Schoeberlein & Haskell 1 Stephanie Schoeberlein, Piper Haskell Professor Wingert Biology 128: Field Natural History 12/4/09

The Effect of White-tailed Deer Management and Fire Suppression on the Northern Hardwood Forests of Pennsylvania

I. Abstract Deer browsing and fire suppression are two factors that are fundamentally altering the composition of Pennsylvanias forests. There is a discernible loss of plant and animal diversity as a result of these issues. Observations garnered from close examination of forested regions at State Game Lands 170, Waggoners Gap, and Route 233 and Ridge Road illuminate the detriments of anti-hunting regulations on white-tailed deer populations and the use of fire suppression as a forest management strategy. Monocultures of red maple and black cherry are abundant in a variety of regions and demonstrate environments lacking in diversity. There is also a vivid browse line in hunting prohibited locations and a propensity for the habitation of invasive plant species. Conversely, areas which permit hunting like Waggoners Gap and areas which have been scorched by fire, such as Route 233 and Ridge Road show signs of improved diversity and forest health. Primary sources of data were gathered through transects conducted at each site.

II. Table of Contents Introduction.2-5 Methods and Materials....6-8 Results..8-11 Discussion12-14 Summary/Conclusion...14 Literature Cited...15-16

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III. Introduction As a keystone species, the habits of the white-tailed deer (Ododcoileus virginiana) severely impact and alter the diversity and composition of forest ecosystems. White-tailed deer populations in the Northeastern United States are at an unprecedented high (Rossell Jr. 2005). Over the past century, deer populations in the United States have drastically risen as a result of human management. In 1906, the State Game Commission of Pennsylvania (PGC) enacted a Buck Law that protected does and juveniles from being hunted (Liscinsky 1989). Since this point in time, management has varied as deer populations have experienced cyclic bloom and crash phases (Wingert 2009). Over the past few decades however, deer populations throughout the Northeast have continued to grow as management methods have favored deer protection (PGC 2008). Similarly, deer populations have thrived as predators such as the gray wolf (Canis lupus) have been extirpated along the East coast through human efforts (Rooney 2001). Current indicators of forest health in areas within Pennsylvania, such as browse lines created by deer herbivory, indicate that white-tailed deer are significantly inflicting damage on forests and limiting wildlife diversity. The understories of Pennsylvanias forests are now dominated by red maple and black cherry seedlings. This observation points to a major change in tree diversity over the past few centuries (Kraft 2004). Pennsylvanias forests are classified as over-mature and trees of the oak and hickory families are prominent among canopies. In light of this dichotomous relationship, future forest canopies will almost exclusively contain red maple and black cherry trees and oak, hickory, and other species will cease to exist in Pennsylvanian woodlands (Long 2007). This shift is largely due to the selective eating habits of the white-tailed deer; red maple and black cherry trees are unpalatable to deer and thus maintain higher rates of survival in spite of heavy deer browsing (Liscinsky 1989). As deer populations increase and approach their

Schoeberlein & Haskell 3 carrying capacity, the damage they exact can be expected to proportionately increase (Miller 2009). Additionally, as forest diversity is diminished, animal diversity will likewise decrease and conditions suited for the growth of invasive plant species will result (Kraft 2004). The greatest projected impact of present deer population management will be a radical restructuring of forest environments throughout Pennsylvania (Rooney 2001). Pennsylvanias deer management program incorporates five goals, all of which have been identified by public consensus (PGC 2008). First, the state assesses the health of local deer herds through reproduction and pregnancy rates. Typically, they calculate the ratio of embryos per doe; a great deal of this data is collected from female carrion and extracted fetuses. A second goal pertains to the healthy management of regional forests. Evaluating and analyzing the existence and success of tree regeneration is crucial in achieving this objective. The Pennsylvania State Game Commission also looks at acceptable levels of human-deer conflict this primarily includes deer/vehicle collisions (DVCs) and agricultural damage sustained by daily deer activity. The introduction of Citizen Advisory Committees (CACs) has aided in illuminating the extent of current human-deer interactions in numerous counties and communities of Pennsylvania. The final goals of Pennsylvanias deer management program involve providing the public with a wealth of information about the influence of deer on forests and providing sufficient recreational opportunities for hunters, birdwatchers, and other wildlife enthusiasts (PGC 2008). In the white-tailed deer investigation, the effects of deer browsing are examined and compared in two locations. Questions that are being evaluated include: How does the presence of hunting affect the existence of deer browse lines? What are the detriments of a perceptible browse line? How does tree composition and forest diversity differ in areas where hunting is

