MacKenzie Nelson

WLF315: Techniques Laboratory

Habitat and perceived predation risk in fox squirrels

Introduction

Predation is one of the strongest selective pressures that impact the adaptations and

behaviors of animals. Not only do prey animals have adaptations to help them avoid or escape

predators, they also are able to react to perceived risks and predators in order to prevent

predation (Lima and Dill 1990). However, perceiving risk requires antipredator behaviors such

as vigilance, which in turn takes away from the behaviors and energy the animal could be using

to forage or reproduce. In this way, antipredator behavior and vigilance is a trade-off between

behaviors that increase individual’s fitness like foraging and behaviors that avoid death by a

predator (Lima and Dill 1990) (Potash et al. 2019) (Ortiz et al. 2019). Antipredator behaviors

allow us to deduce the level of risk an animal is perceiving, and includes decisions about when to

feed, where to feed, what to feed on, and when to change their behavior in response to the risk

(Lima and Dill 1990).

There are several factors that influence an animal’s perception of risk such as their

predator species (including whether it is an avian species or terrestrial species), vegetation cover,

canopy cover, the distance to refuge, their available sightlines, and the microhabitat (Lima and

Dill 1990) (Potash et al. 2019) (Thorson et al. 1998). Vegetation in particular is an important cue

for perceived risk and foraging behavior. For example, research shows that animals that avoid

predators through concealment tend to forage more and perceive less risk when closer to

vegetative cover or refuge (Potash et al. 2019) (Thorson et al. 1998) (Ortiz et al. 2019), while
other species tend to perceive more risk when they are closer to vegetative cover and have

weaker sightlines (Potash et al. 2019).

Since we can’t directly determine how much risk a foraging animal perceives we use

antipredator behaviors as surrogate for perceived risk. According to optimal escape theory, the

distance between a predator and its prey where they prey perceives the risk of be preyed on as

greater than the fitness benefit of staying and foraging is the flight initiation distance (FID)

(Allan et al. 2021). The FID of an animal is determined by their perceived risk ,which, as

established earlier, is determined by environmental factors and the closeness and behavior of the

predator (Lima and Dill 1990). As perceived risk increases, we should expect FID to increase as

well (Allan et al. 2021). There are other measurements that are used to measure perceived risk,

such as giving-up distance (GUD), but for the purposes of this study, only FID was used.

Our study animal, fox squirrels (Sciurus niger), can be found in a variety of forest types

but prefer open fields with scattered open stands of trees and little understory vegetation (Allen

1982). Their diet primarily consists of mast, buds, insects, bulbs, roots, and seeds (Allen 1982).

Our objective with this study was to look at the relationship between the perception of risk in fox

squirrels and how that perception affected their foraging behavior. From this, we formulated two

hypotheses: 1) fox squirrels perceive higher risk if they are farther from refuge because it will

take them longer to flee to it, so they will need to start fleeing from threats sooner, and 2)

squirrels will choose larger trees for refuge because they have less of a chance of being pursued

and caught in taller/larger trees. Using these hypotheses we predicted that 1) Squirrels will have

a larger FID if they are farther from the refuge tree, and 2) The refuge tree will on average have a

greater circumference than the available tree.

Methods

Study Area

The study was conducted on the administration lawn at the University of Idaho. This

habitat is a mowed lawn with intermittent trees, mostly oaks and maples, several walkways

cutting across it, and is surrounded on all sides by low-traffic roads and university buildings. The

data was collected in the afternoons over two days in mid-October, and the weather was warm

and partly cloudy. Because of the time of day and the location of the study site, students were

using the lawn and walking across it frequently while the data was being collected as well.

Data collection

At the study site, we would identify a spot and sit to watch for squirrel within the study

area. Once a squirrel was sighted, we observed it for 1 to 2 minutes, and then a group member

would approach the squirrel at an even pace until the squirrel fled. The location where the group

member stopped and the location where the squirrel fled from were marked with flags, and the

distance between the two, the FID, was measured in meters. The distance between where the

squirrel fled and the refuge tree was also measured in meters, as well as the distance to the next

closest available tree in meters, the circumference at chest height of both trees in centimeters,

and other habitat parameters that will not be addressed in this analysis.

