Predator avoidance of noxious prey, aposematism and defensive mimicry are normally associated with bright, contrasting patterns and colours. However, noxious prey may be unable to evolve conspicuous coloration because of other selective constraints, such as the need to be inconspicuous to their own prey or to specialist predators. Many venomous snakes, particularly most vipers, display patterns that are apparently cryptic, but nevertheless highly characteristic, and appear to be mimicked by other, non-venomous snakes. However, predator avoidance of viper patterns has never been demonstrated experimentally. Here, the analysis of 813 avian attacks on 12 636 Plasticine snake models in the field shows that models bearing the characteristic zigzag band of the adder (Vipera berus) are attacked significantly less frequently than plain models. This suggests that predator avoidance of inconspicuously but characteristically patterned noxious prey is possible. Our findings emphasize the importance of mimicry in the ecological and morphological diversification of advanced snakes.

Summary 1.Aposematism is an antipredatory strategy in which prey animals conspicuously signal their noxiousness to potential predators. This successful strategy is based on avoidance learning of predators. 2. Most species in the viperid genus Vipera have a characteristic dorsal zigzag pattern. It has previously been suggested that the dorsal zigzag pattern of Vipera berus (the adder) has a cryptic function and thus makes the snake less conspicuous to avian predators. However, a recent experiment suggested that the pattern may function as an aposematic signal rather than being cryptic, and therefore induces active avoidance in avian predators. 3. We conducted a field experiment in Doñana national park in Spain, testing whether the zigzag pattern gives survival value to Vipera latastei gaditana against avian predators. We used two sizes of plasticine models with a zigzag pattern vs. without a zigzag pattern to record attacks by avian predators. The background was controlled (white vs. natural) to exclude the possibility that one morph would be more cryptic to predators than the other one. We also tested size-dependent mortality against the signalling and nonsignalling snakes as aposematic signal is expected to enhance the survival of the aposematic species in a size-dependent manner. 4. Predation pressure against snakes was severe, and on average 39% of models were attacked. Coloration did not enhance the survival of juvenile-sized zigzag-patterned snakes, but significantly increased the survival of adult-sized zigzag-patterned snakes on both backgrounds. 5. Our results provide further support for the aposematism hypothesis of zigzagpatterned noxious snakes, although the advantage due to disruptive coloration is not mutually exclusive.

European vipers (genus Vipera) are venomous and often have a distinctive dorsal zigzag pattern. The zigzag pattern of vipers has been suggested to be an example of disruptive colouration which reduces the detectability of a snake. However, recent studies suggest that the patterns have an aposematic function, although those experiments did not exclude the possibility of disruptive colouration. We used plasticine replicas of snakes to examine whether the zigzag pattern of European vipers provides protection from avian predator attacks via disruptive or aposematic function, or if the zigzag pattern might simultaneously serve both antipredatory functions. Experiments were conducted in the Coto Doñana National Park southern Spain. In the experiment, predation pressure caused by birds was compared between zigzag pattern (patterns were painted with and without disruptive effect i.e. breaking body outline or not), classical disruptive colouration (nonrandomly placed patterns that breaks body outline) and control markings (replicas with length wise stripes and models without painted pattern) on natural and controlled backgrounds. We found that zigzag patterned snake replicas suffered less predation than striped ones regardless of the background, providing further evidence that the zigzag pattern of European vipers functions as a warning signal against predators. However, we did not find evidence that the zigzag pattern involves a disruptive effect.

Predation pressure is expected to drive visual warning signals to evolve toward conspicuousness. However, coloration of defended species varies tremendously and can at certain instances be considered as more camouflaged rather than conspicuous. Recent theoretical studies suggest that the variation in signal conspicuousness can be caused by variation (within or between species) in predators' willingness to attack defended prey or by the broadness of the predators' signal generalization. If some of the predator species are capable of coping with the secondary defenses of their prey, selection can favor reduced prey signal conspicuousness via reduced detectability or recognition. In this study, we combine data collected during three large-scale field experiments to assess whether variation in avian predator species (red kite, black kite, common buzzard, short-toed eagle, and booted eagle) affects the predation pressure on warningly and non-warningly colored artificial snakes. Predation pressure varied among locations and interestingly, if common buzzards were abundant, there were disadvantages to snakes possessing warning signaling. Our results indicate that predator community can have important consequences on the evolution of warning signals. Predators that ignore the warning signal and defense can be the key for the maintenance of variation in warning signal architecture and maintenance of inconspicuous signaling.