Schoeberlein & Haskell 4 permitted versus areas where it is prohibited? What are the estimated consequences of current deer management? The debate on whether to let wildfires burn or to use fire suppression as a forest management technique has been raging ever since natural fires have caused problems for civilization. One option is to suppress the expansive fires, which is based on the belief that they only cause damage and no good comes from them. The other alternative has been demonstrated since the time of the Native Americans who absolutely depended on nature, employed fire for a variety of purposes, including increasing the abundance and vigor of plants used for food and forage (Arno 2002). If no action is taken, it departs radically from natural processes and is likely to degrade the forest as a habitat for the historical native plants and animals (Arno 2002). With the progress in science and research methods that have developed over the years, many studies have been performed on this topic bringing new information to light. In some of these forests, attempts to remove fire have led to overly dense, stagnant tree growth, which ultimately leads to fires of extreme severity that are dangerous and costly, if not impossible, to control (Arno 2002). This knowledge of wildfires that has been uncovered in recent years has given us the technical knowledge to manage forests to reduce wildfire hazards and to restore many of the ecological benefits associated with the natural fire regime (Arno 2002). The answer is now clear that fires are an important component of multiple ecosystems and forest values can be damaged as a result of our attempts to exclude fire (Arno 2002). The red maple population has exploded in number and the deliberate suppression of forest fire during this century has, consequently, allowed for th[is] increase in many eastern forests as a later successional species (Abrams 1998). Another key survival adaptation of the red maple occurs as it is a later successional species, [and] can increase in many forest types in

Schoeberlein & Haskell 5 which periodic fire has been suppressed (Abrams 1998). The current fire management technique in Pennsylvania is to suppress forest fires (Grace 2009). Two questions posed in the examination of Pennsylvanian fire suppression methodology include: Are ground fires, surface fires, or crown fires the most beneficial? Which trees are fire dependent and which are fire independent? Several hypotheses and questions regarding the studies on deer management and fire suppression include: 1. As the number of white-tailed deer increases in forest environments, the number of black cherry and red maple trees will increase.
2. As the number of white-tailed deer increases, the diversity of the forest will be reduced. 3. Without fire suppression, the diversity of the forest will be enhanced as natural

management processes are active.

4. Without fire suppression, regenerative and fire dependent tree species such as oak and

pitch pine will thrive.

5. With fire suppression, the number of black cherry and red maple trees will maintain

dominancy in forest understories.

6. The suppression of fire, along with an increase of white-tailed deer will increase the

population of red maple trees in Pennsylvanian forests.

7. How can we go about fixing the adverse relationship between Pennsylvania white-tailed

deer populations and its forests? IV. Methods and Materials

Schoeberlein & Haskell 6 For the investigations into white-tailed deer management and fire suppression in Pennsylvanian forests, two separate types of controlled experiments were conducted; transects and spot boarding. Collectively, four transects were completed at State Game Lands 170 on October 5, 2009 and October 6, 2009. Four sets of spot boarding results were recorded between these dates as well. Similarly, four transects were conducted and four spot boarding results were recorded at Waggoners Gap in Reineman Sanctuary between October 12, 2009 and October 13, 2009. Both of these locations reside in Perry County, Pennsylvania and are part of the North side of Pennsylvanias Kittatinny Ridge. Additionally, four transects were conducted at Route 233 and Ridge Road in Cumberland County, Pennsylvania between October 26, 2009 and October 27, 2009; two transects were done in areas that were burned from a forest fire in the year 2000 and two transects were conducted in areas just outside these burn zones. Each individual transect at each location was performed within an area of 200m2. During this experiment, a runner would start out by walking the tape measure until it was extended to the full distance (200 m2), taking care that his line of travel was as straight and direct as possible. Professor Harold Wingert would then follow the tape measure, naming seedling, sapling, pole, and canopy trees that stood within an arm lengths distance from the tape measure. Recorders copied down Professor Wingerts observations by tallying the number of different tree species and their phase of growth on a chart. While this investigation was taking place, other recorders would measure the Dbh (Diameter at breast height) of canopy trees that Professor Wingert identified, using diameter tape measures. The spot board recordings, which were taken at State Game Lands 170 and Waggoners Gap, were part of a separate study. They were collected by a second group of students and done simultaneous to the transects under Professor Wingert's direction. During this procedure, the group was to find a starting point with relatively flat topography for about 100m within an

