The world in color presents a dazzling dimension of phenotypic variation. Biological interest in this variation has burgeoned, due to both increased means for quantifying spectral information and heightened appreciation for how animals view the world differently than humans. Effective study of color traits is challenged by how to best quantify visual perception in nonhuman species. This requires consideration of at least visual physiology but ultimately also the neural processes underlying perception. Our knowledge of color perception is founded largely on the principles gained from human psychophysics that have proven generalizable based on comparative studies in select animal models. Appreciation of these principles, their empirical foundation, and the reasonable limits to their applicability is crucial to reaching informed conclusions in color research. In this article, we seek a common intellectual basis for the study of color in nature. We first discuss the key perceptual principles, namely, retinal photoreception, sensory channels, opponent processing, color constancy, and receptor noise. We then draw on this basis to inform an analytical framework driven by the research question in relation to identifiable viewers and visual tasks of interest. Consideration of the limits to perceptual inference guides two primary decisions: first, whether a sensory-based approach is necessary and justified and, second, whether the visual task refers to perceptual distance or discriminability. We outline informed approaches in each situation and discuss key challenges for future progress, focusing particularly on how animals perceive color. Given that animal behavior serves as both the basic unit of psychophysics and the ultimate driver of color ecology/evolution, behavioral data are critical to reconciling knowledge across the schools of color research.
Animal culture, defined as "information or behavior-shared within a community-which is acquired from conspecifics through some form of social learning" (1), can have important consequences for the survival and reproduction of individuals, social groups, and potentially, entire populations (1, 2). Yet, until recently, conservation strategies and policies have focused primarily on broad demographic responses and the preservation of genetically defined, evolutionarily significant units. A burgeoning body of evidence on cultural transmission and other aspects of sociality (3) is now affording critical insights into what should be conserved (going beyond the protection of genetic diversity, to consider adaptive aspects of phenotypic variation), and why specific conservation programs succeed (e.g., through facilitating the resilience of cultural diversity) while others fail (e.g., by neglecting key repositories of socially transmitted knowledge). Here, we highlight how international legal instruments, such as the Convention on the Conservation of Migratory Species of Wild Animals (CMS), can facilitate smart, targeted conservation of a wide range of taxa, by explicitly considering aspects of their sociality and cultures. CONSEQUENCES OF SOCIAL KNOWLEDGE An important aspect of social learning is the speed with which new behaviors can potentially spread through populations, with effects that may be positive (e.g., adaptive exploitation of a new food source) or negative (e.g., increasing conflict with humans, such as when sperm whales learn to remove fish from longlines) (2). Transmission can be mediated by an inherent propensity to adopt innovations (e.g., "lobtail" feeding in humpback whales (1)), or curbed by cultural conservatism (e.g., southern resident killer whales' persistent foraging specialization on Chinook salmon (2)). Social learning can result in the emergence of subpopulations with distinctive behavioral profiles, erecting social barriers, as observed in distinct vocal clans of sperm whales (see the Figure). Culturally mediated population structure has important implications for conservation efforts (4), as it can influence species-wide phenotypic diversity and adaptability to changing conditions (5). In some cases, such as humpback or blue whale song, cultural variation can reflect demography and facilitate more efficient, or less invasive, assays of contemporary genetic population structure (1, 4). Most profoundly, culture can play a causal role in establishing and maintaining distinct evolutionary trajectories (6). Another consequence of social learning can be the increased importance of key individuals as repositories of accumulated knowledge, making their targeted protection particularly important for the persistence of social units. For example, the experience of African elephant matriarchs (see
Understanding individual differences in cognitive performance is a major challenge to animal behaviour and cognition studies. We used the Eastern water skink (Eulamprus quoyii) to examine associations between exploration, boldness and individual variability in spatial learning, a dimension of lizard cognition with important bearing on fitness. We show that males perform better than females in a biologically relevant spatial learning task. This is the first evidence for sex differences in learning in a reptile, and we argue that it is probably owing to sex-specific selective pressures that may be widespread in lizards. Across the sexes, we found a clear association between boldness after a simulated predatory attack and the probability of learning the spatial task. In contrast to previous studies, we found a nonlinear association between boldness and learning: both 'bold' and 'shy' behavioural types were more successful learners than intermediate males. Our results do not fit with recent predictions suggesting that individual differences in learning may be linked with behavioural types via high -low-risk/reward trade-offs. We suggest the possibility that differences in spatial cognitive performance may arise in lizards as a consequence of the distinct environmental variability and complexity experienced by individuals as a result of their sex and social tactics.
A crucial problem for most animals is how to deal with multiple types of predator, which differ in their sensory capabilities and methods of prey detection. For animals capable of rapid colour change, one potential strategy is to change their appearance in relation to the threat posed by different predators. Here, we show that the dwarf chameleon, Bradypodion taeniabronchum, exhibits different colour responses to two predators that differ in their visual capabilities. Using a model of animal colour perception to gain a 'predator's eye view', we show that chameleons showed better background colour matching in response to birds than snakes, yet they appear significantly more camouflaged to the snake visual system because snakes have poorer colour discrimination.
Potential prey are often exposed to multiple predators that vary in their foraging tactics and ability to detect prey. For animals that rely on crypsis to avoid predators, one solution is to alter their behaviour or appearance to maximize crypsis in ways that are specific to different types of predator. We tested whether dwarf chameleons ( Bradypodion transvaalense ) showed different behavioural responses, including colour change, towards multiple predators (bird and snake models) that detect and capture prey in different ways, and whether these antipredator responses varied geographically. Chameleons consistently used the same body postures (lateral compression and flipping to the opposite side of the branch) and displayed similar chromatic (colour) contrast against the natural background in response to both predator types. However, they became significantly more achromatically contrasting (brighter) in the presence of the snake compared to the bird. This relative difference in achromatic contrast towards the two types of predator was consistent among populations. There were also significant differences in both absolute achromatic and chromatic contrast among populations despite very similar light environment, background coloration and habitat structure. Our results highlight facultative crypsis as one type of flexible antipredator tactic and emphasize the importance of visual ecology in understanding prey-predator interactions.
Differences between sexes in physiological performance have received little attention in animals. We tested for sex differences in maximum sprint speed and maximal exertion over a range of temperatures in a population of Platysaurus intermedius wilhelmi lizards. We also examined sex-based differences in selected temperature range, mean field body temperatures (T(b)), and thermal activity limits. Finally, we conducted field studies to quantify male and female responses to a potential predator, which may be affected by their respective performance capabilities. Males were faster than females at all temperatures, and body size had no significant effect on sprint speeds. Males and females also selected similar T(b)'s when placed in a thermal gradient, but in the field, male lizards' T(b)'s were different from those of the females. However, predicted sprint speeds for males and females at their field T(b)'s are similar. No significant differences were found between males and females with regard to maximal exertion. When approached in the field, adult male lizards took refuge significantly earlier than did adult females and also fled over shorter distances, suggesting that females rely on crypsis as an escape strategy.
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