(2005). Five replicate measurements were taken from randomly selected regions of each sample, recorded in 1 nm intervals from 300 to 700 nm, and expressed relative to a Spectralon 99% white reflectance standard (Labsphere, Congleton, UK). Fourth, it is at present uncertain whether luminance-based tasks in birds are subserved by the double cones or a weighted sum of single cone output (Jones & Osorio 2004; Osorio & Vorobyev 2005). Furthermore, targets with highly contrasting disruptive patterns survived significantly better than equivalent low-contrast patterns, supporting the disruptive contrast hypothesis. There are many different ways animals and insects can blend in with their surroundings. 6), the prediction was that the non-matching disruptive treatment would have the highest survival of all since these two colour elements had the highest contrast and therefore the strongest disruptive effect. (2005) cannot be considered a test of what we might call the strong theory of maximum disruptive contrast. Long gaps without reduced survival correspond to overnight, when targets were not checked. (2001)) of a typical woodland passerine bird, the blue tit's (Parus caeruleus) double cone photoreceptors (Hart et al. However, it has been argued that concealment may be achieved by strategic use of apparently conspicuous markings. Another factor to bear in mind is that because the mealworm was placed on top of each target, the contrast between the patterns on the ‘wings’ may also be important. In disruptive coloration, the identity and location of a species may be disguised through a coloration pattern. Disruptive eye masks camouflage the eyes of a variety of animals, both invertebrates such as grasshoppers and vertebrates such as fishes, frogs, birds and snakes; some mammals have similar patterns. This is a band of color found on the bodies of birds, fish, and other creatures that conceals the eye, which is usually easy to spot because of its distinctive shape. Coincident disruptive coloration or coincident disruptive patterns are patterns of disruptive coloration in animals that go beyond the usual camouflage function of breaking up the continuity of an animal's shape, to join up parts of the body that are separate. It introduces the different types of camouflage and how they work, including background matching, disruptive coloration and obliterative shading. & Troscianko, T. In press, Stevens, M., Párraga, C. A., Cuthill, I. C., Partridge, J. C. & Troscianko, T. S. In press, Animal coloration: an account of the principal facts and theories relating to the colours and markings of animals, Disruptive coloration and background pattern matching, The predation costs of symmetrical cryptic coloration, A predator's view of animal color patterns, An overview of the relationships between mimicry and crypsis, Progressive background matching in moths, and a quantitative measure of crypsis, Frequency-dependent predation, crypsis and aposematic coloration, The color of light in forests and its implications, Comparing entire colour patterns as birds see them, The visual ecology of avian photoreceptors, Visual pigments, oil droplets, ocular media and cone photoreceptor distribution in two species of passerine: the blue tit (, Discrimination of orientated visual textures by poultry chicks, Animal colour vision—behavioural tests and physiological concepts, Survival analysis: techniques for censored and truncated data, Statistical models and methods for lifetime data, The effects of the light environment on prey choice by zebra finches, Crypsis through disruptive coloration in an isopod, Background-matching and disruptive coloration, and the evolution of cryptic coloration, Optimization of cryptic coloration in heterogeneous habitats, Selection for cryptic coloration in a visually heterogeneous environment, Photoreceptor spectral sensitivities in terrestrial animals: adaptations for luminance and colour vision, Visual ecology and perception of colouration patterns by domestic chicks, The colours of animals: their meaning and use. 2005). 2005). Recently, we have accumulated approximately 5,000 images of animals of many taxa — large and small, wet and dry — but with emphasis on insects, fish, reptiles, amphibians, birds and primates. (2006) for more details). Thayer (1909) argued that disruptive coloration may allow animals found on a range of different backgrounds to achieve camouflage on each, and further, enable them to combine camouflage with other potentially conspicuous forms of coloration (such as warning colours and sexually selected colour patterns). in press), and that the inside treatment can result in straight lines at the edge of the body which may aid detection (Cuthill et al. Disruptive colouration provides camouflage independent of background matching. A classic example of this method is a cheetah whose spots can camouflage them in grass, not only to protect them from prey, but … In press. Avian predation was revealed by complete or nearly complete disappearance of the mealworm. Each block took place on different days and in different locations of the field site, so as to minimize any learning and search image effects. It also accords with Thayer's (1909) idea that disruptive coloration may be a particularly important method of concealment in species which are found on a range of backgrounds and so cannot be perfectly matched to any one