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Animal activity and bait removal experiment in Stockton University’s Forest Management

Area

Kyle Smith

ENVL-2205-002: Ecological Principles Lab

Instructor: Dr. Catherine Tredick

October 8th – 15th, 2020

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Introduction

The anthropogenic fragmentation of once contiguous forested habitats in the U.S. through

rapid urbanization, agricultural conversion, and timber harvesting has decreased forest cover

dramatically and increased the amount of edge habitat (Carman 2013). Fragmentation can attract

edge-adapted invasive species to remaining habitat patches. Native species suited to forested

habitat are thus more susceptible to local extinction, as edge colonizers can often outcompete

such native species (Relyea and Ricklefs 2018). However, forest management treatments of

clear-cutting, thinning, corridor creation, and coppicing can be used as conservation tools to alter

key habitat components of forage and vegetation structure for the benefit of wildlife populations

in remaining forest patches. Silvicultural practices have a strong influence on animal feeding

patterns, brood parasitism and nest predation rates, and small mammal demographics (Zwolak

2009).

Clear-cutting creates a hard-edge effect in formerly contiguous stands, which many

species of bats use for echolocation and insect foraging. A survey study examining the effects of

timber harvesting on bat activity found increased bat activity at timber harvest edges in relation

to harvest and forest interiors (Caldwell et al. (2019). Out of the six bat species studied, Eastern

red bats and hoary bats seemed to benefit most from the creation of forest openings based on

activity levels, though the other four species did not differ significantly in their use of harvest

and control areas. Small-scale clear-cuts provide the necessary edge habitat within large

contiguous forests to suit a wide variety of small and large-bodied bat species. Bat biodiversity

benefits from forest management practices that use a combination of silviculture techniques to

provide edge habitat for forage and maintain adjacent forest stands for roosting.
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Forest songbird nests in fragmented habitats tend to have greater brood parasitism and

predation rates than nests located in contiguous forests. A study on brown-headed cowbird

parasitism of Swainson’s warbler nests found that the probability of brood parasitism was

significantly greater near forest edges and decreased as percent forest cover and understory

vegetation height increased (Benson et al. (2010). An artificial nest predation experiment in

boreal forests found that nest predation rates in habitat corridors were significantly lower than

nest predation rates in clear-cuts, forest edges, and forest interiors (Huhta et al. (2015). The

probability of nest predation similarly decreased as vegetation cover increased. Dense understory

development through canopy tree thinning and habitat corridor creation are both vital to

concealing ground-level songbird nests in fragmented habitats from parasites and predators.

Sustainable silvicultural practices can reset succession in a manner that benefits small

mammal populations. Gasperini et al. (2016) found that coppicing activities of deciduous tree

species had strongly positive effects on the population density of three ground-dwelling rodent

species: A. falvicollis, A. sylvaticus, and M. glareolus. The individual survival rate for generalists

A. falvicollis and A. sylvaticus was also higher in coppiced stands than late successional stands,

though the individual survival rate for specialist M. glareolus was not significantly different

between stands. Coppicing enhances the carrying capacity for rodents in managed stands through

increases in forage and cover availability that mature forests tend to lack. Rotational coppicing of

deciduous tree stands makes habitat suitable for generalist species that best compete in disturbed

forests and for specialist species that best compete in late successional forests.

The aforementioned studies address the effects of silvicultural practices, vegetation

cover, and proximity to the forest edge on mammal activity patterns and avian nest predation. In

this field experiment, our objectives were to evaluate bird and small mammal feeding activity in
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Stockton University’s Forest Management Area and to compare bait removal across forest

management units with different vegetation conditions and proximities to the forest edge. We

sampled vegetation transects, estimated cover plots, and monitored bait stations in clear-cut,

control, thinned, and lowland units. We then performed appropriate statistical analyses to

determine if bait removal was significantly different between distances to the edge along

transects, forest management units, and vegetation conditions. Overall, the purpose of our study

is to compare depredation rates between areas where anthropogenic forest management practices

have altered vegetation conditions and created an edge effect. Our study has major implications

for future nature reserve designs and forest management schemes that protect biodiversity in

habitat fragments as the human population further expands.

