Physical habitat and social conditions across a coral reef shape spatial patterns of intraspecific behavioral variation in a demersal fish

Environmental variation holds a central role in determining the distribution and diversity of species. For any given taxon, the number and type of environmental variables that govern its distribution can range widely, but commonly include both abiotic parameters (e.g. temperature, salinity, wave energy, substratum shelter) and biotic variables such as food and social factors (e.g. conspecific density, local species diversity). Spatial heterogeneity in environmental conditions often equates to variation in habitat quality, which ultimately leads to variation in the distribution of species. However, less commonly considered is that environmental variation can also generate spatial variation in behavior within a species. Similar to other phenotypic traits, behavior can be strongly influenced by variation in habitat, and individuals that experience different physical and social conditions – whether along environmental gradients or among geographically isolated habitats – can differ behaviorally as they cope with disparate local challenges (Reichert 1974; Thompson 1990; reviewed by Foster 1999). Whether arising through