Thursday, February 9, 2017

How Does the Presence of a Predator Impact Whitetail Deer Feeding Behavior?




 Effects of the Addition of Coyote Urine on Whitetail Deer Feeding Behavior at Established Feeding Sites

Jenna Barlow & Nathan Sylte










INTRODUCTION
Whitetail deer (Odocoileus virginianus) are common throughout North America and thrive within diverse habitat regions, many of which have extreme temperature swings between seasons (Rooney 2001). The ability to inhabit regions that experience extreme cold as well as intense warm spells makes the whitetail deer an excellent model homeotherm. Homeotherms are presented with a metabolic challenge during winter months; therefore, whitetail deer must allocate their time in order to consume enough food, but also balance this with rumination, movement, social interactions, and sleeping (Beier and McCullough 1990). Another important part of the activity budget is vigilance.  Increased time spent on vigilance reduces predation risk, but vigilance conflicts with some types of feeding (Benhaiem et al. 2008). Deer have many predators—for example, coyotes that have been observed in packs killing whitetail deer (Gese and Grothe 1995)—and so it is reasonable to expect there will be strong selection for optimizing the ratio of time spent vigilant to time spent feeding. So with the use of predator urine as the apparent risk of a nearby predator, it would be expected that this ratio would shift to more time spent being vigilant and less on feeding.
            The introduction of predator urine to feeding sites has been shown to significantly decrease deer browsing (Swihart et al. 1991). This has important management implications because winter browsing by deer often damages nurseries, natural restoration of timber species, and substantial agriculture crop damage (Swihart et al. 1991; Bellant 1998).  Browsing by deer has also been found to significantly influence forest ecosystems (Rooney and Waller 2003). However, there appears to be little published information on the effect of predator scents on deer feeding and vigilance, though Benhaiem et al. (2008) has found that the more time spent being vigilant when exposed to predation risk can have significant consequences on roe deer energy budgets. Our objective is to study the effects on the whitetail deer activity budget by artificially increasing the apparent predation risk via coyote urine.
METHODS
Four feeding locations baited with corn were established prior to the beginning of the experiment. Sites were established one week in advance on February 29, 2016 to allow deer adequate time to find the sites. From that date on the sites were re-baited with corn every three days. Three sites are located in Dunn County and the fourth is in Chippewa County. All sites are located near woodland edges. Three of the four sites are located at or near a field edge, and the other is in a forest clearing; all sites are at least 200 meters from known deer bedding sites. Our site selection was designed to force the deer out of their comfort zones and into areas farther from cover (and thus be exposed to higher predation risk). Another factor in determining where sites were to be located was the proximity of the sites relative to each other; they were placed at least one mile apart to ensure the same deer were not using multiples of the established feeding sites. Lastly, feeding sites were placed in areas with high whitetail activity, as determined by physical signs such as tracks, heavily used trails, and fecal matter.
            To record data for the experiment, infrared/motion detecting cameras were selected as the most practical option, for they are minimally invasive and can record data for months at a time. On March 7, approximately one week after feeding sites were established, the motion cameras were placed at the four sites. After a 10-day period without coyote urine being present, coyote urine was added to each of the four sites. Several ounces of urine were deposited on a small log that could easily be removed after the 10-day period was up. Five days after the urine was originally added, several more ounces of fresh urine were added to each of the sites. This was done to simulate the frequent presence of a predator. After 10 total days of urine being present, the logs the urine was placed on were removed and the cameras were collected. Unfortunately, the camera at site two turned out to have some technical difficulties and data was unable to be collected from this site. Site two had to be excluded from the experiment.
The variables analyzed in the experiment included total deer activity, percent of deer feeding, percent of deer being vigilant, and time of activity measured by night or day. Total deer activity was measured in each of the two, 10-day periods based on the total number of deer in pictures taken and by the number of pictures taken with deer present. The time of day at which feeding occurred was also measured and was recorded as either day or night, using sunrise and sunset times to establish whether it was day or night. Alertness was measured by counting the number of deer appearing in an alert posture in the pictures. An example of an alert posture includes a deer looking up and away from the feeding site with its ears up. Deer feeding was measured by the amount of deer with their heads in the down posture. The proportion of deer in the picture in an alert posture, as well as the proportion of deer feeding were also taken. Presence of coyote urine represented the independent variable in the experiment. T-tests were used to measure for significant differences in the data. Chi-squared tests were also used in data analysis. Specifically, a Pearson's Chi-squared test with Yates' continuity correction was used.
RESULTS
Overall, whitetail deer spent proportionally more of their time  feeding (50%) compared to the time they spent being in the alert posture (30%, p<0.001). So, in general, deer budgeted more time feeding than openly being vigilant (Figure 1). To support our initial prediction, with the addition of the coyote urine to the sites, the proportion of alertness significantly increased, based on a t-test. The proportion of alertness increased from 0.29 to 0.37 (p=0.018).