situation (see also compromise crypsis; Merilaita et al. Most animals are under persistent risk of predation, and many animals have evolved a range of defensive colour patterns, of which a primary example is camouflage. These were not intended to mimic any real species of moth, but the markings were designed to match the visual texture of mature oak bark. These factors are effective in disruptive camouflage because they exploit edge detection mechanisms that function in early visual processing (Stevens & Cuthill 2006). Survival analysis for both experiments was performed with Cox proportional hazards regression (Cox 1972; Lawless 2002; Klein & Moeschberger 2003), which is ideally suited for censored data and the non-uniform change in predation risk with respect to time of day that are evident in such data (Cuthill et al. This indicates that, as Thayer (1909) and Cott (1940) predicted, disruptive patterns are still effective when some of the pattern elements do not match the background (see also Schaefer & Stobbe (in press)), although it does appear that disruptive patterns are maximally effective when all components match elements present in the background. The disruptive and background-matching treatments had higher survival when both the pattern elements matched the background luminance than when the grey component was brighter than the background. We are grateful to Martin Schaefer Research Group for access to their unpublished manuscript and strongly recommend that readers consult their paper and the accompanying commentary article. Therefore, concealment may still be achieved even when an animal possesses markings not found in the background. Disruptive coloration may enable animals to exploit backgrounds and environments towards which they have only a partial resemblance, and to bear conspicuous markings without paying the full cost of reduced crypsis. The markings necessarily have high contrast and are thus in themselves conspicuous. There were six treatments (figure 1): (a) edge matching, where both the black (4% cone-catch) patches and grey (12% cone-catch) patches matched avian perceived luminance values of oak bark/shadow, and the dark patches overlapped the edge of the ‘wings’ in a disruptive fashion; (b) inside matching, where the colour patches matched the bark/shadow luminance as for the previous treatment, and the shape of the pattern elements matched those in the oak background (as did their general distribution), except that the dark pattern elements did not overlap the wing edges (non-disruptive); (c) edge non-matching (disruptive), where the dark markings (4% cone-catch) matched the luminance of shadow and were placed at the edge of the wings, but the grey components were much lighter than bark (31% cone-catch); (d) inside non-matching (non-disruptive), which had the same luminance patches as treatment (c) but the patches did not overlap the wing edges; (e) average matching, which was a uniformly coloured treatment (no background-matching pattern) where the luminance of the grey was the average of the matching treatment patch luminances (i.e. We’re going to explore five of them: color matching, disruptive coloration, self-decoration, active camouflage… Reflectance of bark (samples from n=30 trees from the study area), as described in Cuthill et al. Furthermore, targets with highly contrasting disruptive patterns survived significantly better than equivalent low-contrast patterns, supporting the disruptive contrast hypothesis. The black pattern components corresponded to a 4% cone-catch, the darkest black that we could print. Disruptive coloration (also known as disruptive camouflage or disruptive patterning) is a form of camouflage that works by breaking up the outlines of an animal, soldier or military vehicle with a strongly contrasting pattern. Figure 2 Survival plot of the experimental treatments (top to bottom: average matching (AM), average non-matching (ANM), edge matching (EM), edge non-matching (ENM), inside matching (IM), inside non-matching (INM)). In the experiment, each replicate for the two-tone treatments had a unique pattern. 1999a,b; Jones & Osorio 2004; Osorio & Vorobyev 2005). Therefore, it would be intriguing to know if there are differences in the survival of disruptively marked individuals and background-matching individuals when the targets are isoluminant and the differences in pattern are solely chromatic. In the experiment, each replicate for the two-tone treatments had a unique pattern.Download figureOpen in new tabDownload PowerPoint. First, when paired with black, light brown patches that contrast with the background would tend towards the common warning (aposematic) coloration of yellow and black. Camouflage is an important strategy in animals to prevent predation. There were several reasons for using greyscale targets. (2006), was measured normal to the image plane using a Zeiss MCS 230 UV–NIR diode array photometer, with illumination by a Zeiss CLX 111 Xenon lamp (Carl Zeiss Group, Jena, Germany) held at 45° to normal. Therefore, the chromatic contrast between the target and bark will have been higher in our non-matching treatments, just as with the luminance. a 50 : 50 blend of the grey values corresponding to the 4 and 12% cone-catches since the dark and light markings were found in approximately equal amounts on each moth); and (f) average non-matching, which was the