‐dependent selection and the benefits of mutual resemblance in spreading the mortality cost of educating predators over a larger prey assemblage. Comimetic species evolve highly similar colour patterns, but such convergence may interfere with intraspecific signalling and recognition in the prey community, especially for species involved in polymorphic mimicry. Using spectrophotometry measures, we investigated the variation in wing coloration among comimetic butterflies from distantly related lineages. We focused on seven morphs of the polymorphic species Heliconius numata and the seven corresponding comimetic species from the genus Melinaea. Significant differences in the yellow, orange and black patches of the wing were detected between genera. Perceptions of these cryptic differences by bird and butterfly observers were then estimated using models of animal vision based on physiological data. Our results showed that the most strikingly perceived differences were obtained for the contrast of yellow against a black background. The capacity to discriminate between comimetic genera based on this colour contrast was also evaluated to be higher for butterflies than for birds, suggesting that this variation in colour, likely undetectable to birds, might be used by butterflies for distinguishing mating partners without losing the benefits of mimicry. The evolution of wing colour in mimetic butterflies might thus be shaped by the opposite selective pressures exerted by predation and species recognition.     

Imperfect Batesian mimicry—the effects of the frequency and the distastefulness of the model

Batesian mimicry is the resemblance between unpalatable models and palatable mimics. The widely accepted idea is that the frequency and the unprofitability of the model are crucial for the introduction of a Batesian mimic into the prey population. However, experimental evidence is limited and furthermore, previous studies have considered mainly perfect mimicry (automimicry). We investigated imperfect Batesian mimicry by varying the frequency of an aposematic model at two levels of distastefulness. The predator encountered prey in a random order, one prey item at a time. The prey were thus presented realistically in a sequential way. Great tits (Parus major) were used as predators. This experiment, with a novel signal, supports the idea that Batesian mimics gain most when the models outnumber them. The mortalities of the mimics as well as the models were significantly dependent on the frequency of the model. Both prey types survived better the fewer mimics there were confusing the predator. There were also indications that the degree of distastefulness of the model had an effect on the survival of the Batesian mimic: the models survived significantly better the more distasteful they were. The experiment supports the most classical predictions in the theories of the origin and maintenance of Batesian mimicry.     

Can experienced birds select for Mullerian mimicry?

Field experiments have shown that avian predators in the wildcan select for similarity of warning signals in aposematic prey(Müllerian mimicry) because a common signal is better protectedthan a signal that is novel and rare. The original theory ofMüllerian mimicry assumes that the mechanism promotingmimicry is predator learning; by sharing a signal, the comimicspecies share the mortality that is due to sampling by inexperiencedpredators. Predation events have not been observed in the wild,and learning experiments with naive bird predators in a laboratoryhave not unambiguously shown a benefit of a uniform signal comparedwith different signals. As predators in the field experimentsare likely to be more experienced compared with previous laboratoryexperiments, we studied selection by experienced predators ona novel imperfect mimic. We trained great tits Parus major toavoid artificial aposematic models and subsequently introducedperfect and imperfect mimics at different frequencies. Birdswith prior experience on the models selected against the imperfectmimics that were at a disadvantage also in a memory test conducteda week after their introduction. Selection against the imperfectmimics was antiapostatic. However, the imperfect mimics alsobenefited from some signal generalization to the models andpossibly gained protection because the birds were familiar withthe alternative cryptic prey that was also present. Our resultssuggest that experienced predators might be more important tothe evolution of mimicry than the learning-based theory assumes.     

Five species of passerine bird differ in their ability to detect Batesian mimics

Multiple predators affect the evolution of aposematic signals in nature and these predators may substantially differ in terms of ecological and cognitive parameters. However, most experimental studies testing the evolution of Batesian mimics use only a single species of predator (usually the great tit or a domestic chick). Therefore, in the present study, we experimentally tested the responses of five passerine predators to an artificially made Batesian mimic (a cockroach equipped with the warning pattern of the red firebug) with respect to their dietary ecology. Half of the individuals of each species were fed on unmodified roaches before the experiment, whereas the other half were fed with mealworms and thus had no previous experience with roaches. We found that Batesian mimics were better protected than inconspicuous prey against inexperienced great tits and robins alone. The other three bird species showed high level of neophobia; therefore, the effect of warning coloration could not be assessed. We also found that experienced birds attacked a greater number of Batesian mimics compared to inexperienced individuals of all tested species, with the exception of blackcaps. In the great tits, robins, and blue tits, a significant number of experienced birds attacked the Batesian mimic, which was possibly the result of a learned search image for a roach. Our results suggest that using a limited array of predators to describe evolutionary processes forming the diversity of antipredatory strategies of the prey may be biased and need not describe the situation occurring in nature.     

