Foraging and scavenging behaviour of the prairie rattlesnake (Crotalus viridis): no evidence that envenomation cues facilitate kleptoparasitism of struck prey

Abstract
Most viperids are ambush predators that primarily use venom to subdue prey, employing a strike-release-trail hunting strategy whereby snakes follow the unique scent of envenomated prey to locate carcasses they have bitten and released. In addition to killing prey, rattlesnakes (like most carnivores) will also opportunistically scavenge carrion. This scavenging strategy likely includes the occasional consumption of carcasses killed by other snakes (i.e., kleptoparasitism). In areas with high densities of other pitvipers, utilizing the unique scent of animals envenomated by other snakes might be a viable alternative foraging strategy. We evaluated this possibility experimentally using a series of captive behavioural trials on prairie rattlesnakes (Crotalus viridis) to determine whether conspecific or heterospecific (C. scutulatus, C. ornatus) envenomation cues might increase the likelihood of kleptoparasitism. Rattlesnakes did not prefer envenomated prey over nonenvenomated prey, nor did they prefer venom cues of one species over another. Although they did frequently scavenge carcasses, in the absence of striking, snakes generally located carcasses using random searching movements instead of scent trails. Additionally, the amount of time rattlesnakes spent investigating carcass trails did not differ significantly among treatments, suggesting that striking, and the resultant formation of a chemical search image of prey, is more crucial to trailing behaviour than venom cues. Moreover, a high degree of behavioural variation among individuals was observed, suggesting that scavenging and kleptoparasitism in rattlesnakes is more complex than previously realized, and making generalizations about these behaviours is challenging. Most viperids are ambush predators that primarily use venom to subdue prey, employing a strike-release-trail hunting strategy whereby snakes follow the unique scent of envenomated prey to locate carcasses they have bitten and released. In addition to killing prey, rattlesnakes (like most carnivores) will also opportunistically scavenge carrion. This scavenging strategy likely includes the occasional consumption of carcasses killed by other snakes (i.e., kleptoparasitism). In areas with high densities of other pitvipers, utilizing the unique scent of animals envenomated by other snakes might be a viable alternative foraging strategy. We evaluated this possibility experimentally using a series of captive behavioural trials on prairie rattlesnakes (Crotalus viridis) to determine whether conspecific or heterospecific (C. scutulatus, C. ornatus) envenomation cues might increase the likelihood of kleptoparasitism. Rattlesnakes did not prefer envenomated prey over nonenvenomated prey, nor did they prefer venom cues of one species over another. Although they did frequently scavenge carcasses, in the absence of striking, snakes generally located carcasses using random searching movements instead of scent trails. Additionally, the amount of time rattlesnakes spent investigating carcass trails did not differ significantly among treatments, suggesting that striking, and the resultant formation of a chemical search image of prey, is more crucial to trailing behaviour than venom cues. Moreover, a high degree of behavioural variation among individuals was observed, suggesting that scavenging and kleptoparasitism in rattlesnakes is more complex than previously realized, and making generalizations about these behaviours is challenging.