Schoeberlein & Haskell 7 approximate 35 degree arc (Wingert 2009). A runner would walk the tape measure out at ten meter intervals, while a stationary sighter and recorder remain[ed] in one place throughout the duration of the sightings (Wingert 2009). A spot board carrier would accompany the runner, transporting a large yellow board marked with twelve equally spaced red spots. The carrier had to hold the board perpendicular to the line of sight from the sighter. The board [was to] be held at the same height from the ground. This [was] accomplished by allowing the weighted line to just touch the ground at each reading (Wingert 2009). At each interval of distance, the sighter would indicate how many whole, partial, or completely obstructed spots were present on the board. The recorder would detail the results in a sighting table. As the runner and spot board carrier moved away from the sighter at ten meter intervals, they were also expected to shift either right or left two meters. The direction of their shifts was held constant throughout the entire process; the process would last until the sighter could no longer identify any degree of the spots on the board. After the transects and spot boarding procedures were completed, students gathered in the lab room and tallied, checked, and combined results to hand in to Professor Wingert. Through the transects, raw data was collected pertaining to the diversity and abundance of tree species and populations in three locations of Pennsylvanias forests. With the data gathered at Route 233 and Ridge Road, evidence of fire-dependent, fire resistant, and fire-susceptible tree species was discovered; the results compared tree diversity in burned and unburned zones of the same forest. Lastly, information was gleaned about the impact and presence of deer browse lines and understory density with the aid of the spot boarding records. For the survey portion of the investigation, a phone interview was conducted with an active hunter. Bernard Haskell was contacted on November 15, 2009 and was asked various

Schoeberlein & Haskell 8 relevant questions about hunting in Vermont. He answered six questions about the forest in which he hunts and about his observations on the deer population and management in Vermont.

V. Results

Figure 1. Transects conducted at State Game Lands 170, detailing a heavy browse line. [Multiply number of red maple trees by 25]

Figure 2. Transects conducted at Waggoners Gap Hawk-Watch, detailing the absence of a perceptible browse line. [Multiply red maple and chestnut oak by two]

TABLE I. Unburned vs. Burned Tree Species at Route 233 and Ridge Road Transect

Species Pine, Pitch Cherry, Black Gum, Black Maple, Red Oak, Chestnut Oak, Black Oak, Red Sassafras

Unburn Burne ed d 42 433 29 43 131 27 4 48 14

36 12 6 19 10 173

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Figure 3. Ratio of Unburned Species from transects at Route 233 and Ridge Road.

Figure 4. Ratio of Burned Species from transects at Route 233 and Ridge Road.

TABLE II. Contingency Table and calculated Degrees of Freedom

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Seedli ng State Game Lands 170 Frequency of Occurrence (Expected) Waggoner's Gap Frequency of Occurrence (Expected) Totals 18 296.9 7 552 273.0 3 570