INTERSEXUAL COMPARISON OF MIMETIC PROTECTION IN THE BLACK SWALLOWTAIL BUTTERFLY,PAPILIO POLYXENES: EXPERIMENTS WITH CAPTIVE BLUE JAY PREDATORS

The black swallowtail butterfly (Papilio polyxenes asterius Stoll), is commonly assumed to exhibit female-limited Batesian mimicry of the aposematic pipevine swallowtail (Battus philenor [L.]), since the dorsal wing surfaces of P. polyxenes females, but not males, resemble those of the model. However, the ventral wing surface is monomorphic and closely resembles that of the model in both sexes. Thus both sexes of P. polyxenes should benefit from mimicry during periods of ventral surface exposure, such as during overnight roosting and other times of high predatory risk. Eight blue jays (Cyanocitta cristata L.) were offered ventrally and dorsally exposed butterfly prey items in an outdoor aviary. Model-conditioned birds refused male and female P. polyxenes equally when the butterflies were presented ventrally. However, significantly more males than females were attacked when the dimorphic dorsum was visible. Both sexes are thus similarly protected when the ventral wing surface is displayed during roosting. The high degree of bird-to-bird variability in response to P. polyxenes mimics suggests that there is a spectrum in ability or willingness of predators to discriminate among mimics of varying similarity to the model. Sexual dimorphism of the dorsal surface of P. polyxenes wings may reflect sexual selection favoring males that are recognizable as satisfactory mates or intrasexual competitors.     

Sensory discrimination and the incipient advantage of mutations

Perception follows logarithmic or power functions, rather than linear functions, of stimulus intensity. A small increment in stimulus strength can be sufficient to elicit discrimination between individuals of different phenotypes when the initial stimulus magnitude is near zero. This may confer an incipient advantage to the mutation that caused it. The psychophysics of signal receivers may be a good predictor of the extent of phenotypic changes. For example, slight similarities in color of Batesian mimics to their distasteful model can be sufficient to cause predators to reject the mimics. Color changes produced by single mutations in incipient Batesian mimics should be more extensive in mimicry complexes where the predators are less sensitive to color differences.     

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.     

Prey community structure affects how predators select for Mullerian mimicry

Müllerian mimicry describes the close resemblance between aposematic prey species; it is thought to be beneficial because sharing a warning signal decreases the mortality caused by sampling by inexperienced predators learning to avoid the signal. It has been hypothesized that selection for mimicry is strongest in multi-species prey communities where predators are more prone to misidentify the prey than in simple communities. In this study, wild great tits (Parus major) foraged from either simple (few prey appearances) or complex (several prey appearances) artificial prey communities where a specific model prey was always present. Owing to slower learning, the model did suffer higher mortality in complex communities when the birds were inexperienced. However, in a subsequent generalization test to potential mimics of the model prey (a continuum of signal accuracy), only birds that had foraged from simple communities selected against inaccurate mimics. Therefore, accurate mimicry is more likely to evolve in simple communities even though predator avoidance learning is slower in complex communities. For mimicry to evolve, prey species must have a common predator; the effective community consists of the predator's diet. In diverse environments, the limited diets of specialist predators could create 'simple community pockets' where accurate mimicry is selected for.     

Protection by association: evidence for aposematic commensalism

Aposematism is a well known and widely used strategy for reducing predation by conspicuous signalling of unprofitability. However, the increased conspicuousness could make this strategy costly if there are no secondary defences to back the signal up. This has made the elucidation of the evolutionary mechanisms for aposematism and that of the closely‐related Batesian and Mullerian mimicry difficult. The present study aims to test whether cryptic and nondefended prey could reduce their predation risk by grouping with aposematic and defended prey. To do this, we used groups of artificial baits that were either cryptic and palatable or conspicuous and unpalatable, along with the corresponding control treatments. These were then presented in mixed and homogeneous treatment groups within a field setting and the local wild bird assemblage was allowed to select and remove baits at will. The results obtained show that undefended non‐aposematic prey can benefit by grouping with aposematic prey, with no evidence that predation rates for aposematic prey were adversely affected by this association. These results provide insights into the evolution of Batesian mimicry. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 81–89.     