Methodology

Site Description

This experiment was completed at Stockton University’s Main Campus in Galloway,

New Jersey. Galloway Township is part of Atlantic County in southern New Jersey. Stockton

University’s Main Campus is situated on 1,600 acres of the Pinelands National Reserve, which is

a sizeable protected swath of the Atlantic Coastal Pine Barrens ecoregion. Based on the 1981-

2010 Climate Normals data provided by the NCEI (n.d.), the mean annual temperature for our

site location is 12.4oC and the mean annual precipitation is 106.05cm. Vegetation cover plots

were sampled and bait stations were monitored in clear-cut upland (burned and unburned),

control upland (burned and unburned), thinned upland (burned and unburned), and unmanaged

lowland units of Stockton’s Forest Management Area. In the upland units of Stockton’s Forest
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Management Area, the primary overstory is dominated by oak-pine and the primary understory is

dominated by Ericaceae spp. In the lowland units of Stockton’s Forest Management Area, the

primary overstory is dominated by either black gum-pine or Atlantic white cedar-red maple and

the primary understory is dominated by Mountain laurel-Ericaceae.

Transect 10 was located in the unmanaged lowland unit of Stockton’s Forest

Management Area adjacent to Lake Pam. The primary overstory was dominated by black gum

with DBH < 10cm and pitch pine with DBH > 10cm for plots 10A-10D. The primary understory

was sparsely dominated by Mountain laurel, Ericaceae, and greenbrier for plots 10A-10C. The

soil in plots 10A-10C had a high moisture content and was rich in organic matter. A transition

from a lowland to upland landscape occurred at plot 10D. The primary overstory shifted to oak-

pine (DBH > 10cm) dominance and the primary understory shifted to dense Ericaceae

dominance. Understory cover and elevation increased, while overstory cover and tree density

decreased with this transition. The soil in plots 10D-10F was drier, sandier, and had a more

definitive layer of leaf litter from inputs of pine needles and deciduous tree leaves.

Field Methods

In order to evaluate differences in avian and small mammal activity based on vegetation

conditions and distances to the forest edge, we sampled vegetation transects and cover plots and

monitored bait stations in clear-cut, thinned, lowland, and control units of Stockton’s Forest

Management Area. Using a tape measure, we measured 50m transects that began with the point

0m on the forest edge of one side of a man-made trail and extended into the forest. The general

location, primary overstory, and primary understory along the transect were recorded. We then
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recorded the plant species and height category (where: (0) = no vegetation, (1) = <1 ft., (2) = 1-2

ft., and (3) = >2 ft.) of the understory vegetation at each meter along the transect. Vegetation

conditions were measured for an additional 5m on either end of the transect (5m across the trail

to the opposite side from point 0m and 5m deeper into the forest from point 50m). We set up 1

m2 quadrat plots on a 10m interval beginning with the point 0m to estimate percent understory

cover and percent canopy cover. A total of 6 quadrat plots were created along each transect and

marked with flags labeled A-F. An outer tree plot was created by recording the number of oaks,

pine, and other species (DBH > 10cm) within 5m of either side of the quadrat plots. Bait stations

were set up by placing one rodent bait block and one boat of bird seed at each quadrat plot (10m

apart). Baits were collected one week later and we recorded whether the baits were removed,

nibbled, or untouched for each quadrat plot. Students sampled a total of 10 vegetation transects

across various units of Stockton’s Forest Management Area and monitored a total of 60 bait

stations.

