Abstract
The risk of predation varies with behavioral cues and body characteristics of potential predators. One such body characteristic is the head/face orientation of the predator. However, a prey individual’s ability to detect the head may be more difficult when the predator’s body is serpentine, with little distinguishing the head from the tail. Here, we tested whether individuals in mixed-species flocks of Carolina chickadees (Poecile carolinensis) and tufted titmice (Baeolophus bicolor) distinguish the head orientation of predator snake models. We conducted behavioral observations at multiple sites each having a bird feeder stocked with seed. Each chickadee and titmouse flock was exposed to two counterbalanced trials: a snake model with head closest to the seed area of the feeder and with tail closest to the seed area of the feeder. Observers recorded the number of seeds taken by each species and also the number of unsuccessful feeder visits. Chickadees and, to a lesser extent, titmice took fewer seeds and had more unsuccessful feeder visits when the head of the snake model was closest to the seed, compared to when the tail was closest to the seed. Titmice, furthermore, had more unsuccessful feeder visits to the black snake model type representing a real snake nest predator for these small songbirds. Therefore, head orientation seems an important factor that some species use to assess predation risk, even for predatory species where head orientation may be a subtle cue.



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Acknowledgments
We thank the staffs of Ijams Nature Center, Norris Dam State Park, and the University of Tennessee Forest Resources, Research, and Education Center for allowing us to carry out these studies at their sites. We are grateful to David Book, Sheri Browning, Gordon Burghardt, Sandy Echternacht, Elizabeth Hobson, Hwayoung Jung, Steven Kyle, the students of the Comparative Animal Behavior Laboratory, and two anonymous reviewers for helpful comments on earlier drafts of this manuscript.
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This study was conducted under approved University of Tennessee IACUC protocol no. 1248.
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Communicated by P. A. Bednekoff
Significance statement
Prey species should pay attention not only to the presence of a predator in the environment, but also to the predator’s behavior. One predator behavioral cue to which prey individuals pay attention is the predator’s head and body orientation. However, this behavioral cue may be difficult to assess if the predator is a snake. Here, we demonstrate experimentally that Carolina chickadees and, to a lesser extent, tufted titmice, forage less from a feeder if the head of a snake model is closer to the feeder than if the tail of a snake model is closer to the feeder. Along with other studies, this work suggests that small songbirds are highly sensitive to the head orientation of potential predators, even in serpentine species where the head location may be difficult to perceive.
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Cantwell, L.R., Johnson, W.T., Kaschel, R.E. et al. Predator-risk-sensitive foraging behavior of Carolina chickadees (Poecile carolinensis) and tufted titmice (Baeolophus bicolor) in response to the head orientation of snake predator models. Behav Ecol Sociobiol 70, 533–539 (2016). https://doi.org/10.1007/s00265-016-2070-x
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DOI: https://doi.org/10.1007/s00265-016-2070-x