A small badge of longevity: opposing survival selection on the size of white and black wing markings

According to handicap principle, exaggerated ornamental traits are supposed to exert costs on their bearers. However, there is much less theoretical and practical consensus about whether and under which conditions ornament expression should positively correlate with survival. We measured age‐related variation and survival selection on the size of white wing patches and black wing tips in a long‐lived monogamous seabird, the common gull Larus canus. Males had larger white patches than females but patch size showed concave relationship with age irrespective of sex, suggesting that white patch size was prone to senescence in both sexes. Extent of wing tip abrasion correlated negatively with the size of white patch, suggesting, in agreement with the Zahavian handicap hypothesis that only individuals with largest ornaments are able of maintaining them and not paying cost of displaying them. Areas of white wing patches and black wing tips correlated negatively. Irrespective of sex, survival selection favored birds with larger white wing patches and smaller black wing tips, which suggests that white and black wing markings may have coevolved as reverse components of a single ornament. Altogether, our results provide an evidence for the case where survival selection on ornamental traits in females is not weaker than in males. Absence of sex differences with respect to most of observed patterns is consistent with a prediction that among monogamous long‐lived species with biparental care, mutual mate choice leads to evolution of elaborate ornamental traits in both sexes.     

EVIDENCE FOR BATESIAN MIMICRY IN A POLYMORPHIC HOVERFLY

Palatable Batesian mimics are avoided by predators because they resemble noxious or defended species. The striking resemblance of many hoverflies to noxious Hymenoptera is a “textbook” example of Batesian mimicry, but evidence that selection by predators has shaped the evolution of hoverfly patterns is weak. We looked for geographical and temporal trends in frequencies of morphs of the polymorphic hoverfly Volucella bombylans that would support the hypothesis that these morphs are Batesian mimics of different bumblebee species. The frequency of the black and yellow hoverfly morph was significantly positively related to the frequency of black and yellow bumblebees across 52 sites. Similarly, the frequency of the red‐tailed hoverfly morph was positively related to the frequency of red‐tailed bumblebees. However, the frequencies of hoverfly morphs were positively spatially autocorrelated, and after controlling for this, only one of the two common hoverfly morphs showed a significant positive relationship with its putative model. We conclude that the distribution of V. bombylans morphs probably reflects geographical variation in selection by predators resulting from differences in the frequencies of noxious bumblebee species.     

PREDATOR PERCEPTION OF BATESIAN MIMICRY AND CONSPICUOUSNESS IN A SALAMANDER

In Batesian mimicry a palatable mimic deceives predators by resembling an unpalatable model. The evolution of Batesian mimicry relies on the visual capabilities of the potential predators, as prey detection provides the selective force driving evolutionary change. We compared the visual capabilities of several potential predators to test predictions stemming from the hypothesis of Batesian mimicry between two salamanders: the model species Notophthalmus viridescens, and polymorphic mimic, Plethodon cinereus. First, we found mimicry to be restricted to coloration, but not brightness. Second, only bird predators appeared able to discriminate between the colors of models and nonmimic P. cinereus. Third, estimates of salamander conspicuousness were background dependent, corresponding to predictions only for backgrounds against which salamanders are most active. These results support the hypothesis that birds influence the evolution of Batesian mimicry in P. cinereus, as they are the only group examined capable of differentiating N. viridescens and nonmimetic P. cinereus. Additionally, patterns of conspicuousness suggest that selection from predators may drive the evolution of conspicuousness in this system. This study confirms the expectation that the visual abilities of predators may influence the evolution of Batesian mimicry, but the role of conspicuousness may be more complex than previously thought.     

