Background complexity can mitigate poor camouflage

Avoiding detection through camouflage is often key to survival. However, an animal's appearance is not the only factor affecting conspicuousness: background complexity also alters detectability. This has been experimentally demonstrated for both artificially patterned backgrounds in the laboratory and natural backgrounds in the wild, but only for targets that already match the background well. Do habitats of high visual complexity provide concealment to even relatively poorly camouflaged animals? Using artificial prey which differed in their degrees of background matching to tree bark, we were able to determine their survival, under bird predation, with respect to the natural complexity of the background. The latter was quantified using low-level vision metrics of feature congestion (or ‘visual clutter’) adapted for bird vision. Higher background orientation clutter (edges with varying orientation) reduced the detectability of all but the poorest background-matching camouflaged treatments; higher background luminance clutter (varying achromatic lightness) reduced average mortality for all treatments. Our results suggest that poorer camouflage can be mitigated by more complex backgrounds, with implications for both camouflage evolution and habitat preferences.

The giant panda is cryptic

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.

Generalist camouflage can be more successful than microhabitat specialisation in natural environments

Background Crypsis by background-matching is a critical form of anti-predator defence for animals exposed to visual predators, but achieving effective camouflage in patchy and variable natural environments is not straightforward. To cope with heterogeneous backgrounds, animals could either specialise on particular microhabitat patches, appearing cryptic in some areas but mismatching others, or adopt a compromise strategy, providing partial matching across different patch types. Existing studies have tested the effectiveness of compromise strategies in only a limited set of circumstances, primarily with small targets varying in pattern, and usually in screen-based tasks. Here, we measured the detection risk associated with different background-matching strategies for relatively large targets, with human observers searching for them in natural scenes, and focusing on colour. Model prey were designed to either ‘specialise’ on the colour of common microhabitat patches, or ‘generalise’ by matching the average colour of the whole visual scenes. Results In both the field and an equivalent online computer-based search task, targets adopting the generalist strategy were more successful in evading detection than those matching microhabitat patches. This advantage occurred because, across all possible locations in these experiments, targets were typically viewed against a patchwork of different microhabitat areas; the putatively generalist targets were thus more similar on average to their various immediate surroundings than were the specialists. Conclusions Demonstrating close agreement between the results of field and online search experiments provides useful validation of online citizen science methods commonly used to test principles of camouflage, at least for human observers. In finding a survival benefit to matching the average colour of the visual scenes in our chosen environment, our results highlight the importance of relative scales in determining optimal camouflage strategies, and suggest how compromise coloration can succeed in nature.

Evolutionary fine-tuning of background-matching camouflage among geographical populations in the sandy beach tiger beetle

Background-matching camouflage is a widespread adaptation in animals; however, few studies have thoroughly examined its evolutionary process and consequences. The tiger beetle Chaetodera laetescripta exhibits pronounced variation in elytral colour pattern among sandy habitats of different colour in the Japanese Archipelago. In this study, we performed digital image analysis with avian vision modelling to demonstrate that elytral luminance, which is attributed to proportions of elytral colour components, is fine-tuned to match local backgrounds. Field predation experiments with model beetles showed that better luminance matching resulted in a lower attack rate and corresponding lower mortality. Using restriction site-associated DNA (RAD) sequence data, we analysed the dispersal and evolution of colour pattern across geographical locations. We found that sand colour matching occurred irrespective of genetic and geographical distances between populations, suggesting that locally adapted colour patterns evolved after the colonization of these habitats. Given that beetle elytral colour patterns presumably have a quantitative genetic basis, our findings demonstrate that fine-tuning of background-matching camouflage to local habitat conditions can be attained through selection by visual predators, as predicted by the earliest proponent of natural selection.

When animal coloration is a poor match

Biologists usually pursue the adaptationist paradigm in trying to explain the functional significance of animal coloration. Here I collate instances in which coloration may be a poor match in the context of background matching, Batesian mimicry, aposematism, and colour polymorphisms. This can occur because of trade-offs with other functions, relaxed selection from predation, or colour trait neutrality. Also, mechanistic, pleiotropic and chance genetic effects can all result in a poor match to the background environment or to signaling efficiently. While biologists implicitly recognise these constraints placed on adaptive coloration, they rarely explicitly acknowledge the heterodox notion that coloration might be under weak selection or no selection at all. Unfortunately, it is difficult to show this definitively, as illustrated in an investigation into the function of colour polymorphisms in coconut crabs.