Data Analysis

After one week, each bait was assigned as depredated (1) if the bait was removed or not

depredated (0) if the bait had been left. Based on our vegetation transect measurements, we

converted the treatment variable of tree density into a categorical variable, where: (0) = 0 trees,

(1) = 1-5 trees, (2) = 6-10 trees, (3) = 11-15 trees, (4) = 16-20 trees, and (5) = >20 trees. Based

on our cover plot estimates, we converted the treatment variables of canopy cover and ground

cover into a categorical variable, where: (0) = 0% cover, (1) = 1-10% cover, (2) = 11-25% cover,

(3) = 26-50% cover, (4) = 51-75% cover, and (5) = >75% cover. Using a pivot table, we

determined how many baits blocks were depredated for each category of the various treatments.
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We then calculated the observed percentage depredated and the expected values for each

category. Using the observed and expected depredation values, we calculated the Chi-sq statistic

for each category and calculated the sum of these values to gain the total Chi-sq statistic for each

of the six treatments (distance to edge, management unit, vegetation height, tree density, %

ground cover, and % canopy cover). The total Chi-sq statistic for each treatment was then

compared to the critical value corresponding to the degrees of freedom in each calculation (df =

number of categories - 1) to determine if statistically significant differences exist between

depredation rates across categories of the treatment. All of our data analysis was conducted using

Microsoft Excel.

Results

We found a total depredation rate of 78.3% in our experiment, as 47 baits were removed

from the 60 stations we monitored. Since the total χ² statistic was less than the critical value for

all six treatments, we must retain the null hypothesis that depredation rates were the same across

categories of each of the treatments (Figure 1).

Statistically significant differences in depredation rates did not exist between distances

along the transects to the forest edge (χ² = 1.13, df = 5, Figure 1). However, forest edges were the

only transect point observed to have a 100.0% depredation rate in this experiment and

depredation rate tended to decrease with increased distance to the forest edge (Figure 2). There

were also no statistically significant differences in depredation rates between clear-cut, control,

thinned, and lowland management units in this experiment (χ² = 0.40, df = 3, Figure 1).

However, we observed a 100.0% depredation rate in only the lowland unit. The depredation rate
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was slightly higher in the thinned units (76.7%) than either the clear-cut or control units (75.0%,

Figure 3).

We found no statistically significant differences in depredation rates across areas of

different vegetation height (χ² = 0.27, df = 2, Figure 1). Depredation rates were the highest in

areas where the average vegetation height was >2 ft. (90.0%) and <1 ft. (78.9%), whereas areas

with an average vegetation height that fell between these categories had the lowest depredation

rate of 73.3% (Figure 4). Depredation rates were not significantly different between areas of

different tree density (χ² = 0.18, df = 3, Figure 1). In general, depredation rate increased with

increased tree density, ranging from 72.7% in plots with no trees to 100.0% in plots with 11-15

trees (Figure 5). We also found no statistically significant differences in depredation rates among

plots with different ground cover percentages (χ² = 1.33, df = 5, Figure 1) or different canopy

cover percentages (χ² = 2.50, df = 5, Figure 1). However, plots with 0-25% ground cover had a

100.0% depredation rate in this experiment and plots in all categories with >26% ground cover

had considerably lower depredation rates of 69.2-75.0% (Figure 6). In contrast, plots in all

categories with >26% canopy cover had the highest depredation rates (85.7-100.0%) and

depredation rates were also relatively high in plots with no canopy cover (71.4%) compared to

plots in categories with 1-25% canopy cover (33.3-40.0%, Figure 7).

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Treatment χ² df Critical Value

Distance to Edge 1.13 5 11.1

Management Unit 0.40 3 7.8

Vegetation Height 0.27 2 6.0

Tree Density 0.18 3 7.8

% Ground Cover 1.33 5 11.1

% Canopy Cover 2.50 5 11.1

Figure 1. Total Chi-sq statistic values for each treatment variable in Stockton’s Forest
Management Area, along with corresponding degrees of freedom and critical values used for
statistical significance determination.

