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Fish scales are bony plates embedded in the skin that vary extensively in shape across taxa. Despite a plethora of hypotheses regarding form–function relationships in scales, we know little about the ecological selective factors that shape their diversity. Here we examine evolutionary patterns of scale morphology using novel three-dimensional topography from the surfaces of 59 species of damselfishes, a prominent radiation of coral reef fishes. We find evidence that scale morphology changes with different flow environments, such that species that spend more time in open-water habitats have smoother scales. We also show that other aspects of ecology lead to highly derived scales. For example, anemonefishes show an evolutionary transition to smaller scales and smaller ctenii (scale spines). Moreover, changes in body shape, which may reflect ecological differentiation, are related to scale shape but not surface properties. We also demonstrate weak evolutionary integration among multiple aspects of scale morphology; however, scale size and shape are related, and scale morphology is correlated between different body regions. Finally, we also identify a relationship between aspects of lateral line pore morphology, such that the number of lateral line pores per scale and the size of those pores are inversely related. Overall, our study provides insights into the multidimensionality of scale evolution and improves our understanding of some of the factors that can give rise to the diversity of scales seen across fishes.

 

Abstract Coral reef fishes constitute one of the most diverse assemblages of vertebrates on the planet. Color patterns are known to serve a number of functions including intra- and inter-specific signaling, camouflage, mimicry, and defense. However, the relative importance of these and other factors in shaping color pattern evolution is poorly understood. Here we conduct a comparative phylogenetic analysis of color pattern evolution in the butterflyfishes (Chaetodontidae). Using recently developed tools for quantifying color pattern geometry as well as machine learning approaches, we investigate the tempo of evolution of color pattern elements and test whether ecological variables relating to defense, depth, and social behavior predict color pattern evolution. Butterflyfishes exhibit high diversity in measures of chromatic conspicuousness and the degrees of fine versus gross scale color patterning. Surprisingly, most diversity in color pattern was not predicted by any of the measures of ecology in our study, although we did find a significant but weak relationship between the level of fine scale patterning and some aspects of defensive morphology. We find that the tempo of color pattern diversification in butterflyfishes has increased toward the present and suggest that rapid evolution, presumably in response to evolutionary pressures surrounding speciation and lineage divergence, has effectively decoupled color pattern geometry from some aspects of ecology. Machine learning classification of color pattern appears to rely on a set of features that are weakly correlated with current color pattern geometry descriptors, but that may be better suited for the detection of discrete components of color pattern. A key challenge for future studies lies in determining whether rapid evolution has generally decoupled color patterns from ecology, or whether convergence in function produces convergence in color pattern at phylogenetic scales.