The effect of alternative prey on the dynamics of imperfect Batesian and Müllerian mimicries

Both Batesian and Müllerian mimicries are considered classical evidence of natural selection where predation pressure has, at times, created a striking similarity between unrelated prey species. Batesian mimicry, in which palatable mimics resemble unpalatable aposematic species, is parasitic and only beneficial to the mimics. By contrast, in classical Müllerian mimicry the cost of predators' avoidance learning is shared between similar unpalatable co-mimics, and therefore mimicry benefits all parties. Recent studies using mathematical modeling have questioned the dynamics of Müllerian mimicry, suggesting that fitness benefits should be calculated in a way similar to Batesian mimicry; that is, according to the relative unpalatability difference between co-mimics. Batesian mimicry is very sensitive to the availability of alternative prey, but the effects of alternative prey for Müllerian dynamics are not known and experiments are rare. We designed two experiments to test the effect of alternative prey on imperfect Batesian and Müllerian mimicry complexes. When alternative prey were scarce, imperfect Batesian mimics were selected out from the population, but abundantly available alternative prey relaxed selection against imperfect mimics. Birds learned to avoid both Müllerian models and mimics irrespective of the availability of alternative prey. However, the rate of avoidance learning of models increased when alternative prey were abundant. This experiment suggests that the availability of alternative prey affects the dynamics of both Müllerian and Batesian mimicry, but in different ways.     

Subtle variation in size and shape of the whole forewing and the red band among co‐mimics revealed by geometric morphometric analysis in Heliconius butterflies

Heliconius are unpalatable butterflies that exhibit remarkable intra‐ and interspecific variation in wing color pattern, specifically warning coloration. Species that have converged on the same pattern are often clustered in Müllerian mimicry rings. Overall, wing color patterns are nearly identical among co‐mimics. However, fine‐scale differences exist, indicating that factors in addition to natural selection may underlie wing phenotype. Here, we investigate differences in shape and size of the forewing and the red band in the Heliconius postman mimicry ring (H. erato phyllis and the co‐mimics H. besckei, H. melpomene burchelli, and H. melpomene nanna) using a landmark‐based approach. If phenotypic evolution is driven entirely by predation pressure, we expect nonsignificant differences among co‐mimics in terms of wing shape. Also, a reinforcement of wing pattern (i.e., greater similarity) could occur when co‐mimics are in sympatry. We also examined variation in the red forewing band because this trait is critical for both mimicry and sexual communication. Morphometric results revealed significant but small differences among species, particularly in the shape of the forewing of co‐mimics. Although we did not observe greater similarity when co‐mimics were in sympatry, nearly identical patterns provided evidence of convergence for mimicry. In contrast, mimetic pairs could be distinguished based on the shape (but not the size) of the red band, suggesting an “advergence” process. In addition, sexual dimorphism in the red band shape (but not size) was found for all lineages. Thus, we infer that natural selection due to predation by birds might not be the only mechanism responsible for variation in color patterns, and sexual selection could be an important driver of wing phenotypic evolution in this mimicry ring.     

REVISING A CLASSIC BUTTERFLY MIMICRY SCENARIO: DEMONSTRATION OF MÜLLERIAN MIMICRY BETWEEN FLORIDA VICEROYS (LIMENITIS ARCHIPPUS FLORIDENSIS) AND QUEENS (DANAUS GILIPPUS BERENICE)

Batesian and Müllerian mimicry relationships differ greatly in terms of selective pressures affecting the participants; hence, accurately characterizing a mimetic interaction is a crucial prerequisite to understanding the selective milieux of model, mimic, and predator. Florida viceroy butterflies (Limenitis archippus floridensis) are conventionally characterized as palatable Batesian mimics of distasteful Florida queens (Danaus gilippus berenice). However, recent experiments indicate that both butterflies are moderately distasteful, suggesting they may be Müllerian comimics. To directly test whether the butterflies exemplify Müllerian mimicry, I performed two reciprocal experiments using red-winged blackbird predators. In Experiment 1, each of eight birds was exposed to a series of eight queens as “models,” then offered four choice trials involving a viceroy (the putative “mimic”) versus a novel alternative butterfly. If mimicry was effective, viceroys should be attacked less than alternatives. I also compared the birds' reactions to solo viceroy “mimics” offered before and after queen models, hypothesizing that attack rate on the viceroy would decrease after birds had been exposed to queen models. In Experiment 2, 12 birds were tested with viceroys as models and queens as putative mimics. The experiments revealed that (1) viceroys and queens offered as models were both moderately unpalatable (only 16% entirely eaten), (2) some birds apparently developed conditioned aversions to viceroy or queen models after only eight exposures, (3) in the subsequent choice trials, viceroy and queen “mimics” were attacked significantly less than alternatives, and (4) solo postmodel mimics were attacked significantly less than solo premodel mimics. Therefore, under these experimental conditions, sampled Florida viceroys and queens are comimics and exemplify Müllerian, not Batesian, mimicry. This compels a reassessment of selective forces affecting the butterflies and their predators, and sets the stage for a broader empirical investigation of the ecological and evolutionary dynamics of mimicry.     