average value of the non-matching treatment patterns (a 50 : 50 blend of grey values corresponding to the 4 and 31% cone-catch values). Large datasets are available through Proceedings B's partnership with Dryad. Different samples, from different trees, were used for each replicate target.Figure 1 Sample stimuli used in the experiment: (a) edge matching; (b) inside matching; (c) edge non-matching; (d) inside non-matching; (e) average matching (average of the two tones in the matching treatments); and (f) average non-matching (average of the two tones in the non-matching treatments). Especially considered in the case of insects, Disruptive coloration, crypsis and edge detection in early visual processing, Concealing-coloration in the animal kingdom: an exposition of the laws of disguise through color and pattern: being a summary of Abbott H. Thayer's discoveries, Aposematism and crypsis combined as a result of distance dependence: functional versatility of the colour pattern in the swallowtail butterfly larva, Tetrachromacy, oil droplets and bird plumage colours, Darwinism. Camouflage strategies interfere differently with observer search images, Contrast, contours and the confusion effect in dazzle camouflage, Disruptive camouflage impairs object recognition, Outline and surface disruption in animal camouflage, Cuttlefish camouflage: context-dependent body pattern use during motion, Behaviourally mediated crypsis in two nocturnal moths with contrasting appearance, Contrasting coloration in terrestrial mammals, Perception of edges and visual texture in the camouflage of the common cuttlefish, Sepia officinalis, Cephalopod dynamic camouflage: bridging the continuum between background matching and disruptive coloration, Concealed by conspicuousness: distractive prey markings and backgrounds, Defining disruptive coloration and distinguishing its functions, Chromaticity in the UV/blue range facilitates the search for achromatically background-matching prey in birds. In the experiment, each replicate for the two-tone treatments had a unique pattern.Download figureOpen in new tabDownload PowerPoint. 1998; Kelber et al. 6), the prediction was that the non-matching disruptive treatment would have the highest survival of all since these two colour elements had the highest contrast and therefore the strongest disruptive effect. If an animal or object possesses markings which match a random sample of the background (crypsis), then some markings will sometimes intersect the outline of the body in a disruptive fashion purely by chance. This was not the case; the disruptive treatment where both pattern elements matched the background, survived best. Enhancement of chromatic contrast increases predation risk for striped butterflies, Empirical tests of the role of disruptive coloration in reducing detectability, Predator perception and the interrelation between different forms of protective coloration, Perception of visual texture and the expression of disruptive camouflage by the cuttlefish, Sepia officinalis. However, when one colour element did not match the background, disruptive placement of colour patches did significantly improve survival over non-disruptive placement. It is notable that the mealworm–wing contrast in the average non-matching treatment (19.5 versus 17.5%) was far smaller than in the average matching treatment (19.5 versus 8%), and yet the survival of the latter treatment was higher (figure 1). It is notable that the mealworm–wing contrast in the average non-matching treatment (19.5 versus 17.5%) was far smaller than in the average matching treatment (19.5 versus 8%), and yet the survival of the latter treatment was higher (figure 1). Overall, our results provide further support for the theory of disruptive coloration, and show that it is a method of concealment far more resilient to potentially negative factors, such as non-background-matching components, than is crypsis alone. Because our targets did not match the colour of bark, the result that targets with two-tone background-matching patterns suffer less predation than monochrome targets without such patterns is an indirect demonstration that pattern detection in birds involves luminance information (Osorio et al. The raccoon butterfly fish uses the black and yellow coloration patterns to make it hard for a predator to single out an individual in a group. The mean perceived luminance (double cone photon catch) of the oak bark samples was 12% of that for the white standard. 2005). The above considerations, combined with the fact that the browns present in oak bark are relatively unsaturated colours, gave us confidence that our greyscale targets would be quite cryptic in those treatments where the grey matched the luminance of oak bark. Large datasets are available through Proceedings B's partnership with Dryad. The higher survival of disruptive (edge) treatments than the non-disruptive (inside) treatments of equivalent background-matching luminance confirms the findings of Cuthill et al. 2003). in press a). This makes it difficult for other animals such as lions to see it. Therefore, concealment may still be achieved even when an animal possesses markings not found in the background. However, it has been argued that concealment may be achieved by strategic use of apparently conspicuous markings. Our results also indicate that optimal concealment is likely to be achieved