Saplin g 612 332.4 26 305.6 638

Pole 15 13.02 10 11.98 25

Canop y 25 27.6 28 25.39 53

Totals 670

616

1286

P<0.05

2= 1039.22; DF=7

Interview Interviewer: Piper Haskell Interviewee: Bernard Haskell, hunter in Norwich, Vermont 1. Do you approve of deer management in forests? Absolutely, it is necessary if we want to have healthy forests and healthy and abundant herds. Deer hunting is big sport here and it makes up a big and lucrative industry.
2. Have you seen a change in the forest over the past few decades as a consequence of

rising deer populations? In Vermont, we actually havent seen as many deer in recent years. The turkey population has sky-rocketed thoughmaybe because they share food?
3. Is there a lot of undergrowth in forests? Or are there discernible absences in

seedling populations? In most forests around Norwich, there is plentiful undergrowth. I dont see any apparent browse lines typically when I hunt throughout the area. 4. What is deer management like in Vermont? It doesnt seem as if there is as much damage from deer populations in Vermont as compared to Pennsylvania. Perhaps because deer population density is lower here. Again, hunting is a big part of deer management in Vermont. Bow hunting is popular, as is black powder and rifle. Rifle hunting season begins today! Theres also a Vermont Fish and Wildlife Commission which tracks deer patterns and forest ecological health.

Schoeberlein & Haskell 11 5. What are the dominant tree species in your area? Mostly hardwood speciesBeeches, Ashes, Maples. 6. How often do you hunt? I go on hunting trips every 3-4 months.

VI. Discussion The results gathered from the various investigations support claims stated in the introduction. The third hypothesis regarding the correlation between increased hunting and improved forest diversity was evidenced in the recordings from the transects conducted at State Game Lands 170. Red maple was consistently present in all four stages of growth, but not excessively so. Black birch was particularly popular in its seedling phase, reaching a count of 525. Sassafras similarly maintained a high number of seedlings and saplings, however no other species of tree yielded comparable results to the black birch. While red maple did not dominant the understory, diversity was marginal as various species of oak and hickory were still missing from the forest. This suggests perhaps, a recent recovery of forest diversity and the possibility that many of the regions native tree species were depleted by deer browsing at some point in the recent past; this agrees with Krafts observations (2004). Within the study of white-tailed deer effects on forest ecology, it is recognized that deer browsing occurs most notably in regions where hunting is prohibited. Data from transects at Waggoners Gap validate observations made by Long and Rooney that a significant restructuring of forest ecology is taking place in regions where hunting of white-tailed deer is prohibited (2007, 2001). Red maple was the prevailing species with fifteen seedlings and 105 saplings in an 800m2 area. Chestnut oak accounted for the largest concentration of saplings at 152. Surprisingly, several species of other trees including chestnut oak and hickory sp were reasonably prevalent in their sapling phase. A starker comparison

Schoeberlein & Haskell 12 occurred in the seedling and pole recordings where all counts of species aside from red maple were negligible. Additionally, three out of four species of oak were absent in the understory and were present only in the canopy of the forest. This data conveys a diminution in forest tree diversity; species originally native to Pennsylvanian forests, such as red and white oaks are now virtually non-existent, agreeing with Long (2007). Deer over-browsing seemed rampant in the area and a browse line was apparent about three to four feet high in the understory growth. The transects at Route 233 and Ridge Road detail the effects of fire suppression as a forest management technique. The burned section of the forest was overwhelmingly populated with fire dependent pitch pine at a count of 433. Sassafras remained a second and less dominant outlier with 173 trees existent in the region. In contrast, red maple was the most abundant tree in the unburned section of the transect with 131 trees; the second most popular tree was red oak at a drastically lower number of 48almost three times less than the population of red maple. Most oak tree species were also more copious in the unburned section indicating that they are highly susceptible to fire. Oak chestnut was diminished in the burned area from 27 to six trees. Similarly, red oak decreased from 48 to ten trees. After compiling collected spot boarding statistics in a contingency table, 2 was equal to 1039.22 and degrees of freedom were seven. In addition, P<0.05; it can be inferred from this P value that some force aside from chance is facilitating the disproportionate and overwhelmingly widespread existence of red maple in Pennsylvanian forests. A conjecture of deer over-browsing is a likely possibility based on above-mentioned results and observations made through other scholarly researches. Spot boarding results reinforced hypotheses from the introduction pertaining to deer over-browsing and were corroborated by the sparse understory. The interview with Bernard Haskell did not corroborate the evidence given above; however, the problem of white-tailed deer does not seem to be as overt in Vermont as it is in

Schoeberlein & Haskell 13 Pennsylvania. Tree diversity also seems to be somewhat different as beeches and ashes account for a large part of tree populations in Vermont forests and are not as predominant in Pennsylvanian forests as red maple and black cherry.

