Evidence of signaling benefits to contrasting internal color boundaries in warning coloration

It has been suggested that the common existence of regular patterning in aposematic prey animals makes them "stand out" from the background, improving detection and recognition. Another suggestion is that internal patterns could have a similar positive effect on predator aversion learning as prey-to-background contrast. We used wild caught blue tits (Cyanistes caeruleus) and artificial prey signals to investigate if internal color boundaries, pattern regularity and pattern symmetry affect learning. Birds in different treatments were trained, on a complex background, to discriminate between artificial prey with different nonrewarding color stimuli with a black pattern and rewarding stimuli without a black pattern, followed by a generalization test. This study provides evidence of learning benefits to internally contrasting patterns as the striped prey stimuli were learned faster than the unstriped. Also, we found no beneficial effects of pattern regularity and symmetry. The birds generalized more between prey with different black patterns than to the profitable prey, suggesting that color is of foremost importance. The generalization test also showed a greater avoidance of striped than that of unstriped prey, suggesting some attention on patterns. Thus, internal patterning may affect signal salience and in some circumstances benefit prey due to both a faster avoidance learning and generalization behavior.