Abstract
Visual camouflage evolution in animals depends on both light's interaction with its surroundings and the limits of what the natural observers of the camouflage can see. Changes in lighting - such as those caused by shifting weather - can quickly alter how animals and their surroundings appear due to the generation of shadows both cast onto objects and from self-shading. The extent and nature of these changes will depend on the three-dimensional (3D) structure of the environment. Despite the apparent effects of lighting on object and background appearance, the enormity of interactions and the diversity of animal camouflage methods and appearances pose challenges to investigating the combined effects of lighting and habitat structure on camouflage. Genetic algorithms and mathematical methods for generating animal patterns provide a potential solution for investigating camouflage's broad feature space by evolving artificial prey under different lighting conditions within different habitats. Here, an online artificial evolution experiment was used to examine the effect of lighting, both direct and diffuse, and habitat geometry on camouflage. Lighting and geometry changed the appearance of the evolved prey, and the predictive power of common measures of camouflage. Crucially, lighting condition systematically altered how contrasting the prey-targets' internal patterning was and interacted with habitat geometry to affect the evolved pattern shapes, colours, and countershading. Our work demonstrates the importance of considering the relative geometry and lighting of an environment when determining the function of animal colouration and the adaptive value of camouflage.