100.0%
90.0%
Observed Depredation Rate

80.0%
70.0%
60.0%
50.0%
40.0%
30.0%
20.0%
10.0%
0.0%
0 10 20 30 40 50
Distance to Edge (m)

Figure 2. Relationship between observed rate of depredation in Stockton’s Forest Management

Area and distance to the forest edge based on combined data from 10 transects. Line of best fit
(dashed) given by the equation y = -0.0037x + 0.8762; shows a weakly negative association (R2
= 0.2733) between bait removal percentage and increasing distance to the forest edge along 50m
transects.
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100.0%
90.0%

Observed Depredation Rate

80.0%
70.0%
60.0%
50.0%
40.0%
30.0%
20.0%
10.0%
0.0%
Clear-cut Control Thinned Lowland
Management Unit

Figure 3. Observed depredation rates in Stockton’s Forest Management Area based on

management unit.

100.0%
90.0%
Observed Depredation Rate

80.0%
70.0%
60.0%
50.0%
40.0%
30.0%
20.0%
10.0%
0.0%
<1 ft. 1-2 ft. >2 ft.
Average Vegetation Height

Figure 4. Observed depredation rates in Stockton’s Forest Management Area based on average
understory vegetation height (feet) by category. Data from Plot 5C was incomplete and has thus
been omitted from this calculation.
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100.0%
90.0%

Observed Depredation Rate

80.0%
70.0%
60.0%
50.0%
40.0%
30.0%
20.0%
10.0%
0.0%
0 1-5 6-10 11-15
Tree Density (number of trees/outer plot)

Figure 5. Observed depredation rates in Stockton’s Forest Management Area based on tree
density (number of trees with DBH > 10cm per outer plot) by category.

100.0%
90.0%
Observed Depredation Rate

80.0%
70.0%
60.0%
50.0%
40.0%
30.0%
20.0%
10.0%
0.0%
0% 1-10% 11-25% 26-50% 51-75% >75%
Ground Cover

Figure 6. Observed depredation rates in Stockton’s Forest Management Area based on ground
cover percentage by category.
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100.0%
90.0%

Observed Depredation Rate

80.0%
70.0%
60.0%
50.0%
40.0%
30.0%
20.0%
10.0%
0.0%
0% 1-10% 11-25% 26-50% 51-75% >75%
Canopy Cover

Figure 7. Observed depredation rates in Stockton’s Forest Management Area based on canopy
cover percentage by category.

Discussion

In this experiment, we observed a general trend of greater bird and small mammal

foraging activity at the forest edge bait stations of our transects in comparison to the forest

interior bait stations (Figure 2). Caldwell et al. (2019) found a similar overall trend of elevated

mammal feeding activity at forest edges in comparison to clear-cut interiors and control forested

interiors. Benson et al. (2010) found that brood parasitism rates of song bird nests were

significantly greater near the forest edge, though we found no significant differences in

depredation rates based on distance to the edge (Figure 1). In our study and the one conducted by

Benson et al. (2010), depredation rates and brood parasitism rates tended to decrease with

increased distance to the forest edge, respectively (Figure 2). The elevated animal activity along

the forest edge is likely the result of the favorable foraging conditions created by the abrupt

landscape transition. Edge habitats create distinct boundaries between adjacent communities
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where species may move between communities or filter into the edge habitat from one of the

communities. Ecotones are capable of supporting a large number of mammal and bird species

from each adjacent habitat and also species adapted to edge habitats (Relyea and Ricklefs 2018).

The edge effect creates a greater resource availability per unit area than either adjacent habitat

contains independently and ecotones are thus desirable foraging areas for many rodent and bird

species (Bolen and Robinson 2003). Our observations support the notion that clear-cutting small

patches within contiguous forests could enhance mammal feeding opportunities at forest edges to

some degree, while still maintaining forested habitat vital for shelter and songbird nest

concealment (Benson et al. (2010); Caldwell et al. (2019).

