Evolution of alternative foraging tactics driven by water temperature and physiological constraints in an amphibious snake

Abstract

Amphibious predatory ectotherms live and forage in two environments (aquatic and terrestrial) that can drastically differ in temperature means and variance across space and time. The locomotor performance of ectotherms is known to be strongly affected by temperature. However, how differences in water temperature may drive the evolution of alternative foraging tactics in amphibious animals remains poorly understood. Fish-eating Viperine snakes Natrix maura occur from high altitude cold water streams to warm shallow lakes, and employ two main feeding strategies: sentinel foraging (underwater sit-and-wait behaviour) and active foraging (fish chasing). Using 272 juvenile snakes we measured: the performance kinetics of diving and swimming in a wide range of water temperatures; basal metabolic levels in relation to body temperature; and the type of foraging mode expressed in water-temperature-acclimated snakes. Individual swimming performances increased with testing temperature (10, 15, 20, 25 or 30°C). Apnoea time followed an opposite trend however, plausibly reflecting the fact that oxygen demands are related to the metabolic rate of ectotherms. That is, snake heart rates increased with body temperature. Snakes acclimated to 10°C water mostly displayed sentinel foraging. By contrast, 20°C and 30°C water-acclimated snakes were extremely active fish chasers. Individual apnoea times at the various testing temperatures were all correlated; as were individual swimming speeds. There was however no clear relationship between an individual's ability to hold its breath and its ability to swim, suggesting that both performance traits may be the target of different selective pressures. Fast swimming speed and long breath holding abilities are likely key determinants of both foraging success and predatory evasion, although in a context dependent manner. Active swimming foraging is likely to be advantageous in warm water (>20°C), while sentinel foraging appears better suited to cold water (<14°C). The physiological aspects of foraging tactics of amphibious snakes combined with field and laboratory observations support the idea that physiological and environmental constraints may generate shifts in habitat use and associated foraging tactics in amphibious ectotherms. Avenues for further research are discussed.