 Across each of the three sites, the presence of urine tended to increase deer activity during the day. In all three sites, the proportion of time spent feeding when there was no urine present was drastically different comparing day and night. This indicates that at each site the deer had a preference to feed at one time versus the other. Whereas when there was urine present the proportion of time spent feeding was often the same amount for both day and night, suggesting the urine affected their willingness to feed. Specifically, shifting deer feed more with the presence of  light. This was especially shown in site 4. There was also always a greater proportion of deer feeding when there was no urine added. There is a pattern that emerges in Figure 3, the higher frequency of feeding either day or night remained the same even with the addition of the urine.  
         
DISCUSSION
            Our hypothesis that the addition of coyote urine would impact whitetail deer feeding behavior with regards to vigilance/alertness and willingness to feed was supported by our results. It should first be noted that deer behaved differently at each site. Night feeding was prefered at site four, while deer prefered to feed during the day at site three.  At site one, the deer tended to feed at night; however, there was more variation in their feeding times. With the presence of coyote urine, deer tended to feed more during daylight hours, while without the presence of urine they fed more at night. This indicates that the presence of coyote urine impacted their willingness to feed at night. This makes sense considering coyotes are more active at night (Andelt 1985). Although the presence of coyote urine did not stop deer from feeding at the sites, it did alter their feeding schedule as well as their budgeted time in feeding and alertness. The fact that the presence of coyote urine didn’t completely stop the deer from feeding at the sites demonstrates how important calorie replacement is for  whitetail deer. 
            Proportionally, whitetail deer spent more time feeding than they did in an alert posture (Figure 1). This is logical considering they would never be able to obtain the necessary calories to survive if they spent a majority of their time in an alert posture instead of feeding. With the presence of coyote urine, the budget for this behavior was altered. Whitetail deer were significantly more alert with the presence of urine than without (Figure 2). Therefore, they did not spend as much time feeding as they otherwise would have. This result does not contradict the study of Swihart et al. which found that the presence of predator urine significantly reduced deer browsing.
            Temperature may have represented an important potential covariate in our experiment. Although the effects temperature had on whitetail feeding behavior were not directly measured in our experiment due to camera complications, temperature has been shown to influence deer activity (Beier and McCullough 1990). It is possible that some of the cooler temperature swings during the second part of the experiment (urine present) influenced the deer to feed more during the day. However, the extent to which temperature influenced whitetail deer feeding behavior in our experiment is uncertain. Future studies may want to better analyze how temperature influences whitetail feeding behavior. It is possible that whitetail may be more vigilant during colder time periods and may spend substantially more time feeding during the day.
            Overall, studying animal behavior is difficult but important. At any point in time there may be many different variables influencing the way an animal is behaving, which makes studying how animals behave in the wild challenging. There is still much knowledge to be gained in the study of homeotherms and homeothermy. Further advancements in the study of homeotherm behavior will only benefit our overall knowledge of homeothermy. 
REFERENCES
Andelt, W. F. (1985). Behavioral Ecology of Coyotes in South Texas. The Wildlife Society, 94, 3-45.
Allen, E.O. 1968. Range use, foods, condition, and productivity of white-tailed deer in Montana. The Journal of Wildlife Management 32:130-141.

Beier, P. and D.R. McCullough. 1990. Factors influencing white-tailed deer activity patterns and habitat use. Wildlife Monographs 109:3-51.

Bellant, J.L., T.W. Seamans, and L.A. Tyson. 1998. Predator urines as chemical barriers to white-tailed deer. Proceedings of the Eighteenth Vertebrate Pest Conference Paper 4.

Benhaiem, S., M. Delon, B. Lourtet, B. Cargnelutti, S. Aulagnier, A.M. Hewison, N. Morellet, and H. Verheyden. 2008. Hunting increases vigilance levels in roe deer and modifies feeding site selection. Animal Behaviour 76:611-618.

Gese, E.M. and S. Grothe. 1995. Analysis of coyote predation on deer and elk during winter in Yellowstone National Park, Wyoming. The American Midland Naturalist 133:36-43.

Rooney, T.P. 2001. Deer impacts on forest ecosystems: a North American perspective.
Forestry 74.3:201-208.

Rooney T.P. and D.M. Waller. 2003. Direct and indirect effects of white-tailed deer in forest ecosystems. Forest Ecology and Management 181:165-176. 

Swihart, R.K., J.J. Pignatello, and M.J.I. Mattina. 1991. Aversive responses of white-tailed deer, Odocoileus virginianus, to predator urines. Journal of Chemical Ecology 17:767-777.








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