Data analysis

The data obtained was analyzed in RStudio in order to test the relationships between the

variables in our hypotheses. The relationship between FID and the distance from the refuge tree

at flight, the first hypothesis, was analyzed using a correlation test and visualized using a
scatterplot. The second hypothesis was analyzed by using a paired t-test to see the relationship

between the circumference of the refuge trees and the available trees.

Results

The data included 50 squirrels observed over October 17th and 18th, 2022 in about 2-hour

sessions in the afternoon both days. The data was collected by about 4-5 groups of 4-5 students

on each day.

For the first hypothesis, the correlation between FID and the distance to the refuge tree

was not significant, with a p-value of 0.283. The correlation coefficient of 0.15 and Figure 1 also

demonstrates a weak positive relationship, but the high p-value indicates it isn’t significant

enough to imply correlation.

Figure 1. Relationship between the distance to the refuge tree and FID of fox
squirrel on the Admin Lawn in October 2022.
 For the second hypothesis, the paired t-

test provided a p-value of 0.041, suggesting the

difference in the means between the

circumferences of the refuge tree and the

available tree were significantly different.

Figure 2 shows that the mean of the

circumference of the refuge tree is greater than

that of the available tree. There is overlap in the

95% confidence intervals of the two means

(Figure 2), but the p-value supports there being

a statistically significant difference between

them.

Figure 2. A comparison of the mean circumference

of the available trees and the refuge trees that the
fox squirrel fled to.

Discussion

The results of the correlation test did not support the first hypothesis. This is surprising,

given how much support there is for a correlation between FID and distance to the refuge tree in

other literature. For two examples of many, Thorson et al. (1998) found a strong response to

changes in the microhabitat, mainly the distance to a tree, in the giving-up density of fox

squirrels. They found that fox squirrel gave up by 97% when they were feeding over 4 meters

away from a tree compared to when they were feeding at the base. Additionally, Dill and
Houtman (1987) also found that FID in response to a motorized cat increased in gray squirrels as

distance to refuge increased. There could be a number of explanations for the discrepancy in our

findings and the finding of most other literature. Firstly, human error or inconsistencies between

the groups in the methodologies and data collection could lead to inaccurate data, which in turn

could lead to inaccurate data analysis. There is also evidence in the literature on antipredator

behavior that urban animals, particularly squirrels, can become habituated to humans. McCleery

(2009) looked at FID, vigilance, and antipredator behavior in fox squirrels across the urban-rural

gradient and found that vigilance decreased as the number of humans present in an area

increased, suggesting more constant stimuli to humans habituated the squirrels to predators in

general. Allan et al. (2021) countered this suggestion, agreeing that animals can be habituated to

constant human disturbance, but highlighting that the lack of antipredator behaviors towards

other predators in urban environments could be explained by a lack of predators in anthropogenic

urban settings. Kiteendorf and Dantzer (2021) agreed with this and found that squirrels in trials

of FID with only human presence displayed less vigilance and antipredator behaviors in urban

spaces than in rural ones but displayed more consistent antipredator behavior towards other

predators across the urban-rural gradient. These findings could indicate that the fox squirrels on

the University of Idaho campus that we studied could have had abnormal FIDs because of their

regular and near constant exposure to people on a day-to-day basis. This could lead to the data

we collected being accurate to reality, but not in line with other findings in the literature and not

applicable to all fox squirrels everywhere.

The results did not disprove our second hypothesis. The paired t-test and Figure 2 showed

weak, but significant differences in the circumferences between the refuge trees that the squirrels

fled to and the next available trees. However, there are other possible explanations for this
pattern. In this paper, we did not analyze the relationship between the position where the squirrel

fled and the distance from it to the refuge and available trees, so we are unsure if distance’s

possible role in the tree choice. Still, it would make sense for fox squirrels to choose larger trees

because they are likely taller, sturdier, and more likely to have cavities for the squirrels to hide

in. This is a factor of the microhabitat that should be studied more in future research to clarify

and confirm the finding of this paper since our data likely had poor precision and the outcome of

the results was not very strong. Additionally, no studies on the relationship between tree choice

and tree size or circumferences could be found, so this is likely an unexplored idea in the

literature.

Literature Cited

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