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 (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 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.

No abstract

The effect of viewing distance on the perception of visual texture is well known: spatial frequencies higher than the resolution limit of an observer's visual system will be summed and perceived as a single combined colour. In animal defensive colour patterns, distance-dependent pattern blending may allow aposematic patterns, salient at close range, to match the background to distant observers. Indeed, recent research has indicated that reducing the distance from which a salient signal can be detected can increase survival over camouflage or conspicuous aposematism alone. We investigated whether the spatial frequency of conspicuous and cryptically coloured stripes affects the rate of avian predation. Our results are consistent with pattern blending acting to camouflage salient aposematic signals effectively at a distance. Experiments into the relative rate of avian predation on edible model caterpillars found that increasing spatial frequency (thinner stripes) increased survival. Similarly, visual modelling of avian predators showed that pattern blending increased the similarity between caterpillar and background. These results show how a colour pattern can be tuned to reveal or conceal different information at different distances, and produce tangible survival benefits.

Aposematic signals are often characterized by high conspicuousness. Larger and brighter signals reinforce avoidance learning, distinguish defended from palatable prey and are more easily memorized by predators. Conspicuous signalling, however, has costs: encounter rates with naive, specialized or nutritionally stressed predators are likely to increase. It has been suggested that intermediate levels of aposematic conspicuousness can evolve to balance deterrence and detectability, especially for moderately defended species. The effectiveness of such signals, however, has not yet been experimentally tested under field conditions. We used dough caterpillar-like baits to test whether reduced levels of aposematic conspicuousness can have survival benefits when predated by wild birds in natural conditions. Our results suggest that, when controlling for the number and intensity of internal contrast boundaries (stripes), a reduced-conspicuousness aposematic pattern can have a survival advantage over more conspicuous signals, as well as cryptic colours. Furthermore, we find a survival benefit from the addition of internal contrast for both high and low levels of conspicuousness. This adds ecological validity to evolutionary models of aposematic saliency and the evolution of honest signalling.

Defended prey often use distinctive, conspicuous, colours to advertise their unprofitability to potential predators (aposematism). These warning signals are frequently made up of salient, high contrast, stripes which have been hypothesized to increase the speed and accuracy of predator avoidance learning. Limitations in predator visual acuity, however, mean that these patterns cannot be resolved when viewed from a distance, and adjacent patches of colour will blend together (pattern blending). We investigated how saliency changes at different viewing distances in the toxic and brightly coloured cinnabar moth caterpillar (Tyria jacobaeae). We found that although the caterpillars' orange-and-black stripes are highly salient at close range, when viewed from a distance the colours blend together to match closely those of the background. Cinnabar caterpillars therefore produce a distance-dependent signal combining salient aposematism with targeted background matching camouflage, without necessarily compromising the size or saturation of their aposematic signal.