Colour pattern variation forms local background matching camouflage in a leaf-mimicking toad.

Optimal camouflage can, in principle, be relatively easily achieved in simple, homogeneous, environments where backgrounds always have the same colour, brightness and patterning. Natural environments are, however, rarely homogenous, and species often find themselves viewed against varied backgrounds where the task of concealment is more challenging. One result of variable backgrounds is the evolution of intraspecific phenotypic variation which may either be generalized, with multiple similarly cryptic patterns, or specialized, with each discrete colour form maximizing concealment against a single component of the background. We investigated the role of phenotypic variation in a highly variable population of the Neotropical toad Rhinella margaritifera using visual modelling and a computer-based detection task. We found that phenotypic variation was not divided into discrete colour morphs, and all toads were well camouflaged against the forest floor. However, although the whole population may appear to consist of random samples from the background, the toads were a particularly close match to the leaf litter, suggesting that they masquerade as dead leaves, which are themselves variable. Furthermore, rather than each colour form being equally effective against a single background, each toad was specialized towards its own particular local surroundings, as suggested by a specialist strategy. Taken together, these data highlight the importance of background matching to a nominally masquerading species, as well as how habitat heterogeneity at multiple spatial scales may affect the evolution of camouflage and phenotypic variation.

Imperfect transparency and camouflage in glass frogs.

Camouflage patterns prevent detection and/or recognition by matching the background, disrupting edges, or mimicking particular background features. In variable habitats, however, a single pattern cannot match all available sites all of the time, and efficacy may therefore be reduced. Active color change provides an alternative where coloration can be altered to match local conditions, but again efficacy may be limited by the speed of change and range of patterns available. Transparency, on the other hand, creates high-fidelity camouflage that changes instantaneously to match any substrate but is potentially compromised in terrestrial environments where image distortion may be more obvious than in water. Glass frogs are one example of terrestrial transparency and are well known for their transparent ventral skin through which their bones, intestines, and beating hearts can be seen. However, sparse dorsal pigmentation means that these frogs are better described as translucent. To investigate whether this imperfect transparency acts as camouflage, we used in situ behavioral trials, visual modeling, and laboratory psychophysics. We found that the perceived luminance of the frogs changed depending on the immediate background, lowering detectability and increasing survival when compared to opaque frogs. Moreover, this change was greatest for the legs, which surround the body at rest and create a diffuse transition from background to frog luminance rather than a sharp, highly salient edge. This passive change in luminance, without significant modification of hue, suggests a camouflage strategy, "edge diffusion," distinct from both transparency and active color change.

Distance-dependent defensive coloration in the poison frog Dendrobates tinctorius, Dendrobatidae.

Poison dart frogs provide classic examples of warning signals: potent toxins signaled by distinctive, conspicuous coloration. We show that, counterintuitively, the bright yellow and blue-black color of Dendrobates tinctorius (Dendrobatidae) also provides camouflage. Through computational modeling of predator vision, and a screen-based detection experiment presenting frogs at different spatial resolutions, we demonstrate that at close range the frog is highly detectable, but from a distance the colors blend together, forming effective camouflage. This result was corroborated with an in situ experiment, which found survival to be background-dependent, a feature more associated with camouflage than aposematism. Our results suggest that in D. tinctorius the distribution of pattern elements, and the particular colors expressed, act as a highly salient close range aposematic signal, while simultaneously minimizing detectability to distant observers.

Optimal background matching camouflage.

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.