Boldness and exploration vary between shell morphs but not environmental contexts in the snail Cepaea nemoralis

Abstract

Understanding the maintenance of among-individual behavioral variation in populations, and predicting its consequences, are key challenges in behavioral ecology. Studying the association between repeatable behaviors and other traits under selection may shed light on the underlying selective pressures. We used the model snail Cepaea nemoralis to examine whether individual behavior is associated with shell morph, a key trait that has been extensively studied in the context of thermal tolerance and predator avoidance, and which is known to be under strict genetic control in this species. We quantified proxies of boldness and exploration in snails of three morphs coming from two habitats with different thermal contexts. We show that both behaviors were repeatable at the among-individual level (within-state Rboldness = 0.22 [95% credible interval: 0.15, 0.29]; Rexploration = 0.20 [0.15, 0.25]). Behavior was associated with shell morph, with the darker morph (five-banded) being consistently shyer and slower to explore. There was no evidence that thermal environment of origin influenced behavior. Snails became faster when test temperature increased; we found no evidence morphs differed in their thermal response. Boldness and exploration were correlated among individuals, forming a syndrome (r = 0.28 [0.10, 0.46]). We discuss what these results may tell us about the type of selection exerted by predators. We also detail how our results hint to a genetic link between shell morph and behavior, and the evolutionary implications of such a link. Finally, we discuss how our findings combined with decades of evolutionary research make C. nemoralis a very valuable model to study the evolution of behavior in response to environmental changes.