Polymorphic Müllerian mimicry and interactions with thermal melanism in ladybirds and a soldier beetle: a hypothesis

It is argued that groups of similarly coloured species of coccinellids are Müllerian mimicry rings. This is based on a synthesis of the literature about the nature of their biology and aposematic colour patterns, their highly developed chemical defence and the responses of bird predators to them. The system of multiple mimicry ‘rings’ is illustrated for the Dutch coccinellid fauna. Some polymorphic species, including Adalia, exhibit red forms and black melanic forms which are apparently components of different putative mimicry rings. A similar reasoning is put forward with regard to the orange and the black forms of the soldier beetle Cuntharis livida. Hypotheses involving spatial variation in comimics, as have been developed to account for some other cases of polymorphic Miillerian mimicry, predict that sympatric polymorphic species exhibiting similar sets of phenotypes will show parallels in their geographical variation. This is tested for A. bipunctata and A. decempunctata in The Netherlands. On this local scale there is no parallel variation; A. bipunctata exhibits marked geographical differentiation whereas A. decempunctata shows a general uniformity in morph frequency. Observations on their population biology show that only in A. bipunctata is there a major spring period of adult reproduction on shrubs exposed to direct sunshine. Previous work has demonstrated an influence of thermal melanism in this period of the life cycle. It is suggested that local responses in species such as A. bipunctata may reflect a partial ‘escape’ from stabilizing aposematic selection. The basis of a steep cline found in C. livida, which opposes one in A. bipunctata, is unknown and unlikely to be related to mimicry. There is some evidence that the polymorphism is influenced by non-random mating. When species and communities of coccinellids are considered on a wide geographical scale many observations about their colour patterns and spatial variation, especially those of Dobzhansky, support an interaction between selection favouring mimetic resemblance and forms of climatic selection, especially thermal melanism. The polymorphism in Adalia is discussed in relation to a system of multiple mimicry rings and to Thompson's recent theoretical treatment of the maintenance of some polymorphisms for warning coloration by a balance between aposematic and apostatic selection. This becomes more tenable in coccinellids because of evidence that bird predators show a variable response to them. Frequency-independent selection arising from thermal melanism can provide the basis of spatial variation in equilibrium points. An alternative to such a hypothesis is one in which differences in unpalatability between species of coccinellids are emphasized (after experiments of Pasteels and colleagues). Some less unpalatable species such as Adalia may have responded to periods of prolonged disruptive selection acting in a frequency-dependent way to promote polymorphic mimicry associated with different modal colour patterns and intermediate in nature between classical Batesian and Müllerian mimicry. The likely occurrence of a supergene controlling polymorphism in some coccinellids is consistent with such an explanation.     

Visual modelling suggests a weak relationship between the evolution of ultraviolet vision and plumage coloration in birds

Birds have sophisticated colour vision mediated by four cone types that cover a wide visual spectrum including ultraviolet (UV) wavelengths. Many birds have modest UV sensitivity provided by violet‐sensitive (VS) cones with sensitivity maxima between 400 and 425 nm. However, some birds have evolved higher UV sensitivity and a larger visual spectrum given by UV‐sensitive (UVS) cones maximally sensitive at 360–370 nm. The reasons for VS–UVS transitions and their relationship to visual ecology remain unclear. It has been hypothesized that the evolution of UVS‐cone vision is linked to plumage colours so that visual sensitivity and feather coloration are ‘matched’. This leads to the specific prediction that UVS‐cone vision enhances the discrimination of plumage colours of UVS birds while such an advantage is absent or less pronounced for VS‐bird coloration. We test this hypothesis using knowledge of the complex distribution of UVS cones among birds combined with mathematical modelling of colour discrimination during different viewing conditions. We find no support for the hypothesis, which, combined with previous studies, suggests only a weak relationship between UVS‐cone vision and plumage colour evolution. Instead, we suggest that UVS‐cone vision generally favours colour discrimination, which creates a nonspecific selection pressure for the evolution of UVS cones.