by twinning crypsis and disruptive coloration since both these forms of camouflage were effective in reducing the risk of predation compared to either non-matching or non-patterned treatments. Because the paper on which patterns were printed absorbs UV, the targets were not ‘bird grey’ (i.e. More specifically, Cott (1940) proposed two key tenets of disruptive theory: first, ‘differential blending’, where some patches on an individual stand out from the background while other patches blend in; and second, ‘maximum disruptive contrast’, where adjacent pattern elements are highly contrasting in tone and some are different from the background. Imperfect camouflage: how to hide in a variable world? This lies roughly as an intermediate between the luminance of the grey patches in the matching (12%) and non-matching (31%) treatments, suggesting that mealworm–wing luminance difference was similar across these treatments. Camouflage strategies interfere differently with observer search images, Contrast, contours and the confusion effect in dazzle camouflage, Disruptive camouflage impairs object recognition, Outline and surface disruption in animal camouflage, Cuttlefish camouflage: context-dependent body pattern use during motion, Behaviourally mediated crypsis in two nocturnal moths with contrasting appearance, Contrasting coloration in terrestrial mammals, Perception of edges and visual texture in the camouflage of the common cuttlefish, Sepia officinalis, Cephalopod dynamic camouflage: bridging the continuum between background matching and disruptive coloration, Concealed by conspicuousness: distractive prey markings and backgrounds, Defining disruptive coloration and distinguishing its functions, Chromaticity in the UV/blue range facilitates the search for achromatically background-matching prey in birds. Figure 2 Survival plot of the experimental treatments (top to bottom: average matching (AM), average non-matching (ANM), edge matching (EM), edge non-matching (ENM), inside matching (IM), inside non-matching (INM)). Animals showed two primary responses to the test stimuli: a disruptive body pattern (corresponding with a high disruptive score) on some, and a mottle/uniform body pattern (corresponding with a low disruptive score) on others (figure 2; one-way ANOVA for correlated samples: F 4,17 = 8.77, p < 0.0001). It also accords with Thayer's (1909) idea that disruptive coloration may be a particularly important method of concealment in species which are found on a range of backgrounds and so cannot be perfectly matched to any one situation (see also compromise crypsis; Merilaita et al. Disruptive coloration may allow animals to exploit backgrounds on which they are not perfectly matched, and to possess conspicuous markings while still retaining a degree of camouflage. However, the targets with the disruptive non-matching patterns survived as well as the non-disruptive matching treatment. The inside luminance matching treatment survived better than the inside non-matching treatment (W1=17.254, p<0.001). The research was supported by a BBSRC grant to I.C.C., T. Troscianko and J. C. Partridge, and by a BBSRC studentship to M.S. Using studies of both real animals and artificial systems, this book synthesises the current state of play in camouflage research and understanding. a 50 : 50 blend of the grey values corresponding to the 4 and 12% cone-catches since the dark and light markings were found in approximately equal amounts on each moth); and (f) average non-matching, which was the average value of the non-matching treatment patterns (a 50 : 50 blend of grey values corresponding to the 4 and 31% cone-catch values). 2005). Calibration of the printer was necessary because a printer's greyscale (here 8 bit, or 0–255) is rarely linearly related to perceived luminance for a human, far less a bird (Westland & Ripamonti 2004; Stevens et al. In all, 266 replicates were censored in the analysis (36.9%): 79 from invertebrate predation, 10 never relocated and the 177 surviving unpredated to the end of the 48 h trials.Figure 2 Survival plot of the experimental treatments (top to bottom: average matching (AM), average non-matching (ANM), edge matching (EM), edge non-matching (ENM), inside matching (IM), inside non-matching (INM)). 2005; Merilaita & Lind 2005, 2006; Stevens et al. We test this counter-intuitive idea that conspicuous patterns might aid concealment, using artificial moth-like targets with pattern elements designed to match or mismatch the average luminance (lightness) of the trees on which they were placed. p. 50, fig. Many butterflies have large, circular patterns on the upper part of their wings. Curves are the probability of surviving bird predation as a function of time (in minutes), based on Kaplan–Meier estimates to account for censoring due to non-avian predation and survival to the end of the study period. The area of the pinhead was negligible (less than 2 mm2). many camouflage patterns exist among all animals, not just the changeable cephalopods. In any one block, 10 replicates of each treatment were randomly allocated, one per tree, along a nonlinear transect of ca 1.5 km by 20 m (targets were placed upon fewer than 5% of available trees), subject to the constraints that no lichen covered the trunk and no young trees of trunk circumference less than 0.9 m were used.
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