Our experiment evaluated differences in depredation rates between forest management

units. We found no significant differences in depredation rates between clear-cut, control,

thinned, and lowland units (Figure 3). Similarly, Huhta et al. (2015) found no significant

differences in nest predation rates between clear-cuts, forest interiors, and forest edges. Clear-cut

and control units in our study had identical depredation rates of 75.0% (Figure 3). Similarly,

Caldwell et al. (2019) found that four of the six bat species in their study did not differ

significantly in their use of clear-cut and control areas. This suggests that some generalist

mammal and predatory bird species may frequently cross forest edges and can equally utilize

clear-cut and forested areas for foraging. However, it should be noted that Gasperini et al. (2016)

found that generalist rodent A. sylvaticus preferred recently coppiced stands to control areas, as

foraging opportunities were more plentiful. The bait monitoring period and limited number of

bait stations in our experiment may account for the lack of significant differences in depredation

rates observed across forest management units. Within the week-long time frame between setting

and monitoring bait stations, rodents or birds could have easily ventured outside of their home
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range foraging areas and discovered the bait stations. Additionally, the 60 bait stations we

created may not have been enough to accurately reflect animal activity patterns in different

treatment units of Stockton’s Forest Management Area.

In our experiment, we also observed the effects of vegetation conditions on small

mammal and avian feeding activity. We found a trend of decreased depredation rates in areas

with >26% ground cover, though no significant differences existed in depredation rates across

ground cover categories (Figures 6 and 1). Huhta et al. (2015) similarly observed a decrease in

nest predation as vegetation cover increased. Benson et al. (2010) found that brood parasitism

rates tended to decrease with increased vegetation height and forest cover. In contrast, our study

found that depredation rates were typically the highest in areas where the average understory

vegetation height was > 2ft. and in areas where the canopy cover was estimated to be >26 %,

though no significant differences existed between categories for either treatment (Figures 4, 7,

and 1). This finding is somewhat contradictory to what we would have expected, as the closed

canopy of climax forests often hinders the development of understory vegetation that would

provide food and shelter for ground-dwelling species (Bolen and Robinson 2003). Birds and

small mammals may have favored foraging in areas of high canopy cover and tall vegetation in

our study because such areas provide concealment from predators while feeding. Our

observation that depredation rates tended to increase with increased tree density supports the

notion that tree thinning may benefit songbird populations by enhancing the development of

understory vegetation and thus nest concealment from predators (Figure 5; Benson et al. (2010).

However, we found no significant differences in depredation rates across tree density categories

and the observed rate of depredation was actually 1.7% higher in thinned units than clear-cut and

control units (Figures 1 and 3). Regional ecological differences between study sites may account
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for the discrepancies between our observations of animal activity based on vegetation conditions

and those of previous studies. It should be noted that our experiment was concentrated in upland

pine-oak forests of the northeastern U.S., which is a region very ecologically different from the

lowland deciduous forests of the southeastern U.S. where the study by Benson et al. (2010) was

conducted.

The findings of this study should be considered by nature reserve designers who seek to

protect biodiversity in habitat patches. The reserve design should maximize the total area to edge

ratio, such that the influence of edge colonizers and brood parasites on native songbird

populations is minimized (Benson et al. (2010). Large reserves should contain several small-

scale clear-cuts among the forested matrix to provide habitat heterogeneity and forage

opportunities for both edge-adapted species and those suited to forest interiors (Caldwell et al.

(2019). Thinned buffer zones should be created between forest edges and interiors to enhance

understory vegetation development. This will increase the carrying capacity of the reserve by

improving shelter and food resource availability. The nests of ground-dwelling bird or rodent

species will also be better concealed from predators in areas with well-developed understories

(Benson et al. (2010). If possible, reserve designers should collaborate with land owners to create

habitat corridors that connect reserves. Habitat corridor creation will decrease the susceptibility

of populations to local extinctions and decrease patch isolation by facilitating gene flow and

population movements between habitat patches in a safe manner (Relyea and Ricklefs 2018).

Habitat corridors may also have significantly lower nest predation rates than treated forest

patches (Huhta et al. (2015). Overall, the forest management practices we studied in Stockton’s

Forest Management Area can all be applied in some capacity to sustaining biodiversity in

fragmented habitats.
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Literature Cited

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Caldwell K, Carter T, Doll J. 2019. A comparison of bat activity in a managed central

hardwood forest. The American Midland Naturalist 181(2): 225–244.

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