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  1. Individual animals behave differently from each other for myriad interrelated intrinsic and extrinsic reasons, and this behavioral variation is the raw substrate for evolutionary change. Behavioral varia- tion can both enhance and constrain long-term evolution (Foster, 2013), and it provides the basic materials on which natural and sexual selection can act. A rich body of historical experimental and conceptual foundations precedes many of the topics discussed. This classic literature is vast and impor- tant, and we encourage the reader to examine it in detail (e.g., Lehrman, 1953; Lorenz, 1971; Schnei- rla, 1966; Waddington, 1959) because we discuss more recent literature. For example, the study of the mechanisms that underlie behavioral variation has a divisive history, which involves carving out the relative contributions of genes and environment to a particular phenotype. Developmental systems and reaction-norm views challenged the issue of gene or environment by arguing that the interplay between genetic substrates and environmental inputs defined adaptive phenotypes across multiple contexts (Fos- ter, 2013; Gottlieb, 1991a, 1991b; Jablonka & Lamb, 2014). Identifying the interactional relationship between components permits researchers to under- stand how behavior becomes organized (Gottlieb, 1991a, 1991b) and can reveal links between indi- vidual variation and population-level persistence, species diversification (or stasis), and community dynamics (reviewed in Dingemanse & Wolf, 2013). Similarly, the study of individual differences has a rich history situated in the areas of behavioral genet- ics, sociobiology, behavioral ecology, developmen- tal psychology, personality theory, and studies of learning and cognition. Each area has its own goals, associated techniques, and levels of explanation. The study of behavioral variation during early develop- ment, for instance, has been documented primarily by psychologists studying proximate mechanisms in laboratory animal models, whereas the study of dif- ferent adult morphs using the adaptationist perspec- tive has been dominated by behavioral ecologists examining natural populations (Foster, 1995). A more complete description of individual differences requires an integrative study of the mechanisms (e.g., developmental, physiological) that guide intra- individual flexibility and the associated adaptive fine tuning of behavioral types. It is through this integra- tion that researchers can make predictions about the response of different individual phenotypes, groups, populations, and species to novel situations (e.g., captive and urban environments). 
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