VII. Summary/Conclusion Many of Pennsylvanias forests are losing diversity as unchecked populations of deer decimate understories and create cascading negative impacts on various other wildlife species that share the same environments. A ban on white-tailed deer hunting in certain areas further contributes to the rapidity and severity of the loss of diversity in forests. An inadequately informed public also counteracts the success of proper deer and forest management as important environmental advocates such as Gary Alt are replaced in positions of authority. Fire suppression illustrates similar effects on diversity; human management of natural fires inhibits forest growth and regeneration. Through investigation of white-tailed deer impacts and fire suppression in Pennsylvanian forests, it is evident that three hypotheses carry great relevancy and veracity. The suppression of fire, along with an increase of white-tailed deer will increase the population of red maple trees in Pennsylvanian forests. Secondly, as the number of white-tailed deer increases in forest environments, the number of black cherry and red maple trees will increase. And lastly, without fire suppression, the diversity of the forest will be enhanced as natural management processes are active. Observations made in these investigations will allow the Pennsylvania Game Commission to assess existing forest management techniques and shed light on projected consequences of the continuation of present methodology. It is lucidly evinced through gathered research that fire suppression and excessive populations of white-tailed deer are catastrophic to the health, diversity, and future of Pennsylvanian forests.

Schoeberlein & Haskell 14 VIII. Literature Cited Arno, Stephen F, and Steven Allison-Brunnell. 2002. Flames in Our Forest. Pages 6, 7, 17, 135, 139. Washington: Island. Print. Grace, James R, dir. 2009. Forest Fire Protection. State Forest Resource Management Plan. Bureau of Forestry. Web. 16 Nov. 2009. <http://www.dcnr.state.pa.us/forestry/sfrmp/foresthealth.htm#ffm>. This website outlines the current white-tailed deer management and forest fire protection techniques employed by PGC. It also gives the history and reasoning behind their stated goals for forestry management. Kraft, Lidia Szabo, Thomas R Crow, and David S Buckley. Effects of harvesting and deer browsing on attributes of understory plants in northern hardwood forests, Upper Michigan, USA. Forest Ecology and Management 199.2-3 (2004): 219-230. ScienceDirect. Web. 1 Dec. 2009. Liscinsky, Steve. How it was: Deer Management in Pennsylvania. N.p.: Pennsylvania Game Commission, n.d. N. pag. Print. Long, Zachary T, Thomas H Pendergast, IV, and Walter P Carson. The impact of deer on relationships between tree growth and mortality in an old-growth beech-maple forest. Forest Ecology and Management 252.1-3 (2007): 230-238. ScienceDirect. Web. 1 Dec. 2009. Miller, Brad F, Tyler A Campbell, and Benjamin R Laseter. White-tailed deer herbivory and timber harvesting rates: Implications for regeneration success. Forest Ecology and Management 258.7 (2009): 1067-1072.

Schoeberlein & Haskell 15 ScienceDirect. Web. 1 Dec. 2009. Pennsylvania's Deer Management Goals: Measuring Progress. N.p.: PGC, 2008. N. pag. Pennsylvania Game Commission. Web. 1 Dec. 2009. <http://www.pgc.state.pa.us/pgc/lib/pgc/deer/pdf/ 2_measuring_progress.pdf>. This source educates readers on basic white-tailed deer management strategies and goals in the state of Pennsylvania. It is informative and relatively detailed in pamphlet format. Rooney, Thomas P, and Donald M Waller. Direct and indirect effects of white-tailed deer in forest ecosystems. Forest Ecology and Management 181.1-2 (2003): 165-176. ScienceDirect. Web. 1 Dec. 2009. Rossell, C Reed, Jr, Bryan Gorsira, and Steven Patch. Effects of white-tailed deer on vegetation structure and woody seedling composition in three forest types on the Piedmont Plateau. Forest Ecology and Management 210.1-3 (2005): 415-424. ScienceDirect. Web. 1 Dec. 2009.