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Camouflage helps animals to hide from predators and is therefore key to survival. Although widespread convergence of animal phenotypes to their natural environment is well‐established, there is a lack of knowledge about how species compromise camouflage accuracy across different background types in their habitat. Here we tested how background matching has responded to top–down selection by avian and mammalian predators using Sahara–Sahel desert rodents in North Africa. We show that the fur colouration of several species has become an accurate match to different types of desert habitats. This is supported by a correlation analysis of colour and pattern metrics, investigation of animal‐to‐background similarities at different spatial scales and is confirmed by modelling of two predator vision systems. The background match was closest across large (or global) spatial scales, suggesting a generalist camouflage tactic for many background types. Some species, may have a better match to the background over small (or focal) spatial scales, which could be the result of habitat choices or differential predation. Nevertheless, predicted discrimination distances of fur colouration were virtually indistinguishable for mammalian and low for avian vision model, which implies effective camouflage. Our study provides one of the best documented cases of multilevel camouflage accuracy in geographically widespread taxa. We conclude that background matching has become an effective and common adaptation against predatory threat in Sahara–Sahelian desert rodents. The authors study provides one of the best documented cases of multilevel camouflage accuracy in geographically widespread taxa. They conclude that background matching has become an effective and common adaptation against predatory threat in Sahara–Sahelian desert rodents.

Evolutionary biologists frequently wish to measure the fitness of alternative phenotypes using behavioral experiments. However, many phenotypes are complex. One example is coloration: camouflage aims to make detection harder, while conspicuous signals (e.g., for warning or mate attraction) require the opposite. Identifying the hardest and easiest to find patterns is essential for understanding the evolutionary forces that shape protective coloration, but the parameter space of potential patterns (colored visual textures) is vast, limiting previous empirical studies to a narrow range of phenotypes. Here, we demonstrate how deep learning combined with genetic algorithms can be used to augment behavioral experiments, identifying both the best camouflage and the most conspicuous signal(s) from an arbitrarily vast array of patterns. To show the generality of our approach, we do so for both trichromatic (e.g., human) and dichromatic (e.g., typical mammalian) visual systems, in two different habitats. The patterns identified were validated using human participants; those identified as the best for camouflage were significantly harder to find than a tried-and-tested military design, while those identified as most conspicuous were significantly easier to find than other patterns. More generally, our method, dubbed the “Camouflage Machine,” will be a useful tool for identifying the optimal phenotype in high dimensional state spaces.

For camouflage to succeed, an individual has to pass undetected, unrecognized or untargeted, and hence it is the processing of visual information that needs to be deceived. Camouflage is therefore an adaptation to the perception and cognitive mechanisms of another animal. Although this has been acknowledged for a long time, there has been no unitary account of the link between visual perception and camouflage. Viewing camouflage as a suite of adaptations to reduce the signal-to-noise ratio provides the necessary common framework. We review the main processes in visual perception and how animal camouflage exploits these. We connect the function of established camouflage mechanisms to the analysis of primitive features, edges, surfaces, characteristic features and objects (a standard hierarchy of processing in vision science). Compared to the commonly used research approach based on established camouflage mechanisms, we argue that our approach based on perceptual processes targeted by camouflage has several important benefits: specifically, it enables the formulation of more precise hypotheses and addresses questions that cannot even be identified when investigating camouflage only through the classic approach based on the patterns themselves. It also promotes a shift from the appearance to the mechanistic function of animal coloration. This article is part of the themed issue ‘Animal coloration: production, perception, function and application’.

Judgements of volume may influence the rate of consumption of alcohol and, in turn, the amount consumed. The aim of the current study was to examine the impact of the size and shape of wine glasses on perceptions of wine volume. Online experiment: Participants (n = 360; recruited via Mechanical Turk) were asked to match the volume of wine in two wine glasses, specifically: 1. the Reference glass holding a fixed reference volume, and 2. the Comparison glass, for which the volume could be altered until participants perceived it matched the reference volume. One of three comparison glasses was shown in each trial: 'wider' (20% wider but same capacity); 'larger' (same width but 25% greater capacity); or 'wider-and-larger' (20% wider and 25% greater capacity). Reference volumes were 125 ml, 175 ml and 250 ml, in a fully factorial within-subjects design: 3 (comparison glass) x 3 (reference volume). Non-zero differences between the volumes with which participants filled comparison glasses and the corresponding reference volumes were identified using sign-rank tests. Participants under-filled the wider glass relative to the reference glass for larger reference volumes, and over-filled the larger glass relative to the reference glass for all reference volumes. Results for the wider-and-larger glass showed a mixed pattern across reference volume. For all comparison glasses, in trials with larger reference volumes participants tended to fill the comparison glass less, relative to trials with smaller reference volumes for the same comparison glass. These results are broadly consistent with people using the relative fullness of glasses to judge volume, and suggest both the shape and capacity of wine glasses may influence perceived volume. Perceptions that smaller glasses contain more than larger ones (despite containing the same volume), could slow drinking speed and overall consumption by serving standard portions in smaller glasses. This hypothesis awaits testing.

Background matching is the most familiar and widespread camouflage strategy: avoiding detection by having a similar colour and pattern to the background. Optimizing background matching is straightforward in a homogeneous environment, or when the habitat has very distinct sub-types and there is divergent selection leading to polymorphism. However, most backgrounds have continuous variation in colour and texture, so what is the best solution? Not all samples of the background are likely to be equally inconspicuous, and laboratory experiments on birds and humans support this view. Theory suggests that the most probable background sample (in the statistical sense), at the size of the prey, would, on average, be the most cryptic. We present an analysis, based on realistic assumptions about low-level vision, that estimates the distribution of background colours and visual textures, and predicts the best camouflage. We present data from a field experiment that tests and supports our predictions, using artificial moth-like targets under bird predation. Additionally, we present analogous data for humans, under tightly controlled viewing conditions, searching for targets on a computer screen. These data show that, in the absence of predator learning, the best single camouflage pattern for heterogeneous backgrounds is the most probable sample.

The giant panda ( Ailuropoda melanoleuca ) is an iconic mammal, but the function of its black-and-white coloration is mysterious. Using photographs of giant pandas taken in the wild and state-of-the-art image analysis, we confirm the counterintuitive hypothesis that their coloration provides camouflage in their natural environment. The black fur blends into dark shades and tree trunks, whereas white fur matches foliage and snow when present, and intermediate pelage tones match rocks and ground. At longer viewing distances giant pandas show high edge disruption that breaks up their outline, and up close they rely more on background matching. The results are consistent across acuity-corrected canine, feline, and human vision models. We also show quantitatively that the species animal-to-background colour matching falls within the range of other species that are widely recognised as cryptic. Thus, their coloration is an adaptation to provide background matching in the visual environment in which they live and simultaneously to afford distance-dependent disruptive coloration, the latter of which constitutes the first computational evidence of this form of protective coloration in mammals.

Movement is the enemy of camouflage: most attempts at concealment are disrupted by motion of the target. Faced with this problem, navies in both World Wars in the twentieth century painted their warships with high contrast geometric patterns: so-called \"dazzle camouflage\". Rather than attempting to hide individual units, it was claimed that this patterning would disrupt the perception of their range, heading, size, shape and speed, and hence reduce losses from, in particular, torpedo attacks by submarines. Similar arguments had been advanced earlier for biological camouflage. Whilst there are good reasons to believe that most of these perceptual distortions may have occurred, there is no evidence for the last claim: changing perceived speed. Here we show that dazzle patterns can distort speed perception, and that this effect is greatest at high speeds. The effect should obtain in predators launching ballistic attacks against rapidly moving prey, or modern, low-tech battlefields where handheld weapons are fired from short ranges against moving vehicles. In the latter case, we demonstrate that in a typical situation involving an RPG7 attack on a Land Rover the reduction in perceived speed is sufficient to make the grenade miss where it was aimed by about a metre, which could be the difference between survival or not for the occupants of the vehicle.

A complete explanation of the diversity of animal colour patterns requires an understanding of both the developmental mechanisms generating them and their adaptive value. However, only two previous studies, which involved computer-generated evolving prey, have attempted to make this link. This study examines variation in the camouflage patterns displayed on the flanks of many felids. After controlling for the effects of shared ancestry using a fully resolved molecular phylogeny, this study shows how phenotypes from plausible felid coat pattern generation mechanisms relate to ecology. We found that likelihood of patterning and pattern attributes, such as complexity and irregularity, were related to felids' habitats, arboreality and nocturnality. Our analysis also indicates that disruptive selection is a likely explanation for the prevalence of melanistic forms in Felidae. Furthermore, we show that there is little phylogenetic signal in the visual appearance of felid patterning, indicating that camouflage adapts to ecology over relatively short time scales. Our method could be applied to any taxon with colour patterns that can reasonably be matched to reaction–diffusion and similar models, where the kinetics of the reaction between two or more initially randomly dispersed morphogens determines the outcome of pattern development.

Static high contrast ('dazzle') patterns, such as zigzags, have been shown to reduce the perceived speed of an object. It has not escaped our notice that this effect has possible military applications and here we report a series of experiments on humans, designed to establish whether dynamic dazzle patterns can cause distortions of perceived speed sufficient to provide effective defence in the field, and the extent to which these effects are robust to a battery of manipulations. Dynamic stripe patterns moving in the same direction as the target are found to increase the perceived speed of that target, whilst dynamic stripes moving in the opposite direction to the target reduce the perceived speed. We establish the optimum position for such dazzle patches; confirm that reduced contrast and the addition of colour do not affect the performance of the dynamic dazzle, and finally, using the CO2 challenge, show that the effect is robust to stressful conditions.

Countershading, a vertical luminance gradient from a dark back to a light belly, is perhaps the most common coloration phenotype in the animal kingdom. Why? We investigated whether countershading functions as self-shadow concealment (SSC) in ruminants. We calculated “optimal” countershading for SSC by measuring illumination falling onto a model ruminant as a function of time of day and lighting environment. Calibrated images of 114 species of ruminant were compared to the countershading model, and phylogenetic analyses were used to find the best predictors of coats’ countershading characteristics. In many species, countershading was close to the model’s prediction of “optimal” countershading for SSC. Stronger countershading was associated with increased use of open lighting environments, living closer to the equator, and small body size. Abrupt transitions from dark to light tones were more common in open lighting environments but unassociated with group size or antipredator behavior. Though the SSC hypothesis prediction for stronger countershading in diurnal species was not supported and noncountershaded or reverse-countershaded species were unexpectedly common, this basic pattern of associations is explained only by the SSC hypothesis. Despite extreme variation in lighting conditions, many terrestrial animals still find protection from predation by compensating for their own shadows.