Quatic Foraging in Garter Snakes: a Comparison of Specialists and Generalists

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Abstract
To examine the assumption, underlying much ecological theory, that a "Jack-of-all-trades is master of none", a comparison was made of the aquatic predation of four colubrid snakes, two aquatic specialists (Thamnophis couchi, T. melanogaster) and two terrestrial-aquatic generalists (T. sirtalis, and T. elegans). Observations were made in the field, then juvenile snakes were compared under controlled laboratory conditions. The specialists and generalists had qualitatively different foraging repertoires. The specialists made lengthy dives, crawled slowly on the underwater substrate and made long-distance underwater responses to prey: orientation, approach, pursuit, and frontal attack. The generalists dived briefly, with rapid serpentine locomotion, or searched "terrestrially" by wandering along the shoreline and snatching prey from the water surface. Underwater, the generalists' only prey-directed response was sudden short-distance attack, and they appeared to rely on chance encounters with prey. There were significant differences between the two pairs. In comparison with the generalists, the specialists 1) captured more fish, 2) spent more time on aquatic search, 3) spent more aquatic search time diving, 4) spent more diving time on the underwater substrate, 5) made more attacks in every category of search in open water, and 6) attacked underwater from a greater distance. Significant differences were not found in attack frequencies in those search categories where underwater vision is probably unimportant (aerial attacks and attacks in underwater crevices), nor in aerial attack distances. The specialists' behavioral superiority in respect of their specialization supports the "Jack-of-all-trades" assumption. The observations suggest that this superiority is associated with superior underwater visual acuity (which may be due to superior visual accommodation). The generalists' principal underwater searching technique (serpentine diving) appears to be energetically costly, and may only be profitable when aquatic prey are especially vulnerable. Nerodia sipedon, member of a genus that is closely related to Thamnophis but more aquatic, had a foraging repertoire qualitatively similar to that of the generalists. The similarity may be associated with N. sipedon's nocturnal habits and presumed reliance on non-visual prey cues.