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

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.

Allopatric divergence and secondary contact with gene flow: a recurring theme in rattlesnake speciation

The study of recently diverged lineages whose geographical ranges come into contact can provide insight into the early stages of speciation and the potential roles of reproductive isolation in generating and maintaining species. Such insight can also be important for understanding the strategies and challenges for delimiting species within recently diverged species complexes. Here, we use mitochondrial and nuclear genetic data to study population structure, gene flow and demographic history across a geographically widespread rattlesnake clade, the western rattlesnake species complex (Crotalus cerberus, Crotalus viridis, Crotalus oreganus and relatives), which contains multiple lineages with ranges that overlap geographically or contact one another. We find evidence that the evolutionary history of this group does not conform to a bifurcating tree model and that pervasive gene flow has broadly influenced patterns of present-day genetic diversity. Our results suggest that lineage diversity has been shaped largely by drift and divergent selection in isolation, followed by secondary contact, in which reproductive isolating mechanisms appear weak and insufficient to prevent introgression, even between anciently diverged lineages. The complexity of divergence and secondary contact with gene flow among lineages also provides new context for why delimiting species within this complex